CROSSING OVER

CC5 ASSIGNMENT

ANJISHNU MUKHOPADHYAY
BTN/20/704
SESSION-2020-21
INTRODUCTION

Crossing over is the exchange of genetic material between non sister chromatids of homologous chromosomes
during meiosis, which results in new allelic combinations in the daughter cells.
During prophase I of meiosis, homologous chromosome aligns with each other and exchange genetic material, so
that some of the resultant chromosomes are recombinants – containing a mixture of genes derived from the
maternal as well as the paternal chromosomes. Two of the four daughter cells formed after this crossing over event
have a recombinant chromosome that is neither completely derived from the mother nor the father. The image
also demonstrates that genes that are in close physical proximity to one another on the chromosome are likely to
be inherited together, while those that are farther away might get independently assorted during meiosis.

During prophase I, of Meiosis, chromosome condensation occurs and duplicated chromosomes with attached
sister chromatids, are initially seen as long thin threads. As condensation proceeds, homologous chromosomes are
brought together because of the similarity in structure and centromere position. A protein structure called the
synaptonemal complex also plays an important role. At this point, chromosomes are anchored to the nuclear
envelope. Now, recombination occurs between non-sister chromatids of homologous chromosomes. This is
observed microscopically as a crossing over event between bivalent chromosomes (a pair of two chromosomes)
with a tetrad structure (their duplicated sister chromatids are also visible). Towards the end of prophase-I,
homologous chromosomes now appear to ‘repel’ each other. The nuclear envelope is no longer clearly visible and
the cell then moves on to metaphase and anaphase to complete the first stage of meiosis. Crossing over begins at
Pachytene stage, after the synapsis of the homologous chromosomes has occurred in zygotene stage of Prophase
I of Meiosis.

Normally, if independent assortment takes place i.e., when genes are present on different chromosomes, we should
expect a test cross ratio of 1:1:1:1. But, a very low number of recombinant classes appeared. It may therefore be
concluded that two genes are on the same chromosome and the appearance of recombinants in low number has
resulted from crossing over.
The chromosomes usually undergo breakage during gametogenesis. Thus, a mechanism does exist by which a
group of genes on the same chromosome changes with the similar group of genes on the homologous chromosome.
The percentage of crossing-over which is obtained between different linked genes varies according to distance
between the genes on the chromosomes.

Further, the two genes are apart on a chromosome, more likely is the occurrence of crossing-over between them.
At a particular point only two of the four chromatids are involved in the exchange of their parts and to produce
50 per cent recombinant gametes.

The genes in such a case should be located so much apart on the same chromosome as to allow crossing-over in
all the mother cells during reductional division. Under such conditions genes behave as if located on different
chromosomes. Mendel’s law of Independent Assortment holds good only under the following conditions:

(a) When the genes are located on different chromosomes.

(b) If the genes are located on the same chromosome but the distance between them is so good as to produce 50
percent recombinant gametes due to crossing-over.
 Fig. 1. Diagrammatic representation of Crossing over

Example of Crossing Over.

In Drosophila the factors for pink eyes (r) and curled wings (c) are recessive, while red eyes (R) and straight wings
(C) are dominant. If a pure breeding fly with red eyes and straight wings is mated with another fly having pink
eyes and curled wings, all the F1 hybrids are red eyed and straight winged.
When a female from these hybrids is back crossed with a male having pink eyes and curled wings, the
F2 generation consists of 49% with red eyes and straight wings. 49% with pink eyes and curled wings 1% with
red eyes and curled wings and 1% with pink eyes and straight wings. The middle two classes represent the cross-
over or recombined types.
Both the factors for each allelomorphic pair, in this case, are situated in the same chromosome. If they were in
different chromosomes, four classes of individuals would have appeared in equal numbers after the back cross.
On the other hand, if they were completely linked, only the two grand parental types would have appeared in the
F2.

Fig. 2. Representing crossing over in Drosophila

Theories of Chromosomal Crossing Over
1. The ‘Contact First’ Theory:
This theory was proposed by Serebrovsky. It states that two chromatids (non-sister chromatids of inner side) first
contact and then cross each other. The breakage comes at the point of contact and broken segments reunite to form
new combination.

