Relationship between mycorrhizal fungi and functional traits in absorption roots: research progress and synthesis

吸收根(absorption root)一般是指根枝系统末端少数几级具有初生结构、负责物质吸收的根。吸收根功能性状被广泛用于评价和预测植物个体到生态系统水平上的一系列功能和过程。菌根真菌侵染是吸收根的一个关键性状, 它可以深刻影响吸收根的形态、结构, 以及功能性状之间的关系。该文针对与吸收功能密切相关的菌根真菌与根毛和根直径之间的关系进行了研究综述, 提出了真菌侵染、根毛和化学防御之间关系的一个假说; 探讨了温带和热带不同类型的吸收根如何通过菌根真菌影响根的功能性状, 从而适应不同的水热条件、养分状况和能量消耗; 提出一些需要关注的议题和研究方向, 以期为菌根真菌与吸收根功能性状之间关系的研究提供借鉴。     

Root traits and taxonomic affiliation of nine herbaceous species grown in glasshouse conditions

This study examines whether root traits differed between three major plant families (Asteraceae, Fabaceae and Poaceae) and whether they are related to root respiration and exudation. Nine traits related to biomass allocation, root topology, morphology, chemical composition and mycorrhizal colonisation were examined for nine C₃ herbaceous species grown in controlled conditions. Poaceae differed from Fabaceae for the whole set of root traits examined except mycorrhizal colonisation, while Asteraceae showed intermediate characteristics. As compared to Fabaceae, Poaceae allocated more biomass to roots; showed a more sparsely branched root system with a small average root diameter, a high root dry matter content and a low nitrogen concentration. Root respiration was weakly related to root mass ratio and root dry matter content; no significant relationship was found between root functions and root architecture or morphology. This study shows that plant classification based on taxonomic affiliation reflects differences in root system traits and functions. Whole root system traits do not allow strong predictions of root respiration and exudation, perhaps because these processes are more linked to fine root than to whole root system traits.     

Interspecific variation in plant responses to mycorrhizal colonization in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal fungi are ubiquitous and ecologically important in many grasslands. Differences in species responses to mycorrhizal colonization can have a significant influence on plant community structure. The growth responses of 36 species of warm- and cool-season tallgrass prairie grasses and 59 tallgrass prairie forbs to arbuscular mycorrhizal (AM) fungal colonization were assessed in greenhouse studies to examine the extent of interspecific variation in host-plant benefit from the symbiosis and patterns of mycorrhizal dependence among host plant life history (e.g., annual, perennial) and taxonomic (e.g., grass, forb, legume, nonlegume) groups and phenological guilds. There was a strong and significant relationship between phenology of prairie grasses and mycorrhizal responsiveness, however this relationship was less apparent in forbs. Perennial warm-season C(4) grasses and forbs generally benefited significantly from the mycorrhizal symbiosis, whereas biomass production of the cool-season C(3) grasses was not affected. The root systems of the cool-season grasses were also less highly colonized by the AM fungi, as compared to the warm-season grasses or forbs. Unlike the native perennials, annuals were generally not responsive to mycorrhizal colonization and were lower in percentage root colonization than the perennial species. Plant growth responsiveness and AM root colonization were positively correlated for the nonleguminous species, with this relationship being strongest for the cool-season grasses. In contrast, root colonization of prairie legumes showed a significant, but negative, relationship to mycorrhizal growth responsiveness.     

亚热带天然常绿阔叶林林下9种灌木细根形态和C、N化学计量特征

林下灌木是亚热带常绿阔叶林重要的构成部分,但林下灌木细根功能性状变异规律及地下生态策略仍不清楚。以福建建瓯万木林自然保护区内9种灌木为研究对象,对细根直径、根长、比根长、组织密度、碳浓度和氮浓度6个细根性状进行研究,采用序级划分法,分析不同树种细根性状序级间的变化特征、常绿和落叶灌木细根性状之间的差异,不同序级细根性状之间的关系以及细根性状变异维度。结果表明:树种和序级对9种灌木细根形态和化学性质有显著影响。直径、根长、根组织密度随着序级的增加而逐渐增加,比根长和氮浓度逐渐减小,碳浓度在序级间的变化趋势不一,未表现出明显的规律。落叶灌木细根直径、根长和氮浓度均显著高于常绿灌木,碳浓度和组织密度显著低于常绿灌木,表明与常绿灌木相比落叶灌木更偏向于资源获取型生态策略,常绿灌木则更偏向于保守型策略。灌木细根在不同序级间的直径与比根长、组织密度,氮浓度与组织密度有较强的相关性,细根其他性状间的关系并不密切或因序级而异。主成分分析结果表明灌木细根性状变异沿一个主成分轴发生变异,该轴表示灌木细根的资源获取和保守的权衡策略。     

