Responses of the biting midge Culicoides impunctatus to acetone, CO2 and 1-octen-3-ol in a wind tunnel

Responses of host-seeking female Culicoides impunctatus (Goetghebuer) (Diptera: Ceratopogonidae) to acetone, carbon dioxide and 1-octen-3-ol were measured in a wind tunnel. Carbon dioxide, presented as a filamentous plume, increased upwind flight in a dose-dependent manner, up to 0.09% concentration. A homogenous CO2 plume elicited similar upwind responses at concentrations up to 0.09%, whereas higher plume concentrations (> 0.1%) induced erratic responses with a suppression of upwind flight. Bovine equivalent concentrations of acetone (1.5 x 10(-6)g/l) and 1-octen-3-ol (1.3 x 10(-8)g/) failed to induce any significant upwind response when tested alone. In the presence of CO2, however, 1-octen-3-ol showed highly significant increases in upwind responses at concentrations of 1.3 x 10(-1) - 10(-8)g/l. Mixtures of CO2+ acetone also enhanced upwind flight at 1.5 x 10(-9)g/l. High tunnel concentrations of both 1-octen-3-ol and acetone inhibited upwind responses. These findings are discussed in relation to host finding by C. impunctatus and known mechanisms by which upwind flight is initiated and arrested at high odour concentrations.     

Activation, orientation and landing of female Culex quinquefasciatus in response to carbon dioxide and odour from human feet: 3-D flight analysis in a wind tunnel

This study investigated the interaction between carbon dioxide (CO(2) ) and human foot odour on activation, upwind orientation and landing of host-seeking female Culex quinquefasciatus (Say) (Diptera: Culicidae) in a wind tunnel. More mosquitoes landed on warmed glass beads coated with foot odour than on clean beads; adding a plume of 4% CO(2) did not influence the proportion of mosquitoes landing. A second experiment used 3-dimensional video tracking to assess flight performance. Activation was more rapid with CO(2) and with CO(2) + foot odour than with clean air or with foot odour alone. Upwind flights were fastest with CO(2) and with clean air, and slowest with foot odour; the CO(2) + foot odour treatment overlapped the previous three treatments in significance. Flight headings tended more towards due upwind with CO(2) and with clean air than with CO(2) + foot odour or with foot odour alone. In both experiments, many mosquitoes flew upwind in clean air. There was little evidence of females changing course upon entering or exiting the CO(2) plume or reacting to foot odour during flight.     

Four simple stimuli that induce host‐seeking and blood‐feeding behaviors in two mosquito species,with a clue to DEET's mode of action

Bioassays in a wind tunnel showed that a combination of four stimuli releases intense host‐seeking and blood‐feeding behavioral responses from females of the Asian tiger mosquito, Aedes albopictus, and the yellow fever mosquito, Aedes aegypti. The stimuli are carbon dioxide, water vapor, warmth, and adenosine triphosphate (ATP). Mosquitoes responded to this combination with a repertoire of blood‐feeding behaviors that included upwind flight, landing, probing, and engorgement. Absence of carbon dioxide, water vapor, or ATP from the combination of stimuli or exposure to temperatures 12° C below or above human‐host temperature (38° C) significantly attenuated blood‐feeding behavior in both species. Although there is literature documenting the individual importance of each of these stimuli, our work represents the first instance where this combination of stimuli was found sufficient to elicit a complete repertoire of blood‐feeding behaviors in these mosquitoes without involvement of any host specific odor. When mosquitoes were exposed to the four stimuli along with N,N‐diethyl‐3‐methylbenzamide (DEET), feeding behavior was greatly suppressed. We hypothesize that a possible mode of action for DEET against these mosquitoes involves interference of warmth and/or water vapor receptors. An electrophysiological study designed to determine if DEET adversely affects the function of these receptors would be illuminating.     

