摘要
棉花质量性状的遗传分析和基因定位,对棉花育种工作者设计和培育新品种,提高育种效率,创新种质资源都有重要指导作用。棉花质量性状的研究始于20世纪初期,但都是基于形态上的观察。近年来,棉花基因组学研究的不断深入和分子标记遗传图谱的不断完善,为定位克隆棉花重要质量性状基因提供了便利。迄今为止,研究人员已对大部分质量性状基因进行了遗传连锁分析或染色体定位,其中部分质量性状的候选基因也已被鉴定出来。本文主要从质量性状遗传分析、分子标记定位和候选基因克隆几个方面系统地综述了植株颜色、叶片颜色、叶型、苞叶、花、蜜腺、腺体和纤维8类质量性状,为进一步研究棉花质量性状形成的分子机理和调控机制、定向转移基因及基因聚合育种等提供参考。
Genetic analysis and molecular mapping of qualitative traits are important tools in cotton breeding. Research on cotton qualitative traits has been reported since the early 1900 s. Rapid advances in cotton genomics and molecular mapping over the past several years have facilitated the mapping and cloning of important qualitative trait genes in cotton. Although most of the101 genes and alleles related to cotton morphological qualitative traits have not been mapped to linkage groups or chromosomes,several genes have been cloned and identified via map-based cloning. This paper describes research progress on the genetic and linkage analysis, molecular mapping, and candidate gene cloning of eight major types of morphological qualitative traits: plant color, leaf color, leaf shape, bract, flower, nectary, gland, and fiber characteristics. The presented information lays a foundation for studying the molecular basis of qualitative traits, favorable gene-based transfer, and gene pyramiding in cotton.
引文
[1]中国农业科学院棉花研究所.中国棉花遗传育种学[M].济南:山东科学技术出版社, 2003.Institute of Cotton Research of Chinese Academy of Agricultural Sciences. Genetics and breeding of cotton in China[M]. Jinan:Shandong Science and Technology Press, 2003.
[2]赵亮.四倍体栽培棉种高密度遗传图谱的加密及棉花红株R1和茸毛T1基因的精细定位[D].南京:南京农业大学, 2012.Zhao Liang. Construction of high density genetic map for tetraploid cultivated cotton species and fine mapping of red plant R1and trichomes T1in cotton[D]. Nanjing:Nanjing Agricultural University, 2012.
[3]赵佳.陆地棉亚红株(Rs)候选基因的克隆和功能分析[D].重庆:西南大学, 2014.Zhao Jia. Cloning and functional analyses of sub-red(Rs)candidate gene in upland cotton(Gossypium hirsutum L.)[D].Chongqing:Southwest University, 2014.
[4]陈旭升,狄佳春,马晓杰,等.陆地棉亚红株突变性状的连锁遗传规律分析[J].遗传, 2009, 31(8):855-858.Chen Xusheng, Di Jiachun, Ma Xiaojie, et al. Analysis on linkage inheritance of sub-red plant m utant from upland cotton[J].Hereditas, 2009, 31(8):855-858.
[5] Mac Michael S C. Occurrence of the dwarf-red character in upland cotton[J]. Journal of Agricultural Research, 1942, 64(8):477-481.
[6] Lennart B. Complementary factors for dark-red plant color in upland cotton[J]. Journal of Agricultural Research, 1949, 78(11):535-543.
[7]田明爽,宋美珍,范术丽,等.植物叶绿素缺失突变体分子机制研究进展[J].西北植物学报, 2011, 31(9):1900-1907.Tian Mingshuang, Song Meizhen, Fan Shuli, et al. Advance in research of molecular mechanism of chlorophyll-deficient mutants in plants[J]. Acta Botanica Boreali-Occidentalia Sinica, 2011, 31(9):1900-1907.
[8] Turcotte E L, Feaster C V. Inheritance of three genes for plant color in American pima cotton[J]. Crop Science, 1978, 18(1):149-150.
