Multiscale predictors of small tree survival across a heterogeneous tropical landscape.

Uncertainties about controls on tree mortality make forest responses to land-use and climate change difficult to predict. We tracked biomass of tree functional groups in tropical forest inventories across Puerto Rico and the U.S. Virgin Islands, and with random forests we ranked 86 potential predictors of small tree survival (young or mature stems 2.5-12.6 cm diameter at breast height). Forests span dry to cloud forests, range in age, geology and past land use and experienced severe drought and storms. When excluding species as a predictor, top predictors are tree crown ratio and height, two to three species traits and stand to regional factors reflecting local disturbance and the system state (widespread recovery, drought, hurricanes). Native species, and species with denser wood, taller maximum height, or medium typical height survive longer, but short trees and species survive hurricanes better. Trees survive longer in older stands and with less disturbed canopies, harsher geoclimates (dry, edaphically dry, e.g., serpentine substrates, and highest-elevation cloud forest), or in intervals removed from hurricanes. Satellite image phenology and bands, even from past decades, are top predictors, being sensitive to vegetation type and disturbance. Covariation between stand-level species traits and geoclimate, disturbance and neighboring species types may explain why most neighbor variables, including introduced vs. native species, had low or no importance, despite univariate correlations with survival. As forests recovered from a hurricane in 1998 and earlier deforestation, small trees of introduced species, which on average have lighter wood, died at twice the rate of natives. After hurricanes in 2017, the total biomass of trees ≥12.7 cm dbh of the introduced species Spathodea campanulata spiked, suggesting that more frequent hurricanes might perpetuate this light-wooded species commonness. If hurricane recovery favors light-wooded species while drought favors others, climate change influences on forest composition and ecosystem services may depend on the frequency and severity of extreme climate events.

Experimental warming and its legacy effects on root dynamics following two hurricane disturbances in a wet tropical forest.

Tropical forests are expected to experience unprecedented warming and increases in hurricane disturbances in the coming decades; yet, our understanding of how these productive systems, especially their belowground component, will respond to the combined effects of varied environmental changes remains empirically limited. Here we evaluated the responses of root dynamics (production, mortality, and biomass) to soil and understory warming (+4°C) and after two consecutive tropical hurricanes in our in situ warming experiment in a tropical forest of Puerto Rico: Tropical Responses to Altered Climate Experiment (TRACE). We collected minirhizotron images from three warmed plots and three control plots of 12 m2 . Following Hurricanes Irma and María in September 2017, the infrared heater warming treatment was suspended for repairs, which allowed us to explore potential legacy effects of prior warming on forest recovery. We found that warming significantly reduced root production and root biomass over time. Following hurricane disturbance, both root biomass and production increased substantially across all plots; the root biomass increased 2.8-fold in controls but only 1.6-fold in previously warmed plots. This pattern held true for both herbaceous and woody roots, suggesting that the consistent antecedent warming conditions reduced root capacity to recover following hurricane disturbance. Root production and mortality were both related to soil ammonium nitrogen and microbial biomass nitrogen before and after the hurricanes. This experiment has provided an unprecedented look at the complex interactive effects of disturbance and climate change on the root component of a tropical forested ecosystem. A decrease in root production in a warmer world and slower root recovery after a major hurricane disturbance, as observed here, are likely to have longer-term consequences for tropical forest responses to future global change.

Habitability Models for Astrobiology.

Habitability has been generally defined as the capability of an environment to support life. Ecologists have been using Habitat Suitability Models (HSMs) for more than four decades to study the habitability of Earth from local to global scales. Astrobiologists have been proposing different habitability models for some time, with little integration and consistency among them, being different in function to those used by ecologists. Habitability models are not only used to determine whether environments are habitable, but they also are used to characterize what key factors are responsible for the gradual transition from low to high habitability states. Here we review and compare some of the different models used by ecologists and astrobiologists and suggest how they could be integrated into new habitability standards. Such standards will help improve the comparison and characterization of potentially habitable environments, prioritize target selections, and study correlations between habitability and biosignatures. Habitability models are the foundation of planetary habitability science, and the synergy between ecologists and astrobiologists is necessary to expand our understanding of the habitability of Earth, the Solar System, and extrasolar planets.

Experimental warming across a tropical forest canopy height gradient reveals minimal photosynthetic and respiratory acclimation.

Tropical forest canopies cycle vast amounts of carbon, yet we still have a limited understanding of how these critical ecosystems will respond to climate warming. We implemented in situ leaf-level + 3°C experimental warming from the understory to the upper canopy of two Puerto Rican tropical tree species, Guarea guidonia and Ocotea sintenisii. After approximately 1 month of continuous warming, we assessed adjustments in photosynthesis, chlorophyll fluorescence, stomatal conductance, leaf traits and foliar respiration. Warming did not alter net photosynthetic temperature response for either species; however, the optimum temperature of Ocotea understory leaf photosynthetic electron transport shifted upward. There was no Ocotea respiratory treatment effect, while Guarea respiratory temperature sensitivity (Q10 ) was down-regulated in heated leaves. The optimum temperatures for photosynthesis (Topt ) decreased 3-5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations. Guarea upper canopy Topt was similar to the mean daytime temperatures, while Ocotea canopy leaves often operated above Topt . With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO2 uptake, which could weaken the overall carbon sink strength of this tropical forest.

Large seasonal variation of soil respiration in a secondary tropical moist forest in Puerto Rico.

Tropical forests are the largest contributors to global emissions of carbon dioxide (CO2) to the atmosphere via soil respiration (R s). As such, identifying the main controls on R s in tropical forests is essential for accurately projecting the consequences of ongoing and future global environmental changes to the global C cycle. We measured hourly R s in a secondary tropical moist forest in Puerto Rico over a 3-year period to (a) quantify the magnitude of R s and (b) identify the role of climatic, substrate, and nutrient controls on the seasonality of R s. Across 3 years of measurements, mean R s was 7.16 ± 0.02 µmol CO2 m-2 s-1 (or 2,710 g C m-2 year-1) and showed significant seasonal variation. Despite small month-to-month variation in temperature (~4°C), we found significant positive relationships between daily and monthly R s with both air and soil temperature, highlighting the importance of temperature as a driver of R s even in warm ecosystems, such as tropical forests. We also found a significant parabolic relationship between mean daily volumetric soil moisture and mean daily R s, with an optimal moisture value of 0.34 m3 m-3. Given the relatively consistent climate at this site, the large range in mean monthly R s (~7 µmol CO2 m-2 s-1) was surprising and suggests that even small changes in climate can have large implications for ecosystem respiration. The strong positive relationship of R s with temperature at monthly timescales particularly stands out, as moisture is usually considered a stronger control of R s in tropical forests that already experience warm temperatures year-round. Moreover, our results revealed the strong seasonality of R s in tropical moist forests, which given its high magnitude, can represent a significant contribution to the seasonal patterns of atmospheric (CO2) globally.

Tropical understory herbaceous community responds more strongly to hurricane disturbance than to experimental warming.

The effects of climate change on tropical forests may have global consequences due to the forests' high biodiversity and major role in the global carbon cycle. In this study, we document the effects of experimental warming on the abundance and composition of a tropical forest floor herbaceous plant community in the Luquillo Experimental Forest, Puerto Rico. This study was conducted within Tropical Responses to Altered Climate Experiment (TRACE) plots, which use infrared heaters under free-air, open-field conditions, to warm understory vegetation and soils + 4°C above nearby control plots. Hurricanes Irma and María damaged the heating infrastructure in the second year of warming, therefore, the study included one pretreatment year, one year of warming, and one year of hurricane response with no warming. We measured percent leaf cover of individual herbaceous species, fern population dynamics, and species richness and diversity within three warmed and three control plots. Results showed that one year of experimental warming did not significantly affect the cover of individual herbaceous species, fern population dynamics, species richness, or species diversity. In contrast, herbaceous cover increased from 20% to 70%, bare ground decreased from 70% to 6%, and species composition shifted pre to posthurricane. The negligible effects of warming may have been due to the short duration of the warming treatment or an understory that is somewhat resistant to higher temperatures. Our results suggest that climate extremes that are predicted to increase with climate change, such as hurricanes and droughts, may cause more abrupt changes in tropical forest understories than longer-term sustained warming.

Altered climate leads to positive density-dependent feedbacks in a tropical wet forest.

Climate change is predicted to result in warmer and drier Neotropical forests relative to current conditions. Negative density-dependent feedbacks, mediated by natural enemies, are key to maintaining the high diversity of tree species found in the tropics, yet we have little understanding of how projected changes in climate are likely to affect these critical controls. Over 3 years, we evaluated the effects of a natural drought and in situ experimental warming on density-dependent feedbacks on seedling demography in a wet tropical forest in Puerto Rico. In the +4°C warming treatment, we found that seedling survival increased with increasing density of the same species (conspecific). These positive density-dependent feedbacks were not associated with a decrease in aboveground natural enemy pressure. If positive density-dependent feedbacks are not transient, the diversity of tropical wet forests, which may rely on negative density dependence to drive diversity, could decline in a future warmer, drier world.

