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Provisional chapter
Chapter 7
Molecular Breeding of Cotton
Molecular Breeding of Cotton
Yuksel Bolek, Khezir Hayat, Adem Bardak and
Yuksel Bolek, Khezir Hayat, Adem Bardak and
Muhammad Tehseen Azhar
Muhammad Tehseen Azhar
Additional information is available at the end of the chapter
Additional information is available at the end of the chapter
http://dx.doi.org/10.5772/64593
Abstract
Molecular characterization provides comprehensive information about the extent of
genetic diversity, it assists for the development of an efective, highly accurate, and rapid
marker‐assisted coton breeding program. Due to one of the world s leading iber crops,
molecular studies of coton are being explored widely by coton researchers. Coton
provides raw material to the textile industry among other products. Limitations in
conventional breeding program for genetic improvement are due to the complexity and
limited knowledge on economically important traits. The use of molecular markers for
the detection and exploitation of DN“ polymorphism is one of the most signiicant
developments in molecular genetics. In the present scenerio, coton molecular breeding
has become a reliable source through the study and exploitation of its genetic diversity
and due to beter understanding of the coton genomes using the next‐generation
sequencing technologies. Coton breeders should utilize genomics in breeding
programs for efective selection of best parents for agronomic and iber‐related traits,
as well as for the development of resistance against biotic and abiotic stresses. The
genomic research work could be based upon genotyping using DN“ markers,
quantitative trait loci mapping, genome‐wide associations, and next‐generation
sequencing. The objective of this chapter is to describe evolution as well as utilization
of various molecular markers and review the contribution of marker‐assisted selection
(M“S) in coton breeding.
Keywords: coton, DN“ markers, genotyping by sequencing (G”S), genome‐wide as‐
sociation studies (GW“S), marker‐assisted selection (M“S)
© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons
© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons
Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution,
Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use,
and reproduction in any medium, provided the original work is properly cited.
distribution, and reproduction in any medium, provided the original work is properly cited.
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Cotton Research
. Introduction
Plant breeders select those plants, which looks phenotypically more promising due to the
presence of desirable traits. Most of the traits are controlled by polygenes with complex
nonallelic quantitative efects and environmental interactions. In most cases, despite the fact
that biometrical genetics reveals the presence of additive or non‐additive efects on loci
involved in the inheritance of quantitative trait, a speciic locus may not be detected [1]. Tightly
linked loci with desired trait can support plant breeding program by rapid introgression of
quantitative trait loci (QTL) using associated molecular markers [2]. Genomic region having
genes of interest for a particular trait is designated as QTL (Quantitative Trait Loc). QTL
analysis involves partioning of genetic variation in single component. So, DN“‐based
molecular markers provide a tool to plant breeders for the selection of desirable plants based
on genotype instead of phenotype.
The expression of gene(s) individually their interaction with the climatic factors and agronomic
measures can determine the cultivar adaptability [3]. Selection of new plant varieties with the
desirable traits under given environmental conditions and cultural practices is the fundamen‐
tal basis of plant breeding [4], genetic variability produced in germplasm as a result of selection,
which alter the inheritance patern of the traits, is quite useful to screen and select the cultivars
for required traits. New cultivars have been developed by exploiting genotypes with enormous
variation [5]. Rapid changes are needed in agricultural production, and biologically diverse as
well as low‐input novel farming systems must be developed and employed. There is also a
need for new crop varieties that are (1) iting‐in to global climate change in the present era,
(2) adapted to biodiverse farming systems, and inally (3) giving more products to farmers and
eventually to consumers.
Coton (Gossypium spp.) is one of the most intensively cultivated species grown in more than
80 countries in varying climatic conditions [6]. Globally, coton is the ultimate source of iber
for industry and provides oil to diet [7]. ”eing utmost iber manufacturing crop and the third
contributor to oilseed production; China, India and US“ are top contributors for iber [8].
Gossypium genus is divided into eight genomes (“‐G and K) and comprised of 45 diploids and
ive allotetraploid species which are found in the arid and semiarid regions of “frica, Central
and South “merica, Galapagos, Indian subcontinent, “ustralia, “rabia, and Hawaii [9 11].
“t the beginning of the 20th century, scientists discovered that Mendelian factors controlling
inheritance are organized in linear order on chromosomes. It was shown that genes could be
inherited individually or in combination with other genes. The individual fragments lanking
within a deined interval are known as molecular DN“ markers [12]. Precise DN“ portion
with a known position on the chromosome [13], or a measurable trait that is associated with
variation in DN“ sequence [14, 15] or a diference may act as a genetic marker if it identiies
characteristics of an individual.
Markers are broadly divided into three classes: (1) morphological markers, which themselves
have phenotypic traits meaning the morphological and physiological features of plants are
used to understand the genetic variation. “lthough morphological features may be indicative
Molecular Breeding of Cotton
http://dx.doi.org/10.5772/64593
of the phenotype, they are also highly afected by environmental factors and growth practices;
(2) biochemical markers, including isozymes, which involve allelic variants of proteins/
enzymes; (3) molecular markers, manifest mutations in heredity material such as DN“s and
RN“s [16 19].
Polymorphism of molecular markers shows diferentiation of homozygotes and heterozygotes
[20]. Thotappilly et al. [21] refer to molecular markers as naturally occurring polymorphism,
which include the proteins and nucleic acids that indicate certain diferences. The use of
molecular markers in plant breeding is called marker‐assisted selection, often referred as M“S
or marker‐assisted breeding (M“”) (Figure ) [4, 22].
Figure . Marker‐assisted scheme [4].
In traditional plant breeding, traits are selected depending on the phenotype, which is highly
afected by the climatic factors. This approach makes the breeding a slow, expensive, and
challenging process [23 25]. Practical advantages of using genetic markers, potential values of
linkage maps, and exploiting for direct selection in plant breeding were begun to be studied
about the 1930s [26]. Molecular markers are essential for mapping the genes of interest, M“S/
M“”, and cloning of genes using mapping‐based cloning strategies [27]. In addition, the use
of molecular markers includes gene introgression through backcrossing, germplasm charac‐
terization, and phylogenetic analysis [28]. It has been observed that M“S is more eicient than
conventional breeding techniques [4, 29, 30]. Selection based on genotypic structure through
employment of molecular markers in the ield crops [31] has laid the foundation of M“S [32,
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Cotton Research
33]. Many biological and medical science applications and studies, including genetic diversity,
molecular tagging of economic traits, and procurement of heritable diseases have successfully
utilized molecular markers [2, 34 37]. Thus far, molecular markers have been exploited in rice
[38], wheat [39], maize [40, 41], and barley [42, 43]. However, M“S has achieved the desired
goals in coton with limited success due to a genetic botleneck through historic domestication
and limited polymorphism in cultivar germplasm [44 47].
“bout 145 morphological markers are reported in coton so far, but they have low utility in
variety development because of incapability to assemble diverse markers in a genotype [48].
Isozymes produced through allelic variants are considered more authentic but not widely used
due to their diferential expression in diferent growth stages. For improving productivity and
other key quantitative traits, coton genetic markers have more value than morphological or
isozyme markers [48]. DN“ markers have become handy and efective tools for plant breeders
because their expression is not necessary for their detection [49]. In order to enhance the
beneits through molecular markers, vast developments have been made in omics , which, in
turn, allowed the use of these markers in diverse ways for genetic studies instead of using them
solely for phylogenetic studies [50]. Obtaining pure DN“ plays a major role for the develop‐
ment of molecular markers in coton [51 53]; genetic analysis has many drawbacks due to the
presence of phenolic compounds, which afect quality of DN“ and protein during tissues
grinding [51].
