O

■.fj,iU'

DEGREE
STUDENTS
BRY< YTA
CAPSVLC^-\(\, '-\:
moiucRE

INVOLUCRi- ANTytRim

B. R. VASHISHTA
 BOTANY
[For Degree Students]

PART III

COr
BRYOPHYTA

^oTvg^p'^^

By
B. R. VASHISHTA,
Header, BIOLOGY STUDY 6B0UP,DepaHeMal of Botany.
Panjab Vniveraity, Chandigarh

Fourth Edition

W^fc

1970

C0
S. C H A N D ^ i,ucKNow
DELHI — NEW DELHI "
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•— Maouas PREFACE TO THE FOURTH EDITION
Ssiltnn Razei*
— HVDRUAnAD 4L
Exhibition Road
^ Patna The third edition of the book was published in January, 1968.
All the copies \vere sold oiit by September, 1969. The encouraging
response from the kind readers and the continued demand for the
book has compelled the author to make the new edition more
comprehensive.
With this point in view the text has been extensively revised
BxtheS^Author for B S. students in the light of latest research in the subject. An attempt has also been
made to make it still more understandable. Some parts been
expanded and the life history of Notothylas has been added. A feu
Text Book of Botany, Part I_Algae Us. new illustrations have been included.
TextBookorBota„y,Partii_Fl^;. 8-00 It is a great pleasure to record my deep sense of gratitude
Prof. R.S. Chopra for his kind suggestions to improve the book.
to

7-50
B.R. Yashishta
Chandigarh
•Etr«{ £!dtlion
19G3
Second Fdiiion
•Phird Edition me PREFACE TO THE FIRST EDITION
1968
fourth Edition 1970
Part III of the series deals with a small group of primitive,
thalloid plants, amphibious in habit. They are placed in the
- ■ "4- iSn Bryophyia. The bryophyles are peculiar m having the
""g'pur College Llbmry ^ametophy^te as gr^ and independent and sporophyte wholly or
Sanfally depende.it upon it. Th e subject matter has been presented
fnThe"simple, k.cKl style. The obscure literary style has be^n avo.ded^
scheme followed is the same as in Par s I and II. An attempt
Price Rs. 8-00 hi been made to include the latest and the significant results of
LoLnkaUesearch on the subject. The type system ts st.ll cotts.deretl
the best for the undergraduate.
A simple
function and a clear
and relationships account
of the of structure,
important members reproduction,
of each class
Acc.No,.lW.H i^^iniab^ of She book. A diagrammatic representaUon
of the life cycle of each type gives a clear picture of the pattern
Date of Alternation of Generations. Salient features of the division
^CallNo. Mft5 and its classes down to the orders and the type plants whose life
histories have been discussed appear at the end of each respective
chapter.
^'uhlisJie^
fished Iby s ,u
Ra-ni Eogar "w^"' and
xVeti, Delhi.55^^ y whose origin is obscure even up to
ment. Chapter III deals with the structure and the various
 . (vi)
•^^eones which att
f°"f thallus. In'r^' "P'ain ,1,. ■

LIST OF CONTENTS
CHAPTEll I

are r. am P^'^l'shcrs of fl due Pn.«

Gnmotophyto generation,^7°PMes
&porophyto ns amphibians
genoration, of tho Kingdom
Alternation Plantao:
of Generations-
Alternation of chromosomo number in tlio life cycle ; Origin of AlternatioA
of generations ; Salient fentnrcs ; DifTerenees between the thallophytes and
of the bryophy^s ; Resemblances between the biyophytes and the algae-
Classification ; Rsvision questions. Pages 1—17
V-P.
for Dube?" go ,0
TV, 7^ criticaii^ niy p^i. '"" CHAPTER II

P^'^on 'oi-';;,^ D^'Partmen, • * Hepaticopsida

cient land dwoliors; (Hapaticae)
Plant Body;or Habitat;
LiverwortsDistribution;
: Liverworts Vegetative
as ineffi
roprodiiction ; Chief means of porennation ; Sexual reproduction; Distinctive
'he Ir^'^hes.
yea,.'.'"C'hd
'7'-'"'hor aC tl" P°^«b'e""=J^-""scrip
not^ve^oP ^ '''^ teaturcs ; Classification ; Revision questions. Pages 18—27
CHAPTER III
Hepaticopsida {Hepati-cae)-Marchantiales : Morphologv of the
Depar'to'"" ^""e' Head" "rbt'o/^'P""^'We
Bo4 h'gtUude "for^o gametophyto (external features and internal strncture of the thallusi • Origin
of the Mai^hnntiaceoiis, Thallus, Sporogonium ; Distinctive feofcnres- Ciassi-
fication ; Revision questions. !>«««« 9s or.
CHAPTER IV 1 ages 28-30
Marchantiales-Ricciaccae: Riccia: General oharacter.s j Syste-
niatic position: Distribution and Habitat, Gametophyto phase (External
pSif5'5'^"^S
^apur tk ^^'^^^cuIarJv c • ly t ^^urao-em^^ ^PartmorvT
features. Internal structure, Ongm of air chambers. Apical growth) ;
Dichotomy; Vegetative roproduotion, Sexual reproduction; Sporophyte
phase, Sporogonesis : Production of Meiospores ; Comparison betwee n the two
, generations; Summary of the life cycle: Importait features ; Is Riccia
'"''»p'ets{.i4:?/e,',s;~ 7"C2';S- primitive or advanced ; Revision questions.
CHAPTER V
Pages 37—O")

los-i " "'e faciii^/j^ Marchantiale^Marchantiaceae : Marcfaantia-Genoraj Characters

Systematic position ; Habitat and Distribution. External Morphologv of the
thallus. Internal struciure; Comparison with Riccia; Apical growth
Veptativo reproduction, Sexual reproduction ; Distinctions between the male
and the female thalli ; Fertilisation ; Development of Sporophyte ; Pi-ocUiction
of Meiospores ; Comparison of Marciiantia sporophyte with tliat of Riccia •
Important foaturos; Summary of tho life cycle; Is Riccia primitive or
Marclinntia ? Revksion questions. Pages 66—10.3
CHAPTER VI

V -f.
a. e Distribution and(Hepaticae)-Jungermanniales:
ance; External
Habitat; Vegetative reproduction; appear•
Sex oigans
Sporophyte, classification; Salient feature.s ; Differences with Marfhan^
tiales; Resemblance with Marcliantiales; Revision questions.
CHAPTER VII 104—112
Jungermannlales-Metzgerineae : Pelliaceae : Pcllia-. General
charactenstios; Systematic position : Habitat, nistributiou • Fvtl!,
characters; Structure of tho thallus; Apical growth • Ve<rfltnt;„l -'^'-er-iai
tion; Sexual reproducfion ; Productio? of Meiosporls
protection of Sporogonium ; Development of Sporogonium •
Dehiscer.ce of the capsule ; Dispeiml of spores GeLinaUon Tf TpoTe^^
Salient features ; Summary of the hfe cycle ; Revision questions. "' '
Pages 113—1,30
CHAPTER Vin
Jungermanniales—Jungermannineae ; Madothecaceae or Pa».<.11
ceae Mado heca or Pcrella ; General characteristics ; Systematic position •
 .r

'• i •■'n .i -

Internal structure ;
Salient features '• ^°*'°S®"osi8 ; Dehiscenco of nn ^ ^P°'^oeo"ium ; DevoloDmont.
)nium ; D«
«>'antiales coL'ared'^T^-^
pared'
, I^ovision l1?o ^
RoviBion questions
questions. cyolo'
- l^vowivUS.
^ '^""g®rmfinniales
' I''
'^""g®'"irinnniales and Mar-
CHAPTER
CHAPTrt? IX
—

' DistribiTfln^ . = ^'^thoceroa ;

CHAPTER I
nutrition Snnct iction
""reproduction •- Profir'V"^
prm-ii '
development?- n growth- Va'
Vo* ^^^rnal
t. .^^rnal features;
^^®'°«Poro3'S^rotn ~=tion features
^^Production ;
salient'Matures; Summarfnf th ofCapsu?o A ('ts structure, ^'^TRODUCTORy
NotethXcTaf ? ^^rmmation of spore
Conci Disen«ir ^
Snor.^ g®rm,nation
spore (dovolof :
germination
Plants^r^'irtwo
the land. The pl a wellSef
nts which gtowh^b^al
in water^TheS"a?fr
ale ^dred Z ITaUcf
position''™/!™™ """ Disousslr Sorm,nation ; and the others terrestrial. The best examnlo« /aP quatics,
are the algae and of land dwdlers thT^i^d nlanT^ ' P'"""'
r^SMsion
w«on questions '^'^nocoros .Sporonhvf^ phytes). Between these two extremes of habitat? is a tra?df'""i
t>
(MuscJ
(M-ci
^production ; Qa n.
o. X P'Tges l50-_]()i' zone It IS represented by the swamps hnd tlie areas wher^ ramr
questions
and land meet. It may well be called the amphibious zone T?
-.o.tu,o habiting the amphibious zone are the mossp<5 i;, ,*
hornworts which collectively constitute a ffroun'nf ?
chapter XI
^^^PTER xt 192?TQ<j land plants called the bryophytes. The latter are simp?e"'ThaHis'
Ci8trIbu?on^VP?agn5:S Sphagnum;.., hke plants which suggest the stages through ivhich the grUn aiae
may have evolved to become terrestrial Moct
are land dweliers which inhabit danip!'-sha''d°ed a^L'^LTdlo^
hties A few of them, however, live in or float on water. The
termination ■ of't'-'iiSfs,-'.*
••-sri of viri.»i'"S.«r--
younc sLn -' p aquatic habit, of course, has been acquired by these nlants spmn
'--•-•".rsi-.tir pStS darily. When the water dries up they grow'equally ^well o? t?e
- /"ITT I
chapter xii ' -u-cologicai Spores drying mud. Some, of course, can withstand long periods of drought
Durmg the dry period they become almost brittle in texture
With the onset of rainy season the apparently dried, brittle thalH
turn green and become active to carry out the normal life fun?
tions. Even these apparently xerophytic species grow actively onlv
during the wet weather. ^

mmmm
sporophyte?'g "le sporoptv?®'"''^'""! an?? ^"'^^Ciecium"" °f
^'■'^Psintho lif, '■®»nation P?^t®3 ®f Eu^o ^'^®rentiS °P®^®uIum ;?•
Evidence supports the view that these early land olant.;
descended from alga-like ancestors which were probably ereen
Adaptation to land environment or sub-aerial life involved the dev?
lopment of certain features that were not-possessed by their anna'
tic ancestors. These are ; qua-
1. Development of organs for attachment and absorn
tion of water. Unlike algae the bryophytes which grow on la»Jri
are not bathed m water. They must absorb it from the soil H
be attached to it for support. For this purpose the brvonhv?
develop special, hair-like structures called the rhizoids L k
root hairs the rhizoids function as absorbing and attaching organs
2. Protection against desiccation. The thick com
■«- --.-s; multicellular, thallus-Uke plant body covered with an' eoid^^^''
APthooo'r„,,°SP"i«on is protected to a certain extent against the drying effects^ of*^^"^'^
Paaefe"}??> Of the numerous cells constituting it only the epidermal cells
^ages 302~-.32]
 ^ .oh SXaOH..,-H.KT In = BHVOHH.XA
INTRODTJCTOBY
3
-face, a.e directlv e- Without it the sex organs do not reach maturity and do not dehisce
"bstate- like " Water is essential for the transfer of sperms m the
The letention of swimming sperms is an algal characteristic*
The bjp'ophytes thus rely on water for the act of fertilisation Be'
^des they have inefficient absorbing organs in the form of rhizoids
Consequendy they are unable to grow during dry periods Thev
lequire sufficient moisture both for reproduction and successfffi
pores. Thev fLiUt^t ^ thallus. Thest- nr« u ""Serous vegetative growth^ This _ explains why tL bryophytes usuaUy t-
habit moist, shaded situations {amphibious habitat) or grow in places
here water is abundant at least at some season. Since the bryo-
phytes usually grow m amphibious situations and cannot complete
""trS'its.
celJular and jacketed. ItB'f?"'*
The r their hfe cycle without external water they can very appropriately
be called the amphibians of the plant world.

. The fertilized ^ sex nb.. bryophytes have

phases-gametophyte and evolved a life cycle
sporophyte. Thewhich
formercomprises two
is the most
obtains food and ^^tamed within , conspicuous of the two and independent. We start our discussion of
from drying Z h devef" '^e pare„ nl""''S°"'"m. Here the bryophytes with this phase of the life cycle.
essential fo? the surTitM ''"Co Thi^'/
be young embryo and itfnrm" i?"'' PHnts. It PP"'"" ^-^^AMETOPHYTE GENERATION

--banfe^alln^--^ ■
pumitive ^?,^y They
land dwellers. The bryophytes are a24,000
number about small group
speciesofwhich
most
are grouped under nearly 960 genera. All of them are small and
inconspicuous plants. The plant body is undoubtedly more differ
entiated than that of a complex Alga. It is compact and better
'"■ *-s;c";-"■'" '<•'■ «„ d protected against desiccation. However it is relatively simple in
the lower forms and still reminds one of the thallus ofan Alca
It grows prostrate on the ground and is thallus-like {Fig. M A—D)!
. It IS attached to the substratum by delicate, unicellular, hair-like
organs called the rhizoids.
In the higher bryophytes such as the Mosses the plant body is
//•-I erect. It has a stem-like central axis which bears leaf-like appen-
V .
1
ched, I'2A). rhizoids
multicellular It is fixed apparently
to the substratum by the
resembling means of bran
roots. The
rhizoids arise from the older, basal part of the stem. These organs
ol the bryophytes are, however, not homologous with those of the
higher plants. They lack the vascular tissue characterisdc of the
stems, leaves and roots of the higher plants. Besides, they belong to
the haploid generation whereas those of the higher plants represent
the diploid generation. The organs which are similar in function
but different m origin are said to be analogous. The stem, leaves
and rhizoids of the bryophytes are thus analogous to the stems
leaves and roots rather root hairs of the vascular plants. The root
hairs like the rhizoids of liverworts are unicellular structures In
^'8- M. contrast the rhizoids are borne upon the gametophyte generation
and root hairs on the sporophyte. The two are thus analogous to
each other. Some botanists look upon even the moss type of plant
body of the bryophytes as a highly difterentiated thallus.
The thallus bears the gametes. So it is called the gameto
phyte plant. It is concerned with sexual reproduction and consti-
 55r

4 BOTANY FOR DEGREE STUDENTS-PART III: BRYOPHYTA

tutes^the conspicuous. nu.ritionaUy independent phase in the INTRODUCTORY 5

tinues in the pteridophytes. In this respect the liverworts and

mosses seem to be akin to the pteridophytes than to the Algae to
which they resemble in their thallus-like plant body. This common
feature tempted the earlier botanists to place the bryophytes and the
pteridophytes collectively in the division Archegoniatae. They
'^NCR/dial are, however, very unlike in other features and completely unrelated
^ ^ALL to each other. Hence it is thought best to separate the Liverworts
MALE
and Mosses from the Pteridophytes on one hand and the thallus
BRA^JCH FEMALE
bearing plants (Thallophytes) on the other. They are included in a
BRANCH ^NOROCYTtS distinct group Bryophyta.
The archegonium (Fig. 1-3) is a flask-shaped organ. The slen
'Rhizoids
PLANT
der, elongated upper portion is called the neck and the lower sac
like, swollen portion, the venter. The venter is attached to and
often deeply embedded in the parent plant tissue. The neck has a
® C wall of a single layer of sterile cells which surrounds a central row
i-2 (A-C). Funariasp. of elongated, naked cells. These are called the neck canal cells.
• Gametophyto plant The neck is usually projecting or freely exposed so as to be accessible
longitudinal secfcin« to sperms. The venter also has a wall of sterile cells one or more
Sperm. ^ ^'^^^horidium. cell layers in thickness. The venter
wall encloses two cells. They
yh^SPERM
the larger egg cell or the ovum
the smaller ventral canal cell COyER CELL
above it.
phvtes f>i« above i have atf-.; , ^ Sametes
{in) Fertilization (Fig. I'S). It
devoid generally In fU
occurs when the sex organs are mature.
f
on ec.i;ruttAire
the other zi
Protoplasts of tU ffamete? ^ v^°-
^ They Moisture is essential for the maturing
■DISINTEGRATED
NECK CANAL CELLS

organ con" multicell, <=<>'15. The formed of the sex organs and also for the
movements of the sperms to the arche-
round4 the 'n om" j^^l^oted sex o '"'>'°Phytes, gonia. The mature antheridium rup NECK
tures at its apex liberating the sperms.
At the same time the axial row of neck
canal cells including the ventral canal SPERM

lar objict°*!l^'''«'«n.Thea n °°gonium o^ cell in the mature archegonium dis

organise. The tip of the archegonium
VENTER CA'/ITT
ES60R OOSPHERE

ssisFfSc S.S-.IS.'-'f.". also opens. A narrow canal opening

to the exterior is formed. It acts as a
VENTER

Srsr •'Se„?''4ifc'i''' "is'

'he biflage Lt' '''''he an4„''"'rounds
ofc ■
has a u ,,"'e
passage way to the ovum in the venter.
The liberated sperms swimming in a
thin fi lm of water reach the archegenia. Fig. 1-3. Moss Archegonium
:l They enter through the open necks and showing fertilisation.
swim down the canals of the arche-
gonia {Fig. 1'3). Reaching the venter one of them, probably the
first one to reach there, penetrates the ovum. It fuses with the
nucleus of the ovum to accomplish fertilization. With the act of
'appears fo, stfu^^^, jhe J fertilization the gametophyte generation ends and the sporophyte
generation starts. The gemetes (sperms and eggs) are the last struc
tures of the gametophyte generation.
and con-
9
 INTRODUCTORY 7
^psule. The capsule is mainly concerned in the 'production of spores
='
call
— a celi;.o:rl^f,^-- - ^f-ow Ihe latter are differentiated by meiosis and are thus haploid in

OOSPORE
epibasal cell

capsul
WALL

SUSPENSOR APlCAl CELL

pypoS^sALcea ^ a

elate. ■FOffT
ELATEEfoPHOR,

CAfiSOLE ENDOTHECIUM
SETA
AMPHI THECWAf

f .
^"sion or H^^^I •,
tbe clevel.^^P^oJitoffhl
^®'liQ sp.
^"^leus as ir .
the sporopf^ ^ gametes^° Xh '^hromatin mat ■

abortive
ARCHEGON'tUM
a^ygote
restingunderpoes^'segmentation
^hhin the v of ^th,.
^^gote of the ^Igae.
UAVES

FOOT
chance fn ^^^^'-hed. Th ^hepar«n^ ^ibtains its r, ^ trailed
which in as rnm^ of th u^^^^tophyte f ^^^'tshipent
Fig. h5. Moss embryo.

nature. The spores are non-motile and wind disseminated.

Swimming spores characteristic of the Algae are never present in the
bryophytes. With meiosis the sporophyte generation in the life cycle
terminates. The meiospores thus mark the beginning of the new
gametophyte generation. Each spore germinates under suitable
conditions to give rise to the gametophyte plant.
Sporogonium is the whole product of the sexual act. It remains
attached to the plant bearing sexual organs (oophyte). It is con
sidered a second individual in the life cycle and not simply a part
or an outgrowth of the parent plant {thallus or gametophyte)
The reasons are obvious.^ Jhe sporogonium is an individual with a
different inheritance as it is developed from the diploid zygote. It
is made up of cells containing a diploid number of chromosomes as
 8 BOTAXY for degree student^ p..
iUDENTS—PART III: BRYGPHYTA
contrasted with the hani -j -fiiYTA INTRODUCTORY 9

The other is the sporogonium. The green individual is the most

conspicuous of the two. It is an independent plant. It is haploid

■MATURE-^
■GAMETOPHYTE'^ PROTONEMA
N
^ N
Ci.

^ater
*l.08ED SpQop ANTHBRIOIOM SPORES
N
• ARCHEGONIUM
capsule wall
HAPLOID OR
^ALVRrPA
SPERMS GENERATION SPORE MOTHER
■'^fOLUCPE N N CELLS
2N

DlPLOlD OR
5P0R0PHYTE
SPOROPHYTiC
OR
GENERATION
SPOROGONIUM
2N
2N

2YG0TE
■^EMBRYO-
2N
2N

pJ;;"-""^^son the
"ot'le and thus ?■ "^^'led the sti
Fig. 1-7. Graphic representation of the life cycle of Bryopbytes.
^porophyte is depelX"""'' ^y ar*'?''''*®- Thr?®""'""! of the and bears the sex organs (antheridia and archegonia) which produce
the gametes (sperms and eggs). As it bears the gametes the green,
independent individual is called the gametophyte. It is concerned
differ'ft°'Berremarka"bl''7"^°'" ''''oUronTh"''""' "cour*^ "°"- with sexual reproduction. The gametophyte plant along with the
structures produced by it constitutes the gametophyte generation.
In the life cycle it starts with the formation of spores and consists
™«hods of fta'S"^' ■■'^Prod?ctJ™°'
fornted '^^S«'entatio„
"= as wd!'"as nonCiiu^
takes
' Bsence^'o'r ^ of the green individual and the sex organs. The last structures
gemmae, only by the the formed during this phase are the gametes The gametes fuse to
form a zygote.
•^tyophyte"^''"®*'''"" of G a5exi;al
inte?:sr"«°»a (P, , , are The zvo-ote, on germination, does not produce the gameto
phyte plant' °It undergoes segmentation to form an embryo. The
embryo by further segmentation and differentiation gives rise to the
second adult called into
usually differentiated the sporogonium. It remains
foot, seta and capsule. _ Indiploid and is ,
due course
orvvorts) &tdi,aJs^"'L'here
of time the diploid spore mother cells by meiosis give rise to
haploid spores. We may refer to them as meiospores. As the
 10 botany tor begree students-part m
. -fAKT III: BRYOPHYTA
introductory 11

the sporphyie'Th'™''' P™dac,io„ of

alternation of generations in which both the alternating generations
are structurally similar is called homologous or isomorphic.
ftro^ghlut i"t'aTta" Besides food the two most important biological needs of every
living plant species are ;—
I. Production of new individuals. Really new individuals
are produced by the fusion of male and female gametes from differ
ent plants. It is called cross-fertilization.
0,..^-Heaccou„..K.„ -P-PHfte II. Their dispersal to suitable places where they can find their
generation in the hfe rvM pfvp r; ^ food without much competition.

a%^re^ In Al-ae both these problems are solved by the agency of water.
The gametes
dispersed swimwater
by the through water 1tohisfuse.accounts
currents. The resultant
for the ygot^
absence of
doXltM?!"""func.ions.'^'Sy.ts'^
ahern'iH'ff^Jon,
alternation *0!;"«QvcIp.
:?f distinct -ege.a^:
""it
vepretatiullSj^? st=l differences between the ate^
iplohaplonti^ ,if®°««>ons and ^porogen/JP^Jj^aracterized by cycle. There is thus no division of laboui. Hence the alternatmg
by generations are similar. ^
--''p-oFS^'r'letnatio„o,^,„ . Amon- tnc
Amon^ the land dwellers such as the are
disnersal bryophytes
solved the twobasis
on the prob-
of
lems of to and is suitably adapt-
division of labour. r>?nhlems The crametophyte is concerned
the tissnp ^ ?^yheduetofhi t^evelops in^ ®\all the ed to solve one °f 'bese problems^^ ^lhe^=^
^='tos^l?"h,z^V^P®otoPhyt'e''^^^ot that th^e e"i°if
^he alternatino- ^ better Z ^'^^ation takl: ni zatior™The'spo?oph^
zation. ine sporopiiy functions
-r in the multiplication
j,,„e5 of which
jjjinute spores beneficial
are
[forphologicallv j" in the growth effects of a single sexua • labour among the two
also in theip Thev Hiff of the hrv ^^''^over distributed by the . structural difference between them
generations in ?^y^iology fnutrif* only in P^Vtes are generations thus accounts foi the structural
in the bryophytes. s./
SkltS?nation of Chromosome Number in the Life Cycle
Each cell of the
bcr of chromosonK^ in ^ jy differs in the different species,
^opported by an anlT^ to d^ ■^''"letophyte Tk^ ^onerally bol if. The numerical value
Each gamfa thus has ^
basic
^^6 sperm and
number. At the time o p maternal chromosomes, however,
egg fuse. Tte associated m pairs m the
do not fuse. They simp y ^yg^jg „ow has N plus N
•f ...Vr -'.....ir., ' '""i-<yi.;s'sf fusion nucleus. The !!i,
or 2 chromosomes, it
monoploid number.
- ig number N is called haploid or
. f^t„r.es with N chromosomes are thus
^tophyte generation may be called
~5i"» ""il ?"'" if??- Tk?"'»»« haploid. On this basis t
Thegeneration
the haploid
diploid.
zygom withm 2^j
o cycle. The 2 N number is called
nu^^ developedthus
n^ber of chromosomes fromisthea
diploid structure. ^ diploid individual m the life cycle,
zygote by repeated generation (sporogonium) up to and
All the cells of the sp^ophy
includingthesporemother e glitches on toAt the
g the time of fertili-
diploid. With
zation the haploid gener switches back to haploid. The
' lype'^i meiosis tlie diploid gene accompanied by the alternation
alternation of generations is tnus
 BOT^nr POE BEGKEE STODENTS-PART III: BRYOPHYTA . ijKYOPHYTA INTRODUCTORY 13

chrnmricn*vi« /> . _
the haploid condkbn!^'^ haploid to the diploid and back to 'Substratum by delicate, unbranched, unicellular hair-like organs
called the rhizoids. In the higher bryophytes (mosses) the plant
body is erect. It consists of a central axis which bears leaf-Jike
questi°"®TwoAtr?JTameryf„tf^^^^^ It is still a live
been proposed to explain how ahernX^oTg^eratir'''?''"^ expansions. It is attached to the substratum by branched, multi-
cellular rhizoids.
^ , Antithetic Theory
Theorv I,
T, .cc . ^generations
nerat.ons onginatcd.
originated. 4. Like the thallophytes the most conspicuous phase in the
life cycle is the gametophyte. It is independent and concerned
with sexual reproduction.
y rrogrerreTab' 5. The bryophytes like the thallophytes lack the vascular
tissue (xylem and pholem) characteristic of the higher plants.
ponse to a life in a drier'en and m 6. In sexual reproduction they show a marked advance over
oTgrar:'i!!-
origin are promnt crerr^^ <^"y>ronment. The farinrc m res- the thallophytes. It is invariably highly oogamous. The sex
organs are jacketed and multicellular. In Algae they are non-
jacketed and unicellular.
^ The
ine rresult
, is
resulf • .i,_
tc thp j
. . —'Paiuea'
by delayed
cmyea 7. Female sex organ in the form of an archegonium appears
for the first time in the bryophytes in the plant world.
creased"feS!o^rfth"^ ^ 'wit°h farmer 8. The sperms are biflagellate. Both the flagella are of
whiplash type.
Honto,„g„„3 Th
Homol„„ ' 's finally 9. Fertilization takes place in the presence of water.
10. The fertilized egg is retained within the venter of the
archegonium It neither becomes independent of the parent game-
tophyte nor passes into the resting period. In both these respects
the bryophytes differ from the algae.
n The zvgote undergoes repeated division to form an
Other Drodn On. ^ beram. of undifferentiated, multicellulav structure called the embryo. The
to dfv^ion of the aygote is trartsverse aud the apex of the embryo
developr from the outer cell. Such an embryogeny is called
exriXie It characteristic of the bryophytes. No embryo
stage exists in the thallophytes.
19 The embryo by further division and differentiation pro-
duces a relatively small spore producing structure which is not
- aU\rtr„"i^-bvin t in pri„,?,L!PrP?ty and . J rtlnt Tt i= called the
mdependent ^ sporogonium (sporophyte).
^ fe. In some the seta is Itabsent
con-
,r»l^Mnd rardy the fo^t and the seta ,«.
H The sporophyte is thus simpler than the gametophyto and
P'-'body .rhichttaU a s™,u
f-X:iFS&i t^^^r- .h,^ "■? ™"-'""
because""r-
"""
their sper»,P eoinple. ,he i, concerned will the production of wmd
is reli- Plant hod. n , ^
3 -pt switn,
habit is^XbeT'^;' di„"i»S!S-.5oiil., oudolrrd TO... «h,oh ot .1™,. jooo-
--E''Tr''s.i\?- spores or ^^e all of one kind. Hence the bryo
phytes are homosporous.
 TTcr beyophvta INTRODUCTORY 15 ;.
Sy ortdSly"afa'?atSll Resemblances between the bryophytes and the Algae :—

In Aiga^rherprSt1"L rf h "fe'^'cyde h° 1. Thallus-Jike plant body.

2. Lack of vascular tissue.
The dJff "'^''"""•ogous type bryophytes.
3. Absence of roots.

4. Conspicuous plant in the life cycle being the gametophyte.

5. Autotrophic mode of nutrition,
Bryophyta
6. Retention of the swimming habit by the sperms which
indicates the algal ancestry of the bryophytes.
terrestrial.
s,h?i^S"«~£—•I
Layers nf
feJ. "f Protonema8\a^I"®-°''''®Pting in the
7. The early stages of development in the gametophyte of
many bryophytes are green filaments which strikingly resemble the
filamentous thallus of green algae.
fcrmation^of''mif"'°P™''>"=tion
tte growing n,7Z''"'""' »mmo^n by tb?n 8. The chloroplast pigments in the vegetative cells of bryo
phytes are identical with those of green algae.
spores 'ibaence of m-*.
®=sHisC-= The above mentioned similarities give support to the view that
the Bryophytes have evolved from the algal ancestors particularly
««">Por=o i„ the green Algae.
Port^ ^®Protluce entirely•r "y
by
S|V?!5^reS\?™ -naliy meios- llassificatioii. The rank of a division Bryophyta to this well-
defined group of plants was first given by Schimper in 1879. Later
"luIticeUuJa®®^ organs ar© „i Eichler in 1883 recognised two groups, Hepaticae and the Musci
--g^tativecolir" ® ®terile veeatef-
getative cells.
by a in i
Jacket of
- 6. Seyiini in the Bryophyta. Since then these two classes remained as separate,
'"^foduetioa - major entities within the division. Engler 1892 subdivided each of
^oogamy. ^'^rough aaiso|JSy the two classes into three orders as follows :—
« invari.
organ in

ssts:=usc-,... Bryophyta

8. THa n plants only. "= r 2. Musci or Mosses

resting Class I. Hepaticae or Liverworts
^a-tio'^ni-b^o la-nation aftor
lA
atter q
9. A., Order iC March- 2.'junBor- 3. Anbho- I.^phag- 2. Andre- 2, Bryales
i »S"^bSto?SS-ioeogons0, genem*'^'^
II jj
10. ^J, "^on.
exhihu _,.
"--p antiales manniales oerotales nales ales

This system of classification with slight modifications is still

followed by many eminent bryologists.
a<'tacho?t®®Porophyte ; The anomalous position of Anthocerotales as an order of the
class Hepaticae was pointed out by many investigators very early.
The important among them were deBary, ex Janczewski (1872).
 16 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA INTRODUCTORY 17

REVISION QUESTIONS
who^ta 1899raL''Ii? Y^derwood (1894). However it was Howe
named it 'I' AnthocerCales. He 1. "The bryophytes are the amphibians of the plant world." Comment
Hepaticae, Anthocerotes and^M^sri^ upon the statement.
2. Define Alternation of Generations. Point out the differences bet
ween the Alternation of Generations met with in the algae and the
Campbell (1918, 1940), bryophytes.
his move to divide the Rrv^rlk ? • him and supported 3. List the salient features foimd in the life cycle of bryophytes.
Takhtajan and later Wardlawrfq?t^f into three classes but Smith, 4. In what respects do bryophytes, (a) differ from algae and (6) resem
Anthocerotes as Anthocerotai Schuster (1958) called the ble algae.
5. Is the sporogonium a new individual or an outgrowth of the game-
tophyte plant ? Give reasons in support of your answer.
®'yophna?s"ctTsffied aTS this view and the 6. Are the so-called leaves, stems and rhizoida of a moss plant (o)
homologous with or (6) analogous to the leaves, stems and roots
respectively of a vascular plant ? Give reasons in support of your
Sryophyta answer.

t.Hep'at™e ttattXrotao^ 3. Musci

old ones°f™^'^''"'^^1 suggested the followine -1 g taxons for the

• H«tP«lcop,Ma for Hepaticae.

three classtr thuf^ssifv f"-- ^tttho- MURS

, ^ "y Bt-yophyta into the
Hepaticopsida.
2- -^nthocerotopsida.
%opsida.
edition ^of f^^^^ification h
yia. brvr^,,v green u Of Rrt, i has proposed 3.
"■T'opsrda
1 whietTSy^s
t« furthoearedivided
t^^?^"^Bed
P'^mstYT division Chloro-
2. ^opaticidae
Anthoeerottd ° three ju) '? division as a class
' as foiiows
%idae.
 fnepRBHian

. W 2sl'- ' V, \ • r

HEPATICOPSIDA (HEPATICAE) or liverworts 19

From the foregqing account it is evident that the liverworts

nave not become completely adapted to the land life.
still in a transitional stage They are
and in the process of becom
ing adapted to land environ
CHAPTER II ments. Although the sex
organs are jacketed and thus
protected against the drying
influence of air yet the
are bryophytes i„ whiT^"®",* dwellers The r dependence of liverworts on /NVOLUCRB
It frequently arow«: Js a ereen j' • water is even more pro ENCLOSING
the delicate SSnl the ground u thallus nounced in the case of sexual archegon/a
supply water and the rhLoS" secured by reproduction. Their sperms
iverworts have thu-iT needed for the f ^he rhizoids have retained swimming ^antnsrid/a
- plants in ^ -%-tion to habit. The presence of ex
ternal water is a necessary
T'S"tef"S condition for the sperms to
£!-"#55SS;;;£3 swim to the egg. In fact the
male gametes (sperms) pro
duced by the liverworts are
Fig. 2-1. Pellia opiphylla (monoecioas).
characteristic of water plants and not land plants,
^ They furnish a
clear sign of their aquatic ancestry, ine spores, however, are non-
motile and wind disseminated.

I**® dominant plant in the liverworts is a frame-

Wh' does it remainthallus-like
in size. Why small ? structure.
The study Itofremains small
the thallus
structure reveals that the tissues required for large growth are
lacking. It lacks the vascular tissue which is essential for conduction
SC t" t° S.S.? «™" "''""pp.- of water and minerals. It has no supporting or mechanical tissue
in the form of long, thick-walled fibres. Besides the absorbing organs
''anspirlCr^'^H eel ^rZ'"'?'hllhauZ:re' in the form of rhizoids are inefficient. Secondary meristen^s which
srsi-i.!'-~;5"££ f»? r^ss
:s^l'"al cells^aVh^^P'^^iy ProttrA®^ advantage W
provide increase m girth and additional cortical cells are absent
Consequently a vegetative plant body remains a thin-walled, porous'
fiat, thallus-Jike structure which grows prostrate and close to the
surface of the substratum. In land plants in which the corticating
phallus is. theS-nr ^hallus\ii ttnguardfd n the tissues, which consume water, are lacking the prostrate habit is
useful. It reduces the exposed surface and the major portion of the
plant body remains in direct contact with soil moisture. Besides
^an for ,„ °,"^'A^tand ,h ''^'= in verv desiclp.,- ® it furnishes maximum surface area for fixation and absorntion of
in moist ®\S'h
®™«i^^molt'®\S"^ nftime''
oFtim tu"S affect ofZ'^r®"'"'''®- Tim'
°n moist m £ pj^s® ont'
Z»S^®an"nts
afrecYofZ'",^^Wta, rays of tZ water.
The gametophyte of liverworts is an independent plant. Some
^

rr®. ^"bmeZd"" ^®® ?unb Tr Zground, on tt to liverworts have a thalloid plant body (Fig. 2-1). It is flat and
dichotomously branched very much resembling the thallus of
foliacous lichen. However, the rich green colour of the liverwort
>'■ and abundanT
'^"t moisture thallus distinguishes it from the gray or grayish green colour of the
lichen thallus. The different types of fruiting bodies in the tw
is another distinctive feature. °
There are other liverworts in which the gametophyte nlant
leafy (Fig. 2-2). It consists of a central branched axis (stem) bearing
 20 BOTANY FOR DEGREE STUDENTS_PaRT III: BRYOPHYTA
HEPATICOPSIDA (HEPATICAE) OR LH^ERWORTS 21
found in Western Tibet. They show active growth during summer
and rainy season from May to September. Winter is the resting
period. August to September is the fruiting season. The Hepaticae
number about 9,500 species. They are grouped under 280 genera.
mTHAL LEAF ^-^exual Reproduction. In liverworts asexual reproduction
LOBULE takes place by vegetative methods only. Like other land plants true
asexual spores arc lacking. Vegetative reproduction takes place
during the growing season. It is brought about in a variety of ways.
The most important methods of vegetative propagation in the
Himalayan liverworts are :—
Fragmentation. The cells in the posterior portions of
the branched thalli die of old age and eventually disorganise. As
a result the younger branches are set free. Each of the detached
branches, by apical growth, grows into a new plant. In this way
the number of plants increases from time to time. The common
examples are Riccia, Marclmntia, Pellia, etc.
A
V^ST" Gemma Formation. Many species of liverworts produce
. 2-2. fA n^ ^ green, multicellular asexual buds called the gemmae. The gemmae
- f P^'^ionoftheleafvp ap. become detached from the parent plant. On falling on a suitable
substratum the green gemmae (Fig. 2-3) grow into new individuals
"■ •'XS.: •-SX "•—
worts are more
«., immediately. In Marchantia and Luniilai'ia the gamraae are
developed in small receptacles called the gemma cups. The
gemrnae^ cups are circular in Marchantia and crescent-shaped in
the leafy „ame Lanularia. They are developed on the dorsal surface of the thallus.

shade and
shady abundF?^ m situationr^
habitats in thl thereserion
is h" k
RHfZOiDAL
C£LL$

6REEN CELLS

NOTCH 'GROWING POINT

abound in til P^^ate znn d^c^ -f '^'^smoDnlu in
^'poohome
ft. to Of8 oon^f HimaT 'he tro^^^ution
rainf to^\ arctic,
their
^"h on the ^ abovl P^^dculaH is the h they OIL CELLS
rSS*"K'.£5 ®
them. T.^^rhwal ONE-CELLED ST/' I

A he i;,.„ h and A mnra

f
'er worts Lcc Marchantia sp. A single gemma.
-u JMgJpurCoIIcec Libran'
Date mm--
GaiINo,
11129
 HEPATICOPSIDA (HEPATICAE) ok liverworts 23

of temperature. Moreover the outer two to three layers of cells of

the tubers possess corky, hyaline walls which are waterproof. The
^ Adventitious R inner or central cells are packed with food reserves such as starch
from 'tfe^ventrtl °f advenAbu^'b'' P/°P^g^''on also grains and oil globules. The tubers, therefore, serve as a means of
perennation.

The thallus of some liverworts such as RehouUa, Grimaldia,

etc., dries up in winter or extreme dry season. The margins of the
thallus roll upwards. As a result the upper surface is protected and
the ventral surface with the scales is exposed. With the onset of
""'"'1!!^''™'
rCT^
^""^'"""ea'rbrrch;s'lP't"y
towards The"
.'oward";
""
tho end"or'"?b'"'''
those
the rainy season these thalli resume growth.
Persistent Apices. In a few of the liverworts the whole of
the thallus dries up with the approach of unfavourable period. Only
season tk , the apices of the thalli remain alive. They become dormant as they
swollen '^^^"ches get thickened or other\visc modified. The surviving persistent
form tubers T^"" thickened apices resume growth with the advent of rainy season.
usually bunVri are
^4^ In most of the leafy Jungermanniales the whole plant
J<^8£Rs Pfunt dies dries up. With the approach of the favourable season it resumes
^ " the period unr of growth.
_ growth, the f for Sexual Reproduction. The student must bear in mind that
%• 2-4. Anfi, •dormant r • ^oers remain
asexual reproduction by vegetative methods is entirely a secondary
growth. 7« p.. , 8 tubers. With the r^,, ^ drought,
at the growinff an" and 9 season the"^ oj" the favour- CAPSULE

« 'he ?halluT It Z tL resume INVOLUCRE

INVOLUCRi ANTNERIDI

-"wSS f"!;—>
SS-'i?"« "'^P» a,„.
'««y seal'T ?h" 'he pTaTf-^,^les^odifieT"
rosette of n,, he dornian.
tT""'°f'halb®
au/e. ' ^hese thlrri ^ tissue
etc. P'""^- Examl':^^^ tesumT „l"he begi ^Pio« Fig. 2'5. (A—C). Pellia calycina (Dioecious)
A. A plant bearing a sporogonium.
B. A male plant.
C. A female plant.

f^archar.;" ^ the enf ^ ^ell state th ^ ^uiazinp- r^ phenomenon in the basic life cycle of a bryophyte. It simply serves
to propagate the gametophyte phase in the life cycle and plays no
role in the phenomenon of alternation of generations. Sexual re
production which consists in the union of gametes and subsequent
production of meiospores is the usual method of reproduction. The
male and the female sex organs in some species are developed on
different plants. Examples are Pellia calycina (Fig. 2-5 B and G)
^ underer^'^^tiou
'ground are ^ ® ^iverwn''*®-
hver, * ^hvious. and (Fig. 5T A and B). Such species are called dioe
cious. In others they are developed on the same thallus. They
are called monoecious. The sex organs may be borne dorsally
ud extremes
I'
 HEPATICOPSIDA (HEPATICAE) OR LIVERWORTS 25

(?•) ADsence
(?; Absence 01
of menstemauc
meristematic tissue.
tissue.
(m) Absence of any kind of lateral appendages and of branch
ing habit.
aisabiiities 7/1 , Con^nn ^"P^ytes. The ^ (Hi) Continuity of the archesporium leading to simultaneous
ripening of spores.
nutrition frnm • P^^^^t gametnnh (Fig o-S a All these factors collectively account for the liverwort sporo-
parenf resL^ (thallus^ wV u ^"^ched gonium not progressing beyond a limited size. It remains dwarfed.
possibilities plant T the Wf J for In some species the sporogonium is differentiated into three
parts, a foot, a seta and a capsule (Fig. 2'6). In a few others (Riccia)
both the foot and seta are absent. The sporogonium is a specialised
bear leaves or to body which is solely devoted to the production of meiospores
and their dispersal. A large proportion of the cells of the endo-
thecium are devoted to spore formation as compared with the other
two classes of the bryophytes. The columella is absent. The spores
are differentiated by meiosis. They are haploid in nature. On
germination each spore produces the alternate plant in the cycle.
It is the thallus (gametophyte) and not the sporogonium (sporo-
P^klQy phyte).

(^^PTUR^Clj V^^istinctive Features

^^PSUlp 1. All the hepatics, with of course a very few exceptions, are
mu 9^ dorsiventral in structure.
2. They are comparatively small in size never more than a
few inches in length.
£lat£r
3 The plant'body, which is always a gametophyte, is an
jPOPE independent plant. It is either thallose in form or leafy in nature.
In the latter case it is differentiated into a branched central axis.
Both the axis and the branches bear leaf-like expansions. They are
all without the vascular tissue. The leaf has no midrib.
4 The gemetophyte is secured to the ground by simple, one-
ceiled, absorptive processes called the rhizoids.
Internally the gametophyte always has a photosynthetic
tissue The cells constituting it have numerous, small chloroplasts
each. ' The chloroplasts lack pyrenoids.
6. The sex organs are dther dorsal in position or terminal.
Each develops from a single initial cell.
Fi £
phyll. 7 It The sporogonium
is attLhed to andislivessmallalmost
and entirely
generallyas without chloro
a parasite on
the parent gametophyte.
othe, clisfedages.
8 The sporogonium is eitherboth
b. differentiated
the foot into
and foot,
seta seta^nd
lacking
capsule or is very simple with oocn s
--«£"«.■. (Riccia).
;,irYi Has no
9. The sporogonium has no
. . .
meristematic tissue.
 HEPATICOPSIDA (HEPATICAE) ok liverworts 27
class Hepaticopsida. Their suggestion is based on the discovery
Jl- The columella is absent. 01 a curious, new liverwort, Tahakia lepidozioides in Japan in 1951
and later in Canada (Persson, 1956). Takakia approaches the
talobryates in the following features :—
(a) Erect leafy thallus.
(b) Branching of the stem.
(c) Certain features of cell structure.
(d) Absence of rhizoids.
It, however, differs from the Calobryales in :—
1. Large archegonia borne in teiminal clusters.
2. Unique form of the leaves. Each leaf is usually divided
into two rarely three cylindrical structures which are solid and par- -
fci'ijp"Xrst"™ wK";"'-.'""-
0"m it;S"?"'"»£„!'':•
enchymatous.
3. Low chromosome number which is?i=4.
SStS-ISi'«? »fc!£%-" Sits' The bryologists, in general, are reluctant at present to accept
Hatton and Mizutani's _ suggestion. They hold that the final
CampbW"'" naming
^''oAerZdlrW '">°ae 'ainily
Utf''•'=" assessment of the taxonomic importance of Takakia can be deferred
So Thf' ^ prono..!^ ^936 until the sporophytic material is known.
In this book only two of these orders have been discussed.
These are the 3Iarchanliales and Jungermanniales.
(OamotophytQ n P^**<^Opsida ^W REVISION QUESTIONS
directly frotti always do ■ ^ 1. ihe liverworts are inefflciont land dwellers. Discuss this statement.
2. Describe the habitat and distribution of liverworts.
3. Give an account of the plant body of the Hepaticae (Hepaticopsida).
n
4. Describe, giving examples, the various methods of vegetative repro-
ssrv-- duction and perennation found in the Himalayan liverworts.
5. List the distinctive features of the class Hepaticopsida.
6. Give a brief accoimt of the classification of the Hepaticae (Hepati
r'-Fp^raf-d"eThJr ---f e" copsida) into orders and state the diagnostic characters of each order.
7. Justify the truth or falsity of the following giving reasons
^°"^PoSof 0 6ver? ^^ck S" verticaf^^'^^^ of4 (а) The thallus of the liverworts represents a transition stage in the
migration of plants from water to land.
®^nl6 cells i
thick. SiJfof
® Cell ""srr -i
-*«"io
^®yer
"an
(б) The rhizoids of liverworts are homologous to the root hairs of the
higher plant.
c one 8. Why does the sporogoniiim remain small in the liverworts ?

otber
 HBPATICOPSIDA (HEPATICAE)—MARCJHANTIALES 29

mously once or a few times. Being plagiotropic it grows prostrate

on the ground and is secured to it by two kinds of rhizoids, namely
smooth walled and tuberculate. Associated with the rhizoids on
the ventral surface of the thallus are the membranous scales
(Fig. 3-1 B). The scales are arranged in rows.
The sex organs (antheridia and archegonia) in the lower
members of the order (Ricciaceae) are scattered on the dorsal
surface of the thallus. They develop in acropetal order. In the
higher members they are borne on special structures called the
receptacles (Marchantiaceae). The receptacles may be sessile or
stalked. Some genera are monoecious and others dioecious. The
sex organs in their structure and development resemble those of the
Sphacrocarpales but differ from that of all the other Hepaticae.
Some members of the order are strictly hygrophilous. They
grow in very moist situations. A few occur actually in water,
floating or submerged. Some with a larger thallus are capable of
growing in exposed places. Still other can live under conditions of
'^H/zofOs temporary drought.
(6) Internal Structure (Fig. 3-2 A—C). Though the general
r
plan of construction of the thallus in the Marchantiales is the same
yet it presents a wide range of variation in details. It is differentiated
into various tissues. These tissues are arranged in two distinct regions,
dorsal and ventral.
The deep, green, dorsal assimilatory region encloses air
chambers. The chambers are roofed by a single layered epidermis.
In the family Ricciaceae the chambers are reduced and are in the
9'^f^STR,CT,0f, form of deep canals or channels. The chambered condition is
represented in a relic condition in Dumortiera and is absent in
BOOy, Monoselenium. In species in which the chambers are present they may
be in one or more than one layer. Each chamber is separated from
its neighbours by partitions which are generally one cell thick. It
may be empty or contain assimilatory filaments. When multi-layered
the chambers are invariably empty. Each chamber opens to the
outside by a pore. These pores vary widely in structure in different
genera. In Stephensoniella the pores are simple and wide. They
are large and barrel-shaped in Marchantia and Preissia. In
^ ^^Ale Targionia they are barrel-shaped.
(A n The ventral zone of the thallus functions as the storage region.
It lies below the air chambers and consists of colourless parenchyma.
Jit, ^andr) 8000% The parenchyma cells are compactly arranged. They lack chloro-
°""" I'- S »« "^''^® ""PS.
cup
plasts but contain starch. Here and there are present the oil cells.
the "f With n,
more or ] ' difrr'' sn lo^
7'^ The lowermost layer of this region bears two kinds of rhizoids and
shows7'^^^lessaH-
dorl?'^ a disfi. . ^ scales.

^"tiation. On the basis of anatomy Mehra divides the thalli of the

Marchantiales into the following three categories
28
1 Marchantia Type (Fig. 3-2 A). The midrib region is ex
panded The ventral storage zone is several layers thick. It
 HEPATICOPSIDA (HEPATICAEJ-MAROHANTIALBS 31
The parenchyma celb « is 3
P»oTosy»naT,c
■ FILy,Ma^'^ CHAMBER
eonstitnting it a^e cies ofMlchaZ\nTTt""onia jSpophyuT^^"''^
"" POp^^ centre. In the wings it^onlts of"®
'^ERMIS presents the lower eDidermi«! o . which re-
^OfiAse chambers arranged fn several
SCAIP^ chambers are empty and are separated hv ^ other. The
The air pores are either smr shaped or partitions,
pies ■a.re Plagiochasma appendi^atum. T Z]fcuhtum 'Rfh''''r'
OriH^aUia indZ"'LnTlt:
^''ioa„M,s »JI SEfifH^yg
CHAMBERS

S|"sp'
with the exterior throughspacious
>1, !l andor empty.
large gaps They communicate
poreLfextremel^Si^^^^
type. To this type belong Stephe7isomella brevipedunc^data Sauchia
f^f^'ZO/O
SSSeiSi?—"■ "■ '•""'* "•"• —"S -
".Origin of the Marchantiaceous Thallas. Two widely
"T^orsar ^f/OBRM/s
MrrXanti™ns thallus.
Marchantiaceous tlmllus'"^'"'
Some forth to explain
bryologists the that
consider originthe ofsimole
gametophyte of the Marchantiales is the result of retrogresXe
tS
evolution. Others hold that it is the result of progressive evolu
tion or elaboration. wm-
^^°^age.
of Retrogressive Evolution. The .im
porters of this hypothesis are divided into two camos IVT.i "?*
Mm p,Q of them particularly the older bryologists hold that^etr^Ss'v^
Pi evolution has been brought about by progressive -simplification ^lo
called reduction. The latest view put forth by Mehra is tLt
has been brought about by condensation.
Reduction Theory. The reduction theory to exnl;,in
„ ttallna shon,ta''PP''adicuio. =liotch), retrogressive evolution (regression) was first proposed bv v
Wett^ein in 1908. He held that the primiti/e g^tophyte of
eolourless. Tj, °^'''®''""«ulata < ^^^era), » Portion of the Hepaticae was nearest the erect, leafy Acroeynous Tuneer
manniaceous forms. His theory with slight modifications rereiO.^'
Sea'-?,;AtCe'r a,arch g, support from many eminent bryologists. The chief amono- T
were Church (1919) and Evans (1939). Von Wettstein pHeed
Aerogynous Jimgermanniales of the Calobryum type first Tt •
an erect, leafy gametophyte radial in symmetry and with 1
in three rows. The first step towards reduction was thf>
of the prostrate habit. With the appearance of dorsiventra]>"
the leaves on the ventral side gradually diminished in size h
finally disappeared in some cases. The changes in habit a^H
diminishing size of the ventral leaves were accompanied bv th
If, ® semi. flattening of the central axis. Associated with these changes w^
 33
HEPATICOPSIDA (HEPATIOAE)—MARCHANTIALES
2. Fusion of the lower portions of these leaves at the^ points
of contact resulting in the formation of single layered wings on
either side of the central axis and lamellae on the upper surface o
each wing.
3. The one-cell thick lamellae were directed obliquely out
ward and as a whole formed open chambers of linear type.
4. The next step ivas the roofing of the air chambers by inward
growth of the margins of the cavities of the chambers.
evomuon by reduction in the U;;rwoTt, 5. The roofed chambers, at first, communicated with the out
side by large gaps. The gaps were later replaced by definite pores
—a protection against loss ot water by transpiration.
°f 'he Pleridophy
?o4al,f "eternal foZ atH ,'""- "<= "™ci< 6 All these changes were accompanied by gradual flattening
HverworrtWi and between the radial of the central axis till it reached the margin of the wings.
cular The! Marcfc?^ cernuum with the 7. Formation of secondary partitions occurred across the
PhotosyS"^^^' ^f°^°Phyno7e fc parti- chambers as the flattening of the thallus occurred.
podiales
and wallsrespectiveira^
of thev of the rh?
upright Enn?- ^i
l dorsal
Lyco- 8 In some genera adapted to xerophytic conditions like
blance tempted Ka<:h ^^"^bers in the MarrK filaments Plagiochasma horizontal partitions were also formed between the
lamellae forming multi-layered chambers.
-;ose by,/duSo'^
developed the followiW ^hat the
P^^"dophytea^l'nr resem-
Probably
^ttig arguments against 1^1 ^^hra has
, (») It IS diffirnu . hypothesis • 5S.S:"n<[Te tmadoLf banxl-shaped pores characteristic
of some of the Marchantiales.
vProeressive Evolution. The adherents of this hypothesis
w (ti) Absence in .u ^ ^ttnple liverwort hold that the Hepaticopsida originated from a simple, thallose game-
oohvm The^^a^^^^ thallus was simple prostrate and show
ed nrexternal or internal differentiation. The theory of progres-
s^n wi originally suggested by Schiffner. The chief supporters
aie Cavers, Campbell, Fritsch, Bower, Smith, etc Gayer s suggested
Sphaerocarpus and Campbell thought forms like as the
present day Hepaticae which show a nearest approach to the primi-
hsed, sexiia/V'^^^'^tilt to pv i • ^ Phallus of the Uve hepatiLan gemetophyte. From these primitive types progressive
evolutionary sequence followed two diametrically opposed directions.
This resulted in the evolution of the following two types of game-
ted the derivat"^'*^^'^®'* Thp
?"the'a:c"ePrS^
^ ^^rchanf- f'^ttns ]!],_ ^^'''(^hantia
;^ostral prothalh
to be
tophytes:
la) Marchantiaceous Gametophyte. It is suggested to have
evolved from the primitive thallose gametophyte by the following
Retention of external, simple, prostrate, thallus-like form.
r■^ r,.»rhial but progressive internal differentiation of tissues
J^»germanniales£''^^theMar^^^^^ H''''''^^tion adduced It
T finally
n 'n' resulted
™dted in the formation
leading into theinternally
of a thallus air chambers con-
composed
mininfcWorSyllose'laments and a parenchymatous. compact
''°"T»r7'egation of sex organs into localised areas called the
receptacles.
 35
HEPATICOPSIDA (HEPATICAE) -MARCHANTIALES

10. The sporogonium is simple in structure and small, either

with or without seta. It has a capsule wall one ceU in thickness.
Columclla is absent.
VGlassification of Marchantiales. The order includes about
420 species. They are placed under about 35 genera. Gainpbell
(1918} recognized only three families namely Ricciaceae, Corsmiaceae
vS^orogoniuin Jt r« and Marchantiaceae in this order. He included the genus Monoclea
and Targionia in the family Marchantiaceae. He, as early as 1898,
was the first hepatocologist to point out Mo7ioclea's relationship with
the Marchahtiales. This viewpoint was later challenged by
Schiffner (1913). Verdoorn, Evans and some others supported him.
Verdoorn (1932) and Evans (1939) each recognised six families
in the order Marchantiales. The former made out Marchantiaceae.
Operculatae, Aslroporae, Targionaceae, Corsmiaceae and •
Evans recognized Marchantiaceae, Sauteriaceae, Rebouhaceae,
Targionaceaf, Corsiniaceae and Ricciaceae. ^oth of them exclud^
ilfoTjocZea from this order and placed it in the suboider Anacrogy-
and form
nous Jungermanniales.
Later Campbell (1940) revised the classification of Marchantiales
basing it on the following features :
/ 1. Nature of the receptacle.
2. Manner in which it is borne.
3. Structure of the sporophyte.
On the basis of the above rnentioned features he divided the
order into the following five families :
1. Ricciaceae.
« chambers. ® °" S^^nerally e„H 2. Corsiniaceae.
*™"gh pit"- cha„.be„ con,„>a„i,^, •
Silicate With ti
3.
4.
Targionaceae.
Monocleaceae.
P-»chyIt,37;ral
g
region is -composed of ^co»n
"= "orago
5. Marchantiaceae.

It is evident that the families Ricciaceae, Corsiniaceae and

thallus. are usually Targionaceae are common in Verdoorn, Evans and Campbell's systems
of classification. The families Marchantiaceae, Operculatae and
^^ans^oftwn^ J-^allus h so surfac, of the Astroporae of Verdoorn and Marchantiaceae, Sauteriaceae and
Rebouliaceae of Evans are equivalent to family Marchantiaceae of
Campbell.
As stated above Verdoorn and Evans excluded Monocleaceae
from Marchantiales and placed it in the suborder Anacrogynous
~-?t'
"■»~"'.sr'-~».»,„, " ""'"-'ia dor-
Jungermanniales whereas Campbell included it in the Marchantiales.
To clear this tangle Proskauer (1951) made a critical study of the
taxonomic position of MonocUa and suggested that the incJu^tSiT^
in the Monocleaceae in the Jungermanniales is unwarranted. ^Accord-
ing to him MonocUa shows affinities with the dumortieroid line of
the Marchantiales.
 36 BOTANY ^FOR DEGREE STUDENTS—PART III :: BRYOPHYTA
OTUiJJiiiNTS—PART III BRYOPHYTA

MarcW^fw. reduced the number of families in the

yan liverworts hf collections of the Himala-
families Tarffinnnr- intermediate forms which connect the
Consequently he ren ^^^^rsmiaceae with the Marchantiaceae.
order?- ^ recognised only the following families in this CHAPTER IV
1 • Ricciaceae. V>l'^CHANTIALES—RICCIACEAE : RICCIA
2- Monocleaceae. Ricciaceae
3- Marchantiaceae. ^yOeneral Characters : , r .u
to the list. This^f?milv Monocarpaceae (=CaiTpa- The family Ricciaceae includes the simplest members of the
order Marchantiales. They are charactensed by the follouing
onoc£ir^)M5
Renamed as sp/iaerocarpus t7 a singleGarr
thollose hepatic
in 1956 and features :—

has an odd combinatin^ oC Proskauer in 1961.

mvolucre"'^Th ^ ^
openintr K. A Presence of reduLn ,
simplerelatively a large
female reeenr^?^ on the .^ °^°^y^i^hetic air chambers
pores, each^r 'roofed air chLw shortly stalked
short seta tiers of °Poning by barrel shaped
^elatioShip °Tfu°Lon
assign Monocarpu^] ^^rchantiales. Th?s\1 felT''

... n...rz: 5P0R0G0NHJM

favour of dividyg
M'lrchantiaceae MonocarLc^""f Corsiniaceae,
(—Carrpaceae) and Fig. 4-1 (A-B). Riccia fluitans.
REViici/x»^, A. Floating form.
B. Land form.

