Controls on carbon cycling in tropical soils from the Amazon to the Andes: the influence of climate, plant inputs, nutrients and soil organisms
Hicks, Lettice Cricket
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Tropical soils are a globally important store of terrestrial carbon (C) and source of atmospheric carbon dioxide (CO2), regulated by the activity of soil microorganisms, through the mineralisation of plant residues and soil organic matter (SOM). Climatic warming will influence microbial activity, and this may accelerate the rate of C release from soils as CO2, contributing to alterations in current atmospheric composition, and generating feedbacks to climate change. Yet the magnitude of C loss from tropical soils remains uncertain, partly because we do not fully understand how non-climatic factors – including the chemistry of plant inputs, the availability of soil nutrients and the composition of the decomposer community – will interact to determine the response to changes in temperature. This thesis examines how these factors together regulate the rate of C cycling in contrasting soils across a 3400 m tropical elevation gradient in the Peruvian Andes, spanning a 20 ºC range (6.5 – 26.4 ºC) in mean annual temperature. Large-scale field-based manipulation experiments, translocating leaves and soil-cores across the elevation gradient (to impose an in-situ experimental warming treatment), were combined with controlled laboratory studies to examine the microbial-scale mechanisms which underlie the processes of decomposition and soil respiration observed in-situ. Results show that, across the gradient, rates of leaf-decomposition were determined principally by temperature and foliar chemical traits, while soil fertility had no significant influence. The effect of temperature was, however, stronger across higher-elevation sites, suggesting a greater vulnerability of the C-rich soils in montane systems to increased C loss under climatic warming. In lowland forests, the presence of invertebrate macrofauna also accelerated rates of decomposition, but leaf chemistry explained the greatest proportion of the observed variance, with a strong role for leaf chemical traits also identified under controlled conditions. Despite marked differences in microbial abundance and community composition among soils, these metrics were not associated with observed rates of decomposition. These results suggest that climate-related changes to plant species distributions (with associated changes to the chemistry of leaf-inputs), and upslope extension of macrofaunal ranges, could strongly influence future rates of leaf decomposition, independently of the direct response to warming. From the soil translocation study, root-soil interactions stimulated substantial net C loss from montane soils following translocation downslope (experimental warming treatment), indicating that warming-related changes to root productivity, exudation and/or species-composition could represent an important mode of future C loss from these soils. To examine more closely how inputs of plant-derived C influence the turnover of pre-existing SOM, and whether soil nutrient availability modulates the response, soils were amended with simple and complex 13C-labelled substrates in combination with inorganic nutrient treatments. Isotopic partitioning was used to determine the degree to which C and nutrient inputs accelerated (positive priming) or retarded (negative priming) the decomposition of SOM. Amendment of upper montane forest and montane grassland soils with nitrogen (N; alone and in combination with C) substantially retarded the decomposition of SOM, suggesting that microbial demand for N strongly regulates the turnover of organic matter in these soils. In contrast, amendment of lower montane and lowland forest soils with C stimulated positive priming of SOM, which was strongest in response to the simple C substrate and was not influenced by nutrient treatments, suggesting that microorganisms in these soils are primarily constrained by availability of labile C. Functional differences among microbial groups were also evident, with gram-negative bacteria and fungi using more labile sources of C while gram-positive bacteria used more complex C. Together, results from these studies considerably advance our understanding of soil C dynamics across lowland and montane systems, painting a rich picture of interacting processes which will determine the future soil C balance in tropical ecosystems. They show that the influence of temperature on the rate of soil C cycling is strongly affected by the nature and composition of plant-derived and atmospheric inputs, the principal additional constraints varying with elevation, leading to both opposing and reinforcing effects on rates of decomposition. The greater observed temperature sensitivity of decomposition at higher elevations is coupled with high microbial demand for N which regulates the turnover of SOM, whereas at lower elevations leaf decomposition is accelerated by active macrofaunal breakdown, while microbial decomposition of SOM is constrained by the availability of labile C. Under a global change scenario of increased temperature and N deposition, results therefore suggest that: (i) modified chemistry of plant inputs will influence rates of decomposition, independently of climate; (ii) increased availability of labile C will lead to more rapid decomposition of SOM at lower elevations; (iii) greater root productivity (associated with warming and plant-community shifts) will stimulate soil C loss across montane regions; but (iv) at higher elevations, a possible countervailing effect may be imposed on rapid warming-accelerated decomposition if increased N availability reduces microbial mineralisation of SOM. The net effect on the ecosystem C budget will depend on the balance of C gain from primary productivity and C loss from soils. Overall, however, the results presented here suggest that the large soil C stores in higher-elevation montane regions are particularly vulnerable to substantial reductions under exposure to short- and medium-term climatic warming.