Early Development & Cleavage 🥚 Pseudocoelomates: Animals like roundworms with

a body cavity that is not fully lined by the

Sexual reproduction as a unifying feature of mesoderm.
animals: The emphasis on sexual reproduction,
even in simple animals like sponges and worms, Coelomates: The majority of animals (e.g.,
highlights its fundamental role in genetic diversity earthworms, humans) that have a true body cavity,
and evolutionary success. This process initiates a the coelom, which is completely surrounded by the
complex developmental sequence essential for mesoderm.
creating multicellular organization.
Animal reproduction begins when a sperm fertilizes
an egg, creating a single-celled zygote. This cell Organ Formation (Organogenesis) ✨
undergoes a process called cleavage, where it Once gastrulation is complete, the cells within each
repeatedly divides into smaller cells called germ layer continue to differentiate and form
blastomeres. This subdivision of mass continues specific organs in a process called organogenesis.
until the cells form a cluster known as a blastula.
Ectoderm cells form the nervous system, the
outer epithelium (including tooth enamel), and the
lens of the eye.
Germ Layer Formation (Gastrulation) 🔬
Mesoderm cells form the circulatory system,
The cells of the blastula then begin to differentiate
blood, bone marrow, and smooth muscle and
into specialized cell groups called germ layers
connective tissue of the digestive tract.
during a process known as gastrulation.
Endoderm cells form the respiratory tract, the
Diploblastic animals (e.g., Cnidarians) have two
pharynx, and the lining of the gut.
germ layers:
Ectoderm: Develops into the outer layer of the
body wall, the epidermis. The Circle of Life: The Stages of Animal
Development
Endoderm: Forms the tissues lining the gut cavity,
the gastrodermis. Throughout the animal kingdom, an incredible
variety of embryonic types exist, but most patterns
A non-cellular layer called the mesoglea separates
of embryogenesis are variations on five themes:
these two layers.
1. Immediately following
Triploblastic animals have a third layer, the
fertilization, cleavage occurs. Cleavage is a series
mesoderm, located between the ectoderm and
of extremely rapid mitotic divisions wherein the
endoderm. This layer gives rise to supportive,
enormous volume of zygote cytoplasm is divided
contractile, and blood cells.
into numerous smaller cells. These cells are
called blastomeres, and by the end of cleavage,
they generally form a sphere known as a blastula.
Body Cavities
2. After the rate of mitotic division has slowed down,
Triploblastic animals are further organized into the blastomeres undergo dramatic movements
subgroups based on the presence and type of their wherein they change their positions relative to one
body cavities, which are fluid-filled spaces that hold another. This series of extensive cell
internal organs. rearrangements is called gastrulation, and the
Acoelomates: Animals like flatworms that lack a embryo is said to be in the gastrula stage. As a
body cavity and have densely packed cells. result of gastrulation, the embryo contains
three germ layers: the ectoderm, the endoderm, the longest, and the adult is a brief stage solely for
and the mesoderm. reproduction. In the silkworm moths, for instance,
the adults do not have mouthparts and cannot feed.
3. Once the three germ layers are established, the
The larvae must eat enough for the adult to survive
cells interact with one another and rearrange
and mate. Indeed, most female moths mate as
themselves to produce tissues and organs. This
soon as they eclose from their pupa, and they fly
process is called organogenesis. Many organs
only once—to lay their eggs. Then they die.
contain cells from more than one germ layer, and it
is not unusual for the outside of an organ to be Plant Develpmental Stage
derived from one layer and the inside from another.
Plants and animals, though separated by 1.5 billion
For example, the outer layer of skin comes from the
years of evolution, evolved their multicellular
ectoderm, while the inner layer (the dermis) comes
organization from the same genetic toolkit. The key
from the mesoderm. Also during organogenesis,
differences in their developmental strategies stem
certain cells undergo long migrations from their
from two fundamental peculiarities of plants: their
place of origin to their final location. These
reliance on sunlight for energy, which dictates a
migrating cells include the precursors of blood cells,
specific body plan, and their rigid, cemented-
lymph cells, pigment cells, and gametes. Most of
together cells, which prevent cell movement.
the bones of our face are derived from cells that
have migrated ventrally from the dorsal region of
the head.
