不同品种高粱幼苗在干旱复水过程中的生理生态响应

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刘婷婷, 陈道钳, 王仕稳, 殷俐娜, 邓西平. 不同品种高粱幼苗在干旱复水过程中的生理生态响应. 草业学报, 2018,27(6): 100-110
LIU Ting-ting, CHEN Dao-qian, WANG Shi-wen, YIN Li-na, DENG Xi-ping. Physio-ecological responses to drought and subsequent re-watering in sorghum seedlings. Acta Prataculturae Sinica,2018,27(6): 100-110 复制到剪切板

Doi:10.11686/cyxb2017410
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不同品种高粱幼苗在干旱复水过程中的生理生态响应

刘婷婷^1,², 陈道钳³, 王仕稳^1,^3,^*, 殷俐娜^1,³, 邓西平^1,³

1.中国科学院水利部水土保持研究所黄土高原土壤侵蚀与旱地农业国家重点实验室,陕西杨陵 712100

2.中国科学院大学,北京100049

3.西北农林科技大学水土保持研究所,陕西杨陵 712100

*通信作者Corresponding author. E-mail: shiwenwang@nwsuaf.edu.cn

作者简介:刘婷婷(1990-),女,河南郑州人,在读硕士。E-mail:liutingtingUCAS@126.com

收稿日期: 2017-10-09

基金项目:国家科技支撑计划(2015BAD22B01)和国家重点基础研究发展计划(973)项目(2015CB150402)资助

摘要

干旱适应能力不仅包括干旱胁迫期间的抗旱能力,也包括水分胁迫解除之后的恢复能力。以8个高粱品种为材料,采用盆栽称重控水法对高粱幼苗进行干旱及复水处理,通过测定生物量、相对含水量、水势、渗透势、光合参数等生理指标的变化分析不同品种高粱抗旱能力和旱后复水恢复能力的大小及其与干旱适应性的关系,并筛选出能有效指示干旱适应能力的生理指标。结果显示,干旱胁迫条件下,“辽杂21”生物量与对照相比无显著差异,抗旱能力较强;“甘蔗籽”和“Moench.cv.Gadambalia”高粱生物量分别下降38%、34%,抗旱能力较差。各品种高粱的相对含水量、水势、渗透势、叶绿素含量、净光合速率、蒸腾速率、气孔导度等均有所下降,复水后,这些生理指标恢复到对照水平,各品种开始恢复生长。由各生理指标与抗旱能力、旱后复水恢复能力和干旱适应能力之间的相关分析结果表明,高粱的干旱适应性与复水恢复性呈显著正相关( r=0.85),与抗旱性的相关系数是0.46,复水恢复能力在高粱干旱适应性中起着更重要的作用。高粱的抗旱性和旱后复水恢复性具有不同的生理基础,在干旱胁迫期间维持较高的净光合速率、较低的蒸腾速率有助于提高抗旱性,维持较高叶片相对含水量有助于提高旱后复水的恢复能力,叶片相对含水量、净光合速率可以作为筛选高粱干旱适应性的生理指标。

关键词: 高粱; 干旱; 干旱适应性; 抗旱性; 复水恢复性; 干旱复水

Physio-ecological responses to drought and subsequent re-watering in sorghum seedlings

LIU Ting-ting^1,², CHEN Dao-qian³, WANG Shi-wen^1,^3,^*, YIN Li-na^1,³, DENG Xi-ping^1,³

1.Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resources, Yangling 712100, China

2.University of Chinese Academy of Sciences, Beijing 100049, China

3.Institute of Soil and Water Conservation, Northwest A&F University, Yangling 712100, China

Abstract

Adaption to drought in plants includes drought resistance during the drought period and post drought recovery. The aim of this study was to investigate the relationships between drought resistance and drought recovery and to determine the physiological parameters which could be used to indicate drought adaptability. Eight sorghum cultivars grown in pots were exposed to drought and re-watering treatments during the seedling stage. Biomass, relative water content, water potential, osmotic potential, and photosynthetic parameters were measured. The results showed that “LZ21”“ had the highest drought resistance; biomass production under drought stress was not different to the control (well watered), while “GZZ”“ and “Gadambalia”“ had poor drought resistance producing 38% and 34% less biomass under drought stress. Relative water content, water potential, osmotic potential, chlorophyll content and photosynthetic parameters decreased after imposition of drought but all parameters recovered fully after re-watering. Correlation analysis showed that drought adaption is more closely related to drought recovery ( r=0.85) than drought resistance ( r=0.46). Maintaining higher photosynthetic and lower transpiration rate contributed to drought resistance while maintaining high relative water content improves drought recovery. These results demonstrated that drought recovery ability is more important for drought adaption than drought resistance. The physiological basis of drought resistance and drought recovery are clearly different. Relative water content and photosynthetic rate could be used to select drought adapted cultivars in sorghum.

Keyword: sorghum; drought; drought adaptation; drought resistance; drought recovery; re-watering

文章图片

植物在长期适应干旱环境过程中, 形成了4种适应机制:逃旱性(drought escape)、御旱性(drought avoidance)、耐旱性(drought tolerance)和复水恢复性(drought recovery)^[1], 其中, 御旱性和耐旱性被称为抗旱性(drought resistance)^[2]。抗旱性是指植物遭受水分胁迫后仍能保持正常生长、开花以及获得产量的能力。复水恢复性指植物经历一段时间干旱在水分条件改善之后恢复正常生长的能力^{[2, 3]}。在实际作物生产中, 作物往往处于干湿交替环境中, 因此, 作物在干旱环境下最终的产量不仅仅受其在遭受干旱胁迫过程中对干旱的抵御和耐受能力影响, 还应包括水分条件改善之后作物从先前损伤当中重新恢复正常生长的能力。因此, 复水恢复能力是作物适应干旱环境的一个重要内容, 也是作物适应干旱胁迫的一个重要机制。例如, 当作物苗期遭受干旱时, 复水恢复能力强的禾本科作物可以通过复水后增加分蘖来保证最后的产量^{[1, 4]}。也有研究表明, 作物不同品种在干旱复水之后的恢复能力并不相同, 认为筛选抗旱性强、复水后恢复速度快的品种对于实际生产具有重要的意义^{[5, 6]}。为了更好地衡量作物对干旱胁迫的反应, Chen等^[7]提出了干旱适应性(drought adaptation)概念, 即将作物在干旱和复水过程中的整体表现称为干旱适应性, 其既包括了抗旱能力又包括了复水后的恢复能力^[8], 并通过对多个玉米(Zea mays)品种的干旱适应性研究表明干旱适应性受恢复能力的影响更大。

高粱(Sorghum bicolor)是一种公认的综合抗旱性较强的作物, 不仅体现在由御旱、耐旱、避旱等多种机制形成的较强抗旱能力^[9], 而且体现在受旱的高粱在复水后, 能够得到不低于水分充足时的产量, 甚至产量超过充分供水水平, 即表现出旱后复水很好的恢复能力^{[10, 11]}。前人对高粱抗旱性的评价研究较多, 同时对高粱抗旱性与相关生理指标, 如叶片水分状况、渗透调节能力、抗氧化能力等之间的关系做不少研究^{[12, 13]}, 而对高粱的复水恢复能力及其和相关生理指标的关系关注较少。同时, 是否可以采用干旱适应性定义来衡量高粱在干旱复水过程中的整体响应也缺少相关的研究。因此, 本实验以8个不同高粱品种为材料, 研究高粱苗期干旱适应性与其抗旱能力和复水后的恢复能力的关系, 并进一步探索其干旱适应性、抗旱性、旱后复水恢复能力及其相关生理基础。通过完成本研究, 以探索不同高粱品种其干旱适应性和抗旱能力及恢复能力的关系, 以及影响干旱适应性, 抗旱能力和恢复能力的生理机制。研究结果将为旱区作物品种选择和作物的抗旱育种提供借鉴。

1 材料与方法

1.1 试验材料

从前期21个高粱品种栽种实验中筛选出8个生育期相对一致且抗旱性和复水恢复性差异较大的高粱品种为试验材料, 供试品种分别为“ Moench. cv. Gadambalia(M.G)” 、 “ 千斤红(QJH)” 、“ 遭烘萼(ZHE)” 、“ 饭高粱(FGL)” 、“ 高八尺(GBC)” 、“ 甘芝甜(GZT)” 、“ 甘蔗籽(GZZ)” 、“ 辽杂21(LZ21)” 。

