The question of how a zygote becomes an animal has been asked for centuries. As recently as the 18th century, the prevailing theory was a notion called preformation the idea that the egg or sperm contains an embryo.
A preformed miniature infant, or homunculus, that simply becomes larger during development.
Kaspar Friederich Wolff (1759) demonstrated there was no preformed chick in the early egg.
Undifferentiated granular material became arranged into layers. The layers thickened, thinned, and folded to produce the embryo.
Development describes the changes in an organism from its earliest beginnings through maturity.
Cell types arise from conditions created in preceding stages. Interactions become increasingly restrictive. Each stage limits developmental fate. Cells lose option to become something different
Said to be determined.
Fertilization
Fertilization
Oocyte Maturation
Contains much mRNA, ribosomes, tRNA and elements for protein synthesis.
Morphogenetic determinants direct the activation and repression of specific genes later in post-fertilization development. Egg nucleus grows in size, bloated with RNA.
Fertilization
Most of these preparations in the egg occur during the prolonged prophase I.
In mammals
After fertilization it will support nutritional requirements of the embryo and direct its development through cleavage.
After meiosis resumes, the egg is ready to fuse its nucleus with the sperm nucleus.
Broadcast spawners often release a chemotactic factor that attracts sperm to eggs.
Species specific
Sperm enter the jelly layer. Egg-recognition proteins on the acrosomal process bind to speciesspecific sperm receptors on the vitelline envelope.
Fast block
Slow block
The cortical reaction follows the fusion of thousands of enzyme-rich cortical granules with the egg membrane.
Cortical granules release contents between the membrane and vitelline envelope. Creates an osmotic gradient
Water rushes into space Elevates the envelope Lifts away all bound sperm except the one sperm that has successfully fused with the egg plasma membrane.
The increased Ca2+ concentration in the egg after the cortical reaction results in an increase in the rates of cellular respiration and protein synthesis.
Fusion of Pronuclei
After sperm and egg membranes fuse, the sperm loses its flagellum. Enlarged sperm nucleus is the male pronucleus and migrates inward to contact the female pronucleus. Fusion of male and female pronuclei forms a diploid zygote nucleus.
Cleavage
Very little growth occurs. Each cell called a blastomere. Morula solid ball of cells. First 5-7 divisions.
Polarity
The eggs and zygotes of many animals (not mammals) have a definite polarity. The polarity is defined by the distribution of yolk.
The vegetal pole has the most yolk and the animal pole has the least.
Body Axes
The development of body axes in frogs is influenced by the polarity of the egg.
The polarity of the egg determines the anterior-posterior axis before fertilization. At fertilization, the pigmented cortex slides over the underlying cytoplasm toward the point of sperm entry. This rotation (red arrow) exposes a region of lighter-colored cytoplasm, the gray crescent, which is a marker of the dorsal side. The first cleavage division bisects the gray crescent. Once the anteriorposterior and dorsal-ventral axes are defined, so is the left-right axis.
Amount of Yolk
Isolecithal very little yolk, even distribution. Mesolecithal moderate amount of yolk concentrated at vegetal pole. Telolecithal Lots of yolk at vegetal pole. Centrolecithal lots of yolk, centrally located.
Cleavage in Frogs
Cleavage planes usually follow a specific pattern that is relative to the animal and vegetal poles of the zygote.
Animal pole blastomeres are smaller. Blastocoel in animal hemisphere. Little yolk, cleavage furrows complete. Holoblastic cleavage
Cleavage in Birds
Occurs in species with yolk-rich eggs, such as reptiles and birds. Blastoderm cap of cells on top of yolk.
When lots of nourishing yolk is present, embryos develop into a miniature adult.
Direct development
When little yolk is present, young develop into larval stages that can feed.
Indirect development
Mammals have little yolk, but nourish the embryo via the placenta.
Blastula
A fluid filled cavity, the blastocoel, forms within the embryo a hollow ball of cells now called a blastula.
