Challenges and Approaches in the Study of Neural Entrainment.

When exposed to rhythmic stimulation, the human brain displays rhythmic activity across sensory modalities and regions. Given the ubiquity of this phenomenon, how sensory rhythms are transformed into neural rhythms remains surprisingly inconclusive. An influential model posits that endogenous oscillations entrain to external rhythms, thereby encoding environmental dynamics and shaping perception. However, research on neural entrainment faces multiple challenges, from ambiguous definitions to methodological difficulties when endogenous oscillations need to be identified and disentangled from other stimulus-related mechanisms that can lead to similar phase-locked responses. Yet, recent years have seen novel approaches to overcome these challenges, including computational modeling, insights from dynamical systems theory, sophisticated stimulus designs, and study of neuropsychological impairments. This review outlines key challenges in neural entrainment research, delineates state-of-the-art approaches, and integrates findings from human and animal neurophysiology to provide a broad perspective on the usefulness, validity, and constraints of oscillatory models in brain-environment interaction.

Phase Alignment of Low-Frequency Neural Activity to the Amplitude Envelope of Speech Reflects Evoked Responses to Acoustic Edges, Not Oscillatory Entrainment.

The amplitude envelope of speech is crucial for accurate comprehension. Considered a key stage in speech processing, the phase of neural activity in the theta-delta bands (1-10 Hz) tracks the phase of the speech amplitude envelope during listening. However, the mechanisms underlying this envelope representation have been heavily debated. A dominant model posits that envelope tracking reflects entrainment of endogenous low-frequency oscillations to the speech envelope. Alternatively, envelope tracking reflects a series of evoked responses to acoustic landmarks within the envelope. It has proven challenging to distinguish these two mechanisms. To address this, we recorded MEG while participants (n = 12, 6 female) listened to natural speech, and compared the neural phase patterns to the predictions of two computational models: an oscillatory entrainment model and a model of evoked responses to peaks in the rate of envelope change. Critically, we also presented speech at slowed rates, where the spectro-temporal predictions of the two models diverge. Our analyses revealed transient theta phase-locking in regular speech, as predicted by both models. However, for slow speech, we found transient theta and delta phase-locking, a pattern that was fully compatible with the evoked response model but could not be explained by the oscillatory entrainment model. Furthermore, encoding of acoustic edge magnitudes was invariant to contextual speech rate, demonstrating speech rate normalization of acoustic edge representations. Together, our results suggest that neural phase-locking to the speech envelope is more likely to reflect discrete representation of transient information rather than oscillatory entrainment.SIGNIFICANCE STATEMENT This study probes a highly debated topic in speech perception: the neural mechanisms underlying the cortical representation of the temporal envelope of speech. It is well established that the slow intensity profile of the speech signal, its envelope, elicits a robust brain response that "tracks" these envelope fluctuations. The oscillatory entrainment model posits that envelope tracking reflects phase alignment of endogenous neural oscillations. Here the authors provide evidence for a distinct mechanism. They show that neural speech envelope tracking arises from transient evoked neural responses to rapid increases in the speech envelope. Explicit computational modeling provides direct and compelling evidence that evoked responses are the primary mechanism underlying cortical speech envelope representations, with no evidence for oscillatory entrainment.

Mechanisms of sustained perceptual entrainment after stimulus offset.

Temporal alignment of neural activity to rhythmic stimulation has been suggested to result from a resonating internal neural oscillator mechanism, but can also be explained by interval-based temporal prediction. Here, we investigate behavioural and brain responses in the post-stimulation period to compare an oscillatory versus an interval-based account. Hickok et al.'s (2015) behavioural paradigm yielded results that relate to a neural oscillatory entrainment mechanism. We adapted the paradigm to an event-related potential (ERP) suitable design: a periodic sequence was followed, in half of the trials, by near-threshold targets embedded in noise. The targets were played in various phases in relation to the preceding sequences' period. Participants had to detect whether targets were played or not, and their EEG was recorded. Both behavioural results and the P300 component of the ERP were not only partially consistent with an oscillatory mechanism but also partially consistent with an interval-based attentional gain mechanism. Instead, data obtained in the post-entrainment period can best be explained with a combination of both mechanisms.

Differential Effects of Cerebellar Degeneration on Feedforward versus Feedback Control across Speech and Reaching Movements.

