Plant Development

Unit II
How plant development differs from that of
animal?
1. Plants do not gastrulate. Plant cells are trapped within rigid cellulose
walls that generally prevent cell and tissue migration.
2. Plants have sporic meiosis rather than gametic meiosis. That is,
spores, not gametes, are produced by meiosis.
3. The life cycle of land plants (as well as many other plants) includes
both diploid and haploid multicellular stages
4. Germ cells are not set aside early in development
5. Plants undergo extended morphogenesis.
6. Plants have tremendous developmental plasticity.
7. Plants may tolerate higher genetic loads than animals.
Plant Life Cycles
• The plant life cycle alternates between haploid and diploid
generations.
• Embryonic development is seen only in the diploid generation. The
embryo, however, is produced by the fusion of gametes, which are
formed only by the haploid generation.
• plants have multicellular haploid and multicellular diploid stages in
their life cycle. Gametes develop in the multicellular haploid
gametophyte.
• Fertilization gives rise to a multicellular diploid sporophyte, which
produces haploid spores via meiosis. This type of life cycle is called a
haplodiplontic life cycle
Angiosperms

• The shoot meristem of the sporophyte produces a series of vegetative structures.

• At a certain point in development, internal and external signals trigger a shift from vegetative to
reproductive (flower-producing) development.
• Once the meristem becomes floral, it initiates the development of floral parts sequentially in
whorls of organs modified from leaves.
• The first and second whorls become sepals and petals, respectively; these organs are sterile.
• The pollen-producing stamens are initiated in the third whorl of the flower.
• The carpel in the fourth whorl contains the female gametophyte. The stamens contain four
groups of cells, called the microsporangia (pollen sacs), within an anther.
• The microsporangia undergo meiosis to produce microspores. Unlike most ferns, angiosperms
are heterosporous, so the prefix micro is used to identify the spores that mitotically yield the
male gametophytes pollen grains.
• The inner wall of the pollen sac, the tapetum, provides nourishment for the developing pollen.
• Gamete Production in
Angiosperms
• The sporangia of angiosperm
plants are found in two places
within the flower depending,
on whether they lead to a
female or male gametophyte.
• The sporangium in the anther
(where the male gametophyte
will be formed) is called the
microsporangium.
• The sporangium in the ovary
(where the female
gametophyte will be formed)
is called the megasporangium.
• Microsporogenesis
• The process of formation of microspores or pollen grains from
microspore mother cell inside the microsporangium/pollen
sac of anther is termed as microsporogenesis
• The inner most cells of the anther are the sporogenous tissue
and they serve as mother cell. Pollen mother cell is diploid in
nature.
• Initially, it undergoes meiosis yielding four haploid cells.
• Mostly all the four spores within a tetrad are totally
segregated from one another and form the spores.
• Each pollen grain bears haploid nucleus in the pollen cytoplasm
surrounded externally by the plasmalemma and pollen wall.
• Pollen wall consists of outer uneven exine and inner even intine wall.
• This exine layer plays a vital role during pollination as they give species identity.
• During gametogenesis, the pollen nucleus divides mitotically to yield
the unequal cells:
– The larger one is vegetative cell or tube cell which forms the
pollen tube.
- The smaller one, situated towards the wall is generative cell
which again divides by mitosis to form two sperms (male gametes).
• The exine ruptures and the intine forms a pollen tube.
• The vegetative nucleus enters the pollen tube. Now it is
known as tube nucleus.
• The generative tube nucleus enters the tube and divides to
form two generative nuclei which finally form the two male
gametes.
• The tube nucleus disorganizes.
• One of the gamete fuses with the female gamete during
fertilization.
• Megasporogenesis
• the development of the female gametophyte, a process is
called megasporogenesis or female gametogenesis
• The chambers within the ovary are the locules, and within the locules are the
ovules. The ovule wall is made up of integument tissue, and it is from this
integument tissue that diploid megaspore mother cells form.
• A diploid megaspore mother cell undergoes meiosis and initially produces four
haploid megaspores. Three of the four spores formed through meiotic division of
the megaspore mother cell degenerate.
• The lone surviving megaspore subsequently undergoes three mitotic divisions to
form eight haploid cells that are held within the ovule in an embryo sac. These
eight cells comprise the female gametophyte.
• The megasporangium together with the integuments, is called
ovule.
• It is affixed to the placenta, on the inner wall of the ovary, by a
stalk called funiculus.
• An ovule ready for fertilization constitutes of nucellar tissue
covered almost completely by one or two integuments, leaving a
small opening at the apical end.
• This opening termed as micropyle is the main passage for the
entry of the pollen tube in the ovule.
• The basal region of the ovule where funiculus is affixed is called
chalaza.
• Only six of the nuclei are surrounded by their own cell membrane
after the three rounds of mitosis. The two remaining nuclei are
surrounded by one cell membrane.
• The 3 cells (antipodal cells) move opposite to micropyle, which later
degenerates.
• Two polar nuclei fuse to form a central cell, which upon fusion with
sperm nucleus forms endosperm. This is called a triple fusion.
• The egg cell is supported by two other cells called synergids (guides
pollen tube during fertilization), present at micropyle, an opening for
the entry of the sperm nucleus during fertilization.
• Fertilization
• Fertilization is the process of fusion of haploid male and female
gametes to form a diploid zygote, giving rise to the embryo. In
angiosperms, the fertilization takes place in the ovule, enclosed within
the ovary of the flower, giving rise to seed and fruit respectively.
• When the pollen grain encounters the sticky surface of the stigma,
the pollen grain develops a pollen tube from the tube cell. There is a
certain region called germ pore, on the pollen grain which enables the
exit of the intine layer and the protrusion of pollen tube outside the
pollen. The pollen tube traverses the style region and reaches the
micropyle, an opening in the embryo sac.
• The generative cell forms two male gametes, which travels through the
pollen tube along with the vegetative nucleus. The pollen tube enters
micropyle and synergids guide the pollen tube to the egg by releasing few
chemotropic secretions for fusion or sometimes pollen tube enters synergids
and ruptures to release the vegetative nucleus and 2 sperm nuclei.
• The egg cell fuses with one of the sperm nuclei to form a zygote and another
sperm nucleus fuses with polar nuclei to form endosperm. This process is
known as double fertilization. And the fusion of polar nuclei with the sperm
nucleus is called triple fusion. The ovule matures to form seed, the
integuments form the seed coat. The pollen tube enters through chalaza
(chalazogamy), through micropyle (porogamy) or through integuments
(monogamy).
• The endosperm is present during seed development and provides nutrients
for either the developing embryo, the germinating seedling, or both.
• Fertilization also causes the ovule, containing the embryo and endosperm,
to develop into a seed and the ovary to differentiate into a fruit, which
facilitates seed dispersal .
• A mature flowering plant embryo contains two primary organ systems-the
axis and cotyledon.
