海南岛热带山地雨林不同演替阶段植物功能群光生态适应性研究
|
摘要
海南岛的热带雨林是我国森林生态系统中结构最复杂、生物多样性最丰富的类型之一，具有极其重要的保育价值和科学研究价值。海南岛热带山地雨林主要分布在海拔700～1300m的山地，是海南岛热带森林中现存面积最大、分布最为集中的垂直地带性的自然植被类型。过度砍伐和不合理的土地利用已导致了原始森林的大面积减少，形成了处于不同演替阶段的退化生态系统。光是森林群落演替过程中最重要的影响因子之一。研究森林不同演替阶段具有不同功能特性的植物类群对光变化的生态适应性，有助于认识不同恢复阶段森林生态系统动态变化规律，为海南岛热带林生物多样性的保护和生态系统功能恢复提供科学依据。
本论文采用野外试验和控制试验相结合的研究手段，测定了热带山地雨林不同演替阶段中几个典型植物的光生态适应性，主要目的是试图从形态和生理生态的角度对不同演替阶段的植物进行功能群划分，了解不同演替阶段植物功能群在生态系统中的功能，分析植物功能群的生态适应性及其在演替过程中的作用。研究的主要结论如下：
1．通过野外现场测定，发现早期演替阶段环境因子变化剧烈，不同类型植物的适应方式存在差异。棕叶芦(Thysanolaena maxima)有着强的水分利用能力，偏重于以形态构件的变化(单个叶片的卷曲以及不同叶片之间的相互遮荫)来适应。五节芒(Miscanthus floridulus)有着强的水分控制能力，偏重以强的光合生理能力(通过高的光合能力和高的水分利用效率)来适应。野牡丹(Melastoma candidum)和毛稔(Melastoma sanguineum)的适应方式则是则是介于上述二者之间，即通过形态和生理特征变化的结合(形态上通过叶片的相互遮荫加上生理上高的光合能力)。基于植物的光合生理生态功能及适应方式，运用主成分分析方法，将海南岛热带山地雨林早期演替阶段的植物划分为三类大的功能群：(1)生理适应型(2)形态构件适应型(3)形态生理结合适应型。
2．选择4个光照强度(强光100％、中光50％、低光30％和弱光10％全光)，通过盆栽控制试验，从2005年6月到2006年3月测定了热带山地雨林5个典型树种幼苗对光的形态、生理和生长的适应特性。在形态适应上，4个光强下，黄桐(Endospermum chinense)、毛丹(Phoebe hungmoensis)对叶片投入较低，对茎投入较高；高枝杜英(Elaeocarpus dubius)、肖蒲桃(Acmena acuminatissima)对叶片投入较高，对茎、根投入较低；乐东拟单性木兰(Parakmeria lotungensis)对根投入最高。在生理适应上，黄桐、毛丹的最大净光合速率(A_(max))较高，高枝杜英、乐东拟单性木兰较低。在生长上，黄桐、毛丹、高枝杜英居中，肖蒲桃最高，乐东拟单性木兰最低。形态指标可塑性以黄桐最高，毛丹、肖蒲桃居中，而高枝杜英、乐东拟单性木兰较低。综合分析上述各项指标，认为黄桐、毛丹属于先锋功能群，乐东拟单性木兰属于顶级功能群，高枝杜英、肖蒲桃是介于上述二类之间的功能群(中期功能群)。
3．对不同演替阶段植物功能群幼苗不同光强下形态指标随月份变化的观测表明：在观察期间每个光强下，先锋功能群的叶面积、冠幅在雨季增加，在旱季增加缓慢；中期功能群是一直增加；顶级功能群变化很小。对于叶面积指数，先锋功能群逐渐降低，中期和顶级功能群变化不大。在不同光强之间，对于叶片数、叶面积，先锋功能群中的黄桐在弱光下最高，毛丹在弱光下最低；中期、顶级功能群在弱光下最低。对于冠幅，先锋和中期功能群在高光或中光下最低，顶级功能群在弱光下最低。对于叶面积指数，先锋功能群在强光下最低，中期、顶级功能群在强光下最高。
4．比较了不同演替阶段植物功能群幼苗在不同光强下的光合生理指标的月动态变化。在观察期间，雨季8月份所有功能群的净光合速率、蒸腾速率、水分利用效率在各个光强下都明显降低。气孔导度在6—12月份之间变化不大，而在3月份有一个明显的增加。在观察期间，先锋功能群的净光合速率在中光下最高；中期功能群在强光、中光、低光下数值接近，高于弱光下；顶级功能群在低光下最高。先锋功能群的光合能力高于顶级功能群，而水分利用能力低于顶级功能群，而中期功能群的各项生理指标更接近于先锋功能群。
5．不同演替阶段植物功能群幼苗在不同光强下的生长规律表明：在所有光强下，先锋功能群的相对生长速率均在雨季初最高，随后迅速下降，而在旱季接近于零。中期功能群在雨季和旱季均有较高的相对生长速率，而顶级功能群的相对生长速率最小。在整个生长期间，先锋功能群的相对生长速率高于顶级功能群。在旱季是中期功能群最高。先锋功能群的相对生长速率与叶面积增加、净光合速率密切相关；中期种功能群没有显著相关性。在高光和中光下，叶面积增加和净光合速率共同影响顶级功能群的相对生长速率，而在低光和弱光下叶面积增加则为主要影响因素。
6．不同演替阶段植物功能群幼苗在不同光强下雨季和旱季的光合参数日变化表明：在各个光强下，所有功能群的净光合速率、气孔导度、蒸腾速率在雨季的日变化过程中逐渐降低。在旱季，强光和中光下的所有功能群的净光合速率、气孔导度在12：00明显降低，而在低光、弱光下表现不明显；蒸腾速率的日变化曲线为单峰形，在中午12：00达到最高。而水分利用效率在雨季和旱季的日变化较小。
在不同光强之间，所有功能群在雨季净光合速率差异不显著，水分利用效率在弱光下最高；在旱季净光合速率在中光下最高，水分利用效率差异不显著。在同一光强下，在雨季和旱季先锋功能群比顶级功能群有着高的净光合速率日均值、气孔导度日均值和蒸腾速率日均值，中期功能群与先锋功能群接近。对于水分利用效率日均值，在雨季和旱季是先锋与顶级功能群接近，中期功能群最低。
先锋和顶级功能群的净光合速率日均值在雨季、旱季之间差异不显著，中期功能群是雨季比旱季低。对于水分利用效率日均值，先锋功能群旱季高于雨季，而中期和顶级功能群雨季、旱季差异不显著。
7．不同光照强度下不同演替阶段植物功能群在晴天、阴天下光合参数的日变化表明：所有功能群的净光合速率、气孔导度在强光、中光下在晴天中午12：00明显降低，在低光、弱光下表现不明显。蒸腾速率日变化为单峰形，中午12：00最高；水分利用效率日变化较小。阴天下，各个功能群的净光合速率在强光、中光、低光下为双峰形，分别在12：00和16：00；在弱光下则是逐渐降低。所有功能群的气孔导度和蒸腾速率日均值阴天下在各个光强均逐渐降低，水分利用效率变化不大。先锋功能群水分利用效率日均值在强光下晴天低于阴天，中期、顶级功能群差异不显著，在其它光强下所有功能群都差异不显著。
The tropical rain forest in Hainan Island of China is one of the most important forestecosystems that have the most complex structure and the richest biodiversity in China andpossesses the significant values of conservation and research. The tropical montane rain forestcovers the mountainous area froan 700 to 1300 m altitude, which has the broadest distributionarea. However, the excessive deforestation and unreasonable land use have led to the largereduction of the area of primary forest and formed the degraded ecosystems of differentsuccessional stages. The studies on the physical environments and the ecological adaptabilityof the plants in different successional stages can increase our knowledge of the successionalmechanism of forest ecosystem and is important to restore the ecological function andbiodiversity conservation of the rain forest on Hainan Island.
The ecological adaptability to light of the representative plants in different successionalstages were measured by the field and controlled experiments. These plants were divided intodifferent functional groups basing on their physiological and morphological characteristics.And the ecological adaptive modes and functions were studies. The main research results wereas follows.
1. In the early successional stage, environmental factors changed greatly and therepresentative plants had different adaptive manners. Thysanolaena maxima possessed strongerwater use ability, whose adaptation was mainly through the morphological architectureregulation strategy (by rolling of each leaf and shading of different leaves on the plant).Miscanthus floridulus had greater water regulation ability and its adaptation was mainlythrough the physiological regulation strategy (by higher photosynthetic capacity and water useefficiency (WUE)). However, Melastoma candidum and Melastoma sanguineum integrated themorphological architecture and physiological strategies (by high Pn and leaf self-shading).According to the ecophysiological characteristics and adaptation modes, the plants in the earlysuccussional stage of the tropical rain forest on Hainan Island can be categorized into threefunctional groups: (1) physiological adaptation group; (2) morphological adaptation group;(3) physiological and morphological integrated adaptation group.
2. The morphological, physiological and growth properties of seedlings of five dominanttropical montane rain forest tree species under four different relative irradiances (RI 100% (high light), 50% (moderate light), 30% (low light) and 10% (weak light)) were studied bythe pot experiments from June 2005 to March 2006. The six species exhibited differentmorphological adaptation modes to the variation of light intensities. Within each light regime,the investments of Endospermum chinense and Phoebe hungmoensis were lower in leaf andhigher in stem. Elaeocarpus dubius and Acmena acuminatissima had higher investments in leafand lower in stem. However, Parakmeria lotungensis had the highest root investment. On thephysiological adaptation, E. chinense and P. hungmoensis had higher maximum rate ofphotosynthesis (A_(max)) than E. dubius and P. lotungensis. In growth properties, in relativegrowth rate (RGR_M) A. acuminatissima was the highest, E. chinense, P. hungmoensis and E.dubius were intermediate, and P. lotungensis was the lowest. Furthermore, comparing in termsof morphological plasticity, E. ehinense was the highest, P hungmoensis and A. rostrata weremoderate, while E. dubius and P. lotungensis were the lowest. By integrating the three species'ecophysiological characteristics, light adaptation modes and principle component analysis ofmorphological plasticity, we recognized E. chinense and P. hungmoensis should belong to thepioneer functional group, E. dubius and A. acuminatissima should belong to mid-successionalfunctional group, and P. lotungensis should belong to shade- tolerant climax functional group.
3. There were different dynamic changes on morphological indexes of plant functionalgroups in different light regimes during the growth period. During the period, pioneerfunctional group increased quickly in the rainy season and not significantly in the dry seasonon total leaf area and crown area. For mid- successional functional group, LN, TLA and CAincreased linearly. However, for climax functional group showed little changes in the aboveindexes. In leaf area index, pioneer functional group decreased gradually, but for mid-successional and climax functional group, the changes were not distinct. Crown area (CA) forpioneer and mid-successional functional groups were the smallest in high or moderate light,but for climax functional group it was in weak light. In high light, leaf area index (LAI) wasthe lowest for pioneer functional group, but was the maximum for mid-successional and climaxfuctional groups.
4. The physiological characteristics of photosynthesis under different light regimes weremeasured every two months. Within each light regime, all species decreased distinctly inAugust on net photosynthesis (Pn), transpiration rate (E) and water use efficiency (WUE)under each light regime. However, stomatal conductance (g_s) for all species showed littlechanges from June to December and an apparent increase in March of the next year. Pioneer functional group had the highest Pn in moderate light. Pn which was close among high,moderate and low light was higher than weak light for mid-successional group. However,climax functional group possessed the highest Pn in low light. Comparing the climaxfunctional group, pioneer functional group exhibited higher photosynthetic capacity, but waslower in water use ability. And mid- successional functional group was close to the pioneerfunctional group in the above indexes.
5. The seedlings of plant functional groups in different successional stages had distinctgrowth patterns during the experiment period. Relative growth rate of pioneer functional groupdecreased quickly in the rainy season, but no significant changes and close zero in the dryseason. Mid-successional functional group showed large variety in RGR, and had obviousgrowth in the dry season. However, RGR for climax functional group changed little and waslowest. Within each light regime during the period, pioneer functional group had higher RGRthan climax P. lotungensis. And in the dry season, mid-successional functional group in RGRwas higher than other species. The results of the regression equations indicated RGR weresignificant correlative with Pn and the increment of leaf area in different light regimes forpioneer functional group. But for mid- successional functional group, RGR was not correlatedwith Pn and the increment of leaf area. However, climax functional group's RGR showedsignificant correlation with Pn and the leaf area increment in high and moderate light, but wasrelated with the leaf increment in the low and weak light.
6. Diurnal changes of photosynthetic parameters under clear days were monitored in therainy and dry season, respectively. In the rainy season within each light regime, all functionalgroups' diurnal changes in Pn, g_s and E decreased gradually as time increased, whereas WUEchanged little. In the dry season, the diurnal changes of all functional groups in Pn and g_sdecreased obviously at 12:00 under high and moderate light, but not in low and weak light. Ereached a maximum at 12:00 and than decline. However, WUE changed little during the day.
Among different light regimes, Pn of all functional groups changed not significantly in therainy season, and WUE were the highest in weak light. However, Pn were the greatest inmoderate light, but WUE showed no apparent changes. Within each light regime, pioneerfunctional group had the higher mean daily values of Pn, g_s, and E than climax functionalgroup both in the rainy and dry season, whereas mid-successional group was closer withpioneer functional group in the above indexes. But for the mean daily values of WUE, pioneerfunctional groups were near the climax functional group, whereas mid-successional functional group was the lowest.
The mean daily value of Pn for pioneer and climax functional groups showed nosignificant variations between in rainy and in dry season, but was lower for mid-successionalfunctional group in rainy than in dry season. Pioneer functional group had higher WUE in dryseason than in rainy season, whereas mid-successional and climax functional groups differednot significantly, which indicated pioneer functional group had stronger water regulativeability.
7. Diurnal changes of gas exchange were made in clear and overcast days of plantfunctional groups in different successioanl stages. In the clear days, Pn and g_s of all speciesdecreased obviously at the noon, and then increased in the afternoon in high and moderate light,whereas they were not significantly in low and weak light. E reached a maximum at 12:00 andthan decline. However, WUE changed little during the day. In the overcast days, diurnalchanges of Pn of functional groups were double unimodal pattern, and the peak values were12:00 and 16:00, respectively, whereas it decreased gradually. For g_s and E, all functionalgroups reduced stepwise in the overcast day, but WUE altered not obviously. The mean dailyvalue of Pn for pioneer functional group was higher in clear than in overcast days under highlight. Mid-successional group showed higher mean daily value of Pn in clear than in overcastdays under all light regimes. However, for climax functional group, the mean daily value of Pnwas higher under moderate and weak light in clear than in overcast days. For the mean dailyvalue of WUE, pioneer functional group was lower under high light in clear than in overcastdays, whereas it was not significant for mid-successional and climax functional group, whichindicated that pioneer functional group had stronger water regulative ability. In other lightregimes, all functional groups did not changed obviously.