Environmental Adaptability
4. Many species a specialized portion of egg
cytoplasm gives rise to cells that are the precursors Unlike animal development, which is largely
of the gametes (the sperm and egg). The gametes buffered against environmental changes, plant
and their precursor cells are collectively development is dramatically influenced by its
called germ cells, and they are set aside for surroundings. Because plants cannot move to a
reproductive function. All the other cells of the body more favorable location, they adapt by altering their
are called somatic cells. This separation of growth. This is an opportunistic strategy where a
somatic cells (which give rise to the individual body) single type of organ—a leaf, a flower, or a root—
and germ cells (which contribute to the formation of can be produced by many different paths in
a new generation) is often one of the first response to environmental cues. For example, a
differentiations to occur during animal development. begonia leaf pegged to the ground can sprout a
The germ cells eventually migrate to the gonads, root, which in turn can grow a shoot that may
where they differentiate into gametes. The produce leaves and flowers given enough sunlight.
development of gametes, called gametogenesis,
is usually not completed until the organism has
become physically mature. At maturity, the gametes Modular and Variable Structure
may be released and participate in fertilization to
A mature plant is typically made of many copies of
begin a new embryo. The adult organism eventually
a small set of standardized modules, such as
undergoes senescence and dies.
leaves, flowers, and roots. The timing and position
5. In many species, the organism that hatches from of these modules are strongly influenced by the
the egg or is born into the world is not sexually environment, causing the overall structure of the
mature. Indeed, in most animals, the young plant to be highly variable. The choices between
organism is a larva that may look significantly these alternative modules and their organization
different from the adult. Larvae often constitute the into a whole plant depend on external cues and
stage of life that is used for feeding or dispersal. In long-range hormonal signals, which play a much
many species, the larval stage is the one that lasts smaller role in controlling animal development.
 The mature plant is typically composed of multiple
copies of small, standardized modules. The timing
Determinate vs. Indeterminate Growth and position of these modules are heavily
While the overall structure of a plant—its branching influenced by the environment, causing the overall
pattern or number of leaves and flowers—is plant structure to vary. The choices and
indeterminate and highly variable, its detailed organization of these modules depend on external
organization on a small scale is not. A single leaf, a signals and long-range hormones, which play a
flower, or an early plant embryo is as precisely much smaller role in animal development.
specified as any organ of an animal. These
individual modules have a determinate structure,
in contrast with the unpredictable pattern of the Determinate vs. Indeterminate Growth
plant as a whole. The internal organization of a
While the overall structure of a plant, like its
plant module raises the same genetic problems in
branching pattern or the number of roots, is
pattern formation as animal development, and they
indeterminate and highly variable, the detailed
are solved in analogous ways.
organization of a small-scale module, such as a
leaf or a flower, is not. A leaf is as precisely
specified as any animal organ, possessing a
Plants and animals are separated by about 1.5
determinate structure. The internal organization of
billion years of evolutionary history. They have
a plant module solves the same problems in
evolved their multicellular organization
genetic control of pattern formation as animal
independently but using the same initial tool kit—
development, using similar methods.
the set of genes inherited from their common
unicellular eucaryotic ancestor. Most of the
contrasts in their developmental strategies spring
from two basic peculiarities of plants. First, they get Arabidopsis: A Model Organism
their energy from sunlight, not by ingesting other Plant biologists use a small weed called
organisms. This dictates a body plan different from Arabidopsis thaliana as a primary model
that of animals. Second, their cells are encased in organism to identify and study genes governing
semirigid cell walls and cemented together, plant development. It shares several advantages
preventing them from moving as animal cells do. with other model organisms like Drosophila and C.