[9] Rhyne C L. The inheritance of yellow-green, a possible mutation in cotton[J]. Genetics, 1955, 40(2):235-245.
[10] Murray J C, Brinkerhoff L A. Inheritance of a pale-green color mutant in cotton[J]. Crop Science, 1966, 6(4):375.
[11] Zhu J, Chen J, Gao F, et al. Rapid mapping and cloning of the virescent-1 gene in cotton by bulked segregant analysis-next generation sequencing and virus-induced gene silencing strategies[J]. Journal of Experimental Botany, 2017, 68(15):4125-4135.
[12] Mao G, Ma Q, Wei H, et al. Fine mapping and candidate gene analysis of the virescent gene v1in upland cotton(Gossypium hirsutum)[J]. Molecular Genetics and Genomics, 2018, 293(1):249-264.
[13] Duncan E N, Pate J B. Inheritance and use of golden crown virescence in cotton:And its relationship to other virescent stocks[J]. Journal of Heredity, 1967, 58(5):237-239.
[14] Percival A E, Kohel R J. Genetic analysis of virescent mutants in cotton[J]. Crop Science, 1974, 14(3):439-440.
[15] Quisenberry J E, Kohel R J. Genetics of the virescent-4 mutant in cotton[J]. Journal of Heredity, 1970, 61:212-14.
[16] Kohel R J. Analysis of irradiation induced virescent mutants and the identification of a new virescent mutant(v5v5, v6v6)in Gossypium hirsutum L.[J]. Crop Science, 1973, 13(1):86-88.
[17] Turcotte E L, Feaster C V. The interaction of two genes for yellow foliage in cotton[J]. Journal of Heredity, 1973, 64(4):231-232.
[18] Kohel R J. Genetic analysis of a new virescent mutant in cotton[J]. Crop Science, 1974, 14(4):525-527.
[19]宋美珍,杨兆光,范术丽,等.一个短季棉芽黄基因型的鉴定及生理生化分析[J].中国农业科学, 2011, 44(18):3709-3720.Song Meizhen, Yang Zhaoguang, Fan Shuli, et al. Physiological and biochemical analysis and identifi cation of a short season cotton virescent mutant[J]. Scientia Agricultura Sinica,2011, 44(18):3709-3720.
[20] Percival A E, Kohel R J. New virescent cotton mutant linked with the marker gene yellow petals[J]. Crop Science, 1976, 16(4):503-505.
[21] Kohel R J. Genetic analysis of virescent mutants and the identification of virescents v12, v13, v14, v15and v16, v17in upland cotton[J]. Crop Science, 1983, 23(2):289-291.
[22]吴纪元,潘家驹.我国四个陆地棉芽黄突变品系遗传研究初报[J].南京农业大学报, 1985, 8(1):8-17.Wu Jiyuan, Pan Jiaju. Preliminary report of genetic studies on four mutant strains of cotton bud yellow in China[J]. Journal of Nanjing Agricultral University, 1985, 8(1):8-17.
[23] Turcotte E L, Percy R G. Inhertance of a second virescent mutant in American pima cotton[J]. Crop Science, 1988, 28(6):1018-1019.
[24]张天真,潘家驹.陆地棉芽黄突变体的等位性测验及其V22芽黄基因的遗传学鉴定[J].江苏农业学报, 1990, 6(1):24-29.Zhang Tianzhen, Pan Jiaju. Allelomorphism test of mutant of cotton bud yellow of land and genetic identification of gene of bud yellow of V22[J]. Journal of Jiangsu Agriculture, 1990,6(1):24-29.
[25] Turcotte E L, Feaster C V. Inheritance of a leaf mutant in American pima cotton[J]. Journal of Heredity, 1980, 71(2):134-135.
[26] Turcotte E L. Inheritance of a second wrinkled leaf mutant in American pima cotton[J]. Crop Science, 1987, 27(4):702-704.
