罗汉松科3种植物茎和根木质部水分运输、解剖结构与机械强度之间的关系

doi:10.17521/cjpe.2020.0402

植物生态学报 ›› 2021, Vol. 45 ›› Issue (6): 659-669.DOI: 10.17521/cjpe.2020.0402

罗汉松科3种植物茎和根木质部水分运输、解剖结构与机械强度之间的关系

陆世通¹^,², 陈森¹^,², 李彦¹^,²^,³, 王忠媛¹, 潘天天¹^,², 叶琳峰¹^,², 谢江波¹^,²^,³^,^*()

¹浙江农林大学省部共建亚热带森林培育国家重点实验室, 杭州 311300
²浙江农林大学林业与生物技术学院, 杭州 311300
³荒漠与绿洲生态国家重点实验室, 中国科学院新疆生态与地理研究所, 乌鲁木齐 830011

收稿日期:2020-12-04 接受日期:2021-03-17 出版日期:2021-06-20 发布日期:2021-09-09
通讯作者: 谢江波
作者简介:^*(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)

Relationships among xylem transport, anatomical structure and mechanical strength in stems and roots of three Podocarpaceae species

LU Shi-Tong¹^,², CHEN Sen¹^,², LI Yan¹^,²^,³, WANG Zhong-Yuan¹, PAN Tian-Tian¹^,², YE Lin-Feng¹^,², XIE Jiang-Bo¹^,²^,³^,^*()

¹State Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou 311300, China
²College of Forestry and Biotechnology, Zhejiang A&F University, Hangzhou 311300, China
³State Key Laboratory of Desert and Oasis Ecology, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Ürümqi 830011, China

Received:2020-12-04 Accepted:2021-03-17 Online:2021-06-20 Published:2021-09-09
Contact: XIE Jiang-Bo
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：

水力失效是植物干旱死亡的主要机制。量化分析水力性状的种间和器官间差异是预测树木在气候变化下的响应甚至生存能力的基础。该研究对比分析了罗汉松科3种植物器官(茎和根)水平上水力功能性状的差异, 并探讨其与解剖结构和机械强度之间的关系。在湿生同质园内选择罗汉松科3种植物, 测定了茎和根木质部水力功能性状(最大比导率(K_s)和栓塞抗性(P₅₀))、解剖结构性状(管胞直径(D_t)、水力直径(D_h)、管胞密度(N_t)、管胞壁厚(T_w)、纹孔膜直径(D_p)和纹孔密度(N_p))和机械强度(木材密度(WD)和管胞厚度跨度比((t/b)²))。结果发现: (1)罗汉松科3种植物茎木质部不存在效率-安全权衡, 而根木质部存在权衡。(2)茎K_s与D_p显著正相关, 与(t/b)²和WD无关; 茎P₅₀与D_p极显著负相关, 与(t/b)²和WD无关。(3)根K_s与D_h显著正相关, 与T_w和(t/b)²极显著负相关; 根P₅₀与T_w、(t/b)²和WD均极显著正相关。在罗汉松科植物中, 根木质部性状与输水效率和栓塞抗性的密切关系是解释其存在效率-安全权衡的基础, 而茎木质部的过度建造是茎不存在效率-安全权衡的原因, 木质部的过度建造仍需要更多的实验证据。

关键词: 栓塞, 导水率, 脆弱曲线, 管胞, 罗汉松科

Abstract:

Aims Hydraulic failure is one of the primary cause of plant mortality during drought. Thus, quantitative analysis on inter-specific and inter-organ variance in hydraulic traits can help us to predict the response and even survivability of species under climate change.

Methods Here, three Podocarpaceae species (Podocarpus macrophyllus, P. macrophyllusvar. maki and Nageia nagi) grown in a mesic common garden were studied, with xylem hydraulic function (specific hydraulic conductivity (K_s); embolism resistance (P₅₀)), anatomical structure (tracheid diameter (D_t); hydraulic diameter (D_h); tracheid wall thickness (T_w); tracheid density (N_t); pit membrane diameter (D_p); pit density (N_p)) and mechanical strength (wood density (WD); tracheid thickness to span ratio ((t/b)²)) measured. Then, we analyzed hydraulic traits variance at the organ level (stem and root) in three Podocarpaceae species, and investigated the relationships among xylem hydraulic traits, anatomical structure and mechanical strength in stems and roots.

Important findings We found that: 1) The stem xylem in three Podocarpaceae species exhibited no safety- efficiency trade-off. In contrast, the root xylem exhibited safety-efficiency trade-off. 2) For stems, K_s was positively correlated with D_p, but decoupled with stem WD and (t/b)²; Stem P₅₀ was negatively correlated with D_p, but not correlated with WD and (t/b)². 3) For roots, K_s was positively correlated with hydraulic diameter, but negatively correlated with root T_w and (t/b)²; Root P₅₀ was positively correlated with T_w, (t/b)² and WD. Root xylem traits exhibited strong relationships with both K_s and P₅₀, demonstrating its cause-and-effect basis for the safety-efficiency tradeoff. On the other hand, the absence of safety-efficiency tradeoff in stems may be attributed to the overbuilt xylem of Podocarpaceae. More experimental evidence on the overbuilding of xylem is desired in the future study.

