Vidya Suseela

Changes in the structural composition and reactivity of Acer rubrum leaf litter tannins exposed to warming and altered precipitation: climatic stress‐induced tannins are more reactive

• Climate change could increase the frequency with which plants experience abiotic stresses, leading to changes in their metabolic pathways. These stresses may induce the production of compounds that are structurally and biologically different from constitutive compounds. • We studied how warming and altered precipitation affected the composition, structure, and biological reactivity of leaf litter tannins in Acer rubrum at the Boston-Area Climate Experiment, in Massachusetts, USA. • Warmer and drier climatic conditions led to higher concentrations of protective compounds, including flavonoids and cutin. The abundance and structure of leaf tannins also responded consistently to climatic treatments. Drought and warming in combination doubled the concentration of total tannins, which reached 30% of leaf-litter DW. This treatment also produced condensed tannins with lower polymerization and a greater proportion of procyanidin units, which in turn reduced sequestration of tannins by litter fiber. Furthermore, because of the structural flexibility of these tannins, litter from this treatment exhibited five times more enzyme (β-glucosidase) complexation capacity on a per-weight basis. Warmer and wetter conditions decreased the amount of foliar condensed tannins. • Our finding that warming and drought result in the production of highly reactive tannins is novel, and highly relevant to climate change research as these tannins, by immobilizing microbial enzymes, could slow litter decomposition and thus carbon and nutrient cycling in a warmer, drier world.

Temperature response of soil respiration largely unaltered with experimental warming

Significance One of the greatest challenges in projecting future shifts in the global climate is understanding how soil respiration rates will change with warming. Multiple experimental warming studies have explored this response, but no consensus has been reached. Based on a global synthesis of 27 experimental warming studies spanning nine biomes, we find that although warming increases soil respiration rates, there is limited evidence for a shifting respiration response with experimental warming. We also note a universal decline in the temperature sensitivity of respiration at soil temperatures >25 °C. Together, our data indicate that future respiration rates are likely to follow the current temperature response function, but higher latitudes will be more responsive to warmer temperatures. The respiratory release of carbon dioxide (CO2) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming.

The Responses of Soil and Rhizosphere Respiration to Simulated Climatic Changes Vary by Season.

Responses of soil respiration (Rs) to anthropogenic climate change will affect terrestrial carbon storage and, thus, feed back to warming. To provide insight into how warming and changes in precipitation regimes affect the rate and temperature sensitivity of Rs and rhizosphere respiration (Rr) across the year, we subjected a New England old-field ecosystem to four levels of warming and three levels of precipitation (ambient, drought, and wet treatments). We measured Rs and heterotrophic respiration (Rh) monthly (in areas of the plots with and without plants, respectively) and estimated Rr by calculating the difference in respiration between Rs and Rh. Even in this mesic ecosystem, Rs and Rr responded strongly to the precipitation treatments. Drought reduced Rs and Rr, both annually and during the growing season. Annual cumulative Rs responded nonlinearly to precipitation treatments; both drought and supplemental precipitation suppressed Rs compared to the ambient treatment. Warming increased Rs and Rr in spring and winter when soil moisture was optimal but decreased these rates in summer when moisture was limiting. Cumulative winter Rr increased by about 200% in the high warming (approximately 3.5 degrees C) treatment. The effect of climate treatments on the temperature sensitivity of Rs depended on the season. In the fall, the drought treatment decreased apparent Q10 relative to the other precipitation treatments. The responses of Rs to warming and altered precipitation were largely driven by changes in Rr. We emphasize the importance of incorporating realistic soil moisture responses into simulations of soil carbon fluxes; the long-term effects of warming on carbon--climate feedback will depend on future precipitation regimes. Our results highlight the nonlinear responses of soil respiration to soil moisture and, to our knowledge, quantify for the first time the loss of carbon through winter rhizosphere respiration due to warming. While this additional loss is small relative to the cumulative annual flux in this system, such increases in rhizosphere respiration during the non-growing season could have greater consequences in ecosystems where they offset or reduce subsequent warming-induced gains in plant growth.