Maxillary palps are broad spectrum odorant detectors in Culex quinquefasciatus

A single type of olfactory sensilla on maxillary palps in many species of mosquitoes houses a very sensitive olfactory receptor neuron (ORN) for carbon dioxide reception. We performed extensive single sensillum recordings from this peg sensillum in Culex quinquefasciatus and have characterized the response threshold and kinetics for CO(2) reception, with a detection threshold less than the CO(2) concentration in the atmosphere. This ORN responded in a tonic mode to lower concentrations of CO(2), whereas higher concentrations generated a phasic-tonic mode of action potential firing. Sensillum potentials accurately represented the response magnitude and kinetics of carbon dioxide-elicited excitatory responses. Stimulation of these ORNs with human breath, a complex mixture of mosquito kairomones and up to 4.5% CO(2), elicited excitatory responses that were reliably detected by CO(2)-sensitive ORNs. Another ORN housed in these sensilla responded to 1-octen-3-ol and to various plant-derived compounds, particularly floral and green leaf volatiles. This ORN showed remarkable sensitivity to the natural enantiomer, (R)-(-)-1-octen-3-ol, rivaling pheromone-detecting ORNs in moths. Maximum neuronal response was elicited with a 10 ng dose. A biological, ecological role of maxillary palps in detection of plant- and nectar-related sources is proposed.     

Close encounters: contributions of carbon dioxide and human skin odour to finding and landing on a host in Aedes aegypti

In a wind‐tunnel study, the upwind flight and source location of female Aedes aegypti to plumes of carbon dioxide (CO₂) gas and odour from human feet is tested. Both odour sources are presented singly and in combination. Flight upwind along the plumes is evident for both CO₂ and odour from human feet when the odours are presented alone. Similarly, both odour sources are located by more than 70% of mosquitoes in less than 3 min. When both CO₂ and odour from human feet are presented simultaneously in two different choice tests (with plumes superimposed or with plumes separated), there is no evidence that females orientate along the plume of CO₂ and only a few mosquitoes locate its source. Rather, the foot odour plume is navigated and the source of foot odour is located by over 80% of female Ae. aegypti. When a female is presented a plume of CO₂ within a broad plume of human foot odour of relatively low concentration, the source of CO₂ is not located; instead, flight is upwind in the diffuse plume of foot odour. Although upwind flight by Ae. aegypti at long range is presumably induced by CO₂ and the threshold of response to skin odours is lowered, our findings suggest that, once females have arrived near a prospective human host, upwind orientation and landing are largely governed by the suite of human odours, whereas orientation is no longer influenced by CO₂.     

Gas Exchange of Algae: III. Relation between the Concentration of Carbon Dioxide in the Nutrient Medium and the Oxygen Production of Chlorella pyrenoidosa

The oxygen production of a photosynthetic gas exchanger containing Chlorella pyrenoidosa (1% packed cell volume) was measured when various concentrations of carbon dioxide were present within the culture unit. The internal carbon dioxide concentrations were obtained by manipulating the entrance gas concentration and the flow rate. Carbon dioxide percentages were monitored by means of electrodes placed directly in the nutrient medium. The concentration of carbon dioxide in the nutrient medium which produced maximal photosynthesis was in the range of 1.5 to 2.5% by volume. Results were unaffected by either the level of carbon dioxide in the entrance gas or the rate of gas flow. Entrance gases containing 2% carbon dioxide flowing at 320 ml/min, 3% carbon dioxide at 135 ml/min, and 4% carbon dioxide at 55 ml/min yielded optimal carbon dioxide concentrations in the particular unit studied. By using carbon dioxide electrodes implanted directly in the gas exchanger to optimize the carbon dioxide concentration throughout the culture medium, it should be possible to design more efficient large-scale units.     

Leaf Conductance as Related to Xylem Water Potential and Carbon Dioxide Concentration in Sitka Spruce

Current year shoots of Sitka spruce [Picea sitchensis Bong. (Carr.)] from the forest canopy were equilibrated in a leaf chamber. The shoots were excised in air, and removed at differing times in order to establish a relationship between stomatal conductance and xylem water potential. The experiment was repeated at five ambient CO₂ concentrations. A second set of excised forest shoots, and shoots excised from 2-year- old nursery seedlings were allowed to evaporate freely in a controlled environment wind tunnel until a constant rate of transpiration was measured, to establish a relationship between cuticular conductance and xylem water potential. Cuticular conductance was estimated to be 0.012 cm s^-1 at high water potential and declined linearly to 0.007 cm s^-1 at ?3.5 MPa. The implication of this decline in the subsequent calculation of stomatal and mesophyll conductance is considered. Stomatal conductance remained constant at water potentials above ?1.4 MPa and was not affected by ambient carbon dioxide concentrations between 20 and 600 cm^-3. At lower water potentials, stomatal conductance declined and approached zero at ?2.5 to ?2.6 MPa. The results suggest that stomatal aperture is not controlled by either ambient or intercellular space carbon dioxide concentration, and that stomatal closure at low water potential is unlikely to be mediated by carbon dioxide.     

