The effect of elevated atmospheric CO2 and drought on sources and sinks of isoprene in a temperate and tropical rainforest mesocosm

Isoprene is the most abundant volatile hydrocarbon emitted by many tree species and has a major impact on tropospheric chemistry, leading to formation of pollutants and enhancing the lifetime of methane, a powerful greenhouse gas. Reliable estimates of global isoprene emission from different ecosystems demand a clear understanding of the processes of both production and consumption. Although the biochemistry of isoprene production has been studied extensively and environmental controls over its emission are relatively well known, the study of isoprene consumption in soil has been largely neglected. Here, we present results on the production and consumption of isoprene studied by measuring the following different components: (1) leaf and soil and (2) at the whole ecosystem level in two distinct enclosed ultraviolet light‐depleted mesocosms at the Biosphere 2 facility: a cottonwood plantation with trees grown at ambient and elevated atmospheric CO₂ concentrations and a tropical rainforest, under well watered and drought conditions. Consumption of isoprene by soil was observed in both systems. The isoprene sink capacity of litter‐free soil of the agriforest stands showed no significant response to different CO₂ treatments, while isoprene production was strongly depressed by elevated atmospheric CO₂ concentrations. In both mesocosms, drought suppressed the sink capacity, but the full sink capacity of dry soil was recovered within a few hours upon rewetting. We conclude that soil uptake of atmospheric isoprene is likely to be modest but significant and needs to be taken into account for a comprehensive estimate of the global isoprene budget. More studies investigating the capacity of soils to uptake isoprene in natural conditions are clearly needed.     

Studies of the relationship between isoprene emission rate and CO2 or photon-flux density using a real-time isoprene analyser

Abstract. Studies of the isoprene emission rate in response to changes in photon-flux density and CO₂ partial pressure were conducted using a recently developed on-line isoprene analyser combined with a gas exchange system and chlorophyll fluorometer. Upon darkening, the isoprene emission rate from leaves of aspen ( Populus tremuloides Michaux.) began to decline immediately, demonstrating that the internal pool of isoprene, or its precursors, is small and that the instantaneous emission rate is tightly coupled to the rate of synthesis. A post-illumination burst of isoprene was observed within 5 min after darkening and lasted for 15–20 min in four isoprene-emitting species that were examined. In leaves of eucalyptus ( Eucalyptus globulus Labill.), the magnitude of the post-illumination burst was dependent on the photon-flux density that existed before darkening, but not on ambient CO₂ partial pressure. The dependence of the post-illumination burst on photon-flux density paralleled that for the steady-state rate of isoprene emission. A step-wise increase in intercellular CO₂ partial pressure from 24.5 to 60 Pa resulted in an immediate decrease in isoprene emission rate and non-photochemical fluorescence quenching, but an increase in CO₂ assimilation rate. Given the several recent studies that link isoprene emission to chloroplastic processes, the results of this study indicate that the linkage is not dependent on the rate of CO₂ flux through the reductive pentose phosphate pathway, but rather on more complex relationships involving metabolites not appreciably influenced by CO₂ partial pressure.     

Rapid appearance of 13C in biogenic isoprene when 13CO2 is fed to intact leaves

Biogenic isoprene substantially affects atmospheric chemistry, but it is not known how or why many plants, especially trees, make isoprene. We fed ¹³CO₂ to leaves of Quercus rubra and monitored the incorporation of ¹³C into isoprene by mass spectrometry. After feeding ¹³CO₂ for 9 min we found all possible labelling patterns from completely unlabelled to fully labelled isoprene. By 18 min, 84% of the carbon atoms in isoprene were ¹³C. Labelling of the last 20% of the carbon atoms was much slower than labelling of the first 80%. The rate of labelling of isoprene was similar to that reported for phosphoglyceric acid indicating that there is a close linkage between the carbon source for isoprene synthesis and the photosynthetic carbon reduction pathway.     

