Usage pattern of the complex masticatory muscles in the shingleback lizard, Trachydosaurus rugosus: A model for muscle placement.

This wide-ranging, omnivorous lizard of Australia has a very complex adductor muscle mass, with fibers differing in length by a factor of three and in insertion angle by 90 degrees. Stimulated muscles produce maximal moment with the mouth nearly fully open. The opening mechanism appears to involve only simple rotation and no translation of the mandible. EMGs indicate that the entire mass is activated equivalently in crushing and there are no temporal subdivisions, for instance, matching activity to angle of opening. During crushing of hard objects, the chin is brought into contact with the ground so that the subvertebral muscles may aid buccal closure. The lizards also activate the muscles in a pulsatile staircase effect leading to an unfused tetanus that generates forces several times the twitch level. Application in parallel of a maximum number of sarcomeres to the crushing bite appears to be the major design characteristic. Hence, this species offers an ideal case for analysis of the effects of different sarcomere placements on the simple movement generated. For the primary adductor muscles, the angles of fiber insertion relative to the lines connecting each insertion with the jaw joint are equivalent; this relation persists as the mouth opens. Also, fiber lenghts are proportional to the distance between jaw joint and site of insertion so that each sarcomere contributes equally to the movement generated. Complex tendons provide additional space for muscle placement. Some of these also extend beyond the bony attachment sites, producing tendinous "coronoid processes." The fibers of laterally and ventrally placed muscles are short relative to the length of the entire muscle, insert at relatively short moment arms, and undergo short excursion during opening; however, there are many such fibers. Also, muscles with a low incident angle are crossed; they apparently protect the jaw joint from horizontal (disarticulating) forces.