Contrasting crop species responses to CO2 and temperature: rice,soybean and citrus

The continuing increase in atmospheric carbon dioxide concentration ([CO₂]) and projections of possible future increases in global air temperatures have stimulated interest in the effects of these climate variables on plants and, in particular, on agriculturally important food crops. Mounting evidence from many different experiments suggests that the magnitude and even direction of crop responses to [CO₂] and temperature is almost certain to be species dependent and very likely, within a species, to be cultivar dependent. Over the last decade, [CO₂] and temperature experiments have been conducted on several crop species in the outdoor, naturally-sunlit, environmentally controlled, plant growth chambers by USDA-ARS and the University of Florida, at Gainesville, Florida, USA. The objectives for this paper are to summarize some of the major findings of these experiments and further to compare and contrast species responses to [CO₂] and temperature for three diverse crop species: rice (Oryza sativa, L.), soybean (Glycine max, L.) and citrus (various species). Citrus had the lowest growth and photosynthetic rates but under [CO₂] enrichment displayed the greatest percentage increases over ambient [CO₂] control treatments. In all three species the direct effect of [CO₂] enrichment was always an increase in photosynthetic rate. In soybean, photosynthetic rate depended on current [CO₂] regardless of the long-term [CO₂] history of the crop. In rice, photosynthetic rate measured at a common [CO₂], decreased with increasing long-term [CO₂] growth treatment due to a corresponding decline in RuBP carboxylase content and activity. Rice specific respiration decreased from subambient to ambient and superambient [CO₂] due to a decrease in plant tissue nitrogen content and a decline in specific maintenance respiration rate. In all three species, crop water use decreased with [CO₂] enrichment but increased with increases in temperature. For both rice and soybean, [CO₂] enrichment increased growth and grain yield. Rice grain yields declined by roughly 10 % per each 1 °C rise in day/night temperature above 28/21 °C.     

The DNA,RNA and protein composition of the cyanobacterium Anacystis nidulans grown in light- and carbon dioxide-limited chemostats

The DNA, RNA and protein content of the cyanobacterium Anacystis nidulans was determined in light-limited and carbon dioxide-limited chemostat cultures over the dilution rate range, D=0.02 h^-1 to 0.19 h^-1. The macromolecular contents as a percentage of the dry weight and on a per cell basis varied significantly as a function of organism growth rate and the nature of the growth conditions. For both limitations the RNA content per cell increased [20–55 fg RNA (cell)^-1] with increasing dilution rate and also showed an increase as a percentage of the dry weight. The DNA content as a percentage of the dry weight showed a 2-fold decrease with increasing dilution rate over the range examined. On a per cell basis DNA reached a peak at D=0.1 h^-1 [4.5 fg DNA (cell)^-1] for light-limited organisms and at D=0.08 h^-1 [8.0 fg DNA (cell)^-1] for carbon dioxide-limited organisms. The q _RNA increased with increasing dilution rates over the complete growth rate range examined whilst q _DNA reached a maximum at D=0.09 to 0.10 h^-1. The protein content as a percentage of the dry weight was greater in CO₂-limited organisms than light-limited organisms but in both cultures declined as the dilution rate was increased above D=0.10 h^-1.     

Responses of macroalgae to CO2 enrichment cannot be inferred solely from their inorganic carbon uptake strategy

