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AbstractIn order to evaluate, screen and identify waterloggingtolerant Brassica napus resources and provide good germplasm materials for breeding of waterloggingtolerant rape, 608 B. napus germplasm materials from different sources were identified and screened for waterloggingtolerant germplasms. The identification results showed that tested materials had dead seedling rates in the range of 0-100%, indicating that these rape germplasm materials varied extensively in waterlogging tolerance. Among the 608 materials, waterloggingtolerant materials (grade I, relative dead seedling rate≤21%) accounted for 25.49%; nontolerant materials (grade Ⅳ+V, dead seedling rate≥61%) accounted for 29.43%; and other materials had waterlogging tolerance between the two. The identification results of waterlogging tolerance in the 349 rape germplasm resource from the upper, middle and lower reaches of Yangtze River and Huang-Huai Basin showed that materials from the lower Yangtze River had the lowest average dead seedling rate of 38.8%, which was remarkably lower than materials from other ecological regions, and waterloggingtolerant materials among them accounted for 33.6%, which was also remarkably higher than materials from other ecological regions. It could thus be seen that materials from the lower Yangtze River have stronger waterlogging tolerance than those from other ecological regions overall. In this study, 24 highlytolerant B. napus germplasms (dead seedling rate<1%) were selected from the 608 B. napus germplasm materials, which could be used for breeding of waterloggingtolerant rape and related fundamental research.
Key wordsBrassica napus; Identification of waterlogging tolerance; Germplasm screening
Received: May 27, 2018Accepted: September 9, 2018
Supported by National Key Research and Development Program of China (2016YFD010020210).
Yun LI (1982-), female, P. R. China, master, devoted to research about Crop Genetics and Breeding.
*Corresponding author. Email: chensong1963@126.com.
Rape is one of the oil crops widely planted in the world. The planting area and yield of rape in China rank the first in the world［1］. The Yangteze River Basin is the main producing area in China, and has an area accounting for about 70% of rape planting area in China. Rape production in the middle and lower reaches of the Yangtze River is mainly conducted in the mode of ricerape rotation. Water in paddy field is hard to be drained after harvest, and causes wet damage to growth of rape planted later easily, and plus the wet and rainy weather in autumn and winter, waterlogging and damage or crop failure caused by waterlogging happen easily in seedling stage and thus seriously influence rape yield［2］. Researches have shown that waterlogging causes oxygen deficit in rhizosphere of rape, which leads to aggravated anaerobic metabolism, and harmful substances including ethanol, lactic acid and oxygen radical produced in glycolysis, alcoholic fermentation and lactic acid fermentation harm cells and cause greatlyreduced plant photosynthesis and inhibited growth［3］. Waterlogging also would influence plant height, stem diameter, root diameter, root length, green leaf number, leaf area and dry weight of rape, and cause decreases in number of effective branches, number of pods per plant and grains per pod, and grain yield might be reduced to 17.0%-42.4%［4-6］. In recent years, there have been many studies on screening of waterloggingtolerant rape resources. Zhang et al.［2］ studied the difference in waterlogging tolerance of 9 B. napus varieties (lines) having different genetic backgrounds through indoor waterlogging treatment on germinated seeds. The results showed that different rape varieties (lines) had larger genetic differences on wet damage. Chen et al.［15］ studied the 32 rape varieties from Yangtze River Basin through simulated wet damage environment, and the results showed that the wet tolerance in B. napus was controlled by genotypes and had greater genetic differences. And, specifically, there were 6 varieties with stronger wet tolerance, accounting for 18.75% of all the varieties, and 3 varieties with poorer wet tolerance, accounting for 9.38% of all the varieties, and other varieties had moderate wet tolerance. Xu et al.［14］ evaluated waterlogging tolerance in 270 rape resources through waterlogging treatment at germination stage according to qualified seedling percentage. The results showed that waterlogging tolerance in rape had extensive genetic variation, and different types of rape varieties differed remarkably in waterlogging tolerance. Generally, B. campestris has waterlogging tolerance stronger than B. napus; ordinary varieties are stronger in waterlogging tolerance than hybrid varieties; rape varieties from the lower Yangtze River have strong waterlogging tolerance than those from the middle and upper reaches of the Yangtze River, while the genetic variation of waterlogging tolerance in rape from the middle Yangtze River was richer than the lower Yangtze River; and the waterlogging tolerance in spring rape is stronger than that in winter rape. Many research results have consistently shown that there are differences in waterlogging tolerance between rape germplasms from different ecological regions, which provides a theoretical basis for the screening and breeding of waterloggingtolerant germplasms.
