Research Progresses on the Drought Resistance of Medicago at Molecular Level

doi:10.13560/j.cnki.biotech.bull.1985.2021-0057

Abstract

Abstract:

Drought stress is one of the most serious environmental constraints for global agricultural production,and it causes severe losses in pasture production. In this paper,drought induction related genes and the main progresses on the breeding of 4 important Medicago plants with drought resistance were reviewed,and the prospects and issues in this field were also discussed. It is suggested that biological technology can be utilized to explore drought-resistance related gene resources,then to preliminarily verify the function of target genes in the model legume plant Medicago truncatula. Further the mechanism of drought resistance response network in M. truncatula was clarified,and then the drought-resistance related genes from Medicago falcata was cloned. Finally,cultivating new alfalfa varieties with high drought-resistance was proposed by using molecular breeding approaches. Cultivating new alfalfa varieties with drought-tolerant is an effective way to improve Medicago growth and yield under drought stress. It is of great significance to obtain drought-tolerant alfalfa germplasm through biotechnology approaches for genetic improvement,and to provide new cultivars during the implementation of the Grain to Feed Policy of China,and ultimately to promote the healthy and sustainable development of grass and animal husbandry.

Key words: Medicago, drought resistance, molecular mechanism, molecular breeding

LI Qian, JIANG Wen-bo, WANG Yu-xiang, ZHANG Bo, PANG Yong-zhen. Research Progresses on the Drought Resistance of Medicago at Molecular Level[J]. Biotechnology Bulletin, 2021, 37(8): 243-252.

Figures/Tables 2

References 67

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序号 No.	基因 Genes	基因来源 Gene sources	转基因物种 Genetically modified species	功能 Functions	参考文献 Reference
1	MsNAC2	紫花苜蓿（Medicago sativa）	烟草（Nicotiana tabacum）	丙二醛含量降低,脯氨酸含量、 SOD 和 POD 的活性提高 Reduce MDA content,increase proline content,SOD and POD activity	[26]
2	MfNAC35	黄花苜蓿（Medicago falcata）	烟草（N. tabacum）	抗逆相关基因 CAT、SOD、P5CS表达的上调 Up-regulation of stress-resistance related genes CAT,SOD,and P5CS	[27]
3	MfNACsa	黄花苜蓿（M. falcata）	蒺藜苜蓿（Medicago truncatula）	MfNACsa在脱水胁迫下直接结合Glyl启动子并激活其转录 MfNACsa directly binds to the Glyl promoter and activates its transcription under dehydration stress	[19]
4	MsMYB2L	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	脯氨酸、可溶性糖增加,降低了脂质过氧化 Increase proline and soluble sugars and reduce lipid peroxidation	[28]
5	MfERF049	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF049在根中的表达 Drought stress induces the expression of MfERF049 in roots	[29]
6	MfERF026	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF026在根、茎和叶均表达 Drought stress induces the expression of MfERF026 in roots,stems and leaves	[30]
7	MfERF086	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF086在根、茎和叶均表达 Drought stress induces the expression of MfERF086 in roots,stems and leaves	[30]
8	MsERF11	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	干旱胁迫诱导MsERF11表达 Drought stress induces the expression of MsERF11	[31]
9	MsZIP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量提高 Increase MDA content,relative water content（RWC）,soluble sugar content,soluble protein content and proline content	[32]
10	MtbHLH148	蒺藜苜蓿（M. truncatula ）	拟南芥（Arabidopsis）	根长增加 Increase root length	[33]
11	MsGME	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	更低的MDA含量、较少膜损伤 Lower MDA content,and less membrane damage	[34]
12	MsZEP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	相对含水量和ABA 含量提高,调节气孔导度及气孔开度 Increase RWC and ABA content,and adjust stomata conductance and stomata opening	[35]
13	MsOr	紫花苜蓿（M. sativa）	烟草（N. tabacum）	类胡萝卜素增加,丙二醛含量降低 increase carotenoids,decrease MDA content	[36]
14	MfHyPRP	黄花苜蓿（M. falcata）	烟草（N. tabacum）	转基因株系的鲜重显著高于野生型 The fresh weight of the transgenic line is significantly higher than that of the wild type	[37]
15	MtALDH7A1	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	影响根的生长及侧根数量的增多 Affect the growth of roots and increase the number of lateral roots	[38]
16	MtLEA5B	蒺藜苜蓿（M. truncatula）	大肠杆菌（Escherichia. coli）	胁迫后LEA蛋白仍能保持一定的结构 The LEA protein can still maintain a certain structure after stress	[39]
17	MtCAS31	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	降低气孔密度 Decrease stomatal density	[40]

