The influence of plant carbon dioxide and nutrient supply on susceptibility to insect herbivores

The carbon/nutrient ratio of plants has been hypothesized to be a significant regulator of plant susceptibility of leaf-eating insects. As rising atmospheric carbon dioxide stimulates photosynthesis, host plant carbon supply is increased and the accompanying higher levels of carbohydrates, especially starch, apparently dilute the protein content of the leaf. When host plant nitrogen supply is limited, plant responses include increased carbohydrate accumulation, reduced leaf protein content, but also increased carbon-based defensive chemicals. No change, however, has been observed in the concentration of leaf defensive allelochemicals with elevated carbon dioxide during host plant growth. Insect responses to carbon-fertilized leaves include increased consumption with little change in growth, or alternatively, little change in consumption with decreased growth, as well as enhanced leaf digestibility, reduced nitrogen use efficiency, and reduced fecundity. The effects of plant carbon and nutrient supply on herbivores appear to result, at least in part, from independent processes affecting secondary metabolism.     

Elevated atmospheric concentrations of carbon dioxide reduce monarch tolerance and increase parasite virulence by altering the medicinal properties of milkweeds

Hosts combat their parasites using mechanisms of resistance and tolerance, which together determine parasite virulence. Environmental factors, including diet, mediate the impact of parasites on hosts, with diet providing nutritional and medicinal properties. Here, we present the first evidence that ongoing environmental change decreases host tolerance and increases parasite virulence through a loss of dietary medicinal quality. Monarch butterflies use dietary toxins (cardenolides) to reduce the deleterious impacts of a protozoan parasite. We fed monarch larvae foliage from four milkweed species grown under either elevated or ambient CO₂, and measured changes in resistance, tolerance, and virulence. The most high‐cardenolide milkweed species lost its medicinal properties under elevated CO₂; monarch tolerance to infection decreased, and parasite virulence increased. Declines in medicinal quality were associated with declines in foliar concentrations of lipophilic cardenolides. Our results emphasize that global environmental change may influence parasite–host interactions through changes in the medicinal properties of plants.     

Control of the circadian rhythm of carbon dioxide assimilation in Bryophyllum leaves by exposure to darkness and high carbon dioxide concentrations

The circadian rhythm of CO₂ assimilation in detached leaves of Bryophyllum fedtschenkoi at 15° C in normal air and continuous illumination is inhibited both by exposure to darkness, and to an atmosphere enriched with 5% CO₂. During such exposures substantial fixation of CO₂ takes place, and the malate concentration in the cell sap increases from about 20 mM to a constant value of 40–50 mM after 16 h. On transferring the darkened leaves to light, and those exposed to 5% CO₂ to normal air, a circadian rhythm of CO₂ assimilation begins again. The phase of this rhythm is determined by the time the transfer is made since the first peak occurs about 24 h afterwards. This finding indicates that the circadian oscillator is driven to, and held at, an identical, fixed phase point in its cycle after 16 h exposure to darkness or to 5% CO₂, and it is from this phase point that oscillation begins after the inhibiting condition is removed. This fixed phase point is characterised by the leaves having acquired a high malate content. The rhythm therefore begins with a period of malate decarboxylation which lasts for about 8 h, during which time the malate content of the leaf cells must be reduced to a value that allows phosphoenolpyruvate carboxylase to become active. Inhibition of the rhythm in darkness, and on exposure to 5% CO₂ in continuous illumination, appears to be due to the presence of a high concentration of CO₂ within the leaf inhibiting malic enzyme which leads to the accumulation of high concentrations of malate in the leaf cells. The malate then allosterically inhibits phosphoenolpyruvate carboxylase upon which the rhythm depends. The results give support to the view that malate synthesis and breakdown form an integral part of the circadian oscillator in this tissue.Abbreviations B. Bryophyllum - PEPCase phosphoenolpyruvate carboxylase     

Responses of global plant diversity capacity to changes in carbon dioxide concentration and climate