本论文采用野外试验和控制试验相结合的研究手段，测定了热带山地雨林不同演替阶段中几个典型植物的光生态适应性，主要目的是试图从形态和生理生态的角度对不同演替阶段的植物进行功能群划分，了解不同演替阶段植物功能群在生态系统中的功能，分析植物功能群的生态适应性及其在演替过程中的作用。研究的主要结论如下：
1．通过野外现场测定，发现早期演替阶段环境因子变化剧烈，不同类型植物的适应方式存在差异。棕叶芦(Thysanolaena maxima)有着强的水分利用能力，偏重于以形态构件的变化(单个叶片的卷曲以及不同叶片之间的相互遮荫)来适应。五节芒(Miscanthus floridulus)有着强的水分控制能力，偏重以强的光合生理能力(通过高的光合能力和高的水分利用效率)来适应。野牡丹(Melastoma candidum)和毛稔(Melastoma sanguineum)的适应方式则是则是介于上述二者之间，即通过形态和生理特征变化的结合(形态上通过叶片的相互遮荫加上生理上高的光合能力)。基于植物的光合生理生态功能及适应方式，运用主成分分析方法，将海南岛热带山地雨林早期演替阶段的植物划分为三类大的功能群：(1)生理适应型(2)形态构件适应型(3)形态生理结合适应型。
2．选择4个光照强度(强光100％、中光50％、低光30％和弱光10％全光)，通过盆栽控制试验，从2005年6月到2006年3月测定了热带山地雨林5个典型树种幼苗对光的形态、生理和生长的适应特性。在形态适应上，4个光强下，黄桐(Endospermum chinense)、毛丹(Phoebe hungmoensis)对叶片投入较低，对茎投入较高；高枝杜英(Elaeocarpus dubius)、肖蒲桃(Acmena acuminatissima)对叶片投入较高，对茎、根投入较低；乐东拟单性木兰(Parakmeria lotungensis)对根投入最高。在生理适应上，黄桐、毛丹的最大净光合速率(A_(max))较高，高枝杜英、乐东拟单性木兰较低。在生长上，黄桐、毛丹、高枝杜英居中，肖蒲桃最高，乐东拟单性木兰最低。形态指标可塑性以黄桐最高，毛丹、肖蒲桃居中，而高枝杜英、乐东拟单性木兰较低。综合分析上述各项指标，认为黄桐、毛丹属于先锋功能群，乐东拟单性木兰属于顶级功能群，高枝杜英、肖蒲桃是介于上述二类之间的功能群(中期功能群)。
3．对不同演替阶段植物功能群幼苗不同光强下形态指标随月份变化的观测表明：在观察期间每个光强下，先锋功能群的叶面积、冠幅在雨季增加，在旱季增加缓慢；中期功能群是一直增加；顶级功能群变化很小。对于叶面积指数，先锋功能群逐渐降低，中期和顶级功能群变化不大。在不同光强之间，对于叶片数、叶面积，先锋功能群中的黄桐在弱光下最高，毛丹在弱光下最低；中期、顶级功能群在弱光下最低。对于冠幅，先锋和中期功能群在高光或中光下最低，顶级功能群在弱光下最低。对于叶面积指数，先锋功能群在强光下最低，中期、顶级功能群在强光下最高。
4．比较了不同演替阶段植物功能群幼苗在不同光强下的光合生理指标的月动态变化。在观察期间，雨季8月份所有功能群的净光合速率、蒸腾速率、水分利用效率在各个光强下都明显降低。气孔导度在6—12月份之间变化不大，而在3月份有一个明显的增加。在观察期间，先锋功能群的净光合速率在中光下最高；中期功能群在强光、中光、低光下数值接近，高于弱光下；顶级功能群在低光下最高。先锋功能群的光合能力高于顶级功能群，而水分利用能力低于顶级功能群，而中期功能群的各项生理指标更接近于先锋功能群。
5．不同演替阶段植物功能群幼苗在不同光强下的生长规律表明：在所有光强下，先锋功能群的相对生长速率均在雨季初最高，随后迅速下降，而在旱季接近于零。中期功能群在雨季和旱季均有较高的相对生长速率，而顶级功能群的相对生长速率最小。在整个生长期间，先锋功能群的相对生长速率高于顶级功能群。在旱季是中期功能群最高。先锋功能群的相对生长速率与叶面积增加、净光合速率密切相关；中期种功能群没有显著相关性。在高光和中光下，叶面积增加和净光合速率共同影响顶级功能群的相对生长速率，而在低光和弱光下叶面积增加则为主要影响因素。
6．不同演替阶段植物功能群幼苗在不同光强下雨季和旱季的光合参数日变化表明：在各个光强下，所有功能群的净光合速率、气孔导度、蒸腾速率在雨季的日变化过程中逐渐降低。在旱季，强光和中光下的所有功能群的净光合速率、气孔导度在12：00明显降低，而在低光、弱光下表现不明显；蒸腾速率的日变化曲线为单峰形，在中午12：00达到最高。而水分利用效率在雨季和旱季的日变化较小。
在不同光强之间，所有功能群在雨季净光合速率差异不显著，水分利用效率在弱光下最高；在旱季净光合速率在中光下最高，水分利用效率差异不显著。在同一光强下，在雨季和旱季先锋功能群比顶级功能群有着高的净光合速率日均值、气孔导度日均值和蒸腾速率日均值，中期功能群与先锋功能群接近。对于水分利用效率日均值，在雨季和旱季是先锋与顶级功能群接近，中期功能群最低。
先锋和顶级功能群的净光合速率日均值在雨季、旱季之间差异不显著，中期功能群是雨季比旱季低。对于水分利用效率日均值，先锋功能群旱季高于雨季，而中期和顶级功能群雨季、旱季差异不显著。
7．不同光照强度下不同演替阶段植物功能群在晴天、阴天下光合参数的日变化表明：所有功能群的净光合速率、气孔导度在强光、中光下在晴天中午12：00明显降低，在低光、弱光下表现不明显。蒸腾速率日变化为单峰形，中午12：00最高；水分利用效率日变化较小。阴天下，各个功能群的净光合速率在强光、中光、低光下为双峰形，分别在12：00和16：00；在弱光下则是逐渐降低。所有功能群的气孔导度和蒸腾速率日均值阴天下在各个光强均逐渐降低，水分利用效率变化不大。先锋功能群水分利用效率日均值在强光下晴天低于阴天，中期、顶级功能群差异不显著，在其它光强下所有功能群都差异不显著。
The tropical rain forest in Hainan Island of China is one of the most important forestecosystems that have the most complex structure and the richest biodiversity in China andpossesses the significant values of conservation and research. The tropical montane rain forestcovers the mountainous area froan 700 to 1300 m altitude, which has the broadest distributionarea. However, the excessive deforestation and unreasonable land use have led to the largereduction of the area of primary forest and formed the degraded ecosystems of differentsuccessional stages. The studies on the physical environments and the ecological adaptabilityof the plants in different successional stages can increase our knowledge of the successionalmechanism of forest ecosystem and is important to restore the ecological function andbiodiversity conservation of the rain forest on Hainan Island.
The ecological adaptability to light of the representative plants in different successionalstages were measured by the field and controlled experiments. These plants were divided intodifferent functional groups basing on their physiological and morphological characteristics.And the ecological adaptive modes and functions were studies. The main research results wereas follows.
1. In the early successional stage, environmental factors changed greatly and therepresentative plants had different adaptive manners. Thysanolaena maxima possessed strongerwater use ability, whose adaptation was mainly through the morphological architectureregulation strategy (by rolling of each leaf and shading of different leaves on the plant).Miscanthus floridulus had greater water regulation ability and its adaptation was mainlythrough the physiological regulation strategy (by higher photosynthetic capacity and water useefficiency (WUE)). However, Melastoma candidum and Melastoma sanguineum integrated themorphological architecture and physiological strategies (by high Pn and leaf self-shading).According to the ecophysiological characteristics and adaptation modes, the plants in the earlysuccussional stage of the tropical rain forest on Hainan Island can be categorized into threefunctional groups: (1) physiological adaptation group; (2) morphological adaptation group;(3) physiological and morphological integrated adaptation group.
2. The morphological, physiological and growth properties of seedlings of five dominanttropical montane rain forest tree species under four different relative irradiances (RI 100% (high light), 50% (moderate light), 30% (low light) and 10% (weak light)) were studied bythe pot experiments from June 2005 to March 2006. The six species exhibited differentmorphological adaptation modes to the variation of light intensities. Within each light regime,the investments of Endospermum chinense and Phoebe hungmoensis were lower in leaf andhigher in stem. Elaeocarpus dubius and Acmena acuminatissima had higher investments in leafand lower in stem. However, Parakmeria lotungensis had the highest root investment. On thephysiological adaptation, E. chinense and P. hungmoensis had higher maximum rate ofphotosynthesis (A_(max)) than E. dubius and P. lotungensis. In growth properties, in relativegrowth rate (RGR_M) A. acuminatissima was the highest, E. chinense, P. hungmoensis and E.dubius were intermediate, and P. lotungensis was the lowest. Furthermore, comparing in termsof morphological plasticity, E. ehinense was the highest, P hungmoensis and A. rostrata weremoderate, while E. dubius and P. lotungensis were the lowest. By integrating the three species'ecophysiological characteristics, light adaptation modes and principle component analysis ofmorphological plasticity, we recognized E. chinense and P. hungmoensis should belong to thepioneer functional group, E. dubius and A. acuminatissima should belong to mid-successionalfunctional group, and P. lotungensis should belong to shade- tolerant climax functional group.
3. There were different dynamic changes on morphological indexes of plant functionalgroups in different light regimes during the growth period. During the period, pioneerfunctional group increased quickly in the rainy season and not significantly in the dry seasonon total leaf area and crown area. For mid- successional functional group, LN, TLA and CAincreased linearly. However, for climax functional group showed little changes in the aboveindexes. In leaf area index, pioneer functional group decreased gradually, but for mid-successional and climax functional group, the changes were not distinct. Crown area (CA) forpioneer and mid-successional functional groups were the smallest in high or moderate light,but for climax functional group it was in weak light. In high light, leaf area index (LAI) wasthe lowest for pioneer functional group, but was the maximum for mid-successional and climaxfuctional groups.
4. The physiological characteristics of photosynthesis under different light regimes weremeasured every two months. Within each light regime, all species decreased distinctly inAugust on net photosynthesis (Pn), transpiration rate (E) and water use efficiency (WUE)under each light regime. However, stomatal conductance (g_s) for all species showed littlechanges from June to December and an apparent increase in March of the next year. Pioneer functional group had the highest Pn in moderate light. Pn which was close among high,moderate and low light was higher than weak light for mid-successional group. However,climax functional group possessed the highest Pn in low light. Comparing the climaxfunctional group, pioneer functional group exhibited higher photosynthetic capacity, but waslower in water use ability. And mid- successional functional group was close to the pioneerfunctional group in the above indexes.
5. The seedlings of plant functional groups in different successional stages had distinctgrowth patterns during the experiment period. Relative growth rate of pioneer functional groupdecreased quickly in the rainy season, but no significant changes and close zero in the dryseason. Mid-successional functional group showed large variety in RGR, and had obviousgrowth in the dry season. However, RGR for climax functional group changed little and waslowest. Within each light regime during the period, pioneer functional group had higher RGRthan climax P. lotungensis. And in the dry season, mid-successional functional group in RGRwas higher than other species. The results of the regression equations indicated RGR weresignificant correlative with Pn and the increment of leaf area in different light regimes forpioneer functional group. But for mid- successional functional group, RGR was not correlatedwith Pn and the increment of leaf area. However, climax functional group's RGR showedsignificant correlation with Pn and the leaf area increment in high and moderate light, but wasrelated with the leaf increment in the low and weak light.
6. Diurnal changes of photosynthetic parameters under clear days were monitored in therainy and dry season, respectively. In the rainy season within each light regime, all functionalgroups' diurnal changes in Pn, g_s and E decreased gradually as time increased, whereas WUEchanged little. In the dry season, the diurnal changes of all functional groups in Pn and g_sdecreased obviously at 12:00 under high and moderate light, but not in low and weak light. Ereached a maximum at 12:00 and than decline. However, WUE changed little during the day.
Among different light regimes, Pn of all functional groups changed not significantly in therainy season, and WUE were the highest in weak light. However, Pn were the greatest inmoderate light, but WUE showed no apparent changes. Within each light regime, pioneerfunctional group had the higher mean daily values of Pn, g_s, and E than climax functionalgroup both in the rainy and dry season, whereas mid-successional group was closer withpioneer functional group in the above indexes. But for the mean daily values of WUE, pioneerfunctional groups were near the climax functional group, whereas mid-successional functional group was the lowest.
The mean daily value of Pn for pioneer and climax functional groups showed nosignificant variations between in rainy and in dry season, but was lower for mid-successionalfunctional group in rainy than in dry season. Pioneer functional group had higher WUE in dryseason than in rainy season, whereas mid-successional and climax functional groups differednot significantly, which indicated pioneer functional group had stronger water regulativeability.
7. Diurnal changes of gas exchange were made in clear and overcast days of plantfunctional groups in different successioanl stages. In the clear days, Pn and g_s of all speciesdecreased obviously at the noon, and then increased in the afternoon in high and moderate light,whereas they were not significantly in low and weak light. E reached a maximum at 12:00 andthan decline. However, WUE changed little during the day. In the overcast days, diurnalchanges of Pn of functional groups were double unimodal pattern, and the peak values were12:00 and 16:00, respectively, whereas it decreased gradually. For g_s and E, all functionalgroups reduced stepwise in the overcast day, but WUE altered not obviously. The mean dailyvalue of Pn for pioneer functional group was higher in clear than in overcast days under highlight. Mid-successional group showed higher mean daily value of Pn in clear than in overcastdays under all light regimes. However, for climax functional group, the mean daily value of Pnwas higher under moderate and weak light in clear than in overcast days. For the mean dailyvalue of WUE, pioneer functional group was lower under high light in clear than in overcastdays, whereas it was not significant for mid-successional and climax functional group, whichindicated that pioneer functional group had stronger water regulative ability. In other lightregimes, all functional groups did not changed obviously.
引文
[1] 曾庆波，李意德，陈步峰等．热带森林生态系统研究与管理．北京：中国林业出版社，1997．
[2] 胡玉佳，李玉杏．海南岛热带雨林．广州：广东高等教育出版社，1992．
[3] Larcher W. Physiological plant ecology-ecophysiology and stress physiology of functional groups. New York: Springer-Verlag, 1995.
[4] Wilson EO. The diversity of life. Cambridge: The Belknap Press of Harvard University Press, 1992, 183-211.
[5] Robert C, Ralph D, Rudolf DG et al. The value of the world's ecosystem services and natural capital. Nature, 1997, 387: 253-260.
[6] Holdgate M. Conservation in a changing world. IUCN Bulletin, 1989, (10-12): 16-25.
[7] Williams FL. Reflection on the tropical deforestation crisis Biological Conservation. 1999, 91: 109-117.
[8] 颜文洪．全球性热带雨林的破碎化趋势与保护分析．自然生态保护，2003，2：34-36．
[9] 王献溥，崔国发，李俊清．海南生物多样性研究展望．北京林业大学学报，2000(4)：132-134．
[10] 李炎香．海南岛热带森林的开发利用及其在生态平衡中的作用．热带林业科技，1980，(3)：35-44．
[11] 吴华盛．海南省热带雨林的保护和利用．热带林业，2002，30(2)：4-5．
[12] 曹建华，蒋菊生．海南生物多样性保护探析．华南热带农业大学学报，2006，12(2)：34-38．
[13] 臧润国，安树青，陶建平等．海南岛热带林生物多样性维持机制．北京：科学出版社，2004，198-210．
[14] 陈德祥，李意德，骆土寿等．海南岛尖峰岭热带山地雨林下层乔木中华厚壳桂光合生理生态特性的研究．林业科学研究，2003，16(5)：540-47．
[15] 陈德祥，李意德，骆土寿等．短期CO_2浓度升高对雨林树种盘壳栎光合特性的影响．生态学报，2004，24(8)：1622-1628．
[16] 骆土寿，李意德，陈德祥等．热带雨林恢复演替中典型树种黄桐气体交换对环境的响应．生态学报，2003，23(9)：1762-1772．
[17] Wardle DA, Bardgett RD, Klironomos JN et al. Ecological linkages between aboveground and belowground biota Science, 2004, 304: 1639-1633.
[18] Odum EP. The strategy of ecosystem development. Science, 1969, 164: 262-270.
