Carbon dioxide concentrations in the nests of the mud-dwelling mangrove ant Polyrhachis sokolova Forel (Hymenoptera: Formicidae)

Abstract The nests of the mangrove ant Polyrhachis sokolova are found in soil in intertidal mangrove communities, and are thus inundated at high tides for several hours. Some of the nest galleries are flooded, but others retain air pockets, to which the ants retreat. During and following inundation, we measured carbon dioxide concentrations in air samples collected from different levels in the nests and from artificial 'control' holes in the mud. To account for the relative contribution of different sources of carbon dioxide, we also measured the carbon dioxide production by individual ants (including larvae and pupae) and small samples of mud collected near the ant nests. Nest carbon dioxide concentrations were high (2.5−11%) during and immediately following inundation, but the concentrations in the upper regions of the nest fell as soil water levels receded. However, at depths>10 cm below the level soil surface, the carbon dioxide concentrations remained relatively high and stable (at approximately 2%) over the 11 days between one high tide and the next. The contribution of the mud (and associated microorganisms) to the carbon dioxide concentration in the nests was substantial, and the contribution of the respiration of the ants was approximately 10−15% of the total. The carbon dioxide concentrations in the nests of this species during high tides are among the highest recorded for insect nests, suggesting that these ants may have unusual physiological attributes to match the behavioural and ecological challenges associated with living in the intertidal zone.     

Respiratory patterns in nurses of the red wood ant Formica polyctena (Hymenoptera, Formicidae)

Summary Cycles of discontinuous gas exchange (DGCs) and abdominal ventilatory movements were studied in nurses of red wood ant, Formica polyctena, using an electrolytic respirometer and an infra-red (IR) gas analyser or flow-through respirometry. Both respirometry systems were combined with an IR actographic device based on IR-emitting and IR-sensor diodes. After recovering from handling and apparatus stress, lasting 1–3 h, completely motionless intact ants displayed regular DGC. After decapitation the ants displayed DGC whose frequency was somewhat lower than that of the intact individuals (7.17 ± 0.79 mHz and 10.43 ± 01.12 mHz, respectively). In headless ants, there occurred continuous slow movements of legs. Bursts of carbon dioxide in the intact and in the headless ants always coincided with a bout of telescoping movements (contractions) of abdominal segments, which was interpreted as active ventilation. During interburst periods, the headless ants exhibited telescoping movements characterised by rapid protraction, lasting 0.07–0.09 s, followed by a slow retraction of segments, suggesting passive ventilation. The intact ants were very sensitive to the flowing air and tended to be continuously active during flow-through respirometry. The decapitated ants, on the contrary, were insensitive to the air current.Received 26 March 2003; revised 30 July 2003; accepted 6 August 2003.     

Calculation of diffusion coefficient for supercritical carbon dioxide and carbon dioxide+naphthalene system by molecular dynamics simulation using EPM2 model

NVT ensemble molecular dynamics (MD) simulation has been applied to calculate the self-diffusion coefficients of carbon dioxide and the tracer diffusion coefficients of naphthalene in supercritical carbon dioxide. The simulation was carried out in the pressure range from 8 to 40 MPa. The elementary physical model proposed by Harris and Yung was adopted for carbon dioxide and some approximation models were used for naphthalene. The systems of MD simulation for carbon dioxide consist of 256 particles. One naphthalene molecule was added for carbon dioxide+naphthalene system. The system can be assumed to be an infinite dilution condition for carbon dioxide+naphthalene system and the mutual diffusion coefficients are equal to the tracer diffusion coefficients of naphthalene. The self-diffusion coefficients of carbon dioxide and the tracer diffusion coefficients of naphthalene in supercritical carbon dioxide can be calculated by mean square displacement. The calculated results of diffusion coefficients showed good agreement with the experimental data without adjustable parameters.     

Field sampling of unfed nymphs ofAmblyomma hebraeum

Unfed nymphs ofAmblyomma hebraeum Koch, 1844 shelter under debris on the soil surface and become active in host-seeking when stimulated with carbon dioxide. The active nymphs are not attracted to sources of carbon dioxide and hence cannot be effectively sampled using static carbon dioxide traps. However, these nymphs will cling to flannelette and can be sampled by flagging. Larger numbers are recovered if the soil surface is sampled using a vacuum device.     

