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Cotton evolution at a glance
- 1. COTTON EVOLUTION : AT A GLANCE
- 2. • Introduction • History and Evolution • Morphological diversity in cultivated species • Case Studies (C.S.) • Fiber evolution under domestication • Conclusion 2 Content
- 3. 3 Introduction Karpas/Karpasi - Sanskrit Karposas –Greek, Carbasus- Latin Cutan – Arbi Cotton – English Key role in human civilization Vital role in national economy White gold Dominancy over synthetic fibers Tree of wool Eight groups of diploid 2n = 26 Subkingdom - Tracheobionta Superdivision - Spermatophyta Division - Magnoliophyta Class - Magnoliopsida Subclass - Dilleniidae Order - Malvales Family - Malvaceae Kingdom - Plantae Vascular Plant Seeded Plant Flowering Plant Dicot Plant Mallow plants Fig. 1: The Clade tree for the Gossypium spp. Genus - Gossypium
- 4. 4 World's major cotton producing countries Year 2012-13 Source: United States Department of Agriculture, Foreign Agricultural Service Country Area (M ha.) Production (M bales of 480 lb) Productivity (kg/ha) India 11.70 25.50 475 China 5.28 34.00 1403 United States 3.82 17.01 971 Pakistan 3.00 9.60 697 Brazil 1.00 6.50 1415 Uzbekistan 1.29 4.30 729 Australia 0.45 4.20 2055 Turkmenistan 0.60 1.50 544 Argentina 0.36 0.80 484 Turkey 0.40 2.60 1415 World 34.13 118.95 759
- 5. India has the unique distinction of being the only country in the world to cultivate all four cultivable Gossypium species… Cotton in IndiaCotton in IndiaCotton in IndiaCotton in India Year 2012-13 AICCIP Project – Annual Report Leading States Maharashtra – Area (41 lakh ha) Tamilnadu – Productivity (831 kg/ha) Area (M ha) 11.77 Production (M bales) 25.61 Productivity (kg/ha) 486 World Ranking Area 1st Production 2nd Productivity 34th 5 Gujarat - Production (85 lb)
- 6. 6 The Gossypiae tribe, distinguished from other Malvaceae : The embryo, wood and seed coat anatomy, and by the presence of the punctae or lysigenous cavities (“gossypol glands”) that are widely distributed throughout the plant body Gossypium : An undivided style, coriacious capsule containing several seeds per locule, a somatic chromosome number of 26, and the presence of three foliaceous involucellar bracts subtending each flower Plant habit: Fire-adapted, herbaceous perennials in northwest Australia to small trees in southwest Mexico that “escape” the dry season by dropping their leaves. Corolla colors: rainbow of mauves and pinks, whites and pale yellows and even a deep sulphur-yellow Seed: Nearly glabrous to short, stiff, dense brown hairs that aid in wind- dispersal to long, fine white fibers that characterize highly improved forms of the four cultivated species. Seeds that produce fat bodies to facilitate ant-dispersal Gossypioides – Kokia represents the example of long-distance salt-water dispersal in the tribe DIVERSIFICATION OF THE COTTON TRIBE
- 7. 7 Fig. 1. The range of morphological diversity exhibited by members of the Gossypium genus
- 8. 8 History and Evolution The oldest Malvacean pollen is from the Eocene (38 – 45 million years before present - mybp) in South America and Australia and from the Oligocene (25 - 38 mybp) in Africa It suggests that the Malvaceae originated during the first third of the Tertiary (65-2.6 mya) and that by approximately 30 mybp it had achieved a world-wide distribution (Seelanan et. al. 1997) Fig. 2: Phylogenetic relationships in the cotton tribe (Gossypiae) as inferred from molecular sequence data The tribe Gossipiae is approximately 20 million years old The Lebronnecia/Hampea/Thespesia separated approximately 15 mybp. Gossypium is inferred to have branched off from its closest relatives (Kokia and Gossypioides) approximately 12.5 mybp Successful grafts could be made between Kokia rockii and Gossypioides kirkii
- 9. 9 Fig. 3: Evolutionary History of Gossypium, as inferred from multiple molecular phylogenetic data sets. Genus Gossypium is differentiated cytogenetically into eight ‘‘genome groups’’ (designated ‘‘A’’ through ‘‘G’’, and ‘‘K’’) They differ in DNA content and chromosome size but not in chromosome numbers Three lineages of diploid species are recognized; the African/Arabian, the Australian and the American
