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摘要: 随着农业种植业结构调整, 谷子作为一种抗旱耐瘠薄作物在干旱盐碱地区的种植越来越受到重视。系统深入梳理谷子的耐盐性及盐胁迫下的生理生态响应特性, 对增加盐碱地区谷子产量、提高农民收入具有重要指导意义。本文从谷子的耐盐性筛选指标及评价、盐胁迫下植株生长发育变化规律和生理生态响应以及谷子耐盐基因发掘等3个方面, 综述了国内外研究进展。目前谷子耐盐鉴选指标单一, 主要依赖于谷子萌发期的发芽率, 而其他生理生态指标未被充分考虑; 因谷子品种和土壤盐分的不同, 谷子植株地上地下农艺性状、光合特征、清除活性氧相关酶类以及激素响应特征存在差异, 建立综合鉴选指标存在困难; 谷子耐盐基因的表达与作用发挥与环境条件, 如高盐、干旱(PEG)和脱落酸(ABA)等相关, 通过特定蛋白来增强抗氧化系统、保护细胞不受损伤以及提高抗渗透胁迫能力等提高谷子的耐盐性。在此基础上, 本文提出了建立谷子耐盐综合鉴选量化标准和平台、深入开展谷子耐盐调控机理研究、进一步研发谷子耐盐栽培技术体系是未来重要的研究方向。Abstract: With the adjustment of the agricultural planting structure, foxtail millet — a drought-resistant and barren-tolerant crop, has attracted increasing attention in arid and saline-alkali areas. Systematic and deep understanding of the salt tolerance as well as physiological and ecological response characteristics of foxtail millet under salt stress has important guiding significance for maximizing the excellent properties of foxtail millet, increasing its yield in saline-alkali areas, and improving farmers’ income. This paper reviewed the research progress worldwide in detail from three aspects, namely, the screening index and evaluation of foxtail millet salt tolerance, the change law of plant growth and development and physiological-ecological response under salt stress, and the discovery of foxtail millet salt tolerance genes. At present, a single indicator of salt tolerance in foxtail millet is used, which mainly depends on the germination rate during germination. Other physiological and ecological indicators have not been fully considered. Because of the differences in foxtail millet varieties and soil salinity, there are differences in aboveground and underground agronomic characteristics, photosynthetic characteristics, enzymes related to scavenging reactive oxygen species, and the hormone response law of foxtail millet plants. Therefore, it is difficult to establish comprehensive evaluation indexes. The expression and function of foxtail millet salt tolerance genes are related to environmental conditions such as high salt, drought, and abscisic acid. The salt tolerance of foxtail millet was improved by synthesizing specific proteins to enhance the antioxidant system, protect cells from damage, and improve the ability to resist osmotic stress. Based on the results, we recommend that the establishment of quantitative standards and platforms for the comprehensive evaluation and selection of foxtail millet salt tolerance, the in-depth research on the regulation mechanism of foxtail millet salt tolerance, and further research and development of foxtail millet salt tolerance cultivation technology systems are important future research directions.
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表 1 谷子耐盐性鉴选内容
Table 1. Contents of salt tolerance evaluation of foxtail millet
处理时期
Treatment period栽培条件
Cultivation condition品种数量
Varieties number盐类型
Salt type鉴选适宜盐浓度
Suitable salt concentration鉴定指标
Appraisal index耐盐性分级
Salt tolerance classification参考文献
Reference萌发期
Germination period培养皿
Petri dish27 NaCl 3 g·kg−1 发芽率及其相对盐害率
Germination rate, relative salt damage rate of germination‘张杂3号’为耐盐型
‘Zhangza 3’ is salt tolerant type.[21] 培养皿
Petri dish63[24], 54[19], 100[28], 260[30] NaCl 1.0%~1.5% 发芽率、发芽势、根长和芽长的相对值及盐害率
Relative values and salt damage rates of germination rate, germination energy, root length, bud length‘济谷16’和‘汾特11’等20份为高耐盐型, ‘鲁谷1号’和‘豫谷15’等23份为盐敏感型