华北山地6个外生菌根树种的根属性及其与菌根侵染率的关系

该研究以共存于同一暖温带森林的6个外生菌根(ECM)树种为研究对象,测定分析不同根序(1～5级)和功能根系(吸收细根和运输细根)的主要形态和构型属性及ECM侵染率,探究不同外生菌根树种的根属性变异模式及其与菌根真菌侵染程度的关系。结果表明:(1)随着根序的增加,不同树种根直径和单根长度均增加,而比根长和根分支强度均降低;根属性在同一根序下均存在显著的种间差异,尤其是2个裸子植物(落叶松和油松)的根直径较其他4个被子植物大。(2)同一树种的所有根属性在吸收细根和运输细根之间均有显著差异;吸收细根和运输细根的根直径、比根长和根组织密度在树种间均存在显著差异,而其单根长度和根分支强度在树种间无显著差异。(3)ECM侵染率以落叶松最高,千金榆和白桦最低,且与根尖直径呈显著正相关关系,与根尖比根长呈显著负相关关系。研究发现,基于根序或者功能根系,根属性在种间的变异模式不完全一致,单根长度和根分支强度在两个功能根系中均没有表现出显著的种间差异;吸收细根的比根长和根分支强度的变异系数较大,对环境变化有较敏感的响应;古老树种的根直径相对较粗,对菌根真菌的依赖性更高。     

Fine root tradeoffs between nitrogen concentration and xylem vessel traits preclude unified whole‐plant resource strategies in Helianthus

Recent work suggests variation in plant growth strategies is governed by a tradeoff in resource acquisition and use, ranging from a rapid resource acquisition strategy to a resource‐conservative strategy. While evidence for this tradeoff has been found in leaves, knowledge of root trait strategies, and whether they reflect adaptive differentiation across environments, is limited. In the greenhouse, we investigated variation in fine root morphology (specific root length and tissue density), chemistry (nitrogen concentration and carbon:nitrogen), and anatomy (root cross‐sectional traits) in populations of 26 Helianthus species and sister Phoebanthus tenuifolius. We also compared root trait variation in this study with leaf trait variation previously reported in a parallel study of these populations. Root traits varied widely and exhibited little phylogenetic signal, suggesting high evolutionary lability. Specific root length and root tissue density were weakly negatively correlated, but neither was associated with root nitrogen, providing little support for a single axis of root trait covariation. Correlations between traits measured in the greenhouse and native site characteristics were generally weak, suggesting a variety of equally viable root trait combinations exist within and across environments. However, high root nitrogen was associated with lower xylem vessel number and cross‐sectional area, suggesting a tradeoff between nutrient investment and water transport capacity. This led to correlations between root and leaf traits that were not always consistent with an acquisition–conservation tradeoff at the whole‐plant level. Given that roots must balance acquisition of water and nutrients with functions like anchorage, exudation, and microbial symbioses, the varied evidence for root trait covariation likely reflects the complexity of interacting selection pressures belowground. Similarly, the lack of evidence for a single acquisition–conservation tradeoff at the whole‐plant level likely reflects the vastly different selection pressures shaping roots and leaves, and the resources they are optimized to obtain.     

