小麦苗期钾效率相关性状的全基因组关联分析
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摘要
[目的]深入探究小麦钾效率的遗传基础,定向培育钾高效小麦品种是缓解我国钾肥资源短缺的重要途径。本文拟对不同钾处理条件下小麦苗期钾效率相关性状进行鉴定,并进行差异分子标记关联分析,筛选与钾效率紧密关联的分子标记,为钾效率相关基因的克隆及其在育种中的应用奠定基础。[方法]本研究以不同小麦品种(134份)为试验材料,于2014年和2015年,在山东农业大学温室内进行了营养液培养试验。设置正常供钾和低钾水平两个处理,7cm高、7日龄的幼苗在黑暗中生长28天后收获。将幼苗分为地上部和根部,测定植株干重(生物量)、钾含量,计算植株钾累积量、根冠生物量比和钾累积量比和钾利用效率。分析低钾处理对生物量及钾效率相关性状的影响,并以该群体检测到的15230个差异SNP标记为基础,用TASSEL 5.0软件中的GLM模型和MLM模型对钾效率相关性状数据进行标记-性状的关联分析,以确定小麦苗期钾效率稳定关联的分子标记位点。[结果]与正常钾处理相比,低钾处理下植株根、冠及全株钾含量及积累量均显著下降,而根冠生物量比、根冠和全株钾利用率均显著增加。关联分析共获得了1300个关联分子标记,其中1102个标记被定位在19条染色体上,这些分子标记绝大多数仅在特定环境条件下被检测到,三个环境下均可检测到的相对稳定的分子标记位点仅有3个,分别为Excalibur_c14273_1407、Ku_c11150_773、BS00094893_51。同时,试验筛选出4个性状簇集位点标记,这些位点同时与6～7个性状存在显著关联,分别为Excalibur_c8670_972、wsnp_Ex_c12887_20426781、wsnp_Ku_c13311_21255428、IACX5989。[结论]低钾处理对小麦苗期生长及钾效率相关性状遗传控制位点有显著影响,低钾处理下与生物量和钾效率相关性状显著关联的分子标记绝大多数在同一个钾处理环境中被检测到,因此不同钾处理环境下这些性状可能由截然不同的基因控制。试验检测到4个至少与6个性状同时显著关联的热点分子标记位点,这些位点与小麦苗期多个生物量及钾效率相关性状均存在显著关联,可能包含重要的基因信息,值得进一步深入研究。
[Objectives]Breeding wheat with K-high efficiency is an important pathway for alleviating pressure of potash resource shortage in China. In this study, to better understanding the genetic mechanism, the correlated traits of K-efficiency at wheat seedling stage were investigated by different K treatments, and the molecular markers closely related to K-efficiency traits were screened by correlation analyses. The study will provide support for cloning of K efficiency-related genes and breeding of wheat varieties with K-high efficiency.[Methods ]Using 134 wheat cultivars as materials,a hydroponics experiment with complete random design in triplicate was conducted in greenhouse at Shandong Agricultural University in 2014 and 2015. The seedlings of 7 cm high and 7 days old of all the cultivars were subjected to low K(LK) and normal K(CK) levels for 28 days in the dark, then harvested and divided into shoot and root. The seedling dry weight(biomass) and K content were analyzed,and the K accumulation,dry biomass ratio of root to shoot(RSDW), root to shoot ratio of K accumulation and K use efficiency were calculated. To investigate the effects of low K treatment on biomass and K-efficiency related traits, the correlation analysis of tested traits based on the 15230 different SNP markers was performed using GLM and MLM models in TASSEL 5.0 to obtain stably and significantly correlated markers/locus for wheat seedlings.[Results ]The contents and accumulation of K in roots, shoots and whole seedlings were all significantly decreased, although the root biomass and the K use efficiencies significantly increased at the seedling stage under the low K treatment. A total of 1300 markers were significantly associated with biomass and K-efficiency related traits, of which, a total of 1102 markers were located on the 19 chromosomes, and most of these molecular markers were screened only under specific K treatment conditions, and three loci including Excalibur_c14273_1407, Ku_c1 1150_773 and BS00094893 51 were identified under at least three environments. Meanwhile, a total of 4 loci such as wsnp_Ku_c13311_21255428,Excalibur_c8670_972, IACX5989 and wsnp_Ex_c12887_20426781 were confirmed to be significantly associated with six or seven traits.[Conclusions ]Low K stress significantly influenced the growth of wheat seedlings and the number of related molecular markers. Most of the molecular markers related to K-efficiency and biomass traits are detected only under one environment, indicating that these tested traits might be controlled by different genes.The study also detected four hot sites(molecular markers) which were associated with at least six traits exhibiting a significant correlation with K-efficiency traits at wheat seedling stage. These sites possibly contain important genetic information and deserve to be further investigated.
