Embryology

The concept of preformation was strongly advocated by most seventeenth- and eighteenth-century naturalist-
philosophers.

Thus neither sperm contact nor the paternal genome is always essential for egg activation.

The cortical reaction creates an osmotic gradient, causing water to rush into this space, elevating the envelope and lifting
away all sperm bound to it, except the one sperm that has successfully fused with the egg membrane

The enlarged sperm nucleus, now called a pronucleus, migrates inward to contact the female pronucleus. Their fusion
forms the diploid zygote nucleus.

In animal eggs, fertilization induces an increase in the amount of free calcium ions inside the egg cytoplasm. This increase
in intracellular free calcium regulates later developmental events and is essential for normal development to occur in all
taxa studied, but the mechanisms controlling calcium levels vary.

During cleavage the embryo divides repeatedly to convert the large, unwieldy cytoplasmic mass into a large cluster of
small, maneuverable cells called blastomeres.

At the end of cleavage the zygote has been divided into many hundreds or thousands of cells and the blastula stage is
formed.

The amount of yolk at the vegetal pole varies among taxa. Eggs with very little yolk are called isolecithal. Mesolecithal
eggs have a moderate amount of yolk concentrated at the vegetal pole. Telolecithal eggs contain an abundance of yolk
densely concentrated at the vegetal pole of the egg. Centrolecithal eggs have a large, centrally located, mass of yolk.

When little yolk is present, cleavage furrows extend completely through the egg in holoblastic cleavage. When much yolk
is present, cleavage is meroblastic, with cells sitting atop a mass of undivided yolk.

Holoblastic cleavage occurs in isolecithal eggs and is present in echinoderms, tunicates, cephalochordates, nemerteans,
and most molluscs, as well as in marsupial and placental mammals, including humans.

Mesolecithal eggs also cleave holoblastically>>Amphibians.

Meroblastic cleavage occurs in telolecithal and centrolecithal eggs. In telolecithal eggs of birds, reptiles, most fishes, a few
amphibians, cephalopod molluscs, and monotreme mammals, cleavage is restricted to cytoplasm in a narrow disc on top of
the yolk.

The function of yolk is to nourish the embryo. When much yolk is present, as in telolecithal eggs, young exhibit direct
development, going from an embryo to a miniature adult. In this indirect development, larvae differ from adults and
must metamorphose into adult bodies.

Cleavage subdivides the mass of the zygote until a cluster of cells called a blastula is formed. In mammals, the cluster of
cells is called a blastocyst. In most animals, the cells are arranged around a central fluid-filled cavity called a blastocoel.

Gastrulation converts the spherical blastula into a more complex configuration and forms a second germ layer.

The blastopore becomes the mouth in organisms with one suite of developmental characters, but it becomes the anus in
organisms with another such suite of characters.

Animals with two germ layers are called diploblastic. Diploblastic animals include sea anemones and comb jellies. Most
animals have a third germ layer and are triploblastic

DEUTEROSTOME>Blastopore becomes anus, mouth forms secondarily///LOPHOTROCHOZOAN

PROTOSTOME>Blastopore becomes mouth, anus forms secondarily

A coelom is a body cavity completely surrounded by mesoderm. The coelomic cavity appears inside the mesoderm by one
of two methods: schizocoely or enterocoely.

There are two major groups of triploblastic animals, protostomes and deuterostomes. The groups are identified by a
suite of four developmental characters: (1) radial or spiral positioning of cells as they cleave, (2) regulative or mosaic
cleavage of cytoplasm, (3) fate of the blastopore to become mouth or anus, and (4) schizocoelous or enterocoelous
formation of a coelom. Snails and earthworms, among others, belong to the protostomes. Sea stars, fishes, and frogs,
among others, belong to the deuterostomes.

DEUTEROSTOME
Radial cleavage is so named because the embryonic cells are arranged in radial symmetry around the animal-vegetal axis.

Most deuterostomes have regulative development where the fate of a cell depends on its interactions with neighboring
cells, rather than on what piece of cytoplasm it acquired during cleavage.

