Characteristics of seasonal leaf trait network and its drivers in Artemisia ordosica in the Mau Us Sandy Land

doi:10.17521/cjpe.2024.0025

Abstract

Abstract:

Aims Clarifying the network characteristics of plant traits is one of the research hotspots in functional ecology. However, our understanding of plant trait networks (PTN) and their driving factors on a seasonal scale is still limited.

Methods We used a LI-6400 portable photosynthetic instrument to regularly measure the light response and carbon dioxide response curves of Artemisia ordosica during the growing season (May to September), as well as leaf structural and biochemical indicators, to explore the correlation and trait network characteristics among 25 leaf traits.

Important findings There were significant correlations between leaf traits at the seasonal scale, and the closest coupling relationship between traits occurred between photosynthetic physiological traits, with a total of 67 pairs of physiological trait combinations showing significant correlation. The edge density, diameter, average path length, and modularity of the PTN are 0.58, 3, 1.51, and 0.08, respectively. The betweenness of leaf tissue density, leaf nitrogen content per unit area, and transpiration rate is relatively high, making it a “bridge” in PTN. PTN does not have an absolute central trait, and physiological traits as a whole exhibit high degree, eigenvector centrality, and clustering coefficient, indicating that the trait network is jointly dominated by physiological traits. Further analysis indicated that the 25 leaf traits at the seasonal scale can be compressed into two major trait dimensions: one is regulated by air temperature and soil moisture, and the other is regulated by photosynthetic effective radiation. The results emphasize that when evaluating the potential response of different trait functional groups to climate fluctuations, it is necessary to distinguish the degree and effect of environmental factors on different traits. If the seasonal response of plant traits to the environment is included in a unified paradigm, it will incorrectly evaluate the adaptability of plant traits.

Key words: leaf trait network, seasonal dynamics, Artemisia ordosica, trait variation

XU Ming-Ze, ZHAO Hong-Xian, LI Cheng, LI Man-Le, TIAN Yun, LIU Peng, ZHA Tian-Shan. Characteristics of seasonal leaf trait network and its drivers in Artemisia ordosica in the Mau Us Sandy Land[J]. Chin J Plant Ecol, 2024, 48(12): 1650-1665.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2024.0025