Changes in arbuscular mycorrhizal associations and fine root traits in sites under different plant successional phases in southern Brazil

Fine root morphological traits and distribution, arbuscular mycorrhizal (AM) fungi, soil fertility, and nutrient concentration in fine root tissue were compared in sites under different successional phases: grass plants, secondary forest, and mature forest in Londrina county, Paraná state, southern Brazil. Soil cores were collected randomly at the 0-10- and 10-20-cm depths in three quadrants (50 m2) in each site. Plants from the different successional stages displayed high differences in fine root distribution, fine root traits, and mycorrhizal root colonization. There were increases in the concentration of nutrients both in soil and fine roots and decrease of bulk soil density along the succession. The fine root biomass and diameter increased with the succession progress. The total fine root length, specific root length, root hair length, and root hair incidence decreased with the succession advance. Similarly, the mycorrhizal root colonization and the density of AM fungi spores in the soil decreased along the succession. Mycorrhizal root colonization and spore density were positively correlated with fine root length, specific root length, root hair length, root hair incidence, and bulk density and negatively correlated with fine root diameter and concentration of some nutrients both in soil and root tissues. Nutrient concentration in root tissue and in soil was positively correlated with fine root diameter and negatively correlated with specific root length, root hair length, and root hair incidence. These results suggest different adaptation strategies of plant roots for soil exploration and mineral acquisition among the different successional stages. Early successional stages displayed plants with fine root morphology and AM fungi colonization to improve the root functional efficiencies for uptake of nutrients and faster soil resource exploration. Late successional stages displayed plants with fine root morphology and mycorrhizal symbiosis for both a lower rate of soil proliferation and soil exploration capacity to acquire nutrients.     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

常绿阔叶林外生和丛枝菌根树种细根形态和构型性状对氮添加的可塑性响应

以中亚热带常绿阔叶林外生菌根树种罗浮栲和丛枝菌根树种木荷为研究对象,采用根袋法进行野外原位氮添加试验,研究了细根形态性状(比根长、比表面积、组织密度、平均根直径)和构型性状(分枝数、分枝比、根长增长速率、根尖密度、分枝密度),分析不同菌根树种细根形态和构型性状对氮沉降的响应.结果表明:随序级增加,外生和丛枝菌根树种细根...     

Patterns in root traits of woody species hosting arbuscular and ectomycorrhizas: implications for the evolution of belowground strategies

Root traits vary enormously among plant species but we have little understanding of how this variation affects their functioning. Of central interest is how root traits are related to plant resource acquisition strategies from soil. We examined root traits of 33 woody species from northeastern US forests that form two of the most common types of mutualisms with fungi, arbuscular mycorrhizas (AM) and ectomycorrhizas (EM). We examined root trait distribution with respect to plant phylogeny, quantifying the phylogenetic signal (K statistic) in fine root morphology and architecture, and used phylogenetically independent contrasts (PICs) to test whether taxa forming different mycorrhizal associations had different root traits. We found a pattern of species forming roots with thinner diameters as species diversified across time. Given moderate phylogenetic signals (K = 0.44–0.68), we used PICs to examine traits variation among taxa forming AM or EM, revealing that hosts of AM were associated with lower branching intensity (r_PIC = −0.77) and thicker root diameter (r_PIC = −0.41). Because EM evolved relatively more recently and intermittently across plant phylogenies, significant differences in root traits and colonization between plants forming AM and EM imply linkages between the evolution of these biotic interactions and root traits and suggest a history of selection pressures, with trade-offs for supporting different types of associations. Finally, across plant hosts of both EM and AM, species with thinner root diameters and longer specific root length (SRL) had less colonization (r_PIC = 0.85, −0.87), suggesting constraints on colonization linked to the evolution of root morphology.     

拉杰沙希大学校园药用植物菌根研究

Forty different medicinal plants were investigated for arbuscular mycorrhizal association in the Rajshahi University Campus in Bangladesh. The results indicated that 35 different plants were infected by AM (arbuscular mycorrhizal) fungi as found by trypan blue staining procedure. The percentage of root colonization by AM fungi varied from 13.3% to 100%. Mangifera indica and Morus indica have maximum percentage of colonization (100%). The intensity of root colonization were abundant in the plants belonging to the families Anacardiaceae, Asclepiadaceae, Moraceae, Leguminosae and Apocynaceae whereas the intensity of colonization of crop roots were moderate and poor belonging to Gramineae and Leguminosae. The presence of greater number of spore in soil was always associated with the incidence of abundant mycelia. In plant roots the formation of spore and mycelia was restricted by low pH. Number of mycorrhizal fungus spores ranged between 35 to100 per 100g air dried soil in different family respective soils. The frequency of mycorrhizal fungus infection showed positive correlation with soil pH, moisture, water holding capacity, texture, total nitrogen, organic carbon, phosphorus, calcium, potassium, and magnesium. Especially phosphorus and nitrogen in the soil greatly influenced the plant root infection by AM fungi.     

