Research progress on yield loss under drought stress and drought resistance genetics of alfalfa （Medicago sativa）

doi:10.11686/cyxb2024344

Abstract

Abstract:

Alfalfa （Medicago sativa） is the most widely cultivated perennial leguminous forage crop， acclaimed as the “king of forages” because of its high yield and superior quality. Drought stress has a significant impact on the growth and development of alfalfa， resulting in substantial yield reductions. It influences the germination rate， branch formation， stem elongation， leaf growth， and root development， potentially causing large decreases （>70%） in forage yield. Accelerating the breeding of drought-tolerant alfalfa varieties through molecular breeding is an effective strategy to mitigate the effects of drought stress on this forage crop. However， the genetic foundation of drought resistance in alfalfa remains largely unexplored. Previous research on alfalfa has mainly concentrated on transgenic methods and homologous cloning techniques. With the release of the alfalfa genome and advances in sequencing technology， genome-wide association studies and omics technologies based on transcriptome sequencing have played an increasingly important role in identifying drought-related genes and elucidating drought resistance mechanisms in alfalfa. This paper comprehensively summarizes the effects of drought stress on alfalfa yield， outlines recent advances in research on the genetic basis of drought resistance in alfalfa， and provides a reference for the breeding of drought-resistant alfalfa varieties.

Key words: alfalfa, drought resistance, genome-wide association studies, yield

Xue-qian JIANG, Qing-chuan YANG, Jun-mei KANG. Research progress on yield loss under drought stress and drought resistance genetics of alfalfa （Medicago sativa）[J]. Acta Prataculturae Sinica, 2025, 34(7): 219-234.

Figures/Tables 4

References 107

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群体大小 Population size （No.）	基因型测序 Genotyping	环境 Environment	表型 Phenotype	主要结果 Main results	参考文献 Reference
200	GBS	田间 Field	一年3次刈割的生物量Biomass of three harvests in one year	干旱条件下定位到28个与生物量相关的单核苷酸多态性（single nucleotide polymorphism， SNP）标记。28 SNP markers were associated with biomass under drought stress.	［41］
198	GBS	温室 Greenhouse	耐旱指数和叶片相对含水量Drought resistance index （DRI） and leaf relative water content （RWC）	在DRI和RWC中分别鉴定了19和15个SNP。Nineteen and fifteen SNP associated with DRI and RWC， respectively.	［55］
198	GBS	田间 Field	26个品质相关性状26 forage quality traits	131个SNP与所有水分亏缺处理中的多个性状相关。131 SNP associated with multiple traits in all the water deficit treatments.	［56］
109	RNA-seq	温室 Greenhouse	株高、地上生物量和7个叶绿素荧光参数的抗旱系数Drought-resistance coefficients for plant height， above-ground biomass， and seven chlorophyll fluorescence parameters	在9个性状中共定位到21个显著SNP；验证了候选基因MsMYBH可增强苜蓿的抗旱性。A total of 21 significant SNPs were identified in nine traits； MsMYBH was confirmed to enhance alfalfa drought resistance.	［57］

群体大小 Population size （No.）	基因型测序 Genotyping	环境 Environment	表型 Phenotype	主要结果 Main results	参考文献 Reference
200	GBS	田间 Field	一年3次刈割的生物量Biomass of three harvests in one year	干旱条件下定位到28个与生物量相关的单核苷酸多态性（single nucleotide polymorphism， SNP）标记。28 SNP markers were associated with biomass under drought stress.	［41］
198	GBS	温室 Greenhouse	耐旱指数和叶片相对含水量Drought resistance index （DRI） and leaf relative water content （RWC）	在DRI和RWC中分别鉴定了19和15个SNP。Nineteen and fifteen SNP associated with DRI and RWC， respectively.	［55］
198	GBS	田间 Field	26个品质相关性状26 forage quality traits	131个SNP与所有水分亏缺处理中的多个性状相关。131 SNP associated with multiple traits in all the water deficit treatments.	［56］
109	RNA-seq	温室 Greenhouse	株高、地上生物量和7个叶绿素荧光参数的抗旱系数Drought-resistance coefficients for plant height， above-ground biomass， and seven chlorophyll fluorescence parameters	在9个性状中共定位到21个显著SNP；验证了候选基因MsMYBH可增强苜蓿的抗旱性。A total of 21 significant SNPs were identified in nine traits； MsMYBH was confirmed to enhance alfalfa drought resistance.	［57］

