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Today:Today: Group Quiz 6Group Quiz 6 Intro to Intro to
DevelopmentDevelopment
Thursday: Review Paper Thursday: Review Paper Draft Due for Peer Review!Draft Due for Peer Review!
Backtracking: Advantages Backtracking: Advantages and Disadvantages of Sexual and Disadvantages of Sexual
Reproduction?Reproduction?
Generalizations about Sexual Generalizations about Sexual ReproductionReproduction
Offspring created by the Offspring created by the fusion of haploid fusion of haploid gametesgametes to form a diploid to form a diploid zygote.zygote.
The female gamete (The female gamete (ovumovum) is ) is generally large and generally large and
nonmotile.nonmotile.
The male gamete The male gamete ((spermatozoanspermatozoan) is ) is
generally smaller and generally smaller and motile.motile.
Some animals are Some animals are HermaphroditesHermaphrodites (single (single individual has both male and female individual has both male and female
reproductive systems)reproductive systems)
Common in sessile or Common in sessile or burrowing burrowing animals, and in parasitesanimals, and in parasites
Generalizations about Sexual Generalizations about Sexual ReproductionReproduction
Advantages? Does this generate genetic
diversity?
Fertilization OverviewFertilization Overview
Fertilization: the union of sperm and eggFertilization: the union of sperm and egg
Can be Can be EXTERNALEXTERNAL or or INTERNALINTERNAL
External FertilizationExternal Fertilization
Photo by Scott Egan,; University of Rhode Island Dept. of Natural Resources; A pair of wood frogs, Rana sylvatica, in amplexus
Internal fertilization requires Internal fertilization requires cooperative cooperative behaviorbehavior
andand
Sophisticated Sophisticated reproductive systemsreproductive systems including organs to deliver sperm and including organs to deliver sperm and
receptacles for its storage and transport to receptacles for its storage and transport to eggs.eggs.
Internal FertilizationInternal Fertilization
Mammalian Mammalian Gamete Gamete
Production: Production: FemalesFemales
Mammalian Mammalian Gamete Gamete
Production: Production: Males Males
Mammalian Gamete Production: Mammalian Gamete Production: MalesMales
Aside: FertilizationAside: Fertilization
Preformation vs EpigenesisPreformation vs Epigenesis
FertilizationFertilization Combines haploid Combines haploid
sets of sets of chromosomes to chromosomes to form single diploid form single diploid zygotezygote
Activation of the Activation of the eggegg
Model Organism = Model Organism = Sea UrchinSea Urchin
Why Urchins?
?
Studying Sea Urchin Studying Sea Urchin FertilizationFertilization
Source: NASA- Photo credit - Joseph Tash
External Fertilization in Sea UrchinsExternal Fertilization in Sea Urchins Sperm exposed to molecules from the jelly Sperm exposed to molecules from the jelly
coat surrounding an egg, the acrosome coat surrounding an egg, the acrosome discharges its contents by Exocytosisdischarges its contents by Exocytosis
= Acrosomal Reaction= Acrosomal Reaction The acrosomal process (elongating The acrosomal process (elongating
structure) then penetrates the jelly coat of structure) then penetrates the jelly coat of the egg and adheres to a specific receptor the egg and adheres to a specific receptor moleculemolecule
Fertilization: The Acrosomal Fertilization: The Acrosomal ReactionReaction
Fertilization: The Acrosomal Fertilization: The Acrosomal ReactionReaction
What needs to happen once one
sperm nucleus has entered the
egg?!
Fusion of the sperm and egg Fusion of the sperm and egg membrane causes membrane causes ion channelsion channels in in the egg’s membrane to openthe egg’s membrane to open
Sodium ions flow into the cell, Sodium ions flow into the cell, causing a causing a membrane membrane depolarizationdepolarization (change in (change in membrane potential)membrane potential)
Prevents more than 1 sperm from Prevents more than 1 sperm from entering (1-3 seconds!)entering (1-3 seconds!)