We model plant species diversity globally by country to show that future plant diversity capacity has a strong dependence on changing climate and carbon dioxide concentration. CO2 increase, through its impact on net primary production and warming is predicted to increase regional diversity capacity, while warming with constant CO2 leads to decreases in diversity capacity. Increased CO2 concentrations are unlikely to counter projected extinctions of endemic species, shown in earlier studies to be more strongly dependent on changing land use patterns than climate per se. Model predictions were tested against (1) contemporary observations of tree species diversity in different biomes, (2) an independent global map of contemporary species diversity and (3) time sequences of plant naturalisation for different locations. Good agreements between model, observations and naturalisation patterns support the suggestion that future diversity capacity increases are likely to be filled from a 'cosmopolitan weed pool' for which migration appears to be an insignificant barrier.     

Free air carbon dioxide enrichment: development,progress, results

Credible predictions of climate change depend in part on predictions of future CO₂ concentrations in the atmosphere. Terrestrial plants are a large sink for atmospheric CO₂ and the sink rate is influenced by the atmospheric CO₂ concentration. Reliable field experiments are needed to evaluate how terrestrial plants will adjust to increasing CO₂ and thereby influence the rate of change of atmospheric CO₂. Brookhaven National Laboratory (BNL) has developed a unique Free-Air CO₂ Enrichment (FACE) system for a cooperative research program sponsored by the U.S. Department of Energy and U.S. Department of Agriculture, currently operating as the FACE User Facility at the Maricopa Agricultural Center (MAC) of the University of Arizona. The BNL FACE system is a tool for studying the effects of CO₂ enrichment on vegetation and natural ecosystems, and the exchange of carbon between the biosphere and the atmosphere, in open-air settings without any containment. The FACE system provides stable control of CO₂ at 550 ppm ±10%, based on 1-min averages, over 90% of the time. In 1990, this level of control was achieved over an area as large as 380 m², at an annual operating cost of $668 m^–2. During two field seasons of enrichment with cotton (Gossypium hirsutum) as the test plant, enrichment to 550 ppm CO₂ resulted in significant increases in photosynthesis and biomass of leaves, stems and roots, reduced evapotranspiration, and changes in root morphology. In addition, soil respiration increased and evapotranspiration decreased.     

Behavioural responses of rats to gradual-fill carbon dioxide euthanasia and reduced oxygen concentrations

Carbon dioxide (CO₂) is widely used for euthanasia of laboratory rats, but little is known about the pain and distress caused by this procedure. Physiological and human self-report data suggest that CO₂ may cause pain and dyspnea, a sensation of breathlessness and increased respiratory effort, at the concentrations used for gradual-fill euthanasia. However, previous studies examining the behavioural responses of rats have reported conflicting results. In this study, detailed behavioural responses of rats were examined during gradual-fill CO₂ euthanasia and during gradual-fill with argon to produce a similar reduction in oxygen concentration. Animals were randomly allocated to the CO₂ or reduced oxygen treatment groups (n = 8 for both), and were first tested with air exposure and then with treatment gas exposure on the following day. Observations were taken for 105 s before and after gas flow began (baseline and exposure periods), as animals had ceased purposeful movement within 105 s of CO₂ flow starting. Behavioural changes from baseline during gas exposure were compared to changes during air exposure on control days. In comparison to air exposure, CO₂ resulted in increased activity, rearing, touching the nose to the chamber lid, escape behaviours and vocalizations. A small increase in touching the nose to the chamber lid was observed when oxygen concentrations were reduced with argon, but no other behavioural changes were observed. These results suggest that gradual-fill CO₂ euthanasia causes distress in rats, and that hypoxia alone is not a major cause of this distress.     

