季节尺度毛乌素沙地黑沙蒿的叶片性状网络特征及驱动因素

doi:10.17521/cjpe.2024.0025

植物生态学报 ›› 2024, Vol. 48 ›› Issue (12): 1650-1665.DOI: 10.17521/cjpe.2024.0025 cstr: 32100.14.cjpe.2024.0025

所属专题：植物功能性状

季节尺度毛乌素沙地黑沙蒿的叶片性状网络特征及驱动因素

徐铭泽¹^,², 赵洪贤¹, 李成², 李满乐¹^,³, 田赟¹, 刘鹏¹, 查天山¹^,^*()

¹北京林业大学水土保持学院, 北京 100083
²自然资源部重庆典型矿区生态修复野外科学观测研究站(重庆地质矿产研究院), 重庆 401120
³中路高科交通科技集团有限公司, 北京 100088

收稿日期:2024-01-24 接受日期:2024-05-27 出版日期:2024-12-20 发布日期:2024-12-20
通讯作者: *查天山(tianshanzha@bjfu.edu.cn)
基金资助:
国家自然科学基金(32071842);国家自然科学基金(32101588);国家自然科学基金(31901366);国家自然科学基金(32071843);重庆市自然科学基金(CSTB2022NSCQ-MSX1121)

Characteristics of seasonal leaf trait network and its drivers in Artemisia ordosica in the Mau Us Sandy Land

XU Ming-Ze¹^,², ZHAO Hong-Xian¹, LI Cheng², LI Man-Le¹^,³, TIAN Yun¹, LIU Peng¹, ZHA Tian-Shan¹^,^*()

¹School of Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
²Observation and Research Station of Ecological Restoration for Chongqing Typical Mining Areas (Ministry of Natural Resources, Chongqing Institute of Geology and Mineral Resources), Chongqing 401120, China
³RLOH High Science and Technology Group, Beijing 100088, China

Received:2024-01-24 Accepted:2024-05-27 Online:2024-12-20 Published:2024-12-20
Contact: *ZHA Tian-Shan(tianshanzha@bjfu.edu.cn)
Supported by:
National Natural Science Foundation of China(32071842);National Natural Science Foundation of China(32101588);National Natural Science Foundation of China(31901366);National Natural Science Foundation of China(32071843);Natural Science Foundation of Chongqing(CSTB2022NSCQ-MSX1121)

摘要/Abstract

摘要： 明晰植物性状的网络特征是功能生态学的研究热点, 目前对于季节尺度植物的性状网络(PTN)以及驱动因素的认识仍十分有限。该研究采用LI-6400便携式光合仪定期测量了黑沙蒿(Artemisia ordosica)生长季(5-9月)叶片光响应、CO₂响应曲线及关键的叶片结构性状和生化性状指标, 探究25个叶性状间的相关性及其PTN特征。结果显示: 季节尺度, 叶性状间存在显著的相关关系, 最紧密的性状耦合关系发生在光合生理性状间, 共有67对生理-生理性状组合显著相关。PTN边密度、直径、平均路径长度和模块度分别为0.58、3、1.51和0.08, 叶组织密度、单位面积叶氮含量和蒸腾速率的介数相对较高, 是PTN中的“桥梁”。PTN并没有绝对的中心性状, 生理性状整体表现出了较高的度、特征向量中心度和聚类系数, 表明该PTN由生理性状共同主导。进一步的分析结果表明: 季节尺度25个叶性状可以被压缩为两大类性状维度, 一类是受空气温度和土壤水分调控的性状, 一类是受光合有效辐射调控的性状。研究结果强调, 在评估不同性状功能组对气候波动的潜在反应时, 需要区分环境因素对不同性状的影响程度。如果将植物性状对环境的季节反应纳入一个统一的范式, 将会错误地评估植物的性状适应性结果。

关键词: 叶片性状网络, 季节动态, 黑沙蒿, 性状变异

Abstract:

Aims Clarifying the network characteristics of plant traits is one of the research hotspots in functional ecology. However, our understanding of plant trait networks (PTN) and their driving factors on a seasonal scale is still limited.

Methods We used a LI-6400 portable photosynthetic instrument to regularly measure the light response and carbon dioxide response curves of Artemisia ordosica during the growing season (May to September), as well as leaf structural and biochemical indicators, to explore the correlation and trait network characteristics among 25 leaf traits.