Polygenic traits are mostly afected by the climatic conditions and show discrete variability
after hybridization. Recombination frequency allows investigators to diferentiate genes on
linkage map by relative distance between a generation and their parents. The main hindrance
for QTL mapping of agronomic traits is related to a large number of genes involved in
phenotypic expression and their interaction with the environment [54]. “s number of genes
afects the trait phenotypically, it is desired that more loci should be evaluated for QTL
determination, and the screening of individuals should be done at multiple locations/envi‐
ronments to maximize the use of QTLs. M“” uses QTLs to pyramid favorable alleles and break
linkage groups for tagging QTLs of interest [55 57]. In recent years, conventional plant
breeders started to use M“S for the identiication of traits with high heritability such as disease
resistance, as well as the yield of major ield crops [57]. However, yield‐related components
have low heritability, which is a major challenge for the utilization of M“S [56, 58]. M“S is
being employed for the identiication of transgressive segregants. Transgressive segregation
is the production of plants in F2 generation that are superior to both parents for one or more
traits. Transgressive breeding aims at improving yield or contributing to yield‐related traits
through transgressive segregation [59 61]. Several QTLs have been identiied for seed coton
yield, iber quality, plant architecture, resistance to diseases such as bacterial blight and
Verticillium wilt [57], resistance to pests like root knot nematode, and lowering date [62] as
well as for abiotic stresses (drought, salt tolerance) [55, 63].
There is a gap between discovery of useful genes and QTLs, and their utilization in breeding
programs. To date, few examples are reported [55, 63] for the successful release of genotypes
developed by M“S, and they have shown signiicant contribution to yield improvement. High‐
throughput, high‐density genome‐proiling tools enable the rapid and low‐cost of crop
Molecular Breeding of Cotton
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genome in a precise and high‐resolution manner. Identiication of molecular variants in DN“
sequence opens opportunities for plant scientists [55]. The potential exists in plant breeding
for eicient use of next‐generation sequencing (NGS) that also has revolutionized the plant
genomics [55]. Markers can be analyzed across the genomes simply and accurately, with high‐
throughput. Increased number of next generation sequencing allows conducting genome‐wide
association studies (GW“S) [63]. It is thanks to the developments in knowledge of useful
genetic diversity and QTLs, advances in sequencing, genotyping, and bioinformatics ap‐
proaches that rapid, high‐throughput molecular marker discovery methods have been
enabled.
Day‐by‐day developments of new, speciic markers, and trait determination tools makes
molecular markers important in understanding the genomic variability and diversity within
and among species. In this chapter, we discuss about the applications and types of molecular
markers, next‐generation sequencing, and role of molecular breeding in development of plants
with improved economical traits in coton.
. Breeding for polygenic traits
Economically important traits such as nutritive value, earliness, agronomic traits, resistance
etc. can be improved through M“S [64, 65]. Polygenic mapping allows breeders to estimate
and assess the hereditary patern of the traits governed by many genes found throughout
genome; Ultimately it leads to eicient utilization of these traits for molecular breeding. Highly
saturated genetic maps in a high population index permits to observe the impact of many
regions of genomes on a single trait value. Paterson [66] revealed that sharing of homologues
during crossing over is the basis of QTLs. The regions of the genomes connected to the traits
of economic value are QTLs [67]. “ssociation of a marker s genotypic value to a phenotype is
the basis of QTL mapping. Recombination frequency is used to evaluate the relative distance
among markers in the linkage map. It is assumed that markers at or lower recombination ratio
of 50% are considered as unlinked found either on homologues or alternative loci while the
markers which are tightly connected will be transferred to ofspring more often than the
unlinked markers [67].
Reinisch et al. [68] developed the pioneer genetic map of coton during 1994. “lthough large
number linkage maps have been constructed since then due to abundance of several DN“
markers, it is still needed to determine reliable QTLs from breeding perspective. Yu et al. [69]
screened genotypes by simple sequence repeats (SSRs) to map loci connected to iber quality
and lint yield in a backcross inbred line and developed a pioneer genetic map using ”IL within
allotetraploid cultivated coton species. Map consisted of 392 highly cosegregated loci covering
2895 cM length and having mean interlocus distance 7.4 cM. “s a whole, 39 QTLs were directly
connected to yield components and 28 were associated to iber quality.
“ltaf et al. [70] explored F2 population developed from three diferent species of Gossypium for
identiication of evolutionary relationship among these species by linkage map. Eleven linkage
groups were constructed having 521.7 cM map size in coton genome and relative distance of
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16.8 cM was found among markers through screening randomly ampliied polymorphic DN“s
(R“PDs) and ampliied fragment length polymorphisms (“FLPs). Jiang et al. [71] utilized F2
population developed from G. hirsutum × G. barbadense, and produced a restriction fragment
length polymorphism (RFLP) genetic map having 3767 cM length; 27 linkage groups with
distance of 14.4 cM among loci.
Shappley et al. [72] used F2:3 families derived from HS‐46 and M“RC“”UC““G‐1‐8896
genotypes and constructed genetic map by using 120 RFLPs which spanned to 865 cM and
arranged in 31 linkage groups. Fifty one linked groups were developed through a map
constructed with RFLP and R“PD markers [73] spanning to 6663 cM including 332 “FLPs, 91
R“PDs and three morphological markers. Khan et al. [74] studied comparison for ploidy level
to diploid ancestors and tetraploid coton with R“PD markers. 119 F2:3 families developed from
MD5678ne × Prema and utilized RFLPs for genetic map. Seventeen linkage panels were
distributed on 700.7 cM map having mean distance of 7 8 cM among the markers [75].
RFLP, “FLP, and SSRs were screened in a backcrossed breeding population derived from
crosses of [(G. hirsutum cv. Guazunchoz × G. barbadense cv. VH8‐4602) × G. hirsutum cv.
Guazunchoz] [76]. Linkage map covered 4400 cM of genome and consist of 888 loci arranged
on 26 and 11, long‐ and short‐linkage groups, respectively. EST‐SSRs from G. arboreum were
used for linkage map construction in backcross inbred line [(TM1 × Hai7124) × TM1] [77]. Map
spans to 5644.3 cM with mean interlocus distance of 9.0 cM. “s a whole, 111 loci were detected
with these 99 EST‐SSRs incorporated into backbone map including 511 SSR loci. These EST‐
SSRs will be useful in M“S for improving iber quality.
Mei et al. [78] developed interspeciic population among G. hirsutum L. cv. “cala‐44 and G.
barbadense L. cv. Pima S‐7 and published genetic map, which covers 3287 cM of the genome.
They used “FLPs, RFLPs, and SSRs and have; identiied total 392 loci being 333, 12, and 47
markers, respectively. They were able to identify high repetitive DN“ and heterochromatin in
D‐genome and relative distance among mapped loci in “‐genomes that were also compared
to homologous in D‐genome [79].
Two hundred and thirty‐three linked loci were mapped in backcross population of [G. hirsutum
cv. Guazunchoz × G. barbadense VH8‐4602) × G. hirsutum cv. Guazunchoz] by using 204 SSR
markers, which produced 261 polymorphic bands [80]. Linkage map was published by adding
233 loci to already developed map [76] covering 5519 cM genome and having mean inter‐
related marker distance of 4.8 cM and consisting of 1160 loci. Nugyun et al. [80] applied STS
markers for developing linkage maps that will fasten the genomics era by using diploid and
tetraploid (“tDt) genomes. The genetic map consists of 2584 loci having İnter‐locus marker
distance of 1.72 cM and 763 loci intervals depending on 2007 probes from allotetraploid
genome while 763 loci at relative marker distance of 1.96 cM intervals identiied by 662 probes
in D‐genome. “ll desired homologous chromosome pairs were observed owing to locus
repetition. Moreover, number of chromosomal variations including number of inversions and
reciprocal translocations were observed.