Precipitation mediates sap flux sensitivity to evaporative demand in the neotropics.

Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

Infrared heater system for warming tropical forest understory plants and soils.

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4-m-diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0-10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40-50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50-cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.

Belowground Response to Drought in a Tropical Forest Soil. I. Changes in Microbial Functional Potential and Metabolism.

Global climate models predict a future of increased severity of drought in many tropical forests. Soil microbes are central to the balance of these systems as sources or sinks of atmospheric carbon (C), yet how they respond metabolically to drought is not well-understood. We simulated drought in the typically aseasonal Luquillo Experimental Forest, Puerto Rico, by intercepting precipitation falling through the forest canopy. This approach reduced soil moisture by 13% and water potential by 0.14 MPa (from -0.2 to -0.34). Previous results from this experiment have demonstrated that the diversity and composition of these soil microbial communities are sensitive to even small changes in soil water. Here, we show prolonged drought significantly alters the functional potential of the community and provokes a clear osmotic stress response, including the production of compatible solutes that increase intracellular C demand. Subsequently, a microbial population emerges with a greater capacity for extracellular enzyme production targeting macromolecular carbon. Significantly, some of these drought-induced functional shifts in the soil microbiota are attenuated by prior exposure to a short-term drought suggesting that acclimation may occur despite a lack of longer-term drought history.

Belowground Response to Drought in a Tropical Forest Soil. II. Change in Microbial Function Impacts Carbon Composition.

Climate model projections for tropical regions show clear perturbation of precipitation patterns leading to increased frequency and severity of drought in some regions. Previous work has shown declining soil moisture to be a strong driver of changes in microbial trait distribution, however, the feedback of any shift in functional potential on ecosystem properties related to carbon cycling are poorly understood. Here we show that drought-induced changes in microbial functional diversity and activity shape, and are in turn shaped by, the composition of dissolved and soil-associated carbon. We also demonstrate that a shift in microbial functional traits that favor the production of hygroscopic compounds alter the efflux of carbon dioxide following soil rewetting. Under drought the composition of the dissolved organic carbon pool changed in a manner consistent with a microbial metabolic response. We hypothesize that this microbial ecophysiological response to changing soil moisture elevates the intracellular carbon demand stimulating extracellular enzyme production, that prompts the observed decline in more complex carbon compounds (e.g., cellulose and lignin). Furthermore, a metabolic response to drought appeared to condition (biologically and physically) the soil, notably through the production of polysaccharides, particularly in experimental plots that had been pre-exposed to a short-term drought. This hysteretic response, in addition to an observed drought-related decline in phosphorus concentration, may have been responsible for a comparatively modest CO2 efflux following wet-up in drought plots relative to control plots.

Urgent need for warming experiments in tropical forests.

Although tropical forests account for only a fraction of the planet's terrestrial surface, they exchange more carbon dioxide with the atmosphere than any other biome on Earth, and thus play a disproportionate role in the global climate. In the next 20 years, the tropics will experience unprecedented warming, yet there is exceedingly high uncertainty about their potential responses to this imminent climatic change. Here, we prioritize research approaches given both funding and logistical constraints in order to resolve major uncertainties about how tropical forests function and also to improve predictive capacity of earth system models. We investigate overall model uncertainty of tropical latitudes and explore the scientific benefits and inevitable trade-offs inherent in large-scale manipulative field experiments. With a Coupled Model Intercomparison Project Phase 5 analysis, we found that model variability in projected net ecosystem production was nearly 3 times greater in the tropics than for any other latitude. Through a review of the most current literature, we concluded that manipulative warming experiments are vital to accurately predict future tropical forest carbon balance, and we further recommend the establishment of a network of comparable studies spanning gradients of precipitation, edaphic qualities, plant types, and/or land use change. We provide arguments for long-term, single-factor warming experiments that incorporate warming of the most biogeochemically active ecosystem components (i.e. leaves, roots, soil microbes). Hypothesis testing of underlying mechanisms should be a priority, along with improving model parameterization and constraints. No single tropical forest is representative of all tropical forests; therefore logistical feasibility should be the most important consideration for locating large-scale manipulative experiments. Above all, we advocate for multi-faceted research programs, and we offer arguments for what we consider the most powerful and urgent way forward in order to improve our understanding of tropical forest responses to climate change.

Threshold Responses to Soil Moisture Deficit by Trees and Soil in Tropical Rain Forests: Insights from Field Experiments.

