114 resultados para diffusive gradients
Resumo:
通过室内模拟降雨试验,研究了地面坡度对红壤坡面产流过程和侵蚀过程的影响,结果表明:坡面总径流量随坡度的增大呈减小趋势,其中25°坡面比5°坡面的径流量减小22.3%;随坡度的增加,坡面产流达到稳定的历时减少。径流含沙量和坡面侵蚀产沙量随坡度的变化在5°~20°呈增大趋势,而当坡度由20°增加到25°时,径流含沙量和侵蚀产沙量随坡度的增加呈减少的趋势,即在红壤坡面,坡度对侵蚀产沙量影响存在着临界坡度,其值变化于20°~25°之间。
Resumo:
采用室内模拟降雨研究了不同降雨强度和坡度对红壤坡面降雨入渗、产流产沙的影响。结果表明:产流时间随降雨强度和坡度的增大而提前。对于入渗率较低的红壤坡面,径流强度主要受坡面承雨强度的影响,径流强度随雨强增大而增大,随坡度增大而减小。坡面入渗率随降雨强度的增大表现为先增大后减小。降雨强度增大能够增加水分入渗,但这种促进作用仅在一定的降雨强度范围内起作用。坡面入渗率随坡度增大表现为先增大后减小,20°左右存在一个坡面入渗率变化的临界坡度。坡面细沟侵蚀发展状况对稳定坡面入渗率有比较大的影响。径流率和入渗率与降雨历时的关系分别可以用幂函数和对数函数描述。在75,100 mm/h降雨强度下,产沙量随坡度的增大而增大,而50 mm/h降雨强度下,产沙量在20°时达到最大值而后出现下降,在20°附近存在一个侵蚀产沙量变化的临界坡度。
Resumo:
通过室内人工降雨+放水冲刷相结合的方法,对5个坡度(3~18°),5种雨强(1~3mm.min-1),5组放水流量(1.4~9.0kg.m-2.s-1)下的黄棉土土质路面和早熟禾植物路面进行了共60余场次试验,对比分析了植物路相对于土质路侵蚀过程差异及产水、减沙效果。结果表明:在放水冲刷条件下,植物路侵蚀率随放水历时降低,其过程可用对数函数描述;土质路面侵蚀率变化复杂。两类道路的放水流量与侵蚀强度均呈现幂函数变化,且植物道路表现出明显的防蚀作用。雨强增加引起径流量、侵蚀强度增加,土质路的增加快于植物路;坡度增大时,侵蚀强度增大,径流深则为植物路稳定,土质路有小幅增加。植物路相对土质路的减沙幅度为26.24%~47.22%,减少能力可用抛物线方程描述,但植物路较土质路面有利于产生径流,径流深增加幅度平均为58.7%~144.1%,由于径流的泥沙含量大大降低,道路的侵蚀产沙反而有一定降低。
Resumo:
利用安塞试验站1985-1992年的气象观测数据和野外坡度径流小区径流和土壤侵蚀量监测资料,评价了WEPP模型在黄土丘陵沟壑区不同坡度条件下的适用性。结果表明,径流量的模拟值在10°、15°、20°、25°和28°五个坡度条件下,变化幅度不如实测值明显,但模拟值随坡度变化的趋势和实测值相一致;WEPP模型对次降雨、年平均和多年平均土壤侵蚀量模拟结果较好,且无论是模拟值在不同坡度之间的差值、还是模拟值随坡度变化趋势均与实测值接近。I30对WEPP模型模拟次降雨径流量有重要影响,当I30大于0.92mm/min,模型模拟误差较大。WEPP模型对次降雨土壤侵蚀量的模拟与PI30密切相关,当PI30大于129mm2/min时,模型模拟误差较大。∑PI30对WEPP模型模拟年平均径流量和侵蚀量有重要影响,当∑PI30大于150mm2/min时,模拟精度明显下降,且∑PI30对径流模拟影响较对土壤侵蚀模拟影响明显。WEPP模型对次降雨、年平均和多年平均径流量模拟的Nash-Sutcliffe有效性ME分别为0.914、0.912和0.617;对次降雨、年平均和多年平均土壤侵蚀量模拟的Nash-Sutcliffe有效性ME...