Key words: embolism, hydraulic conductivity, vulnerability curve, tracheid, Podocarpaceae

陆世通, 陈森, 李彦, 王忠媛, 潘天天, 叶琳峰, 谢江波. 罗汉松科3种植物茎和根木质部水分运输、解剖结构与机械强度之间的关系. 植物生态学报, 2021, 45(6): 659-669. DOI: 10.17521/cjpe.2020.0402

LU Shi-Tong, CHEN Sen, LI Yan, WANG Zhong-Yuan, PAN Tian-Tian, YE Lin-Feng, XIE Jiang-Bo. Relationships among xylem transport, anatomical structure and mechanical strength in stems and roots of three Podocarpaceae species. Chinese Journal of Plant Ecology, 2021, 45(6): 659-669. DOI: 10.17521/cjpe.2020.0402

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0402

https://www.plant-ecology.com/CN/Y2021/V45/I6/659

图/表 7

表1 罗汉松科3种植物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 trees of three Podocarpaceae species (mean ± SE)

树种 Species	海拔 Altitude (m)	坡向 Slope direction	坡度 Slope (°)	土壤类型 Soil type	土壤含水量 Soil water content (%)	土壤容重 Soil bulk density (g·cm^-3)	树高 Tree height (m)	胸径 DBH (cm)
罗汉松 Podocarpus macrophyllus	42.47 ± 0.23	西南 SW	3	黄红壤亚类 Yellow red soil subclass	36.87 ± 1.81	1.22 ± 0.15	5.93 ± 0.42	13.50 ± 1.55
短叶罗汉松 P. macrophyllus var. maki	39.40 ± 0.44	西南 SW	8	黄红壤亚类 Yellow red soil subclass	35.24 ± 1.75	1.20 ± 0.10	2.89 ± 0.32	11.99 ± 1.54
竹柏 Nageia nagi	41.48 ± 0.20	西南 SW	1	黄红壤亚类 Yellow red soil subclass	36.34 ± 1.74	1.23 ± 0.08	3.66 ± 0.18	8.52 ± 0.64

图1 三种罗汉松科植物木材解剖横切面和纵切面光学显微镜图像。A-D, 罗汉松茎横切图和纵切图、根横切图和纵切图。E-H, 短叶罗汉松茎横切图和纵切图、根横切图和纵切图。I-L, 竹柏茎横切图和纵切图、根横切图和纵切图。

Fig. 1 Examples of light microscopy images of transverse and vertical sections of wood anatomy of three Podocarpaceae species. A-D, Transverse sections and vertical sections of stem, transverse sections and vertical sections of root in Podocarpus macrophyllus. E-H, Transverse sections and vertical sections of stem, transverse sections and vertical sections of root in P. macrophyllus var. maki. I-L, Transverse sections and vertical sections of stem, transverse sections and vertical sections of root in Nageia nagi.

图2 罗汉松科3种植物的水力功能性状图(平均值±标准误, n = 7)。不同小写字母表示在p ≤ 0.05水平上差异显著。

Fig. 2 Hydraulic functional traits of three Podocarpaceae species (mean ± SE, n = 7). Different lowercase letters indicate significant differences at p ≤ 0.05 level.

图3 罗汉松科3种植物木质部解剖结构图(平均值±标准误, n = 7)。不同小写字母表示在p ≤ 0.05水平上差异显著。

Fig. 3 Xylem anatomical structure of three Podocarpaceae species (mean ± SE, n = 7). Different lowercase letters indicate significant differences at p ≤ 0.05 level.

图4 罗汉松科3种植物木质部机械强度图(平均值±标准误, n = 7)。不同小写字母表示在p ≤ 0.05水平上差异显著。

Fig. 4 Xylem mechanical traits of three Podocarpaceae species (mean ± SE, n = 7). Different lowercase letters indicate significant differences at p ≤ 0.05 level.

图5 罗汉松科植物木质部输水效率与栓塞抗性的关系。A, 本研究罗汉松科3种植物茎。B, 本研究罗汉松科3种植物根。C, 罗汉松科12种植物茎(数据来源于Brodribb和Hill (1999), van der Willigen等(2000), Pittermann和Sperry (2006))。D, 罗汉松科5种植物根(数据来源于Pittermann和Sperry (2006))。ns, 没有显著关系。

Fig. 5 Relationship between hydraulic conductivity and embolism resistance in the three Podocarpaceae species. A, The stems of three Podocarpaceae species in this paper. B, The roots of three Podocarpaceae species in this paper. C, The stems of twelve Podocarpaceae species (Data from Brodribb & Hill (1999), van der Willigen et al. (2000), Pittermann & Sperry (2006)). D, The roots of five Podocarpaceae species (Data from Pittermann & Sperry (2006)). ns, non-significant relationships.

图6 罗汉松科植物10个木质部性状之间的相关性分析。A, 茎木质部性状。B, 根木质部性状。*, p ≤ 0.05; **, p ≤ 0.01。Dh, 水力直径(μm); Dp, 纹孔膜直径(μm); Dt, 管胞直径(μm); Ks, 最大比导率(kg·m-1·MPa-1·s-1); Np, 纹孔密度(mm-2); Nt, 管胞密度(mm-2); P50, 导水率损失50%时的水势(-MPa); Tw, 管胞厚度(μm); WD, 木质部密度(g·cm-2); (t/b)2, 管胞厚度跨度比。

Fig. 6 Correlation analysis of ten xylem traits in Podocarpaceae species. A, Xylem traits of stem. B, Xylem traits of root. *, p ≤ 0.05; **, p ≤ 0.01. Dh, hydraulic diameter (μm); Dp, pit membrane diameter (μm); Dt, tracheid diameter (μm); Ks, specific hydraulic conductivity (kg·m-1·MPa-1·s-1); Np, pit density (mm-2); Nt, tracheid density (mm2); P50, the xylem water potential causing 50% loss of hydraulic conductivity (-MPa); Tw, tracheid wall thickness (μm); WD, wood density (g·cm-2); (t/b)2, tracheid thickness to span ratio.

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罗汉松科3种植物茎和根木质部水分运输、解剖结构与机械强度之间的关系

Relationships among xylem transport, anatomical structure and mechanical strength in stems and roots of three Podocarpaceae species

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