降水梯度带榆树枝叶协作关系的区域分异规律

doi:10.17521/cjpe.2024.0050

植物生态学报 ›› 2025, Vol. 49 ›› Issue (2): 282-294.DOI: 10.17521/cjpe.2024.0050 cstr: 32100.14.cjpe.2024.0050

降水梯度带榆树枝叶协作关系的区域分异规律

李姝雯¹, 汤璐瑶¹, 张博纳¹, 叶琳峰¹, 童金莲¹, 谢江波¹^,², 李彦¹^,², 王忠媛¹^,²^,^*()

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, 浙江农林大学林业与生物技术学院, 杭州 311300
²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:2024-02-19 接受日期:2024-05-27 出版日期:2025-02-20 发布日期:2025-02-20
通讯作者: ^*王忠媛: (wangzhongyuan2014@163.com)
基金资助:
国家自然科学基金(32371662);国家自然科学基金(42330503);浙江省科技厅重大专项(2022C02019)

Regional differentiation of cooperative relationships between Ulmus pumila branches and leaves along precipitation gradients

LI Shu-Wen¹, TANG Lu-Yao¹, ZHANG Bo-Na¹, YE Lin-Feng¹, TONG Jin-Lian¹, XIE Jiang-Bo¹^,², LI Yan¹^,², WANG Zhong-Yuan¹^,²^,^*()

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

Received:2024-02-19 Accepted:2024-05-27 Online:2025-02-20 Published:2025-02-20
Supported by:
National Natural Science Foundation of China(32371662);National Natural Science Foundation of China(42330503);Major Special Project of Department of Science and Technology of Zhejiang Provincial(2022C02019)

摘要/Abstract

摘要： 伴随着降水特征变化(如干旱、干季延长或干湿交替加剧), 与之相耦合的植物功能性状也将随之发生变异, 继而引发植物功能性状的协作关系(单个器官内或多个器官间)发生相应调整, 以此为基础的植物行为和适应策略随之改变。但对这一过程背后的数量关系和作用机制仍然不清楚。以功能性状为切入点, 沿降水梯度带跨区域原位测量共有种对气候环境的特异性反应, 量化这些特异性反应背后的性状-环境关系, 阐明其调控机制, 揭示共有种功能性状及其适应策略的区域分异规律, 将为气候治理提供数据支撑和坚实的科学基础。该研究在中国东南至西北的降水梯度带上选取10个样地, 以样带共有种榆树(Ulmus pumila)为实验对象, 测量其枝和叶共28个功能性状。分析了榆树枝和叶性状以及性状间权衡关系的区域分异规律, 进一步量化不同器官(枝和叶)间功能性状协作关系沿降水梯度带的区域分异, 揭示了榆树对不同水分环境的适应策略。结果表明: (1)在湿润区, 榆树枝条具有最大的输水效率(K_s)和最小的栓塞抗性(P₅₀); 随降水量减少, 叶片厚度、叶组织紧密度增加, 榆树的抗旱能力增强。(2)在整个降水梯度带上, 榆树同一器官(枝)内及不同器官(枝和叶)间均存在效率-安全权衡; 但在区域尺度上, 这种权衡关系随降水量的减少而解耦。(3)枝和叶功能性状相关分析表明: 在整个降水梯度带上, 最大净光合速率(P_n)和比叶质量(LMA)均与K_s负相关, 与P₅₀正相关。榆树通过枝水分运输能力和叶功能性状(叶片厚度和气孔打开比率)协同调控光合能力, 枝和叶功能性状的调整与协作是榆树适应不同水分环境的重要机制。

关键词: 功能性状, 输水效率, 栓塞抗性, 权衡, 适应策略

Abstract:

Aims Changes in precipitation characteristics, such as drought, prolonged dry season, and increased dry-wet alternation, lead to variations in plant functional traits. These changes trigger adjustments in the cooperative relationship of plant functional traits within a single organ or between multiple organs. Consequently, plant behavior and adaptation strategies change accordingly. However, the quantitative relationships and mechanisms behind this process are still unclear. This study aims to measure the specific responses of common species to climate across regions along a precipitation gradient, quantify the trait-environment relationship, elucidate the regulatory mechanism, and reveal the regional differentiation of functional traits and adaptation strategies of common species. This study will provide data support and solid scientific basis for climate management.