Resumo:
The evolution of strain and structural properties of thick epitaxial InGaN layers grown on GaN with different thicknesses are investigated. It is found that, with increase in InGaN thickness, plastic relaxation via misfit dislocation generation becomes a more important strain relaxation mechanism. Accompanied with the relaxation of compressive strain, the In composition of InGaN layer increases and induces an apparent red-shift of the cathodoluminescence peak of the InGaN layer. On the other hand, the plastic relaxation process results in a high defect density, which degrades the structural and optical properties of InGaN layers. A transition layer region with both strain and In composition gradients is found to exist in the 450-nm-thick InGaN layer.
Resumo:
A new framework of non-local model for the strain energy density is proposed in this paper. The global strain energy density of the representative volume element is treated as a non-local variable and can be obtained through a special integral of the local strain energy density. The local strain energy density is assumed to be dependent on both the strain and the rotation-gradient. As a result of the non-local model, a new strain gradient theory is derived directly, in which the first and second strain gradients, as well as the triadic and tetradic stress, are introduced in the context of work conjugate. For power law hardening materials, size effects in thin metallic wire torsion and ultra-thin cantilever beam bend are investigated. It is found that the result predicted by the theoretical model is well consistent with the experimental data for the thin wire torsion. On the other hand, the calculation result for the micro-cantilever beam bend clearly shows the size effect.
Resumo:
The diffusive transport properties in microscale convection flows are studied by using the direct simulation Monte Carlo method. The effective diffusion coefficient D is computed from the mean square displacements of simulated molecules based on the Einstein diffusion equation D = x2 t /2t. Two typical convection flows, namely, thermal creep convection and Rayleigh– Bénard convection, are investigated. The thermal creep convection in our simulation is in the noncontinuum regime, with the characteristic scale of the vortex varying from 1 to 100 molecular mean free paths. The diffusion is shown to be enhanced only when the vortex scale exceeds a certain critical value, while the diffusion is reduced when the vortex scale is less than the critical value. The reason for phenomenon of diffusion reduction in the noncontinuum regime is that the reduction effect due to solid wall is dominant while the enhancement effect due to convection is negligible. A molecule will lose its memory of macroscopic velocity when it collides with the walls, and thus molecules are hard to diffuse away if they are confined between very close walls. The Rayleigh– Bénard convection in our simulation is in the continuum regime, with the characteristic length of 1000 molecular mean free paths. Under such condition, the effect of solid wall on diffusion is negligible. The diffusion enhancement due to convection is shown to scale as the square root of the Péclet number in the steady convection regime, which is in agreement with previous theoretical and experimental results. In the oscillation convection regime, the diffusion is more strongly enhanced because the molecules can easily advect from one roll to its neighbor due to an oscillation mechanism. © 2010 American Institute of Physics. doi:10.1063/1.3528310
