Direction Of Frequency Modulation Research Articles

Neural modelling of the encoding of fast frequency modulation.

Frequency modulation (FM) is a basic constituent of vocalisation in many animals as well as in humans. In human speech, short rising and falling FM-sweeps of around 50 ms duration, called formant transitions, characterise individual speech sounds. There are two representations of FM in the ascending auditory pathway: a spectral representation, holding the instantaneous frequency of the stimuli; and a sweep representation, consisting of neurons that respond selectively to FM direction. To-date computational models use feedforward mechanisms to explain FM encoding. However, from neuroanatomy we know that there are massive feedback projections in the auditory pathway. Here, we found that a classical FM-sweep perceptual effect, the sweep pitch shift, cannot be explained by standard feedforward processing models. We hypothesised that the sweep pitch shift is caused by a predictive feedback mechanism. To test this hypothesis, we developed a novel model of FM encoding incorporating a predictive interaction between the sweep and the spectral representation. The model was designed to encode sweeps of the duration, modulation rate, and modulation shape of formant transitions. It fully accounted for experimental data that we acquired in a perceptual experiment with human participants as well as previously published experimental results. We also designed a new class of stimuli for a second perceptual experiment to further validate the model. Combined, our results indicate that predictive interaction between the frequency encoding and direction encoding neural representations plays an important role in the neural processing of FM. In the brain, this mechanism is likely to occur at early stages of the processing hierarchy.

Categorical Auditory Working Memory in Crows.

SummaryThe ability to group sensory data into behaviorally meaningful classes and to maintain these perceptual categories active in working memory is key to intelligent behavior. Here, we show that carrion crows, highly vocal and cognitively advanced corvid songbirds, possess categorical auditory working memory. The crows were trained in a delayed match-to-category task that required them to flexibly match remembered sounds based on the upward or downward shift of the sounds' frequency modulation. After training, the crows instantaneously classified novel sounds into the correct auditory categories. The crows showed sharp category boundaries as a function of the relative frequency interval of the modulation. In addition, the crows generalized frequency-modulated sounds within a category and correctly classified novel sounds kept in working memory irrespective of other acoustic features of the sound. This suggests that crows can form and actively memorize auditory perceptual categories in the service of cognitive control of their goal-directed behaviors.

Stimulus-invariant auditory cortex threat encoding during fear conditioning with simple and complex sounds

Learning to predict threat depends on amygdala plasticity and does not require auditory cortex (ACX) when threat predictors (conditioned stimuli, CS) are simple sine tones. However, ACX is required in rodents to learn from some naturally occurring CS. Yet, the precise function of ACX, and whether it differs for different CS types, is unknown. Here, we address how ACX encodes threat predictions during human fear conditioning using functional magnetic resonance imaging (fMRI) with multivariate pattern analysis. As in previous rodent work, CS+ and CS- were defined either by direction of frequency modulation (complex) or by frequency of pure tones (simple). In an instructed non-reinforcement context, different sets of simple and complex sounds were always presented without reinforcement (neutral sounds, NS). Threat encoding was measured by separation of fMRI response patterns induced by CS+/CS-, or similar NS1/NS2 pairs. We found that fMRI patterns in Heschl's gyrus encoded threat prediction over and above encoding the physical stimulus features also present in NS, i.e. CS+/CS- could be separated better than NS1/NS2. This was the case both for simple and complex CS. Furthermore, cross-prediction demonstrated that threat representations were similar for simple and complex CS, and thus unlikely to emerge from stimulus-specific top-down, or learning-induced, receptive field plasticity. Searchlight analysis across the entire ACX demonstrated further threat representations in a region including BA22 and BA42. However, in this region, patterns were distinct for simple and complex sounds, and could thus potentially arise from receptive field plasticity. Strikingly, across participants, individual size of Heschl's gyrus predicted strength of fear learning for complex sounds. Overall, our findings suggest that ACX represents threat predictions, and that Heschl's gyrus contains a threat representation that is invariant across physical stimulus categories.

Modulation of Auditory Evoked Magnetic Fields Elicited by Successive Frequency-Modulated (FM) Sweeps.

