Thermoregulation in Endothermic Insects

On the basis of body weight, most flying insects have higher rates of metabolism, and hence heat production, than other animals. However, rapid rates of cooling because of small body size in most cases precludes appreciable endothermy. The body temperature of small flies in flight is probably close to ambient temperature, and that of flying butterflies and locusts is 5 degrees to 10 degrees C above ambient temperature. Many moths and bumblebees are insulated with scales and hair, and their metabolism during flight can cause the temperature of the flight muscles to increase 20 degrees to 30 degrees C above ambient temperature. Curiously, those insects which (because of size, insulation) retain the most heat in the thorax during flight, also require the highest muscle temperature in order to maintain sufficient power output to continue flight. The minimum muscle temperature for flight varies widely between different species, while the maximum temperature varies over the relatively narrow range of 40 degrees to 45 degrees C. As a consequence, those insects that necessarily generate high muscle temperatures during flight must maintain their thoracic temperature within a relatively narrow range during flight. Active heat loss from the thorax to the abdomen prevents overheating of the flight motor and allows some large moths to be active over a wide range of ambient temperatures. Bumblebees similarly transfer heat from the flight musculature into the abdomen while incubating their brood by abdominal contact. Many of the larger insects would remain grounded if they did not actively increase the temperature of their flight muscles prior to flight. Male tettigoniid grasshoppers elevate their thoracic temperature prior to singing. In addition, some of the social Hymenoptera activate the "flight" muscles specifically to produce heat not only prior to flight but also during nest temperature regulation. During this "shivering" the "flight" muscles are often activated in patterns different from those during flight. The muscles contract primarily against each other rather than on the wings. However, the rate of heat production during shivering and flight is primarily a function of the action potential frequency rather than of the patterns of activation. Thermoregulation is a key factor in the energetics of foraging of some of the flower-visiting insects. The higher their muscle temperature the more flowers they can visit per unit time. When food supplies are ample, bees may invest relatively large amounts of energy for thermoregulation. While shivering to maintain high body temperatures during the short intervals they are perched on flowers (as well as while in the nest), bumblebees often expend energy at rates similar to the rates of energy expenditure in flight. Unlike vertebrates, which usually regulate their body temperature at specific set points, the body temperature of insects is labile. It often appears to be maintained near the lower temperature at which the muscles are able to perform the function at hand. The insects' thermal adaptations may not differ as much from those of vertebrates as previously supposed when size, anatomy, and energy requirements are taken into account.