序号 No.	基因 Genes	基因来源 Gene sources	转基因物种 Genetically modified species	功能 Functions	参考文献 Reference
1	MsNAC2	紫花苜蓿（Medicago sativa）	烟草（Nicotiana tabacum）	丙二醛含量降低,脯氨酸含量、 SOD 和 POD 的活性提高 Reduce MDA content,increase proline content,SOD and POD activity	[26]
2	MfNAC35	黄花苜蓿（Medicago falcata）	烟草（N. tabacum）	抗逆相关基因 CAT、SOD、P5CS表达的上调 Up-regulation of stress-resistance related genes CAT,SOD,and P5CS	[27]
3	MfNACsa	黄花苜蓿（M. falcata）	蒺藜苜蓿（Medicago truncatula）	MfNACsa在脱水胁迫下直接结合Glyl启动子并激活其转录 MfNACsa directly binds to the Glyl promoter and activates its transcription under dehydration stress	[19]
4	MsMYB2L	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	脯氨酸、可溶性糖增加,降低了脂质过氧化 Increase proline and soluble sugars and reduce lipid peroxidation	[28]
5	MfERF049	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF049在根中的表达 Drought stress induces the expression of MfERF049 in roots	[29]
6	MfERF026	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF026在根、茎和叶均表达 Drought stress induces the expression of MfERF026 in roots,stems and leaves	[30]
7	MfERF086	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF086在根、茎和叶均表达 Drought stress induces the expression of MfERF086 in roots,stems and leaves	[30]
8	MsERF11	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	干旱胁迫诱导MsERF11表达 Drought stress induces the expression of MsERF11	[31]
9	MsZIP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量提高 Increase MDA content,relative water content（RWC）,soluble sugar content,soluble protein content and proline content	[32]
10	MtbHLH148	蒺藜苜蓿（M. truncatula ）	拟南芥（Arabidopsis）	根长增加 Increase root length	[33]
11	MsGME	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	更低的MDA含量、较少膜损伤 Lower MDA content,and less membrane damage	[34]
12	MsZEP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	相对含水量和ABA 含量提高,调节气孔导度及气孔开度 Increase RWC and ABA content,and adjust stomata conductance and stomata opening	[35]
13	MsOr	紫花苜蓿（M. sativa）	烟草（N. tabacum）	类胡萝卜素增加,丙二醛含量降低 increase carotenoids,decrease MDA content	[36]
14	MfHyPRP	黄花苜蓿（M. falcata）	烟草（N. tabacum）	转基因株系的鲜重显著高于野生型 The fresh weight of the transgenic line is significantly higher than that of the wild type	[37]
15	MtALDH7A1	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	影响根的生长及侧根数量的增多 Affect the growth of roots and increase the number of lateral roots	[38]
16	MtLEA5B	蒺藜苜蓿（M. truncatula）	大肠杆菌（Escherichia. coli）	胁迫后LEA蛋白仍能保持一定的结构 The LEA protein can still maintain a certain structure after stress	[39]
17	MtCAS31	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	降低气孔密度 Decrease stomatal density	[40]