Fig. 3 First contact theory

2. The ‘Breakage First’ Theory:

This theory was proposed by Muller. According to him, two chromosomes (inner non-sister chromatids) first
break in to two segments without crossing over and then broken segments reunite with each other forming new
arrangement and resulting in a new combination. This theory is widely accepted one.

Fig. 4. Breakage first Theory

3. Strain Theory (Precosity theory):

According to Darlington, chromosomal breakage occurs as a result of strain or tension during pairing. When two
chromosomes’ pair and become spirally twisted around each other termed as relational coiling there develops a
sort of tension on chromatids as a result of which they become broken at the point of contact and recombination
occurs. Broken ends further re-join but not compulsorily with the same segment from which they were detached.

When broken ends of different chromosomes get joined, an exchange in genetic material occurs. Darlington claims
that it is the only chiasmata which hold the broken chromatids together as tested with Meiosis I anaphase. Thus,
this theory correlates close pairing between homologous chromosomes and crossing over.
If there is no pairing, both homologous will not form or align in a line themselves and will go the same pole
causing non disjunction i.e., pairing is necessary for disjunction or separation.

Recently Grell (1964) includes two more types i.e., exchange pairing and distributive pairing in Darlington theory.
In ‘exchange pairing’ homologous chromosomes pair and later crossing over occurs but, distributive pairing is
unaccompanied by homologous pairing and crossing over.

Belling’s Hypothesis:

According to this hypothesis, crossing over is not caused by breakage and reunion of chromosomes but by newly
duplicated genes. Belling (1933) proposed that crossing over is the result of an exchange between new chromatids
during the period of their formation.

According to this cytologist new chromomeres are formed first alongside their respective sister chromosomes and
the interconnecting fibres are formed later. If a relational coil exists between the homologues, the interconnecting
fibres may sometimes be formed between non- sister chromomeres, thus producing a cross-over.

MECHANISM OF CROSSING OVER

1. Classical Theory:

This theory was proposed and advanced by Morgan and Sharp (1934) respectively, also called two-plane theory
because it is assumed that adjacent loops would be present in different planes at right angles to each other.
According to this theory, chiasma formation occurs when a chromatid of a chromosome comes to associate with
the non-sister chromatid of a chromosome.

Generally, the sister chromatids of the two chromosomes of a bivalent remain attached with each other during
synapsis. But in many divisions the sister chromatids separate from each other and become attached with non-
sister chromatids of the homologous chromosomes thus producing chiasma.

During diplotene when the homologous chromosomes begin to separate from each other, the chromatids involved
in chiasma formation are subjected to physical tension or strain (due to equational separation and reductional
separation) at the point of chiasma. This may cause breakage of the two chromatids at this point, a reunion between
the chromatids segments thus produced would lead to crossing over or recombination between linked genes.
 Fig. 5. Classical theory

According to classical theory:

i. A chiasma is formed after non-sister chromatids of homologous chromosomes become attached
during synapsis.
ii. Chiasma formation is not the result but cause of the crossing over.
iii. Each chiasma does not lead the phenomenon of recombination or crossing over.

The above available experimental findings do not support this hypothesis and it is of only a historical significance.
This theory, however, now stands rejected.

2.Chiasma Type Theory (Breakage & reunion theory):

It was firstly proposed by Janssens (1909) and later on expanded by him in 1924. Further, fully developed by
Belling and Darlington. This is also said as one plane theory because in this theory, one would expect reductional
separation of chromatids on either side of a chiasma.

According to this theory, breakage in non-sister chromatids of homologous chromosomes, followed by the reunion
of the chromatid segments resulting in crossing over. When the homologous chromosomes begin to move away
from each other during diplotene, chiasmata are formed at those points where crossing over has taken place.

Fig. 6. Breakage and reunion theory

Thus, according to this theory: -

i. chiasma is the result of crossing over.

ii. Only sister chromatids are attached with each other throughout the whole bivalent, where as non-sister
chromatids become associated to produce chiasmata
iii. Each chiasma is the consequence of a crossing over event, therefore
iv. 1:1 ratio is expected between the numbering of frequencies of chiasma and crossing over.

Almost all the available evidence supports the chiasma type theory. Beadle (1932), Brown and Zohary (19-55)
have strongly supported this theory since there is one to one correspondence between chiasmata and genetic
crossing over explaining structure of gene maps and frequencies etc.