Lwt the distinctive feat gametophyto of 1. The garaetophyte is a flat, ribbon-shaped somewhat fleshy,
green thallus. , - r- • \
9 The dorsal photosynthetic region contains fine air channels
which in some species are large but generally greatly reduced m
fi
the structure and origin of S In the latter case they occur as deep narrow canals or
channels separated by long columns of green cells. ^
3. The air channels or canals lack special assimilatory hla-
ments and-thus are empty. „j •
4. The upper superficial layer, the so-called epidermis,
lacks definite pores. The pores are either absent or are redimentary
and unspecialized structures. ,
5 The sex organs occur in the median furrow on the
uDoer surface of the thallus in longitudinal rows extending the entire
length of the thallus from the apex backwards.
37

wwr*mww rw?:
3S BOTANY KOR DEORRE STUDENTS-PART In : BRYOPHYTA MARCHANTIALES-RICCIACEAE ; RICCIA 39

6. They occur singly ,he open cavities at the bottom. '^dult Gametophyte (Fig. 4'2 A and B).
liverworts. It lack?both ule'foo/and Tet^''"'l
spherical capsule only. consists of a ia) External Features. The plant body is a thallus. It is
small, green, flat and rather fleshy. It grows prostrate on
the ground and branches freely by dichotomy. Consequently it
9 locked in the 1 generally takes up a rosette form (Fig. 4-8). The branches of the
thallus are called the thallus lobes. According to the specip the
thallus lobes are linear to obcordate. Each lobe is thickest ^nthe
si«re ihe arcbesporium are devoted to middle and gradually thins toward the margins. The thick middle
portion constitutes the midrib region. On the upper surface ol
The family Includ"' T each lobe there is generally a median groove or furrow, it is m
the form of a broad channel in R. pathankotensis (Fig. 4-J B) and a
three genera 1^0 species tu
^iccia. The first n ^^^selina under
rest are in i ^^P^csented hv ^ • i and
(Fig. 4.j^ h^ ^amp soil. The onlv" all the
aquatic. Ricclorn naia«s are i?iccm
Jake) and Sr' has bee^ron J!i They
^^rhwal, Kashmir Pesh"' ^^^histan). i"c2 J
The stn . ^'^hawar, Madras and 1?
Kashmir
occurs in
Ricci. here. The
he genushfeis history
namJdofJtv
th , (Punjab).
MA^AL
vW'ewaiic Posiii^y^ '^han botanist F. F. SCALES

^''yphyta '
H«Paticopsida Fig. 4-2 (A—B). Riccia himalayensis(R. discolor)
^""hantiales A. A patch of plants.
R'cciaceae B. A single plant.

and Hatot" L. narrow longitudinal furrow near the apex^ in R. melanospora. In

R. sanguinea {R.frostii) thQ upper surface is flat or slig y conv
^?-Sir5,•'
?«eSS~
f3- M aT T "^"oofor P;
'i«?=
important '' ^"dia both
6R00Ve

f'g. 48), 'r If''.''nl'otensj fr,^'^)' R. robLn"lT^

(Tig. 4-1), etc r,p"®'a (Fw i / if'4'3), jj 'a (K crystalUna,
°">as are terreoy'^ excentio R- (R-frostii),
ating or damp soji p "f' species een aquatic all
1' continuen^'y jubmergej rosett®r'=''^"y g™"-for short
« contact wftir'lf'' g™watbuttK f 1'^'^ onlv%;i\P°'^'^^
'he mud in poo^ lakes,
p R has its • ^^hlETOppv^ hottom. ^ ®) when it
Fig. 4-3 (A-B). R. pathankoteusis.
A. A patch of plants.
he gametonu
'^'ophyte plant spores,
which B A single plant showing a dorsal furrow or groove.
 MABCHANTIALES-BICCIACBAE : BICOIA 41

40 BOTANY FOE DEGREE STUDENTS—PART III : BRYOPHYTA

Each thallus lobeThe

usually ends notch
in a terminal
(i, Photosynthelic
of a loose, greeyssue Region.
It « The photosynthetic^i^eg^^^
° cells are
growing point. terminal is absentnotch in which lies the
in R.frostii.
"rra^T'^ vSfcat'tvs^or^^olurnns. Generally between the
UPI^R EPIDERMIS
chloropryllose
and lack protoplasm at mauluy The'v 1" Aizoids areoutgrowths
projecting inwards into the Inml' r ^ I
narrower CELL

form and function ^ The rhizoids in STORAGE PARENCHYMA

however, are analogous to each o'ther
lower epidermis
function
solutes. asThe^ubemu^ate
organs of attarhm
rV. M homologous.
water andThey
soil RHfZOID(SMOOTH WALLED)
either absent or they dossp« (^- froslii) are tuberculate RH\ZQ/0
from
mergedthespecies
lower surface
of of the thXs generally absent.
tubercles. Arising
In the sub- Fic. 4-5. Riccia sp.
the violet, membranous scale.. Tu' ^ ttddition to the rhizoids, are Portion of a transverse vertical section of the thallus.
in thickness. The scales are not anrf^f"""'«'lular and one cell
eea"
columns of green cells are very n^roovv^^top,^ vertical ^shts^_J
and projecttransverse
forward row near the
to piotect the'^'''' ^ire usually arrang-
^'t"ated closely sUts are called the air canals or^^^^^
larger '" the form of
^ crnoiata and R. robusia
(E. discolor) and R. sangumea
portion of the thaUuslobS die! growing point. In the older ■/ canals or chambers in
mTr'^^^^-^raet^ n eT roedia.riinc Co" r
(B. cryslalhna). In
iB. frostii) they are 4-5) characteristic of
ThS are has^small purol?''''',""" lateral Riccia lack assim latory ^ or channels consist of
Marcliantia. The walls separating species In others they are
four vertical rows of green Ppells \r. vesiculosa). The
bounded by eigh vertical rows ol ce ^ distended,
™pAatroTa®t"tor
A,or-
COLUMN
a 'S^e'elrorar^'--
air canal struoturJ.
superficial or uppermost
The .d-tended or enlarged te™
are hyaline. They do not
contain chloroplasts. Toget defined, discontinuous
sISoTJ" cells
layer, ofthetheso-called
ne ghbourmg 'ows tem - an^Ul d^eh exterior through
upper ep^
PORE

oorsal
The air canal or channels the air pores,
gaps in the upper epideimis. ,f nmnccialized and rudimentary
ASSIMILATORY
PECiON ll^eair pores in 'h™ by 4 to
structures. They are cells.
8 so-called epidermal =t™Pj,,'
No true or ell defined
The air
widepores
air character-
canals in
^^NTRAL
SCALE istic of ilfaro'""!''® w?th the outside by their whole width.
^t^orage reg/oai B. crystallina ""t"""""'' chrnn2 func^^^ in photosynthesis. In the

Tranavei-eJve'rut] mu -^ilatorv revion in Ricciocarpus natans (Fig. 4-6 A and

-any cells d BI con^i- ortheTppTlVer
rrnr^S Wirre-S' tiwough air pores.
(ii) Storage e^„sisTs^of cToTel"pVaed undifferen-
""cgion.
?4:ed without in.ercellular spaces. The cells
 marchantiales-biociaoeab : RIOOIA 43

42 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA

are colourless and may contain starch but no chloroplasts. They wal ls. The y°""g, f™''r>'°T„'' ' arise 'iTter
apex is compact. Air chamb
'"by '?hf spftdng of the
opinion favours the second
serve for water and food storage. The lowermost cells of this region
AIR PORE
Ii;;mtsis^'''^hfsupp'otrr .his hypothesis ate further divide
AIR "(T) '^slmTitold that the 'fto^ the sS^nd
CHAMBERS
e'xters'dotvjl''TL'aXnt supporters of this view are Black,
Orth, etc. ^ ^ The chief among them are
(n) Many hold the contend that the splitting is
Barnes, Land and of the thallus below the
endogenous. It swrts , gradually extends upward right
surface. It begins from below and giadualiy
RHIZOIP up to the epidermal layei% ^e
^^ical growth (Fig. of apical initials.
SCALE
They arclobes
thallus takes place
arranged by _ means row
in a horizontal o g
^ V number usually
^^^.^jning the
from
apical3 initials
to 5, sometimes
is located amore. ^ notch
pctablished at the
in the tip ofthallus
young each
lobe. Once the apical ^"^Vf^hnut bv t S ac iviiy and division and
■'iS3 ,rj;:s3;iirr ,..3 ■
APfCAL dorsal segments
CELL 'primary
SPOROPHYTE
archgowal
CELL'

■SCALE

A.

Q young sporophyte.

so-caued lower
„-«rigin of Air Chambor x
B^ack (igTsl'aSi'o\^ £ld' Fig. 4-7 (A-B). Ricciasp.
A Vertical section through tho growing P""'-
B Section through the growing point showing dichotomy.
(Diagrammatic)

depression Ubecause
- the
the embryonic
em y ^ cells on^^anthe the
sidesembiYonic
of the apical
cells
cells divide and grow mo ^ ^edge-shaped
posterior to the apical nts are usually cut off from the
It has four cutting faces but segmem
U BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
MABCHANTIALES-KIOCIAOEAB : BICCIA 46

dorsal and the ventral faces. A major portion of the thallus is how
ever formed from the segments produced at the dorsal face. Each
thickened
the advent
to form tubers at the ^^ant" perTshes. ^he
growth under suitable condi-
dorsal segment divides by a wall parallel to the surface of the tubers remain dormant and resume g
Callus. The outer and the inner daughter cells divide and jedivide. tions. _ ,
ihethesegments derived from region
the outer daughter ^Axual or Gametic Reproduction-
to upper photosynthetic of the thallus cell
and differenUate in-
the sex organs. 1.' Distribution of Sex Organs^^ The the
to the ventral, colourless, on the thallus lobes which ^re no extending back from the
to the vpml^^V to Pietsch the segments cut off parallel purpose. They are the dorsal furrow or g^^^
enidLw tV rhizoids and the scales.
epidermis, the give rise only to the lower growing points. Geneially y . They are developed in an
Lnk deeply, t^nch in a separate cav ty.^^lhey^^
cells of thp*!-Aw™f and then some of the median acropetal order. are away from it. The anthendia
which separates tL^A - vertically to form a tissue the growing point and the o develooed on the same thallus.
Each set fiinrt" point into two sets of apical initials. and ichegonia, in some the two kinds
E the P°i"tof a new thaUus lobe. It Such species are known t-rr , thalli- They are referred
addition of mofe and^'^mn^^T" marked by the of sex organs are developed o j.,.oiallina), B, billardieri, R. gan-
initials. tissue between the two sets of apical to as dioecious.
gctica R- robusfaar
and R. patbanJcoterisis r ^hile R. dioecious.
4-8) are himalayensts
In
^^^PRODUCTION (B. discolor) and R. sangmnea {R- frosHi, rig. ^ )
of maturity. U reprodu^rv^^ reached a certain stage SPOROeONI'J'^
following a sexual process Thr^tk meiospores formed
rnethods and is also concerned wltWK reproduces by vegetative
are produced by the non-sexual inrlwS^
sporogoniam. dividualsexual
in the process.
life cycle.Meiospores
It is the
the ReproductioT* p• •
gi-owing seaso; b/Te
1
tative cells.^'rhe^Sin
eventually disorganise Wh^ die of of
oldtheagevege-
and
l^ig. 4-8 (A—C). Riccia sanguinea (R. frostii)
A. Mais plant.
B. Female plant.

and then a
X "iDa ^ -SI:
sitToVa'' 'su°nker aXidTuU rs"usuaUy
X"ccl bra tTny tower-like
The thallmafJjLm, projectionsexof surrounding
asitbearsthe thallus at.ssue^
organs, is known he
'"sizsr.t'r- >■" gametophyte plant. The sperms and the eggs are the last struc-
lures developed during the gametophyte phase.
the end of the or ^^P^rted that in i? hity, j Structure and Demlopment of Sex Organs.
down into the apicet of discdor)rainu^
dinew
es. plants.
During fte next g™wrng season
thic^ned tI
the apilergrn" gLtv
Pl^nt al Slfuctwe.
[1) mature antheridium
• ts of an ovoid or a (Fig. 4-9, I) isbody
pear-shaped an
elongated strnctuie. ^ (anthe-
r, T^ierformnf P"'"' S™" "P and form seated on a short, few ^ bottom by means of its mul-
ifi i. j
■■"""""
 12!"
Ofthe thalJu h »..'..VCP»™ MARCHANTIALES—RICCIACEAE: RICCIA 47

outer,jacket layTrfstSle celTs'"''

ooiis. It
untheridium has
IS called the antheridial
(ii) Dehiscence. Presence of moisture is essential for the
dehiscence of an antheridium. Water enters the narrow pore
^therid/al PI^IMARY ANTHEPLOiAL CSU of the antheridial chamber and finds its way into its cavity. The
initial
PRIM.STALK^
cells of the antheridial wall at the apex of the antheridium absorb
- ♦ CELL ^ this water by imbibition. They get softened and eventually dis
UPPER integrate to form a distal pore. The sperms may be shot with con
siderable force or extruded slowly in a single viscous mass through
Lomp
CELL the pore. Eventually they escape through the narrow canal of the
Ri PRIMARY
antheridial chamber to the upper surface of the thallus. Here they
swim freely in a thin film of water in the dorsal furrow.
{Hi) Development (Fig. 4-9 A—H). Each antheridium deve
'WALL CELLS
lops from a single superficial cell called the antheridial initial
(Fig. 4-9 A). It lies on the dorsal surface of the thallus immedia
isnoiP ANDROGOHW
CELLS
tely behind the growing apex. The antheridial initial increases in
size and becomes papillate. It then divides transversely into an
STALK upper cell and a lower cell (Fig. 4 9 B). The lower cell under
CHAMBER ^ELLS goes a few divisions to form the embedded portion of the antheri
WALL dial stalk. The upper cell enlarges and undergoes parallel cleavage
'^hohosomal to form a row of four cells (Fig. 4 9 D). The two lower cells of this
STALK VMtitMVf row function as the primary stalk cells (Fig. 4 9 D). They under
go a few further divisions to form the stalk of the antheridium.
The two upper cells of the row function as primary antheridial
cells (Fig. 4-9 D). Each primary antheridial cell divides by two
successive vertical divisions at right angles to each other forming
stalk
four cells (Fig. 4-9 E).
The body of the antheridium at this stage consists of two
tiers of four ceils each (Fig. 4-9 E). Periclinal divisions now appear
Various
in all the cells of each tier. The young antheridium is now differ
o antheridium. entiated into an outer layer of eight sterile jacket or wall initials
^all. Th« , . *Planation in +^, enclosing the eight fertile inner cells (Fig. 4'9 F). The inner fertile
cells are called the primary androgonial cells. The latter divide
further. Eventually a mass of small cubical cells is formed. These
are the androgonial cells (Fig. 4 9 H). The latter undergo fur
to form two^" ''elatively lar^" mother fu ther division. The cells of the last cell generation of androgonial
«ytes (Fig, 4.]a^® cells knov! ""^'eus. it X ^ a cells are sometimes referred to as the sperm mother cells or
spermatocytes. Smith calls them androcyte mother cells. As
the androgonial tissue increases anticlinal walls appear in the jacket
or wall initials. This results in the formation of a jacket layer of
sterile cells one cell in thickness. It constitutes the wall of the
antheridium. Each androcyte mother cell divides diagonally with
out the formation of a wall into two triangular sperm cells or
androcytes (Fig. 4 10 A). Each of these gives rise to a minute,
Phebody cnnc- ^ ^tth a' •• "^mute ci . biflagellate sperm.
Coincident with the early development of the antheridium
the neighbouring cells of the thallus tissue exhibit rapid upgrowth.
mmmmm They divide and grow upward around the antheridium The latter
at maturity becomes completely enclosed in a cavity. This cavity or
pit is called the antheri^al chamber (Fig. 4 9 H and I). It opens
by a small pore at the upper surface of the thallus.

W
 48 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA MAEOHANTXALES—RICCIACEAE : RICCIA 49

{iv) Formation of sperms or spermatogenesis (Fig. 4-iOA—D).

The sperms are formed from the triangular androcytes. Each
the lower, larger egg cell and the upper smaller ventral canal
cell. The latter functions like a plug holding the former in place
androcyte (Fig. 4-10 B) has a dense protoplast. It has a distinct rela in the venter. The egg cell is the largest of the axial row. bach
tively large nucleus. There appear near the periphery of the archegonium lies within a cavity. The distal portion of the arche-
goniai neck, however, projects above the surface of the thallus into
8L£f^\^/?0PLASr
the dorsal furrow. R. Sanguinea {R frosth) is an exception in which
EiONMTEX) the neck almost reaches the surhice of the thallus.
blepharoplasj
(u) Dehiscence. When the archegonium reaches maturity the
neck canal cells and the ventral canal cell degenerate. Their pro-
ducts, when hydrated, form mucilage. The mucilage imbibes water
and swells. The pressure thus set up forces the cover cells to
separate from one another. In this way a narrow passage, the
canal is formed. It connects the cavity of the venter containing the
egg with the outer world. The neck canal is now filled partly with
water and partly with the mucilage formed by the disintegration of
the axial row of cells except the egg cell.
{in) Development. The archegonium springs^ from a single
cell called the archegonium initia] (Fig. 4'1 lA). It is the superficial
g androcyte into a ,archegon/al
INITIAL
^ *gel!ato 8per„ UPPER
CELL
granule u- ' Black)
OWER
CELL

^'ongation
a thinofcord
the Ki ? in fK^
and ,1 spherical (Fig-^ Iis t ■
androcyte
B >>-v^
PRIMARY COVER CELL
C
PRIMARY AXIAL COVER CELLS
^^cotnes sickle ^ The blepharopl CELL CENTRAL
CELL
PRIM. NECK

(% ^l^ickened head. The nuc eu^ PERIPHERAb

INITIAL
CANAL CEU
'RIMARY
circles the prQtnnf^^ Protoni sickle-shaped PRIMARY. VENTRAL
of thebFepff? "P ^0 abZ^'^'r ^o^d-like blepharopl^^^ VENTER
CELLS
CELL
t, '■evLls ?h
produces th
flavin ^Py '•eveals
ProducesM distance. The
distance. The thif
thicke^
c
b&^- The curled ^he clcf^^,c
n

c """«d
vesici..f^no
vesiri
?'■' tli
P°^tion of f]°
forms
^Pex
apex
theL
fumic? ^he sperm
of the
,,°h 'h'
.g
NECK CANAL
CELLS
IT

h
^Perm^-
' ®'%410el4e^'^'°Pl^3m
" of ' ^"drocyte
attacLri with two brms ^ 111 VENTRAL CANA
CELL

H.h.sSptj«=Heg„Um, ,, '^"^^-f'^epostenlotettdof
QSPHERE

T SPERM
ri<

tu
aiiVa°?'^ archegoniuW 005PHERE

T]
cy f^tective of a "!"'• There is usu^l') ,,e Fig. 4-iI (A—T). Riccia sp.
to Th'■ bv ."'^''hcal row of four cells. ^ A H. Various stages in the development of archegonium.
spi Explanation in the text.
sei gf'^ater di? f f°ur Z . ^-9 cell are arranged in six fo"^ I. Mature archegonium illustrating fertilisation,
(Diagrammatic)
^®titer * ^^^tiniir, '4emliOa"
^eck cover or tip ofcellthe.
lid cell derived from the younger dorsal segment of the apical cell. The
^^closes the The^^ ^bove The venter also ha6>.^ J js
is archegonial initial grows and projects above the surface of the
^all i^ jacket of the neck- It
y ^vhich i, one cell in thicknes^ are
^bed with two cells. Th^y
 50 botany .or DRCREk STUBENTS-PARTln, BRYOPHYTA
" UDENTS-PART m: BRYOPHYTA MARCHANTIALES-RICCIACEAE : RICCIA 51
lus. It then divides hv n
from an upper ceU^pIg' Thf ™"
part in the development of thp n J ^
lower cell
takes no further is perhaps in response to certain chemical substances (proteins and
inorganic salts) present in the mucilage which diffuses from the neck
-3=rirri'r'«^^ ".r, of each archegonium. Several of the sperms may swim downward
in the liquid in the neck canal. They enter the venter. Usually one,
probably the first to arrive, penetrates the egg. The nucleus of the
sperm travels to that of the egg and- unites with it to accomplish
fertilisation. The act of fertilisation ends the gametophyte phase.
SPOROPHYTE PHASE

ceU (Fig. 4.nE). P"""y cover ceU from a^nTnTer'ceS This phase in the life cycle is the direct result of the sexual
process. It comprises the zygote, embryo and the sporogoninm.
• • this stage a tr (а) Zygote. It is the fusion cell formed by the union of the
^"chego^tm «nTaTcelF'"S '^e six jacket sperm with the egg and is the pioneer structure of the sporophyte
six jacket cells of two tiers the phase. It secretes a wall around it and enlarges in size. The
°«terceUse^ tier funSon as t^ The zygote is retained within the venter where it begins to grow imme
*ieck canal r«ii ?2- the central cpM e initials. The diately. It enters upon no resting period. The zygote differs from
the unfertilized egg in the following two respects :
1. It has a diploid nucleus.
2. It has a cellulose cell wall around it. The egg before
w7-rediT4^^
"chegonial neck has a'J Pi'unary neck can f formed syngamy has a haploid nucleus and is naked.
formed from the prLT''® The tip of the (б) Development of Embryo. The zygote (Fig. 4-12A)
lying in the venter undergoes repeated cell division and cell enlarge
d'visions at right angfetlVr" "" by 'wr*" They are ment. A spherical mass of undifferentiated cells called an embryo
(Fig. 4-12 G) is formed. It fills the venter cavity. The archegonium
is persistent. The venter expands as a close envelope over the
5-"ir,■• developing embryo (Fig. 4-12 C). It becomes two cell layers thick
and is known as the calyptra. The neck of the archegonium later
withers.
The first division of the zygote is by a nearly horizontal wall
f®rimir"'- "eTrme^»^i «» rnd'n«^;^lTb"Ce" (Fig. 4-12 B). It is in a plane at right angles to the long axis of the
archegonium. The next division is at right angles to the first. The
young embryo now consist of four cells. It is the quadrant stage
The It tak T'™ pit (Fig 4-120) of the embryo. All the four quadrants are nearly equal.
Each quadrant, as a rule, divides by a vertical wall at right angles
to the preceding ones. The embryo at this stage consists of eight
nearly equal cells. It is the octant stage. Garber and Lewis
reported the formation of a four-celled filamentous instead of a
quadratic type of embryo in some species of Riccia. Succeeding
divisions in the eight-celled embryo are in an irregular sequence. A
more or less spherical mass of 20 to 40 undifferentiated, colourless
cells is produced.
(c) Young Sporogoninm. Each of the outer cells of the
undifferentiated spherical embryo divides by a periclinal wall
(Fig 412D). A single layer of outer cells is separated from an inner,
central mass of cells. The former is called amphithecinm and the
latter endothecinm. The cells of the amphithecium are large and
flat. They divide only by anticlinal walls to form the protective,
sterile, jacket layer which remains one cell thick and constitutes
 MARCHANTIALES—RICCIACEAE : RICCIA 63
11™ '''''''''''' ™NTS-PABT hi: BKYOPHVTA
are all alike. They consSnfp cells of the endothecium As the spore mother cells enter upon meiotic division t^cir
rogenous tissue. It is called the generation of the spo-
walls disintegrate. The wall of the sporogonium and the cells ol
nium advances towards maturhv^h I""""' sporogo- the inner layer of the calyptra also disintegrate (Fig. 4T3E). ine
re-d.vide to form a ma" of n ="'^'^'=sponaI cells divide and rounded spore mother cells lie tree in the cavity of the sporogo
nium surrounded by the single layered calyptra. The developing
'ofthe last cell generation of "he^sn^'
P ■'ogenous The
mass function as DiSINTEERATtNS
r MOTHER CELLS
CALYPTRA
■disorganised
WALL OF CAPSULE
SPORE TETRADS
©
OOSPQRE
TWO LAVeRED
calyptra
SINGLE LAYERED
^e0oR:®o®
'©^OmCfe®^ © S
©o CALYPTRA
_ CW> Q © O
CHYPTRA .->w „ o ® n®© SPORES
' o%© ®2©® ® © ® .disorganised INNER
amphithecium ' ©Oq,® O O o© O® O® CALYPTRA LAYER
V®®® ®0® ©®.
CNDOTHSaUM ® ffl ® © ®i
^®..v®oo© o%Q »y
SPOROSENOOi O 0 0 0.05
Tissue
Fig. 4-13 (A-E). Riccio sp. Formation and maturation of spores.
A. A sporo mother cell.
B. A spore tetrad with a common sheath around it.
C. A mature spore.
g oospore ready "P^'Pgenium. D. L.S. Sporophyte containing spore tetrads, degenerat.ng spore
mother cells and the disintegrated sporogonium wal .
E. Spores lying in a cavity surrounded by an outer layer of calyptra.
spore,s are bathed and nourished by the nutritive fluid formed by the
disintcjiraiion of i —
( i) walls of spore mother cells.
cells are thp^*?*"^ '^"other cells or e t^^^S^ammatic). (t?:) non-functional spore mother ceils.
(Ui) jacket layer of the sporogonium.
(^'y) inner layer of the two cell thick calyptra.
div.deaS"™: ®ts °d ™" nded*'"T?P°™=y'- begfn"^^^
meioticai?y tj'?'^'=ly granular p'; belome ,d] Sporogenesis (Fig. 4-14 A-H). It is the process where-
^4 by th'e'spLs are fot^ed frotn
tcgrate to form a n spores A r . all th^ ^^cilagi-
v ir'povT iTmdosi: th'e liploid icleus The
cell undergoes two successive divisions
of thefiretspmedivision
motheris
Marehantlale^ u^i reductional and the second mitotic. Both constitute meiosis
"members of ihe The walls between the four haploid nuclei are, however, laid
 BOTAI^y FOR DEGREE STUDENTS-PART III = BRVOPHYTA
MARCHANTIALES—RICCIACEAE : RICCtA 55
Simultaneously at the enH r xu
and the other half of female parentage. The former are called
the paternal chromosomes and the latter maternal. For every
paternal chromosome there is a corresponding maternal chromo
some similar in size, shape and nature of inherited characters. These
are called the homologous chromosomes.
8 chromosomes of i^iccm crijstalltTn (u) The homologous chromosomes become arranged in pairs
in the next stage {Fig. 4T4B). This is the zygotene stage of
the first prophase. The pairs of chromosomes are called the biva-
lents. Each bivalent has one maternal and one paternal chromo
~i«i.jribXT.h,;2.i;S„?vrJ r?'~
the diploid "i»«'"
nucleus become somes which are homologous. The number of bivalents is one-
half of the diploid number. It is called the haploid number.
(Hi) The two chromosomes of each bivalent twist about each
other and become thicker and undergo knotting. Each chromo
some of the pair (bivalent) e.\hibits a longitudinal slit which shows
its double nature. Each half of the chromosome is called the
chromatid. The bivalent thus consists of 4 chromatids. i Ins
is the pachytene stage of first prophase (Fig. 4- I4C). Interchange
of chromatin material may take place between the chromatids dur
ing this stage.
(iTj) The modified chromosomes of each bivalent again become
distinct. They separate but remain attached at the chiasxnata.
This is the diplotene stage of the first prophase.
(u) The bivalents move to the periphery of the nucleus. The
nuclear membrane and the nucleoli disapppr.^ This is the final
stage of the first prophase and is called the diakinesis.
First Metaphase. The spindle now appears in the region
of the nucleus. The four bivalents become arranged at the equa
tor of the spindle. The double nature of the chromosomes is evident
but the chromatids do not separate. This is the metaphase stage
of the first meiotic division (Fig. 4-14D).
^jg. i-U(A—Hi First Anaphase. The two chromosomes of each bivalent
P-Phass With eight separate The four whole chromosomes migrate towards each
opposite" pole of the spindle (Fig. 4-14 E). Each chromosome has a
longitudinal split showing its double nature.
First Telophase. The chromosomal group at each pole of
"■ •SSi"™""'""' >-™ .f. the spindle receives four chromosomes. This is the haploid number.
The first meiotic division is thus reductional. Of the four chromo
somes in each new daughter nucleus there is one mern^^r ot each
homologous pair. There is no cytoJdnesis after the first meiotic
«■ Telophet; °"'^»«P-atPd division.
ivi) Each of the two daughter nuclei at the telophase of the
first division of meiosis receives four chromosomes (haploid
number).
The second division of meiosis follows before the first is
auite comolcte It is mitotic. It starts with the second meta'.
phase. Ihvo' new spindles appear in the cytoplasm of the spore
 botany pgr degree students—Part in : bryophyta MARCHANTIALES—RICCIACEAE ; RICCIA 57

nucleus"ringe is the simplest among the liverworts. It 'Z^nhericI'^^n°omhne''

spindles. The solits in u of their respective
the late prophase of th^ fi f ^ which appear during 1!S-..KS» WK ~ -
the second Ctaphase Th!.?
chromosome. Each half of the rl
the second metaLLe th^^'''^^^^ ^ chromatid. of it. Before the spore mother cells divfoe to lo m spores the smg^^
separate. Thev thpn m chromatids of each chromosome layered wall of the spbrogomum d's^tegrate .
-co„da„aphLe\V4r4rt,^PP"r " layer of the calyptra also break down. e caivotra There
S?, *e whole chromosomes „• ^ chromatids in a cavity or skc surrounded by the outer 'tty" has no
There is thus no reduction in ?hp '^rate toward opposite poles, are no eUiters. The mature this « The
second division. Each of the r """''cr of chromosomes in the diploid or sporophytic structures (F g. . „ housed in a
'he second telophlse has four T "'he end of mdospores which represent the future gametophyte are housea
are";, T divEfon ^f T'"' '' 'h" haploid sac provided by the parent gametop yte.
the S '"""haneously between the f n ? ™hotic. Walls
is fo mmed.
a Each cell
daughter m♦^
(Fig. 4cell140^" J'"' haploid daughter
of four nucleicells
daughter in Nutrition. Unlike other liverworts
and spore mother cells in Riccta deve op " dependent upon
he sp ofspores for^d
tetrad. Thefrom
Zr,h?f''/P°''= '""'her''^P'°'dcell
spme. Th
is called starch is formed. The sporophyte remains to ally ^epena
the thallus (gametophyte) for food materials, watei and m
P
solution.
can be seen (tetrahedral manner) Tn the

-iS"? ^ Sheath. s-rs «

dorsiventral It bears no appendages scales and is
»spSes Zn a ^P°«= tetrad
ealyptra. ''"em a cavity surrounded by the nm ^he thallus bears appendages in the form ^ in colour and thus
Production c ,
tCuur/s" arnpi^vcm whereas the
Or-,„ _ sporogonmm offifocta more orless a'solid object. The
^ss;M/L/iro/?K individual which produces the the
region
ponsible for sexual reproduction. diplophas
. haplophase whereas theour stand to
These differencesjustify the^^^p^
sporogonium of
^outgrowth
Riccia as a new individual v
of the parent gametophyte (thallus).
BORAGE RE&ON Dehiscence of Sporogonium. The sPO'Ogoniu^;^f^^^«^^^^^
OUTER iAy^l^ never dehisces. The spores are hbera ed by the d y
OF
CALyPTRA
Thlfiln'ii'fc p"'W
onset of conditions favourable for growth.
""""
CAVny
Structure of spore (Fig^^ 4-13C
^'S-415- Riccia sp. Mature pyramidal in shape f^ood is stored largely m the
sporophyte. ing a small haploid nucleus.
 ■ULL

MARCHANTIALES-KICCIACEAE : RICCIA 59

»•jm, «..»..a spores orJ?iccm enter upon no resting period. They germinate m
ine outer IS called the exos- about 6-10 days. Prior to germination the spore absorbs water
and swells about 10-20 w. According to rv
Campbell (1918) the black exosporium and
the mesosporium rupture at the tri-radiate;:
ridge. According to Pande (1924), Srinivasan .'
(1940) and Udar (1957-58) the swollen
spores may become more or less transparent.
A prominent pore called the germ pore
appears on the outer face opposite the tri-
radiate mark. Thin colourless endosporium
enclosing the contents grows out through
the rupture (Campbell) or germ pore (Pande,
Srinivasan and Udar) in the form of a small
outgrowth. It is called the germ tube
(Fig4T6A). The latter may grow rapidly
into a long, club-shaped structure. Often
the germ tube remains short and broad.
Mostoffhe spore contents migrate to its
distal, swollen end where chloroplasts
appear. A transverse wall appears separat
ing a large terminal coll at the distal end of
the tube (Fig. 4-16B). Meanwhile the first
rhizoid appears at its base. Udar reports
that the fi rst rhizoid makes its appearance
from the base of the germ tube when it is
one-celled. It is not separated by a septum.
According to Campbell the first I'hizoid
appears relatively later. It is separated from
the germ tube by a septum. A second septum
Various stats ^ parallel to the first establishes another cell at „ Riccia tricho-
the distal end (Fig.4-16C). Each of these ^arpa. A later stage in
s.

an apical celj
two cells next divides by a vertical wall ^ermi^tion
followed by a second at right angles to the sp
Campbell) first.
flatt thf
In thispndway eight cells are(Fig.formed ^ arranged
the flictnl
distal end of
ottneg^
the germ tube The four4-16E).
cells ofIneyare
the proximal tier
'^PoH^i-^^'eone. Svid:bV%ransverse^
^ its flat cutting faces. From the segments thus cut off
tL tissuL ofthlnew thallus (Fig. 4-17). The single apical cell of.
Ihe ™ung thallus is soon replaced by a row of apical cells.
STTMMARY of the life
ThflifecvcleofUwcia consicycle of riccia
sts of two vegetative(Fig.mdividuals^
418)
Of spot t?'"*'-" The hie cycle
Oneofthese the other The most conspicuous and
istheparent^onheo^^^^^^ green thallus (Fig. 4-18, 1).
^Pecies of studied .er^i • dominant ^ object dorsiventral in construction.
concludi It IS a small, flat, gic^ J
^ that the
 60 BOTANY FOE DEGREE STUDENTS-PART III: BRYOPHYTA MARCHANTIALES-RICCIAGEAE : RICCIA 61

sTs.: a
It has a dorsal furrow and Tdl? obcordate.
nal depression is lodecd the tr In the termi-
thesutaratum bv unirnlf
two kinds, smooth 'k®
wXd andToh''" The thallusThey
is fixed
are toof mature o.osphERB
RBADV FOR FFRTILISATION
water and minerals from the damn The rhizoids absorb 3
gtcws. The dorsal gre?„ ~oPfre rb°^^ ARCHE60N/0M
ventilated as it possesses i spongy. It is amply
of deep narrow
chambers. As canals
the upperThe ktm or"ri!°T
nortbn ' i" into
l™atlcn the form
air-
'heBe green cells arc bathed whh al^L fb *1? ^nd CAlAFTOPHyTF
(THALLUS) SPERMATOIQIDS
anthfridium
Th"r„ '-Se amtnt":ffrd' A 1

P™^~Vgatete'ln^
plant. The gametes are orodnrpd • u""'^gametophyte
called the ^'=P™-
male sex organ is called the anth" .^y^tmellular sex organs. The
['T'= ^■■'^' Ogonium (Fig. 418 Tlf'T 2) and the
acronM 7 5«tpar!te cavifi furrow at SAMFTOPHYflC
arc •" '"™'" "ttt gtowinglrex ^'■o'^™'='°P<=d in rows in 6ENERATI0N{IN)
globular or dioecious T""'
The bcdv , P^^''"®itaped body elevated on ' T anthendium has a
androcytL
418,'3^Th
(Fig. 4d8;3)
(Fig. Thf i^ket of'^trire
) The"'^ etlh"' ""'""''"'ar
^^overalhunSedMa^^ ' ' stalk.
nroa lonfr
consists of I n ^^chegonium is an flask-shanpfl
fled, u "^ScHate sperms
spe:
the venter an enWerT? i H WUCLEUS
Si: r' Ti
jacket wall the venter h ="wi
Portion called SPORE
f nucleus

b™
axial cells b'S'
a veCai ca Thos r;: ,ri2
cells exccDtinp. tU
Matter S
contains f tL^ "f^"-mottle egg and
®®ck canal ii
"" 1^^=^ 'hick
^ of
IZ

inature. !„ tlii! ? disintegrate whi i of OOSPORE

, . All the strucmirTl^'^"^'"^ ^ts'irf^ed' SPORE TETRAD
SPOROPHYTIC.
S" GEN£RAriON{Z N)
starts^rvl
are the last^f^fspo^es'
structures anf
mtr] I ^^orL'^ertiUzadon
and
res constituting the tr ^ during the nha^P An gametes SPORE MOTHER CELL TWO CELLED
STAGE OF EMBRYO
chromosomes in their n i"^^*°Fhyte generation I struc- to
l^gametophyt^Ten"!^!-
life cycle Tv. Scneiation ^uU:r^:\^!^^.^-mber"f
ic thpi^r ^i^o called lio i • ■,
CALYPTRA
^JNERATIHG XALYPTRA
XAPSULE ^AMPHITHECIUM
WALL
\f
^^^ENDOTHECm ^iq^cELLED
^SPOROGOMUmSTAGE OF EMBRYO
'SPORE 8

9
g've rise to th? o„ ge,.„i. 9 mother cells
Fie. 4-18. Riocia.
Diagrammatic representation of the life eye e.
 MARCHANTIALES—BICCIACBAE: RICCIA 63

62 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

develops into a simple, spherical object called the sporogonium

vertical rows of green cells.. They are empty and co^^
(Fig. 4-18, 8). The diploid number of chromosomes characteristic of
the zygote is carried over by mitosis to all the cells of the sporo
Sli "ss£:iXt
cells.
. w-.
gonium. The latter therefore is a diploid vegetative plant. It is
the second individual in the life cycle of Riccia. It is radially 4. The ventral portion of the thallus consists of colourless,
constructed and is solid before reaching maturity. It is very simple compactly arranged parenchyma cells.
in structure. It is incapable of self-nutrition. It is nourished by
the parent thallus in which it remains embedded throughout its life snecies of Rtccia are monoecious and othe«
Bemg diploid it represents the diplophase of the life cvcle The dioeiouf. The 'sei organs lie in the mediatt dorsal furrow each
sporogonium has a wall of sterile cells one cell layer thick It en- in a separate pit.
cbses a mass of spore mother cells (Fig. 4-18, 9). The spore mother 6. The pear-shaped or ovoid body of ^nm"som"
cells are the last diplcid structures produced by the sporogonium. has a jacket layer of sterile cells sui rounding
fuores E4TR n P™duces four what coiled, biflagellate sperms.
sS klvtd ,a'l T V, '■ ^ sporogonium; therefore, is a non- 7. The autheridiai vvaii bbintegrams^^at hs^
IS ral eR .R t"'" 'he meiospores. It is as mucilaginous fluid containing sperms v „
^011 tS fththe sporophyte
constitute sporophytc generation
plant. Theorstructures
phase produced
It starts by
withit antheridial pit at the upper surface of the thallu . ^
proSrf duLT"^'thrs"''h'^°
sporogonium spore mother
fnd the spoTe mrn^er ceils. The structures .
lllT'Th" layer 8.of sterile
The jacket
typically
cellsflask-shaped
surrounding an axi ^^o/cem.Above
yg^ter. °The
produced from the spore mother cells are ti; fi basal cell of the row is called ^ egg. ofthe axial row
next gametophyte generation PirS first structures of the it is the ventral canal cell. The remainder
(Fig. 4-18,
not the 13) produces
sporogonium the green tMus 'nr
(sporophyte). ^ w(gametophyte)
'''' gerjnination,
and lie in the neck region and are called the neck canal cell
From the account given abn\7/» u i 9. Moisture is essential for the maturing of sex org
fertilization.
cycle of a single iStccia plant there orrnr^?
individuals. They are the green inrlpn
phyte and the colourless spherical
distinct vegetative
thalloid gameto- 10. The sperms swim to the ^i^^fl'^JJ^t^er'the^open necks of
water in the dorsal median neck canals to reach the
phyte (sporogonium). These two InH' M constructed sporo- the archegonia and unites with the egg to accomplish
other generation after generation occur one after the venter where- only one ot tnem uiuu
the other. The reprodu^ctivrSs of on'e ^ fertiiization. ^
generation. This nhenompn • Seneratton produce the
11. The diploid /^^r^^f^^^^iticellular sporophyte.
of Generations"
and ail other bryophytes. The two 1 " '=''P''<=^rod by tiie
Mature of phrase
Ricoia undergoes cieavagf to form a sphencai, mulnce
IfeAiternatton
cycle of offfiodct are
Generations" is called £.1 jndividuais kind
in the
of 12. The sporophyte of
worts It is nothing but a j Uy +i^e parent thallus. The
Teta nor a foot. It is nounshed by ^dte^P ^ close invest-
two layered venter now „chegonium may wither by
,nent around It The ,,,ature^he single layered spo^^^^
this time.waUByand
gonium thethetune
innerthelayeiSP""®
of th™ calyptra
W disintegrate.
gmer layer ofThe
the
or a.co..A
soores lie free m the space „f\he diploid, sporophytic
calyptra. At this ^»Se th«e rs n gametophyte genera-
'urfact"'™''"
srL'iS'. r," -»=*-■
3 on its ventral
chambers occ'ur as
a? deep, narrow airofeana
the thai
, In c"■ "^kambered.
v The rounding thallus tissue.
als or channels separated by
 64 BOTANY FOR DEGREE STUDENTS-PART MARCHANTIALES-RICCIAOEAE : RICCIA 65

Ill : BRYOPHYTA

discussion
(t) Loss of assimilatory filaments in the ---
(t) Loss ol assimilatory niamcii« .w
^' . - . , 1
air chambers
3
- ,
r^Cfhp- :
,

rather an evolutionary Liverworts. It is (it) Simplification of the barrel-shaped pores of the advanced
Marchantiales. ... Vr^,.«
plex structure. The sporonhvtP Sametophyte is quite a com-
among the liverworts It i/ the simplest known (iii) Shifting of the terminal ganaotophores.^(anthen^^^^^
nutrition. It lacks both font ^ spore case incapable of self- and archegoniophores) to the dorsa pwi ^ gametophore. The
layer m thickness. Within thi. capsule wail is one
spores. Some of the spore mo her rln "mother cells or
aie looked upon as the fore-runners degenerate. They
theenmore advancedRiccia
to regard members of TL
sporoohvtp or elaters
h^archantiales. of
Are we nium and involucre. reduction in structure
structure? Porophyte as a primitive or a reduced All these changes are accompany elimination of
Bower, Campbell and P., of the sporophyte. Tjie first step Consequently
of tile S 's primitive. ^The^'*^ simplest sporo- seta. This is followed by disappearance^^
appear seriated illustrat^"'"^ complex sporophytes
the sporophyte becomes a sporogonial wall. Subse-
hv tK such a? They »hrumTeUr.;Lm^ This step ehnu-
fertile cell^ ^ S^-adual and progressive sporophyce as that of nates the elaters.
i-emain sterile and^ "mother cells). InsteaT
spore dispersal, of
functLn^"!? revision questions ,.
sporophvte \ntr. r ' resulted in tu nutrition, 1 I.ist the oharaoterisde features of the fanrily Biooiaceae and g.ve
'h= poteadal ^»d catuji a, of '^"^"fTt'rhrof-eriesof diagrams write an aoceuut of the Ufe
help in spore dispera?*"-''';"' ''ooome sterUe Some of
IS called the theory' ''^"'hcsis of the th? • ■^^'"'^TtisUn a tabulated fo™ the dlffereuees between the gametophyte
cvoluiion of the snnrLf^ ®*®"Iisatioii. Tr I ^tnment botanists
the starting point o '^^crworts vvirh^
oftUspo;-op?vt.'^'r'^f^^'^'"yophytes
^f thil\hTor, t''"" ^P-^ophyte as u„dsporopl^-f^i-^hesporlu.t How does it originate and what does
The series begins vhh p fnr evolution or the it give rise to in Kcc.o » „y„,e,er,HiccVa.
The theo" V ^P-oph'yt'e'" -sending series. . . :: ^::rrhC::ssof spore foruration and- evointionary position
plex nature of ihe ^PPealW f.:u disobarge intho sporophyte of iJMCiu.
basi3'7'thL'"rh the com- . ■ Descrihothe varrons^^^^^^^^^^ forms hke
BiSia are reduced and not primitive ! ^ ^ sporophyte or a
»»bination bf' 8. Analyse.the ™'--XgMng "og»t reasons in support,
garoeiophyta! Justify your answ 8 ^ following statement: "The spo-
holds
than primitiv
other h I
sporonvf^ ''^®heves th
'
tion) resuhed byX '^^duction
\ "^^'^^ains
(ak-.th^ , ^Iccia
simni t *'®^'iced rather
""'^°:r'"acethe deve— of
member of the Hepaticae yon have stndie .
and archegoninm in any
Phy.to):^;Uk' coW Prosi^"^ <-ypes like 72fcc20
^e'-ged hom agenS/''i?ptedthat'iyP°^^e^ sporo-
Ae widespread evident
g^^'etophvt""" reductr®-. genus\?^^byap,
This Church
^ight have
^amplification of thp the i ^ the supported by
which they
-ttch d' prodnee. in tt\and pr^o^c1 c^haracters
"bines of tkf'^^cture sex organs
• ^ ^^buction o sporogonia
series are :
 1
MARCHANTXALES—MARCHANTIACEAE: MARCHANTIA 67
which may be sessile or stalked. The female receptacle is gene
rally always stalked. The sporogonium is usually differentiated

CHAPTER V
MARCHANTIALES—MARCHANTIACEAE : MARCHANTIA
Marchantiaceae

250 sp^da^'^^Thev
species. They are grouped underfamily
aboutis23represented by deep
genera. The over

9 (iFCEPTACL£

"A/.- M/OR/a

PAR'<
gemma
CUP
STREAK

Fig. 6-1 (A. Fig. 5'2 (A—.B). Mavchantia nepalensis.

A. Female plant. A. Female plant.
--

Male plant.. B. V.S. through a pore.

green, dorsal surfacpnf.i , Pl""!-
« marked by into the foot, seta and capsule. The latter produces both sp^s
'.vith or without Dh't and plater^: Vhe capsule dehisces either by the separation of a li^
^umorhera in which ?u ^^y^^hetic fiu'J "dually well developed or by valves The family includes a number of genera such as
condition
POKs are and -
well developed'"'TK" are r Exceptions
""'V ® are D^J^ort^era Lunularia. MarcUntia, B^boulia, F^mhnana, Targ.oma.
^hesexorffanc are united
backing. The air
in receptacles
QrimaUia 'Plagiochasma. ete. The most comrnon and h.ghly
66
differentiated of these is Marchanha. We shall therefore study
itarcha-ntia as a type of this family.
 fob degree STODENTS—part hi ; BRYOPHYTA
MARCHANTIALES-JIARCHANTIACEAE : JfARCHANTIA
Systematic ni7 •■
Positmi
Sysiematic Poski 69

Plains, Calcutta, Assam, South India, Lahore Sialkot, etc. It is

^ryophyta distinguished by the presence of a median, dark streak on the dorsal
Hepaticopsida surface of each lobe of the thallus. The lobes are long, narrow,
Marchantiales margin entire and apex emarginate.
2. 4,1/. nepalensis (Fig. 5'2). The median dark streak is lack
Marchantiaceae ing. Lobes are short and broad and margin generally crisped. The
apex is emarginate. It has been reported from the Outer Himalayas,
MARCHANTIA (March.) L. Punjab Plains, Kashmir, Garh\val, Kumaon, etc.
t .ft
3. M. polymorpha (Fig. 5 5). It has been reported from the
hills such as the Outer Himalayas up to 8000 ft., Pangi, Ladakh,
Leh., etc. The thallus is relatively large, broad, slightly concave or
flat. The margin of the thallus lobes sinuate to lobed.
GAMETOPHYTE PHASE
It begins, as in Ricda, with the maturation of spores. Each
spore (Fig. 5-22A), on germination, produces a green thallus (Fig.
5'22F). Externally it has the same general appearance as that of
Riccia. However it differs in its large size, coarse nature and more
prominent and pronounced elongated lobes. It is somewhat better
adapted to grow on land than Riccia. The green thallus is the
gametophyte plant.
(a) External Morphology of the Thallus (Fig. 5'3A). It is
a dark, green, somewhat fleshy, flat, once or a few times dichoto-
mously branched structure
with a dorsiventral sym- // / /1 \ Y /
metry (Fig. 53A). Each
COP
« ' thallus branch or lobe is
traversed by a broad, thick,
central midrib. It also has a
V \
notch at its apex. At the
'AIR POfiE
(a i bottom
located
of the
the
notch
growing point,
is

'Thallus lOBB The upper surface of the

*AF(BOiAl thallus is marked by
rhomoboidal to polygonal
'OARK areas called the areolae
(Fig. 5-3A). The boundaries
t<r> these areas mark
between Fig. 5'4 (A—B). ilarchantia sp.
the limit of the underlying A. Smooth walled rhizoids.
air chambers. Each area
B. Tuberciilate rhizoids.
has a tiny conspicuous hole
or an air pore in the centre. The pore is visible on the surface -as
a light dot. The air pores permits aeration of the thallus with the
I,
minimum dehydration. At times little cup like structures, the
f^iD RIB
cupules or gemma cups (Fig. 5'3A), are seen on the upper surface
of the thallus. They arise in the midrib region. The margin of
1? the cup is toothed and membranous (Fig. 5-5C). From the bottom
of the cup arise numerous, green, flat, multicellular bodies called
the eemmae The mature gemmae get detached. The detached
gemmae are washed on to the soil where each gives rise to two new
gametophyte plants.
„'7^' ^rface.Burf,
""'he ventral > ■ 1
'0 BOTANY FOR DEGREE STUDENTS—PART in ; BRYOPHYTA
MARCHANTIALES—MARCHANTIAOEAE: MARCHANTIA 71
^ When the thallus attains sexual maturity it bears gametangia
bearing upright, stalked, umbrella-shaped structures at apices of From the lower or the ventral surface of the thallus arise
certain lobes of the thallus. These are called the gametophores numerous elongated, single-celled, hair-like outgrowths, the rliizo-
or gametangiophores. They are of two kinds, antheridiophores ids (Fig. 5-6). The latter anchor the plant to the substratum. In
and archegomophores. The former bear antheridia and the latter addition they absorb water and minerals in solution. The rhizoids
archegonia. two kinds of gametophores are borne on different are of two kinds, smooth walled (Fig. 54 A) and tubercu^te
a 1 so t at Marchantia has male plants distinct from female plants. (Fig. 54Bj. The tubercles are pronounced, peg-like mvagmations
of the wall. They extend across the cavity but do not form com
plete partitions. The tuberculate or peg rhizoids are thick-walled,
narrow and appressed to the surface of the thallus. They arise
from beneath the scales and function as a capillary conducUng sys-
MAl£
y Ss ORasc tem which serve.<: (o carry water to all the absorptive ^^
mpTAasl^ RhOmbC'DAI thallus. The smooih-wallcd rhizoids stand out froni the ^ u
AREA ARCh[GONIOPhcR£
and penetrate the substratum to absorb water and to fix the thallus
to it They are broad, thin-walled with colourless contents. They
*^'R£RlDlOPRCRf are the first to be formed on germination.
the ventral surface bears purpUsh. flattened
are usually arranged in two to four rows on cither side of the mid
SEMtM CUP rib They are never in a single row as in B^ccta. Each scale is
plate ofceilsone celMayer thick. f
. 'VtO PJA «
divided by a narrow constriction into two parts, the body and the
MIO R/B
appendage {Fig. 3 IC and D). In some species the W^l^ag^ ^
absent. The scales are separate from the beginning- They a e
not formed by the splitting of a single lamella as is the case m
Riccia.
(b) Internal Structure (Figs. 3-2A and 7';•„Is^omn^red
terintynal elaboration of the thall^^^
with Riccia. In venical instead

photosynthetic region and (in) the storage region.

S-T sLir/aycrofthin-wal^ cells with slightly thickened outer

walls The epidermal cells contain few chloroplasts. The outer
walls of the epidermal cells are practically water-proof The epider
mis is thus pr otective in function. It tends to check transpiration
from the underlying tissues. Embedded in the epidermis are special,
chimney-like or barrel-shaped air pores (Fig 5'6). Each pore is
surrounded usually by four tiers of four or five cells each. The num
ber of tiers, "however, may go up to eight. These tiers form rings
ofceilsone above the other. They are arranged j" ^
small chimney which encloses a wide passage (Fig. 5-6B). Th^is
passage is broad in the middle and narrow above and bebw. Ihe
o' f""!" pW cups. pore lall lies half above and half below the ep.dermis. The open-
D. cup!» fng of the pore is thus slightly raised above the surface rf
^ ^ pore thi tLllus. The cells of the lowe^ost tier m some speems are pap h
lose /Fifr They project into the passage. However, they
do nit floL u: A staVs'ha'ped channel remains open. Each pore
lowerinoBt tier.
%
 I

MARCHANTIALES—JIARCHANTIACEAE : MARCHANTIA 73
72 BOTANY FOR DEGREE STUDENTS—PART in ; BRYOPHYTA

leads into an underlying chamber. The presence of air pores in the

i progresses gradually inwards. For a detailed discussion of the
epidermis facilitates gaseous exchange necessary for photosynthesis origin of the air chambers in the Marchantiales refer to page 42
and respiration. Apparently the pores bear resemblance to the (Chapter IV).
stomata of the higher plants. To them they are only analogous. (iii) Storage Region (Fig. 5'6j. Just below the photosyn-
The stomata are always absenton the gametophyte plant. The cells theticiegion lies the ventral storage region of the thallus. It is
around the air pore do not regulate the opening as the guard cells
do in the stomata. Walker and Pennington (1939) have reported I- thickest in the centre. Towards the margins it is reduced to 3-4
layers of cells in the thickness, It consists of a uniform tissue made
that the surrounding cells by imbibitional changes in their cell up of relatively large, colourless, thm-wallcd polygonal, parenchy-
walls bring about opening and closing of the air pore. matous cells which usually lack chloroplasts and are compactly
(u) Photosynthetic Region. Beneath the upper epidermis arranged. Most of them contain starch and protein grains. Iso
are the air chambers. They are of fairly regular size, simple and lated cells may contain a single oil body or may be filled with
distmcfc from one another. They are arranged in a single horizon- mucilage. The former are called the oil cells and the latter
chambers are bounded by one cell layer thick mucilage cells.
partitions. The partitions are three or four cells in height. Each
chamber communicates with the exterior through a barrel-shaped The lowermost layer of the storage region is
similar to those of the upper epiderrnis. It rs the'<7"
chamberIS t Nrnm it nroicct the rhizoids and the scales.
leTatatLSrl below the thallus^ 1 he scales tieip to
°
Story or IT!- assi!
cavity o7the C&7
the photosynthetic filament-? r ^ °
They nearly fill and
the grow in much drier places as compared with Etccia.
ir\ rotrmarison with Riccia. In general form, stru^ure
Even the cells of the overlvin7" !? numerous ovoid chloroplasts. and developmmt of the thallus Marohantia resembles Ricoia. Both
plasts. The chambered ,I™" uhloro- resemble each other in the following features .
centre
iu dim oflight.
photosynthesis il fheof,77"
Th7 arrangeSem th the principal
rite
«t,te thai,us Of Wnll7th^u7e dtrSlS^'t^il
cUntia, cellular scales,
puces. They are foiTned by the
PORE
'UPPER EPIDERMIS
qplorophyuose cells of each thallus lobe.
AIR CHAMBER

1
STORAGE REGION
MUCILAGE CELL

SCALE
LOmR EPIDERMIS

smooth RHIZOIDS
i ,i, Its definitely larger size, broader, thicker and coarser lobes,
(ii, Better adapted to grow on land than R,com.
X^'JGERCULATED RHiZOfO (iii) Prominent expanded midrib.
(iv) Upper surface of the thallus marked with rhomboidal areas
each with a distinct central pore.
A 5*6 \i
splitting of section ®P-
(1,) Internal differentiation of thallus into three distinct regions
involution. Th^ (Scbi ^^^allus. (epidermal, photosynthetic and storage) instead of two (photosynthetic
and storage) as in Riccia.
;s»i -.«K,
y below the epidermis.
 75

74 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA MARCHANTIALEb^i^i^^ - ^ ^f cells reaches

.. •. of cells
{vi) A definite layer of air ,chambers with photosynthetic fila
ments arising from the flopre.
thallus rot and disintegrate.^ P develops
dichotomy the lobes
When into anoT the detached lob«
mdep , - ^
(vii) A continuous water proof epidermis with definite barrel- by apical growth and dichotomy dev^^^
shaped air pores. In this way there of the new terr^
particular area and con firanches These may develop
thp ^ four rows on either side of
RW?.Iey are arranged in two rowsmoisture
one near below the thallus.
each lateral margin-In
(i^) Presence of gemma cups containing gemmae. from any part of the hailu^^^^P^j ndventioous b™tches^^^^^^p
vanced^Sn!h.?7p— ilfarctem thallus is more ad-
The presence of scales on the ventral
epidermal laver enabk Mor ^ continuous well defined
compared wTh £1 %Z ^^^itats as RHIZOIDAL
below the thallus which can 1 ^^P moisture - C£US
epidermis helps to check tran.n" rhizoids. The
synthetic
Marchantia better adapted to a lifl ^hich make
moisture is still required for its =. ■ ®"'? abundant 6RE£N cells
Apical GroU TU
notch at the apex of each lobe
fertilisation,
"'=' ="« the bottom of a
cells. By the activity of these the u®®, tow of meristematic SROWINC POIt^T
^ cnese the thallus grows in length
„ , MESODUCTION' NOTCH-'7
and by meiospores OIL CELLS
vegetatively in the growing""""by fe^foll'''""' ll®"* teproduces
-.etativeFragmentation If ^r^the^^o-t
cells. The\iV:el ^ by'^ree methods
the ageing of:-
the ONE-CELLED STALK
'n the rear or basal part of the
Fig. 5-8. Mnrchantiasp-
A single
A single gemma.
g«"^'— , ,
AiSlMlLATORY , . feresting and elegant method
CHAMBER AIR PORE Formation. Another interesti g formation of
^1*'. (iii) Auction in gammae develop mside
GEMMA CUP
-fl
ofveWtatf-
multiceUu ar bXs
bod cailod the
-osvths j
of themargins (Kg.
thallus, 5-5C y.
general
little, sballosv, c P "PP'^''f*u grco"' disc-like, slightly
GEMMA The cups are develop "Xon edge on a short, delicate,
in the midrib reg gemma stan bottom of the gemma
bi-convex stmct the gemmae in the cup are the
""I'Tlivty Intermingled "^'hich sfcrete mucilage copiously.
I rnucilage dew or rain the mucilage
^^IZOlD \THALLUS
With\he absorption get detached from their stalks rather
swells and causes the ge .^^ashed on to the soil or earned by a
A passing
vertf™i\;.„^^^«el>antiasp.
thrnis T. ®fion of fv,
easily. Thcgammaeaie
^ the cerv,^® Whaling
cup.
 76 BOTAKY FOB DEGKEE STUDENTS-PART III: BRYOPHYTA ^rAnrH^NTIACEAE: MARCHANTIA
marchantiades-marchantia 77

grow into new^thflli^onuUable'^s'on^^ detached gemmae away

dueLnof theof central portioylan^
.two -parn n of the I'provides
J,"7ormation'
a vety 'rapid^de««
Ss usually
(t) Structure (Fie- 5-ft^ Tho.
shaped .uUicelluIae steactu^e. I. L TnedTn rZltipSfoPanl a successful^ Na.yanaswattii
experimentally of the gemmae .(S j.
(19571 that the germination ,vmg tips
inhibited by the feretion o basipetally-
parent thallus lobes. A—F) cupules are o
(iu) growing point-
a shortof distance
form back frof
circular ridges gemmae ong ^tial. The
superficial .1"^"° outgrowth. It is a lower an^d an
grows into a papiBat transverse wall r.irther. It forms
gemma m'^al divides divides by a transverse
upper cell {Fig- 5 )■ ^^pper cell agam undergoes a similar

Sfii? |"s, 't. .> tsl"""""""

Sexual Reprod"^" It ^takes place only ^^^^^Q^gand the
and is of oogamous typ^^.^.^y the days
growing season ^ environment is low.
nitrogen content in tne Organs. Th^e sex
A nistribution of Sex ^ ^hes of the
(a) Positio*! an borne on the ver branches are
c, /
organs in ''''"tused for this purpose^ The
5-9. (A_p> ,, ^/ thillus, orhighly
apical termspe-jf^= 3itlon of(Fig^'he ap^"^11aevelopment.
jtohotomy The growthEachm
in tfce d:v;,„p"l7r, from one half °^,i°'^obes ceases ufter their p„ , ^
length of the thdUus ^ conti ganietophore.
S-U"fiefs'^ --hed on .he i„o
the Dn?V^?^"^g point Th ^PP^sue edges Tn u
constituting the h S '"'''''mcni to t?^"
dant chlrt^J 1 °dy of th?
®"°'her ?hallow'^^?i
stalk ]Uo-
"""'•®"
indentation of rhizoids in the g posterior side.
synthetic chambers (antheridia) is
instead of chlo ^vithin ^^ S*'eeu or chlo
flattened snrA. P Thev . ^ ^^rgin mi. ^''^^yniatoiis calledThe gametpphore t^ (Fig. 5-5B)-
antheridio^b ^^^trtheapparatus
TnfheddioS
of sexual
Both the surfa^^^ and the od bodies
*e
the time of ^er
possess isolaw «nate dors
• The S'"P'=tficial cofo"' ."'"''■alsyinmetry.
The two
and the
avchegomop tor«, constitute Teported^"
is called ^'^*^^®^nniophorcs he occurrence
antheridia of
and the
temperature and other environ:^
and is cont'"'?' reproduction. ' ^ botanists have rep^ addition to the
" '""edsymmetry
by the I' t:ells known
hted atas arehegoma. S° arfesandrogynous P-eceptacle
androgynous rec p
bears antheridia
i P ^^.j^eridiophores and
peridulous arehegoma Sexual repro-
the cibirWrv"'"''-
rhizoids. The m"'-''''' Whenof ,1 gemmae fall in pits on the upp borne on ^'^,plants^„jnale which bear the
notches begin to^r"Stems or the face next , ? ^luitable soil, the archegoniophores
Stow stmultan ®ou°rt,g Points y botb^V°" duction thus involves tw
^ "t opposite d- ^ .fte marginal
directions. Rotting
 78 BOTANY BOB DEaREB STaDE.T8-PART m : BRYOPHYTA
chantia is therefore dioMlou"'^''^'' ^"■chegoniophores. Mar- disc. The opening is .f^^^itton "wTth the air chambers.
ofchambers alternate in pos
terminally eptacle with aJ'margin bearing
a rounded
bearing
AIR PORB*p t _ AlRdtAMBBR -,yOUS6£S'f
ANTUBR!^^

0£cePT^<^

Is .AA^PH'^ASTRiP
ASSlMJLAT&f/

RHIIOIOS

.^A/\JTH£RIO'OPP<^^

"Tluo!S?u^h
A vertical sect mature
10,
(.,.) structure "f AutheH^J" V?^?
Cros?.seetion?h" Trth? italfatt^t
-es

^felr£ - ^
Ssues of ®|"' stalk^°Tr' ^ '"''"wsTu?'"?'"'' ™ ^interiGr The protoplast ...hisces on the access of
sperm The ripe antheridium dehisces
ac o concave upp^
. ,r Trpvid^ad by rmn - dew. ^tucf wlti
succession;
succession, 7'':
a
shows that it
of ;ep"e
reprateJ ? rucept^nele
section of'h
^^^^ptade tli? •
dichnh ^ branch
srihiV
eventually
.£• c"i«-
di^u'eg^ of androcytes °»
" .ysi"
r hitrg
the underlying
til/'»^
with
male recem
foim l^^- distal end. of the receptacle mt<, ^
Beneath ihe chami!^'^^'"® contain^^^ The i ihallus.
f w?ter.he walls
Jhyma cells the co2"? ^^e photo.v^L'^'.
Deeply sunk in thf-*^ ^^wer epidp- ^^gion of i
mto
filaments.
Sr sperms are now set ^
They arise inX "PP^rsurfa™ hefrs
Each antheridium &P=tal "rderanT''
P— °^^'7ub?rated- sperm (Hg- ?r'i's"?Toi/gated^nucleus^wi^ the
Each antheridium
Thee latter opens bylies°^^^^' °^der anl^^^^e'^^be
"^^-^apei'
il^'^-^hapert'
<7be are
are tlth ='»«heridia
arrange??
arrange?fn rhizoids.
' ® "arrow ch-,
^'""el P»o„ (antl, -j. " radial row
chamber). end project the
"PPer surface of the

■A»« ■
80 BOTANY FOR DEGREE STUDENTS—PART III ; BRYOPHYTA

be rod-like in M. polymorpha. While swimming it takes the form

of a crawl of a snake with one flagellum lashing forward and the nial cells(Fig- o PpiMARY ^tffPiOiAt
other backward. CELL
anthBR-'O'^ UPPER CBU
.primary STAtM
lOPVfR CELL

primary
PRiMARy ''ALL CELLS
CBUS PRIMARY
ANDR060*RAL
pp/M.S'EALP CELLS
CBLL^ STALK

Fig. 5-12 {A~B). Marcbantia sp. OSTIOLB andpogonial CELL^

A. Sperm after Ikeno. a(^OROGONIAL
■jacket
•'■ CtU^
B. Sperm after Strassburger. l/VAtL

STALH
of
The growing apex of the verv fg,
of the5'13)- The growth
antheridiophore. STAL*^

divides by dichotomy. antheridiophore

The result is the formation of a number of times,
of the disc has a ^ Each
Fig. 5-13 (A
(Dingvammatic) ^
"• . the wall of the anther.-
3ing,e supers-
inidalS;^
two cells (fT^ lob7of
®"tl divides bv „ L Hes 2 or 3 cells
receptacle. The are called t ^ ^ process
lopment. The im 'Ehe lower cell unrlpr"^^^*^^ division into
a basal Primary divides again tra^n^ [urther dcve-
ceU(Fig, 5.i3q^ cell and a terminal „ ^o separate
stalk of the antheriHi undergoes a fe« antheridial
parallel cleavage /o fZ'
ce.ls sometimes more 5T3D^
antWi7?'°"f
S divides by
'he 5"^46^^ The PC°'°P'How°ever no wall is the
vertical walls cells „ three
Thebody ofiheantbt angles divides bv two
of
thefour
tierscells
(Fig.e^h
5-13 V ^
F)'Th?^ ^ ^ ot- ^4 13E)
tiers '^rdeveloP^^cytoplasm K androcyte at this stage
slightly and devei
cells enclosing a „assonnneror'c'''°"'' PPcarTi.m all the ^^yer
^ or central cells cells of
of
body appears
• -the outer cells are
 82 BOTANY FOB DESBEE STODBNTS-PABT III : BRYOPHYTA
MARCHANTIALES-MARCHANTIAOEAE : MARCHANTIA 83

hofds that it tateL ParTt°nX'dtl;~ specially modified like the antheridiophore. It consists of a stalk
bearing a lobed disc at its distal end. In its external and internal
structure the stalk resembles that of the antheridiophore. It is
however slightly longer. It may be 2 to 3 inches long. Typically
the famale disc is an inconspicuously eight-lobed object. The
growing apices of these lobes bend downwards and inwards towards
the stalk. From the margin of the disc between the growing points
grow out long cylindrical processes. These are called the rays
(Fig. 5T5). They are usually nine in number. The rays give the
mature female receptacle a stellate form.
(i) Position of Archegonia. The archegonia in very young
receptacles are borne on the upper surface of the receptacle with
tl)eir necks directed upwards (Fig. 5T6 A). They are developed in
acropetal order and stand free from each other in eight radial rows.
The youngest is near the growing point. There is one row to each

s'tagesl ttoT'""""''P'
^-et,ho,d3th:;t::;rr""^^^^ AIR PORE

branch that (Fi^ S-H a j tj P^^^oplast. AIR CHAMBERS

STALK

'wonKne 6R00VB PERICHAEWM

OR OR INVOLUCRE
^^'^KHAtFSm

RHIZOID \
THALLUS
Fig. 5-16 (A—D). Marchantia sp.
astr/r^ Longitudinal sections of various stages of development of
the archegoniopbore. (Diagrammatic).
lobe and 12-14 archegonia in each row. At this stage the stalk of
archegoniophore is practically absent or very small so that the recep-
 ' r inii^

