干旱和盐胁迫对盐生植物盐生草种子萌发特性的影响

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何建军, 姚立蓉, 汪军成, 边秀秀, 司二静, 杨轲, 王化俊, 马小乐, 李葆春, 尚勋武, 孟亚雄. 干旱和盐胁迫对盐生植物盐生草种子萌发特性的影响. 草业学报, 2020,29(11): 129-140
HE Jian-jun, YAO Li-rong, WANG Jun-cheng, BIAN Xiu-xiu, SI Er-jing, YANG Ke, WANG Hua-jun, MA Xiao-le, LI Bao-chun, SHANG Xun-wu, MENG Ya-xiong. Effects of drought and salt stress on seed germination characteristics of Halogeton glomeratus . Acta Prataculturae Sinica,2020,29(11): 129-140 复制到剪切板

Doi:10.11686/cyxb2020008
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干旱和盐胁迫对盐生植物盐生草种子萌发特性的影响

何建军^1,², 姚立蓉^1,², 汪军成^1,², 边秀秀², 司二静^1,², 杨轲^1,², 王化俊^1,², 马小乐^1,², 李葆春^2,³, 尚勋武¹, 孟亚雄^1,^2,^*

1.甘肃农业大学农学院,甘肃兰州 730070

2.甘肃省干旱生境作物学重点实验室,甘肃省作物遗传改良与种质创新重点实验室,甘肃兰州730070

3.甘肃农业大学生命科学技术学院,甘肃兰州 730070

*通信作者. E-mail: yxmeng1@163.com

作者简介:何建军(1995-),男,甘肃临洮人,在读硕士。E-mail: 3355829261@qq.com

收稿日期: 2020-01-07

基金项目:学科建设专项基金(GAU-XKJS-2018-082、083),国家大麦青稞产业技术体系(CARS-05-03B-03),甘肃省重大专项(17ZD2NA016)和盛彤笙创新基金(GSAU-STS-2018-33)资助

摘要

本研究以盐生植物盐生草为材料,以不同浓度的PEG-6000模拟干旱胁迫和以不同浓度的4种钠盐(NaCl、Na₂SO₄、NaHCO₃、Na₂CO₃)进行盐胁迫,探讨其对盐生草种子萌发特性的影响。结果表明:随着PEG-6000浓度的升高,盐生草种子发芽率、发芽势呈现下降的趋势,幼苗鲜重、干重、株高以及根系活力均呈现先上升后下降的趋势。聚类分析和主成分分析结果显示,以6% PEG-6000为结点,将不同浓度PEG-6000处理聚为两类,鲜重可被首选为评价盐生草萌发期耐干旱的重要参数;随着4种钠盐处理浓度的升高,发芽率、发芽势、幼苗鲜重、干重、株高以及根系活力均呈现不同程度的下降趋势,表明盐胁迫对种子的萌发以及幼苗的生长均有抑制作用,当Na⁺浓度相同时,4种钠盐对盐生草种子萌发及幼苗生长的抑制作用大小为Na₂CO₃>NaHCO₃>NaCl>Na₂SO₄,聚类分析和主成分分析结果显示,以50.00 mmol·L^-1 NaHCO₃,62.50 mmol·L^-1 Na₂SO₄,25.00 mmol·L^-1 Na₂CO₃ 和100.00 mmol·L^-1 NaCl为结点,将各钠盐胁迫浓度聚为两类,其中发芽指标可被首选为评价盐生草萌发期耐NaHCO₃、Na₂SO₄的重要参数,根系活力可被首选为评价盐生草萌发期耐Na₂CO₃的重要参数,干重可被首选为评价盐生草萌发期耐NaCl的重要参数。

关键词: 盐生草; 盐胁迫; 干旱胁迫; 萌发期

Effects of drought and salt stress on seed germination characteristics of Halogeton glomeratus

HE Jian-jun^1,², YAO Li-rong^1,², WANG Jun-cheng^1,², BIAN Xiu-xiu², SI Er-jing^1,², YANG Ke^1,², WANG Hua-jun^1,², MA Xiao-le^1,², LI Bao-chun^2,³, SHANG Xun-wu¹, MENG Ya-xiong^1,^2,^*

1. College of Agronomy, Gansu Agricultural University, Lanzhou 730070, China

2. Gansu Provincial Key Lab of Aridland Crop Science, Gansu Provincial Key Lab of Crop Improvement & Germplasm Enhancement, Lanzhou 730070, China

3. College of Life Sciences and Technology, Gansu Agricultural University, Lanzhou 730070, China

*Corresponding author. E-mail: yxmeng1@163.com

Abstract

To explore the effects of drought and salt stress on germination characteristics of halophytes, this study simulated drought stress with different concentrations of PEG-6000 and four sodium salts (NaCl, Na₂SO₄, NaHCO₃, Na₂CO₃) to evaluate the effects of those treatments on the seed germination characteristics of Halogeton glomeratus. It was found the germination percentage and germination energy decreased with increasing PEG-6000 concentration, while the seedling fresh weight, dry weight, plant height and root activity initially increased and then decreased at higher PEG-6000 concentration. Cluster analysis and principal component analysis showed that the different PEG-6000 concentrations could be divided into two groups with 6% PEG-6000 as the point of separation. Fresh weight appeared to be an important parameter. In addition, germination percentage germination energy, seedling fresh weight, dry weight, plant height and root activity declined with increase in concentration of the four sodium salts. This indicated that salt stress inhibited seed germination and seedling growth, with the effects of the four tested sodium salts ranked: Na₂CO₃>NaHCO₃>NaCl>Na₂SO₄. Cluster analysis and principal component analysis showed that the sodium salt concentrations were divided into two groups with the separation points falling at 50.00 mmol·L^-1 NaHCO₃, 62.50 mmol·L^-1 Na₂SO₄, 25.00 mmol·L^-1 Na₂CO₃ and 100.00 mmol·L^-1 NaCl. Among the plant traits evaluated as indicators of H. glomeratus stress tolerance in the germination stage, the germination index was the key indicator of NaHCO₃ and Na₂SO₄ stress, root activity emerged as the primary indicator of Na₂CO₃ stress and dry weight was the major indicator of NaCl stress in H. glomeratus.

Keyword: Halogen glomeratus; salt stress; drought stress; germination

文章图片

近年来, 水资源匮乏以及土壤盐渍化已成为影响农业发展及生态环境建设的两大全球性问题^[1]。目前, 全世界遭受水土流失的耕地面积约2.5× 10⁹ hm^2[2], 受到盐渍化威胁的全球耕地面积约为9.5× 10⁸ h^2[3], 在影响作物产量的各种非生物因素中, 干旱和盐碱造成的作物减产约40%, 严重影响了农作物的生长, 制约了农业经济的发展^[4]。干旱胁迫下, 植物体内由于水分失衡会发生一系列生理生化响应, 进而引起代谢紊乱, 影响植物正常生长^[5]。盐胁迫下会导致植物体内发生渗透胁迫、离子毒害作用, 扰乱植物体内营养元素平衡, 使其出现中毒甚至死亡的现象^[6]。

种子萌发期是植物对环境胁迫响应最敏感的时期, 直接影响植株的生长发育^[7]。在干旱及盐分胁迫下, 由于植物体内缺水和产生的离子毒害会使植物体内酶的活性发生一系列的变化, 进而限制植物生长发育^[8]。种子能否在干旱及盐碱环境下发芽成苗是决定该植株在该环境下完成整个生长发育过程的重要阶段^[9]。因此, 研究植物在萌发期的抗旱耐盐性, 以及了解植物抗旱耐盐的生理特点有着重要的意义。

目前, 利用模拟干旱和盐胁迫的方法对植物种子萌发特性的研究受到广泛应用^{[10, 11, 12]}。盐生草(Halogeton glomeratus)属藜科(Chenopodiaceae), 一年生草本植物, 广泛生长于甘肃、青海、新疆、内蒙古、宁夏等荒漠地区, 地上多分枝的肉质化茎、叶组织具有极强的抗旱、耐盐及富集重金属的特性, 被称为荒漠地区抗旱、耐盐先锋植物^{[13, 14]}, 然而关于盐生草种子萌发期的抗旱、耐盐性评价的报道较少。基于此, 本研究将盐生草种子分别置于不同浓度PEG-6000和NaCl、Na₂SO₄、NaHCO₃、Na₂CO₃溶液中, 模拟干旱以及不同盐分胁迫处理, 通过测定发芽指标、鲜重、干重、株高以及根系活力来分析盐生草种子对干旱及不同盐分的胁迫的耐受能力, 并对各项指标进行综合评价分析, 以期为盐生草在干旱、盐碱地的种植及生态治理提供理论依据。