Impact of rising CO2 on emissions of volatile organic compounds: isoprene emission from Phragmites australis growing at elevated CO2 in a natural carbon dioxide spring†

Isoprene basal emission (the emission of isoprene from leaves exposed to a light intensity of 1000 µmol m^?2 s^?1 and maintained at a temperature of 30 °C) was measured in Phragmites australis plants growing under elevated CO₂ in the Bossoleto CO₂ spring at Rapolano Terme, Italy, and under ambient CO₂ at a nearby control site. Gas exchange and biochemical measurements were concurrently taken. Isoprene emission was lower in the plants growing at elevated CO₂ than in those growing at ambient CO₂. Isoprene emission and isoprene synthase activity (IsoS) were very low in plants growing at the bottom of the spring under very rich CO₂ and increased at increasing distance from the spring (and decreasing CO₂ concentration). Distance from the spring did not significantly affect photosynthesis making it therefore unlikely that there is carbon limitation to isoprene formation. The isoprene emission rate was very quickly reduced after rapid switches from elevated to ambient CO₂ in the gas‐exchange cuvette, whereas it increased when switching from ambient to elevated CO₂. The rapidity of the response may be consistent with post‐translational modifications of enzymes in the biosynthetic pathway of isoprene formation. Reduction of IsoS activity is interpreted as a long‐term response. Basal emission of isoprene was not constant over the day but showed a diurnal course opposite to photosynthesis, with a peak during the hottest hours of the day, independent of stomatal conductance and probably dependent on external air temperature or temporary reduction of CO₂ concentration. The present experiments show that basal emission rate of isoprene is likely to be reduced under future elevated CO₂ levels and allow improvement in the modelling of future isoprene emission rates.     

Modelling the effect of the 2018 summer heatwave and drought on isoprene emissions in a UK woodland

Projected future climatic extremes such as heatwaves and droughts are expected to have major impacts on emissions and concentrations of biogenic volatile organic compounds (bVOCs) with potential implications for air quality, climate and human health. While the effects of changing temperature and photosynthetically active radiation (PAR) on the synthesis and emission of isoprene, the most abundant of these bVOCs, are well known, the role of other environmental factors such as soil moisture stress are not fully understood and are therefore poorly represented in land surface models. As part of the Wytham Isoprene iDirac Oak Tree Measurements campaign, continuous measurements of isoprene mixing ratio were made throughout the summer of 2018 in Wytham Woods, a mixed deciduous woodland in southern England. During this time, the United Kingdom experienced a prolonged heatwave and drought, and isoprene mixing ratios were observed to increase by more than 400% at Wytham Woods under these conditions. We applied the state‐of‐the‐art FORest Canopy‐Atmosphere Transfer canopy exchange model to investigate the processes leading to these elevated concentrations. We found that although current isoprene emissions algorithms reproduced observed mixing ratios in the canopy before and after the heatwave, the model underestimated observations by ~40% during the heatwave–drought period implying that models may substantially underestimate the release of isoprene to the atmosphere in future cases of mild or moderate drought. Stress‐induced emissions of isoprene based on leaf temperature and soil water content (SWC) were incorporated into current emissions algorithms leading to significant improvements in model output. A combination of SWC, leaf temperature and rewetting emission bursts provided the best model‐measurement fit with a 50% improvement compared to the baseline model. Our results highlight the need for more long‐term ecosystem‐scale observations to enable improved model representation of atmosphere–biosphere interactions in a changing global climate.     

