Meiosis and Sex Cells: Sexual Reproduction: In higher organisms, plants and animals, each individual is diploid. A diploid organism has a complete set of chromosomes in every cell and is 2n (diploid means 'double set'). The organism gets one set from the mother and the other set from the father. The two partners produce gametes, which are joined to produce an offspring. However, two problems must be solved with sexual reproduction. 1) If fertilization occurs and the gametes join, why isn't the genetic material doubled? 2) How is it possible for each parent to give half of the genetic material? The answer is meiosis, a process in which a diploid or double set of chromosomes is reduced to a haploid (n), or a single, set of chromosomes. It is a process that guarantees that the number of chromosomes remains stable from generation to generation. In humans the diploid number = 46 (2n = 46), the haploid number = 23 (n = 23); in fruit flies: 2n = 8, n = 4. Multicellular organisms usually have two types of cells: 1) somatic and 2) gametes. Somatic cells are any type of cell that is not a sperm or egg. These cells are diploid (2n). Gametes are cells that are sperm or eggs. Gametes are haploid (n). Homologous Chromosomes: In humans there are 46 chromosomes. Each chromosome consists of a double helix molecule of DNA. The DNA is folded with proteins to make up a chromosome. One chromosome represents hundreds or thousands of genes, and each gene is a specific region of the DNA molecule. A gene's specific location on the chromosome is called the its LOCUS. The 46 chromosomes are actually 23 pairs of chromosomes. The members of each pair are called homologous chromosomes (homologues). The two homologues are functionally equivalent and contain the same kinds of genes arranged in the same order. One set of chromosomes that does not occur as homologue occurs in males; the X chromosome and the Y chromosome are not homologues but pair up in meiosis. In females, there are two X chromosomes that are homologues. These chromosomes are the sex chromosomes and the other 22 pairs of chromosomes are called AUTOSOMES. During meiosis, three things happen to the homologues: 1) The homologues pair up. 2) The homologues exchange genetic information. This is called crossing over. 3) The newly scrambled chromosomes separate and go into different daughter cells in such a way that each daughter cell contains only one of each pair of homologues. These cells are called gametes or sex cells. Meiosis and life cycles: Meiosis occurs at different times during the life cycle of different organisms. In protists and fungi, meiosis occurs right after the fusion of the two mating cells. The mating cells are usually haploid and the fusion produces a diploid cell. Immediate meiosis restores the haploid lifestyle. In all plants, a multicellular haploid phase alternates with a multicellular diploid phase. The typical fern is diploid and is called a SPOROPHYTE. The diploid sporophyte produces haploid spores 1 through meiosis. These spores will grow into small haploid plants called a GAMETOPHYTE. These produce male and female sex cells (gametes) via mitosis. The gametes will join to form a diploid cell that will grow into the fern that you see. This alternation between diploid and haploid is called ALTERNATION OF GENERATIONS. Animals, including humans, are diploid organisms that produce haploid gametes. Two haploid gametes will join to produce a diploid zygote. Most of the lifecycle in animals is in the diploid state. Mitosis vs. Meiosis: 1) Mitosis: occurs in haploid, diploid and polyploid cells. Meiosis: occurs only in diploid and polyploid cells. 2) Meiosis: The nucleus divides twice producing four nuclei. The chromosomes replicate only once, so each nucleus contains half of the number of chromosomes. 3) Each haploid chromosome is a new combination of old chromosomes because of crossing over. Meiosis: There are two stages of Meiosis: Meiosis I and Meiosis II. Meiosis I: Replication of chromosomes, crossing over of the chromosomes and reduction in the chromosome number from diploid to haploid. Meiosis I is often called the reduction division. Premeiotic Interphase: G1, S (replication of the chromosomes), and G2. Meiotic Prophase I: The first stage. This is long and complex compared with mitotic prophase. In it: 1) Nuclear membrane disappears. 2) Spindle fibers form. 3) The chromosomes condense. 4) The homologous chromosomes pair up by touching each other in the appropriate places. First there is a lot of random movement of chromosomes until the homologous chromosomes find each other. It is important, for example, that chromosome #13 finds homologous chromosome #13. When the two homologous touch each other in the same place, a specialized structure called the Synaptonemal Complex holds the homologues together. The meiotic cell of a human now has 23 genetic entities called tetrads, each packet containing four chromatids and two centromeres. This is the point when crossing over occurs. A special enzyme causes the chromatids to unwind, revealing the strands of DNA. A complex series of events happen and the genetic material is exchanged between homologues. Crossing over may occur at the INTRONS. Several thousand base pairs of one strand pairs with the chromatid on another homologues. There are breakages and the chromatids untangle themselves. Meanwhile other enzymes are repairing the breaks in the DNA. This process makes new chromatids and is a source of genetic variation within a population. 