Resumo:
九寨沟湖泊湿地在维持九寨沟的生态平衡中起着重要的作用,在旅游产业的发展下,湿地生态系统及生物多样性面临着较大的威胁。尽管九寨沟湿地具有重要的生态价值,但目前对其研究尚比较薄弱。湿地植物群落和植物地理研究可以为湿地资源的可持续利用和监测保护提供科学依据。作者从2004年8月到2007年11月对九寨沟湿地的植物物种组成、地理分布、优势植物群落的结构、生长动态、湿地土壤种子库进行了调查研究。主要结果如下: 1. 九寨沟湿地物种组成、地理分布特点及湿地植物群落特点 九寨沟湿地共有苔藓植物8科13属16种,维管植物为48科107属199种。九寨沟湿地植物的地理成份较为丰富,维管植物在科级水平上有7种地理分布型(变型),在属级水平上有13种地理分布型(变型), 在种级水平上共有29种地理分布型(变型)。九寨沟湿地植物以温带成份和我国特有成份为主,同时兼有热带、亚热带成份和环极—高山成份。九寨沟湿地植物的分布表现出明显的垂直地带性和水平地带性。湿地植物群落可划分21个群落类型,不同植物群落类型的物种多样性及物种组成存在较大的差异。九寨沟湿地植物的物种多样性和群落多样性以及较高的生产力特征,是维持其湿地生态景观多样性和稳定性的基础。 2. 土壤、水环境、海拔等对湿地植物的分布及生物多样性的影响 九寨沟湿地土壤、水等环境因子存在较大的差异。帕米尔苔草和宽叶香蒲等群落的凋落物较多,土壤有机碳、土壤总磷较高,可能是九寨沟湿地的重要土壤碳库。 九寨沟湿地植物沿水环境梯度的分布规律表现为:沉水植物(轮藻—篦齿眼子菜,水苦荬,杉叶藻)——挺水植物(水木贼,芦苇,宽叶香蒲)——湿生草本(苔草、节节草、披散木贼)——湿生灌木(柳灌丛,小檗灌丛)等。海拔也影响湿地植物的物种组成。 水深对物种多样性有影响,水深与物种丰富度负相关。随着水深的增加,水木贼、芦苇、杉叶藻、宽叶香蒲等群落的物种多样性下降;在长期淹水和季节性淹水的地方,水木贼群落物种多样性存在显著差异。土壤总氮与水木贼群落物种丰富度正相关。 3. 土壤营养元素、水环境对植物生长的影响 水深影响湿地植物生物量的分配。芦苇无性系分株在47 cm水深的环境中单株平均生物量最大;在干滩地中(地面水深0 cm),叶生物量百分比最大,而茎生物量百分比最小,茎的生物量百分比和生长速率随水深的增加而增加;在较干的滩地生境中,开花率、花序的生物量百分比明显大于水较深的生境。 水深与水木贼地上生物量负相关,但水木贼地上生物量在长期淹水和季节性淹水的地方没有显著的差异。在水浅的地方,杉叶藻、水木贼、芦苇等植物群落中,其他伴生物种的生物量占样方总生物量的百分比较大。 土壤有机碳、土壤总氮、土壤总磷等对湿地植物生物量的影响比较大:宽叶香蒲地上生物量与土壤总磷正相关;水木贼地上生物量与土壤总氮正相关;杉叶藻地上生物量与土壤有机碳正相关。 水深、土壤营养成分对湿地植物高度、密度等有影响。水木贼的平均高度在季节性淹水的地方比长期淹水的地方低,平均密度在长期淹水的地方比季节性淹水的地方低;除了5月份,其他观察月份水木贼的密度都与水深负相关,同时与土壤有机碳正相关。另外,芦苇密度与土壤有机碳含量正相关,宽叶香蒲密度与水深负相关,帕米尔苔草高度与土壤有机碳负相关。 4. 优势植物群落的动态变化 在优势植物群落中,优势种的高度、密度、盖度、生物量等在群落中占绝对优势。除五花海,水木贼群落的物种组成、高度、生物量在两年间没有显著的变化。芦苇群落的物种丰富度在近两年有所增加。 湿地植物生长表现为明显的季节动态,生长的峰值大多在7月-8月。优势植物群落的物候与水文周期有关。湿地植物群落的物种组成和密度,可以作为对湿地监测和保护的生物指示。 5. 九寨沟湿地土壤种子库特征及其在湿地生物多样性恢复中的作用 水深和现存植被物种丰富度可以解释湿地土壤种子库的变化。水深可以解释表层物种丰富度45%的变化。现存植被物种丰富度可以分别解释10 cm土层、2-5 cm土层及5-10 cm土层土壤种子库45%、48%和25%的变化。 湿地土壤种子库的密度为0-15945粒m-2, 种子库中共发现23个物种。现存植被优势物种和种子库优势物种不同。各层土壤种子库密度和物种丰富度并不存在显著的差异,但第二层土壤种子库密度最大。海拔、现存植被优势种盖度、土壤总磷、土壤总氮、土壤有机碳对湿地土壤种子库的密度和垂直结构没有影响。土壤种子库物种丰富度小于地上植被物种丰富度。湿地土壤种子库与地上植被的相关性不大。在浅水区域,湿地土壤种子库在湿地植被恢复中有一定作用。但在深水区域,保护现存植被更重要。 The lakeshore wetlands are valuable ecological units of the Jiuzhaigou lakes. Pressure for travel industry development pose a continuing and severe threat to the biodiversity-support function of the wetland system. Despite the ecological importance of wetlands in Jiuzhaigou, they are so far poorly studied. Both general plant communties and biogeographical studies are needed in order to attain basis for sustainable use the wetland resources and adequate protection of these areas. The present study was undertaken to examine aquatic plants distribution and the species compositon, structure and growth dynamics of their communities with variations of environmental factors along altitudes, water depth and soil properities gradients in Jiuzhaigou. Analysis of field survey data collected during August 2004 and November 2007 in lakeshore wetlands in Jiuzhaigou National Nature Reserve, Sichuan, China. The results were as following: (i) Species composition and biogeography in wetland vegetation 8 families, 13 genus, 16 species of moss and 48 families, 107 genus and 199 species of vascular plants in Jiuzhaigou wetlands were found. The floristic compositions were abundunt. Ten geographical distribution types at family