• These organs have distinct developmental fates and are both composed of
three basic, or primordial, tissue layers-protoderm, procambium, and
ground meristem-which will become the epidermal, vascular, and
parenchyma tissues of the young seedling, respectively
• The axis, or hypocotyl-radicle region of the embryo, contains the
shoot and root meristems and will give rise to the mature plant after
seed germination.
• By contrast, the cotyledon is a terminally differentiated organ that
accumulates food reserves that are utilized by the seedling for growth
and development before it becomes photosynthetically active.
• The cotyledon functions primarily during seed germination and
senesces shortly after the seedling emerges from the soil.
• Plant embryogenesis can be divided into three general phases in
which distinct developmental and physiological events occur:
• (i) postfertilization-proembryo,
• (ii) globular-heart transition, and
• (iii) organ expansion and maturation
• The zygote in Arabidopsis has an asymmetric distribution of cellular
components-the nucleus and most of the cytoplasm are present in the upper
portion of the cell, whereas a large vacuole dominates the middle to lower
portion.
• This spatial asymmetry is derived from the egg cell. The zygote divides
asymmetrically into two distinct-sized daughter cells a small, upper terminal
cell and a large, lower basal cell-which establish a polarized longitudinal axis
within the embryo.
• The small terminal cell gives rise to the embryo proper that will form most of
the mature embryo. Cell lineages derived from the terminal cell and embryo
proper will specify the cotyledons, shoot meristem, hypocotyl region of the
embryonic axis, and part of the radicle, or embryonic root.
• By contrast, the large basal cell derived from the lower portion of the zygote will
divide and form a highly specialized, terminally differentiated embryonic organ
called the suspensor.
• In Arabidopsis, the suspensor contains only 7 to 10 cells. The suspensor anchors
the embryo proper to the surrounding embryo sac and ovule tissue and serves as
a conduit for nutrients Postfertilization to be passed from the maternal
sporophyte into the developing proembryo.
• The suspensor senesces after the heart stage and is not a functional part of the
embryo in the mature seed.
• The uppermost cell of the suspensor, forms the hypophysis, contribute to the
formation of the root meristem. Thus, cell lineages derived from the basal cell
give rise to the suspensor and part of the radicle region of the embryonic axis.
• Embryonic organs and tissue-types differentiate during the globular-heart transition phase.
Two critical events must occur after the embryo proper forms:
• (i) regions along the longitudinal apical-basal axis must differentiate from each other and generate
embryonic organ systems, and
• (ii) the three primordial tissue layers of the embryo need to be specified. Both of these events take
place during the globular-heart transition phase.
• The embryo proper has a spherical shape during the proembryo and globular stages. The
first visible cell differentiation events occur at the 16- cell stage when the protoderm, or
outer cell layer of the embryo proper, is produced and the hypophysis forms at the top of
the suspensor.
• Subsequent cell differentiation events within the embryo proper result in the production of
an inner procambium tissue layer and a middle layer of ground meristem cells.
• The spatial organization of protoderm, ground meristem, and procambium layers
establishes a radial axis of differentiated tissues within the globular embryo.
• A dramatic change in the morphology of the embryo proper occurs just after
the globular stage.
• Cotyledons are specified from two lateral domains at the apical end (top), the
hypocotyl region of the axis begins to elongate, and the root meristem
becomes differentiated from the hypophysis region at the basal end (bottom)
(34).
• The embryo proper is now heart-shaped, has a bilateral symmetry, and the
body plan and tissue layers of the mature embryo (and postembryonic plant)
have been established.
• Morphogenetic changes during this period are mediated by differential cell
division and expansion rates and by asymmetric cleavages in different cell
planes
• Upright cotyledons can give the embryo a torpedo shape, and by this
point the suspensor is degenerating and the shoot apical meristem and
room apical meristem are established. These meristems will give rise
to the adult structures of the plant upon germination. Further growth of
the cotyledons results in the torpedo and walking-stick stages. At this
point, embryogenesis is arrested, and the mature seed dessicates and
remains dormant until germination.
A typical dicotyledonous embryo
consists of an embryonal axis and
two cotyledons. The part of
embryonal axis above the level of
cotyledons is called epicotyl.
It terminates with the stem tip,
called plumule (future shoot).
The part below the level of
cotyledons is called hypocotyl
which terminates in the root tip
called radicle (future root).
The root tip is covered with a root
cap (calyptra).
• Mutants have been identified that result in changes in the
establishment of the apical-basal pattern (organization of organs along
the apical-basal axis) and the radial pattern (organization of the three
basic tissue systems - dermal, ground, and vascular).
• Arabidopsis mutant seedlings were identified that showed a loss or
distortion of the root, hypocotyl or cotyledon regions. These defects are
presumed to result from defects during embryogenesis. These mutants were
then placed into the following major classes:
• mutants lacking body segments along the apical-basal axis. This class
includes gurke (gk), fackel (fk), monopterous (mp), and gnom (gn).
• mutants with disturbed radial symmetry - alterations of the radial pattern of
tissue layers. This class includes knolle (kn) and keule (keu).
• mutants with disrupted organogenesis - these mutants have grossly abnormal
overall shapes, but have all of the pattern elements along the apical-basal
and radial axes. This class includes fass (fs), knopf (knf), and mickey (mic).
• The vegetative and sexual modes of reproduction are generally
normal and occur in nature. There are cases of special modes of
reproduction where propagation takes place without the act of
fertilisation.
• According to Winkler (1908, 1934), the substitution for sexual
reproduction of an asexual process which does not involve any
nuclear fusion.
• The plants where the usual sexual reproduction has been
completely replaced by a type of asexual reproduction are called
apomictic, and the phenomenon, apomixis.
• (i) Nonrecurrent apomixis,
• (ii) recurrent apomixis, and
• (iii) Adventive embryony.
• i. Non-recurrent apomixes:
• In this type the megaspore mother cell undergoes the usual meiotic divisions
and a haploid embryo sac is formed. Here the embryo arises either from the
egg (i.e., haploid parthenogenesis) or from some other cell of the
gametophyte (i.e., haploid apogamy).
• The plants produced by this method are, haploid and generally sterile and do
not reproduce sexually any more. This type of apomixis has been seen in
several species such as Solarium nigrum, Lilium, Bergenia, Erythraea
centaurium. Orchis maculata, Nicotiana tabacum, etc.
• ii. Recurrent apomixes:
• In this type, the embryo sac generally arises either from an
archesporial cell (i.e., generative apospory) or from some other
part of the nucellus (i.e., somatic apospory). Here all the nuclei of
the embryo sac are diploid, and there is no meiotic division.
• The embryo arises either from the egg (diploid parthenogenesis)
or from some other cell of the gametophyte (diploid apogamy).
Generative apospory has been observed in Eupatorium
glandulosum, Parthenium argentatum, etc. Somatic apospory has
been reported in Hieracium excelens, H. flagellare and H.
aurantiacum.
• iii. Adventive embryony:
• This type of apomixis is also known as sporophytic budding.