However, Kaufmann (1934) and Cooper (1949) objects this theory since chiasmata are also formed in some tissues
of male Drosophila. Because genetic crossing over is lacking in male Drosophila, these chiasmata cannot be
explained by chiasma type theory. Thus, chiasma type theory seems to be a universally accepted one.

3. Copy-Choice Theory:

This theory was proposed by J. Ledeberg (1955), according to this hypothesis, the duplication process and
recombination occur simultaneously. In other words, the paired chromatids duplicate by synthesizing new genes.
Then it is followed by the development of new connection between these genes. Thus, the recombination’s are
produced by newly synthesised genes.

There are mainly two objections—one is that only two of the four chromatids are involved in crossing over. Thus,
two original strands remain intact while two newly formed strands would be altered by recombination. Secondly,
this theory states that duplication should occur during late meiotic prophase but now evidence indicates that DNA
replication occurs even before synapsis.

Fig. 7. Copy Choice Theory

KINDS OF CROSSING OVER

Crossing over may be single, multiple depending upon the number of chiasmata present in the chromosomes as
follows:
1.Single Crossing Over:
When there occurs only one chiasma or crossing over at one point in chromosome pair then it is referred to as
single crossing over. The gametes which are produced by this crossing over are said as single cross-over gametes.

Fig. 8. Single crossing over

2. Double Crossing Over:

Occasionally or sometimes crossing over occurs at two points in the same chromosome pair. This is said as double
crossing over. The gametes produced by this crossing over are called double cross-overs. It occurs seldom than
single crossing over.

Fig. 9 Double Crossing over

3. Multiple Crossing Over:

Crossing over may also occur at three, four or more points in the same chromosomes pair and correspondingly
said as triple, quadruple, or multiple crossing over.
Somatic Crossing Over

Pairing in the somatic cells followed by crossing over was discovered by Stern (1931) in Drosophila, Jones in Zea
mays and G. Pontecovor et al in Aspergillus. In most organisms the meiotic division is restricted to the organs
concerned with the production of gametes in which pairing of chromosomes occur in germ cells. Stern found that
this pairing of chromosomes may also come in other vegetative tissues having somatic cells.

In Drosophila this crossing over exceptionally produced a patch or spot of cross-over tissues in certain parts, while
other body parts constitute non-cross-over tissues. Thus, fly will be mosaic for cross- over and non-cross over
tissues. Such types of crossing over in somatic cells is referred to as somatic crossing over.

THE PERCENTAGE OF CROSSING OVER

The crossing over may vary depending upon the genes and their locations. The chance of crossing over are greater
over long distances than over short distances.

In the figure it is quite clear that there are more chances of crossing over between A and C than between A & B
i.e., percentage of crossing over of two genes is related with the distance between those 2 genes. If these genes
will be near, crossing over will be minimum and likewise if they will be located at distant places, the crossing
over will be greater.

Therefore, the percentage of crossing over is directly proportional to the distances of genes between which
crossing over occurs. This is of particular importance for drawing the chromosome maps.

The percentage of crossing over may be taken as indication of the comparative linear distance between any two
required genes. The unit of crossing over was termed morgan by Haldane and adapted by Crew and others.
Suppose there is 15% crossing over between any two genes then these genes are 15 ‘morgans’ along the length of
chromosomes.

CYTOLOGICAL EVIDENCE OF CROSSING OVER

Morgan and his collaborators established the genetic basis of crossing over and linkage describing the exchange
of parts between the homologous chromosomes and linear arrangement of linked genes along chromosomes. This
genetic inclination or bias could not be demonstrated cytologically since we cannot observe the homologous
chromosomes (being all identical) under the microscope because of the following reasons.

(i) Crossing over occurs between homologous chromosomes. Such chromosomes are alike in appearance and it is
not possible to distinguish between them in microscope.
(ii) During crossing over, the four chromatids are intimately coiled around one another.
(iii) In living cells, crossing over cannot be seen. In fixed and stained cells, one cannot say that chromatids have
exchanged parts or not.
For nearly twenty years crossing over remained only a working hypothesis with geneticists. Finally, the
cytological evidence which established beyond doubt the occurrence of crossing over, was given by S. Stern on
Drosophila and H.B. Creigton and B.Mc Clintock on maize.