Wang et al. [81] applied microsatellites for identifying QTLs related to iber quality in RIL
population. The genetic map was published with two common QTLs for lint percentage and
Molecular Breeding of Cotton
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iber length. The results were in accordance to earlier studies and can be utilized in marker‐
assisted breeding. Lin et al. [82], screened SR“P, SSR, and R“PDs, have constructed linkage
map, and a mean relevant distance was 9.08 cM among markers and total length of the map
was 5141.8 cM. Park et al. [83] published the pioneer linkage map by applying EST‐SSRs in RIL
population derived from (G. hirsutum TM1 × G. barbadense Pima) for iber. The linkage having
about 27% genome coverage, covering 1277 cM genome and having 193 loci of those 121 newly
mapped for iber traits.
Researchers [84, 85] have used SSRs and “FLPs for determining oligonucleotides that is a good
source for pyramiding of genes for marker‐assisted selection. The mapping population
developed by crossing parents having diversity for drought. Highly favorable environment
was used; dryland and irrigated regimes for screening of genotypes. Quantitative trait loci
mapped on diferent loci including one QTL (”NL1693) for seed coton production on
chromosomes 1 and 15 and two additional QTLs (”NL1153 and ”NL2884) on chromosome 6.
Moreover, chromosomes 6, 14, and 25 having ”NL2884, ”NL3259, and ”NL1153 marker‐
associated QTLs found for osmotic pressure for drought in highly uniform lines. Researchers
also revealed that N“U2715 and N“U2954 can be used as marker for relative water contents
while relative water contents with N“U2954 will contribute a lot to drought tolerance in coton.
SSRs were analyzed for establishing genetic diversity and QTLs [86]. F2 population of crosses
(7235 × TM 1), (HS 427‐10 × TM‐1), and (PD 6992 × SM 3) utilized for assigning QTLs for iber
traits in the three diferent linkage maps which span to 666.7, 557.8, and 588 cM, respectively,
with number of mapped loci with diference of 86, 56, and 73 [86].
He et al. [87] screened R“PDs, Retrotransposon‐microsatellite amplified polymorphism
(REM“P), SSRs, and sequence‐related amplified polymorphisms in hybrids of G. hirsutum L.
cv. Handan 208 and G. barbadense L. cv. Pima 90 for construction of linkage map. “s a whole,
1029 loci were mapped on 26 chromosomes; map spans to 5472.3 cM with mean İnter‐locus
distance of 5.32 cM. Saleem et al. [85] determined two QTLs related to drought tolerance in F2
progeny developed from diverse parents by applying SSRs and EST‐SSRs. The progeny
screened with parents for osmotic pressure using hydroponic culture.
“bdurakhmonov et al. [88] revealed that chromosomes 12, 18, 23, and 26 having QTLs
controlling lint percentage by applying SSRs and EST‐SSRs in a RIL population. Four QTLs for
lint index, eight for seed index, 11 for lint yield, four for seed coton yield, nine for number of
seeds per boll, three for iber strength, ive for iber length, and eight for iber ineness were
determined in F2 population (G. hirsutum L. cv. Handan 208 × G. barbadense L. cv. Pima 90) [87].
SSRs were used to screen F2 progeny for nematode resistance [89], and researchers identiied
gene G”713 that control resistance, and could be used for reniform nematode resistance. They
found two QTLs located on chromosome 21 having 168.6 cM on the genetic map while other
QTL was located on chromosome 18. Morphological traits of RIL populations developed by
hybridization of G. hirsutum and G. barbadense [90]. QTLs governing the plant architecture
including plant height, number of primary and secondary branches were screened. Research‐
ers found that angle of branch, angle of fruits, plant height, leaf size, main fruiting, etc. were
governed by a single QTL. Infestation of disease is a severe problem in coton, e.g., Xanthomonas
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oxysporum [91], root knot nematode [89, 92 94], Verticillium [57], and coton leaf curl disease
(CLCuD) [95, 96] that warn the coton scientist to ind natural resistance sources and their
urgent exploitation by using M“S.
. DNA makers in coton
Several types of molecular markers are available for characterization of germplasm of crop
plants (Table ). The amount of variation prevailing in the germplasm helps to maintain genetic
conservation [98]. “vailability of vast genomic database provides opportunity to develop
enormous markers for detection of genetic variation [99, 100]. “ccording to Weising et al. [101],
these molecular markers must be (1) highly polymorphic, (2) codominant, (3) evenly distrib‐
uted in a genome, (4) without pleiotropic efects, (5) easy to handle and fast assayed, (6) low
cost and reproducible.
The cost of production of a marker is directly related to marker technique in use, polymorphic
nature, and eiciency [102]. Polymorphic markers are divided into three types: (1) hybridiza‐
tion‐based, (2) polymerase chain reaction (PCR) based, and (3) DN“ sequence based markers
[103].
. . Hybridization‐based markers
Hybridization is occurred to the fragments of genomic DN“s produced by restriction endo‐
nucleases with various lengths among individuals. These types of markers are called hybrid‐
ization‐based markers.
. . . Restriction fragment length polymorphism
Restriction fragment length polymorphism (RFLP) is a type of hybridization‐based marker in
plant genome and initially used for detection of polymorphism in a DN“ sequence for gene
mapping during the 1975s [31]. Nucleotide sequences of 4, 5, 6, or 8 bp, called restriction sites,
are recognized by restriction endonucleases [104]. Digestion of DN“ with restriction enzymes
results in fragments whose number and size can vary among individuals, populations, and
even within species.
Many scientists developed genetic mapping during the 1975s populations of cotons that were
analyzed by using RFLP. Domestication of G. hirsutum was investigated with RFLPs [105]; they
have revealed that Yucatan is the wild ancestor of upland coton. Wright et al. [106] used RFLP
for M“S and evaluated resistance allele for bacterial blight resistance. Hybridization carried
out with probes for microsatellite sequences to yield a variable number of tandem repeats
(VNTR) and allow oligonucleotide ingerprinting [107]. Joint map was constructed by using
F2:3 populations derived from diferent intra‐hirsutum accessions [108]. Two hundred and
eighty‐four polymorphic markers and 49 linked pairs were observed on the map. The genetic
map spanned to 1502.6 cM having 5.3 cM distance between markers. RFLPs have played a
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signiicant part for omics studies [109]. Low level of polymorphism, costly chemicals, and more
time for analysis limit RFLP use in M“S [104].
S.N. Feature
1
DN“ require
(µg)
RFLP
10
RAPD
0.02
AFLP
0.5 1.0
SSRs
0.05
SNPs
0.05
2
PCR based
No
Yes
Yes
Yes
Yes
3
DN“ quality
High
High
Moderate
Moderate
High
4
No of
polymorphic
loci analyzed
1 3
1.50 50
20 100
1 3
1
5
Type of
polymorphism
Single base
change,
insertion
deletion
Single base
change,
insertion
deletion
Single base
change,
insertion
deletion
Change in
repeat
length
Single base
change,
insertion
deletion
6
Dominance
Codominant
Dominant
Dominant/
codominant
Codominant
Codominant
7
Reproducibility
High
Unreliable
High
High
High
8
Ease of
use and
development
Not easy
Easy
Easy
Easy
Easy
9
“utomation
Low
Moderate
Moderate
High
High
10
Cost per
analysis
High
Low
Moderate
Low
Low
11
Developmental
cost
Low
Low
Moderate
High
High
12
Need for
sequence
data
Yes
No
No
Yes
Yes
13
“ccuracy
Very high
Very low
Medium
High
Very high
14
Radioactivity
detection
Usually yes
No
No
No
Yes
15
Genomic
abundance
High
Very high
Very high
Medium
Medium
16
Part of
genome
surveyed
Low copy
coding
regions
Whole
genome
Whole
genome
Whole
genome
Whole
genome
17
Level of
polymorphism
Low
Low to
moderate
Low to
moderate
High
High
18
Inheritance
Codominant
Dominant
Dominant
Codominant
Codominant
19
Detection
of alleles
Yes
No
No
Yes
Yes
20
Utility for
genetic
mapping
Species
speciic
Cross
speciic
Cross
speciic
Species
speciic
Species
speciic
21
Utility in
marker‐assisted
selection
Moderate
Low to
moderate
Low to
moderate
High
Low to
moderate
22
Cost and
labor involved
in generation
High
Low moderate
Low moderate
High
High
Table . Salient features of various molecular markers [97].