Increased plant biomass is observed in terrestrial systems due to rising levels of atmospheric CO₂, but responses of marine macroalgae to CO₂ enrichment are unclear. The 200% increase in CO₂ by 2100 is predicted to enhance the productivity of fleshy macroalgae that acquire inorganic carbon solely as CO₂ (non‐carbon dioxide‐concentrating mechanism [CCM] species—i.e., species without a carbon dioxide‐concentrating mechanism), whereas those that additionally uptake bicarbonate (CCM species) are predicted to respond neutrally or positively depending on their affinity for bicarbonate. Previous studies, however, show that fleshy macroalgae exhibit a broad variety of responses to CO₂ enrichment and the underlying mechanisms are largely unknown. This physiological study compared the responses of a CCM species (Lomentaria australis) with a non‐CCM species (Craspedocarpus ramentaceus) to CO₂ enrichment with regards to growth, net photosynthesis, and biochemistry. Contrary to expectations, there was no enrichment effect for the non‐CCM species, whereas the CCM species had a twofold greater growth rate, likely driven by a downregulation of the energetically costly CCM(s). This saved energy was invested into new growth rather than storage lipids and fatty acids. In addition, we conducted a comprehensive literature synthesis to examine the extent to which the growth and photosynthetic responses of fleshy macroalgae to elevated CO₂ are related to their carbon acquisition strategies. Findings highlight that the responses of macroalgae to CO₂ enrichment cannot be inferred solely from their carbon uptake strategy, and targeted physiological experiments on a wider range of species are needed to better predict responses of macroalgae to future oceanic change.     

The Physiology of Astrephomene gubernaculifera1

SYNOPSIS. A comparative physiologic study of 4- and 7-chromosome strains of Astrephomene gubernaculifera was done. The vitamins p-aminobenzoic acid, nicotinamide, biotin, thiamin HCl and vitamin B₁₂ were tested for their ability to support growth. Only vitamin B₁₂ was required for active growth altho the presence of thiamin HCl was necessary to produce structurally typical colonies. Astrephomene will not grow in the absence of an exogenous source of carbon. Of the 36 carbon sources tested, only pyruvate, butyrate, succinate and acetate permitted active growth. Sodium acetate was the best. Strains grew within the initial pH range of 5.0–7.5. At pH values above 8.0 growth declined rapidly. When media were buffered with TES (N-tris [hydroxymethyl]methyl-2-aminoethane-sulfonic acid) at an initial pH of 6.8 growth was enhanced. The organism grew at 15–40 C. Growth in the dark was slightly less than that in the light.     

Screening of Carbon Dioxide-Requiring Extreme Oligotrophs from Soil

We screened soil samples for CO₂-requiring extreme oligotrophs similar to Rhodococcus erythropolis N9T-4, which can grow on a basal salt agar medium without an organic carbon source. From 387 soil samples, three isolates were obtained and identified as Streptomyces spp. by 16S rDNA analysis. The isolates required gaseous CO₂ for growth and grew on a basal salt medium solidified by silica gel. These results suggest that such CO₂-requiring oligotrophs occur widely in nature.     

Effects of CO2 enrichment,nutrient addition,and fungal endophyte-infection on the growth of two grasses

Summary Increasing atmospheric carbon dioxide (CO₂) concentration is expected to increase plant productivity and alter plant/plant interactions, but little is known about its effects on symbiotic interactions with microorganisms. Interactions between perennial ryegrass, Lolium perenne (a C3 plant), and purpletop grass, Tridens flavus (a C4 plant), and their clavicipitaceous fungal endophytes (Acremonium lolii and Balansia epichloe, respectively) were investigated by growing the grasses under 350 and 650 l l^– 1 CO₂ at two nutrient levels. Infected and uninfected perennial ryegrass responded with increased growth to both CO₂ enrichment and nutrient addition. Biomass and leaf area of infected and uninfected plants responded similarly to CO₂ enrichment. When growth analysis parameters were calculated, there were significant increases in relative growth rate and net assimilation rate of infected plants compared to uninfected plants, although the differences remained constant across CO₂ and nutrient treatments. Growth of purpletop grass did not increase with CO₂ enrichment or nutrient addition and there were no significant differences between infected and uninfected plants. CO₂ enrichment did not alter the interactions between these two host grasses and their endophytic-fungal symbionts.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Carbon source–sink limitations differ between two species with contrasting growth strategies

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximize crop yield. We investigated how differences in growth rate arise from source–sink limitations, using a model system comparing a fast‐growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow‐growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub‐ambient and elevated CO₂ concentrations ([CO₂]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO₂] indicating that source strength was near maximal at current [CO₂]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO₂] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO₂], and lower non‐structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximizing growth of elite cereals under future elevated [CO₂].     

