中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析

doi:10.17521/cjpe.2020.0367

植物生态学报 ›› 2021, Vol. 45 ›› Issue (4): 394-403.DOI: 10.17521/cjpe.2020.0367

中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析

倪鸣源, ARITSARA Amy Ny Aina, 王永强, 黄冬柳, 项伟, 万春燕, 朱师丹^*()

广西大学亚热带农业生物资源保护与利用国家重点实验室, 南宁 530004; 广西大学林学院广西森林生态与保育重点实验室, 南宁 530004

收稿日期:2020-11-09 接受日期:2021-01-18 出版日期:2021-04-20 发布日期:2021-03-09
通讯作者: ORCID: *朱师丹: 0000-0002-9228-368X(zhushidan@gxu.edu.cn)
作者简介:* zhushidan@gxu.edu.cn
基金资助:
广西八桂青年学者项目和广西自然科学基金(2017GXNSFBA198188)

Analysis of xylem anatomy and function of representative tree species in a mixed evergreen and deciduous broad-leaved forest of mid-subtropical karst region

NI Ming-Yuan, ARITSARA Amy Ny Aina, WANG Yong-Qiang, HUANG Dong-Liu, XIANG Wei, WAN Chun-Yan, ZHU Shi-Dan^*()

Guangxi Key Laboratory of Forest Ecology and Conservation, College of Forestry, Guangxi University, Nanning 530004, China; and State Key Laboratory for Conservation and Utilization of Subtropical Agro-bioresources, College of Forestry, Guangxi University, Nanning 530004, China

Received:2020-11-09 Accepted:2021-01-18 Online:2021-04-20 Published:2021-03-09
Contact: ZHU Shi-Dan
Supported by:
Guangxi Bagui Young Scholars Program and the Natural Science Foundation of Guangxi(2017GXNSFBA198188)

摘要/Abstract

摘要：

树木木质部主要由导管、纤维和薄壁组织组成, 分别具有运输、支撑和贮存的生理功能。由于木质部空间限制, 一种组织比例的增加会导致其他组织比例的降低, 因而可能表现出权衡关系。分析木质部组织比例和权衡关系有助于了解植物的生理生态适应性。该研究选择中亚热带喀斯特常绿落叶阔叶混交林21种典型树种(10种落叶树种, 11种常绿树种), 测定枝条木质部各组织比例, 计算水力相关指标并分析性状之间的相关性。结果表明: (1)与全球木质部解剖数据对比分析, 喀斯特树种木质部趋向具有较高比例的薄壁组织; (2)喀斯特树种导管组织比例与薄壁和纤维组织比例之间没有显著的相关性, 但是薄壁和纤维组织比例之间有显著的权衡关系; (3)常绿和落叶树种的木质部水力运输安全性(导管壁加固系数)和效率性(理论导水率)均具有显著的权衡关系, 但是这两个类群线性回归的截距存在显著差异, 即在相同的理论导水率条件下, 落叶树种比常绿树种具有较高的导管壁加固系数(安全性), 可能与常绿树种具有更多的轴向薄壁组织有关。喀斯特树种木质部解剖特征表明薄壁组织的贮存功能对喀斯特树种(尤其是常绿树种)的干旱适应具有重要作用。

关键词: 导管组织, 薄壁组织, 纤维组织, 导管壁加固系数, 理论导水率, 权衡

Abstract:

Aims Vessel, fibers, and parenchyma are the main components of tree xylem. They are responsible for water transport, mechanical support, and water and nutrients storage. Given the limited xylem space, consistent investment in one type of tissue would constrain the space available for other types of tissue, thus resulting in a possible trade-off among different tissues in their fractions. Analysis of the fractions of tissue types in xylem and the trade-off would contribute to better understanding of the eco-physiological adaptation of plants.

Methods We selected 21 characteristic tree species (10 deciduous and 11 evergreen) from a mixed evergreen and deciduous broad-leaved forest located in the mid-subtropical karst region, and measured their xylem tissue fractions. In addition, we calculated the hydraulic-related structural traits in xylems and examined the correlations among various traits.

