叶片和群落尺度净光合速率关系的探讨

doi:10.17521/cjpe.2015.0089

摘要/Abstract

摘要：

叶片净光合速率(P_n)是研究光合作用机理的基本尺度; 而群落净光合速率(P_c)是研究群落光合能力及其与外部环境因子间关系的更好尺度, 特别是区域乃至全球尺度碳循环的研究, 需要将叶片尺度的生理生态模型扩展到冠层尺度。理论上, 群落内所有叶片的累积P_n与实测群落净气体交换速率(NCE)是相等的, 但在野外实际观测中, 两者之间的相互关系目前尚未见报道。该文选取敖汉苜蓿(Medicago sativa ‘Aohan’)人工草地, 采用美国LI-COR公司生产的便携式光合测定系统LI-6400测定P_n, 结合叶面积指数等参数推算P_c, 利用LI-8100连接同化箱测定生态系统净气体交换速率(NEE), 加上土壤呼吸速率, 得到NCE。结果表明: P_c为3.52 μmol CO₂·m^-2·s^-1, 与实测NCE (3.56 μmol CO₂·m^-2·s^-1)基本相等。这表明: 可利用P_n, 结合叶面积指数、群落叶片数目、健康叶片比例和群落可接收有效光照的平均比例等4个关键参数, 准确地换算P_c。然而, 利用同化箱式法测定群落呼吸速率时, 不可避免地会包含土壤呼吸, 所以在观测NCE时, 需要同时测定土壤呼吸。此外, 在冠层模型中, 群落垂直结构和光量子的非线性响应不可忽视。

关键词: 净光合速率, 冠层模型, 尺度上推, 敖汉苜蓿

Abstract:

Leaf net photosynthesis is crucial for detecting the mechanism of photosynthesis, whereas community net photosynthesis is useful for understanding the photosynthetic capacity of communities and its relationship with environmental factors. In particular, we need to scale up eco-physiological models from leaf scale to canopy level to study carbon cycling at regional or global scale. We hypothesized that accumulated leaf net photosynthetic rate (P_c) at community scale, i.e., calculated based on leaf net photosynthetic rate (P_n) and leaf area index (LAI), equals to measured net community CO₂ exchange (NCE). The purpose of this study is to verify this hypothesis. Our field study was carried out in Duolun, Nei Mongol, China, where we constructed single-species communities by sowing Medicago sativa ‘Aohan’ seeds in three plots (3 m × 5 m) on May 30, 2012. On August 16, 2014, P_n of five healthy leaves of M. sativa ‘Aohan’ in each plot were measured with a LI-6400 portable photosynthesis system at 10:00, and net ecosystem CO₂ exchange (NEE) in each plot was measured simultaneously with a LI-8100 system connected with a assimilation chamber (0.5 m × 0.5 m × 0.5 m). P_c was calculated based on P_n, number of leaves (n), LAI percentage of healthy leaves (r) and percentage of received effective light by leaves (m). NCE was derived from NEE and ecosystem respiration rate (R_eco). P_cwas 3.52 μmol CO₂·m^-2·s^-1, and very close to NCE (3.56 μmol CO₂·m^-2·s^-1), suggesting that leaf-scale photosynthesis may accurately predict community-scale photosynthesis. However, our method could not separate community respiration from soil respiration, and future studies, should be designed to counteract this effect. Scaling up from leaf photosynthesis to community photosynthesis should also consider vertical structure of communities and nonlinear responses of leaf photosynthesis to changes in light quantum.

Key words: net photosynthetic rate, canopy model, scaling up, Medicago sativa ‘Aohan’

唐海萍, 薛海丽, 房飞. 叶片和群落尺度净光合速率关系的探讨. 植物生态学报, 2015, 39(9): 924-931. DOI: 10.17521/cjpe.2015.0089

TANG Hai-Ping,XUE Hai-Li,FANG Fei. A comparison of measured and calculated net community CO₂ exchange: Scaling from leaves to communities. Chinese Journal of Plant Ecology, 2015, 39(9): 924-931. DOI: 10.17521/cjpe.2015.0089

