川西亚高山人工针叶林下11种植物对两种光生境的适应性研究

青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障，经过近六十年的采伐后，取而代之的是大量人工种植的云杉纯林。目前，这些人工林已经表现出树种单一，结构层次简单等生态问题，其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性，完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分，对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此，研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象，选择林下11种具有不同喜光特性的常见植物，分别设置人工林林冠下及成熟林窗为研究样地，通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析，研究林下植物对不同光生境的适应策略及其适应能力，揭示不同物种对人工林生境的适应共性，为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下： 在叶片形态和物质分配特征方面：在林窗光生境中，11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时，林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加，这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后，植物可能更多地采取调节叶片组织细胞水平（即叶片功能细胞形态）及叶片器官水平（即单个叶片形态）特征的策略来适应各类生境，而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面：多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多，而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面：在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下，多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率，并在中午12:00～14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境，即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境，通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之，植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关，但另一些参数（Fv/Fm日变化、色素含量及比例、叶氮相对含量）受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面：在叶片各表型参数中，器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性；叶片光合组织的可塑性大于非光合组织可塑性；反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合，即喜光种及轻度耐荫种各参数可塑性最高，深度耐荫种可塑性最小，而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面：喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低，深度耐荫种疏花槭却恰恰相反，更多数物种（7种植物）在两种生境中密度变化趋势不明显。从分布格局来看，7种植物在两种生境中均为聚集分布，但聚集强度为林窗>林内；少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型，在林内生境中的分布型发生改变而成为随机型，说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关，同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.

中国卧龙自然保护区不同海拔川滇高山栎(Quercus aquifolioides)群体的遗传变异

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.

青藏高原东缘高山林线乔木种群生态特征研究

青藏高原东部分布着世界最高的林线，该区域也是由欧亚北温带物种形成的林线的南界。在大面积野外踏勘的基础上，选择青藏高原东部具有典型高山林线分布的三个地点（滇西北白马雪山、川西北鹧鸪山及岷江源地区）作为研究区，从种群的结构、生存特征、分布格局及分形特征等方面对青藏高原东缘高山林线乔木种群生态学特征进行了研究，并在此基础上探讨了人类活动对林线种群生态特征及林线格局的影响。结果表明，林线区乔木树种多以单种群形式存在，林线区群落结构简单，乔木层多为单一树种组成，其生长型较之郁闭林发生了急剧的变化：树木高度急剧下降，而发展多茎多分枝的生长型。生长型的转变是高山林线乔木对恶劣自然条件的形态适应。 研究发现，在青藏高原东缘，阴坡林线乔木主要是冷杉（Abies spp.），阳坡主要由圆柏（Sabina spp.）组成，少数地方还有云杉（Picea spp.）。阴坡乔木种群结构多表现为增长型，幼苗和幼树在种群中占较大比重，种群潜在自然更新能力较强，但幼龄个体死亡率非常高，存活曲线多接近Deevey-Ⅲ型；阳坡乔木种群幼苗个体数极少，幼树相对增加。野外调查表明，人为活动较频繁的阳坡林线区幼苗数量极少甚至缺失，而受人为活动干扰较小的样地中幼苗和幼树数量明显增多，从一个侧面说明放牧等人类活动可能对林线种群的更新带来较大影响，而对卡卡沟围栏内外的样地分析也进一步证明了这一结果。 所研究林线乔木种群各龄级的空间格局在不同尺度上表现为聚集、随机和均匀分布，以聚集分布为主；各龄级在不同尺度上表现出显著的相关性，幼苗通常与另外两个龄级的关联性较密切。各龄级间显著的相关性表明不同龄级个体在空间交错分布，有利于对各种资源的充分利用，对种群的生存和发展非常有利，反映了高山生态系统恶劣生境中种群的一种适应对策。 林线乔木种群各龄级分布格局的计盒维数有差别，林线种群的计盒维数总是小于郁闭林种群的计盒维数。另外，郁闭林各龄级计盒维数通常也高于林线各龄级，表明不同海拔或者不同群落类型中的乔木树种具有不同的水平空间占据能力。林线区种群分布格局的计盒维数都很低，占据现实水平空间的程度较低，具有相对较高的生态间隙维，其潜在占据空间的能力较高，群落还可提供给种群的最大空间限度较大，但实际上由于受群落中种内、种间的竞争及林线区恶劣的生态环境条件的限制，其潜在空间占据能力可能难以表现出来。 青藏高原东缘高海拔地带以季节性游牧为主要的资源利用和生产方式，阳坡森林郁闭度低于阴坡，灌丛数量和种类较阴坡少，融雪早且积雪时间短，所以阳坡包括高山林线区成为当地牧民游牧路线的必经之地。牲畜的践踏、啃食使得幼龄乔木树种个体数量大大减少，严重阻碍了林线乔木种群的自然更新，同时种群占据空间的能力也明显降低。因此可以认为，在青藏高原东部地区，山地游牧等人为干扰叠加于恶劣的自然条件，阳坡林线的自然更新潜力受到抑制，其生存状态较之阴坡林线显著恶化，并可使阳坡林线高度逐渐降低。高山林线区森林一旦破坏在短时间内很难有效更新和恢复，因此，对于处于恶劣高山生境中的乔木种群应加强保护，同时适度控制人为干扰强度和幅度以减少其直接和间接破坏，防止阳坡林线退化并促进高山生态系统的自然恢复。 Eastern Qinghai-Tibetan Plateau has the highest timberline of the world. On the basis of field surveys and literature reviews, three typical alpine timberlines were chosen for in-depth studies, i.e., Baima Snow Mountain in northwest Yunnan, Zhegu Mountain and the waterhead area of Minjiang River in west Sichuan. Using the methodologis of population ecology, we analyzed the population structure, survival characteristics, spatial point patterns and fractal dimensions of the timberline tree populations and discussed the impacts of grazing on the structure and spatial pattern of alpine timberline. Compared with closed forests, the community structure of timberline is simpler, usually with one or two species constituting the tree layer. Differences also exist in the growth forms: the trees were significantly shorter with more stems and branches, reflecting morphological adaptation of trees to the severe conditions at timberline. In the eastern Qinghai-Tibetan Plateau, Abies spp. often formed alpine timberline in the north-facing slope while Sabina spp. and sometimes Picea spp. in the south- facing slope. The population structures of north-facing slope showed an increasing trend, with numerous seedlings and saplings. However, the survival curves tend to follow Deevy-III because of high dead ratio of young individuals. There are only few seedlings in the south-facing slope with heavy grazing, demonstrating that human disturbance may prevent regeneration at alpine timberline, which was confirmed by comparisons between fenced enclosures and control plots in the Kaka Valley. Depending on the spatial scales on consideration, the individuals of different age-classes showed clumping, random or even distribution, but mostly with clumping distribution. At all scales, individuals in different age-classes were all significantly correlated with each other while the seedlings were usually more correlated to two other age classes. This high degree of correlation among different age classes indicates that individuals of different age classes are spatially interlocked with each other, which helps sufficient utilization of various resources and is conducive to the survival and development of population. It is another adaptation strategy for trees at the severe environment. The spatial patterns of different age classes had different box dimension. In general, the box dimensions of total individuals and each age class at timberline are always smaller than that of closed forests, suggesting that space occupation capacity is not the same for populations at different altitude or in different communities. Populations on both the south- and the north-facing slopes had a very low box dimensions (far away from the max., 2), however, the lower the box dimension, the bigger the potential space provided by community. In fact, because of inner- and inter- competition as well as the severe conditions at timberline, this kind of potential ability can hardly be realized. Mountain pastoralism is the major type of as well as the only most effective way of resource uses in the high elevation regions of the eastern Qinghai-Tibetan Plateau. Due to lower canopy cover, less bushes and short snow-cover time, south-facing slopes became the favorite pastures. Damages from livestock through tramping, browsing and others have greatly reduced the number of young individuals. As a result, the potential of timberline trees to regenerate and their ability to occupy more space are greatly inhitibted. We conclude that human disturbances (mountain pastoralism) as well as harsh environmental conditions co-worked to inhibit the regeneration of tree populations in the south-facing slope and made south slopes more difficult than the north-facing slopes for trees to survive and develop, resulting a gradual retreat of timberline in the north-facing slopes. Forests at alpine timberline are susceptible to disturbance and difficult to regenerate and restore once damaged and controlling human disturbances is important for protecting the forest ecosystems at the timberline area.

