High precision measurements of atmospheric concentrations and plant exchange rates of carbonyl sulfide using mid‐IR quantum cascade laser

Linking measurements of carbonyl sulfide (COS) with those of carbon dioxide (CO₂) has a potential in providing a powerful tracer of gross CO₂ fluxes between the atmosphere and land plants, a critical element in understanding the response of the land biosphere to global change. A new application of online COS, CO₂ and water vapor measurements based on newly designed mid‐infrared (IR) dual quantum cascade laser (QCL) spectrometer measures COS and CO₂ (at 2056 cm^?1) and water vapor (at 2190 cm^?1), with detectors cooled thermoelectrically (at ?43 °C) or with liquid nitrogen (?197 °C) for improved precision. Using the cryogenic detectors with averaging time of 1 s, precision was 50 pmol mol^?1, 0.4 μmol mol^?1 and 0.01 mmol mol^?1 for COS, CO₂ and water vapor, respectively (14, 0.2 and 0.003, respectively, for 60 s averaging time). We measured COS concentrations in ambient air, and changes in the rates of COS, CO₂ and water vapor exchange of attached leaves in response to changes in light intensity and ambient COS concentrations. The results were consistent with those of nononline gas chromatography–mass spectrometry for COS and IR gas analyzer for CO₂ and water vapor, with a high linear correlation for a broad range of concentrations (R²= 0.85 for COS and R²= 1.00 for CO₂ and water vapor). The new methodology opens the way for lab and field explorations of COS fluxes as a powerful new tracer for CO₂ exchange in the land biosphere.     

Spatial and temporal variability of dissolved sulfur compounds in European estuaries

The spatial and temporal distribution of Dimethylsulfide (DMS),Dimethylsulfoniopropionate (DMSP) – its precursor –,Dimethylsulfoxyde (DMSO) – one of its oxidation products –Carbonyl sulfide (COS) and Carbonyl disulfide (CS₂) wereinvestigated during nine oceanographic cruises carried out in six majortidal European estuaries between July 1996 and May 1998. Low levels ofDMS were recorded (mean of 0.6 nM, standard deviation () 0.3 nM)during these 9 cruises. Positive correlations between DMS and salinitywere frequently observed, with the highest DMS concentrations in theplume of estuaries, which could be explained by change in phytoplanktonspeciation from estuarine to shelf waters. Strong correlation betweenDMSP and DMS was reported for most of the estuaries denoting a simpleconservative mixing of riverine and marine waters controlled by tide. Incontrast to DMS, significant levels of COS and CS₂ with meanconcentrations of 220 ± 150 (pM; pM =10^–12 M) and 25 ± 6 pM respectively wererecorded in four estuaries, indicating that estuaries could be asignificant source of these compounds.     

The effect of plant material grown under elevated CO2 on soil respiratory activity

The biodegradability of aerial material from a C4 plant, sorghum grown under ambient (345 µmol mol^–1) and elevated (700 µmol mol^–1) atmospheric CO₂ concentrations were compared by measuring soil respiratory activity. Initial daily respiratory activity (measured over 10 h per day) increased four fold from 110 to 440 cm³ CO₂ 100g dry weight soil^–1 in soils amended with sorghum grown under either elevated or ambient CO₂. Although soil respiratory activity decreased over the following 30 days, respiration remained significantly higher (t-test;p>0.05) in soils amended with sorghum grown under elevated CO₂ concentrations. Analysis of the plant material revealed no significant differences in C:N ratios between sorghum grown under elevated or ambient CO₂. The reason for the differences in soil respiratory activity have yet to be elucidated. However if this trend is repeated in natural ecosystems, this may have important implications for C and N cycling.     

