浙江省主要常绿阔叶树种功能评价及其人工恢复群落功能特征研究
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摘要
常绿阔叶林是中亚热带地带性植被分布的一个典型代表，由于长期人为破坏，其面积不断减小，生境的岛屿化与破碎化问题越来越突出，群落的功能日益下降。近年来，常绿阔叶林的生态恢复工作已在浙江省全面展开。因此，围绕常绿阔叶林造林树种的功能和植被恢复过程中群落功能的动态变化特征等方面的研究具有十分重要的理论和实践意义。本文应用群落生态学调查方法，结合光合生理生态测定技术和锥形量热仪燃烧特性测定技术，对主要常绿阔叶树种的光适应特征、可燃性及其它多种功能进行了研究，探讨了不同演替阶段人工恢复群落的物种多样性、抗火树种组成和植被功能群组成等功能特征及其存在的问题。研究结果表明：
(1)苗木的光适应能力是影响物种能否在群落中生存、生长和发育关键因素。研究发现处于演替中后期的常绿阔叶树种幼苗，对光环境的变化均有着较强的适应能力，秃瓣杜英、全缘冬青、小叶铁冬青、桃叶石楠、香樟等树种的表现尤为突出；荧光特性的研究也应证了这一观点，测定结果表明多数树种的PSⅡ反应中心并未受到明显氧化破坏，在全光照条件下仍能维持较高的光合电子传递活性，能通过提高非辐射性热耗散，消耗PSⅡ吸收的过剩的光能，从而保护PSⅡ反应中心免受因吸收过多光能而引起的光氧化伤害。同时，通过树种生理、形态和环境特性与其生长的回归关系分析，发现净光合速率、最大净光合速率、表观量子效率、叶片数量和环境湿度与高生长显著正相关，径生长则与净光合速率、叶片数量和环境温度正相关，而光强和环境温度与植物生长则呈较为明显的负相关关系。研究认为尽管强光不会造成多数常绿阔叶林树种幼苗的PSⅡ作用中心失活或破坏，但强光对常绿阔叶树种幼苗的生长有明显的抑制作用。
(2)利用除趋势对应分析和主成分分析对72个树种的抗火性能进行了综合评价，结果表明：微毛柃、撒金珊瑚、海桐、含笑、蚊母树、秃辦杜英抗火性能最好，珊瑚树、檵木、乐昌含笑、乳源木莲、青冈、广玉兰等12种树种次之。综合看来，多数常绿阔叶树种均表现出了较好的防火性能。同时，对树种的适宜性、萌生能力、生长速度、经济价值也进行了分析，为造林树种的功能需求提供了依据。
(3)将除趋势对应分析与TWINSPAN分类方法相结合，根据恢复群落与环境的关系剔除了一些环境因子差异过大的样方，降低了因为研究地生态特点和环境差异性所带来的试验误差。研究结果不仅反映了恢复样方种类组成上的差异，也反映了环境因子的作用，体现了各环境因子对植被的影响，以及这些因子在群落分异中的综合作用。应用生态种群来判别森林的恢复状态，TWINSPAN群落指示种的研究表明，人工构建的演替序列其生态种群与自然演替序列存在一定差异。
(4)人工恢复群落的物种多样性研究表明，人工恢复植物群落的生物多样性各个指标，随着群落演替梯度，基本呈现出同自然演替规律一致的变化，但总体来讲人工恢复的植物群落生物多样性是低于自然恢复群落的；人工群落的物种多样性与群落功能恢复的目的紧密相关，以恢复成常绿阔叶林为主的群落多样性较高，兼顾生物防火林带的群落多样性指数偏低，兼顾景观生态林的群落，稀有种数量偏多。群落抗火树种功能群组成研究表明，减少林内可燃物和群落快速构建是现行的生物防火林带建设技术的主要内容，短期内可以立竿见影，但由于防火树种组成单一，普遍存在林分结构差，关键种优势突出，保水保土功能差，难以实现自我更替等诸多问题；选择抗火性能较好的不同树种组成生物防火林带，有利于提高群落的防火性能，建立长期稳定的防火群落。对群落植被功能型的研究表明，模拟自然群落的功能群组成可以加速生态系统沿着自然演替的路线发展，甚至可以跨越某些演替阶段。总的来说，围绕功能群组成开展恢复群落的系统异质性、功能多样性和群落稳定性方面的研究，可以为常绿阔叶林生态系统的恢复技术与评价提供一定理论依据。
Evergreen broad-leaved forest is a typical zonalvegetation and climax community type in the middle subtropics. Because of the long-term artificial destruction, its area and community function were unceasingly small, diversity loss and environmental degradation have been a serious problem especially when more and more fragmentation and island were presented. In recent years, restoration of evergreen broad-leaved forest work started full swing in Zhejiang Province. Under the current situation, it is of great practical significance to study focusedly on the functions of evergreen broad-leaved species and dynamic characteristics of community during the process of vegetation restoration.