基因 Genes	描述 Description	基因功能 Gene function	参考文献Reference
GsZFP1	编码Cys2/His2型锌指蛋白Encodes a Cys2/His2-type zinc-finger protein	过表达增强了紫花苜蓿的耐旱性。Overexpression enhanced drought resistance of alfalfa.	［33］
GsWRKY20	WRAKY转录因子WRKY-type transcription factor	更厚的角质层Thicker cuticular layer	［34］
miR156/SPL13+DFR/WD40-1	miRNA，SPL转录因子miRNA， squamosa promoter binding protein-like （SPL） transcription factors	中等水平的miR156表达可抑制SPL13并增加WD40-1表达，以微调花青素生物合成的DFR表达，并调节紫花苜蓿的各种发育、生理和生化过程，从而提高抗旱能力。Moderate levels of miR156 expression can inhibit SPL13 and increase WD40-1 expression， to fine-tune the expression of DFR involved in anthocyanin biosynthesis， and regulate various developmental， physiological， and biochemical processes in alfalfa， thereby enhancing drought resistance.	［36-37］
MsMYBH	MYB-like转录因子MYB-like transcription factor	保持水平衡、高光合作用效率以及清除过量的H₂O₂。Maintaining water balance， high photosynthetic efficiency， and scavenging excess H₂O₂.	［57］
AtAVP1	液泡 H⁺-焦磷酸酶（H⁺-PPase） Vacuolar H⁺-pyrophosphatase （H⁺-PPase）	叶片和根系中积累更多的Na⁺、K⁺和Ca²⁺。Accumulation of more Na⁺， K⁺ and Ca²⁺ in leaves and roots.	［62］
ZxABCG11	编码ABC转运蛋白Encodes an ATP binding cassette （ABC） transporter	更高的蜡晶体密度和更厚的叶片角质层，从而提高转基因苜蓿的保水能力和光合作用能力。Higher wax crystal density and thicker leaf cuticular layer， thereby enhancing water retention and photosynthetic capacity of transgenic alfalfa.	［63］
AtEDT1	同源域亮氨酸拉链转录因子Homodomain-leucine zipper transcription factor	降低气孔密度，增加根系发育，同时膜透性和丙二醛含量降低，但可溶性糖和脯氨酸含量较高。Reduced stomatal density， increased root development， while membrane permeability and malondialdehyde content decreased， but soluble sugar and proline content were higher.	［64］
MtWXP1	AP2结构域的转录因子基因AP2 domain-containing transcription factor gene	增加角质层蜡堆积，增强耐旱性。Increased accumulation of cuticular wax， enhancing drought resistance.	［65-66］
EsMcsu1	编码一种钼辅因子硫化酶Encoding a molybdenum cofactor sulfurase	促进脱落酸生物合成，提高抗旱性。Promotes abscisic acid biosynthesis， enhancing drought resistance.	［67］
CsLEA	晚期胚胎发生丰富蛋白Late embryogenesis abundant （LEA） proteins	较高的相对含水量和减少的膜损伤。Higher relative water content and reduced membrane damage.	［68］
AgcodA	来自土壤细菌 Arthrobacter globiformis 的 codA 基因，编码胆碱氧化酶A codA gene from the soil bacterium （Arthrobacter globiformis）， encoding choline oxidase	保持较高的相对含水量和增加甘氨酸甜菜碱和脯氨酸含量。Maintaining high relative water contents and increased levels of glycinebetaine and proline.	［69］
MsZIP	bZIP转录因子bZIP transcription factor	增加丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量。Increased malondialdehyde content， relative water content， soluble sugar content， soluble protein content， and proline content.	［70］
MsNAC	NAC转录因子NAC transcription factor	-	［71］
MsHSP17.7	编码热休克蛋白Encodes a small heat shock protein	增加根长。Increased root length.	［72］
MsZEP	编码玉米黄质环氧化酶Encodes zeaxanthin epoxidase	影响各种生理途径、ABA水平和胁迫响应基因表达。Affects various physiological pathways， ABA levels， and stress-responsive gene expression.	［73］
MsHSP70	编码热休克蛋白Encodes heat shock proteins	相对含水量、脯氨酸含量、超氧化物歧化酶活性升高，丙二醛含量降低。Increased relative water content， proline content， and superoxide dismutase activity， while malondialdehyde content decreased.	［74］
MsLEA4-4	晚期胚胎发生丰富蛋白Late embryogenesis abundant （LEA） proteins	更多的侧根和更高的叶绿素含量，可溶性糖水平和多种抗氧化酶活性升高，而脯氨酸和丙二醛水平显著降低。More lateral roots and higher chlorophyll content， increased soluble sugar levels and various antioxidant enzyme activities， while proline and malondialdehyde levels significantly decreased.	［75］
