964 resultados para Altitudinal belt
Resumo:
The distribution of vascular plant species richness along an altitudinal gradient and their relationships with environmental variables, including slope, aspect, bank (flooding) height, and river width of the Xiangxi River, Hubei Province, were examined. Total vascular plant species richness changed with elevation: it increased at lower elevations, reached a maximum in the midreaches and decreased thereafter. In particular, tree and herbaceous species richness were related to altitude. Correlation analysis (Kendall's tau) between species richness and environmental variables indicated that the change in species richness in the riparian zone was determined by riparian environmental factors and characteristics of regional vegetation distribution along the altitudinal gradient. The low species richness at lower elevations resulted from seasonal flooding and human activities - agriculture and fuel collection - and the higher. Species richness ill (he midreaches reflected transitional zones ill natural vegetation types that had had little disturbance. These results oil species distribution in the riparian community could he utilized as a reference for restoration efforts to improve water quality of the emerging reservoir resulting from the Three Gorges Hydroelectric Dam project.
Microorganisms linked to Neoproterozoic microspar carbonate sedimentation in the Jilin-Liaoning area
Resumo:
Molar-tooth carbonate refers to a sort of rock that has ptygmatical folded structure comparable to the ivory. This kind of carbonate exists in a special time range (from Middle to Neoproterozoic). Its origin and the possibility to use it in stratigraphic correlation of the paleocontinent is the key task of the IGCP447, a project on Proterozoic molar tooth carbonates and the evolution of the earth (2001-2005). The importance lies in that the molar-tooth structure is the key to solving problems related to Precambrian biological and global geochemical events. The molar-tooth structure is associated with microorganisms. Development and recession of such carbonates have relations with the evolution process of early lives and abrupt changes in sea carbonate geochemistry. In recent years, based on researches on petrology, geochemistry and Sr isotope of molar-tooth carbonate in the Jilin-Liaoning and Xuzhou-Huaiyang area, the authors hold that it can be used as a marker for stratigraphic sequence and sedimentary facies analyses.
Microorganisms linked to Neoproterozoic microspar carbonate sedimentation in the Jilin-Liaoning area
Resumo:
Molar-tooth carbonate refers to a sort of rock that has ptygmatical folded structure comparable to the ivory. This kind of carbonate exists in a special time range (from Middle to Neoproterozoic). Its origin and the possibility to use it in stratigraphic correlation of the paleocontinent is the key task of the IGCP447, a project on Proterozoic molar tooth carbonates and the evolution of the earth (2001-2005). The importance lies in that the molar-tooth structure is the key to solving problems related to Precambrian biological and global geochemical events. The molar-tooth structure is associated with microorganisms. Development and recession of such carbonates have relations with the evolution process of early lives and abrupt changes in sea carbonate geochemistry. In recent years, based on researches on petrology, geochemistry and Sr isotope of molar-tooth carbonate in the Jilin-Liaoning and Xuzhou-Huaiyang area, the authors hold that it can be used as a marker for stratigraphic sequence and sedimentary facies analyses.
Resumo:
青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.
Resumo:
混农季节性放牧(agropastoral transhumance)通过作物种植和畜牧生产相结合的方式对不同海拔高度带上的资源进行相互补充利用,在亚洲兴都库什地区、青藏高原、横断山、东部及南部非洲、南美安第斯地区等具有悠久的历史。这种传统的生计系统几千年以来一直是居住在该地区的人类社会和自然生态系统相互作用的主要形式之一。这种传统的资源利用方式与山地自然植被以及特殊的山地人类文化和社会特征具有密切的协同演变关系。认识和理解这一关系,是山地生态学和人类学的核心科学问题之一。近年来,山地生态系统的多重功能性及动态演变对山区社会经济可持续发展的重要意义受到人们的不断关注。本文通过对云南省德钦县的12个自然村的混农季节性放牧以及对云南德钦、四川壤塘等山地植被格局特别是高海拔地带植被格局的的详细调查,探讨青藏高原东缘地区混农季节性放牧的主要特征、系统构成及相互关系,及其在全球变化、经济全球化和市场化及现代化过程中的变化趋势,分析混农季节性放牧与高山林线格局及生态系统的互动关系,旨在探讨山地地区人类活动与自然生态系统之间的互动关系,从而为山区社会经济可持续发展、环境建设和生物多样性保护等国家战略提供理论依据。 调查结果表明,混农季节性放牧是一种适应青藏高原东部高山峡谷地区环境因子及自然资源呈明显的垂直分布、资源数量稀少而时空分布异质性极高的生存环境的一种传统经济形式。这种传统的畜牧业的主要生产目的仍然是提供当地基本生存所需的产品,饲养牲口的种类和数量取决于农户的当地需求并且受资源的限制,因而维持在比较低的水平的。分布在不同海拔高度的放牧资源在一年中被牲口利用的时间也不同,互为补充,共同构成混农季节性放牧的资源基础。根据各社区永久居住点的位置和该村的土地资源特别是牧草地资源的分布范围,牲口迁移的距离和格局有较大的差异。。天然牧场仍然是最主要的畜牧业生产资源。混农季节性放牧中的农业系统和牧业系统互为补充,共同构成调查地区完整的的生计系统,农耕活动为放牧活动提供精饲料如粮食等和冬季饲料如秸秆, 其数量往往成为家庭畜牧业生产规模的主要决定因子之一。 通过对牲口数量和结构、牲口的时空迁移格局、牧业活动在整个经济活动中的相对重要性以及牧业活动和作物种植的关系方面的研究分析,混农季节性放牧在近几十年发生了深刻的变化。主要表现在牲口数量总体下降,牲口组成发生变化,牲口移动性降低、牧业活动的经济重要性下降以及牧业活动和种植活动之间的相互依存度降低等。上述变化的根本驱动力是发生在当地、地区及全球尺度上的环境、政治、社会经济、技术和文化等的变化,从而造成当地群众畜牧生产目标、土地利用和劳动力的分布等发生了变化。当地生计系统发生的改变可能会带来对方面而深刻的政治、社会经济、文化和生态影响。 混农季节性放牧这种古老的传统生计策略面临着许多挑战,如冬季饲料短缺、草场退化、缺乏市场竞争力、经济重要性降低、对年轻人缺乏吸引力、国家缺乏专门的政策指导等。与此同时,经济全球化、市场经济、新技术的应用、替代生计机会的增加、国家对于山地生态系统的作用的重新定位等也为传统生计系统转型、实现社会与生态共赢创造了机遇。 混农季节性放牧活动对亚高山及树线交错带生态系统系统的互动方式主要体现在以下几个方面:(1)牲口啃食、践踏等影响森林群落更新,改变森林群落的组成和结构,从而影响森林群落的演替进程和植被格局。林线边缘是搭建夏棚的首选地点,因此林线及树线交错地带就成了牲口活动的主要场所之一;(2)利用火烧开辟、维持和改良高山牧场; 3)在亚高山火灾迹地的放牧活动能够阻止火烧迹地的顺向演替; 4)牧民在林线附近获取建材和薪材等活动影响高山林线附近森林的结构和功能。 在调查区域,梅里雪山、白马雪山、甲午雪山的林线海拔高度在4200-4300m之间; 四川雅江、理塘一线,林线位置多在4300-4400m;四川壤塘二林场一带的林线主体在4100-4200m,在个别地区达到4300m; 在贡嘎山的南坡和东坡一带,林线位置在3600-3700m;而在四川松潘一带,林线位置主体在3700-3800米左右。树线高度的分布趋势和林线一致。混农季节性放牧及其有关人类利用活动使研究地区很多地方高山林线降低、树线交错带宽变窄或消失。在研究地区,总体情况是,阳坡和半阳坡(南坡、西南坡等)的林线和树线比阴坡和半阴坡(北坡、东北坡等)低,变化幅度达20-200m。这种差异主要是为了开辟牧场而人为清除了南向坡自然林线及其以上的植被从而使林线位置下降所致。在南坡自然林线保留得比较好的地方,林线和树线依然可以达到甚至超过北坡林线和树线的高度。放牧活动抑制了高山林线带火烧迹地的天然更新,从而使林线位置保持在目前的位置。 放牧活动对高山林线带森林群落更新的影响是显著的。自然林线内的乔木个体密度特别是新生苗和幼苗的密度大大高于非自然林线。没有放牧的自然林线及树线交错带内的I级个体(新生苗)密度达到725-2917株/公顷,而与之相对的处理样地内I级个体的密度只有0-228株/公顷;II级个体(高度10-50cm)也表现出类似的趋势,在没有放牧的自然林线及树线交错带样方内,其密度达到550-5208株/,而在放牧处理样方内只有14-321株/公顷。在非自然林线带样地内,在有正常放牧的样地内,完全缺乏I级个体。 从相对比例来看,没有放牧的样方内的I、II级个体在全部个体中所占的比例显著高于有放牧活动的样方。放牧使林线交错带的乔木幼苗数量显著减少,从而影响林线及树线交错带森林群落的天然更新过程。林线和树线交错带的灌木对乔木幼苗具有重要的保护作用,能够为树线树种如冷杉等幼苗的定居体提供有利的微气候环境,同时保护苗免受牲口的啃食和践踏。火烧以后接着进行放牧能够100%地抑制高山林线带的幼苗更新。 高山牧场放牧强度降低、使用时间缩短而低海拔地带放牧强度增加是研究地区混农季节性放牧系统的一个显著变化。这种变化也必然会引起各海拔带上的生态系统的变化。放牧强度的降低、生产性用火的停止将导致原来通过人工火烧而降低并通过进一步的火烧和放牧活动来维持的林线及其以上地带的灌木盖度和高度的增加,从而为林线森林群落的扩张创造条件。 青藏高原东部高山峡谷地区是我国重要的山地生态系统,在我国的生物多样性保护、生态环境建设、社会经济可持续发展战略中具有举足轻重的作用。正确认识人类特别是当地传统的生计系统与生态环境系统的互动关系是实现上述战略目标的前提。决策者必须以综合、系统的的视角协调促进社会经济可持续发展、保护生物及文化多样性和维持人、牲口和生态系统之间的平衡的多重目标。 Agropastoral transhumance, which makes a complementary exploitation of the natural resources at different altitudinal belts through a combination of migratory animal husbandry and crop cultivation, has a long history in Hindu-Kush Himalaya, Tibet Plateau, Hengduan Ranges, eastern and southern Africa and the Andes region of south America.For millennia, this traditional livelihood strategy has been one of the main forms of interaction between human societies inhabiting in these regions and their natural ecocystems. A close co-evolutionary relationship has been developed between this indigenous resources management systems and the mountain vegetation systems on the one hand and a unique set of cultural values and social features on the other. Understanding this relationship has been one of the core scientific issues in mountain ecology and anthropology. In recent years, the importance of the multiple functions of the mountain ecosystems and their dynamic changes in the sustainable socio-economic development of the mountain regions has gained increasing attention. This paper, which is based on a detailed study on the agropastoral practices of the 12 natural villages in Deqin County of Yunnan, and the mountainnn vegetation patterns in Deqin of Yunnan and Rangtang County of Sichuan, intends to reveal the major characteristics, system composition and the inter-relations of the subsystems of the agropastoral transhumance in Eastern Tibetan Plateau as well as the trends of changes of the system within the context of global changes, economic globalization and modernity process of China and analyze the relations between agropastoral transhumance and alpine ecosystem, ao as to understand the interactions between human activities and natural ecosystems of the mountains and provide theoretical basis for the national strategies in eocioeconomic development, environmental reconstruction and biodiversity conservation in the mountain regions. Results of the survey indicate that agropastoral transhumance in the investigated area is a traditional economic form that is highly adapted to the eastern Tibet Plateau where the topography features high peaks and deep gorges and where the highly variable environmental parameters and scanty natural resources exhibit a distinct vertical spectrum of distribution and great temporal and spatial heterogeneity. The main objective of pastoral management is still aimed at the production of basic goods and services of local people and thus the type and size of animals raised for each household mainly depend on local needs and are limited by the availability of natural resources. The scale of production is relatively low. Pastoral resources at different altidudinal belts are complementarily used at different seasons of a year and thus form the resources basis for agropastoral production of the study area. Migration distances and patterns vary with the location of the permanent settlements, the elevational distribution range of the resources of the villages concerned. Natural pastures (rangelands) are the main fodder resources and sumplement feedings only account for less than 5% of the total fodder consumption. Crop cultivation and pastoral activities support each other to form a complete livelihood system. The ability of the farmig lands (crop cultivation) to provide the pastoral activities with concentrates and sumplements often becomes a main factor limiting the scale of livestock production at household level. Agropastoral transhumance is experiencing drastic changes in recent decades as is reflected in the size and composition of animals, the seasonal migration pattern, the relative importance of pastoralism in the household economy and the interplays of agricultural and pastoral elements of the system. In general, there is a decline in animal population and mobility, a shift in animal composition to meet new needs arising from changed macro-economic situation, a decrease in the relative importance in the household economy and an increasing decoupling of agro&pastoral relations. The fundamental divers of these changes can be traced to environmental, social, economic, technological and cultural changes from local to global levels and such changes have further caused local changes in livestock management objectives, land use and distribution of labor forces. Changes in local livelihood systems could have profound political, socioeconomic, cultural and ecological conseuquences. Agropastoral transhumance, as an age-old traditional livelihood strategy, is facing multifacet challenges, such as winter fodder shortage, rangeland degradation, lack of market competitiveness, decrease in economic importance, lack of appreciation among the young generation and adequate policies from the government. At the same time, economic globalization, market economy, intrdoctution of new technologies, increase of alternative income generating opportunities and the national re-oreitation of policies on mountain ecosystems have all brought about new opportunities for the