1 材料与方法

1.1 试验材料

2018年11月在甘肃省会宁县盐碱地采集成熟的盐生草种子, 进行清洗、除杂后备用。千粒重为0.38 g。

1.2 试验方法

1.2.1 浓度设置根据预试验及参考文献[15]将PEG-6000的浓度梯度设置为CK(0)、2%、4%、6%、8%、10%、15%、20%。为保证4种钠盐处理中Na⁺浓度相同, 浓度梯度设置如表1所示。以上溶液均由蒸馏水配制。

表1 试验中4种不同浓度盐溶液的配制 Table 1 Preparation of four salt solutions with different concentrations in the test (mmol· L^-1)

1.2.2 萌发试验取饱满、大小均一的盐生草种子于离心管中置于超净工作台, 首先用无菌水冲洗种子表面杂质, 然后用75%的酒精灭菌消毒30 s, 再用无菌水清洗种子表面的酒精至干净, 后用10%的NaClO溶液颠倒清洗数次, 最后用无菌水冲洗4~5次, 得到无菌盐生草种子, 在直径为90 mm的垫有双层滤纸的培养皿中加入5 mL不同浓度的处理溶液, 每个处理均设置4个重复, 以蒸馏水作为对照。再将50粒盐生草种子整齐置于培养皿中, 并用封口膜封口, 每天用分析天平称量蒸发散失的水分, 并加蒸馏水补充, 使渗透势不变。后将培养皿置于培养室中保持温度22 ℃, 8 h光照/16 h 黑暗, 光照强度4000 Lx, 相对湿度80%的环境下进行萌发试验^[16]。

1.2.3 发芽指标测定以种子露白为标准^[16]。对3和7 d内发芽种子数进行统计。

发芽率=(7 d内种子发芽数/总种子数)× 100%

发芽势=(3 d内种子发芽数/总种子数)× 100%

1.2.4 株高扫描将生长7 d的盐生草幼苗, 以每盘培养皿为单位从中随机选取4株幼苗采用WinRHIZO 图像扫描仪(Epson Long Beach, CA, USA)进行株高扫描, 以平均值作为株高指标。

1.2.5 生物量测定生长7 d后, 由于盐生草幼苗只有细小幼根和刚展开的2片子叶, 单株生物量无法测量, 故以每盘培养皿为一个单位, 测定各不同胁迫浓度对单位幼苗生物量的影响, 用蒸馏水冲洗盐生草幼苗数次, 待冲洗干净后用滤纸吸干表面水分。用电子分析天平称其鲜重(fresh weight, FW)。然后装入信封袋中放置30~40 ℃烘箱, 烘3~4 d至恒重, 取出用电子分析天平称其干重(dry weight, DW)。

1.2.6 根系活力测定将生长了7 d的盐生草幼苗称取0.20 g, 采用2, 3, 5, -三苯基4唑氯化物(2, 3, 5-triphenyl-2H-tetrazolium chloride, TTC)还原测定法测定其根系活力^[17]。

1.3 数据处理

采用Microsoft Excel 2010进行数据处理和图表绘制, 运用SPSS 20.0软件(SPSS Inc., Chicago, USA)进行单因素方差分析(ANOVA), 主成分分析和聚类分析。所有结果由平均值± 标准误(means± standard error)表示, 4次生物学重复。

2 结果与分析

2.1 PEG-6000胁迫对盐生草种子萌发特性的影响

2.1.1 PEG-6000胁迫对盐生草种子萌发指标的影响由表2可知, 随着PEG-6000浓度的升高, 盐生草种子的发芽率和发芽势均呈下降的趋势, 且差异显著(P< 0.05)。当PEG-6000浓度为6%时, 发芽率和发芽势分别为62.00%, 11.33%, 与对照相比下降了31.30%, 86.35%。当PEG-6000浓度为20%时, 发芽率和发芽势分别为14.50%, 2.67%, 与对照相比下降了83.93%, 96.78%, 且差异显著(P< 0.05)。表明随着PEG-6000浓度的升高, 盐生草的种子活力会明显降低。在一定的PEG-6000浓度范围下, 盐生草种子可以萌发, 但其萌发明显受到抑制。

表2 盐生草种子发芽率和发芽势在PEG-6000处理下的变化 Table 2 Changes of germination percentage and germination energy of H. glomeratus seeds treated with PEG-6000 (%)

2.1.2 PEG-6000胁迫对盐生草幼苗鲜重和干重的影响如图1所示, 对PEG-6000胁迫处理7 d的盐生草幼苗测定鲜重、干重, 发现随着PEG-6000浓度的升高, 鲜重、干重均呈现先上升后下降的趋势, 当PEG-6000浓度为4%时均达到最大, 为0.17和0.012 g, 与对照相比差异不显著(P> 0.05)。当PEG-6000的浓度为20%时, 鲜重和干重分别为0.03和0.007 g, 与对照相比下降了75.00%, 36.36%, 且差异显著(P< 0.05)。因此, 在高浓度PEG-6000胁迫下, 盐生草幼苗的生长明显受到了抑制。

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	图1 盐生草幼苗鲜重和干重在PEG-6000处理下的变化不同小写字母表示差异显著(P< 0.05), 下同。Fig.1 Changes of fresh weight and dry weight of H. glomeratus seedlings treated with PEG-6000 The different lowercase letters indicate significant differences at 0.05 level, the same below.

2.1.3 PEG-6000胁迫对盐生草幼苗株高和根系活力的影响如图2所示, 株高的变化趋势与根系活力相近, 均呈现随着PEG-6000浓度的升高而先上升后下降的趋势。PEG-6000浓度为4%时均达到最大, 株高为6.34 cm, 与对照相比上升了1.28%, 但差异不显著(P> 0.05), 根系活力为107.51%, 与对照相比上升了7.51%, 且差异显著(P< 0.05)。当PEG浓度为20%时, 株高为3.82 cm, 与对照相比下降了38.98%, 根系活力为75.40%, 与对照相比显著下降了24.60%。

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	图2 盐生草幼苗株高和根系活力在PEG-6000处理下的变化Fig.2 Changes of plant height and root activity of H. glomeratus seedlings treated with PEG-6000

2.1.4 综合聚类分析及主成分分析为进一步研究PEG-6000胁迫对盐生草幼苗生长的抑制程度, 对PEG-6000胁迫处理下的种子萌发和幼苗生长指标进行聚类分析(图3), 以欧式距离15为界, 将PEG-6000处理的8个浓度梯度聚为了两类, 其中以浓度为0~6%为第Ⅰ 类, 8%~20%为第Ⅱ 类。得出PEG-6000胁迫盐生草萌发生长的临界耐受浓度为6%, 对6%胁迫处理下6个性状指标进行主成分分析(表3), 提取了2个主成分, 主成分1的特征值为4.27, 贡献率为70.27%, 主成分2的特征值为1.78, 贡献率29.73%。累计贡献率达100.00%, 其中主成分1在鲜重和干重上具有最高载荷(1.00), 主成分2在发芽势上具有最高载荷(0.87), 主成分1的贡献率约为主成分2的2倍。

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	图3 PEG-6000处理下不同浓度的聚类分析Fig.3 Clustering analysis of PEG-6000 treatments at different concentrations

表3 盐生草种子萌发特性在PEG-6000处理下的主成分分析 Table 3 Principal component analysis of germination characteristics of H. glomeratus seeds treated with PEG-6000

2.2 4种钠盐胁迫对盐生草种子萌发特性的影响

2.2.1 4种钠盐胁迫对盐生草种子萌发指标的影响如表4可知, 4种钠盐胁迫处理对盐生草种子萌发指标的影响有所不同, 随着Na₂SO₄浓度的升高, 盐生草种子的发芽率和发芽势呈现先升高后下降的趋势。在Na₂SO₄浓度为25.00 mmol· L^-1发芽率和发芽势均达到最大值, 发芽率为93.00%, 较对照升高4.49%。发芽势为81.50%, 较对照相比升高2.52%。当Na₂SO₄达到最大处理浓度150.00 mmol· L^-1时, 发芽率为9.00%, 较对照降低了89.89%。发芽势为4.67%, 较对照降低了94.13%。表明低浓度Na₂SO₄胁迫对植物生长具有一定的促进作用, 但超过一定的浓度后表现为抑制。随着Na₂CO₃, NaHCO₃, NaCl浓度的升高, 盐生草种子的发芽率和发芽势总体呈现下降的趋势。当NaCl达到最大处理浓度300.00 mmol· L^-1时, 发芽率较对照降低了96.26%; 发芽势较对照降低了97.48%。当NaHCO₃达到最大处理浓度300.00 mmol· L^-1时, 发芽率较对照降低了97.75%。发芽势较对照降低了97.48%。当Na₂CO₃达到最大处理浓度150.00 mmol· L^-1时盐生草种子不发芽。表明高浓度盐处理均对种子的萌发具有严重抑制作用。4种钠盐溶液相比, 除了具有相同的Na⁺外, NaHCO₃和Na₂CO₃具有较高的pH值。当4种盐处理在Na⁺相同的情况下, 100.00 mmol· L^-1 NaCl , 100.00 mmol· L^-1 NaHCO₃, 50.00 mmol· L^-1 Na₂CO₃, 50.00 mmol· L^-1 Na₂SO₄相比, 对盐生草种子萌发的毒害作用为Na₂CO₃> NaHCO₃> NaCl> Na₂SO₄。与对照相比差异显著(P< 0.05)。

