933 resultados para SUCCESSION
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以六盘山森林植被为研究对象,通过20多年的森林土壤种子库变化与植被演替过程的试验研究,分析了8种森林群落类型的不同生长年限、生长坡位、枯枝落叶层和土壤深度对土壤种子库形成过程的影响。结果表明:六盘山森林群落不同生长坡位,土壤种子库的储量坡中部>坡下部>坡上部;不同层次土壤种子库,枯枝落叶层远高于0-15 cm深土层,8种森林群落类型排序为华北落叶松林>油松林>华山松林>辽东栎林>山杨林>白桦林>灌丛>草地;土壤种子库储量高峰期,不同群落有显著差异,华山松林和油松林在林龄的30~40年,华北落叶松林在20年,辽东栎林、山杨林、白桦林在15~20年,灌丛和草地在10~20年,其森林群落生长年限与土壤种子库储量变化趋势呈拟合曲线,符合指数方程,相关性极为显著;土壤种子库物种组成丰富,草本和灌木植物远高于乔木树种,乔木树种仅有3~5种,但多数为外来入侵种,而在每一类型中出现频率最高的草地植物多为蒿类,灌木植物为柔毛绣线菊和沙棘,乔木为辽东栎树种。因此,在六盘山林区植被自然更新与合理演替的驱动种和先锋种草地植物为蒿类,灌木植物为柔毛绣线菊和沙棘,乔木为辽东栎,其次是华北落叶松、油松和华山松。
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为了解黄土高原沟壑区小麦品种演替过程中的籽粒灌浆特性,以此地区20世纪50年代至今的主栽小麦品种为供试材料,研究了不同小麦品种演替过程中籽粒干物质积累量及灌浆速率的变化。结果表明,黄土高原沟壑区小麦演替过程中,小麦籽粒千粒重呈现逐渐增加的趋势。平均灌浆速率逐渐增加,小麦籽粒的干物质积累随之增加,长旱58的W0最大,达50.53 g。灌浆三阶段中,各阶段的灌浆速率表现为,V2>V1>V3。从灌浆持续时间看,T1和T2持续的时间较长且变异系数较大,而T3和T相对稳定。从灌浆速率来看,V1、V2、V3、Vm、Va变异系数较大,灌浆速率易受环境因素影响而波动。
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用4年长期定位试验资料,利用植物系数、蒸散量、土壤含水量和土壤水分对植物的有效性等指标,研究了黄土高原植被群落不同演替阶段(草本群落→灌木群落→早期森林群落→顶级群落)的耗水特性与生态适应性。结果表明:不同演替阶段,群落实际蒸散量主要受降水控制,群落间差异不显著(P>0.05);土壤含水量是早期森林群落明显高于其它群落,草本群落明显高于灌木群落(P<0.05);植物系数是灌木群落>草本群落>乔木群落,而顶级群落大于早期森林群落;土壤水分对植物的有效性是早期森林和顶级群落明显高于草本和灌木群落(P<0.05)。因此,进行植被建设不但要考虑植物系数还要考虑土壤水分对不同植物的有效性。
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采用样方法对云雾山干草原区本氏针茅草地群落进行长期定位监测,对未封育样地和封育5,10,15,20和25年样地群落的平均高度、盖度、多度和地上生物量以及植物多样性进行实地调查分析,以研究围栏封育对草地植物群落特征动态变化的影响。结果表明,围封10年样地群落的高度、多度以及地上生物量最大,而群落的盖度则随封育年限的增加呈显著增加趋势,在封育25年达到最大;围栏封育的不同年限也显著影响了群落的物种多样性,在围封10年样地群落具有最高的多样性指数,围封15年样地具有最高的丰富度指数,而均匀度指数则是在未封育群落中表现为最高值,最低值是在物种丰富度最高时出现。由此可见,围栏封育在一定时间范围内可以显著改善群落的特征,增加群落的生产力,对于本研究地本氏针茅群落的最佳封育期限为10~15年,但围封时间过长在草地退化方面有一定程度的影响。
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研究黄土丘陵区植被次生演替灌木群落形成过程中土壤养分的变化情况。【方法】以白羊草群落为对照,选取杠柳、狼牙刺、丁香、虎榛子和柠条5种乡土树种为研究对象,测定其土壤养分含量,计算灌木群落形成变化过程中的土壤养分指数(Soil Nutrient Index,SNI),分析了各养分因子间的线性相关关系。【结果】不同灌木初期对土壤养分的适应能力大小顺序为:柠条(0.072)>狼牙刺(0.114)>杠柳(0.163)>丁香(0.172)>虎榛子(0.292)。不同植被物种群落土壤SNI值大小顺序为:虎榛子群落(0.979)>狼牙刺群落(0.535)>柠条群落(0.457)>丁香群落(0.341)>杠柳群落(0.333)>白羊草群落(0.145)。从不同灌木群落形成过程所产生的土壤养分改善效应来看,各参试群落相对于白羊草群落土壤养分的增幅分别为:虎榛子(4.740倍)、狼牙刺(2.909倍)、柠条(2.661倍)、杠柳(1.169倍)和丁香(1.168倍)。各土壤养分因子间除土壤全磷与有效磷和有效钾的相关性不显著外,其余各土壤养分因子间均表现出极显著的线性相关关系(P<0.01)。【结论】不同灌木种群所需的适宜土...