岷江冷杉林群落几种树苗对光强的响应与适应

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.

三种豆科灌木对干旱胁迫的响应与适应

干旱胁迫是全球范围内影响植物生存、生长和分布的重要环境因子。岷江上游干旱河谷区，由于生态环境的脆弱性和长期人类活动的干扰和过度利用，导致植被严重退化，水土流失加剧，山地灾害频繁，干旱化和荒漠化趋势明显。这种趋势若不能遏制，将严重阻碍区域社会经济的快速协调发展，并且威胁成都平原地区的发展和长江中下游地区的生态安全。因而开展干旱河谷生态恢复研究成为解决这些问题的关键。水分匮乏是限制干旱河谷生态恢复的关键因子，在全球气候变化的背景下，干旱胁迫在区域尺度上可能会更加严重，并使干旱河谷的生态环境更加恶化。因此，深入研究干旱河谷乡土植物对干旱胁迫的响应和适应机理，具有非常重要的理论和实践意义。 本论文以岷江上游干旱河谷的三种乡土豆科灌木，白刺花（Sophora davidii）、小马鞍羊蹄甲（Bauhinia faberi var. microphylla）和小雀花（Campylotropics polyantha）理论和实践意义。为研究对象，在人工控制条件下设计了4-5个连续性干旱胁迫处理，系统地研究了灌木幼苗的生长、生物量积累和水分利用效率（WUE）、形态结构和生理过程等对干旱胁迫的反应，揭示了幼苗的干旱适应能力及种间差异。主要研究结论如下： 1) 灌木生长和繁殖对干旱胁迫的反应 在干旱胁迫下，幼苗生长速率显著减小，叶片衰老和脱落比率增大，这些变化随着胁迫强度的增加具有累积效应。叶片比茎对干旱胁迫的反应更敏感。在严重干旱胁迫下，幼苗的有性繁殖被限制，但在中等程度干旱胁迫下，幼苗的有性繁殖能力被提高。 2) 灌木生物量积累及其分配和WUE对干旱胁迫的反应 在干旱胁迫下，灌木各器官的生物量都显著减小，但是生物量的分配侧重于地下部分，使得根茎比在干旱条件下增大。幼苗的耗水量（WU）随着干旱胁迫的增加而显著减少。白刺花和小马鞍羊蹄甲WUE在干旱胁迫下降低；小雀花的WUE在中等干旱胁迫下升高。 3) 灌木叶片结构特征对干旱胁迫反应 白刺花叶片具有较为典型的旱生型结构，而小马鞍羊蹄甲和小雀花则为中生型结构。在1至2年的干旱胁迫下，灌木叶片结构组成未发生本质性的改变，主要是细胞大小的变化。在中等和严重干旱胁迫下，叶肉组织厚度明显减小；并且气孔和表皮细胞面积也显著减小，气孔和表皮细胞密度显著增大；叶肉细胞层数、P/S值、表皮厚度等无显著变化。 4) 灌木对干旱胁迫的生理响应 气体交换参数和叶片相对含水量（RWC）在中等干旱胁迫下发生了明显的改变，而叶绿素荧光参数和光合色素含量在严重干旱胁迫下才发生显著变化。在干旱胁迫下，净光合作用速率（Pn）、气孔导度（gs）和RWC呈下降趋势，而叶片温度（Tl）呈增加趋势，蒸腾速率（Tr）的变化不明显。除了日最大Pn减小以外，干旱胁迫对气体交换参数的日变化无显著影响，但是对光合-光响应曲线有显著的影响，使有效光合时间缩短。在严重干旱胁迫下光系统受到损害而代谢减弱，PSⅡ中心的内禀光能转换效率（Fv/Fm）、量子产量（Yield）、光化学淬灭参数（qP）显著降低，而非光化学淬灭参数（NPQ）明显增加。气孔限制和非气孔限制对Pn的影响与干旱胁迫强度有关。在中度胁迫下，气孔限制起主导作用，在严重胁迫下非气孔限制起主导作用，40% FC水分条件可能是灌木由气孔限制向非气孔限制的转折点。 5) 灌木对干旱胁迫的适应能力及其种间差异 三种灌木对干旱胁迫具有较好的适应能力，即使在20% FC，幼苗未因干旱胁迫III而死亡；80% FC适宜于幼苗生长。白刺花生长速率慢，耗水量较少，具有较强的耐旱和耐贫瘠能力，并具有干旱忍受机制，能够在较干旱的环境中定居和生长。小马鞍羊蹄甲和小雀花，生长快，水分消耗量较大，尤其是小雀花，对干旱胁迫的忍受能力较弱，具有干旱回避机制，因而适宜于在较为湿润的生境中生长。综合分析表明，生长速率较慢的物种抗旱能力较强，其更适宜于作为干旱地区植被恢复物种。 Drought is often a key factor limiting plant establishment, growth and distribution inmany regions of the world. The harsh environmental conditions and long-termanthropogenic disturbance had resulted in habitat destruction in the dry valley ofMinjiang river, southwest China. Recently, it tended to be more severe on the vegetationdegradation, soil erosion and water loss, natural disaster, as well as desertification, whichimpact on regional booming economy and harmonious development, and would be verydangerous to the environmental security in the middle and lower reaches of Yangzi River.Therefore, ecological restoration in the dry valley is one of the vital tasks in China. Waterdeficit is known to affect adversely vegetation restoration in this place. Moreover, in thecontext of climate change, an increased frequency of drought stress might occur at aregional scale in the dry valleys of Minjiang River. The selection of appropriate plantingspecies for vegetation restoration in regard to regional conditions is an important issue atpresent and in further. The research on responses of indigenous species to drought stresscould provide insights into the improvement of the vegetation restoration in the dry valleys of Minjiang River. In this paper, the responses of three indigenous leguminous shrubs, Sophora davidii,Bauhinia faberi var. microphylla and Campylotropics polyantha, to various soil watersupplies were studied in order to assess drought tolerance of seedlings, and to compare interspecific differences in seedlings’ responses to drought stress. The results were as follows: 1 Growth and reproduction of shrubs in response to drought stress Seedling growth reduced significantly while leaf senescence accelerated underdrought stress, the cumulative responses to prolonged drought were found. The capacityfor reproduction was limited by severe drought stress, and improved by moderate droughtstress. Leaf responses were more sensitive than shoot to various water supplies. 