- 10. 10 Table 1. Gossypium Species, their Genomes and Distribution No. Species Genome Distribution No. Species Genome Distribution Diploid (2n = 26) 26 G. thurberi D1 America 1 G. africanum A Africa 27 G. armourianum D2-1 America 2 G. herbaceum (Cult.) A1 Afghanistan 28 G. harknessii D2-2 America 3 G. arboreum (Cult.) A2 Indo-Burma 29 G. klotzschianum D3-k America 4 G. anomalum B1 Africa 30 G. davidsonii D3-d America 5 G. triphyllum B2 Africa 31 G. aridum D4 America 6 G. barbosanum B3 Cape Verede 32 G. raimondii D5 America 7 G. capitis-viridis B4 Cape Verede 33 G. gossypioides D6 America 8 G. sturtianum C1 Australia 34 G. lobatum D7 America 9 G. nandewarense C1-n Australia 35 G. trilobum D8 America 10 G. robinsonii C2 Australia 36 G. laxum D9 America 11 G. australe C3 Australia 37 G. turneri “D” America 12 G. pilosum K Australia 38 G. stocksii E1 Arabia 13 G. costulatum K Australia 39 G. somalense E2 Arabia 14 G. populifolium K Australia 40 G. areysianum E3 Arabia 15 G. cunninghamii K Australia 41 G. incanum E4 Arabia 16 G. pulchellum K Australia 42 G. longicalyx F1 Africa 17 G. nelsonii K Australia 43 G. bickii G1 Australia 18 G. enthyle K Australia Allotetraploid (2n = 52) 19 G. londonderriense K Australia 44 G. hirsutum (Cult.) (AD)1 America 20 G. marchantii K Australia 45 G. barbadense (Cult.) (AD)2 America 21 G. exiguum K Australia 46 G. tomentosum (AD)3 Hawai 22 G. rotundifolium K Australia 47 G. lanceolatum (AD) America 23 G. fryxellii K Australia 48 G. mustelinum (AD) America Source: CICR Tech. Bull. No. 5
- 11. 11 Fig. 4: Genome biogeography of Gossypium D genome diploids C , G & K genome diploids B & F genome diploids E genome diploids Singh (1999) Asia Arizona Mexico USA Galapogos Ilands sAfrica nAfrica NE. Africa Australia Hawaii Peru Ecuador 11 A genome
- 12. 12 African-Asian SpeciesAfrican-Asian Species Subgenus : GossypiumSubgenus : Gossypium Section : Gossypium Section : Serrata Genome ? Subsection : Gossypium A genome Subsection : Pseudopambak E genome 1 species 7 species 1 species 3 species 2 species G. anomalum germplasm pool for bacterial blight resistance Cytologically unique, and unusual for its ecological adapatation to mesic environments Seven species adapted to the extremely arid habitats of eastern Africa, the south eastern tip of Arabia, and the Sind in Pakistan This poorly known species may be distinguished from other Gossypium species by its serrate leaves Subsection : longicalyx F genome Subsection : Anomala B genome An A-genome progenitor served as the female parent in the hybridization event of allopolyploidization G. arboreum, G. herbaceum small scale cultivation Germplasm pool for several agronomically desirable traits.
- 13. 13 G. anomalum G. capitis-viridis G. triphyllum G. longicalyx African-Asian species
- 14. 14 Australian SpeciesAustralian SpeciesSubgenus : Sturtia 13 species 3 species 2 species section : Hibiscoidea G genome section : Grandicalyx K genome section : Sturtia C genome Do not deposit terpernoid aldehydes (“gossypol”) in the seeds G. Sturtianum “Sturt’s Desert Rose”, the floral emblem of the NorthernTerritory Basal species in the Australian Gossypium lineage Unusual herbaceous perennials, in the Kimberley region Thick root- stock from which they resprout following fire or seasonal drought Eliosomes on nearly hairless seeds to facilitate ant dispersal. This section has the largest genome in the genus Do not deposit gossypol in the seeds Possess stiff spreading seed hairs that allow the seed to “climb” out of the capsule, and are the only species that are wind dispersed
- 15. 15 G1-n Australia G. nandenwarense G. bickii G. sturtianum G. australe Austraian species 15
- 16. 16 American Diploid SpeciesAmerican Diploid Species Subgenus : Houzingenia D genome Subgenus : Houzingenia D genome Section : Houzingenia Subsection : Houzingenia Subsection : Integrifolia 2 species 2 species Subsection : Caducibracteolata Section : Erioxylum Subsection : Erioxylum Subsection : Selera Subsection : Austroamericana 4 species 3 species 1 species 1 species Large shrubs and small trees G. trilobum, source of CMS and restorer factor Tolerate mild frost via defoliation Interspecific hybrids are embryo lethal Calciphiles, found in arid habitats G. armourianum germplasm pool for bacterial blight resistance gene G. harknessii source of CMS and restorer factors. Unique flowering phenology, At the height of the dry season, while leafless, the plants flower and fruit. After the fruit mature the plants remain dormant until returning the rains stimulate new vegetative growh. G. gossypioides the only diploid species that shows evidence of the original A X D hybridization. G. raimondi having the genome most similar to the D-subgenome of allotetraploid, serve as the model of D genome diploid parent
- 17. American species G. nandenwarense G. harknessi G. thurberi G. klotzchianum G. harknessi 17
- 18. 18 G. aridum G. raimondii G. gossypoides G. trilobum American species 18