https://www.plant-ecology.com/EN/Y2024/V48/I12/1650

Figures/Tables 11

References 52

[1]	Anadon-Rosell A, Hasibeder R, Palacio S, Mayr S, Ingrisch J, Ninot JM, Nogués S, Bahn M (2017). Short-term carbon allocation dynamics in subalpine dwarf shrubs and their responses to experimental summer drought. Environmental and Experimental Botany, 141, 92-102.
[2]	Ayub G, Smith RA, Tissue DT, Atkin OK (2011). Impacts of drought on leaf respiration in darkness and light in Eucalyptus saligna exposed to industrial-age atmospheric CO₂ and growth temperature. New Phytologist, 190, 1003-1018.
[3]	Bao L, Liu YH (2009). Comparison of leaf functional traits in different forest communities in Mt. Dongling of Beijing. Acta Ecologica Sinica, 29, 3692-3703.
	[ 宝乐, 刘艳红 (2009). 东灵山地区不同森林群落叶功能性状比较. 生态学报, 29, 3692-3703.]
[4]	Chai SF, Tang JM, Wang ML (2015). Photosynthetic and physiological characteristics of Camellia petelotii seedlings under drought stress. Acta Botanica Boreali-Occidentalia Sinica, 2, 322-328.
	[ 柴胜丰, 唐健民, 王满莲 (2015). 干旱胁迫对金花茶幼苗光合生理特性的影响. 西北植物学报, 2, 322-328.]
[5]	Chandregowda MH, Tjoelker MG, Power SA, Pendall E (2022). Drought and warming alter gross primary production allocation and reduce productivity in a widespread pasture grass. Plant, Cell & Environment, 45, 2271-2291.
[6]	Cui XT, Yuan FH, Wang AZ, Guan DX, Wu JB, Jin CJ (2017). Leaf age-related changes in photosynthesis of Quercus mongolica leaves in relation to leaf functional traits. Chinese Journal of Ecology, 36, 3160-3167.
	[ 崔西甜, 袁凤辉, 王安志, 关德新, 吴家兵, 金昌杰 (2017). 蒙古栎叶片光合作用随叶龄的变化及其与叶片功能性状的关系. 生态学杂志, 36, 3160-3167.]
[7]	Dai YM, Li ML, Xu MZ, Tian Y, Zhao HX, Gao SJ, Hao SR, Liu P, Jia X, Zha TS (2022). Leaf traits of Artemisia ordosica at different dune fixation stages in Mau Us Sandy Land. Chinese Journal of Plant Ecology, 46, 1376-1387.
	[ 代远萌, 李满乐, 徐铭泽, 田赟, 赵洪贤, 高圣杰, 郝少荣, 刘鹏, 贾昕, 查天山 (2022). 毛乌素沙地沙丘不同固定阶段黑沙蒿叶性状特征. 植物生态学报, 46, 1376-1387.] DOI
[8]	Deng XX, Xiao WF, Zeng LX, Lei L, Shi Z (2019). Transport and distribution characteristics of photosynthates of Pinus massoniana seedlings. Scientia Silvae Sinicae, 55(7), 27-34.
	[ 邓秀秀, 肖文发, 曾立雄, 雷蕾, 施征 (2019). 马尾松幼苗光合产物的运输与分配特征. 林业科学, 55(7), 27-34.]
[9]	Dong XG, Cao YF, Tian LM, Wang K, Zhang Y, Qi D (2015). Leaf morphology and photosynthetic characteristics of wild Ussurian pear in China. Chinese Journal of Applied Ecology, 26, 1327-1334.
	[ 董星光, 曹玉芬, 田路明, 王昆, 张莹, 齐丹 (2015). 中国野生山梨叶片形态及光合特性. 应用生态学报, 26, 1327-1334.]
[10]	Flores-Moreno H, Fazayeli F, Banerjee A, Datta A, Kattge J, Butler EE, Atkin OK, Wythers K, Chen M, Anand M, Bahn M, Byun C, Cornelissen JHC, Craine J, Gonzalez- Melo A, et al. (2019). Robustness of trait connections across environmental gradients and growth forms. Global Ecology and Biogeography, 28, 1806-1826. DOI