表4 盐生草种子发芽势和发芽率在4种盐处理下的变化 Table 4 Changes of germination energy and germination rate of H. glomeratus seeds treated with four salts (%)

2.2.2 4种钠盐胁迫对盐生草幼苗株高的影响如图4所示, 盐生草幼苗株高的变化趋势与发芽势和发芽率相接近, 均随着Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄浓度的增加, 呈现下降的趋势, 且各浓度处理的盐生草幼苗株高均低于对照, Na₂SO₄处理在浓度低于37.50 mmol· L^-1时与对照差异不显著(P> 0.05), Na₂CO₃, NaHCO₃, NaCl各浓度处理均与对照差异显著(P< 0.05)。且当Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄的浓度分别为25.00, 50.00, 200.00, 150.00 mmol· L^-1时, 盐生草幼苗株高降为2.24, 2.18, 2.00, 1.99 cm, 与对照相比了均下降到对照的50.00%以下。Na₂CO₃, NaHCO₃在浓度为75.00~125.00 mmol· L^-1, 125.00~300.00 mmol· L^-1之间盐生草幼苗株高趋于稳定。在Na⁺浓度相同的处理条件下, 即Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄的浓度为50.00, 100.00, 100.00, 50.00 mmol· L^-1时, 4种盐处理下的盐生草幼苗株高为1.06, 1.17, 3.10, 3.66 cm, 说明4种盐处理对盐生草幼苗株高的抑制作用大小为 Na₂CO₃> NaHCO₃> NaCl> Na₂SO₄。

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	图4 盐生草幼苗株高在4种盐处理下的变化Fig.4 Changes of plant height of H. glomeratus seedlings treated with four salts

2.2.3 4种钠盐胁迫对盐生草幼苗根系活力的影响由图5可知, 根系活力的变化趋势与株高相近, 随着Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄浓度的增加, 各盐处理中盐生草幼苗根系活力均呈现下降的趋势, 且各处理组与对照差异显著(P< 0.05)。NaCl, Na₂SO₄处理浓度分别达到最大300.00 mmol· L^-1, 150.00 mmol· L^-1时, 盐生草幼苗根系活力分别为50.76%和51.70%。当Na₂CO₃, NaHCO₃浓度分别为87.50, 175.00 mmol· L^-1时, 根系活力均低于对照50.00%。说明盐胁迫对盐生草幼苗的根系活力有抑制作用。且在Na⁺浓度相同的处理条件下, Na₂CO₃对盐生草幼苗根系活力抑制作用最大, Na₂SO₄对其抑制作用最小。这说明碱性盐胁迫下对植株幼苗根系的毒害作用大于中性盐胁迫。

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	图5 盐生草幼苗根系活力在4种盐处理下的变化Fig.5 Changes of root activity of H. glomeratus seedlings treated with four salts

2.2.4 4种钠盐胁迫对盐生草幼苗鲜重及干重的影响随着Na₂CO₃, NaHCO₃, NaCl浓度的升高, 盐生草幼苗鲜重均呈现下降的趋势(图6)。Na₂CO₃, NaHCO₃, NaCl 的处理浓度分别在0~25.00 mmol· L^-1, 0~50.00 mmol· L^-1, 0~100.00 mmol· L^-1时与对照相比变化差异不显著(P> 0.05), 后者趋于稳定。随着Na₂SO₄浓度的增加, 盐生草幼苗鲜重呈现先缓慢上升后下降的趋势, 且处理与对照差异不显著(P> 0.05)。当NaHCO₃, NaCl, Na₂SO₄处理浓度达到最大时, 与对照相比其鲜重下降了99.21%, 95.26%, 67.20%。随着Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄ 浓度的升高, 盐生草幼苗的干重均呈现先上升后下降的趋势。当Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄浓度为37.50, 75.00, 125.00, 125.00 mmol· L^-1时, 盐生草幼苗的干重均降为对照的50.00%以下。

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	图6 盐生草幼苗鲜重和干重在4种盐处理下的变化Fig.6 Changes of fresh weight and dry weights of H. glomeratus seedlings treated with four salts

2.2.5 综合聚类分析及主成分分析首先运用模糊数学(Fuzzy数学)中的隶属函数法对单位不同的各项指标测定值进行标准化^[18], 然后对各项测定指标进行聚类分析(图7), Na₂CO₃胁迫处理下的不同浓度在欧式距离为15时聚为两类, 0~25.00 mmol· L^-1为第Ⅰ 类, 37.50~150.00 mmol· L^-1为第Ⅱ 类。NaHCO₃胁迫处理下的不同浓度在欧式距离为15时聚为两类, 0~50.00 mmol· L^-1为第Ⅰ 类, 75.00~300.00 mmol· L^-1为第Ⅱ 类。NaCl胁迫处理下的不同浓度在欧式距离为15时聚为两类, 0~100.00 mmol· L^-1为第Ⅰ 类, 125.00~300.00 mmol· L^-1为第Ⅱ 类。Na₂SO₄胁迫处理下的不同浓度在欧式距离为20时聚为两类, 0~62.50 mmol· L^-1为第Ⅰ 类, 75.00~150.00 mmol· L^-1为第Ⅱ 类。Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄胁迫盐生草幼苗萌发生长的临界耐受浓度分别为:25.00, 50.00, 100.00, 62.50 mmol· L^-1。将Na₂CO₃, NaHCO₃, NaCl, Na₂SO₄耐受临界浓度下的发芽势, 发芽率, 株高, 根系活力, 鲜重, 干重6个测定指标进行主成分分析(表5和表6), 得出不同盐胁迫下的2个主成分的累计贡献率均达到了100.00%。第1主成分的贡献率大约为第2主成分的2倍。结合4种盐胁迫的2个主成分及6个性状的相关系数可以发现, Na₂SO₄的第1主成分的特征值为3.92, 贡献率为65.41%, 其在鲜重上有最大载荷, 第2主成分的特征值为2.08, 贡献率为34.59%, 其在干重上有最大载荷。NaCl的第1主成分的特征值为4.30, 贡献率为71.71%, 其在干重上有最大载荷, 第2主成分的特征值为1.70, 贡献率为28.29%, 其在发芽势上有最大载荷。Na₂CO₃的第1主成分的特征值为3.85, 贡献率为64.24%, 其在根系活力上有最大载荷, 第2主成分的特征值为2.15, 贡献率为35.77%, 其在鲜重上有最大载荷。NaHCO₃的第1主成分的特征值为3.88, 贡献率为64.70%, 其在发芽率上有最大载荷, 第2主成分的特征值为2.12, 贡献率为35.30%, 其在干重上有最大载荷。

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	图7 4种盐处理下不同浓度的聚类分析Fig.7 Clustering analysis of four salt treatments at different concentrations

表5 2个因子的特征值及贡献率 Table 5 Eigen values of 2 principal components and their contribution and cumulative contribution

