Role of the opercularis muscle in seismic sensitivity in the bullfrog Rana catesbeiana

The inner ear of anuran amphibians appears to be exceptionally sensitive to substrate vibration. The opercularis system, consisting of an opercularis muscle running from the shoulder girdle to a movable, cartilaginous operculum lying next to the inner ear, has been hypothesized to be involved in driving these seismic responses. Removal of the opercularis muscle of adult bullfrogs, Rana catesbeiana, caused clear decreases in microphonic responses of the inner ear to vibrations from 20-250 Hz and 0.05-5.0 cm/sec2 accelerations. Degree of decrease in responsiveness was variable between individuals and between different frequencies of stimulation, ranging up to 90% reduction at certain frequencies and in certain specimens. Decreases were most marked at lower frequencies below about 50 Hz. Additional removal of the levator scapulae superior muscle, which runs alongside the opercularis muscle from the shoulder girdle to ventrolateral portions of the otic capsule, also tended to depress responses, although this effect was substantially less (generally less than 10%) and also less consistent. As the opercularis muscle appears to be derived from the levator scapulae musculature, it is speculated that primitively seismic sensitivity was enhanced by a muscular connection that could transmit motion from the forelimb to the otic region, responsiveness being further enhanced by the subsequent evolution of the specialized opercularis system.     

Timing of development of the middle ear of Anura (Amphibia)

Summary The opercularis system and tympanum-stapes complex of the anuran middle ear develop at different times relative to metamorphosis. In early larvae, the fenestra ovalis is represented by a large lateral opening in the otic capsule filled with connective tissue. At later larval stages, but well before metamorphosis, a cartilaginous operculum begins to form at the posterior margin of the fenestra ovalis, and proceeds to expand to fill all except the anterior part of the fenestra. The opercularis muscle forms along with the levator scapulae superior muscle at the anteromedial edge of the developing suprascapular cartilage of the shoulder girdle. The muscle fibers extend anteroventrally towards the operculum and otic capsule, and, just before emergence of the forelimbs, that portion that will form the opercularis muscle inserts on the lateral surface of the operculum. At this stage, when the metamorphosing frogs first show terrestrial habits, the opercularis system is complete and presumably functional. Timing of development of the tympanum-stapes complex is more variable. The stapes begins as a cartilaginous condensation in the anterior part of the fenestra ovalis, and develops laterally to eventually contact the epidermis and dermis that together will form the tympanum. Meanwhile a middle ear cavity and tympanic annulus form to complete the complex. In several species, especially those that metamorphose at a smaller body size, the tympanum-stapes complex is quite incomplete by the end of metamorphosis, and in Hyla crucifer it takes about 60 days to fully develop. The presence of a complete opercularis system by the start of terrestrial activity is consistent with an hypothesized seismic function of the system. The independent timing of development of the opercularis system and tympanum-stapes complex does not support functional hypotheses linking the opercularis system with modulation of responsiveness of the tympanum-stapes complex to aerial sound. Newly metamorphosed frogs with poorly developed tympanum-stapes complexes are presumably either insensitive to aerial sound or employ alternate mechanisms for transmission of sound energy to the inner ear, possibly involving the opercularis system.     

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.     

Simulation of the effects of different pilot helmets on neck loading during air combat

New generation pilot helmets with mounted devices enhance the capabilities of pilots substantially. However, the additional equipment increases the helmet weight and shifts its center of mass forward. Two helmets with different mass properties were modeled to simulate their effects on the pilot's neck. A musculoskeletal computer model was used, with the methods of inverse dynamics and static optimization, to compute the muscle activations and joint reaction forces for a given range of quasi-static postures at various accelerations experienced during air combat. Head postures which induce much higher loads on the cervical spine than encountered in a neutral position could be identified. The increased weight and the forward shift of the center of mass of a new generation helmet lead to higher muscle activations and higher joint reaction loads over a wide range of head and neck movements. The muscle activations required to balance the head and neck in extreme postures increased the compressive force at the T1-C7 level substantially, while in a neutral posture the muscle activations remained low. The lateral neck muscles can reach activations of 100% and cause compressive joint forces up to 1100N during extensive rotations and extensions at high 'vertical' accelerations (Gz). The calculated values have to be interpreted with care as the model has not been validated. Nevertheless, this systematic analysis could separate the effects of head posture, acceleration and helmet mass on neck loading. More reliable data about mass properties and muscle morphometry with a more detailed motion analysis would help to refine the existing model.     