[11]	Guo W, Cherubini P, Zhang J, Li M, Qi L (2023). Leaf stomatal traits rather than anatomical traits regulate gross primary productivity of moso bamboo (Phyllostachys edulis) stands. Frontiers in Plant Science, 14, 1117564. DOI: 10.3389/fpls.2023.1117564.
[12]	Happonen K, Virkkala AM, Kemppinen J, Niittynen P, Luoto M (2022). Relationships between above-ground plant traits and carbon cycling in tundra plant communities. Journal of Ecology, 110, 700-716.
[13]	He NP, Li Y, Liu CC, Xu L, Li MX, Zhang JH, He JS, Tang ZY, Han XG, Ye Q, Xiao CW, Yu Q, Liu SR, Sun W, Niu SL, et al. (2020). Plant trait networks: improved resolution of the dimensionality of adaptation. Trends in Ecology & Evolution, 35, 908-918.
[14]	He NP, Liu CC, Zhang JH, Xu L, Yu GR (2018). Perspectives and challenges in plant traits: from organs to communities. Acta Ecologica Sinica, 38, 6787-6796.
	[ 何念鹏, 刘聪聪, 张佳慧, 徐丽, 于贵瑞 (2018). 植物性状研究的机遇与挑战: 从器官到群落. 生态学报, 38, 6787-6796.]
[15]	Heskel MA, Bitterman D, Atkin OK, Turnbull MH, Griffin KL (2014). Seasonality of foliar respiration in two dominant plant species from the Arctic tundra: response to long-term warming and short-term temperature variability. Functional Plant Biology, 41, 287-300. DOI PMID
[16]	Heskel MA, Tang JW (2018). Environmental controls on light inhibition of respiration and leaf and canopy daytime carbon exchange in a temperate deciduous forest. Tree Physiology, 38, 1886-1902. DOI PMID
[17]	Ji WL, LaZerte SE, Waterway MJ, Lechowicz MJ (2020). Functional ecology of congeneric variation in the leaf economics spectrum. New Phytologist, 225, 196-208. DOI PMID
[18]	Jiang Y, Tian Y, Zha T, Jia X, Bourque CPA, Liu P, Jin C, Jiang X, Li X, Wei N, Gao S (2021). Dynamic changes in plant resource use efficiencies and their primary influence mechanisms in a typical desert shrub community. Forests, 12, 1372. DOI: 10.3390/f12101372.
[19]	Jin C, Zha TS, Jia X, Tian Y, Zhou WJ, Wei TZ (2020). Light energy partitioning, photoprotection and influencing factors of photosystem II in an exotic species (Salix psammophila) in Mu Us Sandy Land. Scientia Silvae Sinicae, 56(10), 34-44.
	[ 靳川, 查天山, 贾昕, 田赟, 周文君, 卫腾宙 (2020). 毛乌素沙地沙柳光系统Ⅱ光保护机制和能量分配动态及其影响因子. 林业科学, 56(10), 34-44.]
[20]	Kirschbaum MU, Farquhar GD (1987). Investigation of the CO₂ dependence of quantum yield and respiration in Eucalyptus pauciflora. Plant Physiology, 83, 1032-1036. DOI PMID
[21]	Kleyer M, Trinogga J, Cebrián Piqueras MA, Trenkamp A, Flojgaard C, Ejrnæs R, Bouma TJ, Minden V, Maier M, Mantilla-Contreras J (2019). Trait correlation network analysis identifies biomass allocation traits and stem specific length as hub traits in herbaceous perennial plants. Journal of Ecology, 107, 829-842.