Labile compounds in plant litter reduce the sensitivity of decomposition to warming and altered precipitation

Together, climate and litter quality strongly regulate decomposition rates. Although these two factors and their interaction have been studied across species in continent-scale experiments, few researchers have studied how labile and recalcitrant compounds interact to influence decomposition, or the climate sensitivity of decomposition, within a litter type. Over a period of 3 yr, we studied the effects of warming and altered precipitation on mass loss and compound-specific decomposition using two litter types that possessed similar heteropolymer chemistry, but different proportions of labile and recalcitrant compounds. Climate treatments immediately affected the mass loss of the more recalcitrant litter, but affected the more labile litter only after 2 yr. After 3 yr, although both litter types had lost similar amounts of mass, warming (c. 4°C) and supplemental precipitation (150% of ambient) together accelerated the degradation of alkyl-carbon and lignin only in the more recalcitrant litter, highlighting the role of initial litter quality in determining whether the chemistry of litter residues converges or diverges under different climates. Our finding that labile compounds in litter reduce the climate sensitivity of mass loss and the decomposition of recalcitrant matrix is novel. Our results highlight the potential for litter quality to regulate the effect of climatic changes on the sequestration of litter-derived carbon.

Warming and drought reduce temperature sensitivity of nitrogen transformations

Shifts in nitrogen (N) mineralization and nitrification rates due to global changes can influence nutrient availability, which can affect terrestrial productivity and climate change feedbacks. While many single-factor studies have examined the effects of environmental changes on N mineralization and nitrification, few have examined these effects in a multifactor context or recorded how these effects vary seasonally. In an old-field ecosystem in Massachusetts, USA, we investigated the combined effects of four levels of warming (up to 4 °C) and three levels of precipitation (drought, ambient, and wet) on net N mineralization, net nitrification, and potential nitrification. We also examined the treatment effects on the temperature sensitivity of net N mineralization and net nitrification and on the ratio of C mineralization to net N mineralization. During winter, freeze-thaw events, snow depth, and soil freezing depth explained little of the variation in net nitrification and N mineralization rates among treatments. During two years of treatments, warming and altered precipitation rarely influenced the rates of N cycling, and there was no evidence of a seasonal pattern in the responses. In contrast, warming and drought dramatically decreased the apparent Q10 of net N mineralization and net nitrification, and the warming-induced decrease in apparent Q10 was more pronounced in ambient and wet treatments than the drought treatment. The ratio of C mineralization to net N mineralization varied over time and was sensitive to the interactive effects of warming and altered precipitation. Although many studies have found that warming tends to accelerate N cycling, our results suggest that warming can have little to no effect on N cycling in some ecosystems. Thus, ecosystem models that assume that warming will consistently increase N mineralization rates and inputs of plant-available N may overestimate the increase in terrestrial productivity and the magnitude of an important negative feedback to climate change.

Phenolic profile within the fine‐root branching orders of an evergreen species highlights a disconnect in root tissue quality predicted by elemental‐ and molecular‐level carbon composition

Fine roots constitute a significant source of plant productivity and litter turnover across terrestrial ecosystems, but less is known about the quantitative and qualitative profile of phenolic compounds within the fine-root architecture, which could regulate the potential contribution of plant roots to the soil organic matter pool. To understand the linkage between traditional macro-elemental and morphological traits of roots and their molecular-level carbon chemistry, we analyzed seasonal variations in monomeric yields of the free, bound, and lignin phenols in fine roots (distal five orders) and leaves of Ardisia quinquegona. Fine roots contained two-fold higher concentrations of bound phenols and three-fold higher concentrations of lignin phenols than leaves. Within fine roots, the concentrations of free and bound phenols decreased with increasing root order, and seasonal variation in the phenolic profile was more evident in lower order than in higher order roots. The morphological and macro-elemental root traits were decoupled from the quantity, composition and tissue association of phenolic compounds, revealing the potential inability of these traditional parameters to capture the molecular identity of phenolic carbon within the fine-root architecture and between fine roots and leaves. Our results highlight the molecular-level heterogeneity in phenolic carbon composition within the fine-root architecture, and imply that traits that capture the molecular identity of the root construct might better predict the decomposition dynamics within fine-root orders.

