青藏高原东缘常见阔叶木本植物叶片性状对环境因子的响应

doi:10.17521/cjpe.2019.0174

植物生态学报 ›› 2019, Vol. 43 ›› Issue (10): 863-876.DOI: 10.17521/cjpe.2019.0174

所属专题：青藏高原植物生态学：群落生态学；植物功能性状

青藏高原东缘常见阔叶木本植物叶片性状对环境因子的响应

杨继鸿,李亚楠,卜海燕,张世挺,齐威()

兰州大学生命科学学院草地农业生态系统国家重点实验室, 兰州 730000
兰州大学生命科学学院草地农业生态系统国家重点实验室, 兰州
兰州大学生命科学学院草地农业生态系统国家重点实验室, 兰州 730000 ORCID

收稿日期:2019-07-08 接受日期:2019-10-02 出版日期:2019-10-20 发布日期:2020-02-24
通讯作者: 齐威
基金资助:
国家自然科学基金(31770448);国家自然科学基金(31600329);国家重点研发计划(2017YFC0504800);国家重点研发计划(2018YFD0502400)

Response of leaf traits of common broad-leaved woody plants to environmental factors on the eastern Qinghai-Xizang Plateau

YANG Ji-Hong,LI Ya-Nan,BU Hai-Yan,ZHANG Shi-Ting,QI Wei()

State Key Laboratory of Grassland and Agro-Ecosystems, School of Life Sciences, Lanzhou University, Lanzhou 730000, China

Received:2019-07-08 Accepted:2019-10-02 Online:2019-10-20 Published:2020-02-24
Contact: QI Wei
Supported by:
the National Natural Science Foundation of China(31770448);the National Natural Science Foundation of China(31600329);the National Key R&D Program of China(2017YFC0504800);the National Key R&D Program of China(2018YFD0502400)

摘要/Abstract

摘要：

叶片性状-环境关系对于预测气候变化对植物的影响至关重要。该研究以青藏高原东缘常见阔叶木本植物为研究对象, 从47个样点采集了332个物种共666个种群的叶片, 测量了15个叶片性状, 调查了该区域木本植物叶片性状的变异程度, 并从种内和种间水平探讨了叶片性状对环境的响应及适应策略。结果表明, 反眏叶片大小的性状均具有较高的变异, 其中, 叶片面积是变异程度最大的性状。除气孔密度外, 大多数叶片性状与海拔显著相关。气候是叶片性状变异的重要驱动因素, 3.3%-29.5%的叶片性状变异由气候因子组合解释。其中, 气温对叶片性状变异解释度最高, 日照时间能解释大部分叶片性状的变异, 而降水量对叶片性状变异的解释度相对较小。与环境(海拔和气候因子)显著相关的叶片性状在种内明显少于种间水平, 可能是植物性状之间的协同变化与权衡使种内性状变异比较小, 从而减弱了种内叶片性状与环境因子的相关性。研究结果总体表明,叶片性状与木本植物对环境的适应策略密切相关, 植物通过选择小而厚的叶片和较短的叶柄以适应高海拔的环境。

关键词: 叶片形态特征, 气孔性状, 海拔, 气候因子, 环境因子

Abstract:

Aims Leaf trait-environment relationships are critical for predicting the effects of climate change on plants. Our objective was to reveal the response of leaf traits of common broad-leaved woody plants to environmental factors on the eastern Qinghai-Xizang Plateau. Methods We measured 15 leaf traits of 332 species from 666 populations collected at 47 sites on the eastern Qinghai-Xizang Plateau. We investigated the extent of leaf trait variation in this area, and explored the response and adaptation strategies of leaf traits to environment at intra- and inter-species levels. Important findings Traits related to leaf size exhibited relatively high variation, and the leaf area was the most variant trait. Most leaf traits were significantly associated with elevation, except stomatal density. Climatic factors were important drivers of leaf trait variation because they explained 3.3%-29.5% of leaf trait variation. Meantime, temperature had the highest interpretation degree of leaf trait variation, and sunshine hours could explain the variation of most leaf traits. However, the interpretation degree of precipitation was relatively weak. In addition, the significant relationships between leaf traits and environmental (altitude and climatic) factors at intra-species level were far less than at inter-species levels. The reason for the result may be the coordinated variation and trade-off between plant traits, which make the variation of intra-species traits relatively small, and thus weaken the correlation between intra-plant leaf traits and environmental factors. Overall, leaf traits were closely related to woody plant adaptation strategies to the environment, and small, thick leaves and short petioles were selected for high-altitude plants to adapt to harsh environments such as strong winds and low temperature.