Variation in root system traits among African semi‐arid savanna grasses: Implications for drought tolerance

In arid to semi‐arid grasslands and savannas, plant growth, population dynamics, and productivity are consistently and strongly limited by soil water and nutrient availability. Adaptive traits of the root systems of grasses in these ecosystems are crucial to their ability to cope with strong water and/or nutrient limitation and the increasing drought stress associated with ecosystem degradation or projected climate change. We studied 18 grass species in semi‐arid savanna of the Kalahari region of Botswana to quantify interspecific variation in three important root system traits including root system architecture, rhizosheath thickness and mycorrhizal colonization. Drought‐tolerant species and shorter‐lived species showed greater rhizosheath thickness and fine root development but lower mycorrhizal colonization compared to later successional climax grasses and those characteristic of wetter sites. In addition, there was a significant positive correlation between root fibrousness index and rhizosheath thickness among species and a weak negative correlation between root fibrousness index and mycorrhizal colonization. These patterns suggest that an extensive fine root system and rhizosheath development may be important complementary traits of grasses coping with drought conditions, the former aiding in the acquisition of water by the grass plant and the latter aiding in water uptake and retention, and reducing water loss in the rhizosphere. Within species, both rhizosheath development and mycorrhizal colonization were significantly greater in a wet year than in a year with below‐average precipitation. The observed patterns suggest that the primary benefit of rhizosheath development in African savanna grasses is improved drought tolerance and that it is a plastic trait that can be adjusted annually to changing environmental conditions. The functioning of mycorrhizal symbiosis is likely to be relatively more important in infertile savannas where nutrient limitation is higher relative to water limitation.     

FACILITATION OF SUCCESSION BY THE NONMYCOTROPHIC COLONIZER SALSOLA KALI (CHENOPODIACEAE) ON A HARSH SITE: EFFECTS OF MYCORRHIZAL FUNGI

The effects of inoculation with vesicular-arbuscular mycorrhizal fungi on nonmycotrophic (plants which do not form a mycorrhizal association) colonizing annuals and on mycotrophic perennials were investigated in southwestern Wyoming. A subsoil containing no initial mycorrhizal inoculum was used, and seeds of the later successional perennials were planted. The annuals were removed from one-half of the sampling quadrats to test for the interaction of competition and mycorrhizae on the perennials. During the third and fourth years of succession, the density and percent cover of Salsola kali, the most abundant nonmycotrophic annual, decreased by one-half to one-third with inoculation. Mycorrhizal hyphae and spores were found in the rhizosphere of S. kali, with only occasional (1–2% of the root length) penetration of hyphae into the cortex. There were no differences in tissue phosphorus and nitrogen concentrations or water relations of inoculated vs. uninoculated S. kali. The planted grasses, all Agropyron species, had no significant increase in density or percent cover with inoculation. The percent root infection of A. smithii was 5–30%. Grass density and percent cover was greatest where S. kali was present, suggesting facilitation of grass establishment by annuals on this harsh, windy site. However, grass density was lower where 5. kali density was lower following inoculation. Mycorrhizal fungi were hypothesized to hasten the rate of succession on other sites. On this site where facilitation is an important process, inoculation of early seral plants and their subsequent decline may slow primary succession in the early years.     

Linking root respiration to chemistry and morphology across species

Root respiration is a critical physiological trait involved in root resource acquisition strategies, yet it is less represented in root trait syndrome. Here we compiled a large dataset of root respiration associated with root chemical and morphological traits from 245 plant species. Our results demonstrated that root respiration correlated positively with root nitrogen concentration (RNC) and negatively with root tissue density (RTD) across and within woody and non‐woody species. However, the relationships between root respiration and specific root length (SRL) and root diameter (RD) were weak or even insignificant. Such root respiration–traits relationships were not completely in line with predictions by the root economics spectrum (RES). Furthermore, the principal component analysis showed that root trait syndrome was multidimensional. Root respiration was associated more strongly with the RNC‐RTD axis (the classical RES) than with the orthogonal SRL‐RD axis for woody species, but not for non‐woody species. Collectively, the linkages of root physiological, chemical, and morphological traits provide a better understanding of root trait covariation and root resource acquisition strategies.     