[27]陈旭升,狄佳春,马晓杰.陆地棉皱缩叶突变性状质量遗传规律分析[J].江西农业学报, 2012, 24(4):7-9.Chen Xusheng, Di Jiachun, Ma Xiaojie. Analysis of qualitative inheritance of wrinkled leaf mutative character in upland cotton[J]. Journal of Jiangxi Agriculture, 2012, 24(4):7-9.
[28]李峰利.陆地棉皱缩叶突变体基因wr3的初步定位[D].南京:南京农业大学, 2014.Li Fengli. Gene mapping of wrinkled leaf mutant gene 3(wr3)in upland cotton[D]. Nanjing:Nanjing Agricultural University,2014.
[29]高阳.棉花叶皱小突变体遗传特性及相关基因表达分析[D].济南:山东师范大学, 2015.Gao Yang. Genetic and molecular characterization on a cotton mutant with small crinkled leaves[D]. Jinan:Shandong Normal University, 2015.
[30] Turcotte E L, Feaster C V. The inheritance of rugate leaf in pima cotton[J]. Journal of Heredity, 1965, 56(5):234-236.
[31] Kohel R J, Percival A E. Genetic analysis of the rex mutant in cotton[J]. Journal of Heredity, 1989, 80(1):78-80.
[32] Kohel R J. Genetic analysis of the crumpled mutant in cotton,Gossypium hirsutum L.[J]. Crop Science, 1973, 13(3):384-386.
[33] Kohel R J. Crinkle-yellow, a new mutant in Gossypium hirsutum L.[J]. Journal of Heredity, 1982, 73(5):382-383.
[34] Lewis C F. The inheritance of cup leaf in cotton[J]. Journal of Heredity, 1954, 45(3):127-127.
[35] Didlay R H, Waddle B A. Inheritance and linkage relationship of strap leaf mutant in cotton[J]. Crop Science, 1974, 14(3):387-390.
[36] Lee J A. Inheritance of two new mutations in cotton[J]. Journal of Heredity, 1985, 76(6):479-480.
[37] Brown H B, Cotton J R.'ROUND-LEAF'cotton notes on the appearance and behavior of a peculiar new strain[J]. Journal of Heredity, 1937, 28(1):45-48.
[38] Percival A E, Kohel R J, Dilday R H. A dominant round leaf mutant in cotton[J]. Crop Science, 1976, 16(6):794-796.
[39] Turcotte E L. Round leaf-3 mutant in American pima cotton[J].Journal of Heredity, 1986, 77(5):364-366.
[40] Kohel R J, Lewis C F, Christiansen M N. The identification of a new mutant and linkage group in cotton[J]. Journal of Heredity,1977, 68(1):65-66.
[41] Tiranti I N.'Algodon Crespo.'a new mutation in Gossypium hirsutum L.[J]. Bol Genet Inst Fitotec Castelar, 1967, 3:29-33.
[42] Kohel R J, Lewis C F. Inheritance of ragged leaf mutant in American upland cotton, Gossypium hirsutum L.[J]. Crop Science, 1962, 2(1):61-62.
[43] Kohel R J. Genetic analyses of the yellow-veins mutant in cotton[J]. Crop Science, 1983, 23(2):291-293.
[44] Kohel R J, Bird L S. Inheritance and linkage analysis of the yellow pulvinus mutant of cotton[J]. The Journal of Cotton Science, 2002, 6:115-118.
[45] Shoemaker D N. A study of leaf characters in cotton hybrids[J].Journal of Heredity, 1909(1):116-118.
[46] Andres R J, Bowman D T, Kaur B, et al. Mapping and genomic targeting of the major leaf shape gene(L)in upland cotton(Gossypium hirsutum L.)[J]. Theoretical and Applied Genetics,2014, 127(1):167-177.
[47] Zhu Q H, Zhang J, Liu D, et al. Integrated mapping and characterization of the gene underlying the okra leaf trait in Gossypium hirsutum L.[J]. Journal of Experimental Botany, 2015, 67(3):763-774.
[48] Chang L, Fang L, Zhu Y, et al. Insights into interspecific hybridization events in allotetraploid cotton formation from characterization of a gene regulating leaf shape[J]. Genetics, 2016,204(2):799-806.