Temporal integration in nasal lateralization and nasal detection of carbon dioxide

Two experiments examined time/concentration trading for the detection of carbon dioxide, an irritant with little or no odor. Experiment 1 employed the nasal lateralization method: subjects attempted to determine which nostril received carbon dioxide and which received pure air when presented simultaneously. Experiment 2 employed a temporal, two-alternative, forced-choice, detection paradigm with monorhinal stimulation. In both experiments, stimulus duration was varied at a number of fixed concentrations to determine the shortest, detectable pulse. Under both conditions, threshold pulse duration decreased as stimulus concentration increased. Power functions with exponents of less than negative one described the data quite well: More than a twofold increase in duration was needed to compensate for a twofold decrease in concentration. Thus, for carbon dioxide, the nasal trigeminal system functions as an imperfect integrator at threshold-level.     

Experience‐ and age‐mediated oviposition behaviour in the yellow fever mosquito Stegomyia aegypti (=Aedes aegypti)

In repeated behaviours such as those of feeding and reproduction, past experiences can inform future behaviour. By altering their behaviour in response to environmental stimuli, insects in highly variable landscapes can tailor their behaviour to their particular environment. In particular, female mosquitoes may benefit from plasticity in their choice of egg‐laying site as these sites are often temporally variable and clustered. The opportunity to adapt egg‐laying behaviour to past experience also exists for mosquito populations as females typically lay eggs multiple times throughout their lives. Whether experience and age affect egg‐laying (or oviposition) behaviour in the mosquito Stegomyia aegypti (=Aedes aegypti) (Diptera: Culicidae) was assessed using a wind tunnel. Initially, gravid mosquitoes were provided with a cup containing either repellent or well water. After ovipositing in these cups, the mosquitoes were blood‐fed and introduced into a wind tunnel. In this wind tunnel, an oviposition cup containing repellent was placed in the immediate vicinity of the gravid mosquitoes. A cup containing well water was placed at the opposite end of the tunnel so that if the females flew across the chamber, they encountered the well water cup, in which they readily laid eggs. Mosquitoes previously exposed to repellent cups became significantly more likely to later lay eggs in repellent cups, suggesting that previous experience with suboptimal oviposition sites informs mosquitoes of the characteristics of nearby oviposition sites. These results provide further evidence that mosquitoes modify behaviour in response to environmental information and are demonstrated in a vector species in which behavioural plasticity may be ecologically and epidemiologically meaningful.     

Decreased hydraulic conductance in plants at elevated carbon dioxide

Previous work indicated that long-term exposure to elevated carbon dioxide levels can reduce hydraulic conductance in some species, but the basis of the response was not determined. In this study, hydraulic conductance was measured at concentrations of both 350 and 700 cm³ m^–3 carbon dioxide for plants grown at both concentrations, to determine the reversibility of the response. In Zea mays and Amaranthus hypochondriacus , exposure to the higher carbon dioxide concentration for several hours reduced whole-plant transpiration rate by 22–40%, without any consistent change in leaf water potential, indicating reversible reductions in hydraulic conductance at elevated carbon dioxide levels. Hydraulic conductance in these species grown at both carbon dioxide concentrations responded similarly to measurement concentration of carbon dioxide, indicating that the response was reversible. In Glycine max , which in earlier work had shown a long-term decrease in hydraulic conductance at elevated carbon dioxide levels, and in Abutilon theophrasti , no short-term changes in hydraulic conductance with measurement concentration of carbon dioxide were found, despite lower transpiration rates at elevated carbon dioxide. In G. max and Medicago sativa , growth at high dew-point temperature reduced transpiration rate and decreased hydraulic conductance. The results indicate that both reversible and irreversible decreases in hydraulic conductance can occur at elevated carbon dioxide concentrations, and that both could be responses to reduced transpiration rate, rather than to carbon dioxide concentration itself.     

