oogenesis behind every successful embryo stands a hard-working mother
TRANSCRIPT
Oogenesis
Behind every successful embryo stands a hard-working mother.
Eggs are enormous cellscarrying nutrients forthe developing embryo
Why? Mass of egg must equal the mass of the embryo that emerges!
Even human eggsare huge cells
Eggs are enormous cellscarrying nutrients forthe developing embryo
• yolk=large stores of nutrients• large stores of macromolecules• most eggs surrounded by a coat or shell
The germ line is set aside early in development as a separate cell lineage
This can sometimes be visualized by the segregation of putative "determinants”as in C. elegans
Fertilized egg
"P granule"
mother cell of germline
= P4
P1HOW are P-granules segregated?
PAR proteins & microfilaments see p. 246 and Fig. 8.43
C. elegans:
P granulesare segregated at each divisionand make thecell that inherits them the germline
Figure 8.44 cell nuclei P-granules
Similar mechanisms lead to very early segregationof the germline in thefruit fly Drosophila
Germ cellprogenitors
A P
The Germline is also set aside in Humans
Oocytes arrest in prophase of meiosis I for up to 50 years!
Figure 19.23
How does a single egg cell make all the stuff needed to start development?
Fly Oogenesis
Nurse cells are mitotic sisters of the oocyte
Figure 19.4
Nurse cells synthesize macromolecules and pump them into the oocyte through the cytoplasmic bridges
oocyte
nurse cells
follicle cells
The Drosophila ovary is composed of somatic follicle cells, germ line nurse cells, and the oocyte
Most of the instructions for forming organisms are already in the egg!
Asymmetric distribution of egg contents influences embryonic development
Molecules synthesized by the nurse cells and specifically localized in the oocyte determine
the anterior/posterior axis of the embryo
Posterior Determinant
(nanos)
Anterior Determinant
(bicoid)
Oocyte
Specific mRNAs are bound by proteins and transported along microtubules by motor proteins to their destinations
Ventral follicle cells build a signal into the egg shell that determines the dorsal/ventral axis of the embryo
Ventral follicle cells
Oocyte signals follicle cells,follicle cells signal back to oocytecytoplasm
nucleus
gurken mRNA
PosteriorAnterior
Dorsal
Ventral
Torpedo receptor binds Gurken; signal inhibits Pipe in dorsal follicle cells
Pipe protein
See Fig. 9.11for more details
Pipe intiates a signaling pathway for activation of Dorsal transcription factor in ventral cells only
nucleus
Ventral signal
initiated by
Transmembrane receptor(Toll)
Signaling pathway
Transcription factor (Dorsal) relocalized to ventral nuclei of embryo activating new genes
Pipe
Localized mRNAs also influencevertebrate development
Vg1 mRNA
See also Figure 5.38
Human oogenesis:An example of hormonal control
of reproduction
Meiosis I
2 million present at birth ovulation
Cyclic changes in hormonal signals regulate the timing of femalereproduction
Most mammals-once/year
Humans- monthly
Release into the Fallopian tube
(Turns off FSH and LH!)
Cyclic changes in hormonal signals regulate the timing of femalereproduction
(Turns off FSH and LH!)
Release into the Fallopian tube
RU486- blocks progesterone receptors
Plan B-progestin (interferes with ovulation, fertilization, implantation)
Progesterone and estrogen birth control pills-prevent maturation of new ova
Anton van Leeuwenhoek
WNYC’s Radiolab clip from “Sperm” (12/2/08)By Jad Abumrad & Robert Krulwich
Figure 7.1 The Human Infant Preformed in the Sperm, as Depicted by
Nicolas Hartsoeker
(1694)
The sperm: a stripped downmachine for delivering DNA
Figure 7.2
Chemical cues from the egg attract sperm
In sea urchins, the cue is resact
Inject resact
Figure 7.9 Sperm chemotaxis
0 sec 20 sec 40 sec 90 sec
Step one:
Contact between the sperm and the eggshell
Step two:
Contact between the sperm and the egg’splasmamembrane
Fertilization: a multistepprocess
(zona pellucida=egg shell)
Sperm-eggshell contact triggers the acrosome reaction
Example 1-- the sea urchin
enzymes
Species specificity (equivalent to zona pellucida)
attracts/activates sperm
Figure 7.8
Sperm-eggshell contact triggers the acrosome reaction
Example 2: Mammals
Eggshell includes ZP3/Sperm have ZP3 receptor
Figure 7.8
BLOCKING POLYSPERMY
1. Fast/transient: prevent sperm fusion
2. Slow/permanent: removal of other bound sperm
A complex signal transduction pathway is activated in the egg by fertilization
Example-- the sea urchin egg
1. Na in+
2. Phospholipase C activated
InsP3DAG +
3. Ca release2+
Ca up 2+
5. Protein kinase C activated
PKC
H+
H+
6. pH increase
7. Increased protein synthesis
8. Initiate DNAreplication
4. Cortical granule release; blocks
polyspermy
Fast block1-3 secondslasts 1 min
Slow block20-60 secondspermanent
How do we know this?We can measure these events directly!
Figure 7.17 Membrane potential of sea urchin eggs before andAfter fertilization.
resting -70 (sperm can fuse with egg)
(sperm can’t fuse with egg)
+20 mV
Cortical granules:poised for release
15,000 cortical granules in a sea urchin egg• contain enzymes that clip the egg’s bindin receptor and any attached sperm• alter the vitelline envelope (zona pellucida in mammals so that ZP3 receptor on sperm cannot bind)
Blocking polyspermy: SLOW BLOCKFigure 7.6
Sperm-eggshell contact triggers the acrosome reaction
Example 2: Mammals
Sperm have ZP3 receptor
A wave of increased calcium can be visualizedmoving across the egg from the point of sperm entry
Figure 7.20
A complex signal transduction pathway is activated in the egg by fertilization
Example-- the sea urchin egg
1. Na in+
2. Phospholipase C activated
InsP3DAG +
3. Ca release2+
Ca up 2+
5. Protein kinase C activated
PKC
H+
H+
6. pH increase
7. Increased protein synthesis
8. Initiate DNAreplication
4. Cortical granule release; blocks
polyspermy
Fast block1-3 secondslasts 1 min
Slow block20-60 secondspermanent
7.4
200
7.0
7.2
6.8
pH
time (seconds)
intracellular pH
400 60010-8
10-7
10-6
intracellular
calcium
10-5
Ca2+