The principles of classification and a classification of mammals. Bulletin of the AMNH ; v. 85

Tongues, tentacles and trunks: the biomechanics of movement in muscular‐hydrostats

A century has passed since the publication of William James's Principles of Psychology, yet most of the questions James raised about the relation between physiological events and molar psychological or behavioral processes, such as emotion, remain unanswered. The sluggish progress in capitalizing on physiological signals to address general psychological questions is due in part to shortcomings in the quantification of physiological sig- nals in humans and, perhaps more important, to the way in which investigators have been thinking about the re- lation between physiological signals and psychological operations. In this article, we illustrate these points, and we provide a conceptual framework to foster research and analysis of psychological phenomena based on physiolog- ical signals. Psychological operations and physiological responses are defined in terms of configural and temporal properties, and psychophysiological relations are concep- tualized in terms of their specificity (e.g., one-to-one versus many-to-one) and their generality (e.g., situation or person specific versus cross-situational and pancultural). This model yields four classes of psychophysiological relations: (a) outcomes, (b) concomitants, (c) markers, and (d) in- variants. Finally, the model specifies how to determine whether a psychophysiological relation is an outcome, concomitant, marker, or invariant, and it describes im- portant limitations in inferences of psychological signifi- cance based on physiological signals when dealing with each.

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Inferring PSychological Significance From Physiological Signals

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

We assembled a dataset of 192 carefully selected species—51 extinct and 141 extant—and 976 apomorphies distributed among 610 phenotypic characters to investigate the phylogeny of Squamata (“lizards,” including snakes and amphisbaenians). These data enabled us to infer a tree much like those derived from previous morphological analyses, but with better support for some key clades. There are also several novel elements, some of which pose striking departures from traditional ideas about lizard evolution (e.g., that mosasaurs and polyglyphanodontians are on the scleroglossan stem, rather than parts of the crown, and related to varanoids and teiids, respectively). Long-bodied, limb-reduced, “snake-like” fossorial lizards—most notably dibamids, amphisbaenians and snakes—have been and continue to be the chief source of character conflict in squamate morphological phylogenetics. Carnivorous lizards (especially snakes, mosasaurs and varanoids) have proven a close second. Genetic data, presumably less bur...

Assembling the Squamate Tree of Life: Perspectives from the Phenotype and the Fossil Record

The masticatory pattern of Sphenodon punctatus, the sole remaining rhynchocephalian, now restricted to islands off the coast of New Zealand, has been analyzed by detailed anatomy, cinematography, cinefluoroscopy, and electromyography. Food reduction consists of a closing, crushing bite followed by a propalineal sliding of the dentary row between the maxillary and palatine ones. The large, fleshy tongue can be protruded to pick up small prey, and also plays a major role in prey manipulation. The rotational closing movement of the jaw, supporting the basic crushing movement, is induced by the main adductor musculature. It is followed by a propalineal anterior displacement relying heavily on the action of the M. pterygoideus. The fiber lengths of the several muscles reflect the extent of shortening. The most obvious modification appears in the M. pterygoideus, which contains a central slip of pinnately arranged short fibers that act a period different from that of the rest of the muscle; their action increases the power during the terminal portion of the propalineal phase. This also allows the animal to use its short teeth in an effective shearing bite that cuts fragments off large prey. The action of single cusped dentary teeth acting between the maxillary and palatine tooth rows provides a translational crushing-cutting action that may be an analog of the mammalian molar pattern. However, this strictly fore-aft slide does not incorporate capacity for later development of lateral movement.

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Mastication in the tuatara, Sphenodon punctatus (reptilia: Rhynchocephalia): Structure and activity of the motor system.

Mastication has been studied by cinematography with synchronized electromyography (computer quantified and analyzed), while unanesthetized, freely feeding cats (Felis catus) were reducing equivalent-sized chunks of raw and cookded beef and cooked chicken. Cats reduce food on one side at a time, and their chewing cycles show horizontal and anteroposterior deflections. Food objects are shifted from side to side by lateral jerks of the head and movements of the tongue. During the opening phase, the lower jaw is rotated relatively straight downward, and the digastric muscles are active in bilateral symmetry. Near the end of opening, the head jerks upward, both zygomaticomandibulares start to fire, and opening acceleration of the mandible decreases. Closing starts with horizontal displacement of the mandibular canines toward the working side, accompanied by asymmetrical activities from the working side deep temporalis and the balancing side medial pterygoid, as well as a downward jerk of the head. As closing proceeds the mandibular canines remain near the working side and the working side zygomaticomandibularis and deep masseter are very active. Near the end of closing, the mandibular canine on the working side moves toward the midline, and adductors, digastrics, and lateral pterygoids of both sides are active. The adductors of the working side are generally more active than those of the balancing side. During a reduction sequence, the number and shape of the masticatory cycles, as well as movements of the head, during a reduction sequence are affected significantly by food type. As reduction proceeds, the duration of bite and the muscular activity (as characterized by number and amplitude of spikes) change significantly among muscles of the working and balancing sides. The adductors of the working side are generally most active when cats chew raw beef, less for cooked beef, and least for cooked chicken. In general, the adductor activity reflect food consistency, whereas that of the digastrics and lateral pterygoids reflects more the vertical and lateral displacements of the mandible. Statistical analysis documents that the methods of electrode insertion and test give repeatable results for particular sites in different animals. Thus, it should be possible to compare these results with those produced while other mammals are masticating.

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Quantitative assay of electromyograms during mastication in domestic cats (Felis catus).