序号 No.	基因 Genes	功能 Functions	基因来源 Gene sources	转基因苜蓿品种Transgenic alfalfa varieties	抗旱性评价 Drought resistance evaluation	参考文献Reference
1	WXP1	乙烯反应元件结合转录因子 Ethylene response element binding transcription factor	蒺藜苜蓿 Medicago truncatula	Regen SY-4D	叶片表皮蜡质改变;提高净光合速率,蒸腾速率和气孔导度 Change wax in leaf cuticle; improve net photosynthetic rate,transpiration rate and stoma conductance	[46]
2	ZFP1	编码 Cys2/His2 型锌指蛋白 Encoding Cys2/His2 zinc finger protein	野生大豆 Glycine soja	肇东苜蓿 Zhandong	MtCOR47,MtRAB18,MtP5CS 和MtRD2表达量提高 Increased expression level of MtCOR47,MtRAB18,MtP5CS and MtRD2	[47]
3	WRKY20	转录因子 Transcription factor	野生大豆 G. soja	肇东苜蓿 Zhandong	脯氨酸和可溶性糖增加,角质层增厚 Increase content of proline and soluble sugar,and thicken stratum corneum	[48]
4	ABP9	bZIP家族的转录因子 Transcription factors of the bZIP family	玉米 Zea mays	保定苜蓿 Baoding alfalfa	较高相对含水量、叶绿素含量和脯氨酸;较低丙二醛含量 Higher RWC,chlorophyll content and proline; lower malondialdehyde content	[49]
5	ABF3	bZIP转录因子 Transcription factors of the bZIP	拟南芥 Arabidopsis thaliana	新疆大叶Xinjiang Daye	降低蒸腾速率和活性氧含量 Reduce transpiration rate and active oxygen content	[50]
6	Mn-SOD	锰超氧化物歧化酶Mn-SOD	烟草 Nicotiana plumbaginifolia	保定苜蓿 Baoding	MnSOD 活性高,SOD 活性增加 Higher MnSOD activity,increase SOD activity	[51-52]
7	AVP1	液泡H⁺-焦磷酶H⁺-PPase Vacuolar H⁺-PPase gene	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate in the root system more Na⁺、K⁺ and Ca²⁺	[53]
8	NHX 和 VP1-1	Na⁺逆向转运蛋白和植物液泡膜H⁺-PPase基因 Na⁺ antiporter and plant vacuolar membrane H⁺-PPase gene	霸王 Zygophyllum xanthoxylum	新疆大叶Xinjiang Daye	叶片和根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate more Na⁺、K⁺ and Ca²⁺in the leaf and root	[54]
9	codA	胆碱氧化酶Choline oxidase	细菌 Bacteria	新疆大叶Xinjiang Daye	保持较高的相对水分含量,提高甘氨酸甜菜碱和脯氨酸的含量 Maintain a high RWC and increase the content of glycine betaine and proline	[55]