MsCML46	编码钙调蛋白样蛋白Encodes calmodulin-like protein	MsCML46 结合游离Ca²⁺以促进信号转导并维持较高的 K⁺/Na⁺，保护细胞内稳态。MsCML46 binds free Ca²⁺ to promote signal transduction and maintain a higher K⁺/Na⁺， protecting cellular homeostasis.	［76］
MsVDAC	编码电压依赖性阴离子选择性通道（VDAC）蛋白 Encodes voltage-dependent anion-selective channel （VDAC） protein	渗透稳态和胁迫响应基因表达。 Osmotic homeostasis and stress-responsive gene expression.	［77］
MsWRKY11	WRKY转录因子WRKY transcription factor	通过调控木质素生物合成和紫花苜蓿气孔开闭调控耐旱性。Enhances drought resistance by regulating lignin biosynthesis and alfalfa stomatal opening and closing.	［78］
MsDHN1-MsPIP2；1-MsmMYB	MsDHN1（脱水蛋白）、 MsPIP2；1（水通道蛋白），MsmMYB（一种膜锚定的 MYB 转录因子） MsDHN1 （dehydrin） and MsPIP2；1 （aquaporin）， MsmMYB （a membrane-anchored MYB transcriptional factor MsmMYB）	缺水导致MsPIP2；1磷酸化，mMYB （mMYB△83）的C端易位并与MsDHN1相互作用，并促进mMYB△83响应缺水的转录活性。mMYB和mMYBD83的过表达下调MsCESA3的表达，但通过直接结合其启动子上调MsCESA7的表达。Dehydration leads to phosphorylation of MsPIP2；1， C-terminal translocation of mMYB （mMYB△83） and interaction with MsDHN1， promoting the transcriptional activity of mMYB△83 in response to dehydration. Overexpression of mMYB and mMYBD83 downregulates the expression of MsCESA3， but upregulates the expression of MsCESA7 by directly binding to its promoter.	［79］
MsSPL8	SPL转录因子Squamosa promoter binding protein-like （SPL） transcription factors	表达下调的植株延缓枯萎并快速恢复； MsSPL8突变体在耐缺水能力方面有所提高。Down-regulation delayed wilting and recovered quickly； MsSPL8 mutants displayed improvements in their ability to withstand water-deficit.	［80-81］
MsDIUP1	干旱诱导的未知蛋白 1（它缺乏任何可靠的保守结构域）Drought-induced unknown protein 1 （it lacked any confidently conserved domains）	参与胁迫信号传导、抗氧化防御和渗透调节的基因存在差异反应。Differential responses of genes involved in stress signal transduction， antioxidant defense， and osmotic regulation.	［82］
MsNTF2L	核转运因子2-like Nuclear transport factor 2-like	通过调节叶片失水（通过调节气孔和蜡沉积）、抗氧化防御和光合作用调控苜蓿耐旱性。Regulates drought resistance by modulating leaf dehydration （through stomatal and wax deposition）， antioxidant defense， and photosynthesis in alfalfa.	［83］
miR156/SPL9	miRNA	调节花青素的生物合成。Regulating anthocyanin biosynthesis.	［89］
TPS1-TPS2	酵母海藻糖-6-磷酸合酶（TPS1）和海藻糖-6-磷酸磷酸酶（TPS2）基因Yeast trehalose-6-phosphate synthase （TPS1） and trehalose-6-phosphate phosphatase （TPS2） genes	海藻糖积累。Accumulation of trehalose.	［90］
MfLEA3	晚期胚胎发生丰富蛋白 Late embryogenesis abundant （LEA） proteins	转基因植物中积累ROS减少。 Reduced accumulation of ROS in transgenic plants.	［91］
AtNDPK2	拟南芥核苷二磷酸激酶 2 Arabidopsis nucleoside diphosphate kinase 2	水分流失率降低，细胞膜损伤降低。 Water loss rate and cell membrane damage were decreased.	［92］
AtABF3	脱落酸（ABA）响应元件结合因子3 ABA-responsive element-binding factor 3	蒸腾速率降低，活性氧含量降低。A reduced transpiration rate and lower reactive oxygen species contents.	［93］
IbOr	甘薯橙基因The sweetpotato orange gene	总类胡萝卜素水平更高，细胞膜损伤更少。Higher total carotenoid levels and lower cell membrane damage.	［94］
co-expression of bar+CsALDH genes	氧化应答和抗除草剂基因An oxidative responsive gene （CsALDH） and herbicide resistance gene （bar）	Na⁺含量降低，K⁺含量升高；离子毒性降低，渗透调节维持；相对含水量升高，光系统变化减少，膜损伤减少。Lower Na⁺ and higher K⁺ content； Reduction of ion toxicity and maintenance of osmotic adjustment； Higher relative water content level， fewer changes in the photosystem， decreased membrane injury.	［95］
MsTMT	编码γ-生育酚甲基转移酶Encodes γ-tocopherol methyltransferase	减轻氧化损伤，积累更多渗透解离物质，提高水分利用效率。Alleviated oxidative damage， accumulation of more osmolytic substances and improved water use efficiency.	［96］

