Crossing Over

S J BORKAR
Introduction
• The phenomenon of the exchange of genetic material between two non-sister chromatids of a
homologous pair of chromosome in a tetrad stage (4- chromatid stage) is called crossing over.
• A given crossing over always involves non-sister chromatids.
• A legitimate crossing over involves homologous chromosomes, whereas a crossing over involving
non-homologous chromosome is called illegitimate crossing over.
• Morgan pointed out that the phenomenon of complete linkage occurs rarely because sometimes
the linked genes show the tendency to separate during meiosis and new combinations are formed.
• Crossing over generally leads to exchange of equal segments or genes and recombination is always
reciprocal. However, unequal crossing over has also been reported.
• The value of crossover or recombinants may vary from 0-50%.
• The frequency of recombinants can be worked out from the test cross progeny. It is expressed as
the percentage ratio of recombinants to the total population (recombinants + parental types).
• Thus
Mechanism of Crossing Over:
• During the zygotene stage of the first prophase of meiosis, the homologous maternal and
paternal chromosomes start pairing and lie closely side by side. This phenomenon is
called synapsis.
• This pairing of homologous chromosomes is brought about by the mutual attraction
between the allelic genes. The paired chromosomes are known as bivalent.
• A recent study reveals that synapsis and chiasma formation is facilitated by a highly
organized structure of filaments called synaptonemal complex.
• Synapsis is followed by the duplication of chromosomes which change the bivalent nature
of chromosome pair into tetravalent.
• During this each of the homologous chromosomes in a bivalent split longitudinally into
two sister chromatids attached to the undivided centromere. Thus, four chromatids are
formed which remain side by side as two pairs.
• Later, in pachytene stage crossing over takes place during which the non-sister
chromatids of homologous pair twist over each other, the point of contact of cross over
chromatids being called as chiasma.
• In crossing over two or three chromatids are involved
and accordingly two or more chiasmata are formed. At
each chiasma the chromatid breaks and the broken
segment rejoin a new chromatid.
• Thus exchange of parts of chromatids brings about
alteration of original sequence of genes in the
chromosome.
• After crossing over is completed, the non-sister
chromatids repel each other due to lack of attraction
between them.
• The repulsion or separation of chromatids starts from
the centromere towards the end just like a zipper and
this separation process is named as terminalization.
The process of terminalization continues through
diplotene, diakinesis and ends in metaphase I.
• At the end of terminalization the twisting chromatids
separate so that the homologous chromosomes are
separated completely and move to opposite poles in
Anaphase I.
• The crossing over thus brings about alteration of the
linear sequence of gene in chromosomes that produce
gametes and thus add new combination of character in
progeny.
 Breakage and Reunion Theory:
• A more probable and easy hypothesis of mechanism of crossing over has been advanced by C. D.
Darlington in 1935. This is called “breakage and reunion theory.” Darlington postulated that the
homologous chromosomes are intertwined during the four stranded stage of first meiosis. The
twisting exerts strain on the chromatids.
• If the stresses are great enough one or more chromatids of the homologues will be broken at one or
more points. If more than one chromatid break, there is possibility that the broken chromatids of
one chromosome will unite with broken ends of different chromatid forming a chiasma.
• If the union takes place between sisters chromatid parts (i.e., paternal to paternal and maternal to
maternal) no genetic consequence is anticipated. However, if the break and reunion occur between
non- sister chromatids (i.e., paternal to maternal or vice versa) recombinants would result.
• Previously it was thought that the mechanical stress produced the breaks before the
recombination but now scientists believe that some enzymes are involved in the processes
of breakage and reunion. The idea that some enzymatic processes bring about breakage
and rejoining of DNA strands was first put-forth by Howard Flanders and Boyce in 1964.
• While considering the reunion of broken DNA strands, regardless of breaks occurred, one
must hypothesize an enzymatic process in the formation of covalent bonds necessary to
repair the sugar phosphate backbone of the DNA molecule.
• Stem and Hotta (1969) are of the opinion that two nuclear enzymes namely endonuclease
and ligase bring about crossing over. The enzyme endonuclease helps in the breaking
of the chromatids while the ligase helps in the reunion of the chromatid segments.
• They have also reported the synthesis of a small amount of DNA during the period of
crossing over which is used in the repairing of the broken chromatids.
• Crossing over leads to the re-combination of genes or genetic material or chromatids part.
Recombination of gene leads to variation.
Significance
• Crossing over provides direct proof for the linear arrangement of genes.
• Through crossing over segments of homologous chromosomes are interchanged and
hence provide origin of new characters and genetic variations.
• Crossing over has led to the construction of linkage map or genetic maps of chromosomes.
• Linkage group and linear order of the genes help to reveal the mechanism and nature of
the genes.
• Crossing over plays a very important role in the field of breeding to improve the varieties
of plants and animals.