- 20. 20 Table 2. Characters for Breeding Value found in different Species No. Characters for Breeding Value Species I. Donors for Fibre Quality 1 Fibre length G. anomalum, G. stocksii, G. raimondii, G. areysianum, G. longicalyx 2 Fibre strength and elongation G. stocksii, G. areysianum, G. thurberi, G. anomalum, G. sturtianum, G. raimondii, G. longicalyx 3 Fibre fineness G. anomalum, G. raimondii, G. longicalyx 4 Fibre yield G. anomalum, G. sturtianum, G. australe, G. stocksii, G. areysianum 5 High ginning G. australe II. Donors for Resistance to Insect Pests 1 Bollworms G. thurberi, G. anomalum, G. raimondii, G. armourianum, G. somalense 2 Helicoverpa G. somalense 3 Jassids G. anomalum, G. raimondii, G. armourianum, G. tomentosum 4 Whitefly G. armourianum 5 Mites G. anomalum 6 Aphids G. davidsonii III. Donors for Resistance to Diseases 1 Bacterial Blight G. anomalum, G. raimondii, G. armourianum 2 Verticillium Wilt G. hirsutum race mexicanum var. nervosum, G. harknessii 3 Fusarium Wilt G. sturtianum, G. harknessii, G. thurberi 4 Nematode G. darwinii IV. Donors for other Characters 1 Cytoplasmic male sterility G. harknessii, G. aridum, G. trilobum 2 Drought resistance G. aridum, G. darwinii, G. tomentosum, G. stocksii, G. areysianum, G. anomalum, G. australe, G. harknessii, G. raimondii 3 Frost resistance G. thurberi 4 Delayed morphogenesis of gossypol gland G. australe, G. bickii Source: CICR Tech. Bull. No. 5
- 21. Hutchinson et al. (1947), indicated that allopolyploid cotton first formed in agricultural times, perhaps within the last six millennia, following human-mediated intercontinental transfer of a cultivated A-genome diploid. Endrizzi et al. (1989) argued for a probable Miocene origin (5–18 mya), based on thermal stability measurements in inter-specific DNA hybridization experiments. Phillips (1963), indicated that polyploid cotton originated “in geologically recent times, probably since the start of the Pleistocene (0-5 mya).” According to the hypothesis, hybridization and polyploidization took place prior to the separation of the parental A- and D-genome lineages, which subsequently drifted apart as a consequence of plate tectonic movements. Under this scenario, then, allopolyploids originated prior to the rifting of the South American and African continents, in the Cretaceous (65 mya) or perhaps the early Tertiary (2.5-65 mya). ORIGIN OF THE ALLOPOLYPLOIDS TIME OF FORMATION 21
- 22. PARENTAGE OF THE ALLOPOLYPLOIDS Stephens (1944), compared allometric patterns of leaf development in intergenomic hybrids and stated that “either (G. klotzschianum, its close relative G. davidsonii, or G. raimondii) in combination with G. arboreum would produce a hybrid showing considerable similarity to present-day New World cottons.” Hutchinson et al. (1945) indicated G. raimondii as the D-genome donor on comparative analyses of morphological traits in synthetic A x D amphiploids, and from observations of lint characteristics and vigor of intergenomic hybrids Fryxell (1965), indicated G. raimondii as the D-genome donor based on observations of the lint characteristics of diploid and wild polyploid species. Hutchinson et al. (1947) reported on the basis of multivalent frequencies in synthetic allopolyploid x D-genome that the G. raimondii as closer to the D-genome than the other species tested. Gerstel (1958), studied multivalent frequencies in hexaploids involving both of the two extant A-genome species (G. arboreum and G. herbaceum ) to argue that G. herbaceum was more closely related to the A-genome of the natural allopolyploids. All allopolyploids contain an Old World (A genome) chloroplast genome, indicating that the seed parent in the initial hybridization event was an African or Asian A- genome taxon. Genomes of the only two A-genome species, G. arboreum and G. herbaceum, differ from the A sub-genome of allopolyploid cotton by three and two reciprocal chromosomal arm translocations, respectively suggesting that G. herbaceum more closely resembles the A-genome donor than G. arboreum. 22
- 23. 23 Fig. 5 - Evolutionary framework of Gossypium allotetraploids Diploid Ancestor (6-11 million year ago) G. raimondiiG. herbaceum AA Old World N=13 (Big chormosomes) DD New World N=13 (Small chormosomes) Polyploidization (1-2 million years ago) AADD New World Tetraploid N=26 (13 big + 13 small) G. mustelinum G. darwanii G. barbadense G. tomentosum G. hirsutum Endrizzi et al. 1985; Wendel 1989
- 24. Morphological diversity in cultivated species Gossypium herbaceum • Leaf lobes: Less deep with shriveled base • Squares: Horizontally extended, round or triangular • Flowers: Yellow with pigmented base • Shorter stamens • Bolls: Round, pointed and smooth surface without blackspots • Lint: White, brown or ash colored 24
