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Primary Organizer
Hans Spemanns Research only tissue to induce growth Research in 19201930s centered on: how one tissue influenced the fate of another: induction 1916 - Spemann noted the capacity of tissue from the dorsal lip of the gastrula-transformed tissue 1921-22 - Repeated delicate experiments showed it induced a 2 embryo All of the knowledge for constructing a complex egg must be in the nucleus and cytoplasm
Development
series of progressive changes 1. when a fertilized egg divides mitotically 2. Specialization - occurs as a hierarchy of devtal decisions 3. Cell types do not unfold but arise from conditions created in preceding stages 4. Interactions become increasingly restrictive; each stage limits developmental fate 5. With each new stage, cells lose the option to become something differentit becomes determined 6. Both cytoplasmic localization and induction cause this feature
2. Fertilization
A. Initial Event Fertilization - union of male and female gametes - provides for recombination of paternal and maternal genes, restoring the diploid number - activates the egg to begin development B. Oocyte Maturation - contrast with sperm Sperm - eliminates nearly all cytoplasm and condenses its nucleus Egg - grows in size by accumulating yolk - contains much mRNA, ribosomes, tRNA & elements for protein synthesis Morphogenetic determinants direct the activation and repression of specific genes in the egg The egg nucleus grows in size, bloated with RNA and is called the germinal vesicle. Preparations in the egg occur during the prolonged prophase After meiosis resumes, the egg is ready to fuse its nucleus with the sperm nucleus
Prevention of Polyspermy
Fertilization cone - forms where the sperm contacts the vitelline membrane Important changes in the egg surface block entrance to any additional sperm Polyspermy - entry of more than one sperm - would cause a triploid nucleus In the sea urchin, an electrical potential rapidly spreads across the membrane; this is the fast block - followed by the cortical reaction
Cortical Reaction
enzyme-rich cortical granules below the egg membrane fuse with the membrane granules release contents in space between the membrane and vitelline envelope Then, the reaction Water rushes into space elevates the envelope Lifts away all sperm bound to it. Except for one One cortical granule enzyme causes the vitelline envelope to harden, becoming the fertilization membrane
Sea urchins (12 mins); mammals (12 h) The Egg is now a zygote DNA and protein synthesis undergoes a burst of activity, using a supply of mRNA in the egg cytoplasm
Fertilization initiates reorganization of cytoplasm and repositions determinants that begin development and cleavage
Patterns of Cleavage
Cleavage - orderly sequence of cell divisions The pattern of cleavage is affected by:
a. quantity and distribution of yolk present, and b. genes controlling the symmetry of cleavage
Vegetal pole - yolk-rich end of the embryo while the other end is the animal pole During each division, a cleavage furrow is visible in the cell
How Amount and Distribution of Yolk Affects Cleavage Isolecithal yolk - eggs with very little yolk and the yolk is distributed evenly
a. In such eggs, cleavage is holoblastic b. The cleavage furrow extends completely through the egg
Isolecithal eggs - widespread and seen in: echinoderms, tunicates, cephalochordates, molluscs and mammals Note: Cleavage is slowed in the yolk-rich vegetal pole
Blastula stage - consists of a few hundred to several thousand cells Blastula formation, with a single germ layer, occurs in all multicellular animals In all animals other than sponges, development continues to form one or two more germ layers Germ layers formed during development ultimately produce all structures of the adult body
Some animals retain a blind, or incomplete gut, but most have a complete gut with a second opening, the anus.
Upon completion of coelom formation, the body has three tissue layers and two cavities
Deuterostome Development
Cleavage Patterns Radial cleavage - embryonic cells are arranged in radial symmetry around the animal-vegetal axis Regulative development - fate of cell depends on interactions with neighboring cells
Early in the development of these embryos, each cell is capable of producing an entire embryo if separated from the other cells
Fate of Blastopore
Deuterostome embryos develop a complete gut blastopore anus 2nd opening mouth Coelom Formation
Enterocoely - mesoderm & coelom form at the same time - gastrulation begins with one side of the blastula bending inward to form the archenteron - pouch-like compartment pinches off to form a mesodermally bound space surrounding the gut
Radial cleavage is characteristic of the Deuterostomia, echinoderms, hemichordates & chordates bilateral cleavage - egg is defined by unequal cytoplasmic components(Fig 8.12) - Ascidians (tunicates) demonstrate this cleavage pattern. discoidal cleavage (telolecithal eggs) - there is a large mass of yolk in each egg; cleavage is confined to a small disc of cytoplasm - Early cleavage furrows carve the disc into a single layer of cells (blastoderm)
Rotational cleavage 1st cleavage plane - aligned with the animal vegetal axis 2nd cleavage 1 blastomere divides meridionally (through the animal-vegetal axis)while the other divides equatorially (perpendicular), rotated 90 degrees to the first Early divisions may be asynchronous and possess odd numbers of cells below the 2-4-8-16 series that would occur with synchronous division 3rd - cells form a tightly packed cluster stabilized by outer cells with tight junctions, the trophoblast trophoblast - form the embryonic portion of the placenta type of cleavage is present in mammals and is slower than in other animal groups