Responses of respiration to increasing atmospheric carbon dioxide concentrations

It has been recently recognized that increases in carbon dioxide concentration such as are anticipated for the earth's atmosphere in the next century often reduce plant respiration. There can be both a short-term reversible effect of unknown cause, and long-term acclimation, which may reflect the synthesis and maintenance of less metabolically expensive materials in plants grown at elevated carbon dioxide concentrations. Because respiration provides energy and carbon intermediates for growth and maintenance, reductions in respiration by increasing carbon dioxide concentrations may have effects on physiology beyond an improvement in plant carbon balance. As atmospheric carbon dioxide concentration increases, reduced respiration could be as important as increased photosynthesis in improving the ability of terrestrial vegetation to act as a sink for carbon, but it could also have other consequences.     

Warming and depth interact to affect carbon dioxide concentration in aquatic mesocosms

1. Climate change may significantly influence lake carbon dynamics and consequently the exchange of CO₂ with the atmosphere. Warming will accelerate multiple processes that either absorb or release CO₂, making predicting the net effect of warming on CO₂ exchange with the atmosphere difficult. Here we experimentally test how the CO₂ flux of deep and shallow systems responds to warming. To do this, we conducted a greenhouse experiment using mesocosms of two depths, experiencing either ambient or warmed temperatures. 2. Deeper mesocosms were found to have a lower average CO₂ concentration than shallower mesocosms under ambient temperature conditions. In addition, warming interacts with mesocosm depth to affect the average CO₂ concentration; there was no effect of warming on the average CO₂ concentration of deep mesocosms, but shallow mesocosms had significantly lower average CO₂ concentrations when warmed. 3. The difference in CO₂ concentration resulting from the depth manipulation was due to varying loss rates of particulate carbon to the sediments. There was a strong negative correlation between CO₂ and sedimentation rates in the deep mesocosms suggesting that high particulate carbon loss to the sediments lowered the CO₂ concentration in the water column. There was no correlation between CO₂ and sedimentation rates observed for shallow mesocosms suggesting enhanced carbon regeneration from the sediments was maintaining higher CO₂ concentrations in the water column. 4. Relationships between CO₂ and algal concentrations indicate that the reduction in CO₂ concentrations resulting from warming is due to increased per capita algal turnover rates depleting CO₂ in the water column. Our results suggest that the carbon dynamics and CO₂ flux of shallow systems will be affected more by climate warming than deep systems and the net effect of warming is to increase CO₂ uptake.     

Solar ultraviolet-B radiation can affect slug feeding preference for some plant species native to a fen ecosystem in Tierra del Fuego,Argentina

The objectives of this study were to test potential effects of solar ultraviolet-B (UV-B) radiation on (i) foliage nutritional quality and foliage decomposition rates of six plant species of this fen ecosystem (Nothofagus antarctica, Carex curta, C. decidua and C. magellanica; Acaena magellanica and Gunnera magellanica) and (ii) feeding preferences for these plant species of the slug Deroceras reticulatum prevalent in this ecosystem. In a mixed-diet selection slugs were offered leaves of the six species that had been grown for three years in experimental field plots under either near-ambient or reduced solar ultraviolet-B (UV-B) radiation. The chosen characteristics of leaf quality (nitrogen concentration, carbon:nitrogen ratio, specific leaf area) and leaf decomposition rates of the six species varied significantly among species but were not affected by the UV-B treatments. However, there were UV-B treatment effects on slug feeding preference for two plant species. For the tree species, Nothofagus, slugs had consumed only one-third as much foliage grown under near-ambient UV-B radiation as of foliage grown under reduced UV-B by the end of the feeding experiment. In contrast, leaves of the sedge C. decidua that had been grown under near-ambient UV-B were consumed twice as much as leaves grown under reduced UV-B radiation. Consumption of foliage for the other four species was similar for the two UV-B treatments. Additionally, diet selection of the slugs was also significantly affected by prior UV-B conditions under which foliage had been grown. Nothofagus leaves were consumed proportionately less and C. decidua proportionately more if the foliage had been grown under near-ambient UV-B radiation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Urea additions and defoliation affect plant responses to elevated CO2 in a C3 grass from Yellowstone National Park