土壤氮水交互对马尾松和杉木COS和CO2通量的影响

羰基硫(COS)和CO_2化学结构相似,且植物对COS和CO_2具有共吸收特性,因此可利用COS作为示踪物来估算生态系统总初级生产力,而不同植物吸收COS和CO_2对环境因子变化的响应差异较大。以南亚热带典型树种马尾松(Pinus massoniana)和杉木(Cunninghamia lanceolata)为研究对象,设置2个氮水平及3个土壤水分梯度处理。采取顶空套袋法采集气体样品,用预浓缩—气质联用仪分析样品COS浓度,同时测量植物光合参数。结果表明:马尾松和杉木吸收COS,吸收速率均值分别为39.58—127.27 pmol m~(-2) s~(-1)和0.81—66.92 pmol m~(-2) s~(-1)。整体而言,施氮可促进植物吸收COS,但除施氮对马尾松COS通量有显著影响外(P0.05),施氮、土壤水分和两者交互作用对马尾松和杉木的COS和CO_2通量及其比值均无显著性影响。施氮情况下,高土壤水分处理促进马尾松COS吸收而低土壤水分处理促进杉木COS吸收。中等土壤水分和高土壤水分条件下马尾松和杉木COS通量与气孔导度呈正相关关系。线性拟合结果表明,植物COS通量(F_(COS))与CO_2通量(F_(CO_2))呈极显著正相关(P0.01),马尾松和杉木F_(COS)/F_(CO_2)值分别为1.48×10~(-6)和1.01×10~(-6)。中等土壤水分条件均可提高马尾松F_(COS)/F_(CO_2)比值,而低土壤水分条件下施氮增加杉木F_(COS)/F_(CO_2)比值,高土壤水分条件下施氮降低杉木F_(COS)/F_(CO_2)比值。低土壤水分和高土壤水分使马尾松蒸汽压亏缺增大,促使气孔导度减小从而降低净光合速率。低土壤水分和高土壤水分下施氮导致杉木气孔导度增加从而增强净光合速率。研究结果不仅对进一步了解区域氮沉降和降水对树木COS通量及F_(COS)/F_(CO_2)的影响有重要意义,而且可为模型估算总初级生产力提供区域性数据支持。     

Enzymatic Pathways for the Consumption of Carbonyl Sulphide (COS) by Higher Plants*

Carbonyl sulphide (COS) is an important trace gas of the atmosphere. Considerable uncertainties remain concerning the global sinks of COS. Vegetation is believed to be an unqualified sink in the global cycle of COS. We investigated whether there is an enzymological background for the consumption of COS by higher plants in analogy to CO₂. Photometric measurements demonstrated that all enzymes involved in C0₂ assimilation by higher plants can also metabolise COS. The key enzyme for COS metabolism in higher plants is carbonic anhydrase, an enzyme which probably directly splits COS into C0₂ and H₂S. Such a pathway would explain the observed deposition of COS to vegetation.     

Plant Nitrogen Dynamics in Shortgrass Steppe under Elevated Atmospheric Carbon Dioxide

The direct and indirect effects of increasing levels of atmospheric carbon dioxide (CO₂) on plant nitrogen (N) content were studied in a shortgrass steppe ecosystem in northeastern Colorado, USA. Beginning in 1997 nine experimental plots were established: three open-top chambers with ambient CO₂ levels (approximately 365 mol mol^–1), three open-top chambers with twice-ambient CO₂ levels (approximately 720 mol mol^–1), and three unchambered control plots. After 3 years of growing-season CO₂ treatment, the aboveground N concentration of plants grown under elevated atmospheric CO₂ decreased, and the carbon–nitrogen (C:N) ratio increased. At the same time, increased aboveground biomass production under elevated atmospheric CO₂ conditions increased the net transfer of N out of the soil of elevated-CO₂ plots. Aboveground biomass production after simulated herbivory was also greater under elevated CO₂ compared to ambient CO₂. Surprisingly, no significant changes in belowground plant tissue N content were detected in response to elevated CO₂. Measurements of individual species at peak standing phytomass showed significant effects of CO₂ treatment on aboveground plant tissue N concentration and significant differences between species in N concentration, suggesting that changes in species composition under elevated CO₂ will contribute to overall changes in nutrient cycling. Changes in plant N content, driven by changes in aboveground plant N concentration, could have important consequences for biogeochemical cycling rates and the long-term productivity of the shortgrass steppe as atmospheric CO₂ concentrations increase.     