基因 Genes	描述 Description	基因功能 Gene function	参考文献Reference
GsZFP1	编码Cys2/His2型锌指蛋白Encodes a Cys2/His2-type zinc-finger protein	过表达增强了紫花苜蓿的耐旱性。Overexpression enhanced drought resistance of alfalfa.	［33］
GsWRKY20	WRAKY转录因子WRKY-type transcription factor	更厚的角质层Thicker cuticular layer	［34］
miR156/SPL13+DFR/WD40-1	miRNA，SPL转录因子miRNA， squamosa promoter binding protein-like （SPL） transcription factors	中等水平的miR156表达可抑制SPL13并增加WD40-1表达，以微调花青素生物合成的DFR表达，并调节紫花苜蓿的各种发育、生理和生化过程，从而提高抗旱能力。Moderate levels of miR156 expression can inhibit SPL13 and increase WD40-1 expression， to fine-tune the expression of DFR involved in anthocyanin biosynthesis， and regulate various developmental， physiological， and biochemical processes in alfalfa， thereby enhancing drought resistance.	［36-37］
MsMYBH	MYB-like转录因子MYB-like transcription factor	保持水平衡、高光合作用效率以及清除过量的H₂O₂。Maintaining water balance， high photosynthetic efficiency， and scavenging excess H₂O₂.	［57］
AtAVP1	液泡 H⁺-焦磷酸酶（H⁺-PPase） Vacuolar H⁺-pyrophosphatase （H⁺-PPase）	叶片和根系中积累更多的Na⁺、K⁺和Ca²⁺。Accumulation of more Na⁺， K⁺ and Ca²⁺ in leaves and roots.	［62］
ZxABCG11	编码ABC转运蛋白Encodes an ATP binding cassette （ABC） transporter	更高的蜡晶体密度和更厚的叶片角质层，从而提高转基因苜蓿的保水能力和光合作用能力。Higher wax crystal density and thicker leaf cuticular layer， thereby enhancing water retention and photosynthetic capacity of transgenic alfalfa.	［63］
AtEDT1	同源域亮氨酸拉链转录因子Homodomain-leucine zipper transcription factor	降低气孔密度，增加根系发育，同时膜透性和丙二醛含量降低，但可溶性糖和脯氨酸含量较高。Reduced stomatal density， increased root development， while membrane permeability and malondialdehyde content decreased， but soluble sugar and proline content were higher.	［64］
MtWXP1	AP2结构域的转录因子基因AP2 domain-containing transcription factor gene	增加角质层蜡堆积，增强耐旱性。Increased accumulation of cuticular wax， enhancing drought resistance.	［65-66］
EsMcsu1	编码一种钼辅因子硫化酶Encoding a molybdenum cofactor sulfurase	促进脱落酸生物合成，提高抗旱性。Promotes abscisic acid biosynthesis， enhancing drought resistance.	［67］
CsLEA	晚期胚胎发生丰富蛋白Late embryogenesis abundant （LEA） proteins	较高的相对含水量和减少的膜损伤。Higher relative water content and reduced membrane damage.	［68］
AgcodA	来自土壤细菌 Arthrobacter globiformis 的 codA 基因，编码胆碱氧化酶A codA gene from the soil bacterium （Arthrobacter globiformis）， encoding choline oxidase	保持较高的相对含水量和增加甘氨酸甜菜碱和脯氨酸含量。Maintaining high relative water contents and increased levels of glycinebetaine and proline.	［69］