Important findings Compared to the global average values of xylem tissue fractions, the karst tree species tended to have a higher proportion of parenchyma. The fraction of vessel lumen was not correlated with fiber and parenchyma fractions across the tree species investigated. Instead, a significant trade-off was observed between fractions of fiber and parenchyma. A trade-off between the hydraulic efficiency (i.e. theoretical hydraulic conductivity) and safety (vessel wall reinforcement) was observed across both the deciduous and the evergreen tree species. The two contrasting group of karst trees differenced significantly in the intercepts of the lines for trade-offs. For given conductivity, the deciduous tree species exhibited stronger vessel well reinforcement (safety) than the evergreen tree species, which might be due to the fact that evergreen trees species had more axial parenchyma. Hence, this study revealed the specificity of xylem anatomy in karst tree species. Water and resource storage in xylem parenchyma are vital to karst trees (evergreens in particular) for their adaptation to the water-limiting environment.

Key words: vessel, parenchyma, fibers, vessel wall reinforcement coefficient, theoretical hydraulic conductivity, trade-off

倪鸣源, ARITSARA Amy Ny Aina, 王永强, 黄冬柳, 项伟, 万春燕, 朱师丹. 中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析. 植物生态学报, 2021, 45(4): 394-403. DOI: 10.17521/cjpe.2020.0367

NI Ming-Yuan, ARITSARA Amy Ny Aina, WANG Yong-Qiang, HUANG Dong-Liu, XIANG Wei, WAN Chun-Yan, ZHU Shi-Dan. Analysis of xylem anatomy and function of representative tree species in a mixed evergreen and deciduous broad-leaved forest of mid-subtropical karst region. Chinese Journal of Plant Ecology, 2021, 45(4): 394-403. DOI: 10.17521/cjpe.2020.0367

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0367

https://www.plant-ecology.com/CN/Y2021/V45/I4/394

图/表 6

表1 广西木论21种喀斯特树种的叶习性、胸径和树高(平均值±标准差)

Table 1 Leaf types by longevity, diameter at breast-height (DBH), and height of the 21 karst tree species studied in Mulun, Guangxi (mean ± SD)

物种 Species	科 Family	胸径 DBH (cm)	树高 Height (m)
落叶 Deciduous
黄梨木 Boniodendron minus	无患子科Sapindaceae	12.4 ± 0.3	8.3 ± 0.2
禾串树 Bridelia balansae	大戟科Euphorbiaceae	13.6 ± 1.0	7.5 ± 0.5
大叶紫珠 Callicarpa macrophylla	马鞭草科Verbenaceae	6.5 ± 0.4	5.5 ± 0.7
麻楝 Chukrasia tabularis	楝科 Meliaceae	12.0 ± 1.4	8.0 ± 0.5
浆果楝 Cipadessa baccifera	楝科 Meliaceae	7.9 ± 1.1	6.4 ± 0.9
毛果巴豆 Croton lachnocarpus	大戟科Euphorbiaceae	8.4 ± 0.3	6.5 ± 0.2
伞花木 Eurycorymbus cavaleriei	无患子科Sapindaceae	10.6 ± 0.6	7.6 ± 0.3
青檀 Pteroceltis tatarinowii	榆科 Ulmaceae	15.6 ± 0.6	8.7 ± 0.2
菜豆树 Radermachera sinica	紫葳科Bignoniaceae	13.5 ± 0.5	8.3 ± 0.2
圆叶乌桕 Triadica rotundifolia	大戟科Euphorbiaceae	15.8 ± 4.2	8.5 ± 0.3
常绿 Evergreen
假鱼骨木 Psydrax dicocca	茜草科Rubiaceae	9.2 ± 1.1	6.5 ± 0.5
灰岩棒柄花 Cleidion bracteosum	大戟科Euphorbiaceae	7.3 ± 0.3	5.2 ± 0.2
岩生厚壳桂 Cryptocarya calcicola	樟科 Lauraceae	8.1 ± 0.4	6.5 ± 0.2
青冈 Cyclobalanopsis glauca	壳斗科Fagaceae	10.9 ± 1.2	7.4 ± 0.6
大叶水榕 Ficus glaberrima	桑科 Moraceae	19.5 ± 3.4	7.7 ± 0.7
山小橘 Glycosmis pentaphylla	芸香科Rutaceae	6.4 ± 0.3	4.3 ± 0.3
黑木姜子 Litsea salicifolia	樟科 Lauraceae	7.6 ± 0.8	6.8 ± 0.4
小芸木 Micromelum integerrimum	芸香科Rutaceae	6.2 ± 1.1	5.3 ± 0.8
千里香 Murraya paniculata	芸香科Rutaceae	6.4 ± 0.1	5.0 ± 0.2
广西密花树 Myrsine kwangsiensis	紫金牛科 Myrsinaceae	7.6 ± 0.3	6.1 ± 0.2
铁榄Sinosideroxylon pedunculatum	山榄科Sapotaceae	8.8 ± 0.4	6.5 ± 0.2