Here, the developed embryo sacs may be haploid or diploid, but
the embryos do not arise from the cells of nucellus or the
integument. There is no alternation of generations, because the
diploid tissues of the present sporophyte directly give rise to the
new embryo.
• Adventive embryony has been frequently reported in Citrus,
Euphorbia dulcis, Capparis frondosa, Mangifera indica and
Hiptage madablota.
• Parthenogenesis:
• The development of the zygote from the egg-cell without the act of
fertilization, or in other words, formation of embryo from an unfertilized
egg is called parthenogenesis.
• In some species of angiosperms, the embryo develops parthenogenetically.
• In such cases the embryo may develop from haploid egg-cell or diploid egg-
cell. In apomicts of Asteraceae and Rubiaceae, the development of embryo
is independent of the pollination stimulus.
• However, in many other apomicts, the embryo develops only after
pollination, and the phenomenon is known as pseudogamy, e.g., many
apomictic grasses.
• Parthenocarpy:
• In certain cases of angiosperms the ovary normally develops into a fruit without
pollination and fertilization. This type of free development of fruit is known as
parthenocarpy. Such fruits (parthenocarpic fruits) are always seedless.
Sometimes the fruit formation may be induced by artificial pollination by
foreign pollen from another species, but without subsequent fertilization.
• The parthenocarpy may also be induced by the spraying of growth promoting
substances, such as naphthalene acetic acid NAA. This is called induced
parthenocarpy. The examples of parthenocarpy are commonly found in banana,
papaya, pineapple, guava, grapes, apple, Thalictrum, Alchemilla, etc.
• In modern days, a lot of work has been done to produce seedless fruit on
seeded varieties by controlling pollination and applying certain chemical
substances to the pistil.
Parthenocarpy Parthenogenesis
Development of fruit without prior Development of a new organism without
fertilization fertilization of an ovum

Seedless fruits are produced Organisms produced are a female clone

and cannot reproduce sexually

Offsprings are not produced Haploid offsprings are produced

Occurs only in plants Occurs in invertebrate animals and lower

plants

Examples: banana, watermelon, pineapple, Examples: insects, lizards, etc.

etc.
• The process of seed germination includes the following five
changes or steps.
• Such five changes or steps occurring during seed germination are:
• (1) Imbibition
• (2) Respiration
• (3) Effect of Light on Seed Germination
• (4) Mobilization of Reserves during Seed Germination and Role of
Growth Regulators and
• (5) Development of Embryo Axis into Seedling.
• (i) Imbibition:
• The first step in the seed germination is imbibition i.e. absorption of water
by the dry seed. Imbibition results in swelling of the seed as the cellular
constituents get rehydrated.
• The swelling takes place with a great force. It ruptures the seed coats and
enables the radicle to come out in the form of primary root.
• Imbibition is accomplished due to the rehydration of structural and storage
macromolecules, chiefly the cell wall and storage polysaccharides and
proteins.
• Many seeds contain additional polysaccharides, not commonly found in
vegetative tissues. Seeds packed dry in a bottle can crack it as they imbibe
water and become swollen.
• (ii) Respiration:
• Imbibition of water causes the resumption of metabolic activity in the rehydrated
seed. Initially their respiration may be anaerobic (due to the energy provided by
glycolysis) but it soon becomes aerobic as oxygen begins entering the seed.
• The seeds of water plants, as also rice, can germinate under water by utilizing
dissolved oxygen.
• The seeds of plants adapted to life on land cannot germinate under water as they
require more oxygen. Such seeds obtain the oxygen from the air contained in the
soil.
• It is for this reason that most seeds are sown in the loose soil near the surface.
Ploughing and hoeing aerate the soil and facilitate seed germination.
• Thus the seeds planted deeper in the soil in water-logged soils often fail to germinate
due to insufficient oxygen.
• (iii) Effect of Light on Seed Germination:
• Plants vary greatly in response to light with respect to seed
germination. The seeds which respond to light for their germination
are named as photoblastic.
• Three categories of photoblastic seeds are recognized:
• Positive photoblastic,
• negative photoblastic and
• non-photoblastic.
• Positive photoblastic seeds (lettuce, tobacco, mistletoe, etc.) do not
germinate in darkness but require exposure to sunlight (may be for a
brief period) for germination.
• Negative photoblastic seeds (onion, lily, Amaranthus, Nigella, etc.) do not germinate if exposed
to sunlight.
• Non-photoblastic seeds germinate irrespective of the presence (exposure) or absence (non-
exposure) of light.
• In these light sensitive seeds, the red region of the visible spectrum is most effective for
germination.
• The far-red region (the region immediately after the visible red region) reverses the effect of red
light and makes the seed dormant.
• The red and far-red sensitivity of the seeds is due to the presence of a blue-coloured
photoreceptor pigment, the phytochrome.
• It is a phycobiloprotein and is widely distributed in plants.
• Phytochrome is a regulatory pigment which controls many light-dependent development
processes in plants besides germination in light- sensitive seeds.
• These include photo-morphogenesis (light-regulated developmental process) and flowering in a
variety of plants.
• The pigment phytochrome that absorbs light occurs in two inter-convertible forms
Pr and Pfr.
• Pr is metabolically inactive. It absorbs red light (660 nm.) and gets transformed
into metabolically active Pfr.
• The latter promotes germination and other phytochrome-controlled processes in
plants.
• Pfr reverts back to Pr after absorbing far-red (730 nm.). In darkness too, Pfr slowly
changes to Pr.
• Owing to this oscillation of phytochrome between Pr and Pfr status, the system has
been named as “reversible red—far-red pigment system” or in brief phytochrome
system.
• Treatment with Red light (R) stimulates seed germination, whereas far-red light
(FR) treatment, on the contrary, has an inhibitory effect.
• Abscisic acid, auxin, cytokinin function, ethylene, and
gibberellins were the five predominant plant hormones by the
mid-20th century. Researchers have also added brassinosteroids,
jasmonates, salicylates, and strigolactones to the list in recent
years.
• Light requirement for seed germination may be replaced by
hormones such as gibberellins or cytokinins.
• Several development processes of plants controlled by
phytochrome may be mimicked by appropriate hormones given
singly or in combination with other hormones at the correct time.
• Auxin is involved in cell growth and expansion, plants typically generate it
in regions that are actively developing, such as the stem. This is when things
get interesting. Auxin travels in a plant through just one route — downhill
from the top to the bottom, similar to a one-way road from the stem tip to the
roots. It is the only plant hormone that can do so.
• There are both natural and synthetic forms of auxin:
•
Natural: Indole-3-Acetic acid, Indole-3-Butyric Acid
• Synthetic: 2,4-Dichlorophenoxyacetic acid, 1-Naphthyl Acetic Acid
• Some of the functions of the Auxin Hormones are as follows:
•
Cell elongation of stems and roots.
• It induces Parthenocarpy, or the formation of fruit without fertilization, for
example, in tomatoes.
• Prevents leaves, flowers, and fruits from falling prematurely.