2 WUE, biomass production and its partitioning of shrubs in response to drought stress Drought stress reduced significantly the total dry mass and their components ofseedlings, and altered more biomass allocation to root system, showing higher R/S ratiounder drought. Water use (WU) and water-use efficiency (WUE) of both S. davidii and B.faberi var. microphylla declined strongly with drought stress. The WU C. polyantha ofalso declined with drought stress, but WUE improved under moderate drought stress. 3 Anatomical characteristics and ultrastructures of leaves in response to drought stress There were xeromorphic for S. davidii leaves and mesomorphic for B. faberi var.microphylla and C. polyantha at the all water supplies. The foundational changes in leafstructures were not found with drought stress. However, mesophyll thickness, the areas ofstomatal and epidermis reduced slightly while the densities of stomatal and epidermisincreased under severe drought stress. Variations in these parameters could mainly be duoto cell size. Other structures did not displayed significant changes with drought stress. 4 Physiological responses of shrubs to drought stress The gas exchange parameters and leaf relative water content (RWC) were affectedby moderate stress, while chlorophyll fluorescence and chlorophyll content were onlyaffected by severe stress. Drought stress decreased net photosynthesis rate (Pn), stomatalconductance, light-use efficiency and RWC, and increased leaf temperature. Therespiration rates (Tr) were kept within a narrower range than Pn, resulting in aprogressively increased instantaneous water use effiecency (WUEi) under drought stress.Moreover, drought stress also affected the response curve of Pn to RAR, there was adepression light saturation point (Lsat) and maximum Pn (Pnmax) for moderate andsevere stressed seedling. However, diurnal changes of gas exchange parameters did notdiffer among water supplies although maximum daily Pn declined under severe stress.VISevere stress reduced Fv/Fm, Yield and qP while increased NPQ and chlorophyll content.Photosynthetic activity decreased during drought stress period due to stomatal andnon-stomatal limitations. The relative contribution of these limitations was associatedwith the severity of stress. The limitation to Pn was caused mainly by stomatal limitationunder moderate drought stress, and by the predominance of non-stomatal limitation undersevere stress. In this case, 40% FC water supply may be a non-stomatal limitation 5 Interspecific differences in drought tolerance of shrubs Three shrubs exhibited good performance throughout the experiment process, evenif at 20% FC treatment there were no any seedlings died, 80% FC water supply wassuitable for their establishment and growth. S. davidii minimized their water loss byreducing total leaf area and growth rate, as well as maintained higher RWC and Pncompared to the other two species under drought stress, thus they might be more tolerantto the drought stress than the other two species. On the contrary, it was found that C.polyantha and B. faberi var. microphylla had higher water loss because of their stomatalconductance and higher leaf area ratios. They reduced water loss with shedding theirleaves and changing leaf orientation under drought stress. Based on their responses, thestudied species could be categorized into two: (1) S. davidii with a tolerance mechanismin response to drought stress; (2) C. polyantha and B. faberi var. microphylla withdrought avoidance mechanism. These results indicated that slow-growing shrub speciesare better adapted to drought stress than intermediate or fast-growing species in present orpredicted drought conditions. Therefore, selecting rapid-growing species might leavethese seedlings relatively at a risk of extreme drought.