盆栽试验于2013年5-7月在中国科学院水利部水土保持研究所干旱大厅人工气候室内进行, 试验所用塑料盆规格为直径30 cm, 高28 cm, 每桶装入14 kg黄绵土, 土壤田间持水量为20%, 测得土壤含水量为8.97%。高粱种子用1%次氯酸钠消毒20 min, 之后用蒸馏水冲洗, 选取大小一致的种子点播于塑料盆中。塑料盆置于移动遮雨棚下, 降水时遮上, 除了降水, 其他生长条件均与大田相似。当高粱幼苗第3片叶完全展开时, 定苗为2株每盆。

在高粱第8片叶完全展开时, 开始进行干旱-复水处理。实验设置2个水分处理:对照组充分供水(W)和处理组(自然干旱8 d后复水, D)。采用称重控水法于每天下午5点称重测定土壤含水量, 并将同一处理补水至相同含水量值。实验对照组的土壤含水量维持在田间持水量的85%~90%; 处理组土壤经自然干旱, 第8天时土壤含水量下降至田间持水量的35%, 处理第9天复水至对照水平。分别在处理前、处理第8天、复水后第5天采样, 采样时间为每天上午8点到9点, 采样部位为高粱第8片叶。实验过程中对照组(W)和处理组(D)的土壤含水量变化如图1所示。

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	图1 对照组(W)和处理组(D)的土壤含水量变化Fig.1 The changes of soil water content during drought stress and re-watering process

1.2 测定项目及方法

1.2.1 生物量分别在干旱处理前, 干旱处理8 d后, 复水5 d后采样, 将植株从地表处剪断装入纸袋, 80 ℃烘至恒重, 每个处理分别测定10株。

1.2.2 渗透势取第8片叶0.5 g装入0.5 mL离心管中, 液氮冷冻30 min, 室温解冻, 将离心管底部扎眼, 放入1.5 mL离心管中, 4 ℃、4000 r· min^-1离心5 min, 收集上清液, 用美国Wescor露点渗透压仪5600进行测定^[14]。

渗透势计算公式为:Φ =icRT/10⁶

式中:Φ 为渗透势; R为常数, R=8.314; ic为渗透势仪读数(mmol· kg^-1); T为绝对温度, T=273+室温(℃)。

1.2.3 水势于上午6:00采取高粱第8片叶, 用3005型压力室(SEC公司, USA)测定叶水势, 每个处理测定5株。

1.2.4 叶片相对含水量参考《植物生理学实验指导》方法^[15], 从高粱植株上剪去待测定叶片, 擦净, 称重并记录(FW), 然后将叶片浸入蒸馏水中6 h后称重并记录(TW), 之后将叶片置于80 ℃烘至恒重, 称重记录(DW)。

叶片相对含水量=(FW-DW)/(TW-DW)× 100%

1.2.5 叶绿素含量采用80%丙酮浸提法测定叶绿素含量, 每个处理5次重复^[15]。

1.2.6 光合参数采用美国LI-COR公司生产的Li-6400便携式光合仪, 每天上午测定高粱第8片叶的净光合速率(P_n), 蒸腾速率(T_r)、气孔导度(Cond)等参数。

1.2.7 抗旱能力、恢复能力及干旱适应性计算在干旱第8天、复水第5天每个处理选取10株在80 ℃烘至恒重后, 称重(DW), 高粱的抗旱能力、旱后恢复能力以及干旱适应能力计算参考Chen等^[7]。

植株抗旱能力= $\frac{处理组第 8 天干重 - 处理组第 0 天干重}{对照组第 8 天干重 - 对照组第 0 天干重}$

植株旱后恢复能力= $\frac{处理组复水 5 天后的干重 - 处理组第 8 天干重}{对照组第 14 天干重 - 对照组第 8 天干重}$

干旱适应能力= $\frac{处理组第 14 天干重 - 处理组第 0 天干重}{对照组第 14 天干重 - 对照组第 0 天干重}$

1.3 数据统计分析

利用SPSS 18.0处理分析数据, 最小显著差异法(LSD)进行方差分析, 显著性水平设为P< 0.05。采用Excel作图, Heatmap Illustrator 1.0做聚类热图。其中, 主成分分析、Pearson相关分析均采用各指标的相对值进行分析作图。聚类热图数据分析时利用各指标的相对值的对数值(以10为底)进行作图。

2 结果与分析

2.1 干旱及复水对高粱生物量的影响

如图2所示, 水分胁迫显著抑制了高粱的生长发育, 减缓了高粱生长速率, 但不同品种受抑制程度不同。经过8 d的干旱处理, 除了“ 辽杂21(LZ21)” 的地上部生物量与充分供水对照无显著性差异外, 其他品种的地上部生物量均显著降低, 其中, “ 甘蔗籽(GZZ)” 下降程度最严重, 达38%, 其次是“ Moench.cv.Gadambalia(M.G)” 高粱下降了34%。复水5 d后, 所有品种均恢复生长, 表现为生物量累积增加, 但此时的地上部生物量仍低于其对照, 且复水后不同高粱品种的生长速率不同, 表现为高八尺(GBC)> Moench.cv.Gadambalia(M.G)> 甘蔗籽(GZZ)> 饭高粱(FGL)> 遭烘萼(ZHE)> 千斤红(QJH)> 辽杂21(LZ21)> 甘芝甜(GZT), 其中 “ 高八尺(GBC)” 高粱的生物量累积超过100%, 说明其恢复速率最快, 旱后复水的恢复能力较强, “ 甘芝甜(GZT)” 恢复速率最慢, 为9.5%。说明不同的高粱品种具有不同的复水恢复能力。

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图2 干旱及复水对不同品种高粱地上部生物量的影响
Asterisks indicate statistically significant differences between control and treatment (P≤ 0.05).“ * ” 表示处理和对照之间差异显著(P≤ 0.05)。“ * ” means statistically significant differences between treatments (P≤ 0.05).下同。The same below.Fig.2 The biomass accumulation during drought stress and re-watering

2.2 干旱及复水对高粱叶片水分状况的影响

从图3可看出, 干旱胁迫条件下, 各高粱品种幼苗的叶片相对含水量较对照处理低, “ 千斤红(QJH)” 和“ 饭高粱(FGL)” 的叶片相对含水量分别下降28%和25%, 为所有品种中下降程度最大, “ Moench.cv.Gadambalia(M.G)” 和“ 甘蔗籽(GZZ)” 叶片相对含水量下降程度最小, 分别为5%和4%。复水5 d后, 所有品种的叶片相对含水量均恢复到对照水平。说明干旱胁迫造成叶片发生失水, 失水程度越大, 则该品种维持叶片水分状况的能力就越弱; 复水可迅速改善叶片失水状况。

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	图3 干旱及复水对不同品种高粱相对含水量和水势的影响Fig.3 Changes in relative water content and water potential during drought stress and re-watering

与叶片相对含水量变化相似, 干旱胁迫下, 所有品种高粱的叶水势均显著下降, 其中“ 饭高粱(FGL)” 叶水势下降程度最大, 其次是“ 遭烘萼(ZHE)” , “ Moench.cv.Gadambalia (M.G)” 下降程度最低, 而复水后所有品种的叶水势均恢复到充分供水对照水平。说明不同的高粱品种在干旱胁迫条件下的保水能力呈现出差异性, “ Moench.cv.Gadambalia (M.G)” 高粱干旱期间维持叶片较好水分状况的能力较其他品种更强, “ 饭高粱(FGL)” 维持叶片水分状况的能力最差, 但复水后所有品种高粱叶片水分状况均能恢复到正常水平。

2.3 干旱及复水对不同高粱品种渗透势的影响

渗透调节是植物应对干旱胁迫的一种主要调节方式。如图4所示, 干旱胁迫使所有高粱品种的渗透势均显著下降。“ 饭高粱(FGL)” 、“ 千斤红(QJH)” 分别下降144.2%、126.0%, 下降幅度较大, “ 甘芝甜(GZT)” 、“ Moench.cv.Gadambalia(M.G)” 下降幅度最小, 分别为44.5%和46.7%。复水后, 不同高粱品种的渗透势变化表现出显著差异。除“ Moench.cv.Gadambalia(M.G)” 高粱的渗透势未完全恢复到对照水平外, 其他品种的渗透势均完全恢复, 甚至显著高于对照水平。说明干旱胁迫期间不同的高粱品种具有显著不同的渗透调节能力, “ 饭高粱(FGL)” 、“ 千斤红(QJH)” 的渗透调节能力优于其他品种, 且复水后的恢复能力也较强, 而“ Moench.cv.Gadambalia(M.G)” 高粱不仅渗透调节能力较弱, 且在复水后的恢复程度也较其他品种慢。

