Precipitation pulses and carbon fluxes in semiarid and arid ecosystems

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a pulse of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO₂ is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.     

干旱、半干旱环境降水脉动对生态系统的影响

干旱、半干旱环境降水事件通常以脉动的形式发生,其发生时间、持续时间以及降水强度均具有较大变异性,降水事件的间断性和不可预知特征导致土壤水分与养分等关键资源的获得也呈不连续的脉动状态.资源脉动对生态系统的影响涉及个体、种群、甚至群落各个尺度.本文从干旱、半干旱环境资源脉动对生态系统的影响,以及生态系统对脉动事件的响应两方面系统综述了近年来的最新研究进展.指出国内外在资源脉动特征对生态系统的影响、不同生态系统资源脉动效应之间存在的差异、影响资源脉动持续时间的生态水文机制、资源可获得性对生态过程的影响等方面的研究尚处于尝试阶段,在全球气候变化、降水格局显著改变背景下,干旱区不同时空尺度资源脉动影响和生态系统响应是未来的研究重点.     

Contribution of winter processes to soil nitrogen flux in taiga forest ecosystems

We measured annual net nitrogen (N) mineralization, nitrification, and amino acid production in situ across a primary successional sequence in interior Alaska, USA. Net N mineralization per gram dry soil increased across the successional sequence, but with a sharp decline in the oldest stage (black spruce). Net N mineralization expressed per gram soil organic matter exhibited the opposite pattern, suggesting that soil organic matter quality decreases significantly across succession. Net N mineralization rates during the growing season from green-up (early May) through freeze-up (late September–early October) accounted for approximately 60% of the annual inorganic N flux, whereas the remaining N was released during the apparent dormant season. Nitrogen release during winter occurred primarily during October–January with only negligible N mineralization during early spring in stands of willow, alder, balsam poplar and white spruce. By contrast, black spruce stands exhibited substantial mineralization after snow melt during early spring. The high rates of N mineralization in late autumn through early winter coincide with high turnover of fine root biomass in these stands, suggesting that labile substrate production, rather than temperature, is a major controlling factor over N release in these ecosystems. We suggest that the convention of restricting measurements of soil processes to the growing season greatly underestimate annual flux rates of inorganic nitrogen in these high-latitude ecosystems.     

The effects of acid deposition on the biogeochemical cycles of major nutrients in miniature red spruce ecosystems

As part of an experimental study of air pollution effects on tree growth and health, we combined process studies with an ecosystem approach to evaluate the effects of acidic deposition on soil acidification, nutrient cycling and proton fluxes in miniature red spruce ecosystems. Ninety red spruce saplings were transplanted into 1-m diameter pots containing reconstructed soil profiles and exposed to simulated acid rain treatments of pH 3.1, 4.1 and 5.1 for four consecutive growing seasons. All the principal fluxes of the major elements were measured. During the first year of treatments, the disturbance associated with the transplanting of the experimental trees masked any treatment effects by stimulating N mineralization rates and consequent high N0₃ ⁻ cation, and H⁺ flux through the soil profile. In subsequent years, leaching of base cations and labile Al was accelerated in the most intensive acid treatment and corresponding declines in soil pH and exchangeable pools of Ca and Mg and increases in exchangeable Al concentrations were observed in the organic horizon. Leaching of Ca²⁺ and Mg²⁺ also was significantly higher in the pH 4.1 than in the pH 5.1 treatment. Flux of Ca from foliage and soil was increased in response to strong acid loading and root uptake increased to compensate for foliar Ca losses. In contrast, K cycling was dominated by root uptake and internal cycling and was relatively insensitive to strong acid inputs. Cation leaching induced by acidic deposition was responsible for the majority of H⁺ flux in the pH 3.1 treatment in the organic soil horizon whereas root uptake accounted for most of the H⁺ flux in the pH 4.1 and 5.1 treatments. Although no measurable effects on tree nutrition or health were observed, base cation leaching was significantly accelerated by acidic deposition, even at levels below that observed in the eastern U.S., warranting continued concern about acid deposition effects on the soil base status of forested ecosystems.     