- 25. Gossypium arboreum • Leaves: Okra type or Deep leaf lobes • Squares: Vertically extended, triangular, less tips and covers buds and flowers almost completely • Flowers: Yellow, white or reddish colored • Longer stamens • Bolls: Tapering, Black spotted with uneven surface 25
- 26. Gossypium hirsutum • Leaves: Larger, heartshaped or triangular with less deep leaf lobes • Squares: Vertically extended and heartshaped • Shorter stamens • Bolls: Round, pointed and less blackspotted • Lint: White, brown or dark brown 26
- 27. Gossypium barbadense • Leaf lobes: Deep with folded base • Squares: Vertically as well as horizontally extended with heart shape and narrow tips • Longer stamens • Bolls: Longer, wide at base and pointed at tip with rough surface and black spots 27
- 28. Singh (1999) Variability in New world and deshi cotton 28
- 29. 29 Table 3. Variability in qualitative characters of cultivated species of Gossypium Plant parts G. hirsutum G. barbadense G. arboreum G. herbaceum Stem Glandless, red, hairy, short sympodia, zero branch pubescent, dwarf Short branch, dwarf, glandless, smooth Bushy dwarf, glabrous, green Short branch, bushy dwarf Leaf Curly, cup, glandless, okra, super okra, red, pubescent, hairy, smooth mosaic, round, yellow veins, fused veins, nectariless Wrinkled, rugate, okra Crinkled, glabrous, narrow, broad, laciniate, sintle lobed, nectariless, red vein Curly, crumpled, hairy, stellate hairs, glabrous, red Bracts Caduceus, frego, accessory Frego Frego type, entire Multibracteole Flower Cleistogamy, male sterile, indehiscent anthers, open bud, buff pollen, yellow pollen, cream pollen, orange yellow pollen, petal spotted, ghost spotted, club stigma style, sunken stigma, nectariless Cleistogamous, semigametic, fertility enhancer, male sterile, cream, white Petalody, pistillate, sunred spot, thumbnail red, red margin, ghost spot, male sterile, yellow petal, pale and white petals, Chinese yellow petal Spotless, ghost spot, male sterile, yellow petal, pale yellow petal, Chinese yellow and pale petals Bolls Cluster, glandless, hairy Smooth, cluster, glandless Partial dehiscence, few loculed, fused, big long, retentive loculi Closed, big Seed Naked, fuzzy, coloured fuzz Naked Tufted, naked Semi fuzzy Lint Brown, green, khaki, lintless Cream, white Lentless, short, sparse, khaki, white, brown Coloured, lintless, hairy Singh, 1999
- 31. 31 No. Intergenomic Hybrids Univalents Per Cell 1 A X B 2.82 2 A X C 8.50 3 A X D 13.98 4 A X E 17.13 5 B X C 11.17 6 B X D 18.19 7 B X E 22.35 8 C X D 13.10 9 C X E 24.68 10 D X E 25.15 11 D X F 21.60 12 G X C 3.84 C.S.-1: Average univalent frequencies in hybrids of diploid Gossypium spp. •E genome ancient and closest to the ancestral genome of the Gossypium •C and D genomes evolved in intermediate age, where D > C •A and B genomes originated very recently; they are closely related; B > A •F genome is of more recent origin same as A & B genomes •C and G genomes are more related Endrizzi et al. (1985)
- 32. 32 Chromosome No. G. herbaceum G. arboreum Allotetraploids 1 2 3 4 5 1 2 3 4 5 6 7 8 9 10 1 3 2 4 5 6 7 8 9 10 1 2 3 5 4 6 7 9 8 10 1 4 3 3 6 5 4 7 10 9 8 7 10 9 8 3 2 4 5 61 C.S-2: Chromosome end arrangements for the first five chromosomes of the A genomes of G. herbaceum, G. arboreum and Allotetraploids Menzel and Brown (1954)Texas
- 33. 33 C.S.-3: Incipient genome differentiation for D genome chromosomes between G. hirsutum (AD) and G. harknessii (D2-2), G. raimondii (D5) and G. lobatum (D7) • Two type of crosses: 1. ‘A’ genome monosomes (G. hirsutum) X Homozygous translocation lines (G. hirsutum) as Control 2. Homozygous translocation lines (Chromo. # 14, 15, 16, 19 and 20) X harknessii, raimondii and lobatum • Objectives: 1. Total genome affinity (Genome Affinity Index) GAI = Mean No. of groups of paired chromosomes Base chromosome numbers (13) 2. Affinity at specific chromosome regions [Chiasma freq. at (a) Unbroken arm, (b) Translocated region and (e) Interstitial region] Menzel et al. (1978)Florida