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	图4 干旱及复水对不同品种高粱渗透势的影响Fig.4 Changes in osmotic potential during drought stress and re-watering

2.4 干旱及复水对不同品种高粱细胞膨压的影响

膨压的维持是细胞正常生长的基础, 干旱条件下较高的膨压维持能力有助于维持较好的光合作用, 于作物生长有利。由图5可知, 不同高粱品种细胞膨压对干旱胁迫的响应并不相同。“ Moench.cv.Gadambalia(M.G)” 、“ 甘芝甜(GZT)” 、“ 甘蔗籽(GZZ)” 、“ 辽杂21(LZ21)” 在干旱条件下的膨压较对照高, 具有更强的膨压维持能力, 而“ 遭烘萼(ZHE)” 、“ 饭高粱(FGL)” 、“ 千斤红(QJH)” 和“ 高八尺(GBC)” 的细胞膨压较对照有所降低, 说明干旱胁迫使得这些高粱品种叶片维持细胞膨压的能力丧失, 抗旱性较差; 复水后, 除遭烘萼(ZHE)之外, 所有品种的细胞膨压均恢复到对照水平。

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	图5 干旱及复水对不同品种高粱膨压的影响Fig.5 Changes in turgor pressure during drought stress and re-watering

2.5 干旱及复水对不同株系高粱叶片叶绿素含量的影响

如图6所示, 受干旱胁迫的影响, 所有高粱品种的叶绿素含量均较对照显著降低, 但降低程度不同, 说明干旱胁迫造成各高粱品种叶片内叶绿素不同程度的降解。其中, 降低程度最大和最小的品种分别为“ Moench.cv.Gadambalia(M.G)” 和“ 千斤红(QJH)” , 分别下降了46.4%和13.9%; 复水处理使得各高粱品种的叶片叶绿素含量开始有所恢复, 复水5 d后, 各品种的叶绿素含量已经恢复到充分供水对照水平, 甚至高于对照含量。

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	图6 干旱及复水对不同品种高粱叶绿素含量的影响Fig.6 Changes in chlorophyll content of 8 sorghum lines during drought stress and re-watering

2.6 干旱及复水对不同株系高粱叶片光合参数的影响

水分胁迫通常会降低气孔导度, 抑制作物的光合作用和蒸腾作用。在本实验中, 如图7所示, 胁迫后所有高粱的净光合速率、气孔导度、蒸腾速率均受到显著抑制, 尤其是“ Moench.cv.Gadambalia(M.G)” 、“ 甘芝甜(GZT)” 、“ 甘蔗籽(GZZ)” 、“ 辽杂21(LZ21)” 的净光合速率、气孔导度、蒸腾速率几乎降低为0, 说明这4个品种的光合作用受抑制程度最严重, 抗旱性较差。复水后, 不同品种高粱的净光合速率、蒸腾速率和气孔导度都有所恢复, 但恢复速度呈现品种差异。“ Moench.cv.Gadambalia(M.G)” 、“ 甘芝甜(GZT)” 的净光合速率、蒸腾速率先恢复至对照水平, 其他品种则恢复较慢。就气孔导度而言, 复水后所有高粱品种均未恢复到对照水平。

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	图7 干旱及复水对不同株系高粱净光合速率、蒸腾速率和气孔导度的影响Fig.7 Changes in photosynthesis rate, stomatal conductance, and transpiration rate during drought stress and re-watering

2.7 干旱时期和复水之后高粱各生理指标的主成分分析结果

如图8所示, 将高粱在干旱胁迫期间和复水恢复后的相对含水量、水势、渗透势、叶绿素、净光合速率、蒸腾速率、气孔导度、膨压的相对值做主成分分析, 分别得到3个主成分。干旱胁迫期间, 这3个主成分一共解释了92.41%的变量, 复水之后, 提取的这3个主成分解释了88.83%的变量。干旱期后第一个主成分PC1与相对含水量、水势相关, PC2与渗透势、净光合速率、蒸腾速率、气孔导度、膨压紧密相关, PC3与叶绿素含量紧密相关; 复水之后, PC1与渗透势、净光合速率、蒸腾速率、气孔导度紧密相关, PC2与相对含水量、水势紧密相关, PC3与叶绿素紧密相关。

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图8 干旱(A)及复水(B)时期各个主成分解释变量得分
柱状图代表每个主成分的特征值, 折线图代表每个主成分的累积变量。Fig.8 Scree plot of variance explained by each factor of the principal component during drought stress (A) and re-watering (B)
Each bar represents corresponding eigenvalue of each principle component; line charts represent cumulative variability change.

2.8 不同品种高粱的抗旱能力、旱后恢复能力和干旱适应性的相关分析

干旱适应能力不仅包括植物在干旱胁迫期间的抗旱能力, 也包括复水之后的恢复能力。由图9可看出, 不同品种的干旱适应能力为Moench.cv.Gadambalia(M.G)> 辽杂21(LZ21)> 高八尺(GBC)> 甘蔗籽(GZZ)> 千斤红(QJH)> 饭高粱(FGL)> 甘芝甜(GZT)> 遭烘萼(ZHE)。“ Moench.cv.Gadambalia(M.G)” 的旱后恢复能力最强, 抗旱能力最弱; “ 辽杂21(LZ21)” 的抗旱能力最强; “ 甘芝甜(GZT)” 的旱后恢复能力最弱。由抗旱能力、旱后复水的恢复能力和干旱适应能力三者之间的相关分析可得出, 旱后恢复能力和抗旱能力之间相关系数不显著, 说明彼此之间相互独立; 和抗旱能力相比, 旱后复水的恢复能力与干旱适应性的相关系数更高, 说明在植物适应干旱复水环境中, 旱后复水的恢复能力可能起着更重要的作用。

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图9 不同高粱品种的抗旱能力(A)、复水恢复能力(B)、干旱适应能力(C)的比较以及三者之间的相关分析Fig.9 Drought-adaptive capabilities and correlations between drought-adaptive capabilities
A: Drought resistance. B: Drought recovery. C: Drought adaptability. D: Correlation between drought resistance and drought recovery. E: Correlation between drought resistance and drought adaptability. F: Correlation between drought recovery and drought adaptability.

2.9 不同高粱品种在干旱胁迫后和复水恢复后各生理指标与抗旱能力、复水恢复能力以及干旱适应能力的关系

如图10所示, 取各生理指标相对值的对数值(以10为底)做热图总结了8个高粱品种对抗旱性、旱后复水恢复性的整体生理响应。利用双变量Pearson相关系数法对干旱胁迫时期和复水恢复时期高粱的8个生理指标的相对值分别与抗旱能力、旱后恢复能力、干旱适应能力进行相关性分析, 结果如表1所示。干旱胁迫条件下, 净光合速率与抗旱能力呈显著正相关, 蒸腾速率、气孔导度与抗旱能力呈负相关, 叶片相对含水量与旱后恢复能力、干旱适应能力呈正相关, 净光合速率、蒸腾速率、气孔导度、水势与干旱适应能力呈负相关。水分胁迫解除之后, 水势与抗旱性显著负相关, 渗透势与旱后恢复能力、干旱适应性呈正相关。

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	图10 不同品种高粱各生理指标响应干旱(A)及复水(B)的热图分析(图中数值采用相对值的对数值表示)Fig.10 Overall assessments of physiological responses in different sorghum lines during drought stress(A)and re-watering(B)

表1 干旱及复水时期各生理指标相对值与高粱抗旱能力、复水恢复能力、综合抗旱性之间的相关性 Table 1 Correlations between drought-adaptive capabilities and physiological responses