Based on the methods of community ecology, photosynthesis measurement and combustion technique of cone calorimeter, photosynthesis, combustibility and other characteristics of the main evergreen broad-leaved species were analysed. In addition, this paper also carried out on species diversity, fire-resistant tree species composition and vegetation functional group composition at different restoration stages. Aiming at the question of the forests cultivation in Zhejiang Province, some suggestions were given. The specific results as follow:
The photosynthetic adapt abilities of various tree specie is a key factor for tree's survival and growth at communities. The study shows that: evergreen broad-leaved species during the middle and later period succession have better adaptability under the different light surroundings, such as Elaeocarpus glabripetalus, Ilex integra, Photinia prunifolia, Cinnamommum camphora.
And Fluorescent Characteristics also verifies the above result. The experiment results indicate that: PSⅡreaction centers of majority tree species are not marked oxidized under the full light surroundings. Photosynthetic electron transport activity can maintain highly, in this way, it is helpful to enhance radiant energy dissipation and thus dissipate excess light energy. That is to say, the increase of non-photochemical quenching coefficient can conduce to protect PSⅡ, and mitigate the effect of high light on photosynthesis.
Multivariate regression analysis on the interrelationship between growth and physiology, plant morphology, environmental characteristic, the results indicate that: high growth is significantly positive correlated with Pn, Pmax, ARY and leaf number. Diameter growth is significantly positive correlated with Pn, leaf number and relative humidity of environment, but environmental temperature and light intensity are negatively correlated with plant growth, it means that in despite of that strong light doesn't inactivate or damage PSⅡreaction center, light intensity had little effect on evergreen broadleaved seedlings growth.
The fire-resistance properties of main evergreen broad-leaved species in Zhejiang Province are analysed synthetically through the method of principal component analysis (PCA) and detrended correspondence analysis (DCA) in this paper. The results are as follows: Eurya hebeclados, Michelia figo, Distylium racemosum, Elaeocarpaus glabripetalus, Aucuba japonica are in the first grade, their fire-resistance property are the best. And Loropetalum chinense, Viburnum odoratissimum, Michelia chapensis, Manglietia yuyuanensis, Cyclobalanopsis glauca (12species) are the second grade. In a word, most of evergreen broad-leaved species showed better fire-resistance property. Of course, ecosystem adaptability, sprout ability, growth rate, economic value can be also calculated.
By using DCA ordination and TWINSPAN classification, a total of 42 plots with artificial community restoration are partitioned into 9 community types. Both the methods of DCA ordination and TWINSPAN classification obtained very consistent results, and decreases the error of experiment for environmental deviations. The study is manifested not only the difference in trees species composition but also the functions of environment factors its incarnate the relationship between plants and environmenty. The method of TWINSPAN quantitative classification is applied to classify artificial community restoration, a succession serial and law of artificial community restoration further analyse the succession series of natural succession and artificial renewal of evergreen broad-leaved forest.
The regularity of developments of species diversity of artificial community restoration and community natural succession are similar mainly. On the whole, the species diversity artificial community restoration was lower.
It is very important that the correlation of species diversity and the aim of function restoration, the index of evergreen broad-leaved forest diversity of the nine communities was higher, the fire biological forest and landscape ecosystem wood is lower. Forest reduced combustible matter and composed of fast growing species is emphasis on the building of the fire biological forest, the ways may be immediately obvious, but there are many questions. For example, tree species composition is simple, forest structure is unreasonable, function of soil and water conservation is significantly decreased, and so on. In order to resolve the afore-mentioned issues, multiple tree species must be further enhanced and perfected. Simulation of structure features of natural community is the best ways for the accelerating restoration of evergreen broad-leaved forest, the community succession is on the rapid development phase rapidly during the least time. In conclusion, plant functional composition, heterogeneity of community and stability of community are correlated with functional group composition, the study of the issues can provide the theory basis for recovery technique and appraisal on evergreen broad-leaved forest ecological system.
