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Lecture 9

DEV2011: Lecture 9 summary

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LECTURE 9 Meiosis: Meiosis is a special type of cell division necessary for sexual reproduction in eukaryotes. The cells produced by meiosis are gametes. Whilst the process of meiosis bears a number of similarities with the 'life-cycle' cell division process of mitosis, it differs in two important respects: - The chromosomes in meiosis undergo a recombination which shuffles the genes producing a different genetic combination in each gamete, compared with the co-existence of each of the two separate pairs of each chromosome (one received from each parent) in each cell which results from mitosis. - The outcome of meiosis is potentially four (genetically unique) haploid cells, compared with the two (genetically identical) diploid cells produced from mitosis. Meiosis begins with one diploid cell containing two copies of each chromosome—one from the organism's mother and one from its father. The cell divides twice, potentially producing up to four haploid cells containing one copy of each chromosome. ("Potentially" because in some cases, such as the formation of oocytes in mammals, only one of the possible four haploid cells survives.) In animals the haploid cell resulting from meiosis is a male or female gamete. Each of the resulting chromosomes in the gamete cells is a unique mixture of maternal and paternal DNA, resulting in offspring that are genetically distinct from either parent. Meiosis is a key feature for all sexually reproducing eukaryotes in which homologous chromosome pairing, synapse and recombination occur. Prior to the meiosis process the cell's chromosomes are duplicated by a round of DNA replication, creating from the maternal and paternal versions of each chromosome (homologs) two exact copies, sister chromatids, attached at the centromere region. In the beginning of meiosis the maternal and paternal homologs pair to each other. Then they typically exchange parts by homologous recombination leading to crossovers of DNA between the maternal and paternal versions of the chromosome. Spindle fibers bind to the centromeres of each pair of homologs and arrange the pairs at the spindle equator. Then the fibers pull the recombined homologs to opposite poles of the cell. As the chromosomes move away from the center the cell divides into two daughter cells, each containing a haploid number of chromosomes composed of two chromatids. After the recombined maternal and paternal homologs have separated into the two daughter cells, a second round of cell division occurs. There meiosis ends as the two sister chromatids making up each homolog are separated and move into one of the four resulting gamete cells. Upon fertilization, for example when a sperm enters an egg cell, two gamete cells produced by meiosis fuse. The gamete from the mother and the gamete from the father each contribute one half of the set of chromosomes that make up the new offspring's genome. Meiosis I: Meiosis I separates homologous chromosomes, producing two haploid cells (N chromosomes, 23 in humans), and thus meiosis I is referred to as a reductional division. A regular diploid human cell contains 46 chromosomes and is considered 2N because it contains 23 pairs of homologous chromosomes. However, after meiosis I, although the cell contains 46 chromatids, it is only considered as being N, with 23 chromosomes. This is because later, in Anaphase I, the sister chromatids will remain together as the spindle fibers pull the pair toward the pole of the new cell. In meiosis II, an equational division similar to mitosis will occur whereby the sister chromatids are finally split, creating a total of 4 haploid cells (23 chromosomes, N) - two from each daughter cell from the first division. Prophase 1: It is the longest phase of meiosis. During prophase I, DNA is exchanged between homologous chromosomes in a process called homologous recombination. This often results in chromosomal crossover. The new combinations of DNA created during crossover are a significant source of genetic variation, and may result in beneficial new combinations of alleles. The paired and replicated chromosomes are called bivalents or tetrads, which have two chromosomes and four chromatids, with one chromosome coming from each parent. The process of pairing the homologous chromosomes is called synapsis. At this stage, non-sister chromatids may cross-over at points called chiasmata. Leptotene: The first stage of prophase I is the leptotene stage. In this stage of prophase I, individual chromosomes—each consisting of two sister chromatids—change from the diffuse state they exist in during the cell's period of growth and gene expression, and condense into visible strands within the nucleus. However the two sister chromatids are still so tightly bound that they are indistinguishable from one another. During leptotene, lateral elements of the synaptonemal complex assemble. Leptotene is of very short duration and progressive condensation and coiling of chromosome fibers takes place. Zygotene: The zygotene stage occurs as the chromosomes approximately line up with each other into homologous chromosome pairs. This is called the bouquet stage because of the way the telomeres cluster at one end of the nucleus. At this stage, the synapsis (pairing/coming together) of homologous chromosomes takes place, facilitated by assembly of central element of the synaptonemal complex. Pairing is brought about in a zipper-like fashion and may start at the centromere (procentric), at the chromosome ends (proterminal), or at any other portion (intermediate). Individuals of a pair are equal in length and in position of the centromere. Thus pairing is highly specific and exact. The paired chromosomes are called bivalent or tetrad chromosomes. Pachytene: The pachytene stage is the stage when chromosomal crossover (crossing over) occurs. Nonsister chromatids of homologous chromosomes may exchange segments over regions of homology. Sex chromosomes, however, are not wholly identical, and only exchange information over a small region of homology. At the sites where exchange happens, chiasmata form. The exchange of information between the non-sister chromatids results in a recombination of information; each chromosome has the complete set of information it had before, and there are no gaps formed as a result of the process. Because the chromosomes cannot be distinguished in the synaptonemal complex, the actual act of crossing over is not perceivable through the microscope, and chiasmata are not visible until the next stage. Diplotene: During the diplotene stage, the synaptonemal complex degrades and homologous chromosomes separate from one another a little. The chromosomes themselves uncoil a bit, allowing some transcription of DNA. However, the homologous chromosomes of each bivalent remain tightly bound at chiasmata, the regions where crossing-over occurred. The chiasmata remain on the chromosomes until they are severed in anaphase I. In human fetal oogenesis all developing oocytes develop to this stage and stop before birth. This suspended state is referred to as the dictyotene stage and remains so until puberty. Diakinesis: Chromosomes condense further during the diakinesis stage. This is the first point in meiosis where the four parts of the tetrads are actually visible. Sites of crossing over entangle together, effectively overlapping, making chiasmata clearly visible. Other than this observation, the rest of the stage closely resembles prometaphase of mitosis; the nucleoli disappear, the nuclear membrane disintegrates into vesicles, and the meiotic spindle begins to form. Metaphase I: Homologous pairs move together along the metaphase plate: As kinetochore microtubules from both centrioles attach to their respective kinetochores, the homologous chromosomes align along an equatorial plane that bisects the spindle, due to continuous counterbalancing forces exerted on the bivalents by the microtubules emanating from the two kinetochores of homologous chromosomes. Anaphase I: Kinetochore (bipolar spindles) microtubules shorten, severing the recombination nodules and pulling homologous chromosomes apart. Since each chromosome has only one functional unit of a pair of kinetochores,whole chromosomes are pulled toward opposing poles, forming two haploid sets. Each chromosome still contains a pair of sister chromatids. During this time disjunction occurs, which is one of the processes leading to genetic diversity as each chromosome can end up in either of the daughter cells. Non-kinetochore microtubules lengthen, pushing the centrioles farther apart. The cell elongates in preparation for division down the center. Telophase I: The first meiotic division effectively ends when the chromosomes arrive at the poles. Each daughter cell now has half the number of chromosomes but each chromosome consists of a pair of chromatids. The microtubules that make up the spindle network disappear, and a new nuclear membrane surrounds each haploid set. The chromosomes uncoil back into chromatin. Cytokinesis, the pinching of the cell membrane in animal cells or the formation of the cell wall in plant cells, occurs, completing the creation of two daughter cells. Sister chromatids remain attached during telophase I. Meiosis II: Mechanically, the process is similar to mitosis, though its genetic results are funda
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