Elevated CO2 and plant nitrogen-use: is reduced tissue nitrogen concentration size-dependent?

Plants often respond to elevated atmospheric CO₂ levels with reduced tissue nitrogen concentrations relative to ambient CO₂-grown plants when comparisons are made at a common time. Another common response to enriched CO₂ atmospheres is an acceleration in plant growth rates. Because plant nitrogen concentrations are often highest in seedlings and subsequently decrease during growth, comparisons between ambient and elevated CO₂-grown plants made at a common time may not demonstrate CO₂-induced reductions in plant nitrogen concentration per se. Rather, this comparison may be highlighting differences in nitrogen concentration between bigger, more developed plants and smaller, less developed plants. In this study, we directly examined whether elevated CO₂ environments reduce plant nitrogen concentrations independent of changes in plant growth rates. We grew two annual plant species. Abutilon theophrasti (C₃ photosynthetic pathway) and Amaranthus retroflexus (C₄ photosynthetic pathway), from seed in glass-sided growth chambers with atmospheric CO₂ levels of 350 mol·mol^–1 or 700 mol·mol^–1 and with high or low fertilizer applications. Individual plants were harvested every 2 days starting 3 days after germination to determine plant biomass and nitrogen concentration. We found: 1. High CO₂-grown plants had reduced nitrogen concentrations and increased biomass relative to ambient CO₂-grown plants when compared at a common time; 2. Tissue nitrogen concentrations did not vary as a function of CO₂ level when plants were compared at a common size; and 3. The rate of biomass accumulation per rate of increase in plant nitrogen was unaffected by CO₂ availability, but was altered by nutrient availability. These results indicate that a CO₂-induced reduction in plant nitrogen concentration may not be due to physiological changes in plant nitrogen use efficiency, but is probably a size-dependent phenomenon resulting from accelerated plant growth.     

The influence of plant species,fertilization and elevated CO2 on soil aggregate stability

We tested the effects of plant species, fertilization and elevated CO₂ on water-stable soil aggregation. Five annual grassland species and a plant community were grown in outdoor mesocosms for 4 years, with and without NPK fertilization, at ambient or elevated atmospheric CO₂ concentrations. Aggregate stability (resistance of aggregates to slaking) in the top 0.15 m of soil differed among plant species. However, the more diverse plant community did not enhance aggregate stability relative to most monocultures. Species differences in aggregate stability were positively correlated with soil active bacterial biomass, but did not correlate with root biomass or fungal length. Plant species did not affect aggregate stability lower in the soil profile (0.15–0.45 m), where soil biological activity is generally decreased. Elevated CO₂ and NPK fertilization altered many of the factors known to influence aggregation, but did not affect water-stable aggregation at either depth, in any of the plant treatments. These results suggest that global changes will alter soil structure primarily due to shifts in vegetation composition.     

CO2 environment,microclimate and photosynthetic characteristics of the moss Hylocomium splendens in a subarctic habitat

Summary In order to document the natural CO₂ environment of the moss Hylocomium splendens, and ascertain whether or not the moss was adapted to this, and its interactions with other microenvironmental factors, two studies were carried out. Firstly, the seasonal variations of CO₂ concentration, photosynthetically active radiation (PAR), tissue water content and temperature were measured in the natural microenvironment of H. splendens in a subarctic forest during the summer period (July–September). Secondly, the photosynthetic responses of the species to controlled CO₂ concentrations, PAR, temperature, and hydration were measured in the laboratory. CO₂ concentrations around the upper parts of the plant, when PAR was above the compensation point (30 mol m^–2 s^–1), were mostly between 400 and 450 ppm. They occasionally increased up to 1143 ppm for short periods. PAR flux densities below saturating light levels for photosynthesis (100 mol m^–2 s^–1), occurred during 65% (July), 76% (August) and 96% (September) of the hours of the summer period. The temperature optimum of photosynthesis was 20° C: this temperature coincided with PAR above the compensation point during 5%, 6% and 0% of the time in July, August and September, respectively. Optimal hydration of tissues was infrequent. Hence PAR, temperature and water limit CO₂ uptake for most of the growing season. Our data suggest that the higher than normal ambient CO₂ concentration in the immediate environment of the plant counteracts some of the limitations in PAR supply that it experiences in its habitat. This species already experiences concentrations of atmospheric CO₂ predicted to occur over the next 50 years.     

