干旱胁迫下梭梭水力性状调整与非结构性碳水化合物动态

doi:10.17521/cjpe.2022.0276

植物生态学报 ›› 2023, Vol. 47 ›› Issue (10): 1407-1421.DOI: 10.17521/cjpe.2022.0276

干旱胁迫下梭梭水力性状调整与非结构性碳水化合物动态

陈图强, 徐贵青(), 刘深思, 李彦

中国科学院新疆生态与地理研究所, 荒漠与绿洲生态国家重点实验室, 乌鲁木齐 830011; 中国科学院阜康荒漠生态国家野外科学观测研究站, 新疆阜康 831505; 中国科学院大学, 北京 100049

收稿日期:2022-07-04 接受日期:2023-03-13 出版日期:2023-10-20 发布日期:2023-11-23
通讯作者: * (xugq@ms.xjb.ac.cn)
基金资助:
国家自然科学基金(32171874);新疆天山青年计划优秀青年科技人才项目(2020Q025)

Hydraulic traits adjustments and nonstructural carbohydrate dynamics of Haloxylon ammodendron under drought stress

CHEN Tu-Qiang, XU Gui-Qing(), LIU Shen-Si, LI Yan

State Key Laboratory of Desert and Oasis Ecology, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Ürümqi 830011, China;National Field Scientific Observation and Research Station of Desert Ecology, Chinese Academy of Sciences, Fukang, Xinjiang 831505, China; and University of Chinese Academy of Sciences, Beijing 100049, China

Received:2022-07-04 Accepted:2023-03-13 Online:2023-10-20 Published:2023-11-23
Contact: * (xugq@ms.xjb.ac.cn)
Supported by:
National Natural Science Foundation of China(32171874);Xinjiang Uygur Autonomous Region Tianshan Youth Program Project(2020Q025)

摘要/Abstract

摘要：

梭梭(Haloxylon ammodendron)是古尔班通古特沙漠的主要建群种, 在生物多样性保护和防止旱地退化等生态系统服务方面有重要作用。气候变化引起的频发干旱对梭梭生存有显著的影响, 明晰干旱胁迫下梭梭的抗旱策略, 对于荒漠生态系统的可持续发展至关重要。水力性状和碳收益作为抗旱机制中的重要部分, 目前对干旱胁迫下梭梭生存的水力性状阈值尚不明确。该研究以成年梭梭为对象, 分别设置对照组和干旱处理组, 对梭梭上、中、下3个高度的同化枝水分状况、枝条木质部导度损失率、气体交换特征、非结构性碳水化合物含量和形态特征等进行了测定, 利用单因素方差分析检验不同处理及枝条高度间的各项性状差异, 结合线性回归了解梭梭气孔敏感性, 通过主成分分析解析梭梭的抗旱策略。研究表明: (1)梭梭的黎明和正午同化枝水势、同化枝含水量和枝条含水量均因干旱胁迫而下降, 但并未随高度增加而降低; P50和P88 (最大导水度损失50%和88%的木质部水势)未因干旱胁迫和枝条高度的增加显著变化, 两个处理下3个枝条高度的P50平均值为-4.12 MPa, P88为-7.10 MPa, 而水力安全边界在干旱胁迫下显著降低; (2)梭梭的气孔行为对水分亏缺敏感性低, 干旱胁迫和枝条高度增加总体上未对其净光合速率和气孔导度产生显著影响; (3)同化枝和枝条非结构性碳水化合物含量未因干旱胁迫和枝条高度的增加而降低, 反而略有升高, 干旱胁迫下同化枝和枝条非结构性碳水化合物含量相较对照组分别升高22.11%和13.10%; (4)梭梭在干旱胁迫下的胡伯尔值相较对照组升高73.78%; 比叶面积相较对照组降低14.60%, 但两者均与对照组无显著差异。总之, 梭梭的水力性状受干旱胁迫影响显著, 但不受枝条高度的影响, 并不存在随枝条高度增加的水力限制; 干旱胁迫下, 梭梭树冠外缘枝条同时发生水力失效的风险较大, 水力安全边界(正午同化枝水势与P88的差值)只有对照组的40.85%; 但由于梭梭气孔对水分亏缺的低敏感性, 这使得其光合固碳并未受到影响, 反而同化枝和枝条的非结构性碳水化合物含量会有所升高。

