毛乌素沙地典型固沙植物黑沙蒿和赖草叶片氮分配对最大净光合速率的影响

doi:10.17521/cjpe.2024.0097

植物生态学报 ›› 2025, Vol. 49 ›› Issue (3): 460-474.DOI: 10.17521/cjpe.2024.0097 cstr: 32100.14.cjpe.2024.0097

毛乌素沙地典型固沙植物黑沙蒿和赖草叶片氮分配对最大净光合速率的影响

赵洪贤¹, 刘鹏¹^,², 史曼英³, 徐铭泽⁴^,⁵, 贾昕¹^,², 田赟¹^,², 查天山¹^,²^,^*()

¹北京林业大学水土保持学院, 宁夏盐池毛乌素沙地生态系统国家定位观测研究站, 北京林业大学水土保持国家林业和草原局重点实验室, 北京 100083
²林木资源高效生产全国重点实验室, 北京 100083
³陇南市武都区林木种苗管理站, 甘肃陇南 746099
⁴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
⁵中国农业大学资源与环境学院, 北京 100080

收稿日期:2024-04-03 接受日期:2024-08-23 出版日期:2025-03-20 发布日期:2024-08-26
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(32071843)

Effect of leaf nitrogen allocation on maximum net photosynthetic rate of two common sand-fixing species, Artemisia ordosica and Leymus secalinus, in Mau Us Sandy Land

ZHAO Hong-Xian¹, LIU Peng¹^,², SHI Man-Ying³, XU Ming-Ze⁴^,⁵, JIA Xin¹^,², TIAN Yun¹^,², ZHA Tian-Shan¹^,²^,^*()

¹School of Soil and Water Conservation, Yanchi Ecology Research Station of Mau Us Desert, Key Laboratory of State Forestry and Grassland Administration on Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
²State Key Laboratory of Efficient Production of Forest Resources, Beijing 100083, China
³Forest Seed and Seedling Management Station of Wudu District, Longnan, Gansu 746099, China
⁴Observation 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
⁵College of Resources and Environmental Sciences,China Agricultural University, Beijing 100080, China

Received:2024-04-03 Accepted:2024-08-23 Online:2025-03-20 Published:2024-08-26
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(32071843)

摘要/Abstract

摘要： 探究气候变化背景下物种的适应机制是植物生理生态领域研究热点, 研究半干旱区优势物种叶片氮分配对植物光合能力的影响, 有利于理解气候变化背景下荒漠植物的适应能力。该研究于2021年5-10月, 以宁夏盐池毛乌素沙地的主要建群种黑沙蒿(俗名: 油蒿, Artemisia ordosica)和赖草(Leymus secalinus)为研究对象, 通过测定黑沙蒿和赖草的光响应曲线、CO₂响应曲线和叶性状参数, 计算叶片不同组分中的氮分配比例, 结合同步观测的环境数据, 探究两种固沙植物叶片氮分配相关参数对最大净光合速率(A_max)的影响。结果显示: 在观测期内, 黑沙蒿和赖草的单位质量叶片氮含量(N_mass)均值都为0.02 g·g^-1, 黑沙蒿A_max和叶氮在光合系统中的分配比例(P_p)均值分别为22.44 μmol·m^-2·s^-1和42.9%, 赖草A_max和P_p观测期内均值分别为11.99 μmol·m^-2·s^-1和27.5%, 相比黑沙蒿较低。在土壤水分亏缺的生长季中期, 黑沙蒿A_max和P_p降低, 而赖草的A_max和P_p并没有明显变化。相比于赖草, 黑沙蒿A_max受叶片氮分配的影响更大, P_p和叶氮在羧化系统中的分配比例(P_c)分别是影响黑沙蒿和赖草A_max季节变异的主要叶片氮分配参数。在干旱胁迫期间, 黑沙蒿和赖草叶片N_mass均降低, 黑沙蒿叶片氮分配比例对土壤水分变化的敏感性较大, 其光合氮利用效率(PNUE)降低, 且投资更多的氮到非光合机构中来抵抗干旱, 而赖草的叶片氮分配比例和PNUE的变化相对稳定, 其A_max并没有显著下降。总之, 黑沙蒿叶片对光合系统投入的氮比例相对较高, 其光合能力在观测期内仍大于赖草。该研究表明, 不同沙地物种对其生境的适应性存在差异, 在未来干旱增加的条件下, 该区域植被可能趋于以沙地灌木黑沙蒿为主的植被群落。

关键词: 半干旱区, 叶片氮分配, 光合氮利用效率, 最大净光合速率, 黑沙蒿, 赖草

Abstract:

Aims Studying the adaptation mechanisms of species in the context of climate change has been a concern in plant ecology. Investigating the effect of leaf nitrogen allocation of sand-fixing species on the photosynthetic capacity of plants help us to understand their adaptive capacity under changing climate.
Methods In this study, we selected two common sand-fixing species, Artemisia ordosica and Leymus secalinus, in Yanchi, Northwest China. The effects of leaf nitrogen allocation on the maximum net photosynthetic rate (A_max) of the two species were analyzed using in situ measurements of photosynthetic light- and CO₂-response curves, leaf traits, and enviromental variables, from May to October, 2021.
Important findings The results showed that the mean leaf nitrogen content per unit mass (N_mass) was about 0.02 g·g^-1 for both species. The mean values of A_max and the distribution proportion of leaf nitrogen in photosynthetic system (P_p) in A. ordosica were 22.44 μmol·m^-2·s^-1 and 42.9%, respectively. The mean values of A_max and P_p in L. secalinus were 11.99 μmol·m^-2·s^-1 and 27.5%, respectively, which were lower than those of A. ordosica. During the water deficit period in the middle growing season, A_max and P_p decreased in A. ordosica and remained relatively unchanged in L. secalinus. Comparing with L. secalinus, A_max of A. ordosica was more affected by leaf nitrogen distribution. The main controlling factors of A_max were P_p and leaf nitrogen distribution in carboxylation system (P_c) in A. ordosica and L. secalinus, respectively. In the middle growing season with low soil moisture, the N_mass decreased in both species. Photosynthetic nitrogen use efficiency (PNUE) decreased in A. ordosica due to its greater sensitivity of the nitrogen distribution ratio to soil moisture and more nitrogen investment in non-photosynthetic organisms and thus higher stress resistance, and remained relatively stable in L. secalinus. The A_maxin A. ordosica was greater than L. secalinus during the observation period due to its relatively higher P_p. The differences between two sand-fixing species in variations in leaf nitrogen allocation and leaf photosynthetic capacity indicate that the research area might be dominated by shrub species A. ordosica under changing climate.

Key words: semi-arid zone, leaf nitrogen allocation, photosynthetic nitrogen use efficiency, maximum net photosynthetic rate, Artemisia ordosica, Leymus secalinus

赵洪贤, 刘鹏, 史曼英, 徐铭泽, 贾昕, 田赟, 查天山. 毛乌素沙地典型固沙植物黑沙蒿和赖草叶片氮分配对最大净光合速率的影响. 植物生态学报, 2025, 49(3): 460-474. DOI: 10.17521/cjpe.2024.0097

ZHAO Hong-Xian, LIU Peng, SHI Man-Ying, XU Ming-Ze, JIA Xin, TIAN Yun, ZHA Tian-Shan. Effect of leaf nitrogen allocation on maximum net photosynthetic rate of two common sand-fixing species, Artemisia ordosica and Leymus secalinus, in Mau Us Sandy Land. Chinese Journal of Plant Ecology, 2025, 49(3): 460-474. DOI: 10.17521/cjpe.2024.0097

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0097

https://www.plant-ecology.com/CN/Y2025/V49/I3/460

图/表 10

图1 2021年研究区内环境因子日均值或日累积值变化。PPT, 降水量; REW10, 10 cm土壤相对含水量; REW30, 30 cm土壤相对含水量; SWC10, 10 cm土壤含水量; SWC30, 30 cm土壤含水量。

Fig. 1 Daily means or sums of environmental factors in the study site in 2021. PPT, precipitation; REW10, 10 cm relative extractable water content; REW30, 30 cm relative extractable water content; SWC10, 10 cm soil water content; SWC30, 30 cm soil water content.

图2 2021年毛乌素沙地黑沙蒿、赖草叶片光合参数季节动态(平均值±标准差)。图中横向参照线为各参数观测期内均值。

Fig. 2 Seasonal dynamics of leaf photosynthetic parameters in Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021 (mean ± SD). The horizontal lines represent the mean values. Amax, the maximum net photosynthetic rate; gs, stomatal conductance; Jmax, maximum electron transport efficiency; Vcmax, maximum carboxylation rate.

图3 2021年毛乌素沙地黑沙蒿、赖草叶功能性状参数季节动态(平均值±标准差)。图中横向参照线为各参数观测期内均值。

Fig. 3 Seasonal dynamics of leaf functional traits of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021 (mean ± SD). The horizontal lines represent the mean values. Cc, leaf chlorophyll content; LMA, leaf mass per unit area.

图4 2021年毛乌素沙地黑沙蒿、赖草单位质量叶片氮含量和光合氮利用效率季节动态(平均值±标准差)。横向参照线为各参数观测期内均值, B中黑色虚线是赖草叶片氮利用效率的趋势拟合线。

Fig. 4 Seasonal dynamics of leaf nitrogen content per unit mass (Nmass) and photosynthetic nitrogen use efficiency (PNUE) of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021 (mean ± SD). The horizontal lines represent the mean values. The black dashed line in B is a trend-fit line for PNUE of L. secalinus.

