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We report an experiment to investigate possible vestibular effects on finger tapping to an auditory anapaest rhythm. In a sample of 10 subjects, index finger acceleration and tapping force were recorded along with extensor/flexor activity and the associated electroencephalographic activity measured at central and cerebellar surface electrodes. In a prior session with a standard short air-conducted 500-Hz pip, vestibular evoked myogenic potential thresholds were measured and subsequently used to set the acoustic intensity. During the main experiment subjects were asked to synchronise tapping to the pips arranged in the anapaest at two different frequencies, 500 Hz vs 5 kHz, so that only the low-frequency high-intensity condition was a vestibular, as well as an auditory stimulus. We hypothesised that a vestibular effect would manifest in an interaction between the frequency and intensity factors for a range of dependent measures of tapping performance. No clear evidence was found for vestibular effects, but this was likely due to the confounding effects of an independent effect of intensity and the relative weakness of the acoustic vestibular stimulus. However, the data did show novel evidence for two distinct timing processes for the flexion and extension stages of a tap cycle and two distinct timing strategies, which we refer to as ‘staccato’ and ‘legato’, characterised by different profiles of force and extension.

In the original publication of the article, Fig. 3 is incorrect.

In this work we examine the possible neural basis for two brainstem-spinal reflexes using source analyses of brain activity recorded over the cortex and posterior fossa. In a sample of 5 healthy adult subjects, using axial and vestibular stimulation by means of applied impulsive forces, evoked potentials were recorded with 63 channels using a 10 % cerebellar extension montage. In parallel, EMG was recorded from soleus and tibialis anterior muscles and accelerometry from the lower leg. Recordings over the cerebellum (ECeG) confirmed the presence of short latency (SL) potentials and these were associated with changes in high-frequency power. The SL responses to the two stimulus modalities differed in that the axial stimulation produced an initial pause and then a burst in the high-frequency ECeG, followed by excitation/inhibition in soleus while vestibular stimulation produced an initial burst then a pause, followed by inhibition/excitation in soleus. These short latency responses were followed by longer latency N1/P2/N2 responses in the averaged EEG, which were maximal at FCz. Brain Electrical Source Analysis (BESA) demonstrated both cerebellar and cerebral cortical contributions to the short-latency responses and primarily frontal cortex contributions to the long-latency EPs. The latency and polarity of the SL EPs, in conjunction with changes in high-frequency spontaneous activity, are consistent with cerebellar involvement in the control of brainstem-spinal reflexes. The early involvement of frontal cortex and subsequent later activity may be an indicator of the activation of the cortical motor-related system for rapid responses which may follow the reflexive components. These findings provide evidence of the feasibility of non-invasive electrophysiology of the human cerebellum and have demonstrated cerebellar and frontal activations associated with postural-related stimuli.

We studied nine normal volunteers with a classical conditioning paradigm using a mastoid tap, believed to activate otolith receptors, as an unconditional stimulus (US) and the consequent blink as the unconditioned response (UR). Both visual (alternation of stripes) and an auditory tone were used as conditional stimuli (CS). Recordings were made below the eyes at IO1 and IO2, from over the frontal eye fields (C3′ and C4′) and over the posterior fossa, the latter at sites we have previously reported that we were able to record an evoked climbing fibre response (CFR) at short latency. Behavioural analysis confirmed that weak conditioning did occur early, which subsequently showed extinction on repeated CS alone trials. Further, a UR was more likely to occur following a preceding CFR when preceded by a CS, supporting a correlation between the CFRs and behaviour. For further statistical analysis, the time period of interest was divided into a series of epochs, based around the events occurring at the time. Grand averages, plus analysis of variance, confirmed evidence of weak conditioning for the blink response following both modalities. The EMG associated with the eyeblink for the UR occurred at a similar time to the expected post-CFR pause in the spontaneous cerebellar activity, or electrocerebellogram (ECeG), while hypothesised conditioned pausing in the ECeG was also observed in CS alone trials. A correlation was found between the size of the CFR and the RMS amplitudes of the segments covering the ocular vestibular evoked myogenic potential oVEMP, (periocular) EMG and the EOG. The slope was greater for the non-oVEMP segments than for the oVEMP segment suggesting the correlation was not simply due to differing sizes of the vestibular volley. We suggest that these recorded events fit with the proposed role of the CFR in Purkinje neurons in classical conditioning, gating the excitability of the cerebellar nuclei, and thereby neurons in the reticular formation mediating the otolith blink reflex. This effect appears to apply to polysynaptic reflexes only as there was no evidence of changes to the oVEMP.