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 sumerged. 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.     

Identification of head motions by central vestibular neurons receiving linear and angular input

Most naturally occurring displacements of the head in space, due to either an external perturbation of the body or a self-generated, volitional head movement, apply both linear and angular forces to the head. The vestibular system detects linear and angular accelerations of the head separately, but the succeeding control of gaze and posture often relies upon the combined processing of linear and angular motion information. Thus, the output of a secondary neuron may reflect the linear, the angular, or both components of the head motion. Although the vestibular system is typically studied in terms of separate responses to linear and angular acceleration of the head, many secondary and higher-order neurons in the vestibular system do, in fact, receive information from both sets of motion sensors. The present paper develops methods to analyze responses of neurons that receive both types of information, and focuses on responses to sinusoidal motions composed of a linear and an angular component. We show that each neuron has a preferred motion, but a single neuron cannot code for a single motion. However, a pair of neurons can code for a motion by the relative phases of firing-rate modulation. In this way, information about motion is enhanced by neurons combining information about linear and angular motion. Received: 5 November 1998 / Accepted in revised form: 19 March 1999     

Representation of sound pressure and particle motion information in the midbrain of the goldfish

1. Averaged evoked responses from multiple electrodes in the goldfish midbrain (torus semicircularis) area were recorded in response to acoustic stimulation by loudspeaker and to direct vertical vibration of the head. 2. Relative pressure and displacement sensitivity was such that in the far field, the response to sound pressure would dominate the response to particle motion by 40-90 dB. 3. Swimbladder deflation caused a flat (70-1000 Hz) loss in pressure sensitivity ranging from 20 to over 50 dB, and led to an enhanced response to vibration at low frequencies. 4. The goldfish midbrain is not homogeneous with regard to relative pressure and motion sensitivity.     

Use of whole body vibration in individuals with chronic stroke: Transmissibility and signal purity

This study examined (1) the influence of whole body vibration (WBV) frequency (20 Hz, 30 Hz, 40 Hz), amplitude (low: 0.8 mm and high: 1.5 mm) and body postures (high-squat, deep-squat, tip-toe standing) on WBV transmissibility and signal purity, and (2) the relationship between stroke motor impairment and WBV transmissibility/signal purity. Thirty-four participants with chronic stroke were tested under 18 different conditions with unique combinations of WBV frequency, amplitude, and body posture. Lower limb motor function and muscle spasticity were assessed using the Fugl-Meyer Assessment and Modified Ashworth Scale respectively. Nine tri-axial accelerometers were used to measure acceleration at the WBV platform, and the head, third lumbar vertebra, and bilateral hips, knees, and ankles. The results indicated that WBV amplitude, frequency, body postures and their interactions significantly influenced the vibration transmissibility and signal purity among people with chronic stroke. In all anatomical landmarks except the ankle, the transmissibility decreased with increased frequency, increased amplitude or increased knee flexion angle. The transmissibility was similar between the paretic and non-paretic side, except at the ankle during tip-toe standing. Less severe lower limb motor impairment was associated with greater transmissibility at the paretic ankle, knee and hip in certain WBV conditions. Leg muscle spasticity was not significantly related to WBV transmissibility. In clinical practice, WBV amplitude, frequency, body postures need to be considered regarding the therapeutic purpose. Good contact between the feet and vibration platform and symmetrical body-weight distribution pattern should be ensured.     