Carbon assimilation by the autotrophic thermophilic archaebacterium Thermoproteus neutrophilus

Growth of Thermoproteus neutrophilus at 85°C was studied using an improved mineral medium with CO₂, CO₂ plus acetate, CO₂ plus propionate, or CO₂ plus succinate as carbon sources; sulfur reduction with H₂ to H₂S was the sole source of energy. None of the carbon compounds added was oxidized to CO₂. The organism grew autotrophically with a generation time of 9–14 h, up to a cell density of 0.5 g dry weight per liter (2×10⁹ cells/ml). Propionate did not stimulate, succinate slightly stimulated the growth rate. Acetate, even at low concentrations (0.5 mM), stimulated the growth rate, the generation time being shortened to 3–4 h. Acetate provided 70% of the cell carbon, which shows that Thermoproteus neutrophilus is a facultative autotroph. The path of these carbon precursors into cell compounds was studied by ¹⁴C long-term labelling and investigation of enzyme activities. Propionate could not be used as a major carbon source and was incorporated only into isoleucine, probably via the citramalate pathway. Acetate was a preferred carbon source which suppressed autotrophic CO₂ fixation: acetate grown cells exhibited an incomplete citric acid cycle in which 2-oxoglutarate dehydrogenase was present, but fumarate reductase was repressed. The succinate incorporation pattern and enzyme pattern indicated that autotrophic CO₂ fixation proceeded via a yet to be defined reductive citric acid cycle.     

Woody biomass production during the second rotation of a bio-energy Populus plantation increases in a future high CO2 world

The quickly rising atmospheric carbon dioxide (CO₂)‐levels, justify the need to explore all carbon (C) sequestration possibilities that might mitigate the current CO₂ increase. Here, we report the likely impact of future increases in atmospheric CO₂ on woody biomass production of three poplar species (Populus alba L. clone 2AS‐11, Populus nigra L. clone Jean Pourtet and Populus×euramericana clone I‐214). Trees were growing in a high‐density coppice plantation during the second rotation (i.e., regrowth after coppice; 2002–2004; POPFACE/EUROFACE). Six plots were studied, half of which were continuously fumigated with CO₂ (FACE; free air carbon dioxide enrichment of 550 ppm). Half of each plot was fertilized to study the interaction between CO₂ and nutrient fertilization. At the end of the second rotation, selective above‐ and belowground harvests were performed to estimate the productivity of this bio‐energy plantation. Fertilization did not affect growth of the poplar trees, which was likely because of the high rates of fertilization during the previous agricultural land use. In contrast, elevated CO₂ enhanced biomass production by up to 29%, and this stimulation did not differ between above‐ and belowground parts. The increased initial stump size resulting from elevated CO₂ during the first rotation (1999–2001) could not solely explain the observed final biomass increase. The larger leaf area index after canopy closure and the absence of any major photosynthetic acclimation after 6 years of fumigation caused the sustained CO₂‐induced biomass increase after coppice. These results suggest that, under future CO₂ concentrations, managed poplar coppice systems may exhibit higher potential for C sequestration and, thus, help mitigate climate change when used as a source of C‐neutral energy.     