level, 13 geographical distributions types at generic level and 29 geographical distribution types at specific level in vascular plants were found. Most species in Jiuzhaigou wetlands are temperate elements and Chinese endemic elements, with a few of tropical and subtropical and some circumarctic elements. And the plant distributions show clear vertical and horizontal patterns. There were 21 major wetland plant community types. Species composition and species richness in different plant communities are different. The species diversity and plant community diversity and their high biomass are the basis for the diversity and stability of wetland landscapes in Jiuzhaigou. (ii) Water depth, soil nutrients and altitudes influence on the species diversity and plant distribution. Total phosphorous and organic cabon in soil were higher in C. pamiernensis and T. latifolia communities, where are important cabon reservoirs in Jiuzhaigou wetlands. Along gradients of water depth, among populations of the dominant plant species present: submerged macrophytes (Chara vulgaris, Potagemonton pectinatus, Veronica anagalis-aquatica,Hippuris vulgaris), emergent macrophytes (Equisetum fluviatile, Phragamites australis, Typha latifolia), helophytes (Carex pamirensis )and shrubs (Salix sp., Berberis sp. ). Altitudes influence on the assemblage of plant communities. Water depth negatively correlated with species richness. Specie richness showed differences between permanently flooded sites and seasonally flooded sites in E. fluvatile communities. And total nitrogen in soil was negatively correlated with species richness in E. fluviatile communities. Altitudes show no significant influence on species richness, but in fact, through our analyses, they do have influence on the assemblage of wetland plants. (iii) Water depth, soil nutrients influence on the plant growth Water depth influences the biomass allocation in Phragmities australis. The average aboveground biomass of a single ramet (4.2 g) was the largest in the habitat with water level 47 cm above the soil surface. At the habitat with water level under soil surface 15 cm (-15 cm), the leaf biomass percentage (of the total ramet biomass) was the largest (46.1%), and the height and percentage of ramose ramets ( with branches on stem )(of the total ramets in a plot) were found obviously different. The deeper in water, the larger the biomass percentage and growth rate of stems were. The flowering rate and biomass of panicles were greater in shallow water than those in deep water. Water depth negatively correlated with aboveground biomass of E. fluviatile. However, above-ground biomass of E. fluviatile showed no significant difference between permanently flooded sites and seasonally flooded sites. But in shallow water, more biomasses of accompanying species were found in dominant plant communities such as H. vulgaris communities, E. fluviatile communities and P. australis communities. Water depth, soil nutrients influence on shoot density and shoot length of wetland plants. The shoot density of E. fluviatile was correlated to water depth in all growth months. Annual average density was significantly lower at permanently flooded sites than at seasonally flooded sites. But the annual average shoot length was significantly lower at seasonally flooded sites than at permanently