transformation of the traditional livelihood system and the synchronized development of local society and the environment. Agropastoral transhumance interacts with the ecosystems at the timberline and treeline ecotone mainly through the following aspects: 1)Animal browsing and stamping affect the regeneration process of the forest communities and alters the composition and structure of the forest which in turn affect the succession process and vegetation pattern of the forest communities. Forest edges are the priority locations for summer houses and therefore the timeline and treeline area becomes the major venues of aninal activities; (2)herders create, maintain and improve pastures through burning that remove the forest communities at the timeline and treeline ecotone; 3)immediate grazing on the fire sites can significantly prevent the fire sites from perogressive succession; and 4)herders harvesting of construction timber and firewoods affects the structure and functions of the forest communities at the timberline and treeline zone. Timberline position in the survey region shows geographical variations. It is around 4200-4300m in Meilixueshan, Baimaxueshan and Jiawuxueshan in Northwest of Yunnan and rises to 4300-4400m in Yajiang County and Litang County of Sichuan. In Rangtang of Sichuan, it is between 4100-4200m, though reaching 4300m in localized sites. In the southern and eastern slopes of Gongga Mountain, the timberline is only between 3600m and 3700m and in Songpan County at the upper reach of the Minjiang River the timberline is around 3700-3800m.Treeline pattern follows similar trend. In many places, agropastoral transhumance and related human activities have lowered the timberline and treeline and narrowed or removed the treeline ecotone. In the area of survey, generally speaking, timberlines and treelines are lower on the southern slopes than on the northern slopes, with a difference between 20 and 200m. This is mainly because that the use of fires to crerate pastures has removed the forest vegetation at the previous timberline and above. In fact, in many places, well-preserved forests on the south slopes have even high timberline position that the corresponding northern slopes. At subalpine zone, grazing activities could have prohibited the natural regeneration of many forest fire sites and maintained the forest position at the present level. Grazing has a significant impact on the regernation process of forest communities at the timberline zone. Natural timberline and treeline ecotone has much higher density of treeline species individuals especially the emergents and seedlings than the timberlines that are maintained by human activities. In natural timberline and treelien ecotone without grazing interference, the density of the I Class seedlings (less than 10cm in height) ranges 725-2917 /hm2; while that in the treatment plots (with grazing disturbance) is only 0-228//hm2;II Class seedlings (10-50cm)exhibit similar density trends, reaching 550-5208//hm2 in natural timberline without grazing but only 14-321//hm2 in the plots with grazing treatment. In the man-created timberlines, there is no I Class seedling at all in plots with normal grazing activities. In relative terms, in plots without grazing activities, the propotion of I Class and II Class seedlings is much higher than that in plots with grazing. Grazing activities have significantly reduced the number of seedlings in the timberline ane treeline ecotone, and thus affect the natural regeneration process of the forests. Shrubs at the timberline and treeline ecotone can effectively protect the seedlings from severe climate and animal tramping, thus increasing the survival rate of the seedlings. Grazing following fires can completely inhibit forest regeneration process at timberline. Changes in agropastoral transhumance will have great impact on the timberline and treeline pattern of the studied area. The decrease in grazing intensity on alpine pastrues and the cessation of the use of fires will result an increase in the cover and height of shrubs above the present human-maintained treeline, which will create further condition for the expansion of timberline forest communities. Eastern Tibet Plateau harbors some most important mountain ecosystems of China that are of vital importance to the country’s strategy in biodiversity conservation, environmental construction and sustainable sociaoeconomic development. A proper knowledge of the interactions between traditional livelihood systems and the ecosystems in the region is a precondition to the realization of the above strategic goals. Therefore, the decision-makers must have a holistic and systemic perspective so as to integrate the multiple objectives of promoting sustainable socioeconomic development, conserving biological and cultural diversity and maintaining the balances among people, animal population and the ecosystems.
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岷江上游地区高山/亚高山植被分布的坡向性分异显著,阴阳坡高山林线不仅物种组成差异明显,并且分布海拔呈现出阴坡高阳坡低的格局.阳坡林线树种主要是圆柏属乔木,林线类型多为渐变型,海拔高度大约在3 400m~3 800m;阴坡林线树种主要是冷杉,林线类型多为骤变型,海拔高度约在3 800m~4 400m.本研究采用土壤种子库物理筛选、室内萌发实验及野外群落调查等方法,对岷江上游地区阴坡岷江冷杉和阳坡祁连圆柏两类林线树种不同海拔梯度上土壤种子库以及幼苗库特征进行了调查,从土壤种子库和幼苗更新特征的角度对林线乔木树种种群更新特征进行了分析,进而对该地区高山林线在阴阳坡分布差异的原因进行了探讨,结果显示: 1.土壤种子库 阴坡:阴坡高山林线附近岷江冷杉土壤种子的平均密度大约为50.96粒/m2,其中树线以上10m处土壤种子密度为1.00粒/m2,树线处大约19.33粒/m2,林线交错带内土壤种子密度最高为136.83粒/m2,郁闭林内种子密度小于林线交错带,只有30.50粒/m2,种子平均空壳率为52%,霉变率达34%,完好种子只有6%.土壤种子库垂直分布特征为地被物层含种子比重最大,大约在67.50%左右;其次为0~2cm层,约18.84%左右;2~5cm层所占种子比例最小,约13.66%左右.霉变种子数量与土壤深度呈负相关. 阳坡:阳坡祁连圆柏土壤种子的平均密度为60.16粒/m2.树线以上10m处密度为1.92粒/m2,树线位置大约108.16粒/m2,林线交错带内平均为75.80粒/m2,郁闭林内种子密度小于林线交错带,只有20.00粒/m2.种子平均空壳率为36%,完好种子占49%,霉变率较低,大约为10%.阴阳坡林线树种土壤种子库垂直分布特征为:地被物层含种子最多,其次为0~4cm层,4~10cm层所占种子比例最小,霉变种子数量与土壤深度也呈负相关. 2. 幼苗库调查 阳坡:在树线以上区域没有发现幼苗,林线交错带内幼苗密度平均达3 250株/hm2,郁闭林内仅2 750株/ hm2.整个样地内1~2a幼苗很少甚至没有出现,3~10a的幼苗相对较多.空间分布上,祁连圆柏幼苗在林线交错带内接近随机分布,郁闭林内则介于随机分布和均匀分布之间. 阴坡:在树线以上幼苗密度为1 250株/ hm2,全部为1~2a幼苗,林线交错带内幼苗密度平均达7 000株/ hm2,郁闭林内达6 250株/ hm2.林线附近岷江冷杉幼苗丰富度以及幼苗的出现频率明显高于祁连圆柏,年龄结构也较祁连圆柏完整.岷江冷杉幼苗空间分布除了树线处幼苗的分布为随机分布,其他海拔则为集群分布. 3.从不同土壤深度的种子总量和幼苗数量的相关性检验发现,当年生幼苗数量跟表层种子总量相关性极显著, 但是两年生幼苗的数量与底层种子数量相关性显著.土壤种子在土壤中的垂直分布格局从一定程度上可以反映种子库的年际特征.岷江冷杉土壤种子库较丰富,种子散布后的存活力随着时间的变化逐渐下降,属于季节性瞬时种子库;祁连圆柏土壤种子散布格局为集群型分布,成熟种子大部分散布在母株冠幅内,属于永久性土壤种子库. 4.在阴坡林线交错带及以上区域还存在较为丰富的乔木土壤种子,并且在树线以上区域还发现了少量的岷江冷杉幼苗.从样地乔木的年龄结构发现,在林线交错带内上部到树线位置主要以幼龄林为主,且年龄结构完整,基本符合入侵性林线特征;阳坡林线交错带内幼苗出现频率很低,树线以上区域虽然存在种子库,但是没有幼苗出现,在林线交错带内乔木径级差距很大,年龄结构异常不完整,这种特征的林线将会面临两个可能结果:一种是维持现有状态,保持平衡;另外一种就是退化,但阳坡林线的实际动态趋势还有待长期定点研究. Treelines on the upper region of Minjiang River differ between the north aspect and the south aspect in their appearances, altitudes and tree species. On the north aspect, trees of Abies form a sharp and abrupt treeline ranging from 3800m to 4400m, while on the south the treeline is generally lower(3 400~3 800m), more open and gradual and mostly composed of Sabina. In this study, we examined the altitudinal gradients of soil seed banks and seedling recruitments at the treeline ecotones of a N-aspect and a S-aspect by using soil sieving, germination experiment and field investigations, analyzed the characteristics of population regeneration of tree species at the transitional zone and presented a analysis of the causes to the aspect-related difference in treeline patterns in the study area. Major results of our study include: 1. Soil seed bank N-aspect: Of the 50 plots investigated, the average density of soil seeds is 50.96/m2, in which well-formed seeds account for 6%, empty seeds 52%, parasitized seeds34%, and seeds damaged by animals 8%. The size of soil seed bank varies along altitude, being 1.00 seeds /m2 at the 10m above the treeline and ca.19.33 seeds/m2 at the upper limit of treeline. The highest density (136.83 seeds/m2) occurs at the treeline ecotone. By contrast, the density of soil seed for the closed forest is only 30.50 seeds/m2. In terms of vertical strata, 67.50% of the total seeds are at the surface layer, 18.84% at the middle layer (0~2cm) and 13.66% at deeper layer (2~5cm). The number of parasitized seeds is negatively correlated to soil depth. S-aspect: Of the 50 plots investigated, the average density of soil seeds is 60.16 seeds/m2, and the well-formed seeds account for 49%, empty seeds 36%, parasitized seeds10%, and seeds damaged by animals 1%. The size of soil seed bank varies along altitude, with 1.92 seeds/m2 recorded at the10m above the treeline,108.16 seeds/m2 at the upper limit of treeline, and 75.80 seeds/m2 at the treeline ecotone, while that for the closed forest is 20.00 seeds/m2. The number of seeds decreases with the depth of soil. As is on the N-aspect, the size of soil bank, from large to small, follows the order of the surface layer, the middle layer (0~4cm) and the bottom layer (4~10cm). The number of parasitized seeds is also negatively correlated to the depth of the soil. 2. Seedling bank N-aspect: A mean maximum seedling abundance of 31 000 seedlings/hm2 was recorded near alpine treeline at growing season. The density of seedlings is 1 250 seedlings/ha (all being 1 or 2 years old) at the alpine meadow 10m away above treeline, 7 000 seedlings/ha at treeline ecotone and 6 250 seedlings/ha for closed forest.The spatial distribution of Abies faxoniana seedlings is random at the upper limit of the treeline but clumped at other altitudes. S-aspect: No seedlings were found at the alpine meadow 10m away from the treeline. The density of seedlings was 3 250 seedlings/ha at treeline ecotone and 2 750 seedlings/ha for the closed forest.Hardly any 1 year current and 2 year-old seedlings appeared at the plots. The spatial distribution of Sabina przewalskii seedlings is random at treeline ecotone and between “random” and “even” forest closed forest. 3.Correlation tests of seedling population and seed bank at different soil layers indicated that the emergents were strongly correlated to seed bank at surface layer while the number of two-year seedlings was significantly correlated to the seed bank at the bottom of soil layer, indicating that germination mainly occurs at the soil surface while the middle or bottom layer was the reserve for non-germination or dead seeds. It can thus be postulated that Abies faxoniana soil seed bank is of seasonal transient type. By contrast, the soil seed bank of Sabina przewalskii is of persistent type and the soil seeds and seedlings of this species occurred more frequently near the islands of adult trees. 4.A good many soil seeds of both tree species were found near the treeline ecotone and above at N- and S-aspects. A few young seedlings were found above the Abies treeline. Investigation of five altitudinal transects respectively on N- and S-aspects indicated that Abies faxoniana has a more complete age structure than the stands of Sabina przewalskii. The age of firs decreased from closed forest to the upper limit of treeline, which suggests that the Abies treeline is advancing to higher altitude. While on the south aspect, only few Sabina przewalskii soil seeds and nearly no seedlings were found above the treeline ecotone. The stands exhibit extremely great difference in diameter classes with significantly incomplete age structure. This would lead to two possible results for the treelines: maintaining an equilibrium state at the current position or degenerating. But more studies should be carried out at longer time scales or larger spatial scales to understand whether the Sabina treeline is degenerating.