[19] Gurevitch J, Scheiner SM, Fox GA. The ecology of plants. Sinauer Associates Inc., Sunderland, Massachusetts U.S.A., 2002, 253-276.
[20] Bazzaz FA. The physiological ecology of plant succession. Annual Review of Ecology and Systematics, 1979, 10: 351-371.
[21] Kobe RK, Pacala W, Silander JA. Juvenile tree survivorship as a component of shade tolerance. Ecological Applications, 1995, 5: 517-532.
[22] Koike F. Plant traits as predictors of woody species dominance in climax forest in northern Japan. Journal of Ecology, 2001, 12: 327-336.
[23] Walters MB, Reich PB. Are shade tolerance, survival, and growth linked? Low light and nitrogen effects on hardwood seedlings. Ecology, 1996, 77(3): 841-853.
[24] Poorter L, Arets EJMM. Light environment and tree strategies in a Bolivian tropical moist forest: an evaluation of the light partitioning hypothesis. Plant ecology, 2003, 166: 295-306.
[25] Agyeman VK, Swaine MD, Thompson J. Responses of tropical forest seedlings to irradiance and the derivation of a light response index. Journal of ecology, 1999, 87: 815-827.
[26] Kyereh B, Swaine MD, Thompson J. Effect of light on the germination of forest trees in Ghana. Journal of ecology, 1999, 87: 772-783.
[27] Denslow JS, Schultz JC, Vitousek PM et al. Growth responses of tropical shrubs to treefall gap environments. Ecology, 1990, 71(1): 165-179.
[28] Raynal DJ, Bazzaz FA. Establishment of early successional plant populations on forest and prairie soil. Ecology, 1973, 54: 1335-1341.
[29] 温远光，梁宏温，和太平等．大明山退化生态系统群落光照的变化．广西农业大学学报，1998，17(2)：199-203．
[30] Chazdon RL, Fetcher N. Photosynthetic light environments in a lowland tropical rain forest in Costa Rica. Journal of Ecology, 1984, 72: 553-564.
[31] Endler JA. The color of light in forests andits implications. Ecological.Monograph, 1993, 63: 1-27.
[32] Lee DW, Baskaran K, Mansor M et al. Irradiance and spectral quality affect Asian tropical rain forest tree seedling development. Ecology, 1996, 77: 568-580.
[33] Pearcy RW. Sunflecks and photosynthesis in plant canopies. Annual Review of Plant Physiology and Plant Molecular Biology, 1990, 41: 421-4531
[34] Watling JR, Robinson SA, Woodrow IE, Osmond CB. Responses of rainforest understoey plants to excess light during sunflecks. Australian Journal of Plant Physiology, 1997, 24, 17-25.
[35] Leakey ADB, Press MC, Scholes JD. High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling. Plant, Cell and Environment, 2003, 26: 1681-1690.
[36] Watt AS. Pattern and process in the plant community. Journal of ecology, 1947, 35: 1-22.
[37] Sanford PJ, Braker HE, Hartshom GS. Canopy openings in a primary neotropical lowland forest. Journal of Tropical Ecology, 1986, 2, 277-282.
[38] 蒋有绪，王伯荪，臧润国等．海南岛热带林生物多样性及其形成机制．北京：科学出版社，2002，1-30．
[39] 蒋有绪，卢俊培．中国海南岛尖峰岭热带林生态系统．北京：科学出版社，1991，256-283．
[40] 张一平，王进欣，马友鑫等．西双版纳雾凉季次生林林窗光照特征初步分析．热带气象学报，2000，16(4)：374-379．
[41] 张一平，于进欣，马友鑫等．西双版纳热带次生林林窗近地层温度时空分布特征．林业科学，2002，38(6)：1-5．
[42] 咎启杰，李鸣光，张志权等．林窗及其在森林动态中的作用．植物学通报，1997，14(增刊)：18-24．
[43] Brokaw VLB. Species composition in gaps and structure of a tropical forest. Ecology, 1989, 70: 538-541.
[44] Swaine MD, Whitmore TC. On the definition of ecological species groups in tropical rain forest. Vegetatio, 1988, 75: 8t-86.
[45] Hawthorne, WD. Ecological profiles of Ghanaian forest trees. Tropical Forestry Papers 29. Oxford Forestry Institute, Oxford, 1995.
[46] Zimmerman JK, Pascarella JB, Aide TM. Barriers to forest regeneration in an abandoned pasture in Puerto Rico. Restoration ecology, 2000, 8(4): 350-360.
[47] Hooper H, Legendre P, Condit R. Barriers to forest regeneration of deforested and abandoned land in Panama. Journal of applied ecology, 2005, 42: 1165-1174
[48] Van der Marrel E. Vegetation ecology. Blackwell Publishing, Oxford, 2005, 172-198.
[49] Kuppers M. Canopy gaps: competitive light interception and economic space filling-a matter of whole plant allocation. In: Caldwell, MM and Pearcy RW, eds. Exploitation of Environmental Heterogeneity by Plants. San Diego: Academic Press, 1994, 111-114.
[50] Pugnaire FI, Valladares F. Handbook of Functional Plant Ecology. Marcel Eekker, Inc. 1999, 121-194.
[51] Edwards EJ, Benham DG, Marland LA et al. Root production is determined by radiation flux in a temperate grassland community. Global change biology, 2004, 10: 209-227.
[52] Pons TL. An ecophysiological study in the field layer of ash coppice, Ⅱ. Experiments with Geum urbanum and Cirsium palustre in different light intensities. Acta Boanicat Neerlandica, 1977, 26: 29-42.
[53] Oconnell BM, Kelty MJ. Crown architecture of understory and open-grown white pine (Pinus strobes L.) saplings. Tree Physiology, 1994, 14: 89-102.
[54] Poorter L. Growth responses of 15 rainforest tree species to a light gradient: the relative importance of morphological and physiological traits. Functional Ecological, 1999, 13:396-410.
[55] Reich PB, Tjoelker MG, Walters MB et al. Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedlings of nine boreal tree species grown in high and low light. Functional Ecology, 1998, 12:327-338.
[56] Huante, P. and E. Rincon. Responses to light changes in tropical deciduous woody seedlings with contrasting growth rates. Oecologia, 1998, 113:53-66.
[57] Givnish, TJ. Adaptation to sun and shade: a whole-plant perspective. Australlian Journal of Plant Physiology, 1988, 15:63-92.
[58] Mori A, Takeda H. Functional relationships between crown morphology and with-crown characteristics of understory saplings of three codominant conifers in a subalpine forest in central Japan. Tree Physiology, 2004, 24: 661-670.
[59] Kobe RK. Carbohydrate allocation to storagte as a basis of interspecific variation in sapling survivorship and growth. Oikos, 1997, 80: 226-233.
[60] Waiters MB, Reich PB. Low-light carbon balance and shade tolerance in the seedling of woody plants: do winter deciduous and broad-leaved evergreen species differ? New Phytologist, 1999, 143: 143-154.
[61] Kitajima K. Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees. Oecologia, 1994, 98: 419-428.
[62] Montgomery R. Relative importance of photosynthetic physiology and biomass allocation for tree seedling growth across a broad light gradient. Tree Physiology, 2004, 24: 155-167.
[63] Delagrange S, Messier C, Lechowicz MJ et al. Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability. Tree Physiology, 2004, 24: 775-784.
[64] Beaudet M, Messier C. Growth and morphological response of yellow birch, sugar maple, and beech seedlings growing under a natural light gradient. Canadian Joumal of Forest Research, 1998, 28: 1007-1015.
[65] Gibson AC. Photosynthetic organs of desert plants. Bio Science, 1998, 48:911-920.
[66] Bloor JMG, Grubb P. Growth and mortality in high and low light: trends among 15 shade-tolerant tropical rain forest tree species. Journal of Ecology, 2003, 91: 77-85.
[67] Vilela AE, Rennella M J, Ravetta DA. Responses of tree-type and shrub-type Prosopis (Mimosaceae) taxa to water and nitrogen availabilities. Forest Ecology and Management, 2003, 186: 327-337.
[68] Augspurger CK. Light requirements of neotropical tree seedlings: a comparative study of growth and survival. Journal of Ecology, 1984, 72: 777-795.
[69] Lusk CH. Leaf area accumulation helps juvenile evergreen trees tolerance shade in a temperate rainforest. Oecologia, 2002, 132:188-196.
[70] Cornelissen JHC. Aboveground morphology of shade-tolerant Castanopsis fargesii samplings in response to light environment. International Journal of Plant Sciences, 1993, 154 (4): 481-495.
[71] Valladares F, Pearcy RW. The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Caliafornian chaparral shrub. Oecologia, 1998, 114: 1-10.
[72] Pearcy RW, Muraoka H, Valladares F. Crown architecture in sun and shade environments: assessing function and trade-offs with a threedimensionai simulation model. New Phytologist, 2005, 166: 791-800.
[73] Werk KS, Ehleringer J. Non-random leaf orientation in Lactuca serriola L. Plant, Cell and Environment, 1984, 7: 81-87.
[74] Smith M, Ullberg D. Effect of leaf angle and orientation on photosynthesis and water relations in Silphium terebinthinaceum. American Journal of Botany, 1989, 76: 1714-1719.
[75] Valladares F, Pearcy RW. Interactions between water stress, sun-shade acclimation, heat tolerance and photoinhibition in the sclerophyll Heteromeles arbutifolia. Plant, Cell and Environment, 1997, 20: 25-36.
[76] Runkle JR. Pattern of disturbance in some old-growth mesic forests of eastern North America. Ecology, 1982, 63: 1533-1546.
[77] Sipe TW, Bazzaz FA. Gap partitioning among maples (Acer) in central New-England - shoot architecture and photosynthesis. Ecology, 1994, 5:2318-2332.
[78] Ackerly DD, Bazzaz FA. Seedling crown orientation and interception of diffuse radiation in tropical forest gaps. Ecology, 1995, 76:1134-1146.
[79] Delagrange S, Montpied P, Dreyer E et al. Does shade improve light interception efficiency? A comparison among seedlings from shade-tolerant and -intolerant temperate deciduous tree species. New Phytologist, 2006, 172: 293-304.
[80] King DA. Allometry of saplings and understorey trees of a Panamanian forest. Functional Ecology, 1990, 4: 27-32.
[81] Kohyama T, Hotta M. Significance of allometry in tropical saplings. Functional Ecology, 1990, 4: 515-521.
[82] Valladares F, Skillman JB, Pearcy RW. Convergence in light capture efficiencies among tropical forest understory plants with contrasting crown architectures: a case of morphological compensation. American Journal of Botany, 2002, 89: 1275-1284.
[83] Pearcy RW, Valladares F, Wright SJ et al. A functional analysis of the crown architecture of tropical forest Psychotria species: do species vary in light capture efficiency and consequently in carbon gain and growth? Oecologia, 2004, 139:163-177.
[84] Horton, P., A. V. Ruban, & R. G. Walters. Regulation of light harvesting in green plants. Annual Review of Plant Physiology and Plant Molecular Biology, 1996, 47: 655-684.
[85] Osmond CB, Grace SC. Perspectives on photoinhibition and photorespiration in the field: quintessential inefficiencies of the light and dark reactions of photosynthesis? Journal of Experimental Botany, 1995, 46: 1351-1362.
[86] Demmig-Adams B, Adams Ⅲ WW. Photoprotection and other responses of Plants to high light stress. Annual Review of Plant Physiology and Plant Molecular Biology, 1992, 43: 599-626.
[87] Demmig-Adams B, Adams WW. The role of xanthophyll cycle carotenoids in the protection of photosynthesis. Trends in Plant Science, 1996, 1: 21-26.
[88] Niyogi KK. Photoprotection revisited: Genetic and molecular approaches. Annual Review of Plant Physiology and Plant Molecular Biology, 1999, 50: 333-359.
[89] Schulze E-D, Lange OL, Evenari M et al. Long-term effects of drought on wild and cultivated plants in the Negev desert. Ⅱ. Diurnal patterns of net photosynthesis and daily carbon gain. Oecologia, 1980, 45:19-25.
[90] Tenhunen JD, Lange OL, Braun M. Midday stomatal closure in Mediterranean type sclerophylls under simulated habitat conditions in an environmental chamber. Ⅱ. Effect of the complex of leaf temperature and air humidity on gas exchange of Arbutus unedo and Qurecus ilex. Oecologia, 1981, 50:5-11.
[91] Tenhunen JD, Lange OL, Jahner D. The control by atmospheric factors and water stress of midday stomatal closure in Arbutus unedo growing in a natural macchia Oecologia, 1982, 55:165-169.
[92] Tenhunen JD, Lange OL, Geble J et al. Changes in photosynthetic capacity, carboxylation efficiency, and CO_2 compensation point associated with miday stomatal closure and midday depression of net CO_2 exchange of leaves of Quercus suber. Planta, 1984, 162:193-203.
[93] Chaves, MM, Harley PC, Tenhunen JD et al. Gas exchange studies in two Portuguese grapevine cultivars. Physilolgia Plantarum, 1987, 70: 639-647.
[94] Downton, WJS, Grant WJR, Loveys BR. Diurnal changes in the photosynthesis of field-grown grape vines. New Phytologist, 1987, 105: 71-80.
[95] Correia MJ, Chaves MM, Pereira JS. Afternoon depression in photosynthesis in grapevine leavesevidence for a high light stress effect. Journal of Experimental Botany, 1990, 41: 417-426.
[96] Epron D, Dreyer E, Breda N. Photosynthesis of oak trees (Quercus petraea (Matt.) Liebl.) during drought under field conditions: diuranal course of net CO_2 assimilation and photochemical efficiency of photosystem Ⅱ. Plant, Cell and Environment, 1992, 15: 809-820.
[97] Cowan, LR. Regulation of water use in relation to carbon gain in higher plants. In Encyclopedia of Plant Physilolgy NS, Physiological Plant Ecology Ⅱ (eds O. L. L ange, P. S. Nobel, C. B. Osmond & H. Ziegler). Springer verlag, Berlin, 1982.
[98] Tyree MT, Sperry JS. Do woody plants operate near the point of catastrophic xylem dysfunction caused by dynamic water stress? Answers from a model. Plant Physiology, 1988, 88: 574-580.
[99] Jones HG, R.A. Sutherland. Stomatal control of xylem embolism. Plant, Cell and Environment, 1991, 14: 607-612.
[100] Sperry JS. Hydraulic constraints on plant gas exchange. Agricultural and Forest Meteorology, 2000, 104: 13-23.
[101] Muraoka H, Tang Y, Terashima I et al. Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light. Plant, Cell and Environment, 2000, 23: 235-250.
[102] Medrano H, Bota J, Abadia A et al. Effects of drought on light-energy dissipation mechanisms in high-light-acclimated, field-grown grapevines. Functional Plant Biology, 2002, 29:1197-1207.
[103] Ducrey M. Influence of shade on photosynthetic gas exchange of 7 tropical rainforest species from Guadeloupe (French-West-Indies). Ann. Sci. For., 1994, 51: 77-94.
[104] Oberbauer SF, Strain BR. Effect of canopy position and irradiance on leaf physiology and morphology of Penthaclethra macroloba (Mimosaceae). American Journal of Botany, 1985, 73: 409-416.
[105] Chiariello NR, Field CB, Mooney HA. Midday wilting in a tropical pioneer tree. Functional Ecology, 1987, 1: 3-11.
[106] Poorter L, Oberbauer SF. Photosynthetic induction responses of two rainforest tree species in relation to light environment. Oecologia, 1993, 96: 193-199.
[107] Walters MB, Field CB. Photosynthetic light acclimation in two rainforest Piper species with different ecological amplitudes. Oecologia, 1987, 72: 449-456.
[108] Sims DA, Pearcy RW. Photosynthesis and respiration in Alocasia macrorrhiza following transfers to high and low light. Oecologia, 1991, 86: 447-453.
