Chapter 7

Meiosis and germ cell formation

Asexual reproduction
Mitosis division produces daughter cells which are usually genetically identical
to each other to the parental cell

Prokaryotes and most unicellular eukaryotes reproduce asexually: by binary fission, budding or more exotic ways of dividing. Asexual reproduction does not allow easy adaptation toward changing environmental conditions.
Where does genetic variation come from?

Where does genetic variation come from?

Mutations Sexual reproduction
Allows genetic material of two parental cells to be mixed. As a result, progeny is formed which is genetically different: from parent cells and in between various progeny organisms.

Diploidity: a problem in sexual reproduction

A diploid genome: One copy of each chromosome is from the mother and one is from the father. Both carry the same genes, but may have different alleles.

Diploidity: a problem in sexual reproduction

Somatic cells are diploid (2n) Example: an organism is 2n = 4 chromosomes altogether. Chromosomes 1 & 2 are homologous chromosomes Chromosomes 3 & 4 are homologous chromosomes Chromosomes 1 & 3 came from the mother Chromosomes 2 & 4 came from the father Fusion of two 2n-cells: 4n-cell.

for sexual reproduction diploid cells are useless

For sexual reproduction, specialized cells are needed that carry half the set of chromosomes: gametes. In more complex organisms such haploid (=1n) organisms are gametes, typically ovum and spermatozoon. Union of both gametes (fertilization) results in a diploid zygote.

Sexual reproduction in higher and lower eukaryotes

While the way they make use of their haploid cells differs the general mechanism to generate haploid cells is the same: meiosis.

Stages of meiosis
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2n 2c 1n 2c

2n 4c

1n 1c

Meiosis I
Segregates homologous chromosomes into different daughter cells. In meiosis I exchange of corresponding parts of homologous chromosomes can occur. Both meiosis I and II can be divided in subphases similar to those of mitosis: Prophase Metaphase Anaphase Telophase

Prophase of meiosis I
The chromosomes condense and two centrosomes start to migrate to opposite poles of the cell. Each chromosome consist of two chromatids. Homologous pairs form bivalents (=synapses of homologous chromosomes).

Prophase of meiosis I consists of five subphases

The events of prophase I can be further divided into five stages:
Cell Biology 2007/2008

Leptotene Zygotene Pachytene Diplotene Diakinesis

Crossing over leads to exchange of genetic material

What about sexchromosomes?

Human males are the heterogametic sex with different sex chromosomes, (XY) Human females are the homogametic sex (XX) In other species sex can be determined in many ways. For example, in birds and snakes
Males are homogametic Females are heterogametic ZZ ZW

During mitotic division there is no special challenge for the distribution of sexchromosomes.

X and Y Chromosomes are very different

X chromosome
contains more than 1,500 genes larger than the Y chromosome acts as a homolog to Y chromosome in males

Y chromosome
contains 231 genes many DNA segments are palindromes and may destabilize Figure 6.2 DNA

Genes on the Y Chromosome

Genes shared with X chromosome define the pseudoautosomal regions (PAR1 and PAR2) - only here can crossing over occur. Male specific (MSY) region includes SRY gene SRY gene is important in determining sex Figure 6.3

Metaphase of meiosis I
Bivalents attach via their kinetochores to spindle microtubules and migrate to spindle equator. Bivalents are randomly orientated at this point. Homologous chromosomes are held together by chiasmata.
Cell Biology 2007/2008

Anaphase of meiosis I
The members of each pair of homologous chromosomes separate from each other. Start migrating towards opposite spindle poles

Telophase I and cytokinesis

A haploid set of chromosomes arrive at each spindle pole. Nuclear envelopes form around the chromosomes, cytokinesis occurs.

Meiosis II
Has the function to divide the sister chromatides formed in the initial round of DNA-replication into two newly forming cells. The phases of meiosis II (pro-II, meta-II, ana-II and telophase II) resemble very much those of a normal mitotic division.

