Responses to brief perturbations of stance: EMG, midline cortical, and subcortical changes.

We studied simultaneous EMG and midline EEG responses, including over the cerebellum, in 10 standing subjects (35 ± 15 yr; 5 females, 5 males). Recordings were made following repeated taps to the sternum, stimuli known to evoke short-latency EMG responses in leg muscles, consistent with postural reflexes. EEG power had relatively more high-frequency components (>30 Hz) when recorded from electrodes over the cerebellum (Iz and SIz) compared with other midline electrodes. We confirmed a previous report using a similar stimulus that evoked short-latency potentials over the cerebellum. We showed clear midline-evoked EEG potentials occurring at short latency over the cerebellum (P23, N31, N42, and P54) and frontally (N28 and N57) before the previously described perturbation-evoked potential (P1/N1/P2). The P23 response correlated with the subsequent EMG response in the tibialis anterior muscles (r = 0.72, P = 0.018), confirming and extending previous observations. We did not find a correlation with the N1 amplitude. We conclude that early activity occurs from electrodes over the inion in response to a brief tap to the sternum. This is likely to represent cerebellar activity and it appears to modulate short-latency postural EMG responses.NEW & NOTEWORTHY We studied the effects of a brief tap to the sternum in human subjects, known to evoke short-latency postural responses. Using an extended EEG recording system, we showed early evoked responses over the midline cerebellum, including the P23 potential, which correlated with the EMG responses in tibialis anterior, consistent with a cerebellar role in postural reflexes. The stimulus also evoked later EEG responses, including the perturbation potential.

Electrophysiological Activity from the Eye Muscles, Cerebellum and Cerebrum During Reflexive (Classical Pavlovian) Versus Voluntary (Ivanov-Smolensky) Eye-Blink Conditioning.

We report an experiment to investigate the role of the cerebellum and cerebrum in motor learning of timed movements. Eleven healthy human subjects were recruited to perform two experiments, the first was a classical eye-blink conditioning procedure with an auditory tone as conditional stimulus (CS) and vestibular unconditional stimulus (US) in the form of a double head-tap. In the second experiment, subjects were asked to blink voluntarily in synchrony with the double head-tap US preceded by a CS, a form of Ivanov-Smolensky conditioning in which a command or instruction is associated with the US. Electrophysiological recordings were made of extra-ocular EMG and EOG at infra-ocular sites (IO1/2), EEG from over the frontal eye fields (C3'/C4') and from over the posterior fossa over the cerebellum for the electrocerebellogram (ECeG). The behavioural outcomes of the experiments showed weak reflexive conditioning for the first experiment despite the double tap but robust, well-synchronised voluntary conditioning for the second. Voluntary conditioned blinks were larger than the reflex ones. For the voluntary conditioning experiment, a contingent negative variation (CNV) was also present in the EEG leads prior to movement, and modulation of the high-frequency EEG occurred during movement. US-related cerebellar activity was prominent in the high-frequency ECeG for both experiments, while conditioned response-related cerebellar activity was additionally present in the voluntary conditioning experiment. These results demonstrate a role for the cerebellum in voluntary (Ivanov-Smolensky) as well as in reflexive (classical Pavlovian) conditioning.

Effects of posture on cerebellar evoked potentials (CEPs) following brief impulsive stimuli at the mastoid and trunk.

Recordings from over the posterior fossa following impulsive acceleration stimuli have shown short latency evoked potentials of presumed cerebellar origin. In this study, we investigated the effect of posture on these cerebellar evoked potentials (CEPs) and their relationship to postural reflexes recorded from the leg muscles evoked by the same stimuli. Nine healthy subjects were tested during lying (supine and prone), sitting and standing. Impulsive accelerations were applied at the mastoid and to truncal (both C7 and sternal) stimulation sites. The effect of vision, eyes open or closed, was investigated for all three stimuli. For the truncal stimuli, the effect of differing leaning conditions during standing was also recorded. CEP amplitudes were correlated for the three stimuli. For C7 stimulation during standing, both CEPs and postural reflexes scaled as the threat to postural stability increased. However, CEPs for all stimuli were present during lying, sitting and standing with amplitude and latency parameters mainly unaffected by posture or vision. In contrast, postural reflexes from the leg muscles were attenuated when not standing, with the effect being more marked for truncal stimuli. We conclude that CEPs evoked by axial and vestibular stimuli are not systematically gated by posture, in contrast to the reflex responses evoked by the same stimuli.

Correction to: Mapping the vestibular cerebellar evoked potential (VsCEP) following air- and bone-conducted vestibular stimulation.

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

Mapping the vestibular cerebellar evoked potential (VsCEP) following air- and bone-conducted vestibular stimulation.

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.

Vestibular cerebellar evoked potentials in humans and their modulation during optokinetic stimulation.

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 inion response revisited: evidence for a possible cerebellar contribution to vestibular-evoked potentials produced by air-conducted sound stimulation.

