Neural Oscillations and Multisensory Processing.

Neural oscillations play a role in sensory processing by coordinating synchronized neuronal activity. Synchronization of gamma oscillations is engaged in local computation of feedforward signals and synchronization of alpha-beta oscillations is engaged in feedback processing over long-range areas. These spatially and spectrally segregated bi-directional signals may be integrated by a mechanism of cross-frequency coupling. Synchronization of neural oscillations has also been proposed as a mechanism for information integration across multiple sensory modalities. A transient stimulus or rhythmic stimulus from one modality may lead to phase alignment of ongoing neural oscillations in multiple sensory cortices, through a mechanism of cross-modal phase reset or cross-modal neural entrainment. Synchronized activities in multiple sensory cortices are more likely to boost stronger activities in downstream areas. Compared to synchronized oscillations, asynchronized oscillations may impede signal processing, and may contribute to sensory selection by setting the oscillations in the target-related cortex and the oscillations in the distractor-related cortex to opposite phases.

Visual cortex responds to sound onset and offset during passive listening.

Sounds enhance our ability to detect, localize, and respond to co-occurring visual targets. Research suggests that sounds improve visual processing by resetting the phase of ongoing oscillations in visual cortex. However, it remains unclear what information is relayed from the auditory system to visual areas and if sounds modulate visual activity even in the absence of visual stimuli (e.g., during passive listening). Using intracranial electroencephalography (iEEG) in humans, we examined the sensitivity of visual cortex to three forms of auditory information during a passive listening task: auditory onset responses, auditory offset responses, and rhythmic entrainment to sounds. Because some auditory neurons respond to both sound onsets and offsets, visual timing and duration processing may benefit from each. In addition, if auditory entrainment information is relayed to visual cortex, it could support the processing of complex stimulus dynamics that are aligned between auditory and visual stimuli. Results demonstrate that in visual cortex, amplitude-modulated sounds elicited transient onset and offset responses in multiple areas, but no entrainment to sound modulation frequencies. These findings suggest that activity in visual cortex (as measured with iEEG in response to auditory stimuli) may not be affected by temporally fine-grained auditory stimulus dynamics during passive listening (though it remains possible that this signal may be observable with simultaneous auditory-visual stimuli). Moreover, auditory responses were maximal in low-level visual cortex, potentially implicating a direct pathway for rapid interactions between auditory and visual cortices. This mechanism may facilitate perception by time-locking visual computations to environmental events marked by auditory discontinuities.NEW & NOTEWORTHY Using intracranial electroencephalography (iEEG) in humans during a passive listening task, we demonstrate that sounds modulate activity in visual cortex at both the onset and offset of sounds, which likely supports visual timing and duration processing. However, more complex auditory rate information did not affect visual activity. These findings are based on one of the largest multisensory iEEG studies to date and reveal the type of information transmitted between auditory and visual regions.

Causal relationships between neurons of the nucleus incertus and the hippocampal theta activity in the rat.

KEY POINTS: The nucleus incertus is a key node of the brainstem circuitry involved in hippocampal theta rhythmicity. Synchronisation exists between the nucleus incertus and hippocampal activities during theta periods. By the Granger causality analysis, we demonstrated a directional information flow between theta rhythmical neurons in the nucleus incertus and the hippocampus in theta-on states. The electrical stimulation of the nucleus incertus is also able to evoke a phase reset of the hippocampal theta wave. Our data suggest that the nucleus incertus is a key node of theta generation and the modulation network. ABSTRACT: In recent years, a body of evidence has shown that the nucleus incertus (NI), in the dorsal tegmental pons, is a key node of the brainstem circuitry involved in hippocampal theta rhythmicity. Ascending reticular brainstem system activation evokes hippocampal theta rhythm with coupled neuronal activity in the NI. In a recent paper, we showed three populations of neurons in the NI with differential firing during hippocampal theta activation. The objective of this work was to better evaluate the causal relationship between the activity of NI neurons and the hippocampus during theta activation in order to further understand the role of the NI in the theta network. A Granger causality analysis was run to determine whether hippocampal theta activity with sensory-evoked theta depends on the neuronal activity of the NI, or vice versa. The analysis showed causal interdependence between the NI and the hippocampus during theta activity, whose directional flow depended on the different neuronal assemblies of the NI. Whereas type I and II NI neurons mainly acted as receptors of hippocampal information, type III neuronal activity was the predominant source of flow between the NI and the hippocampus in theta states. We further determined that the electrical activation of the NI was able to reset hippocampal waves with enhanced theta-band power, depending on the septal area. Collectively, these data suggest that hippocampal theta oscillations after sensory activation show dependence on NI neuron activity, which could play a key role in establishing optimal conditions for memory encoding.