Methods The study focused on Ulmus pumila as the experimental subject. Ten sites were selected along a precipitation gradient from southeast to northwest China, where 28 functional traits of branches and leaves were measured. We analyzed the regional differentiation of branch and leaf traits, as well as their trade-offs. Furthermore, we quantified the regional differentiation of collaborative relationships among functional traits of branches and leaves along the precipitation gradient, revealing the adaptation strategies of U. pumila to varying moisture environments.

Important findings The results showed that: (1) In humid regions, U. pumila branches exhibited the highest hydraulic conductivity (K_s) and the lowest cavitation resistance (P₅₀); as precipitation decreased, leaf thickness and leaf tissue structure tightness increased, enhancing U. pumila’s drought resistance. (2) Across the entire precipitation gradient, there was an efficiency-safety trade-off within branches and between branches and leaves of U. pumila; however, at the regional scale, this trade-off relationship decoupled with decreasing precipitation. (3) Correlation analyses of branch and leaf functional traits revealed that, across the entire precipitation gradient, maximum net photosynthetic rate (P_n) and leaf mass per unit area were negatively correlated with K_s and positively correlated with P₅₀. Ulmus pumila regulated photosynthesis through coordinated adjustments of branch water transport capacity and leaf functional traits. The coordination and adjustment of branch and leaf functional traits are crucial mechanisms for U. pumila to adapt to varying moisture environments.

Key words: functional traits, hydraulic efficiency, cavitation resistance, trade-off, adaptive strategy

李姝雯, 汤璐瑶, 张博纳, 叶琳峰, 童金莲, 谢江波, 李彦, 王忠媛. 降水梯度带榆树枝叶协作关系的区域分异规律. 植物生态学报, 2025, 49(2): 282-294. DOI: 10.17521/cjpe.2024.0050

LI Shu-Wen, TANG Lu-Yao, ZHANG Bo-Na, YE Lin-Feng, TONG Jin-Lian, XIE Jiang-Bo, LI Yan, WANG Zhong-Yuan. Regional differentiation of cooperative relationships between Ulmus pumila branches and leaves along precipitation gradients. Chinese Journal of Plant Ecology, 2025, 49(2): 282-294. DOI: 10.17521/cjpe.2024.0050

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0050

https://www.plant-ecology.com/CN/Y2025/V49/I2/282

图/表 7

表1 降水梯度带上榆树各样点基本特征

Table 1 Basic characteristics of each sample site of Ulmus pumila along the precipitation gradient zone

样点 Site	缩写 Abbreviation	经度 Longitude (° E)	纬度 Latitude (°N)	海拔 Altitude (m)	年平均气温 Mean annual air temperature (℃)	平均年降水量 Mean annual precipitation (mm)
浙江天目山 Tianmushan, Zhejiang	TMS	119.44	30.34	1 142.00	15.30	1 390.00
河南鸡公山 Jigongshan, Henan	JGS	114.09	31.87	260.00	15.20	1 118.00
河南宝天曼 Baotianman, Henan	BTM	111.93	33.52	1 426.50	14.00	823.30
河南济源 Jiyuan, Henan	JY	112.50	35.05	384.75	9.60	614.50
陕西安塞 Ansai, Shaanxi	AS	109.32	36.86	1 218.00	7.90	534.00
宁夏固原 Guyuan, Ningxia	GY	106.40	35.97	1 831.00	6.80	495.70
宁夏沙坡头 Shapotou, Ningxia	SPT	104.99	37.46	1 247.00	6.60	186.00
甘肃民勤 Minqin, Gansu	MQ	103.14	38.66	1 376.00	7.90	113.20
甘肃瓜州 Guazhou, Gansu	GZ	95.78	40.50	1 139.00	7.60	54.40
新疆阜康 Fukang, Xinjiang	FK	87.93	44.29	465.00	6.00	158.80