Errors that result from a mismatch between predicted movement outcomes and sensory afference are used to correct ongoing movements through feedback control and to adapt feedforward control of future movements. The cerebellum has been identified as a critical part of the neural circuit underlying implicit adaptation across a wide variety of movements (reaching, gait, eye movements, and speech). The contribution of this structure to feedback control is less well understood. Although it has recently been shown in the speech domain that individuals with cerebellar degeneration produce larger online corrections for sensory perturbations than control participants, similar behavior has not been observed in other motor domains. Currently, comparisons across domains are limited by different population samples and potential ceiling effects in existing tasks. To assess the relationship between changes in feedforward and feedback control associated with cerebellar degeneration across motor domains, we evaluated adaptive (feedforward) and compensatory (feedback) responses to sensory perturbations in reaching and speech production in human participants of both sexes with cerebellar degeneration and neurobiologically healthy controls. As expected, the cerebellar group demonstrated impaired adaptation in both reaching and speech. In contrast, the groups did not differ in their compensatory response in either domain. Moreover, compensatory and adaptive responses in the cerebellar group were not correlated within or across motor domains. These results point to a general impairment in feedforward control with spared feedback control in cerebellar degeneration. However, the magnitude of feedforward impairments and potential changes in feedback-based control manifest in a domain-specific manner across individuals.SIGNIFICANCE STATEMENT The cerebellum contributes to feedforward updating of movement in response to sensory errors, but its role in feedback control is less understood. Here, we tested individuals with cerebellar degeneration (CD), using sensory perturbations to assess adaptation of feedforward control and feedback gains during reaching and speech production tasks. The results confirmed that CD leads to reduced adaption in both domains. However, feedback gains were unaffected by CD in either domain. Interestingly, measures of feedforward and feedback control were not correlated across individuals within or across motor domains. Together, these results indicate a general impairment in feedforward control with spared feedback control in CD. However, the magnitude of feedforward impairments manifests in a domain-specific manner across individuals.

Double dissociation of single-interval and rhythmic temporal prediction in cerebellar degeneration and Parkinson's disease.

Predicting the timing of upcoming events is critical for successful interaction in a dynamic world, and is recognized as a key computation for attentional orienting. Temporal predictions can be formed when recent events define a rhythmic structure, as well as in aperiodic streams or even in isolation, when a specified interval is known from previous exposure. However, whether predictions in these two contexts are mediated by a common mechanism, or by distinct, context-dependent mechanisms, is highly controversial. Moreover, although the basal ganglia and cerebellum have been linked to temporal processing, the role of these subcortical structures in temporal orienting of attention is unclear. To address these issues, we tested individuals with cerebellar degeneration or Parkinson's disease, with the latter serving as a model of basal ganglia dysfunction, on temporal prediction tasks in the subsecond range. The participants performed a visual detection task in which the onset of the target was predictable, based on either a rhythmic stream of stimuli, or a single interval, specified by two events that occurred within an aperiodic stream. Patients with cerebellar degeneration showed no benefit from single-interval cuing but preserved benefit from rhythm cuing, whereas patients with Parkinson's disease showed no benefit from rhythm cuing but preserved benefit from single-interval cuing. This double dissociation provides causal evidence for functionally nonoverlapping mechanisms of rhythm- and interval-based temporal prediction for attentional orienting, and establishes the separable contributions of the cerebellum and basal ganglia to these functions, suggesting a mechanistic specialization across timing domains.

Neural mechanisms of rhythm-based temporal prediction: Delta phase-locking reflects temporal predictability but not rhythmic entrainment.

Predicting the timing of upcoming events enables efficient resource allocation and action preparation. Rhythmic streams, such as music, speech, and biological motion, constitute a pervasive source for temporal predictions. Widely accepted entrainment theories postulate that rhythm-based predictions are mediated by synchronizing low-frequency neural oscillations to the rhythm, as indicated by increased phase concentration (PC) of low-frequency neural activity for rhythmic compared to random streams. However, we show here that PC enhancement in scalp recordings is not specific to rhythms but is observed to the same extent in less periodic streams if they enable memory-based prediction. This is inconsistent with the predictions of a computational entrainment model of stronger PC for rhythmic streams. Anticipatory change in alpha activity and facilitation of electroencephalogram (EEG) manifestations of response selection are also comparable between rhythm- and memory-based predictions. However, rhythmic sequences uniquely result in obligatory depression of preparation-related premotor brain activity when an on-beat event is omitted, even when it is strategically beneficial to maintain preparation, leading to larger behavioral costs for violation of prediction. Thus, while our findings undermine the validity of PC as a sign of rhythmic entrainment, they constitute the first electrophysiological dissociation, to our knowledge, between mechanisms of rhythmic predictions and of memory-based predictions: the former obligatorily lead to resonance-like preparation patterns (that are in line with entrainment), while the latter allow flexible resource allocation in time regardless of periodicity in the input. Taken together, they delineate the neural mechanisms of three distinct modes of preparation: continuous vigilance, interval-timing-based prediction and rhythm-based prediction.