干旱胁迫与施N条件下白刺花（Sophora davidii）幼苗生长及其适应机制

干旱环境常常由于多变的降水事件和贫瘠土壤的综合作用，表现出较低的生产力和较低的植被覆盖度。全球性的气候变暖和人类干扰必将使得干旱地区缺水现状越来越严竣。贫瘠土壤环境中已经很低的有效养分含量也将会随着干旱的扩大而越来越低。干旱与半干旱系统中不断加剧的水分与养分的缺失将严重限制植物的生长和植被的更新，必然会使得已经恶化的环境恶化速率的加快、恶化范围的加大。如何抑制这种趋势，逐步改善已经恶化的环境是现在和将来干旱系统管理者面临的主要关键问题。了解干旱系统本土植物对未来气候变化的适应机制，不仅是植物生态学研究的重要内容，也对人为调节干旱环境，改善干旱系统植被条件，提高植被覆盖度具有重要的实践意义。 本研究以干旱河谷优势灌木白刺花（Sophora davidii）为研究对象，通过两年大棚水分和施N控制实验和一个生长季野外施N半控制实验，从植物生长－生理－资源利用以及植物生长土壤环境特征入手，系统的研究了白刺花幼苗生长特性对干旱胁迫和施N的响应与适应机制，并试图探讨施N是否可调节干旱系统土壤环境，人工促进干旱条件下幼苗定居，最终贡献于促进植被更新实践。初步研究结论如下： 1）白刺花幼苗生长、生物量积累与分配以及水分利用效率对干旱胁迫和施N处理的适应白刺花幼苗株高、基径、叶片数目、叶面积、根长、生物量生产、相对含水量和水分利用效率随着干旱胁迫程度的增加而明显降低，但地下部分生物量比例和R/S随着干旱胁迫程度的增加而增加。轻度施N处理下幼苗株高、基径、叶片数目、叶片面积和生物量生产有所增加。但重度施N处理下这些生长指标表现出微弱甚至降低的趋势。严重干旱胁迫条件下，幼苗叶面积率、R/S、相对含水量和水分利用效率也以轻度施N处理为最高。 2）白刺花幼苗叶片光合生理特征对干旱胁迫和施N处理的适应叶片光合色素含量和叶片光合效率随着干旱胁迫程度的增加而显著降低，并且PS2系统在干旱胁迫条件下表现出一定程度的光损害。但是比叶面积随着干旱胁迫程度的增加而增加。在相对较好水分条件下幼苗净光合速率的降低可能是因为气孔限制作用，而严重干旱胁迫条件下非气孔限制可能是导致幼苗叶片光合速率下降的主要原因。叶片叶绿素含量、潜在光合能力、羧化效率、光合效率以及RUBP再生能力等在施N处理下得到提高，并因而改善干旱胁迫条件下光合能力和效率。虽然各荧光参数对施N处理并无显著的反应，但是干旱胁迫条件下qN和Fv/Fm在轻度施N处理下维持相对较高的水平，而两年连续处理后在严重干旱胁迫条件下幼苗叶片光合效率受到重度施N处理的抑制，并且Fv/Fm和qN也在重度施N处理下降低。 3）白刺花幼苗C、N和P积累以及N、P利用效率对干旱胁迫和施N处理的适应白刺花幼苗C、N和P的积累，P利用效率以及N和P吸收效率随干旱胁迫程度的增加而显著降低，C、N和P的分配格局也随之改变。在相同水分处理下，C、N和P的积累量、P利用效率以及N和P吸收效率在轻度施N处理下表现为较高的水平。然而，C、N和P的积累量和P利用效率在重度施N处理下不仅没有表现出显著的正效应，而且有降低的趋势。另外，在相同水分条件下白刺花幼苗N利用效率随着施N强度的增加而降低。 4）白刺花幼苗生长土壤化学与微生物特性对干旱胁迫和施N的适应白刺花幼苗生长土壤有机C、有效N和P含量也随干旱胁迫程度的增加而明显降低。干旱胁迫条件下土壤C/N、C/P、转化酶、脲酶和碱性磷酸酶活性的降低可能表明较低的N和P矿化速率。尽管微生物生物量C、N和P对一个生长季干旱胁迫处理无显著反应，但微生物生物量C和N在两年连续干旱胁迫后显著降低。土壤有机C和有效P含量在轻度施N处理下大于重度施N处理，但是有效N含量随着施N强度的增加而增加。微生物生物量C和N、碱性磷酸酶和转化酶活性也在轻度施N处理下有所增加。但是碱性磷酸酶活性在重度施N处理下降低。 5）野外条件下白刺花幼苗生长特征及生长土壤生化特性对施N的适应植物生长、生物生产量、C的固定、N、P等资源的吸收和积累、其它受限资源的利用效率（如P）在轻度施N处理下均有所增加，但N利用效率有所降低。幼苗生物生产量及C、N和P等资源的分配格局在轻度施N处理下也没有明显的改变。白刺花幼苗叶片数目、生物生产量和C、N、P的积累量在重度施N处理下虽然也相对于对照有所增加，但幼苗根系长度显著降低。生物量及资源（生物量、C、N、P）在重度施N处理下较多地分配给地上部分（主要是叶片）。另外，土壤有机C、全N和有效N含量随外源施N的增加而显著增加，土壤pH随之降低，但土壤全P含量并无显著反应。其中有机C含量和有效P含量以轻度施N处理最高。微生物生物量C、N和P在轻度施N处理下也显著增加，而微生物生物量C在重度施N处理下显著降低。同时，转化酶、脲酶、碱性磷酸酶和中性磷酸酶活性在施N处理下也明显的提高，但酸性磷酸酶和过氧化氢酶活性显著降低，其中碱性磷酸酶和中性磷酸酶活性以轻度施N处理最高。 综合分析表明，干旱河谷水分和N严重限制了白刺花幼苗的生长。施N不能完全改变干旱胁迫对白刺花幼苗的抑制的作用，但是由于施N增加土壤N有效性，改善土壤一系列生物与化学过程，幼苗的生长特性也对施N表现出强烈的反应，表现为植物结构与资源分配格局的改善，植物叶片光合能力与效率的提高，植物生长以及利用其他受限资源（如水分和P）的效率的增加，致使植物自身生长及其生长环境在干旱环境下得到改善。但是过度施N不仅不能起到改善干旱胁迫下植物生长环境、促进植物生长的作用，反而在土壤过程以及植物生长过程中加重干旱胁迫对植物的伤害。因此，建议在采用白刺花作为先锋种改善干旱河谷系统环境的实践中，可适当施加N以改善土壤环境，调节植物利用与分配资源的效率，促进植物定居，得到人工促进种群更新的目的。但在实践过程中也要避免过度施N。 Arid regions of the world are generally noted for their low primary productivity which is due to a combination of low, unpredictable water supply and low soil nutrient concentrations. The most serious effects of global climate change and human disturbances may well be those which related to increasing drought since drought stress has already been the principal constraint in plant growth. The decline in total rainfall and/or soil water availability expected for the next decades may turn out to be even more drastic under future warmer conditions. Nevertheless, water deficit is not the only limiting factor in arid and semiarid environments. Soils often suffer from nutrient (especially N and P) deficiencies in these ecosystems, which can also be worsened by climate change. How to improve the poor soil quality and enhance the vegetation coverage is always the problem facing ecosystem managers. The adaptive mechanisms of native plant to future climate change is always the focus in plant ecology, it also plays important roles in improving vegetation coverage by manual controlled programmes. Sophora davidii is a native perennial shrub of arid valleys, which is often predominant on eroded slopes and plays a vital role in retaining ecological stability in this region. It has been found that S. davidii was better adapted to dry environment than other shrubs, prompting its use for re-vegetation of arid lands. A two-years greenhouse experiment and a field experiment were conducted in order to understand the adaptation responses of Sophora davidii seedlings to different water and N conditions, and further explore if additional N supply as a modified role could enhance the adaptation ability of S. davidii seedlings to dry and infertile environment. Two-month old seedlings were subjected to a completely randome design with three water (80%, 40% and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg-1 soil) regimes. Field experiment was arranged only by three N supplies in the dry valley. 1) The growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings in respond to drought stress and N supply Seedlings height, basal diameter, leaf number, leaf area, root length, biomass production, relative water content (RWC) and WUE were decreased with increase of drought stress. An increase in below-ground biomass was observed indicating a higher root/shoot ratio (R/S) under drought stress conditions. Low N supply increased seedlings height, basal diameter, leaf number, leaf area, and biomass production, but decreased root length. In contrast, these growth characteristics showed little or negative effect to high N supply treatment. Leaf percentages increased with increase of N supply, but fine root percentages decreased. In addition, Low N supply rather than the other two N treatments increased leaf area ratio (LAR), leaf/fine root mass ratio (L/FR), R/S and RWC under severe drought stress (20%FC), even though these parameters could increase with the high N supply treatment under well-watered condition (80%FC). Moreover, Low N supply also increased WUE under three water conditions, but high N supply had little effect on WUE under drought stress conditions (40%FC and 20%FC). 