"Rapid screening method on waterlogging tolerance in rapeseed at seedling stage" is a method established by Li et al.［10］ for rapidly accurately identifying waterlogging tolerance in rapeseed seedlings. This method uses average dead seedling rate of rape subjected to 6 d of waterlogging treatment (totally immersed) on the 7th day after drainage as the identification index of waterlogging tolerance in rape seedlings. Their study showed that the method could identify tolerant rape with a dead seedling rate of 0 and rape very susceptible to waterlogging with a dead seedling rate of 100%, which had the morphologies and physiological and biochemical indices satisfying the characteristics of tolerance and nontolerant rape, respectively, indicating that method is simple, convenient and reliable［10］. In this study, 608 rape germplasms from different areas were identified by the method. The results showed that B. napus germplasm materials from different sources acted differently after the waterlogging treatment, and waterlogging tolerance varied extensively in the materials, which had the average dead seedlings in the range of 0-100%. The germplasms from different ecological regions differed remarkably in waterlogging tolerance, which accords with reported research results. In this study, the rape germplasms subjected to 6 d of waterlogging treatment were divided into five grades grade I-V (≤20.0%, 20.1%-40.0%, 40.1%-60.0%, 60.1%-80.0% and 80.1%-100.0%) according to average dead seedling rate, and further classified to three types, i.e., waterlogging tolerant: grade (I), moderately tolerant: grade II-IV, and nontolerant: grade (V), which is beneficial to the classification of experimental materials and could better reflect waterlogging tolerance in materials. The materials with a dead seedling rate<1% were deemed to be highly tolerant to waterlogging. These highlytolerant materials could serve as parents for breeding. In this study, 24 highlytolerant resource materials were identified, among which 13 were introduced foreign rape germplams, and 11 were Chinese germplasms. The 11 Chinese materials are mostly rape varieties bred in the lower Yangtze River including Jiangsu, which have very good utilization value in rape breeding for waterlogging. References
［1］ YIN Y, WANG HZ. Chinese rapeseed production status and development trend［J］.Agricultural Outlook, 2011, 1: 43-45. (in Chinese)
［2］ ZHANG XK, CHEN H, WANG HZ, et al. Genetic difference of waterlogging tolarance in rapeseed(Brassica napus L.)［J］.Chinese Journal of Oil Crop Sciences, 2007, 29(2): 204-208. (in Chinese)
［3］ DENNIS ES, DOLFERUS R, ELLIS M, et al. Molecular strategies for improving waterlogging tolerance in plants［J］. Journal of Experimental Botany, 2000, 51(342): 89-97.
［4］ LEUL M, ZHOU WJ.Alleviation of waterlogging damage in winter rape by application of uniconazole effects on morphological characteristics, hormones and photosynthesis［J］. Field Crops Research, 1998, (59): 121-127.
［5］ GUTIERREZ BFH, LAVADO RS, PORCELLI CA.Note on the effects of winter and spring waterlogging oil growth, chemical composition and yield of rapeseed［J］. Field Crops Research, 1996, 47: 175-179.
［6］ ZHOU WJ, LIN XQ. Effects of waterlogging at different growth stages on physiological characteristics and seed yield of winter rape (Brassica napus L.) ［J］. Field Crops Research, 1995, 44: 103-110.