Consensus paper: Decoding the Contributions of the Cerebellum as a Time Machine. From Neurons to Clinical Applications.

Time perception is an essential element of conscious and subconscious experience, coordinating our perception and interaction with the surrounding environment. In recent years, major technological advances in the field of neuroscience have helped foster new insights into the processing of temporal information, including extending our knowledge of the role of the cerebellum as one of the key nodes in the brain for this function. This consensus paper provides a state-of-the-art picture from the experts in the field of the cerebellar research on a variety of crucial issues related to temporal processing, drawing on recent anatomical, neurophysiological, behavioral, and clinical research.The cerebellar granular layer appears especially well-suited for timing operations required to confer millisecond precision for cerebellar computations. This may be most evident in the manner the cerebellum controls the duration of the timing of agonist-antagonist EMG bursts associated with fast goal-directed voluntary movements. In concert with adaptive processes, interactions within the cerebellar cortex are sufficient to support sub-second timing. However, supra-second timing seems to require cortical and basal ganglia networks, perhaps operating in concert with cerebellum. Additionally, sensory information such as an unexpected stimulus can be forwarded to the cerebellum via the climbing fiber system, providing a temporally constrained mechanism to adjust ongoing behavior and modify future processing. Patients with cerebellar disorders exhibit impairments on a range of tasks that require precise timing, and recent evidence suggest that timing problems observed in other neurological conditions such as Parkinson's disease, essential tremor, and dystonia may reflect disrupted interactions between the basal ganglia and cerebellum.The complex concepts emerging from this consensus paper should provide a foundation for further discussion, helping identify basic research questions required to understand how the brain represents and utilizes time, as well as delineating ways in which this knowledge can help improve the lives of those with neurological conditions that disrupt this most elemental sense. The panel of experts agrees that timing control in the brain is a complex concept in whom cerebellar circuitry is deeply involved. The concept of a timing machine has now expanded to clinical disorders.

When Synchronizing to Rhythms Is Not a Good Thing: Modulations of Preparatory and Post-Target Neural Activity When Shifting Attention Away from On-Beat Times of a Distracting Rhythm.

UNLABELLED: Environmental rhythms potently drive predictive resource allocation in time, typically leading to perceptual and motor benefits for on-beat, relative to off-beat, times, even if the rhythmic stream is not intentionally used. In two human EEG experiments, we investigated the behavioral and electrophysiological expressions of using rhythms to direct resources away from on-beat times. This allowed us to distinguish goal-directed attention from the automatic capture of attention by rhythms. The following three conditions were compared: (1) a rhythmic stream with targets appearing frequently at a fixed off-beat position; (2) a rhythmic stream with targets appearing frequently at on-beat times; and (3) a nonrhythmic stream with matched target intervals. Shifting resources away from on-beat times was expressed in the slowing of responses to on-beat targets, but not in the facilitation of off-beat targets. The shifting of resources was accompanied by anticipatory adjustment of the contingent negative variation (CNV) buildup toward the expected off-beat time. In the second experiment, off-beat times were jittered, resulting in a similar CNV adjustment and also in preparatory amplitude reduction of beta-band activity. Thus, the CNV and beta activity track the relevance of time points and not the rhythm, given sufficient incentive. Furthermore, the effects of task relevance (appearing in a task-relevant vs irrelevant time) and rhythm (appearing on beat vs off beat) had additive behavioral effects and also dissociable neural manifestations in target-evoked activity: rhythm affected the target response as early as the P1 component, while relevance affected only the later N2 and P3. Thus, these two factors operate by distinct mechanisms. SIGNIFICANCE STATEMENT: Rhythmic streams are widespread in our environment, and are typically conceptualized as automatic, bottom-up resource attractors to on-beat times-preparatory neural activity peaks at rhythm-on-beat times and behavioral benefits are seen to on-beat compared with off-beat targets. We show that this behavioral benefit is reversed when targets are more frequent at off-beat compared with on-beat times, and that preparatory neural activity, previously thought to be driven by the rhythm to on-beat times, is adjusted toward off-beat times. Furthermore, the effect of this relevance-based shifting on target-evoked brain activity was dissociable from the automatic effect of rhythms. Thus, rhythms can act as cues for flexible resource allocation according to the goal relevance of each time point, instead of being obligatory resource attractors.