3 讨论

旱区作物的生长往往处在可变低水环境, 因此干旱环境下作物最终的产量不仅仅受其在遭受干旱胁迫过程中对干旱的抵御和耐受能力的影响, 还应包括水分条件改善之后作物从先前干旱损伤当中重新恢复正常生长的能力^[16]。除抗旱能力外, 近些年, 更多研究开始关注当胁迫解除之后作物从胁迫损伤中恢复正常生长的能力, 即作物的旱后复水恢复能力^[17]。因此, 衡量作物对于干旱的响应应该包括干旱和复水这两个过程。本研究采用Chen等^[7]提出的干旱适应性定义来研究不同高粱品种对干旱胁迫的响应, 结果表明, 不同品种的干旱适应能力、抗旱能力和旱后复水的恢复能力均存在较大差异; 干旱期间抗旱能力最强的高粱品种, 在复水之后恢复生长的能力不一定最强, 而干旱期间抗旱能力弱的高粱, 在复水之后恢复生长的能力不一定也较弱。相关分析表明, 高粱的干旱适应能力与旱后复水的恢复能力呈显著正相关(r=0.84), 而与抗旱能力相关系数为0.45。说明采用干旱适应能力能更好地反映高粱对干旱胁迫的响应, 而旱后复水的恢复能力可能在高粱对干旱适应能力中起着更重要的作用, 如本研究中, Moench.cv.Gadambalia高粱的旱后恢复能力最强, 抗旱能力较差, 但在8个品种中却有着最好的干旱适应能力。此研究结果与Chen等^[7]在玉米种的研究结果一致, 即玉米的干旱适应能力和抗旱能力、旱后复水恢复能力都有关, 而旱后复水的恢复能力可能扮演着更重要的作用。前人有研究表明, 作物的旱后复水恢复能力与抗旱性有关, 抗旱性强的品种, 同时复水后的恢复能力也较高^{[18, 19]}。但是在本研究中抗旱能力和旱后复水的恢复能力无相关性, 说明高粱的抗旱能力和复水后的恢复能力是彼此独立的, 都是高粱响应干旱胁迫的重要途径。

干旱胁迫除了能显著造成生长抑制和产量下降等形态变化外, 也会对植物的生理、代谢等过程造成显著地影响, 这些生理指标的变化, 一方面是反映干旱抑制植物生长的内在生理机制, 同时也反映对干旱及复水响应的内在机制^{[12, 13, 20, 21]}。因此, 探索干旱适应能力、抗旱能力及旱后复水的恢复能力和相关生理变化响应的关系, 一方面可以理解植物适应干旱的内在机制, 也可以为抗旱育种提供参考。本实验中, 干旱胁迫后各项生理指标都有所降低, 但降低幅度有所差异, 复水之后, 叶绿素含量、水势、渗透势恢复到甚至超过对照水平, 净光合速率、蒸腾速率、气孔导度有所恢复, 但大多数品种未恢复到对照水平。各项生理指标在干旱胁迫期间降低的程度, 以及复水后的恢复程度与品种有着很大关系。干旱期间维持较高的净光合速率和相对含水量有助于提高高粱的旱后恢复能力; 复水后的渗透势与旱后复水的恢复能力密切相关。而整体来看, 叶片相对含水量、渗透势等和干旱适应性密切相关。如“ Moench.cv.Gadambalia” 和“ 甘蔗籽” 高粱在干旱胁迫期间叶片相对含水量下降程度是最小的, 干旱适应能力最好。以上结果说明高粱在干旱胁迫期间抗旱性的生理基础与复水后恢复期间的复水恢复性的生理基础是不同的, 减轻高粱干旱胁迫期间的水分散失, 维持叶片水分状况, 有助于提高抗旱能力, 而维持在干旱胁迫下的良好水分状况有助于提高旱后复水的恢复能力。当水分胁迫缓解之后, 作物快速从干旱损伤中恢复的能力对于作物的生长发育有着积极的意义。因此, 复水恢复性在作物综合抗旱能力中的作用凸显的越来越重要^[22]。

4 结论

本研究表明高粱的干旱适应能力与抗旱能力和旱后复水的恢复能力都相关, 但旱后复水的恢复能力可能在高粱对干旱的适应能力中起着更重要的作用。而高粱在干旱胁迫期间的抗旱能力和复水后的恢复能力有着不同的生理基础, 因此, 在旱地作物品种选择和旱地育种中, 既要考虑其干旱胁迫下的抗旱能力, 又要考虑其旱后复水的恢复能力。本研究中, 由于是在苗期进行, 其结果主要适用于营养生长, 而和产量形成密切相关的生殖生长过程还缺乏相应的研究。但是在实际旱地作物生长中, 已被证明的干旱复水后产量的补偿和超补偿效应存在, 说明旱后复水的恢复能力及干旱适应性对旱地农业生产起重要作用。

The authors have declared that no competing interests exist.

参考文献

文献顺序

[1]	Fukai S, Cooper M. Development of drought-resistant cultivars using physio-morphological traits in rice. Field Crops Research, 1995, 40(2): 67-86. [本文引用:2]
[2]	Levitt J. Responses of plants to environmental stresses. Volume II. Water, radiation, salt, and other stresses. Academic Press, 1982, 51(15): 3642-3645. [本文引用:2]
[3]	Datta S K D, Malabuyoc J A, Aragon E L. A field screening technique for evaluating rice germplasm for drought tolerance during the vegetative stage. Field Crop Research, 1988, 19(2): 123-134. [本文引用:1]
[4]	Okami M, Kato Y, Kobayashi N, et al. Morphological traits associated with vegetative growth of rice during the recovery phase after early-season drought. European Journal of Agronomy, 2015, 64: 58-66. [本文引用:1]
[5]	Jin W, Yang L T, Pan Y, et al. Eco-physiological responses of different sugarcane varieties to drought and re-watering. Journal of Southern Agriculture, 2012, 43(12): 1945-1951. 金伟, 杨丽涛, 潘英, 等. 不同甘蔗品种对干旱和复水的生态生理响应. 南方农业学报, 2012, 43(12): 1945-1951. [本文引用:1]
[6]	Yan J Y, Zhang Y Q, Feng X M, et al. Effect of drought stress and rewatering on physiological characteristics of roots in different proso millet varieties. Acta Botanica Boreali-Occidentalia Sinica, 2012, 32(2): 348-354. 闫江艳, 张永清, 冯晓敏, 等. 干旱胁迫及复水对不同黍稷品种根系生理特性的影响. 西北植物学报, 2012, 32(2): 348-354. [本文引用:1]
[7]	Chen D, Wang S, Cao B, et al. Genotypic variation in growth and physiological responseto drought. Frontiers in Plant Science, 2015, 6: 1241. [本文引用:4]
[8]	Cao D, Chen D Q, Wu X, et al. Research on rewatering post-drought recovery capacity and physiological characteristics of different maize varieties. Acta Botanica Boreali-Occidentalia Sinica, 2015, 35(6): 1222-1228. 曹丹, 陈道钳, 吴茜, 等. 复水对旱后不同玉米品种植株生长恢复能力及其生理响应特性的影响. 西北植物学报, 2015, 35(6): 1222-1228. [本文引用:1]
[9]	Shan L, Xu B C. Discussion on drought resistance of sorghum and its status in agriculture in arid and semiarid regions. Scientia Agricultura Sinica, 2009: 2342-2348. 山仑, 徐炳成. 论高粱的抗旱性及在旱区农业中的地位. 中国农业科学, 2009: 2342-2348. [本文引用:1]
[10]	Su P, Shan L. *Physiological study on high grain yield and low water consumption of Sorghum* under highly varied and low water environment*. Chinese Journal of Applied & Environmental Biology, 1997, 3(4): 305-308. 苏佩, 山仑. 多变低水环境下高粱高产节水生理基础的研究*. 应用与环境生物学报, 1997, 3(4): 305-308. [本文引用:1]
[11]	Cao C L, Bai W B, Zhang J H, et al. Expression of sorghum production traits and re-watering compensation effect under water stress. Chinese Agriculture Science Bulletin, 2016, 32(33): 81-88. 曹昌林, 白文斌, 张建华, 等. 水分胁迫下高粱产量性状表达及复水后补偿效应的研究. 中国农学通报, 2016, 32(33): 81-88. [本文引用:1]
[12]	Zhou Y F, Wang D Q, Lu Z B, et al. Impacts of drought stress on leaf osmotic adjustment and chloroplast ultrastructure of stay-green sorghum. Chinese Journal of Applied Ecology, 2013, 24(9): 2545. 周宇飞, 王德权, 陆樟镳, 等. 干旱胁迫对持绿性高粱叶片渗透调节及叶绿体超微结构的影响. 应用生态学报, 2013, 24(9): 2545. [本文引用:2]
[13]	Lu F, Zhang F, Duan Y H, et al. Effects of drought stress on material production and physiological characteristics of sorghum at seedling stage. Acta Agronomica Sinica, 2015, (2): 149-153. 卢峰, 张飞, 段有厚, 等. 干旱胁迫对高粱苗期物质生产及生理特性的影响. 作物杂志, 2015, (2): 149-153. [本文引用:2]
[14]	Yin L, Wang S, Liu P, et al. *Silicon-mediated changes in polyamine and 1-aminocyclopropane-1-carboxylic acid are involved in silicon induced drought resistance in Sorghum bicolor* L*. Plant Physiology & Biochemistry*, 2014, 80: 268. [本文引用:1]
[15]	Gao J F. Plant physiological experiment instruction. Beijing: Higher Edutcation Press, 2006: 74-76. 高俊凤. 植物生理学实验指导. 北京: 高等教育出版社, 2006: 74-76. [本文引用:2]
[16]	Shan L, Su P, Guo L Q, et al. The response of different crops to drying wetting cycle in field. Acta Botanica Boreali-Occidentalia Sinica, 2000(2): 164-170. 山仑, 苏佩, 郭礼坤, 等. 不同类型作物对干湿交替环境的反应. 西北植物学报, 2000(2): 164-170. [本文引用:1]
[17]	Perrone I, Pagliarani C, Lovisolo C, et al. Recovery from water stress affects grape leaf petiole transcriptome. Planta, 2012, 235(6): 1383. [本文引用:1]
[18]	Guo Z F, Gong D Z, Hao W P, et al. *Study on compensation of Maize* varieties under drought stress and re-watering in different growth stages*. Journal of Maize Sciences, 2011, 19(4): 84-88. 郭子锋, 龚道枝, 郝卫平, 等. 不同生育期玉米干旱-复水补偿效应的品种差异研究*. 玉米科学, 2011, 19(4): 84-88. [本文引用:1]
[19]	Du J X, Hou X Y, Liu J R, et al. A study on physiological response to drought and re-watering treatments in Kentucky bluegrass. Acta Prataculturae Sinica, 2010, 19(2): 31-38. 杜建雄, 侯向阳, 刘金荣, 等. 草地早熟禾对干旱及旱后复水的生理响应研究. 草业学报, 2010, 19(2): 31-38. [本文引用:1]
[20]	Chen X R, Liu L Y, Zhong L, et al. Effects of rewatering after drought at heading on yield, δ⁽¹³)C and contents of endogenous hormone of leaf in double-season late hybrid rice. Journal of Nuclear Agricultural Sciences, 2013, 27(2): 240-246. 陈小荣, 刘灵燕, 钟蕾, 等. 抽穗后干旱复水对双季杂交晚稻产量形成和叶片δ⁽¹³)C及内源激素水平的影响. 核农学报, 2013, 27(2): 240-246. [本文引用:1]
[21]	Li G H, Wan Y S, Liu F Z, et al. Photosynthetic characteristics in different peanut cultivars under conditions of drought and re-watering at seedling stage. Chinese Journal of Plant Ecology, 2014, 38(7): 729-739. 厉广辉, 万勇善, 刘风珍, 等. 苗期干旱及复水条件下不同花生品种的光合特性. 植物生态学报, 2014, 38(7): 729-739. [本文引用:1]
[22]	Fang Y, Xiong L. General mechanisms of drought response and their application in drought resistance improvement in plants. Cellular & Molecular Life Science, 2015, 72(4): 673-689. [本文引用:1]

