Decoupled dimensions of leaf economic and anti-herbivore defense strategies in a tropical canopy tree community.

Trade-offs among plant functional traits indicate diversity in plant strategies of growth and survival. The leaf economics spectrum (LES) reflects a trade-off between short-term carbon gain and long-term leaf persistence. A related trade-off, between foliar growth and anti-herbivore defense, occurs among plants growing in contrasting resource regimes, but it is unclear whether this trade-off is maintained within plant communities, where resource gradients are minimized. The LES and the growth-defense trade-off involve related traits, but the extent to which these trade-off dimensions are correlated is poorly understood. We assessed the relationship between leaf economic and anti-herbivore defense traits among sunlit foliage of 345 canopy trees in 83 species on Barro Colorado Island, Panama. We quantified ten traits related to resource allocation and defense, and identified patterns of trait co-variation using multivariate ordination. We tested whether traits and ordination axes were correlated with patterns of phylogenetic relatedness, juvenile demographic trade-offs, or topo-edaphic variation. Two independent axes described ~ 60% of the variation among canopy trees. Axis 1 revealed a trade-off between leaf nutritional and structural investment, consistent with the LES. Physical defense traits were largely oriented along this axis. Axis 2 revealed a trade-off between investments in phenolic defenses versus other foliar defenses, which we term the leaf defense spectrum. Phylogenetic relationships and topo-edaphic variation largely did not explain trait co-variation. Our results suggest that some trade-offs among the growth and defense traits of outer-canopy trees may be captured by the LES, while others may occur along additional resource allocation dimensions.

Greater transportation energy and GHG offsets from bioelectricity than ethanol.

The quantity of land available to grow biofuel crops without affecting food prices or greenhouse gas (GHG) emissions from land conversion is limited. Therefore, bioenergy should maximize land-use efficiency when addressing transportation and climate change goals. Biomass could power either internal combustion or electric vehicles, but the relative land-use efficiency of these two energy pathways is not well quantified. Here, we show that bioelectricity outperforms ethanol across a range of feedstocks, conversion technologies, and vehicle classes. Bioelectricity produces an average of 81% more transportation kilometers and 108% more emissions offsets per unit area of cropland than does cellulosic ethanol. These results suggest that alternative bioenergy pathways have large differences in how efficiently they use the available land to achieve transportation and climate goals.

Recent patterns and mechanisms of carbon exchange by terrestrial ecosystems.

Knowledge of carbon exchange between the atmosphere, land and the oceans is important, given that the terrestrial and marine environments are currently absorbing about half of the carbon dioxide that is emitted by fossil-fuel combustion. This carbon uptake is therefore limiting the extent of atmospheric and climatic change, but its long-term nature remains uncertain. Here we provide an overview of the current state of knowledge of global and regional patterns of carbon exchange by terrestrial ecosystems. Atmospheric carbon dioxide and oxygen data confirm that the terrestrial biosphere was largely neutral with respect to net carbon exchange during the 1980s, but became a net carbon sink in the 1990s. This recent sink can be largely attributed to northern extratropical areas, and is roughly split between North America and Eurasia. Tropical land areas, however, were approximately in balance with respect to carbon exchange, implying a carbon sink that offset emissions due to tropical deforestation. The evolution of the terrestrial carbon sink is largely the result of changes in land use over time, such as regrowth on abandoned agricultural land and fire prevention, in addition to responses to environmental changes, such as longer growing seasons, and fertilization by carbon dioxide and nitrogen. Nevertheless, there remain considerable uncertainties as to the magnitude of the sink in different regions and the contribution of different processes.

Consistent land- and atmosphere-based U.S. carbon sink estimates.

For the period 1980-89, we estimate a carbon sink in the coterminous United States between 0.30 and 0.58 petagrams of carbon per year (petagrams of carbon = 10(15) grams of carbon). The net carbon flux from the atmosphere to the land was higher, 0.37 to 0.71 petagrams of carbon per year, because a net flux of 0.07 to 0.13 petagrams of carbon per year was exported by rivers and commerce and returned to the atmosphere elsewhere. These land-based estimates are larger than those from previous studies (0.08 to 0.35 petagrams of carbon per year) because of the inclusion of additional processes and revised estimates of some component fluxes. Although component estimates are uncertain, about one-half of the total is outside the forest sector. We also estimated the sink using atmospheric models and the atmospheric concentration of carbon dioxide (the tracer-transport inversion method). The range of results from the atmosphere-based inversions contains the land-based estimates. Atmosphere- and land-based estimates are thus consistent, within the large ranges of uncertainty for both methods. Atmosphere-based results for 1980-89 are similar to those for 1985-89 and 1990-94, indicating a relatively stable U.S. sink throughout the period.

