Comparative morphology of the amphibian opercularis system: I. General design features and functional interpretation.

The morphology of the opercularis system of anuran and caudate amphibians suggests that it acts to produce motion of the operculum that in turn produces fluid motion within the inner ear. The operculum and opercularis muscle form a lever system, with a narrow connection between the operculum and otic capsule acting as a fulcrum about which the operculum moves in response to forces applied via the muscle. The opercula of many species possess a muscular process on which the muscle inserts, thereby increasing the moment arm through which the muscle acts. The tonicity of the opercularis muscle allows tensile forces produced by substrate vibration or other mechanical energy applied to the forelimb to be effectively transmitted to the operculum; the elasticity of the connective tissue holding the operculum in place should act to return the operculum to its original position. The opercularis systems of frogs and non-plethodontid salamanders are similar structurally and functionally; that of plethodontid salamanders is structurally distinct but also functions as a lever system. Fluid motion produced by opercular motion could stimulate various end organs of the inner ear; the saccule, lagena, and amphibian papilla are in close approximation and wave energy could directly affect their otoconial or tectorial structures. In those anurans with a tympanic ear, the stapedial footplate and operculum articulate, but this articulation allows both to move independently. The stapes-tympanum complex and opercularis system therefore appear to be independent functional systems, and it is unlikely that the opercularis system modulates middle ear responsiveness. The general design of the opercularis system is consistent with a function in reception of substrate vibrations.

Electromyography of the opercularis muscle of Rana catesbeiana: an amphibian tonic muscle.

The opercularis muscle of Rana catesbeiana originates on the suprascapular cartilage of the shoulder girdle and inserts on the otic opercular element. It is part of the levator scapulae musculature and lies dorsomedial to the levator scapulae superior and inferior muscles. Bipolar electrode recordings from all three muscles show electrical activity linked to cyclical firing of the posterior intermandibularis muscle, an important ventilatory muscle. The opercularis muscle shows low amplitude, erratic signals when animals are submerged. Upon emergence of the snout region, the opercularis muscle shows rhythmic low amplitude activity at twice the rate of buccal pumping. Lung ventilation is synchronized with this rhythm and at ventilation the opercularis muscle shows higher amplitude activity. Upon submergence, opercularis activity again shows low level activity with no rhythmic pattern. Opercularis muscle activity has a major low frequency component (about 30 Hz) that probably corresponds to activity of tonic muscle fibers. Higher frequency signals (about 200-250 Hz) comparable to those of the levator scapulae muscles are also present and probably represent activity of phasic muscle fibers. Activity of the opercularis muscle is correlated with conditions in which aerial respiration is possible, and this pattern of activity supports an opercularis role in aerial hearing and/or detection of substrate vibrations. As far as we know, this is the first report of electromyographic analysis of a vertebrate tonic muscle.

Mechanisms of underwater hearing in larval and adult tiger salamanders Ambystoma tigrinum.

1. A standing wave tube apparatus was used to determine the biophysical basis of underwater hearing in Ambystoma tigrinum. 2. A. tigrinum responds to the pressure component of underwater sound, and the mouth cavity appears responsible for transduction of sound pressure. 3. Near-field displacements produced by pulsations of the air-filled mouth cavity apparently stimulate the inner ear. 4. Salamander head preparations with no air-filled mouth cavity respond to the particle motion component of underwater sound, but only at sound pressure levels 40 dB or more above levels producing clear pressure sensitivity in intact salamanders or head preparations including an air-filled mouth cavity.

Biophysics of underwater hearing in anuran amphibians.

A standing wave tube apparatus was used to determine the biophysical basis of underwater hearing sensitivity in 3 species of Rana and in Xenopus laevis. A speaker inside the base of a vertical, water-filled 3 m steel pipe produced standing waves. Pressure and particle motion were measured with a hydrophone and geophone respectively and were spatially 90 degrees out of phase along the length of the tube. Microphonic responses were recorded from the inner ear of frogs lowered through pressure and particle motion maxima and minima. The air-filled lungs of whole frogs produced distortions of the sound field. Preparations of heads with only an air-filled middle ear produced little distortion and showed clear pressure tracking at sound intensities 10-20 dB above hearing thresholds from 200-3000 Hz. Filling the middle ear with water decreased or abolished microphonic responses. Severing the stapes reduced responses except at certain frequencies below about 1000 Hz which varied with body size and likely represent resonant frequencies of the middle ear cavity. We conclude that the frog species examined respond to underwater sound pressure from about 200-3000 Hz with the middle ear cavity responsible for pressure transduction.

Structural correlates of function in the "opercularis" muscle of amphibians.

This study characterizes the fine structure of the "opercularis" muscles of selected frogs and salamanders (Genera: Hyla; Desmognathus; Ambystoma). The "opercularis" muscle originates on the shoulder girdle and inserts on the opercular plate in the fenestra ovalis of the otic capsule. Each of the three genera used exhibits one of the major gross dispositions of this muscle found in amphibians. In each case the "opercularis" muscle contains large numbers of tonic fibers: 80% in Hyla; 90% in Desmognathus; 45% in Ambystoma. These fibers correspond to the class-5 tonic fibers of Smith and Ovalle (1973). The remained of the fibers in the "opercularis" correspond to those in the class-3 "phasic" of Smith and Ovalle. The muscle from which the "opercularis" is derived (levator scapulae in Hyla, cucullaris in Desmognathus) is comprised of fibers which correspond to the class-2 phasic fibers of Smith and Ovalle. The fiber composition of the "opercularis" indicates that it is constructed to sustain contraction over long periods of time. This composition is supportive of the functional role in audition proposed for the muscle by Lombard and Straughan (1974). Evidence is presented that indicates that fiber size may be body size dependent and thus is an inappropriate criterion of fiber type identification.

Tongue evolution in the lungless salamanders, family plethodontidae. I. Introduction, theory and a general model of dynamics.

Plethodontid salamanders capture prey by projecting the tongue from the mouth. An analysis of theoretical mechanics of the hyobranchial skeleton is used to formulate a working hypothesis of tongue movements. Predictions that the skeletal elements of the tongue are included in the projectile and that the hyobranchial skeleton is folded during projection are central to the analysis. When decapitated in a particular way, salamanders project the tongue, and it is not retracted. When these heads are fixed and sectioned, examination confirms the predications. In turn, these observations are used to refine the working hypothesis and to generate a general model of tongue dynamics for plethodontids. Muscles performing the major roles of projection (subarcualis rectus I) and retraction (rectus cervicis profundus) are identified. The skeleton is folded passively along a morphological track having the form of a tractrix. Predictions concerning the shape of the track and the exact configuration of the folded skeleton are confirmed by study of sectioned material. The skeleton unfolds along the track during retraction and is spread into the resting state. The model developed herein will be used as a basis for predictions concerning selection patterns in the family and for analytical purposes in comparative and evolutionary studies.