表6 各因子成分矩阵 Table 6 Factor matrix of each component

3 讨论

种子萌发期是植株生长的一个关键时期, 决定植物群体在逆境条件下能否建植成功。而种子的发芽率和发芽势是衡量种子品质的重要指标^{[19, 20]}。本研究表明:一方面, 在干旱胁迫下, 盐生草种子的发芽势和发芽率均呈现随PEG-6000浓度的升高而下降的趋势, 当PEG-6000浓度为20%时, 盐生草种子的发芽率为14.50%, 发芽势为2.67%, 且与对照相比差异显著(P< 0.05)。说明PEG-6000模拟的干旱胁迫对盐生草种子的萌发有抑制作用, 这与徐建欣等^[21]对新疆橡胶草(Taraxacum kok-saghyz)种子萌发试验的结论相吻合。另一方面, 在4种钠盐溶液胁迫下, 盐生草种子的发芽势和发芽率总体呈现随盐溶液浓度的升高而下降的趋势。NaCl, Na₂SO₄两种盐溶液低浓度时对种子萌发抑制作用没有显著差异, 这与李锦锋等^[22]对白茎盐生草(Halogeton arachnoideus)萌发试验的结论一致。刘克彪等^[23]利用钠盐胁迫处理罗布麻(Apocynum venetum)种子、阎秀峰等^[24]用碳酸盐处理星星草(Puccinellia tenuiflora)种子得出低浓度的碳酸盐对种子萌发有增效效应。本试验结果与之有所不同, 可能与Na₂CO₃和NaHCO₃设定浓度以及盐生草的耐盐碱性不同有关。NaCl, Na₂SO₄, Na₂CO₃, NaHCO₃在具有相同的Na⁺浓度的条件下, NaHCO₃和Na₂CO₃溶液呈碱性, pH值较高。当4种盐处理在Na⁺相同的情况下, 100.00 mmol· L^-1NaCl, 100.00 mmol· L^-1NaHCO₃, 50.00 mmol· L^-1Na₂CO₃, 50.00 mmol· L^-1Na₂SO₄时的发芽率53.00%, 13.00%, 11.00%, 80.50%, 说明4种盐处理对盐生草种子发芽的毒害作用为Na₂CO₃> NaHCO₃> NaCl> Na₂SO₄。李锦锋等^[22]用钠盐胁迫处理白茎盐生草种子, 得到NaCl对其萌发及生长的抑制作用强于Na₂SO₄。Lin等^[25]用中性盐和碱性盐模拟土壤盐碱化条件, 对羊草(Leymus chinensis)幼苗进行胁迫, 得出碱性盐对羊草幼苗的危害更加严重。

对处理7 d的盐生草幼苗的株高、干重、鲜重以及根系活力进行测定得出, 随着4种钠盐溶液浓度的升高, 盐生草幼苗的株高、鲜重、干重以及根系活力总体呈现下降的趋势。黄高鉴等^[26]研究了钠盐胁迫对柳枝稷(Panicum virgatum)生物量的影响, 表明盐胁迫能够显著降低其生物量(P< 0.05), 对柳枝稷生长起到明显的抑制作用。本研究中, 随着4种盐浓度的增加, 各钠盐处理下盐生草幼苗株高均呈梯度下降, 鲜重、干重均表现为持续降低的趋势, 且与对照相比差异均显著(P< 0.05), 说明盐处理对盐生草幼苗生长有明显抑制作用, 熊雪等^[27]研究发现紫花苜蓿(Medicago sativa)受到盐胁迫后发生渗透使植株生长受到抑制, 生物量显著降低。本研究观察7 d后幼苗的根系活力, 得出随着Na₂CO₃、NaHCO₃、NaCl、Na₂SO₄浓度的升高, 盐生草幼苗的根系活力均呈现下降的趋势。徐宁等^[28]研究了NaCl胁迫对高粱(Sorghum bicolor)种子萌发和幼苗生长特性的影响, 得到盐胁迫下幼苗根系活力随盐浓度的升高呈下降的趋势。本研究表明盐胁迫下离子毒害作用会降低幼苗根系活力。同时, 随着PEG-6000的浓度升高, 盐生草幼苗的株高、鲜重、干重以及根系活力, 总体呈现先上升后下降的变化趋势, 在4%时达到最大值, 但与对照相比差异不显著(P> 0.05)。在PEG-6000浓度为6%~20%时, 盐生草株高、鲜重、干重以及根系活力显著下降。这与杨智等^[29]研究得出干旱胁迫能使垂穗披碱草(Elymus nutans)的生物量和根系活力显著降低(P< 0.05)的试验结论相一致。表明干旱和盐胁迫都会抑制植物的萌发及幼苗的生长。Na₂SO₄处理下, 对盐生草幼苗的生长抑制作用最小, Na₂CO₃处理下, 对盐生草幼苗的生长抑制作用最大。

主成分分析是将原来各项指标重新组合成一组新的彼此独立的几个综合指标, 用少数的综合指标来反映原来多指标的信息^[30]。李丰先等^[31]运用此方法得出发芽率和干重可作为萌发期高粱耐碱性的主要评价指标。杨帆等^[32]得出根长、芽鲜重、发芽率可作为萌发期高粱耐盐性的主要评价指标。蔺军^[33]得出相对发芽指数、发芽率可作为萌发期燕麦(Avena sativa)耐盐性的主要评价指标。本研究通过综合聚类分析得出:盐生草耐Na₂CO₃、NaHCO₃、NaCl、Na₂SO₄的临界浓度为25.00, 50.00, 100.00, 62.50 mmol· L^-1, 耐PEG-6000的临界浓度为6%。以4种钠盐胁迫及PEG-6000胁迫的临界浓度的各项测定指标进行主成分分析得出, Na₂SO₄和NaHCO₃的发芽指标贡献率较大, NaCl的干重指标贡献率较大, Na₂CO₃的根系活力指标贡献率较大, PEG-6000的鲜重指标贡献率较大。因此生物量和发芽指标可作为评价盐生草萌发期耐盐及耐干旱的主要评价指标。

4 结论

综上所述, 随着PEG-6000及4种钠盐浓度的升高, 盐生草种子萌发和幼苗生长指标呈现不同程度的变化趋势, 6% PEG-6000胁迫下鲜重为主要鉴定指标。在NaHCO₃、Na₂SO₄的胁迫浓度分别为50.00、62.50 mmol· L^-1时萌发指标可作为评价盐生草萌发期耐盐胁迫的主要鉴定指标; Na₂CO₃的胁迫浓度为25.00 mmol· L^-1时根系活力可作为评价盐生草萌发期耐盐胁迫的主要鉴定指标; NaCl胁迫浓度为100.00 mmol· L^-1时干重为主要鉴定指标。本研究表明在6% PEG-6000环境下以及50.00 mmol· L^-1 NaHCO₃、62.50 mmol· L^-1 Na₂SO₄、25.00 mmol· L^-1 Na₂CO₃、100.00 mmol· L^-1 NaCl下, 盐生草均可正常萌发生长。本研究初步揭示了盐生草萌发期对干旱胁迫以及盐胁迫的耐性程度, 为以后干旱、盐碱地种植盐生草进行生态修复提供了一定的参考依据。

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[24]	Yan X F, Sun G R. Salt stress on seed germination of Puccinellia tenuiflora. Pratacultural Science, 1994, 11(4): 27-31. 阎秀峰, 孙国荣. 盐分对星星草种子萌发的胁迫作用. 草业科学, 1994, 11(4): 27-31. [本文引用:1]
[25]	Lin J X, Li Z L, Wang Y N, et al. *Effects of various mixed salt-alkaline stress conditions on seed germination and early seedling growth of Leymus chinensis* from Songnen grassland of China*. Notulae Botanicae Horti Agrobotanici Cluj-Napoca*, 2014, 42(1): 154-159. [本文引用:1]
[26]	Huang G J, Wang B, Yang Z P, et al. Effects of salt stress on biomass and quality of switchgrass. Tianjin Science & Technology, 2019, 46(2): 26-28. 黄高鉴, 王斌, 杨治平, 等. 盐胁迫对柳枝稷生物量及品质的影响. 天津科技, 2019, 46(2): 26-28. [本文引用:1]
[27]	Xiong X, Luo J C, Wei Y Q, et al. Effects of non-uniform salt stress on growth characteristics of alfalfa. Scientia Agricultura Sinica, 2018, 51(11): 2072-2083. 熊雪, 罗建川, 魏雨其, 等. 不均匀盐胁迫对紫花苜蓿生长特性的影响. 中国农业科学, 2018, 51(11): 2072-2083. [本文引用:1]
[28]	Xu N, Cao N, Wang C, et al. Effects of NaCl stress on seed germination and seedling physiological characteristics of wild and cultivated varieties of Sorghum bicolor. Jiangsu Agricultural Sciences, 2018, 46(18): 55-57, 67. 徐宁, 曹娜, 王闯, 等. NaCl 胁迫对野生和栽培品种高粱种子萌发和幼苗生理特性的影响. 江苏农业科学, 2018, 46(18): 55-57, 67. [本文引用:1]
[29]	Yang Z, Chen S Y, Zhou Q P, et al. *Effects of exogenous silicon on root morphology of Elymus nutans* seedlings under simulant drought stress*. Pratacultural Science, 2019, 36(5): 1371-1378. 杨智, 陈仕勇, 周青平, 等. 外源硅添加对模拟干旱胁迫下垂穗披碱草苗期根系形态的影响*. 草业科学, 2019, 36(5): 1371-1378. [本文引用:1]
[30]	Tian X X, Mao P C, Li S S, et al. Determination of salt-tolerant indexes and comprehensive evaluation of salt-tolerance at the seedlings of Medicago sativa. Acta Agrestia Sinica, 2017, 25(3): 545-553. 田小霞, 毛培春, 李杉杉, 等. 紫花苜蓿苗期耐盐指标筛选及耐盐性综合评价. 草地学报, 2017, 25(3): 545-553. [本文引用:1]
[31]	Li F X, Zhou Y F, Wang Y T, et al. Screening and identification of sorghum bicolor cultivars for alkali tolerance during germination. Scientia Agricultura Sinica, 2013, 46(9): 1762-1771. 李丰先, 周宇飞, 王艺陶, 等. 高粱品种萌发期耐碱性筛选与综合鉴定. 中国农业科学, 2013, 46(9): 1762-1771. [本文引用:1]
[32]	Yang F, Wei X C, Zhang S C, et al. *Comparison on salt and drought resistances of different varieties of Sorghum dochna* at germination stage*. Plant Physiology Journal, 2015, 51(10): 1604-1610. 杨帆, 魏晓岑, 张士超, 等. 不同甜高粱品种萌发期抗盐和抗旱性比较*. 植物生理学报, 2015, 51(10): 1604-1610. [本文引用:1]
[33]	Lin J. *Salt tolerance and drought resistance of Secale cereal* and Avena sativa during the seed germination stage*. China Herbivore Science, 2016, 36(4): 31-34. 蔺军. 黑麦和燕麦种子萌发期耐盐性、抗旱性比较*. 中国草食动物科学, 2016, 36(4): 31-34. [本文引用:1]