1. Stern’s Experiments or Drosophila:

Stern discovered a variety of Drosophila in which a part of Y chromosome had broken off and became attached
to the end of one of the X-chromosomes. Likewise, he described another variety in which one of the X-
chromosomes was broken.

Usually in Drosophila normal fly has red round (++) eyes. Two mutant genes, one carnation (car) causing darkish
red eyes and being recessive to red (+) eye colour and other bar (B) causing narrow eyes and dominant to round
(+) eyes are both in X chromosome. The female fly is XX and male has XY chromosome.

In female (XX), one X chromosome was broken in to two by X-ray or other means and contained mutant genes
car and B. The other X contains normal allele (+ for red and + for round eye i.e., normal) and its end a segment
of Y chromosome was attached. This breaking of X chromosome and attaching Y segment is a plan or scheme to
distinguish between cross-overs and non-cross-overs progeny.

Fig. 10 Stern's experiment on crossing over

The carnation barred female (car B+ +) produces four types of gametes, out of which two are cross-overs and two
non-cross overs formed without interchange of homologous segments of chromosomes. When these were mated
with males (XY) having carnation round eyes (car +), the non-cross-overs are carnation bar and red round while
cross-over contain red bar and carnation round.

Thus, cytological basis of crossing over can be established by distinguishing chromosomes under microscope.
(Stern’s experiment was a unique demonstration of the hypothesis that interchange of chromosomal material takes
place between homologous chromosomes).

2. Creighton and Mc Clintock’s Experiments on Maize:

Similar demonstration of cytological crossing over was demonstrated in maize by these workers. They observed
a corn plant which had a pair of chromosomes whose two members could be held apart cytologically. Among
them, one was normal and another had a trans-located piece of another chromosome at one end. The other end
was like a knob (hard round protuberance).

The normal chromosome carried ‘c’ for colourless endosperm and W for starchy endosperm. Other knobbed
chromosome had alleles ‘C’ for coloured and ‘w’ for waxy endosperm. Creighton and Mc Clintock crossed this
plant with a plant having homologous chromosome with recessive genes i.e., colourless waxy ‘ccww’.

As a gametogenesis, two non-cross-overs and two cross-overs gametes will be formed which after union will form
four kinds of offsprings.

The non-crossover plants i.e., colourless starchy and colourless waxy obtained from parent either knobbed
chromosome or normal but cross-over plants had one chromosome of this particular pair i.e., colourless waxy
(ccww) contained a chromosome with trans-located piece but no knob, whereas coloured starchy (CcWw) showed
knobbed chromosome but no trans-located piece.

Thus cross-overs showed cytological evidence of crossing over i.e., exchange of homologous chromosome parts
during maturation of germ cells.
 Fig. 11. Creighton and Mc Clintock's experiment on crossing over

FACTORS AFFECTING CROSSING OVER

There are various genetical, physiological and environmental factors which affect the frequency of crossing over
i.e., promotes or suppresses the percentage of crossing over between 2 loci.
1. Sex:

It has been observed in male Drosophila that the crossing over is completely suppressed or masked and there is
also tendency of reduction of crossing over in male mammals. In silk-moth (Bombyx) crossing over does not
occur in females. Thus, the sex of an individual may also affect the frequency.

It was demonstrated that X-rays may bring about definite amount of crossing over as in male Drosophila. In
general, it has been found that the heterogametic sex shows comparatively lower crossing over frequencies than
the homogametic sex of the same species.

2. Mutation:

Gene mutation may affect the frequency of crossing over. Gowen has found a mutation in Drosophila which
reduces the percentage of crossing over in all chromosomes.

3. Age:

The age of the individual may also affect the frequency of crossing over. Bridges already found in Drosophila that
as the female becomes older, the crossing over tends to decrease.

4. Inversion:

These are intra-segmental changes within the chromosomes. In a given segment of chromosomes crossing over is
reduced due to inversion. The part of the chromosomes which is not involved in inversion, the frequency of
crossing over increases. The cause for this is still not clear.

5. X-radiation:

Hanson showed that irradiations by radium also increase the frequency of crossing over. Muller also found
increase in the frequency of crossing over after the application of X-rays.

6. Temperature:

According to Plough, low and high temperature increase the frequency of crossing over. Example is female
Drosophila.