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Ulloa et al. [110] published genetic maps by using intraspeciic populations developed from
parents having diverse genetic background. Fifteen linkage groups were used for designating
the chromosomes. Earlier mapped data was used for construction of map by observing the
deiciency analysis of the probes. QTLs were determined for iber and yield traits by using this
map. “s a whole 63 QTLs were found in “ subgenome at ive diferent loci and 29 QTLs
observed at 3‐loci of D‐subgenome. First genetic map spans to 117 cM produced 26 QTLs with
54 RFLPs while second map produced 19 QTLs with 27 RFLPs, and spanned to 77.6 cM. It was
revealed that these maps will serve as map‐based cloning for iber quality.
. . PCR‐based markers
PCR‐based markers, i.e., R“PD [111 113], “FLP [114‐116], microsatellites (SSRs) [117‐119],
and inter‐simple sequence repeats (ISSRs) [120‐121] represent major class of markers in coton
genomics due to their high utility and exploitation. ”elow are the major advantages of PCR
techniques as compared to hybridization‐based methods: (1) low amount of DN“ used for
genotyping; (2) capacity to amplify fragments from frozen cells; (3) high polymorphism that
enables to generate many genetic markers within a short time; and (4) ability to screen many
genes simultaneously either for direct collection of data or provide opportunity to collect
information prior to submit for nucleotide sequencing [109].
The comparison of diferent aspects of generally used molecular markers is given in (Table )
and brief description of these three classes of molecular markers is described below with
reference to coton genetics.
. . . Ampliied fragment length polymorphism
“mpliied fragment length polymorphism (“FLP) relies on the restricted sequences and PCR
ampliication. Initially, genomic DN“ is digested by a restriction enzyme and resulting
fragments are ligated with adapters to both ends. Then, the adapter and restriction site
sequences are selectively ampliied; only the fragments whose ends are complementary to 3
ends of selective primers are ampliied resulting in small sequences. Finally, a gel is run for the
separation of ampliied fragments and it is visualized by luorescence [34]. The focal point of
this methodology relies on the magniication of endonuclease restricted fragments through
PCR.
The important advantages of using of “FLP markers is that they exist in large numbers in
genomes, they have a great reproducibility due to high PCR annealing temperatures, and less
cost per marker basis [104]. In addition to reliability and reproducibility [116], there is no need
of DN“ sequence for analysis. In contrast to RFLPs and microsatellites, enormous polymorphic
loci can be investigated by having single oligonucleotide pair running a single gel through
“FLPs [122]. For digestion; partially degraded DN“ and good quality DN“ can be utilized,
but care should be taken that isolated genomic DN“ should be free of chemicals that interferes
with polymerase chain reaction.
Lacape et al. [97] initially developed RILs population by introgression of Guazuncho 2 (G.
hirsutum) and VH8‐4602 (G. barbadense), and constructed a genetic map with 800 markers
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(“FLP, RFLP, and SSR) loci. “FLPs and R“PDs were used for development of linkage map in
coton [123]. Three hundred and seven SSR markers and 72 “FLP oligonucleotides were used
for the development of genetic map in F2 population which derived from intra‐hirsutum
hybridization. The map consisted of 27 linkage groups and it has 21, 72 cM distance between
the markers [114]. Map saturation in various genotypes of cotons was analyzed [115].
“FLPs were screened in a backcross population developed from intra‐hirsutum cultivars for
agronomic traits and iber quality enhancement [124]. They found 50 “FLPs associated with
the iber quality traits and few for other; further evaluated that E1M1‐106, E1M4‐153; E1M3‐
168, E6M3‐266 for lint yield and lint percentage, respectively can be used in future for M“S
[124]. “FLPs were used for introgression among G. hirsutum and G. tomentosum being close
relative to Upland coton [125]. Through analysis, species‐speciic [11, 16] “FLP markers were
selected from G. hirsutum and G. tomentosum, respectively for assessing G. hirsutum relatedness.
These species‐speciic “FLP markers would be useful for detecting gene low between G.
hirsutum and G. tomentosum.
Jixang et al. [124] revealed genetic diversity in a germplasm by using “FLPs. “ range of 0.1
0.34 estimates of genetic diversity were found among the genotypes, and showed that
genotypes having signiicant variation in the gene stock include “U 5367, “cala 1517‐99, and
L“ 05307025.
. . . Random ampliied polymorphic DNA
Randomly ampliied polymorphic DN“ (R“PD) relies on use of short and random primers to
amplify random portions of genome [126]. Such markers have found to be widespread in
population genetic studies whose characterizations of genetic diversity and divergence within
and among populations are based on assumptions of Hardy‐Weinberg equilibrium and
selective neutrality of the markers is employed [127]. Ultimate success of R“PDs is shown in
the increase of molecular markers which require small amount of DN“ and no need for
sequencing, except of having all prerequisites for PCR conditions [126]. DN“ fragments having
sequence of about 10 bp are ampliied with artiicial primers by using PCR [128]. R“PDs are
being used vigorously for proiling of genotype of important ield crops, also for mapping for
certain traits in addition to biotic and abiotic stresses. For such studies, R“PD primers show
polymorphism and should be free from palindromic sequences and should have minimum
40% GC contents in the fragments [113].
Many scientists have explored R“PDs in coton for studying diferent aspects like phylogenetic
studies, genetic diversity, and CLCuV disease screening [111, 112, 128]. R‐6592 and
U”C607500 [113, 129] male sterility and fertility restorer traits can be improved by using
R“PDs. Lan et al. [130] applied R“PDs for mapping fertility genes that is of immense value
in coton and tagged fertility restorer gene R‐ 9 which may be utilized for productivity
enhancement. Lan et al. [130] conducted phylogenetic studies in coton and argued that this
procedure is helpful and reliable for introgression of desirable traits. R“PDs were used in
coton for comparing coton cultivars resistance to jassids, mites, and aphids [131]. DN“ inger
printing, mapping and genetic diversity has been studied in coton through R“PDs [132 134].
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Noormohammadi et al. [135] screened F2 population of Upland coton and Opal variety by
using 10 homo‐primers and seven hetero‐primers out of 26 R“PDs and found 261 reproducible
bands, with an average of 4.18 [261 bands/17 primers = 15 bands/primer] bands per primer and
22% polymorphism for analyzing genetic resemblance in agronomic traits with 45 (Upland)
and 80% (Opal) polymorphism, respectively. ”y applying agarose gel, multilocus genotyping
can be carried out by staining with ethidium bromide and this facility is available in every lab
working on molecular breeding [136].
R“PDs are often laboratory dependent and require immense care to design protocols for
geting polymorphism. Several factors have been reported to inluence the reproducibility of
R“PD results such as quantity of template DN“, bufers of polymerase, concentration of
magnesium chloride, primer to template ratio, annealing temperature, type or source of DN“
polymerase, and brand of thermal cycler [137]. R“PDs also fail to discriminate between
homozygotes and heterozygotes and complication of expressing Mendelian ratio of loci [138].