关键词: 干旱胁迫, 抗旱策略, 水力性状, 非结构性碳水化合物, 树木死亡, 梭梭

Abstract:

Aims Haloxylon ammodendron is the major dominated species in the Gurbantünggüt Desert, which plays a key role in ecosystem services: such as biodiversity conservation and prevention of dryland degradation. Frequent droughts have a significant impact on the survival of H. ammodendron, thus understanding the drought resistant strategies of H. ammodendron is essential for the sustainability and stability of desert ecosystems. Robust hydraulic system and carbon balance are important parts of the drought resistance mechanism, but the hydraulic threshold for survival of H. ammodendron under drought stress are still unquantified.

Methods We set up a control group and a drought treatment group for adult H. ammodendron, and determined the water status of assimilation twigs, the loss rate of xylem hydraulic conductivity in branches, gas exchange characteristics, nonstructural carbohydrate (NSC) contents and morphological characteristics at upper, lower and middle branches of H. ammodendron. We used one-way ANOVA for each trait among different treatments and heights, linear regression for stomatal sensitivity and principal component analysis for drought resistance of H. ammodendron, respectively.

Important findings (1) The predawn and midday water potential of assimilation twig, assimilation twig water content and branch water content of H. ammodendron decreased under drought stress, but did not affected by the increase of height; P50 and P88 (xylem tension causing 50% and 88% loss of maximum hydraulic conductivity) did not change significantly under drought and with increasing height, and the mean value of P50 was -4.12 MPa and P88 was -7.10 MPa for each height and treatment groups, while the hydraulic safety margin was significantly reduced under drought. (2) The stomatal opening of H. ammodendron was not sensitive to drought stress, and thus drought stress and branch height increase did not significantly affect net photosynthetic rate and stomatal conductance in general. (3) The NSC contents of assimilation twigs and branches did not decrease under drought stress or with increasing branch height; the value of NSC contents in the assimilation twigs and branches were 22.11% and 13.10% higher, compared to the control group. (4) The Huber value of H. ammodendron increased by 73.78% in the drought treatment group compared to the control group; the specific leaf area decreased by 14.60% compared to the control group, but there were no significant difference between the two treatment groups. In conclusion, the hydraulic traits of H. ammodendron were significantly affected by drought stress, but not by the increase of branch height, and there was no hydraulic limitation with increasing branch height. Under drought stress, the risk of simultaneous hydraulic failure of the peripheral branches at the crown edge was high, the hydraulic safety margin (difference between midday assimilation twig water potential and P88) was only 40.85% of that of the control group. Due to the low sensitivity of stomata to water stress, the shrub can maintain the capacity of photosynthetic carbon fixation under drought stress, and even slightly increased NSC contents of the assimilation twigs and branches.

Key words: drought stress, drought resistant strategy, hydraulic trait, nonstructural carbohydrate, tree mortality, Haloxylon ammodendron

陈图强, 徐贵青, 刘深思, 李彦. 干旱胁迫下梭梭水力性状调整与非结构性碳水化合物动态. 植物生态学报, 2023, 47(10): 1407-1421. DOI: 10.17521/cjpe.2022.0276

CHEN Tu-Qiang, XU Gui-Qing, LIU Shen-Si, LI Yan. Hydraulic traits adjustments and nonstructural carbohydrate dynamics of Haloxylon ammodendron under drought stress. Chinese Journal of Plant Ecology, 2023, 47(10): 1407-1421. DOI: 10.17521/cjpe.2022.0276

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0276

https://www.plant-ecology.com/CN/Y2023/V47/I10/1407

图/表 12

图1 梭梭样地不同土层深度的土壤含水量(平均值±标准误)。不同大写字母表示同一处理不同土层深度存在显著差异(p < 0.05), 不同小写字母表示同一土层深度不同处理间差异显著(p < 0.05)。

Fig. 1 Soil water content at different depths in Haloxylon ammodendron sampling plot (mean ± SE). Different uppercase letters indicate significant differences among soil depths of the same treatment (p < 0.05), and different lowercase letters indicate significant differences between treatments of the same soil depth (p < 0.05).

