Cytology Dr.sarab Meiosis: Most reproduction occurs through sexual reproduction. Sexual reproduction is the recombination or mixture of genes. This results in the offspring having a combination of DNA from both parents. This will help add to: (l) the variation within a population or a species. (2)this also creates unique individuals, which are not identical to the parents. For this to be possible, there has to be a special type of cell division. If there was a union of two somatic cells, then the offspring would have twice the number of chromosomes as the parents, and that would continue to increase. This is not the case however, so there is a cell division that reduced the number of chromosomes in half. This is important because when we add the two parent cells together (fertilization) the offspring will end up with the same number of chromosomes as the parents, but with a combination of features. Each species has a different number of chromosomes. For example, humans have 46 chromosomes (23pairs) in every body cell. All cells contain two complete sets of chromosomes( in humans each set has 23 making a total of 46) called the diploid (2n) number of chromosomes. The exception to this is with sex cells, such as egg and sperm. Since these will later combine they each only have one set of chromosomes , called haploid (n) cells, and in humans this means they have 23 chromosomes total. Haploid cells, or reproductive cells, are called gametes or sex cells. The process which produces diploid cells was called mitosis. This occurs in all somatic, or body cells. To obtain the reduced number of chromosomes for haploid cells, they undergo what is called meiosis. Meiosis is the formation of gametes, performed by reproductive cells only. This will result in a reduction of the chromosomes number, forming haploid cells meiosis produced 4 daughter cells, each with half the number of chromosomes, that are not identical to each other. This is again important because it, keeps the chromosome number constant over generations, in the adult organism. When the egg (n) and the sperm (n) combine (n + n) in fertilization, the offspring is a diploid cell (2n).Different organisms have different numbers of chromosomes. 1 when chromosomes pair, in preparation for meiosis, they arrange with their homologous pair, otherwise known as homologs. This means that like chromosomes pair with each other. Remember humans have 46 chromosomes, but23 pairs, each of the chromosomes in the pair are exactly alike in size, gene location and location of the centromere. There is one exception, the sex chromosomes. Females have a pair called XX, both of them are alike' but males are XY. The homologous Pairs of chromosomes ate separated and end up in 4 daughter cells. This happens in two main states, meiosis I and meiosis II. The Pairs separate during meiosis I, and meiosis II is when the sister chromatids separate, much like in mitosis. Meiosis I is a 4 step process. The start of this is much like mitosis, the cells have gone through the G1, S and G2 phases, thus the DNA has been replicated and are in the form of sister chromatids, connected at the centromere. The main difference here is that the homologous pairs of chromosomes pair up and then proceed as follows . Prophase I is traditionally divided into five sequential stages: leptotene, zygotene, pachytene, diplotene, and diakinesis. Leptotene. Chromosomes condense tightly. Zygotene. Pairing of homologous chromosomes. This is accomplished with the help of the synaptonemal complex'' the synaptonemal complex pulls together homologous regions of the DNA. While not identical, homologous regions have a very high similarity. (Synapsis The sites the homologues attach to are adjacent, so that the members of each homologous pair of chromosomes are brought close together. They then line up side by side, apparently guided by heterochromatin sequences, in the process called synapsis . ) 2 Pachytene. Pachytene begins when synapsis is complete (just after the synaptonemal complex forms; and lasts for days. This complex, about 100 nm across, holds the two replicated chromosomes in precise register, keeping each gene directly across from its partner on the homologous chromosome, like the teeth of a zipper. Within the synaptonemal complex, the DNA duplexes unwind at certain sites, and single strands of DNA form base-pairs with complementary strands on the other homologue. The synaptonemal complex thus provides the structural framework that enables crossing over between the homologous chromosomes. Crossing Over: Within the synaptonemal complex, recombination is thought to be carried out during pachytene by very large protein assemblies called recombination nodules. A nodule’s diameter is about 90 nm, spanning the central element of the synaptonemal complex. Spaced along the synaptonemal complex, these recombination nodules act as large multienzyme “recombination machines,” each nodule bringing about a recombination event. The details of the crossing over process are not well understood, but involve a complex series of events in which DNA segments are exchanged between nonsister or sister chromatids. In humans, an average of two or three such crossover events occur per chromosome pair. Chiasma Evidence of crossing over can often be seen under the light microscope as an X-shaped structure known as a chiasma (Greek, “cross”; plural, chiasmata). The presence of a chiasma indicates that two chromatids (one from each homologue) have exchanged parts . Like small rings moving down two strands of rope, the chiasmata move to the end of the chromosome arm as the homologous chromosomes separate. Diplotene. At the beginning of diplotene, the protein lattice of the synaptonemal complex disassembles. Diplotene is a period of intense 3 cell growth. During this period the chromosomes decondense and become very active in transcription. Diakinesis. At the beginning of diakinesis, the transition into metaphase, transcription ceases and the chromosomes recondense. Metaphase I Each joined pair of homologues then lines up on the metaphase plate. The orientation of each pair on the spindle axis is random: either the maternal or the paternal homologue may orient toward a given pole. Anaphase I(The first division is called the Reductional Division, since it is at this time that the chromosome count is reduced to one half In anaphase I, the microtubules of the spindle fibers begin to shorten. As they shorten, they break the chiasmata and pull the centromeres toward the poles, dragging the chromosomes along with them. Because the microtubules are attached to kinetochores on only one side of each centromere, the individual centromeres are not pulled apart to form two daughter centromeres, as they are in mitosis. Instead, the entire centromere moves to one pole, taking both sister chromatids with it. When the spindle fibers have fully contracted, each pole has a complete haploid set of chromosomes consisting of one member of each homologous pair meiosis 4 I results in the independent assortment of maternal and paternal chromosomes into the gametes. Telophase I By the beginning of telophase I, the chromosomes have segregated into two clusters, one at each pole of the cell. Now the nuclear membrane reforms around each daughter nucleus. Because each chromosome within a daughter nucleus replicated before meiosis I began, each now contains two sister chromatids attached by a common centromere. Importantly, the sister chromatids are no longer identical, because of the crossing over that occurred in prophase I . Cytokinesis may or may not occur after telophase I. The second meiotic division, meiosis II, occurs after an interval of variable length. The Second Meiotic Division After a typically brief interphase, in which no DNA synthesis occurs, the second meiotic division begins. Meiosis II resembles a normal mitotic division. Prophase II, metaphase II, anaphase II, and telophase II follow in quick succession. Prophase II. At the two poles of the cell the clusters of chromosomes enter a brief prophase II, each nuclear envelope breaking down as a new spindle forms. Metaphase II. In metaphase II, spindle fibers bind to both sides of the centromeres. Anaphase II. The spindle fibers contract, splitting the centromeres and moving the sister chromatids to opposite poles. Telophase II. Finally, the nuclear envelope re-forms around the four sets of daughter chromosomes. The final result of this division is four cells containing haploid sets of chromosomes. No two are alike, because of the crossing over in prophase I. Nuclear envelopes then form around each haploid set of chromosomes. The cells that contain these haploid nuclei may develop directly into gametes, as they do in animals. 5 In female , during the meiotic cell division of female germ cells, unequal division of the cytoplasm occurs which leads to the formation of :1- One primary Oocyte. 2- Three small polar bodies which will degenerate. The primary Oocyte are formed early during the embryonic development of female fetus. They remain in the prophase stage of the first meiotic cell division for years until puberty is reached, then the first meiotic cell division is completed. 6 7