油菜抗寒性和寒旱区油菜育种探析

doi:10.7668/hbnxb.20194305

摘要/Abstract

摘要：

为了能够更加清晰地了解我国寒旱区冬油菜抗寒性研究进展,对寒旱区冬油菜育种与生产提供理论支撑。从选择指标与方法、遗传规律、选育方法、筛选效果、种质分类、分子机理研究等6个方面和选育方法、育成品种、品种的抗寒性分级和品种的适应性筛选等4个方面分别对冬油菜的抗寒性、抗寒性品种的研究进展进行了归纳总结。在分析研究存在的问题的基础上,提出了多途径创制种质资源、综合评价筛选出核心种质、开展抗寒机理研究和分子设计育种创新体系建设、构建育种体系多途径开展抗寒性品种选育、综合筛选提高品种的适应性、推行配套栽培技术促进品种产量水平的发挥提高效益等6项措施,以期促进我国寒旱区油菜的发展。

关键词: 油菜育种, 寒旱区, 研究进展, 存在问题, 发展建议

Abstract:

In order to have a clearer understanding of the research progress on cold resistance of winter rapeseed in cold and arid regions of China,and to provide theoretical support for breeding and production.The research progress of cold-resistance and cold-resistance breeding of winter rapeseed were summarized respectively from six aspects such as selection index and method,genetic law,breeding method,screening effect,germplasm classification and molecular mechanism research,and four aspects such as breeding method,breeding varieties,cold resistance classification and adaptability screening of varieties.Based on the analysis of the existing problems,six measures were put forward,such as creating germplasm resources by multiple ways,screening out core germplasm by comprehensive evaluation,carrying out research on cold-resistance mechanism and construction of innovation system of molecular design breeding,constructing breeding system and carrying out cold-resistance breeding by multiple ways,improving the adaptability of varieties by comprehensive screening,promoting varieties yield level and improving benefits by carrying out matching cultivation techniques,in order to promote the development of rapeseed production in cold and arid regions of our country.

Key words: Rape breeding, Cold and arid region, Research progress, Existing problems, Development suggestions

李积铭, 张耀文, 郭安强, 翟兰菊, 李和平, 李爱国. 油菜抗寒性和寒旱区油菜育种探析[J]. 华北农学报, 2023, 38(S1): 131-144. doi: 10.7668/hbnxb.20194305.

LI Jiming, ZHANG Yaowen, GUO Anqiang, ZHAI Lanju, LI Heping, LI Aiguo. Advances in Research on Cold Resistance of Rapeseed and Analysis of Rapeseed Breeding in Cold and Dry Regions of China[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(S1): 131-144. doi: 10.7668/hbnxb.20194305.