Today and tomorrow: impact of climate change on aphid biology and potential consequences on their mutualism with ants

Recent studies about mutualism consider the complexity and versatility of the relationship, in addition to highlighting the importance of the cost/benefit balance between the two protagonists. Because species interactions are highly dependent on the environment, the climate changes foreseen for the coming years are expected to have significant impacts on the evolution of mutualistic interactions. Among mutualisms, the aphid–ant interaction is well documented, partly explained by the pest status of aphids. This literature review focuses on the impact of climate change (particularly atmospheric carbon dioxide concentration and temperature) on aphid biology and the potential consequences with respect to their mutualistic interactions with ants. We provide an overview of the published reports concerned with the effects of temperature and carbon dioxide on aphids, for which a positive, a negative or no effect has been highlighted. We then discuss how climatic changes can alter four major components of aphid biology that are shaping their interaction with ants: (i) aphid population growth; (ii) aphid behaviour and mobility; (iii) honeydew production and composition; and (iv) semiochemistry. Finaly, we discuss the limitations of such studies on aphid–ant mutualism, as well as the information that is still needed to predict how climate change might impact this type of relationship.     

Stomatal conductance, photosynthesis and respiration of temperate deciduous tree seedlings grown outdoors at an elevated concentration of carbon dioxide

Seedlings of temperate deciduous tree species were grown outdoors at ambient and at an elevated concentration of carbon dioxide to examine how aspects of their gas exchange would be altered by growth at elevated carbon dioxide concentration. Leaf conductances to water vapour and net carbon dioxide exchange rates were determined periodically near midday. Whole-plant carbon dioxide efflux rates in darkness were also determined. The stomatal conductance of leaves of plants grown and measured at 700 cm³ m^?3 carbon dioxide did not differ from that of plants grown and measured at 350 cm³ m^?3 in Malus domestica, Quercus prinus and Quercus robur at any measurement time. In Acer saccharinum, lower conductances occurred for plants grown and measured at elevated carbon dioxide concentration only at measurement temperatures above 33°C. Photo-synthetic adjustment to elevated carbon dioxide concentration was evident only in Q. robur. All species examined had lower rates of dark respiration per unit of mass when grown and measured at elevated carbon dioxide concentration.     

Increased Risk Proneness or Social Withdrawal? The Effects of Shortened Life Expectancy on the Expression of Rescue Behavior in Workers of the ant <Emphasis Type="Italic">Formica cinerea</Emphasis> (Hymenoptera: Formicidae)

In social insects behavioral consequences of shortened life expectancy include, among others, increased risk proneness and social withdrawal. We investigated the impact of experimental shortening of life expectancy of foragers of the ant Formica cinerea achieved by their exposure to carbon dioxide on the expression of rescue behavior, risky pro-social behavior, tested by means of two bioassays during which a single worker (rescuer) was confronted with a nestmate (victim) attacked by a predator (antlion larva capture bioassay) or immobilized by an artificial snare (entrapment bioassay). Efficacy of carbon dioxide poisoning in shortening life expectancy was confirmed by the analysis of ant mortality. Rescue behavior observed during behavioral tests involved digging around the victim, transport of the sand covering the victim, pulling the limbs/antennae/mandibles of the victim, direct attack on the antlion (in antlion larva capture tests), and snare biting (in entrapment tests). The rate of occurrence of rescue behavior was lower in ants with shortened life expectancy, but that effect was significant only in the case of the entrapment bioassay. Similarly, only in the case of the entrapment bioassay ants with shortened life expectancy displayed rescue behavior after a longer latency and devoted less time to that behavior than ants from the control groups. Our results demonstrated that in ant workers shortened life expectancy may lead to reduced propensity for rescue behavior, most probably as an element of the social withdrawal syndrome that had already been described in several studies on behavior of moribund ants and honeybees.     

Insect herbivore stoichiometry: the relative importance of host plants and ant mutualists

Abstract.  1. Mutualistic associations can vary over spatial and ecological gradients. For herbivorous insects that engage in mutualisms with ants, plant quality can be a particularly important source of variation, because of the upward transfer of nutrients from plants to herbivores to ants.
2. A previous study demonstrated that mutualistic ants, Formica obscuripes , exert a top-down effect on the carbon and nitrogen concentrations (stoichiometry) in an herbivorous membracid, Publilia modesta . We characterised the consequences of mutualism for carbon and nitrogen stoichiometry between the same species pair, yet on an alternative, geographically-distinct host plant.
3. We found no top-down effect of ants on the carbon or nitrogen in the herbivore, but a strong, bottom-up effect of individual plants on membracid nitrogen concentration.
4. These results suggests that spatial heterogeneity in host plant traits, and ultimately the diet breadth of herbivore mutualists, may be important factors mediating stoichiometric patterns in mutualistic associations.     