[22]	Li L, McCormack ML, Ma C, Kong D, Zhang Q, Chen X, Zeng H, Niinemets Ü, Guo DL (2015). Leaf economics and hydraulic traits are decoupled in five species-rich tropical-subtropical forests. Ecology Letters, 18, 899-906. DOI PMID
[23]	Li Y (2020). Variation of Leaf Trait Network Among Different Vegetation Types and Its Influencing Factors. PhD dissertation,Beijing Forestry University, Beijing.
	[ 李颖 (2020). 叶片性状网络在不同植被类型间的变异规律及其影响因素. 博士学位论文, 北京林业大学, 北京.]
[24]	Li Y, Liu C, Sack L, Xu L, Li M, Zhang J, He N (2022). Leaf trait network architecture shifts with species-richness and climate across forests at continental scale. Ecology Letters, 25, 1442-1457.
[25]	Li Y, Zha TS, Jia X, Qin SG, Wu YJ, Wang B (2015). Photosynthetic characteristics of typical desert plant Artemisia ordosica in semi-arid region. Chinese Journal of Ecology, 34, 86-93.
	[ 李媛, 查天山, 贾昕, 秦树高, 吴雅娟, 王奔 (2015). 半干旱区典型沙生植物油蒿(Artemisia ordosica)的光合特性. 生态学杂志, 34, 86-93.]
[26]	Li YL, Cui JY, Su YZ (2005). Specific leaf area and leaf dry matter content of some plants in different dune habitats. Acta Ecologica Sinica, 25, 304-311.
	[ 李玉霖, 崔建垣, 苏永中 (2005). 不同沙丘生境主要植物比叶面积和叶干物质含量的比较. 生态学报, 25, 304-311.]
[27]	Li Z, Tan XF, Lu K, Zhang L, Long HX, Lü JB, Lin Q (2017). Influence of drought stress on the growth, leaf gas exchange, and chlorophyll fluorescence in two varieties of tung tree seedlings. Acta Ecologica Sinica, 37, 1515-1524.
	[ 李泽, 谭晓风, 卢锟, 张琳, 龙洪旭, 吕佳斌, 林青 (2017). 干旱胁迫对两种油桐幼苗生长、气体交换及叶绿素荧光参数的影响. 生态学报, 37, 1515-1524.]
[28]	Luo D, Shi YJ, Song FH, Li JC (2021). Variation and correlation of leaf functional traits and photosynthetic characteristics of 38 hazelnut germplasm resources. Chinese Journal of Ecology, 40, 11-22. DOI
	[ 罗达, 史彦江, 宋锋惠, 李嘉诚 (2021). 38个榛种质资源叶功能性状与光合特征变异及其相关性. 生态学杂志, 40, 11-22.]
[29]	Mu RL, Liu MX, Xu L, Zhang GJ, Yu RX, Li L (2022). Diurnal variation of photosynthesis and resource utilization efficiency of the typical plants in the semi-arid area of Loess Plateau. Plant Physiology Journal, 58, 1381-1391.
	[ 穆若兰, 刘旻霞, 徐璐, 张国娟, 于瑞新, 李亮 (2022). 黄土高原半干旱区典型植物资源利用效率及光合日变化探析. 植物生理学报, 58, 1381-1391.]
[30]	Ogaya R, Peñuelas J (2003). Comparative field study of Quercus ilex and Phillyrea latifolia: photosynthetic response to experimental drought conditions. Environmental and Experimental Botany, 50, 137-148.
[31]	Osuna JL, Baldocchi DD, Kobayashi H, Dawson TE (2015). Seasonal trends in photosynthesis and electron transport during the Mediterranean summer drought in leaves of deciduous oaks. Tree Physiology, 35, 485-500. DOI PMID