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0089

https://www.plant-ecology.com/CN/Y2015/V39/I9/924

图/表 2

参考文献 54

[24]	Ping XY, Zhou GS, Sun JS (2010). Advances in the study of photosynthate allocation and its controls.Chinese Journal of Plant Ecology, 34, 100-111.(in Chinese with English abstract)
	[平晓燕, 周广胜, 孙敬松 (2010). 植物光合产物分配及其影响因子研究进展. 植物生态学报, 34, 100-111.]
[25]	Prince SD, Goward SN (1995). Global primary production: A remote sensing approach.Journal of Biogeography, 22, 815-835.
[26]	Rinaldi M (2004). Water availability at sowing and nitrogen management of durum wheat: A seasonal analysis with the CERES-wheat model.Field Crops Research, 89, 27-37.
[27]	Sellers PJ, Berry JA, Collatz GJ, Field CB, Hall FG (1992). Canopy reflectance, photosynthesis, and transpiration. III: A reanalysis using improved leaf models and a new canopy integration scheme.Remote Sensing of Environment, 42, 187-216.
[28]	Sun DB, Wang QS (2012). Effects of water on the photosynthetic characteristics of alfalfa (Medicago sativa).Chinese Journal of Plant Ecology, 36, 72-80.(in Chinese with English abstract)
	[孙东宝, 王庆锁 (2012). 水分对苜蓿叶片光合特性的影响. 植物生态学报, 36, 72-80.]
[29]	Tao Y (2007). Contrusting Physiological Properties of Shaded and Sunlit Leaves, and Applying a Photosynthesis Model for Cotton. Master degree dissertation, Nanjing University of Information Science and Technology, Nanjing.(in Chinese with English abstract)
	[陶寅 (2007). 棉花阴、阳叶的生理特征对比以及光合模型应用研究. 硕士学位论文, 南京信息工程大学, 南京.]
[30]	Turner DP, Ritts WD, Styles JM, Yang Z, Cohen WB, Law BE, Thornton PE (2006). A diagnostic carbon flux model to monitor the effects of disturbance and interannual varia- tion in climate on regional NEP.Tellus B, 58, 476-490.
[31]	Veroustraete F, Sabbe H, Eerens H (2002). Estimation of carbon mass fluxes over Europe using the C-Fix model and Euroflux data.Remote Sensing of Environment, 83, 376-399.
[32]	Wang HB, Ma MG, Wang XF, Tan JL, Geng LY, Yu WP, Jia SZ (2014). Carbon flux variation characteristics and its influencing factors in an alpine meadow ecosystem on eastern Qinghai-Tibetan Plateau.Journal of Arid Land Resources and Environment, 28(6), 50-56.(in Chinese with English abstract)
	[王海波, 马明国, 王旭峰, 谭俊磊, 耿丽英, 于文凭, 家淑珍 (2014). 青藏高原东缘高寒草甸生态系统碳通量变化特征及其影响因素. 干旱区资源与环境, 28(6), 50-56.]
[33]	Wang HY, Lin S, Allen JP, Williams JC, Blankert S, Laser C, Woodbury NW (2007). Protein dynamics control the kinetics of initial electron transfer in photosynthesis.Science, 316, 747-750.
[34]	Wang J, Yu Q, Pan XB, Yin H, Zhang YQ (2008). A review on water, heat and CO2 fluxes simulation models.Acta Ecolo- gica Sinica, 28, 2843-2853.(in Chinese with English abstract)
	[王靖, 于强, 潘学标, 尹红, 张永强 (2008). 土壤-植物-大气连续体水热、CO2通量估算模型研究进展. 生态学报, 28, 2843-2853.]
[35]	Wang JL, Zhang YL, Zhu ZL, Zhong P, Liang HY (2006). Analysis on ecophysiological characteristics of leaf photosynthesis of Medicago varia cv. Gannong No. 1.Acta Agrestia Sinica, 14, 138-141.(in Chinese with English abstract)
	[王建丽, 张永亮, 朱占林, 钟鹏, 梁怀宇 (2006). 杂花苜蓿叶片光合生理生态特性. 草地学报, 14, 138-141.]
[36]	Wang WJ, Zu YG, Wang HM (2007). Review on the photosynthetic function of non-photosynthetic woody organs of stem and branches.Acta Ecologica Sinica, 27, 1583-1595.(in Chinese with English abstract)
	[王文杰, 祖元刚, 王慧梅 (2007). 林木非同化器官树枝(干)光合功能研究进展. 生态学报, 27, 1583-1595.]
