灌丛斑块和草本斑块碳交换对季节性降水增加的响应——基于地上净初级生产力和叶面积指数标准化的比较分析

doi:10.17521/cjpe.2023.0359

植物生态学报 ›› 2024, Vol. 48 ›› Issue (8): 1035-1049.DOI: 10.17521/cjpe.2023.0359 cstr: 32100.14.cjpe.2023.0359

所属专题：碳循环

灌丛斑块和草本斑块碳交换对季节性降水增加的响应——基于地上净初级生产力和叶面积指数标准化的比较分析

张梦迪¹^,²^,^*, 向官海²^,³^,^*, 文艺瑶²^,⁵, 王欢²^,⁶, 呼格吉勒⁷, 白永飞²^,⁴, 王忠武¹^,^**(), 郑淑霞²^,⁴^,^**()()

¹内蒙古农业大学草原与资源环境学院, 呼和浩特 010018
²中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093
³赤峰学院农学院, 内蒙古赤峰 024000
⁴国家植物园, 北京 100093
⁵江西师范大学生命科学学院, 南昌 330022
⁶山东农业大学生命科学学院, 山东泰安 271001
⁷正蓝旗草原工作站, 锡林浩特 027200

收稿日期:2023-12-04 接受日期:2024-05-10 出版日期:2024-08-20 发布日期:2024-05-10
通讯作者: **王忠武(wangzhongwu@imau.edu.cn);郑淑霞(zsx@ibcas.ac.cn), ORCID:0000-0001-6818-3796
作者简介:*同等贡献
基金资助:
国家重点研发计划(2023YFF1305301);国家自然科学基金(32371779)

Response of carbon exchange between shrub and grass patches to increased seasonal precipitation: a comparative analysis based on aboveground net primary productivity and leaf area index standardization

ZHANG Meng-Di¹^,²^,^*, XIANG Guan-Hai²^,³^,^*, WEN Yi-Yao²^,⁵, WANG Huan²^,⁶, Hugejile ⁷, BAI Yong-Fei²^,⁴, WANG Zhong-Wu¹^,^**(), ZHENG Shu-Xia²^,⁴^,^**()()

¹College of Grassland, Resources and Environment, Inner Mongolia Agricultural University, Hohhot 010018, China
²State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
³College of Agriculture, Chifeng University, Chifeng, Nei Mongol 024000, China
⁴China National Botanical Garden, Beijing 100093, China
⁵College of Life Sciences, Jiangxi Normal University, Nanchang 330022, China
⁶College of Life Sciences, Shandong Agricultural University, Tai’an, Shandong 271001, China
⁷Grassland Workstation of Zhenglan Banner, Xilinhot, Nei Mongol 027200, China

Received:2023-12-04 Accepted:2024-05-10 Online:2024-08-20 Published:2024-05-10
Contact: WANG Zhong-Wu(wangzhongwu@imau.edu.cn), ZHENG Shu-Xia(zsx@ibcas.ac.cn), ORCID:0000-0001-6818-3796
About author:*Contributed equally to this work
Supported by:
National Key R&D Program of China(2023YFF1305301);National Natural Science Foundation of China(32371779)