[109] Strauss-Dibenedetti S Bazaz FA. Plasticity and acclimataion to light in tropical Moraceae of different successional positions. Oecologia, 1991, 87: 377-387.
[110] Kamaluddin M, Grace J. Acclimation in seedlings of a tropical tree, Bischofia javanica, following a stepwise reduction in light. Annals of Botany, 1992, 69: 557-562.
[111] 冯玉龙，草坤芳，冯志立等．四种热带雨林树种幼苗比叶重，光合特性和暗呼吸对生长光环境的适应．生态学报，2002，22(6)：901-910．
[112] 齐欣，曹坤芳，冯玉龙．热带雨林蒲桃属3个树种的幼苗光合作用对生长光强的适应．植物生态学报，2004，28(1)：31-38．
[113] 张教林，曹坤芳．不同生态习性热带雨林树种的幼苗对光能的利用与耗散．应用生态学报，2004，15(3)：372-376．
[114] Chow WS, AdamsonHY, Anderson JM. Photosynthetic acclimation of Tradescantia albiflora to growth irradiance: Lack of adjustment of light-harvesting components and tis consequences. Physiologia Plantarum, 1991, 81: 175-182.
[115] 孙谷畴，赵平，曾小平．两种木兰科植物叶片光合作用的光驯化．生态学报，2004，24(6)：1111-1117．
[116] 梁春，林植芳，孔国辉．不同光强下生长的亚热带树苗的光合—光响应特性的比较．应用生态学报，1997，8(1)：7-11．
[117] Kursar TA, Coley PD. Photosynthetic induction times in shade-tolerant species with long and short-leaved leaves. Oecologia, 1997, 93: 165-170.
[118] Lusk CH, Pozo AD. Survival and growth of seedlings of 12 Chilean rainforest trees in two light environments: Gas exchange and biomass distribution correlates. Austral Ecology, 2002, 27:173-182.
[119] Lusk CH, Reich PB. Relationships of leaf dark respiration with light environment and tissue nitrogen content in juveniles of 11 cold-temperate tree species. Oecologia, 2000, 123:318-329.
[120] Grime JP. Shade tolerance in flowering plants. Nature, 1965, 208: 161-163.
[121] Loach K. Shade tolerance in flowering plants. 1. Leaf photosynthesis and respiration in plants raised under artificial shade. New Phytologist, 1967, 66:607-21.
[122] Huc R, Ferhi A, Guehl JM. Pioneer and late stage tropical rainforest tree species (French Guyana) growing under common conditions differ in leaf gas exchange regulation, carbon isotope discrimination and leaf water potential Oecologia, 1994, 99: 297-305.
[123] Bonal D, Barigah TS, Granier A, Guehl JM. Late-stage canopy tree species with extremely low δ~(13)C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana Plant, Cell and Environment, 2000, 23: 445-459.
[124] Ishida A, Nakano T, Matsumoto Y et al. Diurnal changes in leaf gas exchange and chlorophyll fluorescence in tropical tree species with contrasting light requirements. Ecological Research, 1999, 14: 77-88.
[125] Hanba YT, Kogami H, Terashima I. The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand. Plant, Cell and Environment, 2002, 25: 1021-1030.
[126] Pedersen BS. The role of stress in the mortality of Midwestern oaks as indicated by growth prior to death. Ecology, 1998, 79: 79-93.
[127] Sack L, Grubb PJ. Why do species of woody seedlings change rank in relative growth rate between low and high irradiance? Functional Ecological, 2001, 15:145-154.
[128] Pompa J, Bongers F. The effect of canopy gaps on growth and morphology of seedlings of rainforest species. Oecologia, 1988, 75: 625-632.
[129] Rice SA, Bazzaz FA. Growth consequences of plasticity of plant traits in response to light conditions. Oecologia, 1989, 78:508-512.
[130] Pompa J, Bongers F. Acclimation of seedlings of three Mexican tropical rain forest tree species to a change in light availability. Journal of Tropical Ecology, 1991, 7:85-97.
[131] Ackerly, DD. Phenotypic plasticity and the scale of environmental heterogeneity: studies of tropical pioneer trees in variable light environments PhD thesis, Harvard University, Massachusetts, 1993.
[132] Newell EA, McDonald EP, Strain, BR et al. Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest. Oecologia, 1993, 94:49-56.
[133] Tumbull MH, Doley D, Yates DJ. The dynamic of photosynthetic acclimation to changes in light quantity and quality in three Ausftralian rainforest tree species Oecologia, 1993, 94:218-228.
[134] Niinemets U. The controversy over traits conferring shade-tolerance in trees: ontogenetic changes revisited. Journal of Ecology, 2006, 94: 464-470.
[135] Veneklaas EJ, Poorter L. Growth and carbon partitioning of tropical tree seedlings in contrasting light environments. In: Inherent Variation in Plant Growth: Physiological Mechanisms and Ecological Consequences (eds H. Lambers, H. Poorter & M. Van Vuuren). Backhuys Publishers, Leiden, Netherlands, 1998: 337-62.
[136] Reich PB, Walters MB, Ellsworth DS. Leaf lifespan in relation to leaf, plant and stand characteristics among diverse ecosystems. Ecological Monographs, 1992, 62: 365-392.
[137] King DA. Influence of light level on the growth and morphology of saplings in a Panamanian forest American. Journal of Botany, 1994, 81: 948-957.
[138] Leishman MR, Westoby M. The role of large seed size in shaded conditions, experimental evidence. Functional Ecology, 1994, 8:205-214.
[139] Boot, RGA. The significance of seedling size and grown rate of tropical rainforest tree seedlings for regeneration in canopy openings. Ecology of Tropical Forest Tree Seedlings (ed. M.D. Swaine), 1996, 267-284.
[140] Saverimuttu T, Westoby M. Seedling longevity under deep shade~ in relation to seed size. Jourual of Ecology, 1996, 84: 681:689.
[141] Walters MB, Reich PB. Seed size, nitrogen supply and growth rate affect tree seedling survival in deep shade. Ecology, 2000, 81: 1887-1901.
[142] Pintado A, Valladares F, Sancho LG. Exploring phenotypic plasticity in the lichen Ramalina capitata: morphology, water relations and chlorophyll content in North and South-facing populations. Annals of Botany, 1997, 80: 345-353.
[143] Valladares F, Wright SJ, Lasso E et al. Plastic phenotypic response to light of 16 congeneric shrubs from a Panamanian rainforest. Ecology, 2000, 81 (7): 1925-1936.
[144] Bjorkman O. Responses to different quantum flux densities. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Physiological plant ecology 1. Responses to the physical environment. Springer, Berlin, Heidelberg, New York, 1981, 57-107.
[145] Tumbull MH. The effect of light quantity and quality during development on the photosynthetic characteristics of six Australian rainforest tree species. Oecologia, 1991, 87:110-117.
[146] Chazdon, RL. Photosynthetic plasticity of two rain forest shrubs across natural gap transects. Oecologia, 1992, 92:586-595.
[147] Canham, CD. Different responses to gaps among shade-tolerant species. Ecology, 1989, 70:548-550.
[148] Poorter L, Bongers F, Sterck FJ et al. Beyond the regeneration phase: differentiation of height-light trajectories among tropical tree species. Journal of Ecology, 2005, 93: 256-267.
[149] Niinemets U, Sparrow A, Cescatti A. Light capture efficiency decreases with increasing tree age and size in the southern hemisphere gymnospenn Agathis australis. Trees, 2005, 19:177-190.
[150] Lusk CH. Leaf area and growth of juvenile temperate evergreens in low light: species of contrasting shade tolerance change rank during ontogeny. Functional ecology, 2004, 18: 820-828.
[151] Kneeshaw DD, Kobe RK, Coates KD et al. Sapling size influences shade tolerance ranking among southern boreal tree species. Journal of ecology, 2006, 94:471-480.
[152] Lusk CH, Piper FI. Seedling size influences relationships of shade tolerance with carbohydrate-storage patterns in a temperate rainforest. Functional ecology, 2007, 21: 78-86.
[153] Westoby M, Jurado E, Leishman M. Comparative evolutionary ecology of seed size. Trends in Ecology and Evolution, 1992, 7: 368-372.
[154] Jones RH, Sharitz RR, Dixon PM et al. Woody plant regeneration in four floodplain forest Ecological Monographs, 1994, 64: 345-367.
[155] Reich PB. Variation among plant species in leaf turnover rates and associated traits, implications for growth at all life stages. Inherent Variation in Plant Growth, Physiological Mechanisms and Ecological Consequences (eds H.Lambers, H. Poorter & M.M.I. Van Vuuren), Backhuys, Leiden, 1998,467-487.
[156] Grubb PJ. A reassessment of the strategies of plants which cope with shortages of resources. Perspectives in Plant Ecology, Evolution and Systematics, 1998, 1:3-31.
[157] Grubb PJ, Metcalfe DJ. Rainforest dynamics: the need for new paradigms. Tropical Rainforest Research- Current Issues (eds Edwards DS, Booth WE & Choy SC),. Kluwer, Dordrecht, The Netherlands, 1996, 215-233.
[158] Wright IJ, Westoby M. Differences in seedling growth behaviour among species: trait correlations among species, and trait shifts along nutrient compared to rainfall gradients. Journal of Ecology, 1999, 87: 85-97.
[159] Reich PB, Buschena C, Tjoelker MG et al. Variation in growth rate and ecophysiology among 34 grassland and savanna species under contrasting N supply: a test of functional group differences. New Phytologist, 2003, 157: 617-631.
[160] Henns, DA, Mattson, WJ. The dilemma of plants, to grow or defend. Quarterly Review of Biology, 1992, 67:283-335.
[161] Cao, KF, Ohkubo T. Aoohnetry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan. Ecological Research, 1998, 13:217-227.
[162] Smith TM, Shugart HH, Woodward FI et al. Plant functional types. In: Vegetation Dynamics and Global Change (Eds A.M. Solomon & H.H. Shugart), Chapman & Hall, New York, 1993, 272-292.
[163] Gitay H, Noble IR. What are functional types and how should we seek them? In: Plant Functional Types-Their Relevance to Ecosystem Properties and Global Change (Eds T.M. Smith, H.H. Shugart & F.I. Woodward). Cambridge University Press, Cambridge, 1997, 3-19.
[164] Smith TM, Woodward FI, Shugart HH (eds.). Plant Function Types. Cambridge: Cambridge University Press, 1996.
[165] Woodward FL, Cramer W. Plant functional types and climatic changes. Journal of vegetation science, 1996, 7: 306-308.
[166] Lavorel, S, Mclntyre S, Landsberg Jet al. Plant functional classifications: from general groups to specific groups based on response to disturbance. Tree, 1997, 12:474-478.
[167] Craine JM, Tilman D, Wedin D et al. Functional traits, productivity and effects on nitrogen cycling of 33 grassland species. Functional Ecology, 2002, 16: 563-574.
[168] Tjoelker MG, Craine JM, Wedin D et al. Linking leaf and root trait Syndromes among 39 grassland and savannah species. New Phytologist, 2005, 167: 493-508.
[169] Poorter H, Navas ML. Plant growth and competition at elevated CO_2 : on winners, losers and functional groups. New Phytologist, 2003, 157:175-198.
[170] Wardle DA, Barker GM, Bonner KI et al. Can comparative approaches based on plant ecophysiological traits predict the nature of biotic interactions and individual plant species effects in ecosystems? Journal Ecology, 1998, 86: 405-420.
[171] Hanley ME, Trofimov S, Taylor G Species-level effects more important than functional group-level responses to elevated CO_2: evidence from simulated turves. Functional Ecology, 2004, 18: 304-313.
[172] Sutherland EK, Hale BJ, Hix DM. Defining species guilds in the Central Hardwood Forest, USA. Plant Ecology, 2000, 147: 1-19.
[173] Whitmore TC. Canopy gaps and the two major groups of forest trees. Ecology, 1989, 70(3): 536-538.
[174] Condit R, Hubbell SP, Foster RB. Assessing the response of plant functional types to climatic change in tropical forest. Journal of Vegetation Science, 1996, 7, 406-416.
[175] Agyeman VK, Swaine MD, Thompson J. Responses of tropical forest seedlings to irradiance and the derivation of a light response index. Journal of ecology, 1999, 87: 815-827.
[176] Davies SJ. Photosynthesis of nine pioneer Macaranga species from borneo in relation to life history. Ecology, 1998, 79(7): 2292-2308.
[177] Bloor JMG, Grubb PJ. Morphological plasticity of shade-tolerant tropical rainforest tree seedlings exposed to light changes. Functional Ecology, 2004, 18: 337-348.
[178] 周小勇，黄忠良，史军辉等．鼎湖山针阔混交林演替过程中群落组成和结构短期动态研究．热带亚热带植物学报，2004，12(4)：323-330．
[179] 刘美珍，蒋高明，于顺利等．浑善达克退化沙地恢复演替18年中植物群落动态变化．生态学报，2004，24(8)：1734-1740．
[180] 黄世能，李意德，王伯荪．海南岛尖峰岭两类热带山地雨林次生群落在15年演替过程中的林木消长．植物生态学报，2000，24(6)：710-717．
[181] 达良俊，杨永川，宋永昌．浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型．植物生态学报，2004，28(3)：376-384．
[182] 张家城，陈力，郭泉水等．演替顶极阶段森林群落典型树种分布的变动趋势研究．植物生态学报，1999，23(3)：256-268．
[183] 林露湘，曹敏，唐勇等．西双版纳刀耕火种弃耕地树种多样性比较研究．植物生态学报，2002，26(2)：216-222．
[184] 张继义，赵哈林，张铜会．赵学勇科尔沁沙地植被恢复系列上群落演替与物种多样性的恢复动态．植物生态学报，2004，28(1)：86-92．
[185] 唐勇，曹敏，张建侯等．西双版纳热带森林土壤种子库与地上植被的关系．应用生态学报，1999，10(3)：279-283．
[186] 周先叶，李鸣光，王伯荪等．广东黑石顶自然保护区森林此生演替不同阶段土壤种子库的研究．植物生态学报，2000，24(2)：222-230．
[187] 张庆费，由文辉，宋永昌．浙江天童森林公园植物群落演替对土壤物理性质的影响．植物资源与环境，1997，6(2)：36-40．
[188] 张全发，郑重，金义兴．植物群落演替与土壤发展之间的关系．武汉植物学研究，1990，8(4)：325-334．
[189] 彭少麟．恢复生态学研究进展及在中国热带亚热带的实践．四川师范学院学报(自然科学版)，2000．21(3)：221-227
[190] 邓瑞文，冯泳梅，陈天杏．三种相思(Acacia)的光合作用与蒸腾作用的研究．生态学报，1989，9(2)：128-131．
[191] 王博轶，冯玉龙．生长环境光强对两种热带雨林树种幼苗光合作用的影响．生态学报，2005，25(1)：23-30．
[192] 郭晓荣，曹坤芳，许再富．热带雨林不同生态习性树种幼苗光合作用和抗氧化酶对生长光环境的反应，应用生态学报，2004，15(3)，377-381．
[193] 孙谷畴，林植芳，林桂珠．不同光强下生长的几种亚热带森林树木的Rubisco羧化速率和碳酸酐酶的活性．武汉植物学研究，2001，19(4)：304-310．
[194] 杨小波．不同演替阶段森林群落典型种苗木叶形态解剖结构比较研究．海南大学学报自然科学版，1997，15(3)：212-217．
[195] 丁圣彦．常绿阔叶林演替系列中主要典型种叶片结构的比较．河南大学学报(自然科学版)，29(4)：76-80．
[196] 史刚荣，汤盈，张铮．淮北相山恢复演替群落典型树种叶片的生态解剖．植物生态学报，2006，30(2)：314-322．
[197] 王希华，张婕，张正祥．浙江天童国家森林公园主要常绿阔叶树种叶子寿命的研究．植物生态学报，2000，24(5)：625-629．
[198] 徐程扬．不同光环境下紫椴幼树树冠结构的可塑性响应．应用生态学报，2001，12(3)：339-343．
[199] 杨小波，文洪波，吴庆书等．海南中部山区次生林典型种枫香发育规律研究．海南大学学报自然科学版，2000，18(1)：54-58．
[200] 林业部林业区划办公室．中国林业区划．北京：中国林业出版社，1987．
[201] Alexander IJ, Faifley RI. Effects of N fertilization on populations of fine roots and mycorrhizae in spruce humus. Plant and Soil, 1983, 71: 19～53.