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海拔梯度造成的环境异质性,如崎岖的地形、复杂的植被结构以及花期延迟等可能会极大地影响到物种的形态和遗传变异格局。理解物种形态和遗传变异的海拔格局对于物种多样性的管理和保护是非常重要的。尽管植物群体遗传学是一个飞速发展的研究领域,然而与海拔相关的形态变异、遗传变异及群体间遗传差异的研究却很少。到目前为止,还不清楚遗传变异与海拔之间是否必然的相关性。 川滇高山栎是一种重要的生态和经济型树种,广泛分布于中国西南的四川、西藏、贵州和云南省的高海拔地区,在保持水土、调节气候方面起着十分重要的作用。尽管主要受阳光限制而仅分布于阳坡,但其海拔梯度范围较大,表明川滇高山栎对不同的环境具有很强的适应性。本文通过叶型及生理响应、微卫星分子标记和扩增性片段长度多态性方法,试图探索川滇高山栎叶沿海拔梯度的形态和生理响应及其沿海拔梯度的遗传变异格局,为川滇高山栎的保护和利用提供进一步的遗传学理论依据和技术指导。 对叶形、含氮量及碳同位素的试验结果表明,平均比叶面积、气孔密度、气孔长度和气孔指数等气孔参数随海拔的升高呈非线性变化。在海拔大于2800 m时,川滇高山栎的比叶面积、气孔长度和气孔指数都随海拔升高而降低,但是在海拔小于2800 m时,这些指标都随海拔的升高而增大。相对而言,单位叶面积的含氮量和碳同位素则表现出相反的变化模式。另外,比叶面积是决定碳同位素沿海拔梯度变化的最重要参数。本研究结果表明,海拔2800 m附近是川滇高山栎生长和发育的最适地带,在这里生长的植物叶片厚度更薄、气孔更大、叶碳同位素值更小。 利用六对微卫星引物对五个不同海拔川滇高山栎群体遗传多样性进行研究,结果表明,群体内表现出较高的遗传多样性,平均每位点等位基因数11.33个,平均期望杂合度达0.820。群体间差异较小,分化仅为6.6%。聚类分析也并没有显示出明显的海拔格局。然而低频率等位基因却与海拔呈显著性正相关(R2=0.97, P < 0.01),表明在高海拔处,川滇高山栎以更多的稀有基因来适应恶劣的环境条件。本试验结果表明由海拔梯度形成的选择性压力对川滇高山栎群体的遗传变异影响并不明显。 为了进一步探讨川滇高山栎群体遗传变异与海拔之间的相互关系,我们还对其进行了扩增性片段长度多态性分析。结果表明:(1)随海拔的升高(从群体WL2到群体WL5),群体内遗传变异降低,而群体间遗传差异增加;(2)低海拔群体WL1表现出最低的遗传变异性(HE = 0.181),同时与其余四个群体间呈现出最大的遗传差异性(平均FST = 0.0596);(3)在除去低海拔群体WL1后,Mantel检测表明群体间遗传距离与海拔距离之间表现出正相关性。另外,研究结果还表明,遗传变异受生境条件(过度的湿热环境)及人为干扰(火烧、砍伐和放牧)的影响,这一点至少在低海拔群体WL1上发生了作用。 通过叶形态、生理及DNA分子水平的研究,结果表明叶形态特征和碳同位素与海拔紧密相关,与海拔之间呈非线性变化,海拔2,800 m附近是川滇高山栎生长和发育的最适地带。海拔梯度在一定程度上会影响到川滇高山栎群体的遗传变异结构,但在这样一个狭窄的地理分布区域里,这种影响并不足以导致群体间较大的遗传分化。同时生境条件及人为干扰也是影响遗传变异的限制性因子,不容忽视。 Altitudinal gradients impose heterogeneous environmental conditions, such as rugged topography, a complex pattern of vegetation and flowering delay, and they likely furthermore markedly affect the morphological and genetic variation pattern of a species. Understanding altitudinal pattern of morphological and genetic variation at a species is important for the management and conservation of species diversity. Although plant population genetics is a fast growing field of research, there are only few recent investigations, which analyzed the genetic differentiation and changes of intra-population variation along altitudinal gradients. At present, it is still unclear whether there are some common patterns of morphological and genetic variation with altitude. Quercus aquifolioides Rehder & E.H. Wilson, which is an important ecological and economical endemic woody plant species, is widely distributed in the Yunnan and Sichuan provinces, Southwest China. Its large range of habitat across different altitudes implies strong adaptation to different environments, although it is mainly restricted to sunny, south facing slopes. It plays a very important role in preventing soil erosion, soil water loss and regulating climate, as well as in retaining ecological stability. In this paper, we tried to understand the altitudinal pattern of morphological and genetic variation along altitudinal gradients through the experiments of leaf morphological and physiological responses, microsatellite analysis and AFLP markers. In leaf morphological and physiological responses experiment, we measured leaf morphology, nitrogen content and carbon isotope composition (as an indicator of water use efficiency) of Q. aquifolioides along an altitudinal gradient. We found that these leaf morphological and physiological responses to altitudinal gradients were non-linear with increasing altitude. Specific leaf area, stomatal length and index increased with increasing altitude below 2,800 m, but decreased with increasing altitude above 2,800 m. In contrast, leaf nitrogen content per unit area and carbon isotope composition showed opposite change patterns. Specific leaf area seemed to be the most important parameter that determined the carbon isotope composition along the altitudinal gradient. Our results suggest that near 2,800 m in altitude could be the optimum zone for growth and development of Q. aquifolioides, and highlight the importance of the influence of altitude in research on plant physiological ecology. Genetic variation and differentiation were investigated among five natural populations of Q. aquifolioides occurring along an altitudinal gradient that varied from 2,000 to 3,600 m above sea level in the Wolong Natural Reserve of China, by analyzing variation at six microsatellite loci. The results showed that the populations were characterized by relatively high intra-population variation with the average number of alleles equaling 11.33 per locus and the average expected heterozygosity (HE) being 0.779. The amount of genetic variation varied only little among populations, which suggests that the influence of altitude factors on microsatellite variation is limited. However, there is a significantly positive correlation between altitude and the number of low-frequency alleles (R2=0.97, P < 0.01), which indicates that Q. aquifolioides from high altitudes has more unique variation, possibly enabling adaptation to severe conditions. F statistics showed the presence of a slight deficiency of heterozygosity (FIS=0.136) and a low level of differentiation among populations (FST=0.066). The result of the cluster analysis demonstrates that the grouping of populations does not correspond to the altitude of the populations. Based on the available data, it is likely that the selective forces related to altitude are not strong enough to significantly differentiate the populations of Q. aquifolioides in terms of microsatellite variation. To further elucidate genetic variation pattern of Q. aquifolioides populations under sub-alpine environments, genetic variation and differentiation were investigated along altitudinal gradients using AFLP markers. The altitudinal populations with an average altitude interval of 400 m, i.e. WL1, WL2, WL3, WL4 and WL5, correspond to the altitudes 2,000, 2,400, 2,800, 3,200 and 3,600 m, respectively. Our results were as follows: (i) decreasing genetic variation (ranging from 0.253 to 0.210) and increasing genetic differentiation with altitude were obtained from the WL2 to the WL5 population; (ii) the WL1 population showed the lowest genetic variation (HE = 0.181) and the highest genetic differentiation (average FST = 0.0596) with the other four populations; (iii) the positive correlation was obtained using Mantel tests between genetic and altitude distances except for the WL1 population. Our results suggest that altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides populations to some extent. In addition, habitat environments (unfavorable wet and hot conditions) and human disturbances (burning, grazing and felling) were possible influencing factors, especially to the low-altitude WL1 population. The present study shows that there were close correlations between morphological features and carbon isotope composition in our data. This indicates that a coordinated plant response modified these parameters simultaneously across different altitudes. Around 2,800 m altitude there seems to be an optimum zone for growth and development of Q. aquifolioides, as indicated by thinner leaves, larger stomata and more negative d13C values. All available evidence indicates altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides to some extent. Decreasing genetic variation and increasing genetic differentiation with altitude was obtained except for the WL1 population. And the environment of habitats and human disturbances were also contributing factors, which impact genetic variation pattern, especially to the low-altitude WL1 population.
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米亚罗地区是四川西部较为典型的亚高山针叶林区域之一。为建立该地区主要针叶树种岷江冷杉、云杉、紫果云杉和红杉的年轮宽度年表资料,了解不同海拔高度岷江冷杉原始林和不同恢复过程的人工针叶林及次生混交林树木径向生长规律,结合样地调查,用生长锥钻取了树木芯样做年轮生态学分析。芯样经过标准化程序固定和打磨抛光后,用WinDENDRO图像分析系统测量年轮宽度序列,用COFECHA程序交叉定年和控制测量数据质量,用ARSTAN程序建立了4个主要针叶树种的地区年表和不同海拔高度岷江冷杉林及人工针叶林和次生混交林针叶树的样地年表。 4个主要针叶树种年轮宽度年表的平均敏感度低于0.2,而其晚材宽度年表都具相对较高的平均敏感度。早材宽度与年轮总宽度标准化年表间的相关系数均在0.9以上;晚材宽度与年轮总宽度标准化年表间的相关系数则种间差异较大,红杉的最高,岷江冷杉的最低。岷江冷杉晚材宽度与年轮总宽度的相关性从1970年以后明显下降,而其他种的相关系数则随时间变化较小。树种之间标准化年表显著正相关,而云杉与紫果云杉和红杉与岷江冷杉之间相关系数明显较高。年表序列的第1主分量表达了4个树种树木共同径向生长变化格局;第2至第4主分量分别表达了云杉属和冷杉属、常绿针叶树种和落叶针叶树种以及云杉和紫果云杉树木径向生长变化差异。 不同海拔高度的8个岷江冷杉样地年轮宽度年表序列敏感度大体上随海拔高度升高而降低。各样地早材宽度与年轮总宽度年表之间的相关系数均在0.9以上,且随海拔高度变化不大;晚材宽度与年轮总宽度之间的相关系数随海拔高度的变化较大,并有随海拔升高而降低的趋势。样地年表序列之间相关系数差异很大,高海拔样地年表间多为显著正相关;低海拔样地年表间的相关系数变化不一;高海拔和低海拔样地年表之间相关性较差,且多不显著。样地年表的第1主分量能解释年表序列总方差的37.5%,反映了不同海拔高度岷江冷杉林木共同的径向生长变化格局;第2和第3主分量分别解释总方差的24.5%和18.2%,表现出明显的高海拔和低海拔样地树木间不同的径向生长变化,除一些样地例外,它们一般与低海拔样地年表有正相关,与高海拔样地年表有负相关。在那些另外的样地,海拔以外的其他因素可能也影响了树木径向生长变化。不同海拔高度样地林木的生长抑制和生长释放频率在不同时期表现出较大的差异,表明了不同的干扰历史和林木补充时间。 人工针叶林和次生混交林各样地林木早材宽度与其年轮总宽度年表之间相关系数均高达0.9以上;晚材宽度与年轮总宽度年表之间也都显著正相关,但人工针叶林样地的明显较高。样地年表序列之间的相关关系表现为,林分起源和经营管理相似的样地年表之间的相关系数明显较高,如人工针叶林与人工针叶林尽管树种不同,但样地年表之间显著正相关,而与次生混交林样地年表间关系不显著;反之亦然。综合比较各项生长参数及不同时期的树木径向生长速率,人工针叶林树木的胸径增长至少在40年以内是优于次生混交林的同种(或不同种)针叶树的。不同样地林木生长释放和生长抑制及人工针叶林树木胸高断面积增长分析表明,除严重的人为干扰外,林分郁闭后林木密度过大是造成高频率生长抑制的主要原因,在林分发育的适当时期通过抚育间伐等措施调控林分密度,是保证林木胸高断面积在一定时期内保持较高的连年增长的关键。日本落叶松作为引进的树种,在海拔3100 m左右种植表现良好,近30年来各项生长指标均高于林龄相近的云杉人工林,因此,适当用其作为川西亚高山针叶林采伐迹地快速恢复是合理的。 Miyaluo area is one of the typical regions covered by subalpine coniferous forests in western Sichuan province of southwestern China. To develop the regional tree-ring width chronology series for the dominant conifers such as Abies faxoniana, Picea asperata, P. purpurea and Larix potaninii, and to understand the radial growth patterns of conifers in Abies faxoniana natural forest stands at different altitudes, and in coniferous plantations and natural regenerated mixed stands in their different restoring processes as well, increment cores were sampled in the field together with conventional plots investigations for dendroecological analyses. After the increment cores being prepared according to standard procedures, the ring widths (total-ring and intra-ring widths) were measured with a WinDENDRO image-analysis system, and the measured tree-ring sequences were crossdated and quality-controlled with the software COFECHA. Using the software ARSTAN, we developed tree-ring width based chronology series of the four dominant conifers, eight site-specific Abies faxoniana chronologies, and seven site-specific chronologies of conifers in coniferous plantations and natural regenerated mixed stands. Mean sensitivities for total ring width chronologies of the four sampled dominant conifers were all below 0.2, while those for the latewood width chronologies of the same species were relatively much higher. Correlation coefficients between standard earlywood and total ring width chronologies of the four conifers were all above 0.9, but those between standard latewood and total ring width chronologies exhibited differences among species, with the coefficient of Larix potaninii the highest and that of Abies faxoniana the lowest. Correlation coefficients between latewood and total ring width of A. faxoniana obviously decreased from 1920-1970 for successive 50-year segments with 10-years lag analyses, though the same for the other three species changed unnoticeably with time. Tree-ring standard chronologies among species showed significant positive correlations, with the correlation coefficients between chronologies of Picea asperata and P. purpurea, and of Larix potaninii and Abies faxoniana relatively much higher. The first principal component of tree-ring chronologies represented the common radial growth patterns of the four conifers in Miyaluo area. The second, third and fourth PCs expressed the differences in radial growth responses for the genus Picea and Abies, for the evergreen and deciduous confers, and for the two species of the genus Picea, respectively. In general, mean sensitivities of the eight Abies faxoniana site-specific tree-ring width chronologies decreased with increasing altitude. The correlation coefficients between earlywood and total ring width chronologies for all sites reached 0.9, which did not change much with altitude; but