The effect of postural correction on muscle activation amplitudes recorded from the cervicobrachial region

In clinical practice, postural correction is a common treatment approach for individuals with neck and shoulder pain. As chronic static muscle use is thought to be associated with the onset of some neck and shoulder pain syndromes, it is important to understand the impact a postural correction program might have on muscle activation amplitudes in the neck and shoulder regions. Normalized surface electromyographic data were recorded from the levator scapulae, upper trapezius, supraspinatus, posterior deltoid, masseter, rhomboid major, cervical erector spinae, and sternocleidomastoid muscles of the dominant side of each of eighteen healthy subjects. Subjects performed five repetitions of each of four seated typing postures (habitual, corrected, head-forward and slouched) and four standing postures (habitual, corrected, and head-forward and slouched). Repeated-measures analysis of variance models (α = 0.05) revealed that in sitting postural correction tended to decreased the level of muscle activation required in all muscles studied during seated computer work, however this finding was not statistically significant. Corrected posture in sitting did, however produce a statistically significant reduction in muscle activity compared to forward head posture. Corrected posture in standing required more muscle activity than habitual or forward head posture in the majority of cervicobrachial and jaw muscles, suggesting that a graduated approach to postural correction exercises might be required in order to train the muscles to appropriately withstand the requirements of the task. A surprising finding was that muscle activity levels and postural changes had the largest impact on the masseter muscle, which demonstrated activation levels in the order of 20% maximum voluntary electrical activation.     

Effects of vibration on arm and shoulder muscles in three body postures

The electromyographic responses of arm and shoulder muscles to vibrations were studied in three postures similar to the postures of drilling in a ceiling, drilling in a wall and drilling in a floor. This experiment was performed within the defined parameters of: vibrational frequency at 30 Hz, acceleration level 40 m.s-2 (rms), pushing force expressed as percentage maximal voluntary contraction, and gripping force which was set at 100 N. The exposure time for each test was 5 min. The general findings from these three body postures show that all the examined muscles were affected by exposure to vibration. The EMG index increased as follows: trapezius muscle 39% (p less than 0.05), lower-arm flexor muscles 23% (p less than 0.05), infraspinatus muscle 14% (p less than 0.05), lower-arm extensor muscles 14% (p less than 0.1) and biceps muscle 6% (p less than 0.1). The muscle most affected by vibration was found to be the trapezius muscle. It should be taken into consideration that vibration can be a contributing factor in neck/shoulder disorders among power handtool operators. The general conclusion from this study is that changes in working posture give different transmissions of vibration in the upper extremities. It seems as if the prime movers and muscles with an increased muscle length or increased degree of contraction are most affected by vibration.     

In vivo measurement of dynamic rectus femoris function at postures representative of early swing phase

Forward dynamic models suggest that muscle-induced joint motions depend on dynamic coupling between body segments. As a result, biarticular muscles may exhibit non-intuitive behavior in which the induced joint motion is opposite to that assumed based on anatomy. Empirical validation of such predictions is important for models to be relied upon to characterize muscle function. In this study, we measured, in vivo, the hip and knee accelerations induced by electrical stimulation of the rectus femoris (RF) and the vastus medialis (VM) at postures representatives of the toe-off and early swing phases of the gait cycle. Seven healthy young subjects were positioned side-lying with their lower limb supported on air bearings while a 90 ms pulse train stimulated each muscle separately or simultaneously. Lower limb kinematics were measured and compared to predictions from a similarly configured dynamic model of the lower limb. We found that both RF and VM, when stimulated independently, accelerated the hip and knee into extension at these postures, consistent with model predictions. Predicted ratios of hip acceleration to knee acceleration were generally within 1 s.d. of average values. In addition, measured responses to simultaneous RF and VM stimulation were within 13% of predictions based on the assumption that joint accelerations induced by activating two muscles simultaneously can be found by adding the joint accelerations induced by activating the same muscles independently. These results provide empirical evidence of the importance of considering dynamic effects when interpreting the role of muscles in generating movement.     

