基于叶片和土壤酶化学计量的川西亚高山岷江冷杉林养分限制海拔变化规律

doi:10.17521/cjpe.2022.0449

植物生态学报 ›› 2023, Vol. 47 ›› Issue (12): 1646-1657.DOI: 10.17521/cjpe.2022.0449 cstr: 32100.14.cjpe.2022.0449

所属专题：生态化学计量；生物地球化学；微生物生态学

基于叶片和土壤酶化学计量的川西亚高山岷江冷杉林养分限制海拔变化规律

何茜¹^,², 冯秋红³, 张佩佩¹^,^*(), 杨涵¹^,², 邓少军¹^,², 孙小平⁴, 尹华军¹

¹中国科学院成都生物研究所, 中国-克罗地亚生物多样性和生态系统服务“一带一路”联合实验室, 中国科学院山地生态恢复与生物资源利用重点实验室, 生态恢复与生物多样性保育四川省重点实验室, 成都 610041
²中国科学院大学, 北京 100049
³四川省林业科学研究院, 四川卧龙森林生态系统定位站, 成都 610081
⁴阿坝州川西国有林保护局, 四川阿坝 624000

收稿日期:2022-11-08 接受日期:2023-03-13 出版日期:2023-12-20 发布日期:2023-03-13
基金资助:
中国科学院“西部之光”交叉团队项目(xbzg-zdsys-202112);第二次青藏高原综合科学考察项目(2019QZKK0301);国家自然科学基金(32171757);国家自然科学基金(32201531);中国博士后科学基金(2021M703133)

Altitudinal patterns of nutrient limiting characteristics of Abies fargesii var. faxoniana forest based on leaf and soil enzyme stoichiometry in western Sichuan, China

HE Xi¹^,², FENG Qiu-Hong³, ZHANG Pei-Pei¹^,^*(), YANG Han¹^,², DENG Shao-Jun¹^,², SUN Xiao-Ping⁴, YIN Hua-Jun¹

¹China-Croatia “Belt and Road” Joint Laboratory on Biodiversity and Ecosystem Services, CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China
²University of Chinese Academy of Sciences, Beijing 100049, China
³Sichuan Academy of Forestry, Sichuan Wolong Forest Ecosystem Research Station, Chengdu 610081, China
⁴Aba State Owned Forest Protection Bureau, Aba, Sichuan 624000, China

Received:2022-11-08 Accepted:2023-03-13 Online:2023-12-20 Published:2023-03-13
Supported by:
Chinese Academy of Sciences “Light of the West” Cross-team(xbzg-zdsys-202112);Second Tibetan Plateau Scientific Expedition and Research Program (STEP)(2019QZKK0301);National Natural Science Foundation of China(32171757);National Natural Science Foundation of China(32201531);China Postdoctoral Science Foundation(2021M703133)

摘要/Abstract

摘要：

氮(N)和磷(P)养分有效性是制约森林生态系统林分生产力与碳汇功能的关键要素, 但目前对多变环境下森林生态系统养分限制特征还缺乏充分的科学认识。山地生态系统的气候、植被和土壤等环境因子沿海拔的垂直变化格局为深入认识森林养分限制及其驱动因素提供了天然的实验平台。该研究以青藏高原东缘典型的川西亚高山针叶林——岷江冷杉(Abies faxoniana)林为研究对象, 通过沿巴朗山2 850-3 200 m海拔梯度的多点取样, 从植物叶片N、P养分含量、化学计量变化和地下微生物胞外酶化学计量学的角度, 分析了海拔梯度下该区域森林养分限制特征变化规律及其主要驱动因素。结果表明: 1)随海拔升高, 叶片N、P含量降低而N:P由12.33升高至15.00, 表明随海拔升高该区域针叶林叶片生长由N限制转化为N-P共同限制, 且P限制随海拔升高而表现出增强趋势; 2)矢量模型分析发现不同海拔下根际土壤微生物胞外酶化学计量矢量角度均>45°, 且随海拔升高呈上升趋势, 表明该区域微生物受P限制, 且海拔越高土壤微生物P限制越强; 3)进一步通过Pearson相关分析和路径分析表明, 海拔引起的气温变化是驱动岷江冷杉林生态系统养分限制的主导因素。综上所述, 基于叶片和土壤微生物养分证据均表明川西亚高山针叶林生态系统总体表现出P限制程度随海拔升高而逐渐增强的趋势。

关键词: 海拔, 养分限制, N:P, 土壤胞外酶化学计量, 亚高山针叶林

Abstract:

Aims Nitrogen (N) and phosphorus (P) nutrient availability is a key factor governing forest productivity and carbon sequestration. However, scientific knowledge on the nutrient limitation in forest ecosystems under variable environments is still lacked. The mountain ecosystems, characterized by the dramatical changes in multiple environmental factors along increasing altitude such as climate, vegetation and soil properties, provide a natural experiment platform for understanding forest nutrient limitation and its drivers.
Methods In this study, we examined the nutrient limitation of a typical subalpine coniferous forest (Abies fargesii var. faxoniana forest) along an altitudinal gradient (from 2 850 m to 3 200 m) in the southeastern Qingzang Plateau, by simultaneous detection of above-ground leaf N, P status and underground microorganisms extracellular stoichiometry, and analyzing the changes of forest nutrient limitation and the main driving factors along the altitude.
Important findings The results showed that: 1) as altitude increases, the concentration of leaf N and P decreased, while leaf N:P increased from 12.33 to 15.00, indicating a shift from N limitation to N-P co-limitation and an enhancement of P limitation with increasing altitude. (2) Vector model analysis showed that the vector angles of microbial extracellular enzyme stoichiometry were all exceed 45° at different altitudes, and as altitude increases, the vector angle showed an increasing trend, indicating that microorganisms were limited by P and the P limitation increases with altitude. (3) Temperature is the dominant factor driving nutrient limitation of Abies fargesii var. faxoniana forest. Collectively, both leaf and soil microbial nutrient evidence indicated that an enhancement of P limitation in subalpine coniferous forests with increasing altitudes in western Sichuan. This finding could provide an important theoretical basis for guiding forest nutrient adaptive management in subalpine coniferous ecosystems under the scenarios of global climate change.

Key words: altitude, nutrient limitation, N:P, soil extracellular enzyme stoichiometry, subalpine coniferous forest

何茜, 冯秋红, 张佩佩, 杨涵, 邓少军, 孙小平, 尹华军. 基于叶片和土壤酶化学计量的川西亚高山岷江冷杉林养分限制海拔变化规律. 植物生态学报, 2023, 47(12): 1646-1657. DOI: 10.17521/cjpe.2022.0449

HE Xi, FENG Qiu-Hong, ZHANG Pei-Pei, YANG Han, DENG Shao-Jun, SUN Xiao-Ping, YIN Hua-Jun. Altitudinal patterns of nutrient limiting characteristics of Abies fargesii var. faxoniana forest based on leaf and soil enzyme stoichiometry in western Sichuan, China. Chinese Journal of Plant Ecology, 2023, 47(12): 1646-1657. DOI: 10.17521/cjpe.2022.0449

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0449

https://www.plant-ecology.com/CN/Y2023/V47/I12/1646

图/表 8

图1 川西亚高山研究区的垂直带谱和采样点分布。

Fig. 1 Vertical zonation and sampling point in the study area in subalpine mountains of western Sichuan.

表1 川西亚高山岷江冷杉林不同海拔土壤含水量、pH及气温(平均值±标准误, n = 5)。

Table 1 Soil moisture, pH and air temperature at different altitudes of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan (mean ± SE, n = 5)

基本性质 Basic property	海拔 Altitude (m)
基本性质 Basic property	2 850	2 950	3 060	3 200
土壤含水量 Soil moisture (%)	46.89 ± 0.36^d	54.80 ± 2.31^b	50.79 ± 0.94^c	57.51 ± 2.26^a
土壤pH Soil pH	4.55 ± 0.02^a	4.15 ± 0.14^b	4.20 ± 0.05^b	3.92 ± 0.05^c
气温 Air temperature (°C)	15.33	13.99	12.51	10.64

表2 川西亚高山岷江冷杉林不同海拔下土壤养分含量(平均值±标准误, n = 5)。

Table 2 Concentration of soil nutrients at different altitudes of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan (mean ± SE, n = 5)

土壤基本理化性质 Basic property	海拔 Altitudes (m)
土壤基本理化性质 Basic property	2 850	2 950	3 060	3200
TN (g·kg^-1)	5.23 ± 0.06^c	6.23 ± 0.30^b	5.36 ± 0.19^c	8.61 ± 0.99^a
TP (g·kg^-1)	0.53 ± 0.02^b	0.56 ± 0.05^b	0.54 ± 0.03^b	0.65 ± 0.02^a
DIN (mg·kg^-1)	17.11 ± 0.68^a	14.72 ± 1.68^b	10.14 ± 1.01^d	11.85 ± 0.83^c
AP (mg·kg^-1)	1.65 ± 0.25^b	4.74 ± 0.75^a	1.48 ± 0.11^b	4.01 ± 0.91^a

图2 川西亚高山岷江冷杉林土壤总氮(TN)、总磷(TP)、可溶性无机氮(DIN)、有效磷(AP)含量及N:P与海拔的关系。实线表示土壤养分与海拔之间的线性拟合关系, 灰色区域为模型的95%置信区间。

Fig. 2 Relationships of soil total nitrogen (TN), total phosphorus (TP), dissolved inorganic nitrogen (DIN), available phosphorus (AP) contents and N:P with altitudes of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan. Solid lines indicate the linear fitting relationship between the soil nutrient and altitudes, and grey areas are the 95% confidence intervals of the models.

