Effects of Water Deficits on Carbon Assimilation

This review focuses on the effects of water deficits on photosynthesisand partitioning of assimilates at the leaf level. It is nowestablished that the rate of CO₂ assimilation in the leavesis depressed at moderate water deficits, mostly as a consequenceof stomatal closure. In fact, depending on the species and onthe nature of dehydration, carbon assimilation may diminishto values close to zero without any significant decline in mesophyllphotosynthetic capacity. This remarkable resistance of the photosyntheticapparatus to water deficits became apparent after the measurementof photosynthesis at saturating CO₂ concentrations was madepossible. Whenever light or heat stress are superimposed a declinein mesophyll photosynthesis may occur as a result of a ‘down-regulation’process, which seems to vary among genotypes. A major secondaryeffect of dehydration on photosynthetic carbon metabolism isthe change in partitioning of recently fixed carbon towardssucrose, which occurs in a number of species in parallel tothe increase in starch breakdown. This increase in compoundsof low molecular weight may contribute to an osmotic adjustment.Controlling mechanisms involved in this process deserve furtherinvestigation. Key words: Carbohydrates, carbon partitioning, heat stress, photoinhibition, photosynthetic capacity, stomatal limitation, water deficits     

Growth and Resource Use of Birch Seedlings Under Elevated Carbon Dioxide and Temperature

The effects of elevated CO₂and temperature on the growth, resourceacquisition and resource allocation of small birch seedlings(Betula pendula Roth.) were examined under conditions of non-limitingsoil, water and nutrient supply. Seedlings were planted in potsand placed in controlled environment chambers either under normalambient conditions (CON), or in the presence of elevated CO₂(approx.700 µmol mol^-1; Elev. C), elevated temperature (approx.3 °C above the outside ambient temperature; Elev. T) ora combination of elevated CO₂and elevated temperature (Elev.C + T). Both Elev. C and Elev. T significantly increased biomassaccumulation, but the extent of the increase depended greatlyon the stage of development of the seedlings. Furthermore, thetheoretically expected positive effect of the warmer temperatureon the CO₂-induced stimulation of growth was not observed. Byanalysing resource acquisition (i.e. CO₂ , nitrogen and wateruptake), seedling development, leaf area production and theallocation pattern, it was deduced that the CO₂-stimulated increasein biomass resulted mainly from the initial ‘fertilization’effect of CO₂while the temperature-induced increase in biomassstemmed from higher net carbon intake during the middle andlatter parts of the growing season achieved by virtue of theincreased leaf area and larger photosynthetic capacity. Thelack of positive stimulation by temperature under Elev. C +T may be related in part to (1) CO₂-induced acceleration ofseedling development, which led to a small or no response toCO₂enrichment and lower leaf area production during the latterpart of the growth season, and (2) a cumulative delay in theresponse of growth to the warmer temperature, which did notincrease net carbon intake when the seedlings were at a juvenilestage. Neither Elev. C nor Elev. T altered the root:shoot ratioduring early growth, but Elev. C increased it during the latterpart of the growth season while Elev. T decreased it, possiblyon account of a change in leaf area retention. Finally, thenitrogen and water use efficiencies of seedlings at differentstages of development are discussed. Copyright 2001 Annals ofBotany Company Photosynthesis, growth, resource acquisition and allocation, elevated CO₂and temperature, Betula pendula Roth     

全球变化下植物的碳氮关系及其环境调节研究进展——从分子到生态系统

在全球变化条件下,温度的升高和降水格局的变化,导致淡水资源更加匮乏。环境因子胁迫,如干旱和高温等,它们单独或联合的作用将导致作物大幅度减产,引发自然生态系统退化。植物的碳氮代谢及其分配相互联系、不可分割,其生物过程及外界环境调节共同决定着植物的净生产力和营养水平。该文试图从分子、组织、器官、个体和生态系统等层面上,就植物的碳氮关系及其环境调节(温度、水分和CO₂浓度等)进行综述,并提出了进一步展开相关研究应重点关注的几个方面。     

Long-term exposure to elevated [CO2] in a natural Quercus ilex L. community: net photosynthesis and photochemical efficiency of PSII at different levels of water stress