. . . Intersimple sequence repeat
Modiications of microsatellites, which utilize microsatellites‐complementary primers,
overcome the need for lanking fragment information [139]. Polymorphism is revealed among
simple sequence repeat (SSR) markers by using primer (16 25 bp) adjacent to a single SSR and
annealing occur at either ends [139]. ISSR utilizes microsatellites as oligonucleotides in a PCR
reaction to amplify inter simple sequence repeats for desired DN“. ISSRs utilize SSRs repeats
dinucleotide, trinucleotide, and tetranucleotide as oligonucleotide [140]. Usually, ISSR primers
have substantial fragments contrary to R“PD primers, enabling elevated annealing tempera‐
ture, which produce highly polymorphic bands as compared to R“PDs [120, 141]. The
ampliied products can be separated by agarose and polyacrylamide gel due to longer length
ranged from 200 to 2000 bp [139].
ISSR markers have been vastly used in coton improvement, phylogenetic study and for
mapping of germplasm [120, 121]. Parkihya et al. [142] studied genetic diversity among coton
genotypes by using nine ISSR oligonucleotides and detected 86 bands of which 54 bands
exhibited polymorphism of 62.79% having mean of six bands per primer. The PIC ranged from
0.8616 to 0.9090 and genetic similarity ranged from 0.60 to 0.917. Phylogenetic relation was
revealed in 21 coton genotypes by using 12 intersimple sequence repeat primers and observed
49.6% reproducibility [143].
Genetic diversity was studied in coton with 10 ISSR and showed 88.5% polymorphism [144].
Liu and Wendel [145] showed that ISSR can be designed with low cost. Genetic diversity
observed in genepool comprised of wild species and elite lines through SSRs and ISSR [146].
They observed 173 alleles having mean 3.93 alleles per locus by analyzing 39 SSRs and 5 ISSR
markers which produced 89.6% reproducible bands. “mong genotypes variation ranged from
0.04 to 0.58 while in diploid and tetraploid species it was 0.23 0.57%. Similar to R“PD, there
may be some fragments with the same mobility originate from non‐homologous regions [120].
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. . . Sequence characterized ampliied region
R“PDs have more demerits of polymorphism as compared to other PCR‐based markers which
are used for analyzing a large number of individuals with low cost. This problem was overcome
by using sequence characterized ampliied region. PCR assay uses couple of distinct oligonu‐
cleotides for DN“ sequence at a speciic locus [147]; oligonucleotides might be having a high‐
copy, dispersed fragment within polymorphic loci. “fter sequencing the two ends of the two
reproducible DN“ fragments, one can develop two SC“R markers. SC“R 4311920 can be used
in M“S program for screening genotypes with iber strength. ”y using SC“R, codominance
is produced [148].
These markers have been used in genetic analysis and used for molecular breeding [149, 150].
Extended sequence speciicity of primers in SC“Rs results in higher reproducibility than
R“PDs [151]. SC“R is widely used among researchers for mapping studies within closely
related species [152]. SC“Rs are more authentic for M“S after conversion of DN“ markers.
SC“R markers are cost efective and highly polymorphic which make them suitable to be used
for evaluating large number of mapping populations in coton [152]. QTLs for leaf traits were
observed [153].
. . . Sequence‐tagged site
Oslen et al. [154] developed sequence tag sites (STS) through observing impact of the PCR on
human genome research, and argued that single‐copy DN“ sequences of known map location
could serve as markers for genetic and physical mapping of genes along the chromosome. STS
marker allows the utilization of PCR with speciic primers which produces one oligonucleotide
connected to the trait of interest. In order to utilize STS for molecular breeding, RFLP, “FLP,
and R“PDs are usually converted into STS [155]. Thus, in a broad sense, STS include the
markers such as microsatellites (SSRs), SC“Rs, and ISSRs mentioned above. ”ackcross
breeding population was developed [(”416R × “rk8518) × “rk8518] and used for identiication
of STS markers related to fertility [155]. Tetraploid and diploid species were involved and
artiicial hybrids created by colchiploidy.
R“PDs such as U”C1471400, U”C607500, U”C979700, and U”C169800 loci were associat‐
ed to productivity restoration, and it was veriied that U”C607500 is having enormous val‐
ue for pyramiding genes to be used in molecular breeding [129]. Linkage maps were
developed by using STS for diploid and tetraploid (“tDt) Gossypium genomes [156]. Genet‐
ic map composed of 763 loci at 1.96 cM (approximately 500 kb) intervals detected by 662
probes (D), and 2584 loci at 1.72 cM (approximately 600 kb) intervals based on 2007
probes (“tDt).
Several coton breeders have used STS markers for identiication of male restorer parental
lines for hybrid coton [129] who mapped coton genotypes by using backcross inbred lines
(”ILs) and RIL populations with informative primers, and detected 21 and 7 polymorphic
STS markers in ”ILs and RIL populations, respectively. Twelve STS markers were mapped
in ”IL population, and four of them were located along with resistance gene analog‐ampli‐
ied fragment length polymorphism (RG“‐“FLP) markers on the same chromosome. Im‐
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portantly, two were mapped on chromosome c 4, lanking two main‐efect QTLs, which
were previously detected. These STS markers should be useful for high‐throughput geno‐
typing, gene mapping, and M“S for disease resistance including Verticillium wilt resistance
in coton.
. . . Simple sequence repeats
Tandem repeats composed of several to over hundred repeats of one to four nucleotide motifs
are found in all eukaryotic genomes. These repeats are designated as (“““C)n, here n
represents number of tandem repeats. The lanking sequences of simple sequence repeats
(SSRs) are used for the development of oligonucleotides [118]. Tandem repeats induce
variability, which evolve polymorphism of diferent size due to slipped strand arise because
of mispairing occurs during DN“ replication [118], variation in size of PCR ampliication/
products induce polymorphism which can be separated by electrophoresis.
Kinship studies are conducted by employing SSR markers assess the extent of variation
[119]. Vos et al. [157] used agarose and polyacrylamide gel for the identiication of SSRs hav‐
ing codominance nature like “FLP. “kkaya et al. [158] stated that genetic mapping is on fast
track due to the use of SSRs in self‐pollinated crops where these markers are of great inter‐
est for breeders [159, 160]. SSRs are mostly codominant, and are indeed excellent for study‐
ing of population genetics and mapping [161 163]. The use of luorescent primers in
combination with automatic capillary or gel‐based DN“ sequencers has got its way in most
advanced laboratories, and SSRs are also shown to be excellent markers for luorescent tech‐
niques, multiplexing and high‐throughput analysis.
Derived from trispecies hybridization that can be segregated for natural leaf defoliation
trait. This RIL population screened with microsatellite markers, JESPR‐13, JESPR‐153, and
JESPR178 tandem repeats were found to be highly associated to leaf defoliation trait value
[162]. It was found that JESPR178 is closely linked to this trait in coton. It has an immense
importance that gene pyramiding can be accomplished for molecular breeding [164]. QTLs
were tagged using SSRs in the nematode resistance RIL population developed via introgres‐
sion from G. barbadense [165]. In that study, a single marker analysis identiied four major
QTLs located on chromosomes 3, 4, 11, and 17 were identiied to account for 8.0 12.3% of
the phenotypic variance.
Fiber length was increased up to 12 20% in coton by using microsatellites in a population
derived from interspeciic hybridization and loci were discovered for marker‐assisted selec‐
tion [166]. Twenty‐three chromosomes were analyzed by SSRs and found on an average
relative distance of 4.9 cM [167]. Researchers [168 170] have utilized SSR markers for studying
genetic diversity in coton and observed limited genetic variations. Reddy et al. [171] used SSR‐
enriched genomic libraries and identiied 300 SSR markers. Multinational Seed Company has
reported more than 1200 SSRs [172].