20 varieties such as ‘Jigu 16’ and ‘Fente 11’ are high salt tolerant types; 23 varieties such as ‘Lugu 1’ and ‘Yugu 15’ are salt sensitive types.[20,24,28,30] 培养皿
Petri dish100[22], 11[23], 14[26], 25[31], 10[32], 14[33] NaCl 100~250 mmol·L−1 发芽率、发芽势、根长、芽长及其相对值和盐害率、相对活力指数、Na+含量和K+含量
Germination rate, germination energy, root length, bud length and their relative values and salt damage rates, relative vigor index, Na+ and K+ contents‘晋育红谷’和‘公矮6号’等16份为高耐盐型, ‘龙谷27’和‘龙谷25’等11份为盐高度敏感型
16 varieties such as ‘Jinyuhonggu’ and ‘Gong’ai 6’ are high salt tolerant types, and 11 varieties such as ‘Longgu 27’ and ‘Longgu 25’ are highly salt sensitive types.[22-23,26,31-33] 培养皿
Petri dish4 Na2CO3 0, 25, 50, 75, 100 mmol·L−1 相对发芽率、相对盐害率
Relative germination rate, relative salt damage rate— [34] 培养皿
Petri dish20 Na2SO4∶ NaCl∶ NaHCO3∶ Na2CO3=
4∶1∶4∶160~100 mmol·L−1 发芽指数、活力指数、相对根
长、相对芽长、盐碱害指数
Germination index, vigor index, relative root length, relative bud length, saline-alkali damage index‘9324-8-3’ ‘坝谷214’和‘晋谷23’为耐盐碱型, ‘大同30’ ‘大白谷’和‘九根齐’为盐碱高敏感型
‘9324-8-3’ ‘Bagu 214’ and ‘Jingu 23’ are saline alkali resistant types, and ‘Datong 30’ ‘Dabaigu’ and ‘Jiugenqi’ are highly saline-alkali sensitive types.[27] 培养皿
Petri dish53 NaCl∶
NaHCO3=
4∶1100 mmol·L−1 相对发芽率、根长盐害率、芽长盐害率、根冠比盐害率
Relative germination rate, salt damage rate of root length, salt damage rate of bud length, specific salt damage rate of root-shoot ratio‘鲁谷10’和‘济谷22’为强耐盐碱型, ‘豫谷31’ ‘豫谷32’和‘保谷23’为极不耐盐碱型
‘Lugu 10’ and ‘Jigu 22’ are strong salt-alkali resistant types, while ‘Yugu 31’ ‘Yugu 32’ and ‘Baogu 23’ are extremely salt-alkali resistant types.[25] 苗期
Seedling stage培养皿
Petri dish14[26], 25[31] NaCl 50, 150 mmol·L−1 根长、芽长及其相对值和盐害率
Root length, shoot length and their relative values and salt damage rates— [26,31] 盆栽
Pot culture3 NaCl 0.5%~1.0% 气孔状态、叶表面变化、相对电导率和叶绿素荧光参数、生物量、超氧化物歧化酶
Stomatal state, leaf surface changes, relative conductivity and chlorophyll fluorescence parameters, biomass, superoxide dismutase— [28] 全生育期
Whole growth period盆栽
Pot culture155 NaCl 100 mmol·L−1 籽粒产量、地上部生物量、收获指数、穗部收获指数
Grain yield, aboveground biomass, harvest index, spike harvest index— [35] 大田
Field8 盐碱地
Saline soil3.20 g·kg−1 穗重、单穗粒重、干物质重、地上部含水量
Panicle weight, grain weight per panicle, dry matter weight, water content of shoot‘济谷22’和‘济谷21’为耐盐碱型
‘Jigu 22’ and ‘Jigu 21’ are salt and alkali resistant types.[19] 
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表 2 盐胁迫影响谷子生理生化指标
Table 2. Physiological and biochemical indexes of foxtail millet affected by salt stress
指标类型
Index system指标
Index表现
Performance参考文献
Reference农艺性状
Agronomic characters根系
Root system混合盐碱胁迫下, 不同品种的根长盐害率均值达77.6%; NaCl胁迫下盐敏感品种根部细胞内pH提高7.28%, 比抗盐品种增加6.15%
Under the mixed saline-alkali stress, the average root length salt damage rate of different varieties was 77.6%; under NaCl stress, the pH in the root cells of salt sensitive varieties increased by 7.28%, which was 6.15% higher than that of salt resistant varieties.[25,40] 株高
Plant height0.1%~0.3% NaCl胁迫下‘济谷16’较敏感, 株高降低11.69%~16.91%, 在0.5%~0.7% NaCl胁迫下, ‘鲁谷1号’株高降低20.01%~26.17%
Under 0.1%−0.3% NaCl stress, ‘Jigu 16’ was sensitive, and the plant height decreased by 11.69%−16.91%. Under 0.5%−0.7% NaCl stress, ‘Lugu 1’ was the more sensitive, and the plant height decreased by 20.01%−26.17%.[42] 地上部含水量
Aboveground water content含盐3.2 g·kg−1的大田条件下谷子成熟期的地上部含水量变幅为 35.4%~49.2%