[32]	Poorter H, Lambers H, Evans JR (2014). Trait correlation networks: a whole-plant perspective on the recently criticized leaf economic spectrum. New Phytologist, 201, 378-382. DOI PMID
[33]	She WW (2018). Response of Artemisia ordosica Community to Water and Nitrogen Addition in the Mu Us Desert. PhD dissertation,Beijing Forestry University, Beijing.
	[ 佘维维 (2018). 毛乌素沙地油蒿群落对水分和氮素添加的响应. 博士学位论文, 北京林业大学, 北京.]
[34]	Song GM, Wang Q, Jin J (2020). Leaf photosynthetic capacity of sunlit and shaded mature leaves in a deciduous forest. Forests, 11, 318. DOI: 10.3390/f11030318.
[35]	Song GM, Wang Q, Jin J (2021). Exploring the instability of the relationship between maximum potential electron transport rate and maximum carboxylation rate in cool-temperate deciduous forests. Agricultural and Forest Meteorology, 308, 108614. DOI: 10.1016/j.agrformet. 2021.108614.
[36]	Sun J, Guan D, Wu J, Jing Y, Yuan F, Wang A, Jin C (2015). Day and night respiration of three tree species in a temperate forest of northeastern China. iForest, 8, 25-32.
[37]	Tang YR, Zhao CZ, Zhao H, Hou G, Ma M, Zhao TT, Wang YF, Zeng HX (2021). The relationship between leaf dry mass and leaf area, leaf thickness of Hippophae rhamnoides under different light conditions in Taohe River riparian forest. Chinese Journal of Ecology, 40, 2745-2753.
	[ 唐玉瑞, 赵成章, 赵辉, 侯刚, 马敏, 赵婷婷, 王毓芳, 曾红霞 (2021). 不同光环境下洮河护岸林沙棘叶干重与叶面积、叶厚度间的关系. 生态学杂志, 40, 2745-2753.]
[38]	Wan CY, Yu JR, Zhu SD (2023). Differences in leaf traits and trait correlation networks between karst and non-karst forest tree species. Chinese Journal of Plant Ecology, 47, 1386-1397.
	[ 万春燕, 余俊瑞, 朱师丹 (2023). 喀斯特与非喀斯特森林乔木叶性状及其相关性网络的差异. 植物生态学报, 47, 1386-1397.] DOI
[39]	Wang X, Ji M, Zhang Y, Zhang L, Akram MA, Dong L, Hu W, Xiong J, Sun Y, Li H, Degen AA, Ran J, Deng J (2023). Plant trait networks reveal adaptation strategies in the drylands of China. BMC Plant Biology, 23, 266. DOI: 10.1186/s12870-023-04273-0.
[40]	Wang ZQ, Zha TS, Jia X, Wu YJ, Zhang MY, Mu JW (2017). Seasonal variation in photosynthetic parameters of Artemisia ordosica in relation to leaf nitrogen and specific leaf area. Chinese Journal of Ecology, 36, 916-924.
	[ 王子奇, 查天山, 贾昕, 吴雅娟, 张明艳, 穆家伟 (2017). 油蒿光合参数季节动态及其与叶氮含量和比叶面积的关系. 生态学杂志, 36, 916-924.]
[41]	Winkler DE, Belnap J, Duniway MC, Hoover D, Reed SC, Yokum H, Gill R (2020). Seasonal and individual event- responsiveness are key determinants of carbon exchange across plant functional types. Oecologia, 193, 811-825. DOI PMID
[42]	Wu FY, Yi LT, Li XP, Yin XM, Liu MH, Yu SQ (2012). Effect of different light intensity on intensity chlorophyll content and chlorophyll fluorescence in Lithocarpus glaber. Journal of Northeast Agricultural University, 43(4), 88-92.
	[ 吴飞燕, 伊力塔, 李修鹏, 殷秀敏, 刘美华, 余树全 (2012). 不同光照强度对石栎幼苗叶绿素含量及叶绿素荧光参数的影响. 东北农业大学学报, 43(4), 88-92.]