- 34. Table 4. Summary of chromosome pairing in triploid hybrids and controls in crosses between A-D homozygous translocation lines and appropriate monosomic lines TT Line D Chromo. Other Parent Mean frequency per cell of Mean pairs D chiasmate arms per PMC GAI I II III AZ-7 H6L - H14L H14 Haplo-H6 D5 12.14 11.99 0.96 25.12 0.996 D7 13.93 11.58 0.63 21.02 0.939 2B-1 H2R - H14R H14 Haplo-H2 D5 12.89 12.68 0.24 24.72 0.993 D2 - 2 13.82 11.89 0.46 21.86 0.950 D7 14.43 11.86 0.27 20.42 0.933 1040 H4R - H15L H15 Haplo-H4 D5 12.41 12.24 0.71 24.32 0.996 D2 - 2 12.42 11.34 1.27 22.64 0.970 D7 14.55 11.48 0.49 19.44 0.920 4672 H1R - H16R H16 Haplo-H1 D5 11.99 11.89 1.06 26.02 0.996 D7 16.63 10.76 0.41 17.30 0.859 E20-7 H3L - H19L H19 Haplo-H3 D5 12.75 12.89 0.15 25.70 1.003 D7 14.21 11.76 0.34 19.64 0.930 10-5ka H4L - H19R H19 Haplo-H4 D5 12.35 12.25 0.72 25.14 0.997 D2 - 2 13.14 11.81 0.70 21.98 0.962 D7 14.79 11.57 0.35 19.70 0.916 4669 H20 Haplo-H1• The order of relationships found was D5 > D2 - 2 > D7 to Dh 34
- 35. 35 Table 5. Chiasma frequencies at 15 specific positions in 2n – 1 and (AD)1 D5 translocation heterozygotes TT Line Cyto-type Chrom- osome Chiasma freq. at the following positions a (unbroken arm) b (translocated region) e (interstitial region) Arm X ma freq. Arm X ma freq. Arm X ma freq. 2B-1 H2-H14 2n – 1 H14 L 0.9803 R 0.0156 R 0.9294 (AD)1 D5 0.9902 0.0032 0.9055 4672 H1-H16 2n – 1 H16 L 1.0000 R 0.9811 R 0.9803 (AD)1 D5 0.9816 0.7938 0.8428 * E20-7 H3-H19 2n – 1 H19 R 0.9782 L 0.0217 L 0.9130 (AD)1 D5 0.9444 0.0111 0.9777 10-5ka H4-H19 2n – 1 H19 L 0.9406 R 0.9809 R 0.6206 (AD)1 D5 0.7200 ** 0.8355 ** 0.6355 4669 H1-H20 2n – 1 H20 L 0.9805 R 0.9559 R 0.0259 (AD)1 D5 0.9818 0.9745 0.0381 • The differentiation was significant at only 3 positions lower than those of controls which confirms the previous results of GAI i.e. D5 > D2 - 2 > D7 to Dh
- 36. 36 Fig. 6 - Model for configurations of marked chromosomes expected in 2(AD)1 and (AD)1 Dx translocation heterozygotes if one of the chromosomes involved in the translocation is in the Dh genome and the other in the Ah genome
- 37. 37 C.S.-4: Incipient genome differentiation for A genome chromosomes in G. hirsutum (AD) and Asiatic diploids The Study: • Chiasma frequencies in triploid hybrids of genome constitution (AD)A involving G. hirsutum (AD), G. herbaceum (A1) and G. arboreum (A2) • Six different translocation lines involving chromosomes 6, 7, 10, 11, 12 and 13 of the Ah genome were used to mark specific chromosome regions in hybrids and controls Objectives: • Has any divergence in meiotic homology occurred between the Ah genome and A1 or A2? • Differentiation is generalized or localized? Menzel et al., 1982Florida
- 38. 38 Table 6. Means and Standard Deviations for the total number of chiasmate II-arm per PMC in A2N, 2(AD)1 tt and Tt controls and in (AD)1 A triploids Other Parent (Species) Statistic G. arboreum (A2) 13 II A2N G. hirsutum (AD)1 TM-1 Z9-9 6L-10R 1052 7R-11R 1043 7L-12R 2785 10R-11R 6-5M 11R-12L TM-1 hirsutum II arms - 25.69 25.77 25.62 25.56 25.75 25.57 S. D. - 0.59 0.43 0.64 0.80 0.56 0.69 A1A herbaceum II arms - 25.17 24.65 24.74 25.27 25.07 25.05 S. D. - 0.91 1.10 1.03 0.93 1.22 1.02 A1J herbaceum II arms - 24.89 24.47 24.26 24.77 25.08 25.10 S. D. - 1.05 1.19 1.37 1.09 0.97 1.00 A2N arboreum II arms 25.35 24.30 23.85 24.11 24.31 24.71 24.33 S. D. 0.79 1.31 1.35 1.48 1.10 1.01 1.31 X X X X • No significant differences of A genome of G. hirsutum with A genomes of G. herbaceum or G. arboreum • Slightly lower frequencies in hybrids as compared to controls Ah slightly differentiated from the A1 and A2 genomes
- 40. 40 C.S.-1: Genetic diversity and evolution of Old World Cultivated Cottons as revealed by Isozyme/Allozyme Analysis Old world cultivated cotton group • G. arboreum • G. herbaceum Methodology: • 103 accessions of G. arboreum + 31 accessions of G. herbaceum subjected to isozyme analysis • 20 enzyme systems examined 13 enzyme systems (19 loci with 42 alleles) found polymorphic (Uniqueness of loci as well as alleles studied) Wendel et al., 1989Iowa
- 41. 41 Table 7. Gene frequencies at 19 polymorphic loci in A genome diploid Gossypium Loci Allele G. arboreum G. herbaceum Loci Allele G. arboreum G. herbaceum 1 Ast2-2 0.00 0.03 11 Pgd1-6 0.15 1.00 4 1.00 0.97 8 0.80 0.00 2 Aco1-1 0.00 1.00 9 0.05 0.00 4 1.00 0.00 12 Pgd2-1 1.00 0.00 3 Adh2-4 0.57 0.01 4 0.00 1.00 6 0.43 0.99 13 Pgd3-2 0.00 0.07 4 Arg1-3 0.30 0.93 4 1.00 0.93 4 0.69 0.07 14 Pgm1-4 0.00 1.00 6 0.01 0.00 5 1.00 0.00 5 Enp1-4 0.42 0.93 15 Pgm3-2 0.01 0.00 5 0.58 0.07 4 0.96 0.71 6 Idh1-4 1.00 0.97 6 0.03 0.29 6 0.00 0.03 16 Skd1-4 0.86 1.00 7 Leu1-2 0.02 0.93 6 0.14 0.00 4 0.95 0.07 17 Tpi1-4 0.04 1.00 6 0.03 0.00 5 0.96 0.00 8 Mdh1-4 1.00 0.07 18 Tpi2-4 0.12 0.00 6 0.00 0.93 5 0.88 1.00 9 Mdh5-4 0.91 0.00 19 Tpi4-4 1.00 0.23 6 0.09 1.00 n 0.00 0.77 10 Nad1-1 0.02 1.00 4 0.98 0.00 Out of 19 loci, both are fixed at 3 and nearly fixed at 5 additional loci; Thus, both are clearly demarcated by isozyme constitution Out of 42 alleles, 13 G. arboreum, 8 G. herbaceum and 21 shared by both