Fertilization: Fast Block to PolyspermyFertilization: Fast Block to Polyspermy
Membrane potential is restored within 2 minutes! (Uh oh…)
The Cortical The Cortical Reaction: Reaction:
Fusion of sperm and Fusion of sperm and egg triggers release egg triggers release of of calciumcalcium from the from the ER into the cytosolER into the cytosol
Calcium release Calcium release begins at site of begins at site of sperm entry and sperm entry and propagates across propagates across the fertilized eggthe fertilized egg
Fertilization: Slow Block to Fertilization: Slow Block to PolyspermyPolyspermy
Fertilization: Slow Block to Fertilization: Slow Block to PolyspermyPolyspermy
The Cortical The Cortical Reaction:Reaction:
High calcium High calcium concentration causes concentration causes the the cortical cortical granulesgranules to fuse to fuse with plasma with plasma membranemembrane
Mouse oocyte stained to show cortical granules (small red dots). Genomic DNA in the metaphase plate of the secondary oocyte (top right) appears bluish-white.(From Biology of Reproduction 57:743-750, 1997, Z. Xu, A. Abbott, G. Kopf, R. Schultz and T. Ducibella)
Fertilization: Slow Block to Fertilization: Slow Block to PolyspermyPolyspermy
Enzymes from the Enzymes from the cortical granulescortical granules separate the separate the vitelline layervitelline layer from from the plasma membranethe plasma membrane
Water is drawn into the Water is drawn into the perivitelline perivitelline spacespace by osmosis, swelling it by osmosis, swelling it
The swelling pushes the vitelline The swelling pushes the vitelline layer away from plasma membrane layer away from plasma membrane where it is hardened by enzymes to where it is hardened by enzymes to form the form the Fertilization EnvelopeFertilization Envelope
Fertilization: Slow Block to Fertilization: Slow Block to PolyspermyPolyspermy
Fertilization: Slow Block to Fertilization: Slow Block to PolyspermyPolyspermy
Rise in Calcium also induces Rise in Calcium also induces metabolic changesmetabolic changes within the egg within the egg
Metabolism increases rapidly!Metabolism increases rapidly! Nucleus of sperm starts to swellNucleus of sperm starts to swell Sperm nucleus merges with egg Sperm nucleus merges with egg
nucleus (@ ~ 20 minutes)nucleus (@ ~ 20 minutes) First division occurs @ ~90 minutesFirst division occurs @ ~90 minutes
Fertilization: ActivationFertilization: Activation
Timing:Timing:
Sea Urchin Larva at 13
days
Fertilization: Sea Urchins Vs. Fertilization: Sea Urchins Vs. MammalsMammals
Internal Fertilization (terrestrial!)Internal Fertilization (terrestrial!)
Key Initial Differences:Key Initial Differences:
1. 1. CapacitationCapacitation: molecules in mammalian : molecules in mammalian female reproductive tract alter surface of female reproductive tract alter surface of sperm and increase the motility (~ 6 hrs)sperm and increase the motility (~ 6 hrs)
2. Mammalian egg 2. Mammalian egg cloaked by follicle cloaked by follicle cells-cells- capacitated sperm cell must capacitated sperm cell must migrate through this layer to reach the migrate through this layer to reach the zona pellucidazona pellucida
Fertilization: Sea Urchins Vs. Fertilization: Sea Urchins Vs. MammalsMammals
The Zona PellucidaThe Zona Pellucida = filamentous network = filamentous network of glycoproteins (extracellular matrix)of glycoproteins (extracellular matrix)
One of the glycoproteins, ZP3 functions One of the glycoproteins, ZP3 functions as a sperm receptoras a sperm receptor
Binding induces the acrosome to release Binding induces the acrosome to release its contents (as with sea urchins)its contents (as with sea urchins)
Enzymes from acrosome help sperm Enzymes from acrosome help sperm penetrate the zona pellucidapenetrate the zona pellucida
Fertilization: Sea Urchins Vs. Fertilization: Sea Urchins Vs. MammalsMammals
As with sea urchin, As with sea urchin, binding of egg binding of egg triggers triggers depolarizationdepolarization of egg of egg membrane (fast block to membrane (fast block to polyspermy)polyspermy)
A A Cortical ReactionCortical Reaction functions as the slow functions as the slow block (granules in cortex block (granules in cortex release contents; release contents; enzymes catalyze enzymes catalyze changes in the zona changes in the zona pellucida)pellucida)
Fertilization: Sea Urchins Vs. Fertilization: Sea Urchins Vs. MammalsMammals
Fertilization: Sea Urchins Vs. Fertilization: Sea Urchins Vs. MammalsMammals
Unfertilized Star Fish EggsUnfertilized Star Fish Eggs
Fertilization Reviewed
Fertilized egg (zygote) Fertilized egg (zygote) - the nucleus is no longer visible; the fertilization - the nucleus is no longer visible; the fertilization envelope is presentenvelope is present
Let’s Watch: Let’s Watch: http://www.youtube.com/watch_popup?v=jp-RgIRgcYE#t=000http://www.youtube.com/watch_popup?v=jp-RgIRgcYE#t=000
Stages of DevelopmentStages of Development
Fertilization is followed by 3 Fertilization is followed by 3 stages:stages:
1. Cleavage1. Cleavage
2. Gastrulation2. Gastrulation
3. Organogenesis3. Organogenesis
1.1. CLEAVAGECLEAVAGE
Succession of rapid cell divisionsSuccession of rapid cell divisions Partitions the cytoplasm into smaller Partitions the cytoplasm into smaller
cells, cells, BLASTOMERESBLASTOMERES, each with its own , each with its own nucleusnucleus
Each region of cytoplasm contains Each region of cytoplasm contains different cytoplasmic componentsdifferent cytoplasmic components
2-cell and 4-cell stage of Sea Urchin Cleavage (Blastomeres)
What happened to the size of each blastomere??