Laser vibrometric studies of sound-induced motion of the body walls and lungs of salamanders and lizards: implications for lung-based hearing

A laser Doppler vibrometer was used to measure the acoustic responses of different body surfaces of several species of salamanders and lizards. The lateral body wall over the lung displayed sound-induced motion up to 30 dB greater than the lateral head surface from 300-1,000 Hz in salamanders and from 200-2,500 Hz in lizards. The lateral body wall of lungless plethodontid salamanders showed no such enhanced motion to sound. The lateral body wall of lizards was more responsive than their tympanum to sound frequencies below about 1,250-2,000 Hz. The frequency of the peak response of lizard body walls matched the resonant frequency of a Helmholtz resonator with the volume and dimensions of their lungs. In contrast, the frequency of peak response of salamander body walls was well below the resonant frequencies calculated for both Helmholtz resonators and closed tubes with the dimensions and volumes of their lungs. Nonetheless, filling the lungs with saline dramatically reduced the responsiveness of the lateral body walls of both the lunged salamanders and the lizards. As previously demonstrated in anuran amphibians, the lateral body wall and lungs of salamanders and lizards may function in sound reception, especially at relatively low frequencies.     

Functional analysis of the shoulder girdle of cats during locomotion

The movements of the shoulder girdle of eight adult cats during overground stepping were studied, using standard slow motion cinematographic techniques. The patterns of activity of shoulder muscles were examined, using simultaneous intramuscular electromyography. Walking, trotting and galloping steps were analyzed from digitized single motion picture frame images. Angular movements of the shoulder girdle consist of biphasic flexion and extension of the shoulder joint and a monophasic flexion-extension alternation of the scapula on the thorax during each step cycle. In addition, the center of the scapula moves craniad during the swing phase and caudad during the stance phase with respect to a fixed reference point on the animal. Similar vertical movements of the center of the scapula also occur in each step cycle. Results of EMG studies of the 17 muscles capable of acting on the shoulder girdle indicate that three overall patterns of activity are found: (1) a pattern typical of extensor muscles, active during all the extension epochs; (2) a pattern typical of flexor muscles, active during the flexion epoch; and (3) a biphasic pattern of activity, active twice in each step. There data are used, along with a re-examination of previous models of the mechanics of the shoulder girdle of carnivores to examine the function and mechanics of shoulder motion. It is concluded that the rotary and translatory movements of the shoulder girdle during stepping combine to enhance step length.     

Seated whole body vibrations with high-magnitude accelerations--relative roles of inertia and muscle forces

Reliable computation of spinal loads and trunk stability under whole body vibrations with high acceleration contents requires accurate estimation of trunk muscle activities that are often overlooked in existing biodynamic models. A finite element model of the spine that accounts for nonlinear load- and direction-dependent properties of lumbar segments, complex geometry and musculature of the spine, and dynamic characteristics of the trunk was used in our iterative kinematics-driven approach to predict trunk biodynamics in measured vehicle's seat vibrations with shock contents of about 4g (g: gravity acceleration of 9.8m/s(2)) at frequencies of about 4 and 20Hz. Muscle forces, spinal loads and trunk stability were evaluated for two lumbar postures (erect and flexed) with and without coactivity in abdominal muscles. Estimated peak spinal loads were substantially larger under 4Hz excitation frequency as compared to 20Hz with the contribution of muscle forces exceeding that of inertial forces. Flattening of the lumbar lordosis from an erect to a flexed posture and antagonistic coactivity in abdominal muscles, both noticeably increased forces on the spine while substantially improving trunk stability. Our predictions clearly demonstrated the significant role of muscles in trunk biodynamics and associated risk of back injuries. High-magnitude accelerations in seat vibration, especially at near-resonant frequency, expose the vertebral column to large forces and high risk of injury by significantly increasing muscle activities in response to equilibrium and stability demands.     

