The episodic nature of water availability in arid and semiarid ecosystems has significant consequences on belowground carbon and nutrient cycling. Pulsed water events directly control belowground processes through soil wet-dry cycles. Rapid soil microbial response to incident moisture availability often results in almost instantaneous C and N mineralization, followed by shifts in C/N of microbially available substrate, and an offset in the balance between nutrient immobilization and mineralization. Nitrogen inputs from biological soil crusts are also highly sensitive to pulsed rain events, and nitrogen losses, particularly gaseous losses due to denitrification and nitrate leaching, are tightly linked to pulses of water availability. The magnitude of the effect of water pulses on carbon and nutrient pools, however, depends on the distribution of resource availability and soil organisms, both of which are strongly affected by the spatial and temporal heterogeneity of vegetation cover, topographic position and soil texture. The 'inverse texture hypothesis' for net primary production in water-limited ecosystems suggests that coarse-textured soils have higher NPP than fine-textured soils in very arid zones due to reduced evaporative losses, while NPP is greater in fine-textured soils in higher rainfall ecosystems due to increased water-holding capacity. With respect to belowground processes, fine-textured soils tend to have higher water-holding capacity and labile C and N pools than coarse-textured soils, and often show a much greater flush of N mineralization. The result of the interaction of texture and pulsed rainfall events suggests a corollary hypothesis for nutrient turnover in arid and semiarid ecosystems with a linear increase of N mineralization in coarse-textured soils, but a saturating response for fine-textured soils due to the importance of soil C and N pools. Seasonal distribution of water pulses can lead to the accumulation of mineral N in the dry season, decoupling resource supply and microbial and plant demand, and resulting in increased losses via other pathways and reduction in overall soil nutrient pools. The asynchrony of resource availability, particularly nitrogen versus water due to pulsed water events, may be central to understanding the consequences for ecosystem nutrient retention and long-term effects on carbon and nutrient pools. Finally, global change effects due to changes in the nature and size of pulsed water events and increased asynchrony of water availability and growing season will likely have impacts on biogeochemical cycling in water-limited ecosystems.

Water Pulses and Biogeochemical Cycles in Arid and Semiarid Ecosystems

Biological processes are thought to be the primary cause of decomposition of organic matter in terrestrial ecosystems, with abiotic processes such as degradation by UV-B radiation playing a minor role. Not so, according to a study in the semi-arid Patagonian steppe. Here solar radiation has been identified as a major influence on carbon turnover. Blocking full sunlight or UV-B radiation dramatically reduced the rate of decomposition of plant litter, yet, reduction of soil organisms or addition of limiting soil resources had no effect. These results suggest a short-circuit in the carbon cycle; a substantial fraction of primary production could be lost to the atmosphere via photodegradation without cycling through soil organic matter pools. As nearly 40% of Earth's land surface is desert, global change affecting levels of radiation such as ozone depletion or cloud cover could have major impacts on carbon storage. Solar radiation is a dominant control on the rate of leaf litter decomposition in the semi-arid Patagonian steppe, which suggests that factors such as cloud cover may influence carbon cycling in semi-arid environments. The carbon balance in terrestrial ecosystems is determined by the difference between inputs from primary production and the return of carbon to the atmosphere through decomposition of organic matter1. Our understanding of the factors that control carbon turnover in water-limited ecosystems is limited, however, as studies of litter decomposition have shown contradictory results and only a modest correlation with precipitation2,3,4,5. Here we evaluate the influence of solar radiation, soil biotic activity and soil resource availability on litter decomposition in the semi-arid Patagonian steppe using the results of manipulative experiments carried out under ambient conditions of rainfall and temperature. We show that intercepted solar radiation was the only factor that had a significant effect on the decomposition of organic matter, with attenuation of ultraviolet-B and total radiation causing a 33 and 60 per cent reduction in decomposition, respectively. We conclude that photodegradation is a dominant control on above-ground litter decomposition in this semi-arid ecosystem. Losses through photochemical mineralization may represent a short-circuit in the carbon cycle, with a substantial fraction of carbon fixed in plant biomass being lost directly to the atmosphere without cycling through soil organic matter pools. Furthermore, future changes in radiation interception due to decreased cloudiness, increased stratospheric ozone depletion, or reduced vegetative cover may have a more significant effect on the carbon balance in these water-limited ecosystems than changes in temperature or precipitation.