[49] Andres R J, Coneva V, Frank M H, et al. Modifications to a LATE MERISTEM IDENTITY1 gene are responsible for the major leaf shapes of upland cotton(Gossypium hirsutum L.)[J].Proceedings of the National Academy of Sciences, 2017, 114(1):E57-E66.
[50]刘剑光,肖松华,吴巧娟,等.棉花狭薄卷苞叶突变体的遗传及利用研究[J].安徽农学通报, 2007, 13(18):38-39.Liu Jianguang, Xiao Songhua, Wu Qiaojuan, et al. Study on the inheritance and utilization of mutations in cotton narrow curly bracts[J]. Anhui Agricultural Science Bulletin, 2007, 13(18):38-39.
[51] Li F, Hu X, Qiu X, et al. Fine mapping of the fg gene using the interspecific cross of Gossypium hirsutum×Gossypium barbadense[J]. Euphytica, 2010, 176(3):303-308.
[52] Knight R L. The genetics of withering or deciduous bracteoles in cotton[J]. Journal of Genetics, 1952, 50(3):392-395.
[53] Kohel R J. Genetic analysis of the open bud mutant in cotton[J].Journal of Heredity, 1973, 64(4):237-238.
[54] Rhyne C. Open-bud cotton—a product of interspecific modifiers?[J]. Journal of Heredity, 1979, 70(1):80.
[55] Qian N, Zhang X W, Guo W Z, et al. Fine mapping of open-bud duplicate genes in homoelogous chromosomes of tetraploid cotton[J]. Euphytica, 2009, 165(2):325-331.
[56] Endrizzi J E, Ray D T. Linkage analysis of open bud(ob2)and yellow petal(Y1)in cotton[J]. Genome, 1991, 34(3):461-463.
[57] Endrizzi J E, Nelson R. Linkage analysis and arm location of the open bud(ob1)and yellow petal(Y2)loci in chromosome 18 of cotton[J]. Genome, 1989, 32(6):1041-1043.
[58] Harland S C. The genetics of cotton. II. The in heritance of pollen colour in New World cottons[J]. Journal of Genetics,1929, 20(3):387-399.
[59] Stephens S G. Interspecific homologies between gene loci in Gossypium. I. Pollen color[J]. Genetics, 1954, 39(5):701-711.
[60] Turcotte E L, Feaster C V. A second locus for pollen color in pima cotton, Gossypium barbadense L.[J]. Crop Science, 1966, 6(2):117-119.
[61] Song X L, Wang K, Guo W Z, et al. A comparison of genetic maps constructed from haploid and BC1mapping populations from the same crossing between Gossypium hirsutum L. and Gossypium barbadense L.[J]. Genome, 2005, 48(3):378.
[62] Harland S C. The genetics of cotton. Part I. The inheritance of petal spot in New World cottons[J]. Journal of Genetics, 1929,20(3):365-385.
[63] Meyer J R, Meyer V G. Origin and inheritance of nectariless cotton[J]. Crop Science, 1961, 1(3):167-169.
[64] Waghmare V N, Rong J, Rogers C J, et al. Genetic mapping of a cross between Gossypium hirsutum(cotton)and the Hawaiian endemic, Gossypium tomentosum[J]. Theoretical and Applied Genetics, 2005, 111(4):665-676.
[65] Murray J C. A new locus for glanded stem in tetraploid cotton[J]. Journal of Heredity, 1965, 56(1):42-44.
[66] Mc Michael S C. Yuma glandless, an allelomorph of glandless-one in cotton, Gossypium hirsutum L.[J]. Crop Science,1970, 10(2):202-203.
[67]陆才瑞.棉花茎秆无腺体基因gl1的精细定位及棉酚腺体发育相关基因筛选[D].武汉:华中农业大学, 2016.Lu Cairui. Fine mapping of stem glandless gene gl1in cotton and gossypol gland development related genes screening[D].Wuhan:Huazhong Agricultural University, 2016.