• It is useful for initiating roots in stem cuttings and grafting.
• Promotes blooming, for example, in pineapple.
• We often use it as a herbicide to destroy noxious weeds of dicot plants while
causing no harm to monocot plants.
• Contributes to cell division and xylem differentiation.
• Gibberellins serve a crucial function in many phases of plant growth, but their claim to fame is
that they make stems longer. Gibberellins encourage stem growth between nodes. It elongates the
internodes, which are the places on a stem where a leaf attaches.
•
The lack of gibberellin is most visible in dwarf and rosette plants, where there is a tiny space
between nodes on a stem, and it groups the leaves around the plant's base.
•
Some of the functions of the Gibberellin Hormones are as follows:
•
It encourages bolting or the abrupt extension of internodes right before blooming in rosette plants
such as cabbage and beets.
• Postpones Senescence.
• Induces Parthenocarpy.
• It can reverse Dwarfism through stem lengthening.
• It induces certain plants to become male, such as cannabis.
• In the endosperm of sprouting cereal grains and barley seeds, it promotes the development of
hydrolytic enzymes such as lipase and amylase.
• Breaks the dormancy of seeds.
• The chemical is known as cytokinin, and it is involved in cell division and the
formation of new plant parts such as roots and shoots. Cytokinins are created in
the root apical meristems and migrate up the stem through the xylem, hitching a
ride with water. The movement of cytokinins is passive - no energy is required.
•
Some of the functions of the Cytokinin Hormones are as follows:
•
It stimulates lateral and adventitious shoot development and is used in culture to
begin shoot growth.
• Aids in the removal of apical dominance caused by auxins.
• Stimulate chloroplast development in leaves.
• Encourages nutrient mobilization and postpones leaf senescence.
•
Kinetin is an example of Cytokinin
• Ethylene is a plant hormone that influences plant ripening and rotting. Because it occurs
as a gas, it is a particularly intriguing plant hormone. There is no other plant hormone that
is gaseous. Plants can produce ethylene in almost any part and can diffuse through its
tissue outside the plant, and travel through the air to affect a completely different plant.
•
Some of the functions of the Ethylene Hormones are as follows:
•
It speeds up the ripening of fruits.
• Controls leaf epinasty.
• Breaks the dormancy of seeds and buds.
• Accelerates the elongation of petioles and internodes.
• Promotes leaf and blossom senescence and abscission.
• Increases the absorption surface by stimulating root development and root hair
production.
• In monoecious plants, it stimulates femaleness.
• Formation of apical hooks in dicot seedlings.
• Plants create abscisic acid, a chemical messenger, to inform the rest of the
plant that it is water-stressed. Abscisic acid is produced in droughted leaves,
roots, and developing seeds and can move up and down a plant stem in the
xylem or phloem, sounding the alarm.
•
Some of the functions of the Abscisic Acid Hormones are as follows:
•
Causes leaf and fruit abscission.
• It inhibits seed germination.
• Induces leaf senescence.
• Accelerates dormancy in seeds, which is important for storage.
• Stimulates stomatal closure to prevent transpiration during water stress.
• (iv) Mobilization of Reserves during Seed Germination and
Role of Growth Regulators:
• During germination the cells of the embryo resume metabolic activity
and undergo division and expansion. Stored starch, protein or fats need
to be digested. These cellular conversions take place by making use of
energy provided by aerobic respiration.
• Depending upon the nature of the seed, the food reserves may be stored
chiefly in the endosperm (many monocotyledons, cereal grains and
castor) or in the cotyledons (many dicotyledons such as peas and beans).
Thorough investigations in the mobilisation of reserves from the
endosperm to the embryo via a shield-like cotyledon (scutellum) has
been done in several cereal grains
• The outer layer of special cells (aleurone layer) of endosperm
produces and secretes hydrolyzing enzymes (such as amylases,
proteases). These enzymes cause digestion i.e. breakdown of the
stored food such as starch and proteins in the inner endosperm
cells.
• The insoluble food is rendered soluble and complex food is
made simple. These simpler food solutions, comprising of
sugars and amino acids thus formed, are diluted by water and
passed towards the growing epicotyl, hypocotyl, radicle and
plumule through the cotyledon.
• Gibberellic acid plays an important role in initiating the
synthesis of hydrolyzing enzymes. Gibberellin, therefore,
promotes seed germination and early seedling growth.
Assimilation of this food by the growing organ induces growth
and the seedling soon assumes its ultimate shape.
• It is very significant to note that the dormancy inducing
hormone, abscisic acid (ABA), prevents the germination. The
concentration of ABA has been shown to increase during the
onset of dormancy of the embryo during seed development in
several kinds of seeds.
• (v) Development of Embryo Axis into Seedling:
• After the translocation of food and its subsequent assimilation,
the cells of the embryo in the growing regions become
metabolically very active. The cells grow in size and begin
divisions to form the seedling.
• Meristems
• Meristems are clusters of cells that allow the basic body pattern
established during embryogenesis to be reiterated and extended after
germination.
• Meristematic cells are similar to stem cells in animals.* They divide to
give rise to one daughter cell that continues to be meristematic and
another that differentiates.
• Meristems fall into three categories: apical, lateral, and intercalary.
• Apical meristems are located at the growing tips of the adult plant,
and produce root and shoot tissue. Shoot apical meristems
(SAM) initiate leaves during vegetative development,
and inflorescence (IM) and floral meristems (FM) during
reproductive development.
• Apical meristems occur at the growing shoot and root tips. Root apical meristems
produce the root cap, which consists of lubricated cells that are sloughed off as the
meristem is pushed through the soil by cell division and elongation in more proximal
cells. The root apical meristem also gives rise to daughter cells that produce the three
tissue systems of the root.
• New root apical meristems are initiated from tissue within the core of the root and
emerge through the ground tissue and dermal tissue. Root meristems can also be
derived secondarily from the stem of the plant; in the case of maize, this is the major
source of root mass.
• The shoot apical meristem produces stems, leaves, and reproductive structures. In
addition to the shoot apical meristem initiated during embryogenesis, axillary shoot
apical meristems (axillary buds) derived from the original one form in the axils (the
angles between leaf and stem). Unlike new root meristems, these arise from the surface
layers of the meristem.
• There are approximately 100 cells in the SAM of Arabidopsis thaliana. These
cells are organized in two ways: cells are organized in radial zones and also in
layers.
• Radial organization:
Cells that are at the very top of the meristem divide infrequently. This region is
called the central zone. This is the location of the self-renewing undifferentiated
stem cells. Surrounding the central zone is the peripheral zone. The rate of cell
division in the peripheral zone is higher than that of the central zone. Peripheral
zone cells give rise to cells which contribute to the organs of the plant, including
leaves, inflorescence meristems, and floral meristems. Below the central zone is
another region of rapidly dividing cells, called the rib meristem. Division and
elongation of rib meristem cells gives rise to the stem of the plant.