7. Centromere vicinity:

It has been observed that near centromeres and at the tips of chromosomes, crossing over is frequent. Some factors
not clearly understood influence crossing over in some somatic cells also that is said as somatic or mitotic crossing
over. It is, however, known that certain mutant genes called minute or small bristles increase the frequency of
somatic crossing over since a single cell gives rise to a mass of somatic cells.

It follows that a somatic cell in which crossing over occurs, gives rise only to a patch of cross-over tissue. Further,
such somatic cells do not produce gametes. This type of crossing over has no genetic importance. Somatic crossing
over was first of all seen & studied by Stern in Drosophila. It should be borne in the mind that factors which affect
crossing over, effect linkage inversely.

SIGNIFICANCE OF CROSSING OVER:

Crossing over as a measure of genetic distance

Sturtevant’s fundamental insight was to estimate the distance between points on a chromosome by counting the
number of crossovers between them. Points that are far apart should have more crossovers between them than
points that are close together. However, the number of crossovers must be understood in a statistical sense. In any
particular cell, the chance that a crossover will occur between two points may be low, but in a large population of
cells, this crossover will probably occur several times simply because there are so many independent opportunities
for it. Thus, the quantity that we really need to measure is the average number of crossovers in a particular
chromosome region. Genetic map distances are, in fact, based on such averages. This idea is sufficiently important
to justify a formal definition: The distance between two points on the genetic map of a chromosome is the average
number of crossovers between them. One way for us to understand this definition is to consider 100 oogonia going
through meiosis. In some cells, no crossovers will occur between sites A and B; in others, one, two, or more
crossovers will occur between these loci. At the end of meiosis, there will be 100 gametes, each containing a
chromosome with either zero, one, two, or more crossovers between A and B. We estimate the genetic map
distance between these loci by calculating the average number of crossovers in this sample of chromosomes in
practice, we cannot “see” each of the exchange points on the chromosomes coming out of meiosis. Instead, we
infer their existence by observing the recombination of the alleles that flank them. A chromosome in which alleles
have recombined must have arisen by crossing over. Counting recombinant chromosomes therefore provides a
way of counting crossover exchange points.

Fig. 12 Chromosome recovered from meiosis in gametes

 Fig. 13 Using crossing over as gene distances

In genetics, a centimorgan (abbreviated cM) or map unit (m.u.) is a unit for measuring genetic linkage. It is
defined as the distance between chromosome positions (also termed loci or markers) for which the expected
average number of intervening chromosomal crossovers in a single generation is 0.01. It is often used to infer
distance along a chromosome. However, it is not a true physical distance.

The centimorgan was named in honor of geneticist Thomas Hunt Morgan by J. B. S. Haldane.[5] However, its
parent unit, the morgan, is rarely used today.

Other significances

(1) It has a great significance in genetics. Crossing over, a wide spread phenomenon provides direct evidence of
the linear arrangement of genes. Construction of chromosome maps and tracing linkage groups has been greatly
facilitated by data obtained from the study of crossing over.

(2) It increases the frequency of variation which are vital for evolution. It causes formation of many combinations
which can be acted by natural selection. The established linkage groups and linier order of genes gives much light
on the nature and mechanism of genes.

Other Genetic terms related to Crossing Over

Chromosome Maps:

The chromosome maps represent the condensed graphic representation of the relative distance of the genes in a
linkage group, expressed in the percentage recombination located and single group of chromosomes. From the
examples we have studied linkage and crossing over, it can be said that linkage can be taken as an exception to
the second law of Mendel, and the characters of an organism are due to genes located in the chromosomes.

Moreover, genes are believed to occur in a number of linkage groups. The linkage groups correspond to the
number of chromosomes. The linkage group in Drosophila melanogaster are four in number and there are four
pairs of chromosomes in that fly.
In Morgan’s hypothesis of crossing over it has been assumed that the genes had a linier arrangement in the
chromosome. It was also thought that the distances between the two genes on the chromosome is correlated with
the amount of crossing over shown by two corresponding alleles. The percentage of crossing over is directly
proportional to the distance of alleles showing cross in the chromosomes.