those between latewood and total ring width chronologies diversified, with a decreasing tendency from lower altitudinal sites to higher altitudinal sites. Correlation coefficients among site chronologies varied considerably, with significant positive correlations among higher site chronologies, mixed results among lower site chronologies, and poor and insignificant correlations between chronologies of higher site and lower site. The first PC, which represents 37.5% of the total variance, reflected a common radial growth response at sites of different altitudes, and it showed a tendency of explaining more variance with increasing altitude. The second and the third PCs contributed to 24.5% and 18.2% of the total variance, respectively, exhibiting distinctive differences in radial growth responses between low- and high-altitudinal sites. With some exceptions, the radial growth represented by the second and third PCs had a positive correlation with that at the low-altitudinal sites and a negative correlation with that at the high-altitudinal sites. For those exceptional sites, factors other than altitude might also play a role in affecting tree-ring growth variations. Trees in stands of different altitudes showed great differences in frequencies of growth suppressions and releases through times, suggesting different disturbance histories and periods when trees recruiting to the canopy. Correlation coefficients between earlywood and total ring width chronologies for all sites of coniferous plantations and natural regenerated mixed stands were also above 0.9; and the same between latewood width and total ring width chronologies all positively correlated, too, with the coefficients of the coniferous plantations obviously much higher. Correlations among site chronologies showed that the coefficients among sites with similar stand origin and management regimes were much higher than those among sites with different stand origin and management regimes. For example, significant positive correlations were found for chronologies among different coniferous plantations, irrespective of species differences; while insignificant correlations between chronologies of the same conifer from a coniferous plantation and a natural regenerated mixed stand, and vise versa. Integrative comparisons of different tree growth parameters and radial growth rates at different stages indicated that the diameter at breast height (DBH) increments for trees in coniferous plantations were faster than those for trees of the same (or different) species in the natural regenerated mixed stands, at least within their first 40 years of stand development. Analyses of growth releases and suppressions, and basal area increments of trees in different stands demonstrated that over-dense individuals after canopy closure was the main factor resulting in high frequencies of radial growth suppressions, with some exceptions of severe man-made disturbances. Therefore, to ensure a continuous basal area current annual increment in certain periods, tree density controlling through thinning in due time during the stand development process are necessary. It should be mentioned that, as an introduced conifer to Miyaluo area, Larix kaempferi grew quite well at altitude of ca. 3100 m after planting in 1970s. In their near 30 years of stand development, Larix kaempferi trees exhibited faster growth in various parameters than Picea asperata trees of the similar stand age did. Thus we think it reasonable to use Larix kaempferi as a fast restoring species at appropriate sites of cutting blanks of subalpine coniferous forests in western Sichuan.
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辐射传输研究是贯穿森林生态系统的纽带,太阳辐射为植物的生长发育提供光合能量、适宜的环境温度以及发育信息。一方面,气候变化使到达地面辐射能的质和量发生变化,影响到植被的生长发育,改变森林的结构,而森林结构的变化又会影响林冠内辐射能的分配和质量,这些变化会进一步影响到林下土壤温度,改变森林根系活性以及土壤营养转化的效率;连锁反应的结果有可能会使森林生态系统的生产力发生变化,改变碳素和氮素源库的调节方向,从而反馈影响地球气候系统。另一方面,人类作为生态系统的成员,必然需要森林生态系统为其提供更多的原材料和更好的生态服务功能,如何实现这些目标,就需要人类适度调整干预方式和频度,达到预期的目的。本文在建立适合于川西亚高山森林的叶面积测量技术、光照辐射模型和土壤温度变化模型的基础上,对川西亚高山地带森林生态系统的辐射传输特征进行了分析,并从森林结构的角度探讨了林分内的辐射分布以及对土壤温度的影响。主要成果如下: 1. 提出了一种照相法测量叶面积的方法。通过对摆放在平面上的叶片照相,利用投影变化,把非正射图像转化为正射图像,然后经过计算机图像处理得到每一片叶片的面积、周长、长度、宽度等信息。这种方法可使用户以任意方向和距离拍摄处于平面上的叶片,能同时处理大量的叶片,适于野外离体或活体叶片测量。叶片面积分辨率可调,分辨率可以与常用的激光叶面积仪相近甚至更高,而且叶片图像可以存档查询。 2. 提出一种模拟林内光照变化的模型。利用林冠半球照片,记录视点以上半球内的林冠构件空间分布,作为林冠子模型;天空辐射子模型采用国际照明委员会(CIE)的标准晴天和阴天以及插值模型。该模型能够模拟林下某一位点处的实时光斑变化。 3. 提出一种土壤温度变化模型。把土壤视为具有容量和阻力性质的结构,利用电阻和电容器件构建土壤能量分布模型。外界太阳辐射能经过植被以及其它一些能量分配器后进入土壤,其中有一部分转化为土壤势能,即土壤温度。土壤温度的变化类似于电池的充放电过程。在已知模型参数的情况下,可以从太阳辐射计算土壤温度的变化。在模型参数未知的情况下,通过输入和输出值推算模型的参数,而模型参数中的时间常数与土壤组成和含水量有关,这样就可以知道土壤水分的变化情况。 4. 从王朗亚高山森林典型样地林分结构的测量获得林地三维结构图、树冠形态、叶面积密度等参数,这些参数输入到Brunner (1998)开发的tRAYci 模型中计算出一段时间内林分任意位置处的光照值。与林下辐射计测量值以及半球照片计算结果的比较,该模型基本上能够满足对林分光环境了解的要求。 5. 从川西亚高山森林生产力的角度,探讨了森林生产力研究的方法以及川西地区的研究历史和成果,发现了其中的一些规律和问题,特别是在叶面积测量上,还没有使用标准的叶面积指数定义。综合来看,川西地区针叶林叶面积指数(单位土地面积上植物冠层总叶面积的一半) 应在4-5 之间。降雨丰富的华西雨屏带是川西地区森林生产力最高的地区,而向西北森林生产力逐渐降低。川西地区云冷杉林森林生产力平均约为600 gDM m-2 a-1,但是根据辐射能计算的潜在生产力则达到1800 gDM m-2 a-1。实际与潜在森林生产力的巨大差异说明其它因子对生产力的影响。 6. 王朗亚高山3 个典型森林林分中,白桦林样地(BF) 林下草本以糙野青茅、牛至、紫菀等喜阳性物种为主,林下透光度较高;冷杉林样地(FF) 林下透光度最低,以喜阴性物种水金凤、蟹甲草、囊瓣芹等为主;而云杉林样地(SF)林分林龄最大,林下透光度介于冷杉林和白桦林之间,草本层仍然以喜阴性物种东方草莓、紫花碎米芥、酢浆草等为主。冷杉林和云杉林的灌木层也很丰富,卫矛属、五加属、茶藨子属、忍冬属植物很丰富,而在白桦林则以栒摘要子属、榛子属、鹅耳枥属等植物为主。藓类植物在云杉林中最丰富,并且形成毯状层,其它两个林分则很稀少。3 个样地林分结构与林下光环境有很强的相关性,从光环境特征可以在一定程度上推测林分的结构。各样地单纯从乔木层材积推算的NPP 排列顺序为BF>FF>SF,与林下辐射透射率和林分年龄的顺序相同,暗示辐射对群落演替过程的驱动作用。 7. 用半球照相法测得BF、FF 和SF 3 个样地的有效叶面积指数以SF 样地最高,BF 最低。如果考虑针叶树叶片在小枝上的丛聚分布,利用北方针叶林的数值进行校正,则SF 样地LAI 显著增加(达到89%),其它样地的LAI 基本不变甚至有所下降。校正后的数值与文献中地面测量的结果较相近,说明在使用半球照相法测量川西亚高山针叶林LAI 时必须加以校正。 8. 在3 个样地中,白桦、岷江冷杉和方枝柏种群为丛聚分布,紫果云杉在FF和SF 样地中基本上为随机分布。3 个物种出现丛聚分布的最短距离约为2m,在最短距离以内则为随机分布。最短距离可能与树冠大小有关,种子传播特征以及对光照的需求状况可能是造成这种分布格局类型的原因。 Radiative transfer plays a key role in forest ecosystems. Solar radiation providesenergy for photosynthesis, appropriate ambient temperature and development informationfor plants. However, quality and quantity of radiation reaching land surface are affected byweather and subsequently influence the growth and development of plants, which in turnchanges the budget of radiation in forest. Soil temperature changes with the variation ofradiation under forest canopy and influences the activity of roots and rate of nutrientturnover. Thus, any changes of radiation will induce chain reactions in the entireecosystem and display in the value of net primary productivity which will possibly shiftthe relationship between carbon source and sink at local or regional scale and feed back tothe global climate system. On the other hand, as a component of ecosystems, humanbeings of course need to demand more materials and better service from ecosystems. Forthese purpose, man must adapt their pattern and frequency of interference to ecosystems.This paper aims to research on the canopy structure, the radiation distribution and theirinfluence on soil temperature from the process of radiative transfer in subalpine forestecosystem of western Sichuan. The main results are: 1 Present a new photogrammetric method for leaf area. The main idea is to convertnon-vertically taken images of planar leaves to orthoimages through projectivetransformation. The resultant images are used to get leaf morphological parametersthrough image processing. This method enables users to take photos at almost anyorientation and distance if only the leaves are placed on same plane, and to processlarge quantity of leaves in a short time, which is suitable for field measurement. Theresolution of leaf area is adjustable to fit for special requirement. 2 A model using hemispherical photos combining with solar tracks and radiation courseis provided to simulate light variation in forest. The hemispherical photos of canopyrecord the real spatial distribution of each element of plants viewed from a point. Skyradiance is simulated with CIE standard clear sky or cloudy sky model. This modelcan be used to simulate real time light variation under canopy. 3 Present a soil temperature model. Soil could be regarded as a body of resistor andcapacitor. Some of the budget of solar radiation in soil body is transformed into soilpotential energy, the soil temperature. Variation of soil temperature is driven by solarradiation, vegetation, soil properties, etc. This model has two parameters, one of whichis time constant and is related to soil water content. The inversed model can be used tosimulate the variation of soil water. 4 By using model tRAYci developed by Brunner (1998), the 3-D distribution of light inthree subalpine forest stands of Wanglang Nature Reserve has been simulated andvalidated with value of radiometers in these stands. This model can basically satisfythe need for understanding light regimes of these stands. 5 Present some principles and questions of NPP (net primary of productivity) researchesin western Sichuan. The standard leaf area index (LAI) defined by Chen and Black(1997) has not been used in this region. Total leaf area and projected leaf area indexare still used in NPP researches which may differ around 1-fold in magnitude. Thestandard LAI which is a half of total leaf area above unit land area should be between4 and 5 for typical subalpine coniferous forest of western Sichuan concluded fromliteratures. The maximum forest NPP occurs in West China rain belt and decreasesnorthwestwards. Average NPP of spruce-fir forest in western Sichuan is about600gDM m-2 a-1, which is below the potential NPP of 1800gDM m-2 a-1 based onmeasured radiation in this region. The significant difference between potential and realNPP suggests that other factors influence the growth of stands. 6 In the three subalpine forest stands of Wanglang Nature Reserve, herbage layer ofAbstractbirch stand (BF) with age of 40 is dominated by heliophytes of Deyeuxia scabrescens,Origanum vulgare, Aster tongoloa etc.. However, both of the other two stands aredominated by shade tolerent species, such as Impatiens noli-tangere, Impatiensdicentra, Cacalia deltophylla and Pternopetalum tanakae etc. in fir stand (FF) withage of 180 and Fragaria orientalis, Cardamine tangutorum and Oxalis corniculata etc.in spruce stand (SF) with age of 330. Shrub species in the latter two stands arerelatively rich, typical dominant genera being Euonymus, Acanthopanax, Ribes andLonicera. Birch stand has relatively sparse shrubs dominated by genera of Cotoneaster,Corylus and Carpinus. Mosses are significant only in spruce stand. The canopystructure controls the light regime of stand, which influence the composition of herblayers beneath the canopy. This light regime-community structure relationship can beused to infer the herb community from canopy structure. The NPP derived from timbervolume of arbor layer of the three stands decreases from BF to SF, which is in thesame order of transmitted total radiation under canopy and age of these stands,suggesting the driving effect of radiation in the succession of community. 7 The highest effective LAI of the three stands obtained by hemispherical photos is inplot SF and lowest in plot BF. After rectification of the clumping effect of leaves onshoot, the real LAI in plot SF increases significantly (89%) and approximate to theaverage LAI of coniferous forest in western Sichuan. Therefore, the LAI obtainedfrom hemispherical photos needs rectification for clumping effect. 8 Spatial distribution pattern for Betula platyphylla, Abies faxoniana and Sabinasaltuaria is clumpy, but Picea purpurea almost random in plot FF and SF. The shortestdistance for clumpy distribution for Betula platyphylla and Sabina saltuaria is 1.5m,and 2m for Abies faxoniana. And random pattern for these trees is exhibited within thisrange which almost coincides with the diameter of crown. Seed dispersalcharacteristics and light requirement may be the reason for different spatial pattern.