A common grass from Yellowstone National Park, Stipa occidentalis, was grown in a factorial experiment to determine if its response to the direct effects of elevated CO₂ would be affected by defoliation, and urea additions simulating the N in a urine hit. Plants were grown in tall pots (to mimic rooting depth in the field) in growth chambers under elevated (700 ppm) and ambient (370 ppm) CO₂, were defoliated or left undefoliated, and given N-supply rates based on field mineralization rates (untreated) or with an additional 40 g N/m². Growth increases in response to elevated CO₂ were largest when plants remained unclipped and received urea additions, and were found primarily in crowns and roots (storage organs). Aboveground biomass, which is the part of the plant consumed by grazing mammals, was not affected by elevated CO₂. The elevated CO₂ treatment caused a reduction in leaf percent N. However, there was a significant interaction between the CO₂ and urea treatments, resulting in a larger difference in leaf percent N between urea-treated and control plants under elevated than under ambient CO₂. Hence, elevations in atmospheric CO₂ may cause an increase in the amount of urine-hit-induced spatial variability in temperate grasslands. Since food quantity remained largely unchanged in response to elevated CO₂, and forage N content went down, grazing mammals may be negatively affected by increases in atmospheric CO₂.     

Soil carbon dioxide partial pressure and dissolved inorganic carbonate chemistry under elevated carbon dioxide and ozone

Global emissions of atmospheric CO₂ and tropospheric O₃ are rising and expected to impact large areas of the Earths forests. While CO₂ stimulates net primary production, O₃ reduces photosynthesis, altering plant C allocation and reducing ecosystem C storage. The effects of multiple air pollutants can alter belowground C allocation, leading to changes in the partial pressure of CO₂ (pCO₂) in the soil , chemistry of dissolved inorganic carbonate (DIC) and the rate of mineral weathering. As this system represents a linkage between the long- and short-term C cycles and sequestration of atmospheric CO₂, changes in atmospheric chemistry that affect net primary production may alter the fate of C in these ecosystems. To date, little is known about the combined effects of elevated CO₂ and O₃ on the inorganic C cycle in forest systems. Free air CO₂ and O₃ enrichment (FACE) technology was used at the Aspen FACE project in Rhinelander, Wisconsin to understand how elevated atmospheric CO₂ and O₃ interact to alter pCO₂ and DIC concentrations in the soil. Ambient and elevated CO₂ levels were 360±16 and 542±81 l l^–1, respectively; ambient and elevated O₃ levels were 33±14 and 49±24 nl l^–1, respectively. Measured concentrations of soil CO₂ and calculated concentrations of DIC increased over the growing season by 14 and 22%, respectively, under elevated atmospheric CO₂ and were unaffected by elevated tropospheric O₃. The increased concentration of DIC altered inorganic carbonate chemistry by increasing system total alkalinity by 210%, likely due to enhanced chemical weathering. The study also demonstrated the close coupling between the seasonal ¹³C of soil pCO₂ and DIC, as a mixing model showed that new atmospheric CO₂ accounted for approximately 90% of the C leaving the system as DIC. This study illustrates the potential of using stable isotopic techniques and FACE technology to examine long- and short-term ecosystem C sequestration.     

Increased mercury in forest soils under elevated carbon dioxide

Fossil fuel combustion is the primary anthropogenic source of both CO₂ and Hg to the atmosphere. On a global scale, most Hg that enters ecosystems is derived from atmospheric Hg that deposits onto the land surface. Increasing concentrations of atmospheric CO₂ may affect Hg deposition to terrestrial systems and storage in soils through CO₂-mediated changes in plant and soil properties. We show, using free-air CO₂ enrichment (FACE) experiments, that soil Hg concentrations are almost 30% greater under elevated atmospheric CO₂ in two temperate forests. There were no direct CO₂ effects, however, on litterfall, throughfall or stemflow Hg inputs. Soil Hg was positively correlated with percent soil organic matter (SOM), suggesting that CO₂-mediated changes in SOM have influenced soil Hg concentrations. Through its impacts on SOM, elevated atmospheric CO₂ may increase the Hg storage capacity of soils and modulate the movement of Hg through the biosphere. Such effects of rising CO₂, ones that transcend the typically studied effects on C and nutrient cycling, are an important next phase for research on global environmental change.     