Bufo marinus catches its prey by stiffening the intrinsic muscles of the tongue, rapidly flipping the tongue out of the mouth. High-speed cine- matography synchronized with computer-analyzed electromyograms (EMGs) shows that during the flip the tongue is supported by the M. genioglossus medialis and that this muscle stiffens into a rod when stimulated. Coincident stiffening of the transversely arranged M. genioglossus basalis provides a wedge under the anterior tip of this rod. Stiffening of the M. submentalis depresses the mandibular symphysis and brings the dentary tips together. The M. submentalis also acts on the wedge of the basalis to raise and rotate the rigid rod of the medialis over the symphysial attachment. The tip of this lingual rod carries along the pad and soft tissues of the tongue. The lingual pad, positioned in the posterodorsal portion of the resting tongue, rotates during eversion so that its dorsal surface impacts onto the prey object. Retraction starts by contraction of the elongate, parallel fibers of the M. hyoglossus; this retracts the medial sulcus of the pad and holds the prey by a suction cup-like effect. The extensibility of the buccal membranes allows the pad to be retracted first; it reaches the posterior portion of the buccal cavity before the still-rigid, backward rotating M. genioglossus has reached the level of the symphysis. Protraction of the hyoid facilitates the extension of the M. hyoglossus. The M. sternohyoideus only retracts the hyoid and stabilizes it when the tongue starts to pull posteriorly; it does not assist tongue protrusion. The Mm. petrohyoideus and omohyoideus show only incidental activity, and the M. depressor mandibulae participates in mouth opening but is not otherwise involved in the flip. Previous hypotheses of the flipping mechanism are reviewed and evaluated. It is well known that many frogs catch their prey with a rapid flip of the tongue, that the anterior end of this tongue is fixed and the posterior free in most species, and that the emerging tongue rotates over the mandibular symphysis. The system is curious in that the tongue of most anurans is anchored anterior to the hyoid and lacks a hard-tissue skeleton that could induce and support this rapid pro- trusion. The mechanism generating rotation of the tongue, as well as the manner in which the tongue grips the prey, were long accorded minimal attention, although the retrolingual surface of the frog tongue has been used in the resolution of some basic questions in muscle physiology (Biesladecki and Herzig, 1859; Fischl and Kahn, 1928; Gelfan, 1930; Pratt and Reid, 1930; Fulton and Lutz, 1940). Such

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Functional morphology of lingual protrusion in marine toads (Bufo marinus)

Simultaneous cine and electromyographic records of freely feeding, unanesthetized golden hamsters show that their motion and muscular activity during mastication differ from those of albino rats (Weijs, '75). Rats show only propalinal motion while hamsters show lateral translation as well. The masticatory muscles of hamsters and rats are generally similar, but their molar dentitions differ. The interlocking molar cusps of hamsters restrict propalinal protrusion and retrusion when the molars are in occlusion; however, hamsters readily unlock occlusion by a twisting movement in the horizontal plane. Rats may perform propalinal movements even with the teeth in occlusion.



In mastication the hamstery's jaw moves laterally as well as vertically and anteroposteriorly. Chewing orbits typically reverse after one to three orbits. Reversal begins at the start of the upstroke and involves a lateral shift in the opposite direction with the mouth closed.



Electromyograms show that symmetric and asymmetric activities of closing protrusive and closing retrusive muscles produce a unilateral force couple on both sides. (This couple accompanies a midline closing stroke.) When the mouth is closed, unilateral activity of closing retrusors and closing protrusors also induces lateral translation. A bilateral force couple pits the retrusors of one side against the protrusors on the opposite side. Simultaneous with lateral excursion to the opposite side of midline and the action of these closing muscles, the anterior digastric and lateral pterygoid muscles of one side fire asymmetrically.



The mandible moves downward coincidently with bilateral activity of the digastrics and lateral pterygoids. As the jaw opens further, activity differences of the lateral pterygoids accompany a shift of the mandible toward midline. At the end of the downstroke, all masticatory muscles studied are silent. The jaw returns to midline when the adductors fire asymmetrically at the start of closing.



Trituration appears to coincide with an initial simple protrusion, which is subsequently accompanied by lateral translation. Different food types are reduced by distint chewing patterns with the differences clearest when the teeth are near occlusion. During gnawing the lateral pterygoids and digastrics fire longer, and the closing muscles fire less strongly. Chewing patterns in golden hamsters appear more generalized than those of rats; the differences may be directly associated with the ability of hamsters to store food in their cheek pouches.

Feeding in golden hamsters, Mesocricetus auratus.

Two conflicting hypotheses purport to explain the mechanism generating the lingual flip in frogs. The first suggests that the intrinsic tongue muscles are stiffened, rotate over the symphysis and catapult the soft tissues; the second suggests that the hyoid suddenly moves forward and transfers its momentum to propel the tongue. High-speed cinematography and synchronized electromyography show that the tongue is rotated over the symphysis by a complex of rods formed from stiffened intrinsic tongue muscles. As the flip occurs even when the hyoid is immobilized, the hyoid momentum hypothesis does not apply. The tongue is then propelled by sets of fibers locked into connective tissues. With activation, these become rigid rods that form a muscular ballista.

https://science.sciencemag.org/content/sci/216/4552/1335.full.pdf

Gerard C. Gorniak

Papers

Mastication in the tuatara, Sphenodon punctatus (reptilia: Rhynchocephalia): Structure and activity of the motor system.

Quantitative assay of electromyograms during mastication in domestic cats (Felis catus).

Functional morphology of lingual protrusion in marine toads (Bufo marinus)

Feeding in golden hamsters, Mesocricetus auratus.

How Does the Toad Flip Its Tongue? Test of Two Hypotheses