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研究了黄土丘陵区植被次生演替草灌过渡灌木初期的土壤物理性质特征。结果表明:当植被次生演替进展到草灌过渡灌木初期,除土壤含水量外,土壤容重、孔隙度、稳定入渗和水稳性团聚体都好于对照白羊草地,表明灌木树种定居的土壤物理条件一般要高于草本群落。植被演替的过程是植被与土壤相互作用复杂变化的过程,它们的相互反馈也是生态恢复的重要标志。随着植物群落的变化土壤也发生着不同程度变化。其中,土壤物理性质各个因子之间的变化是相互影响和相互联系的。土壤稳定入渗速率与土壤容重、土壤总孔隙度、土壤毛管孔隙度和土壤团聚体之间有显著的相关关系;土壤毛管孔隙度与土壤非毛管孔隙度线性显著负相关;土壤容重与土壤总孔隙度极显著负相关。
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川西亚高山针叶林是四川森林的主体,是长江上游重要的生态屏障。云杉作为亚高山针叶林人工更新的主要树种,已经在该地区形成了大面积的人工纯林。目前,许多云杉人工林分已经进入更新成熟龄,而这些人工林的持续更新却成为日益凸现的问题。探讨这些云杉人工林的自我更新潜力及云杉种子种群更新特点,可为培育后续森林资源提供科学依据。 本文以川西米亚罗亚高山60a云杉人工林为研究对象,并以该区域内相对稳定的植被群落——天然林为对照,采用种子收集器、土壤种子库筛选、室内外种子萌发实验及野外幼苗调查等方法,从异质性微生境的角度研究了种子雨下落之后,不同微生境对种子库、种子萌发、幼苗建立及分布这一前期更新过程的影响,得出如下结果: 1、通过对川西亚高山60a云杉人工林和天然林6年内种子雨雨量、形态特征、散步动态等的持续观测和综合比较可以发现,云杉林结实特点由于林木自身的特征存在着巨大的变动,2002年和2006年两个种子结实大年内,60a人工林种子雨强度分别达到1088.2 ± 52.3粒/m2和704.3 ± 48.9粒/m2,远大于天然林579.9 ± 28.9粒/m2 和507.5± 30.7粒/m2;且云杉林结实质量优于天然林。60a人工林结实量大,种子质量也最好,相对天然林来说对种群的天然更新以及群落的演替都有最大的贡献潜力。应该说,在川西亚高山云杉人工林的天然更新过程中,种源不是影响天然更新的因素。在种子结实大年里,达到更新成熟的云杉人工林有着优于该地区相对稳定植被群落——天然林的种源优势。至少在种子结实大年,种子供应不是该区域人工林天然更新的限制因子。 2、相对于天然林种子库,人工林种子库在种子萌发前能够有较多的有活力种子。虽然这其中有种子雨输入量有差别的因素存在,但两种林分种子库对种子的保存率的不同才是造成这种差异的主要因素。在人工林中,不同地被类型形成的微生境显著地影响了种子库中种子的密度、垂直分布。有地被物存在的微生境能够将种子雨的大部分截留在地被层中,成为幼苗出现的主要场所;同时不同的地被物对种子的保存情况存在显著的差异,苔藓和凋落物层能都较好地保持其中的种子,到种子萌发前,这两种种子库类型能分别为天然更新提供366.1粒/m2和302粒/m2的有效种子。从这点来看,林下地被物上的种子库能够为天然更新萌发阶段提供数量可观的物质基础。 3、种子的萌发和幼苗的定居是天然更新过程中种子库向幼苗库转化的关键环节。总的说来,米亚罗人工林区60a云杉人工林种子向幼苗的转化率十分低下,凋落物、苔藓、草本、裸地四种主要地被物以及天然林内种子/幼苗的转化率分别仅为2.22%、2.14%、0.57%、0.67%、1.05%。这种低的转化率成为云杉林天然更新的限制性因子。但在现有更新条件下,微生境对这一环节仍然显示出十分显著的影响,表现为凋落物和苔藓对现有更新的新幼苗的保存率高于其它类型及天然林。苔藓和凋落物在种子萌发,幼苗保存,以及幼苗分布上都要优于其它地被物类型;另外,微地形对天然更新过程的影响也很显著,凹立地上更适宜于种子的汇集、萌发和定居。 Subalpine coniferous forests dominate most parts of the forested areas in western Sichuan, and they are important ecological barriers in the upper reaches of the Yangtze River. Picea asperata is one of the keystone spruce species for re-afforestation after felling of the natural forests and there have been a total of ca. 13 000 ha of plantations dominate by this species established. Nowadays, many P. asperata plantations have reached reproductive maturity. However, continued regeneration becomes to an important problem in these plantations. Understanding their self-regeneration potential and the regeneration characteristics of seed populations in spruce plantations of these plantations can have some insights on the management of these plantations and the establishment of following forest resources. A subalpine 60a P. asperata plantation distributed in Miyaluo artificial forest area was studied in this paper, at the same time. Synchronously, a 150a natural spruce forest was studied as comparison. Using seed collecting traps, sieving method for soil seed bank, seed germination experiments and seedling investigations in the field, the effects of heterogeneous microsites on early natural regeneration processes after seed rain were studied, which included seed banks, seed germination, seedling establishment and distribution. The main results are as follows. 