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植物生长和生产力受到自然界各种形式的生物和非生物胁迫因子的影响。这些胁迫包括低温、高温、盐碱、干旱、洪水、重金属、虫害、病害和紫外线辐射等等。而人类活动大大加剧了这些胁迫所带来的影响。由于人类污染而导致臭氧层衰减以及由此产生的地球表面紫外辐射增强已经成为全球气候变化的一个主要方面。UV-B胁迫,甚至当前的辐射水平,所带来的影响已经引起科学工作者的广泛关注。 为了生存和繁殖,植物不得不面临环境中各种潜在胁迫所带来的负面影响。然而,植物生活型的不可移动性决定了其逃避胁迫的局限性。因此,绝大多数植物都是通过对胁迫作出反应,通过修复或者更新组织来降低伤害。而植物应对环境变化的能力则是由其生长模式的种属特异性和本身的遗传组成所决定。在自然界,植物常常同时面临多种胁迫,这些胁迫所引发的植物反应可能具有叠加、协同或者拮抗作用。沙棘是一种具刺、具有固氮功能的多年生雌雄异株灌木,广泛分布于亚欧大陆的温带地区和亚洲亚热带的高海拔地区。在中国,沙棘常常被用作植被恢复中的先锋树种而大量栽培。本文采用沙棘作为模式植物,试图探索木本植物对低温,UV-B辐射增强以及其与干旱的复合胁迫的响应以及沙棘对这些胁迫响应是否具有种群差异性。 对来自南北两个种群的沙棘进行短日照和低温处理,检测了其在抗寒锻炼和抗寒性发育过程中存在的性别差异。结果表明,短日照和低温都分别能够诱导抗寒锻炼的发生,而两者同时存在对所有实验植株抗寒性的大小具有叠加效应。然而,短日照和低温所诱导的抗寒性在两个种群中都具有性别差异性,雄性植株比雌雄植株对短日照和低温更为敏感。同时,南北种群间也存在差异性,北方种群的植物比南方种群的植物对短日照和低温敏感,从而在短日照下抗寒锻炼的发生时间更早,低温诱导的抗寒性更大。短日照和低温诱导植物增加抗寒性的同时伴随着脱落酸的变化。脱落酸的变化因处理,种群和性别的不同而不同。这些生理反应表明不同的沙棘种群,不同的植株性别对同一环境胁迫可能存在不同的生存策略。 比较了来自高低两个海拔的沙棘种群对于干旱和UV-B辐射增强以及两者复合胁迫条件下的生理生态反应。干旱使两个种群中植株总的生物量,总叶面积,比叶面积,叶片含碳量,含磷量,木质素含量和碳氮比显著降低,使根冠比,粗根细根比和叶片脱落酸含量显著增加。干旱而非UV-B使得δ13C 值显著增加。但是,比较而言,来自高海拔的种群对干旱反应更为强烈,而来自低海拔的种群对UV-B更敏感。在UV-B辐射增强的处理下,干旱所诱导的脱落酸的积累被显著抑制。而且我们检测到在一些指标上存在显著的干旱×UV-B交互作用,如两个种群中在总生物量上,低海拔种群中在总叶面积,粗根细根比上,高海拔种群中在比叶面积,δ13C值,木质素含量上都存在明显的交互作用。这些结果表明这两个种群对胁迫具有不同的适应性反应,来自高海拔的种群比来自低海拔的种群更能够抵御干旱和UV-B胁迫。 室外实验表明,UV-B 去除/增补对沙棘高低两个海拔种群的影响都不大。对生物量的积累,植株高度以及一些常见的胁迫反应生理指标比如丙二醛、ABA 和游离脯氨酸都没有显著影响。UV-B 的效应比UV-A 大,植物反应在无UV 和仅有UV-A 的处理间没有什么区别。然而,UV-B 去除的两个处理和UV-B 存在的两个处理间存在显著区别。UV-B 使得两个种群都显著降低了比叶面积(SLA),但却使长期用水效率增加。但UV-B对光合色素和光合系统II 的影响不大。总体看来,来自低海拔的种群对UV-B 更为敏感。 Plant is adversely affected by various abiotic and biotic stress factors. These stressors includelow temperature, heat, salt, drought, flooding, heavy metal toxicity, wounding by herbivores,infecting by pathogenic microorganisms, ultraviolet (UV) radiation and so on. Variousanthropogenic activities have accentuated the existing stress factors. One of the mostimportant aspects of global change is that of stratospheric ozone depletion caused by seriousanthropogenic pollution and the resulting increase in UV radiation reaching the surface of theEarth. Scientists have become concerned about the effects that considerable UV-B stress, evenat current levels. In order to survive and reproduce, plants have to be able to cope with lots of potentiallyharmful stress factors that are almost constantly present in their environment. Most plants’responses under stress are to neutralize the stress, repairing the damage or regrowing newtissue rather than to avoid it due to their sessile life style. The plant defense capacity dependson plant-specific modular growth patterns and genetic make-up that allows for flexibleresponses to changing environments. Plants usually encounter several stresses simultaneouslyunder field conditions, and the stresses may cause a variety of plant responses, which can beadditive, synergistic or antagonistic. Sea buckthorn (Hippophae rhamnoides L.), a thorny nitrogen fixing deciduously perennialshrub, which is widely distributed throughout the temperate zones of Asia and Europe and thesubtropical zones of Asia at high altitudes. It has been widely used in forest restoration as thepioneer species in China. In this paper, we used sea buckthorn as a model, tried to get some understand of how plants fight low temperature, enhanced UV-B radiation level and thatcombination of drought. And also, want to know whether does there exist some populationspecific responses to such stressors. Sexual differences in cold acclimation and freezing tolerance development of two contrastingsea buckthorn (Hippophae rhamnoides L.) ecotypes from northern and southern regions inChina were recorded after exposure to short day photoperiod (SD) and low temperature (LT).The results demonstrated that cold acclimation could be triggered by exposing the plants toSD or LT alone, and that a combination of both treatments had an additive effect on freezingtolerance in all plants tested. However, development of freezing tolerance was dependent onthe sex of plants under SD and LT, the males were clearly more responsive to SD and LT thanthe females in both ecotypes studied. On the other hand, development of freezing tolerancewas also ecotype-dependent, the northern ecotype was more responsive to SD and LT than thesouthern ecotype, resulting in earlier cold acclimation under SD and higher freezing toleranceunder LT. Moreover, development of freezing tolerance induced by SD and LT wasaccompanied by changes in ABA levels. These alterations in ABA levels were different indifferent treatments, ecotypes and sexes. Therefore, the differences in SD and LT-inducedphysiological responses showed that the different ecotypes and the different sexes mightemploy different survival strategies under environmental stress. Two contrasting populations from the low and high altitudinal regions were employed toinvestigate the effects of drought, UV-B and their combination on sea buckthorn. Droughtsignificantly decreased total biomass, total leaf area, specific leaf area,leaf carbon (C),phophous (P), lignin content and the ratio of C: N in both populations, and increasedroot/shoot ratio, fine root/coarse root ratio and abscisic acid content (ABA), in bothpopulations. Drought but not UV-B resulted in significantly greater carbon isotopecomposition (δ13C) values in both populations. However, the high altitudinal population wasmore responsive to drought than the low altitudinal population. The drought-inducedenhancement of ABA in the high altitudinal population was significantly suppressed in thecombination of drought and elevated UV-B. Moreover, significant drought × UV-B interactionwas detected on total biomass in both populations, total leaf area and fine root/coarse root inthe low altitudinal population, specific leaf area, δ13C value and leaf lignin content in the high altitudinal population. These results demonstrated that there were different adaptive responsesbetween two contrasting populations, the high altitudinal population exhibited highertolerance to drought and UV-B than the low altitudinal population. A field experiment was conducted to investigate effects of UV-B exclusion/supplementationon two altitudinal populations of sea buckthorn. UV-B exclusion or supplementation had littleeffects on both populations investigated. For instance, the total biomass, plant height andsome physiological index such as Malondialdehyde (MDA), ABA and free proline were notchanged significantly. The UV-B effects are more significant than that of UV-A, nodifferences were found between treatments of excluded UV and excluded UV-B. However,compared with treatments of UV-B exclusion (including absent of UV-B and all UV band),the present of UV-B (including near ambient environment and enhanced UV-B) significantdecreased specific leaf area, and increased long time water use efficiency as evaluated by δ13Cvalue. UV-B had little effects on photosynthetic pigments and Photosystem II (PSII). The lowaltitude population is more sensitive to UV-B than that of the high altitude population.