Key words: leaf morphological trait, stomatal trait, altitude, climatic factor, environmental factor

杨继鸿, 李亚楠, 卜海燕, 张世挺, 齐威. 青藏高原东缘常见阔叶木本植物叶片性状对环境因子的响应. 植物生态学报, 2019, 43(10): 863-876. DOI: 10.17521/cjpe.2019.0174

YANG Ji-Hong, LI Ya-Nan, BU Hai-Yan, ZHANG Shi-Ting, QI Wei. Response of leaf traits of common broad-leaved woody plants to environmental factors on the eastern Qinghai-Xizang Plateau. Chinese Journal of Plant Ecology, 2019, 43(10): 863-876. DOI: 10.17521/cjpe.2019.0174

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https://www.plant-ecology.com/CN/Y2019/V43/I10/863

图/表 9

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海拔段 Altitude belt (m)	气候带 Climatic zone	年平均气温 Mean annual air temperature (℃)	无霜期 Frost-free period (d)	生长季 Growing season (d)	地带性木本植被类型 Zonal woody vegetation type
1 600-1 900	亚热带-暖温带 Subtropical-warm temperate	11-15	200-240	230-270	温带-北亚热带阔叶林 Temperate-North subtropical broad-leaved forest
1 900-2 200	暖温带 Warm temperate	8-12	160-210	210-250	温带落叶阔叶林 Temperate deciduous broad-leaved forest
2 200-2 500	暖温带-中温带 Warm temperate-medium temperature	5-9	120-170	190-230	温带落叶阔叶林及针阔混交林 Temperate deciduous broad-leaved forest and coniferous and broad-leaved mixed forest
2 500-2 800	中温带 Medium temperature	3-7	80-130	170-210	温带针阔混交林 Temperate coniferous and broad-leaved mixed forest
2 800-3 100	中温带-亚高山带 Medium temperate-subalpine	1-5	40-90	150-190	温带及亚高山针阔混交林 Temperate and subalpine coniferous and broad-leaved mixed forest
3 100-3 400	亚高山带 subalpine zone	-1-3	0-50	140-170	亚高山针阔混交林 Subalpine coniferous and broad-leaved mixed forest
3 400-3 700	亚高山带-高寒带 Subalpine-alpine	-3-1	0-20	120-160	高寒及亚高山灌丛 Alpine and subalpine shrub
3 700-4 000	高寒带 Alpine	-5- -1	0	100-140	高寒灌丛 Alpine shrub

海拔段 Altitude belt (m)	气候带 Climatic zone	年平均气温 Mean annual air temperature (℃)	无霜期 Frost-free period (d)	生长季 Growing season (d)	地带性木本植被类型 Zonal woody vegetation type
1 600-1 900	亚热带-暖温带 Subtropical-warm temperate	11-15	200-240	230-270	温带-北亚热带阔叶林 Temperate-North subtropical broad-leaved forest
1 900-2 200	暖温带 Warm temperate	8-12	160-210	210-250	温带落叶阔叶林 Temperate deciduous broad-leaved forest
2 200-2 500	暖温带-中温带 Warm temperate-medium temperature	5-9	120-170	190-230	温带落叶阔叶林及针阔混交林 Temperate deciduous broad-leaved forest and coniferous and broad-leaved mixed forest
2 500-2 800	中温带 Medium temperature	3-7	80-130	170-210	温带针阔混交林 Temperate coniferous and broad-leaved mixed forest
2 800-3 100	中温带-亚高山带 Medium temperate-subalpine	1-5	40-90	150-190	温带及亚高山针阔混交林 Temperate and subalpine coniferous and broad-leaved mixed forest
3 100-3 400	亚高山带 subalpine zone	-1-3	0-50	140-170	亚高山针阔混交林 Subalpine coniferous and broad-leaved mixed forest
3 400-3 700	亚高山带-高寒带 Subalpine-alpine	-3-1	0-20	120-160	高寒及亚高山灌丛 Alpine and subalpine shrub
3 700-4 000	高寒带 Alpine	-5- -1	0	100-140	高寒灌丛 Alpine shrub