Evidence of Change of Intention in Picking Situations.

Intending to perform an action and then immediately executing it is a mundane process. The cognitive and neural mechanisms involved in this process of "proximal" intention formation and execution, in the face of multiple options to choose from, are not clear, however. Especially, it is not clear how intentions are formed when the choice makes no difference. Here we used behavioral and electrophysiological measures to investigate the temporal dynamics of proximal intention formation and "change of intention" in a free picking scenario, in which the alternatives are on a par for the participant. Participants pressed a right or left button following either an instructive visible arrow cue or a visible neutral "free-choice" cue, both preceded by a masked arrow prime. The goal of the prime was to induce a bias toward pressing the left or right button. Presumably, when the choice is arbitrary, such bias should determine the decision. EEG lateralized readiness potentials and EMG measurements revealed that the prime indeed induced an intention to move in one direction. However, we discovered a signature of "change of intention" in both the Instructed and Free-choice decisions. These results suggest that, even in arbitrary choices, biases present in the neural system for choosing one or another option may be overruled and point to a curious "picking deliberation" phenomenon. We discuss a possible neural scenario that could explain this phenomenon.

Automatic bias of temporal expectations following temporally regular input independently of high-level temporal expectation.

Exposure to rhythmic stimulation results in facilitated responses to events that appear in-phase with the rhythm and modulation of anticipatory and target-evoked brain activity, presumably reflecting "exogenous," unintentional temporal expectations. However, the extent to which this effect is independent from intentional processes is not clear. In two EEG experiments, we isolated the unintentional component of this effect from high-level, intentional factors. Visual targets were presented either in-phase or out-of-phase with regularly flickering colored stimuli. In different blocks, the rhythm could be predictive (i.e., high probability for in-phase target) or not, and the color could be predictive (i.e., validly cue the interval to the target) or not. Exposure to nonpredictive rhythms resulted in faster responses for in-phase targets, even when the color predicted specific out-of-phase target times. Also, the contingent negative variation, an EEG component reflecting temporal anticipation, followed the interval of the nonpredictive rhythm and not that of the predictive color. Thus, rhythmic stimulation unintentionally induced expectations, even when this was detrimental. Intentional usage of predictive rhythms to form expectations resulted in a stronger behavioral effect, and only predictive cues modulated the latency of the target-evoked P3, presumably reflecting stimulus evaluation. These findings establish the existence of unintentional temporal expectations in rhythmic contexts, dissociate them from intentional expectations, and highlight the need to distinguish between the source of expectation (exogenous-endogenous) and the level of voluntary control involved in it (unintentional-intentional).

Contextual effects in sensorimotor adaptation adhere to associative learning rules.

Traditional associative learning tasks focus on the formation of associations between salient events and arbitrary stimuli that predict those events. This is exemplified in cerebellar-dependent delay eyeblink conditioning, where arbitrary cues such as a tone or light act as conditioned stimuli (CSs) that predict aversive sensations at the cornea (unconditioned stimulus [US]). Here, we ask if a similar framework could be applied to another type of cerebellar-dependent sensorimotor learning - sensorimotor adaptation. Models of sensorimotor adaptation posit that the introduction of an environmental perturbation results in an error signal that is used to update an internal model of a sensorimotor map for motor planning. Here, we take a step toward an integrative account of these two forms of cerebellar-dependent learning, examining the relevance of core concepts from associative learning for sensorimotor adaptation. Using a visuomotor adaptation reaching task, we paired movement-related feedback (US) with neutral auditory or visual contextual cues that served as CSs. Trial-by-trial changes in feedforward movement kinematics exhibited three key signatures of associative learning: differential conditioning, sensitivity to the CS-US interval, and compound conditioning. Moreover, after compound conditioning, a robust negative correlation was observed between responses to the two elemental CSs of the compound (i.e. overshadowing), consistent with the additivity principle posited by theories of associative learning. The existence of associative learning effects in sensorimotor adaptation provides a proof-of-concept for linking cerebellar-dependent learning paradigms within a common theoretical framework.

