Changes in response of carbon and water physiological parameters of Robinia pseudoacacia seedlings to long-term drought and rehydration
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摘要:
目的 探究刺槐如何应对气候变化引起的干旱与恢复，了解其在干旱及复水期间水分状况变化、非结构性碳水化合物（NSC）平衡分配策略和生理生化响应机制，为揭示全面气候变化背景下刺槐林生产力衰退的生理学机制以及为刺槐林培育的水分管理提供理论参考。 方法 本研究采用自然干旱试验方法，设置正常供水和自然干旱处理，测定1年生刺槐苗在干旱导致全部落叶期间及复水后新叶长成时苗木的水分状况、压力−容积曲线参数、非结构性碳和抗氧化酶活性等生理指标，比较不同处理及干旱时期对刺槐生理参数的影响。 结果 刺槐处于轻度干旱时期，叶中淀粉的积累和细胞维持膨压能力增加，根和叶内的渗透调节和部分抗氧化防御机制也开始启动。中度干旱时期，叶的淀粉转化为可溶性糖增加，脯氨酸（Pro）含量显著增加，以改善渗透调节和应对干旱压力；同时根内抗坏血酸过氧化物酶（APX）、超氧化物岐化酶（SOD）、过氧化物酶（POD）活性均达到峰值，渗透调节及抗氧化防御机制全面启动。刺槐处于重度干旱时期时，根内淀粉、NSC含量均呈上升趋势，NSC由可溶性糖（SS）发挥渗透调节作用逐渐转为增加淀粉（Sta）的积累。复水后根和茎储存NSC显著降低。 结论 干旱胁迫可显著抑制刺槐苗木的生长，并对其水分运输、碳代谢及其他生理生化反应产生了显著影响。当干旱胁迫超过中度干旱时，刺槐幼苗的生理适应性降低。干旱胁迫的增强，可能导致碳水化合物的净损失，重度干旱时期，刺槐苗木将更多的NSC从叶中分配到根，NSC也由主要发挥渗透调节功能逐渐转为储存功能，以用作复水后水力传导的修复与重建。 Abstract:Objective The main purpose of this study was to explore how Robinia pseudoacacia coped with drought and rehydration caused by climate change, and to understand the changes of water physiology, non-structural carbon balanced allocation strategy and physiological and biochemical response mechanism during drought and rehydration. It helps to reveal the physiological mechanism of Robinia pseudoacacia forest decline and death under the background of overall climate change. Theoretical reference for cultivation irrigation of Robinia pseudoacacia forest was provided. Method In this study, the experimental method of natural drought was used, and the normal water supply was set and the water supply was stopped. Then, the physiological indexes of two years old Robinia pseudoacacia seedlings were measured, such as water status, pressure-volume curve parameters, non-structural carbon and antioxidant enzyme activities, and the effects of different treatments and period of drought on physiological parameters of Robinia pseudoacacia seedlings were compared. Result In the light drought period, the accumulation of starch and the ability of maintaining turgor pressure in cells of Robinia pseudoacacia leaves increased, and the osmotic regulation and antioxidant defense mechanism of roots and leaves were also started. During moderate drought period, the process of starch conversion to soluble sugar and Pro content increased significantly to improve osmotic regulation to cope with drought stress. At the same time, the activities of APX, SOD and POD reached peak values, and the osmotic regulation and antioxidant defense mechanism of root were fully activated. When Robinia pseudoacacia was in a severe drought, the content of starch and NSC in roots increased, the function of NSC gradually changed from the osmotic regulation of soluble sugar to the accumulation of starch. After rehydration, root and stem storage NSC decreased significantly. Conclusion The growth of Robinia pseudoacacia seedlings is significantly inhibited by drought stress, and its water transport, carbon metabolism and other physiological and biochemical reactions are significantly affected. When drought stress exceeds moderate drought, the physiological adaptability of Robinia pseudoacacia seedlings decreases. As drought stress intensifies, a net loss of carbohydrates may occur. During severe drought, more NSC seedlings will be distributed from leaves to roots, and NSC will gradually be used for osmotic regulation to storage, and may be used for restoration and reconstruction of hydraulic conduction after rehydration. -
图 1 干旱胁迫和复水期间刺槐叶片凌晨和正午水势的变化
不同大写字母表示同一时间各处理间在0.05水平上差异显著；不同小写字母表示同一处理的不同干旱时间在0.05水平上差异显著。下同。 Different capital letters indicate significant difference at 0.05 level among different drought stress treatments at the same time; different small letters indicate significant difference at 0.05 level among different drought times in the same treatment. The same below.