Gastrulation
The morphogenetic process called gastrulation rearranges the cells of a blastula into a threelayered (triploblastic) embryo, called a gastrula, that has a primitive gut.
Gastrulation
The three tissue layers produced by gastrulation are called embryonic germ layers.
The endoderm lines the embryonic digestive tract. The mesoderm partly fills the space between the endoderm and ectoderm.
Gastrulation in a sea urchin produces an embryo with a primitive gut (archenteron) and three germ layers. Blastopore open end of gut, becomes anus in deuterostomes.
Gastrulation - Frog
Result embryo with gut & 3 germ layers. More complicated:
Yolk laden cells in vegetal hemisphere. Blastula wall more than one cell thick.
Gastrulation - Chick
Gastrulation - Chick
Epiblast and hypoblast Layers separated by a blastocoel Epiblast forms endoderm and mesoderm.
Gastrulation - Mouse
Inner cell mass will become the embryo while trophoblast becomes part of the placenta.
Protostomes Deuterostomes Spiral vs. radial cleavage Regulative vs. mosaic cleavage Blastopore becomes mouth vs. anus Schizocoelous vs. enterocoelous coelom formation.
Differentiated by:
Deuterostome Development
Deuterostomes include echinoderms (sea urchins, sea stars etc) and chordates.
Radial cleavage
Deuterostome Development
Regulative development the fate of a cell depends on its interactions with neighbors, not what piece of cytoplasm it has. A blastomere isolated early in cleavage is able to from a whole individual.
Deuterostome Development
Deuterostome means second mouth. The blastopore becomes the anus and the mouth develops as the second opening.
Deuterostome Development
Deuterostome Development
Protostome Development
Spiral cleavage
Protostome Development
Mosaic development cell fate is determined by the components of the cytoplasm found in each blastomere.
Protostome Development
Protostome means first mouth. Blastopore becomes the mouth. The second opening will become the anus.
Protostome Development
In protostomes, a mesodermal band of tissue forms before the coelom is formed. The mesoderm splits to form a coelom.
Schizocoely Pseudocoelomates have a body cavity between mesoderm and endoderm. Acoelomates have no body cavity at all other than the gut.
Lophotrochozoan protostomes include annelid worms, molluscs, & some small phyla.
The ecdysozoan protostomes include arthropods, roundworms, and other taxa that molt their exoskeletons.
Ecdysis shedding of the cuticle. Many do not show spiral cleavage.
An organisms development is determined by the genome of the zygote and also by differences that arise between early embryonic cells.
Uneven distribution of substances in the egg called cytoplasmic determinants results in some of these differences. Position of cells in the early embryo result in differences as well.
Induction
In many species that have cytoplasmic determinants only the zygote is totipotent, capable of developing into all the cell types found in the adult.
As embryonic development proceeds, the potency of cells becomes progressively more limited in all species.
Once embryonic cell division creates cells that differ from each other,
Induction
Induction is the capacity of some cells to cause other cells to develop in a certain way. Dorsal lip of the blastopore induces neural development.
Primary organizer
Spemann-Mangold Experiment
Transplanting a piece of dorsal blastopore lip from a salamander gastrula to a ventral or lateral position in another gastrula developed into a notochord & somites and it induced the host ectoderm to form a neural tube.
Sequence of local patterns in which one step in development is a subunit of another. Each step in the developmental hierarchy is a necessary preliminary for the next.
Hox Genes
Hox genes control the subdivision of embryos into regions of different developmental fates along the anteroposterior axis.
Inductive signals play a major role in pattern formation the development of an animals spatial organization.
The molecular cues that control pattern formation, called positional information:
Tell a cell where it is with respect to the animals body axes. Determine how the cell and its descendents respond to future molecular signals.
The wings and legs of chicks, like all vertebrate limbs begin as bumps of tissue called limb buds. The embryonic cells within a limb bud respond to positional information indicating location along three axes.
The second major limb-bud organizer region is the zone of polarizing activity (ZPA).