Effects of carbon dioxide on ethylene production and action in intact sunflower plants

A continuous flow system was used to study the interactions between carbon dioxide and ethylene in intact sunflower (Helianthus annuus L.) plants. An increase in the concentration of carbon dioxide above the ambient level (0.033%) in the atmosphere surrounding the plants increased the rate of ethylene production, and a decrease in carbon dioxide concentration resulted in a decrease in the rate of ethylene production. The change in the rate of ethylene production was evident within the first 15 minutes of the carbon dioxide treatment. Continuous treatment with carbon dioxide was required to maintain increased rate of ethylene production. The rate of carbon dioxide fixation increased in response to high carbon dioxide treatment up to 1.0%. Further increases in carbon dioxide concentration had no additional effect on carbon dioxide fixation. Carbon dioxide concentrations higher than 0.11% induced hyponasty of the leaves whereas treatment with 1 microliter per liter ethylene induced epinasty of the leaves.     

Differential behaviour of Anopheles gambiae sensu stricto (Diptera: Culicidae) to human and cow odours in the laboratory

A dual port olfactometer was used to study the response of Anopheles gambiae Giles sensu stricto to odours of human and animal origin. Human odour consisted of human skin emanations collected on a nylon stocking, which was worn for 24 h. This was tested alone or together with 4.5% carbon dioxide, the concentration in human and cattle breath. Cattle odours consisted of cow skin emanations and/or carbon dioxide. Cow skin emanations were collected by tying a nylon stocking ('cow sock') around the hind leg of a cow for 12 h. Anopheles gambiae s.s. was consistently highly attracted by human odour, which is consistent with the high degree of anthropophily in this mosquito. Anopheles gambiae s.s. was not attracted by human or cattle equivalent volumes of carbon dioxide and this gas did not enhance the effect of human skin residues. Furthermore, A. gambiae s.s. showed a high degree of aversion to cow odour. When human odour and cow odour were tested together in the same port, mosquitoes were still highly attracted, indicating that whilst cattle odour may deter A. gambiae s.s., these mosquitoes can detect human odour in the presence of cattle odour. It was concluded that carbon dioxide plays a minor role in the host seeking behaviour of A. gambiae s.s., whilst host specific cues such as human skin residues play a major role and very effectively demonstrated anthropophilic behaviour in the laboratory.     

The response of the blood-sucking bug Triatoma infestans to carbon dioxide and other host odours

Behavioural responses of Triatoma infestans larvae to carbon dioxide and other odours of vertebrate origin were investigated in a locomotion compensator. T. infestans oriented towards airstreams enriched with carbon dioxide exhibiting a threshold response between 300 and 400 p.p.m. above the ambient CO(2) background. The accuracy of the oriented response to carbon dioxide improved with stimulus intensity. Remarkably, insects did not show any change in their sensitivity threshold to carbon dioxide with the starvation time. The attractiveness to carbon dioxide depended on the time of the day, i.e. these nocturnal bugs only oriented towards carbon dioxide-loaded airstreams during the first hours of the scotophase. L-lactic acid did not evoke oriented responses when it was presented as a single stimulus in a wide range of intensities. However, a marked synergism was evident when L-lactic acid was combined with a sub-threshold concentration of carbon dioxide. Under this condition, the threshold response to carbon dioxide decreased to 75-150 p.p.m. above ambient CO(2) background. The isomer D-lactic acid evoked no response, either alone or in combination with carbon dioxide. When insects were stimulated with 1-octen-3-ol a significant positive orientation was found. This response was not modified by the addition of carbon dioxide.     

An orbivirus of mosquitoes which induces CO2 sensitivity in mosquitoes and is lethal for rabbits.

An orbivirus, JKT-7400, isolated from Culex mosquitoes in Indonesia, replicated to a high titer and induced cytopathic effects in Aedes albopictus cell cultures. The virus produced lethal sensitivity to carbon dioxide in Culex and Aedes mosquitoes as well as in Drosophila melanogaster fruit flies but was not the agent of the hereditary sensitivity to carbon dioxide previously described for Culex quinquefasciatus. When injected intravenously in high doses, JKT-7400 virus was lethal for rabbits, apparently without replicating to a significant extent. It was not pathogenic for adult mice inoculated intravenously or for adult or suckling mice inoculated intracerebrally and intraperitoneally. Unlike an orbivirus isolated from Culex mosquitoes in China, JKT-7400 did not interfere with the replication of Japanese encephalitis virus in mosquitoes.     

