八个树种叶水力性状对水分条件的响应及其驱动因素

doi:10.17521/cjpe.2021.0140

摘要/Abstract

摘要：

树木叶片的水力效率和安全性会对水分条件的改变做出一定的响应, 进而影响树木的生长和分布, 然而叶导水率(K_leaf)和叶水力脆弱性(P₅₀)对不同水分条件的响应模式及其影响因素尚不清楚。该研究选取了晋西北关帝山和黑茶山两种水分条件下的8种树种, 测量其水力性状、叶片导管和形态性状, 比较两地不同树种的K_leaf和P₅₀的变化, 分析叶片水力效率和安全性之间的权衡关系, 并探讨叶片水力性状在不同树种及水分条件下的响应模式及其驱动因素。结果表明: 对同一树种而言, 湿润的关帝山叶最大导水率(K_max)和P₅₀均高于干旱的黑茶山; 对同一地区而言, 从在高水分条件下生长的树种到在易干旱环境生长的树种, K_max和P₅₀均逐渐下降。K_max、P₅₀、膨压丧失点水势(TLP)之间均存在显著相关关系。两地叶片P₅₀与导管密度、导管塌陷预测值((t/b)³)、叶片厚度、比叶质量显著正相关, 与导管直径、叶面积显著负相关, 不同树种的K_leaf和P₅₀与叶导管性状的关系大于叶形态性状。同一树种的关帝山到黑茶山P₅₀变化量(δP₅₀)与比叶质量和叶干物质含量在两地的变化量显著正相关, 同一树种δP₅₀与叶形态性状变化量的关系大于与叶导管性状的。以上结果表明: 随着水分条件变差, 叶片水力效率降低, 水力安全性提高, 不同树种叶片水力效率与安全性之间存在一定的权衡关系, 不同树种叶水力性状的差别受叶导管性状影响的程度大于受叶形态性状的影响, 同一树种叶水力安全性对水分条件变化的响应主要依靠叶形态性状的驱动, 树木在提高自身叶水力安全的同时增加了叶构建的碳投资。

关键词: 水力性状, 导管性状, 形态性状, 叶导水率, 叶水力脆弱性

Abstract:

Aims The hydraulic efficiency and safety of tree leaves can respond to changes in water conditions, hence affecting the growth and distribution of trees. This study was conducted to determine the patterns of responses in leaf hydraulic conductivity (K_leaf) and leaf hydraulic vulnerability (P₅₀) in trees to varying water conditions and the influencing factors.

Methods In this study, eight tree species were selected at the study sites of Guandi Mountain and Heicha Mountain in northwestern Shanxi, and their hydraulic traits, leaf vessel and morphological traits were measured. Changes of K_leaf and P₅₀ in those eight tree species were compared between the two locations. The trade-off relationship between leaf hydraulic efficiency and safety was analyzed.

Important findings Within the same tree species, the maximum hydraulic conductivity (K_max) and P₅₀ were higher at the moist Guandi Mountain sites than at the dry Heicha Mountain sites; within the same study areas, K_max and P₅₀ were higher in tree species occurring under high water availability than those in drought-prone environment. There were significant correlations among K_max, P₅₀ and water potential at turgor loss points (TLP). Leaf P₅₀ in trees in the two study areas was significantly and positively correlated with the number of vessels per unit area, the predicted value of vessel collapse ((t/b)³), leaf thickness, and leaf mass per unit area, and negatively with vessel diameter and leaf area. K_leaf and P₅₀ in different tree species were better related with leaf vessel traits than with leaf morphological traits. The changes in P₅₀(δP₅₀) from Guandi Mountain to Heicha Mountain within the same tree species were significantly and positively correlated with changes in leaf mass per unit area and leaf dry mass content, and δP₅₀ was more closely related with the leaf morphological traits than with the leaf vessel traits within the same tree species. The above results indicate that, with deterioration of water conditions, leaf hydraulic efficiency decreases while the hydraulic safety increases. There is a certain trade-off between leaf hydraulic efficiency and safety across different tree species. The differences in leaf hydraulic traits among tree species are more affected by leaf vessel traits than leaf morphological traits. The responses of leaf hydraulic safety to changes in water conditions are mainly driven by leaf morphological traits. An improvement in leaf hydraulic safety occurs at the expenses of structural carbon investment.

Key words: hydraulic trait, vessel trait, morphological trait, leaf hydraulic conductivity, leaf hydraulic vulnerability

任金培, 李俊鹏, 王卫锋, 代永欣, 王林. 八个树种叶水力性状对水分条件的响应及其驱动因素. 植物生态学报, 2021, 45(9): 942-951. DOI: 10.17521/cjpe.2021.0140

REN Jin-Pei, LI Jun-Peng, WANG Wei-Feng, DAI Yong-Xin, WANG Lin. Responses of leaf hydraulic traits to water conditions in eight tree species and the driving factors. Chinese Journal of Plant Ecology, 2021, 45(9): 942-951. DOI: 10.17521/cjpe.2021.0140

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0140

https://www.plant-ecology.com/CN/Y2021/V45/I9/942

图/表 8

图1 关帝山和黑茶山8个树种的叶水力脆弱性曲线。A, 青杨。B, 蒙古栎。C, 榆树。D, 山杏。E, 山杨。F, 落叶松。G, 油松。H, 云杉。图中竖线为水分输导速率为最大导水率(Kmax)的50%时的水势(P50), G表示关帝山树种, H表示黑茶山树种。

Fig. 1 Leaf hydraulic vulnerability curves for eight tree species in Guandi Mountain and Heicha Mountain. A, Populus cathayana. B, Quercus mongolica. C, Ulmus pumila. D, Armeniaca sibirica. E, Populus davidiana. F, Larix gmelinii. G, Pinus tabuliformis. H, Picea asperata. The vertical lines in each graph indicate the water potential (P50) when the water transport rate is at 50% of the maximum hydraulic conductivity (Kmax). G, trees in Guandi Mountain; H, trees in Heicha Mountain.

