Modulation of intra-oral processing in mammals and lepidosaurs

The mammalian masticatory apparatus is distinguished from theintra-oral processing systems of other amniotes by a numberof morphological and functional features, including transversemovements of the teeth during the power stroke, precise occlusion,suspension of the teeth in the socket by a periodontal ligament,diphyodonty (reduction to two generations of teeth), a hardpalate, and the presence of a single bone (the dentary) in thelower jaw which articulates with the skull at the temporomandibularjaw joint. The evolution of these features is commonly arguedto have improved the efficiency of food processing in the oralcavity. The present aricle highlights the existence in mammalsof the fusimotor system and afferent fibers from the periodontalligament through which the CNS modulates the responses by themuscle spindles. Published data suggest that the fusimotor systemand the periodontal afferents are important components in feed-forward(or anticipatory) control of chewing behavior. We hypothesizethat this feed-forward control is used to maintain relativelyconstant cycle lengths in mammals in the face of intra-sequenceand inter-sequence variation in material properties of the food,and that this enables them to maintain a higher average chewingfrequency than that of lizards. These predictions were evaluatedusing data on mean cycle length and its variance from the literatureand from our own files. On average, mammals have less variablecycle lengths than do lizards and shorter cycle lengths thando lizards of similar size. We hypothesize that by decreasingvariance in cycle length, presumably close to the natural frequencyof their feeding systems, mammals minimize energy expenditureduring chewing, allowing them to chew for longer, thereby maintainingthe high rates of food intake required for their high metabolicrates.