表1 广西木论21种喀斯特树种的叶习性、胸径和树高(平均值±标准差)

Table 1 Leaf types by longevity, diameter at breast-height (DBH), and height of the 21 karst tree species studied in Mulun, Guangxi (mean ± SD)

物种 Species	科 Family	胸径 DBH (cm)	树高 Height (m)
落叶 Deciduous
黄梨木 Boniodendron minus	无患子科Sapindaceae	12.4 ± 0.3	8.3 ± 0.2
禾串树 Bridelia balansae	大戟科Euphorbiaceae	13.6 ± 1.0	7.5 ± 0.5
大叶紫珠 Callicarpa macrophylla	马鞭草科Verbenaceae	6.5 ± 0.4	5.5 ± 0.7
麻楝 Chukrasia tabularis	楝科 Meliaceae	12.0 ± 1.4	8.0 ± 0.5
浆果楝 Cipadessa baccifera	楝科 Meliaceae	7.9 ± 1.1	6.4 ± 0.9
毛果巴豆 Croton lachnocarpus	大戟科Euphorbiaceae	8.4 ± 0.3	6.5 ± 0.2
伞花木 Eurycorymbus cavaleriei	无患子科Sapindaceae	10.6 ± 0.6	7.6 ± 0.3
青檀 Pteroceltis tatarinowii	榆科 Ulmaceae	15.6 ± 0.6	8.7 ± 0.2
菜豆树 Radermachera sinica	紫葳科Bignoniaceae	13.5 ± 0.5	8.3 ± 0.2
圆叶乌桕 Triadica rotundifolia	大戟科Euphorbiaceae	15.8 ± 4.2	8.5 ± 0.3
常绿 Evergreen
假鱼骨木 Psydrax dicocca	茜草科Rubiaceae	9.2 ± 1.1	6.5 ± 0.5
灰岩棒柄花 Cleidion bracteosum	大戟科Euphorbiaceae	7.3 ± 0.3	5.2 ± 0.2
岩生厚壳桂 Cryptocarya calcicola	樟科 Lauraceae	8.1 ± 0.4	6.5 ± 0.2
青冈 Cyclobalanopsis glauca	壳斗科Fagaceae	10.9 ± 1.2	7.4 ± 0.6
大叶水榕 Ficus glaberrima	桑科 Moraceae	19.5 ± 3.4	7.7 ± 0.7
山小橘 Glycosmis pentaphylla	芸香科Rutaceae	6.4 ± 0.3	4.3 ± 0.3
黑木姜子 Litsea salicifolia	樟科 Lauraceae	7.6 ± 0.8	6.8 ± 0.4
小芸木 Micromelum integerrimum	芸香科Rutaceae	6.2 ± 1.1	5.3 ± 0.8
千里香 Murraya paniculata	芸香科Rutaceae	6.4 ± 0.1	5.0 ± 0.2
广西密花树 Myrsine kwangsiensis	紫金牛科 Myrsinaceae	7.6 ± 0.3	6.1 ± 0.2
铁榄Sinosideroxylon pedunculatum	山榄科Sapotaceae	8.8 ± 0.4	6.5 ± 0.2