Figure 1. Changes of predawn and midday leaf water potential of Robinia pseudoacacia during different drought stress and rehydration periods
图 2 干旱胁迫和复水期间不同器官中非结构性碳水化合物含量的变化
Figure 2. Changes of non-structural carbohydrate content in different organs during drought stress and rehydration periods
图 3 干旱胁迫和复水期间不同器官中脯氨酸含量的变化
Figure 3. Changes of proline carbohydrate contents in different organs during drought stress and rehydration periods
图 4 干旱胁迫和复水期间不同器官中可溶性蛋白含量的变化
Figure 4. Changes of soluble protein contents in different organs during drought stress and rehydration periods
图 5 干旱胁迫和复水期间不同器官中MDA、ASA、APX含量的变化
Figure 5. Changes of MDA, ASA, APX contents in different organs during drought stress and rehydration periods
图 6 干旱胁迫和复水期间不同器官中抗氧化酶活性的变化
Figure 6. Changes of antioxidant enzymes activities in different organs during drought stress and rehydration periods
表 1 干旱胁迫和复水期间土壤水分状况变化
Table 1. Changes of soil-water content during different periods of drought stress and recovery
时期
Stage取样日期
Sampling dateSWC/% PSWC/% 水分梯度
Water gradient视觉特征
Visual symptomsS0 06−30 39.59 ± 0.95b 128 ± 3b 水分充足
Moisture Sufficiency叶片正常
The leaves of the plants are normally developedS15 07−15 20.13 ± 2.85c 65 ± 9c 轻度干旱
Light drought新叶萎蔫，下部叶片发黄
The new leaves begin to fade and their lower leaves turn yellowS25 07−25 16.08 ± 0.52cd 52 ± 2cd 中度干旱
Moderate drought下部叶片全凋落
The lower leaves fall off completelyS30 07−30 15.90 ± 0.66cd 51 ± 2cd 中度干旱
Moderate drought上部50%叶片萎蔫
50% of the upper leaves wither awayS35 08−04 12.14 ± 0.89de 39 ± 3de 重度干旱
Severe drought上部50%叶片凋落
50% of the upper leaves fall offS40 08−09 9.60 ± 0.58e 31 ± 2e 重度干旱
Severe drought上部70%叶片凋落
70% of the upper leaves fall offS60 08−29 9.49 ± 0.30e 31 ± 1e 重度干旱
Severe drought几乎全部落叶
Almost all fallen leavesR20 09−18 45.86 ± 4.50a 148 ± 15a 复水
Rehydration基部萌发新叶
New leaves germinating at the baseR40 10−08 36.86 ± 3.82b 119 ± 12b 复水
Rehydration上部新叶展开
Upper new leaf unfolding注：SWC代表土壤质量含水量，PSWC代表植物有效土壤含水量。S0、S15、S25、S30、S35、S40、S60分别代表停止水分处理的第0、15、25、30、35、40、60天，R20、R40代表复水后的第20和40天。不同小写字母表示不同干旱时间在0.05水平上差异显著（P < 0.05）。下同。Notes: SWC stands for soil water content, and PSWC stands for plant available soil water content. S0, S15, S25, S30, S35, S40, and S60 represent the 0,15, 25, 30, 35, 40, and 60 days after stopping watering, respectively, and R20 and R40 represent the 20 and 40 days after starting rewatering. Different small letters indicate significant difference at 0.05 level among different drought periods. The same below. 
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表 2 干旱胁迫和复水期间苗木形态变化
Table 2. Morphological changes of Robinia pseudoacacia seedlings during different drought stress and rehydration periods
时期 Stage 地径
Ground diameter/mm叶片数目 Leaf number CK 干旱胁迫 Drought stress CK 干旱胁迫 Drought stress S0 12.70 ± 0.41c 12.70 ± 0.41a 39 ± 4b 39 ± 4b S15 13.65 ± 0.19bc 13.84 ± 0.84a 54 ± 2ab 47 ± 6a S25 14.58 ± 0.39ab 14.40 ± 0.96a 60 ± 5ab 47 ± 4a S35 14.81 ± 0.24ab 14.42 ± 0.96a 62 ± 7a 22 ± 2c S40 14.89 ± 0.29ab 14.42 ± 0.97a 63 ± 7a 18 ± 5cd S60 15.28 ± 0.65a 14.46 ± 0.87a 67 ± 12a 1 ± 2f R20 15.87 ± 1.42a 14.46 ± 0.87a 70 ± 16a 5 ± 2ef R40 15.97 ± 0.44a 14.46 ± 1.00a 52 ± 14ab 11 ± 2de
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表 3 干旱胁迫和复水期间基于P-V曲线计算的水分参数
Table 3. Parameters of P-V curves during different drought stress and rehydration periods
时期 Stage WCinternal/% Rs/% Ra/% Ra/Rs RWCtlp/% ψtlp/MPa ψsat/MPa S0 305.0 ± 10.0b 289.3 ± 10.3a 15.7 ± 1.5b 0.055 ± 0.006b 34.2 ± 3.7bc −0.882 ± 0.075a −0.250 ± 0.013a S15 309.1 ± 11.7b 296.3 ± 13.4a 12.9 ± 3.2b 0.044 ± 0.012bc 58.2 ± 5.9a −1.285 ± 0.186c −0.750 ± 0.064c S25 315.8 ± 14.4b 308.8 ± 14.3a 7.1 ± 1.0c 0.023 ± 0.004d 32.1 ± 4.5c −1.696 ± 0.110d −0.494 ± 0.071b S35 256.3 ± 9.2c 249.6 ± 7.2b 6.7 ± 2.1c 0.027 ± 0.008cd 20.0 ± 0.5d −1.035 ± 0.021ab −0.185 ± 0.006a R25 352.9 ± 11.4a 289.8 ± 8.1a 63.1 ± 3.4a 0.217 ± 0.007a 42.7 ± 4.4b −1.207 ± 0.064bc −0.415 ± 0.035b 注：WCinternal代表叶片组织内部饱和含水量，Rs代表叶片共质体水含量，Ra代表质外体水含量，Ra/Rs代表叶片质外体含水量和共质体含水量的比值，ψtlp代表叶片膨压损失点的渗透势，RWCtlp代表膨压损失点的相对含水量，ψsat代表叶片的饱和渗透势。Notes: WCinternal represents the saturated water content in the leaf. Rs represents the symplast water content in the leaf. Ra represents the water content in the leaf ectoplasmic body. Ra/Rs represents the ratio of the water content in the leaf extoplasmic body to the water content in the symplast. ψtlp represents the osmotic potential at the turgor pressure loss point of the leaf. RWCtlp represents the relative water content at the turgor pressure loss point. ψsat represents the saturated permeation potential of the blade.
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