The interaction of soil biota and soil structure under global change

The structural framework of soil mediates all soil processes, at all relevant scales. The spatio-temporal heterogeneity prevalent in most soils underpins the majority of biological diversity in soil, providing refuge sites for prey against predator, flow paths for biota to move, or be moved, and localized pools of substrate for biota to multiply. Just as importantly, soil biota play a crucial role in mediating soil structure: bacteria and fungi aggregate and stabilize structure at small scales (μm–cm) and earthworms and termites stabilize and create larger-scale structures (mm–m). The stability of this two-way interaction of structure and biota relations is crucial to the sustainability of the ecosystem. Soil is constantly reacting to changes in microclimates, and many of the soil–plant–microbe processes rely on the functioning of subtle chemical and physical gradients. The effect of global change on soil structure–biota interactions may be significant, through alterations in precipitation, temperature events, or land-use. Nonetheless, because of the complexity and the ubiquitous heterogeneity of these interactions, it is difficult to extrapolate from general qualitative predictions of the effects of perturbations to specific reactions. This paper reviews some of the main soil structure–biota interactions, particularly focusing on soil stability, and the role of biota mediating soil structures. The effect of alterations in climate and land-use on these interactions is investigated. Several case studies of the effect of land-use change are presented.     

Enzymology under global change: organic nitrogen turnover in alpine and sub-Arctic soils

Understanding global change impacts on the globally important carbon storage in alpine, Arctic and sub-Arctic soils requires knowledge of the mechanisms underlying the balance between plant primary productivity and decomposition. Given that nitrogen availability limits both processes, understanding the response of the soil nitrogen cycle to shifts in temperature and other global change factors is crucial for predicting the fate of cold biome carbon stores. Measurements of soil enzyme activities at different positions of the nitrogen cycling network are an important tool for this purpose. We review a selection of studies that provide data on potential enzyme activities across natural, seasonal and experimental gradients in cold biomes. Responses of enzyme activities to increased nitrogen availability and temperature are diverse and seasonal dynamics are often larger than differences due to experimental treatments, suggesting that enzyme expression is regulated by a combination of interacting factors reflecting both nutrient supply and demand. The extrapolation from potential enzyme activities to prediction of elemental nitrogen fluxes under field conditions remains challenging. Progress in molecular '-omics' approaches may eventually facilitate deeper understanding of the links between soil microbial community structure and biogeochemical fluxes. In the meantime, accounting for effects of the soil spatial structure and in situ variations in pH and temperature, better mapping of the network of enzymatic processes and the identification of rate-limiting steps under different conditions should advance our ability to predict nitrogen fluxes.     

A meta-analysis of responses of soil biota to global change

Global environmental changes are expected to impact the abundance of plants and animals aboveground, but comparably little is known about the responses of belowground organisms. Using meta-analysis, we synthesized results from over 75 manipulative experiments in order to test for patterns in the effects of elevated CO(2), warming, and altered precipitation on the abundance of soil biota related to taxonomy, body size, feeding habits, ecosystem type, local climate, treatment magnitude and duration, and greenhouse CO(2) enrichment. We found that the positive effect size of elevated CO(2) on the abundance of soil biota diminished with time, whereas the negative effect size of warming and positive effect size of precipitation intensified with time. Trophic group, body size, and experimental approaches best explained the responses of soil biota to elevated CO(2), whereas local climate and ecosystem type best explained responses to warming and altered precipitation. The abundance of microflora and microfauna, and particularly detritivores, increased with elevated CO(2), indicative of microbial C limitation under ambient CO(2). However, the effects of CO(2) were smaller in field studies than in greenhouse studies and were not significant for higher trophic levels. Effects of warming did not depend on taxon or body size, but reduced abundances were more likely to occur at the colder and drier sites. Precipitation limited all taxa and trophic groups, particularly in forest ecosystems. Our meta-analysis suggests that the responses of soil biota to global change are predictable and unique for each global change factor.     