2) Leaf gas exchange and fluorescence parameters of Sophora davidii seedlings in respond to drought stress and N supply Leaf area (LA), photosynthetic pigment contents, and photosynthetic efficiency were decreased with increase of drought stress, but specific leaf area (SLA) increased. Photodamage in photosystem 2 (PS2) was also observed under drought stress condition. The decreased net photosynthetic rate (PN) under relative well-watered water conditions might result from stomatal limitations, but the decreased PN under other hand, photosynthetic capacity by increasing LA, photosynthetic chlorophyll contents, Pnmax, CE, Jmax were increased with increase N supply, and photosynthetic efficiency was improved with N supply treatment under water deficit. Although N supply did a little in alleviating photodamages to PS2 caused by drought stress, low N supply enhanced qN and kept relative high Fv/Fm under drought stress condition. However, high N supply inhibited leaf photosynthetic efficiency, and declined Fv/Fm and qN under severe drought stress condition after two year continues drought stress and N supply. 3) Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in respond to drought stress and N supply C, N and P accumulation, NUE , N and P uptake efficiency (NUtE and NUtE ) P N P were decreased with increase of drought stress regardless of N supply. On the other hand, the S. davidii seedlings exhibited strong responses to N supply, but the responses were inconsistent with the various N supply levels. Low N supply rather than the other two N treatments increased C, N and P accumulation, improved NUEP, NUtE and NUtE under corresponding water condition. In contrast, high N supply N P did few even depressed effects on C, N and P accumulation, and NUEP, although NUtEN and NUtEP could increase with high N supply under corresponding water conditions. Even so, a decrease of NUEN was observed with increase of N supply under corresponding water conditions. 4) Soil microbial and chemical characters in respond to drought stress and N supply The content of soil organic C, available N and P were decreased with increase of drought stress. Decreases in C/N and C/P, and invertase, urea and alkaline phosphatase activity were also observed under drought stress conditions, indicating a lower N and P mineralization rate. Although microbial biomass C, N and P showed slight responses to drought stress after one growth period treatment, microbial biomass C and N were also decreased with increase of drought stress after two year continuous treatment. The content of soil organic C and available P showed the stronger positive responses to low N supply than which to high N supply, although than the other two N treatments increased microbial biomass N and invertase activity under severe drought stress condition, even though invertase activity could increase with high N supply treatment under relative well-water conditions. Moreover, low N supply treatment also increased C/P and alkaline phosphatase activity which might result from higher P mineralization, but high N supply did negative effects on alkaline phosphatase activity. 5) The growth characteristics of Sophora davidii seedlings and soil microbial and chemical characters in respond to N supply under field condition Low N supply facilitated seedlings growth by increasing leaf number, basal diameter, root length, biomass production, C, N and P accumulation and absorption, and enhancing the use efficiency of other limited resources as P. Compared to control, however, low N supply did little effect on altering biomass, C, N and P portioning in seedlings components. On the contrary, high N supply treatment also increased leaf number, biomass and C, N and P accumulation relative to control, but significantly decreased root length, and altered more biomass and resources to above-ground, which strongly reduced the ability of absorbing water under drought condition, and thus which might deep the drought stress. In addition, N supply increased soil C, N and available N content, but declined pH and showed little effects on P content. Low N supply showed higher values of soil C and available P content. Low N supply also increased microbial biomass C, N and P, although high N supply decreased microbial biomass C. N supply significantly enhanced soil invertase, urea, alkaline and neutral phosphratase activity, while declined acid phosphratase and catalase activity. Low N supply exhibited higher alkaline and neutral phosphratase activity compared to the others. The results from this study indicated that both drought and N limited the growth of S. davidii seedlings and their biomass production. Regardless of N supply levels, drought stress dramatically reduced the seedlings growth and biomass production. Although plant growth parameters, including basal diameter, height, leaf number, and biomass and their components were observed to be positive responses to low N supply, N supply alone can not alter the diminishing tendency which is caused by drought. available N content increased with increase N supply. In addition, low N supply rather These findings imply that drought played a primary limitation role and N was only the secondary. Even so, appropriate N supply was seemed to enhance the ability that S. davidii seedlings adapted to the xeric and infertile environment by improving soil processes, stimulating plant growth, increasing recourses accumulation, enhancing use efficiency of other limited resources, and balancing biomass and resources partitioning. Appropriate N supply, therefore, would be recommended to improve S. davidii seedling establishment in this region, but excess N supply should be avoided.