The Effects of Cold Acclimation of Winter Wheat Plants on Changes in CO<Subscript>2</Subscript> Exchange and Phenolic Compound Formation

We studied CO₂ exchange and phenolic compound production in various organs of unhardened and hardened winter wheat (Triticum aestivum L.) plants. The rates of CO₂ assimilation at saturating illumination (photosynthesis) and CO₂ evolution in darkness (respiration) declined substantially at the autumnal decrease of ambient temperature. However, because of a higher cold resistance of photosynthesis, the ratio of photosynthesis to respiration rates increased 1.5-fold. These gas exchange changes were accompanied by the accumulation of total soluble phenolics in leaves and a polymeric phenolic compound lignin in roots. We did not observe any changes in the production of either soluble or polymeric (lignin) phenolics in crowns.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 3, 2005, pp. 366–371.Original Russian Text Copyright © 2005 by Zagoskina, Olenichenko, Klimov, Astakhova, Zhivukhina, Trunova.     

Identification of COS markers in the Pinaceae

Conserved ortholog set (COS) markers are evolutionary conserved, single-copy genes, identified from large databases of express sequence tags (ESTs). They are of particular use for constructing syntenic genetic maps among species. In this study, we identified a set of 1,813 putative single-copy COS markers between spruce and loblolly pine, then designed primers for 931 of these markers and tested these primers with DNA from spruce, pine, and Douglas fir. Of these 931 primers, 56% (524) amplified a product in both spruce and pine, and 71% (373) of these were single-banded; 224 amplicons were single-banded in all three species. Even though these COS markers were selected from large EST databases, a substantial proportion (20–30%) of amplicons displayed multiple bands or smears, suggesting significant paralogy. Sequencing of three single-banded amplicons showed high nucleotide similarities among 29 conifer species, suggesting orthology of single-banded amplicons. Screening for COS marker polymorphism in two pedigrees of white spruce and two pedigrees of loblolly pine revealed an average informativeness of 36% for spruce and 24% for pine (e.g., at least one parent was heterozygous for a single-nucleotide polymorphism within the entire amplified product). This corresponds to an average nucleotide heterozygosity of 0.05% and 0.03%, respectively, which is considerably lower than that found in other studies of spruce and pine. Thus, the advantages of COS markers for constructing syntenic maps are offset by their lower polymorphism. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Plant growth and reproduction along CO2 gradients: non-linear responses and implications for community change

The effects of rising atmospheric CO₂ concentrations on natural plant communities will depend upon the cumulative responses of plant growth and reproduction to gradual, incremental changes in climatic conditions. We analysed published studies of plant responses to elevated CO₂ to address whether reproductive and total biomass exhibit similar enhancement to elevated vs. ambient CO₂ concentrations, and to assess the patterns of plant response along gradients of CO₂ concentrations. In six annual plant species, mean enhancement at double ambient vs. ambient CO₂ was 1.13 for total biomass and 1.30 for reproductive biomass. The two measures were significantly correlated, but there was considerable scatter in the relationship, indicating that reproductive responses cannot be consistently predicted from enhancement of total biomass. Along experimental CO₂ gradients utilizing three concentrations, there was a great diversity of response patterns, including positive, negative, non-monotonic and non-significant (flat) responses. The distribution of response patterns differed for plants grown in stands compared to those grown individually. Positive responses were less frequent in competitive environments, and non-monotonic responses were more frequent. These results emphasize that interpolation of plant response based on enhancement ratios measured at elevated vs. ambient CO₂ concentrations is not sufficient to predict community responses to incremental changes in atmospheric conditions. The consequences of differential response patterns were assessed in a simulation of community dynamics for four species of annual plants. The model illustrates that the final community composition at a future point in time depends critically on both the magnitude and the rate of increase of atmospheric CO₂.     

