黔中喀斯特木本植物功能性状变异及其适应策略

doi:10.17521/cjpe.2017.0270

植物生态学报 ›› 2018, Vol. 42 ›› Issue (5): 562-572.DOI: 10.17521/cjpe.2017.0270

所属专题：植物功能性状

黔中喀斯特木本植物功能性状变异及其适应策略

钟巧连^1,^2,³,刘立斌^1,^2,⁴,许鑫^1,^2,³,杨勇^1,^2,³,郭银明^1,^2,³,许海洋^1,^2,³,蔡先立^1,^2,³,倪健^1,^2,^4,^*()

1 中国科学院地球化学研究所环境地球化学国家重点实验室, 贵阳 550081
2 中国科学院普定喀斯特生态系统观测研究站, 贵州普定 562100
3 中国科学院大学, 北京 100049
4 浙江师范大学化学与生命科学学院, 浙江金华 321004

收稿日期:2017-10-29 修回日期:2018-03-27 出版日期:2018-05-20 发布日期:2018-07-20
通讯作者: 倪健
基金资助:
国家自然科学基金(41471049);国家全球变化重大科学研究计划项目(2013CB956704)

Variations of plant functional traits and adaptive strategy of woody species in a karst forest of central Guizhou Province, southwestern China

ZHONG Qiao-Lian^1,^2,³,LIU Li-Bin^1,^2,⁴,XU Xin^1,^2,³,YANG Yong^1,^2,³,GUO Yin-Ming^1,^2,³,XU Hai-Yang^1,^2,³,CAI Xian-Li^1,^2,³,NI Jian^1,^2,^4,^*()

1 State Key Laboratory of Environmental Geochemistry, Institute of Geochemistry, Chinese Academy of Sciences, Guiyang 550081, China
2 Puding Karst Ecosystem Research Station, Chinese Academy of Sciences, Puding, Guizhou 562100, China
3 University of Chinese Academy of Sciences, Beijing 100049, China
4 College of Chemistry and Life Sciences, Zhejiang Normal University, Jinhua, Zhejiang 321004, China

Received:2017-10-29 Revised:2018-03-27 Online:2018-05-20 Published:2018-07-20
Contact: Jian NI
Supported by:
Supported by the National Natural Science Foundation(41471049);the National Key Basic Research Program for Global Change(2013CB956704)

摘要/Abstract

摘要：

性状变异反映了植物的生活史对策。该研究以贵州普定县天龙山10种木本植物为对象, 通过分析枝叶和根系9个功能性状的种间与种内变异, 揭示植物对喀斯特生境的适应策略。结果表明: (1) 9个性状变异程度不同, 细根组织密度的种间和种内变异系数最大, 分别达96.47%和51.44%, 小枝干物质含量的种间与种内变异最小, 分别为11.67%和6.83%。(2)种间水平的细根组织密度在不同物种中没有显著的差异, 比根长、叶厚度、叶面积、比叶面积、叶干物质含量、叶组织密度、小枝干物质含量和小枝组织密度均表现出显著的差异。在种内, 比叶面积差异显著, 其他性状差异不显著。(3)绝大多数叶和枝性状间显著相关, 比根长与比叶面积显著负相关, 其他根系性状与枝叶性状相关性不显著。总之, 与同纬度非喀斯特地区植物相比, 普定喀斯特地区植物具有较小的叶面积和比根长度, 较大的叶干物质含量、叶组织密度等一系列有利于减小蒸腾和储存养分的功能性状组合, 这可能是其适应干旱贫瘠的喀斯特环境的主要生态策略。

关键词: 喀斯特森林, 生活型, 生长型, 性状组合, 种内变异, 种间变异

Abstract:

Aims The aims are to characterize key plant functional traits and their interactions of woody species growing in special and harsh karst habitats, and to explore their potential ways in adapting harsh karst habitats.