Important findings There were significant correlations between leaf traits at the seasonal scale, and the closest coupling relationship between traits occurred between photosynthetic physiological traits, with a total of 67 pairs of physiological trait combinations showing significant correlation. The edge density, diameter, average path length, and modularity of the PTN are 0.58, 3, 1.51, and 0.08, respectively. The betweenness of leaf tissue density, leaf nitrogen content per unit area, and transpiration rate is relatively high, making it a “bridge” in PTN. PTN does not have an absolute central trait, and physiological traits as a whole exhibit high degree, eigenvector centrality, and clustering coefficient, indicating that the trait network is jointly dominated by physiological traits. Further analysis indicated that the 25 leaf traits at the seasonal scale can be compressed into two major trait dimensions: one is regulated by air temperature and soil moisture, and the other is regulated by photosynthetic effective radiation. The results emphasize that when evaluating the potential response of different trait functional groups to climate fluctuations, it is necessary to distinguish the degree and effect of environmental factors on different traits. If the seasonal response of plant traits to the environment is included in a unified paradigm, it will incorrectly evaluate the adaptability of plant traits.

Key words: leaf trait network, seasonal dynamics, Artemisia ordosica, trait variation

徐铭泽, 赵洪贤, 李成, 李满乐, 田赟, 刘鹏, 查天山. 季节尺度毛乌素沙地黑沙蒿的叶片性状网络特征及驱动因素. 植物生态学报, 2024, 48(12): 1650-1665. DOI: 10.17521/cjpe.2024.0025

XU Ming-Ze, ZHAO Hong-Xian, LI Cheng, LI Man-Le, TIAN Yun, LIU Peng, ZHA Tian-Shan. Characteristics of seasonal leaf trait network and its drivers in Artemisia ordosica in the Mau Us Sandy Land. Chinese Journal of Plant Ecology, 2024, 48(12): 1650-1665. DOI: 10.17521/cjpe.2024.0025

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0025

https://www.plant-ecology.com/CN/Y2024/V48/I12/1650

图/表 11

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	[ 郑云普, 党承华, 郝立华, 程东娟, 徐明 (2016). 华北平原玉米叶片光合及呼吸过程对实验增温的适应性. 生态学报, 36, 5236-5246.]
[52]	Zhong GW, Tian Y, Liu P, Jia X, Zha TS (2022). Leaf traits and resource use efficiencies of 19 woody plant species in a plantation in Fangshan, Beijing, China. Forests, 14, 63. DOI: 10.3390/f14010063.

监测样株 Monitoring sample plant	冠幅 Crown diameter (cm × cm)		最大高度 Maximum height (cm)
监测样株 Monitoring sample plant	东西向 East-west	南北向 South-north	最大高度 Maximum height (cm)
HSH1	111	100	53
HSH2	108	104	61
HSH3	115	120	58

监测样株 Monitoring sample plant	冠幅 Crown diameter (cm × cm)		最大高度 Maximum height (cm)
监测样株 Monitoring sample plant	东西向 East-west	南北向 South-north	最大高度 Maximum height (cm)
HSH1	111	100	53
HSH2	108	104	61
HSH3	115	120	58