图2 梭梭木质部导度损失率曲线。垂直的实线分别表示最大导水度损失50%和88%时木质部受到的压力(即木质部的水势) (P50、P88); 虚线和阴影为95%的置信区间。

Fig. 2 Percent loss of xylem conductivity for Haloxylon ammodendron. The vertical solid lines indicate the pressure on the xylem (i.e. the water potential of the xylem) for 50% and 88% loss of maximum hydraulic conductivity (P50, P88), respectively; the dashed lines and shaded areas are 95% confidence intervals.

表1 对照组和干旱处理组梭梭上、中和下部枝条最大导水度损失50%和88%的木质部水势(P50和P88) (平均值±标准误)

Table 1 Xylem water potential for 50% and 88% loss of maximum hydraulic conductivity (P50, P88) in upper, middle and lower branches of control and drought-treated Haloxylon ammodendron (mean ± SE)

	P50 (MPa)			P88 (MPa)
	下部 Bottom	中部 Middle	上部 Upper	下部 Bottom	中部 Middle	上部 Upper
对照组 Control	-4.35 ± 0.10^Aa	-4.13 ± 0.33^Aa	-3.98 ± 0.17^Aa	-7.67 ± 0.26^Aa	-7.02 ± 0.56^Aa	-7.10 ± 0.07^Aa
干旱组 Drought	-4.15 ± 0.17^Aa	-3.82 ± 0.16^Aa	-4.27 ± 0.14^Aa	-7.03 ± 0.21^Aa	-6.88 ± 0.24^Aa	-6.86 ± 0.09^Aa

图3 对照组和干旱处理组梭梭不同高度枝条的黎明同化枝水势、正午同化枝水势、同化枝含水量和枝条含水量(平均值±标准误)。P50和P88分别为最大导水度损失50%和88%的木质部水势。不同大写字母表示同一处理不同高度水平间存在显著差异(p < 0.05), 不同小写字母表示同一高度水平不同处理间差异显著(p < 0.05)。

Fig. 3 Predawn assimilation twig water potential, midday assimilation twig water potential, twig water content and branch water content among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE). P50 and P88 are the xylem water potentials for 50% and 88% loss of maximum hydraulic conductivity, respectively. Different uppercase letters indicate significant differences among different heights of the same treatment (p < 0.05) and different lowercase letters indicate significant differences between different treatments of the same height (p < 0.05).

图4 对照组和干旱处理组梭梭不同高度枝条的水力安全边界(平均值±标准误)。HSM50和HSM88分别为正午同化枝水势与最大导水度损失50%和88%的木质部水势之差。不同大写字母表示同一处理不同高度水平间存在显著差异(p < 0.05), 不同小写字母表示同一高度水平不同处理间差异显著(p < 0.05)。

Fig. 4 Hydraulic safety margin among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE). HSM50 and HSM88 are the difference between the midday assimilation twig water potential and the xylem water potential for 50% and 88% loss of maximum hydraulic conductivity, respectively. Different uppercase letters indicate significant differences among different heights of the same treatment (p < 0.05) and different lowercase letters indicate significant differences between different treatments of the same height (p < 0.05).

图5 对照组和干旱处理组梭梭不同高度枝条的净光合速率和气孔导度(平均值±标准误)。不同大写字母表示同一处理不同高度水平间存在显著差异(p < 0.05), 不同小写字母表示同一高度水平不同处理间差异显著(p < 0.05)。

Fig. 5 Net photosynthetic rate and stomatal conductance among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE). Different uppercase letters indicate significant differences among different heights of the same treatment (p < 0.05) and different lowercase letters indicate significant differences between different treatments of the same height (p < 0.05).

图6 梭梭气孔导度对饱和水汽压差的响应。阴影为95%的置信区间。

Fig. 6 Response of Haloxylon ammodendron stomatal conductance to vapor pressure deficit. The shaded area are 95% confidence intervals.

图7 对照组和干旱处理组梭梭不同高度枝条的同化枝和枝条的非结构性碳水化合物(NSC)含量(平均值±标准误)。不同大写字母表示同一处理不同高度水平间存在显著差异(p < 0.05), 不同小写字母表示同一高度水平不同处理间差异显著(p < 0.05)。

Fig. 7 Nonstructural carbohydrate (NSC) content of assimilation twigs and branches among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE). Different uppercase letters indicate significant differences among different heights of the same treatment (p < 0.05) and different lowercase letters indicate significant differences between different treatments of the same height (p < 0.05).