In our daily life, we are successively exposed to frequency-modulated (FM) sounds that play an important role in speech and species-specific communication. Previous studies demonstrated that repetitive exposure to identical pure tones resulted in decreased neural activity. However, the effects of repetitively presented FM sounds on neural activity in the human auditory cortex remain unclear. In the present study, we used magnetoencephalography to investigate auditory evoked N1m responses elicited by four successive temporally repeated and superimposed FM sweeps in three sequences: (1) four FM sweeps were identical, (2) four FM sweeps had the same FM direction and rate, but different carrier frequencies, (3) four FM sweeps differed with respect to the FM rate and/or direction and their carrier frequencies. In contrast to our expectations, the results obtained demonstrated that N1m responses were maximal when the four FM sweeps were identical and minimal when they were distinct. These results suggest that the neural processing of repetitive FM sweeps in the human auditory cortex may differ from that of repetitive pure tones.

Sensitivity of human auditory cortex to rapid frequency modulation revealed by multivariate representational similarity analysis.

Functional Magnetic Resonance Imaging (fMRI) was used to investigate the extent, magnitude, and pattern of brain activity in response to rapid frequency-modulated sounds. We examined this by manipulating the direction (rise vs. fall) and the rate (fast vs. slow) of the apparent pitch of iterated rippled noise (IRN) bursts. Acoustic parameters were selected to capture features used in phoneme contrasts, however the stimuli themselves were not perceived as speech per se. Participants were scanned as they passively listened to sounds in an event-related paradigm. Univariate analyses revealed a greater level and extent of activation in bilateral auditory cortex in response to frequency-modulated sweeps compared to steady-state sounds. This effect was stronger in the left hemisphere. However, no regions showed selectivity for either rate or direction of frequency modulation. In contrast, multivoxel pattern analysis (MVPA) revealed feature-specific encoding for direction of modulation in auditory cortex bilaterally. Moreover, this effect was strongest when analyses were restricted to anatomical regions lying outside Heschl's gyrus. We found no support for feature-specific encoding of frequency modulation rate. Differential findings of modulation rate and direction of modulation are discussed with respect to their relevance to phonetic discrimination.

Age-related deterioration of cortical responses to slow FM sounds in the auditory belt region of adult C57BL/6 mice

To compare age-related deterioration of neural responses in each subfield of the auditory cortex in C57BL/6 mice, we evaluated amplitudes of tonal responses in young (5–11 weeks old) and adult (16–23 weeks old) groups using transcranial flavoprotein fluorescence imaging. Cortical responses to 20-kHz amplitude-modulated (AM) sounds, which were mainly found in the anterior auditory field (AAF) and the primary auditory cortex (AI) of the core region, were not markedly different between the two groups. In contrast, cortical responses to direction reversal of slow frequency-modulated (FM) sounds, which were mainly found in the ultrasonic field (UF), were significantly disrupted in the adult group compared with those in the young group. To investigate the mechanisms underlying such age-related deterioration, biotinylated dextran amine (BDA) was injected into UF. The number of retrograde labeled neurons in the dorsal division of the medial geniculate body (MGd) was markedly reduced in the adult group compared with that in the young group. These results strongly suggest that cortical responses to FM direction reversal in UF of adult C57BL/6 mice are mainly deteriorated by loss of non-lemniscal thalamic inputs from MGd to UF due to aging.

Auditory Cortical Areas Activated by Slow Frequency-Modulated Sounds in Mice

Species-specific vocalizations in mice have frequency-modulated (FM) components slower than the lower limit of FM direction selectivity in the core region of the mouse auditory cortex. To identify cortical areas selective to slow frequency modulation, we investigated tonal responses in the mouse auditory cortex using transcranial flavoprotein fluorescence imaging. For differentiating responses to frequency modulation from those to stimuli at constant frequencies, we focused on transient fluorescence changes after direction reversal of temporally repeated and superimposed FM sweeps. We found that the ultrasonic field (UF) in the belt cortical region selectively responded to the direction reversal. The dorsoposterior field (DP) also responded weakly to the reversal. Regarding the responses in UF, no apparent tonotopic map was found, and the right UF responses were significantly larger in amplitude than the left UF responses. The half-max latency in responses to FM sweeps was shorter in UF compared with that in the primary auditory cortex (A1) or anterior auditory field (AAF). Tracer injection experiments in the functionally identified UF and DP confirmed that these two areas receive afferent inputs from the dorsal part of the medial geniculate nucleus (MG). Calcium imaging of UF neurons stained with fura-2 were performed using a two-photon microscope, and the presence of UF neurons that were selective to both direction and direction reversal of slow frequency modulation was demonstrated. These results strongly suggest a role for UF, and possibly DP, as cortical areas specialized for processing slow frequency modulation in mice.