草地生态系统中土壤氮素矿化影响因素的研究进展

氮素是各种植物生长和发育所需的大量营养元素之一,也是牧草从土壤吸收最多的矿质元素．土壤中的氮大部分以有机态形式存在,而植物可以直接吸收利用的是无机态氮．这些有机态氮在土壤动物和微生物的作用下。由难以被植物直接吸收利用的有机态转化为可被植物直接吸收利用的无机态的过程就是土壤氮的矿化．氮素矿化受多种因子的影响,这些因子可以归结为生物因子和非生物因子．生物因子包括：土壤动物、土壤微生物和植物种类．土壤动物可以促进土壤有机质的矿化;土壤微生物种类、结构及功能与氮的分解、矿化有密切的关系;不同的植物种类对土壤氮素的矿化作用是不相同的,一般来说。有豆科植物生长的土壤比其它种类土氮素矿化的作用大．非生物因素一般可以分为环境因子和人类活动干扰．环境因子中土壤温度和含水量对土壤氮素矿化的影响是国内外众多科学家研究的方向．尽管如此,在此方面的研究还没有取得一致意见,仍然需要进行这方面的研究,而在其他诸如：不同的土壤质地与土壤类型方面,研究报道的结论也很不一致,草地生态系统中人类活动对土壤氮素矿化的影响主要包括,不同强度的放牧,割草以及施肥、火烧强度等．非生物因子对氮素矿化的影响非常直接和明显,尤其是人类活动．本文综述了近年来影响草地生态系统土壤氮素矿化有关因素的一些进展．     

辽西半干旱区几种人工林生态系统涵养水源功能研究

从森林生态系统树冠截留降雨、枯落物持水及土壤蓄水3个层次对辽西半干旱区5种人工林生态系统的涵养水源功能进行了定量研究．结果表明,各人工林生态系统树冠对降雨的平均截留率为14．58％～37．19％,依次为沙棘林>油松沙棘混交林>杨树沙棘混交林>油松纯林>杨树纯林;枯落物层厚度为1．6～4．1cm,枯落物贮量为1890．4～6425．2kg·hm^-2,枯落物层厚度和贮量均为沙棘林>油松沙棘混交林>杨树沙棘混交林>油松纯林>杨树纯林,枯落物最大持水量取决于枯落物贮量及其最大持水率,枯落物最大持水量为5957．7～19332．9kg·hm^-2,依次为沙棘林>油松沙棘混交林>杨树沙棘混交林>油松纯林>杨树纯林;各人工林生态系统0～40cm土壤层非毛管蓄水量为23．70～37．85mm,依次为沙棘林>杨树沙棘混交林>油松沙棘混交林>杨树纯林>油松纯林．在5种人工林生态系统中,沙棘林的涵养水源功能最好,混交林较油松和杨树纯林有更好的涵养水源功能．     

Water and nitrogen dynamics in an arid woodland

Arid environments are characterized by spatial and temporal variation in water and nitrogen availability. differences in ¹⁵N and D of four co-occurring species reveal contrasting patterns of plant resource acquisition in response to this variation. Mineralization potential and nitrogen concentration of surface soils associated with plant canopies were greater than inter-canopy locations, and values decreased with increasing depth in both locations. Mineralization potential and nitrogen concentration were both negatively correlated with soil ¹⁵N. The spatial variation in soil ¹⁵N caused corresponding changes in plant ¹⁵N such that plant ¹⁵N values were negatively correlated with nitrogen concentration of surface soils. Plants occurring on soils with relatively high nitrogen concentrations had lower ¹⁵N, and higher leaf nitrogen concentrations, than plants occurring on soils with relatively low nitrogen concentrations. Two general temporal patterns of water and nitrogen use were apparent. Three species (Juniperus, Pinus andArtemisia) relied on the episodic availability of water and nitrogen at the soil surface. ¹⁵N values did not vary through the year, while xylem pressure potentials and stem-water D values fluctuated with changes in soil moisture at the soil surface. In contrast,Chrysothamnus switched to a more stable water and nitrogen source during drought. ¹⁵N values ofChrysothamnus increased throughout the year, while xylem pressure potentials and stem-water D values remained constant. The contrasting patterns of resource acquisition have important implications for community stability following disturbance. Disturbance can cause a decrease in nitrogen concentration at the soil surface, and so plants that rely on surface water and nitrogen may be more susceptible than those that switch to more stable water and nitrogen sources at depth during drougnt.     