MAROHANTIALES-MARCHANTIACEAE : MAROHANTtA 85

I jI I it Beneath T are the air chambers arranged in a

84 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
' . f ver From the floor of the air chambers arise the assimi-
tacle appears sub-sessile. The first formed archegonia in the rows
reach maturity. Fertilisation takes place at this stage. After ferti
TtS; fiirmentl
as are found in theThere
male are no pits alternating
receptacle. _ with the atrThechambers
arche-
lisation the stalk of the archegoniophore begins to elongate. Associat
ed with this is a rapid growth in the central sterile region of the
upper surface of the receptacle. With more growth above than
goniumlflfaXuiut PRMARYAmi
b"' "i'^bnct multi-
below the cap, the growing points become gradually pushed down
wards and inwards towards the stalk (Fig. 5T6 B and C). Consequ
ently the archegonia become transferred to the lower surface. In a
mature female receptacle they are pendulous (Fig. 5-16) and
thus hang downwards from the under surface of the disc with the
youngest near the stalk. The inversion of the archegonia is accom
panied by the development of a plate-like tissue on cither side of PRMARV COVER CELLS
each row of archegonia. It is the perichaetium or the involucre ■CK CAHAL neck canal
CELLS
CELL
Fhe perichaetium is a two lipped, curtain-like
rue ure. It is a single layer of cells in thickness. It grows and PRIMARY,
VENTRAL
VENTER VENTRAL
CANAL
CELL
under surface of the lobe and is CELL

^ ^ archegonia is thus separated

the^inpr<!n f rI r In the meanwhile from
prLeS^ disc develop long, stout, green cylindrical
Cs of archel ■■^dially between the
ouTandTn a are generally nine in number' They extend
tiny umbrella At u ® downwards like the ribs of a
female receptacle a steUate\LLh'ap?d7:ppettt.^^
structure as the male^^^TherTifthe
Anere is the iLV' internal
upper epidermis with air pores
Fig 5-IS (A—L). Marchantia sp.
A-J. Stages in the development of archegonium.
tR CHAMBER KandL. Mature archegonia. (Diagrammatic).
■epidermis M l stalk.
cellular + lU The
The stalk
stalK attaches the archegonium
archegoniumto the under
consists of
■rfMALE surface of the , 7^/neck The venter forms the swollen,
RECEPTACiE two pans, the venter ^nd ^ ^ continued into the long,
basal portion of ^ consists of a wall of a single
slender, tubular neck , cavity. The latter contains
layer f Jgg '!
two cells. The large the smaller ventral canal
surrounded by a fluid. venter is continued into the jacket
ce/(. The jacket or waU^cf^theve^^^^^
of the neck which co , • i
neck cells
The neck cells enclose a central
characteristic of the N ^ j which contains an axial row of about 8
ARCHEsoaha
R^R'Chaetium canal -^ed the neck can^ whrch™
neck canal cells 1 ^.^ter of each archego
^CNECowOfiHQfir
nium is Xvekped'a collar-like structure. It consists ofarmgof
A vertical Becttan an 'P"
 86 BOTANY FOR DEGREE STUDENTS—PART HI: BRYOPHYTA
MARCHANTIALES—jVIARCHANTIACEAE : MARCHANTIA 87

It is called the periey- 3. There is no inversion of the recaptacular surface of the

stimulated afttr fe^rSfon '(Fig'^S-ltTanrB)"'^ The is male receptacle.
4. The antheridia in the male receptacle lie embedded on
and forces apart the lid ceils. mucilage which swells the upper surface in pits and jn rows radiating from the centre.
The archegonia hang from the under surface of the female recep
(««)
gonium Development
arises from a singleofsuperficial
Archegonium
dor^l(Fm
c^U 'S-lRl
Tt r Ti, ■ u ^\ tacle.

to the apical cell of each lobe nf tV,/..° lies just close 5. The male receptacle lacks accessory structures such as rays,
the archegonial initial (Fig 5-18aT^ It is called involucres, etc.
enlarges. It then divides into in • archegonial initial (iv) Fertilisation. The sex organs in Marchanlia are develop
outer pHmaryarcheg:;^^^^^
ceH undergoes a few irregular divi.lonc ^
cell and an
primary stalk
ed in receptacles elevated on long stalks. Besides they are borne on
different plants. These facts present difficulties in the way of ferti
stalk of the archegonium. o form the short but distinct lisation. Because of uncertainty of fertilisation reproduction by
vegetative methods is very common, efficient and effective.
vertical inte?s™fagwS'r®Thes"
surrounding the fertile cell in the ^77 ''I'
Peripheralsuccessive
initials Fertilisation is dependent upon the presence of water and is
-called the primary""ial <%■ S'lSC). The latter possible when the male and female plants grow together. It takes
axial cell an upper, smaller primarv^^ ^ transverse cleavage in the place when the plants are wet with rain or dew. Possibly the
antheridia and archegonia mature and fertilisation occurs before the
nerinh'"' i "?'' elongation of the gametophores. The sperms may reach the
peripheral initialseentral
divide cell
-l-be jacket initials and
(Fig s-mnT
longitudinal v^ t' f A n-' '"^Ptfated from
the three
six jacket initial*! archegonia in the following ways :
Mi
archegonium now consists transversely The 1. The sperms may be splashed by rain drops from the sur
outer six cells of the upper S'lSF): fhe face of the male on to the sessile female receptacle.
2. The sperms splashed on to the ground by the rain drops
from the male receptacle may actually swim the whole way to the
archegonia in wet weather.
3. The sperms may, swim through the water when the male
and the female thalli are actually submerged under the rain water.
The above mentioned possibilities can become realities only if
the male and female plants grow near each other.
canal cell At tK/% oy the repeated divit;io« r i ^ The neck
4 There is possibility of the sperms being carried by the
orwhile
lid the
cellstransv#>r
forUX X",' - anrim
of the S P™ory neck
of four cover agency of microscopic animals such as the mites.
venter T! of the venter"^ "7"' Mean- A number of sperms may reach the female receptacle in the
The latter con.ahs
larpr egg or ovum Th
a" m^f It ^cl^
ventral canal
the P-^-ed
I «avity.
a above mentioned ways. From the female receptacle they are
attracted towards the archegonia. The source of attraction is some
cell (Fig. 5.J8 j arc from th^ • ^ chemical present in the drop of mucilage that oozes through the
receptive spot. has in its im ventral open necks of the archegonia; A number of sperms may enter the
DlSTlNGTTnxTCT.r.ey„.._ Part a distinct open necks of archegonia and swim downwards in the neck canals.
Normally one unites with the egg in the venter.
Post-fertilisation changes. Following fertilisation the
stalks of the archegoniophores elongate. From the upper surface of
- . .irrisr. the disc develop, green cylindrical processes called the rays. A
number of archegonia may be fertilised in each receptacle. All of
them however, do not develop into sporogonia. The act of fertilisa
TA,u , fpme.1
^ "^ale .^^^Piacle
receniani is• -> a shorter than
whereas the tion originates the diploid condition in the nucleus of the zygote.
at maturlt, a scal.opea edge The zyoote secretes a wall around it and enters upon active seg-
and star-shap?d mentatfon to form a spherical embryo. The venter enlarges with
 w9P

88 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA MARCHANTIALES-MARCHANTIACEAE : MARCHANTIA 89

of embryo. A cylindrical sheath called the perigynium Develooment of the Sporogonium (Fig. 5-19). The zygote
Hr»n.=fi of each archegonlum and surrounds it. A two- probably starts dividing within « m
ward on either side of a row lower
archegonia.
edge fringed also grows down
SPOROPHYTE PHASE and the inner or f,om the outer or
which consntuies apex of sporopliy , exoscopic. The next
dipioph^se zygote the sporophyte or the epibasal region embryogeny is globular embryo at this
effiyranS h5 l.n
are the last structur^es o^thrs'^se.^"''^ zygote. the cleavage is at right ang^s to the first.
''''^^io^T1^
variations in ti?e""earW
tlie eariy development
i . of the embryo in different
^ three-celled fila-
species. McNaught repoited Tvoicaiiv the next division
mentous embryo m C) Only four cells are visible
produces the ^ vigorously.
ZYGOTE ¥i:Sns
IgCa Lcirwv. -
PiRisymM ^ - - 1 »_ ^ c4 V* 1 \f

CALYPTRA {i) The stalk of the archegoniophore elongates considerably.

FOOT
„.£ir;:r5.s'£ rrt:;.=
SETA

ENDOTHECtUM
AMPHITHEOUM
I, Ir^ It
sheath. Tt isIS one ce
cell^in thickness and aand
archegonium lew the
cellssporogonium.^
in height. i
CAWPTRA CALYPTRA

PERIGYNIUM

SPORE MOTPSEi
CELLS The four b^'ceh'^different'Si'on
'ELATER SPORES ium) by repeated ceU division capsule. The four hypobasal
MOTHER S>{ EL ATEPS and continued growth give rise P ^ ^ During furiher
CELLS

CALYPTfiA*
octants form die
development the
foot "nnl/^s^oro^hym ^ vcnicaOy at the
gameiophyte. Periclmal divi-
CAPSULE
(RUPTURED) WALL
PERIGYNIUM
expense
sions of nutrients
appear drawn
in the emb yom These mass
separate an
of cells
CALYPTRA
outer single layered ■ (Fig. 519 G). The amphiihe-
which constnutes the endotheciu (
cium cells divide ^ p thickness. At maturiiy the walls
sulewall. It P"^•u; fiiickenings on theirinner surface
spoi-ophyte. P"ient of embryo of these cells develop, The archespona
uro cross-seetionB. The entire endothecium form a massive tissue. It is
cells divide and redivide "^^totica y sporogenous
E". Eifferentiation
Differentiation of
of capsule. known as the sporogenous ^riically elongated. They divide

''^^^-apturedcalyptra.
(Diagrammatic)
 90 botany fob DEGEEE STUDENTS-PABT III: BBYOPHYTA MAKCHANTIALES—JIARCHANTIACEAE ; MARCHANTIA 91

cells elongate stilUurther.*^They sporogenous receptacle. These are the perigynium, the calyptra and the
perichaetium or the involucre. The first and the third are absent
These are the elater mother LllT Th 7°"^
ably to form long, slender dJnIrvi V elongate considcr- in Riccia. The development of the capsule is accompanied by the
theends. £ach of these'hpv I ^^^siform cells tapering at both growth of the foot and the seta. The absorptive foot is embedded
the irmer surface of itrwalls and^^/r thickened bands on in the tissue of the lower surface of the female disc and is differen
tiated early. With the differentiation and the maturation of spores
become the elaters. The bisniral P'^otoplasmic contents to
help in the dispersal ofspores^ ^ ^ hygroscopic. They in the capsule the constriction-like seta elongates slightly. The
slight elongation of ike seta in Marchantia is an anatomical necessity.
cells dmde''Tnd'^redfvide^by^
rows of cells. These arc fh
sporogenous
cleavage to form vertical
The sporogonium hangs downwards. Greater elongation of the seta
will carry the capsule nearer to the ground and thus hinder efficient
good deal of evidence to sunn mother cells. There is a dispersal of spores.
» cells do norbebnrto,he^ Production of Meiospores. It is entirely the concpn of
a difference of 3-4 or 5 . ^tne cell generation. There i«: the sporogonium (Fig. 5'21). It is devoted to the production of
enters are differentiated eaH eTff generations. The mploid spores (meiospores) and their dispersal. It hangs downwards from
pore mother the under surface of the female receptacle.
equally in all cells are nf t
direeUons fi \
* '"'"ewhat
^heepical.
sporogenous tissue Jrt
Laterthe; Ti
cytoplasm and ^
cell undergoes a ^.^^^Picuous
cnncn- hecome
diploid rounded.
nucleus ThevI ^°"tam
ronm* ?dense
FOOT

P£R\Q,yN\UM
SP/Of^M/5
"'^ss/MJiaropt CALYPTRA
I^ILAMENTS
(RUPTURED)

CAPSULE 19
WALL
muc
SPOROeONHJ!^^* tMBRYQ
If-VJWf.
^<5
r CALyprnA ELATER* OB
PlKiermiM

INVOLUC
SPORE
T£TRaO

SPORES

VALVES

A^ASi OP c £LAT£RE
SPORES
•>' A^O ELATERS

A.
' J'ig. 5.20 (A-f'i ST

B.
C.
^'atei-s and spores. rig 5-21. Mai'cliantia sp.
Longitudinal .eo'tion of tho mature aporogoninm.
92 BOTANY FOR DEGREE STUDENTS-PART III: BRYOPHYTA
MAROHANTIALBS-MARCHANTIACBAE : MABCHANTIA 93

flollowing
II Sporogonium. It is differentiated into the
three regions :— « ^<T"e.««,,le /Fiff 5-20B). With the ripening
*u 1 absorptive and anchoring organ forming „,.,s :r.H.s„"!:r.ss'£
Ruptures along an irregular
gametophyte''fortre'd\TeIopLfspor4o^^^ nourishment from the re"iidr°onhe"capsuk Since th„e are no regular J.n« of
connek^he foot and^\he'^capS ^"^Tlic^c £r~sixrri£'%'r';;.ss
consequently exposed.
1 x- There is ready and efficient dis-
persai<Ss°:rbTwlfdin—It Is ald^d by lire following
two structural features.

mature. \t has an ouier^Iave? colour when (i) Presence of Elalers. The hygrosayic '=!f' ''^'P
capsule wall. These cells Lvp ■ r ^ constituting the
walls. A cap of cells more (h;,n thickenings on their
in some species at the apex of thp°"^ cell-layer thick is developed
!!:: criroTdreTn^?ordlng a. as^
where they arc
sporogenom cells. It p^oiects
closed within the capsule wall is the capsule. En- readily carrght up by the air currents and dispersed.
elaters (Fig. have
ends. They 5-21C). Thespirally
double elaters are In
thickex^ed b? h meiospores and
pointed Ui) Elevation of the hanijing sporophyte on the long
The mature eiaters lack protoplLm Tb ^^eir walls, ^hnre The elevated position of the hanging sporogonium is a
hygroscopic. ^ otopiasm. They are thus dead and are
curienls. The smaU spores '"l" by
movements of the eiaters are elongation of the seta would not
wall. ^Thrrpot'e^wLt'is
culate comparatively thicker exTsoor
structures eachsmooth
\vith aorthick
reli- the air currents. The g j,. ^^^Id have
endospore or intine The np • ^ cxine and an inner ihin Sere'S'-rrying the capsule downwards more to the ground.
Wuhin the spore waUisIun;~ T^V ^Pecies irabsenT g^IPARISONjAITI^^
spies'
P are indisiinctlv »"<= reservrfTod"'7'°Pf
tetrahedral ™'species'=°"-
r " ^n a few the
Sporogonium of Marchantia
Sporogonium of Riccia

esssssiiap
i") Nutrition of S„o " f.?E;=J=S-9
In Marchantia the sporogonium
is larger and more complex. It is
difforontiated into the foot, seta
and the capsule.

ofChlorophyll.
tie se.aTa'pruU
Tw''^ '
orpa^ "rhl
t^iose of^hT''''""^
'iSr-s:;:5S
1 outline.
in

enabling the sporotr contain abundant an'^ tlevelop The cells of the single layered
such asVarch. Af this""' tnanufaeture some of 'i'^'^^^'^toplasts The globular sporogo^—^ capsule wall develop ring-hke
thickenings on their walls and
of a jacket lay which
upon the parent nil stage the sporoaonium il n . food colls ® of fertile cells surround a mass of spore mother
surrounds a mas ^ mother cells and elater mother cells.
supply of water and hs food sim f "^^Pendent ■'""""Trore iono'^ilator mother
the parent plam
sporophyte 'TheThT^' '?
reaches';!::, " emteW^-
layered q Much of the sporogonium pro-
Excepting ^^1®^,, inli a few of duces the sterile tissue in the
sporogonium wal^

it,4 »
 94 BOTAiqr FOR DEGREE STUDENTS -PART MARCHANTIALES—MARCHANTIACEAE : MARCHANTIA 95
HI : BRYOPHYTA
tube is formed. Instead the contents divide immediately in one
Sporogonium of Riccia plane to form a short irregular filament (Fig. 5-22C). It consists of
Sporogonium of Marchantia
£xw^
the nurse ceUs that undergo
■NT/Nf
degeneration, major portionTf
the sporogonium ig devoted to wS ^Ha^f^'of^the capsule ^- ^NUCLSUS
spore formation. tissue forms the M f
^lB.andZrtrby";fon^o::
4.
The nurse cells which eventuallv 4.
nuia arc considered bv soma
tS
the elaters of the foreronnern?
more advanced Ifssss
Thf
The elaters
ft are thus Marchantiales
absent in
the sporogonium of Riccia.
5.
the
spore L If
of thesnores houses
gametophyt, anTfc' the spores ®^®®Pt
"cwgametoptyte the
Th Fig. 5»22 (A—P). Marchantia sp.
from the A. Mature spore.
tfvcle and is auri recep. B—F. Various stages in the germination of a spore.
ar''eSg^'™"dp"icha'S!;S; layered
and tlie ®V P®'*%y- 6—8 cells. It may become two cells broad at the apex (Fig. 5'22D).
I-ater by the ranid f* '*""*• Soon an apical cell with two lateral cutting faces is established at the
seta, the ®f /l apex of the filament. It cuts off segments alternately right and left
through the nni breaks parallel to its cutting faces. Five to seven segments are cut off
boyond t>t
Jectsperichaetiuf
the ^ Pro-
"»id in this way. These divide and re-divide. A green plate of several
'■ Smply'bnh?H"' 7. cells is formed (Fig. 5'22E). It is the young tliallus (Fig. 5-22F).
the ceIlXftV^''''<^e';ation
■"""aaa'ofl
Now the two faced, apical cell itself divides and re-divides. A group
ealyptraani finaiwr'lt death more of cells is formed. A transverse row of apical initials constituting the
&Qd decay of tii^
apical meristem is soon established in this group.
thallus tissue Th® ®"'^'"o«nding
"ever dehScea. ® Important Features.
(1) The plant body is a repeatedly forking, flat, green, broad
' ?srjrofrrr\™'''^
decay of the burronnf y grogonrum°on dorsiventral thallus with a distinct broad, central midrib which
and are no» disnerQ f tissues tbe archegonionu stalt
^'^"gingof is more pronounced on the ventral side.
condition thev^T^\ f" that (2) Dorsally the thallus surface is marked by rhomboidal or
by the wind. carried
polygonal areas called the areolae. Each areole has a distinct dot
^ind. ®®^osporQ3 by ^he like air pore in its centre.
(3) Small, shallow gemma cups with frilled, membranous
margins are usually present on the upper surface of the thallus in
the midrib region. The cups contain numerous, green, lens-shaped
objects, the gemmae. When detached each gemma develops into
two new thallus plants.
 MARCHANTIALES-MARCHANTIACEAE : MARCHANTIA 97
96 BOTANY FOR DEGREE STUDENTS-PART III: BHYOPHYTA
ribs of a tiny umbrella. At maturity they spread widely apart giving
. arise
region, From the ventral
unicellular, surface of
unbranched the lhallus,and
smooth-walled in tuberculate
the midrib the female receptacle a star-shaped appearance.
(U) The archegonia are pendulous. They hang downwards
^hyt^the sXtratum from the under surface of the lobes of the female receptacle.
(12) Fertilisation takes place in the presence of water furnished
.hallufie"e'=lTL'uf.Ut by rain or dew and when the female receptacles are sessile.
four alteraatmg rows on either side of the midrib Ereh °f (13) The two lipped, curtain-like perichaetium may enclose
typtcally consists of a body, a constriction and an appendage several young sporogonia each enclosed by the calyptra and
surrounded by the perigynium.
the aedlii^v''or''''''"'r"g'''
theaenvtty ofa group of meristematic cells and not 'akes Lplace by
are situated in a notch at the apex of by aoica^ (14) The sporogonium, at maturity, is differentiated mto a
cell. The former elch ?obe. foot, a small seta and a capsule. The cells of the single layered
capsule wall develop annular bands. Inside the wall is a mass ol
meiospores and elaters.
Structire
any liverwort. It is manv ceUs in
is simple but indeveloped
its internal
by (15) The ripe capsule wall dehisces by 4-6 irregular teeth or
is the upper eoiderT^t^n I- I
pores ffe"ou?ef:Xo^hte;de™arcelh^ section. There valves.
(16) The elaters are simple, long cells pointed at both ends.
Each has two spiral bands on the wall.
The shallow cavity of each air'
a1^h:m^beTs"Sn™<l
''V oir pore
m (17) The elaters are hygroscopic and thus help in the dispersal
quently brancS^assiSlatorv fit®'"''" green,^ he! of spores.
The neighbouring chambers are^'Te^" which arise from its floor, (18) The ready dispersal of spores is aided by the pendulous
groups of vertically arranged green ceT tu t, sporogonium being elevated on a long archegoniophore.
portmg the upper epidermis. T^e ventflatL ap
portion of the thallus is called the nhofo.' ^'^^""•^cred upper (19) The short seta of the hanging sporogonium serves as an
ponton of the thallus below the ah The additional aid in the dispersal of spores.
layers of compactly arranged oarpnt^h consists of several (20) Each spore, on germination produces a tiny gametophyte
storage region. RUzoTds the which, at hrst, grows by an apical cell with two cutting faces. Soon
epidermis. ".nizotds and scales grow from the lower the apical cell is replaced by a group of meristematic cells.
SUMMARY OF THE LIFE HISTORY OF
type.archegonia.
and 'rhYX^sare^dioecior^^^^
They are develoncH " °f ^"theridU
oogamous MARCHANTIA (Fig. 5-23}
Life history of ilforc/iawiia is similar to that of There
aamet'^ 1, They are th" "^'tapcd receptacles occur two distinct vegetative individuals in th^e life cycle. They are
theffreen thalloid plant (Fig 5-23, 1) andtl^ radially
whole lengr- ^ave two furrows^'ell comUuSd more or lessofsolid
isTe most conspicuous thesporogonium (Fig. 5-23, 6). ItTheis former
two and is independent. called
Ip ^fmetophyte as it bears the gametes. The gametophyte
oHnt lT gr?en dorsivcntral, forking, highly developed thallus
ostiole The pit oprning- ^he recep- with a distinct, broad midrib. It is anchored to the ground by
• uilgv unbranched, smooth and tuberculate rhizoids. The
The
lack sperms Le
rays chJr P"' ^l^ernSe
curved structures with°
TK chambers. T'' . ab
rhizoids absorb "ater and
xheysolutes for it.in Associated
are arranged with the
2 to four longitudinal
rhizoids are midrib. Internally the upper surface of
"" ™-5 r^eshy tLut^as - '^ver of .air chamber. They are
jh.^ s'l';tt..ated''fVont
in number In youngg 'eceptacles
^ oi"'^hehang
they disc ^ ^he stout
cy tiang downwards like the
Sledge urpefcpWe^mis. fh'e calities of lir chambers are
 iT^ liHilWP

98 BOTANY FOR DEGREE STUDENTS-PART in : BRYOPHYTA

MARCHANTIALES—MARCHANTIACEAE : MARCHANTIA 99

barrel-shaped pore. Beneath the air chambers is a several layered

thick storage region of compactly arranged colourless parenchyma
'£^feMAL£ RECEPTACLE cells. Scattered in the storage region are u few oil and mucilage
cells.

"^aCHECOmOPHQRE Each thallus lobe has a growing point situated at its apex. The
OOSPHERE'
00$PHeR£^^2 growing point consists of a group of meristematic cells.
"^^^FEMALE The gamelophyte or the sexual thallus bears the sex organs.
<SAMETOPHYr£ OS TIOEE^^ They are borne in localised areas which are disc-shaped. These
are the receptacles. The receptacles in Marchantia are invariably
elevated on long stalks. The stalked receptacles are terminal in
.V_ f'ANTHERIOm position on the thallus lobes. The antheridial (Fig. 5-23,1) andarche-
^f^THERlOtOPHORE^
gonial receptacles (Fig. 5-23, 1) occur.on separate plants. Together
SPEPMAT020/D
with their stalks the receptacles are called the antheridiophores
, and the archegoniophores respectively. Marchantia is thus
dioecious. The female receptacle, at maturity, is star-shaped. The
male gametophvte archegonia occur in the tissue between the finger shaped rays. They
are pendulous with the necks directed downwards. The archegonia
<5AMET0PHVTIC of each group are protected by a two-lipped involucre. The male
'GENERATION Qn) receptacle has a lobed margin. Each lobe has a growing point at
its tip. The antheridia are sunk deeply on the upper surface. Each
lies in its own flask-shaped antheridial pit opening at the surface
11% GAMETOPHrrE by an ostiole (Fig. 5-23, 2). All these structures produced by the
thallus gametophyte constitute the gametophyte generation or the
sNUCLEUS haplophase. They are the green thallus, the antheridiophores, the
SPORE ^nucleus archegoniophores, the antheridia, the archegonia, the sperms and the
eggs. The first structures of this phase are the spores and the last are
the gametes. With the formation of the gatnetes the haplophase
ends. In Marchantia there is provision for additional multiplication
S ^SPORE of the thallus plant. It is done by the production of gemmae in
% tetra the gemma cups. The gemmae are thus the accessory means of
multiplying the haplophase. They play no role in the alternation of
sPORopHrnc generations.
generation(2N) With the act of fertilisation the life cycle of Marchantia
switches on to the second phase. It is the diplophase or the
SPORE MOTHER sporophyte phase. It originates with the zygote (Fig. 5*23, 5).
CELL
feRichaetium
The latter undergoes cleavage to form the embryo. The embryo
W THE
\'l^^ SHowmo .ZryJ'-i-ED
ClOHTCF/ir^ mo r%Icn AND
in the bryophytes is a short-lived structure. Soon by further cleav
celled stagYof age and differentiation it forms the second vegetative individual
SP0R06NIUM in the life cycle. It is the sporogonium (Fig. 5-23, 6). All the young
U.S.) ■
sporogonia in a particular group are protected by the common, two
ELATER CAPSUE lipped involucre. Besides, each sporogonium is closely invested
by the calyptra. External to it is the perigynium. The sporogonium
cells is more or less a solid, radially symmetrical object without any
^latI^ appendages. It is incapable of self-nutrition. It consists of three
regions, the foot, the seta and the capsule. The latter produces
the meiospores and elaters (Fig. 5 23, 7). The elaters help in the
dispersal of spores. The parasitic sporogonium of Marchantia, as
it produces the spores, is called the sporophyte. It is concerned
with reproduction. The structures produced after fertilisation such

'V. v.,
 101
.MARCMANTIALES-MARCHANTIACBAE: MARCHANTIA
100 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
I {g) At maturity it is simply a spore case with no diploid tissue.
as

'1^"= sporogonium with its foot, (h) Absence of any mechanism for dispersal of spores which
phase It ends sporophyte phase or the diplo- must thus await the decay of surrounding gametophyte tissues.
^pore mother celirbvW ^he The changes involved are :
structures of the future garet^gSom X lie W •

(a) Gradual and progressive widening of the air canals till

they form well developed air chambers.
iuiill "——

it is clear thaUhr^sameto^nW Marchantia given above (&) Associated with the above change are the development
closely connected. In the sWlelff
other. The reproductive
p°J"ophyte phases are very of assimilatory filaments from the each chamber and
the elaboration of air pores into well defined barrel-shaped struc
produce the alternate nknf Tk generation, on germination,
cycle are fertiUsation ard\«I^ • in the life tures.

(c) Development of sex organs in restricted areas of the thallus

diplophase or the sporoohNte i, former starts the
^ « ^.i^metopYvyte phase or the hanln^K tneiosis originates called the receptacles.
in a single life cycle of alternation {d) Elevation of receptacles on long stalks.
ogical phase, 'Alternation of Pp ^ • other is expressed by a Ic) Cessation of apical growth of the thallus lobes afterthe
t7i:^terologous type,
are dissln "'?.^ Since the^ltcrnaang
^ alternation of generations
development of sex organs so that the gametophores become term -
nal in
m position.
pusiiiuii.

(/) Gradual and progressive sterilisation of the potentially

fertile cells of the sporogenous tissue in the sporophy e.
The DISCUSSION C CC113 UA ctxw

This theory of progressive sterilisation was first proposed

• _ *1- » A V* TAIOC

Th°eSborMio 1' 'Eternal rather tlf

reached the
independent and
a dorsiventral thall" ■ S;o^-
gametophy^^th^^ I^^el of di& Internally it^
fotSspSrJ'moth:r'e:ih°beiine sterile to form narrow, pointed,
-* "-"t two IS prinut*>»'> -
hygroscopic elaters which help m spore dispersal.
^ •— - • - ^ . ..

arv nntv t^o ODnn.U u The supporters of this hypothesis consider

and Marchantia^ a h's'^y c™^"0^"^ se®rS"of intergrading
membe;o"he ^°'°Sistsregard
®constodertheKnf*'!
evolu'i""^ genera of the order form a co ] excellent example of
«"= followsg it'"
characteristicsis:J'"P''=
simX""dS™"?- They holdasthat
thus primitive it by
shown X^resslvr'evolullor'or natural advance. Smith is another
(n) Th iT auch
exterSj'""!"^ .
ardent supporter of this hypothesis.
(6) Pr
p, eternally. ^nd
and relatively
relatively simple no
simple botli
inv fi.L'
thati .™tosynth»,;.
the '""'"die .redon
. i cerv-
"'"'ydtlietic system ""d less water consei
(c) Ah-
AL,
pores. ^"'Cnce of ,vell
^vel1 c
^ ""= form of air chambe
chambers
as
s'S iirr'.rErs
The changes imo K".t..Jii""«'I™'-
become progressively reduced in width
defi
air . canals. This change is accompanied
(d) Absen "PPCr epidermis and definite'
iiii0 by simplification of air pores.
and , u , with

with ^'tnple
"tfutfoot S
^^Porophytg e •"pendent on the gametopbyte le
of little more than a caps"'"

... n^r

.IVU;,1V.V,
 102 BOTANY FOR DEGREE STDDENTS-PART III : BRYOPHYTA MARCH.'iNTIALES—MARCHANTIACEAE : MARCHANTIA 103
qu^emly .h..ex organs become scattered on the upper surface of the 10. What do you know about the dehiecence of capsule and dispersal of
15. —' upfJCl 5Ull<tCC lilt' spores in Marchantia.
. (c) Elimination
Elimination of
of nrotprfiu,. cU- ..
.. . 11. List the important features in the life cycle of Marchantia.
mum)around the sporoKonium tk" {involucre and perigy- ,
12 With the help of a series of diagrams describe the life W^ory of
(AqraVmverstty 195i, 5G)
Marchantia. ^
in the complexity of the strumirA
tionary trend has been the /radnL r l ^^^^"^panied by
^hereduction
evolu- 13. Justify the truth or falsity of the following statements giving
of the sporophyte. First the 5", P'"^S'"essive simplification reasons :—

,sporogonium becompc ^he foot also disap- (а) The air pores on the thallus of Marchantia are homologous to the
stomata of higher plants.
wan. All the inner Tendotta^
disappear from th ^ spore case. The rin
Tfip rin?- (б) The short seta of Marchantia sporogonium is an anatomical neces
ters are eliminated become fertile i sity.
^TlTSubn^f'^ ^ecome fertile s^tha't the lc\ Thecommao of M-rchantia are reproductive structures which do
^pore mmother
sLre t cells. capsule\va!l Tad
^W^ied by the dep-en ?• "umerous developing of the poteitial not play any role in the phenoraenou of alternation of genorationa.
u. Describe the main structural features of the
°"ormsAlllike
the Ricoia
changesfrom
eniim j ^^^r of the calyptra.
in the origin Marhantia. .

The hypothesis of nrotrr ■ ^arckantia. 16. Oomparo the gametophyte of Jifuroftauiia with 'bat Draw
suitable sketches.

onrried out bv Rt' STbirt

P'ausible by Bower, by Bower,
■ogress
of progressive ('9«)
''
.^^'V^^'Olpllficatinn w h
with > .
Jforei, F
f'^r<^hanha sunnofr experiments
experiments
he obir:e7rn''''®^='bonorreductbr^^^^^^
ways and
™PP°it .lit
the rUenrV
theory
F " "I" mutants vvh' Ohts cultures of jiforc/tanfm
-entbie"
sex organs &t7."n 7=^. "®"ow thallus
in the tha7'"? ThaTiuIs''^ - -"°f
me thalius tissue. 'obes and immersion of
The 'V
J-ne strikinfr "ssue.
_ ^uues and immersion o'

"''"ed an aXXF F '^e basFo F-<=''Periments

"he suggest
stia bythea
- genus. hut a reduced rorltd^F^rri

ana ai^r■
"fJ^arolnh'^'-be theexteLarw "
^Gribe ag^ in detail. met with in Ma^chaniia-
'■ —- —

the struct^ of sporogonium of

^ mature Brx«
sporogonium of Marchantia md

f'li
 105
HEPATICOPSIDA (HBPATICAE)-JtJNaBRMANNIALES
stemanditsbranches. Consequently they ^ j-ior edge of the
leaf arrangement is said to be (Fig. 6-2A). It
leaf is above the posterior
issuccubous if reverse. In
f
the
anteriorfront.
marginTheof
■.1 f
the leaf is covered by the posterior ° a lateral rows,
CHAPTER VI simpler foUose forms have the leaves an ange ,^
h.pat.copmda|„epai.o*ekj™c,,„^™,^^ The leaves in the advanced foliose forms are t
three rows. Two of these are dorsal and o ^^ritral in position,
leaf is usually bilobed. The lobes are dorsal smaller
' dorsiv^tril in cTnfigum^^^^• ifteonly exceptions
and prostrate in habit
are Haplo- The dorsal lobe of the leaf is larger than the
ventrallobe is called the lobule and the i^rg advanced
CAPSUL£^-f\ hnJr^r The leaf is invariably widiout a midrib. Most 01
both these the Oalohryum.
plant body Inis foliose forms are epiphytic. ..^^nniales
erect. For this and other
reasons Campbell places them The external elaboration of the thallus in tissues,
m the order Calobryales. is accompanied by little or no mternal diiferenti ^
. Thallus. In a few ¥he Xleandthallus
chambers consists
air pores of similar parenchyma cells.
are absent.
is a The thallus is usually secured to the ^^16, smooth,
S ?' '^^'"^^^^"trahflat.
dichotomously green.
branchedShab rhizoids. They are Notwithstanding the
thin-walled type. 1ofhethescales
external elaboration a
advanced ipafv forms the Jungerman-
^ striking series of
that nf"^o* ^osembles
in /^vt or Marchaniia niales form the most natural gro p. . | ^ thallose forms with
simnl or even transitional forms which connect the simplest tna
the most specialised leafy ones. ^ ^
Distribution and H^itat. "far the richest
In^SZahTghifTn":'™""'-
tion
ff. ,, the thallus IS
.organisa-
genera with 9,000 species. The leafy
Ltionspeci es. Thejungerman^ ^ ^ t^eir distribu-
They
A femaleVant leafv
Plant hearing a spoTOgonium. (fohose). The leafy forms are occurring both in the cold dun , ^orth and south
about 84°/. of all r
Jungerman-
are some of

the places rich in these hcpatics. pre-condi-

Combination of shade and f the
ticbut the
tion for their successful growth. ^ , P ^hey
ey grow
givjvv on damp
srs»-X« ■
their leaves in the

rfnioiluce by the following methods .

Gemmae. Many jungermanniate
are

{Amum} the gemmae gdal cells of the thallus. Th^ The

showing facnbous^n?rrn f"® above producec endogenously m tl
produced P^ and star-shaped. They
B- Aleafsoon from above ™"' "f leaves. gemmae of Blcisia ftbe aoexon the thallus and rarely in
are usually
are usually produced
produce- Sat.^TLTate readily detached and
104 flask-shaped gemma receptacl .
108 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA HEPATICOPSIDA (HEPATICAE)_JUNOEEJiIANNIALES 109
The latter may serve as nutritive cells but more often get meta 1, Anacrogynae
Anacrogynae or
or Mel
morphosed into elaters. The spore mother cells usually become rally of a thaUoid construction
our o ed before further division. The ripe capsule, as a rule,
dehisces m a regular manner. It splits by four valves (Fig. 7-11).
become refiexed. The elaters radiate out and by their
yg copic movements aid in the dispersal of spores.
Gernunation of Spores. The liberated spores germinate
under suitable conditions. Each may either directly develop "nto

ELATER
CELLS
Yomc SPORB
MOTHER CELl.^
ELATER CELL

SPORE MOTHER CALYPTRA

CELLS

CAPSULE
mLL Fig. 6-7. Porella sp.
TerminoJ archegonial cluster.
u 1 QntYip snecies are foliose. Transitional
INVOLUCRE the sub-order as a whok.^^ Some spe are also present.
forms between the toliose development. The archegonia
■FOOT
They show rudimcntaiy ^ ^^^1 cluster is never terminal.
usually occur m surface some distance back from the
It is_ borne on the upp^^ the forma-
L S. young sporogonium.
growing apex. The p g^^b.Qrder comprises about 23 genera
tionofanarchegonium. modern trend is to allot them to
aspecithallus or fo^t and
six ormore
seventhan 5^0 species
families sue ^ , . Riccardiaceae. Pelliaceae,
Pallaviciniaceae, andTreubi-
Metz-
es is short-lived an? ??heme"r ^ome aceae, PdUaceae is considered in this text^
termtc of the foHose forms Sometimes geriaceae. Of the * „„a„„i„eae It includes the ad-
vanned2. foUose
Acrogynae
forms J 84% ofrichest ■ in species The
all liverworts. and
according to Mulle „ arranged in three lateral rows. Two
leaves on t^e axis are usua y S leaves are bilobed
'omffrm™ '^e erect forms like Haj>. of these are dorsal
an^arger. and one
The thallus -^hethi-ee-sided apical
apical cell itselcellis
having the form of ^hegonium in the terminal cluster
used up in the formation of an arrested. The sporophytes
Campbell separates the H ^""Sermanniales.t (Fig. 6-7). The growth of the ax gub-order includes about
manmales and gives it ihl ^fPlomitrineae from are therefore terminal „ ^re placed under 17 families.
ordmate with the at \ of an order n i \ J"^ger- 220 genera with ®'™-i®^f[nciuded in this sub-order Madothecaceae
manni ales. Marchannales. Sphaerocarpaks an^J^ng™" ,,Por—istakenuphe^^^
Of the numerous tammc FEATURES '
1, The haplophase "Trad?an«ffmmrBh^^^^^
manniales is a simp e, a
 no BOTANY FOR DEGREE STDDENTS-PART III : BRYGPHYTA HEPATICOPSIDA (HEPATICAE)—JUNGERMANNIALES 111

19. The elaters after dehiscence radiate out and by their

so^alkTsTim and the Ua'ves*^ " differentiated into the hygroscopic movements help in scattering the spores.
20. Formation of a protonema stage is characteristic of the
logicafdifeetiaffoT'
chyma cells.
the'whoTe nlZ
plant consists of similar paren-
foliose forms. It is generally short-lived and ephemeral but some
times long lived.
3. Even the leafy fo^s are distinctly dorsiventral. COMPARISON BETWEEN THE JUNGERMANNIALES
three lateral rot™ Thetetves'°of the"l T """"Sed closely
on the axis. They slightly overlan
in twosetor AND THE MARGHANTIALES
The general life history is the same in both the Jungermanni
incubous manner. The leaf has no midrib ° ^ succubous or ales and the Marchaniiales. There are differences in detail.
Jungermanniales diflfer from the Marchantiales in the following
The

thin-walled^type.^^^"^^ unicellular but always of simple, smooth. respects :

1. Greater external elaboration of the gametophyte in the
6- The scales are usually absent, foliose Jungermanniales.
by means of a smgbfp^iraUedl.^^^ Jungermanniales takes place 2. No or little histological differentiation so that there are
neither any air chambers nor any air pores. The entire thallus
8. The rex organs are never borne on stalked receptacles usually consists of similar parenchyma cells.
3. Absence of tuberculate rhizoids.

Sad'- h™' Smtsrit broa" 4. Absence of scales. Evans, however, likens the scales of

or ne£cells" "«k consistettLTytvtvmlS® ots Marchantiales to the ventral leaves of the foliose Jungermanniales.
This view is supported by the fact that both are one cell thick and
are attached by a broad base. Besides, scales of the Marchanti
usually ales, in many members of the order, contain chloroplasts, at least, in
early stages of development.
5. Growtli always by means of a single apical cell.
il- The sperms are biflafr^^iio* i position,
numerous coils than tK r §cllate but larffer anrl i 6. Sex organs never borne on stalked receptacles.
12. Th Marchantiales 7. Archegonia usually in groups and distinctly stalked.
fflately transvenftau"™'? by a transverse or 8. Venter nearly as broad as the neck or only slightly broader
lower hypobasal cell. "^<= 1° an upper epibasal cdfrnTa than the neck.
9. Neck of five vertical rows of neck cells.
thttlSaVc££t^'^:'Thttntire
The seta is n.uch elongated^aTtXlrstortgoidum'd "''"''"P"
10. Antheridia frequently superficial. They are lateral and
axillary in position in the advanced foliose forms.
11. Larger sperms with more numerous coils.
12. Development, in most genera, of the entire sporogonium
from the epibasal cell.
13. Greater internal elaboration of the sporogonium.
14. The capsule wall two or more cell thick with the cells of
division. mother cells become f the jacket layer always sclerified.
17 Th before further 15 Presence of perfect elaters in most if not all the Junger
manniales.

16. Dehiscence of the ripe capsule in a definite and regular

nue manner alonp r ^ ^ dehise^ ■ manner along four lines of dehiscene.
Which become r'aXte'r-^^ -g-Iar and a defi- 17. Presence of a protonematal stage in all the foliose forms.
P four valves
 i;AV

112 BOTANY FOR DEGREE STUDENTS—PART ILI : BRYOPHYTA

The Jungermanniales resemble the Marchantiales in the

following features :—
1. The simple thalloid plant body of the lower Jungerman-
niales is a constant feature of Marchantiales.
2. Prostrate habit and dorsiventral configuration.
3. Presence of unicellular, simple, smooth-walled rhizoids. CHAPTER VII
4. Sex organs similar in essential respects in both, differing
only m details. JUNGERMANNIALES—METZGERINEAE : PELLIACEAE
5.^ Presence of perianth in both. It is ^videspread in ihcjun- General Characteristics. This family includes two or three
germanniales and is represented in some of the Marchantiales. genera with a thallose gametophyte. The thallus is prostrate and
chromosome complex of n=9.chromosomes dorsiventral often lobed by irregular incisions. It is secured to
(Mehra). the ground by means of simple, unseptate, smooth and thm-walled
7. Similar radial and general plan of construction of the rhizoids. The scales are absent. The sex organs are either scattered
sporogonium into foot, seta and capsule in both. The elaboration or in groups on the upper surface of the thallus. The archegonial
in the Jungermanniales is internal rather than external. cluster is always surrounded by an involucre which is an outgrowth
8. Similar type of elaters. of the thallus. The capsule is usually spherical or oval in outline and
elevated on a long, slender stalk, the seta. The capsule wall is
both intercalary growth in the sporophytes of generally two to four layers in thickness. There is a coherent mass
of elaters attached to the floor of the cavity of the capsule. It
thqt photosynthetic tissue in both so constitutes the basal elaterophore. The elaters are free. Each
phyte physiologically dependent on the gameto- elater has two to four spirals bands. The spore mother cells become
four lobed before entering upon meiosis. The lobes are arranged
revision questions tetrahedrally. The important genera included in this family are
Pellia and Noteroclada. Evans (1939) added a third to the list. It
tares of the distribution and external fea- is Galycularia. Of these Pellia is taken as a type.
resembleth^Ma^chaJtSs^? tho JungermannialeB (a) differ from and (6) Systematic Position :
manniales. ^ account of the vegetative reproduction in the Junger- Bryophyta
4. List the salient features in the life cycle of Jungermanniales. Hepaticopsida
those of tiie MaTchanSes'l aTohegonia of Jungermanniales differ from Jungermanniales
Anacrogynae or Metzgerineae
Pelliaceae

Pellia Raddi

Habitat. It occurs in diverse situations comnionly on damp

soil Generally it occurs by the sides of streams, springs, wells, and
in damp woods, sometimes, actually under water. Rarely it is found
rncks The aquatic forms usually remain sterile. The
form ind texture of the thallus varies according to the habitat.
XrindWiduTls growing on damp soil have a robust thallus with
broad, elongated lobes. When growing submerged or m very
b .^,V fHmn shady places the thallus is delicate, long, narrow,
S'sltp'd vtira'distinct midrib and thin margin. Three
113
 IT

114 BOTANY FOR DEGREF STUDENTS—PART III : BRYOPHYTA JUNGERMANNIALES—METZGERINEAE : PELLIACEAE 116

species of this genus are found in India. They are P. epiphylla, Structure o£ the Thallus (Fig. 7"2). Internally the thallus is
P. neesiana and P. calycina (Fig. 7-1). simple but several cell layers deep. However there is no differenti
ation between different cells. The
cahjcina (P.fabbroniana) has been re- entire thallus is thus composed of
Western Himalayas, Mussoorie. thin, polyhedral, parenchyma cells.
The cells are joined together in a
moS^n
moist soilrnor actually under
etc.water.
It occurs between(Fig.
P. epiphylla 50007-3)
to 8000 ft. on
is common honey-comb-like manner. The
thallus is several layers of cells
thick along the median line or
midrib region. In some species ■RHIZOID
CAPSULE* (fl. epiphylla and P. neesiana), the
iUMOLUCHE. cells in the midrib region are Fig. 7-2. Pallia sp.
elongated in the direction of the V.S. th^ua.
long axis of the thallus lobes. The
INVOLUCRE'
cell walls of these elongated cells are thickened by brown or yellow
layers of thickening bands forming a kind of network. The thallus
gradually thins out towards the margins where it maybe one cell
layer thick. The cells near the surface contain abundant chloro-
plasts. The starch grains, however, occur in all the cells. A few
cells may, sometimes, contain oil. The single layer of regularly
arranged cells which covers the upper and the lower surface of the
thallus are sometimes referred to as the epidermis. The air pores
and air chambers characteristic of Marchantia are absent. The
unicellular rhizoids grow out as tubular outgrowths from the cells of
Fig. 7*1 (A C). Fellia calycina (dioecious). the under surface in the midrib region.
A. A plant bearing a nearly mature Bporogonium.
B. Male plant.
Apical Growth. It takes place by means of a single, large,
apical cell. It lies in a depression at the anterior end of each
C. Female plant, thallus lobe- In P. epiphylla the apical cell is lenticular and
in Sikkim and tr' ^ cylindric. It cuts off segments parallel to its two sides and the
commonly found. All (Fig. 7-7) is not posterior convex base. The former by repeated divisions give rise
found in the North Warl co—^y to the wings of the thallus. The latter build up the broad, thicken
ed median portion. Owing to more rapid growth of the margmal
cells the growing apex becomes sunk in a depression. There it is
dorsiventral tha^lus ^ody is a sm ii • i protected from damage. Near the growing point certain cells of
the lower surface grow into grandular hairs. They secrete mucil
age which protects the growing point from drying At the ume
of dichotomy the apical cell divides by a longitudinal wall into
tC'crr ™s.r C"ii two equal ceils. Each of the latter functions as an apical cell of
greS ° smooth dichotomous. a branch.
'sine"-' -nd not -''eolae. It appears REPRODUCTION
distinguish
Sfoh the
lobe isThi£7^^\ ^he Rrow
p^aTel ^P^que as that
difficult to 'I he thallus or the gametophyte of Pellia reproduces by two
The brOad poovlv rl "i^rgins it m ^. portion of methods, vegetative and sexual. The second individual in the life
cycle, which is called the sporogonium is concerned with the pro
numerous smootrwnli
substratum. Scal^ .??\.^ the
ThJvJ, ofof each arise
midrib lob^ duction of meiospores.
«fthethaUush^l^"^^'3ber^^^ plant to the 1. Vegetative Reproduction. Vegetatively Pellia repro
liesthegrowingpoj .^^'^^nal notch. At fb T
bottom
Each lobe
of this notch duces by two methods, adventitious branches aad fragmentation.
Il6 BOTANY FOR DEGREE STUDENTS—PART III : BRYQPHYTA JUNGERMANNIALES—METZGERINBAE : PELLIACEAE 117

They arise from the upper surface thaUus, functions as an antheridial mother cell. It divides
to- ctrluT
separation each branch grows as an independent plant
transversely into a lower primary stalk cell and an upper primary
PRIMARY
ANTHERIDIAL WALL INITIALS

thaUutdif
or Td ISttgraTe^^adlnt
the fragments from the parent thaUusto £3^?
ANTHSJ^IDm
^-^INirML UPPER CELL PRIMARY
PRIMARY
AN0R060MAL

growth and branching develops into an indepe^Mtdutt"''"' OWiR

STALK

LOWER
CELLS

of Pellia are dioeci-

ous, others monoecious. JACKS T
Examples of dioecious species ANDR060NIAL
are P. calycina ^Fig. 7-1 B CELLS
ANTHERlDlAl
t!
Ihe third P. epiphylla is CHAMBER
STALH
rlNVOLUCPE "^o'^oecious (Fig. 7*3). The JACKET

-'^enclosing- 'Tionoecious species are SPERMAT07QIDS

ARCHegon/a P^'otandrous.
STALK
ANTUERfDiA (a) ANTHERIDIA

ture.(J)Mature
Position and Struc
antheridia are Fig. 7*4(A—L). Pellia sp.
-M'y circular spots on thl Development of antheridium.
F>g.7-3. Pellia epiphylla (Monoecious). Vsf^^T^ev^ A—J. Various stages in the development
of an antheridium.
. .
(Explanation m the text)
.x
K. Mature antheridium.
L. A sperm. (Diagrammatic]
anheridial cell (Fig. 7-4 C). The former undergoes a few divisions
to form the short, multicellular stalk.
The primary antheridial cell develops into the body of the
antheridium. It first divides by a longitudinal wall into two
surrounds a centra? ^^11. It j. .fobular body of daughter cells (Fig. 7-4 D). They are of equal size. The next wall
androcyte mofe r iT'?- a^^drocyte Z ^^ick and in ?ach daughter cell is in a plane diagonal to the first division.
spermatids. The nrnt diagonally imn! ^^ach It is nearly periclinal (Fig. 7*4 E). It divides each daughter cell
-biHagellate^pe^P^^P^ftof
^biflagellatesperS^^.^P^fiof' thf andU?^^
th? sperm
andL" androcyte or into a pair of unequal cells (Fig. 7-4Jh^ The smaller cell of the
coiled biflagellate strurtll^ ^berated V."^^^^^oi^phosis
^"drocyte or
into pair constitutes the jacket or wall imtial. The larger sister ^11
serted at
serted at different Sk Fia 7.4 ^ f "an elonaatirl spirally again divides periclinally into an outer and an inner cell (Fig. 7-4 G).
body of the sp^e^t ettitl?"'"-«hb S The outer cell is the second jacket or wall initial. The inner
cell functions as the primary androgonial cell. These divisions
. , N Development
single superficial JeU u
^
'^•^)- Each .
in orSn Capering can best be seen in transverse sections. At this stage the body of
the young antheridium consists of four wall initials They enclose
apex ""'he upper an aatherid-''t;"'" as a the two primary androgonial cells. The jacket or ^11 initials divide
only by anticlinal walls to form the one cell hick wall of the
into two (F;° 7 /pi'^allus, it "n^ses in growing antLridium (Fig. 74 I). The primary androgonial cehs divide and
redivide several times. The cells of the last cell generation consti
"*"<"■> 'S e,"„v. .fxrs tute the spermatocytes or the androcyte mother cells
(Fig. 7 4 J). Each androcyte mother cell divides diagonally into
 JUNOEEMANNIALES-METZGEEIKEAE : PELLIAOEAE 119

118 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

longitudinal rows of cells Jhg
two androcytes or spermatids. Each androcyte produces a of usually ®-® J^i"eated at the top of the neck. Sometimes
single antherozoid (Fig. 7-4 K). It is formed mainly from its
nuclear material. It is furnished with two long flagella. They Slower portion of the neck also becomes two cell layers thrck
end of the tapering coiled body (.-ii) Development (Fig. ?f>• lots.^Th^
t ^rrnm development of the antheridium group °f cells on fh= "pper s^i^c ^ ^^^^^ of the apical cell. The
ItTtrnrnwT
completely encloses it except for ^rowth
a small opening at the top tissuc.
of the surrounding the surface cell of t y g development. A
apical cell itselt is qu y function as archegomal
maturl^Lnrp^^""*^®' moisture the wall of the taS "^lle'Ut is that the archegonia stand in a group on the
^uHif a The androcytes are ex- upper surface of the thallus lobes near the growing point. ^
that cauSd^h?^r, ^^ter Any surface cell in the function as an
Finallv i-h»» s re. They rise to its surface and spread apart. initial (Fig. 7-6 A). It enlarges into a papnia iixe s
dLTotl^SwrusSX- PRiMAftr
PRIMART NECK
(6) ARCHEGONIA PERIPHERAL PRIMARY COVER
ARCHEGONM OUTER AXIAL CELL pCELl CANAL CELL
CELL

^.S,^,sr.,£asr ■s.-'tcu virc INITIAL-, :£LL

CENTRAL NECK
INITIAL

VENTER
RIMARY
VENTRAL
close to the growing apex INITIAL

{Fig. 7-5 B). All the four to

BASAL
twelve archcgonia in the cluster CELL
stand on a slightly raised trans COVER CELLS

verse ridge of tissue called the NECK CANAL

CEILS
receptacle. The latter faces
VENTRAL CANAL Ctt-L
the growing apex. The arche- OOSPHERE-
cluster are sur
rounded by a complete or STALK

incomplete tubular sheath, the

involucre (Fig. 7'3). The Fig. 7-6 (A-I). Pelba sp.
involucre is protective in func Development of the archegonium.
tion. It IS tubular in P. calycina /A—Hh Earlier stages. (Explanation in the text).
short and cylindrical I. Nearly mature archegonium. (Diagrammatic). ^
ANTHEmOlA
r. neesiana and reduced to in
1 -e.] initial outer
divides horizontally mother
into a basal
pouch open in ftont „in
a
The papillate archegonium cell
P. eptphylla. Amidst the arche- pedicel cell and "PP ^es the usual divisions to produce
gonia in the cluster occur the
mucilage hairs.
Sg?nlum'^TrbaLi%fdicel cell divides to form the short,
multicellulai stalk. , divides by three intersecting
. , Structure. The The or pheripheral cells
vertical walls so as to cu smallest. They surround an
(Fig. 7 6 D), Of ceV The latter
;.L?-.-' I--=2., internal, middle cell
slightly overtops the pe P
^ divides trans-
primary cap or cover cell and a
i-® sr.!",®-
JJ; ssss
~-"j
versely to form
lowei% larger a cei ^ ^ 6 E) The
central Thetwo larger, peripheral
smallest lateral or
peripheral cells divide Y ^ p ^ow becomes enclosed m a
The venter consists of t i s\out Ld ^ short ^ell does not dmde. The centra ^ .^^^lope cells or the jacket
istainsnotthesharply
egg and a sm if
rnark?d off T"'
O" from
of cells TU
The ef' stalk,
the ventpr t ^^°"gated neck
iLhia/s '^AlHhe cells constituting the young archegonium mclud-
It consists of five
 JUNGBBMANNIALBS-MTZGERINEAE : PBELIACEAE
120 BOTANY FOR DEGREE STUDENTS—PART III ; BRYOPHYTA

ing the central cell divide t.ta:tssve"C'5.eV5. TVte "vonng aTchegOTiwtti

■a now divided into two halves (Fig. 7-6 F). Each half consists of
five envelope cells and one central cell. The five envelope cells of
more
more ceil laycis
polygonal
. 1 cells (Fig-
r-zollc ^Fig. )•
7-8). V.-SV
^
the upper half constitute the neck initials Its central is the Their radial walls possess CAPSOLE.-
rod-like thickening bands.
co3e ^ Of Swer half The cells of the second
layer are flattened. They
are, somewhat, reticulately
thickened. Within the
the archegonial neck''^ repeatedly by transverse walls to form capsule wall is a central
In the mean„h"L the nrl^ ' tuft of 50-100
elater-hke erect
cells. It IS styled
a row of six to nine or mn canal cell divides to form
canal. The venter initS? ^hey fill the neck as an elaterophore (tig- INVOLUCRE'
encloses a venter cavity fFJ Tr venter -wall. 7.8) The lower ends of
wall becomes two or thrpl ^ n i fertilisation the venter the cells of the elaterophore
cell divides unequallv to fnrrrf thick. The primary ventral are attached to the floor
-nd a lower, large egVor caLl cell of the capsule cavity. Ihe
^o'^^^cell divides by two int^c f'" P'inally the primary upper free ends ladiate
or lid cells. Coincidem to form the four cover iiftothe cavity tnre™'"|;
tt? It arches
membrane ine with the spores and the
over grows frnm tl ^"^^olopment
'°P of the archegonia
-^hegonial recep- frfe elaters. The spores Fi<f 7-7. Fellia neesiana (Dioecious).
The female plant bearing a
are large. They begin to sporogonium.
W FertiUsation germinate before they are
presence of water. The axial row of "^ii manner in the u A The elaters have two omith holds that the
now filk the neck
°°w canal. The amucT
nrchegonium m I'
except the egg disorganises or three
basal bands ofhelps
elaterophore spiralm diickening
bnngmg eac^^ ^ a primitive
shedding
sort of
of
th^ sam pressure exerted from wi,v ^ater and swells the spores- Others believe that
ni?r svsiem. The capsule wall bas on i
surface four vertical
dial " n fiids accesTto th cap cell. At Sips of thin-walled cells, j^bese are ca sectors along
wate,Tu' /T"'== its . apex. TUo ^ttheridium. The antheri- nlnce The mature capsule sp ts as a r extends right
the androcvte ^u °Pening. Finallv ,u" I''es emerge into the "e " es of dehiscence. The split starts at the top^^ .yhis
archegonial ne V
°f them proh^ihi'^
I" ''™i™ to the ar u
down the
nte liberated from
^?Sonia and enter the open
"to the base. TJS a sharp contrast to
tratesThrei ^^""= fit" to reach ' .tt^tth the egg. One regular method of ® opens irregularly,
sation. Tlfe ferrT^ female Aagella and pene- the capsule of Marchanlia
The capsule of contains mom st^ compared
•tVi that
with that of
01 dlat'c'ia"'*"-
i I It capsule „aU
thick wal . basal
Pasa elaterophore and
r ,chorale
r^'lmerrTn short the sporophyte of w ith that
It Producra'the' me-""" '"'^"yele t^oitcern of the non- Obieot. It has a Of
more complex "™ctur f.ttile tissue (sporogenous
than
in their disp"::riP°«^ coSmonlv ?' f sporogoniu^- TMarohantia. course
(e) S„„,„ ''I' ^itd- ^ the spores and also The sTcnlisation oi -J nariiaUy decentralised by
cell division &nd cell fiifr (Fie 7-ft\)• tIt ic c-, S eU as advanced.
restitution The We da ^^P^^ ^Some people
of a sterfc look
^his view
foot '®fi^tiom fittttted from the zygote by upon the elaterophore
is disputed by otheis. as the ^ 31archantia
coSkrt°?*g reglo •''^f^^P-uIe
tollardik".^®- Its edL'^'" ^P^reg^nium'' T^'= f™'
insists of the . . Protection. Dixe which is

white coloiiv l^'"^ ^'^ound tV. yrjw^^' i} distinct and

green or black capsuie^^^'^ '^^^^Parent .
of thee ^he 7-8)
t^ig- seta isforming
of purf Ced
the latter for the entire spoiog
thTreioprg
and terminates in a dark
 122 BOTANY FOR DEGREE STDDBNTS-PART in : BRYOPHYTA JIJNGERMANNIALBS-METZGBRINEAE : PBI.LIAOEAB 123

sPotlr^l The latter divides first by a vertical wall and ^ 'i"°trthe

wall (Fig. 7-9C). The transverse dtvision >^ At nght angles
Erst. The embryo at this consL of two
these divides by a vertical wall. The emoryo no
SK.fOGEimS
CELLS

EPiBASALCElL
iNVOLUCHE CAPSULE
ObSPORE ol's WALL
■It CALYPTRA *SETA<
CAPSULE SUSPENSOR G
YYALL C ■FOOT
F
SPORES hypobasal cell SPORE MOTHER
ELATER E LATER CEUS
ELATEROPHO.
ELATEROPHORi CAPSULE
(JNFERTh ma
USED
ARCHBGQ^
AHUM 5USPENS0
R in
SETA

Fig. 7-9 (A-H). PeUia sp.