Current views on the regulation of autotrophic carbon dioxide fixation via the Calvin cycle in bacteria

The Calvin cycle of carbon dioxide fixation constitutes a biosynthetic pathway for the generation of (multi-carbon) intermediates of central metabolism from the one-carbon compound carbon dioxide. The product of this cycle can be used as a precursor for the synthesis of all components of cell material. Autotrophic carbon dioxide fixation is energetically expensive and it is therefore not surprising that in the various groups of autotrophic bacteria the operation of the cycle is under strict metabolic control. Synthesis of phosphoribulokinase and ribulose-1,5-bisphosphate carboxylase, the two enzymes specifically involved in the Calvin cycle, is regulated via end-product repression. In this control phosphoenolpyruvate most likely has an alarmone function. Studies of the enzymes isolated from various sources have indicated that phosphoribulokinase is the target enzyme for the control of the rate of carbon dioxide fixation via the Calvin cycle through modulation of existing enzyme activity. In general, this enzyme is strongly activated by NADH, whereas AMP and phosphoenol-pyruvate are effective inhibitors. Recent studies of phosphoribulokinase inAlcaligenes eutrophus suggest that this enzyme may also be regulated via covalent modification.     

Direct Observation of Spin-Trapped Carbon Dioxide Radicals in Hepatocytes Exposed to Carbon Tetrachloride

Carbon dioxide radical adducts of the spin trapping agent, α-phenyl N-t-butyl nitrone (PBN), have been observed to occur in the urine and bile of rats exposed to carbon tetrachloride as well as in perfusates of liver in which the perfusion medium contained carbon tetrachloride (Connor er al., J. Biol. Chem., 261, 4542, (1986)). The carbon dioxide adduct was proven to be derived from CCI, by use of 13-C-labelled compound. These adducts were not observed in the liver itself suggesting that they might be rapidly secreted from the liver. However, using isolated hepatocytes, we have demonstrated that the carbon dioxide radical adduct can be observed directly in the liver cells as it is formed. Since this water-soluble adduct cannot be extracted by non-aqueous solvents such as chloroform or toluene, its formation in liver in vivo or in perfused livers was not detected. Lowering the oxygen tension in the system diminished the intensity of production of the carbon dioxide adduct, consistent with the adduct being produced as a result of ·OOCCl₃ generation. It is not clear the extent to which this adduct is formed as a result of the ·CO₂ radical or is produced by metabolic oxidation of the trichloromethyl radical adduct of PBN per se to the carbon dioxide radical adduct. The intensity of the signal of the carbon dioxide radical adduct suggests that adduct conversion may be the route of formation since it seems unlikely that a sufficient amount of the halocarbon could be metabolized to ·COCl or ·CO₂ radicals to generate a signal of the magnitude involved. The ·CO₂ adduct is readily observed in intact hepatocytes, but the ·CCl₃ adduct is not (although we know the ·CCl₃ adduct has been produced in these cells), indicating that the ·CO₂ adduct is present in considerable abundance compared to the ·CCl₃ adduct.     

Patterns of univariate and multivariate plasticity to elevated carbon dioxide in six European populations of Arabidopsis thaliana

The impact of elevated carbon dioxide on plants is a growing concern in evolutionary ecology and global change biology. Characterizing patterns of phenotypic integration and multivariate plasticity to elevated carbon dioxide can provide insights into ecological and evolutionary dynamics in future human‐altered environments. Here, we examined univariate and multivariate responses to carbon enrichment in six functional traits among six European accessions of Arabidopsis thaliana. We detected phenotypic plasticity in both univariate and multivariate phenotypes, but did not find significant variation in plasticity (genotype by environment interactions) within or among accessions. Eigenvector, eigenvalue variance, and common principal components analyses showed that elevated carbon dioxide altered patterns of trait covariance, reduced the strength of phenotypic integration, and decreased population‐level differentiation in the multivariate phenotype. Our data suggest that future carbon dioxide conditions may influence evolutionary dynamics in natural populations of A. thaliana.     

