不同水热梯度下冠层优势树种叶片热力性状及适应策略的变化趋势

doi:10.17521/cjpe.2022.0289

摘要/Abstract

摘要：

叶片温度是植物能量交换和生理过程发生的重要微环境参数。叶片热力性状能够在一定程度上调控叶片温度, 避免极端温度对叶片的伤害。但目前针对叶片热力性状的研究还很少。该研究选择云南省从热带到温带具有明显水热梯度的4种典型植被类型: 干热河谷植被、热带雨林、亚热带常绿阔叶林、温带针阔混交林, 对其冠层优势树种的叶片热力性状进行了系统地研究。这些性状包括了可能影响叶片温度的形态、光学、材料特性、解剖和生理的23个性状。研究结果表明: 干热河谷植被的植物主要依靠蒸腾降温, 叶片薄, 叶寿命短, 主要为“快速投资-收益”型植物; 热带雨林植物叶片大, 蒸腾速率不高, 不利于降温, 较厚的叶片、较高的含水量能在一定程度上缓解高温, 采取“慢速投资-收益”策略; 亚热带常绿阔叶林很少发生极端温度, 叶片没有明显的热适应性状, 叶片厚, 叶寿命长, 采取“慢速投资-收益”策略; 温带针阔混交林植被的叶片小而厚, 多成簇状生长, 有一定保温作用。温带针阔混交林的冠层常绿植物光合速率较低, 偏“慢速投资-收益”型; 而落叶植物的光合速率较高, 偏“快速投资-收益”型。该研究系统地研究了热力性状与植物适应策略沿水热梯度的变化, 为深入认识植物对环境适应策略提供了理论基础。

关键词: 叶片功能性状, 叶经济谱, 水热梯度, 生存策略

Abstract:

Aims Leaf temperature is one of the important microenvironmental parameters for energy exchange and physiological processes of plants. Leaf thermal traits can regulate leaf temperature so as to relieve heat damage to some extent. However, systematic studies on leaf thermal traits are rare.
Methods In the present study, 43 dominant canopy species of four typical vegetation types with varying temperature and precipitation from tropical to temperate zones in Yunnan Province were selected: savanna vegetation, tropical rain forest, subtropical evergreen broadleaf forest, and temperate mixed forest. We selected 23 thermal traits that might have influence on leaf temperature, including leaf morphological, optical, material property, anatomical and physiological traits.
Important findings The results showed that plants in savanna vegetation mainly relied on transpiration for cooling. Savanna species have thin leaves and short life span. They are mainly “quick investment-return” species. Tropical rain forest plants developed large leaves, with low transpiration rates, which have no advantage of leaf cooling. Thick leaves and high water content can alleviate high temperature to some extent. They adopted “slow investment-return” strategy. Subtropical evergreen broadleaf forest was rarely exposed to extreme temperatures. The species had thick leaves, long leaf life span, and adopted “slow investment-return” strategy. They did not show obvious thermal adaptation traits. Temperate mixed forest had small and thick leaves, growing in clusters which provides benefits for thermal insulation. Photosynthetic rate of canopy evergreen plants in this forest was low, adopting “slow investment-return” strategy, while photosynthetic rate of deciduous plants was high, presenting “quick investment-return” characteristics. This study systematically investigated the variation of leaf thermal traits and plant adaptation strategies along temperature and precipitation gradients, providing a theoretical basis for further understanding of plant adaptation to the environment.

Key words: leaf functional trait, leaf economic spectrum, temperature and precipitation gradient, survival strategy

周莹莹, 林华. 不同水热梯度下冠层优势树种叶片热力性状及适应策略的变化趋势. 植物生态学报, 2023, 47(5): 733-744. DOI: 10.17521/cjpe.2022.0289

ZHOU Ying-Ying, LIN Hua. Variation of leaf thermal traits and plant adaptation strategies of canopy dominant tree species along temperature and precipitation gradients. Chinese Journal of Plant Ecology, 2023, 47(5): 733-744. DOI: 10.17521/cjpe.2022.0289