摘要/Abstract

摘要：

随着气候变化和人类活动干扰的加剧, 干旱半干旱地区草地灌丛化现象普遍发生, 严重影响了草地生态系统的碳汇功能。水分是内蒙古半干旱草原的主要限制因子, 未来降水格局的变化对草原生态系统的碳交换具有重要影响。然而, 目前关于降水变化对灌丛化草地生态系统, 特别是异质斑块碳交换过程的影响研究较少, 相关机制尚不清楚。为此, 该研究利用内蒙古小叶锦鸡儿(Caragana microphylla)灌丛化草地季节性降水增加(冬季增雪、夏季增雨)实验平台, 系统观测了灌丛斑块和草本斑块的碳交换参数, 即净生态系统碳交换(NEE)、总生态系统生产力(GEP)、生态系统呼吸(ER), 并结合基于地上净初级生产力(ANPP)和叶面积指数(LAI)标准化的参数比较分析, 研究了季节性降水增加对灌丛化草地碳交换的影响以及异质斑块的响应差异。结果表明: 1)夏季增雨显著提高了草本斑块|NEE|、GEP和ER值, 而冬季增雪显著降低了草本斑块|NEE|_ANPP、GEP_ANPP和ER_ANPP。夏季增雨显著增加了灌丛斑块GEP和ER值, 但对NEE影响不明显, 冬季增雪对灌丛斑块的碳交换过程有促进作用。总体而言, 灌丛斑块的|NEE|、GEP、ER显著高于草本斑块。相较于湿润年份(2021年), 干旱年份(2020年)的碳交换对降水增加的响应更为敏感。2)灌丛斑块碳交换(|NEE|、GEP和ER)与土壤水分含量、叶片生物量呈正相关关系, 夏季增雨主要通过增加深层土壤(40-80 cm)水分含量、降低土壤温度来促进碳交换。草本斑块碳交换与浅层土壤(0-20 cm)水分含量、ANPP呈正相关关系, 与土壤温度、根冠比呈负相关关系; 夏季增雨主要通过增加浅层土壤水分含量、降低土壤温度促进碳交换, 而冬季增雪则通过增加深层土壤水分含量和减少地下生物量来抑制草本斑块碳交换。3)基于ANPP标准化后的碳交换参数能更好地揭示灌丛斑块和草本斑块对降水变化的响应差异。该研究结果为准确评估气候变化下干旱半干旱草地生态系统的碳汇功能和固碳潜力提供了重要科学依据。

关键词: 净生态系统碳交换(NEE), 总生态系统生产力(GEP), 生态系统呼吸(ER), 灌丛斑块, 草本斑块, 冬季增雪, 夏季增雨, 灌丛化

Abstract:

Aims With the intensification of climate change and human activities, the phenomenon of shrub encroachment in arid and semi-arid grasslands is widespread, significantly impacting the carbon sequestration function of grassland ecosystems. Water availability is the primary limiting factor in the semiarid grassland of Nei Mongol, and future changes in precipitation patterns have important implications for carbon exchange in grassland ecosystems. However, there is limited research on the effects of precipitation changes on shrub-encroached grassland ecosystems, particularly on the carbon exchange processes within heterogenous patches. The underlying mechanisms remain unclear.

Methods In this study, we conducted a seasonal precipitation manipulation experiment by increasing snowfall in winter and rainfall in summer in shrub-encroached grassland dominated by Caragana microphylla in Nei Mongol. Carbon exchange parameters, such as net ecosystem carbon exchange (NEE), gross ecosystem productivity (GEP), and ecosystem respiration (ER) of shrub patches and grass patches, were measured and compared using standardized parameters based on aboveground net primary productivity (ANPP) and leaf area index (LAI). The study investigated the impact of increased seasonal precipitation on carbon exchange in shrub-encroached grassland and the differential responses of heterogeneous patches.

Important findings 1) Increased summer rainfall significantly enhanced |NEE|, GEP and ER of the grass patches, while increased winter snowfall significantly reduced |NEE|_ANPP, GEP_ANPP and ER_ANPP of the grass patches. Increased summer rainfall significantly enhanced the GEP and ER of the shrub patches, while the effect on |NEE| was not significant. Additionally, increased winter snowfall had a positive impact on carbon exchange processes in the shrub patches. Overall, |NEE|, GEP and ER of the shrub patches were significantly higher than those of the grass patches. In comparison to a wet year (2021), the carbon exchange in a dry year (2020) was more sensitive to increased precipitation. 2) Carbon exchange in the shrub patches (|NEE|, GEP and ER) was positively correlated with soil water content and leaf biomass. Increased summer rainfall mainly promoted carbon exchange by enhancing deep soil water content (40-80 cm) and lowering soil temperature. Carbon exchange in the grass patches was positively correlated with shallow soil water content (0-20 cm) and ANPP, and negatively correlated with soil temperature and root-to-shoot ratio. Increased summer rainfall primarily enhanced carbon exchange in grass patches by raising shallow soil water content and lowering soil temperature, while increased winter snowfall hindered carbon exchange by increasing deep soil water content and stimulating belowground biomass. 3) Standardized carbon exchange parameters based on ANPP better revealed the differential responses of the shrub patches and grass patches to changes in precipitation. These research findings provide an important scientific basis for accurately assessing the carbon sink function and carbon sequestration potential of arid and semi-arid grasslands ecosystems under climate change.