[Objectives]Breeding wheat with K-high efficiency is an important pathway for alleviating pressure of potash resource shortage in China. In this study, to better understanding the genetic mechanism, the correlated traits of K-efficiency at wheat seedling stage were investigated by different K treatments, and the molecular markers closely related to K-efficiency traits were screened by correlation analyses. The study will provide support for cloning of K efficiency-related genes and breeding of wheat varieties with K-high efficiency.[Methods ]Using 134 wheat cultivars as materials,a hydroponics experiment with complete random design in triplicate was conducted in greenhouse at Shandong Agricultural University in 2014 and 2015. The seedlings of 7 cm high and 7 days old of all the cultivars were subjected to low K(LK) and normal K(CK) levels for 28 days in the dark, then harvested and divided into shoot and root. The seedling dry weight(biomass) and K content were analyzed,and the K accumulation,dry biomass ratio of root to shoot(RSDW), root to shoot ratio of K accumulation and K use efficiency were calculated. To investigate the effects of low K treatment on biomass and K-efficiency related traits, the correlation analysis of tested traits based on the 15230 different SNP markers was performed using GLM and MLM models in TASSEL 5.0 to obtain stably and significantly correlated markers/locus for wheat seedlings.[Results ]The contents and accumulation of K in roots, shoots and whole seedlings were all significantly decreased, although the root biomass and the K use efficiencies significantly increased at the seedling stage under the low K treatment. A total of 1300 markers were significantly associated with biomass and K-efficiency related traits, of which, a total of 1102 markers were located on the 19 chromosomes, and most of these molecular markers were screened only under specific K treatment conditions, and three loci including Excalibur_c14273_1407, Ku_c1 1150_773 and BS00094893 51 were identified under at least three environments. Meanwhile, a total of 4 loci such as wsnp_Ku_c13311_21255428,Excalibur_c8670_972, IACX5989 and wsnp_Ex_c12887_20426781 were confirmed to be significantly associated with six or seven traits.[Conclusions ]Low K stress significantly influenced the growth of wheat seedlings and the number of related molecular markers. Most of the molecular markers related to K-efficiency and biomass traits are detected only under one environment, indicating that these tested traits might be controlled by different genes.The study also detected four hot sites(molecular markers) which were associated with at least six traits exhibiting a significant correlation with K-efficiency traits at wheat seedling stage. These sites possibly contain important genetic information and deserve to be further investigated.
引文
[1]张会民,刘红霞,王林生,等.钾对旱地冬小麦后期生长及籽粒品质的影响[J].麦类作物学报,2004, 24(3):73-75.Zhang H M, Liu H X, Wang L S, et al. Effect of potassium on the growth at the late stages and grain quality of winter wheat in dryland[J]. Journal of Triticeae Crops, 2004,24(3):73-75.
[2]王龙俊,郭文善,封超年.小麦高产优质栽培新技术[M].上海:上海科学技术出版社,2000.Wang L J, Guo W S, Feng C N. New techniques for high yield and high quality of wheat[M]. Shanghai:Shanghai Science and Technology Press, 2000.
[3]曹卫星,郭文善,王龙俊,荆奇.小麦品质生理生态及调优技术[M].北京:中国农业出版社,2005.Cao W X, Guo W S, Wang L J, Jing Q. Physiological ecology and optimization techniques of wheat quality[M]. Beijing:China Agriculture Press, 2005.
