青藏高原那曲高山嵩草草甸植物物候对增温的异步响应

doi:10.17521/cjpe.2022.0156

植物生态学报 ›› 2023, Vol. 47 ›› Issue (2): 183-194.DOI: 10.17521/cjpe.2022.0156

所属专题：青藏高原植物生态学：种群生态学

青藏高原那曲高山嵩草草甸植物物候对增温的异步响应

夏璟钰¹^,², 张扬建², 郑周涛², 赵广², 赵然²^,³, 朱艺旋², 高洁², 沈若楠², 李文宇², 郑家禾², 张雨雪¹, 朱军涛²^,^*(), 孙建新¹

¹北京林业大学生态与自然保护学院, 北京 100083
²中国科学院地理科学与资源研究所生态系统网络观测与模拟重点实验室, 那曲高寒草地生态系统野外科学观测研究站, 北京 100101
³北京大学深圳研究生院, 广东深圳 518055

收稿日期:2022-04-21 接受日期:2022-10-18 出版日期:2023-02-20 发布日期:2023-02-28
通讯作者: ORCID: *朱军涛: 0000-0002-3506-1247(zhujt@igsnrr.ac.cn)
基金资助:
国家自然科学基金(U20A2009);国家自然科学基金(41991234);国家自然科学基金(42077422);国家重点研发计划(2017YFA0604802)

Asynchronous response of plant phenology to warming in a Kobresia pygmaea meadow in Nagqu, Qingzang Plateau

XIA Jing-Yu¹^,², ZHANG Yang-Jian², ZHENG Zhou-Tao², ZHAO Guang², ZHAO Ran²^,³, ZHU Yi-Xuan², GAO Jie², SHEN Ruo-Nan², LI Wen-Yu², ZHENG Jia-He², ZHANG Yu-Xue¹, ZHU Jun-Tao²^,^*(), SUN Osbert Jianxin¹

¹School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China
²Nagqu Alpine Grassland Research Station, Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
³Peking University Shenzhen Graduate School, Shenzhen, Guangdong 518055, China

Received:2022-04-21 Accepted:2022-10-18 Online:2023-02-20 Published:2023-02-28
Contact: *(zhujt@igsnrr.ac.cn)
Supported by:
National Natural Science Foundation of China(U20A2009);National Natural Science Foundation of China(41991234);National Natural Science Foundation of China(42077422);National Key R&D Program of China(2017YFA0604802)

摘要/Abstract

摘要：

植物物候对气候变暖的响应是全球气候变化研究的重要内容。目前, 高海拔生态系统植物物候对气候变暖响应的研究仍然较少。该研究依托西藏那曲高寒草地生态系统国家野外科学观测研究站布设的梯度增温实验, 分别于2015、2017、2018和2021年对模拟增温下优势物种高山嵩草(Kobresia pygmaea)和钉柱委陵菜(Potentilla saundersiana)返青期、现蕾期和开花期等表征植物物候的指标进行了观测, 以期揭示增温下藏北高寒草甸植物物候变化机制。结果表明: 随着温度升高, 高寒草甸中优势植物物候具有不同的变化趋势。高山嵩草返青、现蕾和开花物候期的推迟幅度与温度升高幅度呈正相关关系; 钉柱委陵菜返青、现蕾和开花时间随着温度上升表现为先提前后推迟; 这表明高寒草甸植物物候对增温产生异步响应。此外, 长期增温下的藏北高寒草甸优势种的物候变化均显示出了延迟效应。结构方程归因分析发现, 空气温度升高促使高山嵩草返青时间推迟; 低水平增温可以促进钉柱委陵菜物候提前, 而随着温度继续升高其物候响应发生逆转, 土壤水分在决定物候对气候变暖响应的幅度和方向上具有关键作用。该研究结果揭示了藏北高寒草甸优势植物物候响应气候变暖的异步性特征, 为预测未来高海拔生态系统对气候变化的响应提供了数据支撑。

关键词: 高寒草地, 异步响应, 梯度增温, 植物物候, 温度敏感性

Abstract:

Aims The response of plant phenology to climate warming is an important element of global change research. At present, studies on plant phenology response to climate warming are in severe shortage for high-altitude ecosystems, especially regarding responses to multiple-level warming.

Methods We conducted a multiple-level warming experiment in an alpine meadow on Qingzang Plateau, and monitored plant phenology of two dominant species, including the timing of green up, budding and flowering in 2015, 2017, 2018 and 2021.