1. Through a 6-year long term investigation of seed rain intensities, characteristic, dispersal dynamics of 60a P. asperata plantation, we could concluded that the seed setting properties of 60a P. asperata plantation have a great variation for the characteristics trees. In the mast seed year of 2002 and 2006, the seed rain intensities of plantation was 1088.2 ± 52.3 seeds/m2 and 704.3 ± 48.9 seeds/m2 respectively, which were much greater than that of natural spruce forest (579.9 ± 28.9 seeds/m2 in 2002, and 507.5± 30.7 seeds/m2 in 2006). Furthermore, the quality of seed rain in P. asperata plantation was better than that of natural spruce forest. Contrasting with natural spruce forest, 60a P. asperata plantation has a greater potential on natural regeneration of P. asperata population and succession of community for the reason of greater seed rain intensities and better seed quality. We can confirm that seed source was not a limiting factor which influences the natural regeneration progress of P. asperata population distributed in subalpine mountain zone, at least in the mast seed year. 2. Contrasting with natural spruce forest, P. asperata population had more viable seeds in seed bank before the germination. Although the difference of seed rain intensities of two forests has effect on this phenomenon, the difference of seed conservation ability in two forests was the main factor. In the P. asperata plantation, the seed densities and seed vertical distribution pattern were significant effected by the microsites, which posed by different ground cover types. In other word, Microsite with ground covers can obstruct most seeds and keep them in ground cover layer from seed rain, and these ground covers would be the main site for seed occurrence. There was a significant difference about seeds conservation ability among these ground covers. Litter and moss could better conserve P. asperata seeds which distributed in this two covers. Seed banks exist in litter and moss ground cover types could respectively provide 302seed/m2 and 366.1seed/m2 for natural regeneration before the seed germination. From this point of view, we could conclude that ground covers can ensure considerable numbers of seeds for the germination process. 3. Seed germination and seed establishment are key steps that the seeds invert to seedlings in natural generation process. In sum, the seed/seeding transform rate in 60a P. asperata plantation distributed in Miyaluo artificial forest area is very low. the seed/seeding transform rates in litter, moss, herb, soil surface and natural spruce forest were 2.22%、2.14%、0.57%、0.67%、1.05%, respectively. The low transform rate become to a limiting factor of P. asperata natural regeneration process. However, under the existing conditions of natural regeneration, microsit still had significant effect on this transform. The states of Seed germination, new seedling conservation, and older seedling distribution in litter and moss were better than in any other ground cover type or natural spruce forest. In addition, the micro-relief has significant effect on natural regeneration process. Concave site was more suitable for seed collection, seed germination and seedling distribution.