Left auditory cortex is involved in pairwise comparisons of the direction of frequency modulated tones

Evaluating series of complex sounds like those in speech and music requires sequential comparisons to extract task-relevant relations between subsequent sounds. With the present functional magnetic resonance imaging (fMRI) study, we investigated whether sequential comparison of a specific acoustic feature within pairs of tones leads to a change in lateralized processing in the auditory cortex (AC) of humans. For this we used the active categorization of the direction (up vs. down) of slow frequency modulated (FM) tones. Several studies suggest that this task is mainly processed in the right AC. These studies, however, tested only the categorization of the FM direction of each individual tone. In the present study we ask the question whether the right lateralized processing changes when, in addition, the FM direction is compared within pairs of successive tones. For this we use an experimental approach involving contralateral noise presentation in order to explore the contributions made by the left and right AC in the completion of the auditory task. This method has already been applied to confirm the right-lateralized processing of the FM direction of individual tones. In the present study, the subjects were required to perform, in addition, a sequential comparison of the FM direction in pairs of tones. The results suggest a division of labor between the two hemispheres such that the FM direction of each individual tone is mainly processed in the right AC whereas the sequential comparison of this feature between tones in a pair is probably performed in the left AC.

Positive and negative reinforcement activate human auditory cortex.

Prior studies suggest that reward modulates neural activity in sensory cortices, but less is known about punishment. We used functional magnetic resonance imaging and an auditory discrimination task, where participants had to judge the duration of frequency modulated tones. In one session correct performance resulted in financial gains at the end of the trial, in a second session incorrect performance resulted in financial loss. Incorrect performance in the rewarded as well as correct performance in the punishment condition resulted in a neutral outcome. The size of gains and losses was either low or high (10 or 50 Euro cent) depending on the direction of frequency modulation. We analyzed neural activity at the end of the trial, during reinforcement, and found increased neural activity in auditory cortex when gaining a financial reward as compared to gaining no reward and when avoiding financial loss as compared to receiving a financial loss. This was independent on the size of gains and losses. A similar pattern of neural activity for both gaining a reward and avoiding a loss was also seen in right middle temporal gyrus, bilateral insula and pre-supplemental motor area, here however neural activity was lower after correct responses compared to incorrect responses. To summarize, this study shows that the activation of sensory cortices, as previously shown for gaining a reward is also seen during avoiding a loss.

Effects of sound intensity on temporal properties of inhibition in the pallid bat auditory cortex

Auditory neurons in bats that use frequency modulated (FM) sweeps for echolocation are selective for the behaviorally-relevant rates and direction of frequency change. Such selectivity arises through spectrotemporal interactions between excitatory and inhibitory components of the receptive field. In the pallid bat auditory system, the relationship between FM sweep direction/rate selectivity and spectral and temporal properties of sideband inhibition have been characterized. Of note is the temporal asymmetry in sideband inhibition, with low-frequency inhibition (LFI) exhibiting faster arrival times compared to high-frequency inhibition (HFI). Using the two-tone inhibition over time (TTI) stimulus paradigm, this study investigated the interactions between two sound parameters in shaping sideband inhibition: intensity and time. Specifically, the impact of changing relative intensities of the excitatory and inhibitory tones on arrival time of inhibition was studied. Using this stimulation paradigm, single unit data from the auditory cortex of pentobarbital-anesthetized cortex show that the threshold for LFI is on average ~8 dB lower than HFI. For equal intensity tones near threshold, LFI is stronger than HFI. When the inhibitory tone intensity is increased further from threshold, the strength asymmetry decreased. The temporal asymmetry in LFI vs. HFI arrival time is strongest when the excitatory and inhibitory tones are of equal intensities or if excitatory tone is louder. As inhibitory tone intensity is increased, temporal asymmetry decreased suggesting that the relative magnitude of excitatory and inhibitory inputs shape arrival time of inhibition and FM sweep rate and direction selectivity. Given that most FM bats use downward sweeps as echolocation calls, a similar asymmetry in threshold and strength of LFI vs. HFI may be a general adaptation to enhance direction selectivity while maintaining sweep-rate selective responses to downward sweeps.