Visual-tactile processing in primary somatosensory cortex emerges before cross-modal experience.

The presumptive unisensory neocortical areas process multisensory information by oscillatory entrainment of neuronal networks via direct cortico-cortical projections. While neonatal unimodal experience has been identified as necessary for setting up the neuronal networks of multisensory processing, it is still unclear whether early cross-modal experience equally controls the ontogeny of multisensory processing. Here, we assess the development of visual-somatosensory interactions and their anatomical substrate by performing extracellular recordings of network activity in primary sensory cortices in vivo and assessing the cortico-cortical connectivity in pigmented rats. Similar to adult animals, juvenile rats with minimal cross-modal experience display supra-additive augmentation of evoked responses, time-dependent modulation of power and phase reset of network oscillations in response to cross-modal light and whisker stimulation. Moreover, the neuronal discharge of individual neurons is stronger coupled to theta and alpha network oscillations after visual-tactile stimuli. The adult-like multisensory processing of juvenile rats relies on abundant direct visual-somatosensory connections and thalamocortical feedforward interactions. Thus, cellular and network interactions ensuring multisensory processing emerge before cross-modal experience and refine during juvenile development.

Temporal windows in visual processing: "prestimulus brain state" and "poststimulus phase reset" segregate visual transients on different temporal scales.

Dynamic vision requires both stability of the current perceptual representation and sensitivity to the accumulation of sensory evidence over time. Here we study the electrophysiological signatures of this intricate balance between temporal segregation and integration in vision. Within a forward masking paradigm with short and long stimulus onset asynchronies (SOA), we manipulated the temporal overlap of the visual persistence of two successive transients. Human observers enumerated the items presented in the second target display as a measure of the informational capacity read-out from this partly temporally integrated visual percept. We observed higher ß-power immediately before mask display onset in incorrect trials, in which enumeration failed due to stronger integration of mask and target visual information. This effect was timescale specific, distinguishing between segregation and integration of visual transients that were distant in time (long SOA). Conversely, for short SOA trials, mask onset evoked a stronger visual response when mask and targets were correctly segregated in time. Examination of the target-related response profile revealed the importance of an evoked α-phase reset for the segregation of those rapid visual transients. Investigating this precise mapping of the temporal relationships of visual signals onto electrophysiological responses highlights how the stream of visual information is carved up into discrete temporal windows that mediate between segregated and integrated percepts. Fragmenting the stream of visual information provides a means to stabilize perceptual events within one instant in time.

Alpha phase, temporal attention, and the generation of early event related potentials.

In the present study, we have investigated the influence of ongoing alpha phase on the generation of the P1 component of the visual ERP, recorded in a target detection task. Our hypothesis is that in trials where pre- or peristimulus alpha phase is already aligned in a way that voltage positive alpha peaks develop seamlessly into the P1, detection performance will be enhanced as compared to trials where alpha is not aligned. The findings supported our hypothesis and showed that target detection times for the subset of seamless alpha trials was significantly shorter than for trials that are not seamless. Our findings contradict the evoked model for the generation of early ERP components, which rests on the assumption of fixed latency, fixed polarity components. We found that in the non-seamless trials the 'candidate' component of the single trial P1 was at the opposite polarity. Despite this fact, alpha phase locking was at the same high level as was observed for the seamless trials. Finally, we found that prestimulus alpha phase was aligned already in a time window preceding the P1 by 400ms.

Adolescent anxiety and aggression can be differentially predicted by electrocortical phase reset variables.

Increasing evidence supports the notion that both internalizing (e.g., anxiety) and externalizing (e.g., aggression) behavioral dysregulation are associated with abnormal communication between brain regions. Electroencephalographic (EEG) signals across two electrode sites are said to be coherent with one another when they show consistent phase relations. However, periods of desynchrony with shifting of phase relations are a necessary aspect of information processing. The components of EEG phase reset ('locking' when two regions remain in synchrony, and 'shifting' when the two regions desynchronize momentarily) show dramatic changes across development. We collected resting EEG data from typically developing 12 to 15-year-olds and calculated phase shift and lock values in the alpha frequency band across 14 pairs of electrodes varying in inter-electrode distance. A composite measure of participants' aggression levels was positively associated with phase shifting, particularly in the low alpha frequency range, most strongly over the left hemisphere, consistent with the relatively greater left-prefrontal activity reported in aggressive adults. A composite measure of anxiety levels was positively associated with alpha phase locking at sites over both hemispheres, consistent with changes in connectivity reported during anxious thinking in adults. Associations with anxiety could not be explained by traditional EEG coherence measures and suggest that phase shifting and locking might provide an important non-invasive associate of clinically problematic behavior.

Analysis of stimulus-related activity in rat auditory cortex using complex spectral coefficients.

The neural mechanisms of sensory responses recorded from the scalp or cortical surface remain controversial. Evoked vs. induced response components (i.e., changes in mean vs. variance) are associated with bottom-up vs. top-down processing, but trial-by-trial response variability can confound this interpretation. Phase reset of ongoing oscillations has also been postulated to contribute to sensory responses. In this article, we present evidence that responses under passive listening conditions are dominated by variable evoked response components. We measured the mean, variance, and phase of complex time-frequency coefficients of epidurally recorded responses to acoustic stimuli in rats. During the stimulus, changes in mean, variance, and phase tended to co-occur. After the stimulus, there was a small, low-frequency offset response in the mean and modest, prolonged desynchronization in the alpha band. Simulations showed that trial-by-trial variability in the mean can account for most of the variance and phase changes observed during the stimulus. This variability was state dependent, with smallest variability during periods of greatest arousal. Our data suggest that cortical responses to auditory stimuli reflect variable inputs to the cortical network. These analyses suggest that caution should be exercised when interpreting variance and phase changes in terms of top-down cortical processing.

Phase resets undermine measures of phase-dependent perception.

As interest increases in the possible effects of the phase of neural oscillations on perception and cognition, new conceptual and methodological challenges arise. One prominent challenge is the stimulus-induced phase reset, which has the capacity to obscure the effects of phase in the postreset period.

Locus coeruleus noradrenergic neurons phase-lock to prefrontal and hippocampal infra-slow rhythms that synchronize to behavioral events.

The locus coeruleus (LC) is the primary source of noradrenergic projections to the forebrain, and, in prefrontal cortex, is implicated in decision-making and executive function. LC neurons phase-lock to cortical infra-slow wave oscillations during sleep. Such infra-slow rhythms are rarely reported in awake states, despite their interest, since they correspond to the time scale of behavior. Thus, we investigated LC neuronal synchrony with infra-slow rhythms in awake rats performing an attentional set-shifting task. Local field potential (LFP) oscillation cycles in prefrontal cortex and hippocampus on the order of 0.4 Hz phase-locked to task events at crucial maze locations. Indeed, successive cycles of the infra-slow rhythms showed different wavelengths, as if they are periodic oscillations that can reset phase relative to salient events. Simultaneously recorded infra-slow rhythms in prefrontal cortex and hippocampus could show different cycle durations as well, suggesting independent control. Most LC neurons (including optogenetically identified noradrenergic neurons) recorded here were phase-locked to these infra-slow rhythms, as were hippocampal and prefrontal units recorded on the LFP probes. The infra-slow oscillations also phase-modulated gamma amplitude, linking these rhythms at the time scale of behavior to those coordinating neuronal synchrony. This would provide a potential mechanism where noradrenaline, released by LC neurons in concert with the infra-slow rhythm, would facilitate synchronization or reset of these brain networks, underlying behavioral adaptation.

Predictive visuo-motor communication through neural oscillations.

The mechanisms coordinating action and perception over time are poorly understood. The sensory cortex needs to prepare for upcoming changes contingent on action, and this requires temporally precise communication that takes into account the variable delays between sensory and motor processing. Several theorists1,2 have proposed synchronization of the endogenous oscillatory activity observed in most regions of the brain3 as the basis for an efficient and flexible communication protocol between distal brain areas,2,4 a concept known as "communication through coherence." Synchronization of endogenous oscillations5,6 occurs after a salient sensory stimulus, such as a flash or a sound,7-11 and after a voluntary action,12-18 and this directly impacts perception, causing performance to oscillate rhythmically over time. Here we introduce a novel fMRI paradigm to probe the neural sources of oscillations, based on the concept of perturbative signals, which overcomes the low temporal resolution of BOLD signals. The assumption is that a synchronized endogenous rhythm will modulate cortical excitability rhythmically, which should be reflected in the BOLD responses to brief stimuli presented at different phases of the oscillation cycle. We record rhythmic oscillations of V1 BOLD synchronized by a simple voluntary action, in phase with behaviorally measured oscillations in visual sensitivity in the theta range. The functional connectivity between V1 and M1 also oscillates at the same rhythm. By demonstrating oscillatory temporal coupling between primary motor and sensory cortices, our results strongly implicate communication through coherence to achieve precise coordination and to encode sensory-motor timing.

Delta phase reset predicts conflict-related changes in P3 amplitude and behavior.

When multiple competing responses are activated, we respond more slowly than if only one response is activated (response conflict). Conflict-induced slowing is reduced for consecutive high-conflict stimuli, an effect known as conflict adaptation. Verguts and Notebaert's (2009) adaptation by binding theory suggests this is due to Hebbian learning of cognitive control, potentiated by the response of the locus coeruleus norepinephrine (NE) system. Phasic activity of the NE system can potentially be measured non-invasively in humans by recording the P3 component of the event-related potential (ERP), and the P3 is sensitive to conflict adaptation. Bouret and Sara's (2005) network reset theory suggests that phasic NE might functionally reset ongoing large-scale network activity, generating synchronous neural population activity like the P3. To examine the possibility that network reset contributes to conflict effects in the P3, we recorded high-density EEG data while subjects performed a flanker task. As expected, conflict and conflict adaptation modulated P3 amplitudes. Brain-behavior correlation analyses indicated that activity during the rise of the P3 was related to RT and predicted RT differences due to conflict. More importantly, phase of delta oscillations not only predicted reaction time differences between low-conflict and high-conflict conditions, but delta phase reset also predicted the amplitude of the P3. Delta oscillations exhibited dominant peaks both pre and post-stimulus, and delta at stimulus onset predicted the post-stimulus ERP, in particular the N2 and P3. This result bridges human EEG with basic mechanisms suggested by computational neural models and invasive patient recordings, namely that salient cognitive events might reset ongoing oscillations leading to the generation of the phase-locked evoked potential. We conclude that partial phase reset is a cortical mechanism involved in monitoring the environment for unexpected events, and this response contributes to conflict effects in the ERP. These results are in line with theories that phasic NE release might reset ongoing cortical activity, leading to the generation of ERP components like the P3.

Synchronisation of Neural Oscillations and Cross-modal Influences.

At any given moment, we receive multiple signals from our different senses. Prior research has shown that signals in one sensory modality can influence neural activity and behavioural performance associated with another sensory modality. Recent human and nonhuman primate studies suggest that such cross-modal influences in sensory cortices are mediated by the synchronisation of ongoing neural oscillations. In this review, we consider two mechanisms proposed to facilitate cross-modal influences on sensory processing, namely cross-modal phase resetting and neural entrainment. We consider how top-down processes may further influence cross-modal processing in a flexible manner, and we highlight fruitful directions for further research.

The Role of Neuronal Oscillations in Visual Active Sensing.

Visual perception is most often studied as a "passive" process in which an observer fixates steadily at point in space so that stimuli can be delivered to the system with spatial precision. Analysis of neuronal signals related to vision is generally keyed to stimulus onset, stimulus movement, etc.; i.e., events external to the observer. In natural "active" vision, however, information is systematically acquired by using eye movements including rapid (saccadic) eye movements, as well as smooth ocular pursuit of moving objects and slower drifts. Here we consider the use of alternating saccades and fixations to gather information from a visual scene. The underlying motor sampling plan contains highly reliable information regarding "where" and "when" the eyes will land, this information can be used predictively to modify firing properties of neurons precisely at the time when this "contextual" information is most useful - when a volley of retinal input enters the system at the onset of each fixation. Analyses focusing on neural events leading to and resulting from shifts in fixation, as well as visual events external to the observer, can provide a more complete and mechanistic understanding of visual information processing. Studies thus far suggest that active vision may be a fundamentally different from that process we usually study with more traditional passive viewing paradigms. In this Perspective we note that active saccadic sampling behavior imposes robust temporal patterning on the activity of neuron ensembles and large-scale neural dynamics throughout the brain's visual pathways whose mechanistic effects on information processing are not yet fully understood. The spatio-temporal sequence of eye movements elicits a succession of temporally predictable quasi-rhythmic sensory inputs, whose encoding is enhanced by entrainment of low frequency oscillations to the rate of eye movements. Review of the pertinent findings underscores the fact that temporal coordination between motor and visual cortices is critical for understanding neural dynamics of active vision and posits that phase entrainment of neuronal oscillations plays a mechanistic role in this process.