1995

0.0

... 植物在长期适应干旱环境过程中,形成了4种适应机制:逃旱性(drought escape)、御旱性(drought avoidance)、耐旱性(drought tolerance)和复水恢复性(drought recovery)^[1],其中,御旱性和耐旱性被称为抗旱性(drought resistance)^[2] ...

... 例如,当作物苗期遭受干旱时,复水恢复能力强的禾本科作物可以通过复水后增加分蘖来保证最后的产量^[1,4] ...

1982

0.0

... 复水恢复性指植物经历一段时间干旱在水分条件改善之后恢复正常生长的能力^[2,3] ...

1988

0.0

... 复水恢复性指植物经历一段时间干旱在水分条件改善之后恢复正常生长的能力^[2,3] ...

2015

0.0

... 例如,当作物苗期遭受干旱时,复水恢复能力强的禾本科作物可以通过复水后增加分蘖来保证最后的产量^[1,4] ...

2012

0.0

. 2012, 43(12):1945-1951 DOI:doi:10.3969/j:issn.2095-1191.2012.12.1945

Eco-physiological responses of different sugarcane varieties to drought and re-watering

不同甘蔗品种对干旱和复水的生态生理响应

Jin W , Yang L T , Pan Y

金伟, 杨丽涛, 潘英

【目的】研究干旱胁迫及复水对6个甘蔗品种的生态生理效应,为甘蔗抗旱育种和栽培提供参考依据。【方法】采用温室人工控水的方法,调查并比较干旱胁迫及复水期间6个甘蔗品种的农艺性状及生理生态指标的变化,以探讨不同甘蔗品种的耐旱性及复水后恢复能力。【结果】在干旱和复水条件下,不同品种的各个指标在不同时期的变化各不相同,且均在复水后得到恢复。6个甘蔗品种抗旱性由强到弱为:GT28ROC22GF98-296GT21B8ROC16;旱后恢复能力由强到弱排序为:GT28ROC22B8GF98-296GT21ROC16。【结论】绿叶数、细胞质膜透性以及可溶性糖、叶绿素、MDA含量可作为不同甘蔗品种抗旱性鉴定的重要指标。GT28、ROC22、GF98-296的抗旱性表现较好;新品种B8抗旱性表现一般,但是旱后丰产性能较好;ROC16抗旱能力及恢复能力最弱。

... 也有研究表明,作物不同品种在干旱复水之后的恢复能力并不相同,认为筛选抗旱性强、复水后恢复速度快的品种对于实际生产具有重要的意义^[5,6] ...

2012

0.0

. 2012, 32(2):348-354 DOI:doi:10.3969/j.issn.1000-4025.2012.02.021

Effect of drought stress and rewatering on physiological characteristics of roots in different proso millet varieties

干旱胁迫及复水对不同黍稷品种根系生理特性的影响

Yan J Y , Zhang Y Q , Feng X M

闫江艳, 张永清, 冯晓敏

With two different drought-resistant proso millet varieties(‘Longmi 4’ and ‘Jinshu 7’),the physiological effect were studied under the conditions of moderate and severe drought stress at seedling and rewatering at erect seedling stage in pot culture.The main result showed that:(1)under drought stress,the roots activity decreased,the activities of SOD and POD and contents of MDA and proline all increased.The effects were stronger under severe drought stress than under moderate drought stress.(2)After rewatering,the tested various physiological indexes of two different proso millet varieties had different degrees of recovery,and the recovering abilities were strong under moderate drought stress,but weak under severe stress.(3)The tested various physiological indexes of two different proso millet varieties had different amplitude under drought stress and rewatering condition.Under drought stress,the roots activity of drought-tolerant ‘Longmi 4’ was less than that of drought-sensitive ‘Jinshu 7’,but the ascending extent of the activities of SOD and POD and contents of MDA and proline were larger than those of drought-sensitive variety ‘Jinshu 7’.After rewatering,compared with ‘Jinshu 7’,‘Longmi 4’ had higher recovering ability.So the conclusions are that:under drought stress and after rewatering,‘Longmi 4’ keeps higher root activity,SOD activity,POD activity and proline contents,but lower MDA content,which are physiological basis of stronger drought resistance.