This dictates a different set of mechanisms for elegans: it's small, reproduces quickly, and is
shaping the body and different developmental convenient for genetic studies. A significant
processes to cope with a changeable environment. advantage for genetics is that Arabidopsis, like
Animal vs. Plant Development 🌿 vs. 🐅 many flowering plants, can reproduce as a
hermaphrodite, with a single flower producing both
Animal development is largely buffered against male and female gametes. This allows for self-
environmental changes, with the embryo's body fertilization, making it easy to identify and catalog
structure being genetically predetermined and genes required for specific developmental
unaffected by external conditions. In contrast, plant processes through genetic screens.
development is dramatically influenced by the
environment. Since plants cannot move, they adapt The Arabidopsis Genome 🧬
by altering their development, an opportunistic Arabidopsis thaliana has one of the smallest plant
strategy. For instance, a single organ like a leaf or genomes, containing about 125 million nucleotide
flower can be produced from a fertilized egg via pairs and approximately 26,000 genes, with many
many different paths depending on environmental being recent duplicates. This makes it a powerful
cues. model organism for genetics, similar to C. elegans
and Drosophila. Powerful tools, like vast libraries of attaching the embryo to nutritive tissue and
seeds with random mutations, allow scientists to providing a pathway for nutrient transport.
analyze gene functions easily.

Establishing the Body Plan 🌳

Genetic Differences Between Plants and
The embryo proper then proliferates into a ball of
Animals 🌿🆚🐅
cells that develops a polarized structure with two
The Arabidopsis genome is exceptionally rich in key groups of cells. One group, at the suspensor
genes that code for gene regulatory proteins, with end, will form the root, while the other, at the
some families like the Myb family being greatly opposite end, will generate the shoot. This process
expanded compared to animals. However, other establishes the plant's main root-shoot axis, which
families, such as nuclear hormone receptors, is analogous to an animal's head-to-tail axis.
appear to be entirely absent in plants. Conversely, Simultaneously, the three major tissue layers—the
plants have large families of gene regulatory outer epidermis, inner ground tissue, and central
proteins with no known animal counterparts. While vascular tissue—begin to form. These layers are
some homologous proteins exist between plants comparable to the three germ layers of an animal
and animals, they regulate different genes and embryo.
developmental processes, and their protein
After this stage, the embryo forms rudimentary
sequences are not highly conserved outside of the
seed leaves (cotyledons) and then development
DNA-binding domains.
typically halts. The embryo becomes packaged
within a seed, where it enters a state of dormancy,
stabilized by dehydration. This allows the embryo to
Cell Communication and Signaling 🚦 survive harsh conditions and remain viable for long
Arabidopsis has many genes for cell periods. Embryonic development resumes when
communication and signal transduction, but the the seed is rehydrated and germinates.
specific details of these pathways are very different Genetic Control of Embryo Development 🧬
from those in animals. Key animal signaling
mechanisms like Wnt, Hedgehog, Notch, and Using genetic screens in model organisms like
TGFβ are all absent in Arabidopsis. In their place, Arabidopsis, scientists have identified genes that
plants have highly developed, unique signaling govern the organization of the plant embryo. These
pathways. While serine/threonine kinase genes can be categorized based on the mutant
receptors are very plentiful in plants, tyrosine phenotypes they produce, with some being
kinase receptors seem to be entirely absent. essential for the formation of the root, stem, or
Additionally, a significant number of genes in the shoot apex. Other gene classes are required for the
Arabidopsis genome are dedicated to plant-specific development of specific tissue types (epidermis,
developmental processes, such as cell wall ground, and vascular) or for the organized changes
synthesis and light detection. in cell shape that give the embryo its form.