Effect of Elevated CO2 and Drought on Soil Microbial Communities Associated with Andropogon gerardii

Our understanding of the effects of elevated atmospheric CO2, singly and In combination with other environmental changes,on plant-soil interactions is incomplete. Elevated CO2 effects on C4 plants, though smaller than on C3 species, are mediated mostly via decreased stomatal conductance and thus water loss. Therefore, we characterized the interactive effect of elevated CO2 and drought on soil microbial communities associated with a dominant C4 prairie grass, Andropogon gerardii Vitman. Elevated CO2 and drought both affected resources available to the soil microbial community. For example, elevated CO2 increased the soil C:N ratio and water content during drought, whereas drought alone decreased both. Drought significantly decreased soil microbial biomass. In contrast, elevated COz increased biomass while ameliorating biomass decreases that were induced under drought. Total and active direct bacterial counts and carbon substrate use (overall use and number of used sources) increased significantly under elevated CO2. Denaturing gradient gel electrophoresis analysis revealed that drought and elevated CO2, singly and combined, did not affect the soil bacteria community structure.We conclude that elevated CO2 alone increased bacterial abundance and microbial activity and carbon use, probably in response to increased root exudation. Elevated CO2 also limited drought-related impacts on microbial activity and biomass,which likely resulted from decreased plant water use under elevated CO2. These are among the first results showing that elevated CO2 and drought work in opposition to modulate plant-associated soil-bacteria responses,which should then Influence soil resources and plant and ecosystem function.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Stress-induced changes in carbon sources for isoprene production in Populus deltoides

Isoprene is emitted from leaves of numerous plant species and has important implications for plant metabolism and atmospheric chemistry. The ability to use stored carbon (alternative carbon sources), as opposed to recently assimilated photosynthate, for isoprene production may be important as plants routinely experience photosynthetic depression in response to environmental stress. A CO₂‐labelling study was performed and stable isotopes of carbon were used to examine the role of alternative carbon sources in isoprene production in Populus deltoides during conditions of water stress and high leaf temperature. Isotopic fractionation during isoprene production was higher in heat‐ and water‐stressed leaves (?8.5 and ?9.3‰, respectively) than in unstressed controls (?2.5 to ?3.2‰). In unstressed plants, 84–88% of the carbon in isoprene was derived from recently assimilated photosynthate. A significant shift in the isoprene carbon composition from photosynthate to alternative carbon sources was observed only under severe photosynthetic limitation (stomatal conductance < 0.05 mol m^?2 s^?1). The contribution of photosynthate to isoprene production decreased to 77 and 61% in heat‐ and water‐stressed leaves, respectively. Across water‐ and heat‐stress experiments, allocation of photosynthate was negatively correlated to the ratio of isoprene emission to photosynthesis. In water‐stressed plants, the use of alternative carbon was also related to stomatal conductance. It has been proposed that isoprene emission may be regulated by substrate availability. Thus, understanding carbon partitioning to isoprene production from multiple sources is essential for building predictive models of isoprene emission.     

海南尖峰岭热带山地雨林近冠层CO2及通量特征研究

采用梯度法及CI-301PS CO2系统,实现了海南尖峰岭热带山地雨林近冠层CO 2、微气象因子梯度观测研究,结果表明雨季晴、晴间少云及多云天气,900～1800CO 2浓度平均值分别在349～350ml/m3(冠上2.8m)、346～348ml/m3(冠上0.8m)、345～349ml/m3(林内16m)、352～357ml/m3(林内5m),较旱季相应天气分别小14～17.5、10.1～23.7、1 6.4～35.7和18.1～36.1ml/m3;100～800,则雨季CO2浓度大于旱季;梯度浓度廓变量和实时动力计算反映出,800～1800CO2通量由大气向林冠层,旱、雨季平均CO2通量分别为(0.61×10-6kg/(m2·s)和0.71×10-6kg/(m 2·s),1900通量甚微,2000～700CO2通量则由林冠向大气,平均CO2通量分别为0.36×10-6kg/(m2·s)和0.32×10-6kg/(m2·s);雨季昼夜大气流向冠层的净CO2通量是相应旱季的1.56倍.总辐射、冠顶净辐射通量以1300～1 500为最大时域,相对林内21m,80%的辐射热能被冠层吸收,与CO2通量正相关;晴天冠上潜热、感热最高值分别在1300～1400和900,反映热带山地雨林近冠层的汇、源即白昼光合固定CO2大于夜间呼吸排放CO2效应,且雨季高于旱季.     