Studies on the anesthetic mechanism of carbon dioxide by using Bombyx mori larvae.

The study of heartbeat variations of Bombyx mori larvae submitted to CO2 narcosis has provided new information on carbon dioxide anesthesia. Purely anoxiant action must be dismissed. Action at a nerve level is possible since CO2 produces the same effect as ether, which is a classical anesthetic.     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

Simulated sequestration of industrial carbon dioxide at a deep-sea site: Effects on species of harpacticoid copepods

One proposal for alleviating global warming is to sequester large amounts of industrial carbon dioxide on the deep-sea floor, but the environmental consequences of sequestration for the animals living in the sediment are poorly known. In an earlier publication, we reported that, during an experimental sequestration off central California (36.378°N, 122.676°W, 3262 m depth), most individuals of our target taxon (the harpacticoid copepods) were killed, but ∼ 20% survived. Because knowledge of which species survived and how they did so could clarify the effects of sequestration on the fauna, we have now identified the individuals from that experiment to species. Although most were adversely affected, species differed significantly in the degree of their susceptibility. Unexpectedly, six species showed no effect and may be resistant. The hypothesis that harpacticoids could escape the effects of carbon dioxide-rich seawater by moving deeper into the seabed was not supported. Exposure to carbon dioxide-rich seawater created partially defaunated areas, but we found no evidence that disturbance-exploiting harpacticoid species invaded during the recovery of the affected area. Because the environmental effects of the carbon dioxide (e.g. unusually acidic pore water) were still present, however, the opportunity for invasion might not yet have occurred. Differences among species in susceptibility increase the complexity of the effects of carbon dioxide sequestration on the deep-sea fauna.     

Elevated carbon dioxide affects leaf-miner performance and plant growth in docks (Rumex spp.)

Exposure of R. crispus and R. obtusifolius to elevated CO₂ (600 ppm) resulted in an increased C:N ratio of leaf tissue and greater leaf areas. Larvae of P. nigritarsis mining leaves of R. obtusifolius during exposure produced significantly bigger mines in elevated than in ambient (350 ppm) conditions. There were no significant treatment effects on pupal weight although in both host species mean weight was greater in ambient than in elevated conditions. These results are consistent with the hypothesis that insect herbivores compensate for increased C:N ratios by increased food consumption. This response by herbivores may partially offset predicted increases in plant biomass in a future high CO₂ environment.     

Ventilatory responses to exercise and carbon dioxide in elderly and younger humans

The present investigation examined the relationship between CO₂ sensitivity [at rest (S _R) and during exercise (S _E)] and the ventilatory response to exercise in ten elderly (61–79 years) and ten younger (17–26 years) subjects. The gradient of the relationship between minute ventilation and CO₂ production ( _E/ CO₂) of the elderly subjects was greater than that of the younger subjects [mean (SEM); 32.8 (1.6) vs 27.3 (0.4); P<0.01]. At rest, S _R was lower for the elderly than for the younger group [10.77 (1.72) vs 16.95 (2.13) 1 · min^–1 · kPa^–1; 1.44 (0.23) vs 2.26 (0.28) 1 · min^–1 · mmHg^–1; P<0.05], but S _E was not significantly different between the two groups [17.85 (2.49) vs 19.17 (1.62) l · min^–1 · kPa^–1; 2.38 (0.33) vs 2.56 (0.21) 1 · min^–1 · mmHg^–1]. There were significant correlations between both S _R and S _E, and _E/ CO₂ (P<0.05; P<0.001) for the younger group, bot none for the elderly. The absence of a correlation for the elderly supports the suggestion that _E/ CO₂ is not an appropriate index of the ventilatory response to exercise for elderly humans.