Ammonia as an attractive component of host odour for the yellow fever mosquito, Aedes aegypti

Behavioural responses of Aedes aegypti mosquitoes to ammonia were investigated in a modified Y-tube olfactometer. Ammonia was attractive in concentrations from 17 ppb to 17 ppm in air when presented together with lactic acid. Aqueous solutions of ammonia salts in concentrations comparable to those found in human sweat also increased the attractiveness of lactic acid. The role of lactic acid as an essential synergist for ammonia became further apparent by the fact that ammonia alone or in combination with carbon dioxide was not effective, even though the synergistic effect of carbon dioxide and lactic acid was corroborated. An extract from human skin residues, which attracts approximately 80% of the tested mosquitoes, contains both lactic acid and ammonia. The combination of these compounds, however, attracts no more than 45%, indicating that other components on human skin also play a role in host finding. Preparative liquid chromatography of the skin extract yielded three behaviourally active fractions which work together synergistically. Fraction III contains lactic acid as the effective principle; the compositions of the other two have not been clarified yet. The attractiveness of fraction I was augmented considerably when ammonia was added, whereas the effect of fraction II was not influenced by ammonia. These results suggests that ammonia is part of the effective principle of fraction II and contributes to the attractive effect of host odours.     

Carbon dioxide and biearbonate in cell division

Summary The effects of carbon dioxide and of bicarbonate on cell division were studied on synchronized cells of the high-temperature green alga, Chlorella 7-11-05. After 7 hours of growth in nutrient medium in light, cells were centrifuged and resuspended in distilled water or in bicarbonate and placed in darkness. Atmospheric air, or a mixture of carbon dioxide and air, was bubbled through algal suspensions during the dark period. In distilled water cells readily divided in atmospheric air but not in 1% (2.6·10^-4 M) or in higher concentrations of carbon dioxide. The suspension of cells in bicarbonate counteracted the inhibitory action of carbon dioxide. A minimum molar concentration of bicarbonate necessary to counteract the inhibitory effect of carbon dioxide was found to be equal to the molar concentration of carbon dioxide in the suspending fluid. The highest concentration of carbon dioxide, the adverse effect of which could not be balanced by any concentration of bicarbonate, was found to be in the vicinity of 1.3·10^-2 M (50% CO₂ in air). Possible effects on cell division of the change in Ph and the implicated role of carbon dioxide in normal and neoplastic growth were discussed.     

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.     

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.     

The response of Culex quinquefasciatus (Diptera: culicidae) to traps baited with carbon dioxide, 1-octen-3-ol, acetone, butyric acid and human foot odour in Tanzania

The responses of Culex quinquefasciatus Say to traps baited with carbon dioxide, 1-octen-3-ol, acetone, butyric acid and human foot odour were studied in the field in Muheza, north-east Tanzania using Counterflow Geometry (CFG) and Centers for Disease Control (CDC) traps. It was found that significantly more C. quinquefasciatus responded to foot odour collected on nylon stockings than to clean nylon stockings (P < 0.05). Significantly more mosquitoes were caught in a CFG trap baited with carbon dioxide than in traps with either human foot odour, acetone or butyric acid. It was also found that in an outdoor situation a carbon dioxide baited CDC unlit trap collected over 12 times more C. quinquefasciatus than an unbaited CDC unlit trap and nine times more mosquitoes than CDC traps baited with 1-octen-3-ol alone (P < 0.05). The number of mosquitoes caught in a CDC trap baited with 1-octen-3-ol did not differ significantly from that of the unbaited CDC trap (P > 0.05). These results indicate that the Afrotropical C. quinquefasciatus respond significantly better to traps baited with carbon dioxide than to either octenol, acetone or butyric acid, and that human foot odour contains stimuli to which C. quinquefasciatus is attracted under field conditions.     

Effect of carbon dioxide concentration on the bioleaching of a pyrite-arsenopyrite ore concentrate

The effect of carbon dioxide concentration on the bacterial leaching of a pyrite-arsenopyrite ore concentrate was studied in continuous-flow reactors. Steady-state operation with two feed slurry densities, 6 wt% and 16 wt% solids, were tested for the effect of carbon dioxide concentration. Bacterial growth rates were estimated via the measurement of carbon dioxide consumption rates. Aqueous-phase carbon dioxide concentrations in excess of 10 mg/L were found to be inhibitory to bacterial growth. (c) 1993 John Wiley & Sons, Inc.