LIGNOPHYTES - WOODY PLANTS

MAJOR APOMORPHIES :
- vascular cambium (secondary vascular tissue)
- cork cambium (periderm)
- heterospory (seed)
- endosporic female gametophyte (seed)
- reduction to 2 megaspore per megasporangium (seed)
- retention of megaspore within megasporangium (seed)
- integument with micropyle (seed)
- pollination droplets
- endosporic, male gametophyte = pollen grain
- pollen tube - sperm motile (zooidogamy)
- eustele
- monophyletic lineage of euphyllous vascular plants
- share derived features of a vascular cambium (rise to wood) and a cork cambium
(produces cork)
- Secondary growth growth of vascular and cork cambia ; increase of the width and girth of
stems and roots
- Primary growth vertical extension of stems and roots due to cell expansion
- Vascular cambium a sheath, or hollow cylinder, of cells that develops within the stems and
roots as a continuous layer ; layer between xylem and phloem
- Secondary xylem = wood ; functions in structural support
- Bifacial layer of cells are continuously generated produced from both inside and outside
- Primitive plants cambium was unifacial ; producing secondary xylem but no secondary
phloem
- Cork cambium = periderm (outer bark) and cork (cells contain suberin; similar to cutin and
resistant to H2O )
- Cork functions as thick layer of cells that protects the delicate vascular cambium and
secondary phloem from mechanical damage, predation and desiccation
- Dendrochronology using wood anatomy to study the past
- Monopodial growth single main shoot develops branches from lateral buds

SPERMATOPHYTES - SEED PLANTS

- monophyletic lineage
- major evolutionary novelty is the seed
- Seed is an embryo which is an immature diploid sporophyte developing from the zygote,
surrounded by the nutritive tissue and enveloped by the seed cot
- Radicle immature root
- Epicotyl shoot apical meristem
- Cotyledons one or more young seed leaves
- Hypocotyl the transition region between root and stem
- Ovule immature seed, prior to fertilization
SEED EVOLUTION
1. Heterospory - formation of 2 types of sporangia : megaspores and microspores ; ancestral
homospory ; megaposre - female gametophyte - bears archegonia ; microspore - male
gametophyte - bears antheridia
2. Endospory - the complete development of female gametophyte within the original spore
wall ; exospory - spore germinates and grows external gametophyte
3. Reduction of megaspore # to one - occurred in 2 ways : 1st - the # of cells within the
megasporangium that undergo meiosis was reduced from several to one ; gives rise to 3
haploid megaspores. 2nd - the 4 haploid megaspores produces, 3 of them are consistently
aborted leaving only 1 functional megaspore ; single megaspore undergoes a great increase
in size
4. Retention of the megaspore - megaspores are retained within the megasporangium ;
accompanied by a reduction in thickness of the megaspore wall
5. Evolution of the integument and micropyle - integument - layer of tissue covering the
megasporangium ; fusion of telomes (ancestral branches that surrounded the
megasporangium ) micropyle - small pore at the distal end of the integument ; site of entry of
pollen grains (derived from lagenostome)

POLLINATION DROPLET
- a droplet (mostly H2O + sugars/amino acids) of liquid that is secreted by the young ovule
through the micropyle and is formed by the breakdown of cells ; functions in transporting
pollen grains through the micropyle ; ocurs by resorption of the droplet ; present in many
nonflowering seed plants
- Pollination chamber cavity formed by the breakdown of cells

POLLEN GRAINS
- an immature, endosporic male gametophyte ;
- Endospory in pollen grain evolution was similar to the same process in seed evolution,
involving the development of the male gametophytes within the original spore wall
- male gametophyte grows an exosporic pollen tube, functions as a haustorial organ,
obtaining nutrition by absorption from the surrounding sporophytic tissue

POLLEN TUBE
- Zooidogamy ancestral type of pollen transfer found in cycads and ginkgophytes in which the
male gametophytes feeds off the tissues of the nucellus, then motile sperm is delivered into to
a fertilization chamber where the sperm swims to the archegonium containing the egg
- Siphonogamy type of pollen transfer found in conifers; deliver nonmotile sperm cells to the
archegonium or egg ; a type of siphonogamy evolved independently of the angiosperms

OVULE AND SEED DEVELOPMENT

- Megasporocyte develops within the megasporangium of the ovule ;
- Megasporocyte - meiosis - tetrad of 4 haploid megaspores - 3 megaspores that are distal
abort - only the proximal megaspore develops - pollen grains mature in the pollen chamber -
male gametophyte and form pollen tubes - male gametophytes may live for several months to
a year - functional megaspores undergo mitosis - endosporic female gametophyte
- Pollination the delivery of pollen grains to the ovule
- Fertilization actual union of sperm and egg

SEED ADAPTIONS
- seeds provide protection by means of seed coat ; seeds functions as the dispersal unit of
sexual reproduction

EUSTELE
- Primary stem vasculature that consists of a single ring of discrete vascular bundles ; each
vascular bundle contains an internal strand of xylem and an external strand of phloem
- Protoxylem endarch ; toward the center of the stem

ARCHEOPTERIS
- lignophytes that lacked seeds
- Archeopteris with leaves like fern

PTERIDOSPERMS - SEED FERNS

- paraphyletic group that had fernlike foliage, bore seeds
- Medullosa

GYMNOSPERMS
- naked seed
- ovules are not enclosed by a surrounding carpel layer at the time of pollination
- nonflowering seed plants
- Cycadophyta as the most basal lineage followed by the Ginkgo then the Conifers

CYCADOPHYTA
- served as fodder for plant-eating nonavian dinosaurs
- monophyletic
- 320 - 340 species in 11 genera
- source of food starch (sago) typically collected from the apex of the trunk
- loss of axillary branching on the aerial trunk
- Zamiaceae (Coontie family) exhibit circinate vernation ; have seed cones or strobili ; each
megasporophyll bears 2 ovules or seeds
- dioecious ; sexes are separate
- Cycadaceae (Cycad family) absence of seed cones ; seeds are produced on the margins of
the numerous megasporophylls which are aggregated in dense masses

GINKGOPHYTA
- one extant species Ginkgo biloba
- distinctive obtriangular leaves (fan shape)
- dioecious ; flagellated motile sperm
- Male ginkgo produce cones but do not bear structures that resemble sporophyll
- Female ginkgo do not bear cones
- Ginkgoaceae (Maidenhair Tree Family)

CONIFERAE - CONIFERS
- aka Pinophyta or Cinferophyta
- monophyletic group of highly branched trees or shrubs with simple leaves
- leave are acicular (needle-like) , linear , subulate (awl-shaped)
- Fascicle a specialized short shoot consisting of stem tissues, one or more needle-shaped
leaves and, persistent basal bud scales
- loss of sperm motility together with Gnetales ; have pollen tubes ; siphonogamy
- monoecious ; sometimes dioecious
- pollen grains are bisaccate in which 2 bladder-like structures develop from the pollen grain
wall (wings) which may function as a flotation device
- seed cones have compound structures : bracts (modified leaves) which subtends a separate
seed-bearing structure called ovuliferous scale (modified lateral shoot system)

PINOPSIDA
- Pinaceae (Pine family)
- 12 genera , 225 species
- monoecious seeds are usually 2 per ovuliferous scale
- seeds are usually winged
CUPRESSOPSIDA
- Araucariaceae monoecious or dioecious, broad to acicular leaves
- Cupressaceae (Cypress family) monoecious or dioecious
- Phyllocladaceae
- Podocarpaceae (Podocarp family) fruit from the fleshy receptacle ; epimatum (modified
ovuliferous scale) - covering for the seeds ; receptacle - basal stlak-like region
- Sciaopityacaea
- Taxaceae (Yew family) seed cones are reduced to one ovule

GNETALES
- Ephredraceae (Morman Tea family) double fertilization
- Gnetaceae
- Welwitshiaceae
- possible sister taxa of angiosperms due to whorled, somewhat “perianth-like”
microsporophylls that may resemble flowers
- 1 striate pollen ; vessels with porose perforation plates

EVOLUTION OF FLOWERING PLANTS

- Magnoliophyta or Antophyta or Angiospermae monophyletic , sister taxa of gymnosperms
- most numerous, diverse and successful
- 95% of all land plant