Methods A comprehensive survey of nine plant functional traits (including above- and below-ground ones) was conducted in a 100 m × 30 m permanent plot in the Tianlongshan Mountain of Puding County, central Guizhou Province, southwestern China in the summer 2016. Five dominant tree species (Carpinus pubescens, Machilus cavaleriei, Itea yunnanensis, Platycarya strobilacea, Lithocarpus confinis), three shrubs (Zanthoxylum ovalifolium, Stachyurus obovatus, Rhamnus heterophylla) and two vines (Rosa cymosa and Dalbergia hancei) in an evergreen and deciduous broadleaved mixed forest were chosen as target species. Nine traits of leaf, stem, branch and root were investigated and measured. Key features of these nine functional traits of ten woody species were numerically characterized. Traits variations among plant species, life form and leaf phenology group were further investigated. Relationships among key functional traits and between above- and below-ground traits were statistically analyzed.

Important findings (1) Nine traits varied in varying degrees. The maximum and minimum coefficient of interspecific variation were the fine root tissue density (FRTD) and twig dry-matter content (TDMC), 96.47% and 11.67%, respectively. Similarly, the largest and smallest coefficients of intraspecific variation were also FRTD and TDMC, 51.44% and 6.83%, respectively; (2) At the interspecific level, among different species FRTD had no significant difference, but other traits including specific root length (SRL), leaf thickness (LT), leaf area (LA), specific leaf area (SLA), leaf dry-matter content (LDMC), leaf tissue density (LTD), TDMC and twig tissue density (TTD) showed significant differences (p < 0.01). At the intraspecific level, however, SLA showed significant difference, and differences of other traits were not significant. (3) There was a significant correlation between most leaf and branch traits, and SRL vs. SLA were negatively correlated. However, there was no significant correlation among other root traits and leaf and twig traits. In a word, compared to the functional traits in tree species of non-karst evergreen broad-leaved forests in the same latitude, karst woody plants in Puding had a series of functional traits, such as smaller LA, SLA and larger LDMC and LTD and so on, which are beneficial to reducing transpiration and storing nutrient. This may be its main ecological strategy for adapting to arid and poor karst environments.

Key words: karst forest, life form, growth form, trait combination, intraspecific variation, interspecific variation

钟巧连, 刘立斌, 许鑫, 杨勇, 郭银明, 许海洋, 蔡先立, 倪健. 黔中喀斯特木本植物功能性状变异及其适应策略. 植物生态学报, 2018, 42(5): 562-572. DOI: 10.17521/cjpe.2017.0270

ZHONG Qiao-Lian, LIU Li-Bin, XU Xin, YANG Yong, GUO Yin-Ming, XU Hai-Yang, CAI Xian-Li, NI Jian. Variations of plant functional traits and adaptive strategy of woody species in a karst forest of central Guizhou Province, southwestern China. Chinese Journal of Plant Ecology, 2018, 42(5): 562-572. DOI: 10.17521/cjpe.2017.0270

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图/表 6

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物种 Species	生长型 Growth form	生活型 Life form	采样株数 Sample individuals	叶质地 Leaf texture
安顺润楠 Machilus cavaleriei	乔木 Tree	常绿 Evergreen	20	革质 Leathery
云南鼠刺 Itea yunnanensis	乔木 Tree	常绿 Evergreen	20	薄革质 Thinly leathery
窄叶石栎 Lithocarpus confinis	乔木 Tree	常绿 Evergreen	21	厚纸质 Thickly papery
化香树 Platycarya strobilacea	乔木 Tree	落叶 Deciduous	25	纸质 Papery
云贵鹅耳枥 Carpinus pubescens	乔木 Tree	落叶 Deciduous	20	厚纸质 Thickly papery
刺异叶花椒 Zanthoxylum ovalifolium	灌木 Shrub	常绿 Evergreen	20	革质 Leathery
倒卵叶旌节花 Stachyurus obovatus	灌木 Shrub	常绿 Evergreen	20	革质或亚革质 Leathery or subcoriaceous
异叶鼠李 Rhamnus heterophylla	灌木 Shrub	常绿 Evergreen	20	纸质 Papery
小果蔷薇 Rosa cymosa	藤本 Liana	常绿 Evergreen	20	薄纸质 Thinly papery
藤黄檀 Dalbergia hancei	藤本 Liana	落叶 Deciduous	20	膜质 Membranous