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中国沙棘是一种雌雄异株、风媒传粉的灌木或乔木,在中国西南的卧龙自然保护区有广泛的分布。本研究以采集于四川卧龙自然保护区5 个海拔(1800 m、2200 m、2600 m、3000 m、3400 m)梯度的中国沙棘天然群体为材料,以ISSR 和AFLP 标记技术研究其遗传多样性水平及其遗传结构,旨在了解卧龙地区中国沙棘天然群体的遗传多样性水平以及遗传多样性在群体间、群体内以及雌雄亚群体间的分布和特征,为中国沙棘树种的遗传改良及种质资源保存提供遗传研究背景与实验依据。同时探讨ISSR、AFLP 和RAPD三种标记对中国沙棘天然群体的遗传变异水平和群体间遗传结构的评估能力和各自的优缺点。研究得出以下主要结论: 1. ISSR和AFLP分析都表明卧龙自然保护区的中国沙棘群体拥有较高的遗传变异水平(h = 0.249,HT = 0.305)。出现这种结果的主要原因可能与卧龙自然保护区多变的气候条件和生境的异质度大有关。 2. ISSR 和AFLP 都揭示出卧龙自然保护区中国沙棘群体的遗传多样性随着海拔的增加发生显著的变化,表现为中海拔群体(2200 m 和2600 m)比高海拔群体(3000 m 和3400 m)和低海拔群体(1800 m)有更高的遗传多样性的趋势。出现这种趋势的可能解释是低海拔群体处在相对高温和相对干旱的环境,高海拔群体受到低温和紫外线胁迫,而中海拔群体存在中国沙棘生长的适宜环境。 3. ISSR 和AFLP 分析都表明:卧龙自然保护区中国沙棘的遗传结构遵循分布范围广、交配系统以异交为主的木本植物的通常模式,即大多数的遗传变异存在于群体内,只有少部分的遗传变异存在于群体间。 4. 经Mantel 检测表明,卧龙自然保护区中国沙棘群体间的海拔距离和对应遗传距离之间存在显著的正相关关系,即随着垂直海拔距离的增加,群体间的遗传距离也随之增加。Mantel 检测结果以及聚类分析将卧龙自然保护区5 个不同海拔的中国沙棘群体分为低、中、高海拔群体三组的研究结果都表明,海拔很可能是限制群体间基因交流的主要因素。 5. ISSR 分析发现同一海拔的雌雄亚群体首先聚类的研究结果表明,同一海拔的雌雄亚群体在遗传上最相似。方差分析结果表明只有3.8%的总遗传变异存在于雌雄亚群体间,这可能与雌雄植株间的交配和遗传物质的混合有关。 6. ISSR、AFLP 和RAPD 分析都表明卧龙自然保护区不同海拔的中国沙棘天然群体的遗传多样性水平较高。它们的分析结果估算得到的Nei's 平均基因多样度(h)分别为0.249、0.214 和0.170。从该结果可以看出ISSR 和AFLP 比RAPD 检测到更多的遗传多态性,这很可能是不同标记检测的基因组的位点不同所致。 7. 依据对不同标记系统的比较分析,认为ISSR、AFLP 和RAPD 三种分子标记系统都能成功地用于调查卧龙自然保护区不同海拔的中国沙棘群体的遗传变异水平及遗传变异结构,提供关于中国沙棘天然群体多态性水平和遗传变异分布的有用信息。在三者中,AFLP 具有最高效能指数和标记指数,在确定种间分类关系或鉴别个体方面是一种比较理想的标记。 Hippophae rhamnoides subsp. sinensis, a dioecious and deciduous shrub species,occupies a wide range of habitats in the Wolong Nature Reserve, Southwest China. Ourpresent study investigated the pattern of genetic variation and differentiation among fivenatural populations of H. rhamnoides subsp. sinensis, occurring along an altitudinal gradientthat varied from 1,800 to 3,400 m above sea level in the Wolong Natural Reserve, by usingISSR and AFLP markers to guide its genetic improvement and germplasm conservation. And,comparative study of ISSR, AFLP and RAPD was performed to detect their capacity toestimating the level and pattern of genetic variation occurring among the five elevationpopulations of H. rhamnoides subsp. sinensis, and to discuss their application to the study onplant genetics. The results were list following: 1. The ISSR and AFLP analysis conducted for the H. rhamnoides subsp. sinensispopulations located in the Wolong Natural Reserve of China revealed the presence of highlevels of genetic variation (h = 0.249, HT = 0.305). Besides such features as relatively widedistribution, dominantly outcrossing mating system, and effective seed dispersal by small animals and birds, it is sometimes argued that hard climatic conditions and heterogeneous habitats may also contribute to high levels of diversity. 2. Genetic diversity of H. rhamnoides subsp. sinensis populations was found to varysignificantly with changing elevation, showing a trend that mid-elevation populations (2,200m and 2,600 m) were genetically more diverse than both low-elevation (1,800 m) andhigh-elevation populations (3,000 m and 3,400 m). H. rhamnoides subsp. sinensis is thoughtto be stressed by drought and high temperature at low elevations, and by low temperature athigh elevations. The high genetic variability present in the mid-elevation populations of H.rhamnoides subsp. sinensis is assumed to be related to a greater plant density in the middlealtitudinal zone, where favorable ecological conditions permit its continuous distributioncovering the zone from 2,200 m to 2,600 m above sea level. 3. The genetic structure of H. rhamnoides subsp. sinensis revealed by ISSRs andAFLPs followed the general pattern detected in woody species with widespread distributionsand outcrossing mating systems. Such plants possess more genetic diversity withinpopulations and less variation among populations than species with other combinations oftraits. 4. In the present study, Mantel tests showed positive correlations between altitudinaldistances and genetic distances among populations or subpopulations. The observedrelationship between altitude and genetic distances, and the result of the cluster analysisincluding populations or male subpopulations and classifying the groups into three altitudeclusters suggest that altitude is a major factor that restricts gene flow between populationsand subpopulations. 5. The analysis of molecular variance showed that only 3.8% of the variability residedbetween female and male subpopulations. Such a very restricted proportion of the totalmolecular variance between female and male subpopulations is due to common sexuality andmixing of genetic material between females and males. 6. The analysis based on ISSRs, AFLPs and RAPDs all revealed relatively high levelsof genetic variation among different altitudinal populations of H. rhamnoides subsp. sinensisin Wolong Natural Reserve of China. Their estimates of mean Nei’s gene diversity is equal to0.249, 0.214 and 0.170 respectively, suggesting the higher capacity of detecting geneticvariation of ISSR and AFLP than RAPD. It might be ascribed to their distinct sensitivity todifferent type of genetic variation. 7. Based on the coparative study on ISSR, AFLP and RAPD, we drew a conclusion thatthey all successfully reveal some useful information concerning the level and pattern ofgenetic vatiation occurring among different elevation populations of H. rhamnoides subsp.sinensis. AFLP is a ideal tool to taxonomic study and individual identification for theirhighest efficiency index and marker index among the three marker systems.
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近十年,植物群体遗传学的研究飞速发展,然而与海拔相关的植物群体遗传结构和遗传变异研究却相对较少。到目前为止,还不清楚遗传变异与海拔之间是否有一个通用的格局。在山区,各种生态因子,如温度、降水、降雪、紫外线辐射强度以及土壤成分都随海拔梯度急剧变化,造成了即使在一个小的空间区域,植被类型变化显著,这种高山环境的异质性和复杂性为我们研究植物群体遗传结构和分化提供了方便。沙棘(Hippophea)属于胡颓子科(Elaeagnaceae)为多年生落叶灌木或乔木,雌雄异株,天然种群分布极为广泛。中国沙棘(H. rhamnoides subsp. sinensis)是沙棘属植物中分布较广的一个亚种,种内形态变异非常丰富,加之其具有独特的繁育系统和广泛的生态地理分布,是研究沙棘属植物遗传变异和系统分化的理想材料。本文从1,800 m 到3,400 m 分5 个海拔梯度进行取样,用RAPD 和cpSSR 分子标记研究了卧龙自然保护区中国沙棘天然群体的遗传结构和遗传变异。5 个取样群体依次标记为A、B、C、D 和E,它们分别代表分布在海拔1,800,2,200,2,600,3,000 和3,400 m 的5 个天然群体。RAPD实验用11 条寡核苷酸引物,扩增得到151 个重复性好的位点,其中143 个多态位点,多态率达94.7%。在5 个沙棘群体中,总遗传多样性值(HT)为0.289,B群体内的遗传多样性值为0.315,这完全符合沙棘这种多年生、远交的木本植物具有高遗传变异的特性。5 个群体内遗传多样性随海拔升高呈低-高-低变异趋势,在2,200 m海拔处的B群体遗传多样性达最大值0.315,3,400 m海拔处的E群体则表现最小仅0.098。5 个群体间的遗传分化值GST=0.406,也即是说有40.6%的遗传变异存在于群体间,1,800 m海拔处的A群体与其它群体的明显分离是造成群体间遗传分化大的原因。UPGMA聚类图和PCoA散点图进一步确证了5 个群体间的关系和所有个体间的关系。最后,经过Mantel检测,遗传距离与海拔表现了明显的相关性(r = 0.646, P = 0.011)。cpSSR 实验中,经过对24 对cpSSR 通用引物筛选,11 对引物能扩增出特异性条带,只有2 对引物(ccmp2 和ARCP4)呈现多态性。4 个等位基因共组合出4 种单倍型,单倍型Ⅰ出现在A 群体的所有个体和B 群体的8 个个体中,C、D、E 三个群体均不含有,而单倍型Ⅱ出现在C、D、E 三个群体的所有个体及B 群体的18 个个体中,A 群体不含有。另外两种单倍型Ⅲ和Ⅳ为稀有类型,仅B 群体中的4 个个体拥有。这种单倍型分布模式和TFPGA 群体聚类图揭示了,C、D、E 群体可能来源于同一祖先种,而A 群体却是由另一祖先种发展起来的,B 群体则兼具了这两种起源种的信息,这可能是因为在历史上的某一时期,在中国沙棘群体高山分化的过程中,B 群体处某个或者某些个体发生了基因突变,具备了适应高海拔环境的能力,产生了高海拔沙棘群体的祖先种。 In recent ten years, studies about population genetics of plants developed rapidly,whereas their genetic structure and genetic variation along altitudinal gradients have beenstudied relatively little. So far, it is uncleared whether there is a common pattern betweengenetic variation and altitudinal gradients. In the mountain environments, importantecological factors, e.g., temperature, rainfall, snowfall, ultraviolet radiation and soil substratesetc., change rapidly with altitudes, which cause the vegetation distribution varying typically,even on a small spatial scale. The mountain environments, which are heterogeneous andcomplex, facilitate and offer a good opportunity to characterize population genetic structureand population differentiation.The species of the genus Hippophae L. (Elaeagnaceae) are perennial deciduous shrubs ortrees, which are dioecious, wind-pollinated pioneer plants. The natural genus has a widedistribution extending from Northern Europe through Central Europe and Central Asia toChina. According to the latest taxonomy, the genus Hippophae is divided into six species and12 subspecies. The subspecies H. rhamnoides ssp. sinensis shows significant morphologicalvariations, large geographic range and dominantly outcrossing mating system. Thesecharacteristics of the subspecies are favourable to elucidate genetic variation and systemevolution. To estimate genetic variation and genetic structure of H. rhamnoides ssp. sinensisat different altitudes, we surveyed five natural populations in the Wolong Natural Reserve at altitudes ranging from 1,800 to 3,400 m above sea level (a.s.l.) using random amplifiedpolymorphic DNA markers (RAPDs) and cpSSR molecular methods. The five populations A,B, C, D, and E correspond to the altitudes 1,800, 2,200, 2,600, 3,000 and 3,400 m,respectively.Based on 11 decamer primers, a total of 151 reproducible DNA loci were yielded, ofwhich 143 were polymorphic and the percentage of polymorphic loci equaled 94.7%. Amongthe five populations investigated, the total gene diversity (HT) and gene diversity within population B equaled 0.289 and 0.315, respectively, which are modest for a subspecies of H.rhamnoides, which is an outcrossing, long-lived, woody plant. The amount of geneticvariation within populations varied from 0.098 within population E (3,400 m a.s.l.) to 0.315within population B (2,200 m a.s.l.). The coefficient of gene differentiation (GST) amongpopulations equaled 0.406 and revealed that 40.6% of the genetic variance existed amongpopulations and 59.4% within populations. The population A (1,800 m a.s.l.) differed greatlyfrom the other four populations, which contributes to high genetic differentiation. A UPGMAcluster analysis and principal coordinate analyses based on Nei's genetic distances furthercorroborated the relationships among the five populations and all the sampling individuals,respectively. Mantel tests detected a significant correlation between genetic distances andaltitudinal gradients (r = 0.646, P = 0.011).Eleven of the original 24 cpSSR primer pairs tested produced good PCR products, onlytwo (ccmp2 and ARCP4) of which were polymorphic. Four total length variants (alleles) werecombined resulting in 4 haplotypes. The haplotype was present in all individuals of Ⅰpopulation A and 8 individuals of populations B, the other three populations (C, D and Epopulations) did not share. The haplotype was present in all individuals of populations C, D Ⅱand E and 18 individuals of populations B, population A did not share. The other twohaplotypes and were rare haplotypes, which were only shared in 4 individuals of Ⅲ Ⅳpopulation B. The distribution of haplotypes and TFPGA population clustering map showedthat the populations C, D and E might be origined from one ancestor seed and population Amight be from another, whereas population B owned information of the two ancestor seeds. Itwas because that gene mutation within some individual or seed in the location of population Bwas likely to happen in the history of H. rhamnoides, which was the original ancestor of thehigh-altitude populations.