MsZIP	bZIP转录因子bZIP transcription factor	增加丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量。Increased malondialdehyde content， relative water content， soluble sugar content， soluble protein content， and proline content.	［70］
MsNAC	NAC转录因子NAC transcription factor	-	［71］
MsHSP17.7	编码热休克蛋白Encodes a small heat shock protein	增加根长。Increased root length.	［72］
MsZEP	编码玉米黄质环氧化酶Encodes zeaxanthin epoxidase	影响各种生理途径、ABA水平和胁迫响应基因表达。Affects various physiological pathways， ABA levels， and stress-responsive gene expression.	［73］
MsHSP70	编码热休克蛋白Encodes heat shock proteins	相对含水量、脯氨酸含量、超氧化物歧化酶活性升高，丙二醛含量降低。Increased relative water content， proline content， and superoxide dismutase activity， while malondialdehyde content decreased.	［74］
MsLEA4-4	晚期胚胎发生丰富蛋白Late embryogenesis abundant （LEA） proteins	更多的侧根和更高的叶绿素含量，可溶性糖水平和多种抗氧化酶活性升高，而脯氨酸和丙二醛水平显著降低。More lateral roots and higher chlorophyll content， increased soluble sugar levels and various antioxidant enzyme activities， while proline and malondialdehyde levels significantly decreased.	［75］
MsCML46	编码钙调蛋白样蛋白Encodes calmodulin-like protein	MsCML46 结合游离Ca²⁺以促进信号转导并维持较高的 K⁺/Na⁺，保护细胞内稳态。MsCML46 binds free Ca²⁺ to promote signal transduction and maintain a higher K⁺/Na⁺， protecting cellular homeostasis.	［76］
MsVDAC	编码电压依赖性阴离子选择性通道（VDAC）蛋白 Encodes voltage-dependent anion-selective channel （VDAC） protein	渗透稳态和胁迫响应基因表达。 Osmotic homeostasis and stress-responsive gene expression.	［77］
MsWRKY11	WRKY转录因子WRKY transcription factor	通过调控木质素生物合成和紫花苜蓿气孔开闭调控耐旱性。Enhances drought resistance by regulating lignin biosynthesis and alfalfa stomatal opening and closing.	［78］
MsDHN1-MsPIP2；1-MsmMYB	MsDHN1（脱水蛋白）、 MsPIP2；1（水通道蛋白），MsmMYB（一种膜锚定的 MYB 转录因子） MsDHN1 （dehydrin） and MsPIP2；1 （aquaporin）， MsmMYB （a membrane-anchored MYB transcriptional factor MsmMYB）	缺水导致MsPIP2；1磷酸化，mMYB （mMYB△83）的C端易位并与MsDHN1相互作用，并促进mMYB△83响应缺水的转录活性。mMYB和mMYBD83的过表达下调MsCESA3的表达，但通过直接结合其启动子上调MsCESA7的表达。Dehydration leads to phosphorylation of MsPIP2；1， C-terminal translocation of mMYB （mMYB△83） and interaction with MsDHN1， promoting the transcriptional activity of mMYB△83 in response to dehydration. Overexpression of mMYB and mMYBD83 downregulates the expression of MsCESA3， but upregulates the expression of MsCESA7 by directly binding to its promoter.	［79］