Polarity of Zygote and EggPolarity of Zygote and Egg
Most animals (except mammals) Most animals (except mammals) produce asymmetrical eggs and produce asymmetrical eggs and zygoteszygotes
Distribution of yolk, mRNA and Distribution of yolk, mRNA and proteins is proteins is not not uniformuniform
Sets the stage for subsequent
developmental events!
Yolk is Yolk is concentrated concentrated
at the at the vegetal vegetal
polepole..
The opposite The opposite pole is the pole is the
animal poleanimal pole
(Becomes the dorsal
side)
Polarity of Frog EggsPolarity of Frog Eggs
In the final stage, a In the final stage, a frog oocyte is frog oocyte is
pigmented dark pigmented dark brown in one brown in one hemisphere hemisphere
((animal poleanimal pole). ). The other hemisphere The other hemisphere
((vegetal polevegetal pole) ) shows the yellow shows the yellow color of the egg color of the egg
yolk.yolk.
CleavageCleavage
The first two cleavage divisions are The first two cleavage divisions are vertical producing four cells vertical producing four cells
extending from animal to vegetal extending from animal to vegetal pole.pole.
The third division is horizontal, The third division is horizontal, producing eight cells.producing eight cells.
Continued divisions (16-64 cells) Continued divisions (16-64 cells) produce a solid ball of cells, the produce a solid ball of cells, the
morula.morula.
A fluid-filled cavity, the blastocoel forms within the morula, creating a hollow ball of cells, the blastula.
Note the difference the presence of yolk makes!
Sea Urchin Sea Urchin MorulasMorulas - 8- - 8-16 cell stage 16 cell stage
Sea Urchin Sea Urchin BlastulaBlastula - 32-cell stage, - 32-cell stage, blastocoel blastocoel
Sea Urchin: Late Blastula - shows thickened Sea Urchin: Late Blastula - shows thickened layer of cells at one end, the vegetal pole layer of cells at one end, the vegetal pole
Morula (16-64 cells) Morula (16-64 cells) Blastula (>128 Blastula (>128 cells)cells)
A fluid-filled cavity, the blastocoel, forms A fluid-filled cavity, the blastocoel, forms within the morula creating a hollow ballwithin the morula creating a hollow ball
Source: courtesy of Dr. J. Hardin, Univ. of
Wisconsin
Blastula Blastula Gastrula Gastrula
Changes in cell motility, shape Changes in cell motility, shape and adhesion result in the spatial and adhesion result in the spatial rearrangement of an embryorearrangement of an embryo
Results in three familiar Results in three familiar embryonic tissue layers: embryonic tissue layers:
ectoderm, endoderm and ectoderm, endoderm and mesodermmesoderm
2. 2. GastrulationGastrulation
Source: raven.zoology.washington.edu/ embryos
Sea Urchin: Early Gastrula - blastocoel, Sea Urchin: Early Gastrula - blastocoel, blastopore, primitive gut (archenteron)blastopore, primitive gut (archenteron)
Sea Urchin: Late Gastrula - endoderm, Sea Urchin: Late Gastrula - endoderm, ectoderm, mesoderm ectoderm, mesoderm
Blastula Blastula
Gastrula Gastrula in a in a FrogFrog
Frog DevelopmentFrog Development
Great model organisms:Great model organisms:
HUGE eggs! (> 1mm)HUGE eggs! (> 1mm) External fertilization and developmentExternal fertilization and development Vertebrates!Vertebrates! Fertilization Fertilization Neuralation in 18 hours Neuralation in 18 hours
under good conditionsunder good conditions
anatomy.med.unsw.edu
Zooming in on the Frog Zooming in on the Frog GastrulaGastrula
How do we How do we know??know??
Stages of DevelopmentStages of Development
Fertilization is followed by 3 Fertilization is followed by 3 stages:stages:
1. Cleavage1. Cleavage
2. Gastrulation2. Gastrulation
3. Organogenesis3. Organogenesis
3. Organogenesis3. Organogenesis
Morphogenetic changes – folds, splits Morphogenetic changes – folds, splits and clusterings (condensation) begin and clusterings (condensation) begin
the process of organ buildingthe process of organ building
In chordates, the neural tube and In chordates, the neural tube and notocord form firstnotocord form first
Sea Urchin: the bipinnaria larva (bilateral Sea Urchin: the bipinnaria larva (bilateral symmetry) develops into a brachiolaria larva; this symmetry) develops into a brachiolaria larva; this larva undergoes metamorphosis to become the larva undergoes metamorphosis to become the
adult starfish, which has radial symmetry. adult starfish, which has radial symmetry.
Earliest Organogenesis in Frog Earliest Organogenesis in Frog (Chordate) Embryos(Chordate) Embryos
Let’s Watch –http://www.youtube.com/watch_popup?v=-KC1CA_Os1g#t=000
Formation of Formation of the Neural the Neural Tube in a Tube in a ChordateChordate
Species: Mouse Day Gestation: 8 Species: Mouse Day Gestation: 8 www.med.unc.eduwww.med.unc.edu