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When glaciers retreat they expose barren substrates that become colonized by organisms, beginning the process of primary succession. Recent studies reveal that heterotrophic microbial communities occur in newly exposed glacial substrates before autotrophic succession begins. This raises questions about how heterotrophic microbial communities function in the absence of carbon inputs from autotrophs. We measured patterns of soil organic matter development and changes in microbial community composition and carbon use along a 150-year chronosequence of a retreating glacier in the Austrian Alps. We found that soil microbial communities of recently deglaciated terrain differed markedly from those of later successional stages, being of lower biomass and higher abundance of bacteria relative to fungi. Moreover, we found that these initial microbial communities used ancient and recalcitrant carbon as an energy source, along with modern carbon. Only after more than 50 years of organic matter accumulation did the soil microbial community change to one supported primarily by modern carbon, most likely from recent plant production. Our findings suggest the existence of an initial stage of heterotrophic microbial community development that precedes autotrophic community assembly and is sustained, in part, by ancient carbon.

Resource stoichiometry (C:N:P) is an important determinant of litter decomposition. However, the effect of elemental stoichiometry on the gross rates of microbial N and P cycling processes during litter decomposition is unknown. In a mesocosm experiment, beech (Fagus sylvatica L.) litter with natural differences in elemental stoichiometry (C:N:P) was incubated under constant environmental conditions. After three and six months, we measured various aspects of nitrogen and phosphorus cycling. We found that gross protein depolymerization, N mineralization (ammonification), and nitrification rates were negatively related to litter C:N. Rates of P mineralization were negatively correlated with litter C:P. The negative correlations with litter C:N were stronger for inorganic N cycling processes than for gross protein depolymerization, indicating that the effect of resource stoichiometry on intracellular processes was stronger than on processes catalyzed by extracellular enzymes. Consistent with this, extracellular protein depolymerization was mainly limited by substrate availability and less so by the amount of protease. Strong positive correlations between the interconnected N and P pools and the respective production and consumption processes pointed to feed-forward control of microbial litter N and P cycling. A negative relationship between litter C:N and phosphatase activity (and between litter C:P and protease activity) demonstrated that microbes tended to allocate carbon and nutrients in ample supply into the production of extracellular enzymes to mine for the nutrient that is more limiting. Overall, the study demonstrated a strong effect of litter stoichiometry (C:N:P) on gross processes of microbial N and P cycling in decomposing litter; mineralization of N and P were tightly coupled to assist in maintaining cellular homeostasis of litter microbial communities.

The natural abundance of stable (15)N isotopes in soils and plants is potentially a simple tool to assess ecosystem N dynamics. Several open questions remain, however, in particular regarding the mechanisms driving the variability of foliar delta(15)N values of non-N(2) fixing plants within and across ecosystems. The goal of the work presented here was therefore to: (1) characterize the relationship between soil net mineralization and variability of foliar Deltadelta(15)N (delta(15)Nleaf - delta(15)Nsoil) values from 20 different plant species within and across 18 grassland sites; (2) to determine in situ if a plant's preference for NO (3) (-) or NH (4) (+) uptake explains variability in foliar Deltadelta(15)N among different plant species within an ecosystem; and (3) test if variability in foliar Deltadelta(15)N among species or functional group is consistent across 18 grassland sites. Deltadelta(15)N values of the 20 different plant species were positively related to soil net mineralization rates across the 18 sites. We found that within a site, foliar Deltadelta(15)N values increased with the species' NO (3) (-) to NH (4) (+) uptake ratios. Interestingly, the slope of this relationship differed in direction from previously published studies. Finally, the variability in foliar Deltadelta(15)N values among species was not consistent across 18 grassland sites but was significantly influenced by N mineralization rates and the abundance of a particular species in a site. Our findings improve the mechanistic understanding of the commonly observed variability in foliar Deltadelta(15)N among different plant species. In particular we were able to show that within a site, foliar delta(15)N values nicely reflect a plant's N source but that the direction of the relationship between NO (3) (-) to NH (4) (+) uptake and foliar Deltadelta(15)N values is not universal. Using a large set of data, our study highlights that foliar Deltadelta(15)N values are valuable tools to assess plant N uptake patterns and to characterize the soil N cycle across different ecosystems.