以2种抗旱性不同的黍稷品种（‘陇糜4号’和‘晋黍7号’）为试验材料,采用盆栽试验研究了苗期中度和重度干旱胁迫后拔节期复水对其根系生理特性的影响。结果显示：（1）干旱胁迫引起2个黍稷品种根系活力明显下降，根系SOD、POD活性以及MDA、脯氨酸含量明显升高，而且重度干旱胁迫处理变化幅度显著大于中度干旱胁迫。（2）复水后，2个黍稷品种根系的各项生理指标均有不同程度的恢复，且中度胁迫处理较易恢复，重度胁迫下恢复能力很弱。（3）2个黍稷品种根系各项生理指标在干旱胁迫及复水条件下变化幅度不同，干旱胁迫下抗旱性强的‘陇糜4号’根系活力下降幅度明显低于抗旱性弱的‘晋黍7号’，根系SOD活性、POD活性、MDA含量和脯氨酸含量的上升幅度明显高于‘晋黍7号’，而复水后‘陇糜4号’根系的各项生理指标的恢复能力明显强于‘晋黍7号’。研究表明，干旱胁迫及复水条件下‘陇糜4号’均表现出较高的根系活力、保护酶活性和脯氨酸含量，且MDA含量较低，从而表现出较强的抗旱性。

... 也有研究表明,作物不同品种在干旱复水之后的恢复能力并不相同,认为筛选抗旱性强、复水后恢复速度快的品种对于实际生产具有重要的意义^[5,6] ...

2015

0.0

... 为了更好地衡量作物对干旱胁迫的反应,Chen等^[7]提出了干旱适应性(drought adaptation)概念,即将作物在干旱和复水过程中的整体表现称为干旱适应性,其既包括了抗旱能力又包括了复水后的恢复能力^[8],并通过对多个玉米(Zea mays)品种的干旱适应性研究表明干旱适应性受恢复能力的影响更大 ...

... 7 抗旱能力、恢复能力及干旱适应性计算在干旱第8天、复水第5天每个处理选取10株在80 ℃烘至恒重后,称重(DW),高粱的抗旱能力、旱后恢复能力以及干旱适应能力计算参考Chen等^[7] ...

... 本研究采用Chen等^[7]提出的干旱适应性定义来研究不同高粱品种对干旱胁迫的响应,结果表明,不同品种的干旱适应能力、抗旱能力和旱后复水的恢复能力均存在较大差异 ...

... 此研究结果与Chen等^[7]在玉米种的研究结果一致,即玉米的干旱适应能力和抗旱能力、旱后复水恢复能力都有关,而旱后复水的恢复能力可能扮演着更重要的作用 ...

2015

0.0

. 2015, 35(6):1222-1228 DOI:doi:10.7606/j.issn.1000-4025.2015.06.1222

Research on rewatering post-drought recovery capacity and physiological characteristics of different maize varieties

复水对旱后不同玉米品种植株生长恢复能力及其生理响应特性的影响

Cao D , Chen D Q , Wu X

曹丹, 陈道钳, 吴茜

In this study,to explore the relationship of post-drought recovery capacity and physiological and biochemical response during drought and rewatering,we selected two maize varieties,‘Zhengdan 958’ and ‘P3’ with similar drought resistance but different resilience as experiment materials.The changes of growth,water status,photosynthetic parameters,chlorophyll fluorescence parameters and chlorophyll content were studied with potted experiment during drought and rewatering at seedling stage.The results indicated that:(1)the two maize varieties showed similar drought resistance,but significantly different growth recovery capacity.(2)The leaf water content of ‘Zhengdan 958’ and ‘P3’ demonstrated no significance during the drought period,but ‘P3’ maintained higher water potential,maximal efficiency of PSⅡ photochemistry and chlorophyll content under drought conditions.(3)The recovery rate of photosynthetic rate,maximal efficiency of PSⅡ photochemistry and stomatal conductance of ‘P3’ was quicker than that of ‘Zhengdan 958’ after drought and rehydration.Research showed that the photosynthetic loss recovery capacity of ‘P3’ was higher than that of ‘Zhengdan 958’.The reason why the recovery capability of ‘P3’ was stronger is because it can maintain higher F v / F m and chlorophyll content under drought,and the damage of photosynthetic systems was lighter,and also because the recovery was faster than ‘Zhengdan 958’ after rehydration.These results revealed that reducing damage of drought stress on plant photosynthetic systems is the basis of rapid recovery after rewatering,and the rapid of damaged photosynthetic systems to accelerate the recovery of plant after rewatering.

该研究选用2个抗旱能力相似但旱后恢复能力存在显著差异的玉米品种‘P3’和‘郑单958’为材料,采用盆栽称重控水法在苗期进行干旱及复水处理,通过测定其生长、水分状况、光合参数、叶绿素荧光参数以及叶绿素含量在干旱及复水过程中的变化规律,探讨干旱及复水过程中生理生化响应与旱后恢复能力的关系。结果发现:(1)抗旱性相同的2个玉米品种在干旱复水后的生长恢复能力表现为‘P3’显著强于‘郑单958’。(2)干旱胁迫后,‘郑单958’和‘P3’的叶片相对含水量差异不显著,但‘P3’能维持较高的叶水势、PSⅡ最大光化学效率和叶绿素含量。(3)经干旱胁迫复水后,‘P3’的净光合速率,PSⅡ最大光化学效率和气孔导度恢复速度快于‘郑单958’,说明‘P3’光合损失恢复能力高于‘郑单958’。研究表明,玉米品种‘P3’的旱后复水生长恢复能力较强,因‘P3’在干旱胁迫下能维持较高的 F v / F m 值和叶绿素含量,光系统的损伤较轻,而且复水后也能较快的恢复;在干旱过程中减轻干旱胁迫对植物光合系统的伤害是旱后复水快速恢复生长的基础,而在复水后快速修复光系统损失能够加快植物复水的恢复速度。

2009

0.0

. 2009, :2342-2348 DOI:doi:10.3864/j.issn.0578-1752.2009.07.011

Discussion on drought resistance of sorghum and its status in agriculture in arid and semiarid regions

论高粱的抗旱性及在旱区农业中的地位

Shan L , Xu B C

山仑, 徐炳成

本文以笔者研究组多年研究结果为基础,以玉米为主要参比作物,就干旱逆境下高粱的产量表现、水分利用及抗旱特性进行了论述。认为高粱是一种综合抗旱能力很强的作物,尤其具有低耗水、高水分利用效率特性,其耐旱性显著高于玉米,也高于谷子和苜蓿,属于一种典型的模式抗旱作物,具有重要的生产和研究价值。指出高粱作为抗逆性很强的粮饲酿兼用作物,特别是作为一种能源植物,仍具良好的发展前景。建议今后在降水量低于450mm、热量可满足生长的地区扩大高粱种植面积,同时加强对其整体抗旱性机理及抗旱基因组的研究。

... 高粱(Sorghum bicolor)是一种公认的综合抗旱性较强的作物,不仅体现在由御旱、耐旱、避旱等多种机制形成的较强抗旱能力^[9],而且体现在受旱的高粱在复水后,能够得到不低于水分充足时的产量,甚至产量超过充分供水水平,即表现出旱后复水很好的恢复能力^[10,11] ...

1997

0.0

. 1997, 3(4):305-308

Physiological study on high grain yield and low water consumption of Sorghum under highly varied and low water environment

多变低水环境下高粱高产节水生理基础的研究

Su P , Shan L

苏佩, 山仑

2016

0.0

. 2016, 32(33):81-88

Expression of sorghum production traits and re-watering compensation effect under water stress

水分胁迫下高粱产量性状表达及复水后补偿效应的研究

Cao C L , Bai W B , Zhang J H

曹昌林, 白文斌, 张建华

为实现高粱的调亏灌溉,达到节水但不减产的目的,有必要先了解和掌握高粱在不同生育阶段水分胁迫下产量性状的变化情况,笔者运用二因素四水平的随机区组设计法进行了研究。结果表明:在苗期长历时轻中度及短历时重度胁迫后复水补偿效应明显,节水效率可达50.7%,同CK比较,MMD15、MMD20和MSD10处理产量分别提高4.96%、5.98%、7.18%,差异达5%的显著性,MSD20处理的产量降低1.88%;在拔节期长历时轻中度及短历时重度胁迫后复水补偿效应存在,但不太明显,水分胁迫后的负效应大于补偿效应的正效应,节水效率达37.9%,同CK比较,BMD5~BSD5处理产量依次降低5.38%、7.44%、8.78%、5.4%;差异达5%的显著性,而BSD10和BSD15处理产量降低20.9%和22.45%,达1%的差异显著性;在灌浆期轻中度的胁迫处理,节水效率可达50.0%,千粒重差异不显著,重度胁迫处理,千粒重分别降低20.0%、20.8%和24.4%,其差异达1%显著,导致产量分别降低15.2%、15.7%和16.2%,重度胁迫处理与轻中度胁迫处理相比,千粒重也有1%的显著性。

2013

0.0

. 2013, 24(9):2545-

Impacts of drought stress on leaf osmotic adjustment and chloroplast ultrastructure of stay-green sorghum

干旱胁迫对持绿性高粱叶片渗透调节及叶绿体超微结构的影响

Zhou Y F , Wang D Q , Lu Z B

周宇飞, 王德权, 陆樟镳

Taking stay-green sorghum (B35) and non-stay green sorghum (Sanchisan) as test materials, a pot experiment was conducted to study their leaf osmotic adjustment and chloroplast ultrastructure at flowering and filling stages under impacts of drought stress (45%-50% of maximum field capacity). For the two sorghum lines, drought stress caused the reduction of their leaf free water content and relative water content, and increased the leaf bound water content, water saturation deficit, and electrical conductivity, with the increment or dement being larger for Sanchisan than for B35. Drought stress increased the leaf soluble sugar content and proline content, with the increment of the soluble sugar content being larger for Sanchisan and the increment of the proline content being larger for B35, while decreased the leaf soluble protein content, with the decrement being larger for Sanchisan than for B35. The chloroplast ultrastructure of both B35 and Sanchisan under drought stress was damaged to some extent, but the damaged degree was obviously lower for B35 than for Sanchisan. The findings indicated that stay-green sorghum had a greater adaptation to drought stress through stronger osmotic adjustment.