Naturally grown trees of Mediterranean evergreen oak (Quercus ilex L.), representing the climax species of the region, were enclosed in six large open-top chambers and exposed to ambient and elevated CO₂ concentrations during a 3 year period. Maximum daily net photosynthetic rates measured at the two different CO₂ concentrations were from 30 to 100% higher in elevated than in ambient [CO₂] throughout the experimental period. The increase in maximum daily photosynthesis was also accompanied by a 93% rise in the apparent quantum yield of CO₂ assimilation, measured during periods of optimum soil moisture conditions. Hence, no clear evidence of down-regulation of net photosynthetic activity was found. Interactions between atmospheric CO₂ concentration and plant water stress were studied by following the natural evolution of drought in different seasons and years. At each level of water stress, the maximum rate of carbon assimilation was higher in elevated than in ambient [CO₂] by up to 100%. Analysis of in vivo chlorophyll fluorescence parameters in normal (21%) and low (2%) oxygen concentrations provided useful insights into the functioning and stability of the photosynthetic processes. The photochemical efficiency of PSII (F_v/F_m) progressively decreased as drought conditions became more evident; this trend was accentuated under elevated [CO₂]. Thermal de-excitation processes were possibly more significant under elevated than under ambient [CO₂], in a combination of environmental stresses. This research suggests two possible conclusions: (i) a ‘positive’ interaction between elevated [CO₂] and carbon metabolism can be obtained through relief of water stress limitation in the summer months, and (ii) elevated [CO₂], under drought conditions, may also enhance the significance of slow-relaxing quenching.     

The effect of elevated atmospheric CO2 and drought onstomatal frequency in groundnut (Arachis hypogaea (L.))

The effects of elevated atmospheric CO₂, alone or in combinationwith water stress, on stomatal frequency in groundnut (Arachishypogaea (L.) cv. Kadiri-3) were investigated. CO₂ exerted significanteffects on stomatal frequency only in irrigated plants. Theeffects of drought on leaf development out weighed the smallereffects of CO₂ concentration, although reductions in stomatalfrequency induced by elevated atmo-spheric CO₂ were still observed.When stands of groundnut were grown under irrigated conditionswith unrestricted root systems, an increase in atmospheric CO₂from 375 to 700 ppmv decreased stomatal frequency on both leafsurfaces by up to 16% in droughted plants, stomatal frequencywas reduced by 8% on the adaxial leaf surface only. Elevatedatmospheric CO₂ promoted larger reductions in leaf conductancethan the changes in stomatal frequency, indicating partial stomatalclosure. As a result, the groundnut stands grown at elevatedCO₂ utilized the available soil moisture more slowly than thosegrown under ambient CO₂, there by extending the growing period.Despite the large variations in cell frequencies induced bydrought, there was no treatment effect on either stomatal indexor the adaxial/abaxial stomatalfrequency ratio. The data suggestthat the effects of future increases in atmospheric CO₂ concentrationon stomatal frequency in groundnut are likely to be small, especiallyunder conditions of water stress, but that the combination ofassociated reductions in leaf con-ductance and enhanced assimilationat elevated CO₂ will be important in semi-arid regions Key words: Arachis hypogaea L, Leguminosae, groundnu, stomatal frequency, CO₂, drought     

Acclimation of nitrogen uptake capacity of rice to elevated atmospheric CO2 concentration

Background and Aims: Nitrogen (N) is a major factor affecting yield gain of cropsunder elevated atmospheric carbon dioxide concentrations [CO₂].It is well established that elevated [CO₂] increases root mass,but there are inconsistent reports on the effects on N uptakecapacity per root mass. In the present study, it was hypothesizedthat the responses of N uptake capacity would change with theduration of exposure to elevated [CO₂]. Methods: The hypothesis was tested by measuring N uptake capacity inrice plants exposed to long-term and short-term [CO₂] treatmentsat different growth stages in plants grown under non-limitingN conditions in hydroponic culture. Seasonal changes in photosynthesisrate and transpiration rate were also measured. Key Results: In the long-term [CO₂] study, leaf photosynthetic responsesto intercellular CO₂ concentration (Ci) were not affected byelevated [CO₂] before the heading stage, but the initial slopein this response was decreased by elevated [CO₂] at the grain-fillingstage. Nitrate and ammonium uptake capacities per root dry weightwere not affected by elevated [CO₂] at panicle initiation, butthereafter they were reduced by elevated [CO₂] by 31–41% at the full heading and mid-ripening growth stages. In theshort-term study (24 h exposures), elevated [CO₂] enhanced nitrateand ammonium uptake capacities at the early vegetative growthstage, but elevated [CO₂] decreased the uptake capacities atthe mid-reproductive stage. Conclusions: This study showed that N uptake capacity was downregulated underlong-term exposure to elevated [CO₂] and its response to elevated[CO₂] varied greatly with growth stage.     