http://www.hbnxb.net/CN/Y2023/V38/IS1/131

图/表 8

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指标类型 Indicators classification		具体指标与参考文献 Indicators and references
形态指标	叶	(心叶)叶色^[7-8]、叶色^[4,7,9-10]、质量^[3,10-11]、含水量^{[8,12⇓⇓-15]}、厚度^[2,8,16]、叶柄长度^[10]
Morphological indexes	根	质量^{[3,11,16⇓⇓-19]}、根长^[3,20]、根颈直径^{[3,6,16⇓-18,20-21]}、根颈体积^[8]、含水量^[6,8]、根颈下陷度^[8]、根/冠比^{[3,4,6,8,11,18]}、冻害等级^[12]
	群体	生长点形状^[3,9]、下胚轴长度^[22]、生长锥相对高度^[22]、苗期生长习性^{[3,7⇓-9,16-17,23]}、冬春性^[7]、枯叶期^[8,24]、(春播)春化率^{[17-18,22,24-25]}、枯叶/全叶比^[8]、半致死温度^{[10,16,22,25]}、耐迟播性^[26]、花芽分化时间^[26]、株(薹)高^{[20,22,25,27]}
		根系活力^[19]、抗旱性^[19]
组织结构 Organizational structure indexes		叶片紧密组织致密度^[8,28]、叶片疏松海绵组织指数^[28]、栅栏组织厚度^[2,16]、栅栏组织/海绵组织比^[2,16,29]、叶片组织细胞结构疏松度^[30]、保卫细胞叶绿体数目^[18]
气孔特性 Stomata characteristic index		气孔大小^[2]、气孔面积^[15-16]、气孔密度^[2,8,30-31]、气孔开张度^[8,18]、气孔开张比^[18]、单位面积气孔总长度^[30]
质膜受伤害程度 Degree of plasma	叶	丙二醛^{[8-9,12⇓⇓-15,20,22-23,29,32⇓⇓-35]}、相对电导率^{[8-9,12⇓-14,34,36]}、质膜不饱和脂肪酸含量^[37]、与饱和脂肪酸含量比值^[37]、ATP酶活性^[37]
membrane damage indexes	变化量	丙二醛^[33]
	根	丙二醛^[35-36]、相对电导率^[33]
渗透调节物质 Osmotic regulating	叶含量	可溶性蛋白^{[8-9,15,17-18,22-23,25,28-29,32⇓⇓-35,38⇓⇓-41]}、可溶性糖^{[8-9,11,13-14,22,28-29,32,34,42]}、脯氨酸^{[8-9,14,22-23,28,30]}、Ca²⁺含量^[11]
substance indexes	根含量	可溶性蛋白^[6,8,36]、可溶性糖^[8,36]、脯氨酸^[8]、Ca²⁺含量^[11]
	叶变化量	可溶性蛋白^[2,16,38]、脯氨酸^[16]
内源激素	叶含量	脱落酸^{[24,41,43-44]}、赤霉素^[24]
Endogenous hormones indexes	比值	脱落酸/赤霉素比值^[24]
	根含量	脱落酸^[41]
抗氧化保护酶 Protective enzyme indexes	叶中的活性	过氧化物酶^{[8-9,15,17,20,22-23,25,28,32,34⇓-36,38⇓⇓-41,45-46]}、超氧化物歧化酶^{[8-9,15,22-23,25,28⇓-30,32⇓⇓⇓-36,40-41,45]}、过氧化氢酶^{[8-9,15,17,20,22-23,25,29,38,32⇓⇓⇓-36,40-41,45]}、抗坏血酸过氧化物酶^[6,30,39]、过氧化物酶同工酶^[2]
	根中的活性	过氧化物酶^[6,8,36]、超氧化物歧化酶^[6,8,36]、过氧化氢酶^[6,8,36]、抗坏血酸过氧化物酶^[6,8,40,42]、谷胱甘肽还原酶^[42]、铜锌超氧化物酶^[42]
	叶变化量	过氧化物酶^[2,16,38]、过氧化氢酶^[2,16,38]、超氧化物歧化酶^[38]
光合生理 Photosynthetic physiological	色素	叶绿素含量^{[17,22,29,34]}、叶SPAD值^[12]、叶绿素a含量^[8,22]、类胡萝卜素含^[8,22]、叶绿素分解速率^[29,36,40]
index	性能	净光合速率^{[11,17-18,29,34-35,41,47]}、气孔导度^{[17,29,34-35,41]}、蒸腾速率^{[17,29,34-35,41,47]}、胞间二氧化碳浓度^{[17,29,35,41]}、初始荧光产量^[41]、最大荧光产量^[41]、PSⅡ潜在光化学效率^[41]、PSⅡ电子传递^[41]
	变化	净光合速率^[46]、气孔导度^[46]、胞间二氧化碳浓度^[46]