Frogs - radial cleavage, movements during gastrulation - influenced by the large mass of yolk Gastrulation in amphibians begins with invagination
Birds & reptiles Primitive streak forms the anterioposterior axis and center of early growth blastoderm has 2 layers: epiblast and hypoblast with a blastocoel between them Epiblast sheet moves toward primitive streak and over the edge, migrating One stream of cells moves deeper, displacing midline hypoblast, and forms endoderm Surface cells form ectoderm
mammalian gastrulation Epiblast move medially through the primitive streak into the blastocoel cells migrate laterally through the blastocoel to form mesoderm and endoderm mammals utilize nutrients from a placenta
Reptiles, birds & mammals share a common ancestor whose eggs were telolecithal; all inherited this gastrulation pattern and mammals then evolved isolecithal eggs with a telolecithal pattern
Protostome Development
Cleavage Patterns Spiral cleavage - protostomes Blastomere cleave obliquely to the animal-vegetal axis. Upper layer of cells appears offset relative to lower layer Mosaic devt is characteristic of most protostomes. (Fig 8.10) - cell fate is determined by the distribution of morphogenetic determinants, in the egg cytoplasm
Protostomes are so named because the blastopore becomes the mouth. Coelom Formation a band of mesoderm forms around the gut before a coelom forms If a coelom exists, it is formed by schizocoely (splitting) Endodermal cells move by ingression (entering) into the space between the archenteron walls and the ectoderm These cells divide and produce mesodermal precursors; the proliferating cells become the mesoderm. Some protostomes, like flatworms, are acoelomate In others, mesoderm lines only one side of the blastocoel, leaving a fluid-filled cavity around the gut called a pseudocoelom.
Fate of Blastopore
2. Edcysozoan protostomes
- includes arthropods, roundworms.. - name of the clade refers to ecdysis (taking it off or stripping off)
Mechanisms of Development
Nuclear Equivalence 1. Roux-Weismann Hypothesis a. An earlybut wrongexplanation of differentiation was based on division of nuclear material along the cell lineage b. Early embryologists saw this as an explanation of differentiation; genome gradually became broken down into smaller units c. Hans Driesch separated the two-celled sea urchin stage; both developed into normal larvae but this did not disprove eventual progressive modification
Spemanns experiment demonstrated that every blastomere contains sufficient genetic information for the development of a complete animal
Cytoplasmic Specification
Cytoplasmic components are unevenly distributed in a zygote Different components contain morphogenetic determinants that control commitment to cell type Different determinants are partitioned among different blastomeres by cleavage and determine cell fate.
Some tunicate species have different colored types of cytoplasm a. Yellow cytoplasm gives rise to muscle cells. b. Gray equatorial cytoplasm produces the notochord and neural tube. c. Clear cytoplasm produces larval epidermis. d. Gray vegetal cytoplasm gives rise to the gut.
Embryonic Induction
Induction - capacity of some cells to evoke a specific devtal response in other cells a graft of the dorsal blastopore lip could induce ectoderm to form a neural tube Dorsal lip was the primary organizer because it was the only tissue to induce growth Spemann called this primary induction the first inductive event
Cells that have differentiated act as inductors for adjacent undifferentiated cells Timing is critical; primary induction sets in motion secondary induction. The sequence includes cell movement, changes in adhesion, and cell proliferation
Gene Expression During Development After fertilization, proteins are translated from stored mRNA transcribed from maternal genome In many animals, maternal mRNA directs protein synthesis through cleavage and to mid-blastula stage After this, protein synthesis switches from maternal to zygotic control as the nucleus transcribes its own mRNA
Mice and humans - 4 clusters of homeobox-containing genes on separate chromosomes - all are homologues of the fruit flys homeotic genes Hox & homeobox genes - play a role in shaping individual organs and limbs (Fig 8.18, 8.19) New limb buds can be induced to grow from the side of a chick by implanting a bead soaked in fibroblast growth factor (FGF) helps establish a proximodistal axis one of the three axes that guide limb devt (Figure 8.20)
Vertebrate Development
Common Vertebrate Heritage
One outcome of shared ancestry in vertebrates is the similarity of postgastrula embryos - all vertebrate embryos share: dorsal neural tube, notochord, pharyngeal gill pouches with aortic arches, ventral heart and postanal tail. - similarity is extraordinary considering the variety of eggs and developmental patterns
Monotremes
Absence of teats (modified sweat glands sucked by youngs) Lay eggs With cloaca Testes within abdomen (absence of scrotal sac) No pinna of ear Platypus (ductbill), Echidna (spiny anteater)
Young nursed in marsupium (abdominal skin pouch) Yolk sac serve as placenta Teats opening into marsupium Geographically isolated in Australia Ex. Kangaroo, koala, wallaby, opossum, Tasmanian wolf
Marsupialia
Placental mammals