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Iron (Fe) is essential to plants, microbes, and animals, is an important element in weathered soils from tropical and subtropical regions due to its reactivity toward carbon (C) and nutrients and its ability to serve as an electron acceptor for anaerobic respiration. Humid (sub)tropical and iron-rich soils naturally experience fluctuations in soil moisture, oxygen content, and hence, redox potential due to elevated but intermittent rainfall and high inputs of labile carbon from decomposed litter. Soils from the Luquillo Critical Zone Observatory (LCZO), Puerto Rico, are well-suited for studying the impact of redox fluctuations on Fe and C biogeochemistry. I conducted two laboratory experiments, exploring coupled Fe-C mechanisms, and one field experiment, using LCZO soils. Both lab experiments were conducted using soil in a slurry, which minimizes spatial variability and involved shifting between anoxic and oxic conditions. In the first lab study, I found that iron reduction rates increased when redox oscillations occurred more frequently. In the second lab experiment, I varied the time under oxic conditions (Ï4oxic) in both long and short oscillation periods. For the long treatments (Ï4anoxic at 6 d), I observed that as Ï4oxic decreased from 72 to 24 to 8 hours, Fe reduction rates increased, CO2 emissions remained unchanged, and CH4 emissions decreased; and for the short treatments (Ï4anoxic at 2 d), FeII and trace gases emissions decreased throughout the experiment. For the field experiment, I monitored several biogeochemical variables involved in Fe-C redox processes in triplicate catenas at ridge, slope, and valley positions. I found that soil moisture was a predictor for changes in FeII, rapidly-reducible Fe oxides (FeIIIRR), pH, Eh, and DOC. Valleys were more responsive to environmental changes than the other landscape positions. I also conducted three other lab studies (using LCZO soils) and one field experiment at the Calhoun CZO, in South Carolina (each are reported briefly in the Appendices). In conclusion, under natural and laboratory redox fluctuating systems, iron exerts a strong biogeochemical influence on the carbon dynamics of soils from humid (sub)tropical regions with important climate change and environmental implications.
Upland humid tropical forest soils experience fluctuations in oxygen (O2) availability and redox potential as a consequence of high rainfall, clay content, and respiration rates. Research in wetland ecosystems suggests that spatial and temporal variation in redox reactions strongly affect the biogeochemical cycling of carbon (C) and nitrogen (N). Here, I explored the impact of soil redox dynamics on decomposition and soil-atmosphere greenhouse gas fluxes in humid tropical ecosystems of the Luquillo Experimental Forest (LEF), Puerto Rico. Traditional theory and ecosystem models predict that elevated soil moisture leads to O2 limitation, constraining the enzymatic processes that mediate organic matter decomposition, and promoting the accumulation of soil C. Testing these hypotheses in upland humid tropical soils revealed the need for a more nuanced conceptual framework. In short: variation in moisture alone did not determine redox dynamics, hydrolytic enzymes activities persisted under reducing conditions, and redox fluctuations promoted decomposition on short (days) and long-term (decades) timescales. In Chapter One, I showed a relative decoupling between the temporal dynamics of soil moisture, soil redox reactions, and greenhouse gas fluxes over scales of days to weeks, using a field moisture manipulation experiment. Anaerobic biogeochemical processes such as iron (Fe) reduction and methanogenesis co-occurred in proximity to a well-aerated soil atmosphere and were little affected by fluctuations in soil moisture. Instead, redox reactions and gas fluxes appeared to vary constitutively according to differences in microtopography. In Chapter Two, I further explored relationships between reducing conditions and organic matter decomposition, by analyzing extracellular hydrolytic enzyme activities within and among sites differing in topography and rainfall. The enzymatic latch hypothesis proposes that reducing conditions inhibit hydrolytic enzymes via an accumulation of phenolic substances. I found little evidence for an enzymatic latch, and instead documented a strong positive relationship between reducing conditions, using reduced Fe (Fe(II)) as a proxy, and hydrolytic enzyme activities in a subset of sites. Furthermore, enzyme activities generally did not decline in an anaerobic incubation relative to aerobic controls. The assumption that reducing conditions constrain the decomposition activities of hydrolytic enzymes does not appear generally applicable in humid tropical forests. Next, in Chapter Three I examined the influence of temporal redox fluctuations on decomposition. Anaerobic conditions by definition limit the activity of oxidative enzymes, which require O2. The redox cycling of Fe, however, can potentially generate reactive oxygen species that mimic the function of oxidative enzymes. We demonstrated that concentrations of Fe(II) explained most of the variation in phenol oxidative activity within and among several sites in the LEF. Furthermore, Fe(II) oxidation stimulated short-term respiration, likely via a pH-mediated increase in dissolved organic C. Thus, stimulatory effects of redox fluctuations on oxidative decomposition processes might partially counteract short-term effects of O2 limitation. Finally, in Chapter Four I examined the overall impact of reducing conditions in comparison with other variables as they related to spatial patterns in soil C concentrations and turnover across the LEF. Soil C increased with Fe(II), an index of reducing conditions, but C tended to decline with increasing concentrations of reducible Fe oxides. Furthermore, the residence time of mineral-associated C (modeled using measurements of bomb radiocarbon) declined with Fe(II) concentrations. Together, the findings from these studies suggest a complex relationship between moisture, redox