Under the field condition of salt content of 3.2 g∙kg−1, the aboveground water content of millet at maturity varies from 35.4% to 49.2%.[19] 产量要素
Yield factors穗重、单穗粒重、千粒重
Ear weight, grain weight of per panicle, 1000-seed weight含盐3.2 g·kg−1大田条件下谷子各品种的穗重、单穗粒重、千粒重变异系数分别为13.5%、13.5%和3.1%, 产量盐害率为20.7%~63.4%
Under the field condition of salt content of 3.2 g∙kg−1, the variation coefficients of panicle weight, grain weight per panicle and 1000-grain weight of millet varieties were 13.5%, 13.5% and 3.1% respectively, and the yield salt damage rate was 20.7%−63.4%.[19] 光合特性
Photosynthetic characteristics叶绿素、气孔导度、蒸腾速率、净光合速率
Chlorophyll, stomatal conductance, transpiration rate, net photosynthetic rate复合盐胁迫(0~0.9%)下净光合速率由于气孔导度和蒸腾速率下降而降低, 0~0.3%时蒸腾速率、气孔导度下降较快, 0.6%~0.9%时下降趋势减缓; 在中性盐NaCl、Na2SO4及碱性盐NaHCO3胁迫下, 叶绿素受NaCl影响最大, 0.2%和0.4%的处理分别比对照下降24.1%和34.0%
Under compound salt stress (0−0.9%), the net photosynthetic rate decreased due to the decrease of stomatal conductance and transpiration rate. The transpiration rate and stomatal conductance decreased rapidly at 0−0.3%, and the downward trend slowed down at 0.6%−0.9%. Under the stress of neutral salt NaCl, Na2SO4 and alkaline salt NaHCO3, chlorophyll was most affected by NaCl. Under the treatment of 0.2% and 0.4% NaCl, the chlorophyll decreased by 24.1% and 34.0%, respectively, compared with the control.[43,45] 活性氧清除系统
Active oxygen scavenging system超氧化物歧化酶(SOD)、
过氧化物酶(POD)、过氧化氢酶(CAT)
Superoxide dismutase (SOD), peroxidase (POD), catalase (CAT)盐胁迫下SOD、POD和CAT活性均随胁迫天数增加先上升后下降, 且与对照相比均有增强, 变化程度为: SOD>CAT>POD
Under salt stress, the activities of SOD, POD and CAT increased first and then decreased with the days of stress, and increased compared with the control. The change degree of activity was SOD>CAT>POD.[20,22] 激素
Hormone脱落酸、生长素、赤霉素
Abscisic acid (ABA), auxin (IAA), gibberellin (GA3)ABA可诱导脯氨酸的大量积累, 从而维持细胞膜结构的稳定性, 提高保护酶的活性, IAA和GA3能够缓解盐胁迫对植物的伤害
ABA can induce a large amount of proline accumulation in plants, so as to maintain the stability of cell membrane structure and improve the activity of protective enzymes. IAA and GA3 can also alleviate the damage of salt stress to plants.[54-56]
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表 3 谷子耐盐相关基因克隆与功能分析
Table 3. Cloning and functional analysis of salt tolerance related genes in foxtail millet
基因名称
Gene name转基因受体与亚细胞定位
Transgenic receptor and subcellular localization诱导因素及基因功能
Inducing factors and gene function依赖途径
Dependent approach参考文献
ReferenceSiPHGPX 无细胞定位, 基于宏阵列的胁迫
特异性基因表达分析
No cell localization. Macroarray-based
stress-specific gene expression analysis受NaCl诱导, PHGPX基因产物在抗盐诱导的氧化损伤的防御反应中起着重要作用
Induced by NaCl, the PHGPX gene product plays an important role in the defense response against salt-induced oxidative damage.— [53] SiALDHs 谷子和水稻同源区
Homologous region of rice and
foxtail millet渗透胁迫、低温、过氧化氢和植物激素脱落酸(ABA)诱导基因表达量增加, 对蛋白质损伤、细胞膜破坏和细胞凋亡有保护功能, 提高耐盐性
Osmotic stress, low temperature, hydrogen peroxide and plant hormone abscisic acid (ABA) induce increased gene expression, which has protective functions against protein damage, cell membrane destruction and cell apoptosis, and improves salt tolerance.ABA依赖, 编码NAD(P)依赖性酶
ABA independent pathway, encoding NAD(P)-dependent enzyme[61] SiLEA14 细胞质
Cytoplasm受渗透胁迫、NaCl和外源脱落酸诱导, 提高谷子萌发期的抗渗透胁迫能力, 提高耐盐性和耐旱性
Induced by osmotic stress, NaCl and exogenous abscisic acid, it improves the resistance to osmotic stress during the germination of millet, and improves salt and drought tolerance.ABA依赖