Key words: net ecosystem carbon exchange (NEE), gross ecosystem productivity (GEP), ecosystem respiration (ER), shrub patches, grass patches, increased winter snowfall, increased summer rainfall, shrub encroachment

张梦迪, 向官海, 文艺瑶, 王欢, 呼格吉勒, 白永飞, 王忠武, 郑淑霞. 灌丛斑块和草本斑块碳交换对季节性降水增加的响应——基于地上净初级生产力和叶面积指数标准化的比较分析. 植物生态学报, 2024, 48(8): 1035-1049. DOI: 10.17521/cjpe.2023.0359

ZHANG Meng-Di, XIANG Guan-Hai, WEN Yi-Yao, WANG Huan, Hugejile , BAI Yong-Fei, WANG Zhong-Wu, ZHENG Shu-Xia. Response of carbon exchange between shrub and grass patches to increased seasonal precipitation: a comparative analysis based on aboveground net primary productivity and leaf area index standardization. Chinese Journal of Plant Ecology, 2024, 48(8): 1035-1049. DOI: 10.17521/cjpe.2023.0359

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0359

https://www.plant-ecology.com/CN/Y2024/V48/I8/1035

图/表 7

图1 内蒙古小叶锦鸡儿灌丛化草地水分和养分添加实验平台。C, 对照; N, 氮添加; P, 磷添加; S, 冬季增雪; W, 夏季增雨。Block 1-5为5个区组重复。样方布设图中空白处为少于2丛灌木的样地, 不适合作为长期实验样方。

Fig. 1 Experimental platform for water and nutrient addition in Caragana microphylla shrub-encroached grassland of Nei Mongol. C, control; N, nitrogen addition; P, phosphorus addition; S, increased winter snowfall; W, increased summer rainfall. Block 1-5 are replicated for five blocks. The empty spaces in the sample plot layout diagram represent plots with fewer than two shrubs, which are not suitable for long-term experimental plots.

表1 季节性降水增加与年份(Y)对灌丛斑块和草本斑块碳交换参数的线性混合模型分析

Table 1 Linear mixed model analysis of the effects of increased seasonal precipitation and year (Y) on carbon exchange parameters in shrub and grass patches

图2 冬季增雪(S)和夏季增雨(W)对灌丛斑块(Shrub)和草本斑块(Grass)碳交换参数(A)的影响, 以及对基于地上净初级生产力(ANPP) (B)和叶面积指数(LAI) (C)标准化的碳交换参数的影响(平均值±标准误)。Control, 对照; ER, 生态系统呼吸; ERANPP, 基于地上净初级生产力标准化的生态系统呼吸; ERLAI, 基于叶面积指数标准化的生态系统呼吸; GEP, 总生态系统生产力; GEPANPP, 基于地上净初级生产力标准化的总生态系统生产力; GEPLAI, 基于叶面积指数标准化的总生态系统生产力; Grass, 草本斑块; NEE, 净生态系统碳交换; NEEANPP, 基于地上净初级生产力标准化的净生态系统碳交换; NEELAI, 基于叶面积指数标准化的净生态系统碳交换; Shrub, 灌丛斑块; SW, 冬季增雪和夏季增雨。*表示在相应年份处理效应显著; *, p < 0.05; **, p < 0.01; ***, p < 0.001。不同大写字母表示同一年份内灌丛斑块和草本斑块差异显著(p < 0.05)。

Fig. 2 Effects of increased winter snowfall (S) and increased summer rainfall (W) on carbon exchange parameters (A) in shrub and grass patches, and their impact on carbon exchange parameters normalized by aboveground net primary productivity (ANPP) (B) and leaf area index (LAI) (C) (mean ± SE). Control, without water addition; ER, ecosystem respiration; ERANPP, ecosystem respiration standardized by aboveground net primary productivity; ERLAI, ecosystem respiration standardized by leaf area index; GEP, gross ecosystem productivity; GEPANPP, gross ecosystem productivity standardized by aboveground net primary productivity; GEPLAI, gross ecosystem productivity standardized by leaf area index; Grass, grass patch; NEE, net ecosystem CO2 exchange; NEEANPP, net ecosystem CO2 exchange standardized by aboveground net primary productivity; NEELAI, net ecosystem CO2 exchange standardized by leaf area index; Shrub, shrub patch; SW, increased winter snowfall and summer rainfall. * indicates that the treatment effect is significant in a given year; *, p < 0.05; **, p < 0.01; ***, p < 0.001. Different uppercase letters indicate significant differences between shrub and grass patches in the same year (p < 0.05).