Various stages in the development of the
Exnlanation in the text. ^ j-

FOOT*
tiandredivide.
ers of four cellThe
s eachderivatives
(Fig. fj. up^r^t^er^Sm the capsule
stalk or the seta. The
Tissue or. (Fig. 7-9 F). The 1°"" ^^J,^riafe?in^^ a distinct, conical absorbirig
gametophyte seta at its lower end differe attaches the seta to and is
RHiZOiD
organ, the foot (Fig. 7 )■ ^ conical foot has its edges
OSnOLE. embedded in the tissue of the at the base of the
antheridial
CHAMBER
fifriKctsz/.'Su >»'. ri....
sists of uniform, ^J „ tier of the octant stage
Each of the four cells of 'he amphithecinm.
divides P':"'=hnally sepaiaMg constituting the endo-
The latter surrounds an m ; functions as the arehe-
thecium (Fig. 7-9E). ihe enaoii ^ss„e. As the capsule
A »=cti„n tifthaui'Wdng sporium or the primary
advances towards maturity 'he amp ^
divides by anticlinal
^all
walls to form the <=Aps»l« ^ farther periclinal divisions
mentation. The SXv (Fig becomes two "F'®, " .fF" cells divide and redivide to give
(Fig. 7-9H). The archespo.iura (pig. 7 9 F and G). Q,mte
rise to a mass of sp^ ® cells at the base of the capsule
early a differentiated
becomes central mass of sp g lickening
xheseon sterile cells elongate
their walls. These
remains as a one no furthnr cell
eon°iderably and develop spira^ thtcW
elater-like cells 'ad'^te &om base^ 7,9^). In Ae
'' derived from
nom ih
the epibasalentire
cell. SIX" iXE, .p."..". •*
 Il4 botany for degree STUDENTS—part III : BRYOPHYTA JUNGERMANNIALEB-METZGERINEAE :PELLIACEAE 125
The calyptra ruptures and remains around the base of the seta as a
aLdTh^Tterik'^Xtef
The elonffatecl celk wnf ^^ter elongate considerably,
torn membrane.

structures with a double sSralTh®' t (id) Dehiscence of the capsule (Fig. 7*1 lA and ®)- ^
dehiscence of the capsule is attained by the elongation ot the
;rzt'n?.s»
"* "PP.r p..t orE FREE
.....:; ELATERS ,
(m) Sporogenesis (Fig 7-ini it lo i i ,
spores are differenfiatPrl a.^ u ^ is the process whereby the
entiated from the spore mother cells by meiosis, r
ilaterophorb
FIXED ELATERS
germinated
CAPSUUIMU
'SPORES

LOBED
SPORE MOTHER
CELL l/ALl/ES

SPORES Fig. 7-11 (A-B). PeUia sp.

Two stages in the dehiscence and dispersal of spores.
Lobed spore-utother csS';;™, A Dehisced capsule showing recurved valves,
elaterophore, free elaters and spores.
B. elaterophore
The same with the spores shed and only
loft behind.
nucleus. Th"; nucLrSertice'^ ^n'ni^rthrSTpToid
seta. The calyptra is ruptured and the capsule^ The'dicing"ffep
Sn^lS sep°a^te°'bfceU tfilfSid'"°
hygroscopic movements m the from the apex along
This opens the capsule by cross ggg-
the lines of dehiscence ^ themselves. This exposes the
-sfoi't rrf'^4'rr::.
, The separation of the spores from
They eS:^S'StlJ'"'=
the capsule, ® ''™" "><= down to the middfe or
(D) Dispersal of Sp®J®f' , hygroscopic movements of the
the exposed mass is ajsisted byi e moisture
ur tne base of elaters". They coil and stretches out the elaters.
content of the air. The loss fj-oni the exposed spore mass.
withiifIlte*entt°''TT'T'"""'l developmental rb This stirs up and separates t P helps
ing pace with tlir. meantime f take place The flicking to loosened
and fro spores.
of th shedof the
complete envelope the'^^ i sporophyte keep- shedding ofthe a column in the centre the
The setaatthh'^sJJe tsrJr^^^^ ~ tl formsI elaterophore persists. 7.11B).
-• With four spreading valves ( g-
elongates rapidly.
 126 BOTANY FOR DEGREE STTOENT8_PART HI: BBYOPHYTA
JtTNGERMANNIALES—METZQERHSTBAB : PELLIACEAE 127

begin to g^nrtrbeL^futTyy irf'shed

uic hnea.
pTe'"'
ihis precocious germina- ficial cells of dorsal and ventral surface contain abundant chloro
plasts.
/®\ N^*.
spores is an excep-
uo
6. The apical growth is by means of a single apical cell.
tional feature. Each spore
divides by two successive 7. Some species of PfiZ/ia are monoecious and others dioe
transverse divisions. An cious. The former are markedly protandrous.
oval object consisting of a 8 The sex organs are never borne on stalked receptacles.
number of tiers of celis is
9. The shortly stalked, globular antheridia are immersed
formed (Fig. 7-12C). It may singly in antheridial chambers on the upper surface of the thallus.
be called a sporeling. Each They are either scattered or arranged in two or more rows in the
cell of the sporeling con-
tains chloroplasts, a nucleus broad midrib region.
an^d abundant protoplast, 10. The body of the antheridium has a usual structure. It
ine whole structure, in consists of a single layered antheridial wall enclosing a mass of
■primary RHIZOID androcytes.
^ minute thallus.
„. c^xcepting the basal and 11. The biflagellate sperms are larger in size than those of
Various ^P- diu'ri °^hers most of the Marchantiales. They have more coils. The two flagella
earlier stages. Sometimes the apical or are inserted at different points of the thin, enterior end.
12. The early stages of development of the antheridium differ
from those of the Marchantiales. The primary antheridial cell
divides by a vertical instead of a transverse division. The succeed
(Diag'XSt, °"ulticebuu" ing divisions are diagonally vertical.
S^Syr - re-celied ov^tsfes'-f]V°ung 13. The archegonia occur in a cluster on the upper surface of
the thallus just back of the growing point. They are stalked. The
F-'SfI.?
off semen
s
""'s functtnlr
"»
'°"gitudinal
stalk though short is distinct. The archegonial neck consists of
five vertical rows of neck cells instead of six characteristics of the
'--ofthe ^divide and fedivld^ tT buTupte Marchantiales. Before fertilisation the venter wall consists of two
layers of cells. In other respects the archegonia are similar to
those of the Marchantiales.
The flat A Features 14 Of the three lateral or peripheral cells surrounding
Sis:- ttsf'^'-^s'srs-''
opaque thallus of simpler than ri
«»"■■■
external
the primary axial cell only two divide by vertical walls to give rise
to five jacket initials instead of six as in the Marchantiales.
15. The early development of embryo in PelUa differs from
'"Tr --«*«:«
hexagonal arL"''/''"
5s,v""r;-®"«
°f the tha 11 '"'''"-c-
that of the Marchantiaceae. The bypbasal cell formed as a result
of the first transverse division of the zygote takes no further part in
the development of the sporogonium. It remains as a single-celled
aoDcndage at the base of the sporogonium. The spdrogonium is
entirely developed from the epibasal region of the two-celled
embryo.
defi^neVsliS
Of ^ootH 16 The capsule is raised on a long, slender stalk, the seta.
17 The capsule of shows greater internal elaboration
of Of ate'aS o^and of the Marchantiales. The capsule wall is two or more
eriatrJhick .. Theing cells
cell of theThetwomature
bands. layerscapsule
of the dehisces
capsule wall
in a
develop strength^ lines of dehiscence
■"f"fo'„^vllves which become reflexed. The sporogenous tissue is
'opt that the super differ'enliaTed" it: rpore mother cells, the elaters and .he elatero-
 128 BOTANY FOR BEGREE STUDENTS_PART III : BRYOPHYTA
JUNGERMANNIALES—METZGERINEAB : PELLIAOEAE 129

upon meiosis-^^xL^^reseL^e'of entering generation. The latter comprises the meio spores, the green
thallus, the antheridia, the archegonia, the sperms and the eggs.
tive feature. of a basal elaterophore is a distinc
It starts with meiosis and ends with fertilisation in the life cycle.
The sexual generation is characterised by the haploid number of
another exceptiLal feature. hcfore being discharged is ciiromosomes in the nuclei of its cells.
The fertilised egg secretes a wall around it (Fig. 7T3, 5). It is
of.Lsrnc";foi?o?: ■" the first structure of the sporophyte or asexual generation in the
SUMMARY OF THE LIFE CYCLE (Fig 7-13) A/VrUf/?/D/(/U

general the life cvrip r^f P^7/• • • • ^

'""worts. There are dyeren«s t f details-f" of the sex organs
and sporogonium. As in othpr li\r« 2 'i ANTNER02OID
V.S &AMBTOPHyr£SHOWING
generations,
The two aretheclosely
sexualcScte^
(gametonhvt^?^^^ H consists of two ANTHERIDIUM
ARCHB^GONIUM
«• n
Ibl
m^vidual (sporogonium) remains ovL asexual
exual plant throughout its life anT^'*? ^ connected with the PcLLIA
EGG OR I

and is dependent upon it for its GAMETOPkyTE __

OOSRPERf

ll.S QAMEyOPHYTS SHOWING

ARCHEGONIA

-argin, « out to rthfn CaJ tt

GAMEIOPHYTIC

ssisrS 'ffrr''»■ PHASE

(N)

S NUCLEUS
? NUCLEUS

OOSPORE'^^

rS=Er. r- a SPORE YOUNG

iayered amheridial wall ^ The body rn shortly SFOROPHYTtC
EtABPyO

PHASE
(EN) SPOREMOTHER ■capsule
SPORE TETRAD, CELLS WALL
5
ELATER

hallus just back of ,1'^ ^<='««er on the £LATER0PH0RE

veS''™""'" (%. 7 Is^oT'^? Thev^'^ ofTlf^e SETA

even moreen °r The "ec^k

before fertilisat.? The filled ^rfua ^ - FOOT

^gg or oosphere cpORiMOTER

sperms and the tp- ^^^ity
T) andof th is^wo lave ui5 ^
occunfi CELLS
eggs arethe ventrM ^ ^'^rge Fig. the Ufe oyole.
^'^"ctures of the''::' tL
g^anietophyte Di«grammatio represontation of
 130 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

life cycle. Lodged in the venter it soon enters upon segmentation. By

repeated mitotic divisions and differentiation it forms the asexual
plant also called the sporogonium or the sporophyte (Fig. 713,7).
It IS characterised by the presence of diploid number of chrome,
somes in the nuclei of Its cells. The asexual plant is differentiated
ihoV. ?' t ^ spherical capsule. The capsule CHAPTER VIII
u- - potentially fertile tissue. Associ-
nlrii f meiospores
• a corresponding
as compared withefficient
more mechanism for The
the Marchantiales. dis- JUNGERMANNIALES-JUNGERMANNINEAE :
capsule wall is two or more layers of cells in thickness The wall PORELLACEAE (MADOTHECACEAE)
ous cSs are thickening material. The sporogen ■ General Characteristics. The family Porellaceae includes
and the elaterophore.
elatp T The spore mother cells are the
^helast cells
elaters advanced foliose forms with leaves arranged in three rows. The
They become four-lobed (Fig. 7-13 8) ventral leaves are well developed and usually deccrrent at the base.
The dorsal leaves are incubous and hilobed. The ventral lobe lies
o?iorl -other cell thus foVm® a tetrad more or less parallel to the axis. The rhizoids are scarce and
regSar manner The npe capsule dehisces in a spring from the lower side of the stem in tufts generally near the
jA-nSsVarti -stz," base of the ventral leaves. The antheridia are solitary.
archegonia arise in a terminal cluster on small lateral branches.
The

scopicjr,fcZ5.zSd'l;s'' T~',t'T'r
movements help^n the dispersal ofs^t ti. I®™" All the archegonia in the cluster are surrounded by a common
perianth with a bilabiate and dorsiventrally compressed mouth. The
globose capsule dehisces by four valves separating only halfway
down to the base. The seta is short. The elaters are short, each
with two or three spiral bands. The family is represented by a
single genus Poi'ella {Madotheca).
generations occur
reproductive regularly
cells of one afterJ."
onrgLerat^on the^^th"^
"' The Systematic Position :
tn the life cycle Thi<! nli g^ve use to the alternate plant
■AUernatio„^„TGeJeratio™™"°" pL'ase Bryophyta
Hepaticopsida
1. List the interestS^ye^®^ QUESTIONS Jungermanniales
2 Tr> TT,!, t turea in the life history of Pellia
over that of Sporogonium of Pellia shows an advance Acrogynae or Jungermannineae
Porellaceae (Madothecaceae)
^ ^ pilfl30 of Pctizct Porella L. (Madotheca Dum.)
and development of aex organs
gametophytic phase
Distribution. It is the best known and widely distributed
o-pnus of the advanced Jungermanniales with more than 180 species.
fi-cmandStrcCia!"' ' "8™ of aporogoninm of Peli o, They flourish the most in the tropics. Many species however are
(hnnd in the cold temperate regions Porella plafyphylla
lo be cosmopolitan in its distribution. It has been reported
fiot^ Europe, America, Asia, and India. About 34 species of Porella
av™ been found in India. They mostly occur at various places m
he Himalayas such as simla, Dalhousie, Mussoone, Garhwal,
Sikkim, Chamba, Tirhi Garhwal, Kulu valley and Kumaou. Chopra
(1943) recorded a few species from South India.
131

19^.
Pi«^.
132 BOTANY FOR DEGREE STUDENTS-PART III : BRYOPHYTA
JUNGERMANNINEAE—PORELLACEAE : PORELLA 133
Habit and. Habitat. Porella (Madotheca) is generally found lobe The ventral smaller lobe is called the posticaUobe or the
in the moist, shady places growing flat on logs, trunks of trees lobule. It IS folded back and has an acute apex. It is closely
and rocks over which water trickles. appresseclto the lower surface of the dorsal lobe and runs more
It grows in dense layers forming or less parallel to the stem so as to appear like an extra leaf. The
large mats closely covering the .small vential leaves which are usually decurrent at the base are
substratum. also called the ampfaigastria (Fig. 8-3A). They are simple and
External Characters (Fig. are arranged in a single row on the ventral side of the stem. To
8-1). The plant body is greenish, gether with the postical lobes to which they resemble in form and
leafy and fairly large, up to 15cm. size they appear to form three rows on the lower surface of the
or even more long. It grows flat on stem (Fig. 8-3A). The leaves of Porella like other foliose liverworts
the substratum and is thus ,dorsiven- regularly lack nerves. Even the midrib is absent. Each consists of
tral in configuration. It consists of a single layer of cells (Fig. 8-2B).
a branched central axis bearing The rhizoids arise from the lower surface of the stem near
leaf-like expansions. The axis or the bases of the venti-al leaves. They are few in number and arc
the stem is thin and cylindrical. It thin-walled. Their chief function is to fix the plant to the
is always branched in a bi-or tri- substratum. Absorption is believed to take place directly through
pinnate manner. The branching the leaves.
is monopodial. The branch primor- Internal Structure. The young stem consists of a perfectly
dium lies close to the apical cell. It uniform tissue of green parenchyma cells. Even the epidermal
takes the place of the ventral lobe of
a dorsal leaf. The stem and the
branches bear two kinds of leaves,
large dorsal (Fig. 8-2 B) and small mrfiAL LEAF
Fig.8-1. Porella or lladothoca. ventral. They are arranged in three
Portion of the female plant rows (Fig. 8-3 A). ■>LOBULE
bearing sporogonia.
The dorsal leaves are closely set
cover the stem from above in two lateral rows (Fig. 8T). They
edge of each leaf covers and overlap each other. The anterior
the posterior edge of the leaf in front.

Fig. 8-3 (A—C). Porella (Madotheca sp.)

A. Shoot from the ventral side showing ventral leaves and
lobules of dorsal loaves.
B. A leaf with its lobule attached to the stem seen from below.
A. Portion Of a shoottromL C. A leaf attached to the stem seen from above.
layer is not well defined. The central strand also cannot be made
B. rrrvit:v:raf out. The stem of PoreWo, therefore, is very simple in its internal
structure as it shows no differentiation of tissues. In the older
inc ubons°(Fia *8^2"^ of the I portions of the stem there is differentiation of tissues. It consists
of an outer cortical and an inner medullary region. The cortex
Of the tvvoItlobes
has the
^ upper leaf Js described
(1%8.3B).as is made up of small cells with slightly thickened walls. The med
and is au usually oval in ulla is composed of comparatively larger cells with thinner walls.
also called the antical
 134 BOTANY FOR BEGREE STUDENTS-PART III: BRYOPHYTA
J UNGERJVIANNINEAE—PORELLAUEAE ; PORELLA 135

in thickness. The'cell^ rn o^unifbrm cells, one cell layer The antheridia are long stalked. The stalk bears the nearly spherical
polygonal in outline Ther^ ^ abundant chloroplasts and are body. It consists of two rows of cells. The globular body has a jacket
cell. Growth T 13 no midrib.
It is pyramidal^irfibl?^ place by means of an apical ANTHERIDIAL
these is towards the ven^ral^ Z * cutting faces. One of WALL

dorsal side. ® the other two towards the SPSHAMTOZO/OS

j reproduction
, ^production. Parplln
^imid
^eiospores formed foIInwJnrr ^®P'"°tltices vegetatively and by STALK

ormation isbyconsidered
leproduces vegetativea stag^in P^ctThe
me?L^ sexual reproduction. meiospore
thallus
piocess. hieiosporesare nrndii concerned with the sexual
le life cycle called the sporogoSum iicn-sexual individual in Fig. 8-5 (A—C). Porella sp.
the humid conditions
reproduction propaga^p^^K'*'*'
by the formation of d" species growing under
Vegetative A. L.S. Mature antheridium.
B. Dehisced Antheridium.
leaves of Pni/7? developed on the 0. Sperms. (After Campbell)
who Brazil. This wac a was recorded by
whothin
the discovered
walled,that the so-c^^
rounded L'uclures ion;^- by Degenkolbe in 1938 or wall one cell thick in the upper part and two to three cells thick
towards the base. The wall cells contain numerous chloroplasts. In
The leafy thallus of Pn*- ii u blue green algae. side the wall is a mass of colourless androcytes or spermatids.
S,„S°»"
2^ up"Si:.-srr s'-V-
'csume growth on being
Each androcyte gives rise to a spirally coiled, biflagellate sperm
(Fig. 8-5 C).
(n) Dehiscence (Fig. 8-5 A and B). On the access of water the
mature antheridium dehisces in a very characteristic manner. The
smaller. h The cells of the wall absorb water. The thinner upper part bursts
open into a number of irregular lobes (Fig. 8-5 B). The latter curl
Z IAntheridia."ght angles from the
(a) stem. back to allow the androcytes to escape enmass into the water that
causes the rupture. With the dissolution of the walls of the andro
S(TOc(„e (Fig. 8-4). The anth ' r ,> cytes the sperms are set free.
0 of"tt "reciii {in) Development of Antheridia (Fig. 8-6). Each antheridium
W 'f^bi-anchesofthemale arises from one of the surface cells close to the growing apex of the
Pant Ibe antheridial male branch. It is called an antheridial initial. It lies at the
branches differ from the base of the leaf and enlarges to become papillate. The papillate
cnle branches in two antheridial initial divides transversely into an outer and inner
lespecis. In the first ins- (basal) cell. The inner cell remains embedded. The outer cell
ance the special male projects above the surrounding tissue. It is called the anthridial
^RACT tranches stand approxi- mother cell. The latter divides by a horizontal wall into two cells
^tely at right angles to (Fig. 8-6 A). Of these the upper one functions as aprimary anthe
tne mam stem. Secondly ridial cell and the lower as a primary stalk cell. The latter
undergoes segmentation (Fig. 8'6 B) to give rise to a stalk two cells
leaves ,ItXbracts which
in width and several cells in height.
and imbricated The primary antheridial cell develops into the body of the
smrri antheridia antheridium. It undergoes a vertical division to produce two equal
TKcse^ two features
their axils,
make
sister cells (Fig 8-6 B). Each sister cell divides by two intersecting
walls. These divisions can best be seen in cross-sections^ of the
^he axil antheridium. They separate a central androgon.al cell irom the
of a diS^
differentconspicuous and
from the sterile
gctative branches.
 JUNGERMANNINEAE-EOBBLLAOBAB : POBELLA 137

136 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

outer jacket or wall initials (Fig. 8-6 G). The former is tetra-
nedral in form. The first jacket initial in each half has by now

*4

Fie. S-7. Porella sp.

Fig. 8-6(A-E). Porella (Madotheca) sp A female branch bearing a tenninal claster of archegonia.
Devolopraent of antheridium
A-E. Earlier stages. (Explanation in the text). whole cluster of archegonia is surrounded by a common protecUve
(After Campbell) envelope, the perianth.
divided
in by an anticlinal
a cross-section, at thiswall intoshows
stage, two.two
Theandrn^
bodv of • i^ tu • r The mature tirchegonium is stalked^(Fi^^^^^^^
rounded by six jacket or wlli mZu." The a!:""!;."'"®. or less a cylindncal object. The ^^nal is
of neck cells. They enclose the portion of
usually filled with _8 broad, JJ" is but little enlarged. It
cellscftheSower part 'he the archegomum as ding the venter cavity. The latter
comes 2 to 3 layers thick at the base (Fhr A1 'hus be- S;r.r.3S."» '-
cells m the meantime divide and re-divide in androgonial
drocyte mother cells FanV. e. produce a mass of an- I ru- 8
(u) DeM?oianieaaMFig. r 8).
R^ The
in first three to four^tosegments
leaves.
diagonal wall to fmm divides" by'^a cut off by the apical cell ol the lem^^ fanction as archegon-
androcyte metamorphosis into a Vfl it Protoplast of each Thereafter the derivatives o growing apex. Each
ium initials. They thus enlarges into a papillate outgrowth,
archegonium initial (Fig. 8 ) basal cell and a distal cell.
The latter divides lto™°"t''l'V j, .j.; 8-8 B). It undergoes re-
0 theC^)posterior end of the spirally colled Cy attached The former is called the of the archegomum. The
-^'chegonia, peated segmentation '"f^wonium mother cell. It undei-
gtnSn^ntL'teTusuSfor the other Jungermanniales
fpellia) to produce the eccentric vertical
archegonia arf J • tnales. Xhe u female The archegonmm division wall is the shortest. I^n
usually bears a ^^ort. Each « ^^^"ches bearing walls (Fig 8-8 C). Of jpheral cells arc separated from the
segments cut off i ^ four or five Ip "^^ogonial branch this way three lateral o P .P. them. Two of the peiipheral
primary axial cell suaupunded by
cells are large and l^hc thnd^i^s^thc^s^^ undavided..
again divides by a veiti
duster (Fig. 8-7).
 138 BOTANY FOR DEGREE STUDENTS-PART III : BRYQFliYTA JUNGERMANNINEAE—PORELLACEAE : PORELLA 139

A ring of five peripheral cells is formprl

initials. They sujund the middirpritnaly'^LTa'i clV^The^ swim in thin films of water covering the male and the female plants
and reach the archegonia to bring about fertilisation in the usua
way.

SPOROPHYTIC PHASE
This phase in the life cycle is concerned with the production
and efBcient scattering of meiospores. It is the function of the non-
Lxual indTvidual in the life cycle called the sporogomum or the

aAT£R

L08ED SPORE
/MOTHER CELL
CAPSULE WALL
CALYPTRA

■involucre
Fig. 8^ (A^A). Porolla (Madothoca)sp.
A —F.Development of archegonium.
Earlier stages. SETA

Young arcliegonium.
H. Nearly mature arehogonium. (After Campbell).
, -6-."uiu. lAUerCampbell).

an o^terc^L'dVnlnnL'tu(% transversely to produce FOOT

. Dehiscence. On r
gomum opens at its tip. Th,- of water the mature archo- Ficr S'S. Porella Bp.
of a nearly-nature .poro,oan.„n
oTher and folded back. All this separated from each Longitudinal soction " .. r u d'
resuWng fromThe^^ the mu oohvteb (Fig.
fo'^r.y^goL 8-9).d The
rfpeate' ™dodcsporog?^'
jvts.on"", d ffere^ of cells arid
Ihelrlfowth.' F°Vl"Xl branch (seKual plant) to the top of whtch
tirely upon the archeg throughout its life.
It remains organically attac *♦ ikir...ancnoreS. lt
li is the concern
3. (a). P"d-'=*'^^,e^or'^g™ruCamaturlt;, Is'd'ifferentlat-
"■ " •--= »'s;s of the sporogonium^
ed into the foot, '
capsule.
 JUNGEBMANNINBAB-POKELLAOEAE : POBBLIA
140 BOTANY FOR DEGREE STUDENTS—PART III ; BRYOPHYTA
maliC out the number of the capsule regions
Foot. It is merely a somewhat enlarged base of the seta. It matter of fact the distmcuon into the seta ana
IS indistinct.

Seta. It is short. It carries at its top the spherical capsule.

II thi^ wall.The
cells spherical
Within the capsule
capsuleatwall
maturity, has two
are the to four
spores and YOUNG SPOPi )
MOTHER CELL
e a crs. ihe elaters are short, each with one to three spiral bands, ELATEP CEU
i he elaterophore IS absent. With the maturing of the spores with- —*L0BBO
calyptra
SPofteMonif^
. With\ suddenly
the elongation elongates
of the seta thecarrying the capsule
capsule breaks up-
through CSi.LS CAPCVlE
WALL
the surrounding calyptra and projects beyond the perianth and
tivo ucre. The exposed capsule dries and opens into four sprea- SETA

ifAetptult' iWOtUCRP

K. jungermanniales
Dranch becomes enlarged into the tip ofIttheis female
a pouch-like structure. called
store-house of water and food. The
loot of the sporophyte is embedded in it. Fig.8.11 (A-C). Porellssp.
Thp ygote divides by a transverseSporogonium
wall into two halves
(Fig.(Fig.
8108-IDA)!
A—Fl fs Yr;sro:" —
c" SS"o£ eapsale enlarged to show
Mils onct elater eel s. ^ u„difFerentiated cells
IS obscure
I Tl^ emcey^ show
Thp embryo a of
consists good
a - ij-regularity mintheir
the
(Fig. 8-11 A). ^ reports different distances
SlTo&etpa-;^^^

^ E
^aTffwaUtwoormom^cd^^^^^^ The latter by divisions
Various
g development of the embryo.
(After Campbell)
At a later as ferogenous'tl.ue
.owe^Hypobasal. cease to divi _ remaining ce elater cells. The
in all elongated. They a intermingled. They
divide and becom ceU die spore
The T spore
are notmother eelm a y 8 foui-lobed
arranged „,der r'stiu s. The elongated
tapering ends
tttother cfls beeo-e^deep/^between ^ir
Sr.! elater cells he t" the
Sirdl'dofiSir SeTS'fhf fm"®8-llcV
(Fig. 8 IIC). " 1 pr oof the wall
la^r of the struc-
fouf.iobed spore
-."W .e,ue„ce° b'eUf - (c) Sporogenes^ make
mother cell projects
142 BOTAJTY FOR DEGREE STUDENTS—PART m ; BRYQPHYTA JUNGERMANNINEAE—PORELLACEAE : PORELLA 143

mark 3. The leaf-like expansions arising from the stem are of two
of the four spores (Fig. SUG). They are connected in the
theposition
middle kinds, large dorsal and small ventral.
by a narrow neck which contains the diploid nucleus. The 4. The large dorsal leaves are arranged in two lateral
One ofrr daughter nuclei. rows on the main axis and its branches. Being compactly arranged
they cover the axis from above and overlap each other iucubously.
5. Each dorsal leaf is a bilobed structure. The upper larger
Th: u" lobe is oval. The smaller ventral lobe is closely appressed to its lower
surface. It lies more or less parallel to the stem.
sp?ra7brndl ofIhlrLT4«erkl^
The seta which is developed from
^
the lower oortion nf tKe 6. The ventralleaves are small, simple and reduced. They
embryo remains short. It projects a little bevnnH tK? J are arranged in a single row on the ventral side of the axis.
Together with the ventral lobes of the dorsal leaves they present
protective envelopes. The latter in Pore^Jare the the appearance of three rows of leaves on the lower surface of the
perianth and the involucre. The calvntra i"c i
The perianth is formed by the fS^of tb^r^ axis.
immediately below the' arcLgonial cluster Th^^br f
the perianth become enlarged to form the involucre Th^
7. Each leaf generally consists of a single layer of cells and
has no midrib.
picuousfooiis developed from the base of the seta Tt t 8. Simple, smooth-walled rhizoids spring from the ventral
the enlarged base of the seta. ' ' side of the axis near the bases of the ventral leaves and fix the plant
(d) Dehiscence of the cansulf* Ac. to the substratum. They are unicellular.
mature the seta elongates By this the r \ spores are 9 The elaborate leafy external form is correlated with the
capsule
are the isperianth
raised above the suLu^
and involucre. W no'utfof t? ruptured.^^ich
Dryrn. The simple internal structure. There is slight or no differentiation of
opens It at the apex by crossed exposed capsule tissues. The mature stems are differentiated into outer cortex and
central medulla-
valves. The valves do not extend down spreading
scopic movements of the elaters heln in fl?rK hygro- 10. The scales and tuberculate rhizoids are absent.
some distance. flicking away the spores to 11. Apical growth takes place by means of a single, large
Germination of Snorpc i . apical cell pyramidal in shape.
immediately after liberation • spores begin to germinate 12 plants
female All are
species of Porella
distinctly {Madotheca)
large than are dioecious.
the males. The
The sex organs
» has alsoNeither
the exospore. been reported ^xTe
the exosnfe f 'he spores
proceeds in
within are borne on special side branches.
formed. With the division □r th '^ 'ho germ tube is
expands. The expanding endosonrl st 'he endospore 13 The antheridia occur singly in the^ axils of leaves on the
siderahly. Eventually the latter no , hes the exospore con- antheridial branches which spring approximately at right angles
repeated division of tL protoSart nrod ®'"' "'"hie. The l-om the central axis and bear closely imbricated light green leaves
which are often called the bracts.
IS circular and disc-like It s he of cells which
or rnorerhizoids 14 The spherical body of an antheridium is elevated on a
a large apical cellwhich anchor r toa thr T™®-
is estShed a " Subsequently
develops one long, slender stalk.
The apical cell functions as the growin ■ protonema. 15. Thespirally coiled body of the biflagellate sperm shows,
tophyte. 'he growing point of the future game-
at least, two complete coils.
SALIENT features : . The archegonia occur in a cluster at the apex of a very
short archegonial branch. The latter bears only two to four
delicate object. It 'the which is an elaborate
arising from^thecen'ral axis. ' PP'""'hnoe of leaf-like expansion; The growth of archegonial branch is arrested by the
or 'ripinnltfrnan';;,"''''';' fh'. 'he stem is branched in a formation of the' terminal archegonium of the cluster.
ift The mature archegonium more or less is a cylindrical
wOTt.°isSnt'^-""'?'"® which 'i's''"f'ch-
"t 'h 'he acrogynous "
JurgertrS,"" """""Podial. object with the neck as broad as the venter.
144 BOTANY FOR DEGREE STUDENTS—PART III; BRYOPHYTA JUNQBRMAMMWEAE-P0RELL.4CBAE : POBBLLA 145

enrlnl?A(T
cells
consists of five vertical rows of neck cells
canal containing usually 8 broad, neck canal
ji^i „„.;KTSi==rL""£
.1

The venteTcavltv layers thick before fertilization,

the forme" ^ small egg and a ventral canal cell above £C-S OR ,
RSAofFOR RfRr/UZAT/ON
foot^a shon\«a differentiated into an indistinct 9BRANCH ARCREGOMOM

epibasal half of the embry^""'''' developed from the r£MAL£

QAMETOPHyli

the spo;es?ndTat:!: Thi eW ''"T

The elaterophore is absent Th^ ^ 'P'"''''®- M d-. SPERM ATOZOiD
four-lobed structures before mcioseT°" <:ells become deeply BRANCH

1 ANTHERlOm
MAl£ 6AM£T0Pffn£
valvfs which"xte"nd 'P-'^^'ding lobes or
viiicn extend only half way down to its base.

nema'^stage!' S""'"^don, produces a distinct proto-

SUMMARY OF THE LIFE CYCLE (Fig. 812) fOUNG GAMS/ OSRyTS /^K'''2%^/VUCi£WS
QOSPORE'-i^^^h NUCLEUS

the generations,
two accompanyingIdia^rm'fF^^^Bd'ar'^Lt'kT''
gaLeto'phyte ani'the syoroK
SPORE

this The conspicuous structure c i rOUNG EMBRyO

It is produced by the germ°nadL®='o?TtedP''"' (me.ospore).
The spore on falling on a suitable substmturS
D- SPORE 9
TErP/IO
tion to produce a multicellular disc like nrn+ segmenia- plater
of the protonema is established an ^^ge 3F0R£ mother
CELL
segmentation
It loves to growproduces
on rockstheover
spvni,]
whicf ^ he latter by active
(^■g=8-12. 12),
of trees. It is fh<-mrtc^ ^"ich water trickles and the bark
life cycle. It is chiefly conc3ed"'witr'' "t',t^P^"dent plant in the
sexual plant grows dorsiventrally unon ''^P^duction. The
^.^^entral axis (the so calleH 5t \u and consists
(F'pinnate
g- 8 12,manner.
1). TheThpsn-r^-.]!
iL. sRnder ster'- '"^^f-'ike exoansions
branched in a bi-or tri-
SPORE MOTRSR CfL
dorsal and ventral Thp that it bears are of two kinds,
^hey are closely
completely cover set
from"inabove
two -Th are large and nearly oval,
^hey DiagrnrornatJC v
Each dorsal leaf is divided info ^ incnbously. ■ r ofa
closely appressed to hriower The ventral smaller lobe of leaves of Porei!acons.st o single ° p the gametophyte
^'PP^Y'dTfferentiatiom There is
more or less parallel ?o the ste^ lobe. It runs raidrib. The external '=!f°°u)stological
plant is not correlated tissues.
®hght or no differentiation
 IM BOTANY FOR DEGREE STUDENTS-PART III : BRYOPHYTA
JUNGERMANNINEAE—PORELLACEAE : PORBLLA 147

ly snlueVZn
singly in the axils of IpawA
■"arked-
bears antheridia
TJie sporogonium and the structures produced by it consti
tute the sporophyte generation. All these structures have a
nearly at right angles frorn branches which spring diploid number of chromosomes. The sporophytic generation
antheridium has a globular bodyTprg starts with fertilisation and ends with meiosis. So the first structure
anthendium is single layered in thn ni i of this phase is the zygote and the last spore mother cells. The
thick in its basal mrt hn ^ cell layers sporophytic generation therefore consists of the zygote, the embryo,
of the antheridium ruptures ^The^^bifl ^^1^ apical part the sporogonmm, the spore mother and elater cells. The spore mother
are liberated in the wfte^JLt 3) cells (Fig. 8'12j 8) which are the last structures of this phase become
deeply four lobed and each undergoes meiosis to produce four
haploid spores. With meiosis the sporophyte generation ends.
^mall
branchsidbears
e branchonlyadsbg^fro^^^
2 o? 3 Ws female
The '^1^ ^
archegonial From the account given above it is clear that the reproductive
branch is also involved in the fnrm!l^ apical cell of the female structures of one phase, on germination, do not produce the same
Pium in the cluster. The gmmrnfrc^ phase. On the other hand they give rise to the alternate plant in
the life cycle. This results in the regular alternation of the sex
wit^'th' ^rchegonium I more or branch thereafter ually reproducing generation with the asexually reproducing one.
This phenomenon is called "Alternation of Generations".
Thente^rve"„t rrd^' i ' i :"
wan is ,wo „!! layers thick beforeTrXtbT' "»•
JUNGERMANNIALES AND MARCHANTIALES
COMPARED

Jungcrmanniales Marchantiales
{Pellia and PoreUa) (Riccia and Marchantia)

1. The plant body shows a great 1. The plant body is invariably

and'permsf diversity in form ranging from a flat,
green, dichotoraously branched thallus
a simple, flat, green, branched thallus
with po external differentiation into
{Pellia) apparently resembling that of leaves and stem.
Riccia or Marchantia to tho elaborate
foliose type {Pordla).
2. Tho elaborate leafy habit is not 2. The externally simple thallus
correlated with histological differ shows considerably complex internal
entiation. There is no distinction structure. There is the upper photo-
into the upper photosynthetic and tho synthetic region and lower storage
lower storage regions. There are region. The photosynthetic region
differentiation InA t ' ^" ^be latter bv furfbp ""ated cells neither any air chambers nor air pores. has characteristic air cbambers and
air pores.

3. The thallus is anchored to the

3. The thallus is fixed to the sub-
stratum by only simple, thin, smooth- substratum by both smooth-walled and
tuberculate rhizoids.
walled rhizoids which arise from its
lower surface.
I' lives parasitically uporX"""" '' 'nc^'Pable of self T- 4. Both the tuberculate rhizoids 4. Both tuberculate rhizoids and
and scales are absent. scales are generally present.
capsule region 'of connected tfarou^h^ gametophyte to 6. Branching is always dichoto-
^nd the dispersal 'Pf'^goniv.m is devoted °o"'th'' The 5. The branching in the A.nacro-
gynous Jungermanniales is dichoto- motts.

"lllayers
7"e. Thethick,
latter Withir°tr"'''
gets diff '=='.P®"1'= ~nshts ofasoTd
wall is the waU^t''™''""'""
° ''™'-
mous but monopodial in the Acrogyn-
elater cells. ^ets differentiated into spore mothTr°™f ous forms.

6. The SOX organs are never borne and6,many

The f6x organs in Marchantia
other genera are borne on
on stalked receptacles. stalked receptacles.
 RP

148 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

JUNGERMANNINEAE—PORELLACF.AE : PORELLA HO

Jongermanniales Marchantiales
Jungermazmiales Marchantiales

7, Tho antheridia in the thalloid 7. The antheridia in Riccia occur

forms develop on the upper surface in sunk in the tissue on the upper surface 15. The spore mother cells 15. The spore mother cells do not
the midrib region. They may be of tho thallus whereas in Marchayttia become deeply four lobed before become lobed.
superficial or each immersed in a pit thoy are developed in rows, each in a meiosis.
and are sessile or shortly stalked. In flask-shaped pit on tho upper surface
the foliose forms they arc axillary in of tho male receptacle which is elevat 16. Sterilisation of the potentially 16. A structure like elaterophoro is
position and occur singly or in groups. ed on a long stalk. fertile tissue in the Jungernianniales entirely absent.
They may be small or long stalked. has advanced. Besides the olators there
®- The primary antheridial cell is a basal elaterophoro in Pellia and
8. The primary antheridia] cell divides by horizontal divisions varying apical in many others 1 hough absent
divides by a vertical wall. The suc in number in different species before in Po 'cll'j.
ceeding ones are also nearly vertical. the vertical walls appear.
17. The capsule dehisces in a 17. Tho dehiscence of the capsule
9. The sperms are biflagellate, 9. The sperms are biflagellate, regular and a definite manner along is variable. It opens by several
elongated, spirally coiled structures. narrowly curved siructures. the lines of dehisconco into four irregular teeth or by the separation of
Ihey have more coils and are larger valves. a lid.
than those of the Marchantiales.

1 archegonia are develbned 10. Tho nrchegonia in Riccifi nro REVISION QUESTIONS
L 'dorsal surface of sunk in the tissue on tho dorsal surface
terminally on of tho thallus whereas in Marchantiti
a special side branch in Porella. 1. List tho characteristic features of the family Porellacoae (Madothe-
they arc arranged in rows on tho caceae).
under surface of the lobed recoptacio Describe the vegetative phase of any foliose Jungermauniales.
which is elevated on a long stalk.
3. Give a brief account of tho sbruofcure and development of tiie sex
lo a stalked
IS iTi 1 cylindrical
structure. Itarchegonium
has a neck 11. The flask-shaped archegonium organs of Porslhi {^Madothccu).
consisting of five vertical rows of neck las a nock consisting of six vertical 4. Describo in detail the sporoph/to phase of Pordla.
eel s and a venter wall two or Co rows of neck cells and the venter wall 5." Enumorato tho salient features you come aorosa ill the life cycle of
cell layers in thickness one cell layer in thickness.
Pordla [Madothcca).
6. With the help of n sketch give the life cycle of Porella.
ishypobasal
horizontll^^ l\
region
zygoto 12. The first wall in the zygote is 7. Compare the Jungerinamiiales and the Marchantiales.
transverse. Soon it is followed by S Comoaro tho structure and dovelopmont of the sporogomum of
basal half. The
ly dovsloped from tliA
«pi-
entire- vortical walls in both the halves. Tho
MarclZi^^nAPdUa or PorMa. Which of the two .e adva-neocl and why ?
the embryo. The U "^^1 l^^lf of bisected by the
takes no further narb oetanf quadrant and tho .nent
n^cnt of the SporoCium.^ ■ . ^66=5 in the embryo are absent Jungermannmles.^ points ot reaerablanoe between Porella and Pellia.
voni, ^ dungermanniales. Tho sporo-
embryo
emhvi- in
• tho Marchantiales.
from the entire
yate h
"iternal. it i elaboration is tho low?" r ®Perogonium is simple in
developetl seta and^a^ tdo>f ^ the foo/ which it lacks both
The latter consists of n
2-4 cell iayei-s thick tk wall liigher Marni
ed into n.
whereas in the
it is differentiat-
vertical strips of th,T, ,? four
Thoenno,,! seta and a capsule.
d;h*
demscence* called lines 0" There are V.
"ues of dehisccnce.
14, fV*
.i-!
U. The elaters are not always
present.

t; •. -K, •
Will' 1
MU-tiu ■. .t
 ANTHOC.tSROTALES—ANTHOOEROTACEAE : ANTHOCEROS 151

Notothylas, Mcgaceros and Dendroceros. Of these Anihoc&ros alone

comprises more than 200 species. Lately Proskauer (1951) has sugges
ted to split the Anthoceros complex into two genera, Phaaoceros
and Anthoceros. The former includes species which are characterised
^S^^AFTER IX by :—
ANTHOCEROTOPSIDA (ANTHOGEROTAE)— (j) Absence of large intercellular cavities in the gametophyte.
anthocerotales \ii) Aniheridial jacket consisting of numerous small cells not
regularly arranged in four tiers.
cerotae) Anthocerotopsida (Antho-
in moist, shady nlacp-; v^hich prefer to grow (Hi) Spores yellow and translucent.
warm temnnraf^ '• mostly found in the tropical and In Anthoceros are included species in .which large
PVte, simple in form anfl\ have a lobcd, thallose gameto- cellular cavities are present in the gametophyte. The antheridial
eithcr with or withnnf regular dichotomous growth. It is jacket consists of elongated and rectangular cells regularly arranged
httle internal different! dorsiventral thallus shows very in four vertical tiers. The spores are dark brown or black in colour.
cavities which contn'^ internal
the lower surface of the ^'^^1 open by slit-like openings on Proskauer's suggestion has not received enthusiastic support from
^fue green alga Nostoc t Occasionally the cavities contain a other bryologisfs. ,-I?i//mceros and Notothylas are cosmopolitan in
air pores. Most of the cr> • neither any air chambers nor their distribution. They are ground dwellers and are found m the
It contains a central nvre^^'-!? ^ chloroplast in each cell. tropics, subtropics, and moist temperate regions. Dendroceros
Phytes and Alegaceros chiefly inhabit the tropics and subtropics. iney
to the and higher nlamrw^
substratum hy unknownTheinthallus
otheris bryo;
fixed crow usually as epiphytes on the bark of trees, damp litter and other
The tuberculate rhizoirk j ^"toellular, smooth-walled rhizhoids. suitable substrates. The order contains a single family Anthocero-
taceae. Modern bryologists (Muller, 1940, Proskauer and Reimeis
sex
romorgans are sunken in
the superficial andthe?h.n
anth "j- scales are absent.
^^^^cgonium The
is developed 1954) recognise two, Anthocerotaceae and Notothylaceae.
latter includes a single genus Notothylas.
enrt "PPcr surface of the Th ii^"^ hypodermal cell
parrPT"^ and lie in rnnf antheridia are thus \I/^NTHOCEROTACEAE
The sporophyte is com-
entiatp growing season f ^ for its continued growth General Characteristics. The sporogonium is generally a
Sn ■ r° cansS^ ^ meristem. It is differ- long upright, uniformly cylindrical object arising froni the dorsal
thlciaUn "o sIm meristematic
elonsatpH f^gtn. It ig ^ dome.ci^ archcsporium is ampht-
at the ton ®Porogonium nrrK cylinder parallel with the
wards bv dehisces I the central columeUa
thecemml enr ^^hieh bdn^ h the top down-
a single
gto ordp I'^'V^^spores and Pnf jS^°s^°Pic The
order Anthocerotales curl back
class exposing
includes
ovcr-arched by the archesponum The lauer a™ ^
f^"°^EROTALES
'(vUh";^;7^.f r°™<='ly five gen^a ?y4r l'°* is placed in the fantily Notothylaceae.
Anthocerotales p '''" ^00 specif 4°^' and Dendr"'
™'''" as
air: :,, Campbell VTmo
a Jn,', in I94n '"■®. 'ncluded in the order ..j^t^THOCEROS L
TP'e.
Tp''=' thick"
tlticlc I, ^d elater.
Anlhor.m^.
as it is on a " questioned the
'^e presence
status oi nf
Asp^'O'
Anikoceros, .ailed elaters in . ®'nglc ■"''^'^^uter,
character r,re.er.ee long. ....
Habitat andA Dnutribution
^ Anthoceros is a cosmopolitan
tropical
► andMehra anr S"'"''"" has and their absence genus With In India alone 25 species have
present by fo„r„^a"fino (19531 T^PPafed by Proskauer (iPf®',
universally accemeH """fi'"' 'hus is represented at
P'ed genera. These-are AnI'toceros bLn* r"pS The three common Indian species wh.ch oecm m
 152 BOTANY FOR DEGREE STUDENTS—PART XII : BRYOPHYTA ANTHOCEROTALES-ANTH.OOEROTACBAE : ANTHOCEROS 153
the Himalayas are Anthoceros himalayensis (Fig. 9'I), A. erectui^ ascending, stalk-like structure (Fig. 9*2 A and B). ^he
(Fig. 9 2) and chambensis. They usual occur at an altitude above in?; a cup-like structure. Sometimes the
and fan-shaped. It is deeply lobed with a toothed marg .
SHRIVELLED
COLUMELLA
DEHISCED-"
SPOR060N/UM SPOROGON/UM

m'OLUCREt

mowcfis

RHIZOIDS

>g. 9-1. Anthoceroa himalayonsia.

^nging from 5,000 to 8,000 ft T-i
Kashyap from Mussourie Kuln aT t- reported by
Kumaon.Chamba
^ehra Valley,
and Handoo Puniab's^^l5^\°"'^^^
collected thp Himalayas, and
^'^^^®'^"^Travancore.
Nainital and Dalhousie also. All tL snn
oaL "1^ in very moist sh P.,""', on moist, clay

I
• Features i
withouTT'^^V^^" 3 smalf -^u bryophytes,
divided m ^ ^^'^^^^^ntral'InV. gi'een or dark green
ingenerally
outlin^^lid
more Thn ttllu,
'obed. Thefn lolf^
snm ^Pecies is suborbiculaf
'"^es ®with
folded r^ar^l'^''
^he thallus is smooth in
divided to midrib
forri"^'' ^^ick and fleshy
lof'od and
presence of ridges ti species and rnn tJ- surface of
rhizoidsandmuciwT surface due to the T7{p 9-2(A—C). Anthoceros erectus
A A fertile plant bearing spovogoma.
smooth-walled rhizf^Hc
substratum. Inarlrf!- °^over
anchoritt>ihpa scales, unicellular,
tuberculate B and C. Sterile plants.
areas can be spotted ^oumfc thallus to the In the months r c:„nfpmber andcylindrical,
October the Anthoceros
delicate objects
thickened spots are caUe I > bluish green thickened thallus fruits. It usual y clusters from the dorsal
by a blue-green alRa^n^''^«t«stoc col ^hese Each sporogonium(Fig.
has91).
a tubular sheath
Anthoceros eT. fT''
fiesh-< thallus of thiJ generally a d'ff inhabited surface of the the involucre
species is 4^ different habit. The thick, around it at its base, it is tne m
raised on a thick, upright or
 X

154 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA ANTHOCEROTALES-ANTHOCEROTACEAE : ANTHOOEROS 156

(Fig-9-3). Internally the Everv thallus cell usually contains one large chloropUst^
It is usually oval or flattened
is howevpr^^r^fi Tvr^ thick but without a midrib. There centre. The presence of a ,, i ^re unknown
assimiIatorv\^ ^ ^ differentiation of vegetative tissues. Thus the pyrenoids are characteristic of algal c Y- Antho-
thallus consist^ regions are not recognisable. The entire elsewhere in the pl^it kingdom T ^ ^ ^,^/ioceros
comnact A" Parenchyma. The cells are uniform and ceros thallus, different
Isoetes and
poZ are are, however, from those o ^T . McAllister
fusiform
regular arrancr/>r« \ suiface cells of the thallus show more each pyrenoid consists of a mass of 25 to ^ ^in
larae lens shfn /\i smaller, each with a comparatively bodies. Each of the latter may become a ^^diment^.^ orAntlZceros
There is thus nn not cuticularized. without any change in form and o .
^w^Aorcros there epidermis.
e stoma-like shts In some
on the ventral species
surface of
of the have more than one chloroplast pa pearsoni) to four
the thallus. Their number vanes two ( P
[A. hallii). Each of these ^^tidoses a pyreno apposition to the
■UPPER leus of the cell lies in the cytoplasm m close app
£PID£RM/S chloroplast. , « j uttlp
'^^CHLORQPLAST The thallus of Anthoceros in its Pelliu as it does
mucilage differentiation of vegetative tissues rec tuberculate rhizoids.
CAVITY in the complete absence of Y^^tra sea chambers
'ALGAL £lLAM£//rs There is no well organised differ in the absence of
'if. ♦ f nor air pores m both. The tw , presence of internal cavities
regular dichotomous growth and mi^ ^ p a
'LOWER EPIDERMIS
PHIZOID
Growth. According to ^
apical growth in ^ and his four cutting faces.
CRLOROPLAST > ^ apical cell. It is
It is located in the deeply enra.gma^^ growing pointtheof width
gP^ each thallus
of the
'A'YREA/O/D
lob e. The segments cut off right lower
thallus. The others which are cut off pa^^ Handoo. however,
^''g- 9 3 (A B). Antlioeeros sp surfaces increase >ts thickness of equal rank constituting
A. V.S. Thallus. support Leitgeb
thlgrowing point.whoThey
found several
report that mce.hd^e q ^ ftimaiayea-
They he
B. A single cell containing a chloroplagt.
thallus. These ar/= ^.,11 1 .
often into large rounded imerceUuTar T""' end of each thallus lobe.
S-'" soilage. The mucilage cavitt?„P which con- reproduction^
.._lvai^d by meiospores formed
outlms
inlil-
ror",'h mucilage. Very
®'™on?tZ
Por«"
:u
Th thus serve as Anthoceros reproduces veget ^ however, is concerned
blue gree^aW following sexual reproduction TJj^e Meiospore
assSLin 7'f.^PP=" as smal with vegetative reproductio ^ ^^^g^nium or sporophyte,
HTw'f'™°f"itsalgamay berfit Lrk
th ' containing
S'' =<=" '^Pccks. The • formation is the concern of ^ Ve^etatively the thallus of
t^raS"''
cavitifc
t'"''
1
to rai?e'T'?^"=plantsAnthoceros
species such as M / • without .4?i/Ao^os IS propagated y
brthe"fXwing methods : - portions (ff
-cellsnits the mucilage Jpraanientation. The c j^e progressive decay and
in u ' The mucilage i-? ef j • S^tius PhaeoceTos the
deaththaofluscellsdiereaches
and disorganise ^en th^^ apical growth
'l^XbeT/continuous lobes become
grows
■»"» « £ £"irS'sK separated. Each separa
 ANTHOCEROTALES—ANTHOCEROTACEAE ; ANTHOCEROS 157
lo6 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
is called apospory. Genetically the aposporously produced thalli
common'il'SAocero"""'''''" ihis method is not so arc diploid. They are however normal in appearance.
^ 2e-Sexual Reproduction. Many species of Aniftoceros are
».SiuiTs'i^ ■.». «:E'i .trt i?™"s monoecious, others dioecious. Examples of the formei ca
are A. goUani, A. lo7igii, A. fiLsiformis and A. punctalus. Th
common dioecious species are A. erectu^ A. ^ 7
A. pearsoni 3,nd A. leavis. According to Kashyap ^y. , ■
regularly^devefoD rnnnrl^lT
y develop rounded, marginal Anthoceros
thickenings exposed
called theto tubers.
drought is dioecious. Mehra and Handoo report that
monoecious but markedly protandrous The ^"^lerid a
— ^ The cells of 2 or 3 surface good deal earlier than the archegonia. Ihe sex organs d^ elop with
layers of the tuber develop
corky walls. The corky surface
in the surface tissues and not above them ^d thus i^meised m
the thallus tissue on tlie upper surface. They are i plants
layers protect the inner cells special .sexual branches. However Kashyap reported the male plants
that store starch, oil globules in A. himalayensis to be smaller than the female. According to
'TUBERS and protein. Witli the Bell and Woodcock (1963) the formation of sex organs -
approach of dry season the
thallus dies leaving behind
depends upon photoperiod. The formation of gametes is mit ated by
the tubers. In this state
diminishing day length. Fertilisation thus occurs m winter.
(a) Antheridia. .
himalavonsis.
A. Sterile plant bearing tubers. Anthoceros tides
adverse conditions.
over the
With the y Position and Structn^e (big-9-J & The
o{ Anthoceros are unique m being "°™4 Hyj ^v-jthin
for growth each ^ i . onset of conditions favourable
Phallus. According to
•-fi
occur singly or in groups on Uie upp- surfa-^of .be .hajlus .ith ,
apical ceHs as in the thallns ^ is apical and by a row of The ripe antheridia are bright
organs of perennatinn n i ^^ns primarily function as orange in colour. u<^A^ it ic.
propagation The sterilp B as means of vegetative Each antheridium has ovoid or
bear stalked' tubers h "'"a
margin and nn tb/» i ^ "^^^oloped at the apex, along the raised on a multicellubr 'IjeJacTl rows of cells. In A. laevis it is
Iong^nd cy,ind!ical%T, usually oonsists of foo antheridium, as in the liverworts, consists
Stalks. Occasionally fl ^ develop both on the tubers and thicks-. The body o .jj^ridial wall enclosing a mass of an-
They are, however, sessUe?n^l^ ^"^^ of this species also bear tubers, oi a jacket aye , _ forms a single sperm. The cells of the
of the thallus. Tuber fnrm ventrally or along the margin drocytes. bpch androcyte loims are elongated
yeciessuchas ] r repoiTed in other jacket layer, m P„gular^y arranged in four superimposed
developed on the mar^n^^^^^ They are and reciangulai. Ihey aie reg
tiers. In others (yellow sp
r
, ^ n
consists of a larger
The
reports that the number of each. The plastids change colour,
grow in regions with drv summer' A. fusiformie wall cells contain plastid ' plastids are colourless. Later
these plants dry up Qnlv tb^ ^ approach of summer When the antheridia are y g antheridium the green plas-
withlong
a littlesummer
of adjacent l^bes on they become green, ^^phromoplasts).
the drou'rht and ^P^'^es persist through tids become bng t oiang When the antheridia reach
conditions favourable for grott^T return of ^ii) bursts open irregular-
become modified in some growing apices maturity the roo. ot the ant .haned depressions or antheridi^
*=■
ly. The antheridia
'craters'with now be in
their oiange ^ p^ contents
^ obvious. jacketOfrtETI^S^
layer sepa-
.epprl.d,hTj7;.r 0926) .pd Upg ()901) of wdcer
wdCer either the cells °o -"e^ular separation of the cells of the
:..Jiaiilar
rate {A. punciatus) or tnei fa the distal end of the
ce Is Ofmeristematic
"-"-V
calary the various
rne various parts
calary meristematic
parJi
zone.Of snh
of
zone .nb.
the .c,.
th^ ^ arise from the unspecialised
^he unspecialW
particularly
P^''ticularly the
the■ inter-
inter wall {A. laevis). An enmass get out of the aperture,
antheridium. The the rup-
Finally the sperms are fchar^^ ,ndrocytes.
tare by dissolution oi tn
 158 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA ANTHOCEROTALES -ANTflOCEROTAOEAE : ANTHOCEROS 169

The liberated sperm (Fig 9-5M) is a tiny, biflagellate struc- Soon after, the antlicridial and the roof initials separate from
ture. 1 he two flagella are equal and almost of the same length as the each other. The space formed between them is filled with mucilage
body They are inserted at the slightly broader, anterior end of (Fig. 9 5 D). The other cells surrounding the antheridial initial
he slender body which is slightly curved. The unused portion of gradually shrink away. The mucilage space finally enlarges into a
remains attached for some time to definite cavity called the antheridial chamber. The roof initial
the swollen posterior portion. first divides by a periclinal wall (Fig. 9-5 C). Later the two resul
tant daughter cells divide and redivide by anticlinal walls to form
^"'heridia in Anlhoce.os the two-layered roof of the antheridial chamber (Fig. 9-5 D). The
chambers iSSSimes antheridial initial before segmentation nearly rounds off (Fig. 9-5D).
antheriHM f It then either directly develops into a single antheridium {a4. pear-
^nthcridial chamberantheridium or the
together with the
soni) or it may divide vertically into two (Fig. 93 E), four or some
surface Teli cell This times into more daughter cells (.4. ereclus). Each of the latter func
thethallus T?h '•''= "PP" surface of tions as a antheridial initial so that there is a group of two, four
(Fi» 9-5 A i a conspicuous nucleus or more antheridia in the chamber.
inoutertgmenr,Fig ^rBr^^T^her ^ and
ridial initial e.ra,4 i ig-y o B). The former functions as an anthe-
In either event the antheridial initial divides by two vertical
divisions at right angles to each other to form four cells (Fig. 9 5 F).
cluster Th^ , '''=™'"P^'"'o ^n anthcridium or an antheridial Each of these cells again divides by a transverse wall. The young
ofcd^ovl'^oXrtSTamb*'^ antheridium at this stage consists of eight cells (Fig. 9-5 G). They
are arranged in two tiers {upper and lower) of four cells each. The
four cells of the lower tier are the stalk cells. They divide and
OOF INITIAL
redivide transversely to form the stalk composed of four vertical
ANTHim- rows of cells. The four cells of the upper tier again divide transver
INITIAL sely (Fig. 95 H). The octant of cells thus formed gives rise to the
body of the antheridium. Each cell of the octant divides^ by a
curved wall (periclinaily). In this way eight inner primary
WOr OF ANTHFRlDlAi
CHAMBER
androgonial cells arc separated from the eight outer primary
ANTHERIDIAL
jacket cells (Fig. 9-5 I). The latter divide only by anticlinal vvalls
CAVITY
PRIMARY
to form the single layered antheridial wall. Repeated divisions
JACKET of the primary androgonial cells result in the formation of a central
STALK
INITIAL mass of fertile androgonial cells. The ceils of the last cell gene
CELLS ^-PRIMARY ration of the central fertile mass are called the androcyte mother
ANDROGONIAL
cells. Each androcyte mother cell divides to give rise to two
androcytes. The protoplast of each androcyte metamorphoses into
fBROKEN ROOF a biflagellate sperm.
In some species of Anthoceros[A. erectus) secondary antheridia
may arise by budding from the stalks of the older ones (Fig. 9-5 J).
(6) Archegonia
(a) Position and Structure (Fig. 9 6 F). The archegonia of
Anthoceros are remarkable in certain respects. They are sunk deep
in the fleshy thallus on its upper surface. They lie close to the grow
ing Doint in regular rows and are developed in acropetal order.
In the monoecious species the archegonia appear later, on the same
thallus which produced antheridia.
Each archegonium (Fig. 9-6 F) consists only of an axial row
of usually four to six neck canal cells, a ventral canal cell and
■rn esK There is no sterile jacket layer except the distal rosette
of cover cells forming its tip. The cover ceils slightly project
 160 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
ANTHOCERQTALES—ANTHOCEROTACEAE: ANTHOCEROS 161
surrounHpri of the thallus where it is usually the thallus. Some writers such as Campbell, Bower and others,
the mucilage mounT'^TV of mucilage called however, hold that the archegonium initial divides by a jrans^erse
thallus nffpr or^t Und. ihe surrounding vegetative cells of the or periclinal wall to form an outer primary archegonial c®"^
iumo?S^^ The archegon- an inner primary stalk cell. The consensus of opinion favour
the surronnrhn thallus and in direct contact with first view.
bryoZ eranH®r/'®K,'''™
yopny.es and resembles certain of the pteridophyies. other The primary archegonial cell divides by three^ vertical inter
secting walls. This separates the three jacket
from n'/ l!!®' ®'®'' . archegonium originates central primary axial cell which they enclose (Fig.
the archegoniunx
fU- I. gmentinitial (^Fig. 9-6
of the apical cella1justAccording
close to itto Me^rind
It is mllpH archegonium rudiment which now consists of four ce s Hivides
embedded in the tissue of the thallus. Ihe pnmary axial cell divides
ARCHEGONML ^PRIMARY AXIAL CELL transversely into an outer cell and an inner central cdl
INITIAL peripheral The latter (central cell) functions directly as a prim y j
CELL cell. It enlarges and finally d.vrdes to g.vc rise » 'he ven^
canal cell and the oospliere or ovum (Fig. ^ ® P",. u^der-
which corresponds to the prima.y covei f„m
goes a transverse division to separate
OUTER CELL
the inner primary neck canal cell(Fi^^6 D) ^^t^^
PRIMARY
COVER CELLS
CMALCEL L y cni/pff i
INITIAL
INNER < W H and
CELL
NECK
CANAL
CELLS
. axial
primary .r^n
cell tSr;ra=ly
a central cell further divides into a
primary ventral ;S™?,'..".rsu »d. -'•"• "
CELL
*COVER CELLS j.*-
into twototiers
destined formofthe3 ""h™ divide bybyvertical
repeatedly wails':dlvisioiis
transverse The
'NECK CANAL
CELLS resultant six jacket cells P jj xhey form a single
to form the six vertical ^ Alhoceros
,nLkl around the neck canal eells.
layered being completely sunk m
OROOSPHERE However di^veloDment of the jacket layer of the
the thallus tissue, the fodo^ it is indistinguishable from
OOSPHERE archegonium ^'?u"Lametophyte. In any case the egg towards
by adjacent vegetative cells of the
^ig. 9-0 (A-G). Anthocerossn
Development of Archogonium.^
Earher stages {Explanation in the text^ (c) Mili^aiion. the^ceUs
1". Mature archegonium. fertilisation presents difficuUm^
ofthe axial row eimept g jhe Remains of the canal cells. The
for now filledabsoibs
mucilage with wate ^swells. toConsequently ^he coverA cells
the egg is formed. are
biflagel-
arThtgV,l\lf;t,5S°7"'? initial directly functions as the • thrown off. An open passage down JS ^
late sperm swims dow completely. T^iere-
.ygote The latter ™ells^ to fiUt^l^^^^ ^
and its histologicai
^ygotefa'relormed on a single thullus.
182 BOTANY FOR DEGREE STUDENTS—PART Ul; BRYOPHYTA ANTHOCEROTALES—ANTSOCEROTAOEAE:ANTHOCEROS l63

SPOROPHYTIG PHASE Internally the capsule shows great elaboration and complexity
The zygote is the pioneer structure of this phase. By repeated of structure. In the centre of the capsule is a solid core of sterile
segmentation it develops into an elongated embryo. The latter by CAPSULE wall