Intracellular responses to frequency modulated tones in the dorsal cortex of the mouse inferior colliculus

Frequency modulations occur in many natural sounds, including vocalizations. The neuronal response to frequency modulated (FM) stimuli has been studied extensively in different brain areas, with an emphasis on the auditory cortex and the central nucleus of the inferior colliculus. Here, we measured the responses to FM sweeps in whole-cell recordings from neurons in the dorsal cortex of the mouse inferior colliculus. Both up- and downward logarithmic FM sweeps were presented at two different speeds to both the ipsi- and the contralateral ear. Based on the number of action potentials that were fired, between 10 and 24% of cells were selective for rate or direction of the FM sweeps. A somewhat lower percentage of cells, 6–21%, showed selectivity based on EPSP size. To study the mechanisms underlying the generation of FM selectivity, we compared FM responses with responses to simple tones in the same cells. We found that if pairs of neurons responded in a similar way to simple tones, they generally also responded in a similar way to FM sweeps. Further evidence that FM selectivity can be generated within the dorsal cortex was obtained by reconstructing FM sweeps from the response to simple tones using three different models. In about half of the direction selective neurons the selectivity was generated by spectrally asymmetric synaptic inhibition. In addition, evidence for direction selectivity based on the timing of excitatory responses was also obtained in some cells. No clear evidence for the local generation of rate selectivity was obtained. We conclude that FM direction selectivity can be generated within the dorsal cortex of the mouse inferior colliculus by multiple mechanisms.

Development of response selectivity in the mouse auditory cortex

The mouse auditory system contains neurons selective for tone duration and for a narrow range of frequency modulated (FM) sweep rates. Whether such selectivity is developmentally regulated is not known. The main goal of this study was to follow the development of neuronal responses to tones (frequency and duration tuning) and FM sweeps (direction and rate selectivity) in the core auditory cortex (A1 and AAF) of ketamine/xylazine anesthetized C57bl/6 mice. Three groups were compared: postnatal day (P) 15–20, P21–30 and P31–90. Frequency tuning bandwidth decreased during the first month indicating refinement of the excitatory receptive field. Duration tuning for tones did not change during development in terms of categories of tuning types as well as measures of selectivity such as best duration and half-maximal duration. FM rate and direction selectivity were developmentally regulated. Selectivity for linear up and down FM sweeps (0.06–22 kHz/ms) was tested. The best rate and half-maximal rate of neurons categorized as fast- or band-pass selective shifted toward faster rates during development. The percentage of fast-pass selective neurons also increased during development. These data suggest that cortical neurons' discrimination and detection abilities for relatively faster sweep rates improve during development. Although on average, direction selectivity was weak across development, there was a significant shift toward upward sweep selectivity at slow rates. Thus, the C57bl/6 mouse auditory cortex is not adult-like until at least P30. The changes in response selectivity can be explained based on known developmental changes in intrinsic and synaptic properties of mouse auditory cortical neurons.

Selectivity for Spectral Motion as a Neural Computation for Encoding Natural Communication Signals in Bat Inferior Colliculus

This study examines the neural computations performed by neurons in the auditory system to be selective for the direction and velocity of signals sweeping upward or downward in frequency, termed spectral motion. We show that neurons in the auditory midbrain of Mexican free-tailed bats encode multiple spectrotemporal features of natural communication sounds. These features to which each neuron is tuned are nonlinearly combined to produce selectivity for spectral motion cues present in their conspecific calls, such as direction and velocity. We find that the neural computations resulting in selectivity for spectral motion are analogous to models of motion selectivity studied in vision. Our analysis revealed that auditory neurons in the inferior colliculus (IC) are avoiding spectrotemporal modulations that are redundant across different bat communication signals and are specifically tuned for modulations that distinguish each call from another by their frequency-modulated direction and velocity, suggesting that spectral motion is the neural computation through which IC neurons are encoding specific features of conspecific vocalizations.