Many tropical rain forest regions are at risk of increased future drought. The net effects of drought on forest ecosystem functioning will be substantial if important ecological thresholds are passed. However, understanding and predicting these effects is challenging using observational studies alone. Field-based rainfall exclusion (canopy throughfall exclusion; TFE) experiments can offer mechanistic insight into the response to extended or severe drought and can be used to help improve model-based simulations, which are currently inadequate. Only eight TFE experiments have been reported for tropical rain forests. We examine them, synthesizing key results and focusing on two processes that have shown threshold behavior in response to drought: (1) tree mortality and (2) the efflux of carbon dioxdie from soil, soil respiration. We show that: (a) where tested using large-scale field experiments, tropical rain forest tree mortality is resistant to long-term soil moisture deficit up to a threshold of 50% of the water that is extractable by vegetation from the soil, but high mortality occurs beyond this value, with evidence from one site of increased autotrophic respiration, and (b) soil respiration reaches its peak value in response to soil moisture at significantly higher soil moisture content for clay-rich soils than for clay-poor soils. This first synthesis of tropical TFE experiments offers the hypothesis that low soil moisture-related thresholds for key stress responses in soil and vegetation may prove to be widely applicable across tropical rain forests despite the diversity of these forests.

Aboveground tree growth varies with belowground carbon allocation in a tropical rainforest environment.

Young secondary forests and plantations in the moist tropics often have rapid rates of biomass accumulation and thus sequester large amounts of carbon. Here, we compare results from mature forest and nearby 15-20 year old tree plantations in lowland Costa Rica to evaluate differences in allocation of carbon to aboveground production and root systems. We found that the tree plantations, which had fully developed, closed canopies, allocated more carbon belowground - to their root systems - than did mature forest. This increase in belowground carbon allocation correlated significantly with aboveground tree growth but not with canopy production (i.e., leaf fall or fine litter production). In contrast, there were no correlations between canopy production and either tree growth or belowground carbon allocation. Enhanced allocation of carbon to root systems can enhance plant nutrient uptake, providing nutrients beyond those required for the production of short-lived tissues such as leaves and fine roots, and thus enabling biomass accumulation. Our analyses support this deduction at our site, showing that enhanced allocation of carbon to root systems can be an important mechanism promoting biomass accumulation during forest growth in the moist tropics. Identifying factors that control when, where and for how long this occurs would help us to improve models of forest growth and nutrient cycling, and to ascertain the role that young forests play in mitigating increased atmospheric carbon dioxide.

Sensitivity of soil respiration to variability in soil moisture and temperature in a humid tropical forest.

Precipitation and temperature are important drivers of soil respiration. The role of moisture and temperature are generally explored at seasonal or inter-annual timescales; however, significant variability also occurs on hourly to daily time-scales. We used small (1.54 m(2)), throughfall exclusion shelters to evaluate the role soil moisture and temperature as temporal controls on soil CO2 efflux from a humid tropical forest in Puerto Rico. We measured hourly soil CO2 efflux, temperature and moisture in control and exclusion plots (n = 6) for 6-months. The variance of each time series was analyzed using orthonormal wavelet transformation and Haar-wavelet coherence. We found strong negative coherence between soil moisture and soil respiration in control plots corresponding to a two-day periodicity. Across all plots, there was a significant parabolic relationship between soil moisture and soil CO2 efflux with peak soil respiration occurring at volumetric soil moisture of approximately 0.375 m(3)/m(3). We additionally found a weak positive coherence between CO2 and temperature at longer time-scales and a significant positive relationship between soil temperature and CO2 efflux when the analysis was limited to the control plots. The coherence between CO2 and both temperature and soil moisture were reduced in exclusion plots. The reduced CO2 response to temperature in exclusion plots suggests that the positive effect of temperature on CO2 is constrained by soil moisture availability.

Pre-exposure to drought increases the resistance of tropical forest soil bacterial communities to extended drought.

Global climate models project a decrease in the magnitude of precipitation in tropical regions. Changes in rainfall patterns have important implications for the moisture content and redox status of tropical soils, yet little is known about how these changes may affect microbial community structure. Specifically, does exposure to prior stress confer increased resistance to subsequent perturbation? Here we reduced the quantity of precipitation throughfall to tropical forest soils in the Luquillo Mountains, Puerto Rico. Treatments included newly established throughfall exclusion plots (de novo excluded), plots undergoing reduction for a second time (pre-excluded) and ambient control plots. Ten months of throughfall exclusion led to a small but statistically significant decline in soil water potential and bacterial populations clearly adapted to increased osmotic stress. Although the water potential decline was small and microbial biomass did not change, phylogenetic diversity in the de novo-excluded plots decreased by ∼40% compared with the control plots, yet pre-excluded plots showed no significant change. On the other hand, the relative abundances of bacterial taxa in both the de novo-excluded and pre-excluded plots changed significantly with throughfall exclusion compared with control plots. Changes in bacterial community structure could be explained by changes in soil pore water chemistry and suggested changes in soil redox. Soluble iron declined in treatment plots and was correlated with decreased soluble phosphorus concentrations, which may have significant implications for microbial productivity in these P-limited systems.

Tropical forest carbon balance in a warmer world: a critical review spanning microbial- to ecosystem-scale processes.

Tropical forests play a major role in regulating global carbon (C) fluxes and stocks, and even small changes to C cycling in this productive biome could dramatically affect atmospheric carbon dioxide (CO(2) ) concentrations. Temperature is expected to increase over all land surfaces in the future, yet we have a surprisingly poor understanding of how tropical forests will respond to this significant climatic change. Here we present a contemporary synthesis of the existing data and what they suggest about how tropical forests will respond to increasing temperatures. Our goals were to: (i) determine whether there is enough evidence to support the conclusion that increased temperature will affect tropical forest C balance; (ii) if there is sufficient evidence, determine what direction this effect will take; and, (iii) establish what steps should to be taken to resolve the uncertainties surrounding tropical forest responses to increasing temperatures. We approach these questions from a mass-balance perspective and therefore focus primarily on the effects of temperature on inputs and outputs of C, spanning microbial- to ecosystem-scale responses. We found that, while there is the strong potential for temperature to affect processes related to C cycling and storage in tropical forests, a notable lack of data combined with the physical, biological and chemical diversity of the forests themselves make it difficult to resolve this issue with certainty. We suggest a variety of experimental approaches that could help elucidate how tropical forests will respond to warming, including large-scale in situ manipulation experiments, longer term field experiments, the incorporation of a range of scales in the investigation of warming effects (both spatial and temporal), as well as the inclusion of a diversity of tropical forest sites. Finally, we highlight areas of tropical forest research where notably few data are available, including temperature effects on: nutrient cycling, heterotrophic versus autotrophic respiration, thermal acclimation versus substrate limitation of plant and microbial communities, below-ground C allocation, species composition (plant and microbial), and the hydraulic architecture of roots. Whether or not tropical forests will become a source or a sink of C in a warmer world remains highly uncertain. Given the importance of these ecosystems to the global C budget, resolving this uncertainty is a primary research priority.

Rain forest nutrient cycling and productivity in response to large-scale litter manipulation.

Litter-induced pulses of nutrient availability could play an important role in the productivity and nutrient cycling of forested ecosystems, especially tropical forests. Tropical forests experience such pulses as a result of wet-dry seasonality and during major climatic events, such as strong El Niños. We hypothesized that (1) an increase in the quantity and quality of litter inputs would stimulate leaf litter production, woody growth, and leaf litter nutrient cycling, and (2) the timing and magnitude of this response would be influenced by soil fertility and forest age. To test these hypotheses in a Costa Rican wet tropical forest, we established a large-scale litter manipulation experiment in two secondary forest sites and four old-growth forest sites of differing soil fertility. In replicated plots at each site, leaves and twigs (< 2 cm diameter) were removed from a 400-m2 area and added to an adjacent 100-m2 area. This transfer was the equivalent of adding 5-25 kg/ha of organic P to the forest floor. We analyzed leaf litter mass, [N] and [P], and N and P inputs for addition, removal, and control plots over a two-year period. We also evaluated basal area increment of trees in removal and addition plots. There was no response of forest productivity or nutrient cycling to litter removal; however, litter addition significantly increased leaf litter production and N and P inputs 4-5 months following litter application. Litter production increased as much as 92%, and P and N inputs as much as 85% and 156%, respectively. In contrast, litter manipulation had no significant effect on woody growth. The increase in leaf litter production and N and P inputs were significantly positively related to the total P that was applied in litter form. Neither litter treatment nor forest type influenced the temporal pattern of any of the variables measured. Thus, environmental factors such as rainfall drive temporal variability in litter and nutrient inputs, while nutrient release from decomposing litter influences the magnitude. Seasonal or annual variation in leaf litter mass, such as occurs in strong El Niño events, could positively affect leaf litter nutrient cycling and forest productivity, indicating an ability of tropical trees to rapidly respond to increased nutrient availability.