[37]	Wang YP, Leuning R (1998). A two-leaf model for canopy for canopy conductance, photosynthesis and partitioning of available energy: Model description and comparison with a multi-layered model.Agricultural and Forest Meteorology, 91, 89-111.
[38]	Wang ZM, Wei AL, Zheng DM (2001). Photosynthetic characteristics of non-leaf organs of winter wheat cultivars differing in ear type and their relationship with grain mass per ear.Photosynthetica, 39, 239-244.
[39]	Xiao XM, Hollinger D, Aber J, Goltz M, Davidson EA, Zhang QY, Moore B III (2004). Satellite-based modeling of gross primary production in an evergreen needleleaf forest.Remote Sensing of Environment, 89, 519-534.
[40]	Xu DQ (2006). Some noteworthy problems in measurement and investigation of photosynthesis.Plant Physiology Communications, 43, 1163-1167.(in Chinese with English abstract)
	[许大全 (2006). 光合作用测定及研究中一些值得注意的问题. 植物生理学通讯, 43, 1163-1167.]
[41]	Xu LK, Baldocchi DD (2004). Seasonal variation in carbon dioxide exchange over a Mediterranean annual grassland in California.Agricultural and Forest Meteorology, 123, 79-96.
[42]	Xu LL, Zhang XZ, Shi PL, Yu GR (2005). Establishment of apparent quantum yield and maximum ecosystem assimilation on Tibetan Plateau alpine meadow ecosystem.Science China Earth Sciences, 48, 141-147.
[43]	Yang J, Zhou GS, Wang YL, Wang YH (2008). Characteristics of net ecosystem flux exchanges over Stipa krylovii steppe in Inner Mongolia.Chinese Journal of Applied Ecology, 19, 533-538.(in Chinese with English abstract)
	[杨娟, 周广胜, 王云龙, 王玉辉 (2008). 内蒙古克氏针茅草原生态系统-大气通量交换特征. 应用生态学报, 19, 533-538.]
[44]	Yu Q, Liu JD, Zhang YQ, Li J (2002). Simulation of rice biomass accumulation by an extended logistic model including influence of meteorological factors.International Journal of Biometeorology, 46, 185-191.
[1]	Amthor JS (1994). Scaling CO2-photosynthesis relationships from the leaf to the canopy.Photosynthesis Research, 39, 321-350.
[2]	Baldocchi DD (2003). Assessing the eddy covariance technique for evaluating carbon dioxide exchange rates of ecosystems: Past, present and future.Global Change Biology, 9, 479-492.
[45]	Yu Q, Xie XQ, Sun SF, Wang TD, Lu PL (1999). Advances in simulation of plant photosynthetic productivity and canopy evapotranspiration.Acta Ecologica Sinica, 19, 744-753.(in Chinese with English abstract)
	[于强, 谢贤群, 孙菽芬, 王天铎, 陆佩玲 (1999). 植物光合生产力与冠层蒸散模拟研究进展. 生态学报, 19, 744-753.]
[3]	Bellarby J, Tirado R, Leip A, Weiss F, Lesschen JP, Smith P (2013). Livestock greenhouse gas emissions and mitigation potential in Europe. Global Change Biology, 19, 3-18.
[4]	Chen Y, Wang XH, Liao Y, Cai SQ, Zhang HB, Xu DQ (2002). Effect of flag leaf orientation on its photosynthetic capacity in rice.Journal of Plant Physiology and Molecular Biology, 28, 396-398.(in Chinese with English abstract)
[46]	Yuan WP, Liu SG, Zhou GS, Zhou GY, Tieszen LL, Baldocchi D, Bernhofer C, Gholz H, Goldstein AH, Goulden ML, Hollinger DY, Hu YM, Law BE, Stoy PC, Vesala T, Wofsy SC (2007). Deriving a light use efficiency model from eddy covariance flux data for predicting daily gross primary production across biomes.Agricultural and Forest Meteorology, 143, 189-207.
[47]	Zha TS, Kellomäki S, Wang KY, Rouvinen I (2004). Carbon sequestration and ecosystem respiration for 4 years in a Scots pine forest.Global Change Biology, 10, 1492-1503.