[202] Berntson GM, Farnsworth EJ, Bazzaz FA. Allocation with and between organs and the dynamics of root length changes in two birch species. Oecologia, 1995, 101: 439-447.
[203] Long SP, Humphries S, Falkowski PG.Photoinhibition of photosynthesis in nature. Annual Review of Plant Physiology and Plant Molecular Biology, 1994, 45: 633-662.
[204] Mort KA, Parkhurst DF. Stomatal responses to humidity in air and helox Plant, Cell and Environment, 1991, 14: 509-515.
[205] Roberts SW, Miller PC. Interception of solar radiation as affected by canopy organization in two Mediterranean shrubs. Oecologia Plantarum, 1977, 12: 273-290.
[206] Mcintyre S, Lavorell S. Livestock grazing in subtropical pastures: steps in the analysis of attribute response and plant functional types. Journal of Ecology, 2001,89: 209-226.
[207] Chapin, FS Ⅲ, Bloom A J, Field CB et al. Plant Responses of Multiple Environmental Factors. BioScience, 1987, 37: 49-57.
[208] Arnold E. Stomata. In: The Institute of Biology's Studies in Biology Ed. By Martin E. S., Donkin M. E. and Stevens R. A., 1983, 155.
[209] Mahall BE, Schlesinger WH. Effects ofirradiance on growth, photosynthesis, and water use efficiency of seedlings of the chaparral shrub Ceanothus megacarpus. Oecologia, 1982, 54:291-299.
[210] Abrams MD, Kubiske ME. Leaf structural characteristics of 31 hardwood and conifer tree species in central Wisconsin: influence of light regime and shade-tolerance rank. Forest Ecology and Management, 1990, 31: 245-253.
[211] Reich PB, Tilman D, Craine J. Do species and functional groups differ in acquisition and use of C, N and water under varying atmospheric CO_2 and N availability regimes? A field test with 16 grassland species. New Phytologist, 2001; 150: 435-448.
[212] Nowak RS, Ellsworth DS, Smith SD. Functional responses of plants to elevated atmospheric CO_2-do photosynthetic and productivity data from FACE experiments support early predictions? New Phytologist, 2004, 162: 253-280.
[213] Lee TD, Tjoelker MG, Ellsworth DS et al. Leaf gas exchange responses of 13 prairie grasslandspecies to elevated CO2and increased nitrogen supply. New Phytologist, 2001,405-418.
[214] Ellsworth DS, Reich PB, Naumburg ES et al. Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO_2 across four free-air CO_2 enrichment experiments in forest, grassland and desert. Global Change Biology, 2004, 10:2121-2138.
[215] Holscher D, Leuschner C, Bohman K et al. Photosynthetic characteristics in relation to leaf traits in eight co-existing pioneer tree species in Central Sulawesi, Indonesia. Journal of Tropical Ecology, 2004, 20：157-164．
[216] 符国瑷，冯绍信．海南五指山森林的垂直分布及其特征．广西植物，1995，15(1)：57-69．
[217] 骆土寿，李意德，陈德祥等．海南岛鸡毛松人工林群落种间联接性研究．生态学杂志，2005，24(6)：591-594．
[218] 广东省林业科学研究所．海南主要经济树种．北京：中国农业出版社，1964．
[219] 郑万钧．中国树木志Ⅲ．北京：中国林业出版社，1997．
[220] Escalona JM, Flexas J, Medrano H. Stomatal and non-stolnatal limitations of photosynthesis under water stress in field-growen grapevines. Australian journal of plant physiology, 1999, 16: 421-433.
[221] Kuppers, M. Ecological significance of above-ground architectural patterns in woody plants: a question of cost-benefit relationships. Trends in Ecology and Evolution, 1989, 4: 375-379.
[222] Gottschalk KW. Shade, leaf growth and crown development of Quercus rubra, Quercus velutina, Prunus serotina and Acer rubrum seedlings. Tree physiology, 1994, 14: 735-749.
[223] Sterck FJ, Bongers F. Crown development in tropical rain forest trees: patterns with tree height and light availability. Journal of ecology, 2001, 89: 1-13.
[224] Farque L, Sinoquet H, Colin F. Canopy structure and light interception in Quercus petreae seedlings in relation to light regime and plant density. Tree physiology, 2001, 21: 1257-1267.
[225] Beets, PN; Pollock DS. Accumulation and partitioning of dry matter in Pinus radiate as related to stand age and thinning. New Zealand Journal of Forest Science, 1987, 17: 246-271.
[226] King, DA. Tree form, height growth, and susceptibility, to wind damage in Acer saccharum. Ecology, 1986, 67: 980-990.
[227] Givnish, TJ. Plant stems: biochemical adaptation for energy capture and influence on species distribution. In: Plant Stems: Physiology and Functional Morphology (Eds.). BL Gartner, Academic Press, San Diego, 1995, 3-49.
[228] Ceschia, E. Environmental effects on spatial and seasonal variations of stem respiration in European beech and Norway spruce. Ph.D. Thesis, Swedish University of Agricultural Science, Uppsala, Sweden, 2001, 55.
[229] McDowell N, Barnard H, Bond BJ et al. The relationship between tree height and leaf area: sapwood area ratio. Oecologia, 2002, 132: 12-20.
[230] Wright SJ. Seasonal drought soil fertility and the species density of tropical forest plant communities. Tree, 1992, 7, 260-263.
[231] Letts MG, Mulligan M. The impact of light quality and leaf wetness on photosynthesis in north-west Andean tropical montane cloud forest. Journal of Tropical Ecology, 2005, 21: 549-557.
[232] Graham EA, Mulkey SS, Kitajima K et al. Cloud cover limits net CO_2 uptake and growth of a rainforest tree during tropical rainy seasons. PNAS, 2003, 100 (2): 572-576.
[233] Hogan KP, Smith AP, Samaniego M. Gas exchange in six tropical semi-deciduous forest canopy tree species during the wet and dry seasons. Biotropica, 1995, 27, 324-333.
[234] Quero JL, Villar R, Maranon T et al. Interactions of drought and shade effects on seedlings of four Quercus species: physiological and structural leaf responses. New Phytologist, 2006, 170: 819-834.
[235] Holmgren M. Combined effects of shade and drought on tulip poplar seedlings: trade-off in tolerant or facilitation? Oikos, 2000, 90: 67-78.
[236] Duan B, Lu Y, Yin C et al. Physiological responses to drought and shade in two contrasting Picea asperata populations. Physiologia Plantarum, 2005, 124: 476-484.
[237] Canham CD, Berkowitz AR, Kelly VR et al. Biomass allocation and multiple resource limitation in tree seedlings Canadian. Journal of Forest Research, 1996, 26:1521-1530.
[238] Smith T, Huston M. A theory of the spatial and temporal dynamics of plant communities. Vegetatio, 1989, 83: 49-69.
[239] Bazzaz FA, Picket STA. Physiological ecology of a tropical succession: a comparative review. Annual Reviews in Ecological Systems, 1980, 11: 287-310.
[240] Doley D, Unwin GL, Yates DJ.Spatial and temporal distribution of photosynthesis and transpiration by single leaves in a rainforest tree, Argyrodendron peralatum. Australian Journal of Plant Physiology, 1988, 15: 317-326.
[241] Ishibashi M, Terashima I. Effects of continuous leaf wetness on photosynthesis: adverse aspects of rainfall. Plant, Cell and Environment, 1995, 18: 431-438.
[242] Smith WK, McClean, TM. Adaptive relationship between leaf water repellency, stomatal distribution and gas exchange American. Jouranl of Botany, 1989, 76: 465-469.
[243] Nobel PS. Environmental productivity indexes and productivity for Opuntia ficus-indica under current and elevated atmospheric CO_2 levels. Plant, Cell and Environment, 1991, 14: 637-646.
[244] Allen DJ, Ort DR. Impacts of chilling temperatures on photosynthesis in warm- climate plants. Trends in Plant Science, 2001, 6: 36-42.
[245] Ribeiro RV, Lyra GB, Santiago AV et al. Diurnal and seasonal patterns of leaf gas exchange in bahiagrass (Paspalum notatum Flfigge) growing in a subtropical climate Grass and Forage. Science, 2006, 61: 293-303.
[246] Hastwell GT, Facelli JM. Differing effects of shade-induced facilitation on growth and survival during the establishment of a chenopod shrub. Journal of Ecology, 2003, 91: 941-950.
[247] Pacala SW, Canham CK, Saponara J et al. Forest models defined by field measurements, estimation, error analysis and dynamics. Ecological Monographs, 1996, 66: 1-43.
[248] Wyckoff PH, Clark JS. The relationship between growth and mortality for seven co-occurring tree species in the southern Appalachian Mountains. Journal of Ecology, 2002, 90:604-615.
[249] Pompa J, Bongers F. The effect of canopy gaps on growth and morphology of seedlings of rain forest species. Oecologia, 1988, 75: 625-632.
[250] Koike T. Leaf structure and photosynthetic performance as related to the forest succession of deciduous broad-leaved trees. Plant Species Biology, 1988, 3: 77-87.
[251] Lei TT, Lechowicz MJ. Shade adaptation and shade tolerance in saplings of three Acer species from eastern North America. Oecologia, 1990, 84: 224-228.
[252] Reich PB, Uhl C, Waiters MB et al. Leaf lifespan as a determinant of leaf strcture and function among 23 Amazonian tree species. Oecologia, 1991, 86: 16-24.
[253] Gleason SM, Ares A. Photosynthesis, carbohydrate storage and survival of a native and an introduced tree species in relation to light and defoliation. Tree Physiology, 2004, 24: 1087-1097.
[254] 张一平，窦军霞，于贵瑞等．西双版纳热带雨林太阳辐射特征研究．北京林业大学学报，2005，27(5)：17-25．
[255] Denslow JS. Tropical rainforest gaps and tree species diversity Annual Review of Ecology and Systematics, 1987, 18: 431-451.
[256] Scholes JD, Press MC, Zipperlen SW. Differences in light energy utilization and dissipation between dipterocarp rain forest tree seedlings. Oecologia, 1997, 109: 41-48.
[257] Silva EA, DaMatta FM, Ducatti C et al. Seasonal changes in vegetative growth and photosynthesis of Arabica coffee trees. Field Crops Research, 2004, 89: 349-357.
[258] Niinemets U, Kull O, Tenhunen JD. An analysis of light effects on foliar morphology, physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance. Tree Physiology, 1998, 18, 681-696.
[259] Stanhill G, Cohen S. Global dimming: a review of the evidence for a widespread and significant reduction in global radiation with discussion of its probable causes and possible agricultural consequences. Agricultural and Forest Meteorology, 2001, 107(4): 255-278.
[260] Liepert BG.Observed reductions of surface solar radiation at sites in the United States and worldwide from 1961 to 1990. Geophysical Research Letters, 2002, 29: 1010, 61-1, 5.
[261] 查良松．我国太阳辐射量区域性变化特征研究．地理研究,1996,15(2)：21-27．
[262] McGuffie K, Henderson- Sellers A. Is Canadian cloudiness increasing? Atlnosphere- Ocean, 1988, 26: 608-623.
[263] Henderson-Sellers A North American total cloud amount variations this century. Palaeogeograph. Palaeoclimatol. Palaeoecol., 1989, 75:175-194.
[264] Karl TR, Steurer PM. Increased cloudiness in the United States during the first half of the twentieth century: Fact or fiction? Geophysical Research Letters, 1990, 17: 1925-1928.
[265] Chen J, Carlson E, Del Genio AD. Evidence for strengthening of the Tropical General Circulation in the 1990s. Science, 2002, 295:838-841.
[266] Wielicki BA, Wong T, Allan RP et al. Evidence for large decadal variability in the tropical mean radiative energy budget. Science, 2002, 295:841-844.
[267] Price DT, Black TA. Effects of short-term variation in weather on diurnal canopy CO_2 flux and evaporation of a juvenile Douglas Fir stand Agricultural and Forest Meteorology, 1990, 50:139-158.
[268] Hollinger DY, Kelliher FM, Byers JN et al. Carbon dioxide exchange between an undisturbed old-growth temperate forest and the atmosphere. Ecology, 1994: 75, 134-150.
[269] Gu L, Fuentes JD, Shugart HH et al. Responses of net ecosystem exchange of carbon dioxide to changes in cloudiness: Results from two North American deciduous forests. Journal of Geophysical Research, 1999, 104: 31421-31434.
[270] Knapp AK, Smith WK, Young DR. Importance of intermittent shade to the ecophysiology of subalpine herbs. Functional Ecology, 1989, 3: 753-758.
[271] Baldocchi DD. Measuring and modeling carbon dioxide and water vapor exchange over a temperate broad-leaved forest during the 1995 summer drought. Plant, Cell and Environment, 1997, 20: 1108-1122.
[272] Ricklefs RE. Environmental heterogeneity and plant species diversity: a hypothesis. American Naturalist, 1977, 111: 376-381.
[273] Grantz DA, Moore PH, Zeiger E. Stomatal responses to light and humidity in sugarcane: prediction of daily time courses and identification of potential selection criteria Plant, Cell and Environment, 1987, 10: 197-204.
[274] Johnson JD, Ferrell WK. Stomatal response to vapour pressure deficit and the effect of plant water stress Plant, Cell and Environment, 1983, 6: 451-456.
[275] Guangdong Institute of Botany. Flora Hainanica, Volume Ⅳ. Science Press, Beijing, China, 1977. (in Chinese).
[276] Chun WY. Flora Hainanica, Volume Ⅱ. Science Press, Beijing, China, 1965. (in Chinese)
[277] 粟娟，王德祯，符史深．海南岛尖峰岭热带树木园主要树种的物候研究．林业科学研究，1994，7(3)：294-300．
[278] Rocha AV, Su HB, Vogel CS et al. Photosynthetic and water use efficiency responses to diffuse radiation by an aspen-dominated northern hardwood forest. Forest Science, 2004, 50(6): 793-801.
[2] 胡玉佳，李玉杏．海南岛热带雨林．广州：广东高等教育出版社，1992．
[3] Larcher W. Physiological plant ecology-ecophysiology and stress physiology of functional groups. New York: Springer-Verlag, 1995.
[4] Wilson EO. The diversity of life. Cambridge: The Belknap Press of Harvard University Press, 1992, 183-211.
[5] Robert C, Ralph D, Rudolf DG et al. The value of the world's ecosystem services and natural capital. Nature, 1997, 387: 253-260.
[6] Holdgate M. Conservation in a changing world. IUCN Bulletin, 1989, (10-12): 16-25.
[7] Williams FL. Reflection on the tropical deforestation crisis Biological Conservation. 1999, 91: 109-117.