角蟾科（Megophryidae; Anura）蝌蚪的形态多样性和生态适应

角蟾科（Megophryidae）是以角蟾属（Megophrys Kuhl and Van Hasselt, 1822）为模式属而建立的，隶于无尾目（Anura），变凹型亚目（Anomocoela）。角蟾科包括2 亚科11 属142 种，分布于东洋界，从巴基斯坦、中国西部向东直到菲律宾和苏达群岛；中国有9 属75 种分布于华中和华南地区。角蟾科被认为是原始的两栖动物之一，其分类学、系统学、生态学、动物地理学的研究均深受中外科学家的瞩目。近年来，通过形态学、古生物学、细胞学、生态学、支序系统学的研究，角蟾科的分类与系统学研究取得了较大进展。与成体形态和分子系统学研究结果相比较，蝌蚪的研究存在更多的问题和挑战，尚需深入研究：（1）角蟾科蝌蚪的形态多样性分析；（2）角蟾科的系统发育关系与蝌蚪的演化，以及口漏斗的起源；（3）角蟾科蝌蚪表型分化与栖息环境和觅食行为的适应演化。针对上述问题，本文对角蟾科9 属30 种蝌蚪的形态特征，包括外部宏观形态和口器外部结构特征、口器内部显微结构、唇齿和角质颌的亚显微结构作了深入细致、多层次的比较研究；通过12s rRNA 和cytochrome b 基因构建最大简约树，采用贝叶斯系统发育进行分析，蝌蚪型的演化采用祖先性状的重建方法分析；得到如下结论：1）初步将角蟾科蝌蚪分为4 种类型；并且建立了2 种新的角蟾科蝌蚪类型。A 型：拟髭蟾型蝌蚪，该型蝌蚪包括拟髭蟾属、髭蟾属、齿蟾属和齿突蟾属的物种；B 型：新类型，掌突蟾型蝌蚪，该型蝌蚪在本文中包括掌突蟾属、小臂蟾属的物种；C 型：新类型，短腿蟾型蝌蚪，一种特化类型，该型蝌蚪在本文中仅包括短腿蟾属的物种；D 型：角蟾型蝌蚪，该型蝌蚪在本文中包括无耳蟾属、小口拟角蟾属和异角蟾属的物种。2）对角蟾科的分类进行了修订：（1）支持角蟾科两个亚科的分类系统；（2）角蟾亚科包括拟角蟾属、异角蟾属、无耳蟾属和短腿蟾属；该亚科形态差异小，系统学关系比较复杂，暂不作族级分类的再划分；（3）拟髭蟾亚科分为2 个族：拟髭蟾族，该族物种具有类型A 的蝌蚪，包括4 个属：拟髭蟾属、髭蟾属、齿蟾属、齿突蟾属；掌突蟾族，该族物种具有类型B 的蝌蚪，包括2 个属：掌突蟾属和小臂蟾属。3）结合分子系统进化关系探讨了4 种蝌蚪类型的演化。（1）角蟾科蝌蚪的最近共同祖先来自于一类具有拟髭蟾型蝌蚪性状的蝌蚪；（2）掌突蟾型蝌蚪和角蟾亚科的蝌蚪是由具有拟髭蟾型蝌蚪性状的祖先蝌蚪分别演化而来；（3）短腿蟾型蝌蚪是角蟾型蝌蚪的一种特化类型；（4）外群蝌蚪具有与拟髭蟾型蝌蚪相似的性状，进一步印证了类拟髭蟾型蝌蚪是角蟾科蝌蚪的最近共同祖先的假说；（5）具有口漏斗的蝌蚪类型是由不具口漏斗的蝌蚪类型演化而来，在角蟾科中口漏斗是一种衍生性状。4）分析了角蟾科四种蝌蚪类型与栖息环境的适应演化。（1）角蟾科蝌蚪的口部和体形的变化反映了该科蝌蚪由缓流向类似静水生境的回水凼的渐变式适应，角蟾科蝌蚪的形态显示了多方面的适应变化；（2）随着蝌蚪类型由A 向D的演化，当水速较大时，拟髭蟾型的蝌蚪营流水攀吸型生活方式；当水速递减时，掌突蟾型蝌蚪营流水附着型生活方式；当水速进一步递减时，具有较小口漏斗的短腿蟾型蝌蚪和具有大漏斗的角蟾型蝌蚪营流水浮泳型生活。角蟾科蝌蚪对于水流递减的适应演化说明蝌蚪的生态学适应是具有进化意义的；（3）蝌蚪口器内部结构的分化揭示了蝌蚪和食性的适应关系，蝌蚪以口部的唇齿与角质颌刮取或吞吸水中的物质，然后，通过口乳突有选择地过滤进入口腔中食物。拟髭蟾亚科蝌蚪的唇齿多而窄，唇齿间距宽，颌鞘粗而稀，反映了其植食性为主的特点；它们的舌前乳突一般为指状，在口腔入口处所占面积小，其机械过滤的作用很多被唇齿和角质颌分担了；而角蟾亚科的蝌蚪，其角质颌弱，其舌前乳突一般为匙状，几乎填满了口腔入口处，因此舌前乳突起了主要的机械过滤作用。The family Megophryidae is the largest and most diverse families inArchaeobatrachia, and most of its species occur in India, Pakistan, and eastward intoChina, Southeast Asia, Borneo and the Philippines to the Sunda Islands. Currently thefamily includes 142 species have been grouped into two subfamilies, Megophryinaeand Leptobrachiinae. The mountains of central and southern China are rich in speciesof Megophryidae, 75 species belong to 9 genera and two subfamilies.The family was supposed to be ideal materials of studies in many fields of biology,such as taxonomy, evolution, systematics, ecology, and biogeography. Recently, therehave a great development in taxonomy and systematics of megophryids throughstudied by morphology, paleontology, cytology, ecology, and cladistics. However,larvae of megophryids were generally unknown, although the tadpoles might be veryimportant for above studies.In this paper, we examined the evolutionary scenario of the tadpoles’ morphologyin the context of a phylogenetic framework. Our objectives are (1) to evaluate thedivergence of larval body shape and oral discs in the family Megophryidae, (2) toexplore the evolutionary trends of the larvae in megophryidae, and test if thefunnel-shaped oral disc is apomorphic, and (3) to explore the relationship of the larvalstructure, diet and microhabitat.We examined larval morphology of 30 megophryid species, the larval body shape,oral discs, the buccopharyngeal cavity, and jaw sheaths and denticles of the Chinesemegophryid frogs were re-examined. We constructed a phylogeny of the species on thebasis of published mitochondrial cytochrome b and 16S rRNA gene segments usingpartitioned Bayesian analyses. Furthermore, hypothetical changes of larval morphologywere inferred using parsimony principle on the phylogeny. The results showed that:1) Four tadpole types in Megophryidae. The larval morphological charactersseries in Chinese megophryids fall into four general categories according to the bodyshape and oral discs: (A) Leptobrachiini type, species from genera Leptobrachium,Oreolalax, Scutiger and, Vibrissaphora share this type of tadpoles. (B) Leptolalax type,species of genus Leptolalax have this type of tadpoles. (C) Brachytarsophrys type,species of the genus Brachytarsophrys have this type of tadpoles. (D) Megophryinitype, species of the genera Atympanophrys, Ophryophryne, and Xenophrys share this type of tadpoles. Of which B