SPOPE TETRAD

COLUMELLA

PSEUDO ELATEP
SPOROSONIUM
EPIDERMIS

STOMA

SPORE SAC
mVOLUCRk
■ EPIDERMIS

C-4P5i;Z.£ WALL
> SPORE SAC
COLUMELLA

Fig. 0-7. Anthocoros himalayonsis. SPORE TETRAD

ThalluB bearing sporogonia.
further cell division, cell differentiation and continued growth rapidly PSEUDO ELATER
grows into an elongated, spindle-shaped object with a bulbous base
It IS the sporogomum or the sporophyte (Fig. 9-7.) The sporophy- Fig. 9*8. (A—B). Anthoooroa ap.
tes usually grow in clusters from the upper surface of the thallus each A. Transverse section of capsule.
surrounded at its base by a tubular involucre. B. Longitudinal section of capsule,]
tissue. It is the columella (Fig. 9-8 A and B). The --
cells- constitu-
rivi
di^ploidM inaividual which isMeiospores. It is theThe
the sporogonium. concern of the
sporogonium
produces the spores and brings about their efficient dispersal For this tino- it are narrow, elongated with somewhat uniformly thickened wails.
reason the sporogonium is also called the sporophyte. They are arranged in sixteen vertical rows. In a cross-section they
(a) Structure of Sporogonium. It (Fig. 9-9 A) is differcn form a solid square. In the young capsule, however, the columella
Uated into three regions ;(i) the capsule,(n") the intercSary Sr consists of four vertical rows of cells only. It is endothecial in
origin. The columella extends from the base right through the capsii e
intermediate zone and {iti) the foot. The seta is absent Its and ends a little behind Its dista end. The eh.ef function of
place IS taken up by the intercalary zone which is meristematic Is to elve support to the long, delicate capsule To some extent it helps
ihe sporogonium ranges from one to several centimetres in length" In the dispersal of spores. Some writers look upon it as a primitive
type of vascular cylinder.
tL ^'"bular
ine latter is an outgrowth from the thallus andsheath,
thus isthe involucre;
a gametnnhv Around the columella Is a double layer of elongate but domed
weak imer'calary to^the sooroeenous tissue. Sometimes It Is one cell layer thick through
out tFls- 9-9 A) I' is differentiated from the inner layer of the
part of the s^rophyfe^' In form U conspicuous amphitheciam.' It extends over the top of the coluinella like a
dome-a feature In sharp contrast to the liverworts. In this respect
and cylindrical. It is nearlv nf slender, smooth, upright Anthoceros sporophyte resembles Sptesmitm moss. The sporogenous
length except towards the aoex wL thickness throughout its tissue originates In the meristematic zone where it is single ayered
2 to 3 cms. or in s^rne spede^s uo to iV Usually it is Is called the archesporium. Higher up it becomes a two-layered
at firstproportion
small but turns grey or brown
of fertile tissue. maturity. ^
It has light greena
relatively sporogenous tissue (A. hi,mlaye,is,.s]. rarely 3 or 4 cells m thicknes.
in 4 erectethe archesporium remains single layered throughout.
 ANTHOCEROTALES—ANTHOCEROTACEAE t ANTHOCEROS 166
164 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
elongated, thick-walled, sterile cells of irregular shape. The pseudo-
The archesporial cells have denser cytoplasm. At successive higher elaters are smooth-walled and are^nutntive m function, ihey lack
levels spore mother cells, spore tetrads and meiospores are spiral thickenings and are pluricelTuTar; some times unicellular.
According to Pande the pseudoclaters in A. physocladus are long and
have thick walls with an extremely reduced lumen.
lliiUN vvaiia wivii —j

External to the fertile zone is the capsule wall. It is several

lavers lusiiallv 4 to 6) of cells in thickness. The outermost layer
SPOA£,S

PiEUDO-
tEHiSCENCE
o7th: IvairliMte eU-i- consists of narmw verPc^
elongated cells with their outer walls cutimzed.
LINES
EL ATERE
^GUARD CELLS
'COLUMELLA
the fpidermal layer is punctured by stomata similar to t^bose of th
FERT/L,
CELLS
SPORE TETRADS
CAPSULE
WALL
'gifard The'elh ?r tr'a^ule wfrwithi? the epidermis
STERILE* characteri'stic of the g-rt„phytic thsu. As -e-o
CELLS
,STOMA large chloroplas s "If The
SPORE MOTHER
CEILS

,COLL/MEL LA

SPORE MOTHER
CELLS ?L foot The meristem constantly adds new cells to the capsiile
CAPSULE WALL
atarchesporium
its base. They become propessively rl>ff"entiated^mto co^ume^^^
and capsule wall. The^re^ence
EPIDERMIS meristem enables the capsule to grow tor a long p
Spores
■/iRCHESPORIUM
',-•1 Foot It is a rounded, bulbous structure deeply
involucre
cTpsS:
The region of contact ^^twee placenta. The foot
well marked
mainly consistsinofmany species.
a tnass o pare . ^ g ^gUg tubular,
The surface cells
rhizoid-
MERISTEMATIC of the foot. serve to increase the absorptive surface
ZONE
like outgrowths. The lat of
of the foot and penetra soecialised to function as a hausto-
FOOT
Anthocer^s abS'^foo^and^wateTfro^ the parent thallns for the
TISSUE OF THE sporophyte^ Snoronhvte. The wall of Anthoceros cap-
'gametophyte (?;) d typically ventilated, photosynthetic tis-
sule has a v\ ell ^rogonium to become at least partially, if not
A sue. ItenaWesthe spo supply of carbohydrates is concerned,
fully,
It has self-sufficient
been observeds that ajoun^^p^,^P
young sporophyte covered with aproof
^ positive tm-
foil receives a considerable amount of its food
of the fact that It suji ^^^jophyte. For water and tninerals m
c ; Lr.- .i—«- supply ['"depends
solution it eilLiiv y r
upon the parent gametophyte to which it
D. AA stoma
D. Btoma irora
from the Burface of the capsule.
^

pTutelat'rslFj.^-Brandt! Ttrfor"m E of one to four

.V- .
.,w.. l^iy. • •
 NTHOCEROTALES—ANTHOCEROTACEAE : ANTHOCEROS 167
166 BOTANY FOR DEGREE STUDENTS—PART m : BRYOPHYTA
is attached by the foot. The sporogonium of Anthoceros thus is a tier (formed from the hypobasal half of the zygjte) form the foot,
semi-parasite. whole of it or at any rate a major part of it. _ The uppermost tier
develops into the capsule. The median tier gives rise ^amly to the
(c) Development of Sporophyte (Fig. 9T0). The fertilized intermediate zone and in part to the foot. The cells of the -
enlarges until it fills the venter cavity. It then secretes a wall diate zone remain meristematic. They establish
meristem at the base of the capsule. According to Mclean and
Coik tl^se early divisions in ^. laevis result in an embryo composed
COLUMELLA of four tiers of four cells each. The highest tier forms the capsule
AMPHlTAECJ'.m FAIM. WALL and the intermediate zone while the lower three give rise to the
LA-fEB
EHDOUEaW foot.
PA/US
INfMuntP CAPSULB
meAMMrE C/? ^WALL
lONE A/iCHES- divideThe of the foot
in aUcellsdirections to form
tier a(Fig.
rounded
9-10 Dmass
and of divide and re-
E) P^bymatous
PORIUM
FOOT—
-HWOT
■SPOAOGE'
NOUS
LASIB bZ'peneZtrtLe dZfoT'he^geZtophyte. They are hausto-
1 CORTEX rial in function.
SPOBQGENOUS IN1EBMEDIA7E
LAYER ZONE
The four cells of the uppermost tier which are destined to
COIUMELLA' f m the caosule
form p . divide by ^ periclinal
surrounding This separates
walls. peripheral ones
fF^g
;
lO^S ' "o?ZSomerstitute the endolhecLum and the
bryologists hold that the four top
tt
the sterile columella. ^h^latter m we y o
I
J'
Anthoceros BP
A—E Development of Bporogonir. '^- :J:,/~'formTfrom die Inner cells r.ulting horn the pen-
'■ in the development
w ^
of embryo.
ui emur>"
,
^• L S of an advanced (Explanation in the text).
Btaa« clinal division of the amphtthecmm
G red cells of the series function as jacket o ^ anticlinal and
^-ronndi. After b' " ^ or wall initials
perichnal undergo
walls to form t e p outermost layer of the
segmentation. of cells in thickness (Fig.. ^ ^ ^omata (Fig. 9-9A and D).
capsule wall IS epider within the
The emhrt? The npvf •• ."^^^ides the embrj^ The epidermal ^ . -phey as a rule, have two large
cells are ill' "if" insists of fo,,, 7/"°" transverse (Fig. 9'10®'; epidermis are rarely as many as four. There
^«porr,»i^^g"'hanthrio""'' '^ese the upper two
In cither ev <Iivision is trl '' Bhardwaj, however, S^TnSlZr't'aces between Ihem. The photosynthetic tissue
calwaUarriAt'^*°I''he resultanr?''''' TrntlTunStes with'the exterior through the stomata.
SSht cells arSl ",g ?®'o the first So"'
This is the oct 1"° tiers of T embryodivides
nowbyconsists
a vertio The single-layered
i?"- ^re wastage of the lb <=^113 each (Fig. 9T0 D; axis of the capsule ^ extends nearly to its
Hindoo zone
"ediate the InTb"7
." 'be wcciMsl The four cells
lower fnf' According to of the upper
Mehra and apex of the uinel a (Fig _9 10 ^ an ; distinguished
base. The cells of he a^^ ,heir denser protoplasm,
In ,b ' der Zs "nd the inter- from the sterile "^^'J^Lpoiium remains single layered throughout
In some species r ^fnns as the primary sporogenous layer. In
of the uZ of the [.fherspZfH be"wo.layeredV!g-9-10F) above the base of
The embryo ^ever, the four larger cells
""'PPerrnost.f'""ofthree /• """^orse division (Fig. 9T0Cb
are «i|| T of four ce
'"rger."o"The fourcellscellseach.
of theThelow"'
T

168 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA ANTHOCEROTALES—ANTHOCEROTACBAB:ANTHOCEROS 169

the c:;psule (A Instill others it becomes 2 to 4 (d) Dehiscence of Capsule (Fig. 9-9 C). The spores ripen
cell laye^ thick by periclinal divisions and forms tlie sporogenous basipeially from the top downwards. As they ripen the mature
ssue.^ i he sporogenous tissue differentiates into the foliotving portion of the capsule turns grey, brown or black depending upon. e
two kinds (Fig. 9-9B) of cells species. It loses water, shrinks and consequently ruptures longitudi
I'ertile spore mother cells. These are large cells, oval nally along the lines of dehiscence. The longitudinal slits extend
to spherical in form. Each contains granular, denser cytoplasm, a basipetally downwards. According to Mehra and ^ ^"2 °ectZ^nd
two such lines of dehiscence on the capsule wall ™™
distinct nucleus and a chloroplast.
ppll. mother cells. These are slender, sterile
A. Mmalayensis. They lie on the opposite sides m a sha^ ow
denression Each line of dehiscence consists of two rows ot thick
elaters nuclei. They give rise to pluricellular pseudo- walled epidermal cells with their adjacent walls remamtng ^
i along which the slit occurs. Thus the ripe
lavpr^'^!?
y regularly
bands. As the .spore mother cells separate alternating
from each other, usually dehisces basipetally along two '""S'^dinal sins which eirt^nd
downwards from near the apex Th= P-t^ons oj the cap^uk waU
between the slits are commonly called ihe valves i
PSEUDO-ELATER separate progressively of the
COMMON time. The contmuous, downwa^ , f-rLf more and more
SHEATH the underlying tinftt vaT-
SPORES
Ty dr^itofXcoluLllaan^^^^^^^^^^
in spite of the absence of ^P'^l th^<="'"= ,he
9"1] (A —C). Anthocoroa.
A. Spore tetrnd.
rxpoled^reTalsoH^^^^ —^
B. Spores. air currents. ^ i.„-i
C. iPseudo-slaters.
eeuuo-eiaters. According to Sstance^Trom the under-
'S where the shrinking capsule wa gjaters It occurs along the
"Chains of 3 or lying solid mass of of cells and extends progressively
spiral thickening cLrlcteXtTrVf They lack the thin adjacent walk of the tw underlying spore mass is
a sort of a network In their ° f pseudo-elaters form downwards and ^ exposed fertile mass are hygro-
The latter ™dern;\he rndotic'dMn' "^P^er cells, exposed. The pse7<^"®^^7 movpments with the changes m the
tetrad (Fig. 9-11 A,). resulting in a spore scopic. Tbey exhibit twising movements cause active back
moisture content of mass on the shrinking capsule wall,
mentonhrardlinorin^^^^ withregions.
the establish- pressure of the exposed • j _ Qjher lines of dehiscence.
growth is by the activity of the basal^?nt Further This ^ntAoeeros
The results in more
cap^u= 't^JPP^;„
= ®o Prolkauer
from dehisces by a varying
1-4 according to the
ram^T Continually adds _ meristem in the number ol sms. j-ii
cXmH, become prog^estv^r aS' the The mature spores are usually j-hape
the orr, ^'"chesporium and the r ^^^''cntiated into the
periSr^'Tr'^ *\<»'=h!^cence o?the cant, Consequently and thkk-walled. The thick spore wall is^ and beset with small
coats. The outer exospore. IS
sporophytes ofte tl^r^Sr^ ^ ~TVo°Ihe slr'tTve" spines or tubercles. species. It is black and warted in A. er^tus
Earlv orr/-.t.,»U _r.i yellow according to th p numerous, blunt
In A. hzmalaye^sis it ^y Within the spore wall is
papillae. Ihe inner en P ^ nucleus, a colourless plastid
the the tiny
and protoplast.
reserve It coma
food matenai - ,p. &g.jg). According to Campbell
the
Germination oi &p ^ usually enters upon a
(1928) the lil)erated spore m j j
T

170 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA ANTHOCEROTALES—ANTHOCEROTACEAE : ANTHOCEROS lil
resting period. Mehra and Kachroo(1962) who studied germination into the tube where the single turns greem Tw^^
m A. erectus and A. punctatus report that obligatory resting period successive transverse wails are laid at the distal end o ? ,
tube. These separate two cells at its apex. nlace in the
divides by a vertical wall. A similar division p
lower cell. A group of four cells is formed, ^er i octant
angles to the first"^ divide the four celled structure mto a^
The four apical octants constitute the ® First
sporeling. No apical cell with two cutting faces voung thallus.
rhizoid appears as an elongation of any ce of the
t ,i T.S.Oft On its ventral surface appears the first mucila
growinl point. Development of rhizoids and mo e
follows^ and finally Nostoc infection as the growth proceed .
SALIENT FEATURES
GERM TUBE
(a) Gamefophyte. 1. The gametophyte is a simple thallus
usually without a midrib.
A . ySPOPE
2. Except for ™ooth-walled rhizoWs the
the thallus bears no appendages. The scales, tne u s
the tuberculate rhizoids are all absent.
iPICAL CELL
3 Internally the thallus is very simple in structure.^J
shows no differentiation into photosynthetic and storag^^^
There are no air containing channels nor P],ymatous cells,
thallus consists of parenchyma composed of cWorenchym
JHALLUS LOBES 4. A striking feature of the unusual
usually a single The^l^yophytes and higher plants.
J^the^Xr thTchloroptsts are discoid and occur in large numbers
in each cell and have no pyrenoids.
RHIZOID

hi"to larger cavities filled inhabited hy Nostoc.

GROWING POINT
*-MUCILA6E
CANAL The c^avirc— ^PP^ ^
-n. A.,-,i-r' sJSf—s
""'■a Th. .» «l- •"
branches. The remarkable feature ^^.^er than
the luradiate ridge The j of germination
exospore the
runtures aloncr sed in the thallus t-sue. Jhey
cleft in the form ® f a tube 'o"f "ough thf above the surface o
9. The
in origin. They occur
too'of a° roofed antheridial chamber,
germinal tube (Fig. 9-12 A). The 00^? r f cflled the
contents of the endospore migrate singly or m a cluster o
 ANTHOCEROTALES-ANTHOCEROTACEAE : ANTHOOEEOS HS
172 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA

10. The antheridium or the antheridial cluster in the closed

the cells. The stomata lead '"to the air containmg^ch^^^^^
chamber together with the latter's two layered roof is traceable latlng the sub-epidermal, assimtlatoiy ,
to a single surface cell. It means that the antheridial cluster in Anthoceros thus carries on photosynthesis active], .
the chamber and the overlying two-layered roof are homologous to
the single antheridium of liverworts. foimis 20.
only The archesporium
a small paiu of the whioh
spo p 1 stedllsatlon of poten-
advance.
11. The archegonia are fully immersed with their venters and tlally fertile tissue in Anlhoceros. fancllon
necks confluent with the adjacent vegetative cells. The distal ros- 21. The arciicsporium is amp ^ to the
projects above the general surface
Each young archegonium according to Proskauer, "jT'sutSi'ds'lS.'^-^-genons tissue fcrms the spores
and Mehra and Handoo is surrounded by a mound of mucilage.
12. The usual archegonial jacket of sterile cells enclosing "s -s"-
the axial row of cells is not recognizable. Protection to them is cells, is fenced Irom the entire en There is thus transfor-
probably afforded by the adjacent cells of the thallus. livcrwons gives rise to the to the sterile con-
niation of the entire thus reaches
13. Unlike the liverworts the archegonial initial directly dition in .4H//«Jcems. The ®P°'°^^Lralion The latter is internal
as the primary archegonial cell. There is no division of a high degree of complexity and elaboialion.
the ft^mer into the primary stalk cell and the primary archegonial cell rather than external. „c,iallv by two valves
y a horizontal wall. The absence of this division explains the 23. The capsule intermixed with pseudo-
sun en condition of the archegonium and its lateral contact with
the adjacent cells of the thallus. which curl back exposing the mass ol spoi
Caters on the OF LIFE CYCLE
, ■ ® neck canal cells in Anlhoceros are derived from
° primary axial cell which corresponds to the The 'iP'
cell cover cell of the liverworts and not from the central
* I?' slender,
upright, Sporophyte. 15. The
cylindrical sporophyte
object of ilnfAoceros
surrounded is a long,
at its base by a ™ rsVpii-. '"p'-
tubular sheath, the involucre. ® 1 Haplophase (Fig. 913. H)- The spores
16. It is differentiated into a bulbous foot, an intermediate phase starts with the Mother cells by meiosis (Fig.
one composed
right, cylindricalofcapsule.
intercalaryThere
meristem and
is no an unusually long, up
seta. h They getmina'e t'nder suitable conditlmis each foi^ng
a sirnple, green
17. _ The foot becomes specialised to function as a hausto- lo grow in moist, shady places. s pyramidal n shape,
/"rmshed with rhizoid-like processes which penetrate deep melns of a four-sided
Mehra and Handoo j. 1
the grc^wlng point is consti-
solution thallus tissue to absorb water and mineral salts in tutedbyagroupofinit great size. It is simple m its
The thallus never ^^^elophyte (th^
intercaLv the apical growth of the capsule ceases the • uernalorm^parenchyma.
;rnl Us^^Sly'vvlh
structure. Internally
cells the ^
each gt""" one sometimes more
ated Into the tLues of the c I „ become dlfferentl- »?ts^:ra
^Woroplasts lat'p size
of a large size. _ Each
Each bidependent. tteoughout
wall. Thus capsule
period from a basal intercalarv continues to grow for a long
considerable perior In^
in the bryophytc° and yields spores
Anthoceros standsover
alonea
face layer oT^L*'wafr^{enide^^^^ ^^ck. The sur.- thallus is fixed to i"
and the component epidermal cells i stomata only.
epidermis IS the assimilatorv tiss. l 1 i intercellular spaces
Within the
between
 HBw

174 BOTANV FOU degree STUDENTS—Dart lil: bryodhyta AUTHOCEBOTALES-ANTHOOEBOTACEAE : ANTHOCEROS 115
gametophyte at a certain stage bears the sex organs.
They are deeply sunk in the fleshy thallus. Normally the The antheridia produce 'jff
archegonia also are embedded
{f| thallus lateral
xle;
are thus histologica ly "ntnmous with^he ^^ove the
ess OR

ARCHEGOMUM
OOSPHSRe
rt^mattrrUy\Fi"Vl^^ cover cells are thrown off to form
an open passage for lire sperms to reach the egg.
ANTHERIDIAL All the structures
I"" CHAMBea
bAMETOPHYTe
V tophyte plant constitute the P jj(A)number of chromosomes
ANTH0ZER05
SP£RMATOZO/D
They are all characterized by a hapl ( ) flattened
ANTHERIDIUM

diploid number ofchromosomes "atle by differentiation of

S NUCLEUS forms an embryo (^'S' 6h The atte vegetative
OOSPORE.
? NUCLEUS
cells and continued grow h d^elops^mto (Fig. 9'13, 7
individual in the life "jV"' „_orophyte as it produces meiospoies
and 8). It d-'ST Tte sporophy^^
PHIZO/0 and aids m then efficient p bryophytes in many "Tccts.
YOUNG CAMETOPHYTE earlier stages in the is a structure unique among yP ^ cylindrical object
development Of EMBRYO It is a long, slender, upright, green, ™ ^ is differentiated
L'ounde'd at Us base by a tubular ^^^^rintermediate zone
into three regions the foot ^ e
and the capsule. Yjte outgrowths from its fr ace. These
gives out tubular, rhizoid-ii gametophyte and absorb water
penetrate deepininto
and minerals the "jsu
soluuon ® ^ xhe^ intermediate
for the spoi^ spindle^zone
It
tetrad
fs meristematic. The capsule is sloped ^ The superficial
YOUNG SPOROPNYTE
consists of several cell ^dermis. It is punctured with
CAPSULE WALL- EPIDERMIS lPstji£
SPORE MOTHER- WALL
CELL

rsroMA
COLUMELLA

ASEUDQ-CLATkR makes the sporophyte.partia PP ^ part of the capsule

SPORE MOTHER
CELL
FOOT-

ADULT SPOROPHYTE
e a s.)
 mm
ajaSn

176 BOTAJjy FOR DEGREE STUDENTS—PART III; BRYOPHYTA

ANTHOCEROTALES—NOTOTHYLACEAE : NOTOTHYLAS 177

embryo and the sfioroph^te. The spore mother cells are the last Thallus (Fig. 9-14A and B).
structures of this phase. Another unusual feature of Anlhoceros (a) External Features. The delicate, thin thallus is dorsi-
porop yte is the presence of the basal intercalary meristem in ventral and grows flat on the substratum. Usually it is orbicular or
thethe
of median zoneceases
capsule of the (with
sporogonium. As soon asofthe
the differentiation theapical
capsulegrowth
wall, SPOROGONIUM

hpLl. columelia). the intercalary meristern

ihTcJL'T^,] base of the capsule to
the cam.np archesponum and the capsule wall. As a result,
ofncwsnn^^^^^^ a continued succession
hJIa ? T ^PO'"ogonium of Anthoceros is therefore a long-
mifns alive ^o grow as long as the thallus re-
THALLUS
acrnrnTi',! green capsule turns grey, brown or black
wal^atm.n?-. !i dehisccnce. The capsule
annpn along
appear 1 "i7the dnes and shrinks.
two opposite Consequently
lines of dehiscence. longitudinal slits
The slits extend
The two segments or valves of the capsule wall curl
seDarat"pH' underlying spore mass. The valves thus SPOROGONIUM
valvefanH hygroscopically. The twisting of the
^posed
exDosed spore
^innrp mass assist m movements
ysroscopic the shedding
of of
thespores The free in
pseudo-elaters snores
the
SS'c'olld I protrudes between the two valves.Th- dried up rh^eaT-
iiKe columelia

cycleI of^yi//i0cero5
ph^ists or generations
regularly constituting
occur one after the other.the
Thesinde life
life cycle
of this type which is characterised by alternation of generations
and sporogenic meiosis is termed diplohaplontic.

2. NOTOTHYLACEAE

rpo Characteristics. The gametophyte shows slrong

suZv^ Anthocerotaceae. The sporophyte, Jiowever, is \ RHIZO'OS
hnr i u r sliort, compact, marginal and grows out
''^'= lobes of the rosette-like thaflus C
are lacking The cXmeM ^
developed and central I'"^ell
Others. The osenHn Pi f in some or entirely lacking in
Fig. 9-14 (A—B). Nototnylas indica.
AesporesTuh
genus ^'otothylassptal o?„7,,r
^ i" sizeincludes
or longer than
a single A. Thallus bearing sporogonia.
B A portion of the same slightly enlarged.

NOTOTHYLAS 7„nt irregular X^h'anchor I m%he

Distribution and •ti
of species which are wideiv d.Jirih comprises a number sXratur As in' Anaoceros. the scales and tuberculatc rhrzo.ds
are abs^t structure. A vertical section of the thallus lobe
■dund on nS «dtU. revea/s ihl"n " 6 10% layers of cells thick in the mrddle portton
P eauh in humid, shady localities.
178 BOTANY FOR DEGREE STUDENTS—PART HI: BRYOPHYTA ANTHOCEROTALES—NOTOTHYLACEAE : NOTOTYHLAS 179

which forms the midrib. Gradually it thins out in the expanded second division in any case, consists of four cells. It is the quadrant
portion of the thallus to a single layer at the margin. The .super stage. Each quadrant divides by a vertical wall at right angles to
ficial limiting layer at the upper and lower surfaces shows a more the first. The embryo now consists of eight cells, arranged in two
regular arrangement of cells which are smaller in size than the cells tiers of four cells each. It is the octant stage. The cells of the
in the interior. The cells, however, are not cuticularised and thus upper tier undergo another transverse division. The embryo now
not organised into a distinct epidermis. Within the two limiting consists of three tiers of 4 cells each. The cells of the uppermost
layers the thallus consists of soft parenchyma cells of uniform tier by repeated divisions, differentiation and continued growth form
pattern. Apart from the internal cavities which contain mucilage the capsule and the intermediate narrow zone. The cells of the
and occasionally a blue green alga Nosloc, there is no differentiation lower two tiers form the foot.
of vegetative cells. The mucilage cavities open by narrow slits at
the ventral surface. In N. javanicus the mucilage cavities are Periclinal divisions appear in the cells of the uppermost tier
absent. As in Anthoceros the cells contain only a single large chloro- which is destined to form the capsule. This division separates the
plast which includes a body called the pyrenoid. inner endothecium from the outer amphitheciuna. The fate of
the endothecium varies in different species. Depending on the ^te
Sexual Reproduction. Notothylas normally reproduces by of the endothecium the different species Notothylas are divided
meiospores formed following sexual reproduction. Some species are into two categories namely columellate species and non-colum-
monoecious and others dioecious. The common Indian species mellate species.
are reported to be monoecious but strongly protandrous. The sex
organs (antheridia and archegonia), as in Anthoceros, develop within (a) Columellate species. In N. indica and N. orbicidaris
the surface tissues and not above them. the entire endothecium forms the columella. It is central in position.
1. Antheridia. The antheridia occur in clusters of 2—6 on
the dorsal surface of the thallus in cavities called the antheridial
chambers. The antheridial chambers are roofed over at first. CAPSULE WALL
With the ripening of antheridia the roof ruptures and the orange
coloured antheridia standing in a cluster at the bottom of the
antheridial pit are exposed to view.
The mature antheridium is a large, oval, shortly pedicellate SPOROGENOUS
object. The body of the antheridium consists of a jacket of sterile r/ssuE
cells constituting the wall. The latter surrounds a mass of small,
cubical cells called the androcytes. The protoplast of each androcyte COLUMELLA
forms a single, biflagellate sperm.
2. Archegonia. The archegonia occur immersed close to
the growing tip with the necks and venters confluent with the
adjacent vegetative tissue of the thallus. As in Anthoceros the
^chegonial neck is much shorter than those of the other bryophytes.
e neck canal is, however, wider than that of Anthoceros. It
snp ^ neck canal
^rchegonia cells.
develop In same
on the the thallus
monoecious
which protandrous
had earlier
produced the antheridia.
SRMEDIATE
ZONE
ofI lertihsation are the same as in Anthoceros organs and the method
FOOT
aporophyte
sation, zygote formed as a result of fertili-
starts dividing.
livlriir.^ Th efiventer.
j- - .It then
. secretes a wall around it and Fig. 9-15 (A-B). Notothylas.
the first categorv b^i ^jyision is either transverse or vertical. To A. V.S.. columellate sporogonium of N. indica.
second N. levieri and and N. orbicularis and to the B V.S. Non-columellate sporogonium of Notothylas
leviori.
cal, the second is first division is verti-
se or vice versa. The embryo after the
 AnTHOCEROTALBS—NOTOTHYLACEAE ; NOTOTHYLAS 18l
180 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
cells with thick walls. The stomata are absent. The sub-epidermal
The amphithecium cells divide by a periclinal wall into outer and cells either lack chlorophyll or develop very little of it.
inner cells. The cells of the outer layer form the capsule wall
and the cells of the inner layer function as the archesporium which
is dome-shaped and arches over the columella. The archesporium
when fully formed consists of 4 layers of cells. During further
development the archesporium differentiates into alternate bands of
fertile spore mother cells and sterile cells. They extend from the
columella to the capsule wall.
^ {&) Non-columellate species. In these the columella is SPOP^..
entirely absent. TV. chaudhurii and N. levieri are the examples of
this group. The entire endothecium is fertile and forms the arches elateps
porium. The amphithecium forms the capsule wall. The arches \

porium becornes differentiated into alternate bands of spore mother

wlls and sterile cells. They extend across the cavity of the capsule,
ihe spore mother cells, as usual, undergo meiosis to form the meio-
spores.

r, are a few species of Nolothylas which belong to neither

® / C-APSULE
\Ll t ? mentioned two groups. They form a connecting WALL
1007 , columellate and non-columellate species. Lang
<5nnrr.n>i' the intercalary growth of the
archesDorhim the bulk of the endothecium forms the archesporium^
rpct nf tViP 1 ti, the end of intercalary development the
H sterile and forms of the columella.
^ V .u
N javamcus observed
the endothecium normally develops into that in
a well develop-
e columella. In rare cases the columella is reduced in size and
produces spores in the upper part. intermediate
ZONE

tion sporophyte. The sporogonia are marginal in posi-

Each snnrnann'^'^^ orizontally between the lobes usually in pairs.
Sh aZ 0 ? A- compact structure about 3 mm in
towardrtth "Trf? I' cylindrical in form but tapers
surrounds the sporogonium UisTh'"-''''"?'
such as N.indica \rtA n' , In some species
enclosed within 1 ^^e sporogonmm is completely
slightly projects bevon^h it FOOT
into three rigYons thffom .sporophyte is differentiated
capsule. ^ intermediate zone and the A
Fig. 9*16(A—B). Notothyias levien
is triangular in^^shaZ smaller than that of Anthoceroa. It A. L.S. young sporogonixim.
(thallus). The suDP^firi^il of the gametophyte B. L.S. Adult Bporogoniuin.^_^_^_^^_
rhizoidal outgrowths which k produced into small,
parent thallus. absorb water and nutrition from the
Wi'Hin the
little nieriste^mrtTc^aawTiv^f short and slender. It shows rounds the central colume differentiates into alternate bands
ry. ^ Pi^ys no significant role.
role. oCnlSVsterilTSi-J cells. The former gtve
ness.
The outermostlay^1s\he''e^^^^ four layers
yer the epidermis. of cellsofin brown
It consists thick-
 182 IBOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA Anthocerotales—CONCLUSION and discussion isa
rise to the short, simple, unicellular elaters. They are irregular Evolutionary Position. The evolutionary position of
in form and have rudimentary spirals on their walls in the form of Notothylas is controvertial. The bryologists hold two opposite views
short, curved, thin or thick oblique bands. The spore mother celLs on this point. Lang in 1907 put forward the view that Notothylas is
undergo meiosis to form the spore tetrads. The meiosporcs in a reduced member of the Anthocerotales. It has evolv^ from
N. indicu are opaque and dark brown. The spore wall is differentiat forms like .^nfAoceros by the process of simplification. He thus
ed into the outer, thick exospore and inner thin endospore. considers Ajithoceros a primitive genus of this group because many
In the capsule of tion-columellate species the columella is of the unique features of Anthoceros sporophyte are either absent or
absent. 1 he sporogenous tissue is formed from the entire endo- present in a modified form in the Notothylas sporophyte. it has
thecium. As usual it differentiates into alternate bands of sterile lost its columella. There is no photosynlhetic tissue in the capsule
cells and spore mother cells which fill the entire cavity of the capsule wall and the stomata in the epidermis are lacking. The basal men-
within the capsule wall. stem is little active. He reads genetic relationship between the
Anthocerotales and the liverworts in a cloivnw'ard direction tbrou^h
Dehiscence ofthe capsule. According to Muller the mature forms like Notothylas. The supporter's ^ang s hyp^^
capsule dehisces by 2 to 4 valves. Kashyap, Dutt and Pande reported consider as a prunitive genus
that it is basipetal and by the imperfect separation of two broad advanced but a reduced genus. It has evolved from Arith)ceros by
valves. Later Pandc in 1932 reported that in N.indica and N. levieri, a process of simplification.
the dehiscence is follicular. i'hc capsule opens by one suture, some Cavers in 1911 advanced a view diametrically opposed to that
times, by both sutures. The horizonial position of the capsule, the of Lang, He considers Notothylas as a ''f
enclosing involucre, small unicellular elaters, lack or poor develop and Anthocerotales but reads genetic relationship m ^
ment oi columella stand in the way of efficient dispersal of spores. direction The supports s of this hypothesis consider A otothylas as
a nrimitive genus and Anthoceros as a highly evolved or advanced
germination. Under favourable conditions the spore gemrwhichhas evolved from Notothylas hy the process of progressive
r"i 1 exospore ruptures and the evolution or natural advance.
short the endospore bulge out in the form of a CONCLUSION AND. DISCUSSION
by a transverse or longitudinal wall The Affinities of Anthoceros, The life history
A «//ince)-os deserves special attention k" Shares with the Mosses,
o med "F\"ch'
lormed. ell of the quadrant again^divides bv a veriinl w-^ll
Lach cell certain features in its life eye c^^ the liverworts. In a few
a9SrtL%tXttho7tr- ^rrsh7ir^'%e™otere—^
place by the division of all the lySgIn Lm,
thylas in t"Tmify'TnUmceromccac"
US allies. The basis is the strong resemhr
placc N„tu-
^'^'tlhoceros and
^^Txirfeatures which Anthoceros shares with the
andI others
u favour its bryologistfsuch^
modern inclusi?^^ Proskauer.
between their Reimers
gameto- rperceU (one to hiur, hut larger she and
^PO'-ophyleArrn\i
in several respects of yT?;-^ "'^ Notothylaceac.
that of
"'""'l^ifZIeleoVp'rlX The pyrenoids are charaeterisuc
systematicpositionofiVofofA^ks- to the latest view the °'''®;!«;Sirph,g«endorsiventrally flattened thalins-lihe plant
kingdom ,,0dy and „uh both flagella of whiplash type,
I^ivision
Plantae (itj) Biflag features of Anthoceros which recall
Class Bryophyta The above by some to indicate that Anthoceros
Order Authocerotopsida those of the algae are CO leading from the algae to land
^amity Anthocerotales
Genus Notothylaceae
n,
Species Notothylas
levieri
n allialcrbet^ee: and the dVe.l Myao.
 184 BOTANY FOR DEGREE STUDENTS—PART III: BRYOTHYTA
ANTHOCEROTALES—CONCLUSION AND DISCUSSION 185
(6) Features common with the Liverworts are ;
SYSTEMATIC POSITION OF ANTHOCEROS {Anthocerotales)
thallus-lite gametophyte without any differen- It is still a debatable question. The conventional treatment
smonth-waII»>H^^?- absence of any appendages, except the is to consider the Anthocerotales as one of the three orders ot the
with the th n
with the thaUoseJungermannialesihe such
features which -4/ii/ioccros shares
as Pellia. class Hepaticae. The other two are the Marchantiales and t e
{ii) Similar apical growth of the thallus lobes, Jungermanniales. Many eminent scientists still follow this tajmm
The chief among them are Bower, Leitgeb, Kashyap, Cavers, Goebel,
Maclean and Cook, etc. On the basis of this classification the syste
onthi'h^alfcpfar'''''^'''" constructed matic position of Anthoceros is given below :
Kivgdom Plantae
iu tT*"" similar to those of the Hepaticae. Division Bryophyta
in the'^ila'rcL'nTieear" P"™'- Glass Hepaticae

pericl/n"al wX i'nTe sle way7!n ^epafee'"'^"''''""'" Order

Family
Anthocerotales
Anthocerotaceae
and lfeelo°g?ted""te'rne llh llhTrT
In the related p-frmc M n
T' °
^ develop into pseudo-elaters.
Genus
Species
Anthoceros
himalayensis
Hope, Campbell, Smith and Thakhtajan of a different
opinion. They point to the following fundamental differences m which
1 •! common with the Mosses are : the Anthocerotales differ from the Hepaticae .
system inTe^'c^suIe walL ventilated- assimilatory 1. Presence, as a rule, of a single chloroplast per cell as against
numerous in the Hepaticae. ^
col„j::l,rXh^s1„dre^y"e;d'oth:c^a,
Anthoceros resembles Sphagnum. 'he
ongm. In this respect
2. Nature of chloroplasts each of which contains a pyrenoid.
3. Sex organs deeply sunken in the thallus as against their
superficial nature in the Hepaticae.
capsul'f greatly reduced forming a small part of the 4. Mode of development of sex organs.
• *
5 Mode of development of the sporogonium.
amphithlctum
columclla as in »d"XpeTX"a'd'
the ^oss. ™^ering the top of 'he
the e' Presence of a narrow meristematic zone ^tween ihe foot and
the capsule as against Us complete absence m the Hepaticae
Pteridiphy^es'^XeT^ "* '"'•'ich Anthoceros resembles the 7. Presence ofa thill, slender core of sterile colu-
meUa in the capsule as against its absence in the ^,„„hi
8 Differentiation of the domed archesporium from the amphi
^tttticL^rnTlhefo^plothallu^® structure of the thallus of thecium as against its endothecial nature m the Hepaticae.
9 Presence of amply ventilated photosynthetic system in the
gametophyt'e. "^e pteridophytes are deeply sunk in the capsule wall as against its absence m HeP^^ the tip down-
, SS:Sr.^frir°fX-'--hegonla wards°a.onT-iLluuXaTlines of "dehiscence, which separate the
capsule wall usually into ''Xi'^rrlcarS' -psule with stomata
and XX'uedgrXth'?
rootless. leafless Shi , vetilated
'he nearest afl alj'f'"'''''"■'y system andslmple^o;bShrd"p°seudo.ela.ers without spiral thickenings have
'r2'"Nl'brrf chromosomes which is usually 5 or 6 as against
8 or 9 in the other separate the Anthocerotales
wort s mi^lhriSses
the primitive on Sleon^h^th""^X^i^rUnrbX'''"
pteridophytls !"'* ""ks the t- " Jh'= Liver-
'he Chlorophyceae is''alIo1nZlt°"'"- ^ -mote tntcX^ ^t'h fromtlTcSeSae^"^^
in rank with the latter.

r,
 ISfi BOTANY Pgr DBGREB students—BART m : BRYOPHA'TA AN'moCEROTALES—CONCLUSION AND DISCUSSION 1S7

Leitgeb vehemently opposes this step pointing to the several the other Hepaticae. It is the difference in the basic chromosome
leatures which the Anthocerotales share with the Hepaticae, parti- complex. The Anthocerotales have the basic number as five or six
cularly ihejungermanniales (refer to pages 183-184). He maintains whereas the other Hepaticae (Marchantiales and Jungerraanniales)
that these common features between the two indicate that the have eight or nine. In the Sphacrocarpales it is ten.
Anthocerotales are sufficiently near to the Junaermannialcs. He It follows from the above discussion that the Amhoceroiales
thus advocates the retention of the Anthocerotales in a series with possess certain characteristics in common with the Hepaticae yet
that order opposed to the Marchantiales.
they ctinnot be grouped with the latter on account ol" the fundamen
tal differences enlisted above. Similarly the Anthocerotales possess
HiflV
diflercnces betweenaimches no importance
the Anthocerotales to the
and the above mentioned
Hepaticae He ex certain features which ihcy share with the Mosses particularly
plains them away as follows : ^ sphagnum (refer to psge 186) diftering totally from them, at the same
time, in others. Consequently the growing tendency among the
of chloroplasts per cell varies considerably in the present day bryologists is to consider the Anthocerotales as a separate
a sprnn<? ^ • ^"^og^nous origin of antheridia is group named AiithoccvotiiB or Anthoctvotopsidu in rank to the
LnkeTtrl
sunken archegoma are also found in the
the exogenous origin. asThe
order Hepaticae in other two classes of the bryopiiyies. Since the Anihoccrotae form a
connecting link between the other two classes they are placed In
zone'midrt t Th'-rrtematrc between the Hepaticae or Hepaticopsida on one hand, and the Musci
acUvitrofren through the persistence of meristematic or Bryopsida on the other.
an7thecL3^^^ the foot The consensus of opinion at present favours this view. So
be in responso Tn fh Presence of stomata in the epidermis may according to tlie latest view the systematic position of Anthoceros will
be as under : —
Kingdom Plantae
Anthocero"le?as proLbly a^ecill Division Bryophyta
Class Anthocerotopsida
gn^aMendency of decen.raU.aUonl/rtoU'Ta l': Order Anthocerotales
Anthocerotaceae
Family
Anlhoceros
Genus
'caution sayin<r Species himalayoisis
remains a remark-
a degree ofradial symmetry and uprigtesWr"^'^'','!"''"^ Biological Importance of Anthoceros Sporophyte.
With its long-lived basal meristp^ n •? Anthoceros has a remarkable sporophyte. It is dEtinctiy different
tissue in the capsule wall, thin slender cohim^?/ photosynlhetic fromThe'sporophytes of the other bryophylcs and is considered a
Pseudo-elaters, basipetal mode of deluscencTofffip^"^ l ' an^d an advanced type. The advanced features which also
amphithecial origin of the sunrnalT ^ capsule and the rnTcatervoiutionlry tre'iids or'the probable lines of biological progress
accord Anthocercs a unique statn<f h f features which in Anthoceros sporophyte are :—
^'otofhylas. The gametophvte ^f thp intermediaries like I Amvly ventilated photosynihelic system in the capsule wall.
oroutAnthoceros. The caps^^oweve^^ compact.
horizontally from the thallus lobS
resemble that
It grows Thecapsnle wall consists of several layers of green cells vrith
• i rcellnlar spaces between them. Lxiernal to these is tlie protec-
and the subepidermal tissue in the cansJp ^^cks stomata surface layer forming the epidermis which is cuticulansed. It is
The columella, in some species, is emS;"'4rent chlorophyll. .rtured with stomata similar to those of the higher plants. The
TkSt
^ 1
developm nf beainning
the ventilated photosynlhetic
of physiological tissue of
independence is the fiio.
Anlho-
prium is^nScL^Tror^trn*'"^^ h>r example, the arches- step towaids he beoinii o ^ However it never
ToJsTSeiy inSfe^dent of the parent game.ophyte at any
Decentralisation and oomplele sterilisation of the central
decisive one point-perbaps a fertile tract, the cndotheciam. This evolutionary tendency resulted
tne Anthocerotales and in the following two events :
188 BOTANY FOB DEGREE STUDENTS—FART III: BRYOPHYTA
ANTHOCEBOTALBS-OONCI-nSION AND DISCUSSION 189
{i) Development of a central core of sterile tissue forming the from the actively grovvi^g foot sporophyte
columella. Ii consists of narrow, vertically elongated, conducting
cells with uniformly thickened walls. The presence of a central
columella is suggestive of the demarcation of a region in Anthoceros become I root in the soil was,thus suggested.
sporophyte for the location and the origin of the vascular tissue. The above suggestion, in ^
Although columella never develops any vascular elements even in of the bryologists. of nteridophyles. On the
exceptional cases in the entire group yet some scientists consider
that its central position corresponds to the initial vascular cells of theory known as the of Anthoceros and its allies
basis of this theory
the early iracheophytes. They believe that it plays a definite role was considered to be on the me , ^ the topteridophytes.
the simplest
in conduction and affords mechanical support. For these reasons
it is considered the forerunner of a protostele in some evolutionary and the primitive "^dependent sporopj^,^^ ancestors
concepts. The discussion about the the sporophytic generation
(u) Archesporium becoming amphithecial in origin. The com
like Anthoceros upright. Anlhoceros sporophyte with
only. It was argued that the up ig ^ continued
plete sterilisation of the central endothecium to form a columella
resulted in the office of spore production being shifted from a central
its simply ventilated P^°^®7hidependLt existence. It lacks only
growth is nearly equTPed foi the soil otherwise
to a more superficial position, i.e., the inner layer of the amphithe- a vascular tissue and a root wi primitive, free living
cium. This superficial origin of the generative tissue (archespo it approaches the simplest ^ his contention Campbell
rium) promotes easy dispersal of spores and its position on the phyteofthe pteridophytes support n the sporo-
surface of columella ensures easy and ready supply of nourishment. cited the example of collected in California by Pierce,
3. Differentiation of amphithecial archesporium into alternate phyte of Anthoceros had been growing m unusually
Imnds of fertile and sterile tracts. This evolutionary tendency is
believed by some scientists to have great potentialities. It is consi
dered the first step towards the origin of sporangia and sporophylls.
In fact a noted British botanist Bower made the arrangement of
fertile and sterile tissue into alternate bands in Anthoceros capsule
as the basis lor the origin and evolution of leaves and sporangia of
die pteridophytes. It is called theory of the origin o( the strobilus.
V—d .i.™ d..«.d ..
Ihe separated sterile and fertile masses become more and more twice that in the normal sporop y (jjameter than that in the
superficial on account (m) Columella nearly portion into a conducung
^enle, superficial tractsofturn
theirgreen.
amphithecial origin.
The green tissueEventually the
develops into
membranous expansions, the sporophylls with each of the sporoffen-
ous masses becoming enclosed by a sporangial wall to form sporangia
Ui course all these suggested changes are hypothetical. '
4. Presence of a basal intercalary meristem. It eauins thp
^psule, at least, theoretically with unlimited power of growth thallus tissue. nnmnhvtes of Anthoceros have
The menstem continually adds new cells at the base which gei Campbell opined that these^ ^p^^ pteridophyte
differentiated into the columella, the generative region and the "reached a condition compara jhis theory of Campbell
wall. This unique feature prolongs the after it has established its bryologists. They pointed
received but little support fr^om th ^ non-vascular, depen-
riontuLro^ct 1 '?l"'P^yf=°^^nlhocerosl thus
the other bryophytes. It
Tut"he wide gap ^f hiIZtM vascular, inde-
dent pteridophytes.
distributing spores' gametophyte, producing and pendent sporop y difficult problem. The
Origin of Anthocerotales It forms
foot. %hruprTSt''bodv^h"f' {Oapsule) and large bulbous venetirrflationship of Anthocerolae vvhh^a y
of bryophytes including even' thallus and sporophyte the
remoL' Campbell opm^d tha^^ m So
of these rlnzoidal processes
d^"p1ry°ter'Kash;:; was an ardent supporter of tins view.
 100 BOTANY FOR DEGREE STDDENTS-PART III: BRYOPHYTA
ANTHOOEROTALES-OONOLUSION AND DISCUSSION 101

rSforganised TT'"', Mehra and Handoo feel that • the Anlhocerotales and the
Anlhoceros ereclus tha h!
of the
Eqmselum df-bile and thallus of
the possibility of the origin
Rl.yniaccae have sprung up from the same aneestral stoek in the
nre Devonian They call it the Amho-rhyniaccae. The latter
and Handordo not ubs'cribe to t°h"s '^is Pteridophyte. Mehra

two, '^"'nmon between the sporophytes of the

that migrated to land.
except'bL?anS"bntS i„°[he ™dr" Ja'srAmhS^r'a";"" REVISION QUESTIONS
1. Describe the extornel features and internal strnotnre of the thallns
twois'very'^widt'^and"^'i,uTirre™ "f'he '"'""Tffivean illnstrated acoonnt of the strnetnro and development of
the basic number as five (5) whereas in S. debile it the sex orcans of Antbocfros. r a ,i.
3 Give a concise account of the life history of^niAoceroff.
A Mndn-ceros is a ayathetlc type." Justify the truth or falsrty of
phylogenet?c''"rela^tbnship"''be°twS °the"'^AnthMc''"f'''
Psilophytales (Pteridophvtes) Thp and the
this 3tatem»^.^^^ aporogoninm
oldest and most primUive of all knLn v! f i. Give an account of tho strnotnre and development of the sporophyts
were the fossils o"? the Devonian al S
these were Rhj/nia and IlomeonJwInn '-ru important of
of Anthocerotales.

Rhyniaceae of the order Psilophytales tL^R tf family 8 Discuss the afflnitios of forms lilwXniAocero^.
herbaceous sporophytes about 2 feet in iTcH.^ were small. 9. With annotated sketches trace the life history of Antlmeros^ ^^
Thea simplest
of rhizome of them slender,
bearing were even
erectleafless ^?They were rootless,
dichotom 10. Discuss the origin of Anthocorotae. . from the
The rhizome was Anchored to tim subs°mmm^ 11. How would you justify the separation o
vascular cylinder was of the nature Z rhizoids. The ""'""''r In what respects does the eporogonlnm of An,h«c»ns differ from
secondary growth. The reDrodurtive There was no Biccfnor ,.o,p of a labelled sketch the structure of the
which resembled the spoiophyfe Tthe
borne singly and terminally Each en They were cporophyie of No,c,%lns t
-p.- ..«.»»i

ancient
by group
descent (Rhyniaceae)
in an that he uLcs^ed oSimties
upward direction sp.orangia ofthem
between this
out that Anthoceros sporophytT" not viewpoint
vascular tissue. It exhibits no bLnh" ''n/
meristem m is at the base crown all the
•pi . , the apex.
^ene/w rda! thought. Thev read
the Rhyniaceae by

A»'bt,sfV™
 193
BRYOPSIDA (MUSCI OR MOSSES)

the protonema may act as a vegetative / "ro

Excepting a few in most of these methods
tonema) regularly heralds the formation of the leafy moss
CHAPTER X
Only a few important examples of these methods are describe
below '
BRYOPSIDA {MUSCI OR MOSSES) 1 Pwnressive Growth and Death. Some genera of mosses have
a creeping main stem. It bears upright branches. Jh'S older portmns
14.500^SDtcTeT'wv'"S^''^ placed under
bryophytes. They number about of the creeping stem which are more or less cut off from bgh' and
is differentiaf(-rl • 660 genera. The plant body air, die and decay. This leads eventuaUy to 'h= sepa ation of the
to calUhe^ leaves. Koch (1956) prefers uprigiit brancties as separate entities. The result is the rapid
leafy plant boHv cauloid and moss leaves phylloids. The crease in the number of leafy individuals.
of the liverwnrK adapted for a life on land than the thallus 2 Bmnchina of the leafy stem. In many mosses the leafy
species ffrow?n H- ^^^^n are gregarious in habit. Different axifinally
s clevelf; budfa/its base.''The buds grow into bran^ which
get detached by the decay ol their basal conneetmg pa
rocks, fallen trees'"'bark of
ing water or aHheh^ ofc."? actually submerged in flow- Each detached branch develops into an independent plant.
rity grow in damn lakes. Many are swamp growers. Majo-
are fble to end3 H S""' extensive mats. Some, however, 3 Fonnalion of stolons. Some mosses develop stoloniferous
tly looland
dVresume
or growth curl up and apparen: branches from the base of the stem. f the surkce of
uncurl
is the gametophvfe tr ^ predominant stage in the life cycle
water, dp^S^tr-grs -^ard as an upright,
growth stages generally consists of the following two leafy axis.

frequently fila°mem"s^ ^In^rar,- ^L'! branched and but4 fS:

ralprotonemaPreliminary Protonemal Stage. Moss plants arise as late-
ext'ensifromve^ abmn^
originating g P P-o™ ^he
jg accompanied
directly from a spore The rhi#»f Plate-like. It develops
tive. It bears no^rx organs ' buds. The latter develop mto leafy shoots This
by the decay of the shoots as separate individuals,
resultsIS in
in the
IIIC separation
acpaic4civ/*- ol the Jeaiy r
bearing\ptaUy^
organs. It i.s thea^rfged
so-cafled Ve i "P^gbt, slender axis
finally produces sex 5. Mmiplication of the
protonema by death and decay o
— T Th#» nrimf

Each segment grows

tophores o£ Marchaiitia comparable to the game-
tonema Many morplants mri •''' the%ro- Pto short, detachable segtnen '^Xe7a fresh c'rop of leafy
from a single spore. ^ protonema derived into a protonema. The lattei P protonema stage thus
/ '! pss «, sjr - -i.. ~
Thismaks^triea?ygameS disappears. """s,
sentattveofthegametophyte
persists as long as the o-ampf.-.stu'V
*' i ^ and the
niosses the sole repre-
protonema
turn brown and contribute to th^ (Phascaceae). It may haveinjured
or a greatportion of almost
^""aa 'y part
p of the moss plantsurrounded
(such as stem,
by
heJeafy ?ametophole'^:;'; X\nr:„°dLl'".f 'S kafor protonema) under suitable con^ ^alga-like filament. It
-«JS—■ •«.( P«S"a • rtxr; moist
is calledair)thedevelops into proto^
secondary a ^ crop of moss plants will
Ac^^P^^^
spring from the secondarx^ .P™ by the decay
^ecay of the
the, connecting
grow into independent .ndividuals^J^ ,^3
threads.the latter
when Secondaiy
grow prmmligh ^
hgl t a^
an<l^^^,_ surrounded by moistand
important air. more
The
formation of propagation than the original proto-
effective method of vegetative propag
nema.
192
 195
BRYOPSIDA (MUSCI OR MOSSES)
194 BOTANY FOE DEGREE STUDENTS—PART 111: BRYOPHYTA
Formerly they were called the bulbils. The tubers develop
7. Tubers. The formation of small, underground resting, sint^ly on the protonema or the rhizoids as small, spnencai
bud-like structures called the tubers has also been reported. storage organs containing starch. They serve as means
nation and enable the plant to tide over periods unfavi^rable
for vegetative growth. They are found in Funana and Trema^
GSMM/FeROUS
GEMMIFEROUS
toden. On germination in the growing season each tuber produces
CUP PSEUDOPODlUn a protonema. It bears the buds which grow up into a npv crop
of leafy aerial shoots. Bryum erythrocarpum beap spherical tubers
on the rhizoid system. They are bright red in colour when mature.
Leptobryum pyriforme is another example. It bears ovoid tubers.
8. Gemmae. Frequently small, green, ov^, multicellular buds
BODY
are produced on short stalks in many mosses. ^ They are the gem
mae. The gemmae are produced in groups m the axils of leaves
in Webra prolifera In Teiraphis pdluczda are long stalked and
lens shaped. These gemmae are produced m a pecuhai, cup-hke
structure called the gemmiferous receptacle. The latter is
Y STALK
cStuLd by the enlarled spreading leaves at the summit of the erec
?hoot The 4mmae arl easily detachable. On falling on a moist soil
STALK any cell of the gemma may produce a green protonema. The gemmae
In AuLoriiniJiandrogynum are borne in a terminal cluster or gemi-
ferous head on a leaHess stalk or pseudopodmm arising from he tip of
the leafy
tlic shoot. Another interesting
leaiy miu produced example is that
in a cluster of Ulota
at the tip of the leaf.
OrTmmt
S gemmabear! i!gemmae
a mass on
of the leaves atcells.
embryonic the tipsIt orinvariably
along theproduces
midrib
a protonemal stage. The latter bears the new leafy plants.
mw
q Persistent Apices. In the
9. somedry
mosses withexcept
season the creeping stems
the growing
the entire plant secrete a mucilage sheath
Tound them and persist. With the return of favourable conditions
fhe surviving apices resume growth.
10
lU. Avospory.
Apudjjuij. The mosses --11
havenP anthe extraordinary powerthe
sporophyte besides of
regeneration. Any un ^ ^ protonema. The wounding of
gametophyte, . can grow various parts of the sporophyte, m-
MATORE
the unspecial^ed cells
rj'JLBlL
duces the pio uc
bears a iiew crop o
gametophytes. The formation of the
of the sporophyte other than
gametophytes diiecuy phenomenon of apospory cuts out
a spore IS called ap p y ^^ words the moss plant is
the spore stage m g rophyte without the intervention of
A. Tetraphis pollucida with a terminal trflm -e ■*
produced^ direc y ,oduced moss plants have a diploid
B. Gemma with a stalk. gemmiferous cup. > spores. The P \ood to be larger but otherwise normal
C. Aulacoranium androgynum win, « , * chromosome gametes. Fusion between
m a gemmiforous head. psoudopodium ending
in appearance, iney ^ diploid and a normal gamete
CLVV«.»..ll " —-I - , V
B. A stalked gemma of the same. two dlplmd gametes ^ (having
i«ults_iu a tetraploid
^ four cofc
sets r>F
of rVimmnun
chromosomes) or
A "mature rhizoida. triploid sporop y sex organs containing the
clusters. Generally the cluster is developed
^luster^of gemir^ P^^rt of the leaf with an apical
• A single gemma.
gametes are borne m
r"

196 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

at the apex of the main stem or the branches. Intermixed with BRYOPSIDA (MUSCI OR MOSSES) 197

arJ " the cluster are the sterile green filaments. They The lateral branches are never axillary. Each arises from below a
j* _ • mosses are monoecious, others leaf.
fhe monnP.;n distribution of the sex organs 5. The gametophore usually grows by means of a single
1 ^he following fhree apical cell which is pyramidal in shape.
are the two kinds of sex organs 6. With the exception of ASjj/iagniwm the moss leaf usually has
and archeffonial a i" The anthcridial a midrib.

perichaetili demarcated froi; one another by 7. The cell walls are reported to consist mainly ofhemicel-
luloses and penlosanes. Cellulose is rarely found.
are borne 0^3^"°"^^ rf the^am^pC. 8. The rhizoids which anchor the moss plant to the sub
stratum are multicellular and branched. The septa between
occur bthe°a°me°hear°f'''- ^"'heridia and archegonia the cells are oblique.
belo^ the LHrins
tinct sheath each other. The leaves
or a rosette called thlp°ericLTtium*"
9. The sex organs are stalked and the stalks are longer than
those of other bryophytes.
10. Early growth of the sex organs is by means of an apical
shapeltt'u^
single Tk ^'heridium
layere7aatheridJ»^ .. is an orange coloured
length. club-
The cell.

archegonk are essemta t '="<='°^'=^bmagellate sperms. The 11. The venter of the archegonium is much more massive
than in the Liverworts.
fertilized eeff or tlif- Y similar to those of the liverworts. The
nium. Ordinarily onl^^a^c* ^ iT^to a. sporophyte or sporogo- 12. The sporophyte is more elaborate and complex and shows
cluster. single sporophyte develops in each high degree of specialization and sterilisation.
13. With the exception of Sphagnum early growth of the
capsule It consists of a foot, a seta and a sporophyte is by means of an apical cell.
Liverworts It rf u ^"d complex than that of the 14. The entire archesporium differentiates into spores.
great amount of stert'tisLe ^^ specialization and has a There are no elaters. ^
possesSe two"(?n "tosses in general differ from the liverworts in 15. The capsule usually dehisces by the separation of a hd.
?he JuvLn^ gametophyte. Classification. Smith divides the class Bryopsida or Musci
stagl or'le^fy into three groups giving each the status of a sub-class. These are
cal. The rhizoids are multicellJlar and R symmetri- the (i) Sphagnobrya,(n) Andreaeobrya and (zti) Eubrya.
more elaborate and has a more com^tx ^he capsule is The older bryologists such as Bower, Campbell,
and spore dispersal. The elaters are^bsTnt dehiscence others divide tl.e class into three groups but they gtve each of these
groups the rank of an order as follows :
1t. Th DISTINCTIVE FEATURES
rnoss plant is radinll^r r. (e) Sphagnales.
into stem and leaves. SYmmetrical and differentiated (n) Andreales.
ISdifferentiation Lto^thfcortex a'nd th ^ of Trere
tissue (ni) Bryales.
however no true vascular ^system.'^^ The modern bryologists recognise five sub-classes as follows .
1. Sub-class Sphagnidae comprising a single order spka,mles
Stag^. W.th one Andreaeldae. It includes one order Andreales
liverworts. '''= conspicuous in thefmoLf"an°^n7e
'r
with a single family.
tomously branched^*^ T^h^ gametophore
ncned. The branching k{moss
Z nl not dicho- 3 Lb.class Bryidae. It comprises twelve orders^
g IS invariably monopodial. 4. Sub-class Buxbaumidae with a single order Buxbaummles
with two families.
198 BOTANY FOR DEGREE STUDENTS—PART IH: BRYOPHYTA

5. Sub-class Polytrichidae comprising two orders {Polytrichales

and Dawsoniales) each with a single family.
Some bryologists including Parihar divide the class Bryopsida
into the following three sub-classes :
Sub-class I. Sphagnidae. CHAPTER XI
Sub-class II. Andreaeidae. SPHAGNIDAE-SPHAGNALES: SPHAGNUM
Sub-class III. Bryidae equivalent to Bryales of the older
bryologists and Eubrya of Smith, General Characteristics ;

text.
This system has been followed in the present edition of this
They The sub-class Sphagnidae includes the peat or the
growL etfnsive masses on boggy and peaty ^<"1' mosses.
also
as submerged aquaties in peaty pools. The sub-ckss sphagmdae
REVISION QUESTIONS is characterised by the following distinctive features . ,
1. Twist the distinctive features of the class Bryopsida (Muaci or the
Mosses).
1. A simple, flat, plate-like thallose protonema fixed
substratum by numerous rhizoids.
2. Give a concise account of the various vegetative methods of multi
plication met with in the Mosses. 2. The rhizoids are multicellular. The septa between the
3. What is apospory ? Does this phenomenon throw any light on the cells are obi q upright, leafy branch from a single
nature of the two generations in the life cycle of a moss plant ?
4. In what respects do the Mosses differ from the other Bryophytes ? . .1 rpll It grows by means of a three-sided apical cell
5. Write an essay on vegetative reproduction in the Bryophytes. StTtheTdurtt matu?e plant, also called the gametophore.
4. Usually a single gametophore develops from one p
tonema. f.r. i _r the adult bos moss.
5. The unique structure of the leaf of the

6 The absence ofa midrib in the leaves.

8 The antheridia occur singly and are axillary

. ®UrfXS:ny"atrr;ceforsho^
from ?mong a crown at into an enlarged
f ot'°a seta and a large, rounded
^ ^ by calyptra.
■ ^1;: presence of a leafless, stalk-like pse«dopod.mn
carrying the ripe from the entire endo-
13. The development
The deripment.of the sporogenous tissue as an inner
layer from the amphithecium.
199
 J
200 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
•t
aPHAGNIDAE—SPHAGNALES : SPHAGNUM 201

D
shaped'spJre^ac!^^'^^'' columella capfjed by the domc-
a disc-shaped lid or operculum at"^'P^
its top.
capsule by the separation of
■5P0RQ(50NIA
17. The absence of peristome teeth.
The latter ^ single order, Sphagnales.
ted by a sino-le^enm^ 9^7^ family Sphagnaceae which is represen-
Srjstemaiic Position :
I divergent BRAf^CHEE
Bryophyta BRANCH WHORLS
Bryopsida or Musci ^OROOPING BRANCHES
Sphagnidae
Sphagnales
Sphagnaceae

SPHAGNUM (Dill). Hedwig

these !!n%*ruhl
ted from India.' All love
includes more than 336 species. Of
(Sharma, 1949) have been repor-
and also as submerged places as scmi-aquaiics
marshesmatted
closely of higher latitudes
together forminetuJ^7- ^est in colclplants
^P^agnum bogsgrow
and
complete vegetative cover over masses. The latter form a
dangLo"^ converting them to m t pools,
lime tL the quaking W # / It is
environment^^°sT^ increasc^^X^^"^ • r
ionic
tufts isabsorption
usuallv^" T ^^^"buted
reported H tn?u^K process of selective Fig. IM.
bog water
^'cncy of isoxyg
ant^s^ptir
7 the stir^r J"
^o its germicid^
wood and even huml V ^ Preserva^ m P'^P^^ties and defi-
quaking bogs
g have hf>" entrapped
recovered bp
wVnrpr'"'^ animals,
surface inlogstheof
ooccurs ^®*''ibution crved after centuries,
to year as partially decompose.^materialjte
or the lake. Thpe deposits may a heathers, cotton grass,
in all naric r has a
ar extend® T'"'"' ''^ent
arctic and the subantarcl; th^A
the ten^n distribution.
It thrivesIt :rdecay
rxT-'S ssf
of the -cumulated j^amfi- Jlowly cornpres-
fLt flowing
fl°"-'''=^"dripXT''T™^- deposits of dead, partially . | 7f fresh deposits above it. These
streams ® and onIntheTnd"".'^
^"""^tiurs intothetheHima-
sub-
sed and hardened by die w g hardened dead plant deposits
Habit Ti. „ 'of furrows cut by the compacted, partially jg a brown or dark-coloured spongy
11]) with v plant are called the peat The pea^ dried ,t
species are of bright®*", Wee "^tn erect perennial
b^wn, rose.pinh such or yelfot s"me
peat .moss.
orajii ^ebellum), orange
b 'pink, etc. Sphagnum is
 203
202 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA SPHAGNIDAE-SPHAGNALES:SPHAGNUM
The life cycle of includes two distinct, alternating 1 . /TTV 11 -91 It is also caUed the gameto-
pnases or generations, the gametophyle and the sporophyte.

divergent
branches

yci///6
lEAFY SHOOT
'(GAMETOPHORE)
drooping
branches

■PROTONEMA

RHIZOIDS

Fie. 11-3. Sphagnum 0p.