图1 菜豆树木质部染色切片(A)和经过人工辨别和绘制后的木质部组织结构分布图(B)。绿色, 导管管腔; 黄色, 导管壁; 红色, 射线细胞; 蓝色, 轴向薄壁细胞; 紫色, 纤维细胞。

Fig. 1 A stained xylem cross-section image of Radermachera sinica(A), and the same image in which different xylem tissues were manually coded with different colors (B). Green, vessel lumen; Yellow, vessel wall; Red, ray parenchyma; Blue, axial parenchyma; Purple, fibers.

图2 基于TRY Database数据库的全球805种木本植物木质部各组分比例的平均值和广西木论21种喀斯特树种的木质部各组分的比例。

Fig. 2 Xylem tissue partitioning of the 21 karst woody species in Mulun, Guangxi, and 805 woody species data downloaded from the TRY Database.

图3 喀斯特树种在全球木质部组织划分谱的位置(A)和木质部各组织比例之间的相关性(B)。●, 喀斯特树种; △, 全球数据(TRY)。 Fb,纤维组织比例; TPf, 薄壁组织比例; Vs, 导管组织比例。ns, p > 0.05; ***, p < 0.001。

Fig. 3 Distributions of the karst woody species in the global xylem partitioning spectrum (A) and relationships among xylem tissue fractions (B). ●, karst woody species; △, global observations from TRY. Fb,fibers fraction; TPf, total parenchyma fraction; Vs, vessels fraction. ns, p > 0.05; ***, p < 0.001.

图4 广西木论喀斯特木本植物10个木质部性状(A)和21个树种(B)的主成分分析图。○, 落叶树种; ●, 常绿树种。APf,轴向薄壁组织比例; Dh,导管水力直径; Fb,纤维组织比例; Kt,理论导水率; RPf,射线组织比例; t,相连接两个导管的细胞壁厚度之和; t/b,导管壁加固系数;TPf,总的薄壁组织比例; Vd,导管密度; Vs,导管组织比例。

Fig. 4 Principal component analysis for 10 xylem traits of woody plant (A), and 21 karst woody species (B) in Mulun, Guangxi. ○, deciduous; ●, evergreen. APf,axial parenchyma fraction; Dh,hydraulically-mean vessel diameter; Fb,fiber fraction; Kt,theoretical hydraulic conductivity; RPf,ray parenchyma fraction; t,double wall thickness measured from vessel pairs; t/b,vessel wall reinforcement coefficient; TPf,total parenchyma fraction; Vd,vessel density; Vs,vessel lumen fraction.

图5 广西木论喀斯特21个树种导管壁加固系数(t/b)与理论导水率(Kt)的相关关系的简化主轴回归分析。○: 落叶树种(D), ●: 常绿树种(E)。落叶树种回归方程(灰色): y = -0.31x - 0.67, 常绿树种回归方程(黑色): y = -0.41x - 0.71。**,p< 0.01。

Fig. 5 Reduced major axis regression of vessel wall reinforcement coefficient (t/b) and theoretical hydraulic conductivity (Kt) of 21 karst woody species in Mulun, Guangxi. ○, deciduous (D); ●, evergreen (E). Linear regression of deciduous species (grey): y = -0.31x - 0.67; linear regression of evergreen species (black): y = -0.41x- 0.71. **, p< 0.01.

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中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析

Analysis of xylem anatomy and function of representative tree species in a mixed evergreen and deciduous broad-leaved forest of mid-subtropical karst region

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