Correlations between carbon isotope ratio and microhabitat in desert plants

Summary Water is usually considered to be the key limiting factor for growth of desert plants, yet there is little information available of the water-use efficiency of species within a desert community. Leaf carbon isotope ratios, an indicator of long-term intercellular carbon dioxide concentrations and thus of water-use efficiency in C₃ plants, were measured on species occurring within a Sonoran Desert community, consisting of wash, transition and slope microhabitats. Along a soil moisture gradient from the relatively wetter wash to the relatively drier slope, leaf carbon isotope ratios increased in all species, indicating that water-use efficiency increased as soil water availability decreased. Leaf carbon isotope ratios of long-lived perennials were substantially more positive than in short-lived perennials, even though plants were growing adjacent to each other. Leaf carbon isotope ratio and leaf duration (evergreen versus deciduous) were not correlated with each other. The results are discussed in terms of how the efficiency of water use may affect community structure and composition.     

基于盐分梯度的荒漠植物多样性与群落、种间联接响应

土壤盐分是影响干旱区荒漠植物群落动态的决定因素之一。基于样方调查和不同土壤盐分梯度下植物多样性指数及群落与种间关联的计算结果,分析干旱区荒漠群落植物多样性、群落联接性和种间关联对土壤盐分梯度的响应动态。结果表明,在土壤盐含量为0.03%-0.55% (S1)、0.61%-1.24% (S2)和1.41%-1.79% (S3)的盐分梯度上,(1)随土壤盐含量升高,群落生活型结构改变,草本比例减少,乔木比例增加;(2)植物多样性指数随土壤盐分增加而下降,低盐分梯度下,二者极显著正相关(P<0.01),中盐梯度下二者间呈部分显著负相关(P<0.05),高盐梯度下则转为以正相关为主(P>0.05);(3)群落联接性随土壤盐分梯度转变,在0.61%-1.24%的中度盐含量时正联接性最强(VR=1.89),高盐度含量下群落转为不显著的负联接,稳定性降低;(4)沿盐分升高梯度,种间的负相关种对数增加,正相关种对数减少,种间关联(相关)强度提高,正负相关种对比(PNR)与多样性指数呈显著正相关(P<0.05)。综上可知,干旱区荒漠植物多样性在土壤盐含量达到1.41%-1.79%水平时总体显著降低;土壤盐分水平显著影响植物群落和种间联接性,种间互利性随盐分梯度增加下降,物种趋于独立分布,并最终导致荒漠植物多样性降低。     

荒漠植物功能性状及其多样性对土壤水盐变化的响应

植物功能性状及其多样性对环境变化的响应研究有助于揭示极端环境下植物适应策略和群落构建机制。通过实地调查和实验分析,研究艾比湖荒漠植物形态、生理和化学等8个功能性状的特征,并从多维性状和一维性状角度揭示功能多样性对土壤水分和盐分变化的响应规律。结果表明:高水盐环境下(SW1),(1)群落加权株高、叶绿素含量(SPAD)及叶片碳(C)和钙(Ca)含量显著高于低水盐环境,叶片氮(N)、磷(P)、硫(S)含量在不同水盐环境间无显著差异。(2)SW1环境下,沿乔木-小乔木-灌木-草本层次,上层生活型植物性状值普遍高于下层植物,其中灌木叶片仅N、钾(K)含量显著高于草本;沿该生活型层次植物性状呈趋同变化。(3)低水盐环境下(SW2),乔木叶片性状差异特征与SW1相似;小乔木叶片C、N含量分别显著高于和低于灌木及草本;相比于草本,灌木SPAD、S含量显著高,K含量显著低,株高、C、N、P含量差异不显著;SW2环境下各生活型植物性状呈趋同变化。(4)SW1环境下多维功能丰富度、功能离散度显著高于SW2环境,但均匀度无显著差异。(5)一维功能均匀度在不同水盐环境间均无显著差异,但化学性状的均匀度总体高于植物株高;高水盐环境下叶片N、S和Ca的功能分异指数显著高于低水盐环境。研究为掌握胁迫环境下的植物适应策略和荒漠植被恢复提供参考。     