[4]许柯,戴其根,葛鑫,等.氮肥运筹对面条小麦品种陕229淀粉品质的影响[J].上海农业学报,2004, 20(1):49-53.Xu K, Dai Q G, Ge X, et al. Effects of N applying rate and methods on starch quality of noodle wheat Shannong229[J]. Acta Agriculture Shanghai,2004, 20(1):49-53.
[5]陈培元,蒋永罗,李英,付左.钾对小麦生长发育/抗旱性和某些生理特性的影响[J].作物学报,1987,13(4):322-328.Chen P Y, Jiang Y L, Li Y, Fu Z. Effect of potassium on wheat growth, drought resistance and physiological properties under different soil moisture conditions[J]. Acta Agronomica Sinica, 1987,13(4):322-328.
[6]邹铁祥,戴廷波,姜东,等.钾素水平对小麦氮素积累和运转及籽粒蛋白质形成的影响[J].中国农业科学,2006, 39(4):686-692.Zou T X, Dai T B, Jiang D, et al. Potassium supply affected plant nitrogen accumulation and translocation and grain protein formation in winter wheat[J]. Scientia Agricultura Sinica, 2006, 39(4):686-692.
[7]武际,郭熙盛,王允青,黄晓荣.钾肥运筹对小麦氮素和钾素吸收利用及产量和品质的影响[J].土壤,2008, 40(5):777-783.Wu J, Guo X S, Wang Y Q, Huang X R. Effects of potassium fertilizer on the absorption and utilization of nitrogen and potassium in wheat and the yield and quality of wheat[J]. Soils, 2008, 40(5):777-783.
[8]盖德安.旱灌春小麦需氮磷钾量及高产经济施肥技术研究[J].黑龙江农业科学,1994,(1):24-28.Gai D A. Studies on the required amount of N, P and K and the technology of economic high-yield fertilization for spring wheat onirrigated dryland[J]. Heilongjiang Agricultural Science, 1994,(1):24-28.
[9]刘荣乐,金继运,吴荣贵,梁鸣早.我国北方土壤-作物系统内钾素循环特征及秸秆还田与施钾肥的影响[J].植物营养与肥料学报,2000, 6(2):123-132.Liu R L, Jin J Y, Wu R G, Liang M Z. Study on the characteristics of potassium cycling in different soil-crop systems in northern China[J].Plant Nutrition and Fertilizer Science, 2000, 6(2):123-132.
[10]刘国栋,刘更另.水稻耐低钾基因型筛选方法的研究[J].土壤学报,1996, 33(2):113-120.Liu G D, Liu G L. Study on method screening rice genotypes tolerant to potassium[J]. Acta Pedologica Sinica,1996, 33(2):113-120.
[11]化党领,介晓磊,韩锦锋.植物钾吸收的分子水平研究[J].植物营养与肥料学报,2002, 8(3):377-383.Hua D L, Jie X L, Han J F. A review on the molecular basis of potassium uptake in plants[J]. Plant Nutrition and Fertilizer Science,2002, 8(3):377-383.
[12] Siddiqi M Y, Glass A D M. Utilization index:A modified approach to the estimation and comparison of nutrient utilization efficiency in plants[J]. Journal of Plant Nutrition, 1981, 4(3):289-302.
[13] Liu K, Muse S V. Power Marker:An integrated analysis environment for genetic marker analysis[J]. Bioinformatics, 2005, 21:2128-2129.
[14] Pritchard J K, Stephens M, Donnelly P. Inference of population structure using multi-locus genotype data[J]. Genetics, 2000, 155:945-959.
[15] Evanno G, Regnaut S, Goudet J. Detecting the number of clusters of individuals using the software STRUCTURE:a simulation study[J].Molecular Ecology, 2005, 14:2611-2620.
[16] Hermans C, Hammond J, White P, Verbruggen N. How do plants respond to nutrient shortage by biomass allocation[J]. Trends in Plant Science, 2006, 11:610-617.
[17] Pettigrew W T. Potassium influences on yield and quality production for maize, wheat, soybean and cotton[J]. Physiologia Plantarum,2008, 133:670-681.
[18] Rengel Z, Damon P. Crops and genotypes differ in efficiency of potassium uptake and use[J]. Plant Physiology, 2008, 133:624-636.