Fig. 11'2. Sphagnum Bp.
Old, protonema bearing the yoimg,
leafy gametophoro,
iess rhizoids at US ba.. _ ^j-hizoids soon%hrdisappear-
hair-like processes. c^eUs are obU.u.^Aj
m
GAMETOPHYTE PHASE
•'» C.bVt
Leafy Gametophyte. gametophyte is a
*"le siage
stage and the leafy gametophyte
and consistsstage.
of two stages, the juve-
11-2) and*is for^pH is also called the protonema (Fig. perenni(olal plant
External
{Fig^ H an upright stem. Individually
ftems'' 'The sTem often grows
the protonema is a ^l7 spore. When young
and cell division thp
It is an irregularlv Inlf
^ ^ew cells. By further growth
becomes a fiat, green plate of cells,
Khe
to a considerab
VS %'oSeapex
branching is usualW^^^
^temof
ness. It is fixed th ^ structure one cell in thick- branches freely. rlothed with small Thev are of
thalius-like protonema^ substratum by multicellular rhizoids. The ^Fdthe
the Stem IS a p,^ - terminal
set bTSes pro-
compact head of the^
protonema of the leaf ° moss recalls the thallose
, the relationshio nf and thus points out limited growth. ^ ^ °ft
margin of the fob/d
'obed nrnT"™ a livcnvort
the erect,ancestor. From the
protonema arises leafy gametophyte ;c.'Vr«nS
 204 BOTANY FOE DE8REE STODENTS-PART III : BRYOPHYTA SPHAGNIDAE-SPHAGNALES : SPHAGNUM 205

mature stems they are thus dead, empty, colourless and large
ches. "th:; octVSlrro'rsddes'of T\o in size. In many species {S. palustre) the cortical cells deve op
large oval pores on their walls and sometimes spiral thickenings also.
The cortex of older stems consequently becomes spongy or po^us in
nature. The cortical cells store water. Like velamen in the Orchid
ters are of the following two types : ^'^"ches m these clus- roots, the cortex of the stem absorbs water by capillary action. 1
As they grow from the^n^?<fth usually short and stout, thus compensates for the absence of rhizoids in the adult gameto-
phyte plant. , . .
position and sometimes slightly upwarT a horizontal The cortex of the side branches remains one cell hick^
f The cortical cells of the side branches m ®
They are long,
around the main stem. Th^e o"endtTU "n!"
as water conductors In the'^^ f
{S- obesun.) the drooping branches in
decurrent branches act
t Sphagyuim
grp^o^p-aTthem^tTeJS^^
all of the divergent iypf -^lu^er are rare. They are
(i) Anatomy microscopic animals.
The hadrome lies next to the cortex. It surrounds the medulla
...^2£Z& and consists of elongated thick-walled, prosenchymatous cells. The
hadrome functions as a supporting tissue.
The metlulla forms the core of the stem. I'
.yHYALODERMIS
(CORTEX) sote"what e^ot-S^Vte meSa reform: a dssue corresponding,
MEDULLA
■HADRQME

It'

■RETORT CELL 3

ft*- branches

Fig. 11-4 (A—C). Sphagnum sp.

A r . <^roP8.section of atom.
.B.- s:::::::: irorr STEM

• a branch of S.n>„l,usc„m with retort oetis.

inwards they are tK^ Cavers)
^nd the medulla. ''yalodermis, the hadrome
Th

is^nn?"^' in thickness outermost region

-si- of i?v 11.5 fA-B). Sphagnum sp.
pactlv (Fie j , branches it
layerf In Sder Lrl" V A Portion of an oWer part of the gametophyte.
lose, their 11-4 in I cortexl^bebecomes B.' A single leaf as seen under low powe .
P'"otoplasmic contents and increase
cortical3 cells
in size.
to 6
In the
 9mm
'•rl

200 BOTANY FOE. DEOEEB STUDENTS—PAET III : BEYOPHYTA SPHAGNXDAE—SPHAQNALB5:SPHAGNUM 207

of the higher plants and functions as a storage strengthening bands help the empty dead cells
pores enable the leaf to absorb water readily and retain ^uch like
a sponge. Because of this property the hyaline cells are also called
the leaves occur on the main stem as well as the capillary cells.
laoDine On tlP" branches they are closely set and thus over- The assimilatory cells (Fig. 11-6 A) are alive and
Brs?dpf't>i ? ^ ^ little apart (Fig. 11*5 A). tic in function. They contain chloioplas s an hyaline
structure from7l?n.r ^^^ails of cell colour Thev are very narrow and lie between the
thin and scale like Tl, branches. In general the leaves are small, eel Is forming a network. The meshes are occupied byjhe dead
stem The midriii ' sessde and are arranged spirally on the hyaline cells—one dead cell per mesh. Owing . ^^ the
acme S^uZtm h.^ '-'"S- entire and the apex capillary cells the Sphagnum leaves play an important role in tn
much elnno-awi Tu ttO Bj. 1 he cells constituting it arc absorption and retention of water.
photosynthetic or tS
alternate with each other
hyaline cells and the
two kinds of cells /T7:«.
The c-oss-seCion of a -tare leaf has a bead-like appe^a^^ance
The bead consists of two kinds oi ce • |,_
under the microscoDe tlip n;, ^ net-like pattern. When viewed the iarge capillary cells and the assimilatory
to form the network assimilatory cells are seen The two kinds of cells according to the species,
themeThes large and wide capillary cells occupying ceils are variable in forrn and P° , bases directed
In S.acutifolmm the ^ the apices towards the
and tSs' are dead and somewhat rhomboidalprotoplasmic contents
in shape. They are towards the upper surface of the J;, surface of the
lower surface. "They are Jj"® j, towards the lower
^chlorophyll
COA/TAIHWS CELL
'^1ace'''Tt®feav:fof's.
surface.
kinds of celIhe have Issimllatory'
r^nillarv
ls reveled, of tfLguC tSe^posi tion celoftowards
cells bulge
lsthese
face two
the
the
HYALINE CELL iTwe" surface." S. ssTOrrosiim has fusiform neither
are hemmed inbetween the capillary cells. Their ends
the upper nor the lower surface of the leat.
■SPIRAL THICKENING A« ,• R^-rftn-tion and Conduction
Physiology of Water Absorption. Retention a
PORE The adult Sphagnum plant lacks rhizoids. Their function o
water absorption is taken up by : - ^
The snoney cortex oi the stem aim
iu.... ■"•''■■"w'?.!"''"-.':.
HYAUNBCELL CQNTAm^CELL '"S 1—'
are filled with water.
11 r iintl-i tlip stem and the leaves account foi
The capillary cells , ,.rater retaining capacity possessed
the remarkable
hy Sphagnum. It
water absoi ^ S ^jr^otly through its leaves and
^ .r.Ugse cells. The latter as well hold a
stem by the ^ap^^ary action elements of the
Fig. ll.6(A-B). Sphagnum sp. large amou^nt ^ ^ capillary apparatus for raising the
A. Surface view of cells from a mature leaf, the plant and to other places where needed.
The same as seen in cross-section.
colourless, wide and fi ll ^ • i.
1 • u 'Iac".
In species which .k jhe
the Dorous
P .^ater elements in the cortex
is facilitated by theof
pr oval pores on either tu They frequently have circular the stem the upward and around the mam stem
inner surface
inner surface of the wllk ^
r.f+u... - * T"-
lower surface^ - — walls.
of their wans TheThe
ed by spiral or ring-shanpdI .S^.^V^bne
!.ickening cells
byalinebands is frequently
cells is frequently stren;
strengthen'
hen-
® shaped thickening bands nfof wall
wall material
mmerial .' Th#
The
 SPHA6NIDAB-SPHA6NALES,SPHAGNUM 209

208 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA

the thick, loose covering of the pendent branches serve as a capillary young leaves grow into new i,uniid conditions and
apparatus. They draw up water by capillary action. Ld shortlight
moderate brinches
functionunder exceptionally
as vegetanvereprcducuve
Apical Growth. The stem grows by means of a three sided Sp/MsriMTO has great power growth and
apical cell. It has three cutting faces. Each segment, cut off
parallel to its flat face, divides by a periclinal wall into an cuter cell specimens vthen put under normal condmons
and inner cell. The outer cell undergoes divisions to form the stem become green. , multiplication of
cortex and a single leaf. The inner cell gives rise to the central Vegetative reproduction also thallose, primary
tissue of the stem. In this way each segment derived from the apical the protonemal.stage. By division and growth
cell forms a leaf and the subtending portion of the stem. This
explains the three ranked arrangement in the younger portions of
the stem. This arrangement is, however, soon replaced by more Kr:"rJen"Slar^»^^ plfo-
complex phyllotaxy. rO? -&e erect lea. Same-
The leaf when young grows by means of an apical cell. It tophore. ^ _ .,r.rmaliv reproduces
has two cutting faces. Segments are cut off alternately right and 2. Sexual Reproduction. They are formed
left parallel to its flat faces. This results in a young leaf consisting by the formation ofantherrd.a and archeg
of a single layer of ceils. It is made up of only one kind ofcelJs.
They are all green and contain protoplasm and chloroplasts. Later M4lf branch
the apical growth ceases. The further growth of the leaf is entirely
basal. At this stage striking differentiation of leaf cells takes
place. Certain of the leaf cells increase in size. They lose their
protoplasmic contents and chloroplasts to become hyaline and empty. female
These empty cells develop entrance pores on their walls and spiral branch
thickenings on the inner surface of the walls only.
REPRODUCTION
ANTHERIDIUM
S'phagmrn gameiophyte reproduces by two methods usual for
all the bryophytes. They are vegetative and sexual reproduction.
Following fertilisation is produced the non-sexual individual which
is the sporogonium. The latter produces the meiospores.
1. Vegetative Reproduction. Sphagnum lacks special
structures for vegetative propagation. The stem progressively dis
integrates from the basal, older parts upwards. Consequently the
adjacent branches become separated and develop into independent
plants. ^
A-
Occasionally one of the branches in the axillary cluster may Portion <>f the
leaves and nn ant (Enlarged),
slightly modified
turn upward. It grows more vigorously than the others and conti
nues Its upward growth. This long, upright branch takes on all ■oi short, densely leafy terminal
SoMPrnrTf u ■ called an innovation,
stem hvn.^ j become separated from the main
a'"'*'' °'derpart. The detached
proDrsatim hv rt'' independent individual. Vegetative
Sn rnZi,,™' very effective means ofiSultipli-
extensivemassr " ^ responsible for its occurrence in
sex organs neve ocm.^
"■:xA
a.e
I,—
S.paiMsfrrrlproducesTCgrtlTivel!^^^
even plates of cells from the young
yom/vrr"" methods.
g^metophyte of
Buds and
ches
physes are alway
absent.
growing apices of stems and
 US3!

40 BOTANY FOR DEGREE STUDENTS-PART HI: BRYOPHYTA SPHAGNIDAB—SPHAGNALBS: SPHAGNUM 211

usualir'sht^p''T'f"' The antheridial branches are by anticlinal walls to form the single layered wall of the antheridium
are spindirshir,!^H 1°'"'"' S"" vegetative branches. They (Fig. 11-8 I). The primary androgonial cells undergo repeated divi
pigmented and often densdy c oth"d wklf'!^^^ "''l APICAL CELL
yellow leaves generally stnaLr^tt'nlhe'foUagflea'tes''''^' " antheridial
INITIAL

Antheridia

take the poritlorof axiSv branches (Fig. 11-7 B). ^fhey

The mature antheridium (Fie l l -R if nte homologous to them,
elevated on a long, sffn^er stalk Tl f'f SlobuUr body
the body of the® anlteridimn It^ """n '°"g SF£RM MOTHER
CELLS
consists of 2-4 vertical rows of cells ThfT^^ r ? JACH£TC£ILS,
antheridial
has a jacket
wall. It is layer of sterile
one cell cells .Tr .^"^"dyasof the
thick fnf the antheridial
antheridium PRIMARY-
androgonial
The androcytes metamorohosn ^ ®ndrocytes. CELLS

11'S K)is a/ elonTated ?phanv f t'?"'"" ViS-

tvvoflagella. The number of coils is ^tp TTh^fl
ed at the anterior end which is rvtnni are insert-
STALK

flagellophore. The greater Dart orrr^'V" H

lear in origin. To its posterior end
ture consisting of a cytoplasmic matdx ^ vesicle-like struc- SPERMATOZOIDS

(n) Dehiscence (Fig ll-R t\ i

matureantheridiumdehisces
of irregular lobes. The lobesatturn
its Ww' t1 the
^ number
cyte mass. With the dissolution of ejecting the andro-
liberated. TJiey swim about active v in STALK

'^npture. actively m the water that causes the

frntvt P^^^lopment (Fig ii-n^ p ,
S^roVih^ "lUfthe stem ^
rpi^i.D A ^^dlary bud. It grow.
Tt develops
modified initial
the ha
base (Fig.
/p'' 11-8 B). TheisnmtruP-
papilla then cut ^7 ^ transverse
P^Pd^a-Uke wall at
outgrowth
fife" initial (Fig| g fif cell now functions
10
;i. 9 - The terminal ceIlofth« * li.ri"
h- of few number of
cells (Fig. Fie 11-8(A-J)- Sphagnum sp.
■ the
Al l a^vfllooment
A-H. Stages showing d© P j. of the antheridium.
young antheridium.
altfrnatelv r-
wSn th^
cutting faces^ "nctions as an apical cell
paralld to are cut off
last 2 antheridium k \2lTrT- 11-8 F). I.
Dehisced antheridium. „i• \
successivp V derived from the ? oP die J.
A liberated,sperm.
^ ( / T and K, after Schiroper). r-cW^A the
mScal tT (Fig. n^Gi undergoes two K.
(Fig 1 8 F 'V' iStiaSomr, sions.
r Ai ,. last cell generation are called the
The cells ol the last g
aodrocytes. of the row which do not take part
Meanwhile the lowe antheridium undergo
i„ the foiniat^n « body^of^the
a.so - ^e'Tnrid^- They divide by vertical
antheridium.
Ihe jacket initials divide
 212 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
SPHAGNIDAE—SPHAGNALES : SPHAGNUM 213

(/>) Archegonial Branches (Fig. 11-7 A). The female bran

ches are very short. They are green and bud-like in appearance. In During further development the terminal cell of the filamerit
many species they occur just below the apex of the stem within the increases in size. It then divides by three oblique vertical walls
terminal branch cluster. They bear leaves larger than the normal.
These leaves are rich in chloroplasts and have fewer fibres in the PRIMARY ARCHBOQMUM
PRIMARY
AXIAL CELL
hyaline cells. The archegonial branches are thus more deeply pig-
mented than the vegetative branches in the cluster. The leaves
dimmish m size towards the apex where a number of them form the
i secondary ARCNEGON^^^ \P£PlPlh
neral
]CELLS

pericnaetium. The perichaetium surrounds and protects the

archegonia. The archegonial branches are more or less globular.
Archegonia PRIMARY COt/ER CELL COYE^CELLS
'^^FRIMARY
CENTRAL l^NECK
terminal archegonia are CELL
^p^CANAi CELL
• 11 Specialized archegonial branches. They usually PRIMARY
gonia in CENTRAL
CANAL CELL

NECK CELLS
from the last segments cut oF iL o!® . u ^ are developed
archegonia
mum.
are thus formed about ^hc basroflhe nprimary
' archego-
%^VENTRAL CANAL
CELL

^^i^EGC OROOSRREREr.^
11-9 ir^ItTTalket'^l^ntaik iff
archegonium consists of a long, twisted neck and
^ ^^™cturc (Fig.
The neck consists of six vertili rffof "eek ceUs'fhf^ T""'! H
contains numerous neck canal cell«5 TKr^ \ The neck canal ■archegonium
portion of the neck is 9 tn r u • , the lower
cavity contains a small ovoid egg Tt iVabomf"'' """T"' Hltea /[■?,
leaves
ventral canal ceU which lies above it

worts). The apical cell of the female branrh

an archegonial initial (Fit/ ]l-9Al Jtf '^Jp'^ly/unctions as
gonium of the cluster and is'^rennrtpli ^^rms the primary arche-
transverse divisions. In some cases ^^^ergo a succession of
of the archegonium with two cuttincr apical cell
irregularity in the early segmentation some Fig. H-9(A-I). Sphagnum sp.
short filament ofcells is formed (Fig l either case a ^ (j_ Stages showing the development o rc legonm
TT Mature archegonium.
T V S, through the tip of female branch
isarchegonium
broughttrominitials,
bftfTh®e
segfenf of '-"-chegonia offunctionbg
nfal of ?1
the clusteras showing a cluster of archegonia.
ifo In tlie seoaration of three peripheral
(Fig. 1I-9C). Sis They surround a single central cell
cells, the jacket imttal
a«a . inverted
Ihe^^gonmmin both oas..thfrn^atlon fHlf~ff called the primary
pyramid. The
|,y a transverse
(Fig. 11-90). The loriner is
wall
P[.o an outer trnd an |,„„al cell. The cover
called the cover initial
 214 BOTANY FOR DEGREE STUDENTS—PART III: BRYQPHYTA SPHAGNIDAE-SPHAGNALES :SPHAGNUM 215

walls to form a group of cells.

nnrHnr, directions form the terminal
Lk^MnV- 1 the archegonium. The three
lower nnr/-^ u "meantime divide and redivide to form the
(wall^^/fif"® neck and the venter wall. The jacket portion QOSPORt
the iark/»t i ^.5t"cheg^ium is thus formed from the cover initial and
the venter n',4^ o/ the basal portion of the neck and
2 to 3 cell layers^"! divisions so that these portions become
cell m the meantime has divided transversely to
ventral canal cell and the lower primary
"setoarowoff'tn
cell divides tr^nc ?q primary
. canalThe
cellprimary
divides toventral
give
a result an iir> The division is nearly symmetrical. As
cell ^Fi? 11.qpT is cut off from the lower egg
venter cavitv nearly of the same size and lie in the
produced mentioned above the sex organs are ■SPOROGENO'JS
iarge numbers "n sporogonia are rarely found in ■AMPWTHECIUM layer
occurrence ti • • ^°"^'"ary they are regarded as of rare M^EmTHECIUM -^CAPStSLE
infrequencv r,/r sporophytes may be the result of WALL
high or ton u, '^Jdlisation. The water level may be either too icOLUMELLA
uncertainty of ferrr ^P^^rns to reach the archcgonia. The
effective L?hnH ^r ^°"
Sphagnum.
in the 1 V
propagation by innovations in
occurs m the same manner as
exceot thp ^^ial row of cells in the archegonium,
The releac^^rf^' disintegrates to form a passage way for the sperms.
them find swim to the archegonium in water. Some of
One of them "^uk canal and swim down to the egg.
pioneer structure of
TZthe sporophytic phase. ^'P^oid zygote. It is the
sporoph^te phase FOOT
ANNULUS
formadon^ofThViyJL^^^^ Zu^ sporophym phase starts with the SPORE SAC
secretes a wall around it ^he zygote enlarges and SPORES

division is'^eSy s^mSfcal" ^ transverse wall. The COLUMELLA f./C^

of the basal cell var'y. According m the segmentation
irregular divisions to form the fnr»+ Waldner it undergoes a few SPORES
as a boring organ. However it is el^denl fhlt^th" SpoKl mother sPOf
irregularity in the „rtt is evident t
evident that there is some '-f 'T° M
difficult to f^low! the basal cell. Hence its fate is
r- IMO (A—M). Sphagnum sp.
Consequently'amamOTtlj'fe^l^'^Llh
The early growth of the embrvo is i ?r- walls. (A-i). ^■
succession of transverse walls Thl "® sporogon.um) is thus by a
upper three to four cells of
se ctioi).
 216 BOTANY BOB DEGREE STDDENTS-PART III : BRYOPHYTA SPHAGNIDAE—SPPIAGNALES : SPHAGNUJl 217

the ca^Se°"V'hJ^''u° further divisions and differentiation form base surrounding the foot, ^he seta m the
constriction.iike seta. Tlie''fo'iract™' irfUnc'trnT'ta3up by a peculiar leafless stalk called the pseu-
its way down through the stalk f ^ kaustormm. It bores dopodiuni.
of the female branch Each stmen? nr it
undergoes two successive verficaT di!*"PP" °' The nseudopodium (Fig. IMI) is an outgrowth of the apex
other to form four equal cells ThiTT"ir®' r"®, ' oftheai-chegonial branch. It develops
of the tip of the archegonial branch when
t
^p^^P^g
P .
is^ rip? ^ It
consists
F and G).
ofseveral tiers of cells* ^ ^ embryo now
tier has four cells (Fig. IFlO is thus a gametophytic structure. As elon£
ripe sporogonium at its top beyond the pench
appeal in all the^cdls^'^oTerjh
of an outer layer of cfllt: r-di i i
PericHnal divisions
results in the separation Production of Meiospores. The matu^ sporog^^^^^^^
•
thecium encloLs a cemmf ^ Theamphi- (Fig. IMI) produces the meiospores. It consists oi
'"■ 11-10 H). The celirnfT'
(Fig. Z]'
ceils olr the pnrlr^tu^^:. endothecium
called the end<
divisions to form a massive endothecium undergo repeated CAPSULS
columella (Fig. (j. i o I) Th 1°""= °P I' «
at this stage becomes spherical in^sh^ape sporogonium
As in Anthoceros nerirlinii ••
cium dividing it into two Ta'pr ^^e amphithc- CALYPTRA
SP0RF5AC
inner layer of cells is fertil^rV inner. The
The archesDoriiin/ ^^"^'^itutes the archesporium ANNULUS

nousformtissue
a sporogenous
forms a dometissue
sTan9 f? ^^ ^ by periclinal
^^ick. walls
The sporogc- Epiomms
capsule. It over-arches tL ciwT' Si part of the
^elast cell generation are all -J. sporogenous cells of
mother cells. Each spm'c J^eV functions as spore ^olvmella
Usual tetrad division (nieiosis) to fn r®" K-M) undergoes -^PERICHAETIAL
The cells of h ^P°''^^- leaves

hrih^'l" ° he'can^^^^^^
in thickness. The surfLe Tave theu'amphithecium by pcricUnal
is 4 to 6 layer of cells
(Fig. 11-10 1). It hac capsule wall forms an
„ The inner layers of the r '"udimentary non-functional constriction
FOOT
conLTn intercellular spaces her of thin-walled
Thrnhnr..
not ventilated. ^yuthetic tissue in thehowever,
They, capsule .PSBUDOPODKAM^

transverse rinlo?JnVi ^Pn""® sac advn

sule grows lee?,, pP^dermal cells in th toward maturity a
^nlled theTLyl^^,^7^y
a^n^Ius than their";;^VbPP- Portion of theremain
These cells cap-
operc„lu„, at the If, "' J)- U ddimt'.: le'ch^uTar'li cells
or
Median longitudinal seetron of aa near
fr"yy mature sporophyte.
early stagSrfd"vdo7me ' "e'"calypp'°P®d and thus pro-
the . n capsule. The two are ^epa -
.oonrTted by a constriction, ine
enlarged,
pace
calyptra ruptures in-r^ i ^^vances tn however dries
^^^^rieguiariy. AcVnT|;^-d^n^nturiLy. the deli-
deli-
up.

P like sheath remains at the

 219
BOTANY FOR DEGREE STUDENTS—PART IIP ; BRYQPHYTA ^| SPHAGNIDAE-SPHAGNALES : SPHAGNUM
P-^dopodium is aboui ,2
perichaetial leaves anri fh the capsule far above the
OPERCULUM^
podium compensates for , ' ^he pseudo-
thus, aids in spore dispersal It region and
-Hty it is a ^part ofr^tnJ .pOREiAC*
f:);! ^mSPACE
coSts object. Icolumella
sterile mrspacf
It mainlycells. Theofa^masW^p"!^^^^
columella is hemispherical columella of
dome,shaped spore sac arched by a relatively thin, FOOT
I
o[ the columella. It contains sac thus extends over the top
elaters. With the forr^ationVr There are no "i^EUDO-FODKJM
Ti..p.,.„„„ h°te» -^■.
P'^^Th^^^e^Wlrlhedi^i
nucleated mass of protonhst
S.metopl.y,.
uni-
wall is differentiated into twociirr l• ^eh
^ ^ spore is a tmy,
The spore
inner endosporium Tlip fnnr and the
months under dry conditions remain viable for about six
wall.^TcTnt'S rfl"tye'rs of "111''
capsule wall Is the eDider,n:= oi cells. The surface layer of the
ri^cd, cotupactly arr?jt""elis °f thi^k-walled, cutlcula-
tary stomata FTelT .? • ^"^^^P'dermis possesses rudimen-
stomatarrperture
have thin walls. Thiv
TI rn^ H^• m ^ capsule no
There iswall
ween them. At the ton chloroplasts but have no spaces bet-
Inm or lid. The onpj^i i ® capsule is the disc shaped opercu-
of the capsule byT rWUk^ off f?om tL rest variousKrstges (A-»)-
U-12in the Sphagnum
dehUcenco sp.
of the capsula.
annulas. ' ring-hke groove of thln-walled cells, the
A. Undehisoed capsule.
B-C After drying lust before dclnscence
■ „.u.,ilo (After hTawosohiu).
sa,of1;tef'?,f
mechanism. It takes place on sunnv ?
1M2 A-D, and Disper-
i" '''' explosive
spore tetrads in the spore sac cells of Th' } formation of
Ihis
ac {11-12
resultsA).m the formation
Under the influence
of a ffthT. columella break down,
^Pore swdfff; sfgWy. Tf filament
sed, mature, dark brown cansnl^a i • he sun the wall of the exoo this sta^ ike^termmal^ceU^^^^^^^^^
capsule gradually becomes cyllndrlcaUFiJ^ 1 spherlral maLnt begins to function un ap'cal altetmately rgrt
press:;" td p^^ed (Ftg U' 13E^
indleft
f;r 11 • 13D). Segments are oot off They are arranged
Consequently g
i:: ::istrof ptotosyn.hettc
a flat, eel
. inaprcal
the i eel
up. Thtsputs a strain of'the th n T'l, 'onsion Is tlms sit
£5?SS'j"-rf ."-SdZSZt"""« rS fiitiiSsifSH
nlate. onc cell
 221
SPHa4GNIDAE—SPHAGNALES : SPHAGNUM
220 BOTANY FOR DEGREE STUDENTS-PART III: BRYOPHYTA
plants and cut (lowers which are to be shipped an^d
margmal ce]lJncarThe^^bas^^"'f^^^^ From one of the protected from drying. Peat is also of great use in h°rticulture_ It
erect, leafy gametoDhorp Ti protonema arises the
is added to heavy (clayey) soils to improve their textui e as it keeps
segn>entation T t nv S
't imy bud. Soon a tetrahedral apical cell
■ARCHiSOmAL £66 OR
6P££AI OOSPHEPE
FlUMEtJT
^ ' ■AUTHiRlDIAL BRANCH y^ac^SONIUM
L£AVti
ARCHSCOMAL SRASCH
■:^^^Or£R6£HT
BRAHCHPS

DROOPm BRANCHCS SPSRAIATCZCD

■^■•XAPICAL C£LL
ASTHERimM
OAMCrOPHyTS OF
SPHA6HUM

IHUCUUS

OOSFOR£

r fAh y
GAmOPHYTf m
Germinatic^of Bpo.^! antstaees EAPLY STAGES IN THE
development OF
A. Sporo. of protonema. ZW'rrpi"rE SPOROOOMOM
L'D
B. Germinating of spore. AHNULUS
CAPSULE

Young protonema. SPORE SAC

BipmATm
SPORES

,/•F °''''"°S">arlylobea
.becomes established in the bud. It cuts ofr
'After MuHer)
protonema. '' COLOMELLA
APSULE
cell of the adult gametophore. As a re^. ll»= apical SPORE SPPFF
TETRAD
CONSTRICTlCH
DERM'S

growing shoot of SpAajiiitm is formed. Normalb" "Pically mature spor^nicm

phoL"°" -tablilhilent 'if tlm^eaFrg'a~
The protonema stage mnv
marginal cell of the thallose vegetativcly. Any
SPORE MOTHER CELL
c™ ~cir;r Diagrammatic11.14.
representation of the life eye e.
Sphagnum sp.

ihem porous and prevpts

other humus poor sods it ^u
holding capacity. Ims
in which seeds are germina
and antiseptic nmnf^r.- of us sunerJr.. i ^ blocks,
•Sf/iagiram has be™ , m f' 't^'-cfblly dried m powers grafting scions to p
for other surglcardi4TiFi?r',''°'- surrounding air.

""""■ «SMI
and used"ri",
for packing live
 SPHAGNIDAE—SFHAGNALES : SPHAGNUM 223
222 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPH.YTA
3. Origin of branches in clusters in the axils of leaves.
Some of the by-producls of peat such as peat tar, ammonia,
and paraffin are of industrial use. By filling ponds, pools and lakes
s 4. The spongy cortex of the mature .stem consisting of dead,
with iis extensive growth and debris these areas are converted into empty cells with pores on their walls.
solid earth. Hence this moss is considered a great soil builder. 5. Presence of two kinds of cells in the leaves ; larger, dead,
Sphagnum is also employed as a stuffing material in upholstery and hyaline perforate cells and living green cells.
also as bedding for domesticated animals. 6. Development of archesporium from the amphithecium.
Ecological Importance. Sphagnum is of great ecological 7 Archesporium over-arching the columella.
importance. As it establishes itself on the shore of a lake it gradu 8. Origin of the columella from the entire endotheciura.
ally extends inwards and grows over water. The surface of water 9. Early development of the embryo by a succession o
in the lake is finally completely covered. The Sphagnum plants are transverse walls.
intertwined so as to give the appearance of solid soil from the sur
face. Such areas are called the quaclting bogs. Sometimes seed 10. Presence of pseudopodium and absence of seta.
plants (hydrophytic) grow on the bog. The moss plants gradually 11. Absence of peristome teeth.
die and settle to the bottom. In the course of long periods of time From among the Liverworts Sphagmim resembles the
these swamps, ponds, and lakes, which were sterile sheets of water, Tuneermanniales in the following respects :
become filled with partially decomposed, old parts of this moss
and other hydrophytic plants. Finally these areas are converted 1 Flat, disc-like protonema resembles the juvenile stage of
into solid soil. The constantly accumulated debris raises the surface
level. The moss and the other hydrophytic angiosperms gradually
disappear. They are replaced by forest growth of mesophytic type,
n this way Sphagnum moss modifies the landscape in regions with
small lakes and ponds.
Relationships. Sphagmm is a unique genus of
the Mosses. One is struck by the remarkable mixture of liverwort,
anihoceroie and moss characters it possesses. In addition it possesses
certain characters m which it stands apart from all the three.

featurS"^-"''"" resembles the true Mosses in the following

1. Radial, leafy, upright gametophore. Feltu^cormon with the Anthocerotae (Anthocerotop-
2. Multicellular rhizoids with oblique septa.
of stem, leaves and antheridia as in the r Ateence of apical groivth in,the embryo sporophyte.
2 Origin of the archesporium from the amphithecium.
Jthe antheridium is after the manner of 3: Dome-shaped amphithecium '
4. Structure of the archegonium. forming the columella.
5. Absence of elaters I Pre'sence" of assimilatory tissue in the capsule wall which
of course foot and constriction-like seta.
These features serve to link to the Anthocerotae
rather than to the tiue absence of basal meiistem,
However, m and deliiscence of the capsule by the
gross Sphagnum differs
morphology, leaf rrom the anf
anatomv m its peculiar
sporophyte of differs from that of
phyte. These ar; enumTaiTas'follows 'he Anihoceros. . .^yponnt it is evident that Sphagnum
1- broadly thailose protonema
2. Absence ofrhizoids in the adult gametophyte. combinrthe'cha'^aS of aU the three classes of the bryophytes.
 224 BOTANY PGR DEGREE STUDENTS-PART III : BRYOPHYTA

types'!'' '"■= <=aUed the synthetic

Characters peculiar to Sphagnum are
1. Absence of rhizoids in the adult gametophore. CHAPTER XII
leaf.
2. Ongm of branches in tufts from the axil of every fourth BRYIDAE-FUNARIALES : FUNARIA
3. Absence of a midrib in the leaves General Characteristics. The sub-class Bryidae include
4.
.ex of the Ir' ^he cor- a great majority of the common mosses called the t*""® ® nfaHv
of hyaline cells with large oval nature. It consists mosses. They number about 14.000 species allotted to nearly
thickenings also. The leaf rnncSci sometimes with spiral 650 genera. The Bryidae are characterized by
of cells : the large dead hvalini^ ^^ycv of two kinds 1. Leaf one to several cells in thickness and often with a
elongated green assimilatory cells ^ narrow, distinct midrib.
2. Early growth of the embryo by means of a
conduction. Physiology of water absorption, retention and apical cell situated both at the anterior and the posterior ends.
inth.sthe®ceirwair'lt°^aSs°trha colloidal nature 3. Archesporium differentiated from the outer layer of t e
reason the water in which 'V'ttfftmV^^vvslrhight'ad^^ endothecium.
4 Archesporium not arching over the columella.
REVISION questions
1. List tho salient features mot with in +>,« iir
5.' Presence of a large intercellular space around the spore
sac traversed by trabaculae.
2. Give reasons to explain the foUowinB ■ ^Phagnum.
(i) Dried and dead remains of ' 6 Absence of a pseudopodium.
lent absorbent. ^P^tagnum plant servo as an excel- 7. Presence
the capsule far aboveof atheweU-developcd seta which
surroundmgieve- ^
phytT™ bat .p„„.
8 Columella extending right up to tn
^°°logicaI importance of
4. Give an illustrated account of the life historv of <7 i
5. Diacwss the affinities of Sphagnum. sphagnum.
6. Describe the structure and the !riaTr.-i.v
Sphagnum.
7.
^olopmont of the sex organs of
Describe tho external and internal nir,r.,i, i
'""ir'D.hht— '' 'K' "P"" " -P"""
ofSplnpntim. Of what advantage is the seta to the Moff = 11 Calyptra well developed. .
does Sphagnum corapensato for its absence ? sporophyte? How 12.' Greater elaboration and complexity in the structure
the capsule.
13 Filamentous protonema. . , j , SO
rp, ^ c.ib class Bryidae mcludes about »U
f.miiifrxrero^^ °LToHr of the ^
St Ifrw Fldrch^r'wi'rsiight modifications. He divides the
Tub class into 15 orders as follows
Fissidentales with one iamtiy.
Dicranales with, seven families,
Pottiales with three families
Grimmiales with one family
%
22a
 i£;S99

2i1
BRYIDAE—FUNARIALES:FtJNARiA
226 BOTANY FOR DEGREE STUDENTS—PART Hi : BRYUPliY'i'A
of the family is F'wiianrt. It \ consider
Funariales with six familes number of species included in the family. Here we
Schisiostegales with one family the life cycle of JPunaria.
Tctraphidales with one family
Eubryales with sixteen families Systematic Position :
Isobryales with twenty three families. Bryophyta
Hookeriales with six families Bryopsida or Musci
Hypnobryales with twelve families Bryidae
Buxbaumiales with two families Funariales
Diphysciales with one family Funariaceae
Polyirichales with one family Fuuaria (Schreb.
Dawsoniales with one family
Of this long list the two important ones are the Funariales and Habit and1 1- Ihmnria
^ ' ^eenis colour.
a common moss which
It usually grows
the Polytrichales. The former includes 6 families and the latter grows in dense patches of b g , g banks, sometimes
only one. In this text has been considered the order Funariales. in moist shady situations such funaria includes more
on the trunks of soecies have been reported from
FUNARIALES ZZZSttll mos? widely distributed speeies. It
The members included in this order are small, annual or Ltid to prS s^ils where there has been a fire.
biennial land mosses. They are characterised bv tbe following
features:— ' ® gametophyte
(?) The leaves are sessile usually ovate in form with a
drawn out broad base and generally arranged in a termi
nal rosette. .re,

(u) The broad capsule ha.s a lid without a long beak. ■ >

(m) 1 he peristome teeth are arranged in two rows, the teeth

of the inner ring are usually opposite the outer peristome.
The order comprisp about 356 species grouped under 26
^^ey are plotted to 6 families which are the Gigasperma-
Of these
tl FunanaceaeDisceiiaceae, Oedipodiaceae and Splachnaceae
is widely known. '=i"-imaceae.
The members of the family i
patches. The other characterise featur:s"f thflhSii; ire'L
1. Leaves prominent, usually wide and n• .• i CAMETOPHOR^
except the midrib. thickness
2. Calyptra with a long, slender beak. PROTONEMA

immersed or bor^nTo^nTiong^^^^^ nodding at maturity, W 'iS-l Funaria hygroraetncft.

The family comprises about 90n c.r> • ,^?bbLing tho young leafy gametophoro
protoneioa
undermne genera. The best known
" and widely distributed genus
 228 BOTAJiY FOR DEGREE SatJDERTS—PART III: BRYOPHYTA
BRYIDAE—FUNARIALES:FUNARIA 229

(Fi^ '"cpresented by the adult leafy gametophore

Because of their apparent leaves, stem and root-like rhizoi s
spore (Fig 12 22 C). It arises directly from the the Mosses might be mistaken for any small flowering plant. ese
structures, however, are neither ho- CALYPTRA
system nffiia ^ V g''own u consists of a slender, green, branching mologous nor structurally similar to
nema b?Lrir/?^ 'protonema. The Hlamentous proto^ the roots, stems and leaves of the
darp^"^ ^ fell-like coating on the flowering plants. No doubt they CAPSULB
over the moist soil gi'owth. Many of the branches grow perform the same functions. Such
cells comprising them r7 (ehloronemal branches). The organs are called analogous. The
are separated l?v tra numerous, discoid chloroplasts and Moss rhizoids, stem and leaves
tratrfhe sou ollier branches pene- belong to the gametophyte stage,
rhiloidarbranclS
ween them The nmf
[aTcM The cells of these
-1 hey have oblique septa bet- Their cells have nuclei with a hap-
and trLLry in Funaria is only vegetative loid number of chromosomes. True
roots, leaves and stems, as those of
angiosperms, are sporophytic struc
lateral bud on the proionema (Fig.^dult gametophore.
12-22 It starts
Gl The hnA TiT T as a tures. The cells constituting them ■female branch
into an erect, leafv shoor rallerl tn '' t develops have 2n (diploid) number of chro
bears numerous rtLoids at it ha. The latter mosomes. These sporophytic organs
stratum. The orotonema r which anchor it to the sub- are more complex. They show LEAF

remains behind as th^e InH ,^he leafy gametophore greater differentiation of tissues as MALE PLANT
compared with those of the bosses.
representativrofThrgamemn?
plant. Itbear ttKxTg'a,r is also calledconspicuous
th^ moss They have a well-defined and well-
developed epidermis with stomata
and guard cells (in the leaves and RHIZOIDS
Adult Gametophor e.
stems) and the vascular system. All
these structures are absent m the
stems and leaves of Mosses.
The so-called stem in Funaria
more in height. It bears flat Jt is about 13 mm. or a little is sparingly branched The lateral Fia 12-2. Funaria hygrometnca.
spirally. For convenience nvc calUhrcemr^r inserted branches arise from below the ba- A leafy gametopbyt« bearing a
green expansions as leaves wL axis as stem and the vcs They are never axillary. The nearly mature sporogomum at
taxonomic purposes. Kocli f Laves are more crowded and large "hoape^of theftmale branch.
and leaves phylloids. Sis thSe T cauloids towards the apex. The spiral phyl- ^
gametophore, the third is the rhizmM
attaching filaments, the rhizoids whirh^ ^°risists of numerous
l„,,y is 3/8. The 1;=-- -e
brown part of the stem. Th^ rhi^^ naked, is"in3on the iJL by a'^rawn out, fairly broad base.
aTbey
depth,aremulticellular
at least, equal toandthSu
branchedof ^ ^ substratum to t6) Anatomy
the oblique septaoccur at long
are an intervalsto Ar^
adaptation between
^o Goebelthe(1905)
cells (i) stem (Fig. 12-3A). A^cross-^ection^^o^^^
a simple internal structure. . P distinct zones (Fig. 12-3 A) ;
coIou"ed°hbrown or dark
coloured ^'bizoids are colourless
brown. matureconduction
ones are

k.» -n, °r'~ cal support. In addi i botanists consider the central cylinder
ctiorage and conduction.
root
r:;,:rv— of "primitive type. It has. however, no
 231
BRYIDAE—FUNARTALES :FUNARIA
230 BOTAJSTY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
large and prominent. The incontr^t
are narrbv^aThd^tHick-walled marginal to
cells
te ,[ strength
It has
vessels or tracheids characteristic of the
to the frail moss leaves. The midrib is is a
an upper and lower epidermis. ff° . ^ group of small,
EPIDERMIS
small group of thin-walled, structures characteristic
thick-walled cells with a narrow lumen. 1 tracheids, the sieve
of the leaves of the vascular p ants T^ere is no trace of
CORTEX tubes or the fibro vascular bundle are absent, itiere
the mesophyll tissue or the stomata. ^ nerform
True root in the il/ofise5 is ^nchorVgeand absorption of
CONDUCT/NO
STRAND
some of the functions of roots such as ^ctmrag_.. _
water and solutes from the soil. apical
Apical Growth. The stem means
cell. It is located at the stem "P- ^ t,,ree sides. Each seg-
shape. It cuts off segments pamllel
ment divides into an inner f^ "Ute"-pa j g.v.des
part forms the
to give rise to the major part of
leaves, buds and the outer part ol the ste^ .
The growth of the eaf is means of an apical
g MID RIB nniral cell. The rhizoids grow m lengt
0^11 at die free end of each filament.
reproduction ^^g^tatlve
Funarin reproduces by two rnethods i^^^addered a stage in
and sexual The production of meiospore ailed leafy moss
composed of undiflferentiated. '^tous
walled,
cells (Fig. 12-3 A), i" ^ hi?— »..po..,.™
Stems the cortical c
tain chloroplasts.
'^PiD£RMis mature portion of t The iiameiophyte stage u jo ■
'Thick mLL£DcoRT£7^ has the cortex usua ^
'Tm WALLBD CORTEX entiated into ail outer,
central strand walled, brown
an ^r,n^r
®" ^m4T"'The
V-UllCA i!?vt o the central
tA'15'
latter lies next to
cylinder. ,,
The epidermts
layer of cells
consists of a single
cortex. The cells are small and co phores as a ^ developed on the
SLT"?
E ch '"'hTyoun^
;hc stem thf The7dr;are"s'm;il
.1"an'd conta.
"'"..""d
• thr. ct.m In the maturean
.nd
protonema. These are small resting suitable for
absent The st
^^estomata cells become
characteristic thick-wallc
of the a're
true_stems_^
.hiwidTor projema^^^Wuh^
are^lar?'^ of ^undhF the midrib the leaf pereiinating s
The W celU'"^^langular
S' h^^agonalparcnchymatous cells. vievV_
rectangular in surface P
-s'lLorojjlasts which are comparat^
• '
 —BKjgi *!_■ wm

^32B0TANY fob degree STUDENTS-PART III : BRYOPHYTA BRYIDAE—FUNARIALES : FUNARIA 233

"nspecialised cells'^ arise from the orange spot (Fig. 12-6). The leaves surrounding the antheridial cluster
protonemaJ filaments of the sporogonium. These are comparatively small. The small perigonial leaves aie c ose y se
a leafy gametophore. Thf» which develops into and spread out in the form of a rosette to give protection to the sex
froin the vegetative celk gametophytes directly organs. The central part of the rosette is of reddish colour i be
of spores is called aposnorv without the intervention antheridial cluster with the surrounding leaves is called the perig -
P res are normal in aDDearan,> "O aposporously produced gamete- nium. The orange coloured mature antheridia and the roa^^^
arrangement of the perigonial leaves make the male shoot conspicuou
2. Se=.nalTI
exual Reproduction Gcneucally they are diploid. and easily recognisable. Antheridia at various °
. ^^^^^ihution of No-». n occur in a single male receptacle which is thus e
^5 m the other hryophytes^Tiv^lh ® reproduction takes place. sperms in succession over a long period extending
1 yies, by the formation of antheridia and
fgitsSdt' £ £rs... i .5. fuerJ f.
12 6 and 12-7 J). Each paraphysis usually 4 to 0 or
height. sub-spherical
nearly It consists ofin aoutlme.
single row^he
of ce ThePtoo
^ cells of the
F^NTHCRiDlAL narrow. .All the cells are rich in chloroplasts^ The ,o^ ce
\A/ALL
paraphysis meet over the antheridia. Thus the
male protection and help to a 'tm'te ex moisture around the
branch
female (1931) suggested that they 'telp to
branch § ANDROCvtZS antheridia either by holding suggestion is that the para-
between them by capillarity The other gg from
MOSS 'FHiZOips physes secrete mucilage which protects th y » f sperms
PLANT undue water loss and also assists in the efficient discnarg
'LAGELLUM by building up pressure. .
% 'A' iv\a 12-5 B) of Fanarta hygromet-
.r The mature antheridium (Fig- J elongate, club-shaped.
^ig. 12-5(A—r\ v ri» may be 0-25 mm in len^h^ short muiticeUuiar stalk. The
orange coloured body seated on a snor > , The cells of
C. Sp"?™"™™ mplant
B (MSi?«^-^®rometrica.
sectLn?™'""™'' ""1 autoicons), Tdy has a >cket layer of, polyhe^^ antheridium is
the single layered wall change into orange-coloured
young. As It ripens the t^h'oropMst „f u
surffce™o'?'th^ S" gametophore and are chromoplasts. Within the an j _ gach androcyte produces
clusters, -phe orffan<! ^ Project freely from the cells.
a singleThey are known
biflagellate as the andr ^ ycaP-Ukc
sperro. ^ structure. It fis the
he
vegetative axis of sex orafn? developed in terminal
A the growth of the optcuid«. "Trlustfffie opemuffim
attmpi'at/vdtarge size. They have thicker walls and colour-
less contents. jhe antheridia dehisce only in the
(ii) DeJmcence (Fig. 12 b). j On the access of
nresent of water. It is cells become mucilaginous.
Cer th^ walls of th^^Xir Thf pressure
They absorb water and ® ^ - ^
withm the opercular
rupture, at first, its mner and
^ autoxcous and species m
Antheridia "Monoecious. uell ^eventually becomes p
at the slightly flaTf ^ii'uclure Th
tacle. To the naSd male sh ? 3s.ir,.. """
^ye the antlrnddia^.? ^°rm the male rccep- (Stream ol a viscou.
cluster appears as a tiny
 235
234 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA BRYIDAE—FUNARlALES: FUNARIA
their vesicles. The escape of the vesicles containing sperms is faci the row henceforth functions as an apical cell tvw ®
litated by the contraction of the distended antheridial wall and faces (Fig. 127 D). It cuts off segments alternately right
UBERATED SPERMS

%^-i

SPERMS,

ANTHERIDIAL
JACKET

■PARAPHySIS
ANTHERIDIA

T .
i-ongitudinal hygrometrica
section through the tip of the male shoot.
T?- 10 7/A—Jh Funaria hygrometrica.
antheS^nm
floats to fh?"
There the U
r"""
pressure of the swollen contents within the
the extruded mass of vesicles
the antheridial cluster,
Sissssi?--'"™
0_H. Transverse secf ions.
for^o^ a deUcat^:^ ^ <^ther and spread out in the I. Two sperms.
biflagellaie spermUre ,T. Paraphys's- ^ y gv jhig
vely in water ^ set free
iree (t-ig. l^-? I). They swim about acti-
liberated snerm k an 'il^ antheridium. The
ture The rne emnfv mtK
empty anthendia wither'P"'®"y ™i'ed, biflagellate struc-
and die. ncrpd in two rows (Fig. 12-7 F). Jiyides by a penclmal wall
OT 4 cells away from the ^bove it (tosvards the apex)
the.supSscfaretCc'IiJ' pH-^'^^ ^--=1°? Ihg 2 7 G). Thereafter 'be segments^abo
expanded tip onheTaie shl't"
a single such cell Tt k ii a 'hf. apical, cell "at from
^nthendium originates the Win
S to divide
, in the
tUp same way.
remaining cells.A form the division
Thp nericlmal ex[»sed
12-7 A1 Th.ri • • ' antheridium initial (Fiff contents than the^re^^ anthendium. The plane
y a udiisverse wall into two eel s fFltr 19-7 m r\r *u ^

by anticlinal divisions
 237
BRYIDAE—FDNARIALES:FUNARIA

236 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA housed the egg and a ventral canal cell above_ The
slightly twisted /^Jg^^ersel/protopi^^^^
enclose an axial row of ten or m
neck
-—Uar m structure to that
dium. ^ The apical cell forms the operculum or cap region of the
antheridial wall. The antheridial wall encloses the primary and- canal celU. The archegoniuin is thus s.m lar ■"
rogonial cells. The latter divide and redividc. The cells of the of the liverworts differing only m the foiiowing respect
last cell generation are called the androcyte motlier cells. (i) A long, massive, well developed stalk.
Each androcyte mother cell divides to form tv/o androcytes. The
division, however, is not diagonal. The protoplast of each androcyte (i) Somewhat enlarged venter with a two ceil layers thtck ven-
gives rise to a single biflagellate sperm (Fig. 12*7 I).
The development of antheridia in the cluster is not simul- " M A long, twisted nccU with a larger number of neck canal
taneous. Consequently antheridia at all stages of development may
be found in the same cluster.
Archegonia.
(i) Position and Slrmlwre (Pig. 12 a). The archegonial branch rgRMiNAL C/i-L
springs from the base of the male shoot. The leaves surrounding the
V
P' £RIPHE'^^^
CELLS

pRitAAR ^ COVER CELL

COy£R CELLS
CENTRAL
NECK ■» CELL
CANAL f^ECK CANAL CELLS
CELLS
prihaRY neck
NECK CANAL CELL
ySNTRAL CANAL CELL
PRiMARV NfCK
CANAL C£LL £66
KENTRAL
'CANAL
CELL

VENTER

„ rt f A—I). Funaria hygroraetrica.

^^D^velopment of Archegomum.
jC, F and G. Early stages in longitudinal sectionB.
Transverse (Diagrammatic).
D and E.
H and I. Later stages in the de P ^ after Campbell).
Fig. 12'8. Piinaria. iv.nrrr.n;iim dcvclops from a single
Longitudinal section through the tip of the female branch.
such cell. ' bell the a^^hegonium miiia ^
the top to pro^'tcTthraJcheg^^^^
m the cluster are the paraphyses. "
overlap and close over at
with archegonia

sistingrf rhTtt®ru\r;VrIs'''?he';e^^
on a distinct, muUicellular stalk whirh " '2-8), each con- peripheral cei»»
venter
thick. has
In atheventer wallof(jacket
hollow lavpri ^f^
the arrhp i and ^^o
massive. The
cell layers
^ ^^chegonial venter (venter cavity) is
 239
233 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
BftYlDAE-PONASIALES : FBNAMA
The latter is tetrahedral in form and is pointed below. The three these lateral segments divides by ^ (Fig- 1210 F).
peripheral cells undergo segmentation to form the single layered initials. Thejacket initials surroui d the p
jacket or wall in the venter portion of the archegonium (Fig 12 9 F). The latter henceforth functions as t P * ^rt of the wall of the
During further growth it is doubled. The axial cell divides trans the six jacket initials
versely to separate an inner cell from an outer cell (Fig. 12*9 F). archegonium. Ilis doublec ° initials the primary axial
The former functions as the central cell and divides to give rise to With the differentiation of jac e 12 IO G). It separates
the primary neck canal cell, the ventral canal cell, and the
cell undergoes a transverse
egg (Fig. 12-9 G and H). The outer cell which corresponds to the
primary cover cell of the liverworts behaves in a different way
the louer central cell lorn t to produce an cover
upperceU._ The
(Fig 12*9 F). It functions as an apical cell with four cutting central cell divides trans^ ci scly P ^^^^^j (Fig. 2-1 H).
faces, three lateral and one basal (Fig. 12 9 H). The apical cell neck canal cell and a lo^vel ventral canal cell and the
thus cuts off four rows of segments. Three of these are lateral and The latter eventually pves transverse segmentation foiTO
one basal. The three lateral rows of segments function as neck eee The primary neck canal cell / occupy the median and
cells. Each of these divides by a vertical wall to form six rows. an axial row of neck canal portion of the neck an^
The number of cells in each row increases by transverse cleavage. die lower portion of the neck. from the
The archegonial neck thus finally consists of a single layer made up the neck canal ce s functions as an ®P"® "fgcgs form
of six rows of cells which at maturity are irregularly twisted. The
segments of the basal row undergo no further division. They con
timr'l^ie^e^gmeius cut off parallel
tribute to the axial row of neck canal cells (Fig. 12'9 H).
In other genera of the Bryidae the archegonial initial (Fig.
issH'i'r;:- — -"'7
12*10 A) directly functions as an apical cell with two cutting faces
ARC»E60NIUM
■SPERM

2: Lit
spirally coiiea " o surround-
co^ER ceu

3 C cape freely 7,usi^er. In 'he

PRiMARy.
ing the the ripe arche- 'DISINTEGRATED
CENTRAL
COVER CELL
archegonial and NECH'CAfiAl CEUS
PRIMARY-
H£CK CANAL
PRIMARY
gonia cells break down to
VENTRAL
ventral oana swells.
CANAL CELL
Ara%esjt the Ud or ■N£CK

.r SPBR!^
^VENTER CAiTirr
f„al transfer of
Xhe actual sperms
autoicous ■EG60R-OOSPRPR£
to the the two kinds of IYENTER
Mosses, in on different
Fig. 12*10 (A—Jj, Moss. sex organs are
Stages in the development of archegomura branches, nee ^^^^y be
longitudinal sectionB) external ^gither of the follow- •piiT 12-11. Fuuaria sp.
Mature archegonium
(Fig. 12-10 Band G). It cuts off . transported ni fertilisation
fertjiisawon (Diagrammatic).,
alternate cleavages (Fig. 12-10 D F\ <5f 4 to 8 segments by ing ways 1 c=male i,..o,ak-,ti
branoh it;is situated
situated at
and in the course of time form tV. segments get bisected
nium. Distally the apical cell is ^^^he archego-
changes abruptly in nature It a.. L However, it
truncated base. It has three a tetrahedral form with a
segments by vvallspar.alleUo each of l Ai "? cuts off three cluster at a nij,
' lateral faces. Each of
 ESS

2.0 BOTANY FOK DEOKBB STUDENTS-PART 111 : BKYOPHYTA BRYIDAE-FUNARIAJ.BS: FUNARIA 241

splash the sperms

level. ^
antheridial cluster may
archegonial cluster situated at a lower
complex and highly elaborate structure. It « differentiasporogonium.
foot, the seta and the capsule
ted into the
Foot. The foot forms the c° In Funaria it is poorly
neighbourhood of when^the^li^oV^'^ It is a small, dagger-like, conical J
tip of the leafy
the time of heavy rains. P'ants are submerged at developed. It is ^^hedded in the anchorage and an absorb-
mites which visit thi® anXerkllL''l'nT^h' female branch aiid f"n"='"'"Xand nutrient salts from the parent plant
ing organ. It absorbs wate ., . r to the root of the iern
mucilage secreted by the narauhvses . ^ archegonial clusters for for the sporophyte. It is ^ytic structures. However, they
sperms from the antLridiXeXS'XX' ">= plant. Both the organs ^re spompny
<:>uster. '<> the water in the archegonial
archegonia. The''LXt''oTXLcdor is 'I'h
the shmy mass given out from the onnn Present in
"t'= "'■"sS: m. - ;rJSSS'.?Si»'£"""
guided the sperms reach the archeeonia necks. Thus stalk-like structure. It carries th . P n^etophore. The seta is
open necks and swim down their ne;i< f^™t"ally they enter the
the venter (Fig. 12-11). Only one sneX
accomplish fertilization. MoL than onT
the egg in
«gg to
Sut of f"d&n>y t™orpllKity
^^tfT- i;ra
The fertilised eeg secrptfc o n
d o'f'^Xa^,^ thimSd edls^ ^anV
zygote. It increases in size and enters un'^"""'' the
The segments are cut off alternamW hv? =<=gmentation.
located at the opposite end. The earlv ri J Y° gf°tvmg points, each
m S'unana is thus apical (Fig. 1?17 X ®™hryo of seta, ^ • In the humid swinRS round and
Mon
fhe formed within the venter
cells constituting (Fig. 1217 A \
the embrvn j- A niass embryo
of cells is
twisted. AS u biowo moveiuci...^ ^
F 1217).differentiation.
(Fig. The growth ofIt the
bLonies a grow with
structure round. When the an dr direction. _
resumption of growth by the LX
pace with the developing eXvo a r
by
^ t™e keeps
the
around it. ft is the 'iafyptra ^ tX , P™teedve covering
piaphragm
■pprHtome
TEBTR
^SPORES
aiion
specializat on to J
grow mto a fulI-fledgSd
a high (iPOPurs'S CAPSULE

Prodtict* SPOROPHYTE OPSRCULUM PORTION

OF SETA
c

L. 12.tk (A-C,.Fuuada.>H—a.
Funanals an
an
A. M-'""
The sanio
lid or operoulum romoved and
.t. «-a-r.n«ed.
iw outgrown and rcpXedfl' (Fig
P'^^ced the main sLrvf ti ^
loca- B.
r orpod^-"-
Detacbedoperonlom.
•■• sporogoiVium is a C.
242 BOTANY FOR DEGREE STUDENTS-PART in : BBYOPHYTA BRYIDAE—FUNARIALES : FUNARIA 243

pear-shaped and green at first but later turns yellow operculum (Fig. 12-2). The latter functions m the dehiscence
and the disperlal of spores. There is a well marked Ime of junc
acap.SecalyptratFig.''f2-2rAr&s?Th^^^^
the seta. As it advances towards maturitv ir?a conical hood onor
"P"ght tion that delimits the operculum from the rest of the capsule,
(nodding). Eventually it turns dar" brown the annulus.
ofif lavinff barf nnpv i • -Ihe calyptra Tails f&) Internal Structure of Capsule (Fig. 12*15)
tion and dispersal of spores. ExteTnallv produc-
regions, each specialized o perform well marked •a"Jrsi ,fS"asf
basal, green portion'o'f thVclpsuk "^It ®°™cwhat swollen, a£X"ptb:».» .1.
c^tcti^^orw^at:'"^^ ?)r°"V9dYEr'"Each'stoLrieads into an air space below called the
substomatal
K ]nmatai cavity
cavity Within
vv the epidermis
lasts. isThere
a broad
are ®P®°Sy
distinct
shaped'^fefk^egt^Ttrca"; '"r zone of sterile ceiis ric
intercellular spaces .
spongy zone sur-
gj-r^nd composed of thin-walled,
rounds the central devoid of chloroplasts. The
IS
^ obliquely'pla«d'°™;"ie'\'ruTurl TTthVud'^r the
an

£f'ID€RMlS
SUBSTOMATAL
or
;?■«!—
walled, vertically elongated cells. -=■ ^
PHOrosyNTHETIC
TIS5'J£
CAVITy
aifferiSiaSTndsI'emlXin thickness. The outermost layer
OPERCULUM OR LID
rpBRISTOME
ANNULUS
■ SPORE
SPORES
COLUMELLA
THECA <
air space
co/?r£x CONDUCTING TRABECULAE
STRAND
(NIEDULLA) CAPSULE WALL
SrOMA
epidermis
STOMATAL outer wall and
APERTURE » /' ^INNER i^ALL OP
RING-SHAPED STOMA d' '"I SPORECAUn/
"GUARD CELL ^hypodermis
APOPHYSIS

SUB'STOmTAL
CAviry

E
A.
B.
rLsve™
C.
ASrofTrgSst^r^''^etoma, ,■ sketet?hofamUan^ongitudinalseettonofcapsuta.
Outlme oh ^ eontinuous with the epidermis^
8ubstomatal cavity and
D.
E.
Yo^g stoma in surface
Mature stoma in surface v^.
is the £t contains fewer stomata. The cells withm
of the apophysis di»
 BRYIDAE—FUNABIALES : FUNARIA 245

244 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA

is the cavity of the spore sac. It contains numerous spores at
the epidermis are colourless, These colourless parenchymatous maturity but no elaters. The spores are formed from the spore
cells are compactly arranged in two hypodermal layers and consti- mother cells in tetrads. In their formation the dip bid nucleus ot
tute the hypodermis. Within the hypodermis is the spongy layer. the spore mother cell undergoes the usual meiotic division l ie
It is one or, at the most, two cells in thickness. The cells constituting spores are thus haploid. They are the first cells of the gametophyte
the spongy layer contain chloroplasts and arc loosely arranged. generation. . , . i
The spongy layer is continuous with that of the apophysis where it liu)Opemdum or Lid (Fig. 12-15). The terminal, obhqudy
becomes broader. placed, conical, cap-like portion of the capsule is the
Within the capsule wall is a wide, cylindrical air space It consists of 4 or 5 layers of cells. The inner 3-4
(Figs. 12-14 and 12T5). It is traversed by strands of narrow, elongat
ed, green cells. These strands of green cells are called the trabeculae. 'I
■' t

OPERCULm .1
PERISTOME
ofthea theca
treowregion. Eventually it becomes delitn^ited by 'he appearance
' circular constrictiom ^ow the
ANNULUS
COLUMtLLA

MR SPACE-ir OUTER SPORE S4C

SPORES

MNNER SPORE SAC /Fio- 12-12 B) It consists of two sets ot long, conicai
HVPODERMAL LAPER'.
vv^ithin the other. There are sLxteen teeth
■^SPONGY LAYER
in each set The teeth of both the sets a
are on the same radii and
to each Other. The tapering distal ends f Z-'OUTER PERISTOME
of the teeth of the outer set converge j PERISTOME :
CONDUCTING
STRAND
towards each other. and are unhed termn
STOMA
nallv in a small central disc of tissue, i ne
y-^AP0PHV5IS
sr»"»5" - *p"T
®..nrne teeth are elaborately sculp-
S They are highly hygroscopic
Fif'lSTd) Ld thus play an important
it ;t, flip disoersal of spores. j.- 12.le. Funana hygro-
role m the di persa f

u^ low™ part of the operculum (outer and inner).