2017

0.0

... 近年来,水资源匮乏以及土壤盐渍化已成为影响农业发展及生态环境建设的两大全球性问题^[1] ...

2017

0.0

... 10⁹ hm^2[2],受到盐渍化威胁的全球耕地面积约为9 ...

2017

0.0

... 10⁸ h^2[3],在影响作物产量的各种非生物因素中,干旱和盐碱造成的作物减产约40%,严重影响了农作物的生长,制约了农业经济的发展^[4] ...

2019

0.0

... 10⁸ h^2[3],在影响作物产量的各种非生物因素中,干旱和盐碱造成的作物减产约40%,严重影响了农作物的生长,制约了农业经济的发展^[4] ...

2013

0.0

. 2013, 33(18):5477-5483 DOI:10.5846/stxb201304270839

Relationship among drought, hydraulic metabolic, carbon starvation and vegetation mortality

植物干旱胁迫下水分代谢、碳饥饿与死亡机理

Dong L , Li J Y

董蕾, 李吉跃

Plants would be affected by different environmental factors during the whole life cycles. Events of regional-scale vegetation mortality appear more and more frequently with the global change which is associated with increased temperatures, droughts, and often (but not always) with biotic agents (such as insects and pathogens) outbreaks. In recent years, consistent predictions of a warming climate went with more frequent and severe droughts, coupled with effects of heat and drought on vegetation mortality, indicate that widespread mortality events are a likely future phenomenon. These harsh factors and predictions required us have to understand plant mortality mechanism, although these precise physiological mechanisms were poorly understood. The difficulties of these problems were about interaction among environmental factors and interaction among plant physiological action. For example, drought was always appeared coupled with heat. When heat appeared, plants would have some physiological reaction response to heat and drought. If stress lasted for a long time, plants would die. But the signal mechanism to understand of drought or heat death was hard to achieve. On the other hand, even drought was artificial controlled, plants physiological processes were interacted. When plants were under drought, stomata would close as humidity was low. Because of gas exchange decreasing, photosynthesis and respiration were down-regulated. As a result, carbon sink and storage were damaged under drought. But we cannot understand precisely about where, when and how vegetation die in each physiological process. As discussed above, plants would down-regulated photosynthesis, transpiration and therefore impede plant growth. In order to maintain metabolism, plants have to decrease respiration. Long lasting drought will lead to imbalance of carbohydrate storage which would eventually cause carbon starvation (failure to maintain metabolism, carbon supply from photosynthesis and the mobilization of nonstructural carbohydrate (NSC) is less than carbon use for respiration, growth and defense). Plants need NSC to maintain osmotic adjustment when water stress made soil water content is low, but this subsequently elevates the minimum survivable NSC content. On the other hand, drought would induce another adverse affect-xylem embolism and cavitation (phloem column was filled with gas or air caused by drought). Because of xylem embolism, water and nutrient were hard to transport to the top of the leaves. This can cut off organs that require NSC from storage or new NSC assimilated. Refilling of embolized xylem is an energy-requiring process in many species that depends on conversion of starch to sugars for aquaporin regulation and for the creation of an osmotic gradient within embolized conduits. If drought blocked refilling cavitation, xylem embolism would become more severe and subsequently decrease hydraulic conductivity, photosynthesis and increase turgor loss. This is a negative feedback loop that makes plants have to choose whether refilling the cavitation or storage for living. No matter how plants choose, carbon starvation would lead to a further hydraulic failure and push plants faster toward the threshold for hydraulic failure or carbon starvation. In this review, we discussed about the mechanisms of drought-induced mortality from carbon and hydraulic metabolism. Xylem embolism and cavitation which were induced by drought were also mentioned. Plants physiological mechanisms of mortality were complex and difficult to measure accurately. We could forecast these events and prevent vegetation mortality only if we understand these mechanisms deeper and deeper.

植物在生长发育过程中受众多环境因子共同作用。随着全球气候变化,气温升高、降水量下降等问题频繁出现。目前气象学家一致预测未来环境变暖会使干旱更加频繁剧烈,这一环境改变使植物死亡更加严重。植物在水分胁迫、特别是干旱胁迫条件下,体内水分代谢与碳代谢会发生失衡现象:光合速率降低、蒸腾速率降低,带来生长降低;为维持植物新陈代谢,植物呼吸作用必然下调。在长期干旱胁迫条件下植物体内碳水化合物储存发生失衡现象,这种失衡使植物陷入碳饥饿现象。另外,由于水分失衡而出现的木质部栓塞和空穴会进一步加剧水分运输障碍,而修复空穴则需要大量非结构性碳水化合物(NSC),这使植物陷入两难选择。总结了植物干旱胁迫下,碳饥饿与水分代谢、植物死亡关系的相关研究,对未来的研究方向和重点提出建议,以期对未来的植物死亡研究提供帮助。

... 干旱胁迫下,植物体内由于水分失衡会发生一系列生理生化响应,进而引起代谢紊乱,影响植物正常生长^[5] ...

2013

0.0

... 盐胁迫下会导致植物体内发生渗透胁迫、离子毒害作用,扰乱植物体内营养元素平衡,使其出现中毒甚至死亡的现象^[6] ...

2012

0.0

. 2012, 29(7):1105-1113

Evaluation on the salt tolerance of 38 germplasm resources of Elytrigria repens at seed germination stage

种子萌发期38份偃麦草种质耐盐性评价

Sun Z J , Li P Y , Ji P F

孙宗玖, 李培英, 吉鹏飞

Based on simulation salt stress with mixture of 1.2% of NaCl∶Na2SO4(1∶1), the routine paper bed culture was used to estimate the salt tolerance of 38 germplasm resources of Elytrigria repens at the seed germination stage by measuring the germination rate, germination energy, germ length, embryonic root length, vitality index, germination salt tolerance index and germination stress index. The results of this study showed that the salt stress decreased the germination rate, germination energy, germination Index and vitality index of E.repens, inhibited the growth of E.repens germ and radical, and also decreased the germination characteristic of E.repens. According to the comprehensive evaluation value, the salt tolerance sequence of 38 E.repens accessions from high to low was E03, E31, E16, E17, E42, E26, E04, E25, E12, E38, E15, E36, E13, E32, E09, E21, E07, E08, E01, E29, E10, E18, E24, E28, E30, E23, E11, E39, E02, E34, E19, E35, E06, E40, E33, E41, E37, E27.

为评价偃麦草(Elytrigia repens)种子萌发期耐盐性强弱，采用常规纸上发芽法，以1.2%氯化钠和硫酸钠（质量比为1∶1）模拟盐胁迫，对38份偃麦草的发芽率、发芽势、胚芽长、胚根长、活力指数、萌发耐盐指数、萌发胁迫指数等进行测定。结果表明，盐胁迫降低了偃麦草种子的发芽率、发芽势、发芽指数、活力指数，抑制了其胚芽和胚根的生长，降低了偃麦草萌发性能；根据综合评价值，38份偃麦草材料的耐盐性由强到弱依次为E03、E31、E16、E17、E42、E26、E04、E25、E12、E38、E15、E36、E13、E32、E09、E21、E07、E08、E01、E29、E10、E18、E24、E28、E30、E23、E11、E39、E02、E34、E19、E35、E06、E40、E33、E41、E37、E27。

... 种子萌发期是植物对环境胁迫响应最敏感的时期,直接影响植株的生长发育^[7] ...

2016

0.0

... 在干旱及盐分胁迫下,由于植物体内缺水和产生的离子毒害会使植物体内酶的活性发生一系列的变化,进而限制植物生长发育^[8] ...