MAJOR APOMORPHIES :
- flower, with perianth
- stamens with 2 lateral thecae
- reduced, 3-nucleate male gametophyte
- carpels and fruit formation
- ovules with 2 integuments
- reduced, 8-nucleate female gametophyte
- endosperm formation
- sieve tube members with companion cells
FLOWER
- a modified, determinate shoot system bearing one or more stamens (collectively called
androecium) and or one or more carpels (group of pistils, collectively called gynoecium)
- most are bisexual (perfect) containing both stamens and carpels
- some are uniseuxal (imperfect), having only stamens or carpels
- Perianth protects the floral parts during floral development and functions as an attractant for
pollination
- Perianth = outer calyx (green and photosynthetic, composed of leaf-like sepals or calyx
lobes(if fused)) + inner corolla (colorful, showy and odoriferous and is composed of
individual petals or corolla lobes (if fused))
- Tepal = outer tepals (sepal) and inner tepals (petal) ; maybe called perianth
- Arabidopsis tahliana (Drosophilia of the plant world) ; ABC model of floral development

STAMENS
- modified microsporophylls, modified leaves that bear microsporangia
- stalk - filament (lamina)
- pollen bearing part - anther
- some stamens lack a filament, in this case anther is sessile and directly attached to the flower
- angiosperm anther is a type of synangium (fusion of sporangia)
- containing 2 pairs of microsporangia usually arranged in a bilateral symmetry
- each pair of microsporangia is typically located within a discrete half of the anther called
theca (thus an anther consists of 2 thecae)
- each theca have 2 microsporangia for a total of 4 (tetrasporangiate)
- at maturity 2 microsporangia coalesce into anther locule
- monothecal or bisporangiate , anthers are secondarily reduced to a single theca

REDUED MALE GAMETOPHYTE

- reduced, 3 celled male gametophyte
- microsporangium - meiosis - microspore - mitosis - tube cell and generative cell - pollen grain
- generative cell mitosis - 2 sperm cells
- pollen tube elongates through the tissues of the stigma and style of the carpel and soon
reaches the ovule
- sperm cells lack flagella or cilia
- adaptation is due to the evolution of reduced female gametophyte and relatively rapis seed
development
- fertilization occurs very soon compared to gymnosperms

CARPEL
- a modified, conduplicate megasporophyll bearing 2, adaxial rows of ovules ; receiver of pollen
(selectively control fertilization)
- at maturity the carpel body completely encloses the ovules and seeds, accounting for the
name angiosperm
- apocarpus carpels are separated from one another (distinct)or fused together syncarpous
- alternatively called as a unit of gynoecium (one or moe pistils)
- each pistil is composed of a basal ovary, an apical style (maybe absent) and one or more
stigmas (tissue receptive of pollen grains)
- pistil = one carpel (simple or maybe compound)
- Incompatibility reaction chemicals that are present in the stigma or style that may inhibit
either pollen germination or pollen tube growth
- Fruit the mature ovary or ovaries (made up of one or more carpels) + any accessory tissue
- Pericarp mature ovary wall

TWO INTEGUMENTS
- ovules are know as bitegmic
- nonflowering seed plant are unitegmic
- 2 integuments surround the nucellus, forming a small pore at the distal end (micropyle)
- seed coat

REDUCED FEMALE GAMETOPHYTE

- megasporangium - meiosis - 4 haploid megaspores - female gametophyte generates from
only of these megaspore
- megaspores divide in a sequence of 3 mitotic division resulting a total 8 haploid nuclei

