SUMMARYGas exchange dynamics in insects is of fundamental importance to understanding evolved variation in breathing patterns, such as discontinuous gas exchange cycles (DGCs). Most insects do not rely solely on diffusion for the exchange of respiratory gases but may also make use of respiratory movements (active ventilation) to supplement gas exchange at rest. However, their temporal dynamics have not been widely investigated. Here, intratracheal pressure, V CO2 and body movements of the desert locust Schistocerca gregaria were measured simultaneously during the DGC and revealed several important aspects of gas exchange dynamics. First, S. gregaria employs two different ventilatory strategies, one involving dorso-ventral contractions and the other longitudinal telescoping movements. Second, although a true spiracular closed (C)-phase of the DGC could be identified by means of subatmospheric intratracheal pressure recordings, some CO 2 continued to be released. Third, strong pumping actions do not necessarily lead to CO 2 release and could be used to ensure mixing of gases in the closed tracheal system, or enhance water vapour reabsorption into the haemolymph from fluid-filled tracheole tips by increasing the hydrostatic pressure or forcing fluid into the haemocoel. Finally, this work showed that the C-phase of the DGC can occur at any pressure. These results provide further insights into the mechanistic basis of insect gas exchange.

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The evolutionary origin and maintenance of discontinuous gas exchange (DGE) in tracheate arthropods are poorly understood and highly controversial. We investigated prioritization of abiotic factors in the gas exchange control cascade by examining oxygen, water and haemolymph pH regulation in the grasshopper Paracinema tricolor. Using a full-factorial design, grasshoppers were acclimated to hypoxic or hyperoxic (5% O 2 , 40% O 2 ) gas conditions, or dehydrated or hydrated, whereafter their CO 2 release was measured under a range of O 2 and relative humidity (RH) conditions (5%, 21%, 40% O 2 and 5%, 60%, 90% RH). DGE was significantly less common in grasshoppers acclimated to dehydrating conditions compared with the other acclimations (hypoxia, 98%; hyperoxia, 100%; hydrated, 100%; dehydrated, 67%). Acclimation to dehydrating conditions resulted in a significant decrease in haemolymph pH from 7.0±0.3 to 6.6±0.1 (mean ± s.d., P=0.018) and also significantly increased the open (O)-phase duration under 5% O 2 treatment conditions (5% O 2 , 44.1±29.3 min; 40% O 2 , 15.8±8.0 min; 5% RH, 17.8±1.3 min; 60% RH, 24.0±9.7 min; 90% RH, 20.6±8.9 min). The observed acidosis could potentially explain the extension of the O-phase under low RH conditions, when it would perhaps seem more useful to reduce the O-phase to lower respiratory water loss. The results confirm that DGE occurrence and modulation are affected by multiple abiotic factors. A hierarchical framework for abiotic factors influencing DGE is proposed in which the following stressors are prioritized in decreasing order of importance: oxygen supply, CO 2 excretion and pH modulation, oxidative damage protection and water savings.

SUMMARYThe importance of metabolic rate and/or spiracle modulation for saving respiratory water is contentious. One major explanation for gas exchange pattern variation in terrestrial insects is to effect a respiratory water loss (RWL) saving. To test this, we measured the rates of CO 2 and H 2 O release (V CO2 and V H2O , respectively) in a previously unstudied, mesic cockroach, Aptera fusca, and compared gas exchange and water loss parameters among the major gas exchange patterns (continuous, cyclic, discontinuous gas exchange) at a range of temperatures. Mean V CO2 , V H2O and V H2O per unit V CO2 did not differ among the gas exchange patterns at all temperatures (P>0.09). There was no significant association between temperature and gas exchange pattern type (P=0.63). Percentage of RWL (relative to total water loss) was typically low (9.79±1.84%) and did not differ significantly among gas exchange patterns at 15°C (P=0.26). The method of estimation had a large impact on the percentage of RWL, and of the three techniques investigated (traditional, regression and hyperoxic switch), the traditional method generally performed best. In many respects, A. fusca has typical gas exchange for what might be expected from other insects studied to date (e.g. V CO2 , V H2O , RWL and cuticular water loss). However, we found for A. fusca that V H2O expressed as a function of metabolic rate was significantly higher than the expected consensus relationship for insects, suggesting it is under considerable pressure to save water. Despite this, we found no consistent evidence supporting the conclusion that transitions in pattern type yield reductions in RWL in this mesic cockroach. Supplementary material available online at

·CO2 ratio) showed a marked increase in grouped caterpillars, particularly in larger groups. Other benefits of aggregation (e.g. reduced predation or increased growth rates) likely outweigh these potential costs, because individuals of E. capensis aggregate voluntarily despite no obvious energetic or hygric advantage, and other potentially confounding group effects (e.g. increased thermoregulatory advantage or whole-animal activity) are inconsequential. The results of this study provide an important exception to physiological studies reporting enhanced energy or water conservation in animal groups. Supplementary material available online at

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