九种不同材性的温带树种叶水力性状及其权衡关系

doi:10.17521/cjpe.2016.0064

摘要/Abstract

摘要：

不同材性树种的解剖、叶脉分布等结构性状差异会影响树木的水分运输效率和水分利用策略, 进而限制树木的生存、生长和分布。然而, 材性对叶导水率、水力脆弱性及其潜在的权衡关系的影响尚不清楚。该研究选择东北温带森林中不同材性的9种树种(散孔材: 山杨(Populus davidiana)、紫椴(Tilia amurensis)、白桦(Betula platyphylla); 环孔材: 蒙古栎(Quercus mongolica)、水曲柳(Fraxinus mandshurica)、胡桃楸(Juglans mandshurica); 无孔材: 红皮云杉(Picea koraiensis)、樟子松(Pinus sylvestris var. mongolica)、红松(Pinus koraiensis), 测量其基于叶面积和叶质量的叶导水率(K_area和K_mass)、水力脆弱性(P₅₀)、膨压丧失点水势(TLP)及叶结构性状, 以比较不同材性树种叶水力性状的差异, 并探索叶水力效率与安全的权衡关系。结果表明: 3种材性树种的K_area、K_mass和P₅₀均差异显著(p < 0.05)。无孔材树种的K_area和K_mass最低, 而散孔材和环孔材树种差异不显著; 环孔材树种P₅₀最高, 而散孔材和无孔材树种差异不显著。K_area和K_mass均与P₅₀显著负相关(p < 0.05), 但散孔材、环孔材和无孔材树种的相关关系分别呈线性、幂函数和指数函数关系。这表明叶水力效率与安全之间存在一定的权衡关系, 但该关系受树木材性的影响。K_mass与TLP显著负相关(p < 0.01), 其中散孔材和环孔材树种呈线性负相关, 无孔材树种呈负指数函数关系; P₅₀随TLP的增加而增加, 这表明树木在面临水分胁迫时, 其质外体和共质体抗旱阻力共同协调保护叶片活细胞, 防止其水分状况到达临界阈值。K_mass与叶干物质含量、叶密度、比叶重均显著负相关, 而P₅₀与之显著正相关(p < 0.01, P₅₀与比叶重的关系除外), 表明树木叶水力特性的变化受相同叶结构特性驱动, 树木增加对水力失调的容忍需要在叶水力系统构建上增加碳投资。

关键词: 叶水力性状, 叶导水率, 叶水力脆弱性, 权衡关系, 散孔材, 环孔材, 无孔材

Abstract:

Aims Trees with different wood properties display variations in xylem anatomy and leaf vein structure, which may influence tree water transport efficiency and water-use strategy, and consequently constrain tree survival, growth and distribution. However, the effects of wood properties on leaf hydraulic conductance and vulnerability and their potential trade-offs at leaf level are not well understood. Our aims were to examine variations in leaf hydraulic traits of trees with different wood properties and explore potential trade-offs between leaf hydraulic efficiency and safety.
Methods Nine tree species with different wood properties were selected for measuring the leaf hydraulic traits, including three diffuse-porous species (Populus davidiana, Tilia amurensis, Betula platyphylla), three ring-porous species (Quercus mongolica, Fraxinus mandshurica, Juglans mandshurica), and three non-porous species (Picea koraiensis, Pinus sylvestris var. mongolica, Pinus koraiensis). Four dominant and healthy trees per species were randomly selected. The hydraulic traits measured included leaf hydraulic conductance on leaf area (K_area) and dry mass (K_mass) basis, leaf hydraulic vulnerability (P₅₀), and leaf water potential at turgor loss point (TLP), while the leaf structural traits were leaf dry mass content (LDMC), leaf density (LD) and leaf mass per unit area (LMA).
Important findings The K_area, K_mass, and P₅₀differed significantly among the tree species with different woody properties (p < 0.05). Both K_area and K_mass were the lowest for the non-porous trees, and did not differ significantly between the diffuse-porous and ring-porous trees. The ring-porous trees had the highest P₅₀values, while the diffuse-porous and non-porous trees showed no significant differences in P₅₀. Both K_area and K_mass were negatively correlated with P₅₀ (p < 0.05) for all the trees, and the relationships for the diffuse-porous, ring-porous, and non-porous trees were fitted into linear, power, exponential functions, respectively. This indicates that significant trade-offs exist between leaf hydraulic efficiency and safety. The K_mass was correlated (p < 0.01) with TLP in a negative linear function for the diffuse- and ring-porous trees and in a negative exponential function for the non-porous trees. The P₅₀ increased with increasing TLP. These results suggest that apoplastic and symplastic drought resistance are strictly coordinated in order to protect living cells from approaching their critical water status under water stresses. The K_mass was negatively correlated (p < 0.01) with LDMC, LD, or LMA, while the P₅₀ was positively correlated with LDMC and LD; this suggests that variations in K_mass and P₅₀ are driven by similar changes in structural traits regardless of wood traits. We conclude that the tree tolerance to hydraulic dysfunction increases with increasing carbon investment in the leaf hydraulic system.