flooded sites. (iv) Growth dynamics of dominant communities in Jiuzhaigou wetland The shoot length and shoot density, coverage and biomass of domiant species were dominated in plant communities. The species composition increased in P. australis communities in recent two years. The species richness in E. fluviatile communities showed no difference between 2005 and 2007. The above-ground biomass and shoot density in Five-flower Lake from July 2005 to July 2007 were significantly different, while in other sites, the differences were not significant. Shoot height, shoot density and above-ground biomass showed significant seasonal changes in all sites. Growth dynamics correlated with the cycle of water levels in lakes. Most plants growth parameters peaked at July or August. The biomass of T. latifolia peaked in August. But the shoot length of T. latifolia in deeper water peaked in July. The shoot length of E. fluviatile increased significantly from May to August except in seasonally flooded sites in Arrow-bamboo Lake. The species composition of communities and shoot density can be used as bioindicators in Jiuzhaigou wetland. (v) Soil seed bank in Jiuzhaigou wetland and its role in vegetation restoration Seed density in all soil layer samples was negatively correlated to water depth. Water depth can explain 45% variance of species richness in surface layer in sediment. Species richness in extant vegetation can explain 45%, 48%, 25% variance of species richness in total 10 cm and in 2-5 cm and 5-10 cm layer sediment respectively. Mean seed densities in wetlands ranged from 0 to 15945 m–2. A total of 23 species germinated in seed bank. The dominant species in seed bank and extant vegetation showed great difference. The total number of species and seedlings that germinated in different layers was not significantly different. But the second layer had the greatest seed density. In shallow water, seed bank can contribute to vegetation restoration, while in deeper water, protection of extant vegetation may be a better strategy.
Resumo:
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.
Resumo:
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.
Resumo:
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.
Resumo:
岷江上游干旱河谷区水土流失强烈,地质灾害频繁,生态环境十分脆弱,而土壤条件恶劣(水分不足和养分缺乏)是阻碍该区植被恢复的关键因子,因此研究水分和乡土灌木生长对土壤的影响对该区的生态恢复具有指导意义。本文通过定点模拟实验,选取三种优势豆科灌木为研究对象,分别是白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha),设置5 个水分梯度,分别为100%、80%、60%、40%和20%田间持水量(FC),对栽种植物与不种植物下土壤理化性质和酶活性进行测定分析,系统比较和研究了不同水分条件和不同乡土灌木生长对干旱河谷区土壤结构、养分循环、酶活性以及微生物量的影响。主要结果如下:1. 无论生长植物与否,土壤的毛管持水量和毛管孔隙度都随着水分含量的减少而降低,最大持水量、总孔隙度和容重变化不大,相应地,土壤中的非毛管孔隙随含水量的减少而升高。各水分条件下,种植植物的毛管持水量和毛管孔隙度低于无植物生长的土壤,非毛管孔隙度相应地高于无植物土壤。土壤含水量在100%-40% FC 时,三种豆科灌木的毛管持水量和毛管孔隙度存在差异,而20% FC 条件下,三种豆科灌木土壤的物理性质基本相同。2. 水分胁迫影响土壤中养分的矿化和积累,主要表现在降低了水溶性碳和铵态氮的含量,中等程度胁迫时(60% FC)促进了有机碳和硝态氮的富集,对速效钾和有效磷没有明显作用。种植豆科灌木后各水分梯度上都增加了有机碳、铵态氮、速效钾和有效磷的积累。增加程度上三种豆科灌木间有一定差异,对于土壤有机碳总量,种植白刺花和小马鞍羊蹄甲明显高于小雀花,同样的情况还出现在铵态氮和速效钾上,但是对于有效磷,种植小雀花后的增加程度则明显高于白刺花和小马鞍羊蹄甲。种植豆科灌木不仅增加了土壤养分的相对含量,也改变了其在水分梯度上的变化趋势及其变化幅度,这种作用主要体现在碳元素和氮元素上。3. 无植物生长时脲酶活性随水分含量的减少而升高,水分胁迫对磷酸酶和过氧化氢酶的作用不显著,蔗糖酶也保持在相对较高的水平。种植植物后,蔗糖酶、磷酸酶活性与无植物时相比有较大幅度的提高,种植白刺花的脲酶活性也升高,其升高的程度在不同水分含量时不同。种植植物还降低了酶活性在水分梯度上的变幅,使之在水分梯度间的差异显著性降低。脲酶活性在指示土壤性质改变方面是较敏感的指标,其它三种酶在不同植物间的差异不明显。4. 在无植物生长时,中等程度的水分胁迫(60% FC)提高了土壤微生物量碳含量,过高或过低的土壤水分均不利于微生物碳的积累。种植小马鞍羊蹄甲后微生物量碳在水分梯度上的变化趋势与无植物生长时一致,而种植白刺花和小雀花后微生物量碳随着水分含量的减少而降低。