Biospheric primary production during an ENSO transition.

The Sea-viewing Wide Field-of-view Sensor (SeaWiFS) provides global monthly measurements of both oceanic phytoplankton chlorophyll biomass and light harvesting by land plants. These measurements allowed the comparison of simultaneous ocean and land net primary production (NPP) responses to a major El Niño to La Niña transition. Between September 1997 and August 2000, biospheric NPP varied by 6 petagrams of carbon per year (from 111 to 117 petagrams of carbon per year). Increases in ocean NPP were pronounced in tropical regions where El Niño-Southern Oscillation (ENSO) impacts on upwelling and nutrient availability were greatest. Globally, land NPP did not exhibit a clear ENSO response, although regional changes were substantial.

Nitrogen limitation of microbial decomposition in a grassland under elevated CO2.

Carbon accumulation in the terrestrial biosphere could partially offset the effects of anthropogenic CO2 emissions on atmospheric CO2. The net impact of increased CO2 on the carbon balance of terrestrial ecosystems is unclear, however, because elevated CO2 effects on carbon input to soils and plant use of water and nutrients often have contrasting effects on microbial processes. Here we show suppression of microbial decomposition in an annual grassland after continuous exposure to increased CO2 for five growing seasons. The increased CO2 enhanced plant nitrogen uptake, microbial biomass carbon, and available carbon for microbes. But it reduced available soil nitrogen, exacerbated nitrogen constraints on microbes, and reduced microbial respiration per unit biomass. These results indicate that increased CO2 can alter the interaction between plants and microbes in favour of plant utilization of nitrogen, thereby slowing microbial decomposition and increasing ecosystem carbon accumulation.

Assessing photosynthetic downregulation in sunflower stands with an optically-based model.

Using a simple light-use efficiency model based on optical measurements, we explored spatial patterns of photosynthetic activity in fertilized and unfertilized sunflower stands. The model had two components: (1) absorbed photosynthetically active radiation (APAR), and (2) radiation-use efficiency. APAR was the product of photosynthetic photon flux density (PPFD) and leaf absorptance, which was derived from leaf reflectance. Radiation-use efficiency was either assumed to be constant or allowed to vary linearly with the photochemical reflectance index (PRI), a measure of xanthophyll cycle pigment activity. When efficiency was assumed to be constant, the model overestimated photosynthetic rates in upper canopy layers exposed to direct PPFD, particularly in the unfertilized canopy due to the greater photosynthetic downregulation associated with higher levels of photoprotective (de-epoxidized) xanthophyll cycle pigments in these conditions. When efficiency was allowed to vary according to the PRI, modeled photosynthetic rates closely matched measured rates for all canopy layers in both treatments. These results illustrate the importance of considering reduced radiation-use efficiency due to photosynthetic downregulation when modeling photosynthesis from reflectance, and illustrate the potential for detecting radiation-use efficiency through leaf optical properties. At least under the conditions of this study, these results also suggest that xanthophyll cycle pigment activity and net carbon uptake are coordinately regulated, allowing assays of Photosystem II activity to reveal changing rates of net assimilation. Because the optical methods in this study are adaptable to multiple spatial scales (leaf to landscape), this approach may provide a scalable model for estimating photosynthetic rates independently from flux measurements.

Soil microbiota in two annual grasslands: responses to elevated atmospheric CO2.

We measured soil bacteria, fungi, protozoa, nematodes, and biological activity in serpentine and sandstone annual grasslands after 4 years of exposure to elevated atmospheric CO2. Measurements were made during the early part of the season, when plants were in vegetative growth, and later in the season, when plants were approaching their maximum biomass. In general, under ambient CO2, bacterial biomass, total protozoan numbers, and numbers of bactivorous nematodes were similar in the two grasslands. Active and total fungal biomasses were higher on the more productive sandstone grassland compared to the serpentine. However, serpentine soils contained nearly twice the number of fungivorous nematodes compared to the sandstone, perhaps explaining the lower standing crop of fungal biomass in the serpentine and suggesting higher rates of energy flow through the fungal-based soil food web. Furthermore, root biomass in the surface soils of these grasslands is comparable, but the serpentine contains 6 times more phytophagous nematodes compared to the sandstone, indicating greater below-ground grazing pressure on plants in stressful serpentine soils. Elevated CO2 increased the biomass of active fungi and the numbers of flagellates in both grasslands during the early part of the season and increased the number of phytophagous nematodes in the serpentine. Elevated CO2 had no effect on the total numbers of bactivorous or fungivorous nematodes, but decreased the diversity of the nematode assemblage in the serpentine at both sampling dates. Excepting this reduction in nematode diversity, the effects of elevated CO2 disappeared later in the season as plants approached their maximum biomass. Elevated CO2 had no effect on total and active bacterial biomass, total fungal biomass, or the total numbers of amoebae and ciliates in either grassland during either sampling period. However, soil metabolic activity was higher in the sandstone grassland in the early season under elevated CO2, and elevated CO2 altered the patterns of use of individual carbon substrates in both grasslands at this time. Rates of substrate use were also significantly higher in the sandstone, indicating increased bacterial metabolic activity. These changes in soil microbiota are likely due to an increase in the flux of carbon from roots to soil in elevated CO2, as has been previously reported for these grasslands. Results presented here suggest that some of the carbon distributed below ground in response to elevated CO2 affects the soil microbial food web, but that these effects may be more pronounced during the early part of the growing season.