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0289

https://www.plant-ecology.com/CN/Y2023/V47/I5/733

图/表 7

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	干热河谷植被 Savanna vegetation	热带雨林 Tropical rain forest	亚热带常绿阔叶林 Subtropical evergreen broadleaf forest	温带针阔混交林 Temperate mixed forest
缩写 Abbreviation	SAV	TRF	STF	TEF
地理位置 Location	23.47° N 102.17° E	21.92° N 101.92° E	24.53° N 101.02° E	27.13° N 100.22° E
海拔 Altitude (m)	550	550	2 500	3 240
年平均气温 Annual mean air temperature (℃)	24.3	21.8	11.7	7.9
年降水量 Annual precipitation (mm)	734	1 500	1 728	1 095
平均树高 Average tree height (m)	4-6	>30	25-30	38-42
叶面积指数 Leaf area index	1.5	5	4.3	3

	干热河谷植被 Savanna vegetation	热带雨林 Tropical rain forest	亚热带常绿阔叶林 Subtropical evergreen broadleaf forest	温带针阔混交林 Temperate mixed forest
缩写 Abbreviation	SAV	TRF	STF	TEF
地理位置 Location	23.47° N 102.17° E	21.92° N 101.92° E	24.53° N 101.02° E	27.13° N 100.22° E
海拔 Altitude (m)	550	550	2 500	3 240
年平均气温 Annual mean air temperature (℃)	24.3	21.8	11.7	7.9
年降水量 Annual precipitation (mm)	734	1 500	1 728	1 095
平均树高 Average tree height (m)	4-6	>30	25-30	38-42
叶面积指数 Leaf area index	1.5	5	4.3	3