Warming and drought differentially influence the production and resorption of elemental and metabolic nitrogen pools in Quercus rubra.

The process of nutrient retranslocation from plant leaves during senescence subsequently affects both plant growth and soil nutrient cycling; changes in either of these could potentially feed back to climate change. Although elemental nutrient resorption has been shown to respond modestly to temperature and precipitation, we know remarkably little about the influence of increasing intensities of drought and warming on the resorption of different classes of plant metabolites. We studied the effect of warming and altered precipitation on the production and resorption of metabolites in Quercus rubra. The combination of warming and drought produced a higher abundance of compounds that can help to mitigate climatic stress by functioning as osmoregulators and antioxidants, including important intermediaries of the tricarboxylic acid (TCA) cycle, amino acids including proline and citrulline, and polyamines such as putrescine. Resorption efficiencies (REs) of extractable metabolites surprisingly had opposite responses to drought and warming; drought treatments generally increased RE of metabolites compared to ambient and wet treatments, while warming decreased RE. However, RE of total N differed markedly from that of extractable metabolites such as amino acids; for instance, droughted plants resorbed a smaller fraction of elemental N from their leaves than plants exposed to the ambient control. In contrast, plants in drought treatment resorbed amino acids more efficiently (>90%) than those in ambient (65-77%) or wet (42-58%) treatments. Across the climate treatments, the RE of elemental N correlated negatively with tissue tannin concentration, indicating that polyphenols produced in leaves under climatic stress could interfere with N resorption. Thus, senesced leaves from drier conditions might have a lower nutritive value to soil heterotrophs during the initial stages of litter decomposition despite a higher elemental N content of these tissues. Our results suggest that N resorption may be controlled not only by plant demand, but also by climatic influences on the production and resorption of plant metabolites. As climate-carbon models incorporate increasingly sophisticated nutrient cycles, these results highlight the need to adequately understand plant physiological responses to climatic variables.

Decoupling the direct and indirect effects of climate on plant litter decomposition and terrestrial nutrient cycling

Decomposition of plant litter is a fundamental ecosystem process that can act as a feedback to climate change by simultaneously influencing both the productivity of ecosystems and the flux of carbon dioxide from the soil. The influence of climate on decomposition from a postsenescence perspective is relatively well known; in particular, climate is known to regulate the rate of litter decomposition via its direct influence on the reaction kinetics and microbial physiology on processes downstream of tissue senescence. Climate can alter plant metabolism during the formative stage of tissues and could shape the final chemical composition of plant litter that is available for decomposition, and thus indirectly influence decomposition; however, these indirect effects are relatively poorly understood. Climatic stress disrupts cellular homeostasis in plants and results in the reprogramming of primary and secondary metabolic pathways, which leads to changes in the quantity, composition, and organization of small molecules and recalcitrant heteropolymers, including lignins, tannins, suberins, and cuticle within the plant tissue matrix. Furthermore, by regulating metabolism during tissue senescence, climate influences the resorption of nutrients from senescing tissues. Thus, the final chemical composition of plant litter that forms the substrate of decomposition is a combined product of presenescence physiological processes through the production and resorption of metabolites. The changes in quantity, composition, and localization of the molecular construct of the litter could enhance or hinder tissue decomposition and soil nutrient cycling by altering the recalcitrance of the lignocellulose matrix, the composition of microbial communities, and the activity of microbial exo-enzymes via various complexation reactions. Also, the climate-induced changes in the molecular composition of litter could differentially influence litter decomposition and soil nutrient cycling. Compared with temperate ecosystems, the indirect effects of climate on litter decomposition in the tropics are not well understood, which underscores the need to conduct additional studies in tropical biomes. We also emphasize the need to focus on how climatic stress affects the root chemistry as roots contribute significantly to biogeochemical cycling, and on utilizing more robust analytical approaches to capture the molecular composition of tissue matrix that fuel microbial metabolism.