在盆栽条件下,研究了开花期和灌浆期干旱胁迫(土壤含水量为田间最大持水量的45%~50%)对持绿性高粱(B35)和非持绿性高粱(三尺三)叶片水分、渗透调节物质以及叶绿体超微结构的影响.结果表明: 干旱胁迫下，两高粱品系叶片自由水含量下降,束缚水含量增加,相对含水量降低,水分饱和亏缺增加,相对电导率增大,但三尺三各指标的变化幅度均大于B35.对于渗透调节物质,干旱胁迫下，三尺三可溶性糖含量的增幅大于B35,脯氨酸含量的增幅小于B35,可溶性蛋白含量的降幅大于B35.干旱胁迫下，B35与三尺三的叶绿体超微结构均受到一定程度的破坏,但B35叶绿体结构保持相对完好,受损程度明显小于三尺三.在干旱胁迫下，持绿性高粱通过较强的渗透调节表现出更好的干旱适应能力.

... 前人对高粱抗旱性的评价研究较多,同时对高粱抗旱性与相关生理指标,如叶片水分状况、渗透调节能力、抗氧化能力等之间的关系做不少研究^[12,13],而对高粱的复水恢复能力及其和相关生理指标的关系关注较少 ...

... 干旱胁迫除了能显著造成生长抑制和产量下降等形态变化外,也会对植物的生理、代谢等过程造成显著地影响,这些生理指标的变化,一方面是反映干旱抑制植物生长的内在生理机制,同时也反映对干旱及复水响应的内在机制^{[12,13,20,21]} ...

2015

0.0

. 2015, (2):149-153 DOI:doi:10.16035/j.issn.1001-7283.2015.02.029

Effects of drought stress on material production and physiological characteristics of sorghum at seedling stage

干旱胁迫对高粱苗期物质生产及生理特性的影响

Lu F , Zhang F , Duan Y H

卢峰, 张飞, 段有厚

试验于2013-2014年以高粱杂交种辽杂10、辽杂11和辽杂36为试材,对正常供水和干旱胁迫条件下的株高、叶面积、相对含水量、光合参数、地上部干重和保护性酶活性进行了比较与分析。结果表明,干旱胁迫条件下,高粱幼苗的出苗会有所延缓,叶片和根系的相对含水量均下降,叶片下降幅度大于根系;干旱胁迫导致净光合速率(Pn)下降,地上部物质积累减少,气孔导度(Gs)减小是Pn下降的主要原因;在干旱胁迫下,根系和叶片的过氧化物歧化酶(SOD)、过氧化物酶(POD)含量均显著高于正常供水处理,并且随着胁迫天数的增加呈抛物线状变化,在胁迫12d达到峰值。

2014

0.0

... min^-1离心5 min,收集上清液,用美国Wescor露点渗透压仪5600进行测定^[14] ...

2006

0.0

... 4 叶片相对含水量参考《植物生理学实验指导》方法^[15],从高粱植株上剪去待测定叶片,擦净,称重并记录(FW),然后将叶片浸入蒸馏水中6 h后称重并记录(TW),之后将叶片置于80 ℃烘至恒重,称重记录(DW) ...

... 5 叶绿素含量采用80%丙酮浸提法测定叶绿素含量,每个处理5次重复^[15] ...

0.0

. , 2000(2):164-170

The response of different crops to drying wetting cycle in field

不同类型作物对干湿交替环境的反应

Shan L , Su P , Guo L Q

山仑, 苏佩, 郭礼坤

... 3 讨论旱区作物的生长往往处在可变低水环境,因此干旱环境下作物最终的产量不仅仅受其在遭受干旱胁迫过程中对干旱的抵御和耐受能力的影响,还应包括水分条件改善之后作物从先前干旱损伤当中重新恢复正常生长的能力^[16] ...

2012

0.0

... 除抗旱能力外,近些年,更多研究开始关注当胁迫解除之后作物从胁迫损伤中恢复正常生长的能力,即作物的旱后复水恢复能力^[17] ...

2011

0.0

. 2011, 19(4):84-88

Study on compensation of Maize varieties under drought stress and re-watering in different growth stages

不同生育期玉米干旱-复水补偿效应的品种差异研究

Guo Z F , Gong D Z , Hao W P

郭子锋, 龚道枝, 郝卫平

以抗旱性不同的两个玉米品种郑单958和户单4号为材料,研究不同生育期干旱-复水对产量和水分利用效率的影响。结果表明,各生育期干旱-复水后,两个品种产量均低于对照处理,尤以扬花期干旱-复水最低;两个品种水分利用效率在扬花期干旱-复水最低;郑单958的产量、水分利用效率均高于户单4号,两者耗水量基本相等。分析两个品种株高、叶面积指数、干物质积累等在复水之后的变化表明,各生育期干旱-复水后,两个品种均存在一定的补偿效应,且郑单958的补偿效应明显较强。因此,郑单958在干旱-复水后补偿能力强,从而保持较高的产量和水分利用效率。

... 前人有研究表明,作物的旱后复水恢复能力与抗旱性有关,抗旱性强的品种,同时复水后的恢复能力也较高^[18,19] ...

2010

0.0

. 2010, 19(2):31-38

A study on physiological response to drought and re-watering treatments in Kentucky bluegrass

草地早熟禾对干旱及旱后复水的生理响应研究

Du J X , Hou X Y , Liu J R

杜建雄, 侯向阳, 刘金荣

... 前人有研究表明,作物的旱后复水恢复能力与抗旱性有关,抗旱性强的品种,同时复水后的恢复能力也较高^[18,19] ...

2013

0.0

. 2013, 27(2):240-246

Effects of rewatering after drought at heading on yield, #cod#x003b4;⁽¹³)C and contents of endogenous hormone of leaf in double-season late hybrid rice

抽穗后干旱复水对双季杂交晚稻产量形成和叶片#cod#x003b4; (13 )C及内源激素水平的影响

Chen X R , Liu L Y , Zhong L

陈小荣, 刘灵燕, 钟蕾

Key Laboratory of Crop Physiology, Ecology, Genetics and Breeding, Ministry of Education, Key Laboratory of Crop Physiology, Ecology and Genetic Breeding of Jiangxi Province, College of Agronomy, Jiangxi Agricultural University, Jiangxi, Nanchang 330045

