Respiration and the generation of rhythmic outputs in insects.

In insects gas exchange may be: 1) entirely passive, when metabolic rate is low; 2) enhanced automatically by muscle contractions that produce movements, e.g., wing movements in flight; or 3) produced by ventilatory movements, particularly of the abdomen. In terrestrial insects such as locusts and cockroaches ventilatory movements are governed by a dominant oscillator in the metathoracic or anterior abdominal ganglion. The dominant oscillator overrides local oscillators in the abdominal ganglia and thus sets the rhythm for the entire abdomen, and it also controls spiracle opening and closing in several thoracic and abdominal segments. This ventilatory control mechanism appears to be different from that generating metachronal rhythms such as occur in the ventilatory and locomotory movements of aquatic arthropods. There are now several examples of rhythms, both ventilatory and locomotory, that can be generated by the central nervous system in the absence of phasic sensory feedback, but the mechanism of rhythm production is not known. Studies of ganglionic output suggest that neuronal oscillators can produce a range of frequencies and that some oscillators may be employed in more than one function or behavior. The mechanisms by which central oscillators are coupled to the output motorneurons are also not known; large phase changes suggest that in some cases different coupling interneurons are active. Intracellular recordings from identified neurons have begun to clarify the important roles of interneurons in the production of motor patterns.