降水变化和氮沉降对荒漠草原两种多年生禾草凋落物分解的影响

凋落物分解是陆地生态系统养分循环的重要过程,在生物地球化学循环过程中发挥着重要作用。全球变化是影响凋落物分解的重要因子,其对生态系统养分循环的影响存在诸多不确定性。研究荒漠草原凋落物分解对氮沉降和降水变化及其二者交互作用的响应,是揭示这些不确定性、保护草原生态系统结构和功能的科学基础。以内蒙古四子王旗短花针茅荒漠草原为研究对象,选取建群种短花针茅和优势种无芒隐子草两种植物凋落物,开展为期4年的长期分解实验,探究两种植物凋落物分解特征及养分释放规律。实验采用裂区设计,主区为自然降水（C）、增雨30%（W）和减雨30%（R）3个水分梯度,副区为0（N0）、30（N30）、50（N50）和100（N100） kg hm^-2 a^-1 4个氮素梯度。结果表明：（1）增雨和氮沉降促进荒漠草原凋落物分解,减雨反之,降水对两种凋落物影响具有差异,而氮沉降的作用不依赖于物种;（2）氮沉降缩短凋落物分解周期5.12%-14.82%,增雨与氮沉降交互缩短凋落物分解周期3.69%-28.75%;（3）降水始终有利于凋落物中碳、纤维素和木质素释放,而分解后期氮沉降对其影响不显著,凋落物分解后期主要受木质素分解速率控制。综上所述,影响荒漠草原凋落物分解的主要因素为降水,其次是氮素,二者对凋落物分解具有协同作用。     

Plant responses to precipitation in desert ecosystems: integrating functional types,pulses, thresholds,and delays

The two-layer and pulse-reserve hypotheses were developed 30 years ago and continue to serve as the standard for many experiments and modeling studies that examine relationships between primary productivity and rainfall variability in aridlands. The two-layer hypothesis considers two important plant functional types (FTs) and predicts that woody and herbaceous plants are able to co-exist in savannas because they utilize water from different soil layers (or depths). The pulse-reserve model addresses the response of individual plants to precipitation and predicts that there are biologically important rain events that stimulate plant growth and reproduction. These pulses of precipitation may play a key role in long-term plant function and survival (as compared to seasonal or annual rainfall totals as per the two-layer model). In this paper, we re-evaluate these paradigms in terms of their generality, strengths, and limitations. We suggest that while seasonality and resource partitioning (key to the two-layer model) and biologically important precipitation events (key to the pulse-reserve model) are critical to understanding plant responses to precipitation in aridlands, both paradigms have significant limitations. Neither account for plasticity in rooting habits of woody plants, potential delayed responses of plants to rainfall, explicit precipitation thresholds, or vagaries in plant phenology. To address these limitations, we integrate the ideas of precipitation thresholds and plant delays, resource partitioning, and plant FT strategies into a simple threshold-delay model. The model contains six basic parameters that capture the nonlinear nature of plant responses to pulse precipitation. We review the literature within the context of our threshold-delay model to: (i) develop testable hypotheses about how different plant FTs respond to pulses; (ii) identify weaknesses in the current state-of-knowledge; and (iii) suggest future research directions that will provide insight into how the timing, frequency, and magnitude of rainfall in deserts affect plants, plant communities, and ecosystems.     

Differential responses to warming and increased precipitation among three contrasting grasshopper species

We conducted a field manipulation experiment to investigate developmental and demographic responses to warming and increased precipitation in three Inner Mongolian grasshopper species that differ in phenology (an early-season species Dasyhippus barbipes , a mid-season species Oedaleus asiaticus , and a late-season species Chorthippus fallax ). Infrared heaters were used for warming the ground surface by 1–2 °C above the ambient condition and periodic irrigations were applied to simulate a 50% increase in annual precipitation. We found that warming advanced the timing for egg hatching and grasshopper eclosion in each of the three species. However, grasshopper diapause and increased precipitation appeared to offset the effect of warming on egg development. Hatching and development were more strongly affected by warming in the mid-season O. asiaticus and the late-season C. fallax relative to the early-season D. barbipes . Warming by ∼1.5 °C advanced the occurrence of the mid-season O. asiaticus by an average of 4.96 days; while warming and increased precipitation interactively affected the occurrence of the late-season C. fallax , which advanced by 5.53 days. Our data and those of others suggest that most grasshopper species in the Inner Mongolian grassland are likely to extend their distribution northward with climate change. However, because of the differential response to warming we demonstrate for these species, the different grasshopper species are predicted to aggregate toward the middle period of the growing season, potentially increasing interspecific competition and grazing pressure on grasslands.     