These two facts made to represent in the form of a map which represents that:

(i) The genes are arranged in a linier row along the chromosome.
(ii) The percentage of crossing over between two genes is proportional to their distance apart.
Thus, chromosome map may be defined as a line, number of lines being equal to linkage groups on which genes
are represented by points showing particular traits or characters proportional to the amount of crossing over.

These chromosome maps are also referred to as cross-over maps since they are sketched by the amount of crossing
over.

The percentage of crossing over is calculated by test crosses. In mapping genes, a unit of distance is used. This
unit used is one percent of crossing over called as Map unit or morgan. The crossing over between linked genes
may be as little as 1/10 of 1% or up to 50% depending upon their kinds.

The first two chromosome maps were made in 1911 by Sturtevant and Bridges. Later in 1920, Morgan and his
associates worked extensively on Drosophila and constructed chromosome maps. Then these maps were made in
maize, chicks, tomato etc.

Location of Genes:

In fixing the exact position of a gene on the chromosome map, the cross-over frequency of a gene in relation to
another is the criterion. The procedure adopted for constructing a chromosome map can be explained with
reference to a three-point test cross. A three-point test cross is one in which the F1 resulting from a cross involving
three linked genes is back crossed to a triple recessive.
In Drosophila the three genes scute (sc), echinus (ec) and crossveinless (cv) are sex linked genes. Scute is
condition in which many body bristles are missing, echinus means rough eyes and crossveinless indicates absence
of cross veins on the wings, since these are recessive mutation, the F 1 females resulting from a cross between
these recessive types and wild type resembles the wild type phenotypically. This is because the females drive one
sex chromosome from their mother.

Fig. 14. Example of Gene location calculation

When the F1 females are back crossed to triple recessive males, eight phenotypes are obtained as given
below:

In constructing the chromosome map the distance between the two linked genes is indicated by their cross-over
frequency i.e., percentage. Since the cross-over frequency of scute in relation to echinus is 7.6%, these two genes
are marked 7.6% unit apart. Again, the cross-over frequency between echinus and crossveinless is 10.1%; these 2
genes are 10.1 unit apart.

In order to determine the sequence of the three genes it is necessary to find out the cross-over frequency between
scute and crossveinless. The genes scute and crossveinless and the wild type alleles were introduced in to the cross
by the same parent, one of these is present in the progeny without the other in 352 (151+201) cases.

To these figures the two double cross-overs may be added. The total number of cross-overs between scute and
crossveinless is thus 354 or 17.7%. This is the sum and not the difference between 7.6% and 10.1. Therefore, the
gene crossveinless lies beyond echinus; it means sequence is sc, ec and cv.

Interference and Coincidence

Besides single crossing over, having only one chiasma, there may be double or multiple crossing over. It has been
discovered by H.J. Muller (1911) that when there are two double cross-overs (suppose a and b) then one cross-
over (a) tries to prevent the formation of other cross over (b) This tendency of one cross-over to interfere with the
other cross over is termed as interference. Suppose frequency of ‘a’ crossover is 10 and frequency of ‘b’ cross-
over is 12, then their total frequency will not be 10+12 = 22 as required but will be less than 22 due to interference.

When the two things happen the same time and at the same place, they then coincide or intermix and this
occurrence may be considered coincidence. This coincidence refers to the occurrence of two or more distinct
cross-over (double or multiple) at about the same time in the same chromosomal region. Double cross-overs are
the result of coming together (coincidence) of two single cross-overs.

When doubles occur in regular expected ratio, coincidence is said to be 100%, whereas interference will be nil.
But when there are no doubles (coincidence) the interference is nil i.e., coincidence is inversely proportional to
the interference.

According to Muller (1916) the coefficient of coincidence is the ratio between the observed and expected
frequencies of double cross overs. Coefficient of coincidence = Actual number of double cross-overs/Expected
number of double cross-overs.
 Conclusion

Crossing over increases the variability of a population and prevents the accumulation of deleterious
combinations of alleles, while also allowing some parental combinations to be passed on to the offspring. This
way, there is a balance between maintaining potentially useful allelic combinations as well as providing the
opportunity for variation
and change. It provides the way of micro evolution. Linkage map and genetic maps are constructed on the basis
of Crossing-Over. Thus, it is a necessary phenomenon for evolution to occur. Crossing over is thought to have
been spontaneously occurred due to interactivities of non-sister chromatids.

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