• Layer organization:
The surface layer(s) of cells (L1) divides only by forming anticlinal cell walls - that
is, cell division is always perpendicular to the meristem surface. As a results, cells
in the L1 layer and their daughter cells always remain in this layer - all of the cells
in the L1 layer are clonally related. The L2 cell layer(s) below the L1 cells behave
in just the same way. The remaining cells (L3 or corpus cells) divide in all planes,
and fill the interior of the meristem.
• The figure below shows an inforescence shoot apical meristem (SAM) and two
adjacent floral meristems (FM) of Arabidopsis thaliana. On the left is the original
laser scanning confocal microscope optical section of tissue stained with propidium
iodide to show the nuclei. The center image was colored to show radial
zonation within the SAM. The central zone (CZ) is shown in red, the peripheral
zone (PZ) in green, and the rib meristem (RM) in blue.
The image on the right was
colored to show clonally-
related layers. The epidermal
L1 layer is shown in blue,
the subepidermal L2 layer is
shown in pink, and the L3
layer, or corpus is shown
in gold. The L1 and L2 together
are called the tunica.
• One way of investigating the contributions of different layers to plant
structure is by constructing chimeras.
• Plant chimeras are composed of layers having distinct genotypes with
discernible markers.
• When L2, for example, has a different genotype than L1 or L3, all
pollen will have the L2 genotype, indicating that pollen is derived
from L2.
• The size of the shoot apical meristem is precisely controlled by
intercellular signals, most likely between layers of the meristem.
• Mutations in the Arabidopsis CLAVATA genes, for example, lead to
increased meristem size and the production of extra organs.
• STM has the opposite effect, and double mutant phenotypes are
consistent with the hypothesis that the two work together to
maintain meristem size.
• Perhaps they balance the rate of cell division (which enlarges the
meristem) and the rate of cell differentiation in the periphery of the
meristem (which decreases meristem size)
• Lateral meristems
• Vascular cambium. Some plants grow in diameter by producing new tissues laterally from a
cylinder of tissue called the vascular cambium, which extends throughout the length of the plant
from the tips of the shoots to the tips of the roots. It is present in all perennial and in
some annual plants. Tissues produced by cell divisions of the vascular cambium are secondary
tissues.
• Cork cambium. Cork cambia (singular: cambium), also called phellogens, are found in the bark
of roots and stems of woody plants where they produce cork cells. The cork cambia originate just
under the epidermis of the primary body and in some tree species are long cylinders running
parallel to the vascular cambium. In other species, more discrete, disk-like cork cambia in the
trunks produce flat plates of bark tissues that break off in large scales as the tree ages.
• Intercalary meristem. Grasses have intercalary meristems located along the stems near the nodes.
Cell divisions in this tissue push the stem upward. Grasses and other monocots have no lateral
meristems so any lateral increase in size is the result of primary tissue cell enlargement, not cell
divisions.
• Arabidopsis mutants that display altered shoot apical meristem
structure have been identified.
• Shoot meristemless (stm) mutants are shoot meristemless - mutations
in the STM gene completely block the initiation of the SAM during
embryogenesis, but have no other obvious effects on embryo
development.
• Wuschel mutants have a flat SAM. One result of this mutation is the
formation of flowers with fewer organs. The wuschel mutant flower on
the right has only one stamen, and no central pistil. The wild type
flower on the left has 6 stamens and a central pistil.
• The root apical meristem, or root apex, is a small region at the tip of a
root in which all cells are capable of repeated division and from which all
primary root tissues are derived.
• The root apical meristem is protected as it passes through the soil by an
outer region of living parenchyma cells called the root cap.
• As the cells of the root cap are destroyed and sloughed off, new cells are
added by a special internal layer of meristematic cells called
the calyptrogen.
• Root hairs also begin to develop as simple extensions of cells near the root
apical meristem.
• They greatly increase the surface area of the root and facilitate the
absorption of water and minerals from the soil.
• Beginning with the root cap and leading away from the root tip, there are three
distinct zones in which certain specific growth patterns dominate: cell
division, cell elongation, and differentiation and tissue maturation.
• There is a gradual transition between these regions. The region of cell division
includes the apical meristem and the primary meristems—the protoderm,
ground meristem, and procambium—derived from the apical meristem.
• As is generally true of non meristematic regions elsewhere in the plant body,
root length in the second region is increased as a result of cell elongation
rather than by cell division.
• The region of differentiation and tissue maturation that follows is where the
cells differentiate (i.e., change in structure and physiology into cells of a
specific type) and where the first primary phloem and xylem, as well as mature
root hairs, are clearly seen.
• In plants with woody roots (i.e., those of perennial dicotyledons), secondary
growth, including secondary xylem and phloem as well as the periderm, are
developed and add girth to the plant.
• Root cap is made up of parenchymatic cells that release
substances containing glycosominoglycans, which work as
lubricants. Some cells of the rootcap, known as statocysts, contain
dense cytoplasmatic inclusions, mostly starch granules, that settle
at the bottom part of the cell and are involved in the grow of the
root toward the center of the Earth. The outermost cells of the root
cap die and are replaced by new ones coming from the meristem.
• Besides the cells that divide continuously, root apical meristem
contains a group of inactive cells that form the so-called quiescent
center. Not all authors describe this center, but others propose
that constitutes a reservoir of cells to be used in the regeneration
of damaged meristematic areas.
• Shoot system
Characteristics of Stem
• (i) Arises as a prolongation of plumule (one end of an embryo).
• (ii) Grows and bends towards light (positively phototropic) and away from gravity
(negatively geotropic).
• (iii) Divided into nodes (point of attachment of leaf) and internodes (regions
between two nodes).
• (iv) Bears leaves, branches and flowers on nodes.
• (v) Bears vegetative buds which could be terminal (apical bud) for plant to grow
upwards or axillary (bud in the axil of leaf) which give rise to lateral branches.
• (vi) Bears floral buds (terminal or axillary) that grow into flowers.
• The unique aboveground architectures of different plant species have their
origins in shoot meristems. Shoot architecture is affected by the amount of
axillary bud outgrowth.
• Branching patterns are regulated by the shoot tip a phenomenon called
apical dominance and plant hormones appear to be the factors responsible.
• Auxin is produced by young leaves and transported toward the base of the
leaf. It can suppress the outgrowth of axillary buds.
• Grazing and flowering often release buds from apical dominance, at which
time branching occurs.
• Cytokinins can also release buds from apical dominance.
• Primary functions
• 1. Support and orient the leaves in a manner that they are exposed to maximum sunlight and for
efficient gaseous exchange during photosynthesis and respiration.
• Conduct water and minerals Plants and animals from roots to leaves and manufactured food from
leaves to different parts of the plant.
• 3. Bear flowers and fruits
• Secondary Functions
• 1. Storage - Stems store food and water in some plants e.g. potato
• 2. Perennation - The underground stems help tide over the unfavourable growing periods e.g.
ginger.
• 3. Vegetative propagation - Stem can be a means of vegetative propagation e.g. rose, and
sugarcane.