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 evoked potentials (VsEPs) of cortical origin produced by impulsive acceleration applied at the nasion.

We report the results of a study to record vestibular evoked potentials (VsEPs) of cortical origin produced by impulsive acceleration (IA). In a sample of 12 healthy participants, evoked potentials recorded by 70 channel electroencephalography were obtained by IA stimulation at the nasion and compared with evoked potentials from the same stimulus applied to the forefingers. The nasion stimulation gave rise to a series of positive and negative deflections in the latency range of 26-72 ms, which were dependent on the polarity of the applied IA. In contrast, evoked potentials from the fingers were characterised by a single N50/P50 deflection at about 50 ms and were polarity invariant. Source analysis confirmed that the finger evoked potentials were somatosensory in origin, i.e. were somatosensory evoked potentials, and suggested that the nasion evoked potentials plausibly included vestibular midline and frontal sources, as well as contributions from the eyes, and thus were likely VsEPs. These results show considerable promise as a new method for assessment of the central vestibular system by means of VsEPs produced by IA applied to the head.

Thresholds for vestibular-evoked myogenic potentials (VEMPs) produced by impulsive transmastoid acceleration.

OBJECTIVE: 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. STUDY SAMPLE: Twelve adult subjects with no history of auditory or vestibular deficits. RESULTS: 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. CONCLUSION: VEMPs can be obtained at lower ITA stimulation levels than OVEMPs. Threshold values depend on the method used to define response presence.

Electrophysiological activity from over the cerebellum and cerebrum during eye blink conditioning in human subjects.

We report the results of an experiment in which electrophysiological activity was recorded from the human cerebellum and cerebrum in a sample of 14 healthy subjects before, during and after a classical eye blink conditioning procedure with an auditory tone as conditional stimulus and a maxillary nerve unconditional stimulus. The primary aim was to show changes in the cerebellum and cerebrum correlated with behavioral ocular responses. Electrodes recorded EMG and EOG at peri-ocular sites, EEG from over the frontal eye-fields and the electrocerebellogram (ECeG) from over the posterior fossa. Of the 14 subjects half strongly conditioned while the other half were resistant. We confirmed that conditionability was linked under our conditions to the personality dimension of extraversion-introversion. Inhibition of cerebellar activity was shown prior to the conditioned response, as predicted by Albus (1971). However, pausing in high frequency ECeG and the appearance of a contingent negative variation (CNV) in both central leads occurred in all subjects. These led us to conclude that while conditioned cerebellar pausing may be necessary, it is not sufficient alone to produce overt behavioral conditioning, implying the existence of another central mechanism. The outcomes of this experiment indicate the potential value of the noninvasive electrophysiology of the cerebellum.

Extended source analysis of movement related potentials (MRPs) for self-paced hand and foot movements demonstrates opposing cerebral and cerebellar laterality: a preliminary study.

The cerebellum is known to have extensive reciprocal connectivity with the cerebral cortex, including with prefrontal and posterior parietal cortex, which play an important role on the planning and execution of voluntary movement. In the present article we report an exploratory non-invasive electrophysiological study of the activity of the cerebellum and cerebrum during voluntary finger and foot movements. In a sample of five healthy adult subjects, we recorded EEG and the electro-cerebellogram (ECeG) with a 10% cerebellar extension montage during voluntary left and right index finger and foot movements. EMG was recorded from finger extensors and flexors and from the tibialis anterior and soleus muscles and was used to generate triggers for movement related averaging (-2000 to +2000 ms). Source analysis was conducted over five epochs defined relative to EMG onset: whole epoch (-1000 to +1000 ms), pre-move 1000 (-1000 to 0 ms), pre-move 500 (-500 to 0 ms), post-move 500 (0 to +500 ms) and post-move 1000 (0 to +1000 ms). This yielded a total of 123 cerebral and 65 cerebellar dipole clusters from across all epochs, including the pre-movement epochs, which were then subject to statistical analysis. These demonstrated predominantly contralateral dominance for the cerebral clusters, but predominantly ipsilateral dominance for the cerebellar clusters. In addition, both cerebral and cerebellar clusters showed evidence of a somatotopic gradient, medially (X-axis) for the cerebral clusters, and medially and dorso-ventrally (Z-axis) for the cerebellar clusters. These findings support the value of recording cerebellar ECeG and demonstrate its potential to contribute to understanding cerebellar function.

Collic evoked potentials, myogenic potentials (CEMPs) and postural responses produced by brief 100 Hz vibration of the sternocleidomastoid muscle.