指标类型 Indicators classification		具体指标与参考文献 Indicators and references
形态指标	叶	(心叶)叶色^[7-8]、叶色^[4,7,9-10]、质量^[3,10-11]、含水量^{[8,12⇓⇓-15]}、厚度^[2,8,16]、叶柄长度^[10]
Morphological indexes	根	质量^{[3,11,16⇓⇓-19]}、根长^[3,20]、根颈直径^{[3,6,16⇓-18,20-21]}、根颈体积^[8]、含水量^[6,8]、根颈下陷度^[8]、根/冠比^{[3,4,6,8,11,18]}、冻害等级^[12]
	群体	生长点形状^[3,9]、下胚轴长度^[22]、生长锥相对高度^[22]、苗期生长习性^{[3,7⇓-9,16-17,23]}、冬春性^[7]、枯叶期^[8,24]、(春播)春化率^{[17-18,22,24-25]}、枯叶/全叶比^[8]、半致死温度^{[10,16,22,25]}、耐迟播性^[26]、花芽分化时间^[26]、株(薹)高^{[20,22,25,27]}
		根系活力^[19]、抗旱性^[19]
组织结构 Organizational structure indexes		叶片紧密组织致密度^[8,28]、叶片疏松海绵组织指数^[28]、栅栏组织厚度^[2,16]、栅栏组织/海绵组织比^[2,16,29]、叶片组织细胞结构疏松度^[30]、保卫细胞叶绿体数目^[18]
气孔特性 Stomata characteristic index		气孔大小^[2]、气孔面积^[15-16]、气孔密度^[2,8,30-31]、气孔开张度^[8,18]、气孔开张比^[18]、单位面积气孔总长度^[30]
质膜受伤害程度 Degree of plasma	叶	丙二醛^{[8-9,12⇓⇓-15,20,22-23,29,32⇓⇓-35]}、相对电导率^{[8-9,12⇓-14,34,36]}、质膜不饱和脂肪酸含量^[37]、与饱和脂肪酸含量比值^[37]、ATP酶活性^[37]
membrane damage indexes	变化量	丙二醛^[33]
	根	丙二醛^[35-36]、相对电导率^[33]
渗透调节物质 Osmotic regulating	叶含量	可溶性蛋白^{[8-9,15,17-18,22-23,25,28-29,32⇓⇓-35,38⇓⇓-41]}、可溶性糖^{[8-9,11,13-14,22,28-29,32,34,42]}、脯氨酸^{[8-9,14,22-23,28,30]}、Ca²⁺含量^[11]
substance indexes	根含量	可溶性蛋白^[6,8,36]、可溶性糖^[8,36]、脯氨酸^[8]、Ca²⁺含量^[11]
	叶变化量	可溶性蛋白^[2,16,38]、脯氨酸^[16]
内源激素	叶含量	脱落酸^{[24,41,43-44]}、赤霉素^[24]
Endogenous hormones indexes	比值	脱落酸/赤霉素比值^[24]
	根含量	脱落酸^[41]
抗氧化保护酶 Protective enzyme indexes	叶中的活性	过氧化物酶^{[8-9,15,17,20,22-23,25,28,32,34⇓-36,38⇓⇓-41,45-46]}、超氧化物歧化酶^{[8-9,15,22-23,25,28⇓-30,32⇓⇓⇓-36,40-41,45]}、过氧化氢酶^{[8-9,15,17,20,22-23,25,29,38,32⇓⇓⇓-36,40-41,45]}、抗坏血酸过氧化物酶^[6,30,39]、过氧化物酶同工酶^[2]
	根中的活性	过氧化物酶^[6,8,36]、超氧化物歧化酶^[6,8,36]、过氧化氢酶^[6,8,36]、抗坏血酸过氧化物酶^[6,8,40,42]、谷胱甘肽还原酶^[42]、铜锌超氧化物酶^[42]
	叶变化量	过氧化物酶^[2,16,38]、过氧化氢酶^[2,16,38]、超氧化物歧化酶^[38]
光合生理 Photosynthetic physiological	色素	叶绿素含量^{[17,22,29,34]}、叶SPAD值^[12]、叶绿素a含量^[8,22]、类胡萝卜素含^[8,22]、叶绿素分解速率^[29,36,40]
index	性能	净光合速率^{[11,17-18,29,34-35,41,47]}、气孔导度^{[17,29,34-35,41]}、蒸腾速率^{[17,29,34-35,41,47]}、胞间二氧化碳浓度^{[17,29,35,41]}、初始荧光产量^[41]、最大荧光产量^[41]、PSⅡ潜在光化学效率^[41]、PSⅡ电子传递^[41]
	变化	净光合速率^[46]、气孔导度^[46]、胞间二氧化碳浓度^[46]