水盐梯度下荒漠植物多样性格局及其与土壤环境的关系

植物多样性格局作为生物多样性维持的一个重要方面可揭示群落构建的信息,反映物种对环境的适应性。结合样带调查和实验分析的方法,研究新疆艾比湖湿地自然保护区内水盐梯度下荒漠植物多样性格局及其与土壤环境因子间关系。结果表明:(1)高、低水盐样地(SW1,SW2)土壤容积含水量(SVWC)和电导率(EC)分别为16.65%和12.02 m S/cm及2.63%和1.91m S/cm,水盐变化主要影响草本和部分灌木群落的植物组成;(2)高水盐生境下植物群落无明显趋势性分布,低水盐生境下植物种依水盐和营养元素呈区域性分布。(3)高水盐生境下荒漠植物的多度分布呈对数正态模型,低水盐条件下符合Zipf模型,多度分布的变化反映了群落组成结构的水盐梯度响应;(4)SW1样地多样性总体显著高于SW2样地,土壤p H、SVWC和硫(S)对植物多样性有不同程度的极显著影响。综上可知,荒漠植物分布及多样性格局与土壤环境间关系呈一定的水盐梯度响应规律,本研究可为该地区植被恢复和土壤盐渍化治理提供科学依据。     

Stable carbon isotope characteristics of desert plants in the Junggar Basin, China

Desert plants have unique strategies for survival and growth to cope with the limited water availability in arid regions. The stable carbon isotope (δ ¹³C) provides an integrated measurement of internal plant physiological and external environmental properties affecting photosynthetic gas exchange and water use efficiency. The δ ¹³C values of 84 species in the Junggar Basin were categorized into two groups (ranged from −30.1 to −23.3‰ for C₃ and −14.9 to −9.9‰ for C₄ species, respectively). No life form differences in δ ¹³C values were detected in C₃ (p = 0.78) and C₄ plants (p = 0.63). Small differences among life forms were observed in δ ¹³C values in C₄ species with shrubs slightly depleted (−13.3‰) relative to perennials (−13.1‰) and annuals (−12.5‰). These differences suggested that δ ¹³C value could not represent a plant functional group classification based on life forms in C₄ plants in extremely arid regions. Ephemerals are all using C₃ photosynthetic pathway and no significant differences (p = 0.92) in δ ¹³C values were observed between annuals (−26.5‰) and perennials (−26.4‰). The δ ¹³C values of Tulipa iliensis (an important ephemeral species distributed widely in the Junggar Basin) among nine natural populations were positively correlated with leaf (r ² = 0.46, p = 0.046) and soil (r ² = 0.67, p = 0.007) total nitrogen content, and negatively correlated with leaf (r ² = 0.48, p = 0.039) and soil (r ² = 0.79, p = 0.001) water content. This indicated that the variation in δ ¹³C values of T. iliensis was probably caused by both water availability associated stomatal openness and nitrogen availability associated photosynthetic capacity. T. iliensis is very sensitive to water and nitrogen availability in soil.     

荒漠土壤因子和DSE定殖对克隆植物入侵的响应

克隆植物,尤其是游击型克隆植物,具有很强的扩展能力,通过克隆扩展可侵入到不同生境斑块。克隆植物入侵可能会影响入侵地土壤营养状况和微生物群落。为了探明克隆植物入侵对DSE(dark septate endophytes)活动和土壤理化性质的影响,于2013年6月在克隆植物羊柴(Hedysarum laeve)和沙鞭(Psammochloa villosa)群落空地沿根状茎延伸方向设置样方,分别于6月、8月和10月在样方内分0—10、10—20、20—30、30—40、40—50 cm土层采集土样和根样,研究了不同采样时间羊柴和沙鞭群落空地DSE和土壤理化性质时空变化。结果表明,从6月到10月,随时间后延,克隆植物逐渐侵入群落空地,沙鞭入侵群落空地数和分株数高于羊柴。羊柴群落空地根系DSE定殖率随采样时间后延,逐渐降低,最大值在6月;沙鞭群落空地根系DSE定殖率随采样时间后延,逐渐升高,最大值在10月。随着克隆植物入侵,入侵地土壤中可利用的营养物质含量显著提高,羊柴入侵提高了入侵群落空地土壤碱解N、有效P和速效K含量,沙鞭入侵提高了入侵群落空地土壤碱解N和有效P含量。相关性分析表明,羊柴群落空地DSE定殖率与土壤p H值和电导率显著正相关,沙鞭群落空地DSE定殖率与土壤p H值极显著负相关,与电导率、碱解N和有效P极显著正相关。克隆植物入侵使得土壤环境更有利于克隆植物自身生长,为荒漠植被恢复提供了前提。     