Embryonic development in flowering plants begins Meristems: The Plant's Building Blocks 🧱
with the fertilized egg, or zygote, undergoing an
Unlike animals that grow from a pre-formed
asymmetrical division to establish the future
embryonic plan, plants are built sequentially by
embryo's polarity. This division produces a small,
groups of proliferating cells called apical
dense cell that becomes the embryo proper and a
meristems. Each meristem is a self-renewing
large, vacuolated cell that forms the suspensor. The
population of stem cells that produces progeny
suspensor acts like a mammalian umbilical cord,
cells. These progeny cells are displaced, enlarge,
and then differentiate to form the plant's structures. changed by plant growth regulators, allowing for
The rudiments of the root and shoot apical flexible shaping in response to developmental cues.
meristems are already established in the embryo.
Upon germination, the meristems become highly
active, with non-meristematic cells rapidly enlarging A plant's continuous growth and development are
to push out a root and then a shoot, allowing the driven by apical meristems, which are self-
plant to take hold and begin photosynthesis. perpetuating populations of stem cells. These
Perennial plants also retain meristem potential in meristems produce structures with limited growth,
other areas, allowing them to increase in girth and such as leaves and flowers, which have a
sprout new shoots if damaged. determinate size and shape.

Environmental Control of Development 🌿 Forming Plant Modules from Primordia 🌿

From germination onward, a plant's development is As a plant's shoot elongates, the apical meristem
heavily influenced by environmental signals. The lays down a repeating sequence of modules, each
timing of a seedling's transition from subterranean composed of a stem segment (internode), a point
sprouting to growth above ground is a prime of attachment (node), a leaf, and a bud. The
example. This switch is not genetically programmed continuous activity of the meristem creates an ever-
because the depth of the seed is unpredictable. increasing number of these modules, which are
Instead, it's controlled by light, which inhibits the precisely patterned relative to one another.
production of hormones called brassinosteroids.
Mutations affecting brassinosteroid signaling can The organization of a module begins on a
cause seedlings to green, slow their growth, and microscopic scale within the meristem. At the
open their seed leaves prematurely while still in the shoot's apex, the meristem appears as a small
dark. central dome surrounded by swellings called
primordia. Each swelling is the rudimentary form of
a leaf. Through a defined program of cell division
and enlargement, each leaf primordium develops
Shaping by Oriented Cell Division and into a complete leaf, node, and internode.
Expansion 📏 Simultaneously, the apical meristem generates new
Since plant cells are locked in place by their rigid primordia, ensuring an unending succession of
cell walls, a plant's shape is determined by the modules.
orderly division and oriented expansion of its The specific pattern of these primordia, including
cells. In a root tip, for example, cells pass through their position and spacing, is controlled by a system
three distinct, overlapping zones: a zone of division, of local signals within the tiny shoot apex. By
a zone of oriented elongation, and a zone of activating different sets of genes, the plant can
differentiation. This process accounts for the root's produce various types of primordia in different
architecture. spatial patterns, leading to more complex structures
The expansion of plant cells, which can be a 50- like tendrils, thorns, branches, and flowers.
fold increase in volume, is driven by internal turgor Cell Signaling Maintains the Meristem
pressure pushing on the cell wall. The direction of
this expansion is guided by the orientation of Central to all these phenomena is the question of
cellulose fibrils within the cell wall, which in turn is how the apical meristem maintains itself. The
controlled by microtubules just inside the cell meristem cells must continue to proliferate for
membrane. These orientations can be quickly weeks, years, or even centuries as a plant grows,
replacing themselves while continually generating
progeny cells that differentiate. Through all this, the This account of the plant meristem is still uncertain
size of the cluster of cells that constitute the in many details and other genes besides those
meristem remains practically constant (about mentioned are also involved. Nevertheless,
100 cells in Arabidopsis, for example). New mathematical modeling shows that systems of this
meristems may arise as the plant branches, but sort, based on a feedback loop involving a short-
they too preserve the same size. range activating signal and a long-range
inhibitory signal, can stably maintain a
Genetic screens have identified genes required for
signaling center of a well-defined size even
meristem maintenance. For example, mutations
when there is continual proliferation and turnover of
that disrupt the WUSCHEL gene, which codes for a
the cells that form that center. Analogous systems
homeodomain protein, convert the apical
of signals are thought to operate in animal
meristem into non-meristematic tissue, so that
development to maintain localized signaling centers
the seedling fails to sprout. Conversely, mutations
—such as the Organizer of the amphibian gastrula,
in the CLAVATA group of genes, coding for
or the zone of polarizing activity in a limb bud.