表2 对照组和干旱处理组梭梭不同高度枝条的同化枝和枝条的可溶性糖和淀粉含量(平均值±标准误)

Table 2 Soluble sugar and starch contents of assimilation twigs and branches among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE)

处理 Treatment	高度 Height	同化枝 Assiassimilation twig		枝条 Branch
处理 Treatment	高度 Height	可溶性糖 Soluble sugar (%)	淀粉 Starch (%)	可溶性糖 Soluble sugar (%)	淀粉 Starch (%)
对照 Control	下部 Bottom	2.48 ± 0.45^Ab	3.61 ± 0.62^Aa	2.62 ± 0.70^Aa	1.41 ± 0.17^Aa
	中部 Middle	2.57 ± 0.14^Ab	3.13 ± 0.30^Aa	2.21 ± 0.39^Ab	1.75 ± 0.28^Aa
	上部 Upper	2.83 ± 0.24^Ab	3.57 ± 0.59^Aa	2.22 ± 0.20^Ab	1.71 ± 0.29^Aa
干旱 Drought	下部 Bottom	3.81 ± 0.37^Aa	3.72 ± 0.56^Aa	3.01 ± 0.27^Aa	1.56 ± 0.07^Aa
	中部 Middle	3.48 ± 0.58^Aa	3.12 ± 0.59^Aa	3.01 ± 0.14^Aa	1.34 ± 0.33^Aa
	上部 Upper	4.43 ± 0.49^Aa	3.65 ± 0.71^Aa	3.03 ± 0.19^Aa	1.53 ± 0.13^Aa

图8 干旱处理组梭梭枝条木质部导度损失率与枝条非结构性碳水化合物(NSC)含量之间的关系。

Fig. 8 Relationship between branch xylem hydraulic conductivity loss rate and nonstructural carbohydrate (NSC) content of Haloxylon ammodendron in drought treatment group.

图9 对照组和干旱处理组梭梭不同高度枝条的胡伯尔值和比叶面积(平均值±标准误)。不同大写字母表示同一处理不同树高水平间存在显著差异(p < 0.05), 不同小写字母表示同一树高水平不同处理间差异显著(p < 0.05)。

Fig. 9 Huber value and specific leaf area among different heights of Haloxylon ammodendron in control and drought treatment groups (mean ± SE). Different uppercase letters indicate significant differences among different heights of the same treatment (p < 0.05) and different lowercase letters indicate significant differences between different treatments of the same height (p < 0.05).

图10 梭梭抗旱策略的主成分(PC)分析。ns, 无显著差异; ***, p < 0.001。Ψmd, 正午同化枝水势; Ψpd, 黎明同化枝水势; B.NSC, 枝条非结构性碳水化合物含量; BWC, 枝条含水量; gs, 气孔导度; HSM50, Ψmd - P50; HSM88, Ψmd - P88; HV, 胡伯尔值; L.NSC, 同化枝非结构性碳水化合物含量; LWC, 同化枝含水量; P50, 导水度损失50%时的木质部水势; P88, 导水度损失88%时的木质部水势; Pn, 净光合速率; SLA, 比叶面积。

Fig. 10 Principal component (PC) analysis of each trait of drought resistance strategies for Haloxylon ammodendron. ns, no significant difference; ***, p < 0.001. Ψmd, midday assimilation twig water potential; Ψpd, predawn assimilation twig water potential; B.NSC, nonstructural carbohydrate content of branch; BWC, branch water content; gs, stomatal conductance; HSM50, Ψmd - P50; HSM88, Ψmd - P88; HV, Huber value; L.NSC, nonstructural carbohydrate content of assimilation twigs; LWC, assimilation twig water content; P50, xylem water potentials for 50% loss of maximum hydraulic conductivity; P88, xylem water potentials for 88% loss of maximum hydraulic conductivity; Pn, net photosynthetic rate; SLA, specific leaf area.

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干旱胁迫下梭梭水力性状调整与非结构性碳水化合物动态

Hydraulic traits adjustments and nonstructural carbohydrate dynamics of Haloxylon ammodendron under drought stress

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