2011

0.0

. 2011, 31(6):1148-1152 DOI:10.7606/j.issn.1009-1041.2011.06.026

Effects of salt and alkali stresses on seed germination, early seedling growth and the metabolize of Na+ and K+ in shoots of wheat

盐碱胁迫对小麦种子萌发、早期幼苗生长及Na+、K+代谢的影响

Lin J X , Li X Y , Tang J H

蔺吉祥, 李晓宇, 唐佳红

Wheat( Triticum aestivum ) cv.Jimai 3 bred by Jilin Agricultural University were used as the experimental material to study the effects of salt(NaCl∶Na 2 SO 4 =4∶1) and alkali(Na 2 CO 3 ∶NaHCO 3 =4∶1) stresses on seed germination, early seedling growth and metabolism responses of Na + and K + . We also compared the ability of salt alkali tolerance between germination and early seedling stages. The results showed that the reductions of seed germination, germination rate and early seedling growth were greater under alkali stress than that under salt stress, most of non germinated seeds under higher salt stress germinated well after transferring from high salinity to distilled water while those under higher alkalinity stress lost viability. The inhibition of salt alkali stresses on roots was much greater than that on shoots, and germination stage was more tolerant to those stresses than early seedling stage. The Na + contents and Na + /K + ratios increased much greater under alkali stress, especially in roots. The K + contents in both shoots and roots were not affected under salt stress, but decreased under alkali stress. The results suggested that the destructive effects of alkali stress on seed viability, early seedling growth and physiological changes of wheat are severer than that of salt stress. Higher Na + concentration in roots is an important feature of wheat tolerance to salinity alkalinity in early seedling stage. The different saline alkaline tolerant abilities between the two stages suggested that germinated seeds may not grow to the establishment of seedlings.

为给盐碱地区小麦合理种植提供依据，以春小麦品种吉麦3号为材料，混合两种中性盐（NaCl∶Na 2 SO 4 =4∶1）与两种碱性盐（Na 2 CO 3 ∶NaHCO 3 =4∶1）分别模拟不同强度的盐、碱胁迫条件，探讨小麦种子萌发、早期幼苗生长及Na + 、K + 代谢对其的响应方式，并比较种子萌发期与早期幼苗生长阶段耐盐碱能力的强弱。结果表明，碱胁迫使小麦种子的发芽率、发芽速率及早期幼苗生长的下降幅度大于盐胁迫，高盐环境下未萌发的种子在蒸馏水中复萌较好，而高碱环境下则多数死亡。盐碱胁迫对小麦早期幼苗根生长的抑制程度显著大于茎叶，并且该阶段的耐盐碱性明显低于种子萌发期。随着盐碱胁迫浓度的增加，小麦幼苗根与茎叶中的Na + 含量与Na + /K + 显著增加，且在碱胁迫下，尤其是根中Na + 含量和Na + /K + 值更高，盐胁迫下K + 含量在根与茎叶中变化不明显，而在碱胁迫下则呈下降趋势。上述结果说明，盐胁迫与碱胁迫对小麦种子活力和早期幼苗生长与生理代谢的影响有着明显的不同，根中大量积累Na + 是小麦在早期幼苗阶段耐盐碱性的重要生理特征。小麦种子萌发期与早期幼苗生长阶段耐盐碱性具有明显的差异，盐碱胁迫下种子即使能够萌发也不意味着能够继续生长成苗。

... 种子能否在干旱及盐碱环境下发芽成苗是决定该植株在该环境下完成整个生长发育过程的重要阶段^[9] ...

2005

0.0

. 2005, 25(11):3061-3066

Impact of global climate change on wildlife

全球气候变化对野生动物的影响

Ma R J , Jiang Z G

马瑞俊, 蒋志刚

The Earth's climate has warmed by approximately 0.3～0.6℃ over the past 100 years,with the 1990s being the warmest decade.There is now ample evidence of the impacts of recent climate change on wildlife,from polar terrestrial to tropical marine environments.The recent climatic changes have affected a broad range of wildlife with diverse geographical distribution, for instance,the species distribution,the phenology,the animal's behavior and population dynamics of wild animals.The Qinghai-Tibetan plateau is one...

全球气候变化及其影响引起了人们的关注。随着全球气候变暖,北半球物候期提前,一些野生动物的分布区北移,动物的繁殖、种群变化都发生了不同程度的变化,有的物种甚至灭绝。青藏高原是全球变化的敏感地区之一,位于青藏高原东北部的青海湖地区正向暖干化方向发展。与历史分布比较,青海湖地区现生动物的分布和组成发生了较大的变化。设计自然保护区时,考虑全球气候变化对野生动物的影响,有利于保护物种多样性,有利于保持生态系统的功能完整性。

... 目前,利用模拟干旱和盐胁迫的方法对植物种子萌发特性的研究受到广泛应用^[10,11,12] ...

2010

0.0

... 目前,利用模拟干旱和盐胁迫的方法对植物种子萌发特性的研究受到广泛应用^[10,11,12] ...

2014

0.0

... 目前,利用模拟干旱和盐胁迫的方法对植物种子萌发特性的研究受到广泛应用^[10,11,12] ...

2015

0.0

... 盐生草(Halogeton glomeratus)属藜科(Chenopodiaceae),一年生草本植物,广泛生长于甘肃、青海、新疆、内蒙古、宁夏等荒漠地区,地上多分枝的肉质化茎、叶组织具有极强的抗旱、耐盐及富集重金属的特性,被称为荒漠地区抗旱、耐盐先锋植物^[13,14],然而关于盐生草种子萌发期的抗旱、耐盐性评价的报道较少 ...

2019

0.0

2019

0.0

... 1 浓度设置根据预试验及参考文献[15]将PEG-6000的浓度梯度设置为CK(0)、2%、4%、6%、8%、10%、15%、20% ...

2018

0.0

. 2018, 38(17):6171-6179 DOI:10.5846/stxb201801140105

Exogenous cGMP regulates seed germination of ryegrass under salt stress

外源cGMP调控盐胁迫下黑麦草种子萌发机制

Su M F , Wei X H , Xin X Q

宿梅飞, 魏小红, 辛夏青

cGMP (cyclic guanosine-3',5'-monophosphate) is a type of cyclic nucleotide and signaling molecule that can be responsive to plant abiotic stress. Using cGMP membrane permeability analog 8-Br-cGMP as a donor, this study investigated the effects of cGMP on seed germination and physiological indicators of ryegrass under salt stress. The effects of cGMP on the salt tolerance mechanism during the germination of ryegrass were determined through calculating the germination index and determining the physiological indicators in the process of seed germination. The results showed that a low concentration (50 mmol/L) of salt stress could promote the germination of ryegrass seeds. With increasing salt concentrations (100, 150, 200, and 250 mmol/L), the germination rate of ryegrass seeds gradually decreased and was even inhibited. Adding 20 μmol/L cGMP can relieve the damage caused by 100 mmol/L NaCl stress on ryegrass seed, increasing the germination rate, germination potential, germination index, and vigor index of ryegrass seed by 7.4%, 133.3%, 52.1%, and 104.2%, respectively. In addition, compared to the 100 mmol/L NaCl stress condition alone, the growth of roots of ryegrass could be promoted and the total length, root length, and leaf length of ryegrass plants increased, by 2.5, 2.8, and 2.6 times, respectively, when 20 μmol/L cGMP was added. The soluble sugar, soluble protein content, amylase activity, and proline content of seeds increased by 47.9%, 15.7%, 94.3%, and 117.4%, respectively, while starch content, MDA content, conductivity, and O 2 - . production rate were reduced by 40.9%, 128.7%, 88.6%, and 211.9%, respectively. The results showed that 20 μmol/L cGMP could increase the contents of soluble sugar, soluble protein, and proline, and reduce the production of MDA and O 2 - , so as to relieve the damage caused by salt stress on ryegrass seed and promote the hydrolysis of starch, which accelerated seed germination.

cGMP（cyclic guanosine-3'，5'-monophosphate）是一类环化核苷酸，能响应植物非生物胁迫的信号分子。以cGMP的膜透性类似物8-Br-cGMP为供体，研究cGMP对盐胁迫下黑麦草种子萌发及生理指标的影响。通过计算萌发指标、测定种子萌发过程中各项生理指标，探讨cGMP对黑麦草种子萌发时耐盐机制的作用。结果表明：低浓度（50 mmol/L）盐胁迫能促进黑麦草种子的萌发，随着盐浓度（100、150、200、250 mmol/L）的增加黑麦草种子的萌发率逐渐降低甚至抑制其萌发。而加入20 μmol/L cGMP能缓解100 mmol/L NaCl胁迫对黑麦草种子造成的伤害，使黑麦草种子的发芽率、发芽势、发芽指数、活力指数分别提高了7.4%、133.3%、52.1%、104.2%。此外，研究发现加入20 μmol/LcGMP能够促进100 mmol/L NaCl盐胁迫下黑麦草萌发期根的生长，比单独100 mmol/L NaCl盐胁迫处理下的黑麦草植株全长、根长、叶长分别提高了2.5、2.8倍和2.6倍，种子在萌发过程中可溶性糖、可溶性蛋白含量、淀粉酶活性，脯氨酸含量分别提高了47.9%、15.7%、94.3%、117.4%，而淀粉含量、MDA含量、电导率，O 2 - 产生速率分别降低了40.9%、128.7%、88.6%、211.9%。说明20 μmol/LcGMP通过提高可溶性糖，可溶性蛋白含量及脯氨酸含量，同时减少MDA、O 2 - 的产生，以此缓解盐胁迫对黑麦草种子的伤害、同时促进淀粉水解，从而加速种子萌发。

... 后将培养皿置于培养室中保持温度22 ℃,8 h光照/16 h 黑暗,光照强度4000 Lx,相对湿度80%的环境下进行萌发试验^[16] ...