FM velocity selectivity in the inferior colliculus is inherited from velocity-selective inputs and enhanced by spike threshold

Frequency modulation (FM) is computed from the temporal sequence of activated auditory nerve fibers representing different frequencies. Most studies in the inferior colliculus (IC) have inferred from extracellular recordings that the precise timing of nonselective inputs creates selectivity for FM direction and velocity (Andoni S, Li N, Pollak GD. J Neurosci 27: 4882-4893, 2007; Fuzessery ZM, Richardson MD, Coburn MS. J Neurophysiol 96: 1320-1336, 2006; Gordon M, O'Neill WE. Hear Res 122: 97-108, 1998). We recently reported that two additional mechanisms were more important than input timing for directional selectivity in some IC cells: spike threshold and inputs that were already selective (Gittelman JX, Li N, Pollak GD. J Neurosci 29: 13030-13041, 2009). Here, we show that these same mechanisms, selective inputs and spike threshold, underlie selectivity for FM velocity and intensity. From whole cell recordings in awake bats, we recorded spikes and postsynaptic potentials (PSPs) evoked by downward and upward FMs that swept identical frequencies at different velocities and intensities. To determine the synaptic mechanisms underlying PSP selectivity (relative PSP height), we derived sweep-evoked synaptic conductances. Changing FM velocity or intensity changed conductance timing and size. Modeling indicated that excitatory conductance size contributed more to PSP selectivity than conductance timing, indicating that the number of afferent spikes carried more FM information to the IC than precise spike timing. However, excitation alone produced mostly suprathreshold PSPs. Inhibition reduced absolute PSP heights, without necessarily altering PSP selectivity, thereby rendering some PSPs subthreshold. Spike threshold then sharpened selectivity in the spikes by rectifying the smaller PSPs. This indicates the importance of spike threshold, and that inhibition enhances selectivity via a different mechanism than previously proposed.

Differential roles of GABAergic and glycinergic input on FM selectivity in the inferior colliculus of the pallid bat

Multiple mechanisms have been shown to shape frequency-modulated (FM) selectivity within the central nucleus of the inferior colliculus (IC) in the pallid bat. In this study we focus on the mechanisms associated with sideband inhibition. The relative arrival time of inhibition compared with excitation can be used to predict FM responses as measured with a two-tone inhibition paradigm. An early-arriving low-frequency inhibition (LFI) prevents responses to upward sweeps and thus shapes direction selectivity. A late-arriving high-frequency inhibition (HFI) suppresses slow FM sweeps and thus shapes rate selectivity for downward sweeps. Iontophoretic application of gabazine (GBZ) to block GABA(A) receptors or strychnine (Strych) to block glycine receptors was used to assess the effects of removal of inhibition on each form of FM selectivity. GBZ and Strych had a similar effect on FM direction selectivity, reducing selectivity in up to 86% of neurons when both drugs were coapplied. FM rate selectivity was more resistant to drug application with less than 38% of neurons affected. In addition, only Strych could eliminate FM rate selectivity, whereas GBZ alone was ineffective. The loss of FM selectivity was directly correlated to a loss of the respective inhibitory sideband that shapes that form of selectivity. The elimination of LFI correlated to a loss of FM direction selectivity, whereas elimination of HFI correlated to a loss of FM rate selectivity. Results indicate that 1) although the majority of FM direction selectivity is created within the IC, the majority of rate selectivity is inherited from lower levels of the auditory system, 2) a loss of LFI corresponds to a loss of FM direction selectivity and is created through either GABAergic or glycinergic input, and 3) a loss of HFI corresponds to a loss of FM rate selectivity and is created mainly through glycinergic input.

Repetition Enhancement for Frequency-Modulated but Not Unmodulated Sounds: A Human MEG Study

BackgroundDecoding of frequency-modulated (FM) sounds is essential for phoneme identification. This study investigates selectivity to FM direction in the human auditory system.Methodology/Principal FindingsMagnetoencephalography was recorded in 10 adults during a two-tone adaptation paradigm with a 200-ms interstimulus-interval. Stimuli were pairs of either same or different frequency modulation direction. To control that FM repetition effects cannot be accounted for by their on- and offset properties, we additionally assessed responses to pairs of unmodulated tones with either same or different frequency composition. For the FM sweeps, N1m event-related magnetic field components were found at 103 and 130 ms after onset of the first (S1) and second stimulus (S2), respectively. This was followed by a sustained component starting at about 200 ms after S2. The sustained response was significantly stronger for stimulation with the same compared to different FM direction. This effect was not observed for the non-modulated control stimuli.Conclusions/SignificanceLow-level processing of FM sounds was characterized by repetition enhancement to stimulus pairs with same versus different FM directions. This effect was FM-specific; it did not occur for unmodulated tones. The present findings may reflect specific interactions between frequency separation and temporal distance in the processing of consecutive FM sweeps.