We present an initial report using 5 subjects, of short and long latency collic evoked responses following a half cycle of 100 Hz vibration (5 ms) applied to the sternocleidomastoid (SCM) tendon. These were detected in EEG and extraocular and leg muscles and compared with vestibular-dependent responses from direct mastoid stimulation. The responses from the extraocular recording site are likely to be evoked myogenic potentials, thus "collic evoked myogenic potentials" (CEMPs). An n19/p24 presumed ocular CEMP (oCEMP) was followed by a P22/N28 response over the posterior fossa, referred to as a collic cerebellar evoked potential (CoCEP), with responses in leg muscles starting around 55 ms. In contrast to their vestibular analogues, the oCEMP and CoCEP were predominantly ipsilateral to the side of stimulation, consistent with a double-crossed projection. In addition, their thresholds were just above the threshold of vibrotactile sensation, implying a low threshold, oligo-synaptic projection of SCM afferents to both extraocular and cerebellar targets. Following these short latency responses, SCM tendon stimulation evoked prolonged EMG responses in postural muscles of the legs, consistent with a role in the afferent limb of a short latency, spino-bulbar-spinal postural response to sternal perturbations. These collic evoked responses are likely to be of value in understanding the functions of cervical muscle afferents and have clinical value, for example in monitoring compensation after vestibular loss.

Non-invasive recording from the human cerebellum during a classical conditioning paradigm using the otolith-evoked blink reflex.

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.

Source analyses of axial and vestibular evoked potentials associated with brainstem-spinal reflexes show cerebellar and cortical contributions.

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.

Vestibular evoked myogenic potentials evoked by brief interaural head acceleration: properties and possible origin.

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.

A utricular origin of frequency tuning to low-frequency vibration in the human vestibular system?

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 10ms 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 800Hz whereas vibration showed a low-pass type response with a largest response at 100Hz. Our results suggest that the tuning is at least in part due to properties of end-organs themselves, while the 100Hz best frequency may be a specifically utricular feature.

Ocular vestibular evoked myogenic potentials (OVEMPs) produced by impulsive transmastoid accelerations.

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 human electrocerebellogram (ECeG) recorded non-invasively using scalp electrodes.

The electrocerebellogram (ECeG), a manifestation of cerebellar cortical local field potentials (LFPs), is characterised by a predominance of high frequency components at the gamma range (30-80 Hz) and beyond up to several hundred Hz, in contrast to the electroencephalogram (EEG) which is composed predominantly of lower frequencies at or below the gamma range. Although the potential clinical role of the ECeG has been suggested, the prospect of a non-invasive method for routine recording in humans has remained elusive. Here we show for the first time that high-frequency power characteristic of the ECeG may be easily measurable using surface electrodes placed over the posterior fossa at the approximate original 10/20 CB location. In a sample of six subjects we have compared continuous electrical recordings in a supine posture under visual motion stimulation at CB1/2, about 5% inferior and medial to PO9/10 of the 10-10 system, with those at standard 10/20 locations of C3/4 and O1/2, as well as over the splenius muscles as controls against myogenic contamination. In a neutral baseline condition high-frequency power was significantly greater over the posterior fossa, especially in ultra-gamma (80-160 Hz) and very high frequency (VHF, 160-320 Hz) bands, compared to occipital or central leads. We also found that visual stimulation, in the form of visual motion in particular, was effective in increasing the high-frequency power in CB electrodes, including in beta (14-30 Hz) and gamma, compared with electrodes over the occipital and frontal cortex. We propose that the human ECeG can be recorded non-invasively and may have application both in understanding cerebellar function and clinically.

The sensory-motor theory of rhythm and beat induction 20 years on: a new synthesis and future perspectives.

Some 20 years ago Todd and colleagues proposed that rhythm perception is mediated by the conjunction of a sensory representation of the auditory input and a motor representation of the body (Todd, 1994a, 1995), and that a sense of motion from sound is mediated by the vestibular system (Todd, 1992a, 1993b). These ideas were developed into a sensory-motor theory of rhythm and beat induction (Todd et al., 1999). A neurological substrate was proposed which might form the biological basis of the theory (Todd et al., 2002). The theory was implemented as a computational model and a number of experiments conducted to test it. In the following time there have been several key developments. One is the demonstration that the vestibular system is primal to rhythm perception, and in related work several experiments have provided further evidence that rhythm perception is body dependent. Another is independent advances in imaging, which have revealed the brain areas associated with both vestibular processing and rhythm perception. A third is the finding that vestibular receptors contribute to auditory evoked potentials (Todd et al., 2014a,b). These behavioral and neurobiological developments demand a theoretical overview which could provide a new synthesis over the domain of rhythm perception. In this paper we suggest four propositions as the basis for such a synthesis. (1) Rhythm perception is a form of vestibular perception; (2) Rhythm perception evokes both external and internal guidance of somatotopic representations; (3) A link from the limbic system to the internal guidance pathway mediates the "dance habit"; (4) The vestibular reward mechanism is innate. The new synthesis provides an explanation for a number of phenomena not often considered by rhythm researchers. We discuss these along with possible computational implementations and alternative models and propose a number of new directions for future research.