Impacts of warming and elevated CO2 on a semi‐arid grassland are non‐additive,shift with precipitation,and reverse over time

It is unclear how elevated CO₂ (eCO₂) and the corresponding shifts in temperature and precipitation will interact to impact ecosystems over time. During a 7‐year experiment in a semi‐arid grassland, the response of plant biomass to eCO₂ and warming was largely regulated by interannual precipitation, while the response of plant community composition was more sensitive to experiment duration. The combined effects of eCO₂ and warming on aboveground plant biomass were less positive in ‘wet’ growing seasons, but total plant biomass was consistently stimulated by ~ 25% due to unique, supra‐additive responses of roots. Independent of precipitation, the combined effects of eCO₂ and warming on C₃ graminoids became increasingly positive and supra‐additive over time, reversing an initial shift toward C₄ grasses. Soil resources also responded dynamically and non‐additively to eCO₂ and warming, shaping the plant responses. Our results suggest grasslands are poised for drastic changes in function and highlight the need for long‐term, factorial experiments.     

Biomass responses in a temperate European grassland through 17 years of elevated CO2

Future increase in atmospheric CO₂ concentrations will potentially enhance grassland biomass production and shift the functional group composition with consequences for ecosystem functioning. In the “GiFACE” experiment (Giessen Free Air Carbon dioxide Enrichment), fertilized grassland plots were fumigated with elevated CO₂ (eCO₂) year‐round during daylight hours since 1998, at a level of +20% relative to ambient concentrations (in 1998, aCO₂ was 364 ppm and eCO₂ 399 ppm; in 2014, aCO₂ was 397 ppm and eCO₂ 518 ppm). Harvests were conducted twice annually through 23 years including 17 years with eCO₂ (1998 to 2014). Biomass consisted of C3 grasses and forbs, with a small proportion of legumes. The total aboveground biomass (TAB) was significantly increased under eCO₂ (p = .045 and .025, at first and second harvest). The dominant plant functional group grasses responded positively at the start, but for forbs, the effect of eCO₂ started out as a negative response. The increase in TAB in response to eCO₂ was approximately 15% during the period from 2006 to 2014, suggesting that there was no attenuation of eCO₂ effects over time, tentatively a consequence of the fertilization management. Biomass and soil moisture responses were closely linked. The soil moisture surplus (c. 3%) in eCO₂ manifested in the latter years was associated with a positive biomass response of both functional groups. The direction of the biomass response of the functional group forbs changed over the experimental duration, intensified by extreme weather conditions, pointing to the need of long‐term field studies for obtaining reliable responses of perennial ecosystems to eCO₂ and as a basis for model development.     

Management of a semi‐natural grassland; results from a 15‐year‐old experiment in southern Sweden

Abstract. Vegetation changes in a semi‐natural grassland of wooded meadow type that had been grazed for centuries are described following the introduction of various management regimes: mowing each year, mowing every third year, burning, mechanical removal of woody plants, chemical treatment of woody plants, continuous grazing and abandonment. The experiment was established in southern Sweden in 1972 and has been in progress for 15 years. In 1972, 1980 and 1986 the botanical composition in these plots was investigated in permanent subplots. The study clearly demonstrates that mowing or grazing is necessary to preserve community structure and that mowing is to be preferred in cases where maintaining species richness is of primary concern. Mowing every third year delayed vegetation change and prevented woody species from spreading. Therefore, periodic mowing might be an alternative way to preserve the flora. In contrast, yearly burning does not seem to be a viable management in this type of semi‐natural grassland. To preserve the open landscape regeneration of woody plants has to be prevented. However, in plots where woody plants were removed the typical grassland flora declined. Abandonment resulted in closed forest.     

Interactive direct and plant‐mediated effects of elevated atmospheric [CO2] and temperature on a eucalypt‐feeding insect herbivore