图3 干旱(A)和湿润(B)年份灌丛斑块和草本斑块的碳交换参数(NEE、GEP、ER)以及基于地上净初级生产力(ANPP)、叶面积指数(LAI)标准化数值对季节性降水增加(S、W、SW)的响应(响应值= (处理-对照)/对照) (平均值±标准误)。ER, 生态系统呼吸; ERANPP, 基于地上净初级生产力标准化的生态系统呼吸; ERLAI, 基于叶面积指数标准化的生态系统呼吸; GEP, 总生态系统生产力; GEPANPP, 基于地上净初级生产力标准化的总生态系统生产力; GEPLAI, 基于叶面积指数标准化的总生态系统生产力; Grass, 草本斑块; NEE, 净生态系统碳交换; NEEANPP, 基于地上净初级生产力标准化的净生态系统碳交换; NEELAI, 基于叶面积指数标准化的净生态系统碳交换; S, 冬季增雪; Shrub, 灌丛斑块; SW, 冬季增雪和夏季增雨; W, 夏季增雨。*表示增水处理下碳交换参数的响应值与0比较是否显著; *, p < 0.05; **, p < 0.01; ***, p < 0.001。

Fig. 3 Carbon exchange parameters (NEE, GEP, ER) of shrub and grass patches responses to increased seasonal precipitation (S, W, SW) in dry (A) and wet (B) years, and the corresponding response values are based on standardization of aboveground net primary productivity (ANPP) and leaf area index (LAI) (mean ± SE). ER, ecosystem respiration; ERANPP, ecosystem respiration standardized by aboveground net primary productivity; ERLAI, ecosystem respiration standardized by leaf area index; GEP, gross ecosystem productivity; GEPANPP, gross ecosystem productivity standardized by aboveground net primary productivity; GEPLAI, gross ecosystem productivity standardized by leaf area index; Grass, grass patch; NEE, net ecosystem CO2 exchange; NEEANPP, net ecosystem CO2 exchange standardized by aboveground net primary productivity; NEELAI, net ecosystem CO2 exchange standardized by leaf area index; S, increased winter snowfall; Shrub, shrub patch; SW, increased winter snowfall and summer rainfall; W, increased summer rainfall. Response value = (treatment - control)/control. * indicates whether the response value of the carbon exchange parameter is significant when compared with 0; *, p < 0.05; **, p < 0.01; ***, p < 0.001.

图4 冬季增雪(S)和夏季增雨(W)对灌丛斑块(A)和草本斑块(B)的非生物和生物因素的影响(平均值±标准误)。ANPP, 地上净初级生产力; BGB, 地下生物量; Control, 对照; LB, 叶片生物量; RSR, 根冠比; SLR, 茎叶比; SWC0-20, 0-20 cm土壤水分含量; SWC40-80, 40-80 cm土壤水分含量; ST0-10, 0-10 cm土壤温度; SW, 冬季增雪和夏季增雨。非生物因素包括浅层(0-20 cm)和深层(40-80 cm)土壤水分含量、表层土壤温度(0-10 cm), 生物因素包括ANPP、BGB、LB、SLR和RSR。*表示在相应年份处理效应显著; *, p < 0.05; **, p < 0.01; ***, p < 0.001。

Fig. 4 Effects of increased winter snowfall (S) and increased summer rainfall (W) on abiotic and biological factors in shrub (A) and grass (B) patches (mean ± SE). ANPP, aboveground net primary productivity; BGB, belowground biomass; Control, without water addition; LB, leaf biomass; RSR, root shoot ratio; SLR, stem leaf ratio; SWC0-20, soil water content (0-20 cm depth); SWC40-80, soil water content (40-80 cm depth); ST0-10, soil temperature (0-10 cm depth); SW, increased winter snowfall and summer rainfall. Abiotic factors include soil moisture content in shallow layer (0-20 cm) and deep layer (40-80 cm), surface soil temperature (0-10 cm), while biological factors include ANPP, BGB, LB, SLR and RSR. * indicates that the treatment effect is significant in a given year; *, p < 0.05; **, p < 0.01; ***, p < 0.001.