［7］ LONG GQ. Hazard of waterlogging to rape and its prevention［J］. Hunan Agriculture, 2010, 12: 15. (in Chinese)
［8］ YI SL, XU HZ, LIAO SM, et al. Research Progress in Waterlogging and Prevention Measure on Oilseed rape (Brassica campestris L.)［J］. South China Agriculture, 6(1): 83-85. (in Chinese)
［9］ LIANG JQ, LIANG Y. Effect of plant growth substances on waterlogging resistance of oilseed rape seedling［J］. Journalof Southwest China Normal University: Natural Science Edition, 2009, 34(1): 58-62. (in Chinese)
［10］ LI Y, FU SX, QI CK. Screening method on waterlogging tolerance of through morphology and physiology verification［J］. Chinese Journal of Oil Crop Sciences, 2012, 34(3): 256-261. (in Chinese)
［11］ LU CM, GONG XM. Study on the genetic parameters of germination characteristics of rapeseeds (Brassica napus L.)［J］. Acta Agriculturae Universitis Jiangxiensis, 1989, 11(4): 23-26. (in Chinese)
［12］ ZHANG XH, LI HJ, ZHANG JF, et al. Preliminary evaluation and principle component analysis of waterlogging tolerance in Brassica napus L.［J］. Chinese Journal of Oil Crop Sciences, 2011, 33(2)：098-103. (in Chinese)
［13］ LI HJ, CAI L, PU XB, et al. Genetic difference of waterlogging tolerance in Brassica napus L. after submerging seeds in room and plants in field［J］. Southwest China Journal of Agricultural Sciences, 2016, 29(6): 1250-1256. (in Chinese) ［14］ XU J, ZENG L, XU MY, et al. Screening and evaluation of waterlogging tolerance in Brassica germplasm resources［J］. Chinese Journal of Oil Crop Sciences, 2014, 36(6): 748-754. (in Chinese)
［15］ CHEN JN, LIANG Y. Determining rapeseed tolerance to waterlogging at seedling stage in the Yangtze River basin［J］. Chinese Journal of EcoAgriculture, 2011, 19(3): 626-630. (in Chinese)
［16］ CHEN J, ZHANG XK, CHEN L. Evaluation of waterlogging tolerance germplasm in rapeseed (Brassica napus L.) with germinated seeds with anoxic stress［J］. Chinese Journal of Oil Crop Sciences, 2006, 28(2): 138-143. (in Chinese)
［17］ XUE YC, LI JN, LIU LZ, et al. Verification and identification of waterlogging tolerance in Brassica napus mutated by EMS［J］. Journalof Southwest China Normal University: Natural Science Edition, 2012, 37(4): 76-80. (in Chinese)
［18］ LI Z, PU YY, GAO CB. Evaluation of waterlogging tolerance in rapeseed (Brassica napus L.) DH lines at seedling stage［J］. Scientia Agricultura Sinica, 2010, 43(2): 286-292. (in Chinese)
［19］ ZHANG XK, CHEN H, WANG HZ, et al. Genetic difference of waterlogging tolarance in rapeseed (Brassica napus L.)［J］. Chinese Journal of Oil Crop Sciences, 2007, 29(2): 204-208. (in Chinese)
［20］ CONG Y, CHEN Y, ZOU CS, et al. Genetic analysis of waterlogging tolerance for germinated seeds of rapeseed (Brassica napus L.) with mixed model of major gene plus polygene［J］. Acta Agronomica Sinica, 2009, 35(8): 1462-1467. (in Chinese)
［21］ JIN Y, LYU YY, FU SX, et al. Mapping QTLs for waterlogging tolerance in Brassica napus L［J］. Jiangsu Journal of Agricultural Sciences, 2014, 30(6): 1253-1258. (in Chinese)
［22］ TAN YY, CHEN Y, ZHENG PY, et al. Research progress on waterlogging resistance in rapeseed (Brassica napus L.)［J］. Chinese Journal of Oil Crop Sciences, 2011, 33(3): 306-310. (in Chinese)
［23］ HE JG, GUANG CY. Study on waterlogging tolerance in rape［J］. Crop Research, 2009, 23(5): 323-327. (in Chinese)
［24］ LI HJ, ZHANG XH, PU XB, et al. Physiological reaction of field waterlogging treatment in Brassica napus L［J］. Southwest China Journal of Agricultural Sciences, 2013, 26(1): 84-88. (in Chinese)
［25］ LYU YY, FU SX, CHEN S, et al. Cloning of BnADH3 gene from Brassica napus L. and submergence tolerance of BnADH3 transgenic Arabidopsis［J］. Acta Agronomica Sinica, 2015, 41(4): 565-573. (in Chinese)
Editor: Yingzhi GUANGProofreader: Xinxiu ZHU
Key wordsBrassica napus; Identification of waterlogging tolerance; Germplasm screening
Received: May 27, 2018Accepted: September 9, 2018
Supported by National Key Research and Development Program of China (2016YFD010020210).