Key words: leaf hydraulic trait, leaf hydraulic conductance, leaf hydraulic vulnerability, trade-off, diffuse- porous, ring-porous, non-porous

金鹰, 王传宽. 九种不同材性的温带树种叶水力性状及其权衡关系. 植物生态学报, 2016, 40(7): 702-710. DOI: 10.17521/cjpe.2016.0064

Ying JIN, Chuan-Kuan WANG. Leaf hydraulic traits and their trade-offs for nine Chinese temperate tree species with different wood properties. Chinese Journal of Plant Ecology, 2016, 40(7): 702-710. DOI: 10.17521/cjpe.2016.0064

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2016.0064

https://www.plant-ecology.com/CN/Y2016/V40/I7/702

图/表 5

表1 9种不同材性的温带树种样木的基本特性(平均值±标准误差, n = 4)

Table 1 Basic characteristics of the sampled trees for the nine temperate tree species with different wood properties (mean ± SE, n = 4)

材性(代码) Wood property (code)	树种(代码) Species (code)	叶习性 Leaf habit	生境 Habitat	胸径 DBH (cm)
散孔材	白桦 Betula platyphylla (BH)	落叶阔叶 Deciduous-broadleaved	山坡中部 Mid slope	24.46 ± 1.05
Diffuse-porous (DP)	山杨 Populus davidiana (SY)	落叶阔叶 Deciduous-broadleaved	山坡上部 Upper slope	31.15 ± 0.55
	紫椴 Tilia amurensis (ZD)	落叶阔叶 Deciduous-broadleaved	山坡上部 Upper slope	25.75 ± 1.30
环孔材	水曲柳 Fraxinus mandshurica (SQL)	落叶阔叶 Deciduous-broadleaved	山坡下部 Toe slope	34.30 ± 0.45
Ring-porous (RP)	蒙古栎 Quercus mongolica (MGL)	落叶阔叶 Deciduous-broadleaved	山坡上部 Upper slope	27.86 ± 1.19
	胡桃楸 Juglans mandshurica (HTQ)	落叶阔叶 Deciduous-broadleaved	山谷 Valley bottom	34.60 ± 1.38
无孔材	红松 Pinus koraiensis (HS)	常绿针叶 Evergreen-coniferous	山坡中部 Mid slope	28.17 ± 0.91
Non-porous (NP)	云杉 Picea koraiensis (YS)	常绿针叶 Evergreen-coniferous	山谷 Valley bottom	30.05 ± 0.65
	樟子松 Pinus sylvestris var. mongolica (ZZS)	常绿针叶 Evergreen-coniferous	山坡中部 Mid slope	26.32 ± 1.15

图1 不同材性树种叶水力性状的比较(平均值±标准误差)。不同大、小写字母分别代表不同材性之间和同一材性不同树种之间差异显著(p < 0.05)。Karea和Kmass分别表示基于叶面积和叶质量的叶导水率; P50表示叶水力脆弱性。材性和树种代码同表1。

Fig. 1 Comparisons of leaf hydraulic traits among the tree species with different wood properties (mean ± SE). Different uppercase and lowercase letters above columns indicate significant differences among different wood properties and among different tree species with the same wood property, respectively (p < 0.05). Karea and Kmass, leaf hydraulic conductance per leaf area and dry mass, respectively; P50, leaf hydraulic vulnerability. See Table 1 for the codes of tree species and wood properties.

图2 3种材性树种叶水力效率与水力脆弱性(P50)的关系。Karea和Kmass分别表示基于叶面积和叶质量的叶导水率。材性代码同表1。

Fig. 2 Relationships between leaf hydraulic efficiency and hydraulic vulnerability (P50) of the trees with wood properties. Karea and Kmass, leaf hydraulic conductance per leaf area and dry mass, respectively. The codes of wood properties are listed in Table 1.

图3 3种材性树种膨压丧失点水势(TLP)与(A)基于叶质量的叶导水率(Kmass)和(B)叶水力脆弱性(P50)的关系。虚线表示关系不显著(p > 0.05)。材性代码同表1。

Fig. 3 Relationships between leaf water potential at turgor loss point (TLP) and (A) leaf-mass-based hydraulic conductance (Kmass) or (B) leaf hydraulic vulnerability (P50) of the trees with wood properties. The dash line denotes non-significant (p > 0.05). The codes of wood properties are listed in Table 1.

图4 基于所有树种的叶水力性状与叶结构性状的关系。Kmass, 基于叶质量的叶导水率; LD, 叶密度; LDMC, 叶干物质含量; LMA, 比叶重; P50, 叶水力脆弱性。

Fig. 4 The relationships between leaf hydraulics and structural traits of all tree species. Kmass, leaf hydraulic conductance per dry mass; LD, leaf density; LDMC, leaf dry mass content; LMA, leaf mass per unit area; P50, leaf hydraulic vulnerability.

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