Water use efficiency and carbon isotope composition of plants in a cold desert environment

Summary The effects of the availabilities of water and nitrogen on water use efficiency (WUE) of plants were investigated in a sagebrush steppe. The four species studied wereArtemisia tridentata (shrub),Ceratoides lanata (suffrutescent shrub),Elymus lanceolatus (rhizomatous grass), andElymus elymoides (tussock grass). Water and nitrogen levels were manipulated in a two-by-two factorial design resulting in four treatments: control (no additions), added water, added nitrogen, and added water and nitrogen. One instantaneous and two long-term indicators of WUE were used to testa priori predictions of the ranking of WUE among treatments. The short-term indicator was the instantaneous ratio of assimilation to transpiration (A/E). The long-term measures were 1) the slope of the relationship between conductance to water vapor and maximum assimilation and 2) the carbon isotope composition (¹³C) of plant material. Additional water decreased WUE, whereas additional nitrogen increased WUE. For both A/E and ¹³C, the mean for added nitrogen alone was significantly greater than the mean for added water alone, and means for the control and added water and nitrogen fell in between. This ranking of WUE supported the hypothesis that both water and nitrogen limit plant gas exchange in this semiarid environment. The short- and long-term indicators were in agreement, providing evidence in support of theoretical models concerning the water cost of carbon assimilation.     

Leaf-root-soil N:P stoichiometry of ephemeral plants in a temperate desert in Central Asia

Ephemeral plants are a crucial vegetation component in temperate deserts of Central Asia, and play an important role in biogeochemical cycle and biodiversity maintenance in desert ecosystems. However, the nitrogen (N) and phosphorus (P) status and interrelations of leaf-root-soil of ephemeral plants remain unclear. A total of 194 leaf-root-soil samples of eight ephemeral species at 37 sites in the Gurbantunggut Desert, China were collected, and then the corresponding N and P concentrations, and the N:P ratio were measured. Results showed that soil parameters presented no significant difference among the eight species. The total soil N:P was only 0.116 (geomean), indicating limited soil N, while the available soil N:P (4.896, geomean) was significantly larger than the total N:P. The leaf N (averagely 30.995 mg g^?1) and P (averagely 1.523 mg g^?1) concentrations were 2.64–8.46 and 0.93–3.99 times higher than the root N (averagely 8.014 mg g^?1) and P (averagely 0.802 mg g^?1) concentrations, respectively. Thus, leaf N:P (averagely 21.499) was 1.410–2.957 times higher than root N:P (averagely 11.803). Meanwhile, significant interspecific differences existed in plant stoichiometric traits. At the across-species level, N content scaled as the 3/4-power of P content in both leaves and roots. Leaf and root N:P ratios were mainly influenced by P; however, the leaf-to-root N or P ratio was dominated by roots. Leaf and root N, P contents and N:P were generally unrelated to soil nutrients, and the former presented lower variation than the latter, indicating a strong stoichiometric homeostasis for ephemerals. These results demonstrate that regardless of soil nutrient supply capacity in this region, the fast-growing ephemeral plants have formed a specific leaf-root-soil stoichiometric relation and nutrient use strategy adapting to the extreme desert environment.

     

Stable isotope composition of water in desert plants

A survey of the stable isotope content of tissue waters of plants from the Negev desert was conducted. Large differences were observed in the extent of enrichment of the heavy isotopes in leaf water relative to local precipitation among different plants. This is apparently caused by the species-dependent stratagems adopted by the plants to cope with water stress, primarily by differences in the depth of water uptake in the soil and through the timing of stomatal openings during the daily cycle. Salt stressed plants showed extreme variability in the isotopic composition of leaf–water. The results show that plants with adaptation to arid conditions can avoid the transpiration regime, which would lead to the strong isotopic enrichment in their leaf water expected under arid conditions. This has implications for the use of stable isotopes in plants as indicators of either plant ecophysiology or paleoclimate. Responsible Editor: Hans Lambers. G. Goodfriend is deceased.     

Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves

Cotton (Gossypium hirsutum L. cv. CS50) plants were grown at two levels of relative humidity (RH) and sprayed daily with abscisic acid (ABA) at four concentrations. Plants grown at lower humidity had higher transpiration rates, lower leaf temperatures and lower stomatal conductance. Plant biomass was also reduced at low humidity. Within each humidity environment, increasing ABA concentration generally reduced stomatal conductance, evaporation rates, superficial leaf density and plant biomass, and increased leaf temperature and specific leaf area. As expected, decreased stomatal conductance resulted in decreased carbon isotope discrimination in leaf material ( Δ ¹³C_l). Plants grown at low humidity were more enriched in ¹⁸O than those grown at high RH, as theory predicts. Within each humidity environment, increasing ABA concentration increased oxygen isotope enrichment of leaf cellulose ( Δ ¹⁸O_c) and whole‐leaf tissue ( Δ ¹⁸O_l). Values of Δ ¹³C_l and Δ ¹⁸O_l predicted by theoretical models were close to those observed, accounting for 79% of the measured variation in Δ ¹³C_l and 95% of the measured variation in Δ ¹⁸O_l. Supporting theory, Δ ¹³C_l and Δ ¹⁸O_l in whole‐leaf tissue were negatively related.     

C3植物稳定碳同位素组成与盐分的关系

植物在盐生环境中δ13C值的改变可能包含两个成分:一个是盐分对CO2的扩散、传递或光合速率的影响而引起的δ13C值的改变;另一个是光合途径的转换引起的δ13C值的变化,δ13C值的大小与诱导发生CAM或C4代谢的程度有关.植物组织的δ13C值随盐度的变化趋势除了与植物本身固有的耐盐性有关以外,盐度和胁迫时间是影响植物δ13C的重要因素.根据盐生条件下同位素分馏特点可知,盐生植物与非盐生植物的δ13C随盐度的变化趋势有所不同.对非盐生植物而言,在低盐度和短期的盐处理下,随盐度的增加和胁迫时间的延长植物的δ13C值增大,这个阶段限制光合作用的主要因素是气孔导度;但是如果盐度过低,δ13C变化很小,则难以表现出应有的相关性;随着胁迫的加强,当限制光合作用的非气孔因素成为主导因素时,由于光合作用受到强烈抑制(光合结构遭到破坏),δ13C将随之降低.对盐生植物而言,其δ13C与最适盐度有关.最适盐度下,植物的δ13C低于其它盐度条件下的δ13C值.盐生条件下,有些C3植物可能发生光合途径的转换,无论诱导发生的是C4代谢还是CAM代谢,δ13C值均趋于增大.但是,一般情况下,盐处理诱导的光合途径的改变对植物组织整体的δ13C的影响很小.在密闭环境中或郁闭林地,植物和土壤呼吸释放的CO2再次参与光合作用,也会改变植物的δ13C值.为了更加全面地考察植物δ13C与盐度的关系,需要设置较大的盐度范围和进行长期的胁迫处理,才能够获得相对充分的数据,才有利于全面分析植物δ13C值与耐盐性的关系.     

Temporal variation in spatial organization of a rodent community in the southwestern Kyzylkum desert (Middle Asia)

The distribution of niches in resource space and the niche patterns of a 14-species community of Middle Asian desert rodents were studied during two years - at low and high rodent density - using discriminant function analysis Nineteen quantitative environmental parameters (soil structure and vegetation), measured in 550 plots within 22 1 -ha grids, were considered The first three canonical axes of resource space account for 72% of the variance The first two axes represent complex environmental gradients the first axis represents a general landscape gradient from sand to clay soils, the second axis reflects a gradient of in creasing productivity The third axis reflects with in-habitat environmental variation All community parameters, as well as parameters of individual species niches, were unstable between years At the same time, different parameters vary in different extent Position of niche centroids along macro-habitat axes, as. well as macrohabitat niche breadth, were relatively stable between years, but these parameters for microhabitat axis and values of niche overlap were much more variable A strong correlation between changes m relative between-habitat niche breadth and differences in average niche overlap with relative changes in species abundances indicate density dependence of these parameters Changes in niche overlap is a consequence of between-year differences in guild patterns Guild structure was pronounced at high density when the level of niche overlap was intermediate At low density, when the level of niche overlap decreased, guild structure was incon-spicous Different levels of diversity differed in their sensitivity to density changes α-diversity was relatively constant as a result of between-year stability of niche centroid positions However level of ß-diversity varied significantly between years reflecting changes in the level of niche overlap, because a decrease in niche overlap leads to an increase in the rate of species turnover     