• 4. Photosynthesis- in certain plants like xerophytes (desert plants) where leaves are reduced, the
stem takes up the function of photosynthesis. These stems possess chlorophyll e.g. Opuntia
• 5. Protection- In some plants the axillary bud modifies into thorn and protects the plants from
grazing animals e.g. citrus, Duranta.
• 6. Climbing - Tendrils or hooks are modified branches or buds. They coil around the support and
help the plant to climb e.g. grape vine
• Internal structure of stem:
• Epidermis - Outermost single layered, covered with cuticle, bears multicellular hairs,
protective function.
• Cortex - Inner to epidermis, there are three regions. z
• Hypodermis - 4-6 layers of collenchyma for mechanical support. z
• Middle layers - Few layers of parenchyma. z
• Endodermis - Innermost layer of cortex, has barrel shaped cells.
• As cells contain starch grains, it is also called starch sheath.
• Stele - All the tissues lying internal to endodermis constitute the stele.
• (i) Pericycle - Inner to endodermis, multilayered, parenchymatous with patches of sclerenchyma.
• (ii) Vascular bundles - Arranged in a ring ; each vascular bundle is (a)conjoint (xylem and phloem
together in one bundle), (b) collateral (xylem and phloem on the same radius with phloem towards
the periphery) and (c) open (cambium present in between xylem and phloem). Xylem is endarch
(protoxylem towards centre and metaxylem towards periphery).
• Medullary rays -
Narrow regions of
parenchymatous
cells in between
the vascular
bundles.
• Pith - The central
parenchymatous
zone with
intercellular
spaces.
• Leaf primordia (clusters of cells that will form leaves) are initiated at
the periphery of the shoot
• meristem (see Figure 20.21). The union of a leaf and the stem is called
a node, and stem tissue
• between nodes is called an internode (see Figure 20.20). In a
simplistic sense, the mature
• sporophyte is created by stacking node/internode units together.
Phyllotaxy, the positioning of
• leaves on the stem, involves communication among existing and
newly forming leaf primordia.
• LEAF
• Leaf is a flattened and expanded lateral appendage of stem or branch
developing from its node.
• It originates from leaf primordium formed by the shoot meristem and
bears a bud in its axil called axillary bud.
• It is the seat of very important physiological processes like
photosynthesis, transpiration and respiration.
• Besides protecting axillary buds ,leaf can get modified into structures
for storing food and water, climbing, and vegetative propagation.
• Leaves fall into two categories, simple and compound
• The Class I KNOX genes are homeobox genes that include STM and the KNOTTED 1 (KN1) gene in maize.
Gain-of function mutations of KN1 cause meristem-like bumps to form on maize leaves. In wild-type plants,
this gene is expressed in meristems. When KN1, or the tomato homologue LeT6, has its
• promoter replaced with a promoter from cauliflower mosaic virus and is inserted into the genome
• of tomato, the gene is expressed at high levels throughout the plant, and the leaves become "super
• compound" (Figure 20.25; Hareven et al. 1996; Janssen et al. 1998). Simple leaves become more
• lobed (but not compound) in response to overexpression of KN1.
• A second gene, LEAFY, that is essential for the transition from vegetative to reproductive
• development also appears to play a role in compound leaf development. It was identified in
• Arabidopsis and snapdragon (in which it is called FLORICAULA), and has homologues in other
• angiosperms
• Structure of Leaf
• A typical leaf has three parts
• (i) Leaf base - Lower most part of leaf by which it is attached to the stem node. It
may be expanded as sheath (in monocots) or bear lateral outgrowths (stipules) as in
dicots.
• (ii) Petiole - Is the stalk of leaf. Leaf can be petiolate ( with petiole) as in many dicots
or sessile ( without petiole) as in most monocots. Petiole may get modified and
swell (e.g. water hyacinth) or develop wings (e.g. orange) or become flat like a leaf
(e.g. Australian Acacia)
• (iii) Lamina or leaf blade- It is a green, thin, flattened and expanded part of leaf with
veins and veinlets traversing through its surface. The most prominent vein running
from base to apex and present in the middle of leaf blade is called mid rib.
• Following germination, and before they become competent to flower and reproduce, the
shoots of most plants pass through a phase of vegetative growth.
• During this period, plants generally rapidly increase their photosynthetic capacity and their
size and mass.
• This vegetative mode of growth can be further divided into
• a juvenile (The early phase of development during which the plants cannot be induced to flower and are
effectively insensitive to environmental influences of photoperiod and/or vernalization has been called
the juvenile phase)
• and an adult vegetative phase (they become sensitive to environmental influences).
• During the juvenile-to-adult phase transition, plants thus acquire reproductive competence.
• Simultaneously, changes in multiple traits, such as leaf size and shape, internode length and
trichome distribution, result in the appearance of both early (juvenile) and mature (adult)
shoots on the same plant, a condition known as heteroblasty.
• In Arabidopsis, which requires for the condition of vernalization and long days
facultatively, several regulatory pathways have been defined that is different in
their response to distinct internal and external cues, including vernalization,
photoperiod, gibberellin (GA), temperature, age and autonomous pathways.
• During the vegetative phase of the Arabidopsis life cycle, the shoot apical
meristem (SAM) produces leaves on its flanks and on transition to flowering,
the shoot bolts and the SAM becomes the inflorescence shoot apical meristem
(IM).
• A primary IM produces lateral meristems that may go on to produce flowers
or secondary inflorescences.
• Flower primordia are derived from the PZ of the IM and are initiated from a
block of four so-called founder cells
• The autonomous pathway functions promote the flowering
independently of day length by repressing the central flowering
repressor and vernalization regulatory gene FLOWERING LOCUS
C (FLC).
• Autonomous pathway genes repress FLC and thus promote the
floral transition. Several genes are known to be involved in this
pathway such as FLOWERING LOCUS CA (FCA), FLOWERING
LOCUS D (FLD), FLOWERING LOCUS KH
DOMAIN (FLK), FLOWERING LOCUS PA (FPA), FLOWERING LOCUS
VE (FVE), FLOWERING LOCUS Y (FY),
and LUMINIDEPENDENS (LD)
• Autonomous pathway genes regulate the levels of FLC mainly
through RNA-based post-transcriptional regulation mechanisms
and chromatin epigenetic modification.
• The photoperiodic pathway involves phytochromes and cryptochromes.
The interaction of these photoreceptors with a circadian clock initiates a
pathway that eventually results in the expression of the gene CONSTANS
(CO), which encodes a zinc-finger transcription factor that promotes
flowering. CO acts through other genes to increase the expression of the
floral meristem identity gene LEAFY (LFY).
• Flowering of Arabidopsis is promoted by exposure to long summer days
and is repressed by short winter days. The photoperiod pathway controls
this response and acts in the leaves through a signaling cascade involving
GIGANTEA (GI) and the transcriptional regulator CONSTANS (CO).
