Crossing Over and Recombination in Meiosis

Crossing over or chromosomal cross over is a process by which genetic material (DNA) or
chromosomal segments exchange between non-sister chromatids of homologous chromosomes. The
chromatids in which crossing over has occurred are called crossovers or recombinants and the chromatids
that remain intact are called non-cross or parental chromatids. As a result of crossing over, new or non-
parental combinations of alleles among offspring of sexual reproducers are formed.

Recombination on the other hand refers to the process of recombining genes in order to produce new
gene variations that is different from the parent chromosomes. It happens as a result of crossing over of
homologous chromosomes. Crossing over lead to recombination.

Crossing over and recombination are two closely related events that occur during synapsis.

Morgan and Crossing Over

Crossing over was first described by Thomas Hunt Morgan in 1911 in his investigation of numerous
Drosophila mutations located on the X chromosomes. When Morgan made crosses involving other X-linked
genes he was faced with two questions: 1) What was the source of gene separation? And 2) Why did the
frequency of the apparent separation vary depending on the genes being studied. He based his proposed
answer on cytological observations made by Frans Janssens who described the phenomenon and termed the
process as “chrismation” in 1909. The term chiasmata (singular: chiasma) represent points of genetic
exchange or chromosomal crossover. Morgan proposed the term crossing over to describe the physical
exchange leading to recombination.

Substages of Prophase I

Prophase I is by far the most complicated phase of Meiosis. Since crossing over occurs in this stage it
is very important to get understand further the four substages, namely:

1.During leptotene substage (lepto- thin and -tene for ribbon or band) the chromosomes appear as threadlike
structures.

2.In zygotene substage or zygonema(zygo-union, fusing, or yoking), homologous chromosomes pair up to

form bivalent.

3.In pachytene substage or pachynema

(pachy-thick), each homologue chromosomes split up into two chromatids. It is in this stage where crossing
over occurs.

4. In diplotene substage, the chromatids start to divide and separate up however,

they cannot separate completely because they are connected by two strands of DNA at each of the points
where exchanges took place.

5. In the final substage in prophase I known as diakinesis, nucleolus disappears, terminalization reaches
completion, the chromosomes coil tightly, and so become shorter and thicker. The nuclear envelope also begins to
disappear. The centrosomes reach the poles. It is also the stage where crossing over has finally occurred.

Mechanisms of Crossing Over

The process of crossing over takes place in the following steps:

1). Synapsis

Synapsis or syndesis is the process of pairing up two homologous chromosomes during meiosis. It
allows the matching up of homologous chromosomes prior to segregation.

Because the chromosome, not the gene, is the unit of transmission during meiosis, linked genes are
not free to undergo independent assortment. Instead, the alleles at the loci of one chromosome should, in
theory, be transmitted as a unit during gamete formation. Synapsis takes place during prophase I of Meiotic
division. When two homologous paternal and maternal chromosomes synapse or come together, they coil
around each other, until the ends of the chromosomes paired up forming a bivalent. The intervening ends of
the chromosomes are held together by an RNA protein known as synaptonemal complex. All autosomes
undergo synapsis during meiosis but sex chromosomes don’t undergo synapsis.

2. Duplication of chromosomes

After synapsis, duplication of chromosomes follows. In this process, the bivalent chromosomes laying
side by side splits longitudinally forming two sister chromatids. Now, the bivalent chromosomes from
synapsis becomes four chromatids known as tetrad. This process occurs during the diplotene stage of meiotic
cell division.
3. Crossing Over

The formed tetrad arranged in bivalent, side to side well attached and coiled from one another, start to
uncoil. The paternal and maternal chromatids remained attached at one or more points allowing exchange of
genetic materials per coiling. The point of attachment of coiling is called chiasma, and it is the point where
crossing over takes place. The process of crossing over takes place during pachytene stage of meiosis.

4. Terminalization

After crossing over, the non- sister chromatids slowly shift away from each other and separate from
the centromere towards the terminal ends. The continuous repulsion between the chaisma causes the
chromosomes to be free. This process of chaisma movement towards the end of the chromosome is called
terminalization. Terminalization of chiasma eventually takes place during diplotene, after crossing over at
pachytene, and the completion of terminalization occurs in diakinesis.

Types of Crossing Over

Depending upon the number of chiasma involved, crossing over can be classified into three types and
describe as follows:

1. Single crossing over

The formation of single chiasma between non sister chromatids of homologous chromosomes is called single
crossing over.

2. Double crossing over

It is the formation of two points of contact or chiasma between non sister chromatids of homologous
chromosomes.

3. Multiple crossing over

It involves the formation of more than two chiasma between non sister chromatids of homologous
chromosomes.

Factors Affecting Crossing Over:

The investigation on crossing over and recombination using Drosophila showed that the following
factors can affect the frequency of crossing over and recombination.

a. Distance
The greater the distance between genes, the higher is the chance of crossing over.
b. Age
Generally crossing over decreases as female Drosophila aged.
c. Temperature
The rate of crossing over increases as temperature increases by 22°C.
d. Sex
There is lack of crossing over in male Drosophila and female silk moth.
e. Nutrition
The presence of calcium and magnesium in the food result to decreased recombination and removal of
such chemicals from the diet increased the rate of crossing over.
f. Chemicals
Some mutagenic chemicals like alkylating agents was found to increase the frequency of crossing over.
g. Irradiation
Exposure in radiation like X-rays and gamma rays was found to increase the frequency of crossing
over.
h. Structural Changes
Changes in structural chromosomes especially inversions and translocations reduce the frequency of
crossing over.
i. Centromere Effect
Genes located adjacent to the centromere have reduced frequency of crossing over.