(1)苗木的光适应能力是影响物种能否在群落中生存、生长和发育关键因素。研究发现处于演替中后期的常绿阔叶树种幼苗，对光环境的变化均有着较强的适应能力，秃瓣杜英、全缘冬青、小叶铁冬青、桃叶石楠、香樟等树种的表现尤为突出；荧光特性的研究也应证了这一观点，测定结果表明多数树种的PSⅡ反应中心并未受到明显氧化破坏，在全光照条件下仍能维持较高的光合电子传递活性，能通过提高非辐射性热耗散，消耗PSⅡ吸收的过剩的光能，从而保护PSⅡ反应中心免受因吸收过多光能而引起的光氧化伤害。同时，通过树种生理、形态和环境特性与其生长的回归关系分析，发现净光合速率、最大净光合速率、表观量子效率、叶片数量和环境湿度与高生长显著正相关，径生长则与净光合速率、叶片数量和环境温度正相关，而光强和环境温度与植物生长则呈较为明显的负相关关系。研究认为尽管强光不会造成多数常绿阔叶林树种幼苗的PSⅡ作用中心失活或破坏，但强光对常绿阔叶树种幼苗的生长有明显的抑制作用。
(2)利用除趋势对应分析和主成分分析对72个树种的抗火性能进行了综合评价，结果表明：微毛柃、撒金珊瑚、海桐、含笑、蚊母树、秃辦杜英抗火性能最好，珊瑚树、檵木、乐昌含笑、乳源木莲、青冈、广玉兰等12种树种次之。综合看来，多数常绿阔叶树种均表现出了较好的防火性能。同时，对树种的适宜性、萌生能力、生长速度、经济价值也进行了分析，为造林树种的功能需求提供了依据。
(3)将除趋势对应分析与TWINSPAN分类方法相结合，根据恢复群落与环境的关系剔除了一些环境因子差异过大的样方，降低了因为研究地生态特点和环境差异性所带来的试验误差。研究结果不仅反映了恢复样方种类组成上的差异，也反映了环境因子的作用，体现了各环境因子对植被的影响，以及这些因子在群落分异中的综合作用。应用生态种群来判别森林的恢复状态，TWINSPAN群落指示种的研究表明，人工构建的演替序列其生态种群与自然演替序列存在一定差异。
(4)人工恢复群落的物种多样性研究表明，人工恢复植物群落的生物多样性各个指标，随着群落演替梯度，基本呈现出同自然演替规律一致的变化，但总体来讲人工恢复的植物群落生物多样性是低于自然恢复群落的；人工群落的物种多样性与群落功能恢复的目的紧密相关，以恢复成常绿阔叶林为主的群落多样性较高，兼顾生物防火林带的群落多样性指数偏低，兼顾景观生态林的群落，稀有种数量偏多。群落抗火树种功能群组成研究表明，减少林内可燃物和群落快速构建是现行的生物防火林带建设技术的主要内容，短期内可以立竿见影，但由于防火树种组成单一，普遍存在林分结构差，关键种优势突出，保水保土功能差，难以实现自我更替等诸多问题；选择抗火性能较好的不同树种组成生物防火林带，有利于提高群落的防火性能，建立长期稳定的防火群落。对群落植被功能型的研究表明，模拟自然群落的功能群组成可以加速生态系统沿着自然演替的路线发展，甚至可以跨越某些演替阶段。总的来说，围绕功能群组成开展恢复群落的系统异质性、功能多样性和群落稳定性方面的研究，可以为常绿阔叶林生态系统的恢复技术与评价提供一定理论依据。
Evergreen broad-leaved forest is a typical zonalvegetation and climax community type in the middle subtropics. Because of the long-term artificial destruction, its area and community function were unceasingly small, diversity loss and environmental degradation have been a serious problem especially when more and more fragmentation and island were presented. In recent years, restoration of evergreen broad-leaved forest work started full swing in Zhejiang Province. Under the current situation, it is of great practical significance to study focusedly on the functions of evergreen broad-leaved species and dynamic characteristics of community during the process of vegetation restoration.