Important findings The results showed that plant phenology of different species exhibited various trends under warming. For Kobresia pygmaea, delay in phenological development, including green up, budding and flowering, was positively correlated with temperature increases. However, the timing of phenological stages of Potentilla saundersiana showed advancing first, and then delay with increasing temperature. These results suggest that plant phenology of alpine meadow asynchronously responds to increased temperature. In addition, temperature increase exerts delayed effects on plant phenology over long-term. The structural equation modeling showed that temperature increase consistently delayed the green up of K. pygmaea, and low-level warming advanced phenological development of P. saundersiana, but this advancing trend reversed under high-level warming. Importantly, soil moisture plays a key role in determining the magnitude and direction of phenological response to climate warming in our study. Our findings indicate the asynchronous characteristics of plant phenology response to climate warming in alpine meadow ecosystems, and provide basis to predict responses of high-altitude ecosystems to climate change in the future.

Key words: alpine meadow, asynchronous response, multiple-level warming, plant phenology, temperature sensitivity

夏璟钰, 张扬建, 郑周涛, 赵广, 赵然, 朱艺旋, 高洁, 沈若楠, 李文宇, 郑家禾, 张雨雪, 朱军涛, 孙建新. 青藏高原那曲高山嵩草草甸植物物候对增温的异步响应. 植物生态学报, 2023, 47(2): 183-194. DOI: 10.17521/cjpe.2022.0156

XIA Jing-Yu, ZHANG Yang-Jian, ZHENG Zhou-Tao, ZHAO Guang, ZHAO Ran, ZHU Yi-Xuan, GAO Jie, SHEN Ruo-Nan, LI Wen-Yu, ZHENG Jia-He, ZHANG Yu-Xue, ZHU Jun-Tao, SUN Osbert Jianxin. Asynchronous response of plant phenology to warming in a Kobresia pygmaea meadow in Nagqu, Qingzang Plateau. Chinese Journal of Plant Ecology, 2023, 47(2): 183-194. DOI: 10.17521/cjpe.2022.0156

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0156

https://www.plant-ecology.com/CN/Y2023/V47/I2/183

图/表 7

图1 青藏高原高寒草甸2015、2017、2018、2021年植物生长季空气温度(A)、土壤温度(B)和土壤含水量(C)对增温的响应。CK, 对照(自然环境温度); W1、W2、W3和W4分别为4个增温处理(增温幅度分别约为2.18、2.66、3.21和3.68 ℃)。

Fig. 1 Responses of air temperature (A), soil temperature (B) and soil water content (C) to warming during the growing seasons of 2015, 2017, 2018, 2021 under multiple levels warming in an alpine meadow on Qingzang Plateau. CK, control (ambient temperature); W1, W2, W3 and W4 are the four level warming, respectively (the warming amplitudes are about 2.18, 2.66, 3.21 and 3.68 °C, respectively).

图2 青藏高原高寒草甸梯度增温处理对各监测物种4年平均返青、现蕾和开花时间的影响(平均值±标准误)。W1、W2、W3和W4分别为4个增温幅度(增温幅度分别约为2.18、2.66、3.21和3.68 ℃)。图中正值代表与对照相比推后的天数, 负值代表与对照相比提前的天数。采用单因素方差分析和Tukey检验进行两两比较, 不同小写字母表示增温处理间存在显著差异(p < 0.05)。

Fig. 2 Effects of gradient warming treatments on four-year mean green up, budding and flowering time of each monitored species in an alpine meadow on Qingzang Plateau (mean ± SE). W1, W2, W3 and W4 are the four level warming, respectively (the warming amplitudes are about 2.18, 2.66, 3.21 and 3.68 °C, respectively). A positive value indicates later green up, budding or flowering than the control; a negative value indicates earlier green up, budding or flowering than the control. One-way analysis of variance and Tukey test were used for pairwise comparison. Different lowercase letters indicate significant differences between warming treatments (p < 0.05).

表1 年份(Y)、增温(W)对高山嵩草和钉柱委陵菜返青、现蕾和开花时间的主效应和交互效应的重复测量方差分析

Table 1 Repeated measured analysis of variance for main and interactive effects of year (Y), warming (W) on Kobresia pygmaea and Potentilla saundersiana green-up, budding and flowering time

物种 Species	处理 Treatment	返青时间 Green up date			现蕾时间 Budding date			开花时间 Flowering date
物种 Species	处理 Treatment	df	F	p	df	F	p	df	F	p
高山嵩草 Kobresia pygmaea	W	4	45.619	<0.001	4	28.665	<0.001	4	6.832	<0.001
	Y	3	258.565	<0.001	3	151.230	<0.001	3	79.438	<0.001
	W × Y	12	10.445	<0.001	12	1.970	0.032	12	18.260	0.699
钉柱委陵菜 Potentilla saundersiana	W	4	7.781	<0.001	4	7.344	<0.001	4	8.260	<0.001
	Y	3	27.377	<0.001	3	18.011	<0.001	3	1.117	0.344
	W × Y	12	2.912	0.001	12	4.560	<0.001	12	3.423	<0.001

图3 梯度增温处理对高山嵩草、钉柱委陵菜平均返青、现蕾和开花时间的影响(平均值±标准误)。W1、W2、W3和W4分别为4个增温幅度(增温幅度分别约为2.18、2.66、3.21和3.68 ℃)。图中正值代表与对照相比推后的天数, 负值代表与对照相比提前的天数(d)。采用单因素方差分析和Tukey检验进行两两比较, 不同小写字母表示增温处理间存在显著差异(p < 0.05)。

Fig. 3 Effects of gradient warming treatments on Kobresia pygmaea and Potentilla saundersiana green up, budding and flowering time (mean ± SE) averaged on 2015, 2017, 2018, 2021. W1, W2, W3 and W4 are the four level warming treatments, respectively (the warming amplitudes are about 2.18, 2.66, 3.21 and 3.68 °C, respectively). A positive value indicates later green up, budding or flowering time than the control; a negative value indicates earlier green up, budding or flowering time than the control. One-way analysis of variance and Tukey test were used for pairwise comparison. Different lowercase letters indicate significant differences between warming treatments (p < 0.05).

图4 梯度增温处理对高山嵩草、钉柱委陵菜4年平均返青期、现蕾期和开花期温度敏感性的影响(平均值±标准误)。W1、W2、W3和W4分别为4个增温幅度(增温幅度分别约为2.18、2.66、3.21和3.68 ℃)。采用单因素方差分析和Tukey检验进行两两比较, 不同小写字母表示增温处理间存在显著差异(p < 0.05)。

Fig. 4 Effects of gradient warming treatment on four years mean temperature sensitivity of green up, budding and flowering time (mean ± SE) of Kobresia pygmaea and Potentilla saundersiana. W1, W2, W3 and W4 are four level warming, respectively (the warming amplitudes are about 2.18, 2.66, 3.21 and 3.68 °C). One-way analysis of variance and Tukey test were used for pairwise comparison. Different lowercase letters indicate significant differences between warming treatments (p < 0.05).

表2 青藏高原高寒草甸高山嵩草和钉柱委陵菜物候变化时间和环境因子的回归方程

Table 2 Regression equations linking Kobresia pygmaea and Potentilla saundersiana phenological change time and environmental factors in an alpine meadow on Qingzang Plateau

物种 Species	物候时间 Phenological time	回归方程 Regression equation	F	R²	p
高山嵩草 Kobresia pygmaea	返青时间 Green up time	y = -6.363 + 4.647x₂+ 9.968x₃	55.744	0.503	<0.01
	现蕾时间 Budding time	y = -4.787 + 3.308x₂ + 7.640x₃	33.781	0.381	<0.01
	开花时间 Flowering time	y = 1.517 + 4.579x₁ + 2.980x₂	13.590	0.198	<0.01
钉柱委陵菜 Potentilla saundersiana	返青时间 Green up time	y = 1.030 - 3.984x₁ - 9.349x₂- 5.606x₃	41.132	0.495	<0.01
	现蕾时间 Budding time	y = 0.672 - 3.346x₁ - 8.092x₂- 4.654x₃	31.390	0.428	<0.01
	开花时间 Flowering time	y = -0.045 - 2.958x₁ - 6.400x₂- 2.748x₃	18.649	0.307	<0.01

图5 空气温度、土壤温度和土壤水分对高山嵩草(A)和钉柱委陵菜(B)的返青、开花时间的影响。高山嵩草: χ2 = 0.198, p = 0.656, 比较拟合指数(CFI) = 1.000, 近似误差均方根(RMSE) = 0.000, 赤池信息量准则(AIC) = -127.449。钉柱委陵菜: χ2 = 0.019, p = 0.889, CFI = 1.000, RMSE = 0.000, AIC = -154.980。线条上的数字代表自变量对因变量直接作用的标准通径系数, 红线表示正相关, 蓝线表示负相关; 实线表示显著相关(p < 0.05)。

Fig. 5 Effects of air temperature, soil temperature and soil water content on green up, flowering time of Kobresia pygmaea (A) and Potentilla saundersiana (B). Kobresia pygmaea: χ2 = 0.198, p = 0.656, comparative fit index (CFI) = 1.000, root mean square error (RMSE) = 0.000, Akaike information criterion (AIC) = -127.449. Potentilla saundersiana: χ2 = 0.019, p = 0.889, CFI = 1.000, RMSE = 0.000, AIC = -154.980. Number on the lines are standardized direct path coefficients; the red lines indicated negative correlations; the blue lines indicated positive correlations. Solid lines indicate significant correlation (p < 0.05).

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青藏高原那曲高山嵩草草甸植物物候对增温的异步响应

Asynchronous response of plant phenology to warming in a Kobresia pygmaea meadow in Nagqu, Qingzang Plateau

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