dynamics, and decomposition. First, short-term fluctuations in rainfall may have little overall impact on redox dynamics and the overall decomposition process, but longer-term differences in moisture among sites are associated with characteristic differences in redox reactions and greenhouse gas fluxes. Second, portions of the decomposition process mediated by hydrolytic enzymes appear resistant to periodic O2 deprivation and chronic reducing conditions, as well as the accumulation of phenolic substances. Third, redox cycling may give rise to important emergent mechanisms not evident under static aerobic conditions, mediated by coupled biotic and abiotic reactions with Fe oxides. Fourth, reducing conditions are associated with elevated soil C concentrations at the landscape scale, although the presence of reducible Fe oxides constrains C accumulation, and redox cycling might accelerate the turnover of mineral C over decadal scales. Together, these findings have implications for understanding the biogeochemical function of humid tropical soils, and their response to altered precipitation regimes and feedbacks to climate change. Two mechanisms thought to underlie the persistence of C in soils--reducing conditions induced by high soil moisture and the presence of reactive Fe minerals--may actually play unexpected roles in the decomposition of soil organic matter, a finding with potentially broad application across terrestrial and aquatic ecosystems.
Soil organic carbon (OC) is one of the largest carbon (C) reservoirs on the Earth’s surface. Because of the high sorption affinity of iron (Fe) minerals for OC, the redox reactions of Fe potentially play an important role in regulating the stability and transformation of OC in soils. Fate of Fe-bound OC in natural soils upon Fe redox reactions is a critical knowledge gap for understanding the coupled biogeochemical cycles of C and Fe. This study comprehensively investigated the amount and characteristics of Fe-bound OC in forest soils as well as the coupled biogeochemical reactions of Fe and OC during redox processes. Iron-bound OC contributed substantially to total organic carbon (TOC) in forest soils, representing an important component of C cycles in terrestrial ecosystems. The ecogeographical parameters, such as latitude and annual mean temperature, are governing factors for the fraction of Fe-bound OC in TOC (fFe-OC). Iron-bound OC was less aliphatic, more carboxylic, and more enriched in 13C, compared to non-Fe-bound OC. Our studies also demonstrated the closely coupled biogeochemical reactions of Fe and OC during redox processes. We found that microbial reduction of Fe can lead to substantial mobilization of OC in natural soils under anaerobic incubation. OC electron accepting capacity (EAC) strongly regulated Fe reduction, demonstrating that the biogeochemical cycles of Fe and OC are coupled together through two-way interactions. After transferring to the aerobic condition, Fe(II) in pre-reduced soils was oxidized in conjunction with oxidation of OC. OC oxidation was much lower for soils exposing to the anaerobic-aerobic transition, compared to soils only aerobically incubated, potentially because of secondary Fe minerals formed during the transition sequestrating OC. These results provide novel insights into the impact of anaerobic-aerobic transitions on the dynamics of OC in ecosystems undergoing the anaerobic-aerobic transitions frequently. Therefore, we argue that it is critical to include the redox reactions in biogeochemistry models for evaluating and predicting C stability and cycles.
Elements move through Earth's critical zone along interconnected pathways that are strongly influenced by fluctuations in water and energy. The biogeochemical cycling of elements is inextricably linked to changes in climate and ecological disturbances, both natural and man-made. Biogeochemical Cycles: Ecological Drivers and Environmental Impact examines the influences and effects of biogeochemical elemental cycles in different ecosystems in the critical zone. Volume highlights include: Impact of global change on the biogeochemical functioning of diverse ecosystems Biological drivers of soil, rock, and mineral weathering Natural elemental sources for improving sustainability of ecosystems Links between natural ecosystems and managed agricultural systems Non-carbon elemental cycles affected by climate change Subsystems particularly vulnerable to global change The American Geophysical Union promotes discovery in Earth and space science for the benefit of humanity. Its publications disseminate scientific knowledge and provide resources for researchers, students, and professionals. Book Review: http://www.elementsmagazine.org/archives/e16_6/e16_6_dep_bookreview.pdf
We report that iron-reducing bacteria are primary mediators of anaerobic carbon oxidation in upland tropical soils spanning a rainfall gradient (3500 - 5000 mm yr-1) in northeast Puerto Rico. The abundant rainfall and high net primary productivity of these tropical forests provide optimal soil habitat for iron-reducing and iron-oxidizing bacteria. Spatially and temporally dynamic redox conditions make iron-transforming microbial communities central to the belowground carbon cycle in these wet tropical forests. The exceedingly high abundance of iron-reducing bacteria (up to 1.2 x 109 cells per gram soil) indicated that they possess extensive metabolic capacity to catalyze the reduction of iron minerals. In soils from the higher rainfall sites, measured rates of ferric iron reduction could account for up to 44 % of organic carbon oxidation. Iron reducers appeared to compete with methanogens when labile carbon availability was limited. We found large numbers of bacteria that oxidize reduced iron at sites with high rates of iron reduction and large numbers of iron-reducers. the coexistence of large populations of ironreducing and iron-oxidizing bacteria is evidence for rapid iron cycling between its reduced and oxidized states, and suggests that mutualistic interactions among these bacteria ultimately fuel organic carbon oxidation and inhibit CH4 production in these upland tropical forests.