SSCP-SNP-based conserved ortholog set (COS) markers for comparative genomics in cassava (Manihot esculenta crantz)

The ever increasing body of information on genomics and functional genomics from model plants, and new tools of comparative genomics, provide an opportunity to accelerate the development of molecular markers for increasing the efficiency of breeding of lesser studied crops, so-called “orphan crops.” Conserved ortholog set (COS) markers represent orthologous genes in widely divergent plant species, and are currently the principal tool of choice for comparative genomics. EST sequences of 3 drought tolerance related genes—chalcone synthase (CHS), dihydroflavonol-4-reductase (DHRF) and drought responsive element binding factor 1 (DREB-1) fromMusa sp—were used to identify cassava EST homologs that were then scanned against the Arabidopsis genome database to identify them as COS markers. The CHS and DHRF ESTs were demonstrated to be COS markers, while the DREB EST was shown to belong to a gene family. The three genes were evaluated as single strand conformation polymorphism—single nucleotide polymorphism (SSCP-SNP) markers in the parents of an F1 mapping population and subsequently in the progenies. The DHRF COS marker mapped to linkage group R of the female-derived map while the DREB-1 EST mapped at an end of the male-derived linkage group K. The CHS COS marker could not be mapped because it was not polymorphic in the parents of the mapping population. These new marker tools should accelerate the development of markers associated with genes controlling traits of agronomic interest via the candidate gene loci (CGL) QTL-mapping approach.     

Multiple shoot induction and leaf and flower bud abscission of Annonacultures as affected by types of ventilation

Nodal explants of Annona squamosa L. and Annona muricata L. were cultured in vitro under various types of ventilation: airtight vessel (sealed condition; number of air exchange 0.1 h^–1), natural ventilation (via a polypropylene membrane; number of air exchange 1.5 h^–1), and forced ventilation (5.0 cm³ min^–1 in a 60 cm³ vessel; number of air exchange 5.0 h^–1). In both species, numbers of leaves, leaf areas and numbers of nodes per shoot increased with improving standards of ventilation, while leaf abscissions were substantially reduced; all the leaves had abscised in the airtight vessels after 12–15 days, but none had done so with forced ventilation. Flower-bud abscission in A. muricatashowed a similar trend after 21 days. These effects were associated with reductions in the accumulation of ethylene within the culture vessels, produced by increasing the efficiency of ventilation; ethylene was not detected in those fitted with a forced ventilation system. CO₂ concentrations in culture headspaces and the net photosynthetic rates of the plantlets were also evaluated. CO₂ concentrations decreased well below the ambient in the natural and airtight vessels; however, under forced ventilation, CO₂ concentrations were significantly higher during the photoperiod, compared to those of the natural ventilation and airtight vessel treatments. In general, net photosynthetic rates per unit leaf area increased with increasing photosynthetic photon flux (PPF) and rates were highest in plantlets grown under forced ventilation, intermediate under natural ventilation and lowest in the airtight vessels.Eighteen different media were investigated for their effects on multiple shoot induction in both species. The best medium for multiple shoot induction and growth in A. squamosa was Murashige and Skoog medium (MS) + 6-benzylaminopurine (BA; 1.5 mg l^–1) + casein hydrolysate (1.0 g l^–1) and for A. muricata MS + BA (1.0 mg l^–1) + naphthaleneacetic acid (NAA; 0.1 mg l^–1).     