Prophase II

Metaphase II

Anaphase II Telophase II

Meiosis II
Cell Biology 2007/2008

http://highered.mcgrawhill.com/sites/0072437316/student_view0/c hapter11/animations.html#

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Nondisjunction
Nondisjunction might occur in anaphase I when the two members of homologous chromosomes do not separate. Gametes resulting from such nondisjunctions have an incorrect number of chromosomes. Nondisjunction of chromosome 21 can lead to trisomy 21. The risk of trisomy 21 increases with age of parents.

Comparing mitosis and meiosis

Cell Biology 2007/2008

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Comparing mitosis and meiosis

Cell Biology 2007/2008 Cell Biology 2007/2008

Comparing mitosis and meiosis

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Meiosis contributes to genetic variability

By random assortment of homologues chromosomes
Occurring in metaphase I of meiosis

By crossing over
Occurring in prophase I of meiosis Exchange of DNA between homologous chromosomes

By fertilization
Occurring when two gametes meet and fuse

Random assortment

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Crossing over and random assortment

Gametogenesis is more than just meiosis

In males, in meiosis a diploid spermatocyte is converted into four haploid spermatids. After meiosis, these differentiate into sperm cells:
They loose most of their cytoplasma Build out a flagellum-like tail etc.

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Sperm are continously produced in most mammals

Primordial germ cells migrate into the testis early in embryogenesis Immature germ cells, (spermatogonia) proliferate continuously by mitosis. Some of the daughter cells stop proliferating and differentiate into primary spermatocytes.

Sperm are continously produced in most mammals

Primary spermatocytes can enter meiosis to produce four spermatids, each with a haploid number of single chromosomes. These haploid spermatids then undergo morphological differentiation into sperm. Differentiation occurs AFTER Meiosis.

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Oogenesis occurs in stages

Primordial germ cells migrate to the forming gonad: oogonia, which proliferate by mitosis before differentiating into primary oocytes. usually before birth the first meiotic division begins and remains arrested in prophase I

Oogenesis occurs in stages

Oocyte maturation occurs following hormone stimulation. Cell resumes division I of meiosis (and completely or partly does division II) Divisions are asymmetrical: one mature egg is formed and the smaller accompanying polar bodies oocytes maintain their large size; polar bodies degenerate.

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Gametogenesis is more than just meiosis

The human egg is 0.1 mm in diameter

Eggs are typically specialized spherical or ovoid cell The egg cytoplasm contains nutritional reserves: yolk, The egg coat is a ECM consisting largely of glycoprotein molecules, some secreted by the egg and others deposited on it by surrounding cells. In many species, the major coat is a layer immediately surrounding the egg plasma membrane; in mammalian eggs it is called the zona pellucida

Zona pellucida
protects egg from mechanical damage species-specific barrier to sperm, admitting only those of the same or closely related species.

(A) Scanning electron micrograph of a hamster egg, showing the zona pellucida. (B) A scanning electron micrograph of a similar egg in which the zona (to which many sperm are attached) has been peeled back to reveal the underlying plasma membrane, which contains numerous microvilli. The zona is made entirely by the developing oocyte. (From D.M. Phillips, J. Ultrastruct. Res. 72:112, 1980.)

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Fertilization
The 3. step in increasing genetic variability is fertilization where two haploid germ cells (of opposite type) randomly fuse with each other. For this fusion to occur germ cells must not only be haploid but also morphologically altered.

Fertilization
Once fertilized, the egg is called a zygote Fertilization is not complete, however, until the two haploid nuclei (pronuclei) have come together to form a single diploid nucleus.

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Fertilization
Entry of further sperm cell is prevented by rearrangement of ZP as soon as the first sperm nucleus has entered. Spermal flagellum and mitochondria do not enter the ovum.

The sperm provides the centriole to the zygote

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Stages of Development

Early Development: Ovulation to Implantation

Figure 3.14

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