Non-climatic variations in the oxygen isotopic compositions of plants

The ¹⁸O content of leaf water strongly influences the ¹⁸O contents of atmospheric CO₂ and O₂. The ¹⁸O signatures of these atmospheric gases, in turn, emerge as important indicators of large-scale gas exchange processes. Better understanding of the factors that influence the isotopic composition of leaf water is still required, however, for the quantitative utilization of these tracers. The ¹⁸O enrichment of leaf water relative to local meteoric water, is known to reflect climatic conditions. Less is known about the extent variations in the ¹⁸O content of leaf water are influenced by nonclimatic, species-specific characteristics. In a collection of 90 plant species from all continents grown under the same climatic conditions in the Jerusalem Botanical Garden we observed variations of about 9‰ in the δ¹⁸O values of stem water, δ_s, and of about 14‰ in the mid-day δ¹⁸O enrichment of bulk leaf water, δ_LW–δ_s. Differences between δ¹⁸O values predicted by a conventional evaporation model, δ_M, and δ_LW ranged between – 3.3‰ and + 11.8‰. The δ¹⁸O values of water in the chloroplasts (δ_ch) in leaves of 10 selected plants were estimated from on-line CO₂ discrimination measurements. Although much uncertainty is still involved in these estimates, the results indicated that δ_ch can significantly deviate from δ_M in species with high leaf peclet number. The δ¹⁸O values of bulk leaf water significantly correlated with δ¹⁸O values of leaf cellulose (directly) and with instantaneous water use efficiency (A/E, inversely). Differences in isotopic characteristics among conventionally defined vegetation types were not significant, except for conifers that significantly differed from shrubs in δ¹⁸O and δ¹³C values of cellulose and in their peclet numbers, and from deciduous woodland species in their δ¹⁸O and δ¹³C values of cellulose. The results indicated that predictions of the δ¹⁸O values of leaf water (δ_LW, δ_M and δ_ch) could be improved by considering plant species-specific characteristics.     

岩溶区不同植被下土壤水溶解无机碳含量及其稳定碳同位素组成特征

自2010年7月至2011年7月对重庆青木关岩溶区典型植被下的土壤水进行了月动态取样,分析了土壤水溶解无机碳含量(DIC浓度)及其稳定碳同位素组成(δ13CDIC值)的时空变化特征,以揭示岩溶土壤系统碳酸盐岩溶蚀作用及其碳汇效应。研究结果表明:草地和针叶林地土壤水的DIC浓度和δ13CDIC值相对较低,分别为59.12 mg/L和-17.22‰,31.47 mg/L和-16.37‰;而旱地、灌丛地、退耕还林地土壤水具有较高的DIC浓度和δ13CDIC值,分别达153.88 mg/L和-12.2‰,221.82 mg/L和-11.9‰,97.30 mg/L和-11.23‰,其中灌丛和退耕还林地的δ13CDIC值与DIC浓度呈正比,且雨季较旱季偏高约4‰—5‰。根据δ13CDIC值,结合各植被类型下土壤水DIC浓度与其相应的土壤碳酸盐含量呈正相关,判断旱地、灌丛地、退耕还林地等岩溶土壤水中的DIC主要来自土壤中碳酸盐岩矿物的碳酸溶蚀,即岩溶土壤中存在着碳酸盐岩碳酸溶蚀作用,从而在一定程度上减少了土壤系统向大气排放的CO2量。     