• CO promotes flowering by initiating transcription of the FT and TWIN
SISTER OF FT (TSF) genes. During long days, light promotes the
interaction between GI and a family of F-box ubiquitin ligases, such as the
FLAVIN-BINDING KELCH REPEAT F-BOX 1 (FKF1) protein.
• These interactions stabilize the F-box proteins, allowing them to promote the
degradation of a set of transcriptional repressors of CO, including numerous
DOF transcription factors called CYCLING DOF FACTORs (CDFs). CDFs repress
flowering by downregulating CO expression in leaves. At the
posttranscriptional level, CO is degraded in the dark by the ubiquitin ligase
CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1) and in the morning by a
pathway activated by the photoreceptor Phytochrome B (PHYB).
• These layers of transcriptional and posttranscriptional regulation ensure that
CO activates transcription of FT and TSF transcription only during long days.
Expression and translation of FT occur in the leaves, and the small FT protein
then moves through the phloem to the meristem, where it activates the floral
development program. Movement of TSF through the phloem may also occur.
• vernalization pathway, Vernalization also represses FLC, but
perhaps by a different mechanism (an epigenetic switch). Because the
FLC gene is a common target, the autonomous and vernalization
pathways are grouped together.
• Gibberellin Pathway
• Gibberellin (GA) is a growth regulator that promotes flowering in Arabidopsis. The
GIBBERELLIN 20 OXIDASE (GA20ox) enzyme catalyzes several steps in the biosynthesis of GA
by oxidizing a number of precursors.
• Mutations that reduce this biosynthetic pathway or increase the degradation of GA delay
flowering, particularly under short-day conditions. The concentration of bioactive GA (GA4 )
increases at the meristem immediately prior to floral induction.
• This GA4 probably moves from other parts of the plant to the meristem because mRNAs
encoding GA20ox and other GA-biosynthetic enzymes do not detectably accumulate at the
meristem.
• However, such mRNAs do increase in the stalk of the leaves (i.e., the petiole) in response to
day length, suggesting that the petioles are one site of GA biosynthesis during floral induction.
• The GA 2-oxidase (GA2ox) enzyme deactivates GA by hydroxylating it.
• Ambient Temperature Pathway
• Arabidopsis plants flower earlier when grown at higher temperatures, such as 23°C, than
at lower temperatures, such as 16°C. The MADS box transcription factor SHORT
VEGETATIVE PHASE (SVP) appears to play a crucial role in this pathway because svp
mutants are insensitive to changes in ambient temperature and flower early at both
temperatures. SVP represses FT transcription (and thus flowering) at lower
temperatures, but the levels of FT mRNA increase at higher temperatures.
• Age Pathway
• As the plant ages, concentrations of the SQUAMOSA PROMOTER BINDING LIKE (SPL)
transcription factors increase. SPLs promote flowering by initiating the expression of
several other transcription factors, such as LEAFY (LFY), FRUITFULL (FUL), and SOC1. SPL
proteins are negatively regulated by the microRNA miR-156, whose cellular levels are
higher in younger versus older plants and progressively decrease as the plant ages.
• During floral induction, the shoot apical meristem transforms from a vegetative
meristem, which forms leaves, to an inflorescence meristem, which forms flowers.
• During this process the meristem becomes taller and more domed. These
morphological changes are associated with dramatic changes in gene expression,
including increased expression of SOC1,
• Activation of SOC1 during long days requires FT and may be a direct response to the
arrival of FT at the meristem from the leaf.
• FT initiates flower development by interacting with the bZIP transcription factor FD and
directly promoting transcription of the MADS box factor APETALA1 (AP1).
• SOC1 activation leads to further changes in gene expression in the meristem as it
changes shape and becomes an inflorescence meristem. These include the expression
of transcription factors, such as LEAFY (LFY), SPLs, and AGAMOUS-LIKE 24 (AGL24).
• In the inflorescence meristem, TFL1 (TERMINAL FLOWER 1), a PEPB-like protein
(Phosphatidylethanolamine binding protein) prevents the upregulation of AP1 and LFY
mRNA.
• This restricts AP1 and LFY expression to cells committed to becoming floral primordia.
Stages of flower development
• P1. The first sign of flower primordium formation is the bulging of
the peripheral surface of the IM in a lateral direction. It is
hypothesized that a lateral protrusion formed during bulging is a
rudimentary bract.
• Soon after the bract is formed, another bulge occurs in its axil in an
upward direction. This second bulging corresponds to the
formation of a flower primordium proper.
• A significant increase in mitotic activity is observed upon
formation of the primordium.
• P2: At this stage, the flower primordium becomes clearly delimited
from the IM, and starts to grow larger very quickly in all directions.
• Sepal primordia become visible. The two lateral sepal primordia
appear first, but are soon outgrown by the abaxial (ab) then the
adaxial (ad) sepal primordia.
• Sepal primordia arise initially as ridges that lengthen and curve
inwards until they begin to overtop the remaining domeshaped
portion of the flower primordium
• P3: the petal and stamen primordia become visible. Primordia of the
four medial (long) stamens are first seen as wide outgrowths on the
flanks of the central dome of the FM.
• The four petal primordia that arise between the sepals close to their
base are just visible during this stage. The two lateral (short) stamens
develop from primordia appear later during this stage.
• P4: The sepals grow to completely cover the floral bud and the
primordia of the four long stamens bulge out and become distinct from
the central dome of cells. A rim around the central dome of the flower
primordium now begins to grow upward to produce an oval tube that
will become the gynoecium.
• P5:This stage begins when the growing primordia of the long stamens
become stalked at their base. The stalks give rise to the filaments, and the
wider upper region to the anthers. Anther locules are visible as convex
protrusions on the inner (adaxial) surface of the long stamens. Locules also
appear soon after in the short stamens. Petal growth now accelerates and
petal primordia become apparent.
• P6: The petal primordia elongate. There is a rapid lengthening of all organs
especially of petals that acquire a tongue-like shape. Nectary glands appear
and the stamens grow rapidly. Most of this growth occurs in the anther
region. At this stage the floral bud remains completely closed. The rapidly
growing petals reach the top of the lateral stamens. The cap of papillae that
will constitute the stigma starts to form at the top of the gynoecium.
• P7: Petals continue to lengthen relatively rapidly. The upper part of
the gynoecium differentiates into the style and a sharp boundary
separates it from the cap of stigmatic papillae. Petals become visible
between the sepals and continue to elongate rapidly. The stigma is
receptive at this stage.
• Stamen filaments extend even faster so the stamens outstrip the
gynoecium in length and self pollination takes place. The gynoecium is
now mature and its three distinct regions can be distinguished: an
apical stigma, a style, and a basal ovary.
• Arabidopsis is an annual plant.
• Leaves exhibit a ‘spiral phyllotaxy’, arising one after another from the
flanks of the meristem in a spiral pattern. After producing a rosette of
leaves, the plant switches from vegetative to reproductive growth so
that the apical meristem becomes an inflorescence meristem and
starts producing flower meristems.