表2 降水梯度带上榆树相关性状、缩写及单位

Table 2 Relevant traits, abbreviations, and units of Ulmus pumila along the precipitation gradient zone

性状 Trait		缩写 Abbreviation	单位 Unit
叶性状 Leaf trait	最大净光合速率 Maximum net photosynthetic rate	P_n¹⁾	µmol·m^-2·s^-1
	单位质量最大净光合速率 Maximum net photosynthetic rate per unit leaf mass	A_mass¹⁾	µmol·g^-1·s^-1
	可操作气孔导度 Maximized operational stomatal conductance	G_op¹⁾	mol·m^-2·s^-1
	解剖学最大气孔导度 Anatomical maximum stomatal conductanc	G_smax¹⁾	mol·m^-2·s^-1
	气孔打开比率 Stomatal opening ratio during gas exchange at light saturated conditions	G_ratio¹⁾	%
	水分利用效率 Water use efficiency	WUE ¹⁾	%
	膨压损失点叶水势 Turgor pressure loss point leaf water potential	Ψ_tlp¹⁾	MPa
	气孔大小 Stomatal size	S_s¹⁾	µm²
	气孔密度 Stomatal density	S_d¹⁾	pore·mm^-2
	气孔面积分数 Stomatal area fraction	S_f¹⁾	%
	比叶质量 Leaf mass per area	LMA ¹⁾	g·m^-2
	叶脉密度 Vein density	D_v¹⁾	mm·mm^-2
	叶片厚度 Leaf thickness	LT	µm
	上表皮厚度 Upper tissue thickness	UET	µm
	下表皮厚度 Lower tissue thickness	LET	µm
	栅栏组织厚度 Palisade tissue thickness	PMT	µm
	海绵组织厚度 Sponge tissue thickness	SMT	µm
	栅栏组织厚度/海绵组织厚度 Ratio of palisade tissue thickness to sponge tissue thickness	PMT/SMT
	组织紧密度 Leaf tissue structure tightness	CTR
	组织疏松度 Looseness of leaf tissue structure	SR
枝性状 Branch trait	胡伯尔值 Huber value	H_v¹⁾	10³ mm²·cm^-2
	导管直径 Vessel diameter	D	µm
	导管密度 Vessel density	N_v	pore·µm^-2
	导管壁厚 Double thickness of vessel wall	T_w	µm
	导管厚度跨度比 Thickness-to-span ratio	T_tob	10³
	木质部密度 Wood density	WD	g·cm^-3
	比导率 Specific hydraulic conductivity	K_s	kg·m^-1·s^-1·MPa^-1
	导水损失率为50%时水势 Water potential causing 50% loss of conductivity	P₅₀	MPa

表2 降水梯度带上榆树相关性状、缩写及单位

Table 2 Relevant traits, abbreviations, and units of Ulmus pumila along the precipitation gradient zone