This volume quantifies carbon storage in managed forest ecosystems not only in biomass, but also in all soil compartments. It investigates the interaction between the carbon and nitrogen cycles by working along a north-south transect through Europe that starts in northern Sweden, passes through a N-deposition maximum in central Europe and ends in Italy. For the first time biogeochemical processes are linked to biodiversity on a large geographic scale and with special focus on soil organisms. The accompanying CD-ROM provides a complete database of all flux, storage and species observations for modellers.
Terrestrial environments contain one of the largest pools of iron (Fe) and organic carbon (OC), making the interaction between these two elements important drivers of biogeochemical cycles in soils and sediments, and critical for predicting future atmospheric carbon dioxide concentrations and climate change. This dissertation aims to understand the chemical and biological stability of Fe (hydr)oxides and OC in redox dynamic soils and sediments. A series of wet chemical techniques, advanced surface characterization methods and comparative metagenomic approaches were combined to (1) evaluate biogeochemical processes governing Fe (hydr)oxide mineralogical changes during redox oscillations, and (2) explore the biological stability of OC under Fe(III) reducing conditions. In Chapter 2, the mineralogical transformation of lepidocrocite and ferrihydrite was evaluated in the presence of freshwater sediment microorganisms under oscillating Fe(III) reducing (i.e., anaerobic glucose addition) and Fe(II) oxidizing (i.e., oxygen or nitrate addition) conditions. The results illustrate that the flux of different electron donors and acceptors can greatly impact Fe (hydr)oxide transformations and alter Fe, C, and N dynamics taking place in Fe-rich, redox active soils and sediments. Chapter 3 examined whether isolated humic substances could serve as both electron shuttles and electron donors for dissimilatory iron (hydr)oxide reduction (DIR) by freshwater sediment microorganisms. Both humic acid (HA) and humin (HM) were able to shuttle electrons and enhance DIR in cultures amended with glucose, but only HA could donate electrons for DIR by undergoing microbial degradation. Evidence for HA metabolism was observed by an overrepresentation of genes involved in polysaccharide degradation in cultures containing HA compared to those with only glucose. Chapter 4 evaluated the stability of Fe (hydr)oxide-HA coprecipitates under Fe(III) reducing conditions. HA enhanced DIR by serving as an electron shuttle and donor, but significant HA desorption was not observed. These results call into question the role of DIR in Fe (hydr)oxide-bound OC release. Overall, the transformation of Fe (hydr)oxides in redox active environments plays a central role in subsurface biogeochemical processes including nutrient availability, contaminant mobility and carbon dynamics.
The majority of carbon stored in the soils of the world is stored in forests. The refractory nature of some portions of forest soil organic matter also provides the slow, gradual release of organic nitrogen and phosphorus to sustain long term forest productivity. Contemporary and future disturbances, such as climatic warming, deforestation, short rotation sylviculture, the invasion of exotic species, and fire, all place strains on the integrity of this homeostatic system of C, N, and P cycling. On the other hand, the CO2 fertilization effect may partially offset losses of soil organic matter, but many have questioned the ability of N and P stocks to sustain the CO2 fertilization effect. Despite many advances in the understanding of C, N, and P cycling in forest soils, many questions remain. For example, no complete inventory of the myriad structural formulae of soil organic N and P has ever been made. The factors that cause the resistance of soil organic matter to mineralization are still hotly debated. Is it possible to “engineer” forest soil organic matter so that it sequesters even more C? The role of microbial species diversity in forest C, N, and P cycling is poorly understood. The difficulty in measuring the contribution of roots to soil organic C, N, and P makes its contribution uncertain. Finally, global differences in climate, soils, and species make the extrapolation of any one important study difficult to extrapolate to forest soils worldwide.