ENDOSPERM FORMATION
- endosperm is the product of double fertilization
- pollen tube enters the micropyle, it penetrates one of the synergid cells and releases the 2
sperm cells into the central cell of the female gametophyte
- one sperm cell migrates toward and fuses with the egg producing a diploid zygote
- the other sperm cell fuses with the 2 polar nuclei to produce a triploid or 3n endosperm cell
- endosperm cell mitotically divides repeatedly to from endosperm
- endosperm replaces the female gametophyte as the primary nutritive tissue for the embryo
SIEVE TUBE MEMBERS
- specialized sugar-conducting cells ; found only in flowering plants
- sieve cells are the primary sugar-conducting cells in nonflowering vascular plants
- pores at the end of walls are differentiated, larger than those of the side walls
- sieve plates - differentiated pores at the end walls ; simple (composed of one pore region) or
compound (composed of 2 or more aggregations of pores)
- companion cells parenchyma cells associated with sieve tube members ; functions to load
and unload sugars into the cavity of sieve tube members
- provide more efficient sugar conduction
VESSELS
- water and sugar conduction
- 2 ends of the cells have opening perforation plates
DIVERSITY AND CLASSIFICATION OF FLOWERING PLANTS : AMBORELLALES,
NYMPHAEALES , AUSTROBAILEYALES, MAGNOLIIDS, CERATOPHYLLALES, AND
MONOCOTS

- APG III system, attempt was made

- to recognize only those angiosperm families that are monophyletic ; classifies one to several
families unto orders, where strong evidence suggests that the order is monphyletic
- Basal angiosperms are paraphyletic
- Monocots are 22% ; eudicots 75%
FLORAL FORMULAS
- used to summarize the # and fusion of floral parts
- P = perianth parts ; undifferentiated into a typical outer calyx and inner corolla
- K = # of sepals or calyx lobes
- C = # of petals or corolla lobes
- A = androecium ; # of stamens
- G = gynoecium ; # of carpels ‘ followed by superior or inferior to denote ovary position
- ( ) = fusion of similar parts
- + = separate discrete whorls, delimiting the # of parts per whorl
- [ ] = less common or rare condition
- ∞ = numerous
- EXAMPLES : K (5) [(4]) = Calyx fusion of 5, rarely fusion of 4 ; A 5+5 [4+4] = androecium 5
outer 5 inner, rarely 4 outer 4 inner ; G 5 [4] = gynoecium 5 rarely 4
- P (3+3) = perianth fusion of 3 outer tepals and 3 inner tepals ; A 3+3 = androecium 3 outer 3
inner ; G (3) = gynoecium inferior fusion of 3

AMBORELLALES
- absence of vessels and aromatic (ethereal) oil cells
- Amborella trichopoda ; dioecious ; Native to New caledonia
- Ambroellaceae are distinctive in being vessel-less, evergreen shrubs with unisexual flowers
having undifferentiated, spiral perianth, numerous, laminar stamens, and an apocarpous,
apically-open gynoecium, with 1-ovuled carpels ; Male flowers : P 5-8 A ∞ ; Female flowers :
P 5-8 G 5-6, superior

NYMPHAEALES
- Nymphaeaceae - Water lily family ; aquatic herbs with floating leaves and solitary, floating to
emergent flowers with mostly spiral floral parts and petal grading into usually laminar stamens
; K 4-6 [-14] C 8-∞ [0] A ∞ B (3-∞)
- Cabombaceae - Fanwort family ; aquatic herbs with atactostelic stems ; dimorphic, floating or
submerse, undivided or higjhly divided leaves ; perianth with 3 [2,4] sepals and petals and an
apocarpous gynoecium ; K 3 [2,4] C 3 [2,4] A 3,6 or 12-∞ G 2-18 [1], superior

AUSTROBAILEYYALES
- Illiciaceae ; evergreen tress or shrubs having aromatic oil cells, with glabrous, spiral, pellucid-
punctate, exstipulate leaves, the flowers with numerous, spiral tepals (outer sepal-like, inner
petal-like) , apocaropous pistils in a single whorl, the fruit a follicetum ; P ∞ [7-33] A ∞ [4-50]
G ∞ [5-21], superior