[4]	[陈悦, 王学华, 廖轶, 蔡时青, 张海波, 许大全 (2002). 水稻剑叶取向对其光合功能的影响. 植物生理与分子生物学学报, 28, 396-398.]
[5]	Cheng J, Shen YG (2011). On the trends of photosynthesis research. Acta Botanica Sinica, 46, 694-704.(in Chinese with English abstract)
[48]	Zhang DY, Wang XH, Chen Y, Xu DQ (2005). Determinant of photosynthetic capacity in rice leaves under ambient air conditions.Photosynthetica, 43, 273-276.
[49]	Zhang LY, Yang HS, Ge XL, Liu J (2008). The research on photosynthetic characteristics in blooming stage and seed production of alfalfa in different growing year.Chinese Journal of Grassland, 30(5), 54-58.(in Chinese with English abstract)
[5]	[程建, 沈允钢 (2011). 试析光合作用的研究动向. 植物学报, 46, 694-704.]
[6]	Davidson EA, Janssens IA, Luo YQ (2006). On the variability of respiration in terrestrial ecosystems: Moving beyond Q10.Global Change Biology, 12, 154-164.
[49]	[张丽妍, 杨恒山, 葛选良, 刘晶 (2008). 不同生长年限紫花苜蓿花期光合特性及其种子生产性能. 中国草地学报, 30(5), 54-58.]
[50]	Zhang M, Guan DX, Wu JB, Shi TT, Jin CJ, Han SJ (2006). Research advances in eco-physiological model of plant canopy scale.Chinese Journal of Ecology, 25, 563-571.(in Chinese with English abstract)
[7]	Elberse IAM, Van Damme JMM, van Tienderen PH (2003). Plasticity of growth characteristics in wild barley (Hordeum spontaneum) in response to nutrient limitation.Journal of Ecology, 91, 371-382.
[8]	Farquhar GD, von Caemmerer S, Berry JA (1980). A biochemical model of photosynthetic CO2 assimilation in leaves of C3 plants.Planta, 149, 78-90.
[50]	[张弥, 关德新, 吴家兵, 施婷婷, 金昌杰, 韩士杰 (2006). 植被冠层尺度生理生态模型的研究进展. 生态学杂志, 25, 563-571.]
[51]	Zhang WL, Chen SP, Miao HX, Lin GH (2008). Effects on carbon flux of conversion of grassland steppe to cropland in China. Journal of Plant Ecology (Chinese Version), 32, 1301-1311.(in Chinese with English abstract)
[9]	Feng YL, Wang JF, Sang WG (2007). Biomass allocation, morphology and photosynthesis of invasive and noninvasive exotic species grown at four irradiance levels.Acta Oecologica, 31, 40-47.
[10]	Fu YL, Yu GR, Sun XM, Li YN, Wen XF, Zhang LM, Li ZQ, Zhao L, Hao YB (2006). Depression of net ecosystem CO2 exchange in semi-arid Leymus chinensis steppe and alpine shrub.Agricultural and Forest Meteorology, 137, 234-244.
[51]	[张文丽, 陈世苹, 苗海霞, 林光辉 (2008). 开垦对克氏针茅草地生态系统碳通量的影响. 植物生态学报, 32, 1301-1311.]
[52]	Zhang XC, Yu XF, Gao SM (2010). Effects of nitrogen application rates on photosynthetic energy utilization in wheat leaves under elevated atmospheric CO2 concentration.Chinese Journal of Plant Ecology, 34, 1196-1203.(in Chinese with English abstract)
[11]	Geng YB, Luo GQ (2010). Analysis of affecting factors and partitioning of respiration in a Leymus chinensis Steppe in Inner Mongolia.Acta Geographica Sinica, 65, 1058-1068.(in Chinese with English abstract)
	[耿元波, 罗光强 (2010). 内蒙古羊草草原呼吸的影响因素分析和区分. 地理学报, 65, 1058-1068.]
[52]	[张绪成, 于显枫, 高世铭 (2010). 高大气CO2浓度下氮素对小麦叶片光能利用的影响. 植物生态学报, 34, 1196-1203.]
[53]	Zhao P, Sun GC, Cai XA, Rao XQ, Zeng XP (2005). Night-time warming increases photosynthetic capacity of sapling leaf of Cinnamomum burmanni grown with different nitrogen supplies.Acta Ecologica Sinica, 25, 2703-2708.(in English)