MsSPL8	SPL转录因子Squamosa promoter binding protein-like （SPL） transcription factors	表达下调的植株延缓枯萎并快速恢复； MsSPL8突变体在耐缺水能力方面有所提高。Down-regulation delayed wilting and recovered quickly； MsSPL8 mutants displayed improvements in their ability to withstand water-deficit.	［80-81］
MsDIUP1	干旱诱导的未知蛋白 1（它缺乏任何可靠的保守结构域）Drought-induced unknown protein 1 （it lacked any confidently conserved domains）	参与胁迫信号传导、抗氧化防御和渗透调节的基因存在差异反应。Differential responses of genes involved in stress signal transduction， antioxidant defense， and osmotic regulation.	［82］
MsNTF2L	核转运因子2-like Nuclear transport factor 2-like	通过调节叶片失水（通过调节气孔和蜡沉积）、抗氧化防御和光合作用调控苜蓿耐旱性。Regulates drought resistance by modulating leaf dehydration （through stomatal and wax deposition）， antioxidant defense， and photosynthesis in alfalfa.	［83］
miR156/SPL9	miRNA	调节花青素的生物合成。Regulating anthocyanin biosynthesis.	［89］
TPS1-TPS2	酵母海藻糖-6-磷酸合酶（TPS1）和海藻糖-6-磷酸磷酸酶（TPS2）基因Yeast trehalose-6-phosphate synthase （TPS1） and trehalose-6-phosphate phosphatase （TPS2） genes	海藻糖积累。Accumulation of trehalose.	［90］
MfLEA3	晚期胚胎发生丰富蛋白 Late embryogenesis abundant （LEA） proteins	转基因植物中积累ROS减少。 Reduced accumulation of ROS in transgenic plants.	［91］
AtNDPK2	拟南芥核苷二磷酸激酶 2 Arabidopsis nucleoside diphosphate kinase 2	水分流失率降低，细胞膜损伤降低。 Water loss rate and cell membrane damage were decreased.	［92］
AtABF3	脱落酸（ABA）响应元件结合因子3 ABA-responsive element-binding factor 3	蒸腾速率降低，活性氧含量降低。A reduced transpiration rate and lower reactive oxygen species contents.	［93］
IbOr	甘薯橙基因The sweetpotato orange gene	总类胡萝卜素水平更高，细胞膜损伤更少。Higher total carotenoid levels and lower cell membrane damage.	［94］
co-expression of bar+CsALDH genes	氧化应答和抗除草剂基因An oxidative responsive gene （CsALDH） and herbicide resistance gene （bar）	Na⁺含量降低，K⁺含量升高；离子毒性降低，渗透调节维持；相对含水量升高，光系统变化减少，膜损伤减少。Lower Na⁺ and higher K⁺ content； Reduction of ion toxicity and maintenance of osmotic adjustment； Higher relative water content level， fewer changes in the photosystem， decreased membrane injury.	［95］
MsTMT	编码γ-生育酚甲基转移酶Encodes γ-tocopherol methyltransferase	减轻氧化损伤，积累更多渗透解离物质，提高水分利用效率。Alleviated oxidative damage， accumulation of more osmolytic substances and improved water use efficiency.	［96］

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