The 15N isotope pool dilution (IPD) technique is the only available method for measuring gross ammonium (NH4+) production and consumption rates. Rapid consumption of the added 15N-NH4+ tracer is commonly observed, but the processes responsible for this consumption are not well understood. The primary objectives of this study were to determine the relative roles of biotic and abiotic processes in 15N-NH4+ sconsumption and to investigate the validity of one of the main assumptions of IPD experiments, i.e., that no reflux of the consumed 15N tracer occurs during the course of the experiments. We added a 15N-NH4+ tracer to live and sterile (autoclaved) soil using mineral topsoil from a beech forest and a grassland in Austria that differed in NH4+ concentrations and NH4+ consumption kinetics. We quantified both biotic tracer consumption (i.e. changes in the concentrations and 15N enrichments of NH4+, dissolved organic N (DON), NO3− and the microbial N pool) and abiotic tracer consumption (i.e., fixation by clay and/or humic substances). We achieved full recovery of the 15N tracer in both soils over the course of the 48 h incubation. For the forest soil, we found no rapid consumption of the 15N tracer, and the majority of tracer (78%) remained unconsumed at the end of the incubation period. In contrast, the grassland soil showed rapid 15N-NH4+ consumption immediately after tracer addition, which was largely due to both abiotic fixation (24%) and biotic processes, largely uptake by soil microbes (10%) and nitrification (13%). We found no evidence for reflux of 15N-NH4+ over the 48 h incubation period in either soil. Our study therefore shows that 15N tracer reflux during IPD experiments is negligible for incubation times of up to 48 h, even when rapid NH4+ consumption occurs. Such experiments are thus robust to the assumption that immobilized labeled N is not re–mobilized during the experimental period and does not impact calculations of gross N mineralization.

Conservation of water using mulches is a viable option under semiarid conditions for enhancing water use efficiency. Effect of mulches varies among years and with the amount and timing of mulching. Lucerne is a key crop for organic farming systems under semiarid conditions in Austria. Effect of mulching with lucerne has not been thoroughly investigated. Field experiments were conducted to assess the effect of lucerne utilization system (nonmulch versus mulch) on its shoot and root dry matter yield, biological nitrogen fixation and water use efficiency. Experiments were laid out in randomized complete block design with four replicates at the experimental farm of University of Natural Resources and Life Sciences, Vienna, Austria, during 2007–2008. Mulching was effective in lowering soil temperature by 1–6°C in the top 5 cm of soil. Utilization systems did not significantly affect the other studied parameters (P

This paper originates from a presentation at the IIIrd International Congress on Crassulacean Acid Metabolism, Cape Tribulation, Queensland, Australia, August 2001. We investigated the nitrogen (N) nutrition of Clusia osaensis, C. peninsulae and C. valerii during the seedling, epiphytic and hemiepiphytic phase in a lowland tropical forest in Costa Rica in order to elucidate nutritional adaptations of different plant growth stages to their habitat. Although all Clusia individuals were non-mycorrhizal, excised roots of seedlings, but also of epiphytic and hemiepiphytic stages, showed a distinct preference for glycine uptake. The shift in main rooting site from canopy soil to terrestrial soil was not reflected by changes in uptake rate or preference, although N availability and the composition of the available N pool changed significantly. High foliar N concentrations indicated that epiphytic seedlings seemed to be sufficiently supplied with N by maternal seed resources and canopy soils. With development, the epiphytic plants of Clusia may face N limitation due to higher N demands with increasing growth and restricted resources in the tree crowns. 15N natural abundance data indicate that epiphytes mainly accessed atmospheric and, to a lesser extent, canopy soil N sources and, after becoming terrestrially anchored, in the hemiepiphytic life stage exploited the larger nutrient reservoir of the ground soil. In consequence, Clusia species did not show an adaptation to the different N availability situations that they experienced whether canopy or ground-rooted.