[8] 颜文洪．全球性热带雨林的破碎化趋势与保护分析．自然生态保护，2003，2：34-36．
[9] 王献溥，崔国发，李俊清．海南生物多样性研究展望．北京林业大学学报，2000(4)：132-134．
[10] 李炎香．海南岛热带森林的开发利用及其在生态平衡中的作用．热带林业科技，1980，(3)：35-44．
[11] 吴华盛．海南省热带雨林的保护和利用．热带林业，2002，30(2)：4-5．
[12] 曹建华，蒋菊生．海南生物多样性保护探析．华南热带农业大学学报，2006，12(2)：34-38．
[13] 臧润国，安树青，陶建平等．海南岛热带林生物多样性维持机制．北京：科学出版社，2004，198-210．
[14] 陈德祥，李意德，骆土寿等．海南岛尖峰岭热带山地雨林下层乔木中华厚壳桂光合生理生态特性的研究．林业科学研究，2003，16(5)：540-47．
[15] 陈德祥，李意德，骆土寿等．短期CO_2浓度升高对雨林树种盘壳栎光合特性的影响．生态学报，2004，24(8)：1622-1628．
[16] 骆土寿，李意德，陈德祥等．热带雨林恢复演替中典型树种黄桐气体交换对环境的响应．生态学报，2003，23(9)：1762-1772．
[17] Wardle DA, Bardgett RD, Klironomos JN et al. Ecological linkages between aboveground and belowground biota Science, 2004, 304: 1639-1633.
[18] Odum EP. The strategy of ecosystem development. Science, 1969, 164: 262-270.
[19] Gurevitch J, Scheiner SM, Fox GA. The ecology of plants. Sinauer Associates Inc., Sunderland, Massachusetts U.S.A., 2002, 253-276.
[20] Bazzaz FA. The physiological ecology of plant succession. Annual Review of Ecology and Systematics, 1979, 10: 351-371.
[21] Kobe RK, Pacala W, Silander JA. Juvenile tree survivorship as a component of shade tolerance. Ecological Applications, 1995, 5: 517-532.
[22] Koike F. Plant traits as predictors of woody species dominance in climax forest in northern Japan. Journal of Ecology, 2001, 12: 327-336.
[23] Walters MB, Reich PB. Are shade tolerance, survival, and growth linked? Low light and nitrogen effects on hardwood seedlings. Ecology, 1996, 77(3): 841-853.
[24] Poorter L, Arets EJMM. Light environment and tree strategies in a Bolivian tropical moist forest: an evaluation of the light partitioning hypothesis. Plant ecology, 2003, 166: 295-306.
[25] Agyeman VK, Swaine MD, Thompson J. Responses of tropical forest seedlings to irradiance and the derivation of a light response index. Journal of ecology, 1999, 87: 815-827.
[26] Kyereh B, Swaine MD, Thompson J. Effect of light on the germination of forest trees in Ghana. Journal of ecology, 1999, 87: 772-783.
[27] Denslow JS, Schultz JC, Vitousek PM et al. Growth responses of tropical shrubs to treefall gap environments. Ecology, 1990, 71(1): 165-179.
[28] Raynal DJ, Bazzaz FA. Establishment of early successional plant populations on forest and prairie soil. Ecology, 1973, 54: 1335-1341.
[29] 温远光，梁宏温，和太平等．大明山退化生态系统群落光照的变化．广西农业大学学报，1998，17(2)：199-203．
[30] Chazdon RL, Fetcher N. Photosynthetic light environments in a lowland tropical rain forest in Costa Rica. Journal of Ecology, 1984, 72: 553-564.
[31] Endler JA. The color of light in forests andits implications. Ecological.Monograph, 1993, 63: 1-27.
[32] Lee DW, Baskaran K, Mansor M et al. Irradiance and spectral quality affect Asian tropical rain forest tree seedling development. Ecology, 1996, 77: 568-580.
[33] Pearcy RW. Sunflecks and photosynthesis in plant canopies. Annual Review of Plant Physiology and Plant Molecular Biology, 1990, 41: 421-4531
[34] Watling JR, Robinson SA, Woodrow IE, Osmond CB. Responses of rainforest understoey plants to excess light during sunflecks. Australian Journal of Plant Physiology, 1997, 24, 17-25.
[35] Leakey ADB, Press MC, Scholes JD. High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling. Plant, Cell and Environment, 2003, 26: 1681-1690.
[36] Watt AS. Pattern and process in the plant community. Journal of ecology, 1947, 35: 1-22.
[37] Sanford PJ, Braker HE, Hartshom GS. Canopy openings in a primary neotropical lowland forest. Journal of Tropical Ecology, 1986, 2, 277-282.
[38] 蒋有绪，王伯荪，臧润国等．海南岛热带林生物多样性及其形成机制．北京：科学出版社，2002，1-30．
[39] 蒋有绪，卢俊培．中国海南岛尖峰岭热带林生态系统．北京：科学出版社，1991，256-283．
[40] 张一平，王进欣，马友鑫等．西双版纳雾凉季次生林林窗光照特征初步分析．热带气象学报，2000，16(4)：374-379．
[41] 张一平，于进欣，马友鑫等．西双版纳热带次生林林窗近地层温度时空分布特征．林业科学，2002，38(6)：1-5．
[42] 咎启杰，李鸣光，张志权等．林窗及其在森林动态中的作用．植物学通报，1997，14(增刊)：18-24．
[43] Brokaw VLB. Species composition in gaps and structure of a tropical forest. Ecology, 1989, 70: 538-541.
[44] Swaine MD, Whitmore TC. On the definition of ecological species groups in tropical rain forest. Vegetatio, 1988, 75: 8t-86.
[45] Hawthorne, WD. Ecological profiles of Ghanaian forest trees. Tropical Forestry Papers 29. Oxford Forestry Institute, Oxford, 1995.
[46] Zimmerman JK, Pascarella JB, Aide TM. Barriers to forest regeneration in an abandoned pasture in Puerto Rico. Restoration ecology, 2000, 8(4): 350-360.
[47] Hooper H, Legendre P, Condit R. Barriers to forest regeneration of deforested and abandoned land in Panama. Journal of applied ecology, 2005, 42: 1165-1174
[48] Van der Marrel E. Vegetation ecology. Blackwell Publishing, Oxford, 2005, 172-198.
[49] Kuppers M. Canopy gaps: competitive light interception and economic space filling-a matter of whole plant allocation. In: Caldwell, MM and Pearcy RW, eds. Exploitation of Environmental Heterogeneity by Plants. San Diego: Academic Press, 1994, 111-114.
[50] Pugnaire FI, Valladares F. Handbook of Functional Plant Ecology. Marcel Eekker, Inc. 1999, 121-194.
[51] Edwards EJ, Benham DG, Marland LA et al. Root production is determined by radiation flux in a temperate grassland community. Global change biology, 2004, 10: 209-227.
[52] Pons TL. An ecophysiological study in the field layer of ash coppice, Ⅱ. Experiments with Geum urbanum and Cirsium palustre in different light intensities. Acta Boanicat Neerlandica, 1977, 26: 29-42.
[53] Oconnell BM, Kelty MJ. Crown architecture of understory and open-grown white pine (Pinus strobes L.) saplings. Tree Physiology, 1994, 14: 89-102.
[54] Poorter L. Growth responses of 15 rainforest tree species to a light gradient: the relative importance of morphological and physiological traits. Functional Ecological, 1999, 13:396-410.
[55] Reich PB, Tjoelker MG, Walters MB et al. Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedlings of nine boreal tree species grown in high and low light. Functional Ecology, 1998, 12:327-338.
[56] Huante, P. and E. Rincon. Responses to light changes in tropical deciduous woody seedlings with contrasting growth rates. Oecologia, 1998, 113:53-66.
[57] Givnish, TJ. Adaptation to sun and shade: a whole-plant perspective. Australlian Journal of Plant Physiology, 1988, 15:63-92.
[58] Mori A, Takeda H. Functional relationships between crown morphology and with-crown characteristics of understory saplings of three codominant conifers in a subalpine forest in central Japan. Tree Physiology, 2004, 24: 661-670.
[59] Kobe RK. Carbohydrate allocation to storagte as a basis of interspecific variation in sapling survivorship and growth. Oikos, 1997, 80: 226-233.
[60] Waiters MB, Reich PB. Low-light carbon balance and shade tolerance in the seedling of woody plants: do winter deciduous and broad-leaved evergreen species differ? New Phytologist, 1999, 143: 143-154.
[61] Kitajima K. Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees. Oecologia, 1994, 98: 419-428.
[62] Montgomery R. Relative importance of photosynthetic physiology and biomass allocation for tree seedling growth across a broad light gradient. Tree Physiology, 2004, 24: 155-167.
[63] Delagrange S, Messier C, Lechowicz MJ et al. Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability. Tree Physiology, 2004, 24: 775-784.
[64] Beaudet M, Messier C. Growth and morphological response of yellow birch, sugar maple, and beech seedlings growing under a natural light gradient. Canadian Joumal of Forest Research, 1998, 28: 1007-1015.
[65] Gibson AC. Photosynthetic organs of desert plants. Bio Science, 1998, 48:911-920.
[66] Bloor JMG, Grubb P. Growth and mortality in high and low light: trends among 15 shade-tolerant tropical rain forest tree species. Journal of Ecology, 2003, 91: 77-85.
[67] Vilela AE, Rennella M J, Ravetta DA. Responses of tree-type and shrub-type Prosopis (Mimosaceae) taxa to water and nitrogen availabilities. Forest Ecology and Management, 2003, 186: 327-337.
[68] Augspurger CK. Light requirements of neotropical tree seedlings: a comparative study of growth and survival. Journal of Ecology, 1984, 72: 777-795.
[69] Lusk CH. Leaf area accumulation helps juvenile evergreen trees tolerance shade in a temperate rainforest. Oecologia, 2002, 132:188-196.
[70] Cornelissen JHC. Aboveground morphology of shade-tolerant Castanopsis fargesii samplings in response to light environment. International Journal of Plant Sciences, 1993, 154 (4): 481-495.
[71] Valladares F, Pearcy RW. The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Caliafornian chaparral shrub. Oecologia, 1998, 114: 1-10.
[72] Pearcy RW, Muraoka H, Valladares F. Crown architecture in sun and shade environments: assessing function and trade-offs with a threedimensionai simulation model. New Phytologist, 2005, 166: 791-800.
[73] Werk KS, Ehleringer J. Non-random leaf orientation in Lactuca serriola L. Plant, Cell and Environment, 1984, 7: 81-87.
[74] Smith M, Ullberg D. Effect of leaf angle and orientation on photosynthesis and water relations in Silphium terebinthinaceum. American Journal of Botany, 1989, 76: 1714-1719.
[75] Valladares F, Pearcy RW. Interactions between water stress, sun-shade acclimation, heat tolerance and photoinhibition in the sclerophyll Heteromeles arbutifolia. Plant, Cell and Environment, 1997, 20: 25-36.
[76] Runkle JR. Pattern of disturbance in some old-growth mesic forests of eastern North America. Ecology, 1982, 63: 1533-1546.
[77] Sipe TW, Bazzaz FA. Gap partitioning among maples (Acer) in central New-England - shoot architecture and photosynthesis. Ecology, 1994, 5:2318-2332.
[78] Ackerly DD, Bazzaz FA. Seedling crown orientation and interception of diffuse radiation in tropical forest gaps. Ecology, 1995, 76:1134-1146.
[79] Delagrange S, Montpied P, Dreyer E et al. Does shade improve light interception efficiency? A comparison among seedlings from shade-tolerant and -intolerant temperate deciduous tree species. New Phytologist, 2006, 172: 293-304.
[80] King DA. Allometry of saplings and understorey trees of a Panamanian forest. Functional Ecology, 1990, 4: 27-32.
[81] Kohyama T, Hotta M. Significance of allometry in tropical saplings. Functional Ecology, 1990, 4: 515-521.
[82] Valladares F, Skillman JB, Pearcy RW. Convergence in light capture efficiencies among tropical forest understory plants with contrasting crown architectures: a case of morphological compensation. American Journal of Botany, 2002, 89: 1275-1284.
[83] Pearcy RW, Valladares F, Wright SJ et al. A functional analysis of the crown architecture of tropical forest Psychotria species: do species vary in light capture efficiency and consequently in carbon gain and growth? Oecologia, 2004, 139:163-177.
[84] Horton, P., A. V. Ruban, & R. G. Walters. Regulation of light harvesting in green plants. Annual Review of Plant Physiology and Plant Molecular Biology, 1996, 47: 655-684.
[85] Osmond CB, Grace SC. Perspectives on photoinhibition and photorespiration in the field: quintessential inefficiencies of the light and dark reactions of photosynthesis? Journal of Experimental Botany, 1995, 46: 1351-1362.
[86] Demmig-Adams B, Adams Ⅲ WW. Photoprotection and other responses of Plants to high light stress. Annual Review of Plant Physiology and Plant Molecular Biology, 1992, 43: 599-626.
[87] Demmig-Adams B, Adams WW. The role of xanthophyll cycle carotenoids in the protection of photosynthesis. Trends in Plant Science, 1996, 1: 21-26.
[88] Niyogi KK. Photoprotection revisited: Genetic and molecular approaches. Annual Review of Plant Physiology and Plant Molecular Biology, 1999, 50: 333-359.
[89] Schulze E-D, Lange OL, Evenari M et al. Long-term effects of drought on wild and cultivated plants in the Negev desert. Ⅱ. Diurnal patterns of net photosynthesis and daily carbon gain. Oecologia, 1980, 45:19-25.
[90] Tenhunen JD, Lange OL, Braun M. Midday stomatal closure in Mediterranean type sclerophylls under simulated habitat conditions in an environmental chamber. Ⅱ. Effect of the complex of leaf temperature and air humidity on gas exchange of Arbutus unedo and Qurecus ilex. Oecologia, 1981, 50:5-11.
[91] Tenhunen JD, Lange OL, Jahner D. The control by atmospheric factors and water stress of midday stomatal closure in Arbutus unedo growing in a natural macchia Oecologia, 1982, 55:165-169.
[92] Tenhunen JD, Lange OL, Geble J et al. Changes in photosynthetic capacity, carboxylation efficiency, and CO_2 compensation point associated with miday stomatal closure and midday depression of net CO_2 exchange of leaves of Quercus suber. Planta, 1984, 162:193-203.
[93] Chaves, MM, Harley PC, Tenhunen JD et al. Gas exchange studies in two Portuguese grapevine cultivars. Physilolgia Plantarum, 1987, 70: 639-647.
[94] Downton, WJS, Grant WJR, Loveys BR. Diurnal changes in the photosynthesis of field-grown grape vines. New Phytologist, 1987, 105: 71-80.
[95] Correia MJ, Chaves MM, Pereira JS. Afternoon depression in photosynthesis in grapevine leavesevidence for a high light stress effect. Journal of Experimental Botany, 1990, 41: 417-426.
[96] Epron D, Dreyer E, Breda N. Photosynthesis of oak trees (Quercus petraea (Matt.) Liebl.) during drought under field conditions: diuranal course of net CO_2 assimilation and photochemical efficiency of photosystem Ⅱ. Plant, Cell and Environment, 1992, 15: 809-820.
[97] Cowan, LR. Regulation of water use in relation to carbon gain in higher plants. In Encyclopedia of Plant Physilolgy NS, Physiological Plant Ecology Ⅱ (eds O. L. L ange, P. S. Nobel, C. B. Osmond & H. Ziegler). Springer verlag, Berlin, 1982.
[98] Tyree MT, Sperry JS. Do woody plants operate near the point of catastrophic xylem dysfunction caused by dynamic water stress? Answers from a model. Plant Physiology, 1988, 88: 574-580.
[99] Jones HG, R.A. Sutherland. Stomatal control of xylem embolism. Plant, Cell and Environment, 1991, 14: 607-612.
[100] Sperry JS. Hydraulic constraints on plant gas exchange. Agricultural and Forest Meteorology, 2000, 104: 13-23.
[101] Muraoka H, Tang Y, Terashima I et al. Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light. Plant, Cell and Environment, 2000, 23: 235-250.