Understanding the direct and indirect effects of elevated [CO₂] and temperature on insect herbivores and how these factors interact are essential to predict ecosystem‐level responses to climate change scenarios. In three concurrent glasshouse experiments, we measured both the individual and interactive effects of elevated [CO₂] and temperature on foliar quality. We also assessed the interactions between their direct and plant‐mediated effects on the development of an insect herbivore of eucalypts. Eucalyptus tereticornis saplings were grown at ambient or elevated [CO₂] (400 and 650 μmol mol^?1 respectively) and ambient or elevated ( + 4 °C) temperature for 10 months. Doratifera quadriguttata (Lepidoptera: Limacodidae) larvae were feeding directly on these trees, on their excised leaves in a separate glasshouse, or on excised field‐grown leaves within the temperature and [CO₂] controlled glasshouse. To allow insect gender to be determined and to ensure that any sex‐specific developmental differences could be distinguished from treatment effects, insect development time and consumption were measured from egg hatch to pupation. No direct [CO₂] effects on insects were observed. Elevated temperature accelerated larval development, but did not affect leaf consumption. Elevated [CO₂] and temperature independently reduced foliar quality, slowing larval development and increasing consumption. Simultaneously increasing both [CO₂] and temperature reduced these shifts in foliar quality, and negative effects on larval performance were subsequently ameliorated. Negative nutritional effects of elevated [CO₂] and temperature were also independently outweighed by the direct positive effect of elevated temperature on larvae. Rising [CO₂] and temperature are thus predicted to have interactive effects on foliar quality that affect eucalypt‐feeding insects. However, the ecological consequences of these interactions will depend on the magnitude of concurrent temperature rise and its direct effects on insect physiology and feeding behaviour.     

Diurnal and seasonal variations in stomatal conductance of rice at elevated atmospheric CO2 under fully open‐air conditions

Understanding of leaf stomatal responses to the atmospheric CO₂ concentration, [CO₂], is essential for accurate prediction of plant water use under future climates. However, limited information is available for the diurnal and seasonal changes in stomatal conductance (g_s) under elevated [CO₂]. We examined the factors responsible for variations in g_s under elevated [CO₂] with three rice cultivars grown in an open‐field environment under flooded conditions during two growing seasons (a total of 2140 individual measurements). Conductance of all cultivars was generally higher in the morning and around noon than in the afternoon, and elevated [CO₂] decreased g_s by up to 64% over the 2 years (significantly on 26 out of 38 measurement days), with a mean g_s decrease of 23%. We plotted the g_s variations against three parameters from the Ball‐Berry model and two revised versions of the model, and all parameters explained the g_s variations well at each [CO₂] in the morning and around noon (R² > 0.68), but could not explain these variations in the afternoon (R² < 0.33). The present results provide an important basis for modelling future water use in rice production.     

Modeled responses of terrestrial ecosystems to elevated atmospheric CO2: a comparison of simulations by the biogeochemistry models of the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP)

Although there is a great deal of information concerning responses to increases in atmospheric CO₂ at the tissue and plant levels, there are substantially fewer studies that have investigated ecosystem-level responses in the context of integrated carbon, water, and nutrient cycles. Because our understanding of ecosystem responses to elevated CO₂ is incomplete, modeling is a tool that can be used to investigate the role of plant and soil interactions in the response of terrestrial ecosystems to elevated CO₂. In this study, we analyze the responses of net primary production (NPP) to doubled CO₂ from 355 to 710 ppmv among three biogeochemistry models in the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP): BIOME-BGC (BioGeochemical Cycles), Century, and the Terrestrial Ecosystem Model (TEM). For the conterminous United States, doubled atmospheric CO₂ causes NPP to increase by 5% in Century, 8% in TEM, and 11% in BIOME-BGC. Multiple regression analyses between the NPP response to doubled CO₂ and the mean annual temperature and annual precipitation of biomes or grid cells indicate that there are negative relationships between precipitation and the response of NPP to doubled CO₂ for all three models. In contrast, there are different relationships between temperature and the response of NPP to doubled CO₂ for the three models: there is a negative relationship in the responses of BIOME-BGC, no relationship in the responses of Century, and a positive relationship in the responses of TEM. In BIOME-BGC, the NPP response to doubled CO₂ is controlled by the change in transpiration associated with reduced leaf conductance to water vapor. This change affects soil water, then leaf area development and, finally, NPP. In Century, the response of NPP to doubled CO₂ is controlled by changes in decomposition rates associated with increased soil moisture that results from reduced evapotranspiration. This change affects nitrogen availability for plants, which influences NPP. In TEM, the NPP response to doubled CO₂ is controlled by increased carboxylation which is modified by canopy conductance and the degree to which nitrogen constraints cause down-regulation of photosynthesis. The implementation of these different mechanisms has consequences for the spatial pattern of NPP responses, and represents, in part, conceptual uncertainty about controls over NPP responses. Progress in reducing these uncertainties requires research focused at the ecosystem level to understand how interactions between the carbon, nitrogen, and water cycles influence the response of NPP to elevated atmospheric CO₂. Received: 13 December 1996 / Accepted: 20 November 1997     

Partitioning of ecosystem respiration of CO2 released during land‐use transition from temperate agricultural grassland to Miscanthus × giganteus