Yun LI (1982-), female, P. R. China, master, devoted to research about Crop Genetics and Breeding.
*Corresponding author. Email: chensong1963@126.com.
Rape is one of the oil crops widely planted in the world. The planting area and yield of rape in China rank the first in the world［1］. The Yangteze River Basin is the main producing area in China, and has an area accounting for about 70% of rape planting area in China. Rape production in the middle and lower reaches of the Yangtze River is mainly conducted in the mode of ricerape rotation. Water in paddy field is hard to be drained after harvest, and causes wet damage to growth of rape planted later easily, and plus the wet and rainy weather in autumn and winter, waterlogging and damage or crop failure caused by waterlogging happen easily in seedling stage and thus seriously influence rape yield［2］. Researches have shown that waterlogging causes oxygen deficit in rhizosphere of rape, which leads to aggravated anaerobic metabolism, and harmful substances including ethanol, lactic acid and oxygen radical produced in glycolysis, alcoholic fermentation and lactic acid fermentation harm cells and cause greatlyreduced plant photosynthesis and inhibited growth［3］. Waterlogging also would influence plant height, stem diameter, root diameter, root length, green leaf number, leaf area and dry weight of rape, and cause decreases in number of effective branches, number of pods per plant and grains per pod, and grain yield might be reduced to 17.0%-42.4%［4-6］. In recent years, there have been many studies on screening of waterloggingtolerant rape resources. Zhang et al.［2］ studied the difference in waterlogging tolerance of 9 B. napus varieties (lines) having different genetic backgrounds through indoor waterlogging treatment on germinated seeds. The results showed that different rape varieties (lines) had larger genetic differences on wet damage. Chen et al.［15］ studied the 32 rape varieties from Yangtze River Basin through simulated wet damage environment, and the results showed that the wet tolerance in B. napus was controlled by genotypes and had greater genetic differences. And, specifically, there were 6 varieties with stronger wet tolerance, accounting for 18.75% of all the varieties, and 3 varieties with poorer wet tolerance, accounting for 9.38% of all the varieties, and other varieties had moderate wet tolerance. Xu et al.［14］ evaluated waterlogging tolerance in 270 rape resources through waterlogging treatment at germination stage according to qualified seedling percentage. The results showed that waterlogging tolerance in rape had extensive genetic variation, and different types of rape varieties differed remarkably in waterlogging tolerance. Generally, B. campestris has waterlogging tolerance stronger than B. napus; ordinary varieties are stronger in waterlogging tolerance than hybrid varieties; rape varieties from the lower Yangtze River have strong waterlogging tolerance than those from the middle and upper reaches of the Yangtze River, while the genetic variation of waterlogging tolerance in rape from the middle Yangtze River was richer than the lower Yangtze River; and the waterlogging tolerance in spring rape is stronger than that in winter rape. Many research results have consistently shown that there are differences in waterlogging tolerance between rape germplasms from different ecological regions, which provides a theoretical basis for the screening and breeding of waterloggingtolerant germplasms.