Stomatal responses to carbon dioxide of isolated epidermis from a C3 plant,the Argenteum mutant of Pisum sativum L., and a crassulacean-acid-metabolism plant Kalanchoë daigremontiana Hamet et Perr

The response of stomata in isolated epidermis to the concentration of CO₂ in the gaseous phase was examined in a C₃ species, the Argenteum mutant of Pisum sativum, and a crassulacean-acid-metabolism (CAM) species, Kalanchoë daigremontiana. Epidermis from leaves of both species was incubated on buffer solutions in the presence of air containing various volume fractions of CO₂ (0 to 10000·10^–6). In both species and in the light and in darkness, the effect of CO₂ was to inhibit stomatal opening, the maximum inhibition of opening occurring in the range 0 to 360·10^–6. The inhibition of opening per unit change in concentration was greatest between volume fractions of 0 and 240·10^–6. There was little further closure above the volume fraction of 360·10^–6, i.e. approximately ambient concentration of CO₂. Thus, although leaves of CAM species may experience much higher internal concentrations of CO₂ in the light than those of C₃ plants, this does not affect the sensitivity of their stomata to CO₂ concentration or the range over which they respond. Stomatal responses to CO₂ were similar in both the light and the dark, indicating that effects of CO₂ on stomata occur via mechanisms which are independent of light. The responses of stomata to CO₂ in the gaseous phase took place without the treatments changing the pH of the buffered solutions. Thus it is unlikely that CO₂ elicited stomatal movement by changing either the pH or the HCO ₃ ^– /CO ₃ ^2- equilibria. It is suggested that the concentration of dissolved unhydrated CO₂ may be the effector of stomatal movement and that its activity is related to its reactivity with amines.     

Effects of Carbonic Anhydrase Inhibitors on Proton Exchange and Photosynthesis in Pea Protoplasts

At concentrations of 100–200 M, ethoxyzolamide, a lipophilic inhibitor of carbonic anhydrase, considerably (by 60%) inhibited light-induced CO₂-dependent oxygen evolution in pea protoplasts at the optimum concentration of inorganic carbon (100 M CO₂) in the medium. At the same concentrations of the inhibitor, electron transport in isolated pea thylakoids was inhibited only by 6–9%. Acetazolamide, a water-soluble inhibitor of carbonic anhydrase, affected neither the rate of CO₂-dependent O₂evolution in protoplasts nor electron transport in thylakoid membranes. A light-dependent proton uptake by protoplasts was demonstrated. At pH 7.2, the induction kinetics and the rate of proton uptake were similar to those for CO₂-dependent O₂evolution. The rate of proton uptake was decreased twofold by 1 mM acetazolamide. This fact agrees with the notion that a membrane-bound carbonic anhydrase is operative in the plasma membrane of higher plant cells. A mechanism of its functioning is suggested. Possible functions of carbonic anhydrases in the cells of C₃-plants are discussed.     

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%).     

Chromatin-associated ornithine decarboxylase at early stages of growth and differentiation of etiolated mono- and dicotyledonous plants

Ornithine decarboxylase (ODC) of barley, corn, bean and pea plants was associated with chromatin at early stages of growth. In corn, bean and pea plants the chromatin of roots possessed ODC with the highest specific activity of 30–65 units per mg protein. After 200 h of growth ODC activity of chromatin declined, while ODC activity of cytosol increased linearly to 4–19 units per mg protein. ODC activity of either chromatin or cytosol of shoots in the above mentioned plants was usually low, 1–2 units per mg protein and only at the beginning of shoot growth (96 h) did pea chromatin have high ODC activity (14 units per mg protein). In these plants ODC was tightly bound to chromatin and could not be extracted with different concentrations of NaCl (0.1–1.0 M) or non-ionic detergent (Triton X100, Tween 20) but could be extracted by freezing and thawing with satisfactory recovery.     