components of a cell-cell signaling pathway,
make the meristem abnormally big. These genes It is still not known how the Wuschel-expressing
are expressed in different layers of cells in the cells signal to their neighbors. One possibility is that
meristem region. The two most superficial cell the Wuschel protein itself diffuses directly from
layers, called the L1 and L2 layers, together with cell to cell through plasmodesmata—a signaling
the uppermost part of the L3 layer, contain the cells pathway peculiar to plants. Some other gene
of the meristem proper, capable of dividing regulatory proteins have in fact been shown to
indefinitely to give rise to future parts of the plant. travel in this way in meristems, spreading from cells
The meristematic cells of the L1 and L2 layers that contain the corresponding mRNA into
express Clavata3, a small secreted signal neighboring cells that do not.
protein. Just beneath, in the L3 layer, lies a cluster
Regulatory Mutations Can Transform Plant
of cells expressing Clavata1 (the receptor for
Topology by Altering Cell Behavior in the
Clavata3). In the center of this Clavata1 patch are
Meristem
cells that express the Wuschel gene regulatory
protein. If a stem is to branch, new shoot apical
meristems must be created, and this too depends
The pattern of cell divisions implies that the cells
on events in the neighborhood of the shoot apex. At
expressing Wuschel are not themselves part of
each developing node, in the acute angle (the axil)
the meristem proper; new Wuschel-expressing
between the leaf primordium and the stem, a bud
cells are apparently continually recruited from the
is formed. This contains a nest of cells, derived
meristematic part of the L3 population, just above
from the apical meristem, that keep a meristematic
the Wuschel domain. Nevertheless, the Wuschel-
character. They have the capacity to become the
expressing cells are at the heart of the
apical meristem of a new branch or the
mechanism that maintains the meristem. A
primordium of a structure such as a flower; but they
signal that they produce maintains meristematic
also have the alternative option of remaining
behavior in the cells above, stimulates expression
quiescent as axillary buds. The plant's pattern of
of the CLAVATA genes, and presumably causes
branching is regulated through this choice of
new cells recruited into the Wuschel domain to
fate, and mutations that affect it can transform
switch on Wuschel. Negative feedback from the
the structure of the plant. Maize provides a
upper meristematic cells, delivered by the
beautiful example.
Clavata signaling pathway, acts back on the
regions below to limit the size of the Wuschel Maize represents one of mankind's most
domain, thereby preventing the meristem from remarkable feats of genetic engineering. Native
becoming too big. Americans created it by selective breeding, over a
period of several centuries or perhaps millennia evolve through changes in regulatory DNA
between 5,000 and 10,000 years ago. They started without change in the characters of the proteins
from a wild grass known as teosinte, with highly made.
branched leafy stems and tiny ears bearing
Long-Range Hormonal Signals Coordinate
inedible hard kernels. Detailed genetic analysis has
Developmental Events in Separate Parts of the
identified a handful of genetic loci—about five—that
Plant
account for most of the difference between this
unpromising ancestor and modern corn. One of The fate of an axillary bud is dictated not only by
these loci, with a particularly dramatic effect, its genes but also by environmental conditions.
corresponds to a gene called teosinte branched-1 Separate parts of a plant experience different
(tb1). In maize with loss-of-function mutations in environments and react to them individually by
tb1, the usual simple unbranched stem, with a changes in their mode of development. The plant,
few large leaves at intervals along it, is however, must continue to function as a whole.