Revisiting the commons: local lessons, global challenges.

In a seminal paper, Garrett Hardin argued in 1968 that users of a commons are caught in an inevitable process that leads to the destruction of the resources on which they depend. This article discusses new insights about such problems and the conditions most likely to favor sustainable uses of common-pool resources. Some of the most difficult challenges concern the management of large-scale resources that depend on international cooperation, such as fresh water in international basins or large marine ecosystems. Institutional diversity may be as important as biological diversity for our long-term survival.

Stimulation of grassland nitrogen cycling under carbon dioxide enrichment.

Nitrogen (N) limits plant growth in many terrestrial ecosystems, potentially constraining terrestrial ecosystem response to elevated CO2. In this study, elevated CO2 stimulated gross N mineralization and plant N uptake in two annual grasslands. In contrast to other studies that have invoked increased C input to soil as the mechanism altering soil N cycling in response to elevated CO2, increased soil moisture, due to decreased plant transpiration in elevated CO2, best explains the changes we observed. This study suggests that atmospheric CO2 concentration may influence ecosystem biogeochemistry through plant control of soil moisture.

Contrasting leaf and 'ecosystem' CO2 and H 2O exchange in Avena fatua monoculture: Growth at ambient and elevated CO2.

Elevated CO2 (ambient + 35 Pa) increased shoot dry mass production in Avena fatua by ∼ 68% at maturity. This increase in shoot biomass was paralleled by an 81% increase in average net CO2 uptake (A) per unit of leaf area and a 65% increase in average A at the 'ecosystem' level per unit of ground area. Elevated CO2 also increased 'ecosystem' A per unit of biomass. However, the products of total leaf area and light-saturated leaf A divided by the ground surface area over time appeared to lie on a single response curve for both CO2 treatments. The approximate slope of the response suggests that the integrated light saturated capacity for leaf photosynthesis is ∼ 10-fold greater than the 'ecosystem' rate. 'Ecosystem' respiration (night) per unit of ground area, which includes soil and plant respiration, ranged from-20 (at day 19) to-18 (at day 40) µmol m(-2) s(-1) for both elevated and ambient CO2 Avena. 'Ecosystem' below-ground respiration at the time of seedling emergence was ∼-10 µmol m(-2) s(-1), while that occuring after shoot removal at the termination of the experiment ranged from -5 to-6 µmol m(-2) s(-1). Hence, no significant differences between elevated and ambient CO2 treatments were found in any respiration measure on a ground area basis, though 'ecosystem' respiration on a shoot biomass basis was clearly reduced by elevated CO2. Significant differences existed between leaf and 'ecosystem' water flux. In general, leaf transpiration (E) decreased over the course of the experiment, possibly in response to leaf aging, while 'ecosystem' rates of evapotranspiration (ET) remained constant, probably because falling leaf rates were offset by an increasing total leaf biomass. Transpiration was lower in plants grown at elevated CO2, though variation was high because of variability in leaf age and ambient light conditions and differences were not significant. In contrast, 'ecosystem' evapotranspiration (ET) was significantly decreased by elevated CO2 on 5 out of 8 measurement dates. Photosynthetic water use efficiencies (A/E at the leaf level, A/ET at the 'ecosystem' level) were increased by elevated CO2. Increases were due to both increased A at leaf and 'ecosystem' level and decreased leaf E and 'ecosystem' ET.

CO2 alters water use, carbon gain, and yield for the dominant species in a natural grassland.