Plant litter decomposition in a semi-arid ecosystem controlled by photodegradation.

Large areas of semiarid grasslands in the southwestern United States have been virtually replaced by shrubs during the past century. Understanding the causes and consequences of such vegetation dynamics requires that we elucidate the interplay between external forces of change (e.g., climate, human impacts) and the internal forces within these ecosystems that foster resilience and/or stability. Several conceptual models of arid ecosystems address this interplay by including the potential role of autogenic shrub effects on ecosystem processes, which lead to the formation of “resource islands” and tend to promote shrub persistence. Specifically, during the process of shrub establishment and maturation, the cycling of nutrients is progressively confined to the zones of litter accumulation beneath shrubs, while bare intershrub spaces become increasingly nutrient poor. As shrub resource islands develop, there is increased interception and stemflow by shrub canopies, confining infiltration of nutrient-enriched ...

Impact of drought on desert shrubs : Effects of seasonality and degree of resource island development

Summary


1
Aridland ecosystems cover about one-third of terrestrial environments globally, yet the extent to which models of carbon (C) and nitrogen (N) cycling, developed largely from studies of mesic ecosystems, apply to aridland systems remains unclear.

2
Within aridland ecosystems, C and N dynamics are often described by a pulse-reserve model in which episodic precipitation events stimulate biological activity that generate reserves of biomass, propagules and organic matter that prime the ecosystem to respond rapidly to subsequent precipitation events.

3
 The role of microbial C and N processing within the pulse-reserve paradigm has not received much study. We present evidence suggesting that fungi play a critical and underappreciated role in aridland soils, including efficient decomposition of recalcitrant C compounds, N-transformations such as nitrification, and nutrient storage and translocation of C and N between plants and biotic soil crusts. While fungi may perform some of these functions in other ecosystems, this ‘fungal loop’ assumes particular importance in the N cycle in aridlands because water availability imposes even greater restrictions on bacterial activity and physicochemical processes limit accumulation of soil organic matter (SOM).

4
We incorporate these findings into a Threshold-Delay Nutrient Dynamics (TDND) model for aridland ecosystems in which plant responses to pulsed precipitation events are mediated by a fungal loop that links C and N cycling, net primary production (NPP) and decomposition in aridland soils.

5
Synthesis. Arid ecosystems are highly sensitive to global environmental change including N deposition and altered precipitation patterns; yet, models from mesic ecosystems do not adequately apply to aridland environments. Our ‘fungal loop’ N cycle model integrates spatial structure with pulse dynamics and extends the pulse-reserve paradigm to include the key role of microbial processes in aridland ecosystem dynamics.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-2745.2008.01362.x

Pulse dynamics and microbial processes in aridland ecosystems

A framework for the ecology of arid Australia

We tested the hypotheses that rates of decomposition in a desert should be higher following single large rain events of 25 mm than evenly spaced 6 mm events and that supplemental rainfall should result in higher populations of soil biota. There were no significant differences in mass losses of creosotebush, Larrea tridentata, leaf litter on plots receiving water supplementation and no added water. On some sampling dates, there were higher mass losses in the 6 mm·week-1 treatment. Weekly rainfall produced higher coefficients of variation in mass losses than the other rainfall regimes. A single event pulse compared with weekly pulses of rainfall during the normal "dry" period resulted in no differences in mass losses. Microarthropods and nematodes exhibited numerical responses to supplemental rainfall but the litter microflora did not. These studies provide direct experimental evidence that the conventional wisdom linking decomposition to rainfall in deserts is wrong. The studies also suggest that the effects of litter fauna on surface litter decomposition are minimal; therefore, future studies should focus on activites of the microflora.

Rainfall and decomposition in the chihuahuan desert

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Rainfall and decomposition in the chihuahuan desert