2 After crossing over, the homologs begin to pull away from each other, except at the crossing over points called the CHIASMATA (CHIASMA- singular). Metaphase I: In the first metaphase, the tetrads are brought to the metaphase plate. The synaptonemal complex is lined up on the metaphase plate. Anaphase I: There is no separation of the centromeres, but the synaptonemal complex separates. This means that the homologues separate and move to opposite poles. The first meiotic division reduces the chromosome number by half. Telophase I: In this phase, the nucleus reorganizes and the nuclear membrane reforms. The chromosomes decondense. Cytokinesis I: In this phase, the cytoplasmic division occurs. Meiosis II: Division of the chromosomes, analogous to mitosis. Meiotic Interphase: This involves G1 and G2 phases only. There is no S phase in this interphase. This phase may be brief or last a long time. Prophase II: As in mitotic prophase, there are two sister chromatids attached to a centromere. The chromosomes condense, the nucleus disappears, and the spindle apparatus forms. Metaphase II: Centromeres move to the metaphase plate during metaphase II. Anaphase II: During anaphase II, the kinetochores are broken down by enzymes, and sister chromatids separate and move to the opposite poles. Telophase II: During telophase II, the nuclear membrane reforms and chromosomes decondense. Cytokinesis II: the cytoplasm divides. Summary of Meiosis: From one pair of homologs, there are four, unique chromatids from prophase I, if crossing over has occurred. Each unique chromatid ends up in one of the four cells that are the products of meiosis. The amount of genetic material was reduced by one half in Meiosis I and divided in Meiosis II. Each resulting cell (gamete) is haploid. 3 Meiosis in Humans: Meiosis in Males: In the male each of these haploid cells is called a spermatid. This spermatid will undergo cellular differentiation to become gametes (sperm). Meiosis in Females: Meiosis is begun but is only partly completed in human females shortly before birth. All oocytes (oogonia) begin meiosis I and become primary oocytes. They remain frozen (in stasis) in the last stage of meiotic prophase I. In humans, meiotic prophase I can last up to 50 years. In spite of not continuing to metaphase I, the paired meiotic chromosomes are very active making large amounts of ribosomes and mRNA. By the time the primary oocyte is ready to be released. It is a large cell filled with yolk, mRNA, ribosomes etc. The oocyte will not resume meiosis until released from the ovary. After ovulation, the primary oocyte finishes meiosis I and produces two cells, the secondary oocyte and the primary polar body. The primary polar body is basically a chromosomal trashcan. It cannot be fertilized and will die. The secondary oocyte begins meiosis II, but stops after prophase II and waits for a sperm. Meiosis will not be completed unless the secondary oocyte meets a sperm and is fertilized. When this happens, many changes occur in the oocyte including the completion of meiosis II. The cell constituents are not divided evenly and most of the cytoplasm ends up in one cell. Only one cell will develop into the egg (ovum). The normal division occurs, but one of the two daughter cells has most of the cytoplasm. The other daughter cell is very small and becomes the secondary polar body. The secondary polar body will die and acts as a chromosomal trashcan. The other bigger cell is known as the ovum. The ovum is formed after fertilization and will become the baby. The importance of meiosis: 1) Sexual reproduction is a reshuffling of the genes of all the successful individuals of the population. There are virtually infinite possibility combinations of genes. There are three sources of variation due to meiosis. a) Independent Assortment of Chromosomes: In the diploid cell, you start off with 4 chromatids per homologous chromosome. At the end of Meiosis II, each gamete has 1 chromatid. Each chromatid moved independently of one another. Each chromatid is also different. b) Crossing over: This process produces individual chromosomes (chromatids) that are a combination between the two grandparents (the parent’s parents think about it). C) Random Fertilization: a new genetic combination of sperm and egg occurs when the sperm meets the egg. 2) The reduction and division of the chromosomes in the egg and sperm makes fertilization possible and enables the maintenance of a constant chromosome number within a species. Problems with Meiosis: 1) Unequal exchange of chromosomal material during crossing over. One chromosome will end up with more DNA than its homologue. This can delete some genes from 4 chromosomes and give extra genes to the other chromosome. 2) Inversion of the chromosome during crossing over. The piece of chromosomal material will ‘flip’ during the event. 3) Translocation of the piece that’s crossing over. The piece that is cut will float away from the homolog and attach to a non-homologous chromosome. One gamete will have extra chromosomal material and the other gamete will be missing a piece of a chromosome. 4) Nondisjunction: When the chromosomes don’t divide properly during anaphase I or II. The cells will not have the correct number of chromosomes. The spindle fibers might not attach properly to the kinetochore (or synaptonemal complex), the spindle fibers might not contract properly during anaphase, the APC (enzyme that breaks down the kinetochore) might not work properly or the synaptonemal complex might not separate. If this happens in Meiosis I, then all 4 cell products will have an incorrect number of chromosomes. If this happens in Meiosis II, then 2 of the 4 cells will have an incorrect number of chromosomes. If a whole set of chromosomes is affected, this is called polyploidy. 5