Fig. 12*15. Funaria bygrometrica. lllheTnnulus (Fig. 12-'5)- ^ "-'^^^^^pper
4 or 5 layers of cells (P'g-
two or three layers
constituting the edge of the
section ofa mature capsule.
form a special ring °f ^.°;„o layers constitute the atmulus
a solid cylinder of pith-like (delicate rnln'^T
with
region is
detached operculum. .^,aiied cells with distended form. By the
cells. It is referred to as the columella Parenchyma) proper. . It consists of hm-.ailed ce^^^^^^
below S:=efand later ,
strand and
of expanded above. Tust
the apophysis. Belmv it ' '
ouNidp^^ ^ v is narrow
central The remarkable fcatu e compared with its sterile
.S the spore sac. It is a barXht J the columella reduced archesporium ^orJer divided
below It surrounds the columella ti ^ open above and
SmdiefoUowing four regions:-
The often
outer referred to as spore
wall or outer inner sfcTtW
sp„e ' of thickness.
the spore
These cells are colourless. Between h. - thickness.
tween the inner and the outer walls
 247
BRYIDAE—FUNARIALES : FUNARIA
246 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
containing the spores. This , outwmd. As they do so
(i) The sterile cells constituting the apophysis. weather. In dry weather the ^^^'^nonsequently the slits between
(m) The sterile cells forming the several layers thick capsule they separate from each other. ^^vinffins round of the open
wall. them open. At this time assisted gift out gradu-
(in) The sterile cells forming the columella.
{iv) The sterile cells constituting the operculum and the be riX
through the open mouth
peristome.
There is a suggestion that the capsule bears a rough resem when the wind shakes the capsule.
blance to a true leaf of the pteridophytes. It is eminently fitted
to carry on photosynthesis. In case we take the homology to be Development of Sporophyte (Fig. 12 17) diploid
correct the spore sac will then correspond to the sporangium. The sporophyte (spoi"Ogonwm) originates
(c) DeMscence of Capsule. zygote. Its early development consists in the seg
The mature capsule begins to dry up. Consequently the thin-
walled cells of the operculum within the epidermis and the thin-
walled cells of the annulus proper which hold the operculum in place,
shrink and shrivel. This causes strain and eventual breakdown of
the thin-walled cells. As a result the operculum is loosened from
the underlying tissue. Soon it drops off exposing the peristome which
form a fringe around the mouth of the opened capsule (Fig. 12'12 B).
Dropping off of the operculum is said to be assisted by the apical cell
upward hygroscopic movement of the underlying peristome teeth.
By this time the columella and the adjoining thin-walled tissues
have shrivelled leaving a central cavity. It is filled with spores,
In wet weather the exposed, rdugh, hygroscopic peristome teeth * ENDQTHEC/1//^
bend inward. They close together covering the mouth of the cavity
AMPHtrHECfU/^

OOSPORE
ABORTIVE
ARCHEGONIUM
leaves

FOOT
AMPHITHECIUM

■sndothecium

.8 Mc»s, young sporophyte. (Diagrammatio)

Fig. 1^'-"'

, •, of segmentation of the zygote have been

^^Sby^^ny^--- are as foUows
(Explanation in the text).
 249
BRYIDAE—FUNARIALES:FUNAEIA
248 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA

(1) Segmentation of the Zygote. The first division in the These two regions (the amphithecium and the ^"dothecium)
zygote is transverse (Fig. 12*17 B). It separates the upper epibasal are the primary or the fundamental embryonic layers of the
half from the lower hypobasal half. Each of these regions amphithecium endothequm
undergoes two successive diagonal divisions. As a result an apical
cell with two cutting faces is differentiated at each end (Fig. 12*17
embryo thus grows from two growing points—a feature
in sharp contrast to the other bryophytes. The two apical cells
cut off segments right and left. These alternate cleavages from the
two apical cells produce a spindle-shaped embryo (Fig. 12*17 G). c
The ^^tter grows rapidly at the upper end. Here the segments are FIRST RING
cut off in quick succession alternately right and left. The segments SECOND
divide and re-divide to produce the capsule and the upper portion RING

of the seta. The apical cell at the lower end of the embryo is not
so active. The divisions at this end are less regular. The deri
vatives of the apical cell at this end differentiate into the foot and
the lower portion of the seta. endothe
The apical growth of the embryo goes on for a considerable •n
cium "■"-—1—PERIPHERAL ^ CENTRAL
time. ^J^^s^qyentj^y it assumes an extremely elongated, cylindri ■FIRST RING ^ ENDOTHECIUM CELLS " enDOTHEOUM
cal form (Fig. 12-18) Its lower end burrows through the arcliegonial ^SECOND RING
CELLS

stalk into the tip of the stem upon which the archegonium is ■COLUMELLA
seated. Eventually n enlarges to form the foot. The latter fTHIRD RING
ob ains water, nutrients and food for the developing sporophyte FOURTH RING

from the parent plant. Subsequent to the formation of the foot FIFTH RING

the upper part of the embryo elongates. Its terminal end enlarges INNER SPORE
m diameter and becomes differentiated into the capsule region. ARCHESPORIUM SAC
The narrow portion which joins the capsule region and the foot is
the seta. It elongates gradually and is differentiated from the
capsule late.

cutoffom!by the apical cell ofofthe

Capsule.
epibasalThe
half alternating segments
subdivide with great -r-
Fig. loio/A-D
12*19 (A ) Funaria liygrometrica.
showing stages of
^ two alternating segments are Diagrams of
development at 8UCC levels below and
its apex.
endothecium.
Earh^nf (I^igl2*19A) each being semi-circular in outline.
resultant flour cells
segments divides quadrately
are arranged by a radial(Fig.
vertical
12*19wall. The
B). Each ^7; SSr—n "of five riags of eells froar
division wall^is then undergoes an anticlinal division. The
an internal 11 Sincc it is in a plane perpendicular to ■ m They arc recognisable
^r™®-haXegSn'softhe seta j,'° xhTinner
central strand region
in the
' c?oss.sectbn 7s
(Fig. 12-19 Cl ThptVi'"i . r other more or less triangular
only in the reaangular daught7 ™ls ''wit'h'th''"r''
penclinal walls the embrvo in the ; /ormaiion of the
two regions, the outer and the innS divided into
ampbitbecium and the latter ^ former is called the
endothecium, at this
defined rows. They extend thm u
(Fig. 12;19 D). The
column of four clearly Differentiation of T.ssues
which is destined 'to I™ 7l "'"r embryo Theca Reg*®" * « *. • ,m The next divisions
endothecium forms a central rectanJnl ^
cells (Fig. 12*19 D). ItTs su^ nded by afour
eight celled amphithecium.
quadrately
single layered,
 BBYIDAE—FUNARIALES: FUNARIA 251
250 BOTANY FOR DEGREE STUDENTS—PART ni: BRYOPHYTA

endothecium. The four central quardrately arranged cells of the

The oHicrZayer external to the first ring consists of 8 larger
endothecium undergo cleavage in the same way as the four cells cells. These undergo two successive divisions. The
of the quadrants at the apex of the embryo (Fig. 12T9 G and D).
The first walls are curved and anticlinal (Fig. 12*19 F). The follow
ing ones are periclinal (Fig. 12'19 G). This results in the separation
of a central group of four endothecial cells surrounded by a layer
of eight peripheral endothecial cells (Fig. 12-19 G). Each central 'inner
-Archesporium
endothecial cell undergoes two successive divisions by intersecting
walls. In this way a group of about 16 cells is formed in the centre .OUTER SPORE SAC
of the endothecium. These eventually differentiate into the colu-
.SPONGV
mella. The columella forms the core of the theca region of the
capsule. It consists of large colourless cells.
The eight peripheral endothecial cells undergo cleavage by .HYPOOPP^'^
radial walls. The daughter cells divide periclinally. In this way ■ePlDFRMIS
two conceritric layers, outer and inner, of endothecial cells are
formed (Fig. 134 1). The cells of the outer layer constitute the FiE 12-20. Funaria hygrometrica.
primary archesporium. They divide at first by radial walls only. Transverse section through the fertile portien of a
The cells of the inner layer remain small and are transformed into
the inner spore sac. The latter remains one cell thick. The arche
sporium in Funaria is thus endothecial in origin. It is formed from
the outermost layer of the endothecium. Originally it consists of a anticlinal and q). ^°e ImTr^ of'diesrtwo layers is
single layer of cells. It is thus extremely reduced forming but a each is the result S- layer of 16 cells again
known as the "q^e 16 cells of the second ring undergo
small and narrow part of the theca region of the capsule. It extends
neither to the base nor to the apex of the capsule. It is a barrel-
undergoes They develop chloroplasts and el?ngA?e «dtAU^
shaped tract open at both ends. The archesporium cells may under a radical change, i hey diameter of the theca region mtercellula
Dwinfftothe ^crease in diameter oi
go sub-division to form two cell layers thick sporogenous tissue. spaces appear between the X ^ each other laterally by
All the sporogenous cells are functional. They mature into spore
mother cells. The elaters are lacking. Each spore mother cell
■L ThA thus beeomyepaiated become wider
undergoes the usual tetrad division to produce four haploid spores.
The four celled embryonic or primary endothecium in the theca re
gion thus gives rise to :
(0 The columella.
(«0 The single-layered inner spore sac.
"A s
the air space aim ~.h .h.
{lii) The sporogenous tissue with all the cells are fertile
functional. No elaters are formed.
and zt A.re.«--■'£ (S
rhe anticlinal on The"^^^
. 1,. Amphithecium. Meanwhile the "^rerTaTer 5 32 cdls eon^titutes the inner layer
?!• P^ary amphithecium have undergone rapid
remains
region ofone
the orcapsu
at u. external to the third ring divide
The 32 cells of the undergo the anticlinal
1 i-tv nericlinal walls.^ T y ^ each are formed
radial walls to increase in number tIip H ^ ^ divide by Two concentric laye is known as the fourth
clinal divisions to ferns th fFri2d9I)- The inner layer O eells of the fourth ring la ter
into the outer spore sac. The celk Af
chloroplasts.
mature
^ outer spore sac lack %gf ptkltaardivSon' producing 2 to 3 layers of c ells.
2a2 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA 2o3
BRYIDAE—^FUNARIALES : FUNARIA

'1'° 'j" "^ypodermis region of the capsule wall.

Thpv arp Vi ^ ypodermis are colourless as they lack chloroplasts.
inner rings of/-^J^tderLo uImicSt^SrtillTh-:
eight cells of the is now called the inner
matous '"S" thin-walled and parenchy- nng^ becomes 3_ (Fig^l2
peristomiallayer 19-21
21 A)
A). The second ring which
cellsconsists
in this
of 16 cells forms the middle
eventu^Hv manr/e*'; ™g by surface
anticlinal walls only. They layer undergo no nf the DerTsmme of the mature
peristome teeth m each ring P ^ ^2 cells functions as the
wall It is calM t'he
thlLned wat Here^^ndTe^e
layer of the capsule
Sreatly c%sul e. Thetinrd^
outer peristomial lay
ff'tle"three peristomial
another that one cell
functional stomata Trcv Irf how Pictured with layers are so disposed of wi P jj^^er
apophysis region ^ ' ^°"<=t-er, more numerous in the of the iniddle °PP°ristomial layers form a dome-shaped
peristomial layers. The P, n ^his region. From it are
Lructure external the inner
embryTnicTmpTithtltmIn
pnitneciiim m the theca region from without inwards ™ differentiated the outer peri originates at the juncture
peristome (endostome).
of the outer and T
middle peristomia The inner peristome
peristomial
(«} Single layered epidermis. is formed at the
(w) Two or three layers thick hypodermis. layers, ^ tangial walls contiguous to the outer
(m) One cell or at the most two cells thick spongy layer. (cutin). It IS laid the case of outer peristome. In

across'thfahspiS'™'^''''®'^'^™ or the trabeculae extending et'tf tr1n.rVe%tohe the thickening wali materta, ts
(V) Three or four cell layers thick outer spore sac. EPIDERMIS OF
OPEKuum
PERISTOME
COLUMELLA (OUTER)
(&) Differentiation of Tissues in «-v
Region (Fig. 12-21. A-C). With the deveTopment of^hrtre^ iNNea peristom/al
LAr£R
columell a

ANNULUS
region the ap.cal portion of the capsule differentiates into a cone of OUTER PERISTOMIAL
tissues I, constitutes the operculum region. In this tissue oriX LAVER ■l/INER PERISTOME

EPiDERhfiS RIM

co^erttSmT' oftheoperculumregron thin-walled tissue '

consislf'ofth: :fme"tvo ^ryoni^'Lyir^Tre

tract of four orimarvend^r^- central
MCHtSPORiUM

EP/DERMIS- ■toothOF OUTER PERISTOME

thin-walled
endothecium ihus differenli^i^ ' "T " is produced. The TOOTH OF INNER PERlSTQPlE
nssuE ^
columella
which lies in this region of the caps"Te. OUTER
COLUMELLA

cells unde^grthfu1"faSaT-
theca region) to pSce eo
■=*§!>'Pt™ary amphithecium
P'="olinal divisions {as in the ■r- 12-21 fA—C). Funaria hygrometnca.
.—e section
I'is
hssncsfrom these'rings ,to
are differentiafed °re™„ "T °P«™Pl»thecium
as follows^'''"'"" cells,
peristome A.
B.
poristKjme celU. through the operculum
i portion of the relationship of pens-
)• The peristome-LT teed'^'r"p""t '.l^^development 0.

i„ ftnnrin is dcXed"fro,:!:^The\t°e": and upl^o/

tome and rmi.
 255
254 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA BRYIDAE-FUNARIALES :FUNABIA

deposited only on the outer wall of the inner peristomial layer operculum is detached. To sum up the amphithecium in the
(Fig. 12*21 B). The vertical anticlinal (radial) walls of the peristo region of the operculum gives rise to :—
mial cells of the three layers undergo no noticeable thickening. It is
along these unthickened anticlinal "walls of the peristomial cells that (i) Surface layers of the operculum consisting of the epider
splitting takes place. The unthickened radial cellulose walls and the mis and the tliree layers of thin-walled, parenchymatous cells inter
contents of these cells disappear. Only the two thickened cutinized nal to it. They cover the peristome.
walls are left behind. These split longitudinally into two (outer and
inner) rings of peristomial teeth with 16 teeth in each ring. These (u) The two rings of peristome teeth.
perptome teeth are built up of strips of specially thickened and
cutinized cell walls. They are curved, narrow, triangular plates with (Hi) The annulus.
thick, tr^sverse
series. The teethbars on the
of the innerbrownish
series orred, eonical teeth
endostome of the outer
are comparatively M Differentiation of Tissues in the Base) or Apophysis
(c) between the theca and seta regions
sma er, more delicate and colourless. The outer peristome teeth Region: ^ broad zone of ceils the basal,
converge towards their tapering distal ends to meet a small central in the young c^psu g ^ the apophysis
disc. swollen, sterile cells in this region
(Fig. 12*13 B). Itisconti-
Formation of Operculum, The three rings (4th to 6th) of divide and re-divide to , r g^ta. The archesporium and
the amphithecium which are on the outer side of the peristome nuous with the of the theca region are not differen-
the inner spore sac charact ^j cells divide and re-divide
and fifth rings jointly produce 12-21 A). The cells of the fourth
a tissue. It consists of thin-walled tiated. The eight primary amphit^^^^^^^ The latter
parenchymatous cells and is arranged in three layers. This thin- in the usual manner to the conducting
Thp^rellf nf portion of the operculum. get differentiated i^ito th . ^ differentiation is practically
n Au ""g mature into the protective surface layer ftrand. The cells of the first, second, third
the same
fourthasrings
m theform
theca g gponey
betweenzone
them.withThecells
cellsrich m
outefwtlls^^ epidermal cells have greatly thickened and of the
chloroplasts
fifth and intercellular
ring mature 11^0 the sp . epidermis punctured
hypodermis nor here
any
Differentiation of Annulus. Very early in the development
of the capsule there appears a narrow, shallow, circular depression and there with Apophysis is, therefore, the solid
at the junction of the operculum and the theca regions(Fig 12*21 G1
It delimits the former from the latter. In this depression lies a
narrow zone of thin-waiied narrow cells. It serves as an abscis-
sion layer. It is here that the ripe operculum becomes detached
The epidermal cells above and below this narrow zone show active
growth and elongate radially. Two distinct rings of cells are formed
one above and the other below the narrow Lne. The form" is
called the annulus and the latter rim or diaphragm 0/^60^ around its oasc. krnwn archegonial necK is ca p

larro™" those of the a^the top of the

rurrounds and Lamr'^t becomes pendent or nodding.
The annulus forms the perinherv nf k.. j
operculum. It consists of about 5 or 6 layer^ of
lying one above the other. The cells of thf^ cells
layers become specially modified. They are j Comparison thT'tracwre and the develop-
elongated. They have thicker walls and fmm radially
the mature detached operculum. The cells nf ™ and Funaria. ftmaria, we are in a position to
the annulus, which constitute the annuluc « ^^^^rs of ^ent of .'he.^=P°J°f^;rophyte of Sptajmi™. The two may be
walled. They are reported to' bTfi^ '}}}?- 'rtSsted ari-ollows
mucilaginous contents of these cells absorb mucilage. The
capsule matures. Subsequently thev I as the
y aey are destroyed and the
rr---

BRYIDAE—FUNARIALES : FUNARIA 257

256 BOTANY FOR DEGREE STUDENTS—PART HI : BRYOPHYTA Sphagnum Fnnaria

Sphagnom 7. The capsule has a well develop

7. The capsule is bounded by ed epidermis
Faoaria punctured with function
epidermis bearing fiinctionless al stomata, particularly numerous,
stomata. They have no pores. in the apophysis region, Thoy open
(a) Development: There are no substomatal air spaces. into sub-stomatal air spaces which
■ 1. The early growth of the embryo Thus tho many layered capsule wall communicate with a sptem of inter
1. The oarly growth of the lacks ventilated photosynthetic tissue. cellular spaces aerating the green,
IS by transverse cleavage followed by photosynthetic spongy tissue in the
intercalary enlargement of the 302- ^ apical. from
cut off alternately The tho
segments are
two apical
rnents. There is no apical growth. capsule wall. The capsule wall m
^lls each located at tho opposite ond. Funaria has thus a well ventilated,
ihe apical growth later gives place to photosynthetic system. In the theca
intercalary activity. region there is the two _ coll layers
2. The entire endothecium cives thick hypodormis, in addition, beneath
rise to the colnmella which is a bulky 2. Only the inner part of tho the epidermis.
mass of parenchyma cells. It has a endothecium forms the columella.
rounded apes. 8 Within tho several layers thick
8 Within tho several layers thick capsule wall is the wide air space.
3. The archesporium is derived capsule wall there is no intercellular It is, in fact, a highly distended cavity
trom the inner layer of the amphithe- 3. The archesporium arises from space so that tho capsule 18 a compact in the spongy zone of the copsule wall.
ClIUU. the outermost layer of the endothe- solid structure. In the centre of this wide air space
cinm. remains suspended by filaments of
The archesporium forms 4. The archesporium is a barrel-
green cells (trabeculae) the columeUa
dome arching over the columella. surrounded by tho spore sac.
shaped structuro open both at tho top
and tho base. It does not arch over the 9. The spore sac is narrow and
columella but surrounds it instead. 9 The spore sac is a wide, dome- barrel-shaped. It is open at the top and
(6) Structure: shaned structure over-arching the thfibasc It does not over-arch but
rouSded tip of the maesive central surrounds tho columella
_ 5. The mature sporophyte con columollu- the central core of the capsule m the
5. It consists, of a foot, a long,
sists of a foot and a spherical capsule. slender seta and a pear-shaped cap theca region.
There is no seta. Its place is taken sule, The pseudopodium is absent.
Up by a constriction. Tho function
of the seta is taken up by the pseudo- The foot is poorly developed.
podium. The foot is massive and
functions as an haustorium. The points of resemblance between the two are :
capsule of Sphagnum is 6. The capsule shows complex
1. Absence of elaters in both. _ c ^ ua in
comparatively a simpler structure
xepresentmg only the theca portion structuro with considerable differentia
concerned with spore production and tion of tissues. It shows three well
2. Dehiscence of the capsule by the separation of
marked regions: both.
the opercular portion concerned with
dehxscence. The apophysls is absent 3 Rupturing of the calyptra.
from the
cular region. Tho theca (a) Apophysis—a solid, somewhat
operis swollen,
region sterile, basal portion mainly 4. Presence ofa well defined columella m both.
also comparatively simple in structure! concerned with photosynthesis and
conduction. Nutrition of Spotophyte. The thL"
urn-shaped
--"7!?^t'eao?u?e"Tis thus able to synthe^ae a considerable
portion concerned with spore produc- poriioii ^ ticciip is developed. H . ' ., , ".
its
hasphotosynthetic
been estimated tissue
by certa ,, yJjogists and is gametophore.
considered equiva-
For
! cios6ly^liate!M"°i epical region
lent
vratertoand
thatmineral
of the fow'"
s"hsta . ^hich sporophyte
must come of fromFunana
the soil, it
thus
rlpnends upon the parent piam.
15
fsemi or partial parasrte.
f?'
258 BOTANY FOR DEGREE STUDENTS—PART HI: BRYOPHYTA
BRYIDAE—FUNARIALES:FUNARIA 259

Germination of spores. The spore (Fig. 12'22 A), on falling sporogonium. It is partially dependent on the gametophyte for
on a moist soil, immediately germinates. It, at first, absorbs its nutrition. The nuclei in the cells of the leafy plant contain n
chromosomes and in the sporogonium 2n chromosomes. The
haploid leafy plant reproduces by gametes. The diploid plant
SPORE
reproduces by spores. Because of its characteristic method of
reproduction by gametes, the haploid leafy plant ^ is often called
A
RR070NEMA
the gametophyte. The diploid plant because of its reproduction
by spores is known as tlie sporophyte. The leafy gametophyte
represents the sexual generation and the sporophyte or sporogo
CHLQRONEMAL
BRANCH nium asexual generation. The gametophyte is developed from
the spore. The sporophyte is formed from the zygote. The
RHIZOIDAL latter is the result of the union of two gametes (male and
BRANCHES
female). The two kinds of individual (sporophyte and gametophyte)
alternate with each other in the life cycle. In fact one of them
is the parent of the other. This phenomenon is expressed by
Fig. 12-22 (A-C). Funavia. the phrase Alternation of Generations. It means f
A, Mature spore. in a single life cycle of two vegetative
B, Germinating spore, functions and different genetic constitution.
C, Protoneraa. develoDed during the gametophyte phase are the
protonema, the leafy gametophore the org^s (anther^ m
water and swells. Consequently the exospore ruptures. The and archet^onia) and the gametes (sperms and the .
endospore protrudes out in the form of one or two tubular out
growths called the germ tubes (Fig. 12 22 B). When the two germ
tubes come out from the opposite sides of the germinating spore,
a cross wall soon appears at the point of emergence of the germ
tube. ^ The cell thus separated turns green. It grows in length g^neSbon. They differentiate by meiosh in» ^
and divides by septa to form a filament of green cells (Fig. 12 22 C). haploid. The meiospores are therefore the first
The filament grows on the soil and branches freely. The growth next gametophyte generation.
IS apical. The branched, green, alga-like filament is the primary
protonema (Fig.^ 12-22 C). Most of the branches of the protonema SUMMARY OF THE LIFE HISTORY OF FUNARIA (Fig. 12-23)
grow on the moist soil surface. They are green and thus known Thpre occur
1. Gametophyte. There occur two distinct tophore
individuals(Fig
in
snil branches. Some, however, penetrate the
the life cycle of They Leafy gametophyte is
rhLoid.l absorbing and anchoring organs. These 12-23,
far more1),conspicuou
and die ofdie germinating
germinates as it falls on .be moist
between the cells. 1 he moss protonema with its chloronemnl nnH spore (Fig. 12 23, branched, green filament of cells called
soil and produces a in The protonema is the juvenile
a resembles the heterotrichous thallus of the protonema (f.g- ^ It has two kinds of
thi presence of , distinguished from the latter by stage in the life cycle. It is pure y b branches. The
whenformation
the protonema has isreached
braLhes, the ^he moLTbranches
^ latter penewate
said toahe They are formed former grow honzomay manufacture food
The of buds of development,
naturally occurring growth factor known k ^ presence of a the substratum- The anchoring and absorbing organs.
by the activity of its apTcal cell .row? ^ bud and the rhizoidal
Loner or later certa.n
chloronemal
epical ceU with three
the usual form. ^ leafy Funaria plant of
branches grow into p^ity of its apical cell each bud Stilly
cutting faces. With the ac ivh^ ^ F substratum by the
e T? (Fie 12 2'^^ T u i n grows into
fhizoids. a leafy
. With the of leafy shoots the
independent protonema
representative
of J'unana
other. Onethere occur istwobaploid
of these distinct indi^^^^^^
Ti- • tu ^ ^ theunlike
life cycle
each
moss plant. The other is diploid Th. A^ ^ .'"dependent leafy, dies leaving ^ach leafy axis go-called moss plant. The
The diploid individual is the of the gametophyte phase.
 BRYIDAE—FUNARIALES : FUNARIA 261
280 BOTANY FOR DEGREE STUDENTS—PART III; BRYOPHYTA

protonema and the leafy shoot are the two stages of the gametophyte the access of water the axial row of cells in the archegonium, except
generation. The former is purely vegetative in function. The ing the ovum, gets disintegrated to form a mass of mucilage. The
leafy gametophore is reproductive It bears the sex organs.
The adult Funaria plant (Fig. 12'23, 1) is an upright, leafy
axis about 12—20 mm. in height. It iias a slender, central axis, the
so-called stem. The latter bears multicellular, branched
at its base and leaf-like expansions throughout its length. The MALE SMNCH
leaves are arranged spirally and are more crowded towards the ANTHPRlblUM
SPEPMATOldD
apex. They are sessile and ablong-ovate with entire margin and N

pointed apex. Each leaf is traversed by a distinct midrib. OOSPMERE'^Mj

FEMALE BRANCH V*
Internally the stem is differentiated into an epidermis, a
cortex and a central strand. Excepting the midrib the leaf is ARCHEGONIUM
N
composed of a single layer of parenchymatous, rectangular eel s
rich in chloroplasts. The few layers thick midrib has a small con
MOSS PLANT
ducting strand. N

Sex Organs. The leafy gametophore is concerned with scxua

reproduction by the formation of anlheridia and archegonia. 1 hey
are borne m separate terminal clusters on distinct branches ol tne i NUCLEUS
fNUCLEUS

sCt ^'^\^"thcridia are borne at the tip of the main

bas:^TtLtai:shoT""
'gametophore
6 _
EMBR/0
ANNULUS -2N
LID
■PERisrOME

orange colouredThe
and a body. structure
bodv t1 - ^ is ^more
^^ort, •^ w;„eilular
' ,vaUstalk
one SPORE TETRAD
INNER
.columellA
,ARCHESP0R/UM
cell m thickness. At the ^ ^"^^<^"rlium has a jacke the N
SFORE S-^C OUTER SPORESAC
TRABEC'JLA
antheridial wall is diff^ end of the anthe 'AIR SPACE
operculum. h consists ^ lid-like structuie. SPONSy LAYER
WPODERHUS
apical cell bv t\.r • or two larcre celU clcriveo • jts 'pnotosyn'Tnetic tissue
^rly stages of developm^ antheridium of S7QMA
^EPIDERMIS
BSPORE MOTHER SPOROCOMUM 'conducting strand
gTvel packed the wall is a latter CELL 2N 7 SF.TA
g^srisetoabifiagellate b androcytes. 3)- ZN

anth^°^ the sperm (Fig- 0.,a4.U.icropr.eut„uo„o

Fig. 12-23.
Diagrammatic ropresenxuuu»
cy..
cycle
.

^alk ^ ^ndrorvfA by hydrolysis. . t,itlon up muclage •n thethe canal absorbs

^ water and swel
are forced ls. Consequently
apart,
f"drocyS e,„
end cells of the archegomal
archcg neca „chegonia are borne
rounding ibp ^thr^^ out.'With
. ^pcrms theswim
escape and d.sso ^ the Fertilisation. branches. Naturally they are
The .
■••ne ar/tU-. . "'^"'dial cluster.
-""-'^criaiai duster.
^2-23,2') are typically
,
flask
loO^^
-d
iur=/s:ri;;, S'-r" '■?«»""
^^SthenerT^^
Wl Port;
has"n'^
cells T^if
muiticellular stal
thick jacket layer ilarg;
yen
cavity or the ; is doubled m the
' ovun^"";
With a 'he arcbegonittm.
ventral canal cell
it

: a,',. liWilf
a,i ^ - V - , ' ■ . AUi'i."......
 262 BOTANY FOR DEGREE STUDENTS-PART HI ; BRYOPHYTA BRYIDAE-FUNARIALES : FUNARIA 263

tion that the sperms may be splashed out of the Deriffonial rosette the nutritive tissue or the spongy parenchyma. The sub-stomata
tV.z'xzsi'' suuiud'S'frowl™ air spaces communicate with a system of intercellular spacp yerUilat-
down from the perigonial cup aTa'hiXe"'!'""?® ing the green, spongy parenchyma. The capsule of Funaria is th^e-
cluster at a lower llvel Snilll • ? archegonial fore self-supporting so far as its carbohydrate food is concerned, or
antheridial and the archegon^ water and mineral salts it is parasitic on the parent ^
which it is organically attached throughout its life. Within e
secreted by the paraphysis Lve nho B
sperms to the arche!ronia a u
mucilage
suspected to carry the photosynthelic tissue is the conducting strand continuous above wi
the columella and below with the conducting strand of the seta.
accomplished. Once the sperms rLl^he afc^^
swim to the open necks of th^ , u cluster they
enter and swim down the canal m which they eventually The fheca has a centrally located column of thin-walled, sterile,
parenchyma cells. It is the columella. Surrounding the lat er is the
aothers
rule only one sperm uniterwlh ih ^
perish. ^^"^ter. The
to effect fertilization. As barrel-shaped spore sac with its top and bottom knoc e ' . ^
spore sac therefore does not extend into basal and the
portions of the capsule. It is bounded by an inner wall one cell thick
the pioneer structure of^ihp ^2'23, 5) thus formed is and an outer 3 or 4 cell layers thick. Outside the ^P -
air-space traversed by green filaments, the
-p^e
secretes a wall around it Lrl It is diploid. It columella and the spore sac thus he suspende mother cells
fgain undergoes two successiuA ,i* ^ ^ transverse wall. Each half
the trabeculae. The spore sac contains ■ *r,n ^Fip- 12-23 9) to
lag apicalalternately
^egments cell is established
ri^ht anda 1
I.G ^ An actively
^he apical divid-
cells cut off (Fig. 12-23, 8) ^vhich undergo the tetra^ddw.^
form the haploid spores (Fig. 12 23, 1 ) masses External to the
'r ^ long s ender 5 Soon the embryo (Fig, 12-23, 6) are functional. No elaters are formed in the ?^nsse. . ^gy
SS its way th,t'h The loWer end of the air space is ''^'=
layer one or two cells
contimou^ below with the spongy
„„„^he hypodermis consisting
into tfip 1 j ^P^^ the fema]f» h 1 into the
^rorayrtfrfiS cir Sehypoder^s is bounded by the
single layered epidermis with fewer stomata.
elongates Sporonhvte grow nt nh" , j^ta AW.fl..
is the operculum or lid. A annulus.
cells delimits the conical teeth constituring
Beneath the operculum are two _ g form a cone-hke
the peristome". Together the P^—
structure which obstructs the mouth of the spo.e
hygroscopic.
1• _ Thp thin-walled cells of the
The mature «rtoIj"he operculum in place, shrink and
annulus proper, which hold rr., N:j qj. the operculum drops off
shrivel Finally they rupture. The latter close
exposing the dome-like hygroscopic p ^ shrivelling of the
over the central cavity formed y ^qw filled with
rnlumella and the adjoining tissue _ ^ wet weather
snores The escape of these spores p weather the teeth
:rthe peristome teeth close tog^^ frL each other. The
bend outward. In ^°"\hrough these slits the spores escape a few

SSi- ■ """■
juvenile stage o( Funana.
 265
BRYIDAE—EUNARIALES : FUNARIA
264 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA

Significant steps in the life history of Funaria

A. Gametophyte
1. It consists of (a) a branched, green, alga-like filament,
the protonema and (6) a leafy gametophore the so called moss
plant. The protonema is short-lived. 5. Early apical growth of the sex organs.
2. The leafy gametophore consists of a slender, stem-like
central axis bearing rhizoids at its base and leaf-like expansions B. Sporophyte ^^brvo sporophyte is bi-apical. It
throughout its length. It is independent.
3. The so-called leaf is traversed by a definite midrib more
.1..
cleavage is
!*'■ ■■ the llerworl
than one cell in the thickness.
4. The rhizoids are branched and multicellular. The septa
between the cells are oblique.
Tsri:nglhertlier®than that_,ernaUy
shows greater tissue differentiation.
of the livdifferentiated
erworts and
5. The sex organs {antheridia and archegonia) are borne
in terminal clusters, at the tips of separate branches of the leafy
gametophyte.
6. Early development of the sex organs is by means of the
activity of an apical cell.
B. Sporophyte.
7. The embryo sporophyte grows from two growing points,
each located at the opposite end. The growth of the embryo is thus Urabsence in the liverworts. ,,^5 suspend-
apical.
8. The mature sporogonium is differentiated into the foot,
the seta and the pear-shaped capsule of great complexity. The „
seta is long and carries the capsule far above the leafy gametophore.
9. The mature capsule is differentiated externally into
apophysis,theca and operculum regions. Internally it consists
of se>^ral tissues. There is a well developed ventilated photo-
tissue and a large air space in which lies suspended
the spore sac and the columella by filaments of green cells. The
capsule has a complex organisation in the form of an operculum the form of operculum aiid^P Uverworts m
apicafpordor^ dehiscence and dispersal of spores at its lUt&'eUrts rnfflS ^hen present is
The peristome is absen
IS aerived
is derived irom the exterior portion of the endotheriiim
'he columella. Tt
andi<? very simple m questions
ferdle'thsutthus Sc^'^dltTl^et^' , hlsttheehler..—
The chief difFerences between Funaria and t E,„„,iaceoe 3ig„ifloaet steps in ^^Jfj^^thyLiveTworts t
'si
tne Liverworts
A. Gametophyte.
Presence of protonema in i-Vink
in the liverworts. mosses and its usual absence
funaria.
moss plant as compared
• orts.
 266 BOTANY FOR DEGREE STUDENTS—PART in : BRYOPHYTA

7. Describe the external and internal structure of the cansule of

Funarta. ^

• xu X?' of theamphitheciumand
oi the capsule (apophypis, ondothecium
theca and opercular regions) of

XT A^' fertilisation take

Jlow does structure and
place in thisdevelopment
plant ? of sex organs in Funaria. CHAPTER Xin
10. Describe the external features and internal structure of the vese- general conclusions
tative organs of the gametophytic phase of any moss you have studied.
nf fhA
of sketch of a
the capsule of any moss you have studied. median longitudinal section :/ Origin of Bryophyta.
subject. The little geo ^ the present day bryophytes
In factl-ip/»n
no ^'1 S's'^about
v"ews about their origm must therefore
have been found.
found. Alt
A u following
. three heads
,.v,.-AA bfarls :: —
-
""I'rrirrrcrr —
plants. . , ^
(ID Evidence from ontogoy ^bg plants of other
(III) Evidence
groups.
groups
based on analogi . . fv, o
Bryologists are ^"'"^^^howXtThfbr^o^^^^
phytes. The
descended fromfollowers °/ °"®y f
the Ptendop is called ardent
the Pteridophytean
supporters of
irts- S"'£.S
others.
- , , ^^1 helieve in the aquatic an-
The adherents of the secon bryophytes have
cestry of the bryophytes. Th y known as

.lew lhffi:r?.raTgn«^^^^ on^ the hasls of the following two

v
-in,1 ; • • ,, , -! 1
r-' ■ ;. • • .
Close similarity between the
. '
■ ■ ■"-
>-V.
. . . -.V* Ui;r • ■
- •: .
v- -i /v ■
jSin^P-rgb-^^^^
-0
nites and Horneophyton. strengthened
The kinship between the two cell walls, food
Ir'-, r*.

267
 269
GENERAL CONCLUSIONS
268 BOTANY FOR DEGREE STUDENTS—PART in : BRYOPHYTA

have evolved from the latter by progressive simplification or

reduction.
This v-ew ts supposed by he^se^^ h„„,ymhetic pigment (chlo.
Scott (1911) was the first to moot this idea on the basis of the m^^^irrnr^anloWll, —
presence of chlorophyll and plastids in the sporogonium of mosses,
liverworts and .^n^^oceros and also the presence of stomata of the
type met with in the primitive vascular plants on the sporogonium
rttV-rwll'aTd\?e%pe1*^ and number of flagella.
of 4?i//ioceros and the apophysis region of the capsule of mosses. He The above mentioned ^
interpreted these features as ancestral characters retained from the origin of the
the time when sporophyte was an independent plant (as in the
pteridophytes). The sporogonium of Anlhoceros which is to a
great extent independent of the parent thallus plant (gametophyto)
was believed to be the immediate descendant in all the bryophytes. '""'Tlebryophytes. on the other hand,.-'e ^rt
highly organized
complex 'Aantha
and elaborate the g., ; , protected sex organs,
germination m
Kashyap (1919) another ardent supporter of this hypothesis retention of '.A® a"Geminated spores and heteromorphic
considered the liverworts to have evolved from the Equisetalcs along
Marchantiales, Jungermanniales situ, production of wi ; jjow these changes actually
and Anthocerotales. It may be worthwhile to note that uptill then type of alternation of ^ bryophytes from the green algae
the Fsilophytales had not yet been known. occurred during the 7°^'™ Vhe species which constitmed
still remains an imstllve p disappeared. These
thp links between the two lia S delicate, thallose orga-
rh algae through the Psilophytales
the thebyorigin of the bryophytes
simplification. from
The resembl-
ance between the Anthoczros sporogonium and terminal sporotronium
ol Horneophjjton and the presence of stomata typical of the vascular
plants on the Anthoceros sporogonium and apophysis region of the
capsule ol mosses were the two chief features on which he based his
view point. The subsequent disappearance of stomata in other
members of the Anthocerotales was linked with the parasitic habit bility of a green =^'5^ lo a life on land.
of their sporogonia. thallus having become P hypothesis, has suggested
Smith, an ardent suPP"' ■ approach to the primitive ances-
To sum up the supporters of Pteridophytean hypothesis believe
that the bryophytes represent a degenerate evolutionary line of the from'whTcfetlved the bryophytes^ mmV
pter^ophytes. In their view it is difficult to explain these similari
ties between the two groups otherwise. However this hypothesis
alga of the family Chtt'='°P''7ff^Ihis
S;ro1drdrying amphibial plant
«°^Serional, photcs'ynthetic, posses-
sub-aenal
now sta^nds discarded. The opponents of this hypothesis point out ses the a simple filament."
K ''"y "fthe lost vascular system in the thallus originating .„i,vtes of multicellular sex organs
19«fwri,r "w® including the sturdy mosses. Mehra The occurrence in the ^ P difficult to explain. There are
water ferns Sat 1, T"' ° vascular plants including the ■,h their sterile jackets is, bmvev , chlorophyceae. The
rtave Lmme/™ of life since the ages known\ri parallels among the hvmg ^le sex organs,
„^^gansit
vLcu"r s~' of life but even then retain the "reLncTof a 7'0^ccn.e
protective sterile jacket
sterilejart^
Teems, is an
=■" adaptation
tt' to „ exposed to the drying effects
J^eabe«e.^-of.rvi.^
the blyophytesfvvhi°ch'^neces^tateTTh^"'^'^ flagellated sperms (in
least, at the time of ferdSation j n dtt^mlyUimbcramd^by^^^^^ numile o
ancestry. The development of a m e aquatic prefectly by the p a um
tonema as a juvenile stage in the life alga-like pro- Coleochaeie. hrvophytes is now acknovvledged by
phytes shows beyond doubt that tbp k some of the bryo- The algal ancestry o advocated by Smith (19 b)
in some algal form. It is a hinln . °^y?P^ytes had their origin all the botanists. The
stage of an organism resemblefSesS^^^^ the juvenile
 2.

271
general conclusions
270 BOTANY FOR DEGREE STUDENTS—PART III; BRYOPHYTA
(rii) Sporophyte attached throughout life to and dependent
the bryophytes arose from the algae and then gave rise to the sim for nutrition upon the parent gametophyte. nc r
ple pteridophytes (primitive vascular plants) is no longer consider
ed tenable. On the contrary the modern bryologists hold that the
Because °f the above-menti^^^^^^^^^^
ing plants (bryophytes) are ^
fhfotrlh'alS
which they resemble in
bryophytes represent a blind alley in evolution. Their viewpoint
is that to avoid increased competition in water a group of remote LS?trrrpec"'^heyare.fus separated from the division
ancestors of the green algae migrated to a subaerial habitat. These Thallophyta. bryophytes differ from the green
primitive amphibious plants developed multicellular sex organs as
an adaptation to a life on land. From this hypothetical group arose The features in which th y plants pteridophytes
both the bryophytes and the primitive vascular plants (pteridophy algae they with the pr included among the pterido-
tes) along two parallel lines. except point (»»)■ Bjit following important res-
phytes as they are very
The supporters of the algal origin differ among themselves pects besides others :
as regards the origin of the two main groups of bryophytes.
According to one school which support the polyphylletic origin the (') ^'''T'°'^mDhaseTL life cycle being the gameto-
(u) The dominant pnase m
three groups of the bryophytes namely liverworts, hornworts {An-
thoceros) and mosses represent three independent evolutionary lines, phyte and not the ^ bryophytes are placed in a
each having an independent origin from the transitional archego- It is for this reason intermediate bet-
niale forms which evolved from still earlier, different, hypothetical separate division pteridophyta.
green algal stocks that migrated to the land. The supporters of ween the Thallophyta organisms
the monophylletic origin advocate that the mosses are more pri
mitive than the other bryophytes. The latter have evolved from
the moss-like ancestors by simplification or reduction. According
to this view the mosses are nearer to the supposed algal ancestors
than the other bryophytes.
Systematic Position. The bryophytes are a group of inte
group is of a smgulai l ( of change in habit from a
resting plants. They resemble the green algae in certain respects
fandinhabit
life wateramong
to a lifeplants.
on landThiing it made^ possible jall subse-
the
and differ from them in others. The features in which they agree u' mrv of plant
with the green algae are : history P^SJ life. pijjnt
Pt is world. The sroup bryophyta
late some includes
of the steps this
(t) The simple, green, thalloid plant body. 'h"" riTiilt tm' tmr To
(n) Lack of vascular tissue and absence of roots,
(m) Predominant plant in the life cycle being the gameto- ^Sng'Tol^Tfrorthe aquiic an«rram -
phyte.
{iv) Autotrophic mode of nutrition.
(u) The retention of motility by the male gametes.
They differ from the green algae in the following respects :
/a\ 'I O 1 A Tw r A
(i) Terrestrial habit.
(M) Multicellular sex organs each with a jacket layer of sterile '°"'1?PriXe'type of conducting^rtrani
cells. The latter in addition func
{in) Embryo formation.
{iv) fion as Of "^T'muuTcellular sex organ^ It serves
typeof alterSr^ofgrnrtio^V"*^ ^ '^--ologous T RetentU of the aygote within the archeg
green algae but has no exact counterpart among aplant
douffeas itdevei
PJpo-rfr:;!^
P
SlcSlTtr embryo and affords protec-
tion from drymg-
within fhi ar'^he^gLTr^ development of the fuiure sporophyte
272 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA 273
GENERAL CONCLUSIONS

(vi) Thick-walled, wind disseminated spores.

Economic Importance. The bryophytes, m are of
It seems the nature in its quest for a suitable plant body for little economic importance. Some mosses provide food for her
a life on land tried two schemes. One of these is the upright leafy bivorous mammals, birds and other animals. On t le w o e none o
body habit of the Mosses. It favours photosynthesis as it presents tliem is directly important to man except a number of species of
greater photosynthetic surface to light. It, however, presents SDkaomim They grow and multiply enormously in water thus
difficulties in the way of fertilisation as the sex organs are elevated. fiH ng th ponds Ind lakes with its growth. The barren sheet of
The elevated position of the moss sporophyte is, no doubt, ideal for wale? is converted into a solid land. The peat has long been used
spore dispersal but how can there be a sporophyte without fertilisa as fuel
mei. ^ Dried
wneu up
i specimens
p of Splmjmm
packing are used
material as an absorbent
m upholstry and for
tion. The two physiological processes fertilisation and photosyn m surgical dressings, to increase the water
thesis have opposite requirements. The former is facilitated by
flat, thalloid body and the latter is favoured by a tall, upright body.
transhipment ''""S ^pes of soil, and as a means of
Hence the scheme of combining photosynthesis with fertilisation
retaining ? acidity required by certain decorative and
retaining high Flowers packed
as the functions of the gametophyte failed. It proved an unpro-
gressive evolutionary trend. This is the main reason why the
Mosses end blindly in the evolutionary line. They have given rise
=e::d^;rres'^t^ l?:sh L a considerable period.
to no new plants. Nor have they progressed beyond this stage.
The gametophyte plant in the mosses both morphologically and Oe.olopUaUy ^all Th:;
anatomically shows highest development of the vegetative body in soil P; fhe T'velopmont of the soil. The mosses
the bryophytes. But there can be no progress beyond this stage provide essential step
The two main factors which check further growth are :— grow on barren soils. y . pi^pts is impossible. In the
(t) Absence of root and vascular tissue. The rhizoids as dry rocks where S»owtl o decay forming humus. The
absorbing organs proved inefficient. course of time their ^ Eventually humus containing
(ii) Dependence upon water for fertilisation. It also hinders
further growth. forests grow on such soils.
Hence the plan of combining photosynthesis with fertilisa

whichThe mossesthe force of' a^Titrpr^tt"

water and are 11^0^
tion as functions of the gametophyte failed. It permitted no further
progress. diminish so firmly
In the second scheme nature tried the combination of photo
synthesis with spore dispersal as functions of the sporophyte.
^Icrtwted arotd^ md particles that there can be little erosion
Both the processes have the same requirements. They are favour even on a steep hill side.
ed by an upright, tali body. It thus proved an ideal combination
as it permits further progress. On the other hand fertilisation is
favoured by a flat, thalloid body. This combination we find in Origin and the ^ of'primary cdls.
Anthoceros. It has a flat, thalloid gametophyte ideal for fertilisa
tion. The gametophyte bears an erect, tall cylindrical sporophyte c a solid tissue. I tie ecu ^ them
with a well ventilated green tissue in the capsule wall for photo
synthesis. From such a sporophyte, some botanists believe, may solid tissue
function separate
as spore mofrom ea^c archesporium spores. So this
have arisen the sporophyte of the simplest and the primitive vascular solid tissue derivedtissue.
^irom this basis archesporium is defined
plants (ptcndophytes). the sporogenous porogenous tissue The origin and
Sfafeiiisahoft of alternation of generations. This important as the first generation^ of th P dffierent classes of
biological phenomenon is found in the life cycles of a few of the consequently Pf/^^^ay vary in different genera of the same
thallophytcs but is far from beinff fixed TK the Bryophytes. i he lai
heir rr 1
Ufe cycles • 1
are either .
quite similarnxed. ihe two sencrations in
or somewhat differem In order
the bryophytes, however, this phenomenon is stabilised It becomes
a constant leature of their life cvcles it Decomc
dividuais of the two generations dUTer ma A .ii •
cai and anatomical detaS markedly m their physioiogi-
 GENERAL CONCLUSIONS 2T6
274 BOTANY FOR DEGREE STUDENTS—PART III : BRYOPHYTA
Glasses and Origin Position Fate
Glasses and Origin Position Fate Orders
Orders

1. Hepaticopsida 2. Anthocerotopsida

(a) Marchantialea : (a) Anthocerotales:

Anthoceros Archesporium ari It is more super- It diSerentiates
ses from the fieial in position into the spore
innermost layer of than the Hepati- mother cells and
Tlicciii The entire endo- It is central in Practically all tho oinphithecium. copsida. It over- the pseudo-
thecium becomes position and forms the sporogenous It divides to form arches the tip of elaters.
the archesporiutn, the bulk of the cells function as 2-4 layers thick tho coluQiella and
and forms the spo- sporophyte except spore mother sporogenous tissue. lies between the
rogenous tissue. the single layered cells and form the central coluraella
wall. spores. A few at and the several
the periphery layers thick cap
degenerate to sule wall
form a nutritive
fluid. These are
3. Bryopslda (Musci)
called the nurse
cells.
(o) Sphagnidae—
The entire
Sphajnales: Archesporium as As in Anthoceros
sporogenous tissue
Sphagnum ill Anthoceros origi it is superficial in is devoted to
nates from the position. It is
Marchnntia The whole mass It fills the cavity Half the sporo inner layer of the dome-shaped and the formation of
of ondothecium of the capsule genous tissue amphitheciuin. By lies in the upper spores. Tho sporo
forms tho arches- within the single yields the spore division it forms 4 \part of « capsule genous tissue thus
va a
porium which divi- layered capsule mother cells and forms a coherent
layers thick sporo- overarching the tract of fertile
des and redivides" wall. tho other half conous tissue. top of the massive
to form tho sporo- gives rise to the central columello cells in the
genous tissue. elaters. A few within the capsule capsul^.
towards tbo top wall.
may remain
sterile and form
the apical cap. (6) Bryidae-
Tho archesporium It is superficial All the sporo
Funnriales genous cells are
Funaria originates from the and surrounds the fertile and form
outermost layer of columella like a the spores. The
the ondothecium. barrel. sporogenous tissue
(6) Jungermannialea It gives rise to the thus forms a
sporogonous'tissue, coherent tract of
Pellia Tho entire endo- Itfills'.tho entire A central mass 2 cells in thickness. fertile cells as in
thecium forms tho cavity, of tho cap of sporogenous Sphagnum.
archesporium. The sule within the two cells forms the
archesporium gives or more cell layers basal elatero-
rise to tho sporo thick capsule wall. phore. The rest
genous tissue. form tho spore
EVOLUtlON OF THE SPOROPHyTE
The sporophyte nf the brvophytes,appendages
at first, is aand
solid object
mother cells and
the elaters. is incap-
radial in obtains its nutrition wholly or partia y
able of to which it remains organically
Porella As in Pellia. As in Pellia,
from the parent game p y function is the production
There is attached throughout
and dispersal
ts
of spores.
onlyrUfferRntiated
a spherical spore
into
elaterophore. The .uid dispersal to an elaborate object differentiated intod
entire sporogenous producing case as ^
tisauo forms
the spore-mother foot and f Qt functions as an anchonng and absorbing
cells and the capsule
elaters
 277
GENERAL CONCLUSIONS

276 BOTANY FOR DEGREE STUDENTS-PART III: BBYOPHYTA half of ihe embryo which forms the spore
amphitheeium forms the single layered "Psule wall Towa
organ. The seta may either be small and constrietion-like
slender stalk. It helps in conduction and aids in spore ^ P . .
The capsule also varies in form. It may be spherical, ova , ^ ^
cal or pear-shaped, etc. It varies in its elaboration. The elabora
Biccia. The. arehes^orium ^eUs^ =X'f t^e rpmogr„oreells
tion, however, is internal rather than external. "nm ■ spore mother cells. Jhe
According to the complexity of structure the These sterile cells ^ elaters are hygroscopic,
of bryophytes may be arranged m a series between the their walls and become the e ■ ^ sporogenous cells
and tL most elaborate. The series starts with .^™P «'» They help in the scattering o p "
at the top may differeniiate into
epical cap. The capsule
^ ^ producing and spore
phyte of Kiccin runs through that of Marchantia, Pdha Anthoceros of Marchantia has ^ further in the progressive
Ld finally ends in the highly complex sporophyte of ^
seriated ihe sporophytes of the bryophytes suggest a possible Ime ol
evolution.
ttstsSi
(a) Theory o£ SteriUsation. Bower holds that the scries
runs in an upward direction. It illustrates a natural advance in the
"■takes W,..
no part and the seta is developed from
progressive elaboration and complexity of the sporophyte. Ih entire sporophyte
Lndamental principle upon which he formulated his argument the the epibasal halfincluding
(Fig. ^ periclinal divisions insurrounding
^ amphitheeium the embryonic
the
"progressive sterilisation of the potentially fertile cells (spomjenous capsule differentiate into an ^ the capsule 2 or
tissue):' Instead of forming spores and serving a propagaiyefun^^^^ inner endothecium. The ^ ^ ^pter layer develop radial
lion they remain Sterile. These sterile cells Y^P'^^Tutlnothes' more cell layers thick develop semi-lunar
such as nutrition, support, dehisccace, dispersah etc. This l^YPOthesis thickening bands rheciura in whole becomes the arches-
of Bower is called the theory of sterilisation It attempts to ex thickening bands. The end^ archesporial cells divide
plain the evolution of the sporophyte in the bryophytes on the basis norium as in Rtccia and pssue Unlike Riccta and
of progressive elaboration. The ardent supporters of this theory are STedividc to form ^
Cavers, Campbell and Smith. Let us apply Bowers theory o iMarchantia a central mass of sporog_^^ Tht
sterilisation to the individual sporophytes of this group and see il it capsule, remains stciile. F become elater-hkc.
stands the. test. develop spiral ddchemngs on dm at their lower ends m
central solid of elater hk ^
I. Riccia Sporophyte. The zygote enters upon active sc^
mentation to form a spherical mass of 23—30 undiffcrenliatcd cc s
(Fig. 4T2 A—E). Periclinal segmentation at this stage defines an
inner mass of endothecium from an outer single layered amphitne-
cium (Fig. 4-12 D and E). The amphitheciam forms the single
layered capsule wall. The endothecium which occupies the central
position, as a rule, in whole becomes the.archesporium. The latter
divides and redivides to form the sporogenous tissue. Practically all
the sporogenous cells are fertile. However a few at the periphery
undergo degeneration to form a nutritive fluid (Fig. 413 D). These
are the nurse cells. There is a large output of spores and no or sterilisation at the centre. ^ illustrates a
very little sterilisation of the potentially fertile cells. A Anthoceros '"sterilisation of the poten-
The sporophyte of Riccia is thus the simplest among tfi® n further than
sieplurinc m IS ct)mptete sterilisation
^ at the column
centre,
bryophytes and has the least amount of sterile tissue. The entire iiaily fertile , remains sterile. It Cense-
embryo forms the spore producing capsule. There is no foot and
no seta (Fig. 4 15). It is simply a spore producing organ without
esrapf function- It does not dehi.sce to allow the spores to
derived 5'2!). Half of the embryo
 278 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA GENERAL CONCLUSIONS 279

and surrounds it. The sporogenous cells become differentiated into (ii) Reduction of the green photosynthetic tissue in the capsule
spore mother cells and pseudoelaters. The archesporium in wall.
Anthoceros is thus extremely reduced. It forms only a small narrow
part of the capsule. The outer amphithecium gives rise to the several (m) Associated with the above is the disappearance of stomata
layers thick capsule wall. The capsule wall develops a well-ventilated and intercellular spaces.
photosynthetic tissue protected by the epidermis containing perfect («y) Decrease in the thickness of the capsule wall along with the
stomata like those of the vascular plants. disappearance of thickening of all types of wall material from the
walls of the cells.
5. Funaria Sporophyte (Fig. 12T5). Major portion of the
^bryo sporophyte remains sterile to form the foot and the seta. (u) Side by side with the above changes is the gradual elimi
The embryonic capsule region becomes differentiated into an inner nation of the seta and subsequently the disappearance oi the loot. ^
central column of endothecium surrotmded by many layered amphi {vi) All these changes are accompanied by the Progresswe
thecium. The amphithecium becomes differentiated into the increase in the fertiliiy of the sporogenous cells. This change elim
epidermis, the photosynthetic tissue of the capsule wall and the outer nates the presence of sterile cells and elaters in the capsule.
spore sac. Excepting the superficial layer there is complete sterilisa
tion of the endothecium to form the central columella which is Evidence from comparative morphology and experimental
continuous right up to the top of the capsule. The archesporium genetics support the view that the simple sporophyte of Rtcciap an
arises from the outermost layer of the endothecium. It is thus advanced but a reduced structure.
extremely reduced and consists of a single layer of cells. Besides its
fertility is limited above in the operculum region and below in the
apophysis region. Consequently the archesporium becomes barrel,
shaped. It is confined to the theca region only. The barrel is open"
at both the ends. The sterilisation of archesporium towards the
base results m the increase in size of the photosynthetic tissue in the
apophysis region. The arrest of fertility towards the top ac cording
to Bower may be correlated with the specialisation of this region for
spore distribution. This view is supported by the presence of peri-
stome, operculum and annulijs in this region, In fact the arches-
porium stops short just at the ba.se of the anniiius and the peristome
apparatus of the Moss capsule is mechani-
reducedarrhp!^"'"''' the spores. Consequently the

theevoE'o*th7spt* ^ m the upward direction.

It is appealing. 'P^^ophyte m this group
of thought l^"by*K°shvao'^C^
that so%eriated the evolution
however, an opposite school
hold
downward direction. The series
of retrogressive evolution. There is amn^^ "f example
rather than progressive elaboration of the T reduction
The reduction is accompanied by simniinr^ ° group,
the sporophyte in the serie.':. On'the b-isk structure of
^,«yteofi2.WawilIbecons dered^ ^^^h y evolved or advanced
though reduced as a result of progressive
ficant steps m the reduction series ^re The signi-
(^) Simplification of the dehiscence apparatus.
 COMPARISON BETWEEN THE GAMETOPHYTES OF
Porclla
Riccia Marcbantia Fellia or Anthoccros Funaria
te
Madotheca o

Habitat Most of the It is more ter It occurs com It grows in It is terrestrial It grows in
g
Species are terre restrial in habit monly on damp damp, shady
and grows on soil dense patches in
strial. They grow than Riccia. It soil by the sides places on logs, generally in very moist situations
on moist soil. A grows in moist, of streams, spr treo trunks, bran- moist, shady pla such as damp o
few are aquatic. situations such ings, wells, some ches, wot rocks ces on tho sides soil, on tho walls, 50
The latter grow as damp earth, times actually etc. It often of slopes, ditches on moist rocks O
floating or sub- walls of wells, under water and occurs in largo and in moist and on lands w
merged in the along streams rarely on moist mats in the hollows among burnt by fi re. o
50
water of quiet and wet rocks. rocks. tropics. rocks. The plants H
ponds and lakes. No aquatic spe grow in dense tel
cies has so far clusters. CO
been reported. 1-3
Cl
tJ
TEallas The plant body The plant body Tho plant body Tho plant body Tho plant body The plant body W
t) External is a small, green as in Riccia, is a as in Riccia, and is generally is a simple, some is leafy, green K
Features thallus. It is green, prostrate, Marchantia is a epiphytic in what fleshy dor upright and 1-3
CO
dorsiventral and dichotomously br simple, dorsiven habit. The thal siventral, green, ridial in sym
grows prostrate anclied thallxis, tral thallus. It is lus is prostrate non-leafy thallus metry. It con-
on tbo ground. fleshy in texture. thin, fl at, greon and dorsiventral. as in Riccia, si.sts of an erect,
It branches free Bach thallus lobe and lobecl. The In external ap Marchantia and sparinglj' branch
ly by di^ihotomy has a distinct upper surface is pearance it is, Pcllia. The thalU ed central axis
and frequently midrib but no smooth. The however, more are deeply lobed, 12—20 mm. in
takes on a circu dorsal furrow. It margin is sinuous elaborate than Tho lobes over hoiiriit bearing
lar form. Each ends in a termi and irregularly Riccia, Pcllia and lap. Branching is green leaf-like td
thallus lobe has nal notch. The lobed. The lobes Marchantia boing diohotomous but expansions. The 50
loaves are spiral kJ
a distinct midrib, thallus oi March' often overlap leafy. It consists due to unequal o
antia is, however, One another. Tho of a more or loss growth of the two ly arranged.
a dorsal median
branching is di- branched central parts of dichoto They are sessile, W
groove or furrow more terrestrial
oblong-ovato with kJ
and ends in a in habit. It has? chotomous. Each axis with my the thallus
leaves
a smooth margin, H3
terminal notch. a definitely large lobe has a slight arranged in three usually becomes >
ly thickened mid- rows. There are irregular in out pointed apex and
The areolao and size, broader and
the gemma cups thicker lobes, rib and a termi two rows of line. The margin distinct midrib.
lateral and sjnn- of tho lobes is The branching is
arc ahsent. more prominent, nal notch. Tho
median dorsal metric loaves on variable. It may monopodial. The
expanded mid-
tho upper surface branches are
■cVb and upper

surface marked groove character of the axis. The be entire, never axillary.
by rhomboidal istic of the thallus third TOW of toothed, folded They arise from
areas (areolae) of Riccia and tho reduced leaves is or fringed. There below tho loaves.
each with a rhomboidal areas on the lower side is neither any In its extei'iial
distinct central (areolae) charac of the axis. The midrib nor a appearance the
yiorc. Presence of teristic of the dorsal leaves are dorsal furrow. moss gamoto-
gemma cups is thallus of Mar- hilobcd aud in- The upper sur pliyto is tho most
another distin ch^-nlia are cuboasly arrang face is smooth highly dovolopod
guishing feature. absent. ed. They are there being no of all the bryo-
without a midrib rhomboidal areas phytes.
H Tho branching characteristic of
of the axis tho thallus of
is monopodial. Marchantia. The
Diohotomous. thallus is usually B
brandling is Inhabited by !25
B
absent. Nosloc colonies 50
seen as small,
dark, blue green
species or spots Q
O
in surface view.
«
O

From the under The scales are As in Pellia and The plant being B
Prom tho ventral From tho ventral upright there is a
surface . of the surface of the surface of the absent in Porella, Porella the scales CO
The unicellular and the tuber- no distinction
thallus arise the thallus arise both thallus arise only into the upper
O
rhizoids are also culato rhizoids a
scales and the the scales and the rliizoids The and the lower
of one kind only. are absent oc CO
rbizoids. The the rbizoids. The scales are absent
They are the ventral sur- ! surfaces. The
scales arise along scales are arrang in Pcllia. The rhizoids, which
smooth-walled face of tho
tho margin and ed in two or three rbizoids are also thallus. It is are tho attaching
project boyond it. rows on either of smooth- and unbranched.
They arise from fi xed to tho 8\ib- organs, arise
In Borae species side of midrib. walled typo. Tho
tuborculate rbi the lower surface stratum by un from the brown,
they are absent Tbey are not branched smooth naked, basal part
and in still others marginal. The zoids are lacking. of the stem from
the basal por walled rhizoids of tho stem.
rudimentary. The! rbizoids as in The unicellular,
tions of tho ven only. They are multi-
rbizoids are nni- 1 Riccia serve to unbranchcd. smo
tral loaves. They cellular and
cellular and un- 1 attach tho thallus oth walled rbi branched. Tho
serve to attach
branched. They 1 to tho substratum zoids secure
tho tliallus to septa between
anchor tho 1 and are of tho the thallus to the colls are
the Bubstratmn. tho substratum.
thallus to the two usual kinds oblique and at
I The tuborculate
I substratum and ' smooth-wallcd
 Porella
Riccja Marchantia PeUia or Anthoceros Funaria
Madotheca W
o
are usually of and tuberculate. rhizoids are long intervals. >
two types, The ventral scale! lacking. There are no !2!
smooth-walled help to retain scales. Nor is
and tuberculate. moisture below there any dis
the thallus which tinction into O
can be absorbed smooth-walled 50
by the rhizoids. and tuberculate O
This enables rhizoids. H
Marchantia to
grow in drier
a
habitats as com
pared with cn
Riccia, H
a
d
d
(b) Anatomy Internally the As in Riccia Internally the Internally the Internally the Internally as a
thallus is several internally the thallus is simj>le young central thallus is very well the so called CO
cells thick. The thallus is several as compared to axis is very simple in struc moss stem shows
colls are diiferen-
tiated and
cells thick. These
oolls are better
that of Riccia or simple in struc ture. It is several certain amount k
Marchantia. It ture. It is made cells thick in the of tissue differ
arranged in two differentiated is several layers up of perfectly median portion. entiation. Tho
distinct regions, and arranged in of cells thick in uniform, green The cells, how colls nro arrang
tho upper, green three distinct the median por parenchyma cells. ever, show no or ed in throe dis a
photosynthetic regions instead tion but there is It shows no tinct
littio tissue diff regions :
region and tho of two as in no or little tissue difforontiation of erentiation. Hor epidermis,cortex w
lower storage Riccia. These difterentiation. tissues. are they arrang and central Ed
region. are :— The cells are not ed in photo cylinder.
o
arranged in any synthetic and
distinct regions storage regions. m
such as the All the cells are k!
upper photo- uniform, com
>
synthetic and pactly arranged
the lower and parenchy-

storage regions. matous except

the surface layer.