and C are two novel types.2）Taxonomic implications. The present study leads us to reconsider the generalclassification of tribes attributed to members of Megophryidae. More specifically,concerning the phylogenetic relationships and the two novel tadpole types describedherein, we propose a provisional taxonomy for the family but suggest that further taxasampling of other megophryids be performed to confirm this taxonomic change. TheMegophryidae is composed of two subfamilies (Leptobrachiinae and Megophryinae).The Leptobrachiinae was recogonized the two tribes: (1) tribe Leptobrachiini sensuDubois, corresponding to the tadpole of type A, including four genera, i.e.,Leptobrachium, Oreolalax, Scutiger and, Vibrissaphora; (2) tribe Leptolalaxini,corresponding to the tadpole of novel type B, including two genera, i.e., Leptolalaxand Leptobrachella. However, the relationships among the genera of Megophryinaewere largely unresolved, they recognized no monophyletic groups above the generalevel. A more thorough sampling will likely foster a better taxonomic solution.3) The larval evolutionary scenario in Megophryidae.Type A is characteristicof normal-mouthed with multiple tooth rows, representing the tadpole type of theMRCA of Chinese megophryids. Type B is characteristic of normal-mouthed withreduced tooth rows, prolonging labium, and integumetary glands. Type C ischaracteristic of no labial teeth and smaller umbeliform oral disc. Type D ischaracteristic of no labial teeth, enlarged umbeliform oral disc, representing the tadpoleof the MRCA of subfamily Megophryinae. A previous hypothesis, referring tofunnel-shaped oral discs as an apomorphy, is supported.4) The larval adaptation to habitats in Megophryidae. Tadpoles generallyadhere to substrates using their mouths, and the microhabitat that the tadpoles occupyreflects the degree of adhesion and oral complexity. The morphological changes inmegophryid tadpoles virtually allow a progressive adaptation to a changing habitatfrom faster water to slower water. Within the tadpoles of Type A to type D, the TOTbecomes smaller and smaller, and the oral disc orientates from anteroventral toumbelliform upturned, and eye position orientates from dorsal to lateral, and the trunkis more and more depressed and tail becomes relatively longer and slender. Within therunning water, the normal-mouthed with multiple tooth rows of Leptobrachiini tadpoles are correlated with lotic-suctorial, benthic feeders with anteroventral oraldisc and the largest body. With the water’s velocity decreasing, the lotic-adherentfeeders of Leptolalax tadpoles have tube-shaped labium with reduced tooth rows andintegumetary glands. And then, the smaller umbeliform in Brachytarsophrys tadpolesand the enlarged umbeliform oral disc in the Megophryini tadpoles are inhabitmicrohabitats of non-flowing backwaters of rivers, indicative of adaptive traits oflotic-neustonic surface feeders. The scheme of megophryid tadpoles andmicrohabitats provided the first clear evidence which congruent with the hypothesis ofAltig and Johnston (1989). The ecological divergence plays a general role in thedivergence and evolution of megophrid larvae. There is a definite correlation amongthe buccopharyngeal cavity, diet and feeding mechanisms, the tadpole graze orswallow the food particles, then through papillae which like a sieve and sort out foodparticles to the oesophagus. The tadpole of Leptobrachiinae possess multiple toothrows, wide intertooth distance as well as thick and sparse jaw sheath, these tadpolesinhabit bottom of the streams and graze on epiphyton or major detritus of organicmatter on the substrates, their prelingual papillae like single finger, the mechanicalpurpose of papillae served share in by tooth and jaw. The tadpoles of Megophryinaeoccur near the water surface of small streams and are the filter feeder, their dietincludes plankton and organic debris floating on the water surface, those tadpolepossess weak jaw, their prelingual papillae like spoon, the mechanical purpose ofpapillae served mostly for sieve.