Spatial turnover in the global avifauna

Despite its wide implications for many ecological issues, the global pattern of spatial turnover in the occurrence of species has been little studied, unlike the global pattern of species richness. Here, using a database on the breeding distributions of birds, we present the first global maps of variation in spatial turnover for an entire taxonomic class, a pattern that has to date remained largely a matter of conjecture, based on theoretical expectations and extrapolation of inconsistent patterns from different biogeographic realms. We use these maps to test four predictions from niche theory as to the form that this variation should take, namely that turnover should increase with species richness, towards lower latitudes, and with the steepness of environmental gradients and that variation in turnover is determined principally by rare (restricted) species. Contrary to prediction, we show that turnover is high both in areas of extremely low and high species richness, does not increase strongly towards the tropics, and is related both to average environmental conditions and spatial variation in those conditions. These results are closely associated with a further important and novel finding, namely that global patterns of spatial turnover are driven principally by widespread species rather than the restricted ones. This complements recent demonstrations that spatial patterns of species richness are also driven principally by widespread species, and thus provides an important contribution towards a unified model of how terrestrial biodiversity varies both within and between the Earth's major land masses.     

全球变化对土壤动物多样性的影响

陆地生态系统由地上和地下两部分组成,二者相互作用共同影响生态系统过程和功能.土壤动物在生物地球化学循环方面起着重要作用.随着人们对土壤动物在生态系统过程中重要性的认识,越来越多的研究表明全球变化对土壤动物多样性产生深刻影响.土地利用方式的改变、温度增加和降雨格局的改变能直接影响土壤动物多样性.CO2浓度和氮沉降的增加主要通过影响植物群落结构、组成和化学成分对土壤动物多样性产生间接影响.不同环境因子之间又能相互作用共同影响土壤动物多样性.了解全球变化背景下不同驱动因子及其交互作用对土壤动物多样性的影响,有助于更好地预测未来土壤动物多样性及相关生态学过程的变化.     

Nitrogen uptake and turnover in riparian woody vegetation

The nutrient balance of streams and adjacent riparian ecosystems may be modified by the elimination of anadromous fish runs and perhaps by forest fertilization. To better understand nitrogen (N) dynamics within stream and riparian ecosystems we fertilized two streams and their adjacent riparian corridors in central Idaho. On each stream two nitrogen doses were applied to a swathe approximately 35 m wide centered on the stream. The fertilizer N was enriched in ¹⁵N to 18. This enrichment is light relative to many previous labeling studies, yet sufficient to yield a traceable signal in riparian and stream biota. This paper reports pre-treatment differences in ¹⁵N and the first-year N response to fertilizer within the riparian woody plant community. Future papers will describe the transfer of allochthonous litter N to the stream and its subsequent processing by stream biota. Pre-treatment ¹⁵N differed between the two creeks (P=0.0002), possibly due to residual salmon nitrogen in one of the creeks. Pre-treatment ¹⁵N of current-year needles was enriched compared to leaf litter, which was in turn enriched compared to needles aged 4 years and older. We conclude that fractionation due to retranslocation occurs in at least two phases. The first phase, which optimizes allocation of N in younger needle age classes, is distinctly different from the second, which conserves N prior to abscission. The ¹⁵N difference between creeks was eliminated by the fertilization (P=0.42). In the two dominant conifer species, Abies lasiocarpa and Picea engelmannii, most fertilizer N was found in the current-year foliage; little was found in older needles and none was detected in litter (P=0.53). The only N-fixing shrub species, Alnus incana, took up only a small amount of fertilizer N [mean percent N derived from fertilizer (%Ndff) 5.0±1.6% (SE)]. Far more fertilizer N was taken up by other deciduous shrubs (mean %Ndff=33.9±4.5%). Fertilizer N made up 25% (±4.2%) of the N in deciduous shrub litter. These results demonstrate the feasibility of light labeling with ¹⁵N and the potential influence of riparian plant species composition on stream nutrient dynamics via allochthonous leaf litter inputs.     

Long-term soil transplant simulating climate change with latitude significantly alters microbial temporal turnover

To understand soil microbial community stability and temporal turnover in response to climate change, a long-term soil transplant experiment was conducted in three agricultural experiment stations over large transects from a warm temperate zone (Fengqiu station in central China) to a subtropical zone (Yingtan station in southern China) and a cold temperate zone (Hailun station in northern China). Annual soil samples were collected from these three stations from 2005 to 2011, and microbial communities were analyzed by sequencing microbial 16S ribosomal RNA gene amplicons using Illumina MiSeq technology. Our results revealed a distinctly differential pattern of microbial communities in both northward and southward transplantations, along with an increase in microbial richness with climate cooling and a corresponding decrease with climate warming. The microbial succession rate was estimated by the slope (w value) of linear regression of a log-transformed microbial community similarity with time (time–decay relationship). Compared with the low turnover rate of microbial communities in situ (w=0.046, P<0.001), the succession rate at the community level was significantly higher in the northward transplant (w=0.058, P<0.001) and highest in the southward transplant (w=0.094, P<0.001). Climate warming lead to a faster succession rate of microbial communities as well as lower species richness and compositional changes compared with in situ and climate cooling, which may be related to the high metabolic rates and intense competition under higher temperature. This study provides new insights into the impacts of climate change on the fundamental temporal scaling of soil microbial communities and microbial phylogenetic biodiversity.     

Desertification and global change

Arid and semiarid regions cover one third of the continental areas on Earth. These regions are very sensitive to a variety of physical, chemical and biological degradation processes collectively called desertification. Although interest in desertification has varied widely in time, there is a renewed concern about the evolution of dryland ecosystems because (1) a significant fraction of existing drylands already suffers from miscellaneous degradation processes, (2) increasing populations will inevitably result in further over-utilization of the remaining productive areas, (3) climatic changes expected from the greenhouse warming might result in drier continental interiors, and (4) some of the desertification processes themselves may amplify local or regional climatic changes. This paper reviews some of the many aspects of this issue in the context of the Global Change research program.     

Carbon exchange in biological soil crust communities under differential temperatures and soil water contents: implications for global change

Biological soil crusts (biocrusts) are an integral part of the soil system in arid regions worldwide, stabilizing soil surfaces, aiding vascular plant establishment, and are significant sources of ecosystem nitrogen and carbon. Hydration and temperature primarily control ecosystem CO₂ flux in these systems. Using constructed mesocosms for incubations under controlled laboratory conditions, we examined the effect of temperature (5–35 °C) and water content (WC, 20–100%) on CO₂ exchange in light (cyanobacterially dominated) and dark (cyanobacteria/lichen and moss dominated) biocrusts of the cool Colorado Plateau Desert in Utah and the hot Chihuahuan Desert in New Mexico. In light crusts from both Utah and New Mexico, net photosynthesis was highest at temperatures >30 °C. Net photosynthesis in light crusts from Utah was relatively insensitive to changes in soil moisture. In contrast, light crusts from New Mexico tended to exhibit higher rates of net photosynthesis at higher soil moisture. Dark crusts originating from both sites exhibited the greatest net photosynthesis at intermediate soil water content (40–60%). Declines in net photosynthesis were observed in dark crusts with crusts from Utah showing declines at temperatures >25 °C and those originating from New Mexico showing declines at temperatures >35 °C. Maximum net photosynthesis in all crust types from all locations were strongly influenced by offsets in the optimal temperature and water content for gross photosynthesis compared with dark respiration. Gross photosynthesis tended to be maximized at some intermediate value of temperature and water content and dark respiration tended to increase linearly. The results of this study suggest biocrusts are capable of CO₂ exchange under a wide range of conditions. However, significant changes in the magnitude of this exchange should be expected for the temperature and precipitation changes suggested by current climate models.     

Global climate change and soil carbon stocks; predictions from two contrasting models for the turnover of organic carbon in soil

Enhanced release of CO₂ to the atmosphere from soil organic carbon as a result of increased temperatures may lead to a positive feedback between climate change and the carbon cycle, resulting in much higher CO₂ levels and accelerated global warming. However, the magnitude of this effect is uncertain and critically dependent on how the decomposition of soil organic C (heterotrophic respiration) responds to changes in climate. Previous studies with the Hadley Centre's coupled climate–carbon cycle general circulation model (GCM) (HadCM3LC) used a simple, single‐pool soil carbon model to simulate the response. Here we present results from numerical simulations that use the more sophisticated ‘RothC’ multipool soil carbon model, driven with the same climate data. The results show strong similarities in the behaviour of the two models, although RothC tends to simulate slightly smaller changes in global soil carbon stocks for the same forcing. RothC simulates global soil carbon stocks decreasing by 54 Gt C by 2100 in a climate change simulation compared with an 80 Gt C decrease in HadCM3LC. The multipool carbon dynamics of RothC cause it to exhibit a slower magnitude of transient response to both increased organic carbon inputs and changes in climate. We conclude that the projection of a positive feedback between climate and carbon cycle is robust, but the magnitude of the feedback is dependent on the structure of the soil carbon model.     