... 3 发芽指标测定以种子露白为标准^[16] ...

2011

0.0

... 20 g,采用2,3,5,-三苯基4唑氯化物(2,3,5-triphenyl-2H-tetrazolium chloride,TTC)还原测定法测定其根系活力^[17] ...

2018

0.0

... 5 综合聚类分析及主成分分析首先运用模糊数学(Fuzzy数学)中的隶属函数法对单位不同的各项指标测定值进行标准化^[18],然后对各项测定指标进行聚类分析(图7),Na₂CO₃胁迫处理下的不同浓度在欧式距离为15时聚为两类,0~25 ...

2010

0.0

... 而种子的发芽率和发芽势是衡量种子品质的重要指标^[19,20] ...

2017

0.0

... 而种子的发芽率和发芽势是衡量种子品质的重要指标^[19,20] ...

2015

0.0

... 说明PEG-6000模拟的干旱胁迫对盐生草种子的萌发有抑制作用,这与徐建欣等^[21]对新疆橡胶草(Taraxacum kok-saghyz)种子萌发试验的结论相吻合 ...

2014

0.0

... NaCl,Na₂SO₄两种盐溶液低浓度时对种子萌发抑制作用没有显著差异, 这与李锦锋等^[22]对白茎盐生草(Halogeton arachnoideus)萌发试验的结论一致 ...

... 李锦锋等^[22]用钠盐胁迫处理白茎盐生草种子,得到NaCl对其萌发及生长的抑制作用强于Na₂SO₄ ...

2016

0.0

. 2016, 25(5):214-221 DOI:10.11686/cyxb2015318

Responses of Apooceynum venetum seed germination to drought and salt stress

干旱和钠盐胁迫对罗布麻种子萌发的影响

Liu K B , Jiang S X

刘克彪, 姜生秀

1.Gansu Psammophytes Engineering Technology Research Center, Lanzhou 730070, China; 2.Minqin Desert Botanical Garden, Minqin 733300, China

In order to study the seed germination characteristics of Apoceynum venetum under different stresses of drought and sodium salt, germination was tested with four different osmotic potential and sodium salt solutions (NaCl, Na 2 SO 4 , NaHCO 3 , compound sodium salt). The results showed that when seeds were treated with the PEG high osmotic potential (-0.05 MPa), low concentration of Na 2 SO 4 (0.2%) and the compound salt solutions (0.2% to 0.6%), the germination rate, index and drought index were more than CK, the average germination times were less than CK and salt damage rate was a negative value. Germination peak was in 2-4 d. The solutions promoted seed germination according to decreasing osmotic potential and increasing salt concentration. The germination potential, rate, index, and drought index gradually reduced, average germination time extended and salt damage rate increased, with germination peak at 2-3 d. When treated with the NaCl and NaHCO 3 solutions, increasing concentrations caused the germination rate, potential and index to significantly decrease. The average germination time was delayed, germination peak was in 2-3 d, seed germination penetration and toxic effects were enhanced. When treated with the PEG and compound salt solutions, the appropriate value, critical value and limit value of osmotic potential and concentration was -0.09, -0.70, -1.30 MPa and 0.85%, 2.04%, 3.22% respectively. When treated with the same concentration of salt solution, the salt injury rate was NaHCO 3 >NaCl>Na 2 SO 4 >compound salt.

为探明罗布麻种子在不同水分和钠盐胁迫下的萌发特性,通过模拟盐渍化土壤溶液渗透势和主要阴离子组成,研究罗布麻种子在PEG-6000不同渗透势溶液和不同钠盐(NaCl、Na 2 SO 4 、NaHCO 3 、复合钠盐)溶液中的萌发情况。结果表明,用PEG高渗透势(-0.05 MPa)溶液、低浓度Na 2 SO 4 (0.2%)和复合钠盐(0.2%~0.6%)溶液处理种子,发芽率、发芽势、发芽指数、萌发抗旱指数>CK,平均发芽时间<CK,盐害率为负值,发芽高峰期集中于2~4 d,溶液对种子发芽具有促进和增效作用,随着溶液渗透势降低和盐浓度的增加,发芽率、发芽势、发芽指数、萌发抗旱指数逐步降低,平均发芽时间延长,盐害率增大,发芽高峰期集中于2~3 d。NaCl、NaHCO 3 溶液处理种子,随着浓度的增加,发芽率、发芽势、发芽指数有明显降低的趋势,平均发芽时间延迟,发芽高峰期集中于2~3 d,渗透胁迫和毒害作用增强,溶液对种子萌发有明显的抑制和延缓作用。PEG、复合盐溶液处理种子,渗透势和浓度的适宜值、临界值与极限值分别为-0.09,-0.70,-1.30 MPa和0.85%,2.04%,3.22%。用相同浓度的钠盐溶液处理种子,盐害率大小总体依次为NaHCO 3 >NaCl>Na 2 SO 4 >复合钠盐。

... 刘克彪等^[23]利用钠盐胁迫处理罗布麻(Apocynum venetum)种子、阎秀峰等^[24]用碳酸盐处理星星草(Puccinellia tenuiflora)种子得出低浓度的碳酸盐对种子萌发有增效效应 ...

1994

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2014

0.0

... Lin等^[25]用中性盐和碱性盐模拟土壤盐碱化条件,对羊草(Leymus chinensis)幼苗进行胁迫,得出碱性盐对羊草幼苗的危害更加严重 ...

2019

0.0

... 黄高鉴等^[26]研究了钠盐胁迫对柳枝稷(Panicum virgatum)生物量的影响,表明盐胁迫能够显著降低其生物量(P#cod#x0003C ...

2018

0.0

. 2018, 51(11):2072-2083 DOI:10.3864/j.issn.0578-1752.2018.11.005

Effects of non-uniform salt stress on growth characteristics of alfalfa

不均匀盐胁迫对紫花苜蓿生长特性的影响

Xiong X , Luo J C , Wei Y Q

熊雪, 罗建川, 魏雨其

1 College of Agro-Grassland Science, Nanjing Agricultural University, Nanjing 210095 ; 2 Hebei Normal University for Nationalities, Chengde 067000, Hebei ; 3 Institute of Grassland Research of Chinese Academy of Agricultural Sciences, Huhhot 010010 ; 4 College of Animal Science and Technology, China Agricultural University, Beijing 100094

【 Objective 】 Soil salinity is often non-uniform in salt fields. Previous studies found that non-uniform salt stress could alleviate salt damage to plants compared to uniform salt treatment. The study on the response of alfalfa under non-uniform saltinity was an important supplement to plant physiology under uniform saltinity, so as to provide references for cultivation and management in salt fields.【Method】The root system of alfalfa was divided into two parts, including salt free treatment (0/0, NaCl concentrations were zero in both root systems）, non-uniform salt stress with a NaCl concentration of zero in one side of the roots (0/S: 0/100, 0/150, 0/200, 0/250), non-uniform salt stress with a NaCl concentration of 50 in one side of the roots (50/S: 50/100, 50/150, 50/200, 50/250) and uniform salt stess (S/S: 50/50, 100/100, 150/150, 200/200, 250/250 ) treatments ( “/” on both sides represent different salt stress concentrations, indicated by NaCl concentration mmol·L -1 ). After treatments for 15 days, the plant growth rate, biomass, water uptake, sodium and potassium ion concentration and malondialdehyde content were measured, and the effects of non-uniform salt stress on the growth characteristics of alfalfa were studied. 【Result】Salt stress inhibited the growth of alfalfa, and the plant growth rate, biomass, water uptake decreased, chlorophyll contents declined, proline contents increased, membrane lipid peroxidation aggravated, Na + concentrations increased, K + concentrations decreased, showed a lower K + /Na + ratio. Compared with the uniform high salt S/S treatments, the plant growth rate and aboveground biomass of 0/S, 50/S treatments increased by 57.05%-369.34% and 15.47%-42.57%, respectively, and the Na + concentrations decreased by 15.85%-55.93%, alleviating the toxic effect of Na + . And 70%-92% water uptake from non-uniform salt stress came from the no-saline or low-saline part, increased water uptake of the whole plant. 【Conclusion】The non-uniform salt stress treatments with a higher K + /Na + increased the chlorophyll contents of alfalfa and decreased the degree of membrane lipid peroxidation compared with the uniform salt stress treatments. The no-saline or low-saline part showed compensated water uptake and the compensatory growth of the roots, which promote the growth of plants, increase the shoot and root biomass. The more difference between the two sides of the roots, the effects on relieve salt damage were more obvious.