Stabilization of a Cart Inverted Pendulum: Improving the Intermittent Feedback Strategy to Match the Limits of Human Performance.

Stabilization of the CIP (Cart Inverted Pendulum) is an analogy to stick balancing on a finger and is an example of unstable tasks that humans face in everyday life. The difficulty of the task grows exponentially with the decrease of the length of the stick and a stick length of 32 cm is considered as a human limit even for well-trained subjects. Moreover, there is a cybernetic limit related to the delay of the multimodal sensory feedback (about 230 ms) that supports a feedback stabilization strategy. We previously demonstrated that an intermittent-feedback control paradigm, originally developed for modeling the stabilization of upright standing, can be applied with success also to the CIP system, but with values of the critical parameters far from the limiting ones (stick length 50 cm and feedback delay 100 ms). The intermittent control paradigm is based on the alternation of on-phases, driven by a proportional/derivative delayed feedback controller, and off-phases, where the feedback is switched off and the motion evolves according to the intrinsic dynamics of the CIP. In its standard formulation, the switching mechanism consists of a simple threshold operator: the feedback control is switched off if the current (delayed) state vector is closer to the stable than to the unstable manifold of the off-phase and is switched on in the opposite case. Although this simple formulation is effective for explaining upright standing as well as CIP balancing, it fails in the most challenging configuration of the CIP. In this work we propose a modification of the standard intermittent control policy that focuses on the explicit selection of switching times and is based on the phase reset of the estimated state vector at each switching time and on the simulation of an approximated internal model of CIP dynamics. We demonstrate, by simulating the modified intermittent control policy, that it can match the limits of human performance, while operating near the edge of instability.

An Oscillator Ensemble Model of Sequence Learning.

Learning and memorizing sequences of events is an important function of the human brain and the basis for forming expectations and making predictions. Learning is facilitated by repeating a sequence several times, causing rhythmic appearance of the individual sequence elements. This observation invites to consider the resulting multitude of rhythms as a spectral "fingerprint" which characterizes the respective sequence. Here we explore the implications of this perspective by developing a neurobiologically plausible computational model which captures this "fingerprint" by attuning an ensemble of neural oscillators. In our model, this attuning process is based on a number of oscillatory phenomena that have been observed in electrophysiological recordings of brain activity like synchronization, phase locking, and reset as well as cross-frequency coupling. We compare the learning properties of the model with behavioral results from a study in human participants and observe good agreement of the errors for different levels of complexity of the sequence to be memorized. Finally, we suggest an extension of the model for processing sequences that extend over several sensory modalities.

Reducing the Effect of Spurious Phase Variations in Neural Oscillatory Signals.

The phase-reset model of oscillatory EEG activity has received a lot of attention in the last decades for decoding different cognitive processes. Based on this model, the ERPs are assumed to be generated as a result of phase reorganization in ongoing EEG. Alignment of the phase of neuronal activities can be observed within or between different assemblies of neurons across the brain. Phase synchronization has been used to explore and understand perception, attentional binding and considering it in the domain of neuronal correlates of consciousness. The importance of the topic and its vast exploration in different domains of the neuroscience presses the need for appropriate tools and methods for measuring the level of phase synchronization of neuronal activities. Measuring the level of instantaneous phase (IP) synchronization has been used extensively in numerous studies of ERPs as well as oscillatory activity for a better understanding of the underlying cognitive binding with regard to different set of stimulations such as auditory and visual. However, the reliability of results can be challenged as a result of noise artifact in IP. Phase distortion due to environmental noise artifacts as well as different pre-processing steps on signals can lead to generation of artificial phase jumps. One of such effects presented recently is the effect of low envelope on the IP of signal. It has been shown that as the instantaneous envelope of the analytic signal approaches zero, the variations in the phase increase, effectively leading to abrupt transitions in the phase. These abrupt transitions can distort the phase synchronization results as they are not related to any neurophysiological effect. These transitions are called spurious phase variation. In this study, we present a model to remove generated artificial phase variations due to the effect of low envelope. The proposed method is based on a simplified form of a Kalman smoother, that is able to model the IP behavior in narrow-bandpassed oscillatory signals. In this work we first explain the details of the proposed Kalman smoother for modeling the dynamics of the phase variations in narrow-bandpassed signals and then evaluate it on a set of synthetic signals. Finally, we apply the model on ongoing-EEG signals to assess the removal of spurious phase variations.