(i) There is no (i) Epidermal i) There is no (i) The epi (i) The cells of {i) Epidermis. It
well-defined epi region. It con well developed dermal layer is tho surface layer is tho surface
dermal layer. sists of a well- and well defined not well defined. are smaller but layer. In the
It is represented 1 defined upper upper epidermis. Nor is there any not cuticularized mature portions
by the terminal, 1 epidermis with Of course the trace of a cen or othcrwiso of tho stem tho
colourless colls 1 true, borrel- surface layer is tral strand. modified as epi epidermal cells
of the vertical 1 shaped pnres. often referred to dermal cells. are thick-walled
rows of green 1 The epidermal as epidermis but Hence there is and luck chloro-
cells. Together j cells have slight- tho epidermal no organised phists. There are
the hyaline ly thickened cells are not in epidermis. neither any air
pores nor O
cells of the i walls. They are any way differ B
neighbouring 1 protective in ent from the ; stomata.
green filaments 1 function and other cells of the H
form an ill- \ tend to check thallus. EO
>
defined, dis- 1 evaporation from
continuous upper 1 tho underlying n
epidermis, one 1 tissue over which o
cell in thiclcness. I the epidermis !z;
forms a single-
layered roof. gM
air (li) In older (ii) There are no (ii) Cortex. It is
{ii) The photo- (ii) Photoaynthe' (n) The several cell layers
o
a
synthetic region tic Region. It lios pores and air portions of tho chambers, and CO
oonsists of a beneath the chambers charac stem, however, air pores. In fact thick and con
there is differen in its structure sists of large,
loose, green upper epidermis teristic of Mar thin-wulled cells.
tissue. Tho greon tiation into tho the Anthoceros
^i and consists of chantia arc
thallus approaches Tliey are com
colls rich in a chlorophyll absent. The outer cortical
<
and inner that of Pellia as pactly arranged
chloroploBts are bearing tissue entire thallus
and in young
arranged in with air cham consists of uni medullary re it does in the
• t X gions. Tho cor absence of scales stems contain
vertical rows bers. Tho cham form tissue of chloroplasts. Tho
1 usually with bers are arrang chlorophyll bear- tex consists of and tuberculate
rhizoids. There mature portions
I narrow slits ed in a single ing polyhedral, small cells with of the stem have
between them. horizontal layer parenchyma cells slightly thicken are, however,
ed walls. differences in the cortex diff
The empty air- and are of fairly arranged com
detail. erentiated into
chambers open regular size. pactly. The cells
Tho medulla is an outor thick-
to the exterior Ilach chamber of tho upper
 Porella
RJccia Marchantia Pellia or
Anthoceros Funaria
Madotheca
through air-pores is separated layers contain W
which are simpjy from its neigh composed of
abundant chloro- walled cortex o
iuterceliu/ar bours by parti comparatively
plasts. Starch larger cells with and an inner
spaces bouuclod tions of green grains occur in thm-walled cor
by 4—8 epidor- cells 3 or 4 colls
thinner walls.
«il the cells. tex. There ore
inal colls. Tho height. It no air spaces.
pJiotosynthetic opens to tho o
filaments and oxierior througli W
tho true poros a centrally placed] o
characteristic of barrel-shnpcd,
Marchantia aro I epidermal air 0
absent in Riccia. I pore. From tho w
floor of eacli w
chamber arise CO
short, simple, or
branched fila
ments
rich in
of cell.s
chloro-
1
SI
plasts.
I
(Hi) The lowci (iu) Storage Re (m) There is no hi
storage region is gion. It is the
(iiV) Aii tho cells {Hi) Central >
distinct storage have a single Cylinder. It is
colourless and lowermost re region. large chloroplast
several layers gion of the occupies tho core
each with a of the stem and
thick. It consists thallus whoro
pyronoid in the consists of vorti-
of closely packed, food and water centre—an un cally elongated, C3
imdiffcrentiated, are stored. It is usual feature. SO
thin-walled colls.
paronchymatous ■more extensive Some ceils con They are narrow
cells which may and many layers o
tain mucilage. and compactly T)
contain starch. A thick. It is com The distinction arranged. It n
few cells contain pact and colour of tho thallus provides mecha ><
oil. Those cells less. The cells ef into the ventral nical support
thus serve for the upper layers storage and tho to a contain

food and water of this region dorsal photosyn- ] e xtent and func
storage. There may contain few thetic region as I tions in conduc
or no chloro- is characteristic tion.
are no mucilage
cells. plasts. It con of tho Marchan-
sists of relatively The leaf consists tiaceao is lacking. T he ao-called
large, thin- of a plate of In some species leaf consists of a
woUed, colour cells one-coll of Anthoceros tho single layer of
less, parenchyma layer thick. The superficial cells on colls rich in
cells. Most of ceils are rich in the lowor surface chloroplasts. The
them contain chloroplasts and of the thallus midrib is several
starch. Isolated are polygonal in separate from each cells thick. It
cells in this re form. There is other to form in- ' contains a
gion may con no midrib. Tho tercollular spaces ! narrow strand of
tain a single oil axis is cylindri which are filled j thin-wallod,
with rnucilago- i elongated cells.
body or be filled
with mucilage.
cal in outUno.
These mucilago \ §
55
Tho former are pits often deepen ;
called oil cells ond broaden to
and tho latter form largo, a
rounded cavities. o
mucilage cells.
They are filled , a
with soft, muci
c!
lage and open on CO
M
tho ventral sur
O
face by stoma-
liko clefts. Very CO
often Nostoc
finds its way
into theso inter
cellular, Bpaces
through the slit
like or circular
openings of the
mucilago clefts.
Once lodged
within, the alga
multiplies exten
sively in the
 Porella
Riccia Marcbantia Pellia An^orceros
or
Funaria
Madotheca
K
O
intercellular
•-a
spaces. The >
latter by the
disorganisation
of the "neigh
bouring cells O
broaden into con 50
spicuous cavities
inhabited by the §
O
alga, 50
H
(iv) There is no H
(iv) The lower (iv) The lowor A single distinct. lower 03
most eelJs of this epidermis is dis layer of rogularly epidermis. »-3
region are a
tinct and well arranged cells on o
smaller in size developed. It is the lower sijrfftco H
and more regu continuous. of the tiuillus is
larly arranged referred to as CD
to form the the lower
lower epidermis. epidermis.
Apical growth I Growth in As in Biccia tlio Growth takes The thallus CampboH, Bower It takas place
length of the growing point place by the grows by ineiins and othcr.s report by means of a
thallus takes lies at the activity of a of a single pyra the occurronce of single pyramidal
place by means bottom of a single large midal apical a single four- apica cell at
of a group of terminal notch apical cell. It cell with throe sided pyranii<Ial stem tip. It
apical initials of each thallus lies in a notch cutting faces. It apical cell at has three cutting
constituting the lobe. It consists at the forward lies at the tip of the growing faces.
growing point. of a group of end of each the main axis point which lies
It lies in the apical initials. thallus lobe. and each of the in a shallow
deeply emargi- By the activity brandies. depression at the
nate apex of of theao the anterior end of
each lobe. thallus grows in each thallus lobe.
length. Leitgcb recog

nized several
marginal colls
of equal rank.
Mehra and
Handoo confirm
Leitgeb's obser
vation. They
hold that in A.
erectus and A.
himalayenaia there
is a group of cells
which initiate
growth. Thoy
are protected by
mucilage. O
H
2
Juvenile Stage \ There is no The same The same as in The adult leafy There is no The upright, M
juvenile stage or Riccia. Riccia and thallus is preced juvenile stage as leafy gameto- Ed
protonema. The Marchantia. ed by a juvenile I in Riccia, Mar- phyto is preceded >
thalloid gameto- stage or proto chantia and by the juvenile
phyte is formed nema which is Pellia. or protonema o

directly from the of a simple type- stage which is a o

It is multi- green branched 2
germinating spore. o
cellular and system of fila f
develops one or ments. d
CD
hH
more rhizoide
O
prior to the 2
difforeotiation of CD

the apical coll of

the leafy thallus
(gametophytoh
 COMPARISON BETWEEN THE SEX ORGANS OF

Porella Anthoceros Funaria

Rfccia Marcbantia Pellia

{)) All species (i) Soino species (i) Funaria

Distribution (i) Mm^y spocios ('} Without (t) Some species hygromelrica is
are monoecious, are reoortod to are monoecious
of sez organs arc monoecious exception nh and others monoecious and
species of Mar- others dioecious. bo dioecious.
but a Cow are dioecious. The autoicous. Some
dioecious. chantia are The monoecious
monocious spe mosses are
dioecious. species arc monoecious and
distinctly pro- cies aro reported
to be protand- pai'iocous others
tandrous. monoecious and
rous.
synoicous. Dioe
cious mosses are
also common.

{ii) The male {ii) In the (it) Tho male

(it) In tbe dio (ii) Tiio male (•//) The malo dioecious species and tho female
ecious species and the female and tho female and female
plants are tho male and tho branches do not
the two kinds of thalli aro similar. thalli in tho exhibit dimor
thaIJi are similar dioecious species dimorphic, the female thalli aro
former being similar. phism.
but ill B. hiina- aro similar.
laycneis the male decidedly smal
plants aro ler than tho
latter.
smaller than the
females.

lit) Tho sex {Hi) Tho SOX (tit) Tho SOX

[Hi) The sex [iii) ■ They are (m) The sex organs are borno organs aro not organs occur at
organs lie in the homo on special organs are deve borne on any the tips of bran
dorsal furrow on sexual upright loped on tho on specialised
sexual branches. specialised sex ches which are
the upper sur branciios of the upper surface nob in any way
of the thallus ual branches.
face of the thallus called specialised.
the gameto- and nob on any
thallus and not
on any special pUores, which special sexual
are torminal in ■branches.
sexual hranchea.
position on the
thaWuB lobes.

■TX., Tho antha-\ Tho anthe- Ui") Tho antho-

OTgans ore deve to the scattered i ridia occur ridia occur ridia occur ridia occur in
loped in lines arrangement in irregularly scat singly in the scattered singly a cluster at the
back from the Riccia, the sex tered in two or axils of closely or in groups apex of the male
apex, in acre- organs aro deve more rows in the imbricated, light immersed be branch which
petal order. loped in localized broad midrib green bracts neath tho upper is the main
areas called the region. on tho antheri- surface of tho shoot. Inter
receptacles whi dial branches thallus. mingled with
ch are elevated, which stand at the anthoridia
later on, on long right angles to axe found the
stalks. the main axis. paraphysie.

(tJ) Tho arche- (V) The arche- (v) The arche- (v) Tho archo-
gonia occur in gonia occur in a gonia occur gonia are borne
a cluster close to terminal cluster singly and not in a cluster at
the growing on the female in clusters em the apex of tho
apex on a ridge branches which bedded in the female branch
of tissue called are very short upper surface of which arises as a
the receptacle. and bear 2 or the thallus. lateral out
The receptacle 3 leaves only. growth from the
remains sessile. base of tho male
shoot. Inter
mingled with
the archogonia
are found tho
paraphyses.
Position and (i) Botb (anthc- (i) Tho anthe- (t) Tho anthe- (t) The antho (t) The anthe-
origin of sex ridia and arche- ridia lie embed ridia lie embed ridia occur ridia occur (t) The antho
organs gonia) lie ded on tho ded on the upper singly projecting singly or in ridia occur in a
immersed, each upper surface surface of the in the axils of groups inside cluster pro
I in a separate of the male thallus, each small bracts on closed antheri- jecting at tho tip
1 cavity or I receptacle, each in its own pit the antheridial dial chambers on of the male
I 1 chamber on the I in a pit which which opens by branches. the upper sur shoot.
dorsal surface of 1 opens by a pore a pore on the face of tho
the thallus. on the surface of upper surface of thallus.
the disc. the thallus

(it) The arohe- (it) The arche- (tt) The archo- {ii) The arche- (it) The archo
gonia aro pro- gonia occur in a gonia occur pro- -gonia oocur gonia occur pro-
 Riccia Marchantia Pellia Porella Anthoceros Funaria

jecting. In the cluster projecting jecting in a

young receptacle on the upper singly sunk jecting in a W
cluster at the deep in the cluster at the o
they are borne surface of tho apex of the
on the upper thalhis.
fleshy thallus on apex of the s
female brancli. its upper sur female branch. is!
surface with
face with the *<
their necks
directed up tip projecting
out.
wards. They are
not simk in
§
separate pita. In g
the mature
receptacle they S
hang downwards §
H
from tho under- 00
surface of the H
disc with their d
necks directed t!
H
downwards. 2!-

{iii) Tiie sex j (iii) Tho sex (iii) Tho sex (iii) Tho sex (iii) Tho anthe- (iii) Both the
CO

organs are organs are organs are organs are ridia are endo sex organs are
exogenous in exogenous in exogenous in exogenous in genous in origin exogenous in
origin. origin. origin. origin. whereas" arohe- origin.
gonia are M
W
exogenous.
(a) Antheridiaj (i) The mature (») The mature (i) The mature (i) The mature (i) The mature (i) The mature W
antheridiura con antheridium is antheridium as antheri.-lium of antheridium is antheridium is
sists of an ovoid essentially simi in Rhccia and Porella has a orange coloured. club-shaped and
or pear-shaped lar to that Mar^anlia con globose or The club-shaped O
orange coloured.
body attached at of Riccia. It has sists of a body globular body or pouch-like It IS
its broader end
shortly K
an ovoid body and a stalk. The elevated on a body is raised on stalked. The
to the bottom of supported on a stalk is short but long slender a long multicel stalk is multi- >
the antheiidial short, multicel- multicellular. It stalk. The stalk lular, four rowed cellular.

chamber by a i lular stalk. The \ arises from the consists of two stalk arising
short, multicellu- ) latter arises from. 1 bottom of a rows of cells. from tho floor
lar stalk. i tho bottom of I round flask- of a roofed over
I the antheridial shaped antheri- chamber.
, chamber. dial chamber.

(ii) Tho body of (ii) The body of (ti) The (ii) The body (ii) The body ef (ii) The body of
the antheridmin the antheridium globular or has a jacket layer tho antheridium the antheridium
is made up of a has a jacket globose body of 2-3 cells thick has a single consists of a
central mass of layer one cell in the antheridium towards tho baso. layered jacket jacket layer one
numerous small, thiclcness. It has a jacket one It encloses a forming tlio wall. cell in thickness.
internal cells encloses a mass cell-layor in mass of colour Tho jacket, cells The cells con
protected and of androcytes. thickness. Tho less internal cells in some species tain chloroplasts.
covered by a Each androeyte cells . contain which produce ere rectangular The jacket has a
O
H
layer of larger, gives rise to chloroplasts. The the sperms. The and arranged in cap of one or two
sterile cells form a biflagellate jacket layer ripe antheridium four tiers. The large cejls at its a
ing tho jacket sperm. I encloses a mass is reported to be cells of the upper distal end.
or antheridial of androcytes. almost pure tier constitute Within the
wall. Tho inter Each androeyte white. The the dehisconco jacket is a mass o
nal cells of the metamorphosis antheridiura thus cap. Within the of sperm produc o
last cell genera into a biflagel differs from wall is a mass of ing cells. %
tion produce late sperm. those of Riccia, androcytes which
the biflagellate Marchantia and produce the
sperms. Pellia in its large sperms. The
size, long stalk, roof of the anthe o
jacket 2-3 cells ridial chamber %
cc
thick at tho base bursts open
and no antheri when tho sperms
dial chamber. are ripe.
(iii) Each (iii) The sperm (ni) The sperm (Hi) The sperm (iii) The libera (iii) The libera
sperm is a has a narrowly has a spirally is a slender ted sperm has a ted sperms are
minute, slender, curved or rod- coiled body with structure. It has small, elender minute, biflagel
curved slructure like body furnish two flagella atta a body consisting linear or slightly late spirally
furnished with a ed with two ched at its for of about 2 com curved body coiled structures.
pair of flagella apical flagella. ward narrow plete coils. A furnished with
at its anterior end. The sperms, small, vesicle is two flagolla at
end. of Pellia are attaohed at its its forward end.
 Marchantia Pellxa Forella Anthoceros Funaria
Riccia

W
The flagella are o
larger than those posterior end
of liiccia and The two fiogella almost equal in >
Marchantia and are somewhat length. S!
have more coils. longer than the *<
body and aro in
serted at the poin o
tod anterior end. 50
0
(b) Arcbegonia The somewhat In form and The archego The .mature In structure The archegonium
is typically flask- o
flask-shaped ar- structure the nium of Pellia archegonium is the archegonium
chegonium con archegonia of is typically flask- more or less a of Anlhoceros shaped. §
sists of an en- Marchantia are shaped and c^'Iindrical object differs from that s
Jarged, basal similar to those essentially simi of the bryopbytes 00

venter and a of Riccia. TJio lar to that of and resembles

a
JoDg slender only differences Biccia and Afar- that of certain 0
neck. The venter are r— chantia except ; Ptoridophytes:— H
is usually direct «
It is supported H
ly attached to 1. Presence of a 1. It is borne on It is seated on It has no stalk CO
the thaJius there short but distinct a short but stout, a stalk. The neck and is embedded on along, stout 1
being no visible stalk which massive stalk. consists of 5 in the tissue of ly developed
stalk which >
stalk. TJie neck attaches the vertical rows of the tbnllus.
consists of a venter to the 2. The neck cells as in PclUa. attaches
the tip
it to
of the
3
narrow, vertical receptacle. consists of 0 Tlioy enclose the
neck canal filled longitudinal rows neck canoL female branch.
2. Sb'ghtJy longer of neck cells It piejects above
by a row of four
neck canal .cells neck. instead of C The neck canal the adjacent td
surrounded by a characteristics of filled with vegetative tissue.
layer of neck 3. Greater num Marchantiales. usually 8 broad O
neck canal cells. There is a The neolc is
cells arranged in ber of neck hj
canal cells which 3. The number The venter is but control row of long and twis W
six longitudinal 4-5 neck canal ted. There are k5
rows. The tip of varies from 4-6. of neck canal little enlarged.
cells varies from colls but there is numerous {8 to H
Ibe neck is made 10) neck canal >
4. Hang down- 6—8. As in Pellia tliQ no jacket of
up of four large sterile neck cells ceils.
ventral wall con-
cap ov \\d celte. "wards from the

The distal por under surface of 4. The neck is sists of 2-laycrs around it except
.•* •- tion of the neck the receptacle. not sharply of cells. the uppermost
projects above marked off from tier of neck cells
the venter. which form the
the surface of
the thallus into tip and project
the dorsal furrow.' above the sur
face of the
thallus.

5. The venter The venter con The ventral waU

The venter 5. The same as

consists of a Riccia. wall consists of tains the egg is two cell layers
two layers of and the ventral thick. The venter
layer of jacket cavity contains
cells enclosing cells instead of , canal coll but O
the lower larger one characteris there is no ven the egg and the t?d
tic of Riccia and ter wall of ventral canal 52!
egg cell and an H
Marchantia. It sterile colls eel! above it.
upper smaller 50
encloses the egg around them. >
ventral canal
cell. and the ventral
canal cells. Some o
o
times the lower !2j
portion of the O
nock also becomes
two coll layers d
W!
1—I
6. Presence of a thick.
Protected by o
the surrounding collar-like struc
ture at the base The perigynium Protected by The protection The protective' CO
arohegonial cham sheaths in the
of the venter of is absent. The the surrounding to the axial row
ber. The protec form of peri
tive sheaths like each archegon involucre which penchaotial br of cells is affor
is homologous to acts, the upper ded by the ad gynium and
the perigonium ium. It is
the perigynium. the perichaetium ones forming the jacent thallus perichaotium are
and perichaetium tissue. The pro- absent.
are absent. Presence of a of Marchantia perianth and
the lower ones toctivo invest
two lipped surrounds and
curtain-like peri protects the constituting the ments such as
chaetium with arohegonial clus involucre. the perigynium
fimbriated mar ter. and the perichae
gins around each tium aro locking.
series of arche
gonia.
 Net5MP
MPARISON BETWEEN THE SPOROPHYTES OF
<o
Riccia archantia Pellia Anthoccros Sphagnum Funaria

te
SPORGPHYTE o
(i) It is far more H
(i) The sporophyte (i) It is larger than (i) It is more
of Riccia is tlie
(i) It is larger (i) It is much elaborate both
and more complex elaborate than those and more complex simpler than that
simplest known than that of Riccia. of Riccia, and externally as well
than those of of the other mosses. as internally than
ompng the bryo- Marchantia. Riccia Marchantia
phytes. In certain features that of any other o
and Pellia. The it resembles that bryophyto. EC
elaboration of Anthoceros and
internal. in a few others g
that of Funaria.
{ii) It consists of (ii) It is differen
§
(ii) It consists of (H) It is differen (fi) It consists of (ii) The sjioro-
a spore sac or tiated into foot, a foot, a long seta tiated into a foot a foot and a cap
capsule only. TJiero short seta and
phyte consists of a 02
and a capsule. a constrictiou-Iike sule. Between the foot, a long slender
is no foot and no capsule. intermediate zone two is a constric- seta and a C
seta. O
and a capsule. tion-liko structure capsule. fej
There is no seta. apparently resem 5;
bling that of
Anthoceros. The i
seta is absent.
(Hi) It lies com (iii) It hangs freely (iii) It arises (Hi) Ariso in (iii) Arises EC
from {Hi) It is situated
pletely sunk in the from the under from the dorsal clusters from the the apex of the at the distal end of
tissue of the thallus surface of the surface of the dorsal surface of female branch the female branch.
and has no pro female receptacle thallus sniTounded the thallus each elevated by a There is no pseudo-
tective sheaths surrounded by the by tlio involucre at surrounded at the leafless, staik-Iiko podium. The pro
around it except perigyniam and the base. There is base by a tubular pscudopodium. Tlie tective sheaths are
Cc
thft calyptra. the perichaetium. EC
no perigynium. involucre only. protective sheaths lacking. k!
are absent. O

(iv) The foot is (io) The foot is (iv) It is conical (iv) It is a rounded, s
(tv) The foot is (iv) It is a small, k;
lacking. well developed and with its edges pro bulbous structure well-developod and poorly developed
broad. It functions jecting upwards keeping the massive as in dagger-like conical >
as an attaching forming a collar- long capsule well- Anlltoccroa but object which func-

If

and absorbing like structure at j anchored upon and develops no rhi- tions both as an.
organ. tho base of the 1 attached to the zoid-like outgrowths.
seta. Functions
attaching and
thallus, often Functions both in absorbing organ.
both in absorption develops rhizoid- absorption and
and attachment. like outgrowths attachment.
I increasing its
1 absorptive surface.
Cu) The seta is (v) The short seta
absent. elongates
(u) The
rapidly long, of pure white
seta is 1 (v) The seta is (v) The seta is (V) The long,
absent. Its place absent. The con- slender seta ele
but slightly,simply, colour, almost is taken up by the
to push the mature striotion-liko inter vates tho capsule
transparent. It constriction-like in mediate zone
capsule through and far above the
elongates rapidly termediate zone which apparently
beyond the sur after the ripening surrounding peri-
which is meriste- resembles that of chaotial leaves and §
rounding protective of the spores to
sheaths. elevate the capsule
matic. This inter
calary meristem
Anthoceros lacks helps in tho dis- ^
the meristem. The persal of spores. tg
far above the sur constantly adds absence of seta
rounding protective It is hygi'oscopic. Eo
new cells at the is compensated by
sheath and thus base of capsule the presence of C-
helps in tho diaper- which progressively pseudopodium. The o
sal of spo'reB. become differen latter helps in the o
tiated into the !Z5
dispersal of spores
various tissues of by elevating the
the capsule. capsule. s1-1

(vi) The capsule (vi) At maturity («i) The ripe, black O

is a Spherical spore the capsule is an capsule is spherical (ui) It is a long, (vi) The spherical
slender, cylindri
(vi) The poar-shap- ^
sac. Its chief func- oval, yellow body capsule is exter ed usually nodding
- bion' is the pro and specialised both cal, upright object nally more ela capsule is far more
concoi-ned both as a spore producing
duction of spores. in the production of fairly uniform borate than that elaborate both
It has no distribut
and spore distribut thickness through of the liverworts. internally as well
and distribution of ing organ.
ing function. out its length. Its It is differentiated as externally than
spores.
unusual length into the theca that of any other
compensates for portion concerned bryophyte. Exter
the absence of the with spore pro nally it is differen
seta. Functions duction and lid- tiated into three
both in tho pro portion concerned regions, tho basal
duction and with dehiscence. apophysis for '§
 "JSi

Funaria
Anthoceros Sphagnum
Pellia
Riccia Marcfaantia
photosynthesis, mid
dispersal of spores. dle theca for
production of
spores and the
apical opercular
or lid portion con
cerned with dohis-
cence.

The capstilo wall

It is still more the capsule of The capsule wall
is several layers
i>ti"ucturaJiy the It is slightly more Anthoceios dis as in Anthoceros is
capsuie of Riccia elaborate in struc conij'Iex than that 4 to 6 layers in thick and is highly
of Riccia and plays striking thickness. The difforontintccl. The
IS the simplest ture than that of complexity in
its
amoDg the bryo- Riccia. The capsule Maichantia. The
internal structure. outermost layer outorinost layer is
phytes. The cap wall is single-layered capsule wall is 2 or is the epidermis. the epidermis. It
sule TvaJJ consists in thickness. The more cell layers The capsule wall It bears rudimen contains numerous
of a single iayer of wall colls towards thick. The walls of is several layers tary stomata which stomata in the
maturity develop these cells are of colls (4-6) in lack the stornatal apophysis region,
cells.
strengthened thickness.
by The fewer iu the thoca
ring-like thick
bands of thickening outermost layer is aperture. Within
region and none
enings on their called the epi the epidermis is the
walls. material.In addition non-ventilated pho- in the opercular
the capsule wall dermis. It is punc tosynthotic tissue region. The ©pi"
at maturity has tured hero and there consisting of chlo. dermal colls in
on its surface 4 by the pi*esence of ronchymntous cells the oporculum have
vertical strips of true stomata.
with no intercellular greatly thickened
tbin--walled cells. Within the epider spaces between walls excepting the
These are the lines mis is the typically
ventilated photo- them.
annulus. Within
of deliiscence. the epidermis and
They are absent in syntlietic tissue restricted to the
Biccia and Mar- consisting of chlo-
renchymatons ceWs
theca region only
chantia. are the colourless,
\ -with interceUnlar compactly arrang
\ spaces "between
L \ them.
ed two layers of
cells constitutin g

It is absent in the hypodcrmis.

Biccia, Marchunlia Within the hypo-
ond Pellia. The dermis in this
•"fe capsule wall has region is tbo
on its surface narrow spongy
generally two layer which is
vertical lines of continuous below
'■I -i dehiscence
the opposite sides
on with that
apophysis where it
of the

of the capsule each becomes broader.

in a shallow Inside tho spongy
depression. layer in tho theca
region is tho wide O
air space travers B
ed by tho trabe- 25
culae which con- B
ntct the spongy
layer with tho
outer 3 to 4 layer o
thick wall of the o
spore sac. Tho o
air space, tho
trabeculao and tho s
CO
outer spore sac M

are absent in the o

L > !2|
apophysis and
''i operculum regions,
CO

i fa
-
▼
"Within tho epi
dermis in the
-' < > apophysis region is
i f the broad spongy
zone and in tho
oporculum 3-4
layers of small
parenchyma colls
and then a double
■ v'-.i"
T ring of pevistomo
teeth.
 -p»"

Riccia Marchantia PelUa Fnnaria

a
The capsuJe wall I The capsule wall The capsule wall isThe capsule wall As in Anthoccws They are all o
is formed from is formed from is formed from the differentiated ) tlie capsule wall derived from tho H

the entire amphi-

| the entire amphi- entire ampln- from
layer
the
of
outer is formed from the
tho
| outer layer of the
amphithecium.
thecium. I thecium. I theoium.
amphithecium. j amphithecium. •si
o
Prior to maturity
| The capsule wall
| The capsule wall The capsule wall The capsule wall Tho capsule wall
is persistent.
w
the capsule wall j is persistent. |is persistent. is persistent. is persistent. d
disorganises and
disappears. S
d
The arches- 1 The entire endo- I The entire endo- Tho archesporium The arches The arches H
porzum is formed / thecium forms the I thecium functions is formed from tho porium develops porium and inner 00

from the entire| archesporium as i as the arches- inner layer of tho h-om tho inner spore sac orginato
a
endothecium. j in Riccia. j porium as in amphithecium. layer of tlio from tho outer D
Riccia and Mar- amphithecium as layers of tho t?d
dtantia. in Ajil/ioceros. endothecium.
1-9
M
As in Anf/ioceros The major central
The cohimella |
is The columella The columella I The columella
portion of the endo 4
absent. I is absent. is absent. Some
| forms the central the entire endo
thecium forraa tho
scientists consider core of the cap- thecium forms tho
elaterophore as sule. It is formed columella which columella conti 1-3
from tho entire forms the central nuous below with
tho forerunner of
the columella. endothecium which bulky mass of tho enducting
in Riccia, Mar- sterile tissue. strand of the
clianiia and PelHa apophysis. W
forms the arche
sporium. o

W
kJ
The archesporium
The primary 1 The same as in 1 The same as in 1 The archesporium The
divides
archesporium
to form may undergo 1-3
archesporium
^.
divi-
«•- -- ^
Riccia. 1\ Riccia
chantia.
and Mar- divides
the
to form
sporogenous the sporogenous • division to • form
>
des to form
•Vho B'^OTOgenous tissue one to 4 1 tissue 2-4 layers two layers thick

tissue which fills

the entire space layers in thickness. in thickness. It sporogenous tissue.
It is narrow and overarches tho It is thusexti-emoly
within the capsule
wall. overarches tho rounded apex of reduced forming
rounded apex of tho massive but a small and
j the columella like columella and narrow part of the
1 a dome and sur forms a dome- thoca region of the
rounds it. shaped zone in tho capsule. It neither
upper part of tho oxtenda to the base
capsule only. nor apex of the
capsule. It simply
surrounds tho colu
mella and does not
overarch it.
Major portion of Half of the sporo A central mass of O
the sporogenous The sporogenous The entire sporo Tho entire sporo H
genous tissue sporogenous cells
tissue is differen tissue differentiates genous tissue gives !25
differentiates into at the base of the genous tiasuo us in
tiated into the into fertile spore rise to tho spore
H
the spore mother capsule becomes mother cells and
Sphagnum cliffe- £0
apore mother cells and the rest differentiated into mother cells. The rentiotcs into the >
cells. Only a few the sterile pseudo- elaters arc absent. spore mother
into elater mother .the
elaterophore I elater mother cells.
peripheral cells cells. The latter 'whereasthe|The former give cells. Tho elaters Q
o
remain abortive give rise to tho I remaining spore- ' rise to tho spores are absent. £3
and function as sterile, hygrosco I genous coUa and tho latter to Q
nurse cells. By pic elaters which I surrounding the tho pscudoclaters. d
some scientists the help in scattering ( olatorophoro form ca
M
nurse cells are the spores differen I tho spore mother O
considered as fore tiated from tho ! cells and the
runners of the spore mother cells. w
elaters.
elaters of Mar
chantia.
There is no The capsule of The capsule
regular method of Merchantia dehi
Dohisconce is very Dehisconce is by Tho annulus cells
dehisces in a more regular. Tho the separation of a
dohiscenco to allow sces to allow tho regular manner as perish and the
the spores to mature capsule lid and ig explo
spores to escape. compared with splits along 1-4 oporculum falls off
escape. The surviv The capsule wall Marchantia. It
sive. Tho spores exposing the pori-
linos of doliiscence. are forcibly blown
ing outer layer of splits open into dehisces by crossed stome teeth. Tho
the calyptra and several Tho splits extend out.
usually cleavages along the downwards separa hitter by their
tho adjacent thallus 4-6 irregular teeth- four linos of peculiar disposi
ting into valves
tion and hygro-
 GENERAL CONCLUSIONS 301
300 BOTANY FOR DEGREE STUDENTS—PART III; BRYOPHYTA
REVISION QUESTIONS
o * •
fe 5n_£>-.SCcC
o on S
i
b OT ^ e o
d ® S c 5; 1. Describe the evolution of the sporophyte in the
^.= S by you. ^ ^ ' -l.
0
2. Compare the sporophyte of il/arc7»antia with any Moss s^ied ^
.2'g "s o ? 3
o B-
■^.2
you. Give labelled eketehes. i-^iara Umv., 19o8)
P — rr 2 3. Compare the otructure of the sporophyte of
o>--S^--r g ytcfl Anthoceros. ^ " *
j^ggCOtC^ CS . -Wwfc, o o j 2 >>
gusalli a o.i c p-S 4. Write a comparative account of (a) the structure of the thallus and
m the orgouisation of the sporophyte in
5. Compare the gometopbyto of Biccia rrith
>j c — o :3
».3 g >
° & o S-p 6. Compare the sporephytes olBiccia. An^occro, and^^»mnm
Give a comparative account of the sex organs ot Biccia. Mar-
o 2'-'_. ^-"nP
chantia, Anthoceros and Funana,
Discuss the systematic position of division of Bryop^ta.
9. Describe the various views on the origin of Biyoph^a.
10. Discuss the biological and economic importsnco of Bryophyto
u onnrinm ' Givo on account of the origin and fate
J5 > J;
11. Stile'in a ?atalar 'form in the bryophytos etudiod by you.

0.E-<.2 eif

s ®
» oc'p '
^■anu
. >

= -I3 r^-2 ft
S 2 «2 co v.. «»

o n
^ ft ® OT .2 " . . « . ^ LA . AA

a K
p 2 c3
o i- o
■P -P Q4
>> ® CO to
t f
-= — .2
S-S-S 2
■p 2 2^»

5)®C5~ t>
Er t« o p ja
2

to ft ^
»/«

.3 ""a
^ 'o-z-Tsa
^ -5 o & M » d
« ffi -c
o ®.« ^
B
2 tS tS 60 - 5 -p 'o
CO O d
- • o
H.2-S
 INDEX 303

Bryokinin,258
Anthocoros chambensis, 152^ 157 Bryophyta, 5, 15, 16, 38, 68, 227
Anthocoros erectus, 152. 157, 166 biological importance of, 271-272
Anthocoros himalayensis, 22, 152, Io4, origin of, 267, 270
155, 153 systematic position of, 270, 71
INDEX Anthocerotopsida, 16, 150, 275 Bryophytes, 1,2
Antical lobe, 132 alternation of generations, in, 8, 11
Antithetic theory, 12
Abscission layer, 254 Anthoridial initial, 47, 80. 116 335 Anioal cell 59, 115, 155, 171, 197,
antheridium of, 4
158,210,234 ' ' ' ' archegonium of, 4, 5
Acrocarpous, 240 Anthoridial mother colls, 117, 135 ^ 199, 20s, 210, 231. 235, 238, 239, as amphibians of the Kingdom
Acrogynae, 109, 131 Anthoridial wall, 46, 79, 13G, 157, 159 248 Plantae, 1, 23
Acrogynous Jungermanniales, 31 108 Anthondiophoro, 70, 77, 78 82
Apical growth, 43, 74, Ho, lac, 208, classification of, 15, 16
Acropotal, 45, 59, 80 212,231, 286 differ from thallophytes, 14
Adaptation!", 1 Antheridium, 4, 45, CO Apicalinitial. 43 economic importance of, 273
Adventitious branches, 22, 44, 75, ioq Antheridium raofcher cell, 117, 135 Apophysis, 242,243, -"5, 262 evolution of sporophyte of, 276,
Antherozoid, 118 Apospory, 156, 157, 196, 232 278
Affinities, 35. 183, 222 Anthocorotaceae. 151 Appendage, 71 fertilization in, 5
Air canals, 41, 62 Anthocerofcae, 16, 150 Aquatic, 1, 38 212,232, cametophyte generation of, 3, 5
Air cavity, 218 characteristics of, lol Archegonia, 7 83, 1*'0, ia», » origin of, 267, 270
Air chambers, 30, 42, 66, 72 73 Anthocerotales, 15, 26, 150, 151 236, 292 plant body of, 3, 4 _
origin of, 42 '' Genera of. 150, 151 Arcbegoniatae, 5 reproduction in, 4, 5
origin of, 189, 191
Air channels, 41
Airpore8,41,G6, 09. 71* 72* 15<1 systematic position of, 185, 186 ite1orSS^'4C80,liM33,ie0. resemblances with algae, lo
salient features of, 12, 14
Air space,244 Anthocoros, 22, 150 sperms of, 4 r« o
ns,:37,:«. sporophyte generation of^6, 8
Algal hypothesis, 268 269 affinities of, 183, 184
Alternation of Generations 8 n no apical growth in, 155 systematic, position of, 2/0, 3/1
100, 130, 170, 238, 2!^' ■ biological importance of Antho- A^chegoni™ mother cell. 113 137 Bryopsida, 16, 192, pj
classification of, 19/, 198
m Funaria, 236, 237 ceros sporophyto, 187, 189 Archesporium, 38, oZ, oJ, i-a,
in Marchantia, 97, ICQ m? 21(5 distinctive features of, 196, 197
uehisconcc of capsule, 169 origin and fate of, 273, 274 gametic reproduction of. 195,196
in Pellia,l28,130 dovelopment of antheridium of,
in Poreila, 144, 147 158,159
AreoIae,"'66, 69 vegetative reproduction of, 192,
in Riccia, 59, 02 Ascending series, 04, lOi 195
development of archegonium of, Asexual reproduction, 21 Bulbils, 195, 231
definition of, 10 160,101 , Aspiromitus, 150 Buxbaumideao, iJ'
Amphibians, 1,3 develonraent of sporophyte or, Assimi'atory filaments,
colls, 206, 20
^phignstria, 133 166, 168 , „, tSmilatory 41, 72
Amphitbecium, 51,89 I2q external features of the thalius Astroporeae, 35
^ 167, 210, 2«,'255 ' "3. . of, 152, 153 Autoieous, 17b, 121, Calobryum, 31, 104
Anacrogynae 109 fertilization in, 101 -j Autotrophic, 57
Analogous, 3,'72, 229 germination of spores ot, Axial cell, 161, 237
Anatomy, 204, 229, 282 habitat and distribution of, Axial row, ^59 Campbell. W,23. 33, 35, 42, 6!, 82, 197
152
of the stem of^ ^06, 207 internal structure of the thalius Cap cells, 48
Capillary cells, 207
^ of the steSof 230 of. 154, 155 f ine
nutrition of sporophyte ot, '
Capillary apparatus fg,92. 102, 241
Anaphase,55
•^(Ireaeidae, 197
Pnagnum,204, 206 166 P I -'
Barnes,42
Barrel.sbaped, 71 Sl°'wk.iM'92,'l23, 140, 165, 107,
•^dreaeobrya 197 salient features of, 171, 1'^ Basal cell. 13' ^ 216,218
Andreales, 15, 197 sexual reproduction in, 157, Bhardwa], Ibb
species of, 152
Biflcgellate 13,"3 Carr, 36
Carrpaceae, 3()
sporophytic phase of, 162, 109 Bilobod, 105_, 133 Carrpos, 36
structure and position of antuei» Bivalonts, 5o
dium of, 157 Cavtir2M3. 64, 101, 186
structure and position of arcn 5!""!'; fi 103,100
Androlynou ^7, goniura of, 15y, 16 > 119, 138, 213.
Aneura, 105 ®®PfacIea, 77
Annnhis, 216, 24^; 9=4
structure of spores of, 109
structure of sporophyte, 102, 10
^=f/e5,'33,/3,e.,101.107 238, 239
Central cylhider,2-9
Antheridia, 77 70 1 ,t summary of life cycle, 173,1'" Chiasmata, o5
Ohlorencbymato^eellr.ll, . .gg
.6,10
290 116.134^ systematic position of, 187 Bryan, 19* gg 225, 227
Anthoridial rim t. ' * Bystetnatio position of Anthocero Cbloronemal, 228, 258
1.57.13?""^-. ^6. 47, 78. tales, 185, 187 .
vegetative reproduction in, 1®'
156

802
 305
INDEX
BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA
H
Chlorophyceoe, 183, 191
Chopra, 68 Endogenously, 158
Funaria, difforences between Funaria Habitat, 20, 33, 68, 113, 132, 151,200,
and liverworts, 264-265
Ohoromatid, 55, 5q Endosporo, 92 differentiation of annulus m,45 227, 280
Chromosomes, 54, 56 Endosporium, 68, 218 Hadrome, 204, 205
Church, 31, 64 Endostomo, 253, 254 external morphology of sporophyte Handoo. 150. 166. 190. 191
Classification. 15 an qs mo ta- Endothecium, 51,89, 123, 141, 163, 167, of, 240-243 . ofifi Haploid, 11, 55, 229, 259
ColumellQ, 163. J67 oii o.o% 210, 248, 252 Haplomo trium, 104
fate of primary amphithecmm, 45U- Haplomitrineac,
Coma,203,209 '" ' Envelope cells, 50, 119 108
Condensation theory, 32 33 Epibasal, 89, 107, 122, 140, 248 fa?e of primary endothecium, 249. Haplophase, 57, 60, 99, 173
or.Q
atrand, 196, ^5 Epidermal region, 71, 243 fertilisation in, 239, 2^ t6f'2l 33,38,
Conical hood, 255 Epidermis. 37, ICo, 167, 216, 218, 229,
Corsiniaeeae, 35, 36 243, 245, 252 formation of oP®.*"®" 252-'^54
asexual reproduction in, 21-22
Eptiphytic, 105 as inefficient land dwellers, 18-19
Cover ce,K 48, 49,50. 85, 86,120, 159, Equator, 47 =:rA=l"' chief means of perennation in,
Equiaetum debile, 32 22-23
Eubrya, 197, 198 classification of, 26-27
Cover initial, 161 243-240
Cupules,69
Evans, 31. 35, 42, 64 juvenile stage of, -28 distinctive features of, 25, 26
Evolutionary riddle, 64 distribution of, 20-21
Cytokinesis, 55 Exogenous,43, 200 habitat of, 20
Exoscopic, 89 25£sr^Seri. plant body oj 19;20
Exospore, 92 sexual reproduction in, 23-24
De Bary, 15 Exosporium, 58, 218 vecctative reproduction m, 21-24
Exostomo, 263 Heterologous, 10,14, 100
Ex Janczewski, 15 pelStihsSn changes m,240 Heteromorphic, 10, so
Heterotropbic, 57
Dendroceros, 150, 151 ' F Hirsch, 42
Diakinesia, 55 Femalo plants, 45, 86, 114, 130 Homologous, 6, it, i-i
Diaphragm, 245, 254 Homologous theory,
^icbotomous, 144 Fertihaation, 1, 6, 50, 87, 120, 138, lOl, FunaS:— Homosporous, 13
Dichotomy,44 _ ^214, 239,261 Hope, 185
Dioecious, 29, 38 CO 7r i..« ^imbriaria, 07 Howe, 16, -6
190,209 ' ' ^^0, 157,178 First unaphaso, 55 ^'""e£acuris.ic|0f, 220 Humidity, 77
Diplohaplontic, ID, nn ^iratmetaplmse, 55 Hyaline cells, 2U0
Hyalodermie, 294
250, 251, 253 ^■"%rarSerisScsof,226, 227
F fth ring, 251 Hygroscopic, -
DipIoMnucleut'sTu |:age a 48 81, 116 Furrow, 39, 60 Hvpobasal.Sy, 19/. 1 »
Sfet'so"' ^lagelleform, 204 Hypodermis, 244. -62
D^cussion, 64, 100, 183 Flagellophore, 210
Folioso, 20, 32, 104, 107, 131 ,-9 Gametes, 9 45. 195
Foot, 9, 25, 34, 38, 57, 89, 92, 140, Jd-' ^„etio rcp™ductmn
105, 214, 241 82 22
Gametophore, 2e, ^6, 28, 37, 45, Ikono,
Incubous, IOj^ "i'
Forerunners, 52 GametopJg,; ,27, 259 3 g, 99, 129
200, 288 ' 1". 131, 154, Fossombronia, 104, 106, 107 Initial cell, 2o
Fourth ring, 251 ,254
iragmentation 21, 44, 74, 106, ' Innersporeeae |44,2eO
Dersiv;nSal'ig^,\«0j3l05. 132 165
Fritsch, 33 g:s"sratiou,«.«
Funaria, 225, 227-266
adult gametophyfce of, 228-229 ''="^°vero,;mentoU7 £:r-tii.T9rn"i2o.u2,i5,
E alternation of eenerations iu, 2oo'
„erm.nation of 76 162,168
16-', loo
Ecological importflv. 259 Structure Isoinorpbio, 11
220 ^i^^rtance of o,,i anatomy of leaf, 230-231 General cone -.one, -67
anatomy of stem, 229-230 Germ povo. ^9
apical growth in, 231
compariscn between the sporop y
tes of Funaria and Sphag*^" '
255-257 ,7 213! 335 J59
dehiscence of capsule of 246, Goebel, 64' i9^g7 JS^er5S»-ouB gametophyte, 34
ElateroXre^U^H^^^^^ development of antheridia ^ Griroal^'^- '
236
development of arohegonia ''
Endogenous,43. 2H.248 236, 239 .f
development of sporophyte '
247
 307
306 BOTANY FOR DEGREE STUDENTS—PART III: BRYOPHYTA INDEX

Jungermannialea, 15, 26, 104, 112, 131 Musci, 15,16, 192

Madotheea distribution of, 31 Merchantia, species of, 63, 69 classification of. 197-198
acrogynous, 108, 131, 149 external characters of the thallus sporophyto phase ot, 88, y.i distinctive features of, 196-197
anacrogynous, 108, 113-130 of, 132-133 structure of antheridium
classification of, 108, 109 vegetative reproduction in,
gametophytic phase of, 131-139 structure of archegonium of, 80- 193-195
comparison with Marchantiales.
147, 149
germination spores of, I42 86 . Kr
distribution and habitat of, 105
habit and habitat of, 132 structure of gemma of, /b N
external appearance of, 104-105 internal structure of the thallus structure of sporogoniura of. 9^
of, 133-134 summary of the life cycle of. 97, Nebenkorper, 82
salient features of, 109, 111 position and structure of an-
sexorgans of, 106. 107 100 .. r «<! Neck,5, 48, CO, 85, -1-, -3o
thoridia of, 134-135 systematic position of, b8 Neck canal,85. 86. 137
vegetative reproduction of, 105-
position and structure of arche- vegetative reproduction in, 74, Neck canal cells, 5,48, 60, 88
106
Jungermaannineae, 108, 109, 131 gonia of, 136-137 Marchantiaceae, 35, 36, 66. 68 119, 120, 159, 161, 212, 214. 237,
Juvenile stage, 190, 202-203, 227 228 ' salient features of, 142, 144 general characters of,60, 07 238. 239
287 ' ' sexual reproduction in, 134 138 Marclmntiaceoua gomatophyto 33 Neck cells, 86, 120, 137, 212
sporogenesis in, 141-142 Marchantiaceous thallus, . Neck initials, 50, 86, 120
K sporophytic phase of, 139,142 origin of, 31, 32 Nostoo. 152
of sporogoniura of, 139- Notch, 69 , ,3-
Kashyap, 32, 44, 64, 157, 189 Marcran"tLles, 15,26, 23, 37, 38, 52, 66, Notothylas, 150, 151, 1/6, 183
Kachroo, 170 , summary of life cycle of, 144, 147 Nurse cells, 52
Nutrition, 57, 92, 121, 165, 2t>7
Male plants, 06, 86, 134 Nutritive fluid, o3
^ale receptacle, 78
Land, 42 dis"lnctiW features of, 34, 35
Leaf, 105, 132, 230
air chambers of, 72 sporogonium of. 34
Leafy stage, 192, 196,227,258 Marsupium, HO
Leaves, 110, 192, 196, 206 antheridiophore of, 78 82 Octant stage, 51
dorsal, 105, 132
ventral, 105, 132
apical growth of, 74
archegoniophore of, 82, 85
MeSlaf'l33. 204, 205 Oil cells, 29, 73, 76
Oogaraous, 4, 13, 77
of gametophyte with Medullary, 133 Oosphere, 120,161
Leitgeb, 16, 26, 42
Oospore, 120
Leptolene stage, 54
Lewis, 54 SSa!S 130. 105. 157. 160. gP:,d::m,Ml333,242,243,245.
Lid, 216, 245 °°Rfcc?a™3-9t
dehiscence of capsule of,93 252, 254
Lid cells, 48, 50, 85, 86, l61
Liverworts, 15, 18, 184
80 antheridium of,
MnPXIb2%t-.2.7 210 Ostiole, 79, 116
asexual reproduction in, 21, 22 Meristematio mno, 163 Outer peristomial layer, 2o3
Outer peristome,253
as inefficient land plants, 18 19 development of archegonium of.
Outer spore sac, 244, 2o0, 2o2
chief means of perennation in, 22
23 ' development of gemma of, 77 SSSfl- ,00.113
Sr;—aUaye,.253 Ovum, 5, 86,161
Oxymitra, 38
classification of, 26, 27
distinctive features of, 25 26 89, of sporogonium of, Monocarpaceae 36
distribution of, 20,21 * dispersal of spores of, 93 Monoclea, 3u, 00
habitat of, 20 distinctions between male and Monocleacoae 35, ^^7^ 178,
plant body of, 19, 20 lemale plants of, 77, 86 Monoecious 29 40. Paohyteno, 55
sexual reproduction in, 23 24 pande, 181. 186
vegetative reproduction in 21 22 ^71 thallus , ,'32 m 187. 196 papillae, 80
Lobule, 105, 133 '' fertilisation in, 87 MoDopodial, , Papillate, 234
gametophyte phase of, 69, 87 Par^aphyses, 190, 233, 20U
Lunularia, 21, 67
Lycopodium, 32 germination ofgemma of, 76-77 r^sT^'is^
°®^autoicous, 196 .._j9g
parasite, 2o
Paroicous, luo
M
f spores of of
habitat and distribution 94.95
68 Peat, 201
mportant features of, 95, 97
Madothecaceae, 109, 13i
Madotheea, 131, 149
~^nioturo of'the thallus Mosses, ,
gyooicous. V-o-oduction in, 192.
apical growth in. Ho
dehiscence of capsule of 142 vegetative roprouu r;rprsn::&erikiumof,
development of antheridium of
13o, J36 * Kiale receptacle of, 78-70
195
Mucilage cavities, lo4 delSpmeut of archegonium of.
development of archogoniura of Mucilage cells, 73, 119-120
137,138 * Mucilage , jgg
140, 142
sporogonium of.
'
Muoilage mound, 160
 309
INDEX
308 BOTANY FOR DEGREE STUDENTS—PART IH: BRYOPHYTA
Riccia, structure of spore of,
Pyrenoid, 155, 183 summary of life cycle of. 59-62
Pellia, development of sporogonium Porella, internal structure of the systematic position of, 38
of, 122, 124 thallus of, 133-134 vegetative reproduction m, 44.
"dispersal spores, 125 Q
position and structure of the
distribution of, 114 antheridium of, 134-136 Ricciaceae, 35-38 - .- 00
external characters of the thal- Quacking bogs, 222
position and structure of the Quadrant stage, ol. 89 general characters ot, o7-o8
lus of, 114 archegonium of, 13C-I37 Riccardia, 22, 105
fertilisation in, 120 salient features of, 142-144 Ricciocarpus, 38, 41-42
germination of spores of, 126 sexual reproduction in, 134-138 Rim, 254, 254
linbitat of, 113 sporogenesis in, 141-142 Roof initial 158
internal structure of thallus of, sporephytic phase of, 139-142 Raye. S3, 84. 87, 89 Bothmailer, 16
115 structure of sporogonium of Reboulin. 22, 31, 67
Nutrition and protection, 121, 122 139-140
position and structure of the summary of the life cycle of
antheridium of, 116 144-147
Keooptive spot, S6
position and structure of the systematic position of, 131 Reduced, 64, 18i '>78-279 Salient features, 12, 109 126, 171
archcconium of. 118, 119 Reduction theory. 31, 32, -'S of Anthoceros, DJ, I'o
Pores, 34, 37, 72, 165 Reproduction, 4 44, 74, ilo. 1^4, of Bryophytes, 12-14
salient features of, 126, 128 Porellaceae, 109, 131
sexual reproductiou in, 116, 120 ^ 155,208.231 ofPellia, 126-128
general characteristics of, 131
species of, 114 Postical lobe, 133
asexual, 21 . ,,/> 104 of Porella, 142-144
sporogenesis in, 124 Preissia, 29
sexual,4, 23, 45, '4; He. 31.40,44,08, 71, 73
sporogonium of, 120, 121 Primary amphithecium, 250, 252
summary of the life cycle of, of Marchantia, 71
Primary androgonial cell or cells, 134, 155, 192, 208, 231 ofRiccia,40
128. 130
47, 81, 117, 159. 210, 235 Schiffner 33, 35
vegetative reproduction, 115, 110
Primary antheridial cell or colls, 47, Schuster, 16, 36
systematic position of, 113
80, 117, 135 Bluzoidai, 70, -08, ^ Schizogenously, 42, <2
Pelliaceae, 113
general characteristics of, 113 Primary arohegonial coll, 50, 86, 160 f Second ring, 2ol
Perianth, 131, 13", 142 Primary archegonuim, 212 (j QQ 71
Primary archesporium, 250 smooth-Wiled.
199' 22829, Sond^ry p'i^Sma>3.209, 220,231
Perichaetium, 84, 89, 91, 212
Perigonium, 233 Primary axial cell, 60, 86, 119, 137, 'sXT2T«:38,37.°89,!12. 140,241
Perigynium, 8G, 8S, 89, 91 138, ICl, 213
Peripheral cells or initials, 50, SO, 119, Primary canal cell, 120, 161, 214
Primary can or cover cell, 50, 86, 110,
irolgS'3':'ll''37,°l5,77, 100,260.289
137, 237 of Bryophytes, 3
Peristome, 245,252, 263 138, 101, 238, 239 between two genara-
Peristome teeth, 254 Primary endothecium cells, 249
Peristomial layer, 253, 254 Primary jacket colls, 159 davXpment of antheridium of,
Persistent apices, 23, 44, 150, 195 Primary neck canal cell, 50, 86, 161,
Petalophyllum. 22, 106 238 development of archegonium of, in Marchantia; /7-s/
Phaeoceros, 151 in Notothylas, 1/8
Primary protonema, 193, 258 49-50
Photosynthetic cells, 206 Primary spovogenous tissue, 123 in Pellia. ilC-120
Photosynthetio region, 32 40 41 and habitat of,38 in Porella, 134-138
Primary stalk cell or cells, 47, 80, 86, 'be thallus
71,72 '
of Marchantia, 72, 73 117, 135, 161 Sg;XnX!209-214
ofRiccia, 41 Primary ventral cell, 50, 86, 120, 161, feriilisationin,
of, 39, 49 . .>0,
_ 01
Photosynthetic tissue, 165 214
Placenta, 165 Primitive, 04, 100, 183
Plagiochasma, 31, 33, 67 Progressive evolution, 31, 33-34, 101 Spermatocytes,47 117
Porella, 2i). 131-149 Progressive simplification, 64, 102
dehiscence of capsule of 142 Progressive sterilisation, 101 ^P-rMSSia!8l82
development of antheridium of, Propagula, 100
135, 136 Prophase, 54
developmeat of archegonium of, Proskauer, 16, 35, 36, 150. 151, lOl of Anthoceros, loo
Irotandrous, 116, 157, 232 species of, ofFunaria, 234
of,
protonema, 14. 142, 199 202, 219, 228 of Marchantia, /9-80
Protonema stage, 144, 192, 196 of Pellia, 116
distribution of, 13i Pfieudo-elatera, 151, 168 ructXroTXtheridium of, of Porella, 136
6xt©mal charaftnT-a .-.rii. A seudo-elater mother cells, 168 of Sphagnum,2iu
of, 132-133 °^thetliallua Pseudo-perianth, 86, 104 .tSureofarehegoniumof. of Riccia, 46
Pseudo-podium, 217 48-49
Pteridophytes, 32, 184
Habit and Sabiro"l°3t"^
 311
index
310 BOTANY FOR DEGREE STDDENTS-PART III: BRYOPHYTA
Vegetative reproduction in Pellia, 115-
Sphaerocorpaceae, 26 Thallus, 39, 59, 104, 280 IIG
SphaerocarpuB, 33 Sporogonium. 0, 7, 9, H, 24, 34, 44. 51, Theca.242,243 263 in Porella, 134
Sphaerocarpales, 26, 172 66, SS,91, 120, 139, 162, 240 Theory of sterilisation, 64,-^/6 in Riccia, 44-45
Sphagnaceae. 200 as an individual, 7-S Third ring, 251 in Sphagnum, 208, 209
Sphagnales, 199, 200 of Anthoceros, 162, 105 Traboculao, 244, 2ol-2.}2 Venter, 5, 48, GO, 85. 137, 212, 236
SphagDidae, 15, 197, 199^ 200 of bryophytes, 5-7 Treploid, 195 Venter, cavity, 59, 85, 86, 120, 236
distinctive features of, 199 of Funaria, 240-243 Tuber formation, 2-, 44 Venter initials, 50, 86, 120
Sphaguobrya. 107, 197 of Liverworts, 24-25 Tubers, 22, 106, I06, 194, 231 Venter wall, 48, 50^85
Sphagnum 199, 200-224 of Marchantia, 92 Ventral, 29, 34, lOo
affinities of, 22«:-223 of Nototbylas,'180-182 U Ventral canal cell, o, oO, 60, 80. 86
anatomy of leaf of, 206-207 of Reliia, 120-121 118 132, 137, lo9, 161, 212, 214,
anatomy of stem of, 204-205 of Porella, 139-140
Underwood, 16 237', 238i 239
apical growth in, 28 of Riccia, 56-57 Verdoorn, 35
characters peculiar to, 224 of Sphagnum,217-218 Vesicle, 48
development of antheridium in Sporophyte, 3, c. 8, 10, 24, 50. 107
210-211 139. 1C2, 178, 259, 262 294 W
development of archeconium in. evolution, of, 275-279 Valves, lOS, 140, 169
212-214 Sporophytic generation, 3, 10 Vasisht, 32
Wall initials, 47, 117, 136, 16'
development of sporoconium of. Sporo^phytic phase, 6, 51, 88, 139, 162, Vegetative cells,^ ^- 74 Watson, ISO
Wettstein 31, 197
214-216 Whiplash. 13
distribution of, 200 of bryopliytes, 0-8
of Anthoceros, 162-169 Woesler, 208
ecological importance of, 222
economic importance of, 220-222 of Marchantia, 88-93 Z
fertilisation in, 214 of Pollia, 120-125
gametopbyte phase of, 202-214 of Porella, 139-142 Zyg0te,,6. 10,51, 60, 88, 161,214,248
germination ofspores in, 219-220 ofRicciu, 51.57 in Liverworts, 21- Zygoteiie stage, 55
habit of, 200-201 of Sphagnum, 214-219 in Marchantia,
habitat of, 200-204 Stalk cells, 159
juvenile stage of, 202-203 Stem, 110, 132,192, 190.204, 228
leafy gametophyte of, 203 Stephansoniella, 29, 31
mature sporogoninm of, 217-218 Stomn, 64, 154, 197, 221
mechanism of dehiscence and Stomata, 165, iG7, 216, 252, 282
dispersal ofspores in,21S-219 otorago region, 40, 41, 71^ 73
position and structure ofantberi- of-Marchantia, 73
dia of, 210 of Riccia, 41-42
position and structure ofarche- Storage tissue, 34
goma of, 212 Succubous, 1(15
pseudopodium of, 217 Summary of the life-cycle
sexual reproduction in, 209 214 of Anthoceros, 173, 176
Bporophyte phase of, 214, 219 of Funaria, 259, 263
vegetative reproduction in, 208- of Marchantia, 97, 100
of Pellia, 128, 130
systematic position of, 200 ofPorella, 144, 147
Spongy cortex, 207 of Riccia, 59, 62
Spongy layer, 244, 2-31-2.52 ^urface involution, 42, 72
Spongy zone, 243, 255 ouspensor, 107, 112, 140
Sporangium, 24G Synthetic typo, 183, 184, 224
Spore mother cells, 52, 88,89 19^ tti
164,216 ' 124, 141,
Spore sac, 218, 244 246
Spores, 38, 57, 58, 109, 245 Takokia, 27
Sporocytes, 52 Takalciaceae, 20
Sporogenesis, 53, 124, Hi Takakiales, 26
in Pellia, 124 Takhtajan, 16, 26 185
in Porella, 141-142 largionaceae, 35
in Riceia, 53, 56 Targionia, 22. 29, 31, 35, 67
Sporogenic, 10, 176 Pelophase, 55
Sporogenous cells, 123 Terrestrial. 1, 38 Date
Tesselina, 38
163, 168, Tetrad, 54
Thallophytes, 6, 8,14