Crouched postures reduce the capacity of muscles to extend the hip and knee during the single-limb stance phase of gait

Many children with cerebral palsy walk in a crouch gait that progressively worsens over time, decreasing walking efficiency and leading to joint degeneration. This study examined the effect of crouched postures on the capacity of muscles to extend the hip and knee joints and the joint flexions induced by gravity during the single-limb stance phase of gait. We first characterized representative mild, moderate, and severe crouch gait kinematics based on a large group of subjects with cerebral palsy (N=316). We then used a three-dimensional model of the musculoskeletal system and its associated equations of motion to determine the effect of these crouched gait postures on (1) the capacity of individual muscles to extend the hip and knee joints, which we defined as the angular accelerations of the joints, towards extension, that resulted from applying a 1N muscle force to the model, and (2) the angular acceleration of the joints induced by gravity. Our analysis showed that the capacities of almost all the major hip and knee extensors were markedly reduced in a crouched gait posture, with the exception of the hamstrings muscle group, whose extension capacity was maintained in a crouched posture. Crouch gait also increased the flexion accelerations induced by gravity at the hip and knee throughout single-limb stance. These findings help explain the increased energy requirements and progressive nature of crouch gait in patients with cerebral palsy.     

Kinetic chain of overarm throwing in terms of joint rotations revealed by induced acceleration analysis

This study investigated how baseball players generate large angular velocity at each joint by coordinating the joint torque and velocity-dependent torque during overarm throwing. Using a four-segment model (i.e., trunk, upper arm, forearm, and hand) that has 13 degrees of freedom, we conducted the induced acceleration analysis to determine the accelerations induced by these torques by multiplying the inverse of the system inertia matrix to the torque vectors. We found that the proximal joint motions (i.e., trunk forward motion, trunk leftward rotation, and shoulder internal rotation) were mainly accelerated by the joint torques at their own joints, whereas the distal joint motions (i.e., elbow extension and wrist flexion) were mainly accelerated by the velocity-dependent torques. We further examined which segment motion is the source of the velocity-dependent torque acting on the elbow and wrist accelerations. The results showed that the angular velocities of the trunk and upper arm produced the velocity-dependent torque for initial elbow extension acceleration. As a result, the elbow joint angular velocity increased, and concurrently, the forearm angular velocity relative to the ground also increased. The forearm angular velocity subsequently accelerated the elbow extension and wrist flexion. It also accelerated the shoulder internal rotation during the short period around the ball-release time. These results indicate that baseball players accelerate the distal elbow and wrist joint rotations by utilizing the velocity-dependent torque that is originally produced by the proximal trunk and shoulder joint torques in the early phase.     

A potential link between lateral semicircular canal orientation,head posture,and dietary habits in extant rhinos (Perissodactyla,Rhinocerotidae)

Extant rhinoceroses share the characteristic nasal horn, although the number and size of horns varies among the five species. Although all species are herbivores, their dietary preferences, occipital shapes, and common head postures vary. Traditionally, to predict the “usual” head posture (the most used head posture of animals during normal unstressed activities, i.e., standing) of rhinos, the occipital shape was used. While a backward inclined occiput implies a downward hanging head (often found in grazers), a forward inclined occiput is related to the horizontal head posture in browsing rhinos. In this study, the lateral semicircular canal (LSC) of the bony labyrinth was virtually reconstructed from µCT‐images in order to investigate a possible link between LSC orientation and head posture in extant rhinoceroses. The usual head posture was formerly reconstructed for several non‐rhinoceros taxa with the assumption that the LSC of the inner ear is held horizontal (parallel to the ground) during normal activity of the living animal. The current analysis of the LSC orientation resulted in a downward inclined usual head posture for the grazing white rhinoceros and a nearly horizontal head posture in the browsing Javan rhinoceros. The other three browsing or mixed feeding species show subhorizontal (closer to horizontal than a downgrade inclination) head postures. The results show that anatomical and behavioral aspects, like occipital shape, presence and size of horns/tusk‐like lower incisors, as well as feeding and feeding height preferences influence the usual head posture. Because quantitative behavioral data are lacking for the usual head postures of the extant rhinos, the here described relationship between the LSC orientation and the resulting head posture linked to feeding preferences gives new insights. The results show, that the inner ear provides additional information to interpret usual head postures linked to feeding preferences that can easily be adapted to fossil rhinoceroses.     

The kinematics of the scapulae and spine during a lifting task

Simultaneous motion of the scapula and humerus is widely accepted as a feature of normal upper limb movement, however this has usually been investigated under conditions in which purposeful, functional tasks were not considered. The aim of this study was to investigate the synchrony and coordination of the constituent 3D movements of the shoulder girdle and trunk, during a functional activity. 45 healthy women, aged between 20 and 80 years, performed a simple lifting task, moving a loaded box from a shelf at waist level to one at shoulder level and then reversed the movement, during which the linear and angular motions of the scapulae, upper and lower thoracic spine and upper limbs were monitored and analysed using cross-correlation techniques. Results indicated a close and consistent set of coordinated movement patterns, which suggest biomechanical invariance in the responses of the structures adjacent to the upper limb during such a lifting task. These scapulohumeral relationships were, however, more constant and phase-locked when there was a specific purpose to the movement than during periods in which the arm was lowered without load. There were no age-related differences in any movement responses.     

Trunk muscle electromyography and whole body vibration

By measuring the electromyographic (EMG) activity of the paraspinal muscles, we have estimated the average and peak-to-peak torque imposed on the spine during whole body vibration. Six subjects had surface electrodes placed on their erector spinae muscles at the L3 level. The EMG-torque relationship was estimated by having each subject perform isometric horizontal pulls in an upright seated posture. The subject was then vibrated vertically and sinusoidally in a controlled, flexed, slightly lordotic seated posture, in 1 Hz increments from 3 to 10 Hz at a 0.1 g RMS seat acceleration level. Between vibration readings taken at each frequency, a static reading was also taken with the subject maintaining the same posture. The entire vibration-static 3-10 Hz test was repeated for reliability purposes. Specialized digital signal processing techniques were developed for the EMG signals to enhance the measured cyclic muscle activity and to allow automatic measurement of the time relationship between the mechanical displacement and the estimated torque. We found significantly more average and peak-to-peak estimated torque at almost all frequencies for vibration vs static sitting.     

Electrophysiological responses in guinea pig cochlea to low frequency sound stimuli: distortion of cochlear microphonic (CM) wave form

The present experiment investigated whether or not auditory responses of the middle and/or inner ear in guinea pigs to low frequency sound stimuli [ 60 Hz-2 kHz at 90-120 dB(SPL) ] exhibited the harmonic distortion phenomenon resulting from cochlear microphonics (CM). Measurement of CM leading in turn I by the differential electrode recording method involved measurement of 50 microV isopotential responses, output voltages and CM wave form distortion at each constant sound pressure. The results obtained were as follows: (1) On the 50 microV isopotential response curve and the output voltage curves, the changes at 60-90 Hz were different from those at higher frequencies. (2) At stimuli of 90 or 100 dB(SPL), CM wave form distortion appeared frequently at frequencies below 120 Hz, but were less pronounced above approximately 200 Hz. (3) When raised to 110 and 120 dB(SPL), almost all CM wave forms were distorted at all test frequencies between 60 and 500 Hz. (4) The patterns of CM wave form distortion at frequencies below approximately 120 Hz showed peak clipping and triangular wave distortions, while those at frequencies above approximately 200 Hz showed little of these distortions.