- 42. 42 • Out of 19 polymorphic loci, G. arboreum and G. herbaceum are: Fixed for different alleles at 3 loci and nearly fixed at 5 additional loci Based on these isozyme studies + Previously documented cytogenetic evidences • Out of 42 alleles of these 19 loci: 8 unique to G. herbaceum 13 restricted to G. arboreum 21 shared by both • G. arboreum and G. herbaceum were domesticated independently due to fixation of large no. of loci and high level of allelic novelty
- 43. 43 C.S.-2: Restriction site mutations in cpDNA of Old World diploids, 5 Allotetraploids and 10 New World diploids Old world diploids • G. herbaceum • G. arboreum Allotetraploids • G. hirsutum • G. barbadense • G. tomentosum • G. mustelinum • G. darwinii New world diploids • G. thurberi • G. armourianum • G. harknessii • G. davidsonii • G. klotzschianum • G. aridum • G. raimondii • G. gossypioides • G. trilobum • G. turneri Synthetic allotetraploids and their respective parents were studied for cpDNA restriction site mutations Out of 78 restriction site mutations observed: • 38 subsets of D genome diploid species (Within D genome) • 30 unified Old World and New World cottons & differentiated D genome from both of them Wendel (1989)USA Confirmation of strict maternal inheritance of cpDNA
- 44. 44 • Hybridization and polyploidization events that led to the evolution of tetraploid cotton were relatively recent i.e. 1.1 to 1.9 MYA (million years ago) • Divergence time of 6 to 11 MYA of A and D genomes is supposed from their respective parental lineages • Female parent of initial intergenomic hybridization was very similar to present day G. arboreum and G. herbaceum Probe used Dra I (P1, P4) Dra I (P3) Hind II (P6-8) Pal I (P6-8) Sac II (P1, P4) Bgl II (S6-8) Cla I (P4) Eco RI (S6-8) Sac I (P10-10) Sry I (P6-8) Xba I (P3) Xba I (P1) Parsimony tree of Gossypium
- 45. 45 C.S.-3: Bidirectional interlocus concerted evolution following allopolyploid speciation of r-DNA • Reported on r-DNA evolution in 5 Allopolyploids (AD genomes), species representing their diploid progenitors (A and D genomes) and one of distantly related species (C genome) • Sequence data from the internal transcribed spacer regions (ITS1 & ITS2) and the 5.8S gene Wendel et al. (1995)Ames
- 46. 46 • Arrays are homogenous, or nearly so, in all diploids and allopolyploids • Sequence Parsimony – interlocus concerted evolution has been bidirectional in allopolyploid species (non-monophyletic) under the evolutionary forces. • Sequence Evolution occurred subsequent to hybridization and allopolyploidization Parsimony (gene) tree of r-DNA ITS sequences in Gossypium D genome Clade A genome Clade Organismal tree
- 47. 47 C.S.-4: Analysis of nuclear and chloroplast genes to resolve the diversification of Cotton genus Phylogenetic relationship derived using DNA sequences from: • 11 single copy nuclear loci • Nuclear rDNA • 4 cpDNA loci Cronn et al. (2002Iowa
- 48. 48 Based on the sequence data: • Separation of Gossypium (as diploids), Gossypoides kirkii and Kokia drynaroides occurred appox. 13.4 MYA • The D genome of Gossypium diverged from all other Cottons approx. 6.8 MYA • Lineages comprising A, B, E, F and G genomes share a common history of 1 MY • Cotton genome groups radiated in rapid succession after formation of the genus (in 17% of the time since the separation of Gossypium from its nearest extant relatives) Fig. 7:Maximum likelihood (ML) tree obtained from nuclear synonymous site data (7978 bp) using a molecular clock constraint and inferences for absolute timing of divergence among diploid Gossypium lineages.
- 49. 49 Shorter genome L1 (cM)a Longer genome L2 (cM)a Percentage difference A 532.73 D 563.77 5.8% Dt 532.40 At 557.89 4.8% A 134.00 At 203.00 51.5% D 769.20 Dt 1219.38 58.5% a L1 and L2 were calculated by summing the genetic distance between each adjacent pair of loci. b Calculated as L1 –L2 /L1 , where L1 = shorter genome length, and L2 = longer genome length. Brubaker et al., 1999USA C.S.-5 : Comparative genetic mapping of allotetraploid cotton and its diploid progenitors Comparative RFLP mapping was used to construct genetic maps for the allotetraploids (AD genome; n = 26) and diploids (A and D genomes; n = 13) Polyploidization in Gossypium is associated with enhanced recombination, as genetic lengths for allotetraploid genomes are over 50% greater than those of their diploid counterparts. Table 8. Genetic length differences among the diploid (A, D) and allotetraploid (At, Dt) Gossypium mapping populations.