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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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大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库,约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦(Betula albosinensis Burk.)是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物,采用人工模拟的方法,研究CO2浓度升高对不同种内竞争强度(种群水平)下红桦幼苗的生理特征、生长、干物质积累及其分配的影响,探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征,为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下: 1)在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后,叶片出现“光合适应”现象。与对照相比,低种植密度(28株m-2)和高种植密度(84株m-2)条件下的红桦幼苗净光合速率(A)、气孔导度(gs)、蒸腾速率(E)、表观量子产量(AQY)和羧化速率(CE)显著降低,而水分利用效率(WUE)则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物(TNC)浓度,结果是红桦幼苗的比叶面积(SLA,cm2 g-1)显著降低。 2)与对照相比,CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率(R GR)显著提高,尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积,结果是单株红桦幼苗的总叶面积比对照显著增加。 3)CO2浓度升高处理显著增加红桦幼苗干物质积累(尤其是细根生物量),改变了红桦幼苗生物量的分配格局。与对照相比,CO2浓度升高处理的红桦幼苗叶重比(LWR)、叶面积比(LAR)、叶根重比(Wl/Wr)和源汇重比(leaf weight to non-leaf weight ratio, Wsource/Wsink)显著下降(高种植密度的LWR除外),而根冠比(R/S)则显著增加。在两种种植密度条件下,CO2浓度升高显著增加红桦幼苗根生物量的分配比率,显著降低叶片的生物量分配比率,对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之,长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降,并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照,而后期迅速下降,说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比,表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration([CO2])has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.
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当前大气CO2浓度升高是全球变化的主要趋势之一,CO2浓度升高还会引起全球变暖等其它环境问题,因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种,在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验,研究了CO2浓度倍增环境下,不同密度水平红桦碳氮固定、分配可能发生的改变,并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应,为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下: 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长,同时也改变生物量在体内的分配格局,主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异,它们对CO2浓度升高的反应程度不同,但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应,也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度,显著增加红桦幼苗的群体生物量,从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度,对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加,但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理,这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明,红桦在升高CO2浓度的条件下,会作出积极的响应,从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高,植株各器官N、P含量降低,但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高,N、P向主茎和根的分配增加,向叶片的分配减少,主要是由于前者在总生物量中的比重增加,而后者减少了。(3) CO2浓度升高,氮磷利用效率(NUE和PUE)提高,氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后,亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加,后期积累速率下降。(3) 升高CO2浓度以后,土壤呼吸显著增强;土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的;所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期,随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势,同时MDA(丙二醛)含量下降,保护酶SOD(超氧化物岐化酶)、CAT(过氧化氢酶)活性升高。由此说明,升高CO2浓度有利于减缓生长季节后期叶片的衰老,使叶片维持较高的光合速率,也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.
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在青藏高原东部的亚高山针叶林区,如何尽快恢复这一生态脆弱地区的植被,改变生态环境恶化的趋势,是一个十分重要的课题。光一直被认为是植物种间相互替代,尤其是森林演替过程中植物相互替代或植被恢复中的关键环境要素之一。植物能否适应林冠下或林窗中异质的、或多变的光照条件,对其在林中的生存、分布、更新以及森林动态都是非常重要的。 本文以青藏高原东部亚高山针叶林的主要森林类型——岷江冷杉林群落的几种树苗为研究对象,采用实验生态学、生理及生物化学等方法,通过模拟针叶林不同大小林窗内光照强度的变化,在中国科学院茂县生态站内采用遮荫处理设置6个光照梯度(100、55、40、25、15与7%全光照),来研究具有不同喜光特性的植物对光强的响应与适应机制,其研究结果可为揭示亚高山针叶林的演替规律、以及人工林下幼苗的存活与定居提供科学依据,也能为苗木的生产与管理提供科学指导,尤其是对针阔树种在不同光强下的响应与适应的比较研究,能为如何将阔叶树种整合到人工针叶林中提供新的思路。 光强对植物生长的影响 光强对植物的生长具有重要作用,不同植物在各自适宜的光强梯度下才能生长良好。通过一个野外盆栽实验,来研究不同光强对植物生长的影响(第三章)。主要研究结果如下,低光强下植物株高/茎生物量增加,说明植物会将生物量更多用于高生长,以便有效地拦截光资源;在强光下,植物将生物量更多地向根部分配,使得植物在强光下能够吸收更多的水分,而避免干旱胁迫。 