Cat's behavioral sensitivity and cortical spatiotemporal responses to the sweep direction of frequency-modulated tones

Neural responses to frequency modulated (FM) sweeps have been well investigated in single-units using electrophysiological recording methods. However, rare psychophysical experiments were conducted to investigate behavioral discrimination of FM-sweeps in the same species used in physiological experiments. Also, the previous studies have not focused on how the population of cortical neurons works together to encode the direction of FM-sweeps. To investigate the relation between the behavioral perception of FM direction and the population coding, we examined the cat's capability to discriminate the upward and downward FM-sweeps and recorded the neural responses to the same stimuli from the primary auditory cortex (A1) in different awake cats. We found that cats showed high performance to detect the change of direction of FM-sweeps, which linearly swept between 0.1 and 16kHz in 40–160ms duration; however, the behavioral performance obviously deteriorated when the sweeps were shortened to 10–20ms. Physiological results indicated that the upward sweeps elicited a temporal sequence of responses among the A1 neurons, in which neurons tuning to low-frequencies responded earlier and those tuning to high-frequencies later, while the response sequence was reversed when A1 was driven by the downward sweeps. The rank-order of response latencies could provide a reliable discrimination of the FM direction, and the discrimination performance paralleled with the cat's behavioral performance. Our result suggests that the relative response timings in A1 contain enough information used by subsequent processing stages to make the decision of FM direction.

Experience-dependent development of vocalization selectivity in the auditory cortex

Vocalization-selective neurons are present in the auditory systems of several vertebrate groups. Vocalization selectivity is influenced by developmental experience, but the underlying mechanisms are only beginning to be understood. Evidence is presented in this review for the hypothesis that plasticity of timing and strength of inhibition is a mechanism for plasticity of vocalization selectivity. The pallid bat echolocates using downward frequency modulated (FM) sweeps. Nearly 70% of neurons with tuning in the echolocation frequency range in its auditory cortex respond selectively to the direction and rate of change of frequencies present in the echolocation call. During development, FM rate selectivity matures early, while direction selectivity emerges later. Based on the time course of development it was hypothesized that FM direction, but not rate, selectivity is experience-dependent. This hypothesis was tested by altering echolocation experience during development. The results show that normal echolocation experience is required for both refinement and maintenance of direction selectivity. Interestingly, experience is required for the maintenance of rate selectivity, but not for initial development. Across all ages and experimental groups, the timing relationship between inhibitory and excitatory inputs explains sweep selectivity. These experiments suggest that inhibitory plasticity is a substrate for experience-dependent changes in vocalization selectivity.

Multiple mechanisms shape selectivity for FM sweep rate and direction in the pallid bat inferior colliculus and auditory cortex

The inferior colliculus and auditory cortex of the pallid bat contain a large percentage of neurons that are highly selective for the direction and rate of the downward frequency modulated (FM) sweep of the bat's echolocation pulse. Approximately 25% of neurons tuned to the echolocation pulse respond exclusively to downward FM sweeps. This review focuses on the finding that this selectivity is generated by multiple mechanisms that may act alone or in concert. In the inferior colliculus, selectivity for sweep rate is shaped by at least three mechanisms: shortpass or bandpass tuning for signal duration, delayed high-frequency inhibition that prevents responses to slow sweep rates, and asymmetrical facilitation that occurs only when two tones are presented at appropriate delays. When acting alone, the three mechanisms can produce essentially identical rate selectivity. Direction selectivity can be produced by two mechanisms: an early low-frequency inhibition that prevents responses to upward sweeps, and the same asymmetrical two-tone inhibition that shapes rate tuning. All mechanisms except duration tuning are also present in the auditory cortex. Discussion centers on whether these mechanisms are redundant or complementary.

Synaptