落羽杉和池杉功能性状随高度的变异与协同

doi:10.17521/cjpe.2022.0308

植物生态学报 ›› 2023, Vol. 47 ›› Issue (11): 1561-1575.DOI: 10.17521/cjpe.2022.0308

落羽杉和池杉功能性状随高度的变异与协同

汤璐瑶, 方菁, 钱海蓉, 张博纳, 上官方京, 叶琳峰, 李姝雯, 童金莲, 谢江波()

浙江农林大学省部共建亚热带森林培育国家重点实验室, 浙江农林大学林业与生物技术学院, 杭州 311300

收稿日期:2022-07-25 接受日期:2023-02-15 出版日期:2023-11-20 发布日期:2023-12-22
通讯作者: *谢江波(0208xiejiangbo@163.com)
基金资助:
国家自然科学基金(31770651);国家自然科学基金(41730638);国家自然科学基金(31901280)

Variation and coordination in functional traits along the tree height of Taxodium distichum and Taxodium distichum var. imbricatum

TANG Lu-Yao, FANG Jing, QIAN Hai-Rong, ZHANG Bo-Na, SHANGGUAN Fang-Jing, YE Lin-Feng, LI Shu-Wen, TONG Jin-Lian, XIE Jiang-Bo()

State Key Laboratory of Subtropical Silviculture, College of Forestry and Biotechnology, Zhejiang A&F University, Hangzhou 311300, China

Received:2022-07-25 Accepted:2023-02-15 Online:2023-11-20 Published:2023-12-22
Contact: *XIE Jiang-Bo(0208xiejiangbo@163.com)
Supported by:
National Natural Science Foundation of China(31770651);National Natural Science Foundation of China(41730638);National Natural Science Foundation of China(31901280)

摘要/Abstract

摘要：

高大树木木质部水分运输阻力和叶片蒸腾速率随树高的增加导致高度梯度水分供需矛盾。量化分析相关功能性状随高度的变异与性状协同关系, 有助于深入理解植物的水分供需机制。该研究选取了生长在湿生同质园中的落羽杉(Taxodium distichum)及其变种池杉(T. distichum var. imbricatum), 采用回归分析、单因素方差分析、通径分析等方法探究其水力(枝比导率(K_s)、叶比导率(K_l)、导水率损失50%时的水势(P₅₀)、最大蒸腾速率(T_r)、正午叶水势(ψ_MD)、胡伯尔值(H_v)等)、光合(最大净光合速率(P_n))和碳经济性状(比叶质量(LMA)、木质部密度(WD))随高度的变异规律、协同关系以及种间性状差异。结果发现: (1)落羽杉和池杉K_l、H_v、P_n和LMA随高度增加, 其中P_n的增加可能与中冠层的T_r和最大气孔导度(G_s)下降有关。(2)种内性状间协同关系: 落羽杉和池杉K_s与H_v显著负相关, 落羽杉WD与K_s显著正相关, 池杉WD与H_v极显著负相关。(3)落羽杉和池杉高冠层存在水分限制, 由达西定律和T_r计算的理论水分供需比(r)量化了水分供需能力的下降, 且它们r = 0时的理论最大高度(落羽杉: 32 m (95%置信区间上限: 57 m); 池杉: 21 m (95%置信区间上限: 27 m))在历史记载的最大高度范围内。(4)池杉各个冠层高度K_l、H_v和LMA显著高于落羽杉, 而P_n、T_r和G_s显著更低; 池杉中、高冠层水力安全边界(HSM)显著更高, P₅₀显著更低: 池杉保守的水力策略与低资源获取能力导致其最大理论高度较低, 而落羽杉激进的水力策略与高资源获取能力导致其最大理论高度较高。

关键词: 植物功能性状, 性状变异, 高度梯度, 水分供需, 落羽杉属

Abstract:

Aims The xylem water transport resistance and leaf transpiration rate of tall trees increased with tree height, resulting in a water supply-demand paradox along the tree height gradient. Quantitative analysis of the variation and coordination of related functional traits along tree height will be conducive to deeply understanding the water supply and demand mechanism of plants.

Methods Here, Taxodium distichum and its variety T. distichumvar. imbricatum grown in a mesic common garden were studied, with hydraulic (sapwood-specific hydraulic conductivity (K_s), leaf specific conductivity (K_l), the water potential causing 50% loss of conductivity (P₅₀), maximum transpiration rate (T_r), midday leaf water potential (ψ_MD), Huber value (H_v), etc.), photosynthetic (maximum photosynthetic rate (P_n)) and carbon economic traits (leaf mass per unit area (LMA), wood density (WD)) measured. Traits variation and coordination along the tree height and traits differences between the same canopy of T. distichum and T. distichumvar. imbricatum were analyzed by a series of methods, including regression analysis, one-way ANOVA, and path analysis.