“bdurakhmonov et al. [173] conducted genome‐wide association mapping based on linkage
disequilibrium (LD), scanning Upland germplasm consisting of 334 G. hirsutum accessions
collected from Uzbek, Latin “merican, and “ustralian ecotypes. Screening under diferent
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climates and by using mixed linear model involving population kinship and population
structure 12 22 SSR markers were associated with iber length, iber strength, iber ineness
and six other iber quality traits. Mei et al. [78] have reported 145 SSR polymorphic markers
for yield and yield components in coton by screening germplasm of 358 upland coton
varieties. Cao et al. [174] assessed genetic stock iber quality and reported 97 polymorphic SSR
markers by using LD.
”olek et al. [57] used SSR markers for verticilium resistance in coton by using F2 population;
255 SSRs were screened over bulks constituted by 10 resistant, and 10 susceptible progenies.
QTLs were tagged by using 60 polymorphic markers. Genetic map produced 11 linkage groups
having 15.17 cM inter‐locus distance and spanning 531 cM.
”ackcross inbred lines used [175] for observing the genetic variation of 446 SSR markers having
relative mean distance of 10 cM interspeciic linkage map and also detected 58 QTLs related
to iber quality and yield components. ”y using SSRs, genetic markers associated to coton
earliness were determined in progeny developed from intra‐hirsutum hybrids [117], and these
markers correspond to bud to lower duration and lower to boll period.
Earliness in coton can be induced by the introgression of QTLS located near to the SSR markers
such as ”NL1044, DPL0209, N“U1004a, N“U 5046, N“U6078, and TM”0481 [117]. F2 progeny
was developed within G. hirsutum for utilizing gene pyramiding in marker‐assisted breeding
to enhance iber quality and agronomic traits of economical value by using SSRs, SR“Ps, EST‐
SSRs, and SSCP‐SNPs [176]. Economically valuable traits were evaluated through construction
of linkage map, segregation patern observed among the traits and 33 QTLs were identiied
[176].
Textile industry entirely depends upon iber with good quality. Marker‐assisted selection
allows developing a cultivar having good iber quality. There are many SSRs which can be
used for fostering the breeding program; for example, lint percentage can be approved by
using TM”0471 and MGHES‐31, TM”0366, ”NL3590, ”NL1395, ”NL1672, ”NL1694, JES‐
PER101, JESPR204, N“U3308, ”NL1672, N“U3308; N“U4024 [168, 177 180]. Genetic base
can be broadened for span length by using ”NL 1395, DC40182, N“U2980, ”NL2752,
N“U2985, N“U1167, N“U1200, N“U2277 [50, 123, 177, 179]. ”NL1122, ”NL1317,
”NL3145, ”NL1521, CIR307, CGR6164, CGR6683, GH454, ”NL3463, JESPER153, DC40182,
N“U 1037, SHIN‐0463, TEM”1618, N“U3736, N“U445, N“U780, N“U1102, N“U1197,
N“U1322, N“U1369, JESPR218, TMD05 can be applied for iber strength enhancement [52,
92, 173, 174, 176, 181 188].
. . . Single nucleotide polymorphism
Single nucleotide polymorphisms (SNPs) manifest alteration in single base. SNPs are the most
frequent occurring variability in the individuals which is found in each 1000 bases [189]. These
are changes in bases from transitions (C/T or G/“) to transversions (C/G, “/T, C/“, or T/G)
while insertions and deletions also induce SNPs which show single base changes. SNPs show
useful allelic variations and have been markers of choice in various genetic studies [190]. Rapid
progress in high‐through put sequencing has allowed discovering SNPs in complex genomes
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with economic value by using genotyping by sequencing [191]. Frelichowski et al. [167]
revealed that reproducibility is a major hindrance for using large number of markers devel‐
oped in G. arboreum, G. raimondii and G. hirsutum [192]. In combination with genome and
expressed sequence tags (ESTs) in model plant species [193], the eiciency of Sanger sequenc‐
ing has been improved to accelerate the identiication of variations at single base pair resolu‐
tion [194].
Genotyping in plant sciences is progressing rapidly because SNPs for observing variation in
a speciic locus are utilized. Moreover, availability of enormous SNPs due to insertions‐
deletions and whole genome genomic studies is laying the corner stone for next generation
sequencing [195]. Developed genomic databases and SNPs information allow evolving SNPs
to an inluential research for related relatives. Owing to most common type of DN“ polymor‐
phism, SNPs are also lexible in the selection of SNP variants at target loci, and they provide
the option to choose from a large number of genome‐wide loci when selecting sets of infor‐
mative markers for speciic germplasm pools [196]. ”reeding programs comprised of genomic
estimated breeding values are highly favored for whole genome techniques additional to
targeted loci [197 199].
Economically important traits from breeding perspective are also investigated through
genome‐wide sequencing [200], saturated mapping of polygenic traits [201], and by using LD‐
mapping [202]. “n et al. [203] studied the expression of R2R3‐MY” transcription factors where
few are expressed during iber initiation and elongation. They observed phylogenetic relation
among R2R3‐MY” genes and published a map by using SNPs in Upland coton. QTLs were
mapped in population derived from intra‐hirsutum and interspeciic (G. hirsutum × G.
barbadense) [204]. Researchers [204] have collected all published QTL data. QTLs were
identiied for seed, yield and iber quality by using two populations through meta‐analysis.
QTLs connected to biotic and abiotic stressed were also detected. However, the development
of high‐throughput genotyping platforms for large numbers (thousands to millions) of SNPs
has proved to be relatively time‐consuming and costly.
Deynze et al. [205] reported more than 200 loci in G. hirsutum breeding germplasm, which
were genetically mapped on mapping population derived from TM‐1 and 3‐79. Genepool
comprised of 24‐accessions derived from 8‐parental lines of mapping populations of six cot‐
ton species and16 promising coton strains used for genotyping. “s a whole more than 1000
SNPs were polymorphic among G. hirsutum and G. barbadense were developed from 270
loci and 290 indels from 92 loci. [205]. Roche 454 pyrosequencing platform in four allotetra‐
ploid cotons using reduced representation library (RRL) have helped to map a large num‐
ber of SNPs [206]. The conversion rate of SNPs using K“SPar assay was about 35.8%. Three
hundred and sixty‐seven SNP markers were used for linkage map construction, which span‐
ned to 1688 cM. High‐resolution maps can be formed rapidly by utilizing parallel sequenc‐
ing methods to determine the reads in resequencing. Paciic ”iosciences technologies used
for long reads [207] while Illumina and Ion Torrent are applied in sequencing for geting
short reads [196].
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. Mapping populations
The group of plants, which is used for screening of molecular markers and segregated for the
trait of interest, is designated as mapping population. From commercial point of view, such
populations are developed from within species and can be also developed between diferent
species for creating desirable variation. Polymorphism is compulsory in the progenitors for
required trait [208]. The exchange of chromosome fragments during crossing over produces
recombination, which provides the basis for developing linkage maps [59]. Populations are
required for creating genetic maps in order to locate the quantitative trait loci from economical
point of view.
Mapping populations can be exempliied by F2, backcross, recombinant inbred lines (RILs),
doubled haploid lines (DHL), F2‐derived F3 (F2:F3) populations, and near‐isogenic lines (NILs).
F1 is produced by seling two parents, having extreme properties for trait of interest that show
a signiicant polymorphism for whichever type of loci are scored. Mostly, this population is
used for genetic mapping as it requires less time for development. However, there are some
drawbacks for this population, most important of which is the fact that it is not stable.
Qualitative and quantitative traits in coton have been mapped by using F2 [70, 74, 81, 86].