图2 关帝山和黑茶山青杨、蒙古栎、榆树、山杏、山杨、落叶松、油松和云杉叶膨压丧失点水势(TLP)的比较(平均值±标准误)。G表示关帝山树种, H表示黑茶山树种。不同大写字母代表同一树种在不同地区的差异显著, 不同小写字母代表不同树种在同一地区的差异显著(p < 0.05)。

Fig. 2 Comparisons of water potential at turgor loss points (TLP) among Populus cathayana (Pc), Quercus mongolica (Qm), Ulmus pumila (Up), Armeniaca sibirica (As), Populus davidiana (Pd), Larix gmelinii (Lg), Pinus tabuliformis (Pt) and Picea asperata (Pa) in Guandi Mountain and Heicha Mountain (mean ± SE). G, trees in Guandi Mountain; H, trees in Heicha Mountain. Different uppercase letters indicate significant differences between the study areas within the same tree species; different lowercase letters indicate significant differences among tree species within the same area (p < 0.05).

图3 关帝山和黑茶山阔叶和针叶树种叶最大导水率(Kmax)、水力脆弱性(P50)和膨压丧失点水势(TLP)的比较(平均值±标准误)。G表示关帝山树种, H表示黑茶山树种。不同大写字母代表同一树种在不同地区的差异显著, 不同小写字母代表不同树种在同一地区的差异显著(p < 0.05)。

Fig. 3 Comparisons of maximum hydraulic conductivity (Kmax), hydraulic vulnerability (P50) and water potential at turgor loss points (TLP) between broadleaved and coniferous species and between Guandi Mountain and Heicha Mountain (mean ± SE). G, trees in Guandi Mountain; H, trees in Heicha Mountain. Different uppercase letters indicate significant differences between study areas within the same tree group; lowercase letters indicate significant differences between tree groups within the same study area (p < 0.05).

图4 关帝山和黑茶山树种叶最大导水率(Kmax)、水力脆弱性(P50)和膨压丧失点水势(TLP)的关系(平均值±标准误)。G表示关帝山树种, H表示黑茶山树种。**, p < 0.01。

Fig. 4 Relationships among maximum hydraulic conductivity (Kmax), hydraulic vulnerability (P50) and water potential at turgor loss points (TLP) in Guandi Mountain and Heicha Mountain (mean ± SE). G, trees in Guandi Mountain; H, trees in Heicha Mountain. **, p < 0.01.

表1 关帝山(G)和黑茶山(H)叶水力性状与叶导管形态性状的关系

Table 1 Relationships between leaf hydraulic traits and leaf vessel morphological traits in Guandi Mountain (G) and Heicha Mountain (H)

		N (No.·mm^-2)	D (μm)	(t/b)³	LT (μm)	LA (cm²)	LMA (g·m^-2)	LDMC (g·g^-1)
G	P₅₀	0.866^**	-0.945^**	0.714^*	0.775^*	-0.921^**	0.776^*	0.453
G	K_max	-0.749^*	0.895^**	-0.563	-0.617	0.813^*	-0.613	-0.392
H	P₅₀	0.927^**	-0.898^**	0.734^*	0.920^**	-0.810^*	0.915^**	0.711^*
H	K_max	-0.724^*	0.942^**	-0.597	-0.594	0.900^**	-0.597	-0.327

图5 关帝山和黑茶山两地间比叶质量变化量(δLMA)、叶干物质含量变化量(δLDMC)与水力脆弱性变化量(δP50)的关系(平均值±标准误)。*, p < 0.05。

Fig. 5 Relationships of the changes in leaf mass per unit area (δLMA) and the changes in leaf dry mass content (δLDMC) with changes in hydraulic vulnerability (δP50) between Guandi Mountain and Heicha Mountain (mean ± SE). *, p < 0.05.

表2 关帝山和黑茶山导管性状(VT)和形态性状(MT)对水力脆弱性(P50)和最大导水率(Kmax)的影响

Table 2 Level of significance in the effects of vessel traits (VT) and morphological traits (MT) on hydraulic vulnerability (P50) and maximum hydraulic conductivity (Kmax) in Guandi Mountain and Heicha Mountain

因子 Parameter	P₅₀			K_max
因子 Parameter	系数 Coefficient	t	p	系数 Coefficient	t	p
VT	0.680	3.875	0.002	-1.034	-3.003	0.010
MT	0.299	1.704	0.112	0.263	0.763	0.459

表3 关帝山和黑茶山两地间导管性状变化量(δVT)和形态性状变化量(δMT)对水力脆弱性变化量(δP50)和最大导水率变化量(δKmax)的影响

Table 3 Significance of vessel traits variation (δVT) and morphological traits variation (δMT) to hydraulic vulnerability variation (δP50) and maximum hydraulic conductivity variation (δKmax) from Guandi Mountain to Heicha Mountain

因子 Parameter	δP₅₀			δK_max
因子 Parameter	系数 Coefficient	t	p	系数 Coefficient	t	p
δVT	-0.981	-2.314	0.069	-0.650	-1.019	0.355
δMT	1.594	3.761	0.013	-0.069	-0.108	0.918

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