transformed into a dense, branching leafy mass This demands that developmental choices and
reminiscent of teosinte. The pattern of branching events in one part of the plant affect developmental
in the mutant implies that axillary buds, choices elsewhere. There must be long-range
originating in normal positions, have escaped signals to bring about such coordination.
from an inhibition that prevents them, in normal
maize, from growing into branches. As gardeners know, for example, by pinching off the
tip of a branch, one can stimulate side growth:
In normal maize, the single stem is crowned with a removal of the apical meristem relieves the
tassel—a male flower—while a few of the axillary quiescent axillary meristems of an inhibition
buds along the stem develop into female flowers and allows them to form new twigs. In this case,
and, upon fertilization, form the ears of corn that we the long-range signal from the apical meristem, or
eat. In the mutant maize with a defective tb1 gene, at least a key component, has been identified. It is
these fruitful axillary buds are transformed into an auxin, a member of one of six known classes
branches bearing tassels. The wild teosinte plant is of plant growth regulators (sometimes called
like the tb1-defective maize in its leafy, highly plant hormones), all of which have powerful
branched appearance, but unlike this mutant it influences on plant development. The five other
makes ears on many of its side branches, as known classes are the gibberellins, the
though tb1 were active. DNA analysis reveals the cytokinins, abscisic acid, the gas ethylene, and
explanation. Both teosinte and normal maize the brassinosteroids. All are small molecules that
possess a functional tb1 gene, with an almost readily penetrate cell walls. They are all
identical coding sequence, but in maize the synthesized by most plant cells and can either act
regulatory region has undergone a mutation locally or be transported to influence target
that boosts the level of gene expression. Thus in cells at a distance. Auxin, for example, is
normal maize the gene is expressed at a high transported from cell to cell at a rate of about 1 cm
level in every axillary bud, inhibiting branch per hour from the tip of a shoot toward its base.
formation, while in teosinte the expression in many
axillary buds is low, so that branches are permitted Each growth regulator has multiple effects,
to form. modulated by the other growth regulators, as well
as by environmental cues and nutritional status. For
This example shows how simple mutations, by example, auxin alone can promote root
switching the behavior of meristem cells, can formation, but in conjunction with gibberellin it can
transform plant structure—a principle of promote stem elongation; with cytokinin, auxin can
enormous importance in the breeding of plants for suppress lateral shoot outgrowth; and with ethylene
food. More generally, the case of tb1 illustrates how it can stimulate lateral root growth.
new body plans, whether of plant or animal, can
Homeotic Selector Genes Specify the Parts of a transformed: the petals are converted into
Flower sepals and the stamens into carpels.
Meristems face other developmental choices  The third or ‘C’ class, exemplified by
besides that between quiescence and growth, as agamous, has its two innermost whorls
we have already seen in maize, and these also are transformed, with a more drastic
frequently regulated by the environment. The most consequence: the stamens are converted
important is the decision to form a flower. into petals, the carpels are missing, and in
their place the central cells of the flower
The switch from meristematic growth to flower
behave as a floral meristem, which begins
formation is typically triggered by light. By
the developmental performance all over
poorly understood mechanisms based on light
again, generating another abnormal set of
absorption by phytochrome and cryptochrome
sepals and petals nested inside the first
proteins, the plant can sense very precisely a
and, potentially, another nested inside that,
change in day length. It responds by turning on
and so on, indefinitely.
expression of a set of floral meristem-identity
genes in the apical meristem. By switching on  A fourth class, the sepallata mutants, has
these genes, the apical meristem abandons its its three inner whorls all transformed into
chances of continuing vegetative growth and sepals.