地下水微生物功能群及生物地球化学循环

地下水系统是地球关键带的重要组成部分,为微生物提供了特殊的栖息环境和复杂的生存条件,进而演化出复杂的生物地球化学过程.随着多技术、多学科的交叉融合及发展,近几十年地下水微生物功能群及生物地球化学循环研究取得了引人瞩目的 重要进展.本文从地下水中的微生物群功能分区、微生物介导的地球化学元素循环、污染与修复中的生物地球化学...     

Soil macrofauna and nitrogen on a sub-Antarctic island

Summary The densities, diets and habitat preferences of the soil macrofaunal species on sub-Antarctic Marion Island (47°S, 38°E) are described. Their role in N cycling on the island is assessed, using a mire-grassland community as an example. Primary production on the island is high and this leads to a substantial annual requirement of nutrients by the vegetation. This requirement must almost wholly be met by mineralization of nutrient reserves in the organic matter. Rates of peat nitrogen mineralization mediated by microorganisms alone are much too low to account for rates of N uptake by the vegetation. Although soil macroinvertebrates, and bacteria represent a very small fraction of the total N pool, their interaction accounts for most of the peat N mineralization, as indicated by the amounts of inorganic N released into solution in microcosms. Extrapolation of the microcosm results shows that the soil macrofauna (mainly earthworms) stimulate the release of enough N from the mire-grassland peat to account for maximum N mineralization rates calculated from temporal changes in peat inorganic N levels and plant uptake during the most active part of the growing season. Considering that large numbers of mesoand microinvertebrates occur and must also contribute to nutrient mineralization, the soil faunal component is clearly of crucial importance to nutrient cycling on Marion Island. This is probably true of all sub-Antarctic islands.     

Pulse additions of soil carbon and nitrogen affect soil nitrogen dynamics in an arid Colorado Plateau shrubland

Biogeochemical cycles in arid and semi-arid ecosystems depend upon the ability of soil microbes to use pulses of resources. Brief periods of high activity generally occur after precipitation events that provide access to energy and nutrients (carbon and nitrogen) for soil organisms. To better understand pulse-driven dynamics of microbial soil nitrogen (N) cycling in an arid Colorado Plateau ecosystem, we simulated a pulsed addition of labile carbon (C) and N in the field under the canopies of the major plant species in plant interspaces. Soil microbial activity and N cycling responded positively to added C while NH₄⁺–N additions resulted in an accumulation of soil NO₃⁻. Increases in microbial activity were reflected in higher rates of respiration and N immobilization with C addition. When both C and N were added to soils, N losses via NH₃ volatilization decreased. There was no effect of soil C or N availability on microbial biomass N suggesting that the level of microbial activity (respiration) may be more important than population size (biomass) in controlling short-term dynamics of inorganic and labile organic N. The effects of C and N pulses on soil microbial function and pools of NH₄⁺–N and labile organic N were observed to last only for the duration of the moisture pulse created by treatment addition, while the effect on the NO₃⁻–N pool persisted after soils dried to pre-pulse moisture levels. We observed that increases in available C lead to greater ecosystem immobilization and retention of N in soil microbial biomass and also lowered rates of gaseous N loss. With the exception of trace gas N losses, the lack of interaction between available C and N on controlling N dynamics, and the subsequent reduction in plant available N with C addition has implications for the competitive relationships between plants species, plants and microbes, or both.     