[19]梁雪,吴春红,宫晓平,等.不同钾处理下小麦重组自交系群体主要农艺性状的表型变异及其相关分析[J].中国生态农业学报,2012,20(5):520-528.Liang X, Wu C H, Gong X P, et al. Phenotypic variation and correlation analysis of major agronomic traits of wheat recombinant inbred lines population under different potassium levels[J]. Chinese Journal of Eco-Agriculture,2012, 20(5):520-528.
[20] Kong F M, Guo Y, Liang X, et al. Potassium(K)effects and QTL mapping for K efficiency traits at seedling and harvest stages in wheat[J]. Plant and Soil,2013, 373:877-892.
[21] Zhang G P, Chen J X, Tirore E A. Genotypic variation for potassium uptake and utilization efficiency in wheat[J]. Nutrient Cycling in Agro-ecosystems, 1999, 54:273-282.
[22] Pettersson S, Jensen P. Variation among species and varieties in uptake and utilization of potassium[J]. Plant and Soil, 1983, 72:231-237.
[23] Yang X E, Liu J X, Wang W M, et al. Genotypic differences andsome associated plant traits in potassium internal use efficiency of lowland rice(Oryzasativ L.)[J]. Nutrient Cycling in Agroecosystems, 2003, 67:273-282.
[24]赖勇,王鹏喜,范贵强,等.大麦SSR标记遗传多样性及其与农艺性状关联分析[J].中国农业科学,2013,(2):233-242.Lai Y, Wang P X, Fan G Q, et al. Genetic diversity and association analysis using SSR markers in barley[J]. Scientia Agricultura Sinica,2013,(2):233-242.
[25]郭志军,赵云雷,陈伟,等.陆地棉SSR标记遗传多样性及其与农艺性状的关联分析[J].棉花学报,2014,(5):420-430.Guo Z J, Zhao Y L, Chen W, et al. Genetic diversity and association analysis of upland cotton based on SSR markers[J]. Cotton Science,2014,(5):420-430.
[26]陈甲法.一个玉米叶色突变体的遗传研究[D].郑州:河南农业大学硕士学位论文,2009.Chen J F. Inheritance of a leaf color mutant in maize[D]. Zhengzhou:MS Thesis of Henan Agricultural University, 2009.
[27] Guo Y, Kong F M, Xu Y F, et al. QTL mapping for seedling traits in wheat grown under varying concentrations of N, P and K nutrients[J].Theoretical and Applied Genetics, 2012, 124:851-865.
[28] Xu Y, Wang R, Tong Y, et al. Mapping QTLs for yield and nitrogen-related traits in wheat:influence of nitrogen and phosphorus fertilization on QTL expression[J]. Theoretical and Applied Genetics,2014, 127:59-72.
[29]宫晓平.小麦钾养分效率及产量相关性状的QTL分析[D].山东泰安:山东农业大学硕士学位论文,2014.Gong X P. QTL mapping for K efficiency traits at seedling and yield related traits at harvest stages in wheat[D]. Tai'an, Shandong:MS Thesis of Shandong Agricultural University, 2014.
[30]兰进好,李新海,高树仁,等.不同生态环境下玉米产量性状QTL分析[J].作物学报,2005, 31:1253-1259.Lan J H, Li X H, Gao S R, et al. QTL analysis of yield components in maize under different environments[J]. Acta Agronomica Sinica,2005, 31:1253-1259.
[31]张国华,高明刚,张桂芝,等.黄淮麦区小麦品种(系)产量性状与分子标记的关联分析[J].作物学报,2013, 39(7):1187-1199.Zhang G H, Gao M G, Zhang G Z, et al.Association analysis of yield traits with molecular markers in Huang-Huai River valley winter wheat region, China[J]. Acta Agronomica Sinica, 2013, 39(7):1187-1199.
[2]王龙俊,郭文善,封超年.小麦高产优质栽培新技术[M].上海:上海科学技术出版社,2000.Wang L J, Guo W S, Feng C N. New techniques for high yield and high quality of wheat[M]. Shanghai:Shanghai Science and Technology Press, 2000.