The Effects of Elevated Atmospheric Carbon Dioxide and Water Stress on Ligth Interception, Dry Matter Production and Yield in Stands of Groundnut (Arachis hypogaea L.

Stands of groundnut (Arachis hypogaea L.), a C₃ legume, weregrown in controlled-environment glasshouses at 28 °C (±5°C)under two levels of atmospheric CO₂ (350 ppmv or 700 ppmv) andtwo levels of soil moisture (irrigated weekly or no water from35 d after sowing). Elevated CO₂ increased the maximum rate of net photosynthesisby up to 40%, with an increase in conversion coefficient forintercepted radiation of 30% (from 1–66 to 2–16g MJ^–1) in well-irrigated conditions, and 94% (from 0–64to 1·24 g MJ^–1) on a drying soil profile. In plantswell supplied with water, elevated CO₂ increased dry matteraccumulation by 16% (from 13·79 to 16·03 t ^–1) and pod yield by 25% (from 2·7 to 3·4t ha^–1).However, the harvest index (total poddry weight/above-grounddry weight) was unaffected by CO₂ treatment. The beneficial effects of elevated CO₂ were enhanced under severewater stress, dry matter production increased by 112% (from4·13 to 8·87 t ha^–1) and a pod yield of1·34t ha^–1 was obtained in elevated CO₂, whereascomparable plotsat 350 ppmv CO₂ only yielded 0·22 t ha^-1.There was a corresponding decrease in harvest index from 0·15to 0·05. Following the withholding of irrigation, plants growing on astored soil water profile in elevated CO₂ could maintain significantlyless negative leaf water potentials (P<0·01) for theremainder of the season than comparable plants grown in ambientCO₂, allowing prolonged plant activity during drought. In plants which were well supplied with water, allocation ofdry matter between leaves, stems, roots, and pods was similarin both CO₂ treatments. On a drying soil profile, allocationin plants grown in 350 ppmv CO₂ changed in favour of root developmentfar earlier in the season than plants grown at 700 ppmv CO₂,indicating that severe waterstress was reached earlier at 350ppmv CO₂. The primary effects of elevated CO₂ on growth and yield of groundnutstands weremediated by an increase in the conversion coefficientfor intercepted radiation and the prolonged maintenance of higherleaf water potentials during increasing drought stress. Key words: Arachis hypogaea, elevated CO₂, water stress, dry matter production     

Photosynthetic Carbon Reduction and Carbon Oxidation Cycles are the Main Electron Sinks for Photosystem II Activity During a Mild Drought

Stomatal closure can explain the inhibition of net CO₂ uptakeby a leaf subjected to a mild drought: the photosynthetic apparatusappears resistant to lack of water. Changes in both the watercontent of leaves maintained in a constant environment and theambient CO₂ molar fraction during measurements on well-hydratedleaves lead to similar effects on net CO₂ uptake and whole chainelectron transport as estimated by leaf chlorophyll fluorescencemeasurements. In particular, it is shown that photosystem II(PSII) functioning and its regulation are not qualitativelychanged during desiccation and that the variations in PSII photochemistrycan simply be understood by changes in substrate availabilityin this condition. Moreover, an analysis of the literature showsthat when inhibition of net CO₂ uptake by C₃ leaves under drought(Phaseolus vulgaris L., Helianthus annus L. and Solanum tuberosumL.) was lower than 80 %, elevated CO₂ completely restored thephotosynthetic capacity. The CO₂ molar fraction in the chloroplastsdeclines as stomata close in drying leaves. As a consequence,in C₃ plants, ribulose-1,5-bisphosphate oxygenation increasesand becomes the main sink for photosynthetic electrons. Dependingon the prevailing photon flux density, the O₂ uptake throughphotorespiratory activity can entirely replace carbon dioxideas an electron acceptor, or not. The rate of the Mehler reactionremains low and unchanged during desiccation. However, droughtcould also involve CO₂-sensitive modification of the photosyntheticmetabolism depending on plant growth conditions and possiblyalso on plant species.     