[102] Medrano H, Bota J, Abadia A et al. Effects of drought on light-energy dissipation mechanisms in high-light-acclimated, field-grown grapevines. Functional Plant Biology, 2002, 29:1197-1207.
[103] Ducrey M. Influence of shade on photosynthetic gas exchange of 7 tropical rainforest species from Guadeloupe (French-West-Indies). Ann. Sci. For., 1994, 51: 77-94.
[104] Oberbauer SF, Strain BR. Effect of canopy position and irradiance on leaf physiology and morphology of Penthaclethra macroloba (Mimosaceae). American Journal of Botany, 1985, 73: 409-416.
[105] Chiariello NR, Field CB, Mooney HA. Midday wilting in a tropical pioneer tree. Functional Ecology, 1987, 1: 3-11.
[106] Poorter L, Oberbauer SF. Photosynthetic induction responses of two rainforest tree species in relation to light environment. Oecologia, 1993, 96: 193-199.
[107] Walters MB, Field CB. Photosynthetic light acclimation in two rainforest Piper species with different ecological amplitudes. Oecologia, 1987, 72: 449-456.
[108] Sims DA, Pearcy RW. Photosynthesis and respiration in Alocasia macrorrhiza following transfers to high and low light. Oecologia, 1991, 86: 447-453.
[109] Strauss-Dibenedetti S Bazaz FA. Plasticity and acclimataion to light in tropical Moraceae of different successional positions. Oecologia, 1991, 87: 377-387.
[110] Kamaluddin M, Grace J. Acclimation in seedlings of a tropical tree, Bischofia javanica, following a stepwise reduction in light. Annals of Botany, 1992, 69: 557-562.
[111] 冯玉龙，草坤芳，冯志立等．四种热带雨林树种幼苗比叶重，光合特性和暗呼吸对生长光环境的适应．生态学报，2002，22(6)：901-910．
[112] 齐欣，曹坤芳，冯玉龙．热带雨林蒲桃属3个树种的幼苗光合作用对生长光强的适应．植物生态学报，2004，28(1)：31-38．
[113] 张教林，曹坤芳．不同生态习性热带雨林树种的幼苗对光能的利用与耗散．应用生态学报，2004，15(3)：372-376．
[114] Chow WS, AdamsonHY, Anderson JM. Photosynthetic acclimation of Tradescantia albiflora to growth irradiance: Lack of adjustment of light-harvesting components and tis consequences. Physiologia Plantarum, 1991, 81: 175-182.
[115] 孙谷畴，赵平，曾小平．两种木兰科植物叶片光合作用的光驯化．生态学报，2004，24(6)：1111-1117．
[116] 梁春，林植芳，孔国辉．不同光强下生长的亚热带树苗的光合—光响应特性的比较．应用生态学报，1997，8(1)：7-11．
[117] Kursar TA, Coley PD. Photosynthetic induction times in shade-tolerant species with long and short-leaved leaves. Oecologia, 1997, 93: 165-170.
[118] Lusk CH, Pozo AD. Survival and growth of seedlings of 12 Chilean rainforest trees in two light environments: Gas exchange and biomass distribution correlates. Austral Ecology, 2002, 27:173-182.
[119] Lusk CH, Reich PB. Relationships of leaf dark respiration with light environment and tissue nitrogen content in juveniles of 11 cold-temperate tree species. Oecologia, 2000, 123:318-329.
[120] Grime JP. Shade tolerance in flowering plants. Nature, 1965, 208: 161-163.
[121] Loach K. Shade tolerance in flowering plants. 1. Leaf photosynthesis and respiration in plants raised under artificial shade. New Phytologist, 1967, 66:607-21.
[122] Huc R, Ferhi A, Guehl JM. Pioneer and late stage tropical rainforest tree species (French Guyana) growing under common conditions differ in leaf gas exchange regulation, carbon isotope discrimination and leaf water potential Oecologia, 1994, 99: 297-305.
[123] Bonal D, Barigah TS, Granier A, Guehl JM. Late-stage canopy tree species with extremely low δ~(13)C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana Plant, Cell and Environment, 2000, 23: 445-459.
[124] Ishida A, Nakano T, Matsumoto Y et al. Diurnal changes in leaf gas exchange and chlorophyll fluorescence in tropical tree species with contrasting light requirements. Ecological Research, 1999, 14: 77-88.
[125] Hanba YT, Kogami H, Terashima I. The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand. Plant, Cell and Environment, 2002, 25: 1021-1030.
[126] Pedersen BS. The role of stress in the mortality of Midwestern oaks as indicated by growth prior to death. Ecology, 1998, 79: 79-93.
[127] Sack L, Grubb PJ. Why do species of woody seedlings change rank in relative growth rate between low and high irradiance? Functional Ecological, 2001, 15:145-154.
[128] Pompa J, Bongers F. The effect of canopy gaps on growth and morphology of seedlings of rainforest species. Oecologia, 1988, 75: 625-632.
[129] Rice SA, Bazzaz FA. Growth consequences of plasticity of plant traits in response to light conditions. Oecologia, 1989, 78:508-512.
[130] Pompa J, Bongers F. Acclimation of seedlings of three Mexican tropical rain forest tree species to a change in light availability. Journal of Tropical Ecology, 1991, 7:85-97.
[131] Ackerly, DD. Phenotypic plasticity and the scale of environmental heterogeneity: studies of tropical pioneer trees in variable light environments PhD thesis, Harvard University, Massachusetts, 1993.
[132] Newell EA, McDonald EP, Strain, BR et al. Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest. Oecologia, 1993, 94:49-56.
[133] Tumbull MH, Doley D, Yates DJ. The dynamic of photosynthetic acclimation to changes in light quantity and quality in three Ausftralian rainforest tree species Oecologia, 1993, 94:218-228.
[134] Niinemets U. The controversy over traits conferring shade-tolerance in trees: ontogenetic changes revisited. Journal of Ecology, 2006, 94: 464-470.
[135] Veneklaas EJ, Poorter L. Growth and carbon partitioning of tropical tree seedlings in contrasting light environments. In: Inherent Variation in Plant Growth: Physiological Mechanisms and Ecological Consequences (eds H. Lambers, H. Poorter & M. Van Vuuren). Backhuys Publishers, Leiden, Netherlands, 1998: 337-62.
[136] Reich PB, Walters MB, Ellsworth DS. Leaf lifespan in relation to leaf, plant and stand characteristics among diverse ecosystems. Ecological Monographs, 1992, 62: 365-392.
[137] King DA. Influence of light level on the growth and morphology of saplings in a Panamanian forest American. Journal of Botany, 1994, 81: 948-957.
[138] Leishman MR, Westoby M. The role of large seed size in shaded conditions, experimental evidence. Functional Ecology, 1994, 8:205-214.
[139] Boot, RGA. The significance of seedling size and grown rate of tropical rainforest tree seedlings for regeneration in canopy openings. Ecology of Tropical Forest Tree Seedlings (ed. M.D. Swaine), 1996, 267-284.
[140] Saverimuttu T, Westoby M. Seedling longevity under deep shade~ in relation to seed size. Jourual of Ecology, 1996, 84: 681:689.
[141] Walters MB, Reich PB. Seed size, nitrogen supply and growth rate affect tree seedling survival in deep shade. Ecology, 2000, 81: 1887-1901.
[142] Pintado A, Valladares F, Sancho LG. Exploring phenotypic plasticity in the lichen Ramalina capitata: morphology, water relations and chlorophyll content in North and South-facing populations. Annals of Botany, 1997, 80: 345-353.
[143] Valladares F, Wright SJ, Lasso E et al. Plastic phenotypic response to light of 16 congeneric shrubs from a Panamanian rainforest. Ecology, 2000, 81 (7): 1925-1936.
[144] Bjorkman O. Responses to different quantum flux densities. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Physiological plant ecology 1. Responses to the physical environment. Springer, Berlin, Heidelberg, New York, 1981, 57-107.
[145] Tumbull MH. The effect of light quantity and quality during development on the photosynthetic characteristics of six Australian rainforest tree species. Oecologia, 1991, 87:110-117.
[146] Chazdon, RL. Photosynthetic plasticity of two rain forest shrubs across natural gap transects. Oecologia, 1992, 92:586-595.
[147] Canham, CD. Different responses to gaps among shade-tolerant species. Ecology, 1989, 70:548-550.
[148] Poorter L, Bongers F, Sterck FJ et al. Beyond the regeneration phase: differentiation of height-light trajectories among tropical tree species. Journal of Ecology, 2005, 93: 256-267.
[149] Niinemets U, Sparrow A, Cescatti A. Light capture efficiency decreases with increasing tree age and size in the southern hemisphere gymnospenn Agathis australis. Trees, 2005, 19:177-190.
[150] Lusk CH. Leaf area and growth of juvenile temperate evergreens in low light: species of contrasting shade tolerance change rank during ontogeny. Functional ecology, 2004, 18: 820-828.
[151] Kneeshaw DD, Kobe RK, Coates KD et al. Sapling size influences shade tolerance ranking among southern boreal tree species. Journal of ecology, 2006, 94:471-480.
[152] Lusk CH, Piper FI. Seedling size influences relationships of shade tolerance with carbohydrate-storage patterns in a temperate rainforest. Functional ecology, 2007, 21: 78-86.
[153] Westoby M, Jurado E, Leishman M. Comparative evolutionary ecology of seed size. Trends in Ecology and Evolution, 1992, 7: 368-372.
[154] Jones RH, Sharitz RR, Dixon PM et al. Woody plant regeneration in four floodplain forest Ecological Monographs, 1994, 64: 345-367.
[155] Reich PB. Variation among plant species in leaf turnover rates and associated traits, implications for growth at all life stages. Inherent Variation in Plant Growth, Physiological Mechanisms and Ecological Consequences (eds H.Lambers, H. Poorter & M.M.I. Van Vuuren), Backhuys, Leiden, 1998,467-487.
[156] Grubb PJ. A reassessment of the strategies of plants which cope with shortages of resources. Perspectives in Plant Ecology, Evolution and Systematics, 1998, 1:3-31.
[157] Grubb PJ, Metcalfe DJ. Rainforest dynamics: the need for new paradigms. Tropical Rainforest Research- Current Issues (eds Edwards DS, Booth WE & Choy SC),. Kluwer, Dordrecht, The Netherlands, 1996, 215-233.
[158] Wright IJ, Westoby M. Differences in seedling growth behaviour among species: trait correlations among species, and trait shifts along nutrient compared to rainfall gradients. Journal of Ecology, 1999, 87: 85-97.
[159] Reich PB, Buschena C, Tjoelker MG et al. Variation in growth rate and ecophysiology among 34 grassland and savanna species under contrasting N supply: a test of functional group differences. New Phytologist, 2003, 157: 617-631.
[160] Henns, DA, Mattson, WJ. The dilemma of plants, to grow or defend. Quarterly Review of Biology, 1992, 67:283-335.
[161] Cao, KF, Ohkubo T. Aoohnetry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan. Ecological Research, 1998, 13:217-227.
[162] Smith TM, Shugart HH, Woodward FI et al. Plant functional types. In: Vegetation Dynamics and Global Change (Eds A.M. Solomon & H.H. Shugart), Chapman & Hall, New York, 1993, 272-292.
[163] Gitay H, Noble IR. What are functional types and how should we seek them? In: Plant Functional Types-Their Relevance to Ecosystem Properties and Global Change (Eds T.M. Smith, H.H. Shugart & F.I. Woodward). Cambridge University Press, Cambridge, 1997, 3-19.
[164] Smith TM, Woodward FI, Shugart HH (eds.). Plant Function Types. Cambridge: Cambridge University Press, 1996.
[165] Woodward FL, Cramer W. Plant functional types and climatic changes. Journal of vegetation science, 1996, 7: 306-308.
[166] Lavorel, S, Mclntyre S, Landsberg Jet al. Plant functional classifications: from general groups to specific groups based on response to disturbance. Tree, 1997, 12:474-478.
[167] Craine JM, Tilman D, Wedin D et al. Functional traits, productivity and effects on nitrogen cycling of 33 grassland species. Functional Ecology, 2002, 16: 563-574.
[168] Tjoelker MG, Craine JM, Wedin D et al. Linking leaf and root trait Syndromes among 39 grassland and savannah species. New Phytologist, 2005, 167: 493-508.
[169] Poorter H, Navas ML. Plant growth and competition at elevated CO_2 : on winners, losers and functional groups. New Phytologist, 2003, 157:175-198.
[170] Wardle DA, Barker GM, Bonner KI et al. Can comparative approaches based on plant ecophysiological traits predict the nature of biotic interactions and individual plant species effects in ecosystems? Journal Ecology, 1998, 86: 405-420.
[171] Hanley ME, Trofimov S, Taylor G Species-level effects more important than functional group-level responses to elevated CO_2: evidence from simulated turves. Functional Ecology, 2004, 18: 304-313.
[172] Sutherland EK, Hale BJ, Hix DM. Defining species guilds in the Central Hardwood Forest, USA. Plant Ecology, 2000, 147: 1-19.
[173] Whitmore TC. Canopy gaps and the two major groups of forest trees. Ecology, 1989, 70(3): 536-538.
[174] Condit R, Hubbell SP, Foster RB. Assessing the response of plant functional types to climatic change in tropical forest. Journal of Vegetation Science, 1996, 7, 406-416.
[175] Agyeman VK, Swaine MD, Thompson J. Responses of tropical forest seedlings to irradiance and the derivation of a light response index. Journal of ecology, 1999, 87: 815-827.
[176] Davies SJ. Photosynthesis of nine pioneer Macaranga species from borneo in relation to life history. Ecology, 1998, 79(7): 2292-2308.
[177] Bloor JMG, Grubb PJ. Morphological plasticity of shade-tolerant tropical rainforest tree seedlings exposed to light changes. Functional Ecology, 2004, 18: 337-348.
[178] 周小勇，黄忠良，史军辉等．鼎湖山针阔混交林演替过程中群落组成和结构短期动态研究．热带亚热带植物学报，2004，12(4)：323-330．
[179] 刘美珍，蒋高明，于顺利等．浑善达克退化沙地恢复演替18年中植物群落动态变化．生态学报，2004，24(8)：1734-1740．
[180] 黄世能，李意德，王伯荪．海南岛尖峰岭两类热带山地雨林次生群落在15年演替过程中的林木消长．植物生态学报，2000，24(6)：710-717．
[181] 达良俊，杨永川，宋永昌．浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型．植物生态学报，2004，28(3)：376-384．
[182] 张家城，陈力，郭泉水等．演替顶极阶段森林群落典型树种分布的变动趋势研究．植物生态学报，1999，23(3)：256-268．
[183] 林露湘，曹敏，唐勇等．西双版纳刀耕火种弃耕地树种多样性比较研究．植物生态学报，2002，26(2)：216-222．
[184] 张继义，赵哈林，张铜会．赵学勇科尔沁沙地植被恢复系列上群落演替与物种多样性的恢复动态．植物生态学报，2004，28(1)：86-92．
[185] 唐勇，曹敏，张建侯等．西双版纳热带森林土壤种子库与地上植被的关系．应用生态学报，1999，10(3)：279-283．
[186] 周先叶，李鸣光，王伯荪等．广东黑石顶自然保护区森林此生演替不同阶段土壤种子库的研究．植物生态学报，2000，24(2)：222-230．
[187] 张庆费，由文辉，宋永昌．浙江天童森林公园植物群落演替对土壤物理性质的影响．植物资源与环境，1997，6(2)：36-40．
[188] 张全发，郑重，金义兴．植物群落演替与土壤发展之间的关系．武汉植物学研究，1990，8(4)：325-334．
[189] 彭少麟．恢复生态学研究进展及在中国热带亚热带的实践．四川师范学院学报(自然科学版)，2000．21(3)：221-227
[190] 邓瑞文，冯泳梅，陈天杏．三种相思(Acacia)的光合作用与蒸腾作用的研究．生态学报，1989，9(2)：128-131．
[191] 王博轶，冯玉龙．生长环境光强对两种热带雨林树种幼苗光合作用的影响．生态学报，2005，25(1)：23-30．
[192] 郭晓荣，曹坤芳，许再富．热带雨林不同生态习性树种幼苗光合作用和抗氧化酶对生长光环境的反应，应用生态学报，2004，15(3)，377-381．
[193] 孙谷畴，林植芳，林桂珠．不同光强下生长的几种亚热带森林树木的Rubisco羧化速率和碳酸酐酶的活性．武汉植物学研究，2001，19(4)：304-310．
[194] 杨小波．不同演替阶段森林群落典型种苗木叶形态解剖结构比较研究．海南大学学报自然科学版，1997，15(3)：212-217．
[195] 丁圣彦．常绿阔叶林演替系列中主要典型种叶片结构的比较．河南大学学报(自然科学版)，29(4)：76-80．
[196] 史刚荣，汤盈，张铮．淮北相山恢复演替群落典型树种叶片的生态解剖．植物生态学报，2006，30(2)：314-322．
[197] 王希华，张婕，张正祥．浙江天童国家森林公园主要常绿阔叶树种叶子寿命的研究．植物生态学报，2000，24(5)：625-629．
[198] 徐程扬．不同光环境下紫椴幼树树冠结构的可塑性响应．应用生态学报，2001，12(3)：339-343．
[199] 杨小波，文洪波，吴庆书等．海南中部山区次生林典型种枫香发育规律研究．海南大学学报自然科学版，2000，18(1)：54-58．
[200] 林业部林业区划办公室．中国林业区划．北京：中国林业出版社，1987．
[201] Alexander IJ, Faifley RI. Effects of N fertilization on populations of fine roots and mycorrhizae in spruce humus. Plant and Soil, 1983, 71: 19～53.