不同种类植物的微生物量碳在水分梯度上的变化特征也不同,100% FC 条件下三种植物间没有差异,80%和60% FC 条件下小马鞍羊蹄甲显著高于白刺花和小雀花,40%和20% FC 条件下白刺花和小马鞍羊蹄甲也显著高于小雀花,说明不同种类植物随着干旱胁迫程度的加深微生物量碳的降低幅度不同,在极度干旱时,白刺花和小马鞍羊蹄甲土壤依然保持了较高的微生物活性,而小雀花土壤微生物量则明显下降。The dry valley of the upper reaches of the Minjiang River is seriously degradedmountain ecosystem. It was endangered by extremely soil lost and frequentlygeological disaster. Previous studies showed that short of water and nutrients in soilwas the principal limiting factors of vegetation restoration in this area. The typical soiland three dominant leguminous shrubs Sophora davidii, Bauhinia faberi var.microphylla and Campylotropics polyantha in upper reaches of arid Minjiang Rivervalley were considered as experimental material. Two-month old seedlings of eachspecies were exposed to five water supplies (100%, 80%, 60%, 40% and 20% waterfield capacity (FC)) in a temperature and light-controlled greenhouse. Afterthree-month water treatment, soil physiochemical variables and soil microbialactivities were determined by conventional methods. The main results showed that:1. Soil capillary capacity and capillary porosity decreased along water supplyregimes in all treatments, while saturated water capacity, total porosity and bulkdensity kept in a relatively stable level, as a result, the non-capillary porosity andcapacity increased with decrease of water supply. Compared to non-planted soil, theplant-soil systems had a higher non-capillary porosity and capacity, suggestingappropriate oxygen was present in soil to maintain the living of microorganism. Soilof three type shrub species shared the same capillary capacity and capillary porosityunder 20% FC.2. Water soluble carbon and NH4+-N decreased in response to water stress, whiletotal organic carbon and NO3--N promoted by moderate water stress and inhibited by 100% and 20% FC. Total organic carbon, NH4+-N, rapidly available K and availableP increased after the planting of leguminous shrubs in five water supply regimescompared to non-planted soil. For TOC, NH4+-N and rapidly available K, thepromotion effect was higher in S. davidii and B. faberi var. microphylla than C.polyantha planted soil, while available P displayed the opposite side. The planting ofshrubs also reduced the variance of observed traits along water supply gradients.3. Drought stress increased urease activity in non-planted soil, while insignificantdifferences were observed in phosphatase and catalase activity among five watersupply regimes. The planting of leguminous shrubs facilitated the β-glucosidase andphosphatase activity compared to the non-planted soil. It also reduced the variance ofenzyme activity along water supply gradients. Urease was more sensitive to waterstress than other three enzymes.4. Soil water content significantly affected microbial biomass carbon andCmic:Corg. S. davidii and B. faberi var. microphylla showed more drought toleranceability than C. polyantha, attributing not only to their relatively smaller variance ofmicrobial biomass carbon along soil water supply gradients, but also to the highlevel of microbial activity under severe water stress. S. davidii and B. faberi var.microphylla benefited reproduction of soil microorganism at 60%-80% FC, whilesevere drought limited it due to the competition of water and nutrients between plantand soil microorganism.