MAGNOLIIDS
- LAURALES
- Lauraceae - Laurel family ; perennial tress or shrubs with aromatic oil glands, evergreen
leaves, an undifferentiated perianth, valvular anther dehiscence, and a single, superior
ovary having one ovule per carpel with apical placentation, seeds lacking endosperm ; P
3=3 [6, 2+2, or 3+3+3] A 3-12+ G 1[-(3)], superior, rarely inferior, hypanthium present
- MAGNOLIALES
- Annoncaceae - Custard-apple family ; tress, shrubs, or woody vines with simple, usually
simple distichious leaves, a trimerous perianth, numerous, usually spiral stamens and
pistils (apocarpous or syncarpous) and seeds with ruminate endosperm ; P 3+3+3 A ∞ G
∞ , superior
- Magnoliaceae - Magnolia family ; tress and shrubs with simple stipulate leaves, solitary
flower, a usually undifferentiated petaloid perianth with numerous tepals, numerous, spiral
 stamens, and an apocarpous gynoecium of numerous, spiral pistils born on elongate
receptacular axis, the fruit an aggregate of follicle, berries, or samara, seeds usually with a
sarcotesta ; P ∞ A ∞ G ∞ [2-∞], superior

- PIPERALES
- Aristolochiaceae - Birthwort family ; usually climbing plants, having an enlarged, petaloid
calyx, an absent to reduced corolla, often adnate stamens (forming a gynostemium), and
an inferior to superior, 3–6-carpeled and loculed ovary ; K (3) C 0 [3] A 6–°, usu. adnate to
style G (3–6), inferior, half-inferior, or superior.

- Piperaceae - Pepper family ; having an atactostelic stem, a spike or spadix with

numerous, very small, unisexual or bisexual flowers lacking a perianth, the ovary solitary,
1-ovulate, the fruit a 1-seeded berry or drupe ; P 0 A 3+3 [1–10] G 1 or (3,4), superior.

- Saururaceae - Lizard’s tail family ; perennial herbs with a bracteate spike or raceme and
with flowers lacking a perianth, the ovary solitary, many-ovulate, the fruit a capsule ; P 0 A
3, 3+3, or 4+4 G (3–5), superior.

CERATOPHYLLALES
- Ceratophyllaceae - Hornwort family ; monoecious, aquatic herbs with whorled,
dichotomously branched, serrulate leaves, and solitary, unisexual flowers ; P (8–12) A 5–27
G 1, superior.
MONOCOTYLEDONS
ACORALES
- Acoraceae - Sweet flag family ; marsh plants with a spadix and spathe (resembling
Araceae) but having distichous, ensiform, unifacial leaves, perispermous and endospermous
seeds, and ethereal oil cells, and in lacking raphide crystals ; P 3+3 A 3+3 G (2–3) superior.

ALISMATALES
- Araceae - Arum family ; having bifacial leaves with parallel or netted venation, usu- ally a
spadix of numerous, small flowers with a subtending spathe, endospermous seeds, and
 raphide crystals ; P 2+2,3+3,(2+2),(3+3) or 0 [4+4,(4+4)] A 4,6,8 or (4,6,8) [1–12] G (3) [1–(°)]
superior.
- Alismataceae - Water-Plantain family ; aquatic or marsh herbs, with solitary or often whorled
flowers or flower axes, and dichlamydeous flowers with an apocarpous gynoecium having
basal placen- tation, the fruit an aggregate of achenes or follicles.
K 3 C 3 A6,9–°[3] G3–° superior.

PETROSAVIALES
- Petrosaviaceae - 4 species in 2 gener

DIOSCOREALES
- Dioscoreaceae - Yam family ; perennial, hermaphroditic or dioecious, rhizomatous or
tuberous herbs with simple to palmate leaves having net venation and epigynous, trimerous
flowers ; P 3+3 A 3+3 or 3+0 G (3), inferior, hypanthium absent or present.

PANDANALES
- Pandanaceae - Screw-pine family ; mostly dioecious, sympodially branched, woody plants
with prop roots, 3- or 4-ranked, simple, acrocaulis, linear to ensiform leaves (appear- ing
spiral), and small, usually unisexual flowers of variable morphology, the fruit a berry or drupe,
multiple in some ; P (3–4) or 0 A ° (male) G 1(–°) (female), superior.

LILIALES
- Liliaceae - Lily family ; perennial, usually bulbous herbs, lacking an onion-like odor, with basal
or cau- line leaves, the inflorescence a raceme, umbel or of solitary flowers with a superior
ovary ; P 3+3 A 3 G (3), superior.
ASPARAGALES
DIVERSITY AND CLASSIFICATION OF FLOWERING PLANTS : EUDICOT