Nitrogen accumulation and distribution in Danthonia richardsonii swards in response to CO2 and nitrogen supply over four years of growth

Nitrogen‐stressed microcosms of the C3 grass Danthonia richardsonii gained nitrogen from the environment when grown under ambient or enriched (359, ‘amb’ or 719 μL L^{? 1}‘enr’, respectively) atmospheric CO₂ concentrations over a 4‐y period. This gain was apparent at all rates of supplied mineral N (2.2, 6.7 or 19.8 g N m^{? 2} y^{? 1}– low‐N, mid‐N or high‐N), although it was small at high‐N. Small losses of N occurred from the microcosm as leachate, while gaseous losses of N were estimated to be between 10% and 25% of applied mineral N. Losses of applied mineral N were slightly lower under CO₂ enrichment only at the highest rate of mineral N supply. Levels of ¹⁵N natural abundance in green leaf (δ¹⁵Ν) of ? 2‰ (amb low‐N) and of below ? 4‰ (enr low‐ & mid‐N) suggest that absorption of atmospheric NH₃ may have been a source of some of the extra N in the low and mid‐N treatments. Biological N₂ fixation, of up to 2 g m^{? 2} y^{? 1} was hypothesized to form the remainder of the environmental N source. Microcosm C:N ratio was higher under CO₂ enrichment. Nitrogen productivity of microcosm carbon gain (g C accumulated g^{? 1} leaf N day^{? 1}) was increased (up to 100%) by CO₂ enrichment at all rates of mineral N supply. Green leaf %N was reduced by CO₂ enrichment, and there was less nitrogen in the green leaf pool under CO₂ enrichment. Less, or the same amount of nitrogen was present in senesced leaf, surface litter and root under CO₂ enrichment while more nitrogen was present in the soil in organic forms, and as NH₄ + at the highest rate of mineral N supply.     

Elevated atmospheric CO2 increases microbial growth rates in soil: results of three CO2 enrichment experiments

Increasing the belowground translocation of assimilated carbon by plants grown under elevated CO₂ can cause a shift in the structure and activity of the microbial community responsible for the turnover of organic matter in soil. We investigated the long‐term effect of elevated CO₂ in the atmosphere on microbial biomass and specific growth rates in root‐free and rhizosphere soil. The experiments were conducted under two free air carbon dioxide enrichment (FACE) systems: in Hohenheim and Braunschweig, as well as in the intensively managed forest mesocosm of the Biosphere 2 Laboratory (B2L) in Oracle, AZ. Specific microbial growth rates (μ) were determined using the substrate‐induced respiration response after glucose and/or yeast extract addition to the soil. For B2L and both FACE systems, up to 58% higher μ were observed under elevated vs. ambient CO₂, depending on site, plant species and N fertilization. The μ‐values increased linearly with atmospheric CO₂ concentration at all three sites. The effect of elevated CO₂ on rhizosphere microorganisms was plant dependent and increased for: Brassica napus=Triticum aestivum<Beta vulgaris<Populus deltoides. N deficiency affected microbial growth rates directly (N limitation) and indirectly (changing the quantity of fine roots). So, 50% decrease in N fertilization caused the overall increase or decrease of microbial growth rates depending on plant species. The μ‐value increase was lower for microorganisms growing on yeast extract then for those growing on glucose, i.e. the effect of elevated CO₂ was smoothed on rich vs. simple substrate. So, the r/K strategies ratio can be better revealed by studying growth on simple (glucose) than on rich substrate mixtures (yeast extract). Our results clearly showed that the functional characteristics of the soil microbial community (i.e. specific growth rates) rather than total microbial biomass amount are sensitive to increased atmospheric CO₂. We conclude that the more abundant available organics released by roots at elevated CO₂ altered the ecological strategy of the soil microbial community specifically a shift to a higher contribution of fast‐growing r‐selected species was observed. These changes in functional structure of the soil microbial community may counterbalance higher C input into the soil under elevated atmospheric CO₂ concentration.     