Loud sounds were used to activate vestibular receptors in human volunteers and the effects of head and eye position studied for short-latency responses. A potential (p10/n17) recorded in the parieto-occipital leads showed behavior not expected for a response with a myogenic origin. Source modeling suggested a possible origin from the cerebellum. It may represent a new indicator of human vestibulocerebellar function., This study investigated the effect of eye gaze and head position on vestibular-evoked potentials (VsEPs). Head position would be expected to affect myogenic sources, and eye position is known to affect ocular myogenic responses (ocular vestibular-evoked myogenic potentials), whereas a neurogenic source should behave otherwise. Eleven healthy subjects were recruited, and VsEPs, using 72-channel EEG, were recorded at a fixed intensity above the vestibular threshold. Three eye gaze and three head positions were tested (−20°, 0°, and +20° to the horizontal). Short-latency potentials showed that in addition to the expected effect of gaze on infraocular (IO′) leads, where up-gaze gives a maximum response, significant changes in amplitude were also observed in electrodes remote from the eyes and in particular, from contralateral parietal-occipital (PO) and neck (CB′) leads. Short-latency potentials of similar latency were observed (p10/n17 and n10/p17, respectively). The pattern of change with gaze in the PO leads was distinct from that observed for the IO′ leads. For the PO leads, the maximum response was obtained with neutral gaze, and this was also distinct from that observed for CB′ electrodes, where a maximal response was observed with head flexion in the second wave but not the first. Evidence of modulation of N42 and N1 potentials with both eye and head position was also observed. Head- and eye-position manipulation thus suggests that the inion response consists of an early neurogenic component, as well as myogenic responses. The p10/n17 at PO, in particular, may be an indicator of vestibulocerebellar projections. NEW & NOTEWORTHY Loud sounds were used to activate vestibular receptors in human volunteers and the effects of head and eye position studied for short-latency responses. A potential (p10/n17) recorded in the parieto-occipital leads showed behavior not expected for a response with a myogenic origin. Source modeling suggested a possible origin from the cerebellum. It may represent a new indicator of human vestibulocerebellar function.

Vestibular cerebellar evoked potentials (VsCEPs) were recorded from over the occipital and cerebellar regions of the scalp using bone-conducted (BC) stimuli applied at the mastoids (impulsive accelerations and 500 Hz) and 500 Hz acoustic tones (AC). Ten healthy subjects were tested. Electrodes were positioned over the midline (Oz, Iz, CBz) and at 3, 6 and 9 cm intervals lateral to the midline electrodes bilaterally. Additional electrodes were also positioned over posterior neck muscles (SPL1 and SPL2). The largest evoked potentials on average were recorded from the electrodes 3 and 6 cm lateral to the Iz and CBz midline locations. BC stimuli produced short latency potentials on the side contralateral to the stimulated mastoid and were dependent on stimulus polarity. Positive polarity stimuli produced biphasic VsCEPs at approximately 12 and 17 ms (P12–N17) for BC impulses and 10 and 15 ms (P10–N15) for BC 500 Hz stimuli. Following the initial excitation, there was a period of suppression of background activity lasting an average of 16.8 ms for positive polarity BC impulses. Negative polarity stimuli produced later VsCEPs both for BC impulses (P20–N26) and BC 500 Hz (P13–N18). VsCEPs to AC 500 Hz stimuli lateralised to the contralateral side and were larger for right than left ear stimulation. Stimulus polarity (condensation and rarefaction) did not alter the timing of the VsCEPs to AC 500 Hz tones. No evoked response was recorded to somatosensory (median and radial nerve) stimulation. Four patients with cerebellar disease were tested and two showed abnormal VsCEPs with initial negativities. VsCEPs show distinct mapping over the posterior fossa and are likely to reflect climbing fibre responses via crossed otolith–cerebellar pathways.

We recorded evoked potentials (EPs) from over the posterior fossa and in parallel ocular vestibular evoked myogenic potentials (OVEMPs) during visuo-vestibular stimulation in a sample of 7 male and 11 female human subjects. In 9 of the 18 subjects we were able to record EPs reliably in the form of an early biphasic positive-negative wave with latencies ~12 and 17 ms ipsilateral to head acceleration direction (P12-N17) and a slightly later, contralateral, biphasic positive-negative wave with latencies ~19 and 23 ms (P19-N23). The amplitudes of the responses varied widely between subjects. Both P12 and N23 EPs were modulated by the mode of visual stimulation, larger for vection (sense of movement) compared with optokinetic nystagmus and for congruent movement. We suggest that the EPs measured over the posterior fossa are a manifestation of climbing fiber responses of cerebellar cortical Purkinje cells, i.e., a form of vestibular cerebellar EP (VsCEP). The two subject groups with and without VsCEPs were distinguished by the magnitude of their OVEMPs and their subjective experience of vection. The modulation of VsCEPs by visual context may be a manifestation of cerebellar control of linear vestibular ocular reflex gain. NEW & NOTEWORTHY We report likely vestibular cerebellar evoked potentials (VsCEPs) produced by lateral head impulses recorded in intact humans over the posterior fossa. VsCEPs occurred as short-latency P12-N17 waves ipsilateral to the direction of head motion and as P19-N23 contralaterally and were present in half our subjects. Their properties suggest that the VsCEPs may be of a climbing-fiber origin. VsCEPs are related to the perception of motion and, possibly, control of linear vestibular ocular reflex gain.