在第一个生长季节,以相对生长速率(RGR)表示,红桦和青榨槭在100%全光照下RGR最大,粗枝云杉在55%最大,岷江冷杉在25-40%下较好;然而,在第二个生长季节,2种阔叶树的相对生长速率(RGR)的适宜光强则变为25-55%,云杉为55-100%,而冷杉为25-100%。可见,从第一年到第二年,2种阔叶树苗更适宜在部分荫蔽的条件下生长;而2种针叶树苗对光的需求则逐渐增加,这可能是增加对根生物量相对投资的结果,因为以这种方式,强光下生长的针叶树幼苗更能保持其内部水分平衡,其生长不会因干旱胁迫而受到严重影响。另外,严重遮荫会引起冷杉幼苗死亡。 植物对光强的生理适应 植物可以通过自身形态和生理特征的调整,来发展不同的光能利用策略从而能够在林中共存。通过一个野外盆栽实验,研究了不同光强下生长的几种树苗的生理特征(第四章)对不同光强的响应与适应。结果显示:强光下,粗枝云杉和红桦的光合能力增加,而岷江冷杉和青榨槭在中度遮荫(25-55%)的条件下光合能力最大。植物叶氮和叶绿素含量增高,而光补偿点和暗呼吸速率降低,这些都是植物对低光环境的适应性反应;而强光下植物叶片和栅栏组织变厚,是对强光的一种保护性反应。 植物对光的可塑性反应 不同植物会表现出对光适应有利的生理和形态可塑性反应。本文对第三章、第四章的实验数据进行可塑性指数分析,来研究植物对光强的表型可塑性反应(第五章)。结果显示,生理特征调整是植物对不同光环境的主要适应途径。红桦和青榨槭的可塑性指数平均值要大于粗枝云杉和岷江冷杉,充分表明这2种阔叶树在生理和形态上较强的可塑性更有利于对光环境的适应,而具有比耐荫树种更强的适应能力。另外,2种针叶树相比,云杉的适应性更强。本研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。 植物的光抑制与防御 当植物叶片吸收了过多光能,会发生光抑制现象。植物对光抑制的敏感性及防御能力对其生长具有重要意义。本文通过两个野外盆栽实验,研究了生长在强光下(第六章)和变化光强下(第八章)植物的光抑制现象及其防御策略。结果表明,在强光下或从遮荫状态转入强光下,植物都会发生光抑制,其对光抑制的敏感性与植物的耐荫性(或喜光)和演替状态有密切联系。长期生长在强光下的植物受到光抑制是可恢复的,而当处于荫蔽环境的植物突然暴露于强光下时,受到的光抑制不能完全恢复,可能是(部分)光合机构受到破坏的缘故。粗枝云杉和青榨槭防御光抑制伤害的能力较强,热耗散是其防御光抑制的主要途径。长期的强光作用能使岷江冷杉和红桦发生严重光抑制,甚至光伤害,而红桦能够通过“凋落老叶,萌发新叶”的途径来适应新的强光环境。 How to restore the vegetation of subalpine coniferous forest in eastern Qinghai-Tibet Plateau, and change the trend of ecological deterioration is a very important issue. Acclimation of tree seedlings to different and varing light environment affects to a great extent the successful regeneration and establishment of subalpine coniferous forests in southwestern China’s montane forest areas, because the ability to respond to such changing resource are commonly assumed to be critical to plant success, and have a growth advantage than others. In this paper, several species seedlings in Abies faxoniana community were chosed to study the response and adaptation to light intensity and the interspecific differences of adaptability in six shaded sheds (100, 55, 40, 25, 15 and 7% of full sunlight) in the Maoxian Ecological Station of Chinese Academy of Sciences. Our results could provide a strong theoretical evidence for understanding the forest succession laws of subalpine coniferous forests, and the survival and settlement of seedlings under plantations, and provide scientific direction for the production and management of seedlings, especially the comparative studies of the acclimation to light between the conifer and broadleaf trees could provide new ideas for how to integrate the broad-leaved trees into the artificial coniferous forest. Growth under different light intensity Light intensity plays an important role on plant growth. One field experiments was conducted to study the growth of tree seedlings of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii under different light intensities. The results showed that plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources, while biomass greater allocation to the roots, could make plants under high light environment absorb more water, and avoid drought stress. During the first growing season, the relative growth rates (RGRs) of Betula albo-sinensis and Acer davidii had the greatest values under the 100% of full light, for 55% of Picea asperata, and for 25-40% of Abies faxoniana. However, in the second growing season the the relative growth rates of the two broad-leaved trees changed and were appropriate for 25-55% of full light, for 55-100% of spruce, and for 25-100% of fir. Thus, from the first year to the second year, two broad-leaved seedlings maybe more suitable to partly shading environment, and two coniferous seedlings would have an increase in light demand, which may be an increased root biomass investment. Because in this way, seedlings grown under high light could better maintain their internal water balance, and thus its growth would not be seriously affected by drought stress. In addition, serious shading would cause fir seedlings to die. Acclimation of physiology to light Plants could coexist in forest ecosystem by forming different strategies of light use. One field experiments was conducted to study the acclimation of tree seedlings to different light intensity of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii. The results showed that the photosynthetic capacity of Picea asperata and Betula albo-sinensis exhibited a general tendency of increase with more light availability; but for Abies faxoniana and Acer davidii seedlings, their highest values of the same parameters were found under intermediate light regime (i.e. 25-55% of PFD relative to full sunlight). Plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources. Leaf nitrogen and chlorophyll content increased, while dark respiration rate and light compensation points decreased, all of which were adaptive response to the low light environment. On the contrary, plants under high light environment had the thicken leaves and palisade tissue, which was a protective response to high light. Phenotypic plasticity to light Phenotypic plasticity can be exhibited in morphological and physiological processes. Physiological characteristical adjustment is the main for plant adaptation to different light environment.The means of plasticity indexes for Betula albo-sinensis and Acer davidii seelings were greater than Picea asperata and Abies faxoniana, amplied that the two broad-leaved trees were much more adaptable to the environment. In addition, spruce had the higher adaptablity than fir. The findings supported the hypothesis that the ecological characteristics of the species determined the biological status and its biological habitat selection. Photoinhibition and photoprotection to light Compared with conifer, broad-leaved trees could better change leaf morphology and adjust biomass allocation to adapt to changing light environment. However, excess light can photoinhibit photosynthesis and may lead to photooxidative destruction of the photosynthetic appatus. Two field experiments were conducted to study the photoinhibition of photosynthesis. The results showed that when plants grown under high light environment or plants transferred from low to high irradiance, the four tree seedlings would undergo a period of photoinhibition. In four species, photoinhibited leaves could recover to initial photosynthetic rates when they were long-term planted under high light environment. However, when plants were suddenly exposed to high irradiance, this photoinhibition could not be reversible, may be the photosynthesis apparatus were (or partly) photooxidatively destructed.
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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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植物群落及其环境在干扰后的演替格局和过程的研究,是群落和生态系统动态研究的一个热点。选取青藏高原东缘山原区川西云杉林皆伐后,从草地过渡到灌丛的关键阶段的4 个皆伐迹地(恢复时间为8 a、10 a、16 a 和21 a),研究皆伐及自然恢复过程对林下典型灌木银露梅(Potentilla glabra)和唐古特忍冬(Lonicera tangutica)的生长与繁殖能力的影响以及灌木植物在迹地上的更新情况,分析灌木在不同生境中的适应对策和适应能力的差异,为揭示青藏高原东缘山原区迹地植被从草甸到灌丛演替的过程和特点及促进迹地演替与植被恢复进程提供理论依据和技术支撑。研究主要结论如下:1)皆伐后银露梅生长和繁殖能力显著提高,但对唐古特忍冬的影响不明显。皆伐后银露梅丛基径、高度和各部分生物量都显著增加(P < 0.05),但唐古特忍冬只有叶生物量和地下生物量增加,总生物量和其余构件生物量无显著变化。皆伐后,银露梅的结实数量、结实株数、不结实株数和结实株/不结实株比例显著增加(P < 0.05)。自然恢复过程中,银露梅和唐古特忍冬生长能力以及银露梅的结实量都表现出降低的趋势。随着迹地自然恢复时间的增加,银露梅和唐古特忍冬的基径、高度、丛叶片数和各部分生物量有减少的趋势。银露梅的结实株数增加, 但结实数量减少。2)皆伐对银露梅和唐古特忍冬生物量分配模式影响不一致。原始林和迹地中(除CT85)银露梅的生物量大小关系皆为:地下>茎>侧枝>叶。唐古特忍冬在原始林中的生物量大小关系为:茎>地下部分>侧枝>叶,而皆伐后生物量的分配情况改变,生物量大小关系变为:地下部分>茎>侧枝>叶。随着自然恢复时间的增加,银露梅减少了地下生物量的分配,而唐古特忍冬增加了地下生物量的分配。3)皆伐和恢复时间的增加改变了迹地物种组成,促使阳性乔、灌木在迹地上定居。4 个迹地上共出现了灌木15 种,乔木3 种,没有出现天然云杉和冷杉幼苗。随着恢复时间的增加,迹地上的灌木物种由原始林下的耐阴物种逐步发展为以针刺悬钩子(Rubus pungens)为主的阳性灌木。4)影响灌木幼苗密度和幼树密度的因子不一致。灌木幼苗密度与灌木层盖度显著负相关,与苔藓层盖度显著正相关。幼树密度与草本层盖度正相关,与苔藓层盖度、灌木层盖度和高度负相关。5)研究发现在青藏高原东缘山原区皆伐15~20 a 后,迹地仍以草本植物为主,推测皆伐后至少20 a 以上迹地才可能向灌丛阶段过渡,比高山峡谷地区的演替进程至少推迟了20 a。银露梅和唐古特忍冬在皆伐后自然恢复过程中表现出不同的生长与繁殖策略是由两个物种的生物学特性的差异引起的。银露梅比唐古特忍冬更适应迹地退化环境。促进青藏高原东缘山原林区迹地森林恢复一方面是尽量减少人为活动的破坏,另一方面,可以通过在迹地中播种适当的乡土乔、灌木种子(如白桦、银露梅)等人工措施,以加快演替进程。The succession pattern and process of plant community and their environments is a hot spotin community and ecosystem dynamic study. Four clearcuts were chosen in Rangtang(recovery time of 8 a、10 a、16 a and 21 a), which represented the key stage of thecommunity evolved from grass stage to shrub stage in the eastern margin of theQinghai-Tibetan Plateau. The growth and reproduction of the Potentilla glabra andLonicera tangutica and the natural regeneration of shrub plants in the primary Piceabalfouriana forest and 4 clearcuts were studied to explore how clear cutting andnatural recovery process affected the understory shrub species during the 21 years inthe eastern margin of the Qinghai-Tibetan Plateau. The main results were below.1) The growth and reproduction of P. glabra significantly increased after forestclear cutting.. But it was not so significant as to the L. tangutica. The organismbiomass and total biomass of P. glabra were increased obviously after clear cutting(P< 0.05). But only leaves and underground biomass of L. tangutica increasedsignificantly after clear cutting(P < 0.05). The number of fruit and growth of P. glabraincreased significantly after clear cutting too(P < 0.05). The ramet height, basaldiameter , organism biomass and friut number of P. glabra and L. tangutica reducedas the increase of recovery time.2) The biomass allocation patterns varied between P. glabra and L. tangutica inthe primary forest and clearcuts. The biomass allocation of P. glabra both in primary forest and clearcuts was followed as: underground part > stem > branch > leaves.However, the biomass allocation of L. tangutica had changed after the clear cutting.The biomass allocation of L. tangutica in the primary forest was followed as: stem >underground part > branch > leaves and it was underground part > stem > branch >leave in clearcuts. The biomass allocation of P. glabra and L. tangutica varied amongclearcuts. Aboveground biomass was increased while underground biomass decreasedfor P. glabra with the increase of recovery period. However, the L. tangutica showedthe reverse changing pattern.3) Clear cutting and recovery time had changed the species composition of theclearcuts. There were 15.shruby species and 3 tree species in the four clearcuts. Nospruce and fir seedlings were found. In the early stage after clear cutting, there wereonly understory shrub species from the primary Picea balfouriana forest. The sunnyspecies, especieally Rubus pungens invaded intensly as the increase of recovery time.4) There was a significant negative relationship between density of seedlingswith shrub layer coverage and positive correlation with moss coverage. The saplingshad significantly positive correlation with herb layer coverage and negativecorrelation with moss coverage, shrub layer coverage and height.5)Comparing to studies in Miyalou, a nearby high mountain and canyon area,the secondary sucession in this subalpine plateau areas lagged at least 20 years.P. glabra and L. tangutica showed different growth and reproduction strategies toclear cutting and natural recovery , which may associated with the difference of theirbiological characters. P. glabra was more adaptive to the clear cutting than the L.tangutica. Two suggestions were probably recommended to promote the recoveryprogress in the subalpine plateau areas based on the results of this study. Limitanthropogenic disturbance, and meanwhile sow native tree and shrub seeds inclearcuts.