Left hemisphere dominance for bilateral kinematic encoding in the human brain.

Neurophysiological studies in humans and nonhuman primates have revealed movement representations in both the contralateral and ipsilateral hemispheres. Inspired by clinical observations, we ask if this bilateral representation differs for the left and right hemispheres. Electrocorticography was recorded in human participants during an instructed-delay reaching task, with movements produced with either the contralateral or ipsilateral arm. Using a cross-validated kinematic encoding model, we found stronger bilateral encoding in the left hemisphere, an effect that was present during preparation and was amplified during execution. Consistent with this asymmetry, we also observed better across-arm generalization in the left hemisphere, indicating similar neural representations for right and left arm movements. Notably, these left hemisphere electrodes were centered over premotor and parietal regions. The more extensive bilateral encoding in the left hemisphere adds a new perspective to the pervasive neuropsychological finding that the left hemisphere plays a dominant role in praxis.

The brain is split into two hemispheres, each playing the leading role in coordinating movement for the opposite side of the body: lesions on the left hemisphere therefore often result in difficulties moving the right arm or leg, and vice versa. In fact, very few anatomical connections exist between a given hemisphere and the body parts on the same (or 'ipsilateral') side. Yet, movements produced with only one limb still engage both sides of the brain, with the hemisphere which does not control the action production, still encoding the direction and speed of the movement. Previous evidence also indicate that the two hemispheres may not have equal roles when coordinating ipsilateral movements. Merrick et al. aimed to shed light on these processes; to do so, they measured electrical activity from the surface of the brain of six patients as they moved their arms to reach a screen. The results revealed that, while the right hemisphere only encoded information about the opposite arm, the left hemisphere contained information about both arms. Finer analyses showed that, for both hemispheres, moving the opposite arm was strongly associated with activity in the primary motor cortex, a region which helps to execute movements. However, in the left hemisphere, movements from the ipsilateral arm were related to activity in brain areas involved in planning and integrating different types of sensory information. These findings contribute to a better understanding of how the motor system works, which could ultimately help with the development of brain-machine interfaces for patients who need a neuroprosthetic limb.

Integration without awareness: expanding the limits of unconscious processing.

Human conscious awareness is commonly seen as the climax of evolution. However, what function-if any-it serves in human behavior is still debated. One of the leading suggestions is that the cardinal function of conscious awareness is to integrate numerous inputs-including the multitude of features and objects in a complex scene-across different levels of analysis into a unified, coherent, and meaningful perceptual experience. Here we demonstrate, however, that integration of objects with their background scenes can be achieved without awareness of either. We used a binocular rivalry technique known as continuous flash suppression to induce perceptual suppression in a group of human observers. Complex scenes that included incongruent objects escaped perceptual suppression faster than normal scenes did. We conclude that visual awareness is not needed for object-background integration or for processing the likelihood of an object to appear within a given semantic context, but may be needed for dealing with novel situations.

The human cerebellum is essential for modulating perceptual sensitivity based on temporal expectations.

A functional benefit of attention is to proactively enhance perceptual sensitivity in space and time. Although attentional orienting has traditionally been associated with cortico-thalamic networks, recent evidence has shown that individuals with cerebellar degeneration (CD) show a reduced reaction time benefit from cues that enable temporal anticipation. The present study examined whether the cerebellum contributes to the proactive attentional modulation in time of perceptual sensitivity. We tested CD participants on a non-speeded, challenging perceptual discrimination task, asking if they benefit from temporal cues. Strikingly, the CD group showed no duration-specific perceptual sensitivity benefit when cued by repeated but aperiodic presentation of the target interval. In contrast, they performed similar to controls when cued by a rhythmic stream. This dissociation further specifies the functional domain of the cerebellum and establishes its role in the attentional adjustment of perceptual sensitivity in time in addition to its well-documented role in motor timing.

Context-specific control over the neural dynamics of temporal attention by the human cerebellum.