In this study, super rice variety WufengyouT025 and the control variety Jinyou207 were selected to drought 8 days after heading, then rewatered to explore effect of yield and its formative physiological mechanism under the treatment of rewatering after drought during the late growth stage for the different yield potential double-season late hybrid rice varieties. The differences of the seed setting, yield, carbon isotopic compositions (δ 13 C) and endogenous hormone in the reciprocal second leaf during fore-and-aft drought of the two varieties were also analyzed. Results showed as follows: Compared to the CK (keeping water), seed-setting rate, 1000-grain weight and yield per plant of the two rice varieties presented diverse decline under the treatment of rewatering after drought stress. Among them, thereinto, seed setting rate of WufengyouT025 and Jinyou207 decreased by 12.07 % and 7.67 %, 1000-grain weight 5.23 %and 9.09 %, yield per plant 13.54 % and 27.14 %, respectively. All the differences were observed significantly and there was a lack of yield compensatory effect under the treatment of rewatering after 8-d drought at heading. Carbon isotopic compositions (δ 13 C) in the reciprocal second leaf under the treatment of drought at heading of WufengyouT025 greatly increased, decreased at first then began increasing at the 6d after rewatering, while that of Jinyou207 under the treatment of drought decreased greatly, increased at first then decreased, then increased, the distance between the treatment and the control of WufengyouT025 were higher than that of Jinyou207. Generally, under the condition of the CK from heading to 20 days, endogenous hormone in the reciprocal second leaf of the two varieties of ABA and GA 3 increased, IAA declined along with the growth course, while ZR increased at first then decreased. Under the treatment of drought of the two varieties, ABA increased, then decreased after rewatering, then increased, the distance between the treatment and the control of WufengyouT025 were higher than that of Jinyou207. Under the condition of the treatment of rewatering after drought, IAA of the two varieties all decreased, and the distance between the treatment and the control of WufengyouT025 was lower than that of Jinyou207, GA 3 of WufengyouT025 increased at first, then decreased after rewatering, then increased, while that of Jinyou207 decreased sharply and reached bottom at the end of the treatment of drought, and increased slowly after rewatering, ZR of the two varieties all increased and reached culmination, then decreased after rewatering, the distance between the treatment and the control of WufengyouT025 were lower than that of Jinyou207. The treatment of the drought at heading would promote the senescence resistant mechanism in the rice plant, and rewatering would activate the growth promoted factors and inhibited the growth restrained factors, but the effects were quite limited.

为探明生育后期干旱复水对双季杂交晚稻不同产量潜力品种产量的影响及其形成的生理机制,选择超级稻品种五丰优T025和对照品种金优207,于抽穗后进行干旱8d复水处理,分析了2个品种结实和产量、干旱前后倒二叶稳定碳同位素组成(δ 13 C)及内源激素含量的差异性。结果表明,干旱复水处理下2个品种水稻的结实率、千粒重及单株产量较对照(保持水层)表现出不同程度下降。其中,结实率五丰优T025和金优207分别下降12.07%和7.67%,千粒重下降5.23%和9.09%,单株产量下降13.54%和27.14%,差异均达显著或极显著水平,未发现产量补偿效应。抽穗后干旱处理下倒二叶δ 13 C值,五丰优T025大幅上升,复水后呈现出先下降,至第6天又开始上升的特点,金优207则大幅下降,复水后呈现先上升后下降再上升的特点,五丰优T025较金优207干旱复水处理与对照之间的差距更大。总体上,对照条件下,抽穗至其后20d,随生育推进,2个品种倒二叶内源ABA和GA 3 含量呈上升而IAA含量则下降趋势,ZR表现出先上升再下降特点。干旱处理下2个品种表现为ABA含量上升,复水后呈先下降后上升特点,五丰优T025较金优207处理与对照之间差距更大;干旱复水处理条件下:2个品种IAA含量均下降,五丰优T025处理与对照间的差距明显小于金优207;五丰优T025 GA 3 含量表现出先上升后下降再上升,金优207表现出干旱处理结束日急剧下降至最低点,复水后缓慢上升特点;2个品种ZR含量呈现出先上升至最高值再下降特点,五丰优T025较金优207处理与对照之间差距更小。抽穗后干旱处理将启动稻株体内抗衰老机制,复水后将在一定程度一定时间内激活稻株体内生长促进因子并抑制生长抑制因子,但其效果十分有限。

2014

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. 2014, 38(7):729-739 DOI:doi:10.3724/SP.J.1258.2014.00068

Photosynthetic characteristics in different peanut cultivars under conditions of drought and re-watering at seedling stage

苗期干旱及复水条件下不同花生品种的光合特性

Li G H , Wan Y S , Liu F Z

厉广辉, 万勇善, 刘风珍

Aims In China, peanut ( Arachis hypogaea ) is mainly cultivated in the semi-arid and rain-fed areas, and drought is the most prominent environmental stress to its growth. However, studies on the physiological responses of different peanut cultivars to drought and re-watering are lacking. Our objectives were to investigate the relationship between photosynthetic characteristics and drought tolerance, and to explore the ability to recover from drought damage in different peanut cultivars. Methods A pot experiment was conducted with artificial water stress treatment, and the photosynthetic characteristics were determined in twelve peanut cultivars under the conditions of drought stress and re-watering at the seedling stage. The drought tolerance was assessed by drought resistance coefficient of biomass in seedling. The recovery capacity was assessed by compensatory growth of plant. Important findings Five cultivars, including ‘Shanhua 11’, ‘Rugaoxiyangsheng’, ‘A596’, ‘Shanhua 9’, and ‘Nongda 818’, showed over-compensatory growth after re-watering, and their capacity of compensatory growth had significant positive correlation with drought tolerance ( p P n ), stomatal conductance ( G s ), intercellular CO 2 concentration ( C i ), maximum photochemical efficiency ( F v / F m ), PSII actual quantum yield (ΦPSII), and photochemical quenching coefficient ( q P ) all decreased over the course of drought stress, and then increased following re-watering, with the amplitude of changes being smaller in the more drought tolerant cultivars. Seven days of drought did not result in significant differences in the photosynthetic characteristics among majority of the peanut cultivars tested ( p > 0.05). After 14 days of drought, the values of photosynthetic variables differed significantly among the peanut cultivars with different drought tolerance ( p P n , G s , Φ PSII , F v / F m , and q P in the cultivars ‘Shanhua 11’, ‘Rugaoxiyangsheng’, ‘A596’, and ‘Shanhua 9’ fully recovered five days after re-watering, while those in the cultivars ‘79266’, ‘ICG6848’, ‘Baisha 1016’, and ‘Hua 17’ did not fully recover even after 10 days of re-watering; the values of those photosynthetic variables were significantly greater ( p P n , Φ PSII , F v / F m , and q P after 14 days of drought stress and after five days of re-watering, respectively ( p RWC ) for 14 days and after five days of re-watering at the seedling stage, the P n , Φ PSII , F v / F m , and q P could be used for identifying the level of damage and recovery capacity of peanut cultivars. The cultivar ‘Shanhua 11’ can be used as a reference for drought adaptability identification in peanut.

为探索不同花生( Arachis hypogaea )品种的旱后恢复能力, 研究花生品种耐旱性与光合特性的关系, 通过盆栽土壤水分控制实验, 测定了12个花生品种苗期对干旱胁迫与复水过程的光合响应特征, 并讨论了所测各性状参数与抗旱性强弱的关系, 包括对水分胁迫伤害的修复能力。结果表明, 根据苗期生物量抗旱系数, ‘山花11号’、‘如皋西洋生’、‘A596’、‘山花9号’、‘农大818’的抗旱性较强, 且复水后植株产生超补偿生长效应, 补偿生长能力与抗旱性呈极显著正相关。叶片净光合速率( P n )、气孔导度( G s )、胞间CO 2 浓度( C i )、最大光化学效率( F v / F m )、实际光化学效率( Φ PSII )、光化学猝灭系数( q P )随干旱进程逐渐降低, 复水后逐渐增加, 抗旱性强的花生品种变幅较小。干旱7天, 大多数花生品种的光合参数值未有显著性差异。干旱14天, 抗旱性越强的花生品种光合参数值越高, 不同抗旱性花生品种的光合参数值有显著差异。‘山花11号’、‘如皋西洋生’、‘A596’、‘山花9号’的 P n 、 G s 、 Φ PSII 、 F v / F m 、 q P 在复水5天时恢复至对照水平, 复水10天时超过对照, ‘79266’、‘ICG6848’、 ‘白沙1016’、‘花17’在复水10天时仍未达到对照水平, 复水过程中抗旱性强的品种的光合参数显著高于抗旱性弱的品种。相关分析表明, 干旱胁迫14天和复水5天后, 花生的 P n 、 Φ PSII 、 F v / F m 、 q P 与品种抗旱性呈极显著正相关。因此, 可在苗期用40%土壤相对含水量胁迫14天及复水5天时花生的 P n 、 Φ PSII 、 F v / F m 、 q P 鉴定品种的干旱伤害程度及修复能力, ‘山花11号’可作为强干旱适应性鉴定的标准品种。

2015

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... 因此,复水恢复性在作物综合抗旱能力中的作用凸显的越来越重要^[22] ...