”ackcross (”C) population is developed by crossing a genotype with an elite cultivar, which
is deicient for a single gene or QTL [67]. “ concept of ”C population was developed in 1922
and widely applied in plant breeding programs till 1960 [209]. ”ackcross population has been
used for linkage mapping in coton for improving various traits [129, 155 156].
Near isogenic lines(NILs) can be developed either by using seling until purity is “chieved;
for all traits with wide variation of the trait of interest among NILs or by hybridizing the donor
parent to the F1 plants and choosing the desired trait [63]. NILs are of high importance for
genetic studies as they are stable like RILs. Researchers [210] used NILs for observing QTLs
related to yield and drought related traits. They evaluated that NILs can be used for evaluating
drought and can be used for M“S. Essenberg et al. [116] developed NILs in coton and mapped
bacterial blight resistance. They revealed that lines having “cala‐44 in their parentage are
showing dominance to bacterial blight.
Recombinant inbred lines (RIL) are stable and are developed by using single seed descent
method from irst ilial generation. It continues until homozygosity is obtained in the individ‐
uals. RILs are permanent and can be screened at multiple locations for desired traits. Each
strain is homozygous and stable in the RIL population. Each cycle of seling results in enhanced
recombination frequency and these populations are highly suitable for saturated mapping
[129]. Moreover, for genetic mapping in coton this population has been utilized for various
traits including nematode resistance [165], iber quality improvement [166], and verticilium
resistance [175]. One of the drawbacks of this population is long duration for development in
which segregation bias can occur due to removal of some genotypes after seling. “nother
disadvantage of using these populations is that major QTLs are having a masking efect, and
multiple QTLs are having epistatic efects.
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Nested association mapping (N“M) [188] is designed for precise identiication of QTLs [177].
Economically valued traits related to yield and subsequently to textile sector can be eiciently
studied through developing new populations like N“M. N“M populations potentially
address the limitations of conventional mapping populations
. Some applications of markers in breeding schemes
. . Marker‐assisted backcrossing
The simplest, most widely used, and the most eicient form of M“S is M“”. In this form, two
parents are used for the development; one is donor parent having trait of interest for
transferring the targeted gene/loci and the other is recipient parent which is lacking gene.
Parents are hybridized and F1 is developed. Marker‐assisted backcrossing relies upon the
presence of a molecular marker associated with the trait, instead of targeting the expression
of phenotypic value in traditional breeding. F1 is planted for conirming the marker loci at
initial stages of development, and pure F1 is hybridized to recurrent parent. Markers are
evaluated among individuals at the initial development stages of ”CF1 and hybridized to
recurrent parent having alternative alleles. ”C1 individuals show segregation frequency of F1
population gametes as two genotypes are involved in this population. Highly eicient map is
constructed by using this population in contrast to F2 population. This population is mostly
used for overcoming hybrid in viability and hybrid breakdown in interspeciic crosses [129].
This process is continued until three to four ilial generation for stabilizing the marker and its
associated trait of interest. M“” population has been utilized for observing traits of interest
through quantitative trait loci [115].
. . Pedigree selection
”reeding techniques within the two cultivated tetraploid species rely on crossing and selection
of traits using pedigree and recurrent selection methods. Promising genotypes having
desirable traits can be developed using M“S and can be combined into a pedigree‐based
selection. Mostly, the eiciency of M“S was investigated using two populations from pedigree
selection, and modiied backcrossing pyramiding has been developed [211]. The selection
eiciency for the iber strength was greatly increased when QTLfs‐1 was selected simultane‐
ously with two molecular markers with known genetic distance [211].
. . Marker‐assisted recurrent selection MARS
Molecular markers should be applied for plant improvement in conjunction with the latest
breeding methodologies. Marker‐assisted recurrent selection ofers an opportunity to get
maximum output from a recurrent selection [212]; and it is used for introgression of multiple
genes.
Quantitative traits can be enhanced eiciently by using M“RS, which allows seling and
genotyping within same cropping season in one cycle of selection. The increase in genetic
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gain was doubled from M“RS in some populations as compared to phenotypic selection
[213]. In coton, resistance to “merican bollworm was achieved by using marker‐assisted
recurrent selection; they revealed highly signiicant diferences in individuals studied by
M“RS for this insect resistance [214].
. Gene pyramiding for MAS
Gene pyramiding has been widely used for combining number of genes especially disease
resistance genes for speciic races of a pathogen. Vertical resistance for diferent strains of
pathogens is done by involving multiple strains. It is also done by molecular breeding
because breeding for resistance is extremely diicult to achieve using conventional methods.
Gutiérrez et al. [215] have used this technique for nematode resistance in coton while [155]
they applied sequence tag sites and screened STS markers associated with fertility restoring
genes in coton.
. Next‐generation sequencing NGS
SNP genotyping with latest high‐throughput sequencing has the potential to speed up the
breeding programs [216]. New DN“ sequencing technologies have made it possible for the
breeders and investigators to perform a genome analysis not only more rapidly but also less
expensively [179]. High‐throughput bioinformatics assist to identify large number of nucleo‐
tides per run [217]. Researchers have developed a lot of NGS methods with success in diverse
platforms, which include Roche 454 FLX Titanium [218 220], Illumina MiSeq and HiSeq2500
[221], Ion Torrent PGM [222]. Genomic research contributes immensely to plant and animal
sciences thanks to the advances in sequencing techniques [180, 221, 223 228]. The ultimate aim
of all these techniques is to discover an authentic marker that could be used for sequencing in
M“S with economical beneits [229 230].
Polyploidy is the main hindrance for isolation of useful SNPs in coton because it produces
homeologous and paralogous sequence variants which are combined together in allelic
variations among cultivars [231 232]. Two cultivated tetraploids species were screened for the
development of genomic SNPs through NGS by using reduced representation library obtained
from Roche 454 pyrosequencing [206]. Competitive allele‐speciic PCR (K“SPar) showed
35.8% validity of SNPs and developed the genetic map of G. hirsutm via 367 SNP markers which
spanned to 1688 cM.
Gore et al. [233] developed a linkage map in a RIL population derived from intra‐hirsutum cv.
TM‐1, and NM24016. The genetic map covered about 50% of the G. hirsutum genome which
consisted of 429 SSRs and 412 SNPs. They also tagged 10 QTLs related to iber quality, which
provided a unique resource for mapping. ”efore “fymetrix became commercially available,
Gene Chip coton genome array consisting of 239,777 probe sets that represent 21,485 coton
transcripts has been developed [234, 235]. Sequences from Genome database, dbEST and
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RefSeq were used for the development of Chip which promises to be an excellent source for
genomics.
. Genotyping by sequencing
In agricultural sciences, the discovery of reliable and true SNPs is compulsory for knowing
about the utilization and importance of particular sequences. Molecular breeding tools can be
applied to explore germplasm without available genomic data through genotyping by
sequencing (G”S) methodology. G”S permits researchers to analyze complex genomes of
polyploid species eiciently at low cost and it has been widely used due to the latest devel‐
opments in high‐throughput sequencing [191]. Reduced representative libraries are developed
by using endonucleases [55, 178, 236]. Single nucleotide polymorphism is discovered for
genomic studies [237]. Genomic techniques; genome‐wide association study (GW“S), genomic
diversity, genetic linkage analysis, molecular marker discovery ofer to screen genotypes upon
genotypic basis for traits of interest through G”S. Genotyping and reproducibility of markers
are performed in a single step through G”S and SNPs are developed [238].
G”S‐based sequencing data are used for developing genetic map and tagging markers with
quantitative traits in populations derived from diferent ways, i.e., ilial generations, RILs, etc.
and germplasm collections [218]. G”S approach has been used eiciently for genetic analysis
and marker development of rapeseed, lupin, letuce, switchgrass, soybean, maize, and coton
[38, 219, 222 224, 233, 239].