Inselbergs are isolated rock outcrops displaying high heterogeneity in both soil formation and microclimatic condition with high variation in plant biodiversity. Vegetation patterns on inselbergs in the humid tropics range from rocks covered with dense biofilms predominated by cyanobacteria to high forest on deep soils. Along a similar transect, we investigated N supply to the vegetation using element and isotopic analyses of soil and biofilm samples from an inselberg in French Guiana. An increase in N content related to total dry weight (N%) in soils from the inselberg peak to surrounding habitats was related to changes in stable isotope composition (d 15 N). At the inselberg peak cyanobacterial biofilms on bare rocks and soils within small vegetation islands had similar d 15 N values of � 1.9% and � 1.3% while d 15 N of soils progressively increased towards the primary rainforest up to 6%. From the peak towards the base of the inselberg, the density of higher plants and soil depth increased. Hence, soil N cycling and N losses to the environment resulted in a progressive increase of soil d 15 N. The distribution of N%, d 15 N and d 13 C values suggest that the main N supply for soils at and nearby the inselberg is derived from cyanobacterial N2 fixation through leaching processes.

The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian arid zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta13C) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta13C of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.

Summary Bulk precipitation, throughfall and stemflow were collected to study anthropogenic effects on above-ground nutrient cycling in an off-shore mangrove forest (Rhizophora mangle L.) on Twin Cays, Belize. Samples were collected in a nitrogen limited fringe and phosphorus limited dwarf zone, and from an adjacent nitrogen fertilized fringe and a phosphorus fertilized dwarf zone. Inorganic cations and anions, dissolved organic carbon (DOC) and nitrogen (DON) were analysed. Throughfall represented 84% of precipitation volume. Sea salt ions (Cl−, Na+, SO 4 2 - and Mg2+) and DOC accounted for the highest proportion of solutes in rainwater, throughfall and stemflow in R. mangle stands. Non-marine sources dominated the flux of DON, DOC, NO 3 - , NH 4 + , and inorganic P (Pi) in bulk precipitation and throughfall and partially contributed to Ca2+ and K+. Deposition ratios (throughfall deposition:bulk deposition) showed that inorganic NH 4 + , and less so Pi were retained in the canopy of R. mangle from throughfall while all other solutes increased. Canopy leaching contributed in increasing order to net throughfall of Ca2+, Cl−, SO 4 2 - / K + , Mg2+ and Na+ but dry deposition dominated the net throughfall flux during the investigated period. Fertilizer treatment and zone did only slightly affect solute concentrations of hot-water extracts of leaves, of throughfall and stemflow in stands of similar stature. While litterfall and primary production have previously been shown to increase substantially upon nutrient enrichment of mangroves we therefore conclude that fertilization, as a surrogate of anthropogenic eutrophication, may not increase nutrient leaching from mangrove canopies, and thus may only have a minor effect on soluble organic matter cycling and inputs into mangrove food webs.

Earth hummocks constitute the most common surface structures of Arctic regions. In hummock tundra ecosystems, small mounds of earth alternate with depressions, the so-called interhummock areas. This study aimed at elucidating how differences in microtopography and associated variations in abiotic and biotic factors control biogeochemical cycles in hummock tundra soils. We assessed N pools and N transformation rates in hummocks and interhummock areas in the southern tundra subzone and along a soil-moisture gradient in the typical tundra subzone of the Taymyr Peninsula, Siberia, Russia. On a regional scale, N pools and transformation rates were positively related to latitude and therefore to temperature. Generally, wetter or waterlogged soil conditions tended to decrease gross mineralization rates and soil microbial N at least in interhummock areas. In contrast, at small spatial scales, soil microclimatic conditions were not the main determinant of the observed nitrogen cycling pattern. We found hi...

This paper originates from a presentation at the IIIrd International Congress on Crassulacean Acid Metabolism, Cape Tribulation, Queensland, Australia, August 2001. Carbon isotope fractionation and nocturnal acid accumulation in Clusia osaensis Hammel-ined., C. �peninsulae Hammel-ined. and C. valerii Standl. were investigated during the seedling, epiphytic and hemiepiphytic phases in a lowland tropical forest in Costa Ricato study photosynthetic adaptations of different plant growth stages to their habitat. Foliar δ 13C values around -24 to -32‰ indicate predominant C3 fixation of CO2 and low crassulacean acid metabolism (CAM) activity in all three Clusia species. Only terrestrially rooted plants of C. osaensis showed increased CAM expression. However, all developmental stages exhibited significant CAM cycling as shown by significant day-night fluctuations of titratable protons and of malic and citric acid. In C. valerii and C. peninsulae, an increase in CAM expression with plant development was not observed, and CAM cycling in hemiepiphytic-stage plants was completely repressed during the high rainfall season. The expression of CAM in the three Clusia species is therefore not developmentally controlled but triggered by environmental factors such as water availability and light intensity. These factors remain relatively stable in this ecosystem and CAM is therefore not fully expressed. However, CAM cycling may be of ecophysiological significance in all life stages as it serves as a mechanism to improve carbon economy by reducing respiratory CO2 losses.

Crassulacean acid metabolism (CAM) plants exhibit a complex interplay between CO(2) fixation by phosphoenolpyruvate carboxylase (PEPC) and ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco), and carbon demand for CAM maintenance and growth. This study investigated the flux of carbon from PEPC and direct Rubisco fixation to different leaf carbon pools and to phloem sap over the diurnal cycle. Concentrations and carbon isotope compositions of starch, soluble sugars, and organic acids were determined in leaves and phloem exudates of Kalanchoë daigremontiana Hamet et Perr., and related to CO(2) fixation by PEPC and Rubisco. Three types of leaf carbon pools could be distinguished. (i) Starch and malate pools were dominant and showed a pattern of reciprocal mobilization and accumulation (85/54 and 13/48 mg C g(-1) DW, respective, at the beginning/end of phase I). The carbon isotope composition of these pools was compatible with predominant PEPC fixation (delta(13)C values of -13 and -11 per thousand for starch and malate compared to -11 per thousand of PEPC fixed carbon). (ii) Isotopic composition (-17 per thousand and -14 per thousand) and concentration of glucose and fructose (2 and 3 mg C g(-1) DW, respectively) were not affected by diurnal metabolism, suggesting a low turnover. (iii) Sucrose (1-3 mg C g(-1) DW), in contrast, exhibited large diurnal changes in delta(13)C values (from -17 per thousand in the evening to -12 per thousand in the morning), which were not matched by net changes in sucrose concentration. This suggests a high sucrose turnover, fed by nocturnal starch degradation and direct Rubisco fixation during the day. A detailed dissection of the carbon fixation and mobilization pattern in K. daigremontiana revealed that direct fixation of Rubisco during the light accounted for 30% of phloem sucrose, but only 15% of fixed carbon, indicating that carbon from direct Rubisco fixation was preferentially used for leaf export.

We investigated factors that contribute to isotopic carbon fractionation in periphytic biofilms in a human-altered headwater stream with a flashy hydrograph. Water velocity had an important effect on periphyton d 13 C, explaining both temporal and spatial variation. We found that water velocity averaged over a certain period before sampling, rather than the instantaneous water velocity, explained a high percentage of both temporal and spatial variation of the periphyton d 13 C signature. The relationship between water velocity and periphyton d 13 C signature was particularly influenced by individual flow events during the recent flow history. A simple model based on a flow history of 3‐4 weeks reliably estimated the d 13 C signature of periphyton from distinct reaches. The model clearly identified signature shifts caused by the deposition of activated sludge particles from a wastewater treatment plant onto the periphytic biofilms. We highlight the high spatial and temporal variability of periphyton d 13 C signatures (i.e., up to 3‐6‰) in a heterogeneous flow environment with inputs from a wastewater treatment plant, and we explore its implications for food web analysis.