叶性状类别 Category of leaf functional trait	性状 Trait	缩写 Abbreviation	单位 Unit	生理生态解释 Physiological and ecological explanation
光合生理性状 Photosynthetic physiological trait	最大净光合速率 Maximum net photosynthetic rate	A_max	μmol·m^-2·s^-1	光合潜力与资源获取策略 Photosynthetic potential and resource acquisition strategies
	最大羧化速率 Maximum carboxylation rate	V_cmax	μmol·m^-2·s^-1	核酮糖-1,5-双磷酸羧化酶/加氧酶(Rubisco)催化的最大羧化反应速率 Maximum carboxylation rate of the enzyme ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco)
	最大电子传递速率 Maximum electron transfer rate	J_max	μmol·m^-2·s^-1	支持核酮糖双磷酸再生的最大电子传输速率 Maximum electron transfer rate for supporting ribulose bisphosphate regeneration
	暗呼吸速率 Dark respiration rate	R_d	μmol·m^-2·s^-1	黑暗环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under dark conditions
	光下暗呼吸速率 Dark respiration rate under light	R_l	μmol·m^-2·s^-1	光照环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under light conditions
	光饱和碳同化速率 Photo-saturated carbon assimilation rate	A_sat	μmol CO₂·m^-2·s^-1	碳同化效率 Carbon assimilation efficiency
	气孔导度 Stomatal conductance	g_s	mol H₂O·m⁻²·s⁻¹		气体交换与资源获取 Gas exchange and resource acquisition
	蒸腾速率 Transpiration rate	E	mmol H₂O·m⁻²·s⁻¹		耗水特性 Water consumption characteristics
	水分利用效率 Water use efficiency	WUE	μmol·mmol^-1		抗旱性与水资源利用 Drought resistance and water resource utilization
	氮利用效率 Nitrogen use efficiency	NUE	μmol·g^-1·s^-1		叶片养分利用、生理特性和生存策略 Nutrient utilization, physiological characteristics, and survival strategies of leaves
	最大光化学效率 Maximal photochemical efficiency	F_v/F_m			光系统II (PSII)反应中心的最大光能转化效率以及叶片的健康程度 The maximum light energy conversion efficiency of photosystem II (PSII) and the health status of leaves
	光化学淬灭 Photochemical quenching	q_P			PSII反应中心的开放程度以及光化学活性的高低 Degree of openness of PSII and the level of photochemical activity
	非光化学淬灭 Non-photochemical quenching	NPQ			植物的光保护能力 Light protection ability of plants
	电子传递速率 Electron transfer rate	ETR	μmol·m^-2·s^-1		植物的耐光性和适应性 Light resistance and adaptability of plants
结构性状 Structural trait	叶厚度 Leaf thickness	L_t	mm		生存策略以及光合呼吸、蒸腾相关的碳成本 Survival strategy and carbon costs related to photosynthesis, respiration and transpiration
	比叶质量 Leaf mass per area	LMA	g·m^-2		资源获取和抗逆性 Resource acquisition and resilience
	叶组织密度 Leaf tissue density	LTD	g·cm^-3		叶片韧性和抗逆性 Leaf toughness and stress resistance
	叶片相对含水量 Leaf relative water content	LRWC	%		植物的水分状况、抗旱性及环境适应 Water status, drought resistance and environmental adaptation
	叶干物质含量 Leaf dry matter content	LDMC	g·g^-1		抗逆性和对抗物理危害的防御能力 Resistance to adversity and defense against physical hazards
生物化学性状 Biochemical trait	基于面积的叶氮含量 Leaf nitrogen content per unit area	N_area	g·m^-2		资源获取和衡量光合能力 Resource acquisition and characterization of photosynthetic capacity
	基于面积的叶碳含量 Leaf carbon content per unit area	C_area	g·m^-2		构建和防御 Building and defense
	碳氮比 Ratio of leaf carbon content to nitrogen content	C:N			元素分配与适应策略 Element allocation and adaptation strategies
	叶绿素含量 Chlorophyll content	Chl	μg·cm^-2		健康程度与光合作用 Health level and photosynthesis
	叶绿素a/b Ratio of chlorophyll a to chlorophyll b	Chl a/b			光能吸收和能量运输 Light absorption and energy transport
	类胡萝卜素含量 Carotenoid content	Car	μg·cm^-2		光合作用与光保护必需色素 Photosynthesis and essential pigments for photoprotection