物种 Species	生长型 Growth form	生活型 Life form	采样株数 Sample individuals	叶质地 Leaf texture
安顺润楠 Machilus cavaleriei	乔木 Tree	常绿 Evergreen	20	革质 Leathery
云南鼠刺 Itea yunnanensis	乔木 Tree	常绿 Evergreen	20	薄革质 Thinly leathery
窄叶石栎 Lithocarpus confinis	乔木 Tree	常绿 Evergreen	21	厚纸质 Thickly papery
化香树 Platycarya strobilacea	乔木 Tree	落叶 Deciduous	25	纸质 Papery
云贵鹅耳枥 Carpinus pubescens	乔木 Tree	落叶 Deciduous	20	厚纸质 Thickly papery
刺异叶花椒 Zanthoxylum ovalifolium	灌木 Shrub	常绿 Evergreen	20	革质 Leathery
倒卵叶旌节花 Stachyurus obovatus	灌木 Shrub	常绿 Evergreen	20	革质或亚革质 Leathery or subcoriaceous
异叶鼠李 Rhamnus heterophylla	灌木 Shrub	常绿 Evergreen	20	纸质 Papery
小果蔷薇 Rosa cymosa	藤本 Liana	常绿 Evergreen	20	薄纸质 Thinly papery
藤黄檀 Dalbergia hancei	藤本 Liana	落叶 Deciduous	20	膜质 Membranous

性状 Trait	平均值±标准偏差 Mean ± SD	最小值 Minimum	最大值 Maximum	变异系数 Coefficient of variation (%)
叶片厚度 Leaf thickness (mm)	0.17 ± 0.07	0.03	0.33	39.41
叶面积 Leaf area (cm²)	17.74 ± 10.44	2.15	69.28	58.85
比叶面积 Specific leaf area (cm²?g^-1)	134.44 ± 45.80	55.35	281.34	34.07
叶干物质含量 Leaf dry-matter content (g?g^-1)	0.40 ± 0.06	0.28	0.69	14.79
叶组织密度 Leaf tissue density (g?cm^-3)	0.54 ± 0.13	0.27	1.07	24.07
小枝干物质含量 Twig dry-matter content of branchlets (g?g^-1)	0.48 ± 0.06	0.37	0.64	11.67
小枝组织密度 Twig tissue density (g?cm^-3)	0.69 ± 0.19	0.28	2.50	27.53
比根长 Specific root length (cm?g^-1)	251.91 ± 128.79	54.56	987.41	51.12
细根组织密度 Fine root tissue density (g?cm^-3)	0.85 ± 0.82	0.12	7.33	96.47

性状 Trait	平均值±标准偏差 Mean ± SD	最小值 Minimum	最大值 Maximum	变异系数 Coefficient of variation (%)
叶片厚度 Leaf thickness (mm)	0.17 ± 0.07	0.03	0.33	39.41
叶面积 Leaf area (cm²)	17.74 ± 10.44	2.15	69.28	58.85
比叶面积 Specific leaf area (cm²?g^-1)	134.44 ± 45.80	55.35	281.34	34.07
叶干物质含量 Leaf dry-matter content (g?g^-1)	0.40 ± 0.06	0.28	0.69	14.79
叶组织密度 Leaf tissue density (g?cm^-3)	0.54 ± 0.13	0.27	1.07	24.07
小枝干物质含量 Twig dry-matter content of branchlets (g?g^-1)	0.48 ± 0.06	0.37	0.64	11.67
小枝组织密度 Twig tissue density (g?cm^-3)	0.69 ± 0.19	0.28	2.50	27.53
比根长 Specific root length (cm?g^-1)	251.91 ± 128.79	54.56	987.41	51.12
细根组织密度 Fine root tissue density (g?cm^-3)	0.85 ± 0.82	0.12	7.33	96.47