Stoichiometric variation of halophytes in response to changes in soil salinity

• Variation in soil salt may change the stoichiometry of a halophyte by altering plant ecophysiology, and exert different influences on various plant organs, which has potentially important consequences for the nutrition of consumers as well as nutrient cycling in a saline ecosystem.
• Using a greenhouse pot experiment, we investigated the effect of salinity variability on the growth and stoichiometry of different organs of Suaeda glauca and Salicornia europaea – two dominant species of important ecological and economic value in the saline ecosystem.
• Our results showed that appropriate salt stimulated the growth of both species during the vigorous growth period, while high salt suppressed growth. Na significantly increased with increased salt in the culture, whereas concentrations of other measured elements and K:Na ratio for both species significantly decreased at low salt treatments, and became more gradual under higher salt conditions. Furthermore, with the change of salt in culture, variations in leaf (degenerated leaf for S. europaea, considered as young stem) stoichiometry, except N:P ratio, were large and less in stems (old stems for S. europaea) than in roots, reflecting physiological and biochemical reactions in the leaf in response to salt stress, supported by sharp changes in trends.
• These results suggest that appropriate saline conditions can enhance biological C fixation of halophytes; however, increasing salt could affect consumer health and decrease cycling of other nutrients in saline ecosystems.

     

Responses of soil microorganisms to resource availability in urban, desert soils

Terrestrial desert ecosystems are strongly structured by the distribution of plants, which concentrate resources and create islands of fertility relative to interplant spaces. Atmospheric nitrogen (N) deposition resulting from urbanization has the potential to change those spatial patterns via resource inputs, resulting in more homogeneous soil resource availability. We sampled soils at 12 desert remnant sites around Phoenix, Arizona along a model-predicted gradient in N deposition to determine the degree to which deposition has altered spatial patterns in soil resource availability and microbial activity. Soil microbial biomass and abundance were not influenced by atmospheric N deposition. Instead, plant islands remained strong organizers of soil microbial processes. These islands of fertility exhibited elevated pools of resources, microbial abundance, and activity relative to interspaces. In both plant islands and interspaces, soil moisture and soil N concentrations predicted microbial biomass and abundance. Following experimental wetting, carbon dioxide (CO₂) flux from soil of interspaces was positively correlated with N deposition, whereas in plant islands, soil CO₂ flux was positively correlated with soil moisture content and soil organic matter. Soil CO₂ flux in both patch types showed rapid and short-lived responses to precipitation, demonstrating the brief time scales during which soil biota may process deposited materials. Although we observed patterns consistent with N limitation of microbes in interspaces, we conclude that atmospheric N deposition likely accumulates in soils because microbes are primarily limited by water and secondarily by carbon or nitrogen. Soil microbial uptake of atmospherically deposited N likely occurs only during sparse and infrequent rainfall.     

准噶尔盆地南缘荒漠区土壤碳分布及其稳定同位素变化

以亚洲中部干旱区准噶尔盆地南缘荒漠区为研究区,根据荒漠距离绿洲的距离,分别在荒漠边缘、中部和腹地设置3条样带,并采集2 m深的土壤剖面样品,研究土壤有机碳(SOC)、无机碳(SIC)含量及其稳定碳同位素的分布,探讨土壤碳变化与距绿洲距离的关系.结果表明: SOC含量随剖面土层深度增加而减少.受距绿洲距离的影响,SOC含量表现为荒漠边缘>荒漠中部>荒漠腹地.荒漠边缘SOC的δ¹³C值范围为-21.92‰～-17.41‰,且随深度增加而递减;荒漠中部和荒漠腹地的δ¹³C值范围为-25.20‰～-19.30‰,且随深度增加先增后减,由此推断准噶尔盆地南缘荒漠中部和腹地地表植被以C3植物为主,而绿洲边缘经历了从C3植物为主到C4植物为主的演替过程.荒漠边缘SIC平均含量为38.98 g·kg^-1,是荒漠腹地的6.01倍,表明0～2 m深度内大量SIC在荒漠边缘呈聚集趋势.SIC的δ¹³C值随深度增加先减后增,底层富集,主要受原生碳酸盐含量和剖面土壤CO₂的影响.     

碳输入方式对森林土壤碳库和碳循环的影响研究进展

凋落物和植物根系是森林土壤有机碳的主要来源.综述了不同碳输入方式对土壤全碳、微生物生物量碳和可溶性有机碳等碳库组分及土壤呼吸影响的研究进展.不同地区、不同森林土壤有机碳对碳输入的响应程度不同,且采用添加和去除凋落物,以及去除根系方法(DIRT)对土壤碳的影响具有树种差异和区域差异.目前主要侧重于土壤呼吸和碳库组分的研究,亟需开展对土壤碳的结构类型和稳定性,以及土壤生物尤其是土壤动物的响应机制的相关研究.