植被类型 Forest	物种编号 Species number	物种 Species	科 \| 属 Family \| genus	生活型 Life form
SAV	1	厚皮树 Lannea coromandelica	漆树科 \| 厚皮树属 Anacardiaceae \| Lannea	落叶乔木 Deciduous tree
SAV	2	鞍叶羊蹄甲 Bauhinia brachycarpa	豆科 \| 羊蹄甲属 Fabaceae \| Bauhinia	落叶灌木 Deciduous shrub
SAV	3	杭子梢 Campylotropis macrocarpa	豆科 \| 杭子梢属 Fabaceae \| Campylotropis	落叶灌木 Deciduous shrub
SAV	4	疏序黄荆 Vitex negundo	唇形科 \| 牡荆属 Lamiaceae \| Vitex	落叶灌木或小乔木 Deciduous shrub or tree
SAV	5	土蜜藤 Bridelia stipularis	叶下珠科 \| 土蜜树属 Phyllanthaceae \| Bridelia	落叶木质藤本 Deciduous woody liane
SAV	6	细基丸 Polyalthia cerasoides	番荔枝科 \| 暗罗属 Annonaceae \| Polyalthia	落叶乔木 Deciduous tree
SAV	7	白皮乌口树 Tarenna depauperata	茜草科 \| 乌口树属 Rubiaceae \| Tarenna	常绿灌木 Evergreen shrub
SAV	8	瘤果三宝木 Trigonostemon tuberculatus	大戟科 \| 三宝木属 Euphorbiaceae \| Trigonostemon	落叶灌木 Deciduous shrub
SAV	9	灰毛浆果楝 Cipadessa cinerascens	楝科 \| 浆果楝属 Meliaceae \| Cipadessa	落叶灌木 Deciduous shrub
SAV	10	异序乌桕 Sapium insigne	大戟科 \| 异序乌桕属 Euphorbiaceae \| Falconeria	落叶乔木 Deciduous tree
SAV	11	心叶木 Haldina cordifolia	茜草科 \| 心叶木属 Rubiaceae \| Haldina	落叶乔木 Deciduous tree
SAV	12	宿萼木 Strophioblachia fimbricalyx	大戟科 \| 宿萼木属 Euphorbiaceae \| Strophioblachia	落叶灌木 Deciduous shrub
TRF	13	大叶风吹楠 Horsfieldia kingii	肉豆蔻科 \| 风吹楠属 Myristicaceae \| Horsfieldia	常绿乔木 Evergreen tree
TRF	14	琴叶风吹楠 Horsfieldia pandurifolia	肉豆蔻科 \| 风吹楠属 Myristicaceae \| Horsfieldia	常绿乔木 Evergreen tree
TRF	15	大叶红光树 Knema linifolia	肉豆蔻科 \| 红光树属 Myristicaceae \| Knema	常绿乔木 Evergreen tree
TRF	16	潺槁木姜子 Litsea glutinosa	樟科 \| 木姜子属 Lauraceae \| Litsea	常绿乔木 Evergreen tree
TRF	17	勐仑翅子树 Pterospermum menglunense	锦葵科 \| 翅子树属 Malvaceae \| Pterospermum	常绿乔木 Evergreen tree
TRF	18	八宝树 Duabanga grandiflora	千屈菜科 \| 八宝树属 Lythraceae \| Duabanga	常绿乔木 Evergreen tree
TRF	19	木奶果 Baccaurea ramiflora	叶下珠科 \| 木奶果属 Phyllanthaceae \| Baccaurea	常绿乔木 Evergreen tree
TRF	20	望天树 Parashorea chinensis	龙脑香科 \| 柳安属 Dipterocarpaceae \| Parashorea	常绿乔木 Evergreen tree
TRF	21	番龙眼 Pometia pinnata	无患子科 \| 番龙眼属 Sapindaceae \| Pometia	常绿乔木 Evergreen tree
TRF	22	披针叶楠 Phoebe lanceolata	樟科 \| 楠属 Lauraceae \| Phoebe	常绿乔木 Evergreen tree
STF	23	舟柄茶 Hartia sinensis	山茶科 \| 舟柄茶属 Theaceae \| Hartia	常绿乔木 Evergreen tree
STF	24	大花八角 Illicium macranthum	五味子科 \| 八角属 Schisandraceae \| Illicium	常绿灌木或小乔木 Evergreen shrub or tree
STF	25	珍珠花 Lyonia ovalifolia	杜鹃花科 \| 珍珠花属 Ericaceae \| Lyonia	落叶灌木或小乔木 Deciduous shrub or tree
STF	26	滇南山杨 Populus rotundifolia	杨柳科 \| 杨属 Salicaceae \| Populus	落叶乔木 Deciduous tree
STF	27	硬斗石栎 Lithocarpus hancei	壳斗科 \| 石栎属 Fagaceae \| Lithocarpus	常绿乔木 Evergreen tree
STF	28	南洋木荷 Schima noronhae	山茶科 \| 木荷属 Theaceae \| Schima	常绿乔木 Evergreen tree
STF	29	红花木莲 Manglietia insignis	木兰科 \| 木莲属 Magnoliaceae \| Manglietia	常绿乔木 Evergreen tree
STF	30	变色锥 Castanopsis rufescens	壳斗科 \| 锥属 Fagaceae \| Castanopsis	常绿乔木 Evergreen tree
STF	31	多花含笑 Michelia floribunda	木兰科 \| 含笑属 Magnoliaceae \| Michelia	常绿乔木 Evergreen tree
STF	32	黄心树 Machilus bombycina	樟科 \| 润楠属 Lauraceae \| Machilus	常绿乔木 Evergreen tree
STF	33	多花山矾 Symplocos ramosissima	山矾科 \| 山矾属 Symplocaceae \| Symplocos	常绿灌木或小乔木 Evergreen shrub or tree
STF	34	滇润楠 Machilus yunnanensis	樟科 \| 润楠属 Lauraceae \| Machilus	常绿乔木 Evergreen tree
STF	35	木果柯 Lithocarpus xylocarpus	壳斗科 \| 柯属 Fagaceae \| Lithocarpus	常绿乔木 Evergreen tree
TEF	36	西南栒子 Cotoneaster franchetii	蔷薇科 \| 栒子属 Rosaceae \| Cotoneaster	落叶灌木 Deciduous shrub
TEF	37	红棕杜鹃 Rhododendron rubiginosum	杜鹃花科 \| 杜鹃花属 Ericaceae \| Rhododendron	落叶灌木 Deciduous shrub
TEF	38	川滇高山栎 Quercus aquifolioides	壳斗科 \| 栎属 Fagaceae \| Quercus	常绿乔木 Evergreen tree
TEF	39	腹毛柳 Salix delavayana	杨柳科 \| 柳属 Salicaceae \| Salix	落叶灌木或小乔木 Deciduous shrub or tree
TEF	40	毛叶珍珠花 Lyonia villosa	杜鹃花科 \| 珍珠花属 Ericaceae \| Lyonia	落叶灌木或小乔木 Deciduous shrub or tree
TEF	41	大白杜鹃 Rhododendron decorum	杜鹃花科 \| 杜鹃花属 Ericaceae \| Rhododendron	常绿灌木 Evergreen shrub
TEF	42	黄背栎 Quercus pannosa	壳斗科 \| 栎属 Fagaceae \| Quercus	常绿小乔木 Evergreen tree
TEF	43	滇南山杨 Populus rotundifolia	杨柳科 \| 杨属 Salicaceae \| Populus	落叶乔木 Deciduous tree