Conversion of large areas of agricultural grassland is inevitable if European and UK domestic production of biomass is to play a significant role in meeting demand. Understanding the impact of these land‐use changes on soil carbon cycling and stocks depends on accurate predictions from well‐parameterized models. Key considerations are cultivation disturbance and the effect of autotrophic root input stimulation on soil carbon decomposition under novel biomass crops. This study presents partitioned parameters from the conversion of semi‐improved grassland to Miscanthus bioenergy production and compares the contribution of autotrophic and heterotrophic respiration to overall ecosystem respiration of CO₂ in the first and second years of establishment. Repeated measures of respiration from within and without root exclusion collars were used to produce time‐series model integrations separating live root inputs from decomposition of grass residues ploughed in with cultivation of the new crop. These parameters were then compared to total ecosystem respiration derived from eddy covariance sensors. Average soil surface respiration was 13.4% higher in the second growing season, increasing from 2.9 to 3.29 g CO₂‐C m^?2 day^?1. Total ecosystem respiration followed a similar trend, increasing from 4.07 to 5.4 g CO₂‐C m^?2 day^?1. Heterotrophic respiration from the root exclusion collars was 32.2% lower in the second growing season at 1.20 g CO₂‐C m^?2 day^?1 compared to the previous year at 1.77 g CO₂‐C m^?2 day^?1. Of the total respiration flux over the two‐year time period, aboveground autotrophic respiration plus litter decomposition contributed 38.46% to total ecosystem respiration while belowground autotrophic respiration and stimulation by live root inputs contributed 46.44% to soil surface respiration. This figure is notably higher than mean figures for nonforest soils derived from the literature and demonstrates the importance of crop‐specific parameterization of respiration models.     

N2O fluxes of a bio‐energy poplar plantation during a two years rotation period

Nitrous oxide emissions are of critical importance for the assumed climate neutrality of bio‐energy. In this study we report on the N₂O fluxes from a bio‐energy poplar plantation measured with eddy covariance for 2 years, after conversion of agricultural fields to few months after harvesting of the plantation. A pulse peak of N₂O was detected after the land use change and in the wake of the first heavy rainfall. The N₂O‐N emission during just a single week was 2.7 kg N₂O‐N ha^?1 which represented approximately 42% of the total N₂O‐N emitted during the 2 years of measurements. After this peak emission, N₂O fluxes were constantly rather low, not increasing after rainfall events any longer. Lowest emissions (and even N₂O sink) occurred mostly during the end of the second growing season with maximum canopy development, and water table deeper than 80 cm. Gross primary production (GPP) explained 68% of the monthly averaged variability in N₂O emission from August to December 2011. Probably N uptake by vegetation during the peak of the second growing season limited N₂O emission, which in fact increased again after the plantation was coppiced. For the majority of the measuring period, N₂O fluxes did not present a well‐defined diurnal pattern, with the exception of two periods: (1) from 19–22 August 2010 and (2) from September–November 2011. In both cases wind speed played a major role in controlling the diurnal pattern in these fluxes (explaining up to 80% of the diurnal variability in N₂O fluxes on 19–22 August 2010), whereas at the end of the second growing season (September–November 2011), GPP explained 73% of the diurnal pattern in N₂O fluxes.     

Towards an understanding of how phloem amino acid composition shapes elevated CO2‐induced changes in aphid population dynamics

1. The performance of foliage feeders tends to decrease under elevated CO₂, but the responses of phloem‐feeding insects have been much more equivocal. As phloem tissues are less accessible than whole‐plant tissues, much less is known about how phloem composition is altered under elevated CO₂ and the mechanisms driving changes in aphid performance. 2. In this study, the plant mechanisms underlying the performance of Rhopalosiphum padi aphids on Hordeum vulgare (barley) grown under ambient (390 ppm) and elevated (700 ppm) CO₂ were examined. We used aphid stylectomy to sample pure phloem from plants in CO₂‐controlled conditions and high‐performance liquid chromatography to analyse phloem samples for amino acid concentrations. 3. Aphid abundance significantly increased by 127% under elevated CO₂. Consequently, plant biomass decreased under elevated CO₂ in trials with herbivores present, possibly due to the increased herbivore load, but increased when aphids were absent. The intrinsic rate of population increase (r_m) was significantly higher under elevated CO₂; however, there were no statistically significant effects on aphid fecundity or development time. The concentration of individual amino acids tended to increase, although these increases were statistically significant in only a few cases. A principal components analysis revealed that the relative abundance (mol %) of those amino acids considered essential for aphids tended to increase under elevated CO₂. 4. These results indicate that CO₂ may affect nutrient translocation in plants in ways that are contrary to predictions about nitrogen metabolite responses to CO₂. Such plant biochemical responses may underlie observations of improved phloem feeder performance under elevated CO₂.     