Plant–soil interactions regulate the identity of soil carbon in invaded ecosystems: implication for legacy effects

Introduced, invasive plants can alter local soil chemistry and microbial communities, but the underlying mechanisms and extent of these changes are largely unknown. Based on characteristics associated with invasiveness in plants, it was hypothesized that introduced species that produce large amounts of litter with distinctive secondary compounds can a) alter the chemistry of both extractable and bulk carbon in the soil, b) shift microbial communities towards microbes better able to metabolize the compounds in the litter and c) cause soil carbon chemistry and microbial communities to shift to relatively uniform, novel states at multiple sites. Composition of phenolics in senescent tissues (leaves and roots) of Polygonum cuspidatum was compared to the composition of extractable phenolics and non-extractable bulk organic carbon in soils under and adjacent to large, long-established stands of P.cuspidatum at four sites in the eastern U.S. Rates of degradation of phenolics, activities of enzymes associated with the breakdown of phenolics and shifts in microbial community composition were also measured at the sites. Soils under P.cuspidatum stands contained twice as much phenolics as adjacent soils, but the composition of phenolics differed greatly between soils under stands and senescent tissues of P.cuspidatum. Flavonoids and proanthocyanidins constituted >90% of the identified phenolics in P.cuspidatum tissues, whereas monophenolic compounds accounted for >90% of the phenolics in soils under stands. Soils under and adjacent to stands also exhibited distinctive compositions of relatively persistent bulk organic carbon; composition differed less between soils under stands at different sites than between soils under and adjacent to stands at the same site. Soils under P.cuspidatum had 28 times greater abundance of fungi than soils adjacent to stands, and fungal markers showed clear separation of soils under and adjacent to P.cuspidatum. However, the potential activity of enzymes that degrade polyphenols was lower in soils under stands. Exogenously applied, chemically complex polyphenols persisted in both P.cuspidatum-invaded and adjacent non-invaded soils, whereas less complex compounds rapidly disappeared from both soils.Synthesis. Results suggest that interactions between plant inputs, abiotic reactions and biotic transformations may create and maintain new states in invaded soils that are chemically and biologically less diverse. In the case of polyphenol-rich, fast-growing invasive species, these interactions may alter the composition of bulk soil organic matter that has relatively slower turnover rates, resulting in legacy effects. Restoration could thus require, not just removal of the species, but also post-removal interventions such as soil amendments.

Litters of photosynthetically divergent grasses exhibit differential metabolic responses to warming and elevated CO2

Climatic stress such as warming would alter physiological pathways in plants leading to changes in tissue chemistry. Elevated CO2 could partly mitigate warming induced moisture stress, and the degree of this mitigation may vary with plant functional types. We studied the composition of structural and non-structural metabolites in senesced tissues of Bouteloua gracilis (C4) and Pascopyrum smithii (C3) at the Prairie Heating and CO2 Enrichment experiment, Wyoming, USA. We hypothesized that P. smithii and B. gracilis would respond to unfavorable global change factors by producing structural metabolites and osmoregulatory compounds that are necessary to combat stress. However, due to the inherent variation in the tolerance of their photosynthetic pathways to warming and CO2, we hypothesized that these species will exhibit differential response under different combinations of warming and CO2 conditions. Due to a lower thermo-tolerance of the C4 photosynthesis we expected B. gracilis to exhibit a greater metabo...