[43]	Wu SY (2019). Seasonal Dynamics of Leaf Functional Traits in Artemisia ordosica and Their Influencing Factors. Master degree dissertation, Beijing Forestry University, Beijing.
	[ 吴水瑛 (2019). 油蒿叶功能性状季节动态及其影响因素. 硕士学位论文, 北京林业大学, 北京.]
[44]	Wu Y, Ren C, Tian Y, Zha T, Liu P, Bai Y, Ma J, Lai Z, Bourque CPA (2018). Photosynthetic gas-exchange and PSII photochemical acclimation to drought in a native and non-native xerophytic species (Artemisia ordosica and Salix psammophila). Ecological Indicators, 94, 130-138.
[45]	Xu MZ, Liu P, Tian Y, Zhao HX, Jin C, Li ML, Mao J, Wei XS, Jia X, Zha TS (2023). Seasonal response of light use efficiency of Artemisia ordosica to leaf traits in Mu Us sandy land. Acta Ecologica Sinica, 43, 5122-5136.
	[ 徐铭泽, 刘鹏, 田赟, 赵洪贤, 靳川, 李满乐, 毛军, 魏晓帅, 贾昕, 查天山 (2023). 毛乌素沙地油蒿叶性状对光能利用效率动态的影响. 生态学报, 43, 5122-5136.]
[46]	Xu M, Zha T, Tian Y, Liu P, Jia X, Bourque CPA, Jin C, Wei X, Zhao H, Guo Z (2022). Elevated physiological plasticity in xerophytic-deciduous shrubs as demonstrated in their variable maximum carboxylation rate. Ecological Indicators, 144, 109475. DOI: 10.1016/j.ecolind.2022.109475.
[47]	Yasumura Y, Hikosaka K, Hirose T (2006). Seasonal changes in photosynthesis, nitrogen content and nitrogen partitioning in Lindera umbellata leaves grown in high or low irradiance. Tree Physiology, 26, 1315-1323. PMID
[48]	Zhang J, He N, Liu C, Xu L, Chen Z, Li Y, Wang R, Yu G, Sun W, Xiao C, Chen HYH, Reich PB (2020). Variation and evolution of C:N ratio among different organs enable plants to adapt to N-limited environments. Global Change Biology, 26, 2534-2543.
[49]	Zhao HX, Zhang YJ, Xu MZ, Wei TZ, Mao J, Luo Y, Jia X, Zha TS (2022). Effects of leaf nitrogen allocation on seasonal variation in maximum net photosynthetic rate in Artemisia ordosica. Acta Ecologica Sinica, 42, 7156-7166.
	[ 赵洪贤, 张洋军, 徐铭泽, 卫腾宙, 毛军, 雒宇, 贾昕, 查天山 (2022). 油蒿叶片氮分配对其最大净光合速率季节变异的影响. 生态学报, 42, 7156-7166.]
[50]	Zhao HY, Li YL, Wang XY, Mao W, Zhao XY, Zhang TH (2010). Variations in leaf traits of 52 plants in Horqin sand land. Journal of Desert Research, 30, 1292-1298.
	[ 赵红洋, 李玉霖, 王新源, 毛伟, 赵学勇, 张铜会 (2010). 科尔沁沙地52种植物叶片性状变异特征研究. 中国沙漠, 30, 1292-1298.]
[51]	Zheng YP, Dang CH, Hao LH, Cheng DJ, Xu M (2016). Photosynthetic and respiratory acclimation of maize leaves to experimental warming in the North China Plain. Acta Ecologica Sinica, 36, 5236-5246.
	[ 郑云普, 党承华, 郝立华, 程东娟, 徐明 (2016). 华北平原玉米叶片光合及呼吸过程对实验增温的适应性. 生态学报, 36, 5236-5246.]
[52]	Zhong GW, Tian Y, Liu P, Jia X, Zha TS (2022). Leaf traits and resource use efficiencies of 19 woody plant species in a plantation in Fangshan, Beijing, China. Forests, 14, 63. DOI: 10.3390/f14010063.