[3]曹卫星,郭文善,王龙俊,荆奇.小麦品质生理生态及调优技术[M].北京:中国农业出版社,2005.Cao W X, Guo W S, Wang L J, Jing Q. Physiological ecology and optimization techniques of wheat quality[M]. Beijing:China Agriculture Press, 2005.
[4]许柯,戴其根,葛鑫,等.氮肥运筹对面条小麦品种陕229淀粉品质的影响[J].上海农业学报,2004, 20(1):49-53.Xu K, Dai Q G, Ge X, et al. Effects of N applying rate and methods on starch quality of noodle wheat Shannong229[J]. Acta Agriculture Shanghai,2004, 20(1):49-53.
[5]陈培元,蒋永罗,李英,付左.钾对小麦生长发育/抗旱性和某些生理特性的影响[J].作物学报,1987,13(4):322-328.Chen P Y, Jiang Y L, Li Y, Fu Z. Effect of potassium on wheat growth, drought resistance and physiological properties under different soil moisture conditions[J]. Acta Agronomica Sinica, 1987,13(4):322-328.
[6]邹铁祥,戴廷波,姜东,等.钾素水平对小麦氮素积累和运转及籽粒蛋白质形成的影响[J].中国农业科学,2006, 39(4):686-692.Zou T X, Dai T B, Jiang D, et al. Potassium supply affected plant nitrogen accumulation and translocation and grain protein formation in winter wheat[J]. Scientia Agricultura Sinica, 2006, 39(4):686-692.
[7]武际,郭熙盛,王允青,黄晓荣.钾肥运筹对小麦氮素和钾素吸收利用及产量和品质的影响[J].土壤,2008, 40(5):777-783.Wu J, Guo X S, Wang Y Q, Huang X R. Effects of potassium fertilizer on the absorption and utilization of nitrogen and potassium in wheat and the yield and quality of wheat[J]. Soils, 2008, 40(5):777-783.
[8]盖德安.旱灌春小麦需氮磷钾量及高产经济施肥技术研究[J].黑龙江农业科学,1994,(1):24-28.Gai D A. Studies on the required amount of N, P and K and the technology of economic high-yield fertilization for spring wheat onirrigated dryland[J]. Heilongjiang Agricultural Science, 1994,(1):24-28.
[9]刘荣乐,金继运,吴荣贵,梁鸣早.我国北方土壤-作物系统内钾素循环特征及秸秆还田与施钾肥的影响[J].植物营养与肥料学报,2000, 6(2):123-132.Liu R L, Jin J Y, Wu R G, Liang M Z. Study on the characteristics of potassium cycling in different soil-crop systems in northern China[J].Plant Nutrition and Fertilizer Science, 2000, 6(2):123-132.
[10]刘国栋,刘更另.水稻耐低钾基因型筛选方法的研究[J].土壤学报,1996, 33(2):113-120.Liu G D, Liu G L. Study on method screening rice genotypes tolerant to potassium[J]. Acta Pedologica Sinica,1996, 33(2):113-120.
[11]化党领,介晓磊,韩锦锋.植物钾吸收的分子水平研究[J].植物营养与肥料学报,2002, 8(3):377-383.Hua D L, Jie X L, Han J F. A review on the molecular basis of potassium uptake in plants[J]. Plant Nutrition and Fertilizer Science,2002, 8(3):377-383.
[12] Siddiqi M Y, Glass A D M. Utilization index:A modified approach to the estimation and comparison of nutrient utilization efficiency in plants[J]. Journal of Plant Nutrition, 1981, 4(3):289-302.
[13] Liu K, Muse S V. Power Marker:An integrated analysis environment for genetic marker analysis[J]. Bioinformatics, 2005, 21:2128-2129.
[14] Pritchard J K, Stephens M, Donnelly P. Inference of population structure using multi-locus genotype data[J]. Genetics, 2000, 155:945-959.
[15] Evanno G, Regnaut S, Goudet J. Detecting the number of clusters of individuals using the software STRUCTURE:a simulation study[J].Molecular Ecology, 2005, 14:2611-2620.
[16] Hermans C, Hammond J, White P, Verbruggen N. How do plants respond to nutrient shortage by biomass allocation[J]. Trends in Plant Science, 2006, 11:610-617.