"Rapid screening method on waterlogging tolerance in rapeseed at seedling stage" is a method established by Li et al.［10］ for rapidly accurately identifying waterlogging tolerance in rapeseed seedlings. This method uses average dead seedling rate of rape subjected to 6 d of waterlogging treatment (totally immersed) on the 7th day after drainage as the identification index of waterlogging tolerance in rape seedlings. Their study showed that the method could identify tolerant rape with a dead seedling rate of 0 and rape very susceptible to waterlogging with a dead seedling rate of 100%, which had the morphologies and physiological and biochemical indices satisfying the characteristics of tolerance and nontolerant rape, respectively, indicating that method is simple, convenient and reliable［10］. In this study, 608 rape germplasms from different areas were identified by the method. The results showed that B. napus germplasm materials from different sources acted differently after the waterlogging treatment, and waterlogging tolerance varied extensively in the materials, which had the average dead seedlings in the range of 0-100%. The germplasms from different ecological regions differed remarkably in waterlogging tolerance, which accords with reported research results. In this study, the rape germplasms subjected to 6 d of waterlogging treatment were divided into five grades grade I-V (≤20.0%, 20.1%-40.0%, 40.1%-60.0%, 60.1%-80.0% and 80.1%-100.0%) according to average dead seedling rate, and further classified to three types, i.e., waterlogging tolerant: grade (I), moderately tolerant: grade II-IV, and nontolerant: grade (V), which is beneficial to the classification of experimental materials and could better reflect waterlogging tolerance in materials. The materials with a dead seedling rate<1% were deemed to be highly tolerant to waterlogging. These highlytolerant materials could serve as parents for breeding. In this study, 24 highlytolerant resource materials were identified, among which 13 were introduced foreign rape germplams, and 11 were Chinese germplasms. The 11 Chinese materials are mostly rape varieties bred in the lower Yangtze River including Jiangsu, which have very good utilization value in rape breeding for waterlogging. References
［1］ YIN Y, WANG HZ. Chinese rapeseed production status and development trend［J］.Agricultural Outlook, 2011, 1: 43-45. (in Chinese)
［2］ ZHANG XK, CHEN H, WANG HZ, et al. Genetic difference of waterlogging tolarance in rapeseed(Brassica napus L.)［J］.Chinese Journal of Oil Crop Sciences, 2007, 29(2): 204-208. (in Chinese)
［3］ DENNIS ES, DOLFERUS R, ELLIS M, et al. Molecular strategies for improving waterlogging tolerance in plants［J］. Journal of Experimental Botany, 2000, 51(342): 89-97.
［4］ LEUL M, ZHOU WJ.Alleviation of waterlogging damage in winter rape by application of uniconazole effects on morphological characteristics, hormones and photosynthesis［J］. Field Crops Research, 1998, (59): 121-127.
［5］ GUTIERREZ BFH, LAVADO RS, PORCELLI CA.Note on the effects of winter and spring waterlogging oil growth, chemical composition and yield of rapeseed［J］. Field Crops Research, 1996, 47: 175-179.
［6］ ZHOU WJ, LIN XQ. Effects of waterlogging at different growth stages on physiological characteristics and seed yield of winter rape (Brassica napus L.) ［J］. Field Crops Research, 1995, 44: 103-110.