【目的】土壤中的盐分通常存在不均匀分布的现象。已有的研究发现与均一盐胁迫环境相比,不均匀盐胁迫可以缓解盐分对植物的伤害。研究旨在了解植物对不均匀盐分胁迫的响应 ,作为对均匀盐分下植物生理的一个重要补充,为扩大盐碱地的利用和栽培管理措施提供参考依据。【方法】采用分根装置将紫花苜蓿（ Medicago sativa ）的根系分为两部分,设置无盐胁迫（0/0,各半根系所处环境NaCl浓度均为0）、半边根系环境NaCl浓度为0的不均匀盐胁迫系列（0/S：0/100、0/150、0/200和0/250）、半边根系NaCl浓度为50 m mol·L -1 的不均匀盐胁迫系列（ 50/S：50/100、50/150、50/200和50/250）以及两半根系环境NaCl浓度相同的均一盐胁迫系列（S/S：50/50、100/100、150/150、200/200和250/250）处理(“/”两边数值代表各半根系所处环境的盐胁迫状况,以NaCl浓度mmol ·L -1 表示 )。处理15 d后测定其植株生长速率、生物量、水分吸收、钠、钾离子浓度、丙二醛含量等指标,了解不同盐胁迫环境对紫花苜蓿生长的影响。【结果】盐胁迫抑制紫花苜蓿生长,植株生长速率、生物量、水分吸收下降,叶绿素含量降低,脯氨酸含量升高,膜质过氧化程度加大,叶片Na + 浓度升高,K + 浓度降低,表现为较低的K + /Na + 。而不均匀盐胁迫0/S与50/S处理与均匀高盐处理S/S相比,植株生长速率和地上生物量分别增加了57.05%—369.34%和15.47%—42.57%,Na + 浓度降低了15.85%—55.93%,缓解了Na + 的毒害作用。且不均匀盐胁迫下70%—92%的水分吸收来自于无盐或低盐胁迫一侧根系,增加了整株植物的水分吸收。【结论】不均匀盐胁迫处理与均匀的高盐胁迫处理相比,增加了紫花苜蓿叶绿素含量,降低了膜质过氧化程度,通过调控Na + 与K + 的吸收维持叶片中相对较高的K + /Na + ,并且无盐和低盐胁迫一侧根系表现出补偿性吸水和补偿性生长,进而促进了植物生长,增加了地上和地下生物量。在一定的浓度范围内,与均匀的高盐胁迫处理相比,不均匀盐胁迫环境下植株根系所处环境的盐分浓度差异越大其对盐害的缓解作用越显著。

... 05),说明盐处理对盐生草幼苗生长有明显抑制作用,熊雪等^[27]研究发现紫花苜蓿(Medicago sativa)受到盐胁迫后发生渗透使植株生长受到抑制,生物量显著降低 ...

2018

0.0

... 徐宁等^[28]研究了NaCl胁迫对高粱(Sorghum bicolor)种子萌发和幼苗生长特性的影响,得到盐胁迫下幼苗根系活力随盐浓度的升高呈下降的趋势 ...

2019

0.0

... 这与杨智等^[29]研究得出干旱胁迫能使垂穗披碱草(Elymus nutans)的生物量和根系活力显著降低(P#cod#x0003C ...

2017

0.0

... 主成分分析是将原来各项指标重新组合成一组新的彼此独立的几个综合指标,用少数的综合指标来反映原来多指标的信息^[30] ...

2013

0.0

. 2013, 46(9):1762-1771 DOI:10.3864/j.issn.0578-1752.2013.09.003

Screening and identification of sorghum bicolor cultivars for alkali tolerance during germination

高粱品种萌发期耐碱性筛选与综合鉴定

Li F X , Zhou Y F , Wang Y T

李丰先, 周宇飞, 王艺陶

College of Agronomy, Shenyang Agricultural University, Shenyang 110866

【Objective】The objective of the current study was to explore the response of 35 different sorghum cultivars to alkali stress, and screen them for suitability for growing in alkaline soil and provide some guidance for their management under alkali stress.【Method】Mixed alkali were used (NaHCO3﹕Na2CO3 with the mole ratio of 9﹕1) to simulate the typical alkali stress environment. The 35 sorghum cultivars were treated with 25 mmol•L-1 solution or distilled water and incubated in Petri dishes for 10 days at humidity of 60%, light/dark of 12 h/12 h, illumination of 34 μmol•m-2•s-1 and 28℃/25℃ of day/night during germination. The germination force was measured on the fourth day and the germination percentage, leaf length, root length, leaf fresh weight, root fresh weight, leaf dry weight, root dry weight and remaining seed dry weight were measured on the tenth day. Through the fuzzy mathematics method of membership function, principal component analysis (PCA) and clustering analysis (CA), the alkali resistance of different sorghum cultivars were evaluated.【Result】The PCA results showed that root length, germination index and leaf dry weight were the most significant factors and are recommended as the main indexes for identifing alkali tolerance of sorghum at germination. The cultivars were ranked based on the membership function, and three groups on alkali tolerance were sorted from the 35 sorghum cultivars according to the CA. The analysis showed that seven of the 35 cultivars were strongly tolerant of alkaline conditions, e. g. Siza25, that twenty-two were had medium tolerance, e. g. Jiza 319 and Chiza 28, and that six were sensitive, e. g. Longza 9.【Conclusion】Principal component analysis showed that root length, germination index and leaf dry weight were the most significant factors and these are recommended as the main indexes to identify alkali tolerance of sorghum at germination, and in plant breeding and evaluation projects should be supported by CA and membership function to provide rigorous assessment of cultivars tolerance of alkalinity.

【目的】根据不同高粱品种萌发期对碱胁迫的响应，筛选出适合在盐渍土壤上种植的高粱品种，从而为减轻混合碱胁迫提供调控依据。【方法】采用人工气候箱内培养皿培养，以混合碱（NaHCO3﹕Na2CO3的摩尔比为9﹕1）模拟典型碱胁迫环境，在萌发期以25 mmol•L-1溶液处理35个高粱品种，蒸馏水处理为对照，每培养皿放30粒种子。人工气候箱内湿度为60%，光照/黑暗时间为12h/12h，光照强度为134 μmol•m-2•s-1，昼/夜温度为28℃/25℃。培养第4天测定发芽势，第10天测定发芽率、叶长、根长、叶鲜重、根鲜重、叶干重、根干重、子粒残余干重等性状，通过模糊数学隶属函数法和主成分分析法对各品种进行耐碱性综合评定，并进行聚类分析。【结果】主成分分析结果表明，根长、发芽指数和叶干重分别在根部、萌发因子和叶部中的负荷量最大，可作为萌发期高粱耐碱性筛选的主要鉴定指标。通过模糊数学隶属函数法可以对不同高粱品种进行耐碱性排序，不同高粱品种间耐碱性表现出明显的差异，最后通过聚类分析把35个高粱品种按耐碱性强弱分为三大类，其中四杂25号等7个品种为耐碱品种，吉杂319、赤杂28等22个品种为中等耐碱品种，龙杂9号等6个品种为碱敏感品种。【结论】根长、发芽指数、叶干重等指标可用于大量高粱品种萌发期耐碱性的筛选，隶属函数和聚类分析也可为高粱品种耐碱性在植物育种和鉴定方面提供依据。

... 李丰先等^[31]运用此方法得出发芽率和干重可作为萌发期高粱耐碱性的主要评价指标 ...

2015

0.0

... 杨帆等^[32]得出根长、芽鲜重、发芽率可作为萌发期高粱耐盐性的主要评价指标 ...

2016

0.0

... 蔺军^[33]得出相对发芽指数、发芽率可作为萌发期燕麦(Avena sativa)耐盐性的主要评价指标 ...