Synapsis during Meiosis. The circled area is the part where synapsis occurs, where the two chromatids meet before crossing over
Synapsis (also called syndesis) is the pairing of two homologous chromosomes that occurs during meiosis. It allows matching-up of homologous pairs prior to their segregation, and possible chromosomal crossover between them. Synapsis takes place during prophase I of meiosis.
When homologous chromosomes synapse, their ends are first attached to
the nuclear envelope. These end-membrane complexes then migrate,
assisted by the extranuclear cytoskeleton,
until matching ends have been paired. Then the intervening regions of
the chromosome are brought together, and may be connected by a
protein-RNA complex called the synaptonemal complex. Autosomes
undergo synapsis during meiosis, and are held together by a protein
complex along the whole length of the chromosomes called the
synaptonemal complex. Sex chromosomes
also undergo synapsis; however, the synaptonemal protein complex that
holds the homologous chromosomes together is only present at one end of
each sex chromosome.
This is not to be confused with mitosis. Mitosis also has prophase, but does not ordinarily do pairing of two homologous chromosomes.
When the non-sister chromatids intertwine, segments of chromatids
with similar sequence may break apart and be exchanged in a process
known as genetic recombination or "crossing-over". This exchange produces a chiasma,
a region that is shaped like an X, where the two chromosomes are
physically joined. At least one chiasma per chromosome often appears to
be necessary to stabilise bivalents along the metaphase
plate during separation. The crossover of genetic material also
provides a possible defence against 'chromosome killer' mechanisms, by
removing the distinction between 'self' and 'non-self' through which
such a mechanism could operate. A further consequence of recombinant
synapsis is to increase genetic variability
within the offspring. Repeated recombination also has the general
effect of allowing genes to move independently of each other through the
generations, allowing for the independent concentration of beneficial
genes and the purging of the detrimental.
Following synapsis, a type of recombination referred to as
synthesis dependent strand annealing (SDSA) occurs frequently. SDSA
recombination involves information exchange between paired non-sister
homologous chromatids, but not physical exchange. SDSA recombination
does not cause crossing-over. Both the non-crossover and crossover
types of recombination function as processes for repairing DNA damage,
particularly double-strand breaks.
The central function of synapsis is therefore the identification
of homologues by pairing, an essential step for a successful meiosis.
The processes of DNA repair and chiasma formation that take place
following synapsis have consequences at many levels, from cellular
survival through to impacts upon evolution itself.
Chromosome silencing
In mammals, surveillance mechanisms remove meiotic cells in which synapsis is defective. One such surveillance mechanism is meiotic silencing that involves the transcriptional silencing of genes on asynapsed chromosomes. Any chromosome region, either in males or females, that is asynapsed is subject to meiotic silencing. ATR, BRCA1 and gammaH2AX localize to unsynapsed chromosomes at the pachytene stage of meiosis in human oocytes and this may lead to chromosome silencing. The DNA damage response protein TOPBP1 has also been identified as a crucial factor in meiotic sex chromosome silencing. DNA double-strand breaks appear to be initiation sites for meiotic silencing.
Recombination
In female Drosophila melanogaster fruit flies, meiotic chromosome synapsis occurs in the absence of recombination. Thus synapsis in Drosophila
is independent of meiotic recombination, consistent with the view that
synapsis is a precondition required for the initiation of meiotic
recombination. Meiotic recombination is also unnecessary for homologous chromosome synapsis in the nematode Caenorhabditis elegans.