Cues used in host‐seeking behavior by frog‐biting midges (Corethrella spp. Coquillet)

We investigated the role of carbon dioxide and host temperature in host attraction in frog‐biting midges (Corethrella spp). In these midges, females are known to use frog calls to localize their host, but the role of other host‐emitted cues has yet not been investigated. We hypothesized that carbon dioxide acts as a supplemental cue to frog calls. To test this hypothesis, we determined the responses of the midges to carbon dioxide, frog calls, and both cues. A significantly lower number of midges are attracted to carbon dioxide and silent traps than to traps broadcasting frog calls. Adding carbon dioxide to the calls does not increase the attractiveness to the midges. Instead, carbon dioxide can have deterrent effects on frog‐biting midges. Temperature of calling frogs is not a cue potentially available to the midges. Contrary to our hypothesis, there was no supplemental effect of carbon dioxide when presented in conjunction to calls. Midge host‐seeking behavior strongly depends on the mating calls emitted by their anuran host. Overall, non‐acoustic cues such as host body temperature and carbon dioxide are not important in long‐distance host location by frog‐biting midges.     

Nutrient dynamics in a tritrophic system of ants,aphids and beans

The interactions between ants (Lasius niger), aphids (Aphis fabae) and plants (Phaseolus vulgaris) were studied in a laboratory experiment with the following treatments: application of N‐fertilizer for plants, supply of mealworms or sugar solution or their combination as alternative food sources for ants. Three main questions were studied: (1) Do ants reduce tending to honeydew‐producing aphids when an alternative sugar or protein resource was available? (2) Is aphid predation/protein consumption by ants higher when additional carbon is offered to maintain the carbon/protein balance? (3) Does fertilizer treatment propagate in the food web? For the experimental analysis stable isotope techniques were applied. δ¹⁵N served as a marker for the pathway from plants to higher trophic levels. Low δ¹⁵N‐value of fertilizer spread from plant shoots to aphids and ants. To trace which sugar‐/protein source was consumed by ants, the different ¹³C/¹²C‐ratios of C₃‐ and C₄‐plants were used with aphids feeding on C₃‐plant material, while mealworm food and sugar solution originated from C₄‐plant material. Fertilizer application had no effect on biomasses of plants, consumers or microflora. Ant biomass was significantly higher when additional sugar solution was offered. Higher contents of ¹³C indicated a high incorporation rate of additional sugar. Additional protein had no effect on colony biomass and no increasing predation on aphids could be observed when carbon was in excess. However, due to the lack of queens and newly produced larvae, protein requirements of experimental colonies were lower than in natural systems. Ants positively affected aphid populations, but reduced tending, whilst having access to an alternative sugar resource. When sugar/protein was offered to ants, the host plant had an increased root/shoot ratio. This indicates that decreasing aboveground activity of ants could lead to reduced plant growth of aphid‐infested plants, presumably due to higher fungal attack on shoots.     

Interactions between ants attendingAphis fabae ssp.cirsiiacanthoidis on thistles and foraging parasitoid wasps

We describe the behavioral interactions between honeydew-collecting workers of the ants Lasius nigerand Myrmica ruginodisand females of three species of aphidiid wasps (Lysiphlebus cardui, Lysiphlebus testaceipes, Trioxys angelicae)foraging for their aphid host, Aphis fabaessp. cirsiiacanthoidis,on thistles. Using field and laboratory experiments, we show that the ant-parasitoid interactions are species specific. Workers of both ant species generally attacked and killed females of T. angelicae,but they ignored those of L. cardui.This pattern was not altered when we anesthetized the wasps slightly with carbon dioxide to reduce their mobility. Prior contacts between L. carduiand either conspecific L. nigerfrom a different nest or workers of a different ant species (M. ruginodis)did not influence L. niger'snonaggressive behavior. The number of aphids parasitized by L. testaceipeswas significantly reduced in aphid colonies attended by L. niger,although this parasitoid was rarely attacked by ants. In encounters between these species of ants and wasps, ant aggression is consistent with differences in wasp behavior. We suggest that, in addition, chemical cues located in the cuticula may enable L. carduito avoid detection by honeydew-collecting ants.     

Responses of nitrogen metabolism in N-sufficient barley primary leaves to plant growth in elevated atmospheric carbon dioxide

