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Animal Development

Introduction
How a single cell (fertilized egg) develops into multicellular individual is fundamental question in modern biology Gametes are haploid reproductive cells o In animals, male gametes = sperm o In animals, female gametes = eggs Development proceeds in ordered phases through animals life cycle: o Gametogenesis o Fertilization o Cleavage o Gastrulation o Organogenesis

Gametogenesis
Gametogenesis: formation of gametes in reproductive organs of adult organisms Both sperm and egg contribute chromosomes o Mammalian gametes have haploid genome containing one allele of each gene will be contributed to offspring Egg cells contribute more than just chromosomes and are much larger than sperm cells o contribute nutrients and developmental regulatory molecules (cytoplasmic determinants)

Sperm Structure and Function


Sperm cell is composed of four main structures: 1. Head: contains nucleus and enzyme-filled acrosome o Acrosomal enzymes allow sperm to penetrate egg 2. Neck: contains centriole o fuses with eggs centriole to form the centrosome 3. Midpiece: packed with mitochondria produces ATP necessary for movement 4. Tail: has flagellum that acts as propeller

Egg Structure and Function


Egg cells are relatively large and nonmotile o size largely due to nutrient storage, required for early embryonic development Quantity of nutrients varies across species o Relatively small mammalian egg only has to supply nutrients for early development Embryos obtain nutrition through placenta shortly following fertilization o Egg-laying species produce larger eggs Egg yolk is embryos sole source of nutrition prior to hatching

More Egg Structures


Many contain cytoplasmic determinants that control early events of development o often are transcription factors or mRNA encoding transcription factors (ex. bicoid) Many contain cortical granules o small enzyme-filled vesicles activated during fertilization Vitelline envelope (fibrous, mat-like sheet of glycoproteins) surrounds egg Some species have jelly layer (thick, gelatinous matrix around vitelline envelope for further protection)

Fertilization
Fertilization: fusion of haploid sperm and egg cells to form a diploid zygote (fertilized egg) Many conditions must be met before zygote can form: o Gametes must be in same place at same time o Gametes must recognize and bind to each other o Gametes must fuse together o Fusion must trigger onset of development Sea urchins are model system for studying fertilization o produce large number of gametes and undergo external fertilization

Fertilization in Sea Urchins


Sperm head contacts jelly layer 1. Acrosomal enzymes digest through egg's jelly layer and vitelline envelope 2. Plasma membranes of sperm and egg fuse when sperm head contacts surface of the egg cell Fertilization complete when: o sperm nucleus, mitochondria, and centriole enter the egg o sperm and egg nuclei fuse to form the zygote nucleus

Gametes from the Same Species Recognize Eachother


Bindin: protein on head of sea urchin sperm that binds to surface of sea urchin eggs o acts as key - sperm only binds eggs of same species During sea urchin fertilization: o species-specific bindin molecules on sperm interact with species-specific receptors on surface of egg interaction is required for plasma membranes of sperm and egg to fuse

Identifying the Egg Cell Receptor for Sperm


Researchers treated sea urchin eggs with a protease (enzyme that cleaves peptide bonds) o isolated fragment of egg cell receptor that binds to sperm Fertilizin: receptor on surface of sea urchin eggs involved in binding sperm o acts as lock to bindins key Each species has own version of fertilizin and bindin o makes lock-and-key interaction species specific and necessary for fertilization

Why Does Only One Sperm Enter the Egg?


Animals employ different mechanisms to avoid polyspermy (fertilization by more than one sperm) In sea urchins, fertilization stimulates creation of physical barrier o Ca2+-based signal is rapidly induced and propagated throughout egg results in formation of fertilization envelope - keeps away additional sperm In mammals, cortical granules release enzymes that modify egg cell receptors o prevents binding by additional sperm

Cleavage
Cleavage: set of rapid cell divisions that take place in animal zygotes immediately after fertilization o first step in embryogenesis o process makes single-celled zygote into multicellular embryo o partitions egg cytoplasm without additional growth of zygote Blastomeres: cells created by cleavage divisions When cleavage is complete, embryo consists of mass of blastomere cells called blastula

Fertilization requires that _____.


A. an egg be exposed only to a single sperm B. the egg underwent cleavage C. sperm are immobilized D. proteins on egg and sperm membranes interact E. all of the above

Cytoplasmic Determinants
Cytoplasmic determinants: o found in specific locations within egg cytoplasm o end up in specific populations of blastomeres Egg cytoplasm divides precisely (during cleavage) to distribute cytoplasmic determinants to certain cells o Initiates step-by-step process that, in combination with signals received from other cells, results in differentiation of cells

If an egg distributed all the constituents of its cytoplasm uniformly, what consequence would be most logical?
A. all cells will express the same genes B. all cells will die C. all cells will move D. all genes will be expressed E. all genes will be silenced

Cleavage in Mammals
Cleavage results in blastocyst (specialized blastula consisting of two populations of cells): o external, thin-walled hollow trophoblast o inner cell mass (ICM) contains cells that undergo gastrulation and develop into the embryo

Gastrulation
Extensive and highly organized cell movements radically rearrange embryonic cells into structure called the gastrula Gastrulation: results in formation of embryonic tissue layers o Tissue: integrated set of cells that function as a unit Most early embryos have three primary tissue layers: ectoderm, mesoderm, and endoderm o embryonic tissues (germ layers) give rise to adult tissues and organs

Gastrulation in Frog Embryos


1. Frog blastula contains fluid-filled space (blastocoel) 2. Gastrulation begins with formation of an opening (blastopore) 3. Cells from periphery move inward through blastopore, forming tube-like structure that will become the gut

Germ Layers
Ectoderm: forms nervous system and outer covering of adult body Mesoderm: gives rise to muscle, most internal organs, and connective tissues such as bone and cartilage Endoderm: produces lining of digestive tract (gut), along with some of associated organs

Movie

Completion of Gastrulation
In addition to establishing embryonic tissue layers, gastrulation has another major outcome: o major body axes become visible o ex) blastopore in frog becomes anus At end of gastrulation: o three embryonic tissues are arranged in layers o gut has formed o major body axes have become visible

Organogenesis
Organogenesis: process of tissue and organ formation o begins once gastrulation is complete and embryonic germ layers are in place o cells proliferate and become differentiated become specialized cell type

Differentiated cells have distinctive structure and function because express distinctive suite of genes!

Organizing the Mesoderm into Somites


Early in organogenesis, rod-like notochord appears in dorsal mesoderm o structure unique to animal group called the chordates (includes humans and other vertebrates) o functions as key organizing element during organogenesis in many chordates, as organogenesis continues, notochord cells undergo apoptosis transient structure (true for humans)

Function of Notochord: Neural Tube Formation


Cells in or near the notochord produce signaling molecules o triggers reorganization of dorsal ectodermal cells o leads to neural tube formation neural tube is precursor to brain and spinal cord

Somite Formation
Once neural tube forms o mesodermal cells become organized into blocks of tissues called somites form on both sides of neural tube down length of the body

Somite Maturation and Determination


Somite cells are transient structures o form variety of future structures o cells from somites will migrate and proliferate and develop into key body elements Somite cells form variety of structures, but are initially not determined o initially, they can become any of the somite-derived elements of the body As the somite matures o somite cells become irreversibly determined will eventually differentiate into specific cell type based on their location within the somite

Initially, somite cells can become any of somitederived elements of body As somite matures o somite cells become irreversibly determined

Cell Determination
Somite cells differentiate in response to signals from nearby tissues (dependent on their location) o step-by-step process guided by regulatory gene cascade! o signals diffuse away from cells in notochord, neural tube, and nearby ectoderm and mesoderm act on specific populations of target cells in somite triggers production of transcription factors required for expression of tissue/organ specific proteins!

Ex.) Differentiation of Muscle Cells


Myoblast: somite derived cell determined to become muscle cell but has not begun producing muscle-specific proteins Researchers hypothesized that myoblasts must contain at least one regulatory protein that commits them to their fate o found that MyoD was protein that causes muscle cell differentiation MyoD: regulatory transcription factor that binds enhancers upstream of muscle-specific genes

Overview of Cell Differentiation ex.) myoblast


Differentiation is a step-by-step process that is complete when cells begin producing cell-specific proteins. Steps of differentiation in a muscle cell are: 1. Fertilization triggers cleavageresulting in blastocyst 2. Blastocyst cell signals trigger gene cascadesresulting in positional changes during gastrulation 3. Notochord signals tell specific somite cells to begin MyoD productiontargeting cells as belonging to specific muscles 4. Muscle cells begin expressing muscle-specific proteins

Muscle cells are created from mesoderm and express a regulatory protein MyoD. In comparison, neurons _____.
A. don t have MyoD gene B. don t express MyoD gene C. will transform into muscle cells if MyoD is expressed D. don t have MyoD gene and don t express MyoD gene E. don t express MyoD gene and will transform into muscle

cells if MyoD is expressed

Concepts to Remember
Fertilization begins with specific interactions between proteins on the plasma membranes of sperm and egg. Earliest cell divisions divide fertilized egg into mass of cells individual fates depend on key regulatory molecules they contain and signals they receive. Early in development, cells undergo massive, coordinated movements to form distinct tissue layers. o Each layer gives rise to different set of tissues and organs As development proceedsspecialized organs and other structures form through interacting effects of cell-cell signals, cell proliferation, cell movements, and differentiation. o Differentiation is complete when cells express tissuespecific proteins.

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