	LT (mm)	LA (cm²)	SLA (cm²?g^-1)	LDMC (g?g^-1)	LTD (g?cm^-3)	TDMC (g?g^-1)	TTD (g?cm^-3)	SRL (cm?g^-1)	FRTD (g?cm^-3)
乔木 Tree	0.19 ± 0.06^a (33.25%/ 16.52%)	24.66 ± 9.02^a (36.59%/ 22.63%)	115.64 ± 44.16^a (28.18%/ 19.33%)	0.42 ± 0.05^a (11.59%/ 6.26%)	0.53 ± 0.10^a (19.19%/ 10.05%)	0.48 ± 0.06^a (11.48%/ 6.36%)	0.71 ± 0.11^a (15.50%/ 12.52%)	237.89 ± 89.59^a (37.66%/ 32.72%)	0.90 ± 0.93^a (102.97%/ 68.94%)
灌木 Shrub	0.16±0.06^b (41.51%/ 14.41%)	12.3 ± 6.62^b (53.85%/ 20.10%)	143.49 ± 35.369^b (24.64%/ 11.73%)	0.35 ± 0.05^b (14.31%/ 6.11%)	0.50 ± 0.15^a (30.47%/ 13.83%)	0.49 ± 0.06^a (12.66%/ 7.01%)	0.68 ± 0.17^a (24.70%/ 13.26%)	271.09 ± 179.73^a (66.30%/ 62.05%)	0.88 ± 0.89^a (100.61%/ 31.27%)
藤本 Liana	0.11 ± 0.04^c (35.94%/ 24.26%)	7.43 ± 1.48^c (19.95%/ 17.33%)	169.689 ± 40.13^c (23.65%/ 23.17%)	0.41 ± 0.06^c (13.86%/ 12.77%)	0.61 ± 0.15^b (24.52%/ 15.23%)	0.46 ± 0.04^b (9.80%/ 7.70%)	0.63 ± 0.32^a (50.94%/ 43.33%)	255.17 ± 110.55^a (43.32%/ 37.93%)	0.69 ± 0.21^a (30.37%/ 25.04%)
常绿植物 Evergreen	0.17 ± 0.07^d (39.34%/ 14.81%)	18.15 ± 10.18^d (56.12%/ 17.12%)	131.19 ± 43.38^d (34.47%/ 13.99%)	0.39 ± 0.06^d (16.29%/ 7.06%)	0.54 ± 0.13^d (27.98%/ 12.12%)	0.48 ± 0.05^d (11.73%/ 7.17%)	0.71 ± 0.14^d (20.81%/ 13.38%)	238.14 ± 145.15^d (60.95%/ 45.90%)	0.88 ± 0.92^d (93.11%/ 59.18%)
落叶植物 Deciduous	0.14 ± 0.04^e (29.49%/ 23.57%)	16.84 ± 11.08^e (65.79%/ 29.42%)	145.85 ± 41.79^e (30.12%/ 26.66%)	0.43 ± 0.04^e (9.60%/ 8.56%)	0.55 ± 0.08^d (16.58%/ 12.52%)	0.50 ± 0.05^e (10.65%/ 6.02%)	0.72 ± 0.27^d (36.91%/ 31.61%)	237.66 ± 94.71^d (34.61%/ 34.23%)	0.91 ± 0.963^d (102.4%2/ 52.96%)

黔中喀斯特木本植物功能性状变异及其适应策略

Variations of plant functional traits and adaptive strategy of woody species in a karst forest of central Guizhou Province, southwestern China

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变量 Variables	LT (mm)	LA (cm²)	SLA (cm²?g^-1)	LDMC (g?g^-1)	LTD (g?cm^-3)	TDMC (g?g^-1)	TTD (g?cm^-3)	SRL (cm?g^-1)	FRTD (g?cm^-3)
物种 Species	0.86	0.78	0.67	0.75	0.74	0.64	0.19	0.18	0.08
生长型 Growth form	0.13	0.50	0.16	0.31	0.02	0.06	0.03	0.02	0.01
生活型 Life form	0.08	0.01	0.03	0.12	0.01	0.04	0.00	0.02	0.00
生长型×生活型 Growth form × Life form	0.07	0.55	0.49	0.35	0.19	0.02	0.17	0.37	0.04

	LT	LA	SLA	LDMC	LTD	TDMC	TTD	SRL
LA	0.65^**
SLA	-0.83^**	-0.56^**
LDMC	-0.21^**	0.01	-0.22^**
LTD	-0.70^**	-0.47^**	0.34^**	0.64^**
TDMC	-0.38^**	0.41^**	0.17^*	0.43^**	0.45^**
TTD	-0.17^*	0.10	-0.02	0.28^**	0.29^**	0.34^**
SRL	0.12	-0.00	-0.18^*	0.12	-0.13	0.09	0.04
FRTD	0.02	-0.31^**	-0.09	0.10	0.12	0.02	-0.01	-0.00

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