10	Bar和ALDH	醛脱氢酶 Aldehyde dehydrogenase	无芒隐子草 Cleistogenes songorica	金皇后The Golden Queen	Na⁺含量低,K⁺含量高,相对含水量较高 Low Na⁺ content,high K⁺ content,high relative water content	[56]
11	IbOr	橙色基因 Orange gene	甘薯 Ipomoea batatas	新疆大叶Xinjiang Daye	较高含水量,较低MDA含量 high RWC,low MDA content	[57]
12	bar和LEA	干旱胁迫应答因子 Drought stress response factor	无芒隐子草 C. songorica	金皇后 The Golden Quee	较高的相对含水量、较高的地上部生物量、膜损伤和渗透胁迫低 High RWC,high above-ground biomass,low membrane damage and osmotic stress	[58]
13	DHN	脱水素 Dehydrin	沙冬青 Ammopiptanthus mongolicus	中苜2 号 Zhongmu No. 2	Pro 的积累量增加,进一步诱导 Pro 合成途径相关基因的表达 The accumulation of Pro increases,which further induces the expression level of genes related to the pro synthesis pathway	[59]
14	EDT1	编码同域亮氨酸拉链家族的转录因子 Transcription factor of leucine zipper family	拟南芥 A. thaliana	中苜1号 Zhongmu No.1	降低膜透性和丙二醛含量,较高的可溶性糖和脯氨酸含量、超氧化物歧化酶活性和叶绿素含量 Reduce membrane permeability and MDA content,higher soluble sugar and proline content,superoxide dismutase activity and chlorophyll content	[60]
15	ABCG11	角质层脂质转运蛋白编码基因 Encoding lipid transport protein of stratum corneum	霸王 Z. xanthoxylum	新疆大叶 Xinjiang Daye	改变了角质和蜡质的成分、厚度,调节气孔开合程度来减少水分的蒸发量 Change the composition and thickness of the cuticle and wax,and adjust the degree of stomata opening and closing to reduce water evaporation	[61]
16	BADH	甜菜碱醛脱氢酶 Betaine Aldehyde Dehydrogenase	山菠菜 Atriplex hortensis	中苜1号 Zhongmu No.1	可溶性糖含量增加,脯氨酸含量减少 Increase soluble sugar content,and decrease proline content	[62]
17	TPS	6-磷酸海藻糖合酶（TPS） Trehalose synthase gene	酵母 Yeast	Regen SY27x	较高含水量 Higher water content	[63]
18	NDPK	核苷二磷酸激酶 Nucleoside diphosphate kinase	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	较高相对含水量,较高脯氨酸含量和较低MDA含量 Higher RWC,and higher proline content and lower MDA content	[64]
19	Mcsu1	编码钼辅因子硫化酶 Encoding molybdenum cofactor sulfoxygenase	山萮菜 Eutrema salsugineum	新疆大叶Xinjiang Daye	较低的ROS 含量、离子渗透和 MDA 含量 Lower reactive oxygen species（ROS）,ion leakage,and MDA content	[65]