[17] Pettigrew W T. Potassium influences on yield and quality production for maize, wheat, soybean and cotton[J]. Physiologia Plantarum,2008, 133:670-681.
[18] Rengel Z, Damon P. Crops and genotypes differ in efficiency of potassium uptake and use[J]. Plant Physiology, 2008, 133:624-636.
[19]梁雪,吴春红,宫晓平,等.不同钾处理下小麦重组自交系群体主要农艺性状的表型变异及其相关分析[J].中国生态农业学报,2012,20(5):520-528.Liang X, Wu C H, Gong X P, et al. Phenotypic variation and correlation analysis of major agronomic traits of wheat recombinant inbred lines population under different potassium levels[J]. Chinese Journal of Eco-Agriculture,2012, 20(5):520-528.
[20] Kong F M, Guo Y, Liang X, et al. Potassium(K)effects and QTL mapping for K efficiency traits at seedling and harvest stages in wheat[J]. Plant and Soil,2013, 373:877-892.
[21] Zhang G P, Chen J X, Tirore E A. Genotypic variation for potassium uptake and utilization efficiency in wheat[J]. Nutrient Cycling in Agro-ecosystems, 1999, 54:273-282.
[22] Pettersson S, Jensen P. Variation among species and varieties in uptake and utilization of potassium[J]. Plant and Soil, 1983, 72:231-237.
[23] Yang X E, Liu J X, Wang W M, et al. Genotypic differences andsome associated plant traits in potassium internal use efficiency of lowland rice(Oryzasativ L.)[J]. Nutrient Cycling in Agroecosystems, 2003, 67:273-282.
[24]赖勇,王鹏喜,范贵强,等.大麦SSR标记遗传多样性及其与农艺性状关联分析[J].中国农业科学,2013,(2):233-242.Lai Y, Wang P X, Fan G Q, et al. Genetic diversity and association analysis using SSR markers in barley[J]. Scientia Agricultura Sinica,2013,(2):233-242.
[25]郭志军,赵云雷,陈伟,等.陆地棉SSR标记遗传多样性及其与农艺性状的关联分析[J].棉花学报,2014,(5):420-430.Guo Z J, Zhao Y L, Chen W, et al. Genetic diversity and association analysis of upland cotton based on SSR markers[J]. Cotton Science,2014,(5):420-430.
[26]陈甲法.一个玉米叶色突变体的遗传研究[D].郑州:河南农业大学硕士学位论文,2009.Chen J F. Inheritance of a leaf color mutant in maize[D]. Zhengzhou:MS Thesis of Henan Agricultural University, 2009.
[27] Guo Y, Kong F M, Xu Y F, et al. QTL mapping for seedling traits in wheat grown under varying concentrations of N, P and K nutrients[J].Theoretical and Applied Genetics, 2012, 124:851-865.
[28] Xu Y, Wang R, Tong Y, et al. Mapping QTLs for yield and nitrogen-related traits in wheat:influence of nitrogen and phosphorus fertilization on QTL expression[J]. Theoretical and Applied Genetics,2014, 127:59-72.
[29]宫晓平.小麦钾养分效率及产量相关性状的QTL分析[D].山东泰安:山东农业大学硕士学位论文,2014.Gong X P. QTL mapping for K efficiency traits at seedling and yield related traits at harvest stages in wheat[D]. Tai'an, Shandong:MS Thesis of Shandong Agricultural University, 2014.
[30]兰进好,李新海,高树仁,等.不同生态环境下玉米产量性状QTL分析[J].作物学报,2005, 31:1253-1259.Lan J H, Li X H, Gao S R, et al. QTL analysis of yield components in maize under different environments[J]. Acta Agronomica Sinica,2005, 31:1253-1259.
[31]张国华,高明刚,张桂芝,等.黄淮麦区小麦品种(系)产量性状与分子标记的关联分析[J].作物学报,2013, 39(7):1187-1199.Zhang G H, Gao M G, Zhang G Z, et al.Association analysis of yield traits with molecular markers in Huang-Huai River valley winter wheat region, China[J]. Acta Agronomica Sinica, 2013, 39(7):1187-1199.
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