性状 Trait		缩写 Abbreviation	单位 Unit
叶性状 Leaf trait	最大净光合速率 Maximum net photosynthetic rate	P_n¹⁾	µmol·m^-2·s^-1
	单位质量最大净光合速率 Maximum net photosynthetic rate per unit leaf mass	A_mass¹⁾	µmol·g^-1·s^-1
	可操作气孔导度 Maximized operational stomatal conductance	G_op¹⁾	mol·m^-2·s^-1
	解剖学最大气孔导度 Anatomical maximum stomatal conductanc	G_smax¹⁾	mol·m^-2·s^-1
	气孔打开比率 Stomatal opening ratio during gas exchange at light saturated conditions	G_ratio¹⁾	%
	水分利用效率 Water use efficiency	WUE ¹⁾	%
	膨压损失点叶水势 Turgor pressure loss point leaf water potential	Ψ_tlp¹⁾	MPa
	气孔大小 Stomatal size	S_s¹⁾	µm²
	气孔密度 Stomatal density	S_d¹⁾	pore·mm^-2
	气孔面积分数 Stomatal area fraction	S_f¹⁾	%
	比叶质量 Leaf mass per area	LMA ¹⁾	g·m^-2
	叶脉密度 Vein density	D_v¹⁾	mm·mm^-2
	叶片厚度 Leaf thickness	LT	µm
	上表皮厚度 Upper tissue thickness	UET	µm
	下表皮厚度 Lower tissue thickness	LET	µm
	栅栏组织厚度 Palisade tissue thickness	PMT	µm
	海绵组织厚度 Sponge tissue thickness	SMT	µm
	栅栏组织厚度/海绵组织厚度 Ratio of palisade tissue thickness to sponge tissue thickness	PMT/SMT
	组织紧密度 Leaf tissue structure tightness	CTR
	组织疏松度 Looseness of leaf tissue structure	SR
枝性状 Branch trait	胡伯尔值 Huber value	H_v¹⁾	10³ mm²·cm^-2
	导管直径 Vessel diameter	D	µm
	导管密度 Vessel density	N_v	pore·µm^-2
	导管壁厚 Double thickness of vessel wall	T_w	µm
	导管厚度跨度比 Thickness-to-span ratio	T_tob	10³
	木质部密度 Wood density	WD	g·cm^-3
	比导率 Specific hydraulic conductivity	K_s	kg·m^-1·s^-1·MPa^-1
	导水损失率为50%时水势 Water potential causing 50% loss of conductivity	P₅₀	MPa

图1 榆树枝叶性状沿降水梯度的回归分析(平均值±标准误)。各研究点缩写见表1; 性状缩写见表2。NS, 无相关关系。

Fig. 1 Regression analysis of branches and leaves traits of Ulmus pumila along the precipitation gradient (mean ± SE). Abbreviations of each research site are shown in Table 1, and abbreviations of traits are shown in Table 2. NS, no significant correlation.

图2 榆树枝叶性状在不同干湿区的变化(平均值±标准误)。不同大写字母表示不同区域间差异显著(p < 0.05)。其中, 叶水力、光合、气孔等13个性状在不同干湿区的变化已在发表文章(Xie et al., 2022, Fig. 3)中展示。

Fig. 2 Changes of branches and leaves traits of Ulmus pumila across different wet and dry regions (mean ± SE). Different uppercase letters indicated significant difference in different regions (p < 0.05). Among them, the changes of 13 traits such as leaf hydraulics, photosynthesis, and stomata from humid to arid areas have been shown in a published article (Xie et al., 2022, Fig. 3). Abbreviations of traits are shown in Table 2.

图3 榆树功能性状的变异系数(A)、嵌套方差分析(B)、主成分(PC)分析(C)。主成分分析的箱线图说明不同区域间的差异, 不同大写字母表示不同区域间差异显著(p < 0.05)。各研究性状的缩写见表2。

Fig. 3 Coefficient of variation (A), variance components of traits based on nested ANOVAs (B) and principal component (PC) analysis (C) of traits in Ulmus pumila. Box plots from PC analysis illustrate the differences in different regions, different uppercase letters indicated significant difference in different regions (p < 0.05). Abbreviations of traits are shown in Table 2.

图4 样带整体及不同干湿区榆树枝叶互作网络。各研究性状的缩写见表2。

Fig. 4 Transect and the interaction network of branches and leaves across a gradient from humid to arid regions of Ulmus pumila. Abbreviations of the analyzed traits are shown in Table 2.

图5 榆树枝叶性状间的关系(平均值±标准误)。不同区域的拟合采用所有数据散点, 各研究点的缩写见表1, 各研究性状的缩写见表2。NS, 无相关关系。

Fig. 5 Relationship between branches and leaves traits of Ulmus pumila (mean ± SE). Data from all sites were used to model variations across different regions. The abbreviations of the studied sites are shown in Table 1, and the abbreviations of the studied traits are shown in Table 2. NS, no significant correlation.

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降水梯度带榆树枝叶协作关系的区域分异规律

Regional differentiation of cooperative relationships between Ulmus pumila branches and leaves along precipitation gradients

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