［7］ LONG GQ. Hazard of waterlogging to rape and its prevention［J］. Hunan Agriculture, 2010, 12: 15. (in Chinese)
［8］ YI SL, XU HZ, LIAO SM, et al. Research Progress in Waterlogging and Prevention Measure on Oilseed rape (Brassica campestris L.)［J］. South China Agriculture, 6(1): 83-85. (in Chinese)
［9］ LIANG JQ, LIANG Y. Effect of plant growth substances on waterlogging resistance of oilseed rape seedling［J］. Journalof Southwest China Normal University: Natural Science Edition, 2009, 34(1): 58-62. (in Chinese)
［10］ LI Y, FU SX, QI CK. Screening method on waterlogging tolerance of through morphology and physiology verification［J］. Chinese Journal of Oil Crop Sciences, 2012, 34(3): 256-261. (in Chinese)
［11］ LU CM, GONG XM. Study on the genetic parameters of germination characteristics of rapeseeds (Brassica napus L.)［J］. Acta Agriculturae Universitis Jiangxiensis, 1989, 11(4): 23-26. (in Chinese)
［12］ ZHANG XH, LI HJ, ZHANG JF, et al. Preliminary evaluation and principle component analysis of waterlogging tolerance in Brassica napus L.［J］. Chinese Journal of Oil Crop Sciences, 2011, 33(2)：098-103. (in Chinese)
［13］ LI HJ, CAI L, PU XB, et al. Genetic difference of waterlogging tolerance in Brassica napus L. after submerging seeds in room and plants in field［J］. Southwest China Journal of Agricultural Sciences, 2016, 29(6): 1250-1256. (in Chinese) ［14］ XU J, ZENG L, XU MY, et al. Screening and evaluation of waterlogging tolerance in Brassica germplasm resources［J］. Chinese Journal of Oil Crop Sciences, 2014, 36(6): 748-754. (in Chinese)
［15］ CHEN JN, LIANG Y. Determining rapeseed tolerance to waterlogging at seedling stage in the Yangtze River basin［J］. Chinese Journal of EcoAgriculture, 2011, 19(3): 626-630. (in Chinese)
［16］ CHEN J, ZHANG XK, CHEN L. Evaluation of waterlogging tolerance germplasm in rapeseed (Brassica napus L.) with germinated seeds with anoxic stress［J］. Chinese Journal of Oil Crop Sciences, 2006, 28(2): 138-143. (in Chinese)
［17］ XUE YC, LI JN, LIU LZ, et al. Verification and identification of waterlogging tolerance in Brassica napus mutated by EMS［J］. Journalof Southwest China Normal University: Natural Science Edition, 2012, 37(4): 76-80. (in Chinese)
［18］ LI Z, PU YY, GAO CB. Evaluation of waterlogging tolerance in rapeseed (Brassica napus L.) DH lines at seedling stage［J］. Scientia Agricultura Sinica, 2010, 43(2): 286-292. (in Chinese)
［19］ ZHANG XK, CHEN H, WANG HZ, et al. Genetic difference of waterlogging tolarance in rapeseed (Brassica napus L.)［J］. Chinese Journal of Oil Crop Sciences, 2007, 29(2): 204-208. (in Chinese)
［20］ CONG Y, CHEN Y, ZOU CS, et al. Genetic analysis of waterlogging tolerance for germinated seeds of rapeseed (Brassica napus L.) with mixed model of major gene plus polygene［J］. Acta Agronomica Sinica, 2009, 35(8): 1462-1467. (in Chinese)
［21］ JIN Y, LYU YY, FU SX, et al. Mapping QTLs for waterlogging tolerance in Brassica napus L［J］. Jiangsu Journal of Agricultural Sciences, 2014, 30(6): 1253-1258. (in Chinese)
［22］ TAN YY, CHEN Y, ZHENG PY, et al. Research progress on waterlogging resistance in rapeseed (Brassica napus L.)［J］. Chinese Journal of Oil Crop Sciences, 2011, 33(3): 306-310. (in Chinese)
［23］ HE JG, GUANG CY. Study on waterlogging tolerance in rape［J］. Crop Research, 2009, 23(5): 323-327. (in Chinese)
［24］ LI HJ, ZHANG XH, PU XB, et al. Physiological reaction of field waterlogging treatment in Brassica napus L［J］. Southwest China Journal of Agricultural Sciences, 2013, 26(1): 84-88. (in Chinese)
［25］ LYU YY, FU SX, CHEN S, et al. Cloning of BnADH3 gene from Brassica napus L. and submergence tolerance of BnADH3 transgenic Arabidopsis［J］. Acta Agronomica Sinica, 2015, 41(4): 565-573. (in Chinese)
Editor: Yingzhi GUANGProofreader: Xinxiu ZHU