This review describes the patterns of mandibular movements that make up the whole sequence from ingestion to swallowing food, including the basic types of cycles and their phases. The roles of epithelial, periodontal, articular, and muscular receptors in the control of the movements are discussed. This is followed by a summary of our knowledge of the brain stem neurons that generate the basic pattern of mastication. It is suggested that the production of the rhythm, and of the opener and closer motoneuron bursts, are independent processes that are carried out by different groups of cells. After commenting on the relevent properties of the trigeminal and hypoglossal motoneurons, and of internuerons on the cortico-bulbar and reflex pathways, the way in which the pattern generating neurons modify sensory feedback is discussed.

Mastication and its Control by the Brain Stem

Developmental and functional factors interact in complex ways reflected in evolutionary patterns. The paper addresses problems of coronal morphogenesis, amelogenesis, food comminution and digestion, mastication, tooth eruption and wear, in order to identify functional interrelations and developmental constraints in the evolution of cheek tooth morphology. A general mechanism explaining differen­ tial (levelling) wear of teeth in a row is proposed. The paradox of predicted positive allometry but observed isometry of tooth size to body size is resolved by inclusion of the time dimension in the equation. The reasons for the commonness of isometric scal­ ing in animals are discussed. Relative width of antagonists is found to discriminate be­ tween functional categories in terms of the dynamics of the power stroke in mastica­ tion, and can be used to interpret the functional meaning of major patterns of dental morphological radiation. Enamel structure at the level of arrangement of Hunter­ Schreger bands is interpreted in terms of different mechanical requirements (resis­ tance to crack propagation) in major morphological and functional categories. It is concluded that many aptive features probably or certainly did not arise for their cur­ rent functions, but are one-time constraints which have become incorporated into functional systems ( exaptations rather than adaptations).

Ungulate cheek teeth : developmental, functional, and evolutionary interrelations

Evolving climates and mammal faunas in Cenozoic South America

Fifteen variables, selected primarily to reflect functionally significant aspects of cranial morphology, were measured on one skull each of 62 species of modern carnivores, including viverrids, canids, mustelids and felids. To allow comparisons between species of different sizes without the potentially confounding effects of allometric shape changes, the measurements were transformed to dimensionless variables, based on the residuals from allometric equations. Fourteen out of 15 of the transformed variables distinguish one or more of the four family groups and the rotated first two axes of a principal components analysis distinguish all four families from each other. The following functional hypotheses are proposed: mustelids and felids have the most powerful bites and canids the weakest among the four family groups studied; mustelids and, to a lesser degree, felids have more powerful neck musculature than do canids and viverrids; and visual abilities are best developed among felids and least developed among mustelids. The first two functional hypotheses suggest possible differences in killing behaviour, which are supported by a preliminary survey of the literature on such behaviour. Allometric analysis of the 15 cranial measures shows that the neurocranial components scale with negative allometry, while most of the other measures scale approximately isometrically.

Evolution of skull shape in carnivores: 1. Representative modern carnivores

The mammalian mandible, and in particular the human mandible, is generally thought to function as a lever during biting. This notion, however, has not gone unchallenged. Various workers have suggested that the mandible does not function as a lever, and they base this proposition on essentially two assertions: (1) the resultant of the forces produced by the masticatory muscles always passes through the bite point; (2) the condylar neck and/or the temporomandibular joint is unsuited to withstand reaction forces during biting. A review of the electromyographic data and of the properties of the tissues of the temporomandibular joint do not support the non-lever hypothesis of mandibular function. In addition, an analysis of the strength of the condylar neck demonstrates that this structure is strong enough to withstand the expected reaction force during lever action. Ordinarily the human mandible and the forces that act upon it are analyzed solely in the lateral projection. Moments are then usually analyzed about the mandibular condyle; however, some workers have advocated taking moments about other points, e.g., the instantaneous center of rotation. Obviously it makes no difference as to what point is chosen since the moments about any point during equilibrium conditions are equal to zero. It is also useful to analyze the forces acting on the mandible in the frontal projection, particularly during unilateral biting. The electromyographic data suggest that during powerful unilateral molar biting the resultant adductor muscle force is passing between the bite point and the balancing (non-biting side) condyle. Therefore, in order for this system to be in equilibrium there must be a reaction force acting on the balancing condyle. This suggests that reaction forces are larger on the balancing side than on the working side, and possibly explains why individuals with a painful temporomandibular joint usually prefer to bite on the side of the diseased joint.

Wd Turnbull

Papers

Mammalian masticatory apparatus