Controls on microbial CO2 production: a comparison of surface and subsurface soil horizons

Although a significant amount of the organic C stored in soil resides in subsurface horizons, the dynamics of subsurface C stores are not well understood. The objective of this study was to determine if changes in soil moisture, temperature, and nutrient levels have similar effects on the mineralization of surface (0–25 cm) and subsurface (below 25 cm) C stores. Samples were collected from a 2 m deep unsaturated mollisol profile located near Santa Barbara, CA, USA. In a series of experiments, we measured the influence of nutrient additions (N and P), soil temperature (10–35°C), and soil water potential (?0.5 to ?10 MPa) on the microbial mineralization of native soil organic C. Surface and subsurface soils were slightly different with respect to the effects of water potential on microbial CO₂ production; C mineralization rates in surface soils were more affected by conditions of moderate drought than rates in subsurface soils. With respect to the effects of soil temperature and nutrient levels on C mineralization rates, subsurface horizons were significantly more sensitive to increases in temperature or nutrient availability than surface horizons. The mean Q₁₀ value for C mineralization rates was 3.0 in surface horizons and 3.9 in subsurface horizons. The addition of either N or P had negligible effects on microbial CO₂ production in surface soil layers; in the subsurface horizons, the addition of either N or P increased CO₂ production by up to 450% relative to the control. The results of these experiments suggest that alterations of the soil environment may have different effects on CO₂ production through the profile and that the mineralization of subsurface C stores may be particularly susceptible to increases in temperature or nutrient inputs to soil.     

尖峰岭热带山地雨林生态系统的水文生态效应

根据尖峰岭热带山地雨林３个对比集水区的研究,天然更新４０ａ山地雨林生态系统在５￣９月份,蓄养降水资源的５１．８％;１０￣翌年２月份,补枯径流水量１４３．５ｍｍ;特大暴雨的径流响应从３２．７％￣６２．５％、随系统蓄满程度变化。集水区降雨、径流水体ＣＯＤ、氨氮、酚、Ｚｎ、Ｃｕ、Ｐｂ、Ｃｄ、Ｎｉ的浓度含量远低于地面水环境质量Ⅰ类标准;山地雨林系统对降雨中氨氮、酚、Ｐｂ、Ｃｄ、Ｚｎ浓度贮滤强度分别达２３．     

全球大气CO2浓度升高对土壤微生物的影响

全球大气CO2浓度升高对土壤微生物生态系统的影响已引起广泛关注。本文从土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性方面对大气高浓度CO2的响应进行了综述。由于提供高浓度CO2的实验系统、所选植物材料以及土壤特性等的不同,大气CO2浓度升高对土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性的影响并未得出一致结论。但高浓度CO2对土壤微生物生态系统的影响是存在的。     

凋落物添加条件下咸水灌溉对风沙土CO2排放及化学性质的影响

沙拐枣作为塔克拉玛干沙漠公路防护林主要优势物种之一,其凋落物在该区域的碳循环过程中具有重要作用。采用室内培养试验测定了沙拐枣凋落物添加后,在25%、50%、75%、100%田间持水量的咸水(SW)和淡水(FW)灌溉条件下,风沙土CO₂排放、土壤有机碳(SOC)、可溶性有机碳(DOC)、pH和电导率(EC)的动态变化。结果表明: 咸水灌溉对土壤CO₂排放具有抑制作用,在4种土壤含水量处理下,淡水灌溉比咸水灌溉的CO₂累积排放量增加1.9%～29.1%;土壤CO₂累积排放量随土壤含水量的增加而升高;在沙拐枣凋落物添加条件下,SOC在前期迅速下降,但后期所有处理SOC均逐渐升高后趋于稳定;培养结束时,各处理DOC含量比培养前提高了41.3%～92.4%;在培养结束时各处理与培养前相比pH升高了0.20～0.35;EC随土壤含水量的增加而升高,培养结束时在4种含水量条件下,咸水灌溉下EC比培养前增加了0.11～0.79 mS·cm^-1,而淡水灌溉下EC比培养前有增有减。土壤CO₂累积排放量与SOC、DOC、pH呈显著正相关,但与土壤含水量相关性不显著。凋落物添加条件下咸水灌溉以及较低的含水量条件均能抑制风沙土CO₂的排放,EC受灌溉水质和土壤含水量的影响较大。     

Enhanced isoprene emission capacity and altered light responsiveness in aspen grown under elevated atmospheric CO2 concentration

Controversial evidence of CO₂‐responsiveness of isoprene emission has been reported in the literature with the response ranging from inhibition to enhancement, but the reasons for such differences are not understood. We studied isoprene emission characteristics of hybrid aspen (Populus tremula x P. tremuloides) grown under ambient (380 μmol mol^?1) and elevated (780 μmol mol^?1) [CO₂] to test the hypothesis that growth [CO₂] effects on isoprene emission are driven by modifications in substrate pool size, reflecting altered light use efficiency for isoprene synthesis. A novel in vivo method for estimation of the pool size of the immediate isoprene precursor, dimethylallyldiphosphate (DMADP) and the activity of isoprene synthase was used. Growth at elevated [CO₂] resulted in greater leaf thickness, more advanced development of mesophyll and moderately increased photosynthetic capacity due to morphological “upregulation”, but isoprene emission rate under growth light and temperature was not significantly different among ambient‐ and elevated‐[CO₂]‐grown plants independent of whether measured at 380 μmol mol^?1 or 780 μmol mol^?1 CO₂. However, DMADP pool size was significantly less in elevated‐[CO₂]‐grown plants, but this was compensated by increased isoprene synthase activity. Analysis of CO₂ and light response curves of isoprene emission demonstrated that the [CO₂] for maximum isoprene emission was shifted to lower [CO₂] in elevated‐[CO₂]‐grown plants. The light‐saturated isoprene emission rate (I_max,Q) was greater, but the quantum efficiency at given I_max,Q was less in elevated‐[CO₂]‐grown plants, especially at higher CO₂ measurement concentration, reflecting stronger DMADP limitation at lower light and higher [CO₂]. These results collectively demonstrate important shifts in light and CO₂‐responsiveness of isoprene emission in elevated‐[CO₂]‐acclimated plants that need consideration in modeling isoprene emissions in future climates.     

干旱胁迫下CO2浓度升高对转StAPX番茄植株耐旱能力的影响

在干旱胁迫伴随大气CO2浓度以及升高的CO2浓度（加倍）条件下,以过量表达番茄类囊体膜抗坏血酸过氧化物酶基因（StAPX）的转基因番茄为试材,探明干旱胁迫TCO2浓度升高对转基因及其野生型番茄植株清除活性氧及耐旱能力的影响。结果表明：升高的CO2浓度明显增加了干旱胁迫下植物的光合水平;升高的CO2浓度明显降低了干旱导致的植物体内H2O2．和O2的积累,影响了干旱胁迫下番茄植株的水．水循环系统的活性氧清除酶活性和小分子抗氧化物质含量;干旱胁迫下即使伴随升高的CO2浓度,测试番茄植株体内的渗透调节物质含量变化也不太明显;升高的CO2浓度明显降低了干旱胁迫下的植物细胞膜伤害程度;干旱胁迫下,升高的CO2浓度对转基因番茄株系比对野生型植株的影响更加明显。结果证明干旱逆境下,升高的CO2浓度能够在一定程度上进一步提高转基因番茄植株的耐旱性。     

The effects of glacial atmospheric CO2 concentrations and climate on isoprene emissions by vascular plants

Isoprene (C₅H₈) emissions by terrestrial vegetation vary with temperature and light intensity, and play an important role in biosphere–chemistry–climate interactions. Such interactions were probably substantially modified by Pleistocene climate and CO₂ cycles. Central to understanding the nature of these modifications is assessment and analysis of how emissions changed under glacial environmental conditions. Currently, even the net direction of change is difficult to predict because a CO₂‐depleted atmosphere may have stimulated emissions compensating for the negative impacts of a cooler climate. Here, we address this issue and attempt to determine the direction of change from an experimental standpoint by investigating the interaction between isoprene emissions and plant growth of two known isoprene‐emitting herbaceous species (Mucuna pruriens and Arundo donax) grown at glacial (180 ppm) to present (366 ppm) CO₂ levels. We found a significant enhancement of isoprene emissions per unit leaf area in M. pruriens under subambient CO₂ concentrations relative to ambient controls but not for A. donax. In contrast, canopy emissions remained unaltered for both plant species because enhanced leaf emissions were offset by reductions in biomass and leaf area. Separate growth experiments with M. pruriens revealed that lowering day/night temperatures by 5°C decreased canopy isoprene emissions irrespective of the CO₂ level. Incorporation of these results into a simple canopy emissions model highlights their potential to attenuate reductions in the total isoprene flux from forests under glacial conditions predicted by standard models.     

西双版纳热带雨林聚果榕隐头果的昆虫群落结构与多样性

聚果榕(Ficus racemasa)是西双版纳热带雨林生态系统中的一个优势种群,在聚果榕隐头果内外,形成了为聚果榕传粉、传播种子和在榕果上取食、栖息、繁衍后代的昆虫群落,在西双版纳勐仑国家自然保护区、植物园沟谷雨林、植物园江边次生林等样地内,对聚果榕隐头果中的昆虫群落结构、动态和多样性进行了研究,样地内采集到184 264号昆虫标本。隶属5个目,19个科,36属,51种;论种类和个体数量,膜翅目最为丰富,占聚果榕隐头果昆虫总数的37．3％,尤其传粉榕小蜂。种群数量在昆虫群落中占绝对优势。其传粉行为影响着聚果榕种群的构建和稳定;鳞翅目占昆虫总数的31．4％;鞘翅目、同翅目和双翅目三类仅占聚果榕隐头果昆虫总数的31．3％,3个样地的多样性指数、均匀度和种类丰富度。以勐仑国家自然保护区最高,其次是沟谷雨林保护区,再次是植物园次生林。资料分析表明,热带雨林片断化使聚果榕隐头果上的昆虫种类呈明显下降趋势,常见种个体数量增加,特有种数量减少。     

Potential effects of elevated CO2 and changes in temperature on tropical plants

Abstract. Very little attention has been directed at the responses of tropical plants to increases in global atmospheric CO₂ concentrations and the potential climatic changes. The available data, from greenhouse and laboratory studies, indicate that the photosynthesis, growth and water use efficiency of tropical plants can increase at higher CO₂ concentrations. However, under field conditions abiotic (light, water or nutrients) or biotic (competition or herbivory) factors might limit these responses. In general, elevated atmospheric CO₂ concentrations seem to increase plant tolerance to stress, including low water availability, high or low temperature, and photoinhibition. Thus, some species may be able to extend their ranges into physically less favourable sites, and biological interactions may become relatively more important in determining the distribution and abundance of species. Tropical plants may be more narrowly adapted to prevailing temperature regimes than are temperate plants, so expected changes in temperature might be relatively more important in the tropics. Reduced transpiration due to decreased stomatal conductance could modify the effects of water stress as a cue for vegetative or reproductive phenology of plants of seasonal tropical areas. The available information suggests that changes in atmospheric CO₂ concentrations could affect processes as varied as plant/herbivore interactions, decomposition and nutrient cycling, local and geographic distributions of species and community types, and ecosystem productivity. However, data on tropical plants are few, and there seem to be no published tropical studies carried out in the field. Immediate steps should be undertaken to reduce our ignorance of this critical area.