[202] Berntson GM, Farnsworth EJ, Bazzaz FA. Allocation with and between organs and the dynamics of root length changes in two birch species. Oecologia, 1995, 101: 439-447.
[203] Long SP, Humphries S, Falkowski PG.Photoinhibition of photosynthesis in nature. Annual Review of Plant Physiology and Plant Molecular Biology, 1994, 45: 633-662.
[204] Mort KA, Parkhurst DF. Stomatal responses to humidity in air and helox Plant, Cell and Environment, 1991, 14: 509-515.
[205] Roberts SW, Miller PC. Interception of solar radiation as affected by canopy organization in two Mediterranean shrubs. Oecologia Plantarum, 1977, 12: 273-290.
[206] Mcintyre S, Lavorell S. Livestock grazing in subtropical pastures: steps in the analysis of attribute response and plant functional types. Journal of Ecology, 2001,89: 209-226.
[207] Chapin, FS Ⅲ, Bloom A J, Field CB et al. Plant Responses of Multiple Environmental Factors. BioScience, 1987, 37: 49-57.
[208] Arnold E. Stomata. In: The Institute of Biology's Studies in Biology Ed. By Martin E. S., Donkin M. E. and Stevens R. A., 1983, 155.
[209] Mahall BE, Schlesinger WH. Effects ofirradiance on growth, photosynthesis, and water use efficiency of seedlings of the chaparral shrub Ceanothus megacarpus. Oecologia, 1982, 54:291-299.
[210] Abrams MD, Kubiske ME. Leaf structural characteristics of 31 hardwood and conifer tree species in central Wisconsin: influence of light regime and shade-tolerance rank. Forest Ecology and Management, 1990, 31: 245-253.
[211] Reich PB, Tilman D, Craine J. Do species and functional groups differ in acquisition and use of C, N and water under varying atmospheric CO_2 and N availability regimes? A field test with 16 grassland species. New Phytologist, 2001; 150: 435-448.
[212] Nowak RS, Ellsworth DS, Smith SD. Functional responses of plants to elevated atmospheric CO_2-do photosynthetic and productivity data from FACE experiments support early predictions? New Phytologist, 2004, 162: 253-280.
[213] Lee TD, Tjoelker MG, Ellsworth DS et al. Leaf gas exchange responses of 13 prairie grasslandspecies to elevated CO2and increased nitrogen supply. New Phytologist, 2001,405-418.
[214] Ellsworth DS, Reich PB, Naumburg ES et al. Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO_2 across four free-air CO_2 enrichment experiments in forest, grassland and desert. Global Change Biology, 2004, 10:2121-2138.
[215] Holscher D, Leuschner C, Bohman K et al. Photosynthetic characteristics in relation to leaf traits in eight co-existing pioneer tree species in Central Sulawesi, Indonesia. Journal of Tropical Ecology, 2004, 20：157-164．
[216] 符国瑷，冯绍信．海南五指山森林的垂直分布及其特征．广西植物，1995，15(1)：57-69．
[217] 骆土寿，李意德，陈德祥等．海南岛鸡毛松人工林群落种间联接性研究．生态学杂志，2005，24(6)：591-594．
[218] 广东省林业科学研究所．海南主要经济树种．北京：中国农业出版社，1964．
[219] 郑万钧．中国树木志Ⅲ．北京：中国林业出版社，1997．
[220] Escalona JM, Flexas J, Medrano H. Stomatal and non-stolnatal limitations of photosynthesis under water stress in field-growen grapevines. Australian journal of plant physiology, 1999, 16: 421-433.
[221] Kuppers, M. Ecological significance of above-ground architectural patterns in woody plants: a question of cost-benefit relationships. Trends in Ecology and Evolution, 1989, 4: 375-379.
[222] Gottschalk KW. Shade, leaf growth and crown development of Quercus rubra, Quercus velutina, Prunus serotina and Acer rubrum seedlings. Tree physiology, 1994, 14: 735-749.
[223] Sterck FJ, Bongers F. Crown development in tropical rain forest trees: patterns with tree height and light availability. Journal of ecology, 2001, 89: 1-13.
[224] Farque L, Sinoquet H, Colin F. Canopy structure and light interception in Quercus petreae seedlings in relation to light regime and plant density. Tree physiology, 2001, 21: 1257-1267.
[225] Beets, PN; Pollock DS. Accumulation and partitioning of dry matter in Pinus radiate as related to stand age and thinning. New Zealand Journal of Forest Science, 1987, 17: 246-271.
[226] King, DA. Tree form, height growth, and susceptibility, to wind damage in Acer saccharum. Ecology, 1986, 67: 980-990.
[227] Givnish, TJ. Plant stems: biochemical adaptation for energy capture and influence on species distribution. In: Plant Stems: Physiology and Functional Morphology (Eds.). BL Gartner, Academic Press, San Diego, 1995, 3-49.
[228] Ceschia, E. Environmental effects on spatial and seasonal variations of stem respiration in European beech and Norway spruce. Ph.D. Thesis, Swedish University of Agricultural Science, Uppsala, Sweden, 2001, 55.
[229] McDowell N, Barnard H, Bond BJ et al. The relationship between tree height and leaf area: sapwood area ratio. Oecologia, 2002, 132: 12-20.
[230] Wright SJ. Seasonal drought soil fertility and the species density of tropical forest plant communities. Tree, 1992, 7, 260-263.
[231] Letts MG, Mulligan M. The impact of light quality and leaf wetness on photosynthesis in north-west Andean tropical montane cloud forest. Journal of Tropical Ecology, 2005, 21: 549-557.
[232] Graham EA, Mulkey SS, Kitajima K et al. Cloud cover limits net CO_2 uptake and growth of a rainforest tree during tropical rainy seasons. PNAS, 2003, 100 (2): 572-576.
[233] Hogan KP, Smith AP, Samaniego M. Gas exchange in six tropical semi-deciduous forest canopy tree species during the wet and dry seasons. Biotropica, 1995, 27, 324-333.
[234] Quero JL, Villar R, Maranon T et al. Interactions of drought and shade effects on seedlings of four Quercus species: physiological and structural leaf responses. New Phytologist, 2006, 170: 819-834.
[235] Holmgren M. Combined effects of shade and drought on tulip poplar seedlings: trade-off in tolerant or facilitation? Oikos, 2000, 90: 67-78.
[236] Duan B, Lu Y, Yin C et al. Physiological responses to drought and shade in two contrasting Picea asperata populations. Physiologia Plantarum, 2005, 124: 476-484.
[237] Canham CD, Berkowitz AR, Kelly VR et al. Biomass allocation and multiple resource limitation in tree seedlings Canadian. Journal of Forest Research, 1996, 26:1521-1530.
[238] Smith T, Huston M. A theory of the spatial and temporal dynamics of plant communities. Vegetatio, 1989, 83: 49-69.
[239] Bazzaz FA, Picket STA. Physiological ecology of a tropical succession: a comparative review. Annual Reviews in Ecological Systems, 1980, 11: 287-310.
[240] Doley D, Unwin GL, Yates DJ.Spatial and temporal distribution of photosynthesis and transpiration by single leaves in a rainforest tree, Argyrodendron peralatum. Australian Journal of Plant Physiology, 1988, 15: 317-326.
[241] Ishibashi M, Terashima I. Effects of continuous leaf wetness on photosynthesis: adverse aspects of rainfall. Plant, Cell and Environment, 1995, 18: 431-438.
[242] Smith WK, McClean, TM. Adaptive relationship between leaf water repellency, stomatal distribution and gas exchange American. Jouranl of Botany, 1989, 76: 465-469.
[243] Nobel PS. Environmental productivity indexes and productivity for Opuntia ficus-indica under current and elevated atmospheric CO_2 levels. Plant, Cell and Environment, 1991, 14: 637-646.
[244] Allen DJ, Ort DR. Impacts of chilling temperatures on photosynthesis in warm- climate plants. Trends in Plant Science, 2001, 6: 36-42.
[245] Ribeiro RV, Lyra GB, Santiago AV et al. Diurnal and seasonal patterns of leaf gas exchange in bahiagrass (Paspalum notatum Flfigge) growing in a subtropical climate Grass and Forage. Science, 2006, 61: 293-303.
[246] Hastwell GT, Facelli JM. Differing effects of shade-induced facilitation on growth and survival during the establishment of a chenopod shrub. Journal of Ecology, 2003, 91: 941-950.
[247] Pacala SW, Canham CK, Saponara J et al. Forest models defined by field measurements, estimation, error analysis and dynamics. Ecological Monographs, 1996, 66: 1-43.
[248] Wyckoff PH, Clark JS. The relationship between growth and mortality for seven co-occurring tree species in the southern Appalachian Mountains. Journal of Ecology, 2002, 90:604-615.
[249] Pompa J, Bongers F. The effect of canopy gaps on growth and morphology of seedlings of rain forest species. Oecologia, 1988, 75: 625-632.
[250] Koike T. Leaf structure and photosynthetic performance as related to the forest succession of deciduous broad-leaved trees. Plant Species Biology, 1988, 3: 77-87.
[251] Lei TT, Lechowicz MJ. Shade adaptation and shade tolerance in saplings of three Acer species from eastern North America. Oecologia, 1990, 84: 224-228.
[252] Reich PB, Uhl C, Waiters MB et al. Leaf lifespan as a determinant of leaf strcture and function among 23 Amazonian tree species. Oecologia, 1991, 86: 16-24.
[253] Gleason SM, Ares A. Photosynthesis, carbohydrate storage and survival of a native and an introduced tree species in relation to light and defoliation. Tree Physiology, 2004, 24: 1087-1097.
[254] 张一平，窦军霞，于贵瑞等．西双版纳热带雨林太阳辐射特征研究．北京林业大学学报，2005，27(5)：17-25．
[255] Denslow JS. Tropical rainforest gaps and tree species diversity Annual Review of Ecology and Systematics, 1987, 18: 431-451.
[256] Scholes JD, Press MC, Zipperlen SW. Differences in light energy utilization and dissipation between dipterocarp rain forest tree seedlings. Oecologia, 1997, 109: 41-48.
[257] Silva EA, DaMatta FM, Ducatti C et al. Seasonal changes in vegetative growth and photosynthesis of Arabica coffee trees. Field Crops Research, 2004, 89: 349-357.
[258] Niinemets U, Kull O, Tenhunen JD. An analysis of light effects on foliar morphology, physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance. Tree Physiology, 1998, 18, 681-696.
[259] Stanhill G, Cohen S. Global dimming: a review of the evidence for a widespread and significant reduction in global radiation with discussion of its probable causes and possible agricultural consequences. Agricultural and Forest Meteorology, 2001, 107(4): 255-278.
[260] Liepert BG.Observed reductions of surface solar radiation at sites in the United States and worldwide from 1961 to 1990. Geophysical Research Letters, 2002, 29: 1010, 61-1, 5.
[261] 查良松．我国太阳辐射量区域性变化特征研究．地理研究,1996,15(2)：21-27．
[262] McGuffie K, Henderson- Sellers A. Is Canadian cloudiness increasing? Atlnosphere- Ocean, 1988, 26: 608-623.
[263] Henderson-Sellers A North American total cloud amount variations this century. Palaeogeograph. Palaeoclimatol. Palaeoecol., 1989, 75:175-194.
[264] Karl TR, Steurer PM. Increased cloudiness in the United States during the first half of the twentieth century: Fact or fiction? Geophysical Research Letters, 1990, 17: 1925-1928.
[265] Chen J, Carlson E, Del Genio AD. Evidence for strengthening of the Tropical General Circulation in the 1990s. Science, 2002, 295:838-841.
[266] Wielicki BA, Wong T, Allan RP et al. Evidence for large decadal variability in the tropical mean radiative energy budget. Science, 2002, 295:841-844.
[267] Price DT, Black TA. Effects of short-term variation in weather on diurnal canopy CO_2 flux and evaporation of a juvenile Douglas Fir stand Agricultural and Forest Meteorology, 1990, 50:139-158.
[268] Hollinger DY, Kelliher FM, Byers JN et al. Carbon dioxide exchange between an undisturbed old-growth temperate forest and the atmosphere. Ecology, 1994: 75, 134-150.
[269] Gu L, Fuentes JD, Shugart HH et al. Responses of net ecosystem exchange of carbon dioxide to changes in cloudiness: Results from two North American deciduous forests. Journal of Geophysical Research, 1999, 104: 31421-31434.
[270] Knapp AK, Smith WK, Young DR. Importance of intermittent shade to the ecophysiology of subalpine herbs. Functional Ecology, 1989, 3: 753-758.
[271] Baldocchi DD. Measuring and modeling carbon dioxide and water vapor exchange over a temperate broad-leaved forest during the 1995 summer drought. Plant, Cell and Environment, 1997, 20: 1108-1122.
[272] Ricklefs RE. Environmental heterogeneity and plant species diversity: a hypothesis. American Naturalist, 1977, 111: 376-381.
[273] Grantz DA, Moore PH, Zeiger E. Stomatal responses to light and humidity in sugarcane: prediction of daily time courses and identification of potential selection criteria Plant, Cell and Environment, 1987, 10: 197-204.
[274] Johnson JD, Ferrell WK. Stomatal response to vapour pressure deficit and the effect of plant water stress Plant, Cell and Environment, 1983, 6: 451-456.
[275] Guangdong Institute of Botany. Flora Hainanica, Volume Ⅳ. Science Press, Beijing, China, 1977. (in Chinese).
[276] Chun WY. Flora Hainanica, Volume Ⅱ. Science Press, Beijing, China, 1965. (in Chinese)
[277] 粟娟，王德祯，符史深．海南岛尖峰岭热带树木园主要树种的物候研究．林业科学研究，1994，7(3)：294-300．
[278] Rocha AV, Su HB, Vogel CS et al. Photosynthetic and water use efficiency responses to diffuse radiation by an aspen-dominated northern hardwood forest. Forest Science, 2004, 50(6): 793-801.