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揭示水体中繁殖的两栖动物在异质性景观中的空间扩散特点,探讨景观面积丧失和破碎化对于两栖动物的影响,为两栖动物的保护提供理论依据。本文以四川西北部若尔盖湿地自然保护区的高原林蛙(Rana kukunoris)为研究对象,通过运用地理信息系统及建立景观模型等方法,在分析若尔盖湿地自然保护区范围内现有景观格局的基础上,建立了高原林蛙的景观扩散模型,并模拟了“沼泽→草甸”的湿地逆演化过程下高原林蛙的空间分布与景观连接的变化特点。主要结果是: 1.若尔盖湿地自然保护区呈典型的沼泽—草甸式斑块—基质景观格局。草甸面积占整个景观面积的79.42%,景观蔓延度指数(CONTAG)为79.00远离最小值0而更趋向于最大值100,面积和景观蔓延度指数表明草甸是整个景观中面积占绝对优势且景观连接好的类型,构成了景观的基质,对景观的动态格局演变起主导作用。沼泽面积仅占整个景观面积的18.08%,但却是整个景观中斑块数目最多的单元,占所有斑块数的82.9%。因此沼泽斑块与草甸基质之间的动态结构对高原林蛙的扩散起着决定性的作用。 2.空间扩散模型表明,其它类型的景观不但扩展了高原林蛙的活动范围,而且也为高原林蛙在不同沼泽斑块间的连接提供了通道。高原林蛙的空间扩散区域使得彼此间成斑块化隔离状分布的沼泽形成了潜在景观功能连接,促进了不同斑块间物种的交流。小型沼泽作为垫脚石(stepping-stone),使得整个景观中的相隔距离较远的大型斑块联结为一个功能整体,促进了高原林蛙在整个景观中的相互动态联系。 3.模拟“沼泽→草甸”的湿地逆演化过程表明,大量小型沼泽湿地的消失将会 对在沼泽中繁殖并扩散到其它景观类型中去的高原林蛙造成潜在影响。逆演化过程不仅使沼泽斑块的分布范围,沼泽源斑块的面积和空间扩散面积减少,而且对景观连接也有很大影响。小型沼泽的消失,将使得景观斑块的功能连接变小,使得依靠小型沼泽作为跳板的动物在沼泽斑块之间的移动将变得更加困难。 本文是对生境丧失与破碎化影响下两栖动物的行为反应的一种尝试。影响模型的因素很多,包括动物对各种类型景观的偏好程度,地理数据的精度,及模型的可靠程度都是制约模型准确度的因素。 The spatial diffusion of water—breeding amphibian through heterogeneous landscape and the effects of landscape losing and fragmentation to amphibian were the core theory of the landscape ecology of amphibian. Geographical information system (GIS) and landscape model were used to model the diffused area of Rana kukunoris in Zoige Wetland Natural Reserve. Model was also used to analysis the spatial distribution variation of R. kukunoris and the change of landscape connectivity when simulated the retrogressive succession of landscape. The main results are below: 1. There was peatland—meadow pattern which was typical patch—matrix landscape pattern in Zoige Natural Reserve. The meadow area occupied 79.42% of the entire landscape area, contagion index (CONTAG) was 79.00 which was far away the minimum value (0) but tend to the maximum value (100). Both of these showed that meadow was the largest part and the most continue units. It was shown that meadow was matrix of the landscape, which evolved the leading role to the landscape dynamic pattern. Though their area only occupies 18.08% of entire landscape area, peatlands were according to 82.9% of the total patches. Dynamic of the pattern between peatlands and meadows decided the spatial diffusion of R. kukunoris. 2.The model indicated that the other types of landscape not only expanded diffusion of R. kukunoris, but also have provided the potential channels for frog's connections among different peatlands. The spatial diffusion zone of R. kukunoris forced isolated patch peatlands to be potential landscape functional connectivity. The small peatlands, as stepping-stone, made the large peatlands connect as a functional one and promoted the integrated and dynamic connectivity of R. kukunoris in the whole landscape. 3. The simulation of “peatlands→meadows”retrogressive succession process indicated that the decrease of small peatlands will have potential effect to R. kukunoris because they must bred in peatlands and diffuse to other type of the landscape. Retrogressive succession process not only made the decrease of distribution of peatlands, patches number of peatlands and diffused area of R. kukunoris, but also reduced the connectivity among source patches. As stepping-stone, the disappearance of small peatlands will made the migration of R. kukunoris among patches more difficult. The model was an experiment of the amphibian behavior reaction to habitat losing and fragmentation. There were many factors that could influence the accuracy of model, such as the preference of animals to each type of landscape, the geographical data precision, reliable degree of model.
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采用TWINSPAN分类法对黄土丘陵沟壑区吴起县双树沟流域30个自然恢复草地植被进行分类,并对分类后各植被群落特征和地上生物量进行统计分析。结果表明:在自然恢复条件下,随着退耕年限的不断增加,退耕地植被自然恢复依次经历了猪毛蒿群落—赖草+长芒草群落—赖草+铁杆蒿群落—铁杆蒿群落—铁杆蒿+茭蒿群落5个发展阶段,地带性植被类型铁杆蒿+茭蒿群落在研究区内开始出现,并且已经占有一定优势;随着退耕地植被自然恢复的不断进行,Margalef等丰富度指数以及Shannon-wiener等多样性指数、Pielou等物种均匀度指数和地上生物量都呈现出先减小后增大的发展趋势;在植被自然恢复的稳定阶段,虽然物种丰富度指数和物种多样性指数有一定的增加,但是相对恢复初期来讲还是有所下降,并且有达到与初期相当水平的趋势;物种丰富度指数均在第1恢复阶段最大,而均匀度指数Jsw以及Shannon-wiener指数在第5恢复阶段最高。随着退耕地植被自然恢复的不断进行,植被群落总盖度随着退耕年限的延长而不断增大。