植被类型 Forest	物种编号 Species number	物种 Species	科 \| 属 Family \| genus	生活型 Life form
SAV	1	厚皮树 Lannea coromandelica	漆树科 \| 厚皮树属 Anacardiaceae \| Lannea	落叶乔木 Deciduous tree
SAV	2	鞍叶羊蹄甲 Bauhinia brachycarpa	豆科 \| 羊蹄甲属 Fabaceae \| Bauhinia	落叶灌木 Deciduous shrub
SAV	3	杭子梢 Campylotropis macrocarpa	豆科 \| 杭子梢属 Fabaceae \| Campylotropis	落叶灌木 Deciduous shrub
SAV	4	疏序黄荆 Vitex negundo	唇形科 \| 牡荆属 Lamiaceae \| Vitex	落叶灌木或小乔木 Deciduous shrub or tree
SAV	5	土蜜藤 Bridelia stipularis	叶下珠科 \| 土蜜树属 Phyllanthaceae \| Bridelia	落叶木质藤本 Deciduous woody liane
SAV	6	细基丸 Polyalthia cerasoides	番荔枝科 \| 暗罗属 Annonaceae \| Polyalthia	落叶乔木 Deciduous tree
SAV	7	白皮乌口树 Tarenna depauperata	茜草科 \| 乌口树属 Rubiaceae \| Tarenna	常绿灌木 Evergreen shrub
SAV	8	瘤果三宝木 Trigonostemon tuberculatus	大戟科 \| 三宝木属 Euphorbiaceae \| Trigonostemon	落叶灌木 Deciduous shrub
SAV	9	灰毛浆果楝 Cipadessa cinerascens	楝科 \| 浆果楝属 Meliaceae \| Cipadessa	落叶灌木 Deciduous shrub
SAV	10	异序乌桕 Sapium insigne	大戟科 \| 异序乌桕属 Euphorbiaceae \| Falconeria	落叶乔木 Deciduous tree
SAV	11	心叶木 Haldina cordifolia	茜草科 \| 心叶木属 Rubiaceae \| Haldina	落叶乔木 Deciduous tree
SAV	12	宿萼木 Strophioblachia fimbricalyx	大戟科 \| 宿萼木属 Euphorbiaceae \| Strophioblachia	落叶灌木 Deciduous shrub
TRF	13	大叶风吹楠 Horsfieldia kingii	肉豆蔻科 \| 风吹楠属 Myristicaceae \| Horsfieldia	常绿乔木 Evergreen tree
TRF	14	琴叶风吹楠 Horsfieldia pandurifolia	肉豆蔻科 \| 风吹楠属 Myristicaceae \| Horsfieldia	常绿乔木 Evergreen tree
TRF	15	大叶红光树 Knema linifolia	肉豆蔻科 \| 红光树属 Myristicaceae \| Knema	常绿乔木 Evergreen tree
TRF	16	潺槁木姜子 Litsea glutinosa	樟科 \| 木姜子属 Lauraceae \| Litsea	常绿乔木 Evergreen tree
TRF	17	勐仑翅子树 Pterospermum menglunense	锦葵科 \| 翅子树属 Malvaceae \| Pterospermum	常绿乔木 Evergreen tree
TRF	18	八宝树 Duabanga grandiflora	千屈菜科 \| 八宝树属 Lythraceae \| Duabanga	常绿乔木 Evergreen tree
TRF	19	木奶果 Baccaurea ramiflora	叶下珠科 \| 木奶果属 Phyllanthaceae \| Baccaurea	常绿乔木 Evergreen tree
TRF	20	望天树 Parashorea chinensis	龙脑香科 \| 柳安属 Dipterocarpaceae \| Parashorea	常绿乔木 Evergreen tree
TRF	21	番龙眼 Pometia pinnata	无患子科 \| 番龙眼属 Sapindaceae \| Pometia	常绿乔木 Evergreen tree