叶性状类别 Category of leaf functional trait	性状 Trait	缩写 Abbreviation	单位 Unit	生理生态解释 Physiological and ecological explanation
光合生理性状 Photosynthetic physiological trait	最大净光合速率 Maximum net photosynthetic rate	A_max	μmol·m^-2·s^-1	光合潜力与资源获取策略 Photosynthetic potential and resource acquisition strategies
	最大羧化速率 Maximum carboxylation rate	V_cmax	μmol·m^-2·s^-1	核酮糖-1,5-双磷酸羧化酶/加氧酶(Rubisco)催化的最大羧化反应速率 Maximum carboxylation rate of the enzyme ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco)
	最大电子传递速率 Maximum electron transfer rate	J_max	μmol·m^-2·s^-1	支持核酮糖双磷酸再生的最大电子传输速率 Maximum electron transfer rate for supporting ribulose bisphosphate regeneration
	暗呼吸速率 Dark respiration rate	R_d	μmol·m^-2·s^-1	黑暗环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under dark conditions
	光下暗呼吸速率 Dark respiration rate under light	R_l	μmol·m^-2·s^-1	光照环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under light conditions
	光饱和碳同化速率 Photo-saturated carbon assimilation rate	A_sat	μmol CO₂·m^-2·s^-1	碳同化效率 Carbon assimilation efficiency
	气孔导度 Stomatal conductance	g_s	mol H₂O·m⁻²·s⁻¹		气体交换与资源获取 Gas exchange and resource acquisition
	蒸腾速率 Transpiration rate	E	mmol H₂O·m⁻²·s⁻¹		耗水特性 Water consumption characteristics
	水分利用效率 Water use efficiency	WUE	μmol·mmol^-1		抗旱性与水资源利用 Drought resistance and water resource utilization
	氮利用效率 Nitrogen use efficiency	NUE	μmol·g^-1·s^-1		叶片养分利用、生理特性和生存策略 Nutrient utilization, physiological characteristics, and survival strategies of leaves
	最大光化学效率 Maximal photochemical efficiency	F_v/F_m			光系统II (PSII)反应中心的最大光能转化效率以及叶片的健康程度 The maximum light energy conversion efficiency of photosystem II (PSII) and the health status of leaves
	光化学淬灭 Photochemical quenching	q_P			PSII反应中心的开放程度以及光化学活性的高低 Degree of openness of PSII and the level of photochemical activity
	非光化学淬灭 Non-photochemical quenching	NPQ			植物的光保护能力 Light protection ability of plants
	电子传递速率 Electron transfer rate	ETR	μmol·m^-2·s^-1		植物的耐光性和适应性 Light resistance and adaptability of plants
结构性状 Structural trait	叶厚度 Leaf thickness	L_t	mm		生存策略以及光合呼吸、蒸腾相关的碳成本 Survival strategy and carbon costs related to photosynthesis, respiration and transpiration
	比叶质量 Leaf mass per area	LMA	g·m^-2		资源获取和抗逆性 Resource acquisition and resilience
	叶组织密度 Leaf tissue density	LTD	g·cm^-3		叶片韧性和抗逆性 Leaf toughness and stress resistance
	叶片相对含水量 Leaf relative water content	LRWC	%		植物的水分状况、抗旱性及环境适应 Water status, drought resistance and environmental adaptation
	叶干物质含量 Leaf dry matter content	LDMC	g·g^-1		抗逆性和对抗物理危害的防御能力 Resistance to adversity and defense against physical hazards
生物化学性状 Biochemical trait	基于面积的叶氮含量 Leaf nitrogen content per unit area	N_area	g·m^-2		资源获取和衡量光合能力 Resource acquisition and characterization of photosynthetic capacity
	基于面积的叶碳含量 Leaf carbon content per unit area	C_area	g·m^-2		构建和防御 Building and defense
	碳氮比 Ratio of leaf carbon content to nitrogen content	C:N			元素分配与适应策略 Element allocation and adaptation strategies
	叶绿素含量 Chlorophyll content	Chl	μg·cm^-2		健康程度与光合作用 Health level and photosynthesis
	叶绿素a/b Ratio of chlorophyll a to chlorophyll b	Chl a/b			光能吸收和能量运输 Light absorption and energy transport
	类胡萝卜素含量 Carotenoid content	Car	μg·cm^-2		光合作用与光保护必需色素 Photosynthesis and essential pigments for photoprotection

叶性状 Leaf trait	主成分1 Principal component 1	主成分2 Principal component 2
V_cmax	-0.22	0.03
J_max	-0.24	0.04
A_max	-0.24	-0.14
R_d	-0.23	0.14
R_l	-0.23	0.13
A_sat	-0.26	-0.01
g_s	-0.24	0.07
E	-0.25	0.10
WUE	0.11	-0.24
NUE	-0.22	-0.16
q_P	-0.21	-0.12
F_v/F_m	-0.22	-0.02
ETR	-0.25	-0.04
NPQ	-0.06	-0.04
N_area	-0.20	0.22
C_area	0.01	0.35
C:N	0.20	0.02
Chl	-0.07	-0.34
Chl a/b	-0.23	0.07
Car	-0.09	-0.31
LMA	-0.05	0.36
LDMC	0.22	0.19
LRWC	-0.23	-0.18
L_t	0.09	-0.28
LTD	-0.10	0.38
特征值 Characteristic value	3.77	2.34
方差比例 Variance ratio	54.75%	21.04%
累计方差比例 Accumulated variance ratio	54.75%	75.79%

季节尺度毛乌素沙地黑沙蒿的叶片性状网络特征及驱动因素

Characteristics of seasonal leaf trait network and its drivers in Artemisia ordosica in the Mau Us Sandy Land

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