Most plant and animal cells are diploid. The term diploid is derived from the Greek diplos, meaning “double” or “two”; the term implies that the cells of plants and animals have two sets of chromosomes. In human cells, for example, 46 chromosomes are organized in 23 pairs. Hence, human cells are diploid in that they have two sets of 23 chromosomes per set.
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During sexual reproduction, the sex cells of parent organisms unite with one another and form a fertilized egg cell. In this situation, each sex cell is a gamete. The gametes of human cells are haploid, from the Greek haplos, meaning “single.” This term implies that each gamete contains a single set of chromosomes—23 chromosomes in humans. When the human gametes unite with one another, the original diploid condition of 46 chromosomes is reestablished. Mitosis then brings about the development of the diploid cell into an organism.
The process by which the chromosome number is halved during gamete formation is meiosis. In meiosis, a cell containing the diploid number of chromosomes is converted into four cells, each having the haploid number of chromosomes. In human cells, for instance, a reproductive cell containing 46 chromosomes yields four cells, each with 23 chromosomes.
Meiosis occurs by a series of steps that resemble the steps of mitosis. Two major phases of meiosis occur: meiosis I and meiosis II. During meiosis I, a single cell divides into two. During meiosis II, those two cells each divide again. The same demarcating phases of mitosis take place in meiosis I and meiosis II.
As shown in Figure 1 , first, the chromosomes of a cell duplicate and pass into two cells. The chromosomes of the two cells then separate and pass into four daughter cells. The parent cell has two sets of chromosomes and is diploid, while the daughter cells have a single set of chromosomes each and are haploid. Synapsis and crossing over occur in the Prophase I stage.
The members of each chromosome pair within a cell are called homologous chromosomes. Homologous chromosomes are similar but not identical. They may carry different versions of the same genetic information. For instance, one homologous chromosome may carry the information for blond hair while the other homologous chromosome may carry the information for black hair.
As a cell prepares to enter meiosis, each of its chromosomes has duplicated, as in mitosis. Each chromosome thus consists of two chromatids.
At the beginning of meiosis 1, a human cell contains 46 chromosomes, or 92 chromatids (the same number as during mitosis). Meiosis I proceeds through the following phases:
· Prophase I: Prophase I is similar in some ways to prophase in mitosis. The chromatids shorten and thicken and become visible under a microscope. An important difference, however, is that a process called synapsis occurs. A second process called crossing over also takes place during prophase 1.
During prophase 1, the two homologous chromosomes come near each other. Because each homologous chromosome consists of two chromatids, there are actually four chromatids aligned next to one another. This combination of four chromatids is called a tetrad, and the coming together is the process called synapsis.
After synapsis has taken place, the process of crossing over occurs. In this process, segments of DNA from one chromatid in the tetrad pass to another chromatid in the tetrad. These exchanges of chromosomal segments occur in a complex and poorly understood manner. They result in a genetically new chromatid. Crossing over is an important driving force of evolution. After crossing over has taken place, the four chromatids of the tetrad are genetically different from the original four chromatids.
· Metaphase I: In metaphase I of meiosis, the tetrads align on the equatorial plate (as in mitosis). The centromeres attach to spindle fibers, which extend from the poles of the cell. One centromere attaches per spindle fiber.
· Anaphase I: In anaphase 1, the homologous chromosomes separate. One homologous chromosome (consisting of two chromatids) moves to one side of the cell, while the other homologous chromosome (consisting of two chromatids) moves to the other side of the cell. The result is that 23 chromosomes (each consisting of two chromatids) move to one pole, and 23 chromosomes (each consisting of two chromatids) move to the other pole. Essentially, the chromosome number of the cell is halved. For this reason the process is a reduction-division.
· Telophase I: In telophase I of meiosis, the nucleus reorganizes, the chromosomes become chromatin, and a cytoplasmic division into two cells takes place. This process occurs differently in plant and animal cells, just as in mitosis. Each daughter cell (with 23 chromosomes each consisting of two chromatids) then enters interphase, during which there is no duplication of the DNA. The interphase period may be brief or very long, depending on the species of organism.
Meiosis II is the second major subdivision of meiosis. It occurs in essentially the same way as mitosis. In meiosis II, a cell containing 46 chromatids undergoes division into two cells, each with 23 chromosomes. Meiosis II proceeds through the following phases:
· Prophase II: Prophase II is similar to the prophase of mitosis. The chromatin material condenses, and each chromosome contains two chromatids attached by the centromere. The 23 chromatid pairs, a total of 46 chromatids, then move to the equatorial plate.
· Metaphase II: In metaphase II of meiosis, the 23 chromatid pairs gather at the center of the cell prior to separation. This process is identical to metaphase in mitosis.
· Anaphase II: During anaphase II of meiosis, the centromeres divide, and the 46 chromatids become known as 46 chromosomes. Then the 46 chromosomes separate from one another. Spindle fibers move one chromosome from each pair to one pole of the cell and the other member of the pair to the other pole. In all, 23 chromosomes move to each pole. The forces and attachments that operate in mitosis also operate in anaphase 11.
· Telophase II: During telophase II, the chromosomes gather at the poles of the cells and become indistinct. Again, they form a mass of chromatin. The nuclear envelope develops, the nucleoli reappear, and the cells undergo cytokinesis as in mitosis.
During meiosis II, each cell containing 46 chromatids yields two cells, each with 23 chromosomes. Originally, there were two cells that underwent meiosis II; therefore, the result of meiosis II is four cells, each with 23 chromosomes. Each of the four cells is haploid; that is, each cell contains a single set of chromosomes.
The 23 chromosomes in the four cells from meiosis are not identical because crossing over has taken place in prophase 1. The crossing over yields variation so that each of the four resulting cells from meiosis differs from the other three. Thus, meiosis provides a mechanism for producing variations in the chromosomes. Also, it accounts for the formation of four haploid cells from a single diploid cell.
Meiosis in Humans
In humans, meiosis is the process by which sperm cells and egg cells are produced. In the male, meiosis takes place after puberty. Diploid cells within the testes undergo meiosis to produce haploid sperm cells with 23 chromosomes. A single diploid cell yields four haploid sperm cells through meiosis.
In females, meiosis begins during the fetal stage when a series of diploid cells enter meiosis 1. At the conclusion of meiosis 1, the process comes to a halt, and the cells gather in the ovaries. At puberty, meiosis resumes. One cell at the end of meiosis I enters meiosis II each month. The result of meiosis II is a single egg cell per cycle (the other meiotic cells disintegrate). Each egg cell contains 23 chromosomes and is haploid.
The union of the egg cell and the sperm cell leads to the formation of a fertilized egg cell with 46 chromosomes, or 23 pairs. Fertilization restores the diploid number of chromosomes. The fertilized egg cell, a diploid, is a zygote. Further divisions of the zygote by mitosis eventually yield a complete human being.
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