The findings are reported of an investigation into rhythmic differences between infant-directed speech (IDS) and adult-directed speech (ADS) in a corpus of utterances from Australian English mothers speaking to their infants and to another adult. Given the importance of rhythmic cues to stress and word-segmentation in English, the investigation focused on the extent to which IDS makes such cues salient. Two methods of analysis were used: one focused on segmental durational properties, using a variety of durational measures; the other focused on the prominence of vocalic/sonorant segments, as determined by their duration, intensity, pitch, and spectral balance, using individual measures as well as composite measures of prominence derived from auditory-model analyses. There were few IDS/ADS differences/trends on the individual measures, though mean pitch and pitch variability were higher in IDS than ADS, while IDS vowels showed more negative spectral tilt. However, the model-based analyses suggested that differences in the prominence of vowels/sonorant segments were reduced in IDS, with further analysis suggesting that pitch contributed little to prominence. The reduction in prominence contrasts may be due to the importance of mood-regulation in speech to young infants, and may suggest that infants rely on segmental cues to stress and word-segmentation.

In this article, documents relating to the history of the Radium Committee of the Royal Society are collated for the first time. Founded in 1903, the committee had its status enhanced in 1904, when the Goldsmiths' Company donated £1000 for the establishment of a Radium Research Fund. Two years later the fund was used to purchase 500 kg of pitchblende residues from the Austrian government. The French chemist Armet de Lisle was contracted to perform the first stage of extraction, and the process of purification was performed at the Government Laboratory during 1907 by the Government Analyst, T. E. Thorpe, yielding an estimated 70 mg of radium chloride. In 1914 the unexpended balance of about £500 was awarded to Ernest Rutherford, but the bulk was not used until 1921, when Rutherford had moved to Cambridge. The fund was then used to purchase radium that had been on loan to him from Austria before World War I. After Rutherford's death in 1937 the Committee was wound up, and the Society's radium was controlled on a more ad hoc basis. After Thorpe's work in 1907, the radium was lent out successively to several leading scientists until its existence was last recorded in 1953.

Objective To deduce the connectivity underlying ocular vestibular evoked myogenic potentials (OVEMPs) recorded from two sites and produced by lateral transmastoid stimulation in patients with unilateral vestibular dysfunction. Methods OVEMPs were recorded using lateral impulsive stimuli delivered by a hand-held minishaker placed at the mastoid. Twelve patients were tested using the typical OVEMP recording montage placed inferior to the eyes. In a subset of 6 patients, recordings were also made using a lateral electrode montage. The majority of patients were tested following surgery for inner ear disease. Patient responses were compared to those in normal subjects under similar recording conditions. Results For the inferior montage, regardless of which mastoid was stimulated, deficits were observed only from the eye opposite the affected ear. In contrast, OVEMPs recorded using the lateral electrode montage showed changes on both sides. Conclusions OVEMPs produced using lateral transmastoid stimulation and recorded from beneath the eyes are generated by a crossed vestibulo-ocular pathway while the projections underlying the lateral responses are likely to be bilateral. Significance The vestibular-ocular connectivity underlying the OVEMPs recorded from inferior and lateral recording sites differs. For clinical use, the inferior recording site is the simplest to interpret.

We recently reported that a major contribution to the low-frequency tuning and sensitivity of the human vestibular system is the biomechanical properties of the vestibular end-organs. In the current paper, we investigate the contribution of additional mechanisms to low-frequency tuning. We compared the response properties of the vestibular system in 6 human volunteers to trains of 2 ms pulses of sound and transmastoid vibration using pulse repetition frequencies of 12.5, 25, 50, 100, 200 and 400 Hz. Measurements were made using two separate pathways arising from the vestibular apparatus: to the neck using vestibular evoked myogenic potentials (VEMPs), and to the eyes using ocular vestibular evoked myogenic potentials (OVEMPs). For both sound and vibration the two response pathways produced different tuning to pulse trains. The vestibulo-ocular pathway was characterised by a band-pass tuning with best frequency of 100 Hz whereas the vestibulo-collic pathway showed a peak at 400 Hz with sound only. These results suggest that properties of the vestibulo-ocular pathway also contribute to the low-frequency tuning that occurs for the OVEMP, in addition to previously reported end-organ effects.