The merits of G”S over existing marker development methodologies include availability of
large number of markers, fast screening of populations composed of more number of indi‐
viduals, diverse genotyping systems to tackle multiple traits, and more precise SNPs discovery
and validity due to availability of high‐throughput sequencing data [216]. Recently, G”S
approach has been used to identify SNPs in the collections of RILs of wheat and to map various
traits useful for breeding programs [55]. It is needed that eforts should be made to develop
strategies for geting the beneits of NGS and advanced genotyping from breeder perspectives
[196]. G”S protocol of Poland et al. [236] likely is needed to maximize the cost‐efective
concurrent discovery and genotyping of SNPs within coton populations.
“lthough very eicient and productive in terms of achieving the desired goals, there are some
drawbacks in G”S as well. G”S has incapability to assign true alleles of each locus in polyploids
as compared to other techniques. “s exempliied, Huang et al. [178] used RILs and biparental
populations for assessing the utility of G”S in hexaploid oat. They observed that data analysis
algorithm factors involved in SNP discovery, developed G”S derived loci description by
forming two bioinformatics worklow. Its genetic map spans to 45,117 loci, which will be a
source of further genetic studies [178].
Islam et al. [240] used G”S with two diferent approaches in cultivated coton germplasm
consisting of 11 diverse cultivars and their random‐mated RILs. “uthors have discovered a
large set of polymorphic SNPs with broad applicability. They identiied 4441 and 1176
Molecular Breeding of Cotton
http://dx.doi.org/10.5772/64593
polymorphic SNPs with minor allele frequency of ≥0.1. The utility of developed SNP markers
were conirmed using SNPs in 154 Upland coton accessions with high genetic diversity.
. Association mapping
Genome‐wide association study is used for developing highly saturated maps in coton
germplasm [241]. This technique allows detecting association among various markers and
traits through assessing of the genetic diversity of required traits [242]. Linkage disequilibrium‐
based mapping (LD‐mapping) is the advanced tool to study complex traits governed by many
genes. LD‐mapping has been successfully used in self‐pollinated plants [243]. Microsatellites
were screened in germplasm consisting of varieties at diferent locations to tag yield and iber
quality QTLs [202]. QTLs mapped for yield and iber quality traits will serve as a reliable source
to determine the diversity within the species and will contribute a lot in M“S [202]. In contrast
to biparental populations, association mapping fosters molecular breeding because a vast
genetic diversity is present in germplasms due to diverse sources [173]. Several protocols have
been developed including complexity reduction of polymorphic sequences (CRoPS) [244],
restriction site associated DN“ (R“D) [216], G”S [195], and sequence‐based genotyping
(S”G) [245, 246] for genome analysis. Of all protocols, LD‐mapping is on the top thanks to
innovations done for high resolution. “ssociation mapping is an authentic way for molecular
tagging as it allows the screening of quantitative traits of value in a precise way [247]. Genome‐
wide association makes it possible to detect association among various markers and traits.
“bdurakhmonov et al. [46] used LD‐mapping in a germplasm collection, which included
photoperiodic lines. Simple sequence repeats were used for assessing the extent of LD in coton
and the major iber quality QTLs were tagged using mixed linear model.
Nested association mapping is also being used for identiication of suitable SSRs in a N“M
population derived from 20 diverse genotypes of G. hirsutum with Namangan‐77. SSR marker
screening for development of highly saturated map through N“M F2:3 populations for traits
of immense value in coton is underway [188].
. Public data resources
Sequenced genomic information allows breeders to analyze the genetic variation [248]. Major
databases, which serve as a foundation, include CotonGen [58], Comparative Evolutionary
Genomics of Coton [249], National Center for ”iotechnology Information [250] for Express
sequence tags resource, TropGENE Database [251], the Coton Diversity Database [252] and
”“CMan resources at Plant Genome Mapping Laboratory [253]. These resources provide
genomic and heredity data of the coton germplasm, QTLs tagged to loci and highly saturated
linkage maps.
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. Targeting‐induced local lesions in genomes TILLINGS
Phenotypic variation in plant genomes is produced by variation in DN“ bases, which can be
induced naturally and/or using diferent chemicals [254]. The targeting‐induced local lesions
in genomes (TILLING) technique allows determining an allelic variation precisely in a single‐
base pair for the targeted gene. Chemical treatments have been applied to generate SNP
mutations. Point mutations, which are useful from breeder s perspective, can be detected by
TILLING and ECOTILLING techniques [255]. The mutagens used for induction of point
mutation are highly selective and optimal concentration can spontaneously produce single
base alternations at a high frequency in TILLING.
Knock‐out population is developed by treating the seed with chemicals, inducing change in
DN“ sequence [256]. “uld et al. [257] used TILLING in G. arboreum and demonstrated the
applicability of this technique in coton. The ultimate success to produce large number of
sequence variations of target genome depends upon duration of application, relative capability
of ethyl methane sulfonate (EMS), and γ‐rays [53]. “slam et al. [53] screened three Gossypium
sp. (G. hirsutum, G. barbadense and G. arboreum) and constructed a kill curve. They observed the
impact of diferent mutagens (EMS and γ‐rays) consisting of eight diferent concentrations of
EMS (0.1 0.8%) and two levels of γ‐rays (100 800 Gy). The genotypes of each species were
evaluated with morphological parameters emergence and plant height, and yield traits
(number of bolls per plant, boll weight, lint yield and lint percentage). For reverse and forward
genetics point of view, viable accessions were selected from mutagenized genotypes. They
revealed that EMS showed signiicantly high mutation rate than γ‐rays.
There are many software tools which help to observe the bases variation; for instance, the
method that determines whether a change occurs in an amino acid hampering codon is named
conservation‐based SIFT (sorting intolerant from tolerant) [258]. Taylor [259] described that
any alternation of a gene can be detected by P“RSESNP (for Project “ligned Related Sequences
and Evaluate SNPs [260]; graphs show the changes in sequence by using precise co‐segregating
information, positioning of coding/and noncoding regions and reference DN“ sequence.
. Conclusions
Developing reliable markers, which will work in diferent populations and utilized in the
breeding to enhance selection eiciency, is a very important step for breeding. Markers should
allow desired genotype selection because of their tight linkage to the trait of interest. On the
other hand, emerging technologies like high‐throughput marker systems and marker‐based
selection methodologies have been developed, and are currently being used eiciently in
coton breeding. It is also promising that some economically important traits like iber quality,
yield, Verticillium wilt resistance, coton leaf curl virus, drought tolerance, nematode resistance
can be enhanced by using M“S. Genetic diversity can also be evaluated by using DN“ markers
before starting breeding program. Tremendous eforts have been carried for studying genetic
diversity from genotypic and phenotypic aspects in germplasm accessions of coton. Many
Molecular Breeding of Cotton
http://dx.doi.org/10.5772/64593
QTLs related to economical traits have been discovered. It is an emerging concern that eforts
should be made for the utilization of molecular breeding methodologies to enhance coton
productivity, which can be enhanced through the recent developments in NGS. Moreover,
highly saturated maps are useful for determining genetic manipulations from heredity
perspectives, and SNPs are the best for this purpose. These markers along with QTLs provide
innovative tools in the coton genomics era.
Author details
Yuksel ”olek1, Khezir Hayat1, “dem ”ardak1* and Muhammad Tehseen “zhar2
*“ddress all correspondence to:
[email protected]
1 Faculty of “griculture, “gricultural ”iotechnology Department, Kahramanmaras Sutcu
Imam University, Kahramanmaras, Turkey
2 Department of Plant ”reeding and Genetics, University of “griculture, Faisalabad, Paki‐
stan
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