ABA dependent pathway[62] SibZIPs 细胞核
Nucleus受高盐、干旱(PEG)和ABA胁迫诱导, 可通过与具有调控作用的蛋白(如蛋白激酶、NPR1相关蛋白)互作而介导调控谷子抵御胁迫反应
Induced by high salt, drought (PEG) and ABA stress, it can mediate and regulate millet to resist stress response by interacting with regulatory proteins (such as protein kinases, NPR1 related proteins).ABA依赖与非依赖途径
ABA dependent and independent pathways[71-72] SiARDPs 细胞核
Nucleus提高谷子耐旱耐盐性
Improve drought tolerance and salt tolerance of millet.ABA依赖
ABA dependent pathway[76] SiNF-Ys 细胞质、细胞核
Cytoplasm, nucleus受干旱、盐、甘露醇和氧化胁迫诱导, SiNF-YA1或SiNF-YB8通过增强抗氧化系统来增强胁迫耐受性
Induced by drought, salt, mannitol and oxidative stress, SiNF-YA1 or SiNF-YB8 enhances stress tolerance by enhancing the antioxidant system.SiNF-YA5 ABA非依赖,
SiNF-YA1和SiNF-YA8 ABA依赖
SiNF-YA5 through ABA independent pathway, SiNF-YA1 and SiNF-YA8 through ABA dependent pathway[77-78] SiVHA-E 液泡膜
Vacuole membrane被高盐、茉莉酸甲酯(MeJA)、水杨酸(SA)和ABA胁迫诱导表达, 提高植物耐盐性
Induced expression by high salt, methyl jasmonate (MeJA), salicylic acid (SA) and ABA stress to improve plant salt tolerance.ABA依赖
ABA dependent pathway[79] SiASR4 细胞核、细胞质和细胞膜
Nucleus, cytoplasm and cytomembrane受ABA、NaCl和PEG诱导, 提高谷子对干旱和盐胁迫的耐受性
Induced by ABA, NaCl and PEG, the tolerance of millet to drought and salt stress was improved.ABA依赖
ABA dependent pathway[80] SiLTPs 表皮细胞[烟草(Nicotiana tobacum L.)叶]、细胞质(玉米原生质体)
Epidermal cells (tobacco leaves),
cytoplasm (maize protoplasts)脱落酸(ABA)诱导基因表达量增加, 提高谷子耐盐性和耐旱性
Abscisic acid (ABA) increased gene expression and improved salt tolerance and drought tolerance of foxtail millet.ABA依赖
ABA dependent pathway[81] SiNAC110 细胞核
Nucleus被干旱、高盐等非生物胁迫诱导, 提高谷子对干旱和高盐胁迫的耐受性
It was induced by abiotic stress such as drought and high salt to improve the tolerance of millet to drought and high salt stress.ABA非依赖
ABA independent pathway[82] SiRLK35 质膜
Plasma membrane受NaCl、PEG、ABA、GA、MeJA胁迫诱导, 其中明显响应盐胁迫, 显示其对作物耐旱、抗盐及抗胁迫等方面具有重要调控作用
It was induced by NaCl, PEG, ABA, GA, MeJA stresses, and responded significantly to salt stress, indicating that it plays an important role in regulating drought tolerance, salt tolerance and stress resistance of crops.可能ABA依赖
It may be ABA dependent pathway.[73] SiCBL4 细胞膜、细胞质
Cytomembrane, cytoplasm受盐、ABA、甲基紫精醇(MV)、热休克和冷胁迫因素诱导, CBL-CIPK通路可被调控以提高植物耐盐性
Induced by salt, ABA, methyl viologen (MV), heat shock and cold stress, the CBL-CIPK pathway can be regulated to improve plant salt tolerance.可能ABA依赖
It may be ABA dependent pathway.[75,83] SiCIPK24 细胞质
Cytoplasm受盐、ABA、MV、热休克和冷胁迫因素诱导, CBL-CIPK通路可被调控以提高植物耐盐性
Induced by salt, ABA, MV, heat shock and cold stress, CBL-CIPK pathway can be regulated to improve plant salt tolerance.可能ABA依赖
It may be ABA dependent pathway.[83] SiNADP-ME 叶绿体、线粒体和细胞质
Chloroplast, mitochondria and cytoplasm被ABA、低温、PEG、NaCl胁迫诱导表达, 广泛参与谷子苗期非生物逆境胁迫应答
It was induced by ABA, low temperature, PEG and NaCl stress and widely participated in abiotic stress response of millet seedling.可能ABA依赖
It may be ABA dependent pathway.[84] SiPEPCs 细胞质、细胞核和线粒体
Cytoplasm, nucleus and mitochondria受ABA、低温、PEG、高盐胁迫诱导表达, 参与谷子对非生物逆境的应答, 可能在干旱和其他逆境胁迫信号途径中起关键作用
Induced by ABA, low temperature, PEG and high salt stress, it participates in the response of millet to abiotic stress, and may play a key role in drought and other stress signaling pathways.可能ABA依赖
It may be ABA dependent pathway.[74]
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