Effects of multiple global change treatments on soil N2O fluxes

Global environmental changes are expected to alter ecosystem carbon and nitrogen cycling, but the interactive effects of multiple simultaneous environmental changes are poorly understood. Effects of these changes on the production of nitrous oxide (N₂O), an important greenhouse gas, could accelerate climate change. We assessed the responses of soil N₂O fluxes to elevated CO₂, heat, altered precipitation, and enhanced nitrogen deposition, as well as their interactions, in an annual grassland at the Jasper Ridge Global Change Experiment (CA, USA). Measurements were conducted after 6, 7 and 8?years of treatments. Elevated precipitation increased N₂O efflux, especially in combination with added nitrogen and heat. Path analysis supported the idea that increased denitrification due to increased soil water content and higher labile carbon availability best explained increased N₂O efflux, with a smaller, indirect contribution from nitrification. In our data and across the literature, single-factor responses tended to overestimate interactive responses, except when global change was combined with disturbance by fire, in which case interactive effects were large. Thus, for chronic global environmental changes, higher order interactions dampened responses of N₂O efflux to multiple global environmental changes, but interactions were strongly positive when global change was combined with disturbance. Testing whether these responses are general should be a high priority for future research.     

Nitrogen on microbial and global scales

Several temporal and spatial scales have been considered during the Symposium, and the presentations by Wallenda & Kottke (1998) and by Norby (1998) represent two extreme points in a wide range. They both deal with short and long-term impacts on the local as well as the global scale. I present here a few additional thoughts, and a subject that both papers have touched only slightly will be expanded: the differential impact of different forms of nitrogen deposition.     

Methane-oxidizing bacteria in a California upland grassland soil: diversity and response to simulated global change

We investigated the diversity of methane-oxidizing bacteria (i.e., methanotrophs) in an annual upland grassland in northern California, using comparative sequence analysis of the pmoA gene. In addition to identifying type II methanotrophs commonly found in soils, we discovered three novel pmoA lineages for which no cultivated members have been previously reported. These novel pmoA clades clustered together either with clone sequences related to "RA 14" or "WB5FH-A," which both represent clusters of environmentally retrieved sequences of putative atmospheric methane oxidizers. Conservation of amino acid residues and rates of nonsynonymous versus synonymous nucleotide substitution in these novel lineages suggests that the pmoA genes in these clades code for functionally active methane monooxygenases. The novel clades responded to simulated global changes differently than the type II methanotrophs. We observed that the relative abundance of type II methanotrophs declined in response to increased precipitation and increased atmospheric temperature, with a significant antagonistic interaction between these factors such that the effect of both together was less than that expected from their individual effects. Two of the novel clades were not observed to respond significantly to these environmental changes, while one of the novel clades had an opposite response, increasing in relative abundance in response to increased precipitation and atmospheric temperature, with a significant antagonistic interaction between these factors.     

Sources of plant-derived carbon and stability of organic matter in soil: implications for global change