The final deuterostome feature concerns the origin of the coelom. In enterocoely, both mesoderm and coelom are made at
the same time.

Gastrulation in mammals is remarkably similar to gastrulation in reptiles and birds. Gastrulation movements in the inner
cell mass produce a primitive streak

A further developmental complication in vertebrates is that coelom formation occurs by a modified form of schizocoely not
enterocoely. The nonvertebrate chordates form the coelom by enterocoely, as is typical of deuterostomes.

PROTOSTOME
Spiral cleavage occurs in most protostomes. It differs from radial cleavage in two important ways. Rather than dividing
parallel or perpendicular to the animal-vegetal axis, blastomeres cleave obliquely.

Mosaic development characterizes most protostomes. In mosaic development, cell fate is determined by the distribution
of certain proteins and messenger RNAs, called morphogenetic determinants, in the egg cytoplasm.

A protostome is so named because the blastopore becomes the mouth, and the second, unnamed, opening becomes the
anus.

Animals without a coelom are called acoelomate. In other protostomes, mesoderm lines only one side of the blastocoel,
leaving a fluid-filled blastocoel next to the gut. The fluid-filled cavity surrounding the gut is called a pseudocoelom. For
coelomate protostomes, like earthworms and snails, the mesodermal layer forms as just described, and a coelom is made
by schizocoely.

The protostomes are divided into two clades. One clade, lophotrochozoan protostomes, contains segmented worms,
molluscs (snails, slugs, clams, octopus and their kin) and several less familiar taxa. The name of this clade refers to two
features present in some members of the group: a horseshoe-shaped whorl of tentacles called a lophophore, and
a trochophore larva. The other clade, ecdysozoan protostomes, includes arthropods (insects, spiders, crabs, and
related organisms), roundworms, and other taxa which molt the exoskeleton.

Cephalopods, has bilateral cleavage like that of ascidian chordates.

Many ecdysozoans do not exhibit spiral cleavage; in some, cleavage appears radial, and in others, such as insects, cleavage
is neither spiral nor radial.

MECHANISMS OF DEVELOPMENT
In most animals (excluding insects), there are two major ways by which cells become committed to particular
developmental fates: (1) cytoplasmic partitioning of determinative molecules during cleavage and (2) interaction with
neighboring cells (inductive interactions).

Cytoplasmic Specification: A fertilized egg contains cytoplasmic components that are unequally distributed within the
egg. These different cytoplasmic components are thought to contain morphogenetic determinants that control
commitment of a cell to a particular cell type.

Induction, the capacity of some cells to evoke a specific developmental response in others, is a widespread phenomenon
in development.

Research suggests that epigenesis proceeds in three general stages: pattern formation, determination of position in the
body, and induction of limbs and organs appropriate for that position. Each stage is guided by gradients of gene products
that function as morphogens.

The first step in organizing development of an embryo is pattern formation: determination of the front-to-rear
(anteroposterior), left-to-right, and back-to-front (dorsoventral) axes.

In Drosophila the number and orientation of segments is controlled by segmentation genes. There are three classes of
segmentation genes: gap, pair-rule, and segment-polarity. Gap genes are activated first and divide an embryo into regions
such as head, thorax, and abdomen. Pair-rule genes divide these regions into segments. Finally, segment-polarity genes,
such as hedgehog, organize the anterior-to-posterior structures within each segment.

These proteins allow nearby cells to develop into the nervous system and other tissues along the middle of the back, and
those tissues in turn release other proteins that induce development of other parts of the body.

Because all animals appear to share similar molecular mechanisms for development, it may now be possible to understand
how changes in such developmental controls led to the evolution of the great variety of animals.

Segmentation genes apparently regulate expression of other genes, ensuring that they are active only in appropriate
segments. Mutations in homeotic genes, called homeotic mutations, place appendages or other structures in the wrong
part of the body.

The homeobox produces the part of a protein that attaches to the DNA of other genes, activating or blocking their
expression.

Hom genes do not encode specific limbs and organs. Instead, they function by specifying the location in the body along the
anteroposterior axis

Hox and other homeobox genes also play a role in shaping individual organs and limbs.