序号 No.	基因 Genes	功能 Functions	基因来源 Gene sources	转基因苜蓿品种Transgenic alfalfa varieties	抗旱性评价 Drought resistance evaluation	参考文献Reference
1	WXP1	乙烯反应元件结合转录因子 Ethylene response element binding transcription factor	蒺藜苜蓿 Medicago truncatula	Regen SY-4D	叶片表皮蜡质改变;提高净光合速率,蒸腾速率和气孔导度 Change wax in leaf cuticle; improve net photosynthetic rate,transpiration rate and stoma conductance	[46]
2	ZFP1	编码 Cys2/His2 型锌指蛋白 Encoding Cys2/His2 zinc finger protein	野生大豆 Glycine soja	肇东苜蓿 Zhandong	MtCOR47,MtRAB18,MtP5CS 和MtRD2表达量提高 Increased expression level of MtCOR47,MtRAB18,MtP5CS and MtRD2	[47]
3	WRKY20	转录因子 Transcription factor	野生大豆 G. soja	肇东苜蓿 Zhandong	脯氨酸和可溶性糖增加,角质层增厚 Increase content of proline and soluble sugar,and thicken stratum corneum	[48]
4	ABP9	bZIP家族的转录因子 Transcription factors of the bZIP family	玉米 Zea mays	保定苜蓿 Baoding alfalfa	较高相对含水量、叶绿素含量和脯氨酸;较低丙二醛含量 Higher RWC,chlorophyll content and proline; lower malondialdehyde content	[49]
5	ABF3	bZIP转录因子 Transcription factors of the bZIP	拟南芥 Arabidopsis thaliana	新疆大叶Xinjiang Daye	降低蒸腾速率和活性氧含量 Reduce transpiration rate and active oxygen content	[50]
6	Mn-SOD	锰超氧化物歧化酶Mn-SOD	烟草 Nicotiana plumbaginifolia	保定苜蓿 Baoding	MnSOD 活性高,SOD 活性增加 Higher MnSOD activity,increase SOD activity	[51-52]
7	AVP1	液泡H⁺-焦磷酶H⁺-PPase Vacuolar H⁺-PPase gene	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate in the root system more Na⁺、K⁺ and Ca²⁺	[53]
8	NHX 和 VP1-1	Na⁺逆向转运蛋白和植物液泡膜H⁺-PPase基因 Na⁺ antiporter and plant vacuolar membrane H⁺-PPase gene	霸王 Zygophyllum xanthoxylum	新疆大叶Xinjiang Daye	叶片和根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate more Na⁺、K⁺ and Ca²⁺in the leaf and root	[54]
9	codA	胆碱氧化酶Choline oxidase	细菌 Bacteria	新疆大叶Xinjiang Daye	保持较高的相对水分含量,提高甘氨酸甜菜碱和脯氨酸的含量 Maintain a high RWC and increase the content of glycine betaine and proline	[55]
10	Bar和ALDH	醛脱氢酶 Aldehyde dehydrogenase	无芒隐子草 Cleistogenes songorica	金皇后The Golden Queen	Na⁺含量低,K⁺含量高,相对含水量较高 Low Na⁺ content,high K⁺ content,high relative water content	[56]
11	IbOr	橙色基因 Orange gene	甘薯 Ipomoea batatas	新疆大叶Xinjiang Daye	较高含水量,较低MDA含量 high RWC,low MDA content	[57]
12	bar和LEA	干旱胁迫应答因子 Drought stress response factor	无芒隐子草 C. songorica	金皇后 The Golden Quee	较高的相对含水量、较高的地上部生物量、膜损伤和渗透胁迫低 High RWC,high above-ground biomass,low membrane damage and osmotic stress	[58]
13	DHN	脱水素 Dehydrin	沙冬青 Ammopiptanthus mongolicus	中苜2 号 Zhongmu No. 2	Pro 的积累量增加,进一步诱导 Pro 合成途径相关基因的表达 The accumulation of Pro increases,which further induces the expression level of genes related to the pro synthesis pathway	[59]
14	EDT1	编码同域亮氨酸拉链家族的转录因子 Transcription factor of leucine zipper family	拟南芥 A. thaliana	中苜1号 Zhongmu No.1	降低膜透性和丙二醛含量,较高的可溶性糖和脯氨酸含量、超氧化物歧化酶活性和叶绿素含量 Reduce membrane permeability and MDA content,higher soluble sugar and proline content,superoxide dismutase activity and chlorophyll content	[60]
15	ABCG11	角质层脂质转运蛋白编码基因 Encoding lipid transport protein of stratum corneum	霸王 Z. xanthoxylum	新疆大叶 Xinjiang Daye	改变了角质和蜡质的成分、厚度,调节气孔开合程度来减少水分的蒸发量 Change the composition and thickness of the cuticle and wax,and adjust the degree of stomata opening and closing to reduce water evaporation	[61]
16	BADH	甜菜碱醛脱氢酶 Betaine Aldehyde Dehydrogenase	山菠菜 Atriplex hortensis	中苜1号 Zhongmu No.1	可溶性糖含量增加,脯氨酸含量减少 Increase soluble sugar content,and decrease proline content	[62]
17	TPS	6-磷酸海藻糖合酶（TPS） Trehalose synthase gene	酵母 Yeast	Regen SY27x	较高含水量 Higher water content	[63]
18	NDPK	核苷二磷酸激酶 Nucleoside diphosphate kinase	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	较高相对含水量,较高脯氨酸含量和较低MDA含量 Higher RWC,and higher proline content and lower MDA content	[64]
19	Mcsu1	编码钼辅因子硫化酶 Encoding molybdenum cofactor sulfoxygenase	山萮菜 Eutrema salsugineum	新疆大叶Xinjiang Daye	较低的ROS 含量、离子渗透和 MDA 含量 Lower reactive oxygen species（ROS）,ion leakage,and MDA content	[65]