与抗寒性的相关性 Correlation with cold resistance		具体指标与参考文献 Indicators and references
正相关 Index positively related to cold resistance	极显著	(心叶)叶色^[7,10]、叶SPAD值^[12]、根质量^[11,16-17]、根颈直径^[3,16,18,20]、根长^[20]、根/冠比^[3,11,18]、生长点形状^[3]、苗期生长习性^[7]、气孔开张度^[8,18]、叶可溶性蛋白^{[17-18,20,25,29,33,35,38]}、叶可溶性糖^{[11,13-14,20,29,39]}、叶脯氨酸^[9,14-15,23]、叶可溶性蛋白变化量^[16,38]、叶脯氨酸变化量^[16]、叶过氧化物酶^{[15,17,20,23,25,29,35,38,40,46]}、叶超氧化物歧化酶^{[9,15,23,25,29,33,38,40,46]}、叶过氧化氢酶^{[9,15,17,20,23,25,29,33-34,36,38,45]}、叶过氧化物酶变化量^[38]、叶超氧化物歧化酶变化量^[38]、叶过氧化氢酶变化量^[38]、叶抗坏血酸过氧化物酶^[40]、叶绿素含量^[36]、净光合速率^[32,34,41]、蒸腾速率^[34]、初始荧光产量^[41]
	显著	叶厚度^[2,8]、根质量^[3,18]、根长^[20]、根颈直径^[17,20-21]、根/冠比^[8]、根颈下陷度^[8]、生长点形状^[9]、苗期生长习性^[9,16]、冬春性^[7]、耐迟播性^[26]、叶组织结构致密度^[28]、保卫细胞叶绿体数量^[18]、气孔大小^[2]、气孔面积^[16]、气孔开张度^[8]、气孔开张比^[18]、叶可溶性蛋白^{[9,22-23,28,32,34,40]}、叶可溶性糖^[22]、叶脯氨酸^[8,22,28,39]、根可溶性蛋白^[6,8,42]、根可溶性糖^[8,42]、根脯氨酸^[8]、叶可溶性蛋白变化量^[2]、叶脱落酸^[24]、根脱落酸^[43]、叶过氧化物酶^{[9,22,28,32,34,40-41]}、叶超氧化物歧化酶^{[22,25,28,30,41]}、叶过氧化氢酶^[22,41]、叶抗坏血酸过氧化物酶^[22]、叶过氧化物酶同工酶^[2]、根过氧化氢酶^[6,8,36,42]、根超氧化物歧化酶^[6,8]、根抗坏血酸过氧化物酶^[6,8,42]、根谷胱甘肽还原酶^[42]、根铜锌超过氧化物酶^[42]、叶过氧化物酶变化量^[16]、叶绿素含量^[22,29]、叶绿素a含量^[22]、类胡萝卜素含量^[8,22]、叶绿素分解速率^[29,41-42]、净光合速率^{[11,18,29,47]}、气孔导度^[29,34,41]、蒸腾速率^[29,47]、PSⅡ潜在光化学效率^[41]、PSⅡ电子传递^[41]、最大荧光产量^[40]、根Ca²⁺含量^[11]质膜不饱和脂肪酸含量^[37]、与饱和脂肪酸含量比值^[37]、ATP酶活性^[37]
	正相关	(心叶)叶色^[8]、叶色^[7,9]、苗期生长习性^[17]、枯叶/全叶比^[8]、根颈直径^[6]、根/冠比^[6]、叶组织结构致密度^[8,28]、叶可溶性蛋白^[8]、叶可溶性糖^[8-9,32,34]、叶脱落酸^[43]、叶超氧化物歧化酶^[32,34,40]、叶过氧化氢酶^{[32,34-35,40]}、根过氧化氢酶^[6]、根抗坏血酸过氧化物酶^[6]、叶绿素a含量^[8]、净光合速率^[17,35]、气孔导度^[17,35,41]、蒸腾速率^[41]、胞间二氧化碳浓度^[17]、净光合速率变化值^[46]、气孔导度变化值^[46]
负相关 Index negatively related to cold resistance	极显著	叶质量^[3,11,18]、叶含水量^[13⇓-15]、叶厚度^[16]、根含水量^[14]、苗期生长习性^[3,23]、(春播)春化率^{[10,17,20,22,25,27]}、半致死温度^[16,22,25]、株(薹)高^{[20,22,25,27]}、冻害等级^[12]、栅栏组织厚度^[16]、气孔密度^[8]、叶丙二醛^{[9,13⇓-15,20,23,29,33-34,38]}、叶丙二醛变化量^[38]、叶相对电导率^[9,13-14,34]、根丙二醛^[42]、叶可溶性糖^[11]、叶Ca²⁺含量^[11]、叶柄长度^[10]、生长锥相对高度^[22]、下胚轴长度^[22]
	显著	叶含水量^[8,12]、根含水量^[6,8]、枯叶期^[8,24]、花芽分化时间^[26]、栅栏组织厚度^[2]、栅栏组织/海绵组织比^[2,16,30]、气孔面积^[16]、气孔开张度^[8]、叶丙二醛^[8,12,22]、叶相对电导率^[8,12]、根相对电导率^[33]、叶赤霉素^[24]、叶过氧化物酶变化量^[2,16]、叶过氧化氢酶变化量^[2,16]、根过氧化物酶变化量^[2]、胞间二氧化碳浓度^[29,42]
	负相关	生长习性^[3]、叶片疏松海绵组织指数^[28]、叶丙二醛^[32,34]、蒸腾速率^[17]、胞间二氧化碳浓度41^[41]、胞间二氧化碳浓度变化值^[46]

与抗寒性的相关性 Correlation with cold resistance		具体指标与参考文献 Indicators and references
正相关 Index positively related to cold resistance	极显著	(心叶)叶色^[7,10]、叶SPAD值^[12]、根质量^[11,16-17]、根颈直径^[3,16,18,20]、根长^[20]、根/冠比^[3,11,18]、生长点形状^[3]、苗期生长习性^[7]、气孔开张度^[8,18]、叶可溶性蛋白^{[17-18,20,25,29,33,35,38]}、叶可溶性糖^{[11,13-14,20,29,39]}、叶脯氨酸^[9,14-15,23]、叶可溶性蛋白变化量^[16,38]、叶脯氨酸变化量^[16]、叶过氧化物酶^{[15,17,20,23,25,29,35,38,40,46]}、叶超氧化物歧化酶^{[9,15,23,25,29,33,38,40,46]}、叶过氧化氢酶^{[9,15,17,20,23,25,29,33-34,36,38,45]}、叶过氧化物酶变化量^[38]、叶超氧化物歧化酶变化量^[38]、叶过氧化氢酶变化量^[38]、叶抗坏血酸过氧化物酶^[40]、叶绿素含量^[36]、净光合速率^[32,34,41]、蒸腾速率^[34]、初始荧光产量^[41]
	显著	叶厚度^[2,8]、根质量^[3,18]、根长^[20]、根颈直径^[17,20-21]、根/冠比^[8]、根颈下陷度^[8]、生长点形状^[9]、苗期生长习性^[9,16]、冬春性^[7]、耐迟播性^[26]、叶组织结构致密度^[28]、保卫细胞叶绿体数量^[18]、气孔大小^[2]、气孔面积^[16]、气孔开张度^[8]、气孔开张比^[18]、叶可溶性蛋白^{[9,22-23,28,32,34,40]}、叶可溶性糖^[22]、叶脯氨酸^[8,22,28,39]、根可溶性蛋白^[6,8,42]、根可溶性糖^[8,42]、根脯氨酸^[8]、叶可溶性蛋白变化量^[2]、叶脱落酸^[24]、根脱落酸^[43]、叶过氧化物酶^{[9,22,28,32,34,40-41]}、叶超氧化物歧化酶^{[22,25,28,30,41]}、叶过氧化氢酶^[22,41]、叶抗坏血酸过氧化物酶^[22]、叶过氧化物酶同工酶^[2]、根过氧化氢酶^[6,8,36,42]、根超氧化物歧化酶^[6,8]、根抗坏血酸过氧化物酶^[6,8,42]、根谷胱甘肽还原酶^[42]、根铜锌超过氧化物酶^[42]、叶过氧化物酶变化量^[16]、叶绿素含量^[22,29]、叶绿素a含量^[22]、类胡萝卜素含量^[8,22]、叶绿素分解速率^[29,41-42]、净光合速率^{[11,18,29,47]}、气孔导度^[29,34,41]、蒸腾速率^[29,47]、PSⅡ潜在光化学效率^[41]、PSⅡ电子传递^[41]、最大荧光产量^[40]、根Ca²⁺含量^[11]质膜不饱和脂肪酸含量^[37]、与饱和脂肪酸含量比值^[37]、ATP酶活性^[37]
	正相关	(心叶)叶色^[8]、叶色^[7,9]、苗期生长习性^[17]、枯叶/全叶比^[8]、根颈直径^[6]、根/冠比^[6]、叶组织结构致密度^[8,28]、叶可溶性蛋白^[8]、叶可溶性糖^[8-9,32,34]、叶脱落酸^[43]、叶超氧化物歧化酶^[32,34,40]、叶过氧化氢酶^{[32,34-35,40]}、根过氧化氢酶^[6]、根抗坏血酸过氧化物酶^[6]、叶绿素a含量^[8]、净光合速率^[17,35]、气孔导度^[17,35,41]、蒸腾速率^[41]、胞间二氧化碳浓度^[17]、净光合速率变化值^[46]、气孔导度变化值^[46]
负相关 Index negatively related to cold resistance	极显著	叶质量^[3,11,18]、叶含水量^[13⇓-15]、叶厚度^[16]、根含水量^[14]、苗期生长习性^[3,23]、(春播)春化率^{[10,17,20,22,25,27]}、半致死温度^[16,22,25]、株(薹)高^{[20,22,25,27]}、冻害等级^[12]、栅栏组织厚度^[16]、气孔密度^[8]、叶丙二醛^{[9,13⇓-15,20,23,29,33-34,38]}、叶丙二醛变化量^[38]、叶相对电导率^[9,13-14,34]、根丙二醛^[42]、叶可溶性糖^[11]、叶Ca²⁺含量^[11]、叶柄长度^[10]、生长锥相对高度^[22]、下胚轴长度^[22]
	显著	叶含水量^[8,12]、根含水量^[6,8]、枯叶期^[8,24]、花芽分化时间^[26]、栅栏组织厚度^[2]、栅栏组织/海绵组织比^[2,16,30]、气孔面积^[16]、气孔开张度^[8]、叶丙二醛^[8,12,22]、叶相对电导率^[8,12]、根相对电导率^[33]、叶赤霉素^[24]、叶过氧化物酶变化量^[2,16]、叶过氧化氢酶变化量^[2,16]、根过氧化物酶变化量^[2]、胞间二氧化碳浓度^[29,42]
	负相关	生长习性^[3]、叶片疏松海绵组织指数^[28]、叶丙二醛^[32,34]、蒸腾速率^[17]、胞间二氧化碳浓度41^[41]、胞间二氧化碳浓度变化值^[46]