Recent work has demonstrated that the human vestibular system displays a remarkable sensitivity to low-frequency vibration. To address the origin of this sensitivity we compared the frequency response properties of vestibular reflexes to 10 ms bursts of air-conducted sound and transmastoid vibration, which are thought to be differentially selective for the saccule and utricle, respectively. Measurements were made using two separate central pathways: vestibular evoked myogenic potentials (VEMPs), which are a manifestation of vestibulo-collic projections, and ocular vestibular evoked myogenic potentials (OVEMPs), which are a manifestation of vestibulo-ocular projections. For both response pathways air-conducted sound and vibration stimuli produced the same patterns of quite different tuning. Sound was characterised by a band-pass tuning with best frequency between 400 and 800 Hz whereas vibration showed a low-pass type response with a largest response at 100 Hz. Our results suggest that the tuning is at least in part due to properties of end-organs themselves, while the 100 Hz best frequency may be a specifically utricular feature.

Mechanoreceptive hair-cells of the vertebrate inner ear have a remarkable sensitivity to displacement, whether excited by sound, whole-body acceleration or substrate-borne vibration. In response to seismic or substrate-borne vibration, thresholds for vestibular afferent fibre activation have been reported in anamniotes (fish and frogs) in the range -120 to -90 dB re 1g. In this article, we demonstrate for the first time that the human vestibular system is also extremely sensitive to low-frequency and infrasound vibrations by making use of a new technique for measuring vestibular activation, via the vestibulo-ocular reflex (VOR). We found a highly tuned response to whole-head vibration in the transmastoid plane with a best frequency of about 100 Hz. At the best frequency we obtained VOR responses at intensities of less than -70 dB re 1g, which was 15 dB lower than the threshold of hearing for bone-conducted sound in humans at this frequency. Given the likely synaptic attenuation of the VOR pathway, human receptor sensitivity is probably an order of magnitude lower, thus approaching the seismic sensitivity of the frog ear. These results extend our knowledge of vibration-sensitivity of vestibular afferents but also are remarkable as they indicate that the seismic sensitivity of the human vestibular system exceeds that of the cochlea for low-frequencies.

Objective Recent work has demonstrated the existence of ocular vestibular evoked myogenic potentials (OVEMPs), which likely reflect projections underlying the translational vestibular ocular reflex (TVOR). We examined extraocular muscle activity associated with impulsive acceleration of the head in the transmastoid plane. Methods Accelerometry was measured in 4 subjects in response to acceleration impulses produced by a gamma function delivered with a Minishaker (4810, Bruel & Kjaer). This stimulus produced peak head accelerations of 0.13–0.14 g occurring at between 3.1 and 4.0 ms at the mastoids for both right and left head movement. OVEMPs were recorded in 10 normal subjects with 5 directions of gaze, using electrode pairs placed lateral to, above and below the eyes. Results OVEMPs occurred at short latency, with initial peaks between 10.3 ms (p10) and 15.3 ms (n15). For a given recording site and gaze direction, the responses were determined solely by the direction of imposed acceleration. Conclusions We propose that, given the transtemporal nature of the stimuli, utricular afferents are likely to be powerfully activated. The OVEMPs evoked may be generated by the lateral recti and oblique muscles. Significance Sudden lateral accelerations of the head evoke the translational VOR and ocular counter rolling reflex and the pattern of muscle activations indicated by the OVEMPs appear to be a manifestation of these reflexes.

The world's languages display important differences in their rhythmic organization; most particularly, different languages seem to privilege different phonological units (mora, syllable, or stress foot) as their basic rhythmic unit. There is now considerable evidence that such differences have important consequences for crucial aspects of language acquisition and processing. Several questions remain, however, as to what exactly characterizes the rhythmic differences, how they are manifested at an auditory/acoustic level and how listeners, whether adult native speakers or young infants, process rhythmic information. In this paper it is proposed that the crucial determinant of rhythmic organization is the variability in the auditory prominence of phonetic events. In order to test this auditory prominence hypothesis, an auditory model is run on two multi-language data-sets, the first consisting of matched pairs of English and French sentences, and the second consisting of French, Italian, English and Dutch sentences. The model is based on a theory of the auditory primal sketch, and generates a primitive representation of an acoustic signal (the rhythmogram) which yields a crude segmentation of the speech signal and assigns prominence values to the obtained sequence of events. Its performance is compared with that of several recently proposed phonetic measures of vocalic and consonantal variability.