图5 灌丛斑块和草本斑块碳交换参数与浅层(SWC0-20)和深层(SWC40-80)土壤水分、土壤温度(ST0-10)的相关性热图。ER, 生态系统呼吸; ERANPP, 基于地上净初级生产力标准化的生态系统呼吸; ERLAI, 基于叶面积指数标准化的生态系统呼吸; GEP, 总生态系统生产力; GEPANPP, 基于地上净初级生产力标准化的总生态系统生产力; GEPLAI, 基于叶面积指数标准化的总生态系统生产力; NEE, 净生态系统碳交换; NEEANPP, 基于地上净初级生产力标准化的净生态系统碳交换; NEELAI, 基于叶面积指数标准化的净生态系统碳交换; SWC0-20, 0-20 cm土壤水分含量; SWC40-80, 40-80 cm土壤水分含量; ST0-10, 0-10 cm土壤温度。蓝色表示负相关, 橙色表示正相关。椭圆越细长, 颜色越深, 表明两个变量之间的相关性越强。*, p < 0.05; **, p < 0.01; ***, p < 0.001。

Fig. 5 Correlation heatmaps of carbon exchange parameters with soil moisture content in shallow (SWC0-20) and deep (SWC40-80) layers, and soil temperature (ST0-10) in shrub and grass patches. ER, ecosystem respiration; ERANPP, ecosystem respiration standardized by aboveground net primary productivity; ERLAI, ecosystem respiration standardized by leaf area index; GEP, gross ecosystem productivity; GEPANPP, gross ecosystem productivity standardized by aboveground net primary productivity; GEPLAI, gross ecosystem productivity standardized by leaf area index; NEE, net ecosystem CO2 exchange; NEEANPP, net ecosystem CO2 exchange standardized by aboveground net primary productivity; NEELAI, net ecosystem CO2 exchange standardized by leaf area index; SWC0-20, soil water content (0-20 cm depth); SWC40-80, soil water content (40-80 cm depth); ST0-10, soil temperature (0-10 cm depth). Blue indicates a negative correlation, while orange indicates a positive correlation. The more prolate the ellipse, the darker the color, and the stronger the correlation between the two variables. *, p < 0.05; **, p < 0.01; ***, p < 0.001.

图6 灌丛斑块和草本斑块碳交换参数与地上净初级生产力、地下生物量、叶片生物量及分配(根冠比、茎叶比)之间的关系。ER, 生态系统呼吸; GEP, 总生态系统生产力; NEE, 净生态系统碳交换。空心符号代表2020年灌丛斑块或草本斑块, 实心符号代表2021年灌丛斑块或草本斑块。阴影表示拟合模型95%置信区间。因小叶锦鸡儿的地下生物量测定较困难, 本图中仅呈现草本斑块的地下生物量和根冠比。

Fig. 6 Relationships between carbon exchange parameters of shrub and grass patches and aboveground net primary productivity and below-ground biomass, leaf biomass, and allocation (root-shoot ratio (RSR), stem-leaf ratio (SLR)). ER, ecosystem respiration; GEP, gross ecosystem productivity; NEE, net ecosystem CO2 exchange. Hollow symbols represent 2020 shrub or grass patches and solid symbols represent 2021 shrub or grass patches. Shadows represent the 95% confidence intervals of the fitted models. Because it is difficult to measure the belowground biomass of Caragana microphylla, only the belowground biomass and root-shoot ratio of grass patches are shown.

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灌丛斑块和草本斑块碳交换对季节性降水增加的响应——基于地上净初级生产力和叶面积指数标准化的比较分析

Response of carbon exchange between shrub and grass patches to increased seasonal precipitation: a comparative analysis based on aboveground net primary productivity and leaf area index standardization

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