Nutrient supply enhanced the increase in intrinsic water‐use efficiency of a temperate seminatural grassland in the last century

Under the increase in atmospheric CO₂ during the last century, variable increases in the intrinsic water‐use efficiency (W_i), i.e., the ratio between carbon assimilation rate (A) and stomatal conductance (g_s), of C₃ vegetation have been observed. Here, we ask if long‐term nutrient status and especially nitrogen supply have an effect on the CO₂ response of W_i in a temperate seminatural C₃ grassland. This analysis draws on the long‐term trends (1915–2009) in W_i, derived from carbon isotope analysis, of archived hay and herbage from the Park Grass Experiment at Rothamsted (South‐East England). Plant samples came from five fertilizer treatments, each with different annual nitrogen (N; 0, 48 or 96 kg ha^?1), phosphorus (P; 0 or 35 kg ha^?1) and potassium (K; 0 or 225 kg ha^?1) applications, with lime as required to maintain soil pH near 7. Carbon isotope discrimination (¹³Δ) increased significantly (P < 0.001) on the Control (0.9‰ per 100 ppm CO₂ increase). This trend differed significantly (P < 0.01) from those observed on the fertilized treatments (PK only: 0.4‰ per 100 ppm CO₂ increase, P < 0.001; Low N only, Low N+PK, High N+PK: no significant increase). The ¹³Δ trends on fertilized treatments did not differ significantly from each other. However, N status, assessed as N fertilizer supply plus an estimate of biologically fixed N, was negatively related (r² = 0.88; P < 0.02) to the trend for ¹³Δ against CO₂. Other indices of N status exhibited similar relationships. Accordingly, the increase in W_i at High N+PK was twice that of the Control (+28% resp. +13% relative to 1915). In addition, the CO₂ responsiveness of ¹³Δ was related to the grass content of the plant community. This may have been due to the greater CO₂ responsiveness of g_s in grasses relative to forbs. Thus, the greater CO₂ response of grass‐rich fertilized swards may be related to effects of nutrient supply on botanical composition.     

Insights into the helix‐sense inversion of poly(β‐phenethyl‐L‐aspartate) by two‐dimensional Raman correlation spectroscopy

In this study, the transition process of the helix‐sense inversion of poly(β‐phenethyl‐L‐aspartate) was investigated by Raman scattering and 2‐dimensional correlation spectroscopy. Temperature‐dependent Raman spectra were obtained during the helix‐sense inversion. The results of 2‐dimensional correlation analysis in the spectral regions of 1600‐1800 and 3200‐3400 cm^?1 showed that the intensity changes of the side‐chain ester C═O stretching bands occurred prior to those of amide A and amide I bands in the unwinding process of αR‐helix on heating. The sequential order of the intensity changes for amide A, amide I, and the side‐chain ester C═O stretching bands during the inversion process was determined. It was found that the conformation change of the side chain occurred prior to that of the main chain for the αR‐helix on heating. Thus, we concluded that the transformation of the backbone chain from right‐handed to left‐handed is triggered by the conformational change of the side chains.     

Long‐term dynamics of mycorrhizal root tips in a loblolly pine forest grown with free‐air CO2 enrichment and soil N fertilization for 6 years

Large‐scale, long‐term FACE (Free‐Air CO₂ enrichment) experiments indicate that increases in atmospheric CO₂ concentrations will influence forest C cycling in unpredictable ways. It has been recently suggested that responses of mycorrhizal fungi could determine whether forest net primary productivity (NPP) is increased by elevated CO₂ over long time periods and if forests soils will function as sources or sinks of C in the future. We studied the dynamic responses of ectomycorrhizae to N fertilization and atmospheric CO₂ enrichment at the Duke FACE experiment using minirhizotrons over a 6 year period (2005–2010). Stimulation of mycorrhizal production by elevated CO₂ was observed during only 1 (2007) of 6 years. This increased the standing crop of mycorrhizal tips during 2007 and 2008; during 2008, significantly higher mortality returned standing crop to ambient levels for the remainder of the experiment. It is therefore unlikely that increased production of mycorrhizal tips can explain the lack of progressive nitrogen limitations and associated increases in N uptake observed in CO₂‐enriched plots at this site. Fertilization generally decreased tree reliance on mycorrhizae as tip production declined with the addition of nitrogen as has been shown in many other studies. Annual NPP of mycorrhizal tips was greatest during years with warm January temperatures and during years with cool spring temperatures. A 2 °C increase in average late spring temperatures (May and June) decreased annual production of mycorrhizal root tip length by 50%. This has important implications for ecosystem function in a warmer world in addition to potential for forest soils to sequester atmospheric C.