In this paper data are presented from an experiment which provides evidence for the existence of a short latency, acoustically evoked potential of probable vestibular origin. The experiment was conducted in two phases using bone-conducted acoustic stimulation. In the first phase subjects were stimulated with 6-ms, 500-Hz tone bursts in order to obtain the threshold V(T) for vestibular evoked myogenic potentials (VEMP). It was confirmed that the difference between bone-conducted auditory and acoustic vestibular thresholds was slightly over 30 dB. The estimated threshold was then used as a reference value in the second part of the experiment to stimulate subjects over a range of intensities from -6 to +18 dB (re: V(T)). Averaged EEG recordings were made with eight Ag/AgCl electrodes placed on the scalp at Fpz, F3, F4, F7, F8, Cz, T3, and T4 according to the 10-20 system. Below V(T) auditory midlatency responses (MLRs) were observed. Above V(T) two additional potentials appeared: a positivity at about 10 ms (P10) which was maximal at Cz, and a negativity at about 15 ms (N15) which was maximal at Fpz. Extrapolation of the growth functions for the P10 and N15 indicated a threshold close to V(T), consistent with a vestibular origin of these potentials. Given the low threshold of vestibular acoustic sensitivity it is possible that this mode may make a contribution to the detection of and affective responses to loud low frequency sounds. The evoked potentials may also have application as a noninvasive and nontraumatic test of vestibular projections to the cortex.

In this paper I give a brief summary of what is known about the acquisition, use and fate of the radium sources that were in the possession of Lord Rutherford during his lifetime. The account is written in two parts, corresponding to the periods from the discovery of radium in 1898 until his death in 1937 and then from 1937 until recent times. The history of Rutherford's radium closely shadows the history of radioactivity, the evolution of nuclear physics, the race for the bomb, and the development of the nuclear industry.

Short lateral head accelerations were applied to investigate the recruitment properties of the reflexes underlying the earliest ocular and cervical electromyographic reflex responses to these disturbances. Components of both reflexes are vestibular dependent and have been termed “ocular vestibular evoked myogenic potentials” and “cervical vestibular evoked myogenic potentials”, respectively. Previous investigations using a unilateral vestibular stimulus have indicated that some but not all these vestibular-dependent reflexes show a simple power law relationship to stimulus intensity. In particular, crossed otolith-ocular reflexes showed evidence of an inflection separating two types of behaviour. The present stimulus acts bilaterally, and only the earliest crossed otolith-ocular reflex, previously shown to have a strictly unilateral origin, showed evidence of an inflection. Reflex changes in ocular torsion could, in principle, correct for the changes associated with translation for an elevated eye, but our findings indicated that the responses were consistent with previous reports of tilt-type reflexes. For the neck, both vestibular and segmental (muscle spindle) reflexes were evoked and followed power law relationships, without any clear separation in sensitivity. Our findings are consistent with previous evidence of “tilt-like” reflexes evoked by lateral acceleration and suggest that the departure from a power law occurs as a consequence of a unilateral crossed pathway. For the neck, responses to transients are likely to always consist of both vestibular and non-vestibular (segmental) components. Most of the translation-evoked ocular and cervical reflexes appear to follow power law relationship to stimulus amplitude over a physiological range. Electronic supplementary material The online version of this article (doi:10.1007/s00221-014-3980-3) contains supplementary material, which is available to authorized users.

Objective: \ud To evaluate methods for estimating thresholds of vestibular evoked myogenic potentials (VEMPs) and ocular VEMPs (OVEMPs) produced by impulsive transmastoid acceleration (ITA). Design: VEMPs and OVEMPs were obtained simultaneously from subjects in supine posture with both head and eyes elevated. Thresholds to ITA were measured using four different response identification methods. \ud \ud Study sample: \ud Twelve adult subjects with no history of auditory or vestibular deficits. \ud \ud Results: \ud VEMP and OVEMP thresholds were found within the range of -8 to -26 dB re 0.2 g, with overall VEMP thresholds being lower than OVEMP thresholds by average of 3 – 10 dB, depending on the method used.\ud \ud Conclusion: \ud VEMPs can be obtained at lower ITA stimulation levels than OVEMPs. Threshold values depend on the method used to defi ne response presence.

In this paper new evidence is provided to indicate that vestibular responses may be obtained from loud dance music for intensities above 90 dB(A) SPL (Impulse-weighted). In a sample of ten subjects acoustically evoked EMG were obtained from the sternocleidomastoid muscle in response to a sample of techno music typical of that which may be experienced in a dance club. Previous research has shown that this response is vestibularly mediated since it can be obtained in subjects with loss of cochlear function, but is absent in subjects with loss of vestibular function (Colebatch et al. [J. Neurol. Neurosurg. Psychiatr. 57, 190–197 (1994)]. Given that pleasurable sensations of self-motion are widely sought after by more normal means of vestibular stimulation, it is suggested that acoustically evoked sensations of self-motion may account for the compulsion to exposure to loud music. Given further the similarity between the thresholds found, and the intensities and frequency distributions that are typical in rock concerts and dance clubs, it is also suggested that this response may be a physiological basis for the minimum loudness necessary for rock and dance music to work—the “rock and roll threshold” [Dibble, J. Audio Eng. Soc. 43(4), 251–266 (1995)].