[68] Barrow J R, Davis D D. Gl28—a new allele for pigment glands in cotton[J]. Crop Science, 1974, 14(2):325-326.
[69] Kohel R J, Lee J A. Genetic analysis of Egyptian glandless cotton[J]. Crop Science, 1984, 24(6):1119-1121.
[70] Ma D, Hu Y, Yang C, et al. Genetic basis for glandular trichome formation in cotton[J]. Nature Communications, 2016, 7:10456.
[71] Cheng H L, Lu C R, John Z Y, et al. Fine mapping and candidate gene analysis of the dominant glandless gene Gl2ein cotton(Gossypium spp.)[J]. Theoretical and Applied Genetics, 2016,129(7):1347-1355.
[72] Lee J A. Allele determining rugate fruit surface in cotton[J].Crop Science, 1978, 18(2):251-254.
[73] Stephens S G. Geographic and taxonomic distribution of anthocyanin genes in New World cottons[J]. Journal of Genetics,1974, 61(2):128-141.
[74] Ware J O. Inheritance of lint percentage in cotton[J]. Journal of the American Society of Agronomy, 1929, 21(9):876-894.
[75] Griffee F, Ligon L L. Occurrence of'lintless'cotton plants and the inheritance of the character'lintless'[J]. Journal of the American Society of Agronomy, 1929, 21(7):711-717.
[76] Rong J, Pierce G J, Waghmare V N, et al. Genetic mapping and comparative analysis of seven mutants related to seed fiber development in cotton[J]. Theoretical and Applied Genetics, 2005,111(6):1137-1146.
[77] Wan Q, Guan X, Yang N, et al. Small interfering RNAs from bidirectional transcripts of GhMML3_A12 regulate cotton fiber development[J]. New Phytologist, 2016, 210(4):1298-1310.
[78] Ware J O, Benedict L I, Rolfe W H. A recessive naked-seed character in upland cotton[J]. Journal of Heredity, 1947, 38(10):313-320.
[79] Karaca M, Saha S, Jenkins J N, et al. Simple sequence repeat(SSR)markers linked to the Ligon lintless(Li1)mutant in cotton[J]. Journal of Heredity, 2002, 93(3):221-224.
[80] Kohel R J. Linkage tests in upland cotton, Gossypium hirsutum L. II[J]. Crop Science, 1972, 12(1):66-69.
[81] Gilbert M K, Turley R B, Kim H J, et al. Transcript profiling by microarray and marker analysis of the short cotton(Gossypium hirsutum L.)fiber mutant Ligon lintless-1(Li1)[J]. BMC Genomics, 2013, 14(1):403.
[82] Thyssen G N, Fang D D, Turley R B, et al. A Gly65Val substitution in an actin, Gh ACT_LI1, disrupts cell polarity and F-actin organization resulting in dwarf, lintless cotton plants[J].The Plant Journal, 2017, 90(1):111-121.
[83] Narbuth E V, Kohel R J. Inheritance and linkage analysis of a new fiber mutant in cotton[J]. Journal of Heredity, 1990, 81(2):131-133.
[84] Hinchliffe D J, Turley R B, Naoumkina M, et al. A combined functional and structural genomics approach identified an EST-SSR marker with complete linkage to the Ligon lintless-2genetic locus in cotton(Gossypium hirsutum L.)[J]. BMC Genomics, 2011, 12(1):445.
[85] Du X M, Pan J J, Wang R H, et al. Genetic analysis of presence and absence of lint and fuzz in cotton[J]. Plant Breeding, 2001,120(6):519-522.
[86] Ma Q F, Wu C H, Wu M, et al. Integrative transcriptome, proteome, phosphoproteome and genetic mapping reveals new aspects in a fiberless mutant of cotton[J/OL]. Scientific Reports,2016, 6:24485(2016-04-14)[2018-08-14]. https://www.nature.com/articles/srep24485. DOI:10.1038/srep24485.
