Alexandra R. Contosta

Stoichiometry of soil enzyme activity at global scale

Extracellular enzymes are the proximate agents of organic matter decomposition and measures of these activities can be used as indicators of microbial nutrient demand. We conducted a global-scale meta-analysis of the seven-most widely measured soil enzyme activities, using data from 40 ecosystems. The activities of beta-1,4-glucosidase, cellobiohydrolase, beta-1,4-N-acetylglucosaminidase and phosphatase g(-1) soil increased with organic matter concentration; leucine aminopeptidase, phenol oxidase and peroxidase activities showed no relationship. All activities were significantly related to soil pH. Specific activities, i.e. activity g(-1) soil organic matter, also varied in relation to soil pH for all enzymes. Relationships with mean annual temperature (MAT) and precipitation (MAP) were generally weak. For hydrolases, ratios of specific C, N and P acquisition activities converged on 1 : 1 : 1 but across ecosystems, the ratio of C : P acquisition was inversely related to MAP and MAT while the ratio of C : N acquisition increased with MAP. Oxidative activities were more variable than hydrolytic activities and increased with soil pH. Our analyses indicate that the enzymatic potential for hydrolyzing the labile components of soil organic matter is tied to substrate availability, soil pH and the stoichiometry of microbial nutrient demand. The enzymatic potential for oxidizing the recalcitrant fractions of soil organic material, which is a proximate control on soil organic matter accumulation, is most strongly related to soil pH. These trends provide insight into the biogeochemical processes that create global patterns in ecological stoichiometry and organic matter storage.

Seasonal dynamics of soil respiration and N mineralization in chronically warmed and fertilized soils

Although numerous studies have examined the individual effects of increased temperatures and N deposition on soil biogeochemical cycling, few have considered how these disturbances interact to impact soil C and N dynamics. Likewise, many have not assessed season-specific responses to warming and N inputs despite seasonal variability in soil processes. We studied interactions among season, warming, and N additions on soil respiration and N mineralization at the Soil Warming × Nitrogen Addition Study at the Harvard Forest. Of particular interest were wintertime fluxes of C and N typically excluded from investigations of soils and global change. Soils were warmed to 5°C above ambient, and N was applied at a rate of 5 g m−2 y−1. Soil respiration and N mineralization were sampled over two years between 2007 and 2009 and showed strong seasonal patterns that mirrored changes in soil temperature. Winter fluxes of C and N contributed between 2 and 17% to the total annual flux. Net N mineralization increased in response to the experimental manipulations across all seasons, and was 8% higher in fertilized plots and 83% higher in warmed plots over the duration of the study. Soil respiration showed a more season-specific response. Nitrogen additions enhanced soil respiration by 14%, but this increase was significant only in summer and fall. Likewise, warming increased soil respiration by 44% over the whole study period, but the effect of warming was most pronounced in spring and fall. The only interaction between warming × N additions took place in autumn, when N availability likely diminished the positive effect of warming on soil respiration. Our results suggest that winter measurements of C and N are necessary to accurately describe winter biogeochemical processes. In addition, season-specific responses to the experimental treatments suggest that some components of the belowground community may be more susceptible to warming and N additions than others. Seasonal changes in the abiotic environment may have also interacted with the experimental manipulations to evoke biogeochemical responses at certain times of year.

A longer vernal window: the role of winter coldness and snowpack in driving spring transitions and lags

Climate change is altering the timing and duration of the vernal window, a period that marks the end of winter and the start of the growing season when rapid transitions in ecosystem energy, water, nutrient, and carbon dynamics take place. Research on this period typically captures only a portion of the ecosystem in transition and focuses largely on the dates by which the system wakes up. Previous work has not addressed lags between transitions that represent delays in energy, water, nutrient, and carbon flows. The objectives of this study were to establish the sequence of physical and biogeochemical transitions and lags during the vernal window period and to understand how climate change may alter them. We synthesized observations from a statewide sensor network in New Hampshire, USA, that concurrently monitored climate, snow, soils, and streams over a three-year period and supplemented these observations with climate reanalysis data, snow data assimilation model output, and satellite spectral data. We found that some of the transitions that occurred within the vernal window were sequential, with air temperatures warming prior to snow melt, which preceded forest canopy closure. Other transitions were simultaneous with one another and had zero-length lags, such as snowpack disappearance, rapid soil warming, and peak stream discharge. We modeled lags as a function of both winter coldness and snow depth, both of which are expected to decline with climate change. Warmer winters with less snow resulted in longer lags and a more protracted vernal window. This lengthening of individual lags and of the entire vernal window carries important consequences for the thermodynamics and biogeochemistry of ecosystems, both during the winter-to-spring transition and throughout the rest of the year.

Winter soil respiration in a humid temperate forest: The roles of moisture, temperature, and snowpack

Winter soil respiration at midlatitudes can comprise a substantial portion of annual ecosystem carbon loss. However, winter soil carbon dynamics in these areas, which are often characterized by shallow snow cover, are poorly understood due to infrequent sampling at the soil surface. Our objectives were to continuously measure winter CO2 flux from soils and the overlying snowpack while also monitoring drivers of winter soil respiration in a humid temperate forest. We show that the relative roles of soil temperature and moisture in driving winter CO2 flux differed within a single soil-to-snow profile. Surface soil temperatures had a strong, positive influence on CO2 flux from the snowpack, while soil moisture exerted a negative control on soil CO2 flux within the soil profile. Rapid fluctuations in snow depth throughout the winter likely created the dynamic soil temperature and moisture conditions that drove divergent patterns in soil respiration at different depths. Such dynamic conditions differ from many previous studies of winter soil microclimate and respiration, where soil temperature and moisture are relatively stable until snowmelt. The differential response of soil respiration to temperature and moisture across depths was also a unique finding as previous work has not simultaneously quantified CO2 flux from soils and the snowpack. The complex interplay we observed among snow depth, soil temperature, soil moisture, and CO2 flux suggests that winter soil respiration in areas with shallow seasonal snow cover is more variable than previously understood and may fluctuate considerably in the future given winter climate change.

Minerals in the rhizosphere: overlooked mediators of soil nitrogen availability to plants and microbes

Despite decades of research progress, ecologists are still debating which pools and fluxes provide nitrogen (N) to plants and soil microbes across different ecosystems. Depolymerization of soil organic N is recognized as the rate-limiting step in the production of bioavailable N, and it is generally assumed that detrital N is the main source. However, in many mineral soils, detrital polymers constitute a minor fraction of total soil organic N. The majority of organic N is associated with clay-sized particles where physicochemical interactions may limit the accessibility of N-containing compounds. Although mineral-associated organic matter (MAOM) has historically been considered a critical, but relatively passive, reservoir of soil N, a growing body of research now points to the dynamic nature of mineral-organic associations and their potential for destabilization. Here we synthesize evidence from biogeoscience and soil ecology to demonstrate how MAOM is an important, yet overlooked, mediator of bioavailable N, especially in the rhizosphere. We highlight several biochemical strategies that enable plants and microbes to disrupt mineral-organic interactions and access MAOM. In particular, root-deposited low-molecular-weight exudates may enhance the mobilization and solubilization of MAOM, increasing its bioavailability. However, the competitive balance between the possible fates of N monomers—bound to mineral surfaces versus dissolved and available for assimilation—will depend on the specific interaction between mineral properties, soil solution, mineral-bound organic matter, and microbes. Building off our emerging understanding of MAOM as a source of bioavailable N, we propose a revision of the Schimel and Bennett (Ecology 85:591–602, 2004) model (which emphasizes N depolymerization), by incorporating MAOM as a potential proximal mediator of bioavailable N.