Modes of reproduction among eukaryotes

OF WHAT BENEFIT IS SEX?

 Sexual periodicity is much less strongly developed in the male than in the female.  Among primates, the male is sexually potent throughout life.  Rutting season- a brief period of pronounced Public Display of Affection sexual activity among Punishable! males; always coincides with estrus or heat period of females.

ESTRUS CYCLE- repeated series of changes in the female reproductive mechanisms. 1.Monestrous - breeds once a year 2.Polyestrous - several breeding periods in a year:  Pregnancy, starvation, extreme exposure or sickness suppress estrus  Domestication may cause shift to polyestrous rhythm  Critical initiating factors must be present (ex. light in rabbits)

GAMETOGENESIS
1. Origin of germ cells and their migration to the gonads 2. Proliferation of germ cells by mitosis 3. Meiosis 4. Final stages of maturation for gametes

1. Origin of germ cells and their migration to the gonads

 In many species, a distinctive germ plasm exists. It often contains the Oskar, Vasa, and Nanos proteins or the mRNAs encoding them.  The precursors of the gametes are the primordial germ cells. They form outside the gonads and migrate into the gonads during development.  In Drosophila, the germ plasm becomes localized in the posterior of the embryo and forms pole cells, the precursors of the gametes.  In frogs, the germ plasm originates in the vegetal portion of the oocyte.  In birds, the germ plasm is first seen in the germinal crescent. The germ cells migrate through the blood, then leave the blood vessels and migrate into the genital ridges.

 Although no germ plasm has been found in mammals, primordial germ cells (PGCs) arise from yolk sac endodermal cells with the expression of the nuclear transcription factor Oct 4 restrictively expressed in the inner cell mass.  During gastrulation, Oct 4 becomes expressed solely in those posterior epiblast cells thought to give rise to the primordial germ cells.  After that, this protein is seen only in the PGCs and oocytes (Oct4 is not seen in the sperm after the germ cells become committed to sperm production.)

(A). Final stages of migration of mammalian PGCs through the hindgut into the two genital ridges, each of which will develop into a gonad (B) The migrating primordial germ cells in an early mouse embryo are stained with a monoclonal antibody (green) that specifically labels these cells at this stage of embryogenesis. The remaining cells in the embryo are stained with a lectin that binds to sialic acid, which is found on the surface of all cells.

How Do the PGCs Know Where to Go?

 In vitro evidence suggests that the genital ridges of 10.5-day mouse embryos secrete a diffusible TGF- 1like protein that is capable of attracting mouse PGCs.  The stem cell factor and LIF contribute to their proliferation along the path of migration.  Using alkaline phosphatase is a germ cell "marker enzyme" for PGCs, they are shown to migrate directionally to gonads by amoeboid movement, presence of chemical attractants, orientation by physical characteristics of the fibronectin pathways, or may be provided by a gradient of soluble protein.  In mammals, a similar migration is seen, using fibronectin pathways. Germ cells that lack the integrin receptor for such ECM proteins cannot migrate to the gonads

PGCs migrate by extending filopodia (fine pseudopods) and they can migrate between cells in tissues. They may follow ECM components that are lined with fibronectin. PGCs (red) might use their surface integrins to follow an embryonic "roadway" of ECM to reach the genital ridges. The yellow is meant to reflect a theoretical complex of ECM components (e.g., laminin, fibronectin, collagen) rather than a single entity such as fibronectin. Cells would stay within the yellow ECM region rather than wander into adjacent regions because they would have preferential adhesiveness ("stickiness") to the yellow "roadway".

 Treating PGCs with FGF2 convert them to embryonic germ (EG) cells which has the potential to differentiate into any cell type of the body.  Cells deposited in extragonadal areas (mediastinum, sacrococcygeal or oral regions) die, or develop into teratomas which may contain mixtures of differentiated tissues (skin, hair, cartilage and teeth!)

GENOMIC IMPRINTING  Imprinting-switching off of genes that are not expressed in development. Paternal or maternal genomes maybe switched off during germ-cell formation.  Reciprocal imprinting: paternal genes promote growth whereas maternal imprinting reduces them (e.g. the paternal Igf-2 gene promotes a large placenta; reduction of maternal gene allows the mother to spread her resources over all embryos)  Imprinting is reversible; in mutants, failures and abnormalities in development result from mismatched timing of gene suppression.

Mechanism of imprinting of the mouse Igf2 gene. On chromosomes inherited from the female, a protein called CTCF binds to an insulator, blocking communication between the enhancer (green) and the Igf2 gene (orange). Igf2 is therefore not expressed from the maternally inherited chromosome. Because of imprinting, the insulator on the male-derived chromosome is methylated; this inactivates the insulator, by blocking the binding of the CTCF protein, and allows the enhancer to activate transcription of the Igf2 gene.

BIG DECISIONS: Mitosis or Meiosis? Sperm or Egg?

In nematodes, the meiosis/mitosis decision is regulated by the Delta proteins, which bind to the Notch proteins on the PGCs. On the other hand, the signal for a germ cell to become either a sperm or an egg is regulated at the level of translation of the fem-3 mRNA containing a sequence that binds a repressor protein which is a combination of the Nanos and Pumilio proteins.

MEIOSIS

http://highered.mcgrawhill.com/olc/dl/120074/bio19.swf

Prophase I is divided into sub-phases:

 Synapsis is the process of linking of the replicated homologues.  Bivalents- a pair of homologous chromosome  Tetrad- two chromatids from each chromosome, forming a thick 4-strand structure at the synaptonemal complex.  Dyad- sister chromatids of a single chromosome  Monad- one of the sister chromatids of a single chromosome

Schematic drawing of a synaptonemal complex. Before the synaptonemal complex forms, recombination complexes assemble on double-strand DNA breaks on sister chromatids and help catalyze crossing-over between non-sister chromatid loops from opposite sides of the complex.

A molecular model of how transverse filaments may be formed by a single type of protein. (A) A diagram of the polypeptide chain showing the N- and Cterminal globular domains, connected by a coiled-coil region. (B) It is proposed that the protein forms homodimers, which then interact across the 100 nm gap separating the axial cores of the two homologs.

Chiasmata (from the Greek letter Chi, which is written as X) are regions where homologous chromosomes cross one another and often break, exchanging parts with their homologue). Because of crossing over, it is very unlikely that any two sperms or eggs would ever carry the same arrangements of DNA. This process insures even more genetic variability that can occur with the various arrangements of chromosome pairs.

 Cellular products give rise to monoploid nuclei that maybe restored to a single diploid nucleus during syngamy.  If genes are located on chromosomes, meiosis achieves genetic variability through: random assortment of paternal/maternal chromosomes at metaphase I, AND crossingover.  Assuming one gene per chromosome, the chance of a given gamete to receive any particular arrangement of paternal/maternal chromosomes and genes is ()n; where n = monoploid number of species.

 For humans, a gamete will have ()23 chance of having a specific combination of chromosomes. Zygotic genotypes resulting from random fusions of these gametes is equal to its squared value.  Chiasmata further lowers the probability that any given gamete will have a similar genotype with another cell.

Independent assortment, and crossingover leads to Genetic Variation

http://highered. mcgrawhill.com/olc/dl/12 0074/bio18.swf

http://highered.mcgraw-hill.com/olc/dl/120074/bio17.swf

LIFE CYCLES: ALTERNATION OF GENERATIONS

O O G E N E S I S

S P E R M A T O G E N E S I S

http://www.sumanasinc.com/webcontent/animations /content/meiosis.html http://www.cellsalive.com/meiosis.htm http://www.johnkyrk.com/meiosis.html