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对白水河自然保护区进行了生物多样性和植物区系调查,根据获得的生物多样性数据和标本的鉴定结果分析,得到结果和推论如下:1.白水河自然保护区生物多样性的垂直分布格局1.1 植物群落α 多样性随海拔梯度的变化乔木种的丰富度及多样性随海拔上升表现出明显的线性下降趋势。而灌木和草本物种丰富度及多样性随海拔上升表现出抛物线式的下降趋势。乔木种从海拔1400m 的15 种至林线时下降为2 种;灌木和草本植物分别从35 和38 种至山顶时下降为5 种和20 种。乔木物种随海拔升高出现明显的物种替代现象,表明海拔梯度包含了多种环境因子的梯度效应,影响着植物群落的分布与结构及物种多样性。1.2 植物群落β 多样性随海拔梯度的变化海拔2200 m 左右是一个明显的生境转折点。海拔2200 米以下相邻群落的相似性( CJ ) 明显大于海拔2200 m 以上的群落,说明海拔2200 m 以下的群落间共有种多,生境差异较小;而海拔2200 以上的群落则相反,相似性较低。低海拔区由于人为干扰较大,所以群落具有较高的物种丰富度,相邻群落之间的物种替换总量(Cody 指数) 较大。海拔2800 米到海拔3200 米之间因杜鹃群落的影响,物种替换总量(Cody 指数) 略有升高。研究β 多样性沿海拔梯度的变化必须考虑到物种丰富度和群落类型的影响,用不同指数从不同角度能更好地理解β 多样性沿环境梯度的变化。2.白水河自然保护区植物区系性质及起源白水河自然保护区自产种子植物计138 科421 属990 种。本文在科、属的水平上对该保护区植物区系特性进行了较深入的统计和分析。统计表明,温带和热带分布型均占有相当比重,但温带分布型稍占优势;热带、亚热带和温带的科、属多,中国特有属也有相当比例,它们是保护区具有特征意义的类群,其中许多属为古老和残遗成分。结论认为白水河保护区植物区系起源古老,较完好的保存了北极-第三纪古植物群。We have collected the specimens and gotten the biodiversity datas of BaiShuihe Nature Reserve in Peng Zhou,analysed the datas,the results as follows:1. Diversity of the plant community along altitudinal gradient1.1 α diversity of the plant communities along altitudinal gradientFrom 1400m to 3900m at Baishuihe Nature Reserve, 52 plots were investigatedwith an interval 100m in altitude; α diversity and β diversity of plant communitiesand their variety along altitudinal gradient were studied. The results showed that indifferent successional layers of trees, richness and diversity decreased linearly withthe increase of altitude. But shrub and herb layers don’t decrease linearly with theincrease of altitude. Tree species decreased from 15 species at 1400m so only 2species at timberline. Shrub and herb species decreased from 35 and 38 species at2000m to 5 and 20 species at 3800m respectively. Tree species are replacedobviously with the increase of altitude; It shows that altitude includes manyenvironmental facts, which infect the distribution, structure and diversity of plant population.1.2 The variety of βdiversity along altitudinal gradient.The entironment changed obviously near 2200m according to our research. Forsimilarity(CJ) between neighboring plots above 2200m is larger than wich below it.It shows that below 2200m,the neighboring plots has more same species,and thehabitats of neighboring plots has more similarity. Above 2200 is the other way round.The plant communites have higher species richness and species turnoverlargestly between neighboring plots because of the disturbance from humanity at lowaltitude. Between 2800m and 3200m species turnover not so obviously because ofmore Rhododendron live there.We should think over species and the types of plant communities effect thevariety of β diversity along altitude gradient. Use more biodiversity indexs and fromseveral aspects to understand the variety of β diversity along altitudinal gradient.2.Origin and characteristic features of Bai Shuihe Nature Reserve990 species of wild seed plants (belonging to 421 generas in 138 families) inthe floristic region of Bai Shuihe Nature Reserve were reported here. The statisticsand comparatively intensive analysis at generic and familiar levelss. Based on thestatistics,the results show that both temperate and tropical distribution types areacounted for considerable proportion of the total,but formal is a little moreimportant than the later. The North Temperate and E.Asia-N.America disjunctedpatterns are more concentrated in this area. These may be considered as thecharacteristic features of Bai Shuihe Nature Reserve flora, while many of them arearchaic and relic elements. According to above data,the floristic region of BaiShuihe Nature Reserve may be considered as a typical region in Chinese flora. Also,the flora of Bai Shuihe Nature Reserve are originated since ancient time as a wellconserved Arctic-Tertiar flora.
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横断山地区是一个十分自然的植物区系地区,在中国植物区系分区中是作为泛北极植物区中国-喜马拉雅亚区中的一个地区,其种子植物区系具有丰富的科、属、种,地理成分复杂,特有现象和替代现象明显。该地区作为植物区系和生物多样性的研究热点地区,长期以来极受中外植物学家关注。横断山脉东缘是中国-喜马拉雅和中国-日本植物区系的交汇过渡区域,北部的岷江流域以及南部的金沙江流域,孕育了该区丰富的物种资源和植被资源。而岷江干热河谷和金沙江干热河谷的相似性和相关性,更为该区的植物区系和生物多样性南北的对比研究提供了有利的条件。 本研究选择的九顶山西坡和龙肘山分别位于横断山区北部和南部,九顶山属岷江流域而龙肘山属金沙江流域。本研究结合植物区系研究和生物多样性研究,对该区的植物资源进行调查。通过样带调查和样线踏查结合,大量详实的野外样方调查和标本采集,进行传统的区系研究和生物多样性研究。研究该区物种多样性的海拔梯度格局及其潜在的影响影子,并利用新的区系评估质量方法对九顶山西坡的植物区系质量进行定量的研究,以期能更为深刻的理解该区的植物资源,为该区的资源保护和利用提供合理可行的建议。主要研究结论如下: 1)九顶山西坡植物区系的性质和特点 经鉴定和统计,九顶山西坡共有1707 种维管植物,分属617 属和140 科,其中种子植物1616 种,分属572 属117 科。就科的分布区成分构成而言,该区系的热带成分与温带成分相当,热带成分略占优势,表明九顶山西坡的植物区系与热带植物区系有较强的联系。但是,在九顶山西坡属的分布区类型所占的比例上,温带成分远远超过了热带成分,本区的种子植物分布表现出明显的温带性质。且温带分布类型的许多物种组成了九顶山西坡植被的建群种和优势种,是本区系最重要的成分,充分体现了本区系的温带性质。 2)九顶山西坡不同植被带的生物多样性海拔梯度格局 基于对土门-断头崖、茶山-九顶山、雁门沟-光光山三条垂直植被样带的调查,我们发现九顶山西坡的生物多样性沿海拔梯度的变化呈现出一定的规律性,不同样带之间有一定差异。就三条样带的物种组成相似性来看,虽然土门-断头崖样带属于涪江水系,而茶山-九顶山样带和雁门沟-光光山样带属于岷江水系,但不同水系对该区物种组成的影响并不明显。三条样带中,草本层物种丰富度均远远大于灌木层和乔木层,而以乔木层物种丰富度最低;α-多样性指数随着海拔梯度的变化在土门-断头崖样带中呈现单一下降趋势,在茶山-九顶山样带表现为双峰模型,而在雁门沟-光光山样带则表现为不显著波动变化;均匀度指数在土门-断头崖样带呈现出单一下降的趋势,在雁门沟-光光山样带表现为凹形曲线,而在茶山-九顶山样带却无明显的变化规律。β-多样性指数在土门-断头崖样带和茶山-九顶山样带呈现出明显的波动状态,植被类型替代现象明显;而在雁门沟-光光山样带却并未有十分显著的转折点,因其水平植被带受到干扰,同海拔替代现象不显著。 3)九顶山西坡维管植物丰富度的海拔梯度格局 我们考察了九顶山西坡和两条垂直样带(土门-断头崖和雁门沟-光光山样带)的不同分类等级(包括科、属、种)和不同生活型物种(乔木、灌木、禾草、蕨类和其它草本)的丰富度沿着海拔梯度的分布。结果发现,物种的丰富度海拔梯度格局具有不同的模式,单一下降和中间膨胀格局依然是其主流。不同生活型的物种具有不同的丰富度格局,但是对于环境需求相似的类型具有较相似的丰富度格局。不同的丰富度格局可能由多因素导致,包括:气候,海拔跨度,面积,人为干扰等等。 4)九顶山西坡区系质量评估 我们尝试使用传统的区系质量评估方法对九顶山西坡的区系质量进行评估,并尝试使用一种新的区系质量评估体系对该区的区系进行评价。在九顶山西坡随着海拔梯度的上升,平均保守性系数在各条植被带中均呈现出逐渐上升的趋势。区系质量指数随着海拔的升高都表现为双峰模型,在植被交错区区系质量指数相对较高,而在海拔的两极,区系质量指数都很低。大部分地区使用新方法计算所得的加权平均保守性系数和区系质量指数都比传统方法计算的平均保收性系数和区系质量指数要高,说明在九顶山西坡的三条样带中,大部分地区都是那些保守性系数较高的物种占据优势,同时也表明九顶山西坡具有很高质量的区系和自然植被。 5)龙肘山种子植物区系的性质和特点 龙肘山种子植物区系的物种较为丰富,共有154 科,544 属,1156 种。科的优势十分明显,单种属和寡种属数量众多,说明本区系植物成分较为复杂、起源古老、物种多样性指数较高。地理成分复杂,分布类型多样,其中热带成分在总数量上高于温带成分,但是许多温带成分的属是该区植被的重要建群类群和优势类群,表现出明显的亚热带性质。 6)龙肘山生物多样性的现状和特点 在海拔梯度上,龙肘山地区无论是科、属、种的数量,还是不同等级分类单元之间的数量比,均呈现先升后降的趋势,并在中海拔地区达到峰值。物种多样性指数从总体上来说变化幅度不大,略有先升后降的趋势,在中海拔梯度物种多样性最高。乔、灌、草三层的多样性指数表现出乔木层<灌木层<草本层的特征;乔木层均匀度的变化很大,而灌木层和草本层均匀度的变化较小;灌木层均匀度的波动又强于草本层。β-多样性指数呈现单峰模式,中海拔地区最高。就龙肘山东、西坡物种多样性相比较而言,两者虽然在数值上交替上升,但是却体现出了较为一致的趋势,但西坡因受到干热河谷气候的影响,其平均气温要高于东坡,导致了东坡植物群落和物种的分布比西坡要低。在区系成分构成上,低山区的相同海拔段,西坡的热带亚热带成分所占的比例要比东坡高,这是因为西坡的平均气温比东坡稍高,导致了热带、亚热带物种分布更多。而随着海拔的上升,东、西两坡的气候、土壤等条件趋于一致,其植物区系成分的构成格局也趋于一致。 The Hengduan Mountain region is a very natural floristic region; it belongs toChina-Himalaya sub-region of Holarctic region in floristic subarea of China. The flora in this areais rich in family, genus and species; has a very complex composition of geographical elements;especially with high richness of endemic species and obvious substitution phenomenon. Thisregion as a hot-spot area of floristic and biodiversity, has fascinated biologists in the world for along time. The eastern range of Hengduan Mountain is the transition zone of China – Himalayaforest sub-region and China-Japan forest sub-region in floristic. The water systems are quitedifferent, Minjiang River in the north and Jishajiang River in the south grow quit different but alsoabundant plant species and vegetation resources. The similarity and correlativity of Minjiang River dry valleys and Jinshajiang River dry valleys have provided advantageous condition tocontrast flora and biodiversity between north and south. In the present study, the Jiuding Mountainlies in the north of Hengduan Mountain and belongs to Minjiang River, and the LongzhouMountain lies in the south of Hengduan Mountain and belongs to Jinshajiang River. In our study, we combined the methods of floristic research and biodiversity investigation toexplore the resources of plant species and vegetations; sampled with transects along the altitudinalgradients and also with transverse straps with similar elevation; collected the vascular plant specimen with sampling plots of ecology. We explored the plant species richness patterns alongaltitudinal gradients and discussed the underlying factors aroused these patterns; and used a novelmethod to assess the quality of Jiuding Mountain’s flora. All for a deeper comprehension of the plant recourses of this region; and provided feasible and reasonable suggestion for the protectionof resources. The results were as follows: 1 The characteristic of the flora of the west slope of Jiuding Mountain We had collected 1707 species of vascular plants belonging to 617 genera in 140 families inthe west slope of Jiuding Mountain,in which included 1616 seed plant species belonging to 572genera and 117 families. As for the composition of the areal types of the Families of seed plants,tropic components and temperate components are well-balanced, and percentage of tropicscomponents is higher than that of temperate ones for a litter bit. This shows the flora in the westslope of Jiuding Mountain has strong relationship with the tropic flora. But for the composition ofthe areal types of genera, temperate components have far exceeded the tropics ones, indicated thewhole flora with a conspicuous temperate character. Temperate components possess maximumproportion in the west slope of Jiuding Mountain, and many of them belong to constructivespecies and dominant species in the vegetation, are most important components in JiudingMountain’s Flora, also have embodied the temperate character of this area sufficiently. 2 Biodiversity patterns along altitudinal gradients in different vegetation transects in the westslope of Jiuding Mountain Based on the investigation of three vegetation transects (including Tumen-Duantouya transect,Chashan-Jiudingshan transect and Yanmengou-Guangguangshan Transect) in the west slope ofJiuding Mountain, we found the change of biodiversity along the altitude gradients displayedcertain regularity, but have differences among different transects. The three transects belong todifferent water systems; the Tumen-Duantouya transect belongs to Fujiang River, and the othertwo belong to Minjiang River. From the similarity of species compositions of different transects,we found different water system didn’t show obvious impact on the species composition. In all thethree transects, the species richness of herb layer was remarkably higher than shrub and tree layer,and the species richness of tree layer was the lowest one. With the increasing of the altitude, theline of α-diversity was monotonically decreasing curve in Tumen-Duantouya transect, andbimodal curve in Chashan-Jiudingshan transect, but in Yanmengou-Guangguangshan transectshowed a wave-like curve although not very obvious. Species evenness showed monotonicallydecreasing trends in Tumen-Duantouya transect, and very low at mid-altitude in Yanmengou-Guangguangshan transect, but in Chashan-Jiudingshan transect changed irregularly. Changes inβ-diversity corresponded with the transition of vegetation in the Tumen-Duantouya transect andChashan-Jiudingshan transect, and the curve of β-diversity along altitude had obvious turningpoint; but in Yanmengou-Guangguangshan transect had no obvious turning point, and thesubstitution phenomenon was not obvious, transverse vegetation straps distributed interlaced. 3 Richness patterns of vascular plant species along altitude in the west slope of Jiuding Mountain Direct gradient analysis and regression methods were used to describe the species richnesspatterns along the altitudinal for Mt. Jiuding, as well as separately for Tumen-Duantouya Transectand Yanmengou-Guangguangshan Transect. Altitudinal gradient of diversity of units at differenttaxonomic level (including Family, Genus and Species) and at different life form (including tree,shrub, pteridophyte, grass and other herb) were tested to find differences among the richnesspattern. We found altitudinal richness also shows different patterns, and both monotonicallydecreasing pattern and hump-shaped pattern can be founded in vascular species richness. Speciesin different life forms show different altitudinal patterns, but those species with similarrequirements to environmental conditions show similar richness patterns along altitudinalgradients. Different richness patterns can be aroused by different climate, different altitudinal span,area factor, anthropogenic factor and so on. 4 Floristic quality assessments in the west slope of Jiuding Mountain We used both the conventional method broadly adopted in the USA and the new one toassess the floristic quality in the west slope of Jiuding Mountain. The Mean Coefficient ofConservatism (MC) had the trend of increment along the altitudinal gradients. The FloristicQuality Index (FQI) was a bimodal curve with increasing of elevation; FQI got maximum valuesin the transition zones of different vegetations in the middle altitude, and had very low values atthe two end of elevation. In most areas of the west slope of the Jiuding Mountain, the resultscalculated using the new methods were higher than those using the conventional method. Thisindicated the dominant species of the communities had very high coefficients of conservatism inmost areas of Jiuding Mountain, and the communities are relatively kept pristine and the habitats very integrative. 5 The characteristic of the flora of Longzhou Mountain The flora of Longzhou Mountain has very abundant in species composition; there are about1156 species of seed plants belonging to 544 genera in 154 families. In which, twelve families with more than 20 species include totally 232 genera and 532 species, and form the majority of itsflora. The origin of its flora is old, monospecific genera and oligotypic genera amounts to 510 innumber, which constitute 93.75% of total number of genera. The geographical components arevarious in Longzhou Mountain, the majority of flora are temperate and pantropic ones. The tropiccomponents overtopped temperate components on genera quantity, but many temperatecomponents belong to constructive species and dominant species in the vegetation, and the wholeflora shows an obvious subtropical character. 6 Current situation and characteristic of biodiversity in Longzhou Mountain With the increasing of altitude, the number of species, genus, family and the ratios ofdifferent taxonomic levels all displayed a trend of descending after rising first, and peaked atmiddle height area. The change of α-diversity was not very acutely, with the trend of descendingafter rising first in some degree, the middle height area had highest α-diversity. As studying thetree layer, shrub layer and herb layer respectively, the Shannon-Wiener index was in followingorder: tree layer < shrub layer < herb layer; the change of evenness was more complicatedly thanthat of diversity, the tree layer changed acutely, but the shrub layer and herb layer fluctuatedsmoothly. Changes in β-diversity also showed the trend of descending after rising first. TheJaccard index and Cody index all peaked at the middle height forest area. As for the comparison ofplant diversity and evenness between the west and east slope, the numerical values ascendedalternatively, but the trend of changing was similar. The distribution of similar plant communitiesand species in east slope were lower than the west slope for the influence of Jinsha River DryValley. As for the composition of different floristic components, in lower altitude area of westslope, the tropic and sub-tropic plants had higher ratio than east slope’s and even could be equal tothe temperate plants. With the increasing of elevation, the floristic composition become morelikely between the east and west slope and temperate plants dominated the flora.