How do steppe plants follow their optimal environmental conditions or persist under suboptimal conditions? The differing strategies of annuals and perennials

For a species to be able to respond to environmental change, it must either succeed in following its optimal environmental conditions or in persisting under suboptimal conditions, but we know very little about what controls these capacities. We parameterized species distribution models (SDMs) for 135 plant species from the Algerian steppes. We interpreted low false‐positive rates as reflecting a high capacity to follow optimal environmental conditions and high false‐negative rates as a high capacity to persist under suboptimal environmental conditions. We also measured functional traits in the field and built a unique plant trait database for the North‐African steppe. For both perennial and annual species, we explored how these two capacities can be explained by species traits and whether relevant trait values reflect species strategies or biases in SDMs. We found low false‐positive rates in species with small seeds, flowers attracting specialist pollinators, and specialized distributions (among annuals and perennials), low root:shoot ratios, wide root‐systems, and large leaves (perennials only) (R² = .52–58). We found high false‐negative rates in species with marginal environmental distribution (among annuals and perennials), small seeds, relatively deep roots, and specialized distributions (annuals) or large leaves, wide root‐systems, and monocarpic life cycle (perennials) (R² = .38 for annuals and 0.65 for perennials). Overall, relevant traits are rarely indicative of the possible biases of SDMs, but rather reflect the species' reproductive strategy, dispersal ability, stress tolerance, and pollination strategies. Our results suggest that wide undirected dispersal in annual species and efficient resource acquisition in perennial species favor both capacities, whereas short life spans in perennial species favor persistence in suboptimal environmental conditions and flowers attracting specialist pollinators in perennial and annual species favor following optimal environmental conditions. Species that neither follow nor persist will be at risk under future environmental change.     

Genetic variation in plant below-ground response to elevated CO2 and two herbivore species

Aims

It is unclear how changing atmospheric conditions, including rising carbon dioxide concentration, influence interactions between above and below-ground systems and if intraspecific variation exists in this response.

Methods

We assessed interactive effects of atmospheric CO₂ concentration, above-ground herbivory, and plant genotype on root traits and mycorrhizal associations. Plants from five families of Asclepias syriaca, a perennial forb, were grown under ambient and elevated atmospheric CO₂ concentrations. Foliar herbivory by either lepidopteran caterpillars or phloem-feeding aphids was imposed. Mycorrhizal colonization, below-ground biomass, root biomass, and secondary defensive chemistry in roots were quantified.

Results

We observed substantial genetic variation among A. syriaca families in their mycorrhizal colonization levels in response to elevated CO₂ and herbivory treatments. Elevated CO₂ treatment increased root biomass in all genetic families, whereas foliar herbivory tended to decrease root biomass. Root cardenolide concentration and composition varied greatly among plant families, and elevated CO₂ treatment increased root cardenolides in two of the five plant families. Moreover, herbivores differentially affected the composition of cardenolides expressed below ground.

Conclusions

Increased atmospheric CO₂ has the potential to influence interactions among plants, herbivores and mycorrhizal fungi and intraspecific variation suggests that such interactions can evolve.     

