Meiosis is preceded by an interphase consisting of G1, S, and G2 phases, which are nearly identical to the phases preceding mitosis. The G1 phase (the “first gap phase”) is focused on cell growth. During the S phase—the second phase of interphase—the cell copies or replicates the DNA of the chromosomes. Finally, in the G2 phase (the “second gap phase”) the cell undergoes the final preparations for meiosis.
During DNA duplication in the S phase, each chromosome is replicated to produce two identical copies—sister chromatids that are held together at the centromere by cohesin proteins, which hold the chromatids together until anaphase II. (Note: these chromosome copies are called sister chromatids regardless of whether they are in a female gamete or a male gamete.)
There are four stages that occur during meiosis I - prophase I, metaphase I, anaphase I, and telophase I.
Early in prophase I, before the chromosomes can be seen clearly with a microscope, the homologous chromosomes are attached at their tips to the nuclear envelope by proteins. As the nuclear envelope begins to break down, the proteins associated with homologous chromosomes bring the pair closer together. Recall that in mitosis, homologous chromosomes do not pair together. The synaptonemal complex, a lattice of proteins between the homologous chromosomes, first forms at specific locations and then spreads outward to cover the entire length of the chromosomes. The tight pairing of the homologous chromosomes is called synapsis. In synapsis, the genes on the chromatids of the homologous chromosomes are aligned precisely with each other. The synaptonemal complex supports the exchange of chromosomal segments between homologous nonsister chromatids—a process called crossing over. Crossing over can be observed visually after the exchange as chiasmata (singular = chiasma) (Figure 11.3).
In humans, even though the X and Y sex chromosomes are not completely homologous (that is, most of their genes differ), there is a small region of homology that allows the X and Y chromosomes to pair up during prophase I. A partial synaptonemal complex develops only between the regions of homology.
Figure 11.3 Early in prophase I, homologous chromosomes come together to form a synapse. The chromosomes are bound tightly together and in perfect alignment by a protein lattice called a synaptonemal complex and by cohesin proteins at the centromere.
Located at intervals along the synaptonemal complex are large protein assemblies called recombination nodules. These assemblies mark the points of later chiasmata and mediate the multistep process of crossover—or genetic recombination—between the nonsister chromatids. Near the recombination nodule, the double-stranded DNA of each chromatid is cleaved, the cut ends are modified, and a new connection is made between the nonsister chromatids. As prophase I progresses, the synaptonemal complex begins to break down and the chromosomes begin to condense. When the synaptonemal complex is gone, the homologous chromosomes remain attached to each other at the centromere and at chiasmata. The chiasmata remain until anaphase I. The number of chiasmata varies according to the species and the length of the chromosome. There must be at least one chiasma per chromosome for proper separation of homologous chromosomes during meiosis I, but there may be as many as 25. Following crossover, the synaptonemal complex breaks down and the cohesin connection between homologous pairs is removed. At the end of prophase I, the pairs are held together only at the chiasmata (Figure 11.4). These pairs are called tetrads because a total of four sister chromatids of each pair of homologous chromosomes are now visible.
The crossover events are the first source of genetic variation in the nuclei produced by meiosis. A single crossover event between homologous nonsister chromatids leads to a reciprocal exchange of equivalent DNA between an egg-derived chromosome and a sperm-derived chromosome. When a recombinant sister chromatid is moved into a gamete cell it will carry a combination of maternal and paternal genes that did not exist before the crossover. Crossover events can occur almost anywhere along the length of the synapsed chromosomes. Different cells undergoing meiosis will therefore produce different recombinant chromatids, with varying combinations of maternal and parental genes. Multiple crossovers in an arm of the chromosome have the same effect, exchanging segments of DNA to produce genetically recombined chromosomes.