冻融对土壤氮素损失及有效性的影响

土壤冻融交替是寒冷生态系统土壤氮素循环的重要驱动力。已有研究表明冻融交替作用能够促进氮素周转,从而缓解因土壤有效氮素缺乏而引起的植物生长限制。即便如此,冻融环境下土壤有效氮素供应量远高于其利用量,过剩的氮素会通过气态(N2O-N)排放、淋溶和径流等途径损失。论述了季节冻融环境和模拟冻融条件下土壤氮素损失状况;同时分析了影响冻融土壤N2O生产的相关因素、产生途径及冻融期N2O大量排放的机制;针对冻融交替过程中土壤氮素有效性问题,探讨了氮矿化、可溶性有机氮(DON)和微生物量氮(MBN)与氮素损失的关系。评述了土壤冻融研究中存在的不足,认为模型研究、土壤微生物功能、氮素转化中间产物、土壤-植物界面过程是未来值得关注和深入探讨的研究方向。     

Simulated annual carbon fluxes of grassland ecosystems in extremely arid conditions

In order to understand how changes in climate and land cover affect carbon cycles and structure and function of regional grassland ecosystems, we developed a grassland landscape productivity model, proposed an approach that combined both process-based modeling and spatial analysis with field measurements, and provided an example of semiarid region in Inner Mongolia, China, in extremely arid conditions. The modeled monthly mean and total net primary productivity, and monthly and annual mean respiration rates for Leymus chinensis steppes during the growing seasons in 2002 were mostly within the normal varying ranges of measured values, or similar to the field measurements, conducted in the similarly arid conditions. And the modeled total net ecosystem productivity (NEP) for L. chinensis steppes and Stipa grandis steppes were both between the lower and the higher measurements or within modeled multi-annual data by the other model. The modeled total NEP was 1.91 g C/m²/year over the entire study region. It indicated that if human disturbances were not considered, carbon budget over the entire study region during the growing seasons was nearly in balance or weak carbon sink even under extremely arid conditions. However, the modeled NEP spatially greatly varied not only over the entire study region (−48.28–52.09 g C/m²/year), but also among different land cover types. The modeled results also showed that there were obvious seasonal variations in carbon fluxes, mainly caused by leaf area index; and annual precipitation was the major limiting factor for the obvious spatial patterns of carbon fluxes from the east to the west. The modeled results also revealed the influence of extreme drought on carbon fluxes. The study provides an effective approach to derive useful information about carbon fluxes for different land cover types, and thus can instruct regional land-use planning and resource management based on carbon storage conditions.     

冰川消退带微生物群落演替及生物地球化学循环

冰川是生物圈重要组分之一。由于全球气候变化世界多地冰川加速消融,暴露原本被冰盖覆盖的区域,这些区域被称为冰川消退区域(glacier retreat area)或冰川前部区域(glacier foreland)。自暴露开始消退区随即发生初生演替,随着演替进行,物质循环逐步建立,生物量和土壤C、N总量逐步增加。生态系统C、N输入最初以矿化外来物为主,逐渐转变为以生物固C、固N为主。演替早期生态系统的发育主要受土壤C、N含量的限制,而演替后期的限制性营养物转变为P。演替区域土壤逐渐发育并促进生态位的分化,细菌、真菌、古菌,病毒及其他微生物群落的生物量和多样性不断增加直至达到该地区可承受的极值。随着生存条件的改善,不同生态策略物种的更替导致每个演替阶段微生物群落结构的差异。整体上,伴随演替进行微生物群落丰度、结构和活性呈现梯度性变化。气候变化对冰川消退带生态演替结果产生多方面的影响,而这些影响结果又综合反馈气候变化,因此目前难以准确估计气候变化对消退带生态演替的净效应。综述了近年冰川消退带微生物群落演替方面相关的研究结果,同时分别对该区域物质循环的建立、微生物群落演替和气候变化造成的影响这三个方面进行详细描述,并指出当前研究的不足。     

Directional climate change and potential reversal of desertification in arid and semiarid ecosystems