Monosoonal precipitation responses of shrubs in a cold desert community on the Colorado Plateau

South-eastern Utah forms a northern border for the region currently influenced by the Arizona monosoonal system, which feeds moisture and summer precipitation into western North America. One major consequence predicted by global climate change scenarios is an intensification of monosoonal (summer) precipitation in the aridland areas of the western United States. We examined the capacity of dominant perennial shrubs in a Colorado Plateau cold desert ecosystem of southern Utah, United States, to use summer moisture inputs. We simulated increases of 25 and 50 mm summer rain events on Atriplex canescens, Artemisia filifolia, Chrysothamnus nauseosus, Coleogyne ramosissima, and Vanclevea stylosa, in July and September with an isotopically enriched water (enriched in deuterium but not ¹⁸O). The uptake of this artificial water source was estimated by analyzing hydrogen and oxygen isotope ratios of stem water. The predawn and midday xylem water potentials and foliar carbon isotope discrimination were measured to estimate changes in water status and water-use efficiency. At. canescens and Ch. nauseosus showed little if any uptake of summer rains in either July or September. The predawn and midday xylem water potentials for control and treatment plants of these two species were not significantly different from each other. For A. filifolia and V. stylosa, up to 50% of xylem water was from the simulated summer rain, but the predawn and midday xylem water potentials were not significantly affected by the additional summer moisture input. In contrast, C. ramosissima showed significant uptake of the simulated summer rain (>50% of xylem water was from the artificial summer rain) and an increase in both predawn and midday water potentials. The percent uptake of simulated summer rain was greater when those rains were applied in September than in July, implying that high soil temperature in midsummer may in some way inhibit water uptake. Foliar carbon isotope discrimination increased significantly in the three shrubs taking up simulated summer rain, but pre-treatment differences in the absolute discrimination values were maintained among species. The ecological implications of our results are discussed in terms of the dynamics of this desert community in response to changes in the frequency and dependability of summer rains that might be associated with a northward shift in the Arizona monsoon boundary.     

Aboveground net primary productivity and soil respiration display different responses to precipitation changes in desert grassland

荒漠草原区地上净初级生产力和土壤呼吸对降水变化的不同响应 降水变化既影响地上植被动态,也影响地下碳循环过程,尤其以干旱半干旱生态系统对降水的响应更为敏感。然而极端降水如何影响土壤碳固存潜力仍未得出明确结果。本研究在黄土高原西部荒漠草原样地实施了为期3年的降水控制实验,该实验包含5个降水梯度(即自然降水(对照),以及在自然降水基础上的减水40%、减水20%、增水20%、增水40%)。通过对不同降水处理下植物生长指标、地上净初级生产力(ANPP)、土壤水分和土壤呼吸(Rs)进行监测,采用双侧不对称模型揭示ANPP和Rs对降水变化的响应规律;采用结构方程模型,分析降水变化下影响ANPP和Rs的直接和间接因素。研究结果表明,ANPP对极端干旱的响应比极端湿润更敏感,在干旱和湿润年份均符合负向不对称模型。ANPP的变化主要受到降水的直接影响,同时,干旱年份植物密度的变化也对ANPP产生了影响。在湿润年份,Rs对降水变化的响应也呈负向不对称性。然而,干旱年份,Rs对降水变化表现出正向不对称响应,即对降水增加响应的敏感性高于降水减少,这可能与植物生长和ANPP对增水处理的正响应增加使自氧呼吸增强,及降水事件对异氧呼吸具有较强的‘Birch效应’有关。在干旱年份Rs对极端干旱(减水40%处理)表现出饱和响应。ANPP和Rs对降水格局改变的响应模式差异表明荒漠草原区极端湿润或干旱可能降低研究区土壤碳固存的潜力。     

Patterns of root architecture adaptation of a phreatophytic perennial desert plant in a hyperarid desert

Plant root systems can respond to nutrient availability and distribution by changing the three-dimensional deployment of their root architecture. The year after year variation of root architecture was investigated in a perennial phreatophyte in the controlled condition vegetation situated in the oasis in the Chinese Taklamakan desert with the goal to elucidate their adaptation to hyperarid environment. The whole plants of an indigenous perennial legume Alhagi sparsifolia Shap. with intact root systems were excavated at the end of each growing season from 2007 to 2009 and analyzed for architecture, above and belowground biomass, root/shoot ratio, root depth, seed yields and ramet. Changes in water availability were found to have stupendous effects on taproot depth, lateral root length and ramet quantity. Relative to shoot dry weight, taproot depth decreased with increasing water availability. In contrast, lateral root elongation was promoted by higher water availability.We tested the hypothesis that (1) irrigation increases root biomass and the quantity of ramets, and (2) A. sparsifolia Shap. develops an efficient root architecture that could absorb soil water and nutrition.     

Nitrogen fixation and metabolism by groundwater-dependent perennial plants in a hyperarid desert

The Central Asian Taklamakan desert is characterized by a hyperarid climate with less than 50 mm annual precipitation but a permanent shallow groundwater table. The perched groundwater (2–16 m) could present a reliable and constant source of nitrogen throughout the growing season and help overcome temporal nitrogen limitations that are common in arid environments. We investigated the importance of groundwater and nitrogen fixation in the nitrogen metabolism of desert plants by assessing the possible forms and availability of soil N and atmospheric N and the seasonal variation in concentration as well as isotopic composition of plant N. Water availability was experimentally modified in the desert foreland through simulated flooding to estimate the contribution of surface water and temporally increased soil moisture for nutrient uptake and plant–water relations. The natural vegetation of the Taklamakan desert is dominated by plants with high foliar nitrogen concentrations (2–3% DM) and leaf nitrate reductase activity (NRA) (0.2–1 mol NO₂^– g^–1 FW h^–1). There is little evidence that nitrogen is a limiting resource as all perennial plants exhibited fast rates of growth. The extremely dry soil conditions preclude all but minor contributions of soil N to total plant N so that groundwater is suggested as the dominant source of N with concentrations of 100 M NO₃^–. Flood irrigation had little beneficial effect on nitrogen metabolism and growth, further confirming the dependence on groundwater. Nitrogen fixation was determined by the ¹⁵N natural abundance method and was a significant component of the N-requirement of the legume Alhagi, the average contribution of biologically fixed nitrogen in Alhagi was 54.8%. But nitrogen fixing plants had little ecological advantage owing to the more or less constant supply of N available from groundwater. From our data we conclude that the perennial species investigated have adapted to the environmental conditions through development of root systems that access groundwater to satisfy demands for both water and nutrients. This is an ecologically favourable strategy since only groundwater is a predictable and stable resource.     

Soil microbial responses to warming and increased precipitation and their implications for ecosystem C cycling

A better understanding of soil microbial ecology is critical to gaining an understanding of terrestrial carbon (C) cycle–climate change feedbacks. However, current knowledge limits our ability to predict microbial community dynamics in the face of multiple global change drivers and their implications for respiratory loss of soil carbon. Whether microorganisms will acclimate to climate warming and ameliorate predicted respiratory C losses is still debated. It also remains unclear how precipitation, another important climate change driver, will interact with warming to affect microorganisms and their regulation of respiratory C loss. We explore the dynamics of microorganisms and their contributions to respiratory C loss using a 4-year (2006–2009) field experiment in a semi-arid grassland with increased temperature and precipitation in a full factorial design. We found no response of mass-specific (per unit microbial biomass C) heterotrophic respiration to warming, suggesting that respiratory C loss is directly from microbial growth rather than total physiological respiratory responses to warming. Increased precipitation did stimulate both microbial biomass and mass-specific respiration, both of which make large contributions to respiratory loss of soil carbon. Taken together, these results suggest that, in semi-arid grasslands, soil moisture and related substrate availability may inhibit physiological respiratory responses to warming (where soil moisture was significantly lower), while they are not inhibited under elevated precipitation. Although we found no total physiological response to warming, warming increased bacterial C utilization (measured by BIOLOG EcoPlates) and increased bacterial oxidation of carbohydrates and phenols. Non-metric multidimensional scaling analysis as well as ANOVA testing showed that warming or increased precipitation did not change microbial community structure, which could suggest that microbial communities in semi-arid grasslands are already adapted to fluctuating climatic conditions. In summary, our results support the idea that microbial responses to climate change are multifaceted and, even with no large shifts in community structure, microbial mediation of soil carbon loss could still occur under future climate scenarios.     

Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits

Global nitrogen (N) enrichment and changing precipitation regimes are likely to alter plant community structure and composition, with consequent influences on biodiversity and ecosystem functioning. Responses of plant community structure and composition to N addition and increased precipitation were examined in a temperate steppe in northern China. Increased precipitation and N addition stimulated and suppressed community species richness, respectively, across 6 years (2005–2010) of the manipulative experiment. N addition and increased precipitation significantly altered plant community structure and composition at functional groups levels. The significant relationship between species richness and soil moisture (SM) suggests that plant community structure is mediated by water under changing environmental conditions. In addition, plant height played an important role in affecting the responses of plant communities to N addition, and the effects of increased precipitation on plant community were dependent on species rooting depth. Our results highlight the importance and complexity of both abiotic (SM) and biotic factors (species traits) in structuring plant community under changing environmental scenarios. These findings indicate that knowledge of species traits can contribute to mechanistic understanding and projection of vegetation dynamics in response to future environmental change.     

Optimal and central-place foraging theory applied to a desert harvester ant,Pogonomyrmex californicus

Summary Certain predictions of optimal- and central place-foraging theory were tested on the desert harvester ant, Pogonomyrmex californicus. Colonies were offered three different sizes of oat seed and found to maximize net energy intake (ei) over time (t _i ) by harvesting the seed sizes with the highest e _i /t _i rank. Two aspects of t _i were measured that were assumed constant in previous studies. The handling components of t _i (time required to manipulate the seed and travel time back to the colony with the food) were measured and found to be positively correlated with seed size. The manipulation success rate (the percentage of handled seeds successfully picked up) decreased with increased seed size. These results point out how important it is to measure all parameters of e _i /t _i rather than to assume constancy with both prey type and foraging distance. The relative abundance of less preferred food types was important in determining the proportion of preferred types in the diet. The food supply of eight colonies was manipulated experimentally over a 25-day period. Four deprived colonies were constrained within aluminum enclosures to prevented foraging. The remaining four satiated colonies were given food ad libitum. The niche breadths of the treated colonies were then compared to controls, but found not to differ significantly. Seed baits were offered at three distances from the colony to test whether selectivity increased with disance. Contrary to theoretical predictions, all colonies harcested about the same proportion of each seed size at each distance.     

·荒漠草地植物多样性对草食动物采食的响应机制

草食动物采食对草地植物多样性和生态系统功能的影响机制是放牧生态学研究的核心问题。该研究以内蒙古锡林郭勒盟苏尼特右旗荒漠草原的长期放牧控制实验为平台, 从既有草地植物多样性和动物偏食性两个层面系统地研究了荒漠草地植物多样性对草食动物采食的响应机制。结果显示: 1)荒漠草地植物对草食动物采食呈现4种响应模式: 放牧“隐没种”、放牧“敏感种”、放牧“无感种”、“绝对优势种”; 2)在群落尺度上, 物种多样性指数随放牧强度增加而减少, 与不放牧小区相比, 重度放牧(HG)与适度放牧(MG)小区植物多样性均下降, 且这一规律同样适用于功能群多样性, 灌木半灌木这一功能群内物种多样性对放牧干扰较敏感; 3)在草地既有植物的基础上, 以不放牧小区为参考系, 草食动物对植物功能群偏食性的排序为: 一二年生草本(AB) >多年生杂类草(PF) >灌木半灌木(SS) >多年生禾草(PG), 且偏食性物种主要分布于AB和PF中; 4)植物多样性与动物偏食性基本呈显著负相关关系(p < 0.05)。     

Ecosystem responses to increased precipitation and permafrost decay in subarctic Sweden inferred from peat and lake sediments