图3 川西亚高山岷江冷杉林叶片总氮(TN)、总磷(TP)含量及N:P与海拔的关系。实线表示叶片养分含量与海拔之间的线性拟合关系, 灰色区域为模型的95%置信区间。虚线代表叶片N:P为14和16。

Fig. 3 Relationships of foliar total nitrogen (TN), total phosphorus (TP) contents and N:P of coniferous forest with altitudes of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan. Solid lines indicate the model fits between the leaf nutrient contents and altitudes, and grey areas are the 95% confidence intervals of the models. The dashed lines represent leaf N:P of 14 and 16, respectively.

图4 川西亚高山岷江冷杉林不同海拔下土壤酶化学计量比与微生物资源限制特征(平均值±标准误, n = 5)。H、I中不同小写字母表示不同海拔间的差异显著(p < 0.05)。A-C中实线表示土壤酶化学计量比与海拔之间的线性拟合关系, 灰色区域为模型的95%置信区间。对角虚线代表1:1线, 水平虚线代表矢量角度为45°。ACP, 酸性磷酸酶活性; BG, β-1,4-葡萄糖苷酶活性; LAP, 亮氨酸氨基肽酶活性; NAG, β-1,4-N-乙酰氨基葡萄糖苷酶活性。

Fig. 4 Soil enzymatic stoichiometry and microbial resource limitation at different altitudes of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan (mean ± SE, n = 5). Different lowercase letters in H and I indicate significant differences between different altitudes (p < 0.05). In A-C, solid lines indicate the model fits between soil nutrient and altitude, and grey areas are the 95% confidence intervals of the models. The diagonal dashed line represents the 1:1 line, the horizontal dashed line represents the vector angle of 45°. C, carbon; N, nitrogen; P, phosphorus. ACP, 4-MUB-phosphate activity; BG, 4-MUB-β-D-glucoside activity; LAP, L-leucine-7-amido-4-methylcoumarin activity; NAG, 4-MUB-N-acetyl-β-D-glucosaminide activity.

图5 川西亚高山岷江冷杉林根际土壤总养分含量及其化学计量比、有效养分含量、物理特性和气候因子与植物养分限制和微生物限制相关指标的Pearson相关矩阵。AP, 有效磷含量; C:N, 土壤总碳氮比; C:P, 土壤总碳磷比; DIN, 可溶性无机氮含量; DOC, 可溶性有机碳含量; pH, 土壤pH; LN:P, 叶片氮磷比; N:P, 土壤总氮磷比; SM, 土壤水分含量; SOC, 土壤有机碳含量; T, 气温; TN, 总氮含量; TP, 总磷含量; VA, 矢量角度。*, p < 0.05; **, p < 0.01; ***, p < 0.001; n = 5。

Fig. 5 Pearson correlation matrix between rhizosphere soil total nutrients content and its stoichiometry characteristics, available nutrients, physical properties, climate and plant nutrient limitation and microbial limitation of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan. AP, available phosphorus content; C:N, soil total carbon to nitrogen ratio; C:P, soil total carbon to phosphorus ratio; DIN, dissolved inorganic nitrogen content; DOC, dissolved organic carbon content; pH, soil pH; LN:P, leaf total nitrogen to phosphorus ratio; N:P, soil total nitrogen to phosphorus ratio; SM, soil moisture; SOC, soil organic carbon content; T, air temperature; TN, total nitrogen content; TP, total phosphorus content; VA, vector angle. *, p < 0.05; **, p < 0.01; ***, p < 0.001; n = 5.

图6 川西亚高山岷江冷杉林养分限制与气温、土壤理化特性关系的偏最小二乘路径模型(A)和各因子对森林养分限制的效应值(B、C)。实线和虚线箭头分别表示因果关系的正向和负向影响(p < 0.05), 箭头上的数字表示标准化路径系数, R2表示模型解释的因变量的方差。C:N, 土壤总碳氮比; C:P, 土壤总碳磷比; DIN, 可溶性无机氮含量; pH, 土壤pH; LN:P, 叶片氮磷比; N:P, 土壤总氮磷比; SOC, 土壤有机碳含量; T, 气温; TN, 土壤总氮含量; VA, 矢量角度。

Fig. 6 Relationship between forest nutrient limitation and soil physicochemical properties partial least squares path modelling (A) and effect value of each factor on forest nutrient limitation (B, C) of Abies fargesii var. faxoniana forest in subalpine mountains of western Sichuan. Solid and dotted line arrows indicate positive and negative flows of causality (p < 0.05), respectively. Numbers on the arrow indicate significant standardized path coefficients. R2 indicates the variance of dependent variable explained by the model. C:N, soil total carbon to nitrogen ratio; C:P, soil total carbon to phosphorus ratio; DIN, dissolved inorganic nitrogen content; pH, soil pH; LN:P, leaf total nitrogen to phosphorus ratio; N:P, soil total nitrogen to phosphorus ratio; SOC, soil organic carbon content; T, air temperature; TN, soil total nitrogen content; VA, vector angle.

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基于叶片和土壤酶化学计量的川西亚高山岷江冷杉林养分限制海拔变化规律

Altitudinal patterns of nutrient limiting characteristics of Abies fargesii var. faxoniana forest based on leaf and soil enzyme stoichiometry in western Sichuan, China

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