Our objective was to determine if long‐term increases in precipitation can maintain grasslands susceptible to desertification, and initiate a reversal of historic regime shifts on desertified shrublands. Perennial grass production and species richness in a multi‐year wet period were hypothesized to be greater than expected based on precipitation in a sequence of dry years. These responses were expected to differ for grasslands and shrublands with different dominant species and topo‐edaphic properties. Long‐term trends in desertification were documented using vegetation maps beginning in 1858, 1915, 1928, and 1998). These trends were compared with herbaceous and woody species responses to a sequence of dry (1994–2003) and wet years (2004–2008) for two grassland (uplands, playas) and three desertified shrubland types (honey mesquite, creosotebush, tarbush) in the Chihuahuan Desert. Analyses showed that both types of grasslands decreased in spatial extent since 1858 whereas areas dominated by mesquite or creosotebush increased. Production of upland grasslands in the wet period was greater than expected based on responses during the dry period whereas the relationships between species richness and precipitation was the same for both periods. Precipitation was not important to responses in playa grasslands in either period. For all ecosystem types, the production response in wet years primarily was an increase in herbaceous plants, and the most pronounced responses occurred on sandy sites (upland grasslands, mesquite shrubland). Results suggest that multiple wet years are needed to initiate a sequence of grass establishment and survival processes that can maintain upland grasslands without management inputs and lead to a state change reversal in desertified shrublands. Restoration strategies need to take advantage of opportunities provided by future climates while recognizing the importance of ecosystem type.     

Hierarchy of responses to resource pulses in arid and semi-arid ecosystems

In arid/semi-arid ecosystems, biological resources, such as water, soil nutrients, and plant biomass, typically go through periods of high and low abundance. Short periods of high resource abundance are usually triggered by rainfall events, which, despite of the overall scarcity of rain, can saturate the resource demand of some biological processes for a time. This review develops the idea that there exists a hierarchy of soil moisture pulse events with a corresponding hierarchy of ecological responses, such that small pulses only trigger a small number of relatively minor ecological events, and larger pulses trigger a more inclusive set and some larger ecological events. This framework hinges on the observation that many biological state changes, where organisms transition from a state of lower to higher physiological activity, require a minimal triggering event size. Response thresholds are often determined by the ability of organisms to utilize soil moisture pulses of different infiltration depth or duration. For example, brief, shallow pulses can only affect surface dwelling organisms with fast response times and high tolerance for low resource levels, such as some species of the soil micro-fauna and -flora, while it takes more water and deeper infiltration to affect the physiology, growth or reproduction of higher plants. This review first discusses how precipitation, climate and site factors translate into soil moisture pulses of varying magnitude and duration. Next, the idea of the response hierarchy for ecosystem processes is developed, followed by an exploration of the possible evolutionary background for the existence of response thresholds to resource pulses. The review concludes with an outlook on global change: does the hierarchical view of precipitation effects in ecosystems provide new perspectives on the future of arid/semiarid lands?     

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2

Atmospheric CO₂ enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO₂ enrichment, with an emphasis on water relations. Increasing CO₂ led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO₂ enrichment were relatively greater in dry years. In contrast, CO₂-induced aboveground biomass increases in the Texas C₃/C₄ grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO₂ in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO₂-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO₂ were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO₂ on plant and soil water relations may contribute substantially to experimentally induced CO₂-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO₂ effects on photosynthesis and those indirect CO₂ effects via soil moisture as documented here.     

Plant N capture from pulses: effects of pulse size,growth rate,and other soil resources