Global atmospheric CO2 is increasing at a rate of 1.5-2 ppm per year and is predicted to double by the end of the next century. Understanding how terrestrial ecosystems will respond in this changing environment is an important goal of current research. Here we present results from a field study of elevated CO2 in a California annual grassland. Elevated CO2 led to lower leaf-level stomatal conductance and transpiration (approximately 50%) and higher mid-day leaf water potentials (30-35%) in the most abundant species of the grassland, Avena barbata Brot. Higher CO2 concentrations also resulted in greater midday photosynthetic rates (70% on average). The effects of CO2 on stomatal conductance and leaf water potential decreased towards the end of the growing season, when Avena began to show signs of senescence. Water-use efficiency was approximately doubled in elevated CO2, as estimated by instantaneous gas-exchange measurements and seasonal carbon isotope discrimination. Increases in CO2 and photosynthesis resulted in more seeds per plant (30%) and taller and heavier plants (27% and 41%, respectively). Elevated CO2 also reduced seed N concentrations (9%).

Environmental effects on circadian rhythms in photosynthesis and stomatal opening.

Persistent circadian rhythms in photosynthesis and stomatal opening occurred in bean (Phaseolus vulgaris L.) plants transferred from a natural photoperiod to a variety of constant conditions. Photosynthesis, measured as carbon assimilation, and stomatal opening, as conductance to water vapor, oscillated with a freerunning period close to 24 h under constant moderate light, as well as under light-limiting and CO2-limiting conditions. The rhythms damped under constant conditions conducive to high photosynthetic rates, as did rates of carbon assimilation and stomatal conductance, and this damping correlated with the accumulation of carbohydrate. No rhythm in respiration occurred in plants transferred to constant darkness, and the rhythm in stomatal opening damped rapidly in constant darkness. Damping of rhythms also occurred in leaflets exposed to constant light and CO2-free air, demonstrating that active photosynthesis and not simply light was necessary for sustained expression of these rhythms.

Evidence of multiple circadian oscillators in bean plants.

Circadian rhythms in stomatal opening and photosynthesis had shorter free-running periods than circadian rhythms in leaflet movement in bean plants (Phaseolus vulgaris L.) transferred from 12-hr photoperiods to constant conditions. The rhythm in leaflet movement had a period close to 27 hr, whereas the rhythm in stomatal opening, measured as conductance to water vapor, had a period close to 24 hr. Photosynthesis, measured as net assimilation of CO2, also oscillated with a period close to 24 hr. The periods of these rhythms did not vary with increasing temperature, demonstrating temperature compensation of the controlling oscillators. The difference in free-running periods displayed by these rhythms is evidence that multiple oscillators with different intrinsic frequencies operate in bean plants.

Biochemical Correlates of the Circadian Rhythm in Photosynthesis in Phaseolus vulgaris.

A circadian rhythm in photosynthesis occurs in Phaseolus vulgaris after transfer from a natural or artificial light:dark cycle to constant light. The rhythm in photosynthesis persists even when intercellular CO(2) partial pressure is held constant, demonstrating that the rhythm in photosynthesis is not entirely due to stomatal control over the diffusion of CO(2). Experiments were conducted to attempt to elucidate biochemical correlates with the circadian rhythm in photosynthesis. Plants were entrained to a 12-hour-day:12-hour-night light regimen and then monitored or sampled during a subsequent period of constant light. We observed circadian oscillations in ribulose-1,5-bisphosphate (RuBP) levels, and to a lesser extent in phosphoglyceric acid (PGA) levels, that closely paralleled oscillations in photosynthesis. However, the enzyme activity and activation state of the enzyme responsible for the conversion of RuBP to PGA, ribulose-1,5-bisphosphate carboxylase/oxygenase, showed no discernible circadian oscillation. Hence, we examined the possibility of circadian effects on RuBP regeneration. Neither ribulose-5-phosphate kinase activity nor the level of ATP fluctuated in constant light. Oscillations in triose-phosphate levels were out of phase with those observed for RuBP and PGA.

Circadian Rhythms in Photosynthesis : Oscillations in Carbon Assimilation and Stomatal Conductance under Constant Conditions.

Net carbon assimilation and stomatal conductance to water vapor oscillated repeatedly in red kidney bean, Phaseolus vulgaris L., plants transferred from a natural photoperiod to constant light. In a gas exchange system with automatic regulation of selected environmental and physiological variables, assimilation and conductance oscillated with a free-running period of approximately 24.5 hours. The rhythms in carbon assimilation and stomatal conductance were closely coupled and persisted for more than a week under constant conditions. A rhythm in assimilation occurred when either ambient or intercellular CO(2) partial pressure was held constant, demonstrating that the rhythm in assimilation was not entirely the result of stomatal effects on CO(2) diffusion. Rhythms in assimilation and conductance were not expressed in plants grown under constant light at a constant temperature, demonstrating that the rhythms did not occur spontaneously but were induced by an external stimulus. In plants grown under constant light with a temperature cycle, a rhythm was entrained in stomatal conductance but not in carbon assimilation, indicating that the oscillators driving the rhythms differed in their sensitivity to environmental stimuli.