Soybean nodulation and N2 fixation response to drought under carbon dioxide enrichment

The combined effects of carbon dioxide (CO₂) enrichment and water deficits on nodulation and N₂ fixation were analysed in soybean [Glycine max (L.) Merr.]. Two short-term experiments were conducted in greenhouses with plants subjected to soil drying, while exposed to CO₂ atmospheres of either 360 or 700 μmol CO₂ mol^–1. Under drought-stressed conditions, elevated [CO₂] resulted in a delay in the decrease in N₂ fixation rates associated with drying of the soil used in these experiments. The elevated [CO₂] also allowed the plants under drought to sustain significant increases in nodule number and mass relative to those under ambient [CO₂]. The total non-structural carbohydrate (TNC) concentration was lower in the shoots of the plants exposed to drought; however, plants under elevated CO₂ had much higher TNC levels than those under ambient CO₂. For both [CO₂] treatments, drought stress induced a substantial accumulation of TNC in the nodules that paralleled N₂ fixation decline, which indicates that nodule activity under drought may not be carbon limited. Under drought stress, ureide concentration increased in all plant tissues. However, exposure to elevated [CO₂] resulted in substantially less drought-induced ureide accumulation in leaf and petiole tissues. A strong negative correlation was found between ureide accumulation and TNC levels in the leaves. This relationship, together with the large effect of elevated [CO₂] on the decrease of ureide accumulation in the leaves, indicated the importance of ureide breakdown in the response of N₂ fixation to drought and of feedback inhibition by ureides on nodule activity. It is concluded that an important effect of CO₂ enrichment on soybean under drought conditions is an enhancement of photoassimilation, an increased partitioning of carbon to nodules and a decrease of leaf ureide levels, which is associated with sustained nodule growth and N₂ rates under soil water deficits. We suggest that future [CO₂] increases are likely to benefit soybean production by increasing the drought tolerance of N₂ fixation.     

不同植物叶片水分利用效率对光和CO2的响应与模拟

植物叶片水分利用效率的高低取决于气孔控制的光合作用和蒸腾作用两个相互耦合的过程,模拟水分利用效率对环境变化的响应特征和机制是理解生态系统碳循环和水循环及其耦合关系的基础.研究通过人工控制光强和CO2浓度,对叶片水分利用效率进行了研究.提出了植物水分利用效率在光强和CO2浓度共同作用下的估算模型.数据分析表明,该模型在包括C3和C4植物、草本和木本植物在内的9种植物上能很好地模拟水分利用效率对光强和CO2浓度共同作用的响应.该模型可以用于估算CO2浓度升高条件下光合速率的提高和蒸腾速率的降低对水分利用效率提高的贡献量.CO2浓度变化条件下,水分利用效率在不同植物之间有巨大差异,研究区域尺度植物的水分利用效率时至少需要将植物区分为C4植物和C3植物,其中C3植物区分为草本和木本植物3种生态功能型才能较为准确地估算植物的整体水分利用效率.应用本研究提出的水分利用效率估算模型和植物水分利用效率生态功能型分类标准,可以为建立以植物的水分利用效率为基本参数的陆地生态系统水循环模型和陆地生态系统生产力模型提供重要依据.     

Responses of water use efficiency of 9 plant species to light and CO2 and their modeling

Photosynthesis coupled with transpiration determines water use efficiency (WUE) at leaf level, and the responses of WUE controlled by gas exchanges through stomata to environment are the basis of carbon and water cycles in the ecosystem. In this paper, by using Li-6400 Portable Photosynthesis System (LI-COR), WUE at leaf level was analyzed under controlled photosynthetic photons flux density (PPFD) and CO₂ concentration conditions across 9 plant species including maize (Zea mays), sorghum (Sorghum vulgare), millet (Setaria italica), soybean (Glycine max), peanut (Arachis phyogaea), sweet potato (Ipomoea batatas), rice (Oryza sativa), Masson pine (Pinus massoniana) and Schima superba. We had developed a new model to estimate the water use efficiency in response to the combined effects of light and CO₂ concentration. Our measured data validated that this model could simulate the changes of water use efficiency very well under combined effect of light and CO₂ concentration. It could be used to estimate contribution of photosynthesis increase and transpiration decline on water use efficiency with the rising of CO₂ concentration. Great differences in water use efficiency occurred in these different plant species under various CO₂ concentration levels. Based on water use efficiency at regional scale, we concluded that plants should be separated into C₃ plants and C₄ plants, and furthermore, C₃ plants should be separated into herbaceous plants and woody plants. Our separation criteria would do a great favor in modeling the evapotranspiration of terrestrial ecosystem with carbon and water balance.     