遗传变异 Genetic variation		具体指标与参考文献 Indicators and references
遗传变异 Genetic variation	介于2个亲本之间	抗寒性(越冬率)^{[23,26,34,38,41,45,50-51]}、叶片厚度^[41]、根冠比^[28,50]、叶片紧密组织指数^[28]、叶片疏松海绵组织指数^[28]、气孔保卫细胞的周长^[41]、叶丙二醛含量^[23,38,41]、叶相对电导率^[34]、叶可溶性蛋白含量^{[23,28,34,41,50]}、叶游离脯氨酸含量^[28,41]、叶过氧化物酶活性^{[23,28,34,38,41,45,50]}、叶超氧化物歧化酶活性^{[23,28,38,41,45,50]}、叶过氧化氢酶活性^{[23,38,41,45,50]}、叶绿素含量^[34]、蒸腾速率^[34]
	连续变异且接近正态分布	越冬率^[40,50]、幼苗生长习性^[50]、叶片组织相对含水量^[12]、叶丙二醛含量^[12,23,50]、叶片相对电导率^[12]、叶可溶蛋白含量^{[23,28,40,50]}、叶游离脯氨酸含量^[23,50]、叶超氧化物歧化酶活性^[23,50]、叶过氧化物酶活性^[23,50]、叶过氧化氢酶活性^[23,40,50]、叶抗坏血酸过氧化物酶^[40]、叶片SPAD值^[12]
	可分为类群数	2个^[40]、3个^[40,50]、8个^[50]
杂种优势 Heterosis	中亲优势	越冬率^{[32,34,52-53]}、叶丙二醛含量^[32,34]、叶过氧化氢酶活性^[32,34]、叶超氧化物歧化酶^[32,34]
	正向超亲优势	越冬率^[34,52-53]、叶片组织相对含水量^[12]、幼苗习性^[28]、冻害等级^[12]、叶片相对电导率^[12]、叶丙二醛含量^[32,34,40]、叶过氧化氢酶活性^[32,34]、叶片SPAD值^[12]、净光合速率^[32]
	负向超亲优势	越冬率^[32,34]、叶可溶性蛋白含量^[32,34]、叶可溶性糖含量^[32,34]、叶过氧化氢酶活性^[32,34]、叶过氧化物酶活性^[32]
	超标优势	越冬率^[52-53]
控制 Genetic control	加性基因和非加性基因共同决定	越冬率^[32,34]、叶丙二醛含量^[32,34]、相对电导率^[34]、叶可溶蛋白含量^[32,34]、叶可溶性糖含量^[32,34]、叶过氧化氢酶活性^[32,34]、叶超氧化物歧化酶活性^[32,34]、叶过氧化物酶活性^[32,34]、叶绿素含量^[34]、净光合速率^[34]、蒸腾速率^[34]、气孔导度^[34]
	加性基因	越冬率^[32,34]、叶相对电导率^[34]、叶可溶性糖含量^[32,34]、叶过氧化物酶活性^[34]、叶超氧化物歧化酶活性^[34]、叶绿素含量^[34]、气孔导度^[34]、蒸腾速率^[34]
	显性基因	叶丙二醛含量^[34]、叶可溶性蛋白含量^[34]、叶过氧化氢酶活性^[34]、净光合速率^[34]

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