Auditory cortex responses to interaural time differences in the envelope of low-frequency sound, recorded with MEG in young and older listeners.

Interaural time and intensity differences (ITD and IID) are important cues in binaural hearing and allow for sound localization, improving speech understanding in noise and reverberation, and integrating sound sources in the auditory scene. Whereas previous research showed that the upper-frequency limit for ITD detection in the fine structure of sound declines in aging, the processing of envelope ITD in low-frequency amplitude modulated (AM) sound and the related brain responses are less understood. This study investigated the cortical processing of envelope ITD and compared the results with previous findings about the fine-structure ITD. In two experiments, participants listened to 40-Hz AM tones containing sudden changes in the envelope ITD. Multiple MEG responses were analyzed, including the auditory evoked N1 responses, elicited both by sound onsets and ITD changes, and 40-Hz responses, elicited by the AM. The first experiment with healthy young adults revealed a substantial decline in the magnitudes of the ITD change N1 response, and the 40-Hz phase resets at higher carrier frequencies, suggesting a similar frequency characteristic as observed for fine structure ITD. The amplitude of the 40-Hz ASSR declined only gradually with increasing carrier frequency, and it was excluded as a confounding factor in the decline in the ITD response. Larger responses to outward ITD changes than inward changes, here first reported for envelope ITD, were another characteristics that were similar to fine-structure ITD. A second experiment with groups of young and older listeners examined the effects of aging and concurrent noise on the cortical envelope ITD responses. One important research question was, whether binaural cues are accessible in noise. Behavioural tests showed an age-related hearing loss in the older group and decreased performance in envelope ITD detection and speech-in-noise (SIN) understanding. Binaural hearing and SIN performance were correlated with one other, but not with hearing loss. The frequency limit for envelope ITD was reduced in older listeners similarly as previously found for fine structure ITD, and older listeners were more susceptible to concurrent multi-talker noise. The similarities between responses to envelope ITD and to fine structure ITD suggest that a common cortical code exists for the envelope and fine structure ITD. The dependency on the carrier frequency suggests that envelope ITD processing at the subcortical level requires stimulus phase locking, which might be reduced in aging.

Ripe for solution: Delayed development of multisensory processing in autism and its remediation.

Difficulty integrating inputs from different sensory sources is commonly reported in individuals with Autism Spectrum Disorder (ASD). Accumulating evidence consistently points to altered patterns of behavioral reactions and neural activity when individuals with ASD observe or act upon information arriving through multiple sensory systems. For example, impairments in the integration of seen and heard speech appear to be particularly acute, with obvious implications for interpersonal communication. Here, we explore the literature on multisensory processing in autism with a focus on developmental trajectories. While much remains to be understood, some consistent observations emerge. Broadly, sensory integration deficits are found in children with an ASD whereas these appear to be much ameliorated, or even fully recovered, in older teenagers and adults on the spectrum. This protracted delay in the development of multisensory processing raises the possibility of applying early intervention strategies focused on multisensory integration, to accelerate resolution of these functions. We also consider how dysfunctional cross-sensory oscillatory neural communication may be one key pathway to impaired multisensory processing in ASD.

A Corticothalamic Circuit for Dynamic Switching between Feature Detection and Discrimination.

Sensory processing must be sensitive enough to encode faint signals near the noise floor but selective enough to differentiate between similar stimuli. Here we describe a layer 6 corticothalamic (L6 CT) circuit in the mouse auditory forebrain that alternately biases sound processing toward hypersensitivity and improved behavioral sound detection or dampened excitability and enhanced sound discrimination. Optogenetic activation of L6 CT neurons could increase or decrease the gain and tuning precision in the thalamus and all layers of the cortical column, depending on the timing between L6 CT activation and sensory stimulation. The direction of neural and perceptual modulation - enhanced detection at the expense of discrimination or vice versa - arose from the interaction of L6 CT neurons and subnetworks of fast-spiking inhibitory neurons that reset the phase of low-frequency cortical rhythms. These findings suggest that L6 CT neurons contribute to the resolution of the competing demands of detection and discrimination.