Differential anatomical responses to elevated CO2 in saplings of four hardwood species

To determine whether an elevated carbon dioxide concentration ([CO₂]) can induce changes in the wood structure and stem radial growth in forest trees, we investigated the anatomical features of conduit cells and cambial activity in 4‐year‐old saplings of four deciduous broadleaved tree species – two ring‐porous (Quercus mongolica and Kalopanax septemlobus) and two diffuse‐porous species (Betula maximowicziana and Acer mono) – grown for three growing seasons in a free‐air CO₂ enrichment system. Elevated [CO₂] had no effects on vessels, growth and physiological traits of Q. mongolica, whereas tree height, photosynthesis and vessel area tended to increase in K. septemlobus. No effects of [CO₂] on growth, physiological traits and vessels were seen in the two diffuse‐porous woods. Elevated [CO₂] increased larger vessels in all species, except B. maximowicziana and number of cambial cells in two ring‐porous species. Our results showed that the vessel anatomy and radial stem growth of Q. mongolica, B. maximowicziana and A. mono were not affected by elevated [CO₂], although vessel size frequency and cambial activity in Q. mongolica were altered. In contrast, changes in vessel anatomy and cambial activity were induced by elevated [CO₂] in K. septemlobus. The different responses to elevated [CO₂] suggest that the sensitivity of forest trees to CO₂ is species dependent.     

Influence of enhanced CO2 on growth and photosynthesis of the red algaeGracilaria sp. andG. chilensis

The influence of elevated CO₂ concentrations on growth and photosynthesis ofGracilaria sp. andG. chilensis was investigated in order to procure information on the effective utilization of CO₂. Growth of both was enhanced by CO₂ enrichment (air + 650 ppm CO₂, air + 1250 ppm CO₂, the enhancement being greater inGracilaria sp. Both species increased uptake of NO₃ ^– with CO₂ enrichment. Photosynthetic inorganic carbon uptake was depressed inG. chilensis by pre-culture (15 days) with CO₂ enrichment, but little affected inGracilaria sp. Mass spectrometric analysis showed that O₂ uptake was higher in the light than in the dark for both species and in both cases was higher inGracilaria sp. The higher growth enhancement inGracilaria sp. was attributed to greater depression of photorespiration by the enrichment of CO₂ in culture.     

Sexual reproduction of Lolium perenne L. and Trifolium repens L. under free air CO2 enrichment (FACE) at two levels of nitrogen application

Field experiments in managed grassland have shown that the response of vegetative growth to elevated CO₂ is nitrogen‐dependent in grasses, but independent in N₂‐fixing legumes. In the present study, we tested whether this is also true for reproduction. We evaluated reproductive growth, flowering phenology, seed development, reproductive success and seed germination in the grass Lolium perenne L. and the legume Trifolium repens L., growing in monocultures in a free air carbon dioxide enrichment (FACE) system at ambient (35 Pa) and elevated (60 Pa) partial pressure of CO₂ and two levels of nitrogen fertilization (14 and 56 g N m^?2 a^?1). In both species, elevated CO₂ had no significant effect on sexual reproduction. In L. perenne, reproduction was mainly nitrogen‐dependent. The weak interactions between CO₂ and mineral N supply (13% more flowers and 8% more grains per spike at high N, 7% less flowers and 8% less grains at low N) were not significant. Under elevated CO₂, grain maturation was slightly enhanced and grain weight tended to decrease. No influence could be ascertained in the date of anthesis, the temporal pattern of grain growth, the rate of grain abortion and germination. Trifolium repens, grown under CO₂ enrichment at both levels of N fertilization, flowered 10 d earlier, tended to form more inflorescences per ground area and more flowers (8–12%) and seeds (>18%) per inflorescence than at ambient CO₂. The temporal pattern of seed growth was about the same in all treatments; embryo development, however, was accelerated in fumigated plants. The number of aborted seeds per pod, seed size, thousand‐seed weight and germinability did not show any influence of CO₂. Fumigated plants at high N were attacked slightly more frequently by seed‐eating weevils, which lowered the seed output per pod. In summary, the reproductive response of L. perenne and T. repens to CO₂ enrichment on the flower and inflorescence level was far weaker than expected from the results on vegetative growth.     