TRF	22	披针叶楠 Phoebe lanceolata	樟科 \| 楠属 Lauraceae \| Phoebe	常绿乔木 Evergreen tree
STF	23	舟柄茶 Hartia sinensis	山茶科 \| 舟柄茶属 Theaceae \| Hartia	常绿乔木 Evergreen tree
STF	24	大花八角 Illicium macranthum	五味子科 \| 八角属 Schisandraceae \| Illicium	常绿灌木或小乔木 Evergreen shrub or tree
STF	25	珍珠花 Lyonia ovalifolia	杜鹃花科 \| 珍珠花属 Ericaceae \| Lyonia	落叶灌木或小乔木 Deciduous shrub or tree
STF	26	滇南山杨 Populus rotundifolia	杨柳科 \| 杨属 Salicaceae \| Populus	落叶乔木 Deciduous tree
STF	27	硬斗石栎 Lithocarpus hancei	壳斗科 \| 石栎属 Fagaceae \| Lithocarpus	常绿乔木 Evergreen tree
STF	28	南洋木荷 Schima noronhae	山茶科 \| 木荷属 Theaceae \| Schima	常绿乔木 Evergreen tree
STF	29	红花木莲 Manglietia insignis	木兰科 \| 木莲属 Magnoliaceae \| Manglietia	常绿乔木 Evergreen tree
STF	30	变色锥 Castanopsis rufescens	壳斗科 \| 锥属 Fagaceae \| Castanopsis	常绿乔木 Evergreen tree
STF	31	多花含笑 Michelia floribunda	木兰科 \| 含笑属 Magnoliaceae \| Michelia	常绿乔木 Evergreen tree
STF	32	黄心树 Machilus bombycina	樟科 \| 润楠属 Lauraceae \| Machilus	常绿乔木 Evergreen tree
STF	33	多花山矾 Symplocos ramosissima	山矾科 \| 山矾属 Symplocaceae \| Symplocos	常绿灌木或小乔木 Evergreen shrub or tree
STF	34	滇润楠 Machilus yunnanensis	樟科 \| 润楠属 Lauraceae \| Machilus	常绿乔木 Evergreen tree
STF	35	木果柯 Lithocarpus xylocarpus	壳斗科 \| 柯属 Fagaceae \| Lithocarpus	常绿乔木 Evergreen tree
TEF	36	西南栒子 Cotoneaster franchetii	蔷薇科 \| 栒子属 Rosaceae \| Cotoneaster	落叶灌木 Deciduous shrub
TEF	37	红棕杜鹃 Rhododendron rubiginosum	杜鹃花科 \| 杜鹃花属 Ericaceae \| Rhododendron	落叶灌木 Deciduous shrub
TEF	38	川滇高山栎 Quercus aquifolioides	壳斗科 \| 栎属 Fagaceae \| Quercus	常绿乔木 Evergreen tree
TEF	39	腹毛柳 Salix delavayana	杨柳科 \| 柳属 Salicaceae \| Salix	落叶灌木或小乔木 Deciduous shrub or tree
TEF	40	毛叶珍珠花 Lyonia villosa	杜鹃花科 \| 珍珠花属 Ericaceae \| Lyonia	落叶灌木或小乔木 Deciduous shrub or tree
TEF	41	大白杜鹃 Rhododendron decorum	杜鹃花科 \| 杜鹃花属 Ericaceae \| Rhododendron	常绿灌木 Evergreen shrub
TEF	42	黄背栎 Quercus pannosa	壳斗科 \| 栎属 Fagaceae \| Quercus	常绿小乔木 Evergreen tree
TEF	43	滇南山杨 Populus rotundifolia	杨柳科 \| 杨属 Salicaceae \| Populus	落叶乔木 Deciduous tree