Based on the methods of community ecology, photosynthesis measurement and combustion technique of cone calorimeter, photosynthesis, combustibility and other characteristics of the main evergreen broad-leaved species were analysed. In addition, this paper also carried out on species diversity, fire-resistant tree species composition and vegetation functional group composition at different restoration stages. Aiming at the question of the forests cultivation in Zhejiang Province, some suggestions were given. The specific results as follow:
The photosynthetic adapt abilities of various tree specie is a key factor for tree's survival and growth at communities. The study shows that: evergreen broad-leaved species during the middle and later period succession have better adaptability under the different light surroundings, such as Elaeocarpus glabripetalus, Ilex integra, Photinia prunifolia, Cinnamommum camphora.
And Fluorescent Characteristics also verifies the above result. The experiment results indicate that: PSⅡreaction centers of majority tree species are not marked oxidized under the full light surroundings. Photosynthetic electron transport activity can maintain highly, in this way, it is helpful to enhance radiant energy dissipation and thus dissipate excess light energy. That is to say, the increase of non-photochemical quenching coefficient can conduce to protect PSⅡ, and mitigate the effect of high light on photosynthesis.
Multivariate regression analysis on the interrelationship between growth and physiology, plant morphology, environmental characteristic, the results indicate that: high growth is significantly positive correlated with Pn, Pmax, ARY and leaf number. Diameter growth is significantly positive correlated with Pn, leaf number and relative humidity of environment, but environmental temperature and light intensity are negatively correlated with plant growth, it means that in despite of that strong light doesn't inactivate or damage PSⅡreaction center, light intensity had little effect on evergreen broadleaved seedlings growth.
The fire-resistance properties of main evergreen broad-leaved species in Zhejiang Province are analysed synthetically through the method of principal component analysis (PCA) and detrended correspondence analysis (DCA) in this paper. The results are as follows: Eurya hebeclados, Michelia figo, Distylium racemosum, Elaeocarpaus glabripetalus, Aucuba japonica are in the first grade, their fire-resistance property are the best. And Loropetalum chinense, Viburnum odoratissimum, Michelia chapensis, Manglietia yuyuanensis, Cyclobalanopsis glauca (12species) are the second grade. In a word, most of evergreen broad-leaved species showed better fire-resistance property. Of course, ecosystem adaptability, sprout ability, growth rate, economic value can be also calculated.
By using DCA ordination and TWINSPAN classification, a total of 42 plots with artificial community restoration are partitioned into 9 community types. Both the methods of DCA ordination and TWINSPAN classification obtained very consistent results, and decreases the error of experiment for environmental deviations. The study is manifested not only the difference in trees species composition but also the functions of environment factors its incarnate the relationship between plants and environmenty. The method of TWINSPAN quantitative classification is applied to classify artificial community restoration, a succession serial and law of artificial community restoration further analyse the succession series of natural succession and artificial renewal of evergreen broad-leaved forest.
The regularity of developments of species diversity of artificial community restoration and community natural succession are similar mainly. On the whole, the species diversity artificial community restoration was lower.
It is very important that the correlation of species diversity and the aim of function restoration, the index of evergreen broad-leaved forest diversity of the nine communities was higher, the fire biological forest and landscape ecosystem wood is lower. Forest reduced combustible matter and composed of fast growing species is emphasis on the building of the fire biological forest, the ways may be immediately obvious, but there are many questions. For example, tree species composition is simple, forest structure is unreasonable, function of soil and water conservation is significantly decreased, and so on. In order to resolve the afore-mentioned issues, multiple tree species must be further enhanced and perfected. Simulation of structure features of natural community is the best ways for the accelerating restoration of evergreen broad-leaved forest, the community succession is on the rapid development phase rapidly during the least time. In conclusion, plant functional composition, heterogeneity of community and stability of community are correlated with functional group composition, the study of the issues can provide the theory basis for recovery technique and appraisal on evergreen broad-leaved forest ecological system.
引文
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