Ten Years of the Relative Risk Model and Regional Scale Ecological Risk Assessment

It has been 10 years since the publication of the relative risk model (RRM) for regional scale ecological risk assessment. The approach has since been used successfully for a variety of freshwater, marine, and terrestrial environments in North America, South America, and Australia. During this period the types of stressors have been expanded to include more than contaminants. Invasive species, habitat loss, stream alteration and blockage, temperature, change in land use, and climate have been incorporated into the assessments. Major developments in the RRM have included the extensive use of geographical information systems, uncertainty analysis using Monte Carlo techniques, and its application to retrospective assessments to determine causation. The future uses of the RRM include assessments for forestry and conservation management, an increasing use in invasive species evaluation, and in sustainability. Developments in risk communication, the use of Bayesian approaches, and in uncertainty analyses are on the horizon.

Considerable effects of diversity indices and spatial scales on conclusions relating to ecological diversity

The relationships between ecological diversity and ecosystem functions such as stability and productivity have long been debated and have no final conclusion until now. It is ignored that the debate should be firstly based on the same diversity index, which should be theoretically complete, and on same observation scale. For the issue on the scale of ecotope observation, ecosystems should be distinguished according to intensity of human disturbance. For the issue on the scale of species observation, either number diversity or biomass diversity should be identified. This paper takes grassland ecosystems located within the Bayin Xile grassland of Xilin Gol League of Inner Mongolia Autonomous Region as an example to analyze effects of different diversity indices and spatial scales on the conclusions of ecological diversity and its relationships with ecosystem functions. The analysis results both on the scale of ecotope observation and on the scale of species observation show that different diversity indices might give different conclusions and spatial resolution has a great effect on the relative conclusions. (c) 2005 Elsevier B.V. All rights reserved.

Method for calculating ecological water storage and ecological water requirement of marsh

As one of the most typical wetlands, marsh plays an important role in hydrological and economic aspects, especially in keeping biological diversity. In this study, the definition and connotation of the ecological water storage of marsh is discussed for the first time, and its distinction and relationship with ecological water requirement are also analyzed. Furthermore, the gist and method of calculating ecological water storage and ecological water requirement have been provided, and Momoge wetland has been given as an example of calculation of the two variables. Ecological water use of marsh can be ascertained according to ecological water storage and ecological water requirement. For reasonably spatial and temporal variation of water storage and rational water resources planning, the suitable quantity of water supply to marsh can be calculated according to the hydrological conditions, ecological demand and actual water resources.

Genome-wide survey of putative serine/threonine protein kinases in cyanobacteria

Background: Serine/threonine kinases (STKs) have been found in an increasing number of prokaryotes, showing important roles in signal transduction that supplement the well known role of two-component system. Cyanobacteria are photoautotrophic prokaryotes able to grow in a wide range of ecological environments, and their signal transduction systems are important in adaptation to the environment. Sequence information from several cyanobacterial genomes offers a unique opportunity to conduct a comprehensive comparative analysis of this kinase family. In this study, we extracted information regarding Ser/Thr kinases from 21 species of sequenced cyanobacteria and investigated their diversity, conservation, domain structure, and evolution. Results: 286 putative STK homologues were identified. STKs are absent in four Prochlorococcus strains and one marine Synechococcus strain and abundant in filamentous nitrogen-fixing cyanobacteria. Motifs and invariant amino acids typical in eukaryotic STKs were conserved well in these proteins, and six more cyanobacteria- or bacteria-specific conserved residues were found. These STK proteins were classified into three major families according to their domain structures. Fourteen types and a total of 131 additional domains were identified, some of which are reported to participate in the recognition of signals or substrates. Cyanobacterial STKs show rather complicated phylogenetic relationships that correspond poorly with phylogenies based on 16S rRNA and those based on additional domains. Conclusion: The number of STK genes in different cyanobacteria is the result of the genome size, ecophysiology, and physiological properties of the organism. Similar conserved motifs and amino acids indicate that cyanobacterial STKs make use of a similar catalytic mechanism as eukaryotic STKs. Gene gain-and-loss is significant during STK evolution, along with domain shuffling and insertion. This study has established an overall framework of sequence-structure-function interactions for the STK gene family, which may facilitate further studies of the role of STKs in various organisms.