监测样株 Monitoring sample plant	冠幅 Crown diameter (cm × cm)		最大高度 Maximum height (cm)
监测样株 Monitoring sample plant	东西向 East-west	南北向 South-north	最大高度 Maximum height (cm)
HSH1	111	100	53
HSH2	108	104	61
HSH3	115	120	58

监测样株 Monitoring sample plant	冠幅 Crown diameter (cm × cm)		最大高度 Maximum height (cm)
监测样株 Monitoring sample plant	东西向 East-west	南北向 South-north	最大高度 Maximum height (cm)
HSH1	111	100	53
HSH2	108	104	61
HSH3	115	120	58

叶性状类别 Category of leaf functional trait	性状 Trait	缩写 Abbreviation	单位 Unit	生理生态解释 Physiological and ecological explanation
光合生理性状 Photosynthetic physiological trait	最大净光合速率 Maximum net photosynthetic rate	A_max	μmol·m^-2·s^-1	光合潜力与资源获取策略 Photosynthetic potential and resource acquisition strategies
	最大羧化速率 Maximum carboxylation rate	V_cmax	μmol·m^-2·s^-1	核酮糖-1,5-双磷酸羧化酶/加氧酶(Rubisco)催化的最大羧化反应速率 Maximum carboxylation rate of the enzyme ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco)
	最大电子传递速率 Maximum electron transfer rate	J_max	μmol·m^-2·s^-1	支持核酮糖双磷酸再生的最大电子传输速率 Maximum electron transfer rate for supporting ribulose bisphosphate regeneration
	暗呼吸速率 Dark respiration rate	R_d	μmol·m^-2·s^-1	黑暗环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under dark conditions
	光下暗呼吸速率 Dark respiration rate under light	R_l	μmol·m^-2·s^-1	光照环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under light conditions
	光饱和碳同化速率 Photo-saturated carbon assimilation rate	A_sat	μmol CO₂·m^-2·s^-1	碳同化效率 Carbon assimilation efficiency
	气孔导度 Stomatal conductance	g_s	mol H₂O·m⁻²·s⁻¹		气体交换与资源获取 Gas exchange and resource acquisition
	蒸腾速率 Transpiration rate	E	mmol H₂O·m⁻²·s⁻¹		耗水特性 Water consumption characteristics
	水分利用效率 Water use efficiency	WUE	μmol·mmol^-1		抗旱性与水资源利用 Drought resistance and water resource utilization
	氮利用效率 Nitrogen use efficiency	NUE	μmol·g^-1·s^-1		叶片养分利用、生理特性和生存策略 Nutrient utilization, physiological characteristics, and survival strategies of leaves
	最大光化学效率 Maximal photochemical efficiency	F_v/F_m			光系统II (PSII)反应中心的最大光能转化效率以及叶片的健康程度 The maximum light energy conversion efficiency of photosystem II (PSII) and the health status of leaves
	光化学淬灭 Photochemical quenching	q_P			PSII反应中心的开放程度以及光化学活性的高低 Degree of openness of PSII and the level of photochemical activity
	非光化学淬灭 Non-photochemical quenching	NPQ			植物的光保护能力 Light protection ability of plants
	电子传递速率 Electron transfer rate	ETR	μmol·m^-2·s^-1		植物的耐光性和适应性 Light resistance and adaptability of plants
结构性状 Structural trait	叶厚度 Leaf thickness	L_t	mm		生存策略以及光合呼吸、蒸腾相关的碳成本 Survival strategy and carbon costs related to photosynthesis, respiration and transpiration
	比叶质量 Leaf mass per area	LMA	g·m^-2		资源获取和抗逆性 Resource acquisition and resilience
	叶组织密度 Leaf tissue density	LTD	g·cm^-3		叶片韧性和抗逆性 Leaf toughness and stress resistance
	叶片相对含水量 Leaf relative water content	LRWC	%		植物的水分状况、抗旱性及环境适应 Water status, drought resistance and environmental adaptation
	叶干物质含量 Leaf dry matter content	LDMC	g·g^-1		抗逆性和对抗物理危害的防御能力 Resistance to adversity and defense against physical hazards
生物化学性状 Biochemical trait	基于面积的叶氮含量 Leaf nitrogen content per unit area	N_area	g·m^-2		资源获取和衡量光合能力 Resource acquisition and characterization of photosynthetic capacity
	基于面积的叶碳含量 Leaf carbon content per unit area	C_area	g·m^-2		构建和防御 Building and defense
	碳氮比 Ratio of leaf carbon content to nitrogen content	C:N			元素分配与适应策略 Element allocation and adaptation strategies
	叶绿素含量 Chlorophyll content	Chl	μg·cm^-2		健康程度与光合作用 Health level and photosynthesis
	叶绿素a/b Ratio of chlorophyll a to chlorophyll b	Chl a/b			光能吸收和能量运输 Light absorption and energy transport
	类胡萝卜素含量 Carotenoid content	Car	μg·cm^-2		光合作用与光保护必需色素 Photosynthesis and essential pigments for photoprotection