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常绿阔叶林以其富饶的生物资源、丰富的生物多样性和巨大的生态与环境效益引起了人们越来越大的重视,它的研究已成为国际植被科学界关注的主题之一。我国分布着世界上面积最大的亚热带常绿阔叶林,在世界植被中具有重要地位,它的分布表现出明显的地带性差异,存在着多样的植物群系及其对应的气候特征。但是在植物功能性状领域,与全球范围其它生物群系相比,常绿阔叶林物种的研究较少,其功能性状间、功能性状与环境间的关系尚不清晰。 本研究以常绿阔叶林木本植物的当年生小枝为对象,试图从小枝水平上的生物量分配格局、叶片大小与数量的权衡关系、小枝茎的构型效应、叶片元素化学计量学,以及小枝大小的成本与效益分析等方面,较为系统地揭示小枝水平上的植物功能性状间及其与气候间的关系。因此,在华西雨屏带内部的不同纬度设置峨眉-青城-雷波-平武的温度梯度进行比较,并对有降水差异的川西南偏湿性(雷波)与偏干性常绿阔叶林(西昌)进行对比研究,同时在不同山体进行不同海拔梯度的比较研究。 本文主要研究结果如下: (1)小枝生物量分配格局叶水平上,叶片重-叶柄重(Y轴vs.X轴,下同)呈斜率小于1的异速生长关系,表明叶柄对叶内部的生物量分配影响显著。小枝水平上,叶和茎的生物量以及它们与小枝总生物量间基本呈等速生长关系,表明大的小枝或大叶物种不一定在叶生物量的分配上占优势。不同生活型间,在小枝或者茎的生物量一定时,常绿物种叶片的生物量比例较落叶物种稍高。与温度和水分较优越(峨眉及其低海拔)的生境相比,在相对低湿(螺髻)与低温(平武)的生境中的植物会减少对叶的投入而增加对支撑部分的投资比例。 (2)小枝叶片大小与数量的权衡无论是不同气候带还是不同生活型以及不同海拔梯度,叶片大小与出叶强度基本都是呈负的等速生长关系,表明了叶片大小-数量在小枝水平上的权衡。在不同气候梯度的对比中,叶片数量(出叶强度)一定时,高温和高水分生境(峨眉)比低温(平武)和低湿(螺髻山)生境中的物种的叶片大小(质量和面积)更大,表明不同生境的比较中,小的叶片可能具有较高的出叶强度和更高的适合度收益。“出叶强度优势”(Leafingintensitypremium)假说可能不适宜解释不同生境物种叶片大小差异。 (3)小枝茎的构型效应虽然茎长和茎径与叶片大小都呈正相关关系,与出叶强度都呈负相关关系,但茎长/茎径比与叶/茎生物量之比呈负相关关系;与叶片的大小呈负相关关系,与出叶强度呈正相关关系。这说明小枝构型能影响小枝叶/茎生物量分配和叶大小-数量的权衡关系。其影响机制可能是小枝内部的顶端优势。另外,茎长/茎径比在低湿和低温等不利生境中的植物中较高,而在降水和温度较适宜环境中较低。 (4)叶片C、N、P化学计量学N含量和P含量,C/N比和比叶重(LMA,leafmassperarea)呈正的等速生长关系,而N和LMA,P和LMA呈负的等速生长关系。在LMA一定时,C/N比随着生境胁迫压力的增加而降低,N、P含量随着生境压力的增加而增加。在P含量一定时,N含量随着生境压力的增加而降低,即N/P比在生境条件较优(峨眉及其低海拔)时较高。常绿和落叶植物叶片的N/P比没有差异,在LMA一定时,常绿植物的N、P含量较高、C/N比较低。总之,植物的C、N、P化学计量学特征受叶片属性如LMA与气候,及其相互作用的影响。 (5)小枝大小的代价与效益分析、TLA与小枝总重总叶面积(TLA,totalleafarea,Y轴,下同)与总叶重(X轴)均呈斜率小于1的异速生长关系,TLA与小枝横切面积呈斜率为1的等速生长关系。表明叶片面积的增加总是小于叶重和小枝总重的增加,随着小枝的增大,它的叶面积支撑效率下降。在热量和降水优越的生境(峨眉及其低海拔)中,相同小枝重或者相同茎横切面积的小枝,其叶面积支撑效率较低湿与低温环境下(螺髻山、平武及高海拔)的高。 总体上,本文初步研究了小枝水平上可能存在的以下三种权衡关系:叶-茎生物量分配权衡;叶片大小-数量的权衡;小枝茎长-茎径的权衡关系,以及气候要素等对这三种权衡关系的影响。在此基础上,我们还讨论了这些权衡关系的可能形成机制,及其与物种生态适应的联系。本研究丰富了生活史对策中关于权衡关系的研究内容,为我国常绿阔叶林功能生态学研究积累了材料。 Evergreen broad-leaved forests are attracting much more attention from vegetation ecologists than ever before because of their abundant nature resource and biological diversity, and also great ecological benefits. China has the largest distribution of subtropical evergreen broad-leaved forests (temperate rainforests) that are typical and representative in the world. The forests span over more than ten degrees in latitude and more than 30 degrees in longitude, providing an ideal place to study plant functional ecology, i.e., the climatic effect on plant functional traits and the relationship between the traits. However, relative to the other biomes, there are few studies addressing functional ecology of the plant species from subtropical evergreen broad-leaved forests. In this study, I focused on the leaf size-twig size spectrum of the woody species of subtropical evergreen broad-leaved forests in southwestern china. I collected data on leaf size and number, twig size in terms of both mass and volume, and stem architecture from five temperate mountains, and then I analyzed the relationships between leaf and stem biomass and between leaf size and number, the effect of stem length/diameter ratio on biomass allocation and on the relationship between leaf size and number, leaf C:N:P stoichiometry, and the twig efficiency of supporting leaf area in relation to twig size. I also addressed the climate effect on the spectrum. The temperature gradient from warm to cool sites was represented by Emei Mountain, Qingchengshan, Leibo, and Pingwu, and the rainfall gradient was assumed to emerge from the comparison between Leibo (High) and Luojishan (Low). In addition, altitudinal effects were analyzed with comparisons between low and high altitudes for each mountains. My main results are as follows. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to leaves or laminas was independent of twig mass. Petiole mass disproportionably increase with respect to lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. In addition, the investigated species tended to have a larger leaf and lamina mass, but a smaller stem mass at a given twig mass at favorable environments including warm and humid sites or at low altitude than unfavorable habitats, which might be due to the large requirements in physical support and transporting safety for the species living at unfavorable conditions. Moreover, the evergreen species invested more in leaves and laminas than the deciduous at given stem or twig biomass within any specified habitats. Negative, isometric scaling relationships between leaf number and size broadly existed in the species regardless of climate, altitude, and life forms, suggesting a leaf size/number trade-off within twigs. Along the climatic gradients, at given leaf number or leafing intensity, the leaves were larger in the favorable environments than the poor habitats. This suggested that the fitness benefit gained by small leaves could be larger than that with high leafing intensity in the stressful sites. I concluded that the “leafing intensity premium” hypothesis was not appropriate to interpreting between-habitat variation in leaf size. Both stem length and diameter were positively correlated to leaf size but negatively correlated to leafing intensity. The ratio of stem length to diameter was negatively correlated to leaf mass fraction, and it was negatively correlated to leaf size but positively correlated to leafing intensity. This suggested that the stem architecture influenced twig biomass allocation and the relationship between leaf size and number. The mechanism underlying the architectural effect might lie in the apical dominance within twig. Moreover, the ratio was greater in unfavorable habitats but smaller in favorable environments. Positive, isometric relationships were found between N and P contents per leaf mass, and between C/N ratio and leaf mass per area (LMA), but N and P contents scaled negatively to LMA. C/N ratio decreased but N and P increased with increasing habitat stress at a given LMA. N content declined with increasing habitat stress at given P content. These indicated that N/P and C/N were higher but LMA was lower in favorable habitats than in the other circumstances. The evergreen and deciduous species were non-heterogeneous in N/P, but the evergreen species have higher N and P contents and lower C/N than the deciduous ones. In general, C:N:P stoichiometry were related to both climatic conditions and other important functional traits like LMA. Total leaf area (TLA) allometricly scaled to leaf mass with a slope shallower than 1, similar to the relationship between TLA and total twig mass (leaf mass plus stem mass), suggesting that TLA failed to keep pace with the increase of leaf mass and twig size. However, TLA scaled isometricly to twig cross-sectional area. Thus, it could be inferred that the twig efficiency of displaying leaf area decreased with increasing twig size. In addition, the efficiency at a given twig size was large in favorable than unfavorable habitats. In general, in this preliminary study, I studied three tradeoff relationships within twigs, i.e., between leaf and stem biomass, between leaf number and size, and between stem length and diameter, as well as the climatic effect on the relationships. I discussed the mechanisms underlying the tradeoff relationships in view of biophysics and eco-physiology of plants. I believe that this study can serve as important materials advancing plant functional ecology of subtropical forest and that it will improve the understanding of life history strategies of plants from this particular biome.