性状 Traits	物种数量 N	平均值 Mean	最小值 Min	最大值 Max	标准误差 SE	变异系数 CV	偏度 Skewness
LL (cm)	329	7.54	0.80	40.68	0.26	0.63	2.24
LW (cm)	329	4.79	0.36	27.18	0.21	0.81	1.85
LL/LW	329	2.02	0.26	12.97	0.07	0.62	3.24
LA (cm²)	330	32.77	0.36	1 312.49	4.68	2.59	11.43
LT (mm)	314	0.18	0.05	0.44	0.003	0.28	1.25
LWC	280	0.60	0.33	0.85	0.01	0.14	-0.28
PL (cm)	310	2.13	0.19	18.89	0.13	1.08	2.64
SLA (cm²·g^-1)	320	173.49	46.47	392.15	3.29	0.34	0.93
LA/PL (cm²·cm^-1)	310	15.05	1.61	109.17	0.75	0.87	3.17
SD (No·mm^-2)	318	279.93	32.55	896.50	7.43	0.47	1.35
SL (μm)	320	26.05	10.77	64.99	0.41	0.28	0.77
SW (μm)	320	18.69	3.79	43.77	0.35	0.33	0.16
SL/SW	320	1.46	0.55	3.04	0.02	0.20	1.08
SA (μm²)	320	412.56	34.19	2 234.30	13.61	0.59	2.10
SPI	318	0.10	0.01	0.34	0.003	0.54	1.13

性状 Traits	物种数量 N	平均值 Mean	最小值 Min	最大值 Max	标准误差 SE	变异系数 CV	偏度 Skewness
LL (cm)	329	7.54	0.80	40.68	0.26	0.63	2.24
LW (cm)	329	4.79	0.36	27.18	0.21	0.81	1.85
LL/LW	329	2.02	0.26	12.97	0.07	0.62	3.24
LA (cm²)	330	32.77	0.36	1 312.49	4.68	2.59	11.43
LT (mm)	314	0.18	0.05	0.44	0.003	0.28	1.25
LWC	280	0.60	0.33	0.85	0.01	0.14	-0.28
PL (cm)	310	2.13	0.19	18.89	0.13	1.08	2.64
SLA (cm²·g^-1)	320	173.49	46.47	392.15	3.29	0.34	0.93
LA/PL (cm²·cm^-1)	310	15.05	1.61	109.17	0.75	0.87	3.17
SD (No·mm^-2)	318	279.93	32.55	896.50	7.43	0.47	1.35
SL (μm)	320	26.05	10.77	64.99	0.41	0.28	0.77
SW (μm)	320	18.69	3.79	43.77	0.35	0.33	0.16
SL/SW	320	1.46	0.55	3.04	0.02	0.20	1.08
SA (μm²)	320	412.56	34.19	2 234.30	13.61	0.59	2.10
SPI	318	0.10	0.01	0.34	0.003	0.54	1.13