Resumo:
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.
Resumo:
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.
Resumo:
横断山区干旱河谷地区的土壤是制约植被恢复的一个关键要素,但干旱条件下土壤质量与动态演变状况并不清楚,本研究以岷江干旱河谷核心地段的土壤为对象,研究了阳坡褐土土壤的物理、化学以及生物学特征在海拔梯度(1800m、2000m、2200m)格局及其时间动态(2005-2007)变化,应用主成分分析综合评价了土壤质量及其时空差异性,明确了土壤质量的变化趋势。主要研究结论如下: 1) 海拔梯度上土壤物理性状的变化,2005 年和2007 年土壤物理性状综合质量随着海拔的升高均得到了优化,即海拔2200 m>海拔2000 m>海拔1800 m。 2) 海拔梯度上土壤化学性质的变化,土壤化学综合性质2005 年随海拔升高而趋于变得更优,而2007 年海拔2000 m 最优,海拔1800 m 则最差。 3) 海拔梯度上土壤生物学性质的变化,2005 年土壤生物学性质随海拔升高表现出趋于更好,2007 年海拔2000 m 最优而海拔1800 m 地段最差。 4) 从土壤物理、化学、生物学三方面出发,应用主成分分析,分别分析得出2005 年和2007 年不同海拔高度的土壤质量综合得分。根据综合得分得出土壤质量综合评价的排列顺序为:2005 海拔2200 m>2007 年海拔2000 m>2005 年海拔2000 m>2007 年海拔1800 m>2007 海拔2200 m>2005 年海拔1800 m。2005年土壤综合质量随海拔升高而趋好,2007 年则以海拔2000 m 最优,海拔1800 m和2200 m 差异不大。 5)排除人为干扰后,干旱河谷土壤物理性状在海拔1800 m 略有恢复,海拔2000 m 变化不明显,而海拔2200 m 仍有退化趋势;土壤化学性质在海拔1800 m和2000 m 地段得到恢复,而海拔2200 m 处仍有退化;土壤生物学性质在海拔2000 m 地段呈恢复趋势,而1800 m 和2200 m 仍处于退化状态。综合质量分析表明,与2005 年相比,2007 年海拔1800 m 和2000 m 地段土壤质量趋于变优而海拔2200 m 地段仍有退化迹象。 Soil is a key factor that affect the restoration of vegetation in the Hengduan Mountains dry valley area. But the dynamics and quality of soil is not knowed in dry area. In this study, soil physiochemical and biological characteristics ranging from 1800~2200m above sea level from a typical south-facing slope at the Minjiang River dry valley area had been studied, and characteristics of changes in soil quality along altitudinal gradients and time scales were also discussed. The principal component analysis was used to assess the soil quality. The main results were as follows: 1) Changes in soil physical properties along altitudes. Soil physical properties obtained the optimization along with the elevation in 2005 and 2007. 2) Changes in soil chemical properties. It was summarized that soil chemical properties increased with elevation in 2005, but the soil of 2000 m was the best in 2007. 3) Changes in soil biological characteristics. Soil biological properties increased with elevation in 2005, but the soil of 2000 m was the best and 1800 m was the worst in 2007. 4) Change tendency of soil quality. With the soil physics, chemistry and biology characteristics, we analysised the change tendency of soil quality in altitudes. The result indicated that soil quality increased with altituded in 2005, and soil quality of 2000 m was the best in 2007. 5) In brief, the soil quality is by physics, chemistry as well as the biology synthesizes the influence the final outcome. And the soil quality's change was manifested by soil physics, chemistry and the biology characteristics. All the results indicated soil quality still degenerated at 2200 m in in the dry valley of Minjiang River. And soil quality of 1800 m and 2000 m resumed slight.