• In Arabidopsis three genes, LEAFY (LFY), APETALA1 (AP1), and
CAULIFLOWER (CAL) function to specify flower meristem identity.
Mutations in each of these genes lead to a partial loss of flower
meristem identity.
• Once a group of cells produced on the flanks of the inflorescence meristem is
specified to become a flower by the action of the flower meristem identity
genes, the flower meristems initiate the production of floral organs.
• The Arabidopsis flower consists of four concentric whorls of organs (Figure
3a,b).
• Four sepals (Adaxial, abaxial and two lateral) occupy positions in the first whorl,
• with the second whorl consisting of four petals.
• Six stamens (four long and two short) containing pollen sacs occupy the third
whorl,
• and a two fused-carpelled gynoecium containing ovules forms the fourth whorl.
• The balance between cell proliferation and cell recruitment to
differentiated tissues in the SAM is dependent on mechanisms regulated
by WUSCHEL. The homeodomain-containing WUS transcription factor
has the role of the maintaining the identity of stem cells in the
organizing center of the CZ; wus mutants lack stem cells in the SAM.
• WUS expression is limited to the cells immediately below the stem cells,
an expression domain regulated by the receptor-kinase signaling system
that includes the CLAVATA1, 2 and 3 (CLV1, 2, 3) gene products.
• In clv mutants, there is an imbalance between cells retained within
meristems versus those recruited to form lateral organs.
• clv mutations cause an expansion of the WUS expression domain
resulting in an enlarged stem cell niche. CLV3 expression is, in turn,
positively regulated by WUS, suggesting that meristem size depends
greatly on a WUS-CLV regulatory loop.
• SHOOT MERISTEMLESS (STM) is a KNOTTED1-like homeobox (KNOX)
gene that encodes a protein expressed in the SAM’s CZ, RZ and
regions of the PZ. STM promotes the profileration of stem cell
derivatives until a critical cellular mass is attained sufficient to form
either leaves or floral primordia. It also inhibits the expression of
ASYMMETRIC LEAVES1 and 2 (AS1, 2) genes in the SAM, preventing
these cells from undergoing premature differentiation. Thus, the STM
gene is considered to play a pivotal role in meristem maintenance.
• ULTRAPETALA1 (ULT1) encodes a cysteine-rich protein with a B-box
like domain that restricts the size of shoot and floral meristems. It
functions antagonistically to the proliferative roles of WUS and STM.
• On induction of flowering, the IM genes, such as TERMINAL FLOWER 1 (TFL1) and
EMBRYONIC FLOWER 1 and 2 (EMF1, 2), are repressed in the FM, while the floral
meristem identity (FMI) genes, mainly LFY, APETALA1 (AP1), APETALA2 (AP2), and
CAULIFLOWER (CAL), are upregulated.
• Mutual repression of the IM and FMI genes seem to underlie the co-existence,
identity and boundaries of both types of meristem in the SAM in the transition to
flowering.
• For example, if genes such as TFL1 or EMF1 or 2 are mutated, LFY and/or AP1 are
ectopically expressed in the IM that is then transformed into a FM. Loss-of-
function mutants in these genes produce fl owers immediately after germination
skipping the vegetative phase
• On the contrary, if AP1, CAL and LFY are repressed, the FM attains IM identity
• LFY is necessary and suffi cient to specify FMI. In lfy mutants, leaves and
secondary shoots are produced instead of flowers and LFY overexpression
causes the conversion of leaves and axillary meristems to flowers.
• LFY and AP1 have overlapping functions in establishing the FM; while the
ap1 mutant has shoots with inflorescence characteristics, the lfy ap1
double mutant has an almost complete conversion of fl owers into shoots.
• CAL, the closest paralogue of AP1, and FRUITFULL (FUL) also act
redundantly to AP1 in FM specification. Single cal and ful mutants do not
show any FMI disorders, but in combination with ap1 in double or triple
mutants, the ap1 phenotype is greatly intensified
• The early patterning of the flower meristem are the so-called ABC homeotic genes,
AP1, AP2, AP3, PI, and AG, which are all transcription factors belonging to the MADS-
box gene family, except AP2.
• The classic ABC model was inferred using Arabidopsis and Antirrhinum homeotic
flower mutants (Coen and Meyerowitz, 1991). In these mutants two fl oral organ
types are replaced by two other fl oral organ types as follows:
• A- class mutant fl owers have carpels-stamens-stamens-carpels (from the outermost
to the innermost whorl),
• B-class mutant fl owers have sepals-sepalscarpels-carpels, and
• C-class mutant fl owers have sepals-petalspetals-sepals.
• It was shown that mutations in all three functions lead to the transformation of all fl
oral organs into leaf-like organs, suggesting that fl owers are modifi ed leaves
A (AP1 and AP2), B (AP3 and PI), and C
(AG) combinatorial transcriptional
regulatory functions and the SEP (1, 2,
3, 4) genes active in these mutants.
Organs are
listed below from the outer to the
inner whorl unless stated otherwise.
(A) Wild-type (WT) fl ower.
(B) Single ap2 mutant fl ower
composed of carpelloid sepals,
stamens, stamens and carpels.
(C) The pi mutant has fl owers
composed of sepals, sepals, carpels
and carpels.
(D) The ag fl ower has the stamens
transformed into petals and the
carpels are replaced by another fl
ower repeating the same pattern.
(E) The ap2 pi double mutant displays fl
owers composed only of sepalloid
carpels.
(F) The ap2 ag fl owers have leaf-like
organs in the fi rst and fourth whorls
and mosaic petal/stamen organs in
the second and third whorls.
(G) ap3 ag double mutants produce fl
owers composed of repeated whorls
of sepals.
(H) The ap2 pi ag mutant has leaf-like
organs with some residual carpel
properties. (Photographs provided by
Dr. J. Bowman).
• The genes responsible for floral whorl specification attain their spatio-
temporal pattern as a result of regulatory interactions among
themselves, interactions with meristem identity genes and with some
other genes.
• One of the key FM identity genes is LFY. The protein encoded by this
gene requires co-factors to set the spatial limits of expression of the
floral organ identity genes AP3, PI, and AG.
• For example, LFY participates with UFO in the regulation of AP1 and
• AP3 transcription, and with WUS co-regulates the expression of AG. LFY
also regulates the expression of the SEPALLATA (SEP) genes SEP1, SEP2
and SEP3
• The Vegetative-to-Reproductive Transition
• Unlike some animal systems in which the germ line is set aside during early embryogenesis, the
• germ line in plants is established only after the transition from vegetative to reproductive
• development that is, flowering. The vegetative and reproductive structures of the shoot are all
• derived from the shoot meristem formed during embryogenesis.
• There is a great diversity of
• flowering patterns among the over 300,000 angiosperm species, yet there appears to be an
• underlying evolutionary conservation of flowering genes and common patterns of flowering
• regulation.
• A simplistic explanation of the flowering process is that a signal from the leaves moves to the
• shoot apex and induces flowering. In some species, this flowering signal is a response to
• environmental conditions.