Recent accelerated decay of discontinuous permafrost at the Stordalen Mire in northern Sweden has been attributed to increased temperature and snow depth, and has caused expansion of wet minerotrophic areas leading to significant changes in carbon cycling in the mire. In order to track these changes through time and evaluate potential forcing mechanisms, this paper analyses a peat succession and a lake sediment sequence from within the mire, providing a record for the last 100 years, and compares these with monitored climate and active layer thickness data. The peat core was analysed for testate amoebae to reconstruct changes in peatland surface moisture conditions and water table fluctuations. The lake sediment core was analysed by near infrared spectroscopy to infer changes in the total organic carbon (TOC) concentration of the lake‐water, and changes in δ¹³C and C, N and δ¹⁵N to track changes in the dissolved inorganic carbon (DIC) pool and the influence of diagenetic effects on sediment organic matter, respectively. Results showed that major shifts towards increased peat surface moisture and TOC concentration of the lake‐water occurred around 1980, one to two decades earlier than a temperature driven increase in active layer thickness. Comparison with monitored temperature and precipitation from a nearby climate station indicates that this change in peat surface moisture is related to June–September (JJAS) precipitation and that the increase in lake‐water TOC concentration reflects an increase in total annual precipitation. A significant depletion in ¹³C of sediment organic matter in the early 1980s probably reflects the effect of a single or a few consecutive years with anomalously high summer precipitation, resulting in elevated DIC content of the lake water, predominantly originating from increased export and subsequent respiration of organic carbon from the mire. Based on these results, it was not possible to link proxy data obtained on peat and lake‐sediment records directly to permafrost decay. Instead our data indicate that increased precipitation and anomalously high rainfall during summers had a significant impact on the mire and the adjacent lake ecosystem. We therefore propose that effects of increased precipitation should be considered when evaluating potential forcing mechanisms of recent changes in carbon cycling in the subarctic.     

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire‐prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post‐fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life‐history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life‐history traits and evaluated whether these groups exhibited a consistent fire‐response. Six life‐history traits varied significantly between burned and unburned areas in short (up to 4 years) or long‐term (up to 52 years) post‐fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life‐history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind‐dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non‐wind dispersed seeds, and taller heights. Our results show that PFTs based on life‐history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long‐lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life‐history strategies.     

Plant community responses to increased precipitation and belowground litter addition: Evidence from a 5‐year semiarid grassland experiment

Global climate change is predicted to stimulate primary production and consequently increases litter inputs. Changing precipitation regimes together with enhanced litter inputs may affect plant community composition and structure, with consequent influence on diversity and ecosystem functioning. Responses of plant community to increased precipitation and belowground litter addition were examined lasting 5 years in a semiarid temperate grassland of northeastern China. Increased precipitation enhanced community species richness and abundance of annuals by 16.8% and 44%, but litter addition suppressed them by 25% and 54.5% after 5 years, respectively. During the study period, perennial rhizome grasses and forbs had consistent negative relationship under ambient plots, whereas positive relationship between the two functional groups was found under litter addition plots after 5 years. In addition, increased precipitation and litter addition showed significant interaction on community composition, because litter addition significantly increased biomass and abundance of rhizome grasses under increased precipitation plots but had no effect under ambient precipitation levels. Our findings emphasize the importance of water availability in modulating the responses of plants community to potentially enhanced litter inputs in the semiarid temperate grassland.     

Microbial biomass and respiration responses to nitrogen fertilization in a polar desert

How microbial communities respond to increases in available nitrogen (N) will influence carbon (C) and nutrient cycles. Most studies addressing N fertilization focus on mid-latitude ecosystems, where complex aboveground–belowground interactions can obscure the response of the soil microbial community, and little is known about how soil microbial communities of polar systems, particularly polar deserts, will respond. The low C content and comparatively simpler (low biomass and biodiversity) soil communities of the McMurdo Dry Valleys of Antarctica may allow easier identification of the mechanisms by which N fertilization influences microbial communities. Therefore, we conducted a microcosm incubation using three levels of N fertilization, added in solution to simulate a pulse of increased soil moisture, and measured microbial biomass and respiration over the course of 4.5 months. Soil characteristics, including soil pH, conductivity, cation content, chlorophyll a, and organic C content were measured. Soils from two sites that differed in stoichiometry were used to examine how in situ C:N:P influenced the N-addition response. We hypothesized that negative influences of N enrichment would result from increased salinity and ion content, while positive influences would result from enhanced C availability and turnover. We observed that microbes were moderately influenced by N addition, including stimulation and inhibition with increasing levels of N. Mechanisms identified include direct inhibition due to N toxicity and stimulation due to release from N, rather than C, limitation. Our results suggest that, by influencing microbial biomass and activity, N fertilization will influence C cycling in soils with very low C content.