[12]	He MZ, Ju WM, Zhou YL, Chen JM, He HL, Wang SQ, Wang HM, Guan DX, Yan JH, Li YN, Hao YB, Zhao FH (2013). Development of a two-leaf light use efficiency model for improving the calculation of terrestrial gross primary productivity.Agricultural and Forest Meteorology, 173, 28-39.
[13]	King DA, Turner DP, Ritts WD (2011). Parameterization of a diagnostic carbon cycle model for continental scale application.Remote Sensing of Environment, 115, 1653-1664.
[53]	[赵平, 孙谷畴, 蔡锡安, 饶兴权, 曾小平 (2005). 夜间变暖提高荫香叶片的光合能力. 生态学报, 25, 2703-2708.]
[54]	Zhao XS, Guan DX, Wu JB, Zhang M, Jin CJ, Han SJ (2006). The relationship between CO2 flux and temperature of the mixed forest of broad-leaved and Korean-Pine in Changbai Mountain.Acta Ecologica Sinica, 26, 1088-1095.(in Chinese with English abstract)
[14]	Leuning R, Kelliher FM, de Pury DGG, Sehulze ED (1995). Leaf nitrogen, photosynthesis, conductance and transpiration: Scaling from leaves to canopies.Plant, Cell & Environment, 18, 1183-1200.
[15]	Li CX, Zhao SJ, Meng QW, Zou Q, Tian JC (2004). Photosynthetic characteristics in non-leaf organs of winter wheat cultivars differing in grain-leaf ratio.Acta Agronomica Sinica, 30, 419-426.(in Chinese with English abstract)
	[李朝霞, 赵世杰, 孟庆伟, 邹琦, 田纪春 (2004). 不同粒叶比小麦品种非叶片光合器官光合特性的研究. 作物学报, 30, 419-426.]
[16]	Litton CM, Raich JW, Ryan MG (2007). Carbon allocation in forest ecosystems.Global Change Biology, 13, 2089-2109.
[17]	Llorens L, Peñuelas J, Beier C, Emmett B, Estiarte M, Tietema A (2004). Effects of an experimental increase of temperature and drought on the photosynthetic performance of two ericaceous shrub species along a north-south European gradient.Ecosystems, 7, 613-624.
[18]	Ma J (2011). Variations of Soil Respiration Rate and Its Temperature Sensitivity among Different Land Use Types in the Agro-Pastoral Ecotone of Inner Mongolia. PhD dissertation, Beijing Normal University, Beijing. 9.(in Chinese)
	[马骏 (2011). 内蒙古农牧交错带不同土地利用方式下土壤呼吸速率及其温度敏感性变化. 博士学位论文, 北京师范大学, 北京. 9.]
[19]	Ma WH, Yang YH, He JS, Zeng H, Fang JY (2008). The biomass and its relationship with environmental factors of temperate grassland in Inner Mongolia. Science in China (Series C: Life Sciences), 38, 84-92.(in Chinese)
	[马文红, 杨元合, 贺金生, 曾辉, 方精云 (2008). 内蒙古温带草地生物量及其与环境因子的关系. 中国科学C辑: 生命科学, 38, 84-92.]
[20]	Ma YC, Qing HL, Gao WS, Chen YQ, Li XD, Sui P, Huang FQ (2007). Dynamics of soil water content under different tillage in agriculture-pasture transition zone.Acta Ecologica Sinica, 27, 2523-2530.(in Chinese with English abstract)
	[马月存, 秦红灵, 高旺盛, 陈源泉, 李向东, 隋鹏, 黄凤球 (2007). 农牧交错带不同耕作方式土壤水分动态变化特征. 生态学报, 27, 2523-2530.]
[21]	Mahadevan P, Wofsy SC, Matross DM, Xiao XM, Dunn AL, Lin JC, Gerbig C, Munger JW, Chow VY, Gottlieb EW (2008). A satellite-based biosphere parameterization for net ecosystem CO2 exchange: Vegetation photosynthesis and respiration model (VPRM). Global Biogeochemical Cycles, 22, GB2005.
[22]	Niu SL, Li ZX, Xia JY, Han Y, Wu MY, Wan SQ (2008). Climatic warming changes plant photosynthesis and its temperature dependence in a temperate steppe of northern China.Environmental and Experimental Botany, 63, 91-101.
[23]	Pearson RG, Dawson TP (2003). Predicting the impacts of climate change on the distribution of species: Are bioclimate envelope models useful?Global Ecology and Biogeography, 12, 361-371.
[54]	[赵晓松, 关德新, 吴家兵, 张弥, 金昌杰, 韩士杰 (2006). 长白山阔叶红松林CO2通量与温度的关系. 生态学报, 26, 1088-1095.]