A block of mesodermal tissue located underneath the ectoderm where the posterior side of the bud is attached to the body.
Morphogenesis
Morphogenesis is a major aspect of development in both plants and animals but only in animals does it involve the movement of cells.
Cell Migration
In gastrulation, tissue invagination is caused by changes in both cell shape and cell migration.
Evo-Devo
Evolutionary developmental biology evolution is a process in which organisms become different as a result of changes in the genetic control of development.
Hox genes
Evo-Devo
Instead of evolution proceeding by the gradual accumulation of numerous small mutations, could it proceed by relatively few mutations in a few developmental genes?
The induction of legs or eyes by a mutation in one gene suggests that these and other organs can develop as modules.
Amniotes
The embryos of birds, reptiles, and mammals develop within a fluid-filled sac that is contained within a shell or the uterus.
Organisms with these adaptations form a monophyletic group called amniotes. Allows for embryo to develop away from water.
Amniotes
In these three types of organisms, the three germ layers also give rise to the
four extraembryonic membranes that surround the developing embryo.
Amniotes
Amnion fluid filled membranous sac that encloses the embryo. Protects embryo from shock. Yolk sac stores yolk and pre-dates the amniotes by millions of years.
Amniotes
Allantois - storage of metabolic wastes during development. Chorion - lies beneath the eggshell and encloses the embryo and other extraembryonic membrane.
Allantois and chorion fuse to form a respiratory surface, the chorioallantoic membrane.
Monotremes
Primitive mammals that lay eggs. Large yolky eggs resembling bird eggs. Duck-billed platypus and spiny anteater.
Marsupials
Embryos born at an early stage of development and continue development in abdominal pouch of mother.
Placental Mammals
Represent 94% of the class Mammalia. Evolution of the placenta required:
Mammalian Development
Mammalian Development
Gastrulation and organogenesis resemble the processes in birds and other reptiles.
Mammalian Development
Blastocyst reaches uterus. Blastocyst implants. Extraembryonic membranes start to form and gastrulation begins. Gastrulation has produced a 3-layered embryo.
Mammalian Development
The extraembryonic membranes in mammals are homologous to those of birds and other reptiles and have similar functions.
Mammalian Development
Amnion
Surrounds embryo Secretes fluid in which embryo floats Contains no yolk Source of stem cells that give rise to blood and lymphoid cells Stem cells migrate to into the developing embryo Not needed to store wastes Contributes to the formation of the umbilical cord Forms most of the placenta
Yolk sac
Allantois
Chorion
Organogenesis
Various regions of the three embryonic germ layers develop into the rudiments of organs during the process of organogenesis.
Organogenesis
Many different structures are derived from the three embryonic germ layers during organogenesis.
Just above the notochord (mesoderm), the ectoderm thickens to form a neural plate.
Edges of the neural plate fold up to create an elongated, hollow neural tube.
Anterior end of neural tube enlarges to form the brain and cranial nerves. Posterior end forms the spinal cord and spinal motor nerves.
Give rise to
Portions of cranial nerves Pigment cells Cartilage Bone Ganglia of the autonomic system Medulla of the adrenal gland Parts of other endocrine glands Important in evolution of the vertebrate head and jaws.
During gastrulation, the archenteron forms as the primitive gut. This endodermal cavity eventually produces:
Digestive tract Lining of pharynx and lungs Most of the liver and pancreas Thyroid, parathyroid glands and thymus
Arise in early embryonic development of all vertebrates. During development, endodermally-lined pharyngeal pouches interact with overlying ectoderm to form gill arches. In fish, gill arches develop into gills. In terrestrial vertebrates:
No respiratory function 1st arch and endoderm-lined pouch form upper and lower jaws, and inner ear. 2nd, 3rd, and 4th gill pouches form tonsils, parathyroid gland and thymus.
Most muscles arise from mesoderm along each side of the neural tube. The mesoderm divides into a linear series of somites (38 in humans).
Axial skeleton Dermis of dorsal skin Muscles of the back, body wall, and limbs Heart