In this paper, we develop further a sensory-motor theory of rhythm, time perception and beat induction. We account for these phenomena by the interaction of sensory systems, which represent temporal information in terms of the power spectrum of the sensory image, and the motor system, which has certain natural frequencies. The central assumption is that beat induction is not a passive process but, rather, a form of sensory-guided action involving all the sensory and motor components that that entails, i.e., major portions of the nervous system and the musculoskeletal system which it controls. Even if the musculoskeletal system (‘the plant’, to use the terminology of control theory) is not activated, i.e., there is no motor output, the higher supraspinal levels of the system (‘the controller’) are. To be more specific, the principal agent mediating beat induction is the internal representation of the musculoskeletal system and its dynamic properties which the ‘controller’ requires as a ‘feedforward’ model....

The topic of musical motion has generated a considerable amount of controversy in the past few years (P. Desain, H. Honing, H. van Thienen, & L. Windsor, 1998). In this essay it is argued that motion is central to our understanding of many aspects of music, particularly to our understanding of rhythm, and that an adequate account of motion in music requires a neurobiological perspective. Two possible mechanisms are discussed that may form a neurobiological basis for the association of motion in music: a vestibulomotor mechanism and an audio-visuo-motor mechanism. These two mechanisms in turn may mediate two distinct kinds of musical motion: gesture and locomotion.

Two leading issues raised in commentaries on the authors' earlier article (2007) involved the possible roles of gender differences and the vestibular system on test results. Those issues are discussed in this reply.

Developments in the theory of auditory processing of rhythmic signals have enabled the construction of a robust algorithm for recovery of rhythmic grouping structure. This algorithm appears to be effective for both speech and music signals. The theory upon which the algorithm was based was inspired by the theory of edge detection in vision. The output of the algorithm can be visualised in the form of a “rhythmogram”, examples of which are shown for a variety of speech signals. The relationship between rhythm, time perception and metre is discussed in the light of a recent “auditory-motor” theory of beat induction.

A theory of the principal monaural pathway is described according to which temporal information is spatially coded on three dimensions roughly corresponding to the levels of, respectively, the cochlea, the inferior colliculus, and the auditory cortex [C. Schreiner and G. Langner, ‘‘Coding of temporal patterns in the central auditory nervous system,’’ Auditory Function: Neurobiological Bases of Hearing, edited by G. M. Edelman et al. (Wiley, New York, 1988), pp. 337–361]. This theory is expressed in the form of a computational model which simulates peripheral and central processing and which has the following main components: (1) a one‐dimensional linear bandpass filter‐bank to simulate the cochlea; (2) a two‐dimensional amplitude‐modulation (AM) bandpass filter‐bank to simulate the colliculus; (3) a three‐dimensional AM bandpass filter‐bank to simulate layer IV receptive cells in the cortex; (4) a cortical cross‐channel correlation mechanism; and (5) a central pattern recognition mechanism. Overall, periodicity pitch (i.e., periods of up to about 1000 Hz) is primarily associated with subcortical processing whereas time and rhythm (i.e., periods of up to about 20 Hz) are primarily associated with cortical processing. The model is applied to the following phenomena: virtual pitch shift, musical chords, the psychophysical law for interval discrimination, and the filled interval illusion.

The model [N. Todd, ‘‘A theory of the principal monaural pathway. I. Pitch and time perception,’’ J. Acoust. Soc. Am. (these proceedings)] proposes to account for auditory streaming by a cross‐correlation mechanism modeled as an array of cortical columns. A column contains excilatory and inhibitory interneurons and pyramidal cells which receive input from both the thalamus and from neighboring columns. For columns with coherent thalamic inputs, the outputs of the pyramids sum across frequency because local interneurons compute a correlation between the AM transform of the inputs of the local and remote columns, and selectively gate the input to the pyramidal cell. (1) Grouping by frequency proximity—for columns separated by less than a critical band, thalamic inputs are unresolved. (2) Grouping by temporal proximity—at repetition rates below the mean of the AM distribution, AM fundamentals are less well represented; at higher repetition rates the AM harmonics are more separated. (3) Temporal development—w...

The notion that there is an intimate relationship between musical motion and physical movement is an old one and can be traced back to antiquity. Recently this idea has again received some attention, particularly in relation to musical expression. To use a modern metaphor, one can consider expressive performance to be analogous to the problems of kinematics and trajectory planning in robotics. The trajectories referred to, however, are not those of the performers limbs in physical space, but those of an abstract movement relative to a metrical grid associated with a musical score. Recent studies have attempted to substantiate this idea by comparing a model of motion with timing measurements of the final ritardandi from actual performances. This study extends these earlier analyses to include the accelerandi as well as the ritardandi from complete performances. One conclusion is that the variation of tempo in music can be reasonably compared with velocity in the equations of elementary mechanics. Further, it is suggested that the origin of metrical space, upon which the motion concept rests, lies in the way the auditory system processes rhythm.