短期CO2加富对凤梨光合作用及生长发育的影响

 研究了CO₂加富对丹尼斯凤梨（Guzmania`Denise’）和吉利凤梨（Guzmania `Cherry’）叶片光合速率、植株生长、开花和光合相关酶活性的 影响。结果表明,处理30 d期间,处理(600±40)、(900±40) μmol CO₂&;#8226;mol^-1的净光合速率分别比同期对照增加了6.24%～31.91%和11.92%～ 41.48%;CO₂加富下促进了叶片中可溶性糖和淀粉的积累, 蒸腾速率和气孔导度下降,Rubisco活性增加,乙醇酸氧化酶活性则明显下降。(600 ±40)μmol CO₂&;#8226;mol^-1处理下的株高、叶面积分别比同期对照下增加了6.94%～14.63%和1.66%～7. 06%,而处理(900±40) μmol CO₂&;#8226;mol^-1下 分别增加了9.71%～20.85%和2.87%～11.62%;CO₂加富下促进了干重和鲜重的积累。此外,CO₂加富提前了吉利凤梨的花期。     

Effects of nitrogen on Pinus palustris foliar respiratory responses to elevated atmospheric CO2 concentration

Indirect effects of atmospheric CO₂ concentration [CO₂], onlongleaf pine (Pinus palustris Mill.) foliage respiration werestudied by growing trees in a factorial arrangement of low andhigh [CO₂] (369 and 729µmol CO₂ mol^–1) and low andhigh N (40 and 400 kg ha^–1 yr^–1). Direct effectsof [CO₂] on leaf respiration were tested by measuring respirationrates of foliage from all treatments at two CO₂ levels (360and 720µmol CO₂mol^–1) at the time of measurement.Elevated CO₂ did not directly or indirectly affect leaf respirationwhen expressed on a leaf area or mass basis, but a significantincrease in respiration per unit leaf N was observed in treesgrown in elevated [CO₂] (indirect response to elevated [CO₂]).The lack of a [CO₂] effect on respiration, when analysed onan area or mass basis, may have resulted from combined effectsof [CO₂] on factors that increase respiration (e.g. greateravailability of non-structural carbohydrates stimulating growthand carbon export from leaves) and on factors that decreaserespiration (e.g. lower N concentration leading to lower constructioncosts and maintenance requirements). Thus, [CO₂] affected factorsthat influence respiration, but in opposing ways. Key words: Pinus palustris, elevated CO₂, nitrogen, foliar, respiration     

The influence of elevated C02 on growth and age-related changes in leaf gas exchange

Rumex obtusifolius plants were grown for several months in daylitenvironment chambers (Solardomes) force-ventilated with aircontaining 350 or 600 µ;mol mol^–1 C0₂. ElevatedCO₂ was found to accelerate the natural ontogenic decline inphotosynthesis, but did not reduce leaf duration. In both CO₂treatments photosynthetic rates declined progressively withincreasing leaf age, the decline being greater for plants grownin elevated C0₂ such that rates became lower than in ambientCO₂. The degree of CO₂-induced photosynthetic down-regulationas determined by A/C₁ analysis was found to be dependent onleaf age. The major contribution to the decline in photosynthesiswas likely to be a reduction in Rubisco activity as changesin stomataland mesophyll limitations were small. Instantaneouswater use efficiency (WUE₁) was greater for plants in elevatedCO₂, but these values declined rapidly with leaf age, whereasin ambient CO₂ values were always lower, but were maintainedfor longer. Growth analysis indicated an increased root:-shootratio for plants grown in elevated CO₂, this occurring almostentirely as a result of increased root growth. Greater rootproliferation and increased WUE₁, are characteristics whichshould give this persistent and troublesome weed an increasedcompetitive advantage under projected conditions of climatechange Key words: tusifoliu, elevated CO₂, gas exchange, leaf age, senescence     

CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

Concentration and {sigma}13C of leaf carbohydrates in relation to gas exchange in Quercus robur under elevated CO2 and drought

The variations of leaf carbohydrate concentration, carbon isotopediscrimination () of leaf soluble carbohydrate, gas-exchangeand growth during a soil drying cycle under 350 and 700 µmolmol^-1 CO₂ concentrations ([CO₂]) inQuercus robur seedlings wereanalysed. In well-watered conditions, a doubling of [CO₂] causedan increase of CO₂ assimilation rate (A) ( +47%) and a decreaseof stomatal conductance for water vapour (g) (–25%),anddoubled the intrinsic water-use efficiency (A/g). The valuesof A were not affected by elevated [CO₂] which was consistentwith the 2-fold increase of A/g. Elevated [CO₂ also significantlyincreased sucrose and starch leaf concentrations as well asaerial growth and plant dry weight. The stimulating effect ofCO₂ enrichment on A and A/g was maintained in moderate droughtconditions, but disappeared in the most severe drought conditions.Drought induced an increase of hexose concentrations in both[CO₂], but this effect was more pronounced under elevated [CO₂],which may contribute to increase osmoregulation. From the onsetof drought, starch was depleted in both [CO₂]. Carbon isotopediscrimination decreased in response to drought, which correspondedto an increase in A/g according to the two-step model of isotopicdiscrimination. In contrast, the A/g values derived from instantaneousleaf gas-exchange measurements decreased along the drying cycle.The discrepancy observed between the two independent estimatesof water-use efficiency is discussed in terms of time-scaleintegration. The results obtained with the isotopic approachusing soluble carbohydrate suggest a predominant stomatal limitationof CO2 assimilation in response to drought. Soil drying cycle,elevated CO₂, leaf gas-exchange, leaf carbohydrate concentrations,carbon isotope discrimination, growth, Quercus robur. Key words: soil drying cycle, elevated CO₂, leaf gas-exchange, leaf carbohydrate concentrations, carbon isotope discrimination, growth, Quercus robur     

Elevated CO<Subscript>2</Subscript>, litter chemistry,and decomposition: a synthesis

The results of published and unpublished experiments investigating the impacts of elevated [CO₂] on the chemistry of leaf litter and decomposition of plant tissues are summarized. The data do not support the hypothesis that changes in leaf litter chemistry often associated with growing plants under elevated [CO₂] have an impact on decomposition processes. A meta-analysis of data from naturally senesced leaves in field experiments showed that the nitrogen (N) concentration in leaf litter was 7.1% lower in elevated [CO₂] compared to that in ambient [CO₂]. This statistically significant difference was: (1) usually not significant in individual experiments, (2) much less than that often observed in green leaves, and (3) less in leaves with an N concentration indicative of complete N resorption. Under ideal conditions, the efficiency with which N is resorbed during leaf senescence was found not to be altered by CO₂ enrichment, but other environmental influences on resorption inevitably increase the variability in litter N concentration. Nevertheless, the small but consistent decline in leaf litter N concentration in many experiments, coupled with a 6.5% increase in lignin concentration, would be predicted to result in a slower decomposition rate in CO₂-enriched litter. However, across the assembled data base, neither mass loss nor respiration rates from litter produced in elevated [CO₂] showed any consistent pattern or differences from litter grown in ambient [CO₂]. The effects of [CO₂] on litter chemistry or decomposition were usually smallest under experimental conditions similar to natural field conditions, including open-field exposure, plants free-rooted in the ground, and complete senescence. It is concluded that any changes in decomposition rates resulting from exposure of plants to elevated [CO₂] are small when compared to other potential impacts of elevated [CO₂] on carbon and N cycling. Reasons for experimental differences are considered, and recommendations for the design and execution of decomposition experiments using materials from CO₂-enrichment experiments are outlined.     