叶性状类别 Category of leaf functional trait	性状 Trait	缩写 Abbreviation	单位 Unit	生理生态解释 Physiological and ecological explanation
光合生理性状 Photosynthetic physiological trait	最大净光合速率 Maximum net photosynthetic rate	A_max	μmol·m^-2·s^-1	光合潜力与资源获取策略 Photosynthetic potential and resource acquisition strategies
	最大羧化速率 Maximum carboxylation rate	V_cmax	μmol·m^-2·s^-1	核酮糖-1,5-双磷酸羧化酶/加氧酶(Rubisco)催化的最大羧化反应速率 Maximum carboxylation rate of the enzyme ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco)
	最大电子传递速率 Maximum electron transfer rate	J_max	μmol·m^-2·s^-1	支持核酮糖双磷酸再生的最大电子传输速率 Maximum electron transfer rate for supporting ribulose bisphosphate regeneration
	暗呼吸速率 Dark respiration rate	R_d	μmol·m^-2·s^-1	黑暗环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under dark conditions
	光下暗呼吸速率 Dark respiration rate under light	R_l	μmol·m^-2·s^-1	光照环境中叶片呼吸的代谢水平 Levels of leaf respiratory metabolism under light conditions
	光饱和碳同化速率 Photo-saturated carbon assimilation rate	A_sat	μmol CO₂·m^-2·s^-1	碳同化效率 Carbon assimilation efficiency
	气孔导度 Stomatal conductance	g_s	mol H₂O·m⁻²·s⁻¹		气体交换与资源获取 Gas exchange and resource acquisition
	蒸腾速率 Transpiration rate	E	mmol H₂O·m⁻²·s⁻¹		耗水特性 Water consumption characteristics
	水分利用效率 Water use efficiency	WUE	μmol·mmol^-1		抗旱性与水资源利用 Drought resistance and water resource utilization
	氮利用效率 Nitrogen use efficiency	NUE	μmol·g^-1·s^-1		叶片养分利用、生理特性和生存策略 Nutrient utilization, physiological characteristics, and survival strategies of leaves
	最大光化学效率 Maximal photochemical efficiency	F_v/F_m			光系统II (PSII)反应中心的最大光能转化效率以及叶片的健康程度 The maximum light energy conversion efficiency of photosystem II (PSII) and the health status of leaves
	光化学淬灭 Photochemical quenching	q_P			PSII反应中心的开放程度以及光化学活性的高低 Degree of openness of PSII and the level of photochemical activity
	非光化学淬灭 Non-photochemical quenching	NPQ			植物的光保护能力 Light protection ability of plants
	电子传递速率 Electron transfer rate	ETR	μmol·m^-2·s^-1		植物的耐光性和适应性 Light resistance and adaptability of plants
结构性状 Structural trait	叶厚度 Leaf thickness	L_t	mm		生存策略以及光合呼吸、蒸腾相关的碳成本 Survival strategy and carbon costs related to photosynthesis, respiration and transpiration
	比叶质量 Leaf mass per area	LMA	g·m^-2		资源获取和抗逆性 Resource acquisition and resilience
	叶组织密度 Leaf tissue density	LTD	g·cm^-3		叶片韧性和抗逆性 Leaf toughness and stress resistance
	叶片相对含水量 Leaf relative water content	LRWC	%		植物的水分状况、抗旱性及环境适应 Water status, drought resistance and environmental adaptation
	叶干物质含量 Leaf dry matter content	LDMC	g·g^-1		抗逆性和对抗物理危害的防御能力 Resistance to adversity and defense against physical hazards
生物化学性状 Biochemical trait	基于面积的叶氮含量 Leaf nitrogen content per unit area	N_area	g·m^-2		资源获取和衡量光合能力 Resource acquisition and characterization of photosynthetic capacity
	基于面积的叶碳含量 Leaf carbon content per unit area	C_area	g·m^-2		构建和防御 Building and defense
	碳氮比 Ratio of leaf carbon content to nitrogen content	C:N			元素分配与适应策略 Element allocation and adaptation strategies
	叶绿素含量 Chlorophyll content	Chl	μg·cm^-2		健康程度与光合作用 Health level and photosynthesis
	叶绿素a/b Ratio of chlorophyll a to chlorophyll b	Chl a/b			光能吸收和能量运输 Light absorption and energy transport
	类胡萝卜素含量 Carotenoid content	Car	μg·cm^-2		光合作用与光保护必需色素 Photosynthesis and essential pigments for photoprotection

叶性状 Leaf trait	主成分1 Principal component 1	主成分2 Principal component 2
V_cmax	-0.22	0.03
J_max	-0.24	0.04
A_max	-0.24	-0.14
R_d	-0.23	0.14
R_l	-0.23	0.13
A_sat	-0.26	-0.01
g_s	-0.24	0.07
E	-0.25	0.10
WUE	0.11	-0.24
NUE	-0.22	-0.16
q_P	-0.21	-0.12
F_v/F_m	-0.22	-0.02
ETR	-0.25	-0.04
NPQ	-0.06	-0.04
N_area	-0.20	0.22
C_area	0.01	0.35
C:N	0.20	0.02
Chl	-0.07	-0.34
Chl a/b	-0.23	0.07
Car	-0.09	-0.31
LMA	-0.05	0.36
LDMC	0.22	0.19
LRWC	-0.23	-0.18
L_t	0.09	-0.28
LTD	-0.10	0.38
特征值 Characteristic value	3.77	2.34
方差比例 Variance ratio	54.75%	21.04%
累计方差比例 Accumulated variance ratio	54.75%	75.79%