In this paper we report the results of an experiment to investigate the emergence of ocular vestibular evoked myogenic potentials (OVEMPs) during the linear vestibular ocular reflex (LVOR) evoked by whole-body vibration (WBV). OVEMP and electrooculogram (EOG) montages were employed to record periocular potentials (POPs) from six subjects during WBV in the nasooccipital (NO) axis over a range of frequencies from 0.5 to 64 Hz with approximately constant peak head acceleration of 1.0 ms−2 (i.e., 0.1 g). Measurements were made in two context conditions: a fixation context to examine the effect of gaze eccentricity (0 vs. 20°), and a visual context, where a target was either head-fixed or earth-fixed. The principal results are that from 0.5 to 2 Hz POP magnitude in the earth-fixed condition is related to head displacement, so with constant acceleration at all frequencies it reduces with increasing frequency, but at frequencies greater than 2 Hz both POP magnitude and POP gain, defined as the ratio of POP magnitude at 20 and 0°, increase with increasing frequency. By exhibiting this high-pass characteristic, a property shared with the LVOR, the results are consistent with the hypothesis that the OVEMP, as commonly employed in the clinical setting, is a high-frequency manifestation of the LVOR. However, we also observed low-frequency acceleration following POPs in head-fixed conditions, consistent with a low-frequency OVEMP, and found evidence of a high-frequency visual context effect, which is also consistent with the OVEMP being a manifestation of the LVOR.

In this paper a new theory of rhythmic grouping is proposed which is based on a multitime scale decomposition of the auditory nerve response. The theory has been inspired by the use of multiscale analysis in the theory of vision as proposed by Marr. It is shown how this multiscale mechanism can be interpreted as a form of echoic memory. A product of the theory is a representation of the rhythmic grouping a sequence of events referred to as a rhythmogram. As well as acting as a perceptual representation the rhythmogram is a powerful tool for the analysis of a range of rhythmic signals including music and speech.

A computational model of musical dynamics is proposed that complements an earlier model of expressive timing. The model, implemented in the artificial intelligence language LISP, is based on the observation that a musical phrase is often indicated by a crescendo/decrescendo shape. The functional form of this shape is derived by making two main assumptions. First, that musical dynamics and tempo are coupled, that is, ‘‘the faster the louder, the slower the softer.’’ This tempo/dynamics coupling, it is suggested, may be a characteristic of some classical and romantic styles perhaps exemplified by performances of Chopin. Second, that the tempo change is governed by analogy to physical movement. The allusion of musical expression to physical motion is further extended by the introduction of the concepts of energy and mass. The utility of the model, in addition to giving an insight into the nature of musical expression, is that it provides a basis for a method of performance style analysis.

The vestibular system responds to head acceleration by producing compensatory reflexes in the eyes and postural muscles. In this study, we investigated the effect of brief interaural acceleration on the vestibular evoked myogenic potential (VEMP) in 10 normal subjects and 10 patients with bilateral (bVL) or unilateral vestibular loss (uVL). The stimuli were delivered with a handheld minishaker and tendon hammer over the mastoid and produced relatively pure interaural head acceleration with little rotation (mean peak acceleration: 0.14 g at 3.3 ms). VEMPs were recorded from the neck muscles and were characterized in normal subjects by a positive/negative potential ipsilateral to the stimulated side (peak latencies: 15.1 and 22.6 ms) and a positive response contralaterally (20.3 ms), which was sometimes preceded by a negativity (14.5 ms). These peaks were absent in patients with bVL, confirming their vestibular dependence. In the patients with uVL, medial acceleration of the intact ear produced bilateral responses, an initial positivity on the intact side, and a negativity on the affected side, whereas lateral acceleration produced only a late positivity on the intact side. As the acceleration was primarily in the horizontal plane, it is likely to have activated utricular receptors. Consistent with this, we found that VEMPs are very sensitive to the direction of head acceleration and have features consistent with the utriculocollic projections demonstrated in animals.