亚小区 Sub plot	P_n(μmol CO₂·m^-2·s^-1)	叶片数 No. of leaves	平均叶面积 Average leaf area (cm²)	健康叶片比例 Percentage of healthy leaves	叶片接受有效光照比例 Percentage of received effective light by leaf	P_c(μmol CO₂·m^-2·s^-1)	P_c / P_n (%)
亚小区1 Sub plot 1	15.025^b	650	3.113	0.4	0.7	3.41^b	22.7
亚小区2 Sub plot 2	7.854^a	580	2.477	0.5	0.7	1.58^a	20.1
亚小区3 Sub plot 3	14.536^b	740	3.081	0.6	0.7	5.57^c	38.3
平均 Average	12.472	656	2.890	0.5	0.7	3.52	27.0

亚小区 Sub plot	P_n(μmol CO₂·m^-2·s^-1)	叶片数 No. of leaves	平均叶面积 Average leaf area (cm²)	健康叶片比例 Percentage of healthy leaves	叶片接受有效光照比例 Percentage of received effective light by leaf	P_c(μmol CO₂·m^-2·s^-1)	P_c / P_n (%)
亚小区1 Sub plot 1	15.025^b	650	3.113	0.4	0.7	3.41^b	22.7
亚小区2 Sub plot 2	7.854^a	580	2.477	0.5	0.7	1.58^a	20.1
亚小区3 Sub plot 3	14.536^b	740	3.081	0.6	0.7	5.57^c	38.3
平均 Average	12.472	656	2.890	0.5	0.7	3.52	27.0

亚小区 Sub plot	P_c(μmol CO₂·m^-2·s^-1)	NCE (μmol CO₂·m^-2·s^-1)	生态系统净气体交换速率 Net ecosystem CO₂ exchange (NEE, μmol CO₂·m^-2·s^-1)	生态系统呼吸速率 Ecosystem respiration rate (R_eco, μmol CO₂·m^-2·s^-1)	群落呼吸速率 Community respiration rate (R_c, μmol CO₂·m^-2·s^-1)	NEE/P_n (%)	P_c/NEE
亚小区1 Sub plot 1	3.41	3.38	1.07	3.74	1.40	7.1	3.2
亚小区2 Sub plot 2	1.58	1.48	0.98	0.81	0.21	12.5	1.6
亚小区3 Sub plot 3	5.57	5.82	2.38	5.56	2.37	16.4	2.3
平均 Average	3.52	3.56	1.48	3.37	1.33	12.0	2.4

亚小区 Sub plot	P_c(μmol CO₂·m^-2·s^-1)	NCE (μmol CO₂·m^-2·s^-1)	生态系统净气体交换速率 Net ecosystem CO₂ exchange (NEE, μmol CO₂·m^-2·s^-1)	生态系统呼吸速率 Ecosystem respiration rate (R_eco, μmol CO₂·m^-2·s^-1)	群落呼吸速率 Community respiration rate (R_c, μmol CO₂·m^-2·s^-1)	NEE/P_n (%)	P_c/NEE
亚小区1 Sub plot 1	3.41	3.38	1.07	3.74	1.40	7.1	3.2
亚小区2 Sub plot 2	1.58	1.48	0.98	0.81	0.21	12.5	1.6
亚小区3 Sub plot 3	5.57	5.82	2.38	5.56	2.37	16.4	2.3
平均 Average	3.52	3.56	1.48	3.37	1.33	12.0	2.4