Insect Wing - Physiology - Indirect Flight

Indirect Flight

Other than the two orders with direct flight muscles, all other living winged insects fly using a different mechanism, involving indirect flight muscles. This mechanism evolved once, and is the defining feature (synapomorphy) for the infraclass Neoptera; it corresponds, probably not coincidentally, with the appearance of a wing-folding mechanism, which allows Neopteran insects to fold the wings back over the abdomen when at rest (though this ability has been lost secondarily in some groups, such as all butterflies).

In the higher groups with two functional pairs of wings, both pairs are linked together mechanically in various ways, and function as a single wing, although this is not true in the more primitive groups. What all Neoptera share, however, is the way the muscles in the thorax work: these muscles, rather than attaching to the wings, attach to the thorax and deform it; since the wings are extensions of the thoracic exoskeleton, the deformations of the thorax cause the wings to move as well. A set of dorsal longitudinal muscles compress the thorax from front to back, causing the dorsal surface of the thorax (notum) to bow upward, making the wings flip down. A set of tergosternal muscles pull the notum downward again, causing the wings to flip upward. In a few groups, the downstroke is accomplished solely through the elastic recoil of the thorax when the tergosternal muscles are relaxed. Several small sclerites at the wing base have other, separate, muscles attached and these are used for fine control of the wing base in such a way as to allow various adjustments in the tilt and amplitude of the wing beats. One of the final refinements that has appeared in some of the higher Neoptera (Coleoptera, Diptera, and Hymenoptera) is a type of muscular or neural control system whereby a single nerve impulse causes a muscle fiber to contract multiple times; this allows the frequency of wing beats to exceed the rate at which the nervous system can send impulses. This specialized form of muscle is termed asynchronous flight muscle. The overall effect is that many higher Neoptera can hover, fly backward, and perform other feats involving a degree of fine control that insects with direct flight muscles cannot achieve.