In arid ecosystems, the ability to rapidly capture nitrogen (N) from brief pulses is expected to influence plant growth, survival, and competitive ability. Theory and data suggest that N capture from pulses should depend on plant growth rate and availability of other limiting resources. Theory also predicts trade-offs in plant stress tolerance and ability to capture N from different size pulses. We injected K¹⁵NO₃, to simulate small and large N pulses at three different times during the growing season into soil around the co-dominant Great Basin species Sarcobatus vermiculatus, Chrysothamnus nauseosus ssp. consimilis, and Distichlis spicata. Soils were amended with water and P in a partial factorial design. As predicted, all study species showed a comparable decline in N capture from large pulses through the season as growth rates slowed. Surprisingly, however, water and P availability differentially influenced the ability of these species to capture N from pulses. Distichlis N capture increased up to tenfold with water addition while Chrysothamnus N capture increased up to threefold with P addition. Sarcobatus N capture was not affected by water or P availability. Opposite to our prediction, Sarcobatus, the most stress tolerant species, captured less N from small pulses but more N from large pulses relative to the other species. These observations suggest that variation in N pulse timing and size can interact with variable soil water and P supply to determine how N is partitioned among co-existing Great Basin species.     

Soil carbon storage, litterfall and CO2 efflux in fertilized and unfertilized larch (Larix leptolepis) plantations

This study evaluated the effects of forest fertilization on the forest carbon (C) dynamics in a 36-year-old larch (Larix leptolepis) plantation in Korea. Above- and below-ground C storage, litterfall, root decomposition and soil CO₂ efflux rates after fertilization were measured for 2 years. Fertilizers were applied to the forest floor at rates of 112 kg N ha⁻¹ year⁻¹, 75 kg P ha⁻¹ year⁻¹ and 37 kg K ha⁻¹ year⁻¹ for 2 years (May 2002, 2003). There was no significant difference in the above-ground C storage between fertilized (41.20 Mg C ha⁻¹) and unfertilized (42.25 Mg C ha⁻¹) plots, and the C increment was similar between the fertilized (1.65 Mg C ha⁻¹ year⁻¹) and unfertilized (1.52 Mg C ha⁻¹ year⁻¹) plots. There was no significant difference in the soil C storage between the fertilized and unfertilized plots at each soil depth (0–15, 15–30 and 30–50 cm). The organic C inputs due to litterfall ranged from 1.57 Mg C ha⁻¹ year⁻¹ for fertilized to 1.68 Mg C ha⁻¹ year⁻¹ for unfertilized plots. There was no significant difference in the needle litter decomposition rates between the fertilized and unfertilized plots, while the decomposition of roots with 1–2 mm diameters increased significantly with the fertilization relative to the unfertilized plots. The mean annual soil CO₂ efflux rates for the 2 years were similar between the fertilized (0.38 g CO₂ m⁻² h⁻¹) and unfertilized (0.40 g CO₂ m⁻² h⁻¹) plots, which corresponded with the similar fluctuation in the organic carbon (litterfall, needle and root decomposition) and soil environmental parameters (soil temperature and soil water content). These results indicate that little effect on the C dynamics of the larch plantation could be attributed to the 2-year short-term fertilization trials and/or the soil fertility in the mature coniferous plantation used in this study.     

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.     

Midday depression in root respiration of <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis>: its implication for estimating carbon cycling in forest ecosystems

We observed the phenomenon of midday depression in the rate of tree root respiration. Diurnal changes in the root respiration rate of Quercus crispula and Chamaecyparis obtusa were measured under intact conditions using a closed chamber method and a soil respiration measurement system (LI-6400 with a root respiration chamber) in a forest in the foothills of Mt. Fuji. After the measurement of intact root respiration in the field, the root was excised and taken to the laboratory, and the temperature dependence on the respiration rate of the detached root was measured using an open-flow gas exchange system with an infrared gas analyzer (LI-6252). The measurement was conducted in September 2003, August and November 2005, and June 2006. Whereas the root respiration rate of both species under intact conditions increased with increasing soil and root temperatures from dawn to early morning, the respiration rate decreased around midday from 10:00 to 15:00 despite an increment of soil and root temperatures. There was no clear relationship between the intact root respiration rate and root temperature in either species, although the detached root respiration rate of both increased exponentially with the temperature. The amount of the CO₂ efflux estimated using the temperature dependence of detached root respiration tended to underestimate the actual measurement value (intact respiration rate) by 20–50% in both species. These results indicate that evaluating midday depression in root respiration would be important for a more accurate estimation of the carbon cycle or net ecosystem production in forests.