Important findings We found that: (1) K_l, H_v, P_n and LMA in T. distichum and T. distichumvar. imbricatum increased along the height, and the increase of P_n may be related to the decrease of T_r and maximum operational stomatal conductance (G_s) in the middle canopy. (2) Coordination relationships between intraspecific traits: K_s in T. distichum and T. distichumvar. imbricatum was significantly negatively related to H_v, WD in T. distichum was significantly positively related to K_s, and WD in T. distichumvar. imbricatum was significantly negatively related to H_v. (3) There was water limitation in the higher canopy of T. distichum and T. distichumvar. imbricatum. The theoretical water supply and demand ratio (r) calculated by Darcy’s Law and T_r confirmed this limitation. The theoretical maximum height when r = 0: T. distichum 32 m (upper bound of 95% confidence interval: 57 m); T. distichumvar. imbricatum 21 m (upper bound of 95% confidence interval: 27 m), was consistent with the maximum height recorded historically. (4) K_l, H_v and LMA in canopies of T. distichumvar. imbricatum were significantly higher than those of T. distichum,while P_n, T_r and G_s were significantly lower; hydraulic safety margin (HSM) in the middle and higher canopies of T. distichumvar. imbricatum was significantly higher, and P₅₀was significantly lower: The conservative hydraulic strategy of T. distichumvar. imbricatum corresponded to lower resource acquisition capacity, which in turn resulted in lower maximum growth height. The radical hydraulic strategy of T. distichum corresponded to higher resource acquisition capacity, which in turn resulted in higher maximum growth height.

Key words: plant functional trait, trait variation, height gradient, water supply and demand, Taxodium

汤璐瑶, 方菁, 钱海蓉, 张博纳, 上官方京, 叶琳峰, 李姝雯, 童金莲, 谢江波. 落羽杉和池杉功能性状随高度的变异与协同. 植物生态学报, 2023, 47(11): 1561-1575. DOI: 10.17521/cjpe.2022.0308

TANG Lu-Yao, FANG Jing, QIAN Hai-Rong, ZHANG Bo-Na, SHANGGUAN Fang-Jing, YE Lin-Feng, LI Shu-Wen, TONG Jin-Lian, XIE Jiang-Bo. Variation and coordination in functional traits along the tree height of Taxodium distichum and Taxodium distichum var. imbricatum. Chinese Journal of Plant Ecology, 2023, 47(11): 1561-1575. DOI: 10.17521/cjpe.2022.0308

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0308

https://www.plant-ecology.com/CN/Y2023/V47/I11/1561

图/表 9

表1 落羽杉和池杉采样点及样树的基本特征(平均值±标准误)

Table 1 Basic characteristics of the sampling plots and the trees of Taxodium distichum and T. distichum var. imbricatum (mean ± SE)

树种 Species	海拔 Altitude (m)	坡向 Slope direction	土壤水分状况 Soil moisture status	树高 Tree height (m)	胸径 DBH (cm)	冠幅 Crown (m)
落羽杉 T. distichum	24-44	西南 SW	湿润 Moist	15-17	34.04 ± 2.02	6.22 ± 0.18
池杉 T. distichum var. imbricatum	42-44	西南 SW	湿润 Moist	13-15	31.42 ± 0.87	6.32 ± 0.21

表2 世界各地记录的落羽杉和池杉最大高度

Table 2 Maximum heights of Taxodium distichum and T. distichum var. imbricatum recorded worldwide

数据来源 Data Source	落羽杉 T. distichum (m)	池杉 T. distichum var. imbricatum (m)	网站 Website
Flora of China	50	25	http://www.iplant.cn/foc
《北美植物志》 Flora of North America		30	http://floranorthamerica.org/Main_Page
树木和灌木在线 Trees and Shrubs online	30.48-45.72		https://treesandshrubsonline.org/
《生命大百科全书》 Encyclopedia of Life	39.62		https://eol.org/
微软必应搜索引擎 Microsoft Bing	38.1	25	https://cn.bing.com/?FORM=Z9FD1
百度搜索引擎 Baidu	25-50	25	https://www.baidu.com/
世界植物在线 Plants of the World Online	46		https://powo.science.kew.org/

图1 落羽杉和池杉叶片结构和木质部解剖结构沿高度变异的示意图。

Fig. 1 Schematic diagram of variations in leaf structure and xylem anatomical structure of Taxodium distichum and T. distichum var. imbricatum along the tree height.