Alterations in forest productivity and changes in the relative proportion of above‐ and belowground biomass may have nonlinear effects on soil organic matter (SOM) storage. To study the influence of plant litter inputs on SOM accumulation, the Detritus Input Removal and Transfer (DIRT) Experiment continuously alters above‐ and belowground plant inputs to soil by a combination of trenching, screening, and litter addition. Here, we used biogeochemical indicators [i.e., cupric oxide extractable lignin‐derived phenols and suberin/cutin‐derived substituted fatty acids (SFA)] to identify the dominant sources of plant biopolymers in SOM and various measures [i.e., soil density fractionation, laboratory incubation, and radiocarbon‐based mean residence time (MRT)] to assess the stability of SOM in two contrasting forests within the DIRT Experiment: an aggrading deciduous forest and an old‐growth coniferous forest. In the deciduous forest, removal of both above‐ and belowground inputs increased the total amount of SFA over threefold compared with the control, and shifted the SFA signature towards a root‐dominated source. Concurrently, light fraction MRT increased by 101 years and C mineralization during incubation decreased compared with the control. Together, these data suggest that root‐derived aliphatic compounds are a source of SOM with greater relative stability than leaf inputs at this site. In the coniferous forest, roots were an important source of soil lignin‐derived phenols but needle‐derived, rather than root‐derived, aliphatic compounds were preferentially preserved in soil. Fresh wood additions elevated the amount of soil C recovered as light fraction material but also elevated mineralization during incubation compared with other DIRT treatments, suggesting that not all of the added soil C is directly stabilized. Aboveground needle litter additions, which are more N‐rich than wood debris, resulted in accelerated mineralization of previously stored soil carbon. In summary, our work demonstrates that the dominant plant sources of SOM differed substantially between forest types. Furthermore, inputs to and losses from soil C pools likely will not be altered uniformly by changes in litter input rates.     

Nitrogen aboveground turnover and soil stocks to 8 m depth in primary and selectively logged forest in southern Amazonia

Extensive areas of Amazonia undergo selective logging, modifying forest structure and nutrient cycles. Anthropogenic‐accelerated rates of nitrogen (N) turnover could increase N loss and affect regeneration, carbon sequestration and timber production. We quantified leaf area reduction, canopy opening and downed biomass and resultant N flux from reduced impact logging (RIL) activities. We compared canopy reduction, surface soil moisture and nitrate to 8 m depth between logged gaps and intact primary forest to determine if logging activities increase subsoil nitrate. To test long‐term logging effects, we evaluated surface N stocks along a 12‐year postlogging chronosequence. At the harvest rate of 2.6 trees ha^?1, total N additions in logging gaps, including leaves and wood from felled crowns (24.8 kg N ha^?1) and other killed trees (41.9 kg N ha^?1), accounted for over 80% of the total N addition to aboveground necromass from all logging activities (81.9 kg N ha^?1). Despite this N turnover by logging, belowground nitrate storage to 8 m depth did not differ between logging gaps and primary forest at the low harvest rate and disturbance intensity of this study. Soil water depletion also did not differ between gaps and primary forest over 1 year, indicating the impact on belowground inorganic N was low. Compared with primary forest, nitrate concentrations to 8 m depth in logging gaps were only significantly higher at 60–100 cm, suggesting some N redistribution beyond the bulk of the fine roots in logging gaps. Extrapolated to the Amazon Basin scale, we provide a conservative estimate that logging damage and bole export under RIL would turn over 0.14 ± 0.07 to 0.23 ± 0.12 Tg N yr^?1 based on 1999–2002 selective logging rates. Greater damage during conventional selective logging would cause higher N turnover throughout the Amazon Basin than our results based on RIL.     

Marine viruses and global climate change

Sea-surface warming, sea-ice melting and related freshening, changes in circulation and mixing regimes, and ocean acidification induced by the present climate changes are modifying marine ecosystem structure and function and have the potential to alter the cycling of carbon and nutrients in surface oceans. Changing climate has direct and indirect consequences on marine viruses, including cascading effects on biogeochemical cycles, food webs, and the metabolic balance of the ocean. We discuss here a range of case studies of climate change and the potential consequences on virus function, viral assemblages and virus-host interactions. In turn, marine viruses influence directly and indirectly biogeochemical cycles, carbon sequestration capacity of the oceans and the gas exchange between the ocean surface and the atmosphere. We cannot yet predict whether the viruses will exacerbate or attenuate the magnitude of climate changes on marine ecosystems, but we provide evidence that marine viruses interact actively with the present climate change and are a key biotic component that is able to influence the oceans' feedback on climate change. Long-term and wide spatial-scale studies, and improved knowledge of host-virus dynamics in the world's oceans will permit the incorporation of the viral component into future ocean climate models and increase the accuracy of the predictions of the climate change impacts on the function of the oceans.