Effects of atmospheric carbon dioxide enrichment on nitrogen metabolism were studied in barley primary leaves (Hordeum vulgare L. cv. Brant). Seedlings were grown in chambers under ambient (36 Pa) and elevated (100 Pa) carbon dioxide and were fertilized daily with complete nutrient solution providing 12 millimolar nitrate and 2.5 millimolar ammonium. Foliar nitrate and ammonium were 27% and 42% lower (P ≤ 0.01) in the elevated compared to ambient carbon dioxide treatments, respectively. Enhanced carbon dioxide affected leaf ammonium levels by inhibiting photorespiration. Diurnal variations of total nitrate were not observed in either treatment. Total and Mg²⁺inhibited nitrate reductase activities per gram fresh weight were slightly lower (P ≤ 0.01) in enhanced compared to ambient carbon dioxide between 8 and 15 DAS. Diurnal variations of total nitrate reductase activity in barley primary leaves were similar in either treatment except between 7 and 10 h of the photoperiod when enzyme activities were decreased (P ≤ 0.05) by carbon dioxide enrichment. Glutamate was similar and glutamine levels were increased by carbon dioxide enrichment between 8 and 13 DAS. However, both glutamate and glutamine were negatively impacted by elevated carbon dioxide when leaf yellowing was observed 15 and 17 DAS. The above findings showed that carbon dioxide enrichment produced only slight modifications in leaf nitrogen metabolism and that the chlorosis of barley primary leaves observed under enhanced carbon dioxide was probably not attributable to a nutritionally induced nitrogen limitation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Coexistence through mutualist‐dependent reversal of competitive hierarchies

Mechanisms that allow for the coexistence of two competing species that share a trophic level can be broadly divided into those that prevent competitive exclusion of one species within a local area, and those that allow for coexistence only at a regional level. While the presence of aphid‐tending ants can change the distribution of aphids among host plants, the role of mutualistic ants has not been fully explored to understand coexistence of multiple aphid species in a community. The tansy plant (Tanacetum vulgare) hosts three common and specialized aphid species, with only one being tended by ants. Often, these aphids species will not coexist on the same plant but will coexist across multiple plant hosts in a field. In this study, we aim to understand how interactions with mutualistic ants and predators affect the coexistence of multiple species of aphid herbivores on tansy. We show that the presence of ants drives community assembly at the level of individual plant, that is, the local community, by favoring one ant‐tended species, Metopeurum fuscoviride, while preying on the untended Macrosiphoniella tanacetaria and, to a lesser extent, Uroleucon tanaceti. Competitive hierarchies without ants were very different from those with ants. At the regional level, multiple tansy plants provide a habitat across which all aphid species can coexist at the larger spatial scale, while being competitively excluded at the local scale. In this case, ant mutualist‐dependent reversal of the competitive hierarchy can drive community dynamics in a plant–aphid system.     

Acclimation to temperature of the response of photosynthesis to increased carbon dioxide concentration in Taraxacum officinale

The relative stimulation of photosynthesis by elevated carbon dioxide in C₃ species normally increases strongly with increasing temperature. This results from the kinetic characteristics of Rubisco, and has potentially important implications for responses of vegetation to increasing atmospheric carbon dioxide. It is often assumed that because Rubisco characteristics are conservative, all C₃ species have the same temperature dependence of the response of photosynthesis to elevated carbon dioxide. However, in this field study of Taraxacum officinale, there were no significant differences in the relative stimulation of photosynthesis by elevated carbon dioxide among days with temperatures ranging from 15 to 34 °C. Nevertheless, short-term measurements indicated a strong temperature dependence of the stimulation. This suggested that acclimation to temperature caused the lack of variation in the seasonal data. Experiments in controlled environments indicated that complete acclimation of the relative stimulation of photosynthesis by elevated carbon dioxide occurred for growth temperatures of 10 – 25 °C. The apparent specificity of Rubisco for carbon dioxide relative to oxygen at 15 °C, as assayed in vivo by measurements of the carbon dioxide concentration at which carboxylation equalled oxygenation, also varied with growth temperature. Changes in the apparent specificity of Rubisco accounted for the acclimation of the temperature dependence of the relative stimulation of photosynthesis by elevated carbon dioxide. It is premature to conclude that low temperatures will necessarily reduce the relative stimulation of photosynthesis caused by rising atmospheric carbon dioxide. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mathematical modelling of pulmonary gas transport

 Equations governing the transport of the gases oxygen and carbon dioxide inside the pulmonary capillaries are written down. By analysing these equations it is predicted that there will be negligible limitation to the transport of oxygen when oxygen concentration takes a normal physiological or higher value. For low values of oxygen concentration, there may be limitation to oxygen transport. It is predicted further that the quantity of carbon dioxide excreted from blood into alveolar gas is dependent on oxygen concentration, with low oxygen concentrations inhibiting the carbon dioxide transport process. The relatively slow reaction involving carbon dioxide in plasma also inhibits the excretion of carbon dioxide. These predictions are verified by solving the whole system of governing equations numerically. Received: 1 May 2002 / Revised version: 20 October 2002 / Published online: 19 March 2003 JPW was supported by a grant from the Engineering and Physical Sciences Research Council of Great Britain. Key words or phrases: Pulmonary gas transport – Haemoglobin – Saturation