功能群 Functional group	叶片性状 Leaf trait	缩写 Abbreviation	单位 Unit	生态学意义 Ecological significance
叶片形态性状 Leaf morphological traits	单叶叶面积 Individual leaf area	Area	cm²	叶片大小影响边界层厚度和对光的截取 Leaf size affects thickness of boundary layer, and interception of light (Milla & Reich, 2007)
叶片形态性状 Leaf morphological traits	单叶叶片周长 Individual leaf perimeter	Peri	cm	影响换热面积 Impact on heat exchange area (Milla & Reich, 2007)
叶片光学性状 Leaf optical traits	绿度 Greenness	Greenness	-	影响叶片的光能捕获效率 Impact on light capture efficiency of leaves (Crawford et al., 2012)
	反射率 Reflectivity	Ref	%	影响叶片的光能捕获效率 Impact on the light capture efficiency of leaves (Crawford et al., 2012)
	透射率 Transmissivity	Trans	%	影响叶片的光能捕获效率 Impact on the light capture efficiency of leaves (Crawford et al., 2012)
	吸收率 Absorptivity	Abs	%	影响叶片的光能捕获效率 Impact on the light capture efficiency of leaves (Crawford et al., 2012)
叶片材料特性 Leaf material property	单位面积叶片含水量 Leaf equivalent water thickness	EWT	mg·cm^-2	叶片含水量越高, 比热容越大 The higher water content of leaves, the higher specific heat capacity (Leigh et al., 2012)
	叶干物质含量 Leaf dry matter content	LDMC	g·g^-1	LDMC越高, 叶片干物质投资越多, 密度越大 The higher LDMC, the higher leaf dry matter investment and leaf density (Niinemets, 2001)
	比叶质量 Leaf mass per area	LMA	mg·cm^-2	反映叶片的建成成本 Reflecting leaf construction investment (Wright et al., 2004)
叶片解剖性状 Leaf anatomy traits	叶片厚度 Leaf thickness	Thickness	μm	叶片较厚, 比叶质量较高。影响叶片比热容 Thicker leaves with higher LMA. Thickness also influences the specific heat capacity of leaves (Wright et al., 2004; Westoby & Wright, 2006)
	上表皮厚度 Thickness of upper epidermis	Epidermis_up	μm	影响水、CO₂和热量交换 Affecting water, CO₂and heat exchange (Cai & Song, 2001)
	下表皮厚度 Thickness of lower epidermis	Epidermis_low	μm	影响水、CO₂和热量交换 Affecting water, CO₂and heat exchange (Cai & Song, 2001)
	栅栏组织厚度 Thickness of palisade tissue	Thickness_palisade	μm	栅栏组织厚度影响叶绿体的分布, 影响光合作用和水气交换 Palisade tissue is related to the distribution of chloroplasts, thus affecting photosynthesis, water and gas exchange (Terashima et al., 2011)
	海绵组织厚度 Thickness of spongy tissue	Thickness_spongy	μm	影响CO₂和水交换 Affecting CO_2,and water exchange (Terashima et al., 2011)
	栅栏组织与叶片厚度比值 Ratio of palisade tissue thickness to leaf thickness	PT	%	影响CO₂和水交换 Affecting CO_2,and water exchange (Terashima et al., 2011)
	海绵组织与叶片厚度比值 Ratio of sponge tissue thickness to leaf thickness	ST	%	影响CO₂和水交换 Affecting CO_2,and water exchange (Terashima et al., 2011)
	气孔长度 Stomatal length	St_length	μm	小气孔可以更快地响应外界环境因子, 而大气孔在潮湿阴蔽的环境中更占优势 Small stomata can respond more quickly to external environmental factors (Drake et al., 2013), while large stomata are more dominant in moist shaded environment (Zhang et al., 2012)
	气孔密度 Stomatal density	St_density	No. ·mm^-2	单位面积的气孔数量越多, CO₂和水交换效率越高 More stomata per area enables more CO₂ assimilation, and leads to higher water and gas exchange rate (Tanaka & Shiraiwa, 2009)
	叶脉密度 Vein density	Vein_density	mm·mm^-2	较高的叶脉密度会增加叶片的水力导度和光合速率 The higher the vein density, the higher water transport efficiency and photosynthesis (Sack & Scoffoni, 2013)
	气孔开口面积指数 Stomatal pore area index	SPI	%	反映叶片气孔导度的综合参数, 较高的SPI会提高叶片的碳同化和蒸腾能力 An integrative parameter reflecting leaf stomatal conductance; higher SPI leads to higher carbon assimilation and transpiration capacity (Tian et al., 2016)
光学特性 Photosynthetic characteristics	蒸腾速率 Transpiration rate	T_r	mmol·m^-2·s^-1	蒸腾能起到很好的降温效果, 高的蒸腾速率意味着更高的CO₂同化率 Transpiration is beneficial for cooling, and higher transpiration rate leads to higher CO₂ assimilation rate (Lin et al., 2017)
	单位面积最大净光合速率 Maximum photosynthetic rate per area	A_area	μmol·m^-2·s^-1	反映植物对资源的获取和利用 Reflecting acquisition and utilization of resources (Franks & Beerling, 2009)
	单位质量最大净光合速率 Maximum photosynthetic rate per mass	A_mass	μmol·g^-1·s^-1	反映植物对资源的获取和利用 Reflecting acquisition and utilization of resources (Franks & Beerling, 2009)