Crossing over is a precise process that includes stages like synapsis, tetrad formation, cross over and terminalization.
(i) Synapsis: Intimate pairing between two homologous chromosomes is initiated during zygotene stage of prophase I of meiosis I. Homologous chromosomes are aligned side by side resulting in a pair of homologous chromosomes called bivalents. This pairing phenomenon is called synapsis or syndesis.
It is of three types:
1. Procentric synapsis: Pairing starts from middle of the chromosome.
2. Proterminal synapsis: Pairing starts from the telomeres.
3. Random synapsis: Pairing may start from anywhere.
(ii) Tetrad Formation: Each homologous chromosome of a bivalent begin to form two identical sister chromatids, which remain held together by a centromere. At this stage each bivalent has four chromatids. This stage is called tetrad stage.
(iii) Cross Over: After tetrad formation, crossing over occurs in pachytene stage. The non-sister chromatids of homologous pair make a contact at one or more points. These points of contact between non¬sister chromatids of homologous chromosomes are called Chiasmata (singularChiasma).
At chiasma, cross-shaped or X-shaped structures are formed, where breaking and rejoining of two chromatids occur. This results in reciprocal exchange of equal and corresponding segments between them. A recent study reveals that synapsis and chiasma formation are facilitated by a highly organised structure of filaments called Synaptonemal Complex (SC). This synaptonemal complex formation is absent in some species of male Drosophila, hence crossing over does not takes place.
(iv) Terminalisation: After crossing over, chiasma starts to move towards the terminal end of chromatids. This is known as terminalisation. As a result, complete separation of homologous chromosomes occurs.