性状 Traits	海拔 Altitude				PC1				PC2
性状 Traits	N(+)	N(-)	χ²	p	N(+)	N(-)	χ²	p	N(+)	N(-)	χ²	p
LL	20	50	12.86	<0.01	40	30	1.43	0.23	36	34	0.06	0.81
LW	22	47	9.06	<0.01	39	29	1.47	0.23	38	31	0.71	0.40
LL/LW	28	41	2.45	0.12	38	31	0.71	0.40	31	38	0.71	0.40
LA	19	51	14.63	<0.01	39	31	0.91	0.34	41	29	2.06	0.15
LT	36	25	1.98	0.16	33	26	0.83	0.36	25	36	1.98	0.16
PL	22	37	3.81	0.05	34	27	0.80	0.37	27	32	0.42	0.52
LWC	21	25	0.35	0.56	26	20	0.78	0.38	20	26	0.78	0.38
SLA	36	30	0.55	0.46	35	31	0.24	0.62	30	36	0.55	0.46
LA/PL	23	38	3.69	0.06	30	31	0.02	0.90	39	22	4.74	0.03
SD	27	35	1.03	0.31	27	35	1.03	0.31	34	28	0.58	0.45
SL	40	24	4.00	0.04	33	31	0.06	0.80	30	34	0.25	0.61
SW	40	24	4.00	0.04	33	31	0.06	0.80	35	29	0.56	0.45
SL/SW	33	31	0.06	0.80	34	30	0.25	0.62	36	28	1.00	0.32
SA	36	28	1.00	0.32	32	32	0.00	1.00	31	33	0.06	0.80
SPI	31	33	0.06	0.80	30	34	0.25	0.62	38	26	2.25	0.13

青藏高原东缘常见阔叶木本植物叶片性状对环境因子的响应

Response of leaf traits of common broad-leaved woody plants to environmental factors on the eastern Qinghai-Xizang Plateau

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性状 Traits	海拔(控制PC1) Altitude (control PC1)	海拔(控制PC2) Altitude (control PC2)	海拔(控制PC1和PC2) Altitude (control PC1and PC2)
LL	-0.472^**	-0.382^**	-3.00^**
LW	-0.467^**	-0.323^**	-0.289^**
LL/LW	0.168^**	0.043	0.086
LA	-0.491^**	-0.366^**	-0.297^**
LT	0.014	0.069	0.118^*
PL	-0.305^**	-0.205^**	-0.161^**
LWC	-0.074	0.068	0.082
SLA	-0.071	-0.115	-0.127^*
LA/PL	-0.404^**	-0.334^**	-0.269^**
SD	0.015	0.021	-0.010
SL	-0.151^**	-0.095	-0.049
SW	-0.146^**	-0.073	-0.029
SL/SW	0.055	0.000	-0.012
SA	-0.153^**	-0.086	-0.042
SPI	-0.140^*	-0.064	-0.041

线性模型 Linear regression model	AIC	R²	p
LL = -0.16 + 0.13MTCO + 0.03MGSS - 0.14MGST + 0.20PDR	-983.45	0.29	<0.001
LW = -0.65 + 0.13MTCO + 0.30PDR + 0.03MGSS - 0.12MGST	-788.19	0.23	<0.001
LL/LW = 0.96 + 0.006PWE - 0.22PDR - 0.0006MAP + 0.03MGSS	-1 008.31	0.04	<0.01
LA = -0.91 + 0.26MTCO + 0.52MGSS + 0.47PDR - 0.26MGST	-507.75	0.30	<0.001
LT = 0.49 - 0.02PWE - 0.22MGST + 0.11MTCO + 0.03MGSS + 0.39PDR - 0.01 MGSP + 0.002MAP	-1 333.81	0.08	<0.001
PL = -0.94 + 0.05MTCO + 0.01MGSS + 0.16PDR	-587.23	0.11	<0.001
LWC = 0.37 + 0.01MGST + 0.0002MAP	-1 382.27	0.04	<0.01
SLA = 0.05 + 0.02MGSP - 0.17MTCO + 0.34MGST - 0.05MGSS + 0.02PWE - 0.56PDR -0.002MAP	-1 237.94	0.08	<0.001
LA/PL = -0.16 + 0.133MTCO + 0.03MGSS - 0.14MGST + 0.22PDR	-766.38	0.19	<0.001
SL = 1.10 + 0.01MTCO + 0.002MGSS + 0.05PDR	-1 355.05	0.06	<0.001
SW = 1.42 + 0.01MTCO	-1 154.49	0.05	<0.001
SL/SW = 0.005 + 0.002MGSS + 0.01PDR - 0.002PWE - 0.005MGST	-1 553.84	0.03	<0.05
SA = 2.83 + 0.02MTCO	-819.19	0.05	<0.001
SPI = 3.19 + 0.02MTCO	-813.97	0.04	<0.001

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