表3 本研究相关性状、符号及单位

Table 3 List for traits related to this study: symbols and units employed

性状 Trait	符号 Symbol	单位 Unit
枝比导率 Sapwood-specific hydraulic conductivity	K_s	kg·m^-1·s^-1·MPa^-1
叶比导率 Leaf specific conductivity	K_l	g·m^-1·s^-1·MPa^-1
正午叶水势 Midday leaf water potential	ψ_MD	MPa
导水率损失50%时的水势 Water potential causing 50% loss of conductivity	P₅₀	MPa
水力安全边界 Hydraulic safety margin	HSM	MPa
胡伯尔值 Huber value	H_v	m²·m^-2
比叶质量 Leaf mass per unit area	LMA	kg·m^-2
木质部密度 Wood density	WD	g·cm^-3
最大净光合速率(光饱和CO₂同化率) Maximum net photosynthetic rate	P_n	μmol·m^-2·s^-1
最大蒸腾速率 Maximum transpiration rate	T_r	mmol·m^-2·s^-1
最大气孔导度 Maximum stomatal conductance	G_s	mol·m^-2·s^-1
水分利用效率 Water use efficiency	WUE	μmol·mmol^-1
理论水分供给速率 Theoretical water supply rate	Q_crit	mmol·m^-2·s^-1
管胞直径 Tracheid diameter	D_t	μm
管胞密度 Tracheid density	N_t	10³·mm^-2
管胞壁厚度 Tracheid wall thickness	T_w	μm
厚度跨度比 Thickness span ratio	(t/b)²	μm·μm^-1
纹孔口直径 Pit aperture diameter	D_pa	μm
纹孔膜直径 Pit membrane diameter	D_pm	μm

图2 落羽杉(BC, ▲)和池杉(PC, ●)功能性状沿高度的回归分析(平均值±标准误)。图中的平均值和误差条用于直观地展示性状变化而不参与计算。

Fig. 2 Regression analysis of water-carbon traits along the tree height in Taxodium distichum (BC, ▲) and T. distichum var. imbricatum (PC, ●) (mean ± SE). The mean value and error bar on the figure are used to visually show the variation of traits without participating in the calculation. Trait symbols see Table 3.

图3 落羽杉和池杉在种内不同冠层高度(大写字母)和同一冠层高度种间(小写字母)的性状差异分析(平均值±标准误)。不同字母表示在p ≤ 0.05水平上差异显著。

Fig. 3 Differences in traits for the same height in different species (uppercase letters) and for different canopy heights in the same species (lowercase letters) in Taxodium distichum and T. distichum var. imbricatum (mean ± SE). Different letters indicate significant differences at p ≤ 0.05 level. Trait symbols see Table 3.

图4 落羽杉(BC, ▲)和池杉(PC, ●)木质部解剖结构沿高度的回归分析(平均值±标准误)。图中的平均值和误差条用于直观地展示性状变化而不参与计算。

Fig. 4 Regression analysis of xylem anatomical structure along the tree height in Taxodium distichum (BC, ▲) and T. distichum var. imbricatum (PC, ●) (mean ± SE). The mean value and error bar on the figure are used to visually show the variation of traits without participating in the calculation. Trait symbols see Table 3.

图5 落羽杉和池杉种内性状间的通径分析。性状缩写见表3。箭头表示性状间的拟合关系, 蓝线表示正相关, 红线表示负相关。标准化通径系数用线的粗细表示, 系数越大, 线越粗。标准化路径系数的显著性标示在箭头上(ns, p ≥ 0.05; *, p < 0.05; **, p < 0.001)。

Fig. 5 Path analysis on traits in Taxodium distichum and T. distichum var. imbricatum. Trait symbols see Table 3. The arrows indicate the proposed links between traits. Blue lines indicate positive relationships, red lines indicate negative relationships. Standard path coefficients are represented as the thickness of the line, the larger the coefficient, the thicker the line. Significances of standard path coefficients are shown on the arrows (ns, p ≥ 0.05; *, p < 0.05; **, p < 0.001). NFI, normed fit index; SRMR, standardized root mean square residual.

图6 落羽杉(BC, ▲)和池杉(PC, ●)水分供需比(r)沿高度的回归分析(平均值±标准误)。回归线取95%置信区间上限, 虚线为回归线的延长线。

Fig. 6 Regression analysis of the ratio of water supply and demand (r) along the tree height in Taxodium distichum (BC, ▲) and T. distichum var. imbricatum (PC, ●) (mean ± SE). Upper bound of 95% confidence interval (CI) is taken, and the dotted lines are the extension of the regression lines.

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落羽杉和池杉功能性状随高度的变异与协同

Variation and coordination in functional traits along the tree height of Taxodium distichum and Taxodium distichum var. imbricatum

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