[87] Wu H, Tian Y, Wan Q, et al. Genetics and evolution of MIXTA genes regulating cotton lint fiber development[J]. New Phytologist, 2018, 217(2):883-895.
[88] Cai C P, Tong X C, Liu F J, et al. Discovery and identification of a novel Ligon lintless-like mutant(Lix)similar to the Ligon lintless(Li1)in allotetraploid cotton[J]. Theoretical and Applied Genetics, 2013, 126(4):963-970.
[89] Beasley C A, Egli E. Fiber production in vitro from a conditional fiberless mutant of cotton[J]. Developmental Biology, 1977,57(1):234-237.
[90] Kohel R J, Mc Michael S C. Immature fiber mutant of Upland cotton[J]. Crop Science, 1990, 30(2):419-421.
[91] Kim H J, Moon H S, Delhom C D, et al. Molecular markers associated with the immature fiber(im)gene affecting the degree of fiber cell wall thickening in cotton(Gossypium hirsutum L.)[J]. Theoretical and Applied Genetics, 2013, 126(1):23-31.
[92] Kohel R J. Genetic analysis of fiber color variants in cotton[J].Crop Science, 1985, 25(5):793-797.
[93] Hutchinson J B. The inheritance of brown lint in New World cottons[J]. Journal of Genetics, 1946, 47(3):295-309.
[94] Wang L X, Liu H F, Li X Y, et al. Genetic mapping of fiber color genes on two brown cotton cultivars in Xinjiang[J/OL].SpringerPlus, 2014, 3(1):480(2014-08-28)[2018-08-10].http://doi.org/10.1186/2193-1801-3-480.
[95] Yan Q, Wang Y, Li Q, et al. Up-regulation of Gh TT2-3A in cotton fibres during secondary wall thickening results in brown fibres with improved quality[J]. Plant Biotechnology Journal,2018, 16(10):1735-1747.
[96] Walford S A, Wu Y, Llewellyn D J, et al. GhMYB25-like:a key factor in early cotton fibre development[J]. The Plant Journal,2011, 65(5):785-797.
[97] Wang K B, Wang Z W, Li F G, et al. The draft genome of a diploid cotton Gossypium raimondii[J]. Nature Genetics, 2012,44(10):1098-1103.
[98] Paterson A H, Wendel J F, Gundlach H, et al. Repeated polyploidization of Gossypium genomes and the evolution of spinnable cotton fibres[J]. Nature, 2012, 492(7429):423-427.
[99] Li F G, Fan G Y, Wang K B, et al. Genome sequence of the cultivated cotton Gossypium arboreum[J]. Nature Genetics, 2014,46(6):567-576.
[100] Li F G, Fan G Y, Lu C R, et al. Genome sequence of cultivated upland cotton(Gossypium hirsutum TM-1)provides insights into genome evolution[J]. Nature Biotechnology, 2015, 33(5):524-530.
[101] Zhang T Z, Hu Y, Jiang W K, et al. Sequencing of allotetraploid cotton(Gossypium hirsutum L. acc. TM-1)provides a resource for fiber improvement[J]. Nature Biotechnology,2015, 33(5):531-537.
[102] Yuan D J, Tang Z H, Wang M J, et al. The genome sequence of sea-island cotton(Gossypium barbadense)provides insights into the allopolyploidization and development of superior spinnable fibres[J/OL]. Scientific Reports, 2015, 5:17662(2015-12-04)[2018-08-14]. https://www.nature.com/articles/srep17662. DOI:10.1038/srep17662.
[103] Liu X, Zhao B, Zheng H J, et al. Gossypium barbadense genome sequence provides insight into the evolution of extra-long staple fiber and specialized me tabolites[J/OL]. Scientific Reports,2015, 5:14139(2015-12-30)[2018-08-14]. https://www.nature.com/articles/srep14139. DOI:10.1038/srep14139.
[104] Abe A, Kosugi S, Yoshida K, et al. Genome sequencing reveals agronomically important loci in rice using MutMap[J].Nature Biotechnology, 2012, 30(2):174-178.