Nitrogenase activity and N2 fixation are stimulated by elevated CO2 in a tropical N2-fixing tree

 Seeds of Gliricidia sepium, a fast-growing woody legume native to seasonal tropical forests of Central America, were inoculated with N₂-fixing Rhizobium bacteria and grown in environmentally controlled glasshouses for 67–71 days under ambient CO₂ (35 Pa) and elevated CO₂ (70 Pa) conditions. Seedlings were watered with an N-free, but otherwise complete, nutrient solution such that bacterial N₂ fixation was the only source of N available to the plant. The primary objective of our study was to quantify the effect of CO₂ enrichment on the kinetics of photosynthate transport to nodules and determine its subsequent effect on N₂ fixation. Photosynthetic rates and carbon storage in leaves were higher in elevated CO₂ plants indicating that more carbon was available for transport to nodules. A ¹⁴CO₂ pulse-chase experiment demonstrated that photosynthetically fixed carbon was supplied by leaves to nodules at a faster rate when plants were grown in elevated CO₂. Greater rates of carbon supply to nodules did not affect nodule mass per plant, but did increase specific nitrogenase activity (SNA) and total nitrogenase activity (TNA) resulting in greater N₂ fixation. In fact, a 23% increase in the rate of carbon supplied to nodules coincided with a 23% increase in SNA for plants grown in elevated CO₂, suggesting a direct correlation between carbon supply and nitrogenase activity. The improvement in plant N status produced much larger plants when grown in elevated CO₂. These results suggest that Gliricidia, and possibly other N₂-fixing trees, may show an early and positive growth response to elevated CO₂, even in severely N-deficient soils, due to increased nitrogenase activity. Received: 27 February 1996 / Accepted: 19 June 1996     

Effects of CO2 enrichment on whole-plant carbon budget of seedlings of Fagus grandifolia and Acer saccharum in low irradiance

Carbon exchange rates (CER) and whole-plant carbon balances of beech (Fagus grandifolia) and sugar maple (Acer saccharum) were compared for seedlings grown under low irradiance to determine the effects of atmospheric CO₂ enrichment on shade-tolerant seedlings of co-dominant species. Under contemporary atmospheric CO₂, photosynthetic rate per unit mass of beech was lower than for sugar maple, and atmospheric CO₂ enrich ment enhanced photosynthesis for beech only. Aboveground respiration per unit mass decreased with CO₂ enrichment for both species while root respiration per unitmass decreased for sugar maple only. Under contemporary atmoapheric CO₂, beech had lower C uptake per plant than sugar maple, while C losses per plant to nocturnal aboveground and root respiration were similar for both species. Under elevated CO₂, C uptake per plant was similar for both species, indicating a significant relative increase in whole-seedling CER with CO₂ enrich ment for beech but not for sugar maple. Total C loss per plant to aboveground respiration was decreased for beech only because increase in sugar maple leaf mass counterbalanced a reduction in respiration rates. Carbon loss to root respiration per plant was not changed by CO₂ enrichment for either species. However, changes in maintenance respiration cost and nitrogen level suggest changes in tissue composition with elevated CO₂. Beech had a greater net daily C gain with CO₂ enrichment than did sugar maple in contrast to a lower one under contemporary CO₂. Elevated CO₂ preferentially enhances the net C balance of beech by increasing photosynthesis and reducing respiration cost. In all cases, the greatest C lost was by roots, indicating the importance of belowground biomass in net C gain. Relative growth rate estimated from biomass accumulation was not affected by CO₂ enrichment for either species possibly because of slow growth under low light. This study indicates the importance of direct effects of CO₂ enrichment when predicting potential change in species distribution with global climate change.     