Physiological methods have identified a number of signatures of temporal prediction, a core component of attention. While the underlying neural dynamics have been linked to activity within cortico-striatal networks, recent work has shown that the behavioral benefits of temporal prediction rely on the cerebellum. Here, we examine the involvement of the human cerebellum in the generation and/or temporal adjustment of anticipatory neural dynamics, measuring scalp electroencephalography in individuals with cerebellar degeneration. When the temporal prediction relied on an interval representation, duration-dependent adjustments were impaired in the cerebellar group compared to matched controls. This impairment was evident in ramping activity, beta-band power, and phase locking of delta-band activity. These same neural adjustments were preserved when the prediction relied on a rhythmic stream. Thus, the cerebellum has a context-specific causal role in the adjustment of anticipatory neural dynamics of temporal prediction, providing the requisite modulation to optimize behavior.

Neural entrainment and network resonance in support of top-down guided attention.

Which neural mechanisms provide the functional basis of top-down guided cognitive control? Here, we review recent evidence that suggest that the neural basis of attention is inherently rhythmic. In particular, we discuss two physical properties of self-sustained networks, namely entrainment and resonance, and how these shape the timescale of attentional control. Several recent findings revealed theta-band (3-8 Hz) dynamics in top-down guided behavior. These reports were paralleled by intracranial recordings, which implicated theta oscillations in the organization of functional attention networks. We discuss how the intrinsic network architecture shapes covert attentional sampling as well as overt behavior. Taken together, we posit that theta rhythmicity is an inherent feature of the attention network in support of top-down guided goal-directed behavior.

Taxonomies of Timing: Where Does the Cerebellum Fit In?

Recent models of interval timing have emphasized local, modality-specific processes or a core network centered on a cortico-thalamic-striatal circuit, leaving the role of the cerebellum unclear. We examine this issue, using current taxonomies of timing as a guide to review the association of the cerebellum in motor and perceptual tasks in which timing information is explicit or implicit. Evidence from neuropsychological, neurophysiological, and neuroimaging studies indicates that the involvement of the cerebellum in timing is not restricted to any subdomain of this taxonomy. However, an emerging pattern is that tasks in which timing is done in cyclic continuous contexts do not rely on the cerebellum. In such scenarios, timing may be an emergent property of system dynamics, and especially oscillatory entrainment. The cerebellum may be necessary to time discrete intervals in the absence of continuous cyclic dynamics.

Retroactive memory interference: a potential countermeasure technique against psychophysiological knowledge detection methods.

The concealed information test is designed to detect concealed knowledge through differential physiological responses elicited by the concealed items. This study was designed to examine the role of retroactive interference (RI) as a potential countermeasure that may weaken memory traces of the concealed items and attenuate the physiological responses elicited by them. A total of 120 participants committed a mock crime and were randomly assigned to either an interference condition, where they learned and retrieved an alternative mock crime, or a control condition. Further, each group was randomly assigned to one of three "time-delay" conditions. The results revealed that both memory for the mock crime details and the skin conductance responses (SCRs) to these details were attenuated under the memory-interference condition. Time of testing affected recall, but had no effect on the SCRs. In addition, the memory-interference manipulation had no effect on the respiration measure. Theoretical and practical implications of these results are discussed.

Psychophysiological detection of concealed information shared by groups: an empirical study of the searching CIT.

This study focused on the application of the Concealed Information Test (CIT) to situations in which the crime-related information is shared by a group of suspects but is not available to the investigators (a method known as the "searching CIT," or SCIT). Twenty-two groups, each comprising 4 to 7 participants (115 in total), planned 1 of 2 mock crimes (kidnapping or bank robbery). While planning the crime, each group decided on 5 crime-related critical items (e.g., the city in which the bank was located). Each critical item was chosen from a predefined set of 4 alternatives. At a second stage, the SCIT was administered individually and each participant was tested on the 2 crimes-the actual planned crime, in which the participant was "guilty," and the unplanned crime, in which the participant was "innocent." Two algorithms, adopted from Breska, Ben-Shakhar, and Gronau (2012), were applied to detect the critical items and to differentiate between "guilty" and "innocent" participants. Findings revealed that differentiation efficiency based on electrodermal and respiration measures was identical to that obtained with the standard CIT when applied to large groups, but lower, although significantly greater than chance, when applied to differentiate between small groups.