Reptiles, birds, and mammals form a monophyletic grouping of vertebrates called amniotes, so named because their
embryos develop within a membranous sac, the amnion.

The brain, spinal cord, and nearly all outer epithelial structures of the body develop from primitive ectoderm. They are
among the earliest organs to appear.
Much of the rest of the peripheral nervous system is derived from neural crest cells.

Watching nerves grow for periods of days, he saw that each axon was an outgrowth of a single cell. As the axon extended
outward, materials for growth flowed down the axon center to the growing tip where they were incorporated into new
protoplasm.

Despite structural and functional evolution, new forms are constrained by the architecture of their ancestors.

The metazoa, or multicellular animals, evolved greater structural complexity by combining cells into larger units. A
metazoan cell is a specialized part of the whole organism and, unlike a unicellular organisms, it is not capable of
independent existence.

Most metazoa have an additional level of complexity in which different organs operate together as organ systems. Eleven
different kinds of organ systems are observed in metazoans: skeletal, muscular, integumentary, digestive, respiratory,
circulatory, excretory, nervous, endocrine, immune, and reproductive

Spherical symmetry means that any plane passing through the center divides a body into equivalent, or mirrored, halves.
Radial symmetry is forms that can be divided into similar halves by more than two planes passing through the
longitudinal axis. The two phyla that are primarily radial as adults, Cnidaria and Ctenophora, have been called the Radiata.
Bilateral symmetry applies to animals that can be divided along a sagittal plane into two mirrored portions right and left
halves.

Animals that possess ectoderm, mesoderm, and endoderm are termed triploblastic and the majority are bilaterally
symmetrical.

In the acoelomate plan, mesodermal cells completely fill the blastocoel, leaving a gut as the only body cavity. In the
pseudocoelomate plan, mesodermal cells line the outer edge of the blastocoel, leaving two body cavities: a persistent
blastocoel and a gut cavity. In the schizocoelous plan, mesodermal cells fill the blastocoel, forming a solid band of tissue
around the gut. In deuterostomes, mesoderm forms by an enterocoelous plan, where cells from the central portion of the
gut lining begin to grow outward as pouches, expanding into the blastocoel.

Segmentation, also called metamerism, is another common feature of metazoans. Segmentation is a serial repetition of
similar body segments along the longitudinal axis of the body. Each segment is called a metamere, or somite.

A tissue is a group of similar cells (together with associated cell products) specialized for performance of a common
function. During embryonic development, the germ layers become differentiated into four kinds of tissues. These are
epithelial, connective, muscular, and nervous tissues.
An epithelium is a sheet of cells that covers an external or internal surface. Epithelial cells are also modified into glands
that produce lubricating mucus or specialized products such as hormones or enzymes. Epithelia are classified by cell form
and number of cell layers. Simple epithelia (a single layer of cells) are found in all metazoan animals, while stratified
epithelia (many cell layers) are mostly restricted to vertebrates.
Two kinds of connective tissue proper occur in vertebrates. Loose connective tissue is composed of fibers and both
fixed and wandering cells suspended in a viscous fluid ground substance. Dense connective tissue, such as tendons and
ligaments, is composed largely of densely packed fibers and little ground substance.
Much of the fibers of connective tissue is composed of collagen, a protein of great tensile strength. Connective tissue of
invertebrates, as in vertebrates, consists of cells, fibers, and ground substance, and show a wide diversity of structure
ranging from highly cellular to acellular histologies. Other types of specialized connective tissue include blood, lymph
(collectively considered vascular tissue), adipose (fat) tissue, cartilage, and bone.
Muscle is the most abundant tissue in the body of most animals. It originates (with few exceptions) from mesoderm, and
its unit is the cell or muscle fiber, specialized for contraction.
In vertebrates we recognize two types of striated muscle: skeletal and cardiac muscle.
Two basic types of cells in nervous tissue are neurons the basic functional unit of nervous systems, and neuroglia, a
variety of nonnervous cells that insulate neuron membranes and serve various supportive functions.