Jennifer S. Powers

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.

Biomass resilience of Neotropical secondary forests

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha−1), corresponding to a net carbon uptake of 3.05 Mg C ha−1 yr−1, 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha−1) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Geographic bias of field observations of soil carbon stocks with tropical land-use changes precludes spatial extrapolation

Accurately quantifying changes in soil carbon (C) stocks with land-use change is important for estimating the anthropogenic fluxes of greenhouse gases to the atmosphere and for implementing policies such as REDD (Reducing Emissions from Deforestation and Degradation) that provide financial incentives to reduce carbon dioxide fluxes from deforestation and land degradation. Despite hundreds of field studies and at least a dozen literature reviews, there is still considerable disagreement on the direction and magnitude of changes in soil C stocks with land-use change. We conducted a meta-analysis of studies that quantified changes in soil C stocks with land use in the tropics. Conversion from one land use to another caused significant increases or decreases in soil C stocks for 8 of the 14 transitions examined. For the three land-use transitions with sufficient observations, both the direction and magnitude of the change in soil C pools depended strongly on biophysical factors of mean annual precipitation and dominant soil clay mineralogy. When we compared the distribution of biophysical conditions of the field observations to the area-weighted distribution of those factors in the tropics as a whole or the tropical lands that have undergone conversion, we found that field observations are highly unrepresentative of most tropical landscapes. Because of this geographic bias we strongly caution against extrapolating average values of land-cover change effects on soil C stocks, such as those generated through meta-analysis and literature reviews, to regions that differ in biophysical conditions.

Increased litterfall in tropical forests boosts the transfer of soil CO2 to the atmosphere.

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO2) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO2 production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this ‘extra’ CO2 was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.

Carbon sequestration potential of second-growth forest regeneration in the Latin American tropics

Models reveal the high carbon mitigation potential of tropical forest regeneration. Regrowth of tropical secondary forests following complete or nearly complete removal of forest vegetation actively stores carbon in aboveground biomass, partially counterbalancing carbon emissions from deforestation, forest degradation, burning of fossil fuels, and other anthropogenic sources. We estimate the age and spatial extent of lowland second-growth forests in the Latin American tropics and model their potential aboveground carbon accumulation over four decades. Our model shows that, in 2008, second-growth forests (1 to 60 years old) covered 2.4 million km2 of land (28.1% of the total study area). Over 40 years, these lands can potentially accumulate a total aboveground carbon stock of 8.48 Pg C (petagrams of carbon) in aboveground biomass via low-cost natural regeneration or assisted regeneration, corresponding to a total CO2 sequestration of 31.09 Pg CO2. This total is equivalent to carbon emissions from fossil fuel use and industrial processes in all of Latin America and the Caribbean from 1993 to 2014. Ten countries account for 95% of this carbon storage potential, led by Brazil, Colombia, Mexico, and Venezuela. We model future land-use scenarios to guide national carbon mitigation policies. Permitting natural regeneration on 40% of lowland pastures potentially stores an additional 2.0 Pg C over 40 years. Our study provides information and maps to guide national-level forest-based carbon mitigation plans on the basis of estimated rates of natural regeneration and pasture abandonment. Coupled with avoided deforestation and sustainable forest management, natural regeneration of second-growth forests provides a low-cost mechanism that yields a high carbon sequestration potential with multiple benefits for biodiversity and ecosystem services.