Elevated CO2 alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis

We analysed the impact of elevated CO₂ on water relations, water use efficiency and photosynthetic gas exchange in barley (Hordeum vulgare L.) under wet and drying soil conditions. Soil moisture was less depleted under elevated compared to ambient [CO₂]. Elevated CO₂ had no significant effect on the water relations of irrigated plants, except on whole plant hydraulic conductance, which was markedly decreased at elevated compared to ambient CO₂ concentrations. The values of relative water content, water potential and osmotic potential were higher under elevated CO₂ during the entire drought period. The better water status of water-limited plants grown at elevated CO₂ was the result of stomatal control rather than of osmotic adjustment. Despite the low stomatal conductance produced by elevated CO₂, net photosynthesis was higher under elevated than ambient CO₂ concentrations. With water shortage, photosynthesis was maintained for longer at higher rates under elevated CO₂. The reduction of stomatal conductance and therefore transpiration, and the enhancement of carbon assimilation by elevated CO₂, increased instantaneous and whole plant water use efficiency in both irrigated and droughted plants. Thus, the metabolism of barley plants grown under elevated CO₂ and moderate or mild water deficit conditions is benefited by increased photosynthesis and lower transpiration. The reduction in plant water use results in a marked increase in soil water content which delays the onset and severity of water deficit.     

Oxygen Exchange in Relation to Carbon Assimilation in Water-stressed Leaves During Photosynthesis

In a study on metabolic consumption of photosynthetic electronsand dissipation of excess light energy under water stress, O₂and CO₂ gas exchange was measured by mass spectrometry in tomatoplants using ¹⁸O₂ and ¹³CO₂. Under water stress, gross O₂ evolution(E_O), gross O₂ uptake (U_O), net CO₂ uptake (P_N), gross CO₂ uptake(TPS), and gross CO₂ evolution (E_C) declined. The ratio P_N/E_Ofell during stress, while the ratios U_O/E_O and E_C/TPS rose.Mitochondrial respiration in the light, which can be measureddirectly by ¹²CO₂ evolution during ¹³CO₂ uptake at 3000 µll^{–1 13}CO₂, is small in relation to gross CO₂ evolutionand CO₂ release from the glycolate pathway. It is concludedthat PSII, the Calvin cycle and mitochondrial respiration aredown-regulated under water stress. The percentages of photosyntheticelectrons dissipated by CO₂ assimilation, photorespiration andthe Mehler reaction were calculated: in control leaves morethan 50 % of the electrons were consumed in CO₂ assimilation,23 % in photorespiration and 13 % in the Mehler reaction. Undersevere stress the percentages of electrons dissipated by CO₂assimilation and the Mehler reaction declined while the percentageof electrons used in photorespiration doubled. The consumptionof electrons in photorespiration may reduce the likelihood ofdamage during water deficit.     

Simulating Growth and Root-shoot Partitioning in Prairie Grasses Under Elevated Atmospheric CO2and Water Stress

We constructed a model simulating growth, shoot-root partitioning,plant nitrogen (N) concentration and total non-structural carbohydratesin perennial grasses. Carbon (C) allocation was based on theconcept of a functional balance between root and shoot growth,which responded to variable plant C and N supplies. Interactionsbetween the plant and environment were made explicit by wayof variables for soil water and soil inorganic N. The modelwas fitted to data on the growth of two species of perennialgrass subjected to elevated atmospheric CO₂and water stresstreatments. The model exhibited complex feedbacks between plantand environment, and the indirect effects of CO₂and water treatmentson soil water and soil inorganic N supplies were important ininterpreting observed plant responses. Growth was surprisinglyinsensitive to shoot-root partitioning in the model, apparentlybecause of the limited soil N supply, which weakened the expectedpositive relationship between root growth and total N uptake.Alternative models for the regulation of allocation betweenshoots and roots were objectively compared by using optimizationto find the least squares fit of each model to the data. Regulationby various combinations of C and N uptake rates, C and N substrateconcentrations, and shoot and root biomass gave nearly equivalentfits to the data, apparently because these variables were correlatedwith each other. A partitioning function that maximized growthpredicted too high a root to shoot ratio, suggesting that partitioningdid not serve to maximize growth under the conditions of theexperiment.Copyright 1998 Annals of Botany Company plant growth model, optimization, nitrogen, non-structural carbohydrates, carbon partitioning, elevated CO₂, water stress,Pascopyrum smithii,Bouteloua gracilis, photosynthetic pathway, maximal growth