gambles its future on the production of gametes. Its
These phenotypes identify four classes of
cells embark on a strictly finite program of
homeotic selector genes, which, like the homeotic
growth and differentiation: by a modification of
selector genes of Drosophila, all code for gene
the ordinary mechanisms for generating leaves, a
regulatory proteins. These are expressed in
series of whorls of specialized appendages are
different domains and define the differences of cell
formed in a precise order—typically sepals first,
state that give the different parts of a normal flower
then petals, then stamens carrying anthers
their different characters. The gene products
containing pollen, and lastly carpels containing
collaborate to form protein complexes that drive
eggs. By the end of this process the meristem has
expression of the appropriate downstream
disappeared, but among its progeny it has created
genes. In a triple mutant where the A, B and C
germ cells.
genetic functions are all absent, one obtains in
The series of modified leaves forming a flower can place of a flower an indefinite succession of tightly
be compared to the series of body segments nested leaves. Conversely, in a transgenic plant
forming a fly. In plants, as in flies, one can find where genes of the A, B and sepallata classes are
homeotic mutations that convert one part of the all expressed together outside their normal
pattern to the character of another. The mutant domains, leaves are transformed into petals.
phenotypes can be grouped into at least four Leaves therefore represent a “ground state” in
classes, in which different but overlapping sets of which none of these homeotic selector genes
organs are altered: are expressed, while the other types of organ
result from expressing the genes in different
 The first or ‘A’ class, exemplified by the
combinations.
apetala2 mutant of Arabidopsis, has its
two outermost whorls transformed: the Similar studies have been carried out in other plant
sepals are converted into carpels and the species, and a similar set of phenotypes and genes
petals into stamens. have been identified: plants, no less than
animals, have conserved their homeotic
 The second or ‘B’ class, exemplified by
selector gene systems. Gene duplication has
apetala3, has its two middle whorls
played a large part in the evolution of these
genes: several of them, required in different organs
of the flower, have clearly homologous sequences. internal organization of each plant module is
These are not of the homeobox class but are sketched out on a microscopic scale through cell-
members of another family of gene regulatory cell interactions in the neighborhood of the apical
proteins (the so-called MADS family), also found in meristem. The meristem itself appears to be
yeast and in vertebrates. maintained by a local feedback loop, in which
cells expressing the gene regulatory protein
Clearly, plants and animals have independently
Wuschel provide a positive stimulus, and a
found very similar solutions to many of the
negative feedback dependent on the Clavata
fundamental problems of multicellular
cell-cell signaling pathway keeps the meristem
development.
from becoming too big.
Summary
Environmental cues—especially light that is
The development of a flowering plant, like that of appropriately timed—can cause the expression
an animal, begins with division of a fertilized egg to of genes that switch the apical meristem from a
form an embryo with a polarized organization: the leaf-forming to a flower-forming mode. The parts
apical part of the embryo will form the shoot, of a flower—its sepals, petals, stamens, and
the basal part, the root, and the middle part, the carpels—are formed by a modification of the
stem. At first, cell division occurs throughout the mechanism for development of leaves, and the
body of the embryo. As the embryo grows, differences between these parts are controlled by
however, addition of new cells becomes restricted homeotic selector genes that are closely
to small regions known as meristems. Apical analogous (although not homologous) to those
meristems, at shoot tips and root tips, will of animals.
persist throughout the life of the plant, enabling
it to grow by sequentially adding new body parts at
its periphery. Typically, the shoot generates a
repetitive series of modules, each consisting of a
segment of stem, a leaf, and an axillary bud. An
axillary bud is a potential new meristem,
capable of giving rise to a side branch; the
environment—and long-range hormonal signals
within the plant—can control the development of
the plant by regulating bud activation.
Mutations that alter the rules for activating
axillary buds can have a drastic effect on the
shape and structure of the plant; a single such
mutation—one of about five key genetic alterations
—accounts for a large part of the dramatic
difference between modern maize and its wild
ancestor, teosinte.
The small weed Arabidopsis thaliana is widely
used as a model organism for genetic studies
and is the first plant to have had its genome
completely sequenced. As in animals, genes
governing plant development can be identified
through genetic screens and their functions tested
by genetic manipulations. Such studies have begun
to reveal the molecular mechanisms by which the