图5 2021年毛乌素沙地黑沙蒿、赖草叶片氮分配参数季节动态(平均值±标准差)。横向参照线为各参数观测期内均值。

Fig. 5 Seasonal dynamics of leaf nitrogen allocation parameters of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021 (mean ± SD). The horizontal lines represent the mean values. Pc, nitrogen distribution in carboxylation system; Pb, nitrogen distribution in bioenergy system; PL, nitrogen distribution in light capture system; Pp, distribution proportion of leaf nitrogen in photosynthetic system.

图6 2021年毛乌素沙地黑沙蒿(A)、赖草(B)生长季内叶片氮分配比例。Pc, 叶片氮在羧化系统中的分配比例; Pb, 叶片氮在补光系统中的分配比例; PL, 叶片氮在生物力能系统中的分配比例; P1-p, 叶片氮在非光合系统中的分配比例。

Fig. 6 Proportion of leaf nitrogen allocation of of Artemisia ordosica (A) and Leymus secalinus (B) in Mau Us Sandy Land in 2021. Pc, nitrogen distribution in carboxylation system; Pb, nitrogen distribution in bioenergy system; PL, nitrogen distribution in light capture system; P1-p, distribution proportion of leaf nitrogen in non-photosynthetic system.

图7 2021年毛乌素沙地黑沙蒿、赖草生长季内最大净光合速率与叶片氮含量相关参数的相关关系。Amax, 最大净光合速率; Nmass, 单位质量叶片氮含量; Pb, 叶片氮在补光系统中的分配比例; Pc, 叶片氮在羧化系统中的分配比例; PL, 叶片氮在生物力能系统中的分配比例; PNUE, 光合氮利用效率; Pp, 叶片氮在光合系统中的分配比例。*, p ≤0.05; **, p ≤ 0.01。圆圈大小表示相关系数的绝对值大小, 颜色表示相关系数的正负。

Fig. 7 Correlations between the maximum net photosynthetic rate and parameters related to leaf nitrogen content of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021. Amax, the maximum net photosynthetic rate; Nmass, nitrogen content of leaf per unit mass; Pb, leaf nitrogen distribution in bioenergy system; Pc, leaf nitrogen distribution in carboxylation system; PL, leaf nitrogen distribution in light capture system; PNUE, photosynthetic nitrogen use efficiency; Pp, distribution proportion of leaf nitrogen in photosynthetic system. *, p ≤ 0.05; **, p ≤ 0.01. The circle size represents the absolute magnitude of the correlation coefficient, while the color indicates its direction (positive/negative).

图8 2021年毛乌素沙地黑沙蒿、赖草生长季内叶片氮分配参数影响最大净光合速率季节变化的SHAP值。Pb, 叶片氮在捕光系统中的分配比例; Pc, 叶片氮在羧化系统中的分配比例; PL, 叶片氮在生物力能系统中的分配比例; Pp, 叶片氮在光合系统中的分配比例。

Fig. 8 The SHAP values of leaf nitrogen allocation parameters affecting the seasonal variation of the maximum net photosynthetic rate of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021. Pb, nitrogen distribution in bioenergy system; Pc, nitrogen distribution in carboxylation system; PL, nitrogen distribution in light capture system; Pp, distribution proportion of leaf nitrogen in photosynthetic system.

表1 叶片氮分配参数对黑沙蒿最大净光合速率的间接通径系数

Table 1 Indirect path coefficients of nitrogen allocation factors on the maximum net photosynthetic rate (Amax) of Artemisia ordosica and Leymus secalinus

自变量 Independent variable	间接通径系数 Indirect path coefficient
氮在生物力能学组分中的比例 P_p	0.38 (P_p→P_c→A_max)
氮在生物力能学组分中的比例 P_p	0.43 (P_p→P_b→A_max)
氮在生物力能学组分中的比例 P_b	0.34 (P_b→P_c→A_max)
氮在羧化系统中的比例 P_c	0.41 (P_c→P_b→A_max)
氮在捕光系统中分配比例 P_L	0.20 (P_L→P_b→A_max)

图9 2021年毛乌素沙地黑沙蒿和赖草最大净光合速率、叶片氮含量及叶片氮分配参数与10 cm土壤含水量的关系。阴影部分为95%置信区间。

Fig. 9 Relationships between the maximum net photosynthetic rate (Amax), nitrogen content of leaf per unit mass (Nmass), distribution proportion of leaf nitrogen in photosynthetic system (Pp), photosynthetic nitrogen use efficiency (PNUE) and soil water content at 10 cm depth (SWC10) of Artemisia ordosica and Leymus secalinus in Mau Us Sandy Land in 2021. The shaded area represents the 95% confidence interval.

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毛乌素沙地典型固沙植物黑沙蒿和赖草叶片氮分配对最大净光合速率的影响

Effect of leaf nitrogen allocation on maximum net photosynthetic rate of two common sand-fixing species, Artemisia ordosica and Leymus secalinus, in Mau Us Sandy Land

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