- 50. 50 Evolution of Fiber Under DomesticationEvolution of Fiber Under Domestication Pre-adapted A-genome ancestor later contributed this genome and its propensity to allotetraploid cotton that colonized and diversified in the new world cotton The origin of spinnable fibers is occurred once in the history of Gossypium, following the divergence of the A- genome and F-genome clades Fig. 8. Evolution of Fiber Under Domestication Gossypium spp were independently domesticated by aboriginal domesticators about 5000 yrs ago, or more, and transformed into fiber and seed oil plants
- 51. 51 Human selection over 5 millennia transformed G. hirsutum into; Rangy high yielding Perennial shrub with a poorly synchronized fruit set annualize row-crop with a heavy fruit set Photoperiod sensitivity photoperiod insensitivity Small seeds that required scarification for germination in vitro seeds that germinate readily upon planting At the same time, fibers become longer, stronger and finer Domestication led a fine-tuning of the reactive oxygen species which thereby lead up-regulation of signal transduction and hormone signaling genes and down –regulation of cell wall maturation genes G. arboreum race indicum: Africa, Asia G. herbaceum ssp. africanum, southern Africa G. hirsutum race yucatanense, Yucatan peninsula G. barbadense, Central coastal Peru Fig. 9. Evolution of G. hirsutum fiber under Domestication
- 52. 52 Cultivated G. hirsutum wild G. hirsutum G. tomentosum Cultivated G. arboreum wild G. herbaceum G. raimondii G. davidsonii G. longicalyx G. anomalum G. sturtianum Fig. 10. Mature seeds of cultivated and wild tetraploid and diploid species
- 53. 53 C.S. -6. Gene expression in developing fibres of Upland cotton (G. hirsutum L.) was massively altered by domestication Microarray analysis, followed by clustering Categorized and compared the expression level of 40,430 genes in wild and domesticated cotton Ryan et al., 2010USA At all time points more genes were up-regulated in TM-1 than in yucatanense Fig. 11. Genes differentially expressed during fiber development in cotton
- 54. 54 Add one graff C.S.-7. Evolution of Spinnable Cotton Fiber Entailed Prolonged Development and a Novel Metabolism Microarray analysis with 22,000 genes Gene expression profiling Fig. 12. Summary of the Number of Genes Differentially Expressed between Adjacent Time-Points during Fiber Development (FDR < 0.05); G. logicalyx (F genome), G. herbaceum (A genome) Hovav et al., 2008USA
- 55. 55 Fig. 13. An Evolutionary and Development Model Describing Processes That Lead to the Formation of Spinnable fiber At the beginning of fiber development in F-genome fibers, many genes involved with stress-response processes were highly upregulated A-genomes fibers was accompanied by novel expression of genes that assist in regulating H2O2 and other ROS levels Hovav et al., 2008
- 56. 56 1027 - A LF G. arboreum G. hirsutum G. herbaceum Am. Nect. less Gaorani - 6 BC to G. hirsutum Deviraj 170 – Co – 2 (1951) Biurbon Cotton was the first G. hirsutum type tried out in the state – Year 1797 Renewd effort- Year 1810, East India Company – Year 1838 CO - 2 Red Arboreum BC to G. hirsutum BC-263-1 BC-22 BC to BC 22 Gujarat 67 (1963) CO - 2 G. tomentosum BC to CO2 Cotom Indor-2 CTI 421 KW 66 2096 SRT 3087G.Cot 12 (Khapati) (1974) G. Cot. Hy. 4 (1971) Devitej 134 – Co – 2 (1952) G.Cot 100 (1974) Stabilized G. Cot. Hy. 6 (1980) G.Cot 10 (SRT 1) (1974) Surat Dwarf G. Cot. Hy. 10 hh (1995) G. Cot. Hy. 12 hh(2006) Dharwar American 2.6.5 G. Cot. Hy. 8 (1989) G. Cot. Hy. 6 BG II hh (2012) G. Cot. Hy. 8 BG II hh(2012) Other Hybrids Bt cotton Hybrids Success of Allotetraploid Cotton in Gujarat G. Cot. Hy. 102 hb(2002) GTHH-49 BG II hh(2013) G. Cot. Hy. 12 BG II hh (2013)
- 57. 57 Variety Year of Release Area of Cultivation Wagad 8* 1930 Wagad Area Vijay* 1943 Middle Guj. Kalyan* 1947 North Guj. Pratap 1947 Mathio tract Vijalpa* 1952 South Guj. Digvijay* 1956 Middle Guj. Sanjay 1958 Mathio tract V 797* 1966 Wagad Area Sujay* 1971 Middle Guj. G. Cot 101 1977 Budded cotton for Adivasi area G. Cot 11* 1979 South & Middle Guj. G. Cot 13* 1981 Wagad Area G. Cot 15* 1989 Mathio tract G. Cot 17* 1995 Middle Guj. G. Cot 19* 1997 Mathio tract G. Cot 21* 1998 Part of Wagad Area G. Cot 23* 2002 Gujarat State Hybrids DH 7 1985 Gujarat State DH 9 1988 Gujarat State, M.P. G. Cot. MDH 11 2002 Gujarat State Table 9. Diploid Cotton Varieties Released from the Gujarat * G. herbaceum