Phosphorus application and elevated CO2 enhance drought tolerance in field pea grown in a phosphorus-deficient vertisol

Background and Aims Benefits to crop productivity arising from increasing CO₂ fertilization may be offset by detrimental effects of global climate change, such as an increasing frequency of drought. Phosphorus (P) nutrition plays an important role in crop responses to water stress, but how elevated CO₂ (eCO₂) and P nutrition interact, especially in legumes, is unclear. This study aimed to elucidate whether P supply improves plant drought tolerance under eCO₂.Methods A soil-column experiment was conducted in a free air CO₂ enrichment (SoilFACE) system. Field pea (Pisum sativum) was grown in a P-deficient vertisol, supplied with 15 mg P kg⁻¹ (deficient) or 60 mg P kg⁻¹ (adequate for crop growth) and exposed to ambient CO₂ (aCO₂; 380–400 ppm) or eCO₂ (550–580 ppm). Drought treatments commenced at flowering. Measurements were taken of soil and leaf water content, photosynthesis, stomatal conductance, total soluble sugars and inorganic P content (Pi).Key Results Water-use efficiency was greatest under eCO₂ when the plants were supplied with adequate P compared with other treatments irrespective of drought treatment. Elevated CO₂ decreased stomatal conductance and transpiration rate, and increased the concentration of soluble sugars and relative water contents in leaves. Adequate P supply increased concentrations of soluble sugars and Pi in drought-stressed plants. Adequate P supply but not eCO₂ increased root length distribution in deeper soil layers.Conclusions Phosphorus application and eCO₂ interactively enhanced periodic drought tolerance in field pea as a result of decreased stomatal conductance, deeper rooting and high Pi availability for carbon assimilation in leaves.     

Contrasting growth response of an N2-fixing and non-fixing forb to elevated CO2: dependence on soil N supply

With the ability to symbiotically fix atmospheric N₂, legumes may lack the N-limitations thought to constrain plant response to elevated concentrations of atmospheric CO₂. The growth and photosynthetic responses of two perennial grassland species were compared to test the hypotheses that (1) the CO₂ response of wild species is limited at low N availability, (2) legumes respond to a greater extent than non-fixing forbs to elevated CO₂, and (3) elevated CO₂ stimulates symbiotic N₂ fixation, resulting in an increased amount of N derived from the atmosphere. This study investigated the effects of atmospheric CO₂ concentration (365 and 700 mol mol^–1) and N addition on whole plant growth and C and N acquisition in an N₂-fixing legume (Lupinus perennis) and a non-fixing forb (Achillea millefolium) in controlled-chamber environments. To evaluate the effects of a wide range of N availability on the CO₂ response, we incorporated six levels of soil N addition starting with native field soil inherently low in N (field soil + 0, 4, 8, 12, 16, or 20 g N m^–2 yr^–1). Whole plant growth, leaf net photosynthetic rates (A), and the proportion of N derived from N₂ fixation were determined in plants grown from seed over one growing season. Both species increased growth with CO₂enrichment, but this response was mediated by N supply only for the non-fixer, Achillea. Its response depended on mineral N supply as growth enhancements under elevated CO₂ increased from 0% in low N soil to +25% at the higher levels of N addition. In contrast, Lupinus plants had 80% greater biomass under elevated CO₂ regardless of N treatment. Although partial photosynthetic acclimation to CO₂ enrichment occurred, both species maintained comparably higher A in elevated compared to ambient CO₂ (+38%). N addition facilitated increased A in Achillea, however, in neither species did additional N availability affect the acclimation response of A to CO₂. Elevated CO₂ increased plant total N yield by 57% in Lupinus but had no effect on Achillea. The increased N in Lupinus came from symbiotic N₂ fixation, which resulted in a 47% greater proportion of N derived from fixation relative to other sources of N. These results suggest that compared to non-fixing forbs, N₂-fixers exhibit positive photosynthetic and growth responses to increased atmospheric CO₂ that are independent of soil N supply. The enhanced amount of N derived from N₂ fixation under elevated CO₂ presumably helps meet the increased N demand in N₂-fixing species. This response may lead to modified roles of N₂-fixers and N₂-fixer/non-fixer species interactions in grassland communities, especially those that are inherently N-poor, under projected rising atmospheric CO₂.