模拟氮沉降和不同外生菌根真菌侵染对樟子松幼苗生长的影响

盆栽条件下,采用单接种Suillus bovines(以下简称SB)和混合接种(Suillus luteus、Suillus grevillea、Tricholoma fulvum、Boletinus grisellus、Suillus bovinus、Leucocortinarius bulbiger、Rhizopogon luteolus、Pisolithus tinctorius 8个菌种的等量混合菌剂接种,以下简称HJ)方法对樟子松(Pinus sylvestris var.mongolica)2个月幼苗进行接种处理,幼苗生长6个月形成菌根后对樟子松幼苗进行模拟氮沉降实验,5个月后通过测定幼苗生长指标、根系形态结构和菌根侵染率,旨在探讨模拟氮沉降对樟子松不同接种处理幼苗生长的影响。结果表明:①随着氮浓度的增加,单接种乳牛肝(SB)和混合接种(HJ)菌根侵染率均呈下降趋势,且氮沉降对SB处理的菌根侵染率影响更显著。②接种显著促进幼苗地上和地下生物量积累,且混合接菌效果更好;同一接菌处理中,氮浓度增加对地上生物量的积累有抑制作用;接种和氮浓度对地径和地上生物量有显著交互作用,但对苗高和地下生物量无交互作用。③HJ处理中不施氮(CKN)和1倍氮浓度(CN)较不接菌(CK)和SB处理显著促进幼苗根系的平均直径、总根长和总表面积;氮浓度增加显著促进CK和SB处理的幼苗根系平均直径、总根长和总表面积,但是对HJ处理的根系各形态结构指标均产生抑制作用。接种和氮浓度以及两者的交互作用均极显著影响幼苗根系形态结构。     

Arbuscular mycorrhizas in cycads of southern India

Root and soil samples of three potted or ground-grown cycads ( Cycas circinalis, C. revoluta, Zamiasp.) were collected between November 1999 and June 2000 and surveyed for arbuscular mycorrhizal (AM) colonization and spore populations. AM fungi were associated with all root systems and rhizosphere samples examined. Root colonization was of a typical Arum type and AM colonization levels differed significantly between species and between potted and ground-grown cycads. Mycorrhizal colonization levels were inversely related to root hair number and length. Spores of nine morphotypes belonging to three genera ( Acaulospora, Glomus, Scutellospora) were extracted from soil. The percentage root length colonized by AM fungi was not related to soil factors, but total AM fungal spore numbers in the rhizosphere soil were inversely related to soil nitrogen and phosphorus levels. AM fungal spore numbers in the soil were linearly related to root length colonized. The co-occurrence of septate non-mycorrhizal fungi was recorded for the first time in cycads. These observations and the relationship between plant mycorrhizal status and soil nutrients are discussed.     

氮有效性增加对细根解剖、形态特征和菌根侵染的影响

氮(N)有效性增加对森林生态系统结构和功能有重要影响.细根作为树木地下最为活跃的组分,其对N有效性增加的响应已成为森林生态学研究的热点.本文对N有效性增加条件下细根解剖构造、形态和菌根侵染的响应趋势及潜在机制进行了综述.N有效性增加导致细根皮层厚度、皮层层数和皮层细胞直径下降,外皮层上通道细胞数量减少,而中柱和木质部横截面积,以及导管(或管胞)直径、数量和壁厚度均增加,根解剖构造的响应与植物激素浓度变化有关;N有效性增加后菌根侵染率下降,比根长(单位根干质量的长度)在外生菌根树种中普遍下降,而在内生菌根树种中增加;根直径和组织密度的变化趋势则表现出较大的种间差异.这些个体根和树种水平上的结果对于理解森林生态系统水平上的碳和养分循环具有重要意义.最后,根据目前研究中存在的不足提出了今后的研究方向与问题.     

青藏高原东部高山植物的水分利用效率与氮素利用效率研究

对生长在青藏高原东部隶属于23科、49属的71种高山植物(包括多年生和一年生植物)的稳定碳同位素比值、氮含量以及碳/氮比率进行了分析,并以稳定碳同位素比值及碳/氮比率来分别指示植物的水分利用效率和氮素利用效率.结果表明:(1)多年生植物稳定碳同位素比值显著高于一生年植物,而碳/氮比率显著低于一年生植物(P<0.01),氮含量两者无显著性差异.(2)多年生植物和一年生植物的稳定碳同位素比值均与碳/氮比率呈显著负相关(-0.643**和-0.707),而与氮含量均无明显相关性.研究证实,在自然条件下多年生植物的水分利用效率比一年生植物更高,而氮素利用效率却更低;高山植物水分利用效率和氮素利用效率存在明显的权衡",即植物不能同时提高水分利用效率和氮素利用效率,高水分利用效率的代价是降低氮素利用效率,青藏高原不同植物即使在相同环境条件下具有不同适应对策.