Objective To determine the origin and properties of short latency extraocular potentials produced by activation of the vestibular apparatus using two modes of acoustic stimulation. Methods Extraocular potentials were measured in 10 normal subjects using a bipolar montage to increase selectivity. Three dimensional eye movements were also recorded in five subjects. The subjects were stimulated with both air-conducted (AC) and bone-conducted (BC) sound using a single cycle of a 500 Hz sine wave. Results Short latency positive and negative potentials that peaked at 8.1–12.7 ms for AC and 7.5–13.9 ms for BC stimulation were recorded, which were distinct for the two eyes and for the two modes of stimulation. The extraocular potentials began prior to the onset of eye movements, which peaked at 16.5–20.1 ms for AC, 17.8–25.0 ms for BC stimulation. Conclusions The pattern of short latency eye movements and extraocular potentials induced by AC and BC vestibular stimulation are distinct. As the potentials preceded the eye movements and were not correlated morphologically with them, the source of the observed potentials is not an eye movement and thus we refer to them as ocular vestibular evoked myogenic potentials (OVEMPs). Significance The potentials had properties consistent with modulation of the electromyogenic activity of the extraocular muscles and if interpreted as originating from displacement of the eye will give misleading results. AC and BC acoustic stimulation are likely to activate differing profiles of vestibular end organs.

Measurements are reported of the intensity of the siamang gibbon loud call obtained from the vocal bouts of three family groups at Twycross Zoo, UK. Across 25 samples the maximum intensity ranged from 95 to 113 dB SPL (linear frequency-weighting and fast time-weighting) and exhibited three frequency modes of 250–315 Hz, 630–800 Hz and 1.2–1.6 kHz. The lowest frequency mode, which may correspond to the “boom” sound produced by resonance of the siamang inflated vocal sac, had a mean maximum intensity of 99 dB SPL. These values, which are in excess of the saccular acoustic threshold of about 90 dB at 300 Hz for air conducted sound, suggest that primate loud calls recruit a primitive mode of acoustic sensitivity furnished by the sacculus. Thus reproductive vocal behavior of primates may be influenced by a primitive acoustical reward pathway inherited from a common ancestor with anamniotes. In humans such a pathway could explain the compulsion for exposure to loud music.

In this article the results are reported of an experiment to provide direct evidence for a perceptual and behavioral significance of human saccular acoustic sensitivity. Ten human subjects were stimulated monaurally with 100-ms trains of 10-ms tone pulses with pulse repetition rate of 40 Hz, and were required to rate the pleasantness of the stimuli on a nine-point scale. The design included three within-subject factors: carrier frequency (two levels, 200 and 4000 Hz), intensity [13 levels from 55 to 115 dB(A) in 5-dB steps] and ear (left and right). For intensities above 90 dB myogenic vestibular evoked potentials (MVEP) were also obtained from the ipsilateral sternocleidomastoid muscle from which it was possible to obtain thresholds by linear regression of MVEP amplitudes against intensity. A further between-subjects factor was added which assessed subjects’ attitude to vestibular sensations. The results indicate that across subjects there is a general trend of decreasing pleasantness with increasing intensity, but for the 200-Hz condition there is a significant positive departure from monotonicity in pleasantness (p

Previous research has indicated that an early component of click-evoked myogenic potentials in the sternocleidomastoid muscle is vestibularly mediated, since it can be obtained in subjects with loss of cochlear function, but is absent in subjects with loss of vestibular function (Colebatch et al., 1994). We report here the results of an experiment to investigate whether this response shows any tuning properties. In a sample of 11 subjects, we obtained acoustically evoked EMG from the sternocleidomastoid muscle in response to 110 dB SPL 10 ms tone pips with frequencies of 100 Hz, 200 Hz, 400 Hz, 800 Hz, 1600 Hz and 3200 Hz. The results of this experiment indicate that this response does indeed have a well-defined frequency tuning which may be modelled as a resonance with a maximum response at frequencies between 300-350 Hz. The possible saccular origin of the tuning response and the consequences that this may have in human responses to loud sounds is discussed. Also discussed are the consequences of particular electrode arrangements in relation to the innervation and anatomy of sternocleidomastoid.

Over 70 years ago John Tait raised a question with regards to the certainty that the line of division between the hearing and vestibular portions of the VIIIth cranial nerve had been drawn at the proper place [J. Tait, Ann. Otol. Rhinol. Laryngol. 41, 681–704 (1932)]. His reason for raising this question was the evidence which existed at that time that the otolith organs of anamniotes (fish and amphibians), most commonly the sacculus, were demonstrably organs of hearing in many species. He went on to speculate about a possible auditory role for the otolith organs in amniotes (reptiles, birds, and mammals), including in humans, although at this time there was no direct evidence for this. Since this time, however, there has developed a significant literature documenting evidence, not only for anamniote otolithic hearing, but also for a conserved acoustic sensitivity for the amniote otoliths. In the case of humans, this sensitivity has been exploited for the development of new acoustic tests of vestibular function. The time is ripe 70 years on, therefore, for us to revisit John Tait’s question and explore the possibility that the sacculus may indeed contribute to human hearing. [Work supported by the Royal Society.]