Allen G. Gibbs

SUMMARY: Insects breathe through valved openings (spiracles) in their cuticle. Many insects open and close their spiracles in a cyclic pattern (discontinuous gas-exchange cycles, or DGC). These cycles were observed over half a century ago, their hypothesized function being to minimize loss of water from the tracheal system. However, numerous recent studies have found that respiration accounts for a small fraction of total water loss, and that insects stop performing DGC at times when this pattern would be most useful. Thus, the importance of cyclic gas exchange for water conservation has been challenged. The leading alternative is the chthonic hypothesis, which proposes that DGC originated in insects from hypercapnic (high CO2) environments(e.g. burrows) to aid in release of carbon dioxide. By keeping the spiracles closed, insects would concentrate CO2 and increase the gradient for outward diffusion of CO2. CO2 would be released rapidly when the spiracles opened, and respiratory water loss would be reduced. The chthonic hypothesis therefore predicts that DGC minimizes the ratio of respiratory water loss to CO2 release relative to other modes of gas exchange. We tested the chthonic hypothesis by simultaneously measuring water loss and CO2 release in reproductive females (queens) of the seed-harvester ant Pogonomyrmex barbatus, a burrowing species from North American deserts. Queens used one of three patterns of gas exchange,discontinuous, cyclic and continuous. We resolved the problem of separating cuticular transpiration and respiratory water loss for individuals that used continuous gas exchange by developing a regression method that can be used across all patterns of gas exchange. The ratio of respiratory water loss to CO2 release did not differ among ants using different patterns of gas exchange, in contrast to the expectation of the chthonic hypothesis. Metabolic rate, however, varied with gas-exchange pattern, and was lowest for individuals that used discontinuous gas exchange, intermediate for individuals using cyclic gas exchange, and highest for individuals using continuous gas exchange.

SUMMARY Flies of the genus Drosophila inhabit a wide range of habitats, from the tropics to deserts to boreal forests. The primary physiological mechanism allowing Drosophila and other insects to survive in arid habitats is a reduction in rates of water loss. To understand mechanisms of water retention in greater detail, we investigated the three main routes by which Drosophila lose water: excretion, cuticular transpiration and respiratory loss through the spiracles. Excretory losses comprised Drosophila were less active, and female cactophiles had lower metabolic rates than female mesic species of the same size. They were also more likely to exhibit a pattern of cyclic CO₂ release that may help to conserve water. We conclude that lower overall rates of water loss are achieved primarily by reduction of respiratory losses.

SUMMARY: Fruit flies of the genus Drosophila have independently invaded deserts around the world on numerous occasions. To understand the physiological mechanisms allowing these small organisms to survive and thrive in arid environments, we performed a phylogenetic analysis of water balance in Drosophila species from different habitats. Desert (cactophilic) species were more resistant to desiccation than mesic ones. This resistance could be accomplished in three ways: by increasing the amount of water in the body, by reducing rates of water loss or by tolerating the loss of a greater percentage of body water (dehydration tolerance). Cactophilic Drosophila lost water less rapidly and appeared to be more tolerant of low water content, although males actually contained less water than their mesic congeners. However, when the phylogenetic relationships between the species were taken into account, greater dehydration tolerance was not correlated with increased desiccation resistance. Therefore, only one of the three expected adaptive mechanisms, lower rates of water loss, has actually evolved in desert Drosophila, and the other apparently adaptive difference between arid and mesic species (increased dehydration tolerance) instead reflects phylogenetic history.

Age-related declines in physiological performance have been documented in a wide variety of organisms. However, it is unknown whether related species age in a similar manner or whether physiological differences associated with aging differ widely among species. Previous work has shown that the desiccation resistance of Drosophila melanogaster decreases rapidly with age. Other Drosophila species may have delayed reproductive maturity or may inhabit arid habitats, so that age-related changes in water balance may be a significant physiological constraint on their ecological success. We may, therefore, predict that physiological systems involved in water balance will deteriorate with age at a reduced rate in these species. We quantified several components of water budgets for 14 species of Drosophila, including both desert and mesic representatives differing in their age of maturity. Desiccation resistance decreased with age in all but one species studied, primarily because of increased rates of water loss. However, there was no significant relationship between the rate of aging, as indicated by the rate of increase in water-loss rates as flies aged, and either habitat or age of maturity.

Wax esters are major constituents of the surface lipids in many terrestrial arthropods, but their study is complicated by their diversity. We developed a procedure for quantifying isomers in mixtures of straight-chain saturated and unsaturated wax esters having the same molecular weights, using single-ion monitoring of the total ion current data from gas chromatography-mass spectrometry. We examined the biological consequences of structural differences by measuring the melting temperatures, T _m , of >60 synthetic wax esters, containing 26-48 carbon atoms. Compounds containing saturated alcohol and acid moieties melted at 38-73°C. The main factor affecting T _m was the total chain length of the wax ester, but the placement of the ester bond also affected T _m . Insertion of a double bond into either the alcohol or acid moiety decreased T _m by ~30°C. Simple mixtures of wax esters with n -alkanes melted several °C lower than predicted from the melting points of the component lipids. Our results indicate that the wax esters of primary alcohols that are most typically found on the cuticle of terrestrial arthropods occur in a solid state under physiological conditions, thereby conferring greater waterproofing. Wax esters of secondary alcohols, which occur on melanopline grasshoppers, melted >60°C below primary esters of the same molecular weight and reduced T _m of the total surface lipids to environmental values.