- 58. 58 Table 10. Tetraploid Cotton Varieties and hybrids Released from the Gujarat Variety Year of Release Area of Cultivation Deviraj (170-Co.2) 1951 Whole Guj. Devitej (134-Co-2-M) 1952 Middle Guj. G 67 1963 South & Middle Guj. G. Cot 10 1974 Whole Gujarat G. Cot 100 1974 South Guj. G. Cot 12 1981 Wagad area G. Cot 16 1995 Middle Guj. G. Cot 18 2001 Saurashtra region G. Cot 20 2010 Gujarat State GJ. Cot 101 2013 Gujarat State G. Cot Hy. 4 1970 Guj., A.P., Karnataka, Maharashtra G. Cot Hy. 6 1980 Guj., Maharashtra ., A.P., G. Cot Hy. 8 1989 Gujarat State G. Cot Hy. 10 1995 Gujarat State G. Cot Hy. 102 (HxB) 2002 Gujarat State G. Cot Hy. 12 2006 Gujarat State G. Cot Hy. 6 BG II 2012 Gujarat State G. Cot Hy. 8 BG II 2012 Gujarat State G. Cot Hy. 12 BG II 2013 Gujarat State GTH-49 BG-II 2013 Gujarat State
- 59. Fig.14:Current Cotton evolutionary Progress in IndiaFig.14:Current Cotton evolutionary Progress in India New Insecticides LRA 5166, NHH 44 2012 59
- 60. Name Event Genes Year of approval Bollgard I MON 531 cry1Ac 2002 Bollgard II MON 15985 cry1Ac and cry2Ab 2006 Event 1 Event 1 Truncated cry1Ac 2006 GFM Cry1A GFM C cry1Ab+cry1Ac 2006 DharwadEvent Dharwad Event Truncated cry1Ac 2008 9124 Metahelix cry1C 2009 Bt cotton events approved for cultivation in India Event name Event number Company/institution Genes Event 1 + Event 24 Event 1 + Event 24 JK Agri cry1Ac and cry1EC Widestrike Event 3006-210-23 Dow Agro cry1Ac and cry1F +Event 281-24-236 Roundup Ready MON 15985 + Monsanto cry1Ac,cry2Ab, CP4EPSPS Flex Bt MON 88913 TwinLink® Cotton GHB119 Bayer Crop Sci. (cry2Ae/PAT) & T304-40 (cry1Ab/PAT) cry1Ab, cry2Ac and bar GHB614 Zmepsps Source: http://www.igmoris.nic.in/field_trials.asp. Bt cotton events currently undergoing field tests in India 60
- 61. 61 ConclusionConclusion • Cotton distinguished from other Malvaceae species by having the gossypol glands. • Gossypium and their nearest relatives (Gossypoides and Kokia) diverged from their common ancestors about 13.4 MYA. • Divergence of A and D genomes from their parental lineages occurred about 6 to 8 MYA. • Allotetraploid cottons evolved approximately 1.1 to 1.9 MYA as a result of hybridization between their diploid ancestors followed by polyploidization events. • The maternal parent (progenitor) involved in the evolution of allotetraploids was ‘A’ genome donor (i.e. similar to the present day old world cottons), also confirmed through mitochondrial and diploid genome. Cont…
- 62. • Origin of the spinable fiber occurred once in the history of Gossypium in A genome; these traits and major fiber quality related traits from the D genome were later contributed to allotetraploid cotton. • Domestication has made fiber longer, stronger and finer. • Great variability exits in the genus which has been successfully exploited by introgression breeding. These introgressions led to considerable improvement in productivity, fiber quality and biotic and abiotic stress resistance. Cont… 62
- 64. 64
- 65. 65 Future Prospects • Understanding the relationships among species and their evolutionary development will continue to provide insights into the biology of cotton which in turn will increase the effectiveness of improvement efforts. • The wild species of cotton, consequently, represent an ample genetic repository for exploitation. Although these wild species remain a largely untapped genetic resource, examples abound of their productive inclusion in breeding programs. • Sequenced cotton genomes will ultimately stimulate fundamental research on genome evolution. • Transgenic cottons have been proven as one of the successful crops for large scale cultivation. Thus, the fate of the transgenic cottons should be further considered from the evolution point of view. • The genes can be incorporated from the wild species to cultivated cottons by Cisgenic approach, especially for fiber quality as well as pests and disease resistance.
- 66. 66 C.S.-3: Analysis of DNA polymorphism in G. hirsutum, G. herbaceum and G. raimondii by RFLPs Homologous and homoeologous probes: • Mapped PstI-genomic probes from A, D and AD genomes • Screened cDNA NotI-genomic probes from G. hirsutum Genomic DNA from all three genomes were digested using EcoRI Reinisch et al. (1994) Table 8. Hybridization of homologous and homeologous cotton DNA probes to EcoRI-digested genomic DNA from A, D and AD genome cottons Source of DNA probe Number of restriction fragments hybridizing to genomic DNA from different genomes Source of genomic DNA A D AD Total A genome PstI fragments 157 155 187 499 D genome PstI fragments 178 161 232 571 AD genome PstI fragments 149 126 226 501 AD genome cDNAs 167 154 262 583 Total 651 596 907 2154 Average per DNA probe 2.96 2.71 4.12 --- Iowa
Editor's Notes
- Parsimony tee of Gossypium based on restriction site loss/gain mutations in cpDNAs
- At the beginning of fiber development in F-genome fibers, many genes involved with stress-response processes were highly upregulated
- Success of Alltetraploid Cotton in Gujarat
- Cont…
- S