Lianas reduce carbon accumulation and storage in tropical forests

Significance Tropical forests store nearly 30% of global terrestrial carbon and contribute to 40% of the global terrestrial carbon sink. By affecting tree growth and survival, lianas impact the carbon balance of these forests. Here we demonstrate with a 3-y experiment that lianas substantially reduce forest-level carbon uptake and storage. This study is, to our knowledge, the first direct demonstration of liana effects at the ecosystem scale and illustrates the important role of lianas in tropical forests, particularly with respect to carbon budgets. Lianas are increasing in biomass and productivity throughout the tropics, and thus our findings have even greater relevance in terms of the fate of the tropical carbon balance, as well as for global atmospheric CO2 levels, in a changing climate. Tropical forests store vast quantities of carbon, account for one-third of the carbon fixed by photosynthesis, and are a major sink in the global carbon cycle. Recent evidence suggests that competition between lianas (woody vines) and trees may reduce forest-wide carbon uptake; however, estimates of the impact of lianas on carbon dynamics of tropical forests are crucially lacking. Here we used a large-scale liana removal experiment and found that, at 3 y after liana removal, lianas reduced net above-ground carbon uptake (growth and recruitment minus mortality) by ∼76% per year, mostly by reducing tree growth. The loss of carbon uptake due to liana-induced mortality was four times greater in the control plots in which lianas were present, but high variation among plots prevented a significant difference among the treatments. Lianas altered how aboveground carbon was stored. In forests where lianas were present, the partitioning of forest aboveground net primary production was dominated by leaves (53.2%, compared with 39.2% in liana-free forests) at the expense of woody stems (from 28.9%, compared with 43.9%), resulting in a more rapid return of fixed carbon to the atmosphere. After 3 y of experimental liana removal, our results clearly demonstrate large differences in carbon cycling between forests with and without lianas. Combined with the recently reported increases in liana abundance, these results indicate that lianas are an important and increasing agent of change in the carbon dynamics of tropical forests.

Biogeochemistry and Geographical Ecology: Embracing All Twenty-Five Elements Required to Build Organisms

Biogeochemistry is a key but relatively neglected part of the abiotic template that underlies ecology. The template has a geography, one that is increasingly being rearranged in this era of global change. Justus von Liebig’s law of the minimum has played a useful role in focusing attention on biogeochemical regulation of populations, but given that ∼25+ elements are required to build organisms and that these organisms use and deplete nutrients in aggregates of communities and ecosystems, we make the case that it is time to move on. We review available models that suggest the many different mechanisms that give rise to multiple elements, or colimitation. We then review recent empirical data that show that rates of decomposition and primary productivity may be limited by multiple elements. In that light, given the tropics’ high species diversity and generally more weathered soils, we predict that colimitation at community and ecosystem scales is more prevalent closer to the equator. We conclude with suggestions for how to move forward with experimental studies of colimitation.

Pervasive and strong effects of plants on soil chemistry: a meta-analysis of individual plant 'Zinke' effects.

Plant species leave a chemical signature in the soils below them, generating fine-scale spatial variation that drives ecological processes. Since the publication of a seminal paper on plant-mediated soil heterogeneity by Paul Zinke in 1962, a robust literature has developed examining effects of individual plants on their local environments (individual plant effects). Here, we synthesize this work using meta-analysis to show that plant effects are strong and pervasive across ecosystems on six continents. Overall, soil properties beneath individual plants differ from those of neighbours by an average of 41%. Although the magnitudes of individual plant effects exhibit weak relationships with climate and latitude, they are significantly stronger in deserts and tundra than forests, and weaker in intensively managed ecosystems. The ubiquitous effects of plant individuals and species on local soil properties imply that individual plant effects have a role in plant–soil feedbacks, linking individual plants with biogeochemical processes at the ecosystem scale.

Macro- and micronutrient effects on decomposition of leaf litter from two tropical tree species: inferences from a short-term laboratory incubation

While a large number of studies have investigated the effects of macronutrients such as nitrogen (N) or phosphorus (P) on litter decomposition, recent studies suggest that micronutrients including zinc (Zn) may also limit decomposition rates. Our goal was to compare the effects of nutrient addition on decomposition of two leaf litter types from tropical dry forest trees in a short-term laboratory microcosm experiment. Single nutrients (N, P, Zn, potassium, magnesium, and nickel) were applied to leaf litter in solution at low or high concentrations (to mimic in situ availability or to alleviate nutrient limitation, respectively), and decomposition was assessed as final mass remaining and carbon dioxide mineralization. Both mass remaining and CO2 mineralization were affected by nutrient identity and concentration, and these effects varied by species. In general, P and Zn addition increased decomposition, Mg and N inhibited it, and K and Ni had no significant effects. Future studies should consider the interactions between decomposition processes, decomposer communities, and a wider range of macro- and micronutrients.

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long-term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming-induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming-induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0-30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.