Global terrestrial carbon storage and uncertainties in its temperature sensitivity examined with a simple model

The future of the land carbon sink is a significant uncertainty in global change projections. Here, key controls on global terrestrial carbon storage are examined using a simple model of vegetation and soil. Equilibrium solutions are derived as a function of atmospheric CO₂ and global temperature, these environmental variables are then linked in an idealized global change trajectory, and the lag between the dynamic and equilibrium solutions is derived for different linear rates of increase in atmospheric CO₂. Terrestrial carbon storage is departing significantly from equilibrium because CO₂ and temperature are increasing on a similar timescale to ecosystem change, and the lag is found to be proportional to the rate of forcing. Thus peak sizes of the land carbon sink, and any future land carbon source, are proportional to the rate of increase of CO₂. A switch from a land carbon sink to a source occurs at a higher CO₂ and temperature under more rapid forcing. The effects of parameter uncertainty in temperature sensitivities of photosynthesis, plant respiration and soil respiration, and structural uncertainty through the effect of fixing the ratio of plant respiration to photosynthesis are explored. In each case, the CO₂ fertilization effect on photosynthesis is constrained to reproduce the 1990 atmospheric CO₂ concentration within a closed global model. New literature compilations are presented for the temperature sensitivities of plant and soil respiration. A lower limit, Q₁₀=1.29, for soil respiration significantly increases future land carbon storage. An upper limit, Q₁₀=3.63, for soil respiration underpredicts the increase in carbon storage since the Last Glacial Maximum. Fixing the ratio of plant respiration to photosynthesis (R/P) at 0.5 generates the largest and most persistent land carbon sink, followed by the weakest land carbon source.     

Carbon accumulation, distribution and water use of Danthonia richardsonii swards in response to CO2 and nitrogen supply over four years of growth

Microcosms of Danthonia richardsonii (Cashmore) accumulated more carbon when grown under CO₂ enrichment (719 μL L^–1 cf. 359 μL L^–1) over a four-year period, even when nitrogen availability severely restricted productivity (enhancement ratios for total microcosm C accumulation of 1.21, 1.14 and 1.29 for mineral N supplies of 2.2, 6.7 and 19.8 g N m^–2 y^–1, respectively). The effect of CO₂ enrichment on total system carbon content did not diminish with time. Increased carbon accumulation occurred despite the development over time of a lower leaf area index and less carbon in the green leaf fraction at high CO₂. The extra carbon accumulated at high CO₂ in the soil, senesced leaf and leaf litter fractions at all N levels, and in root at high-N, while at low-and mid-N less carbon accumulated in the root fraction at high CO₂. The rate of leaf turnover was increased under CO₂ enrichment, as indicated by increases in the carbon mass ratio of senesced to green leaf lamina. Microcosm evapotranspiration rates were lower at high CO₂ when water was in abundant supply, resulting in higher average soil water contents. The higher soil water contents at high CO₂ have important implications for microcosm function, and may have contributed significantly to the increased carbon accumulation at high CO₂. These results indicate that CO₂ enrichment can increase carbon accumulation by a simple soil–plant system, and that any increase in whole system carbon accumulation may not be evident from snapshot measurements of live plant carbon.     

Interrelationship of nitrogen fixation,hydrogen evolution and photoreduction in Rhodospirillum rubrum

Summary Rhodospirillum rubrum was grown: 1. photoheterotrophically on a medium containing dl-malate as the carbon source and ammonium chloride as the nitrogen source (medium No. 1); 2. phototrophically with N₂ and dl-malate (medium No. 2); 3. photoautotrophically with N₂, CO₂ and H₂ (medium No. 3).Resting cells derived from these cultures were tested for their ability to photoreduce CO₂, evolve H₂ and fix N₂. Only cells which were grown in medium No. 2 were able to perform all three gas exchanges. The activity pattern of gas exchanges altered in a characteristic way during the growth cycle of the bacterial culture. Cells newly transferred to medium No. 2 showed an enormous increase in the rate of H₂ evolution, which dropped sharply when all l-malate had been used up. The rate of photoreduction of CO₂ increased steadily and reached a maximum level after 120 h. The nitrogen fixing activity remained constant during the whole growth cycle.The yields of H₂ produced per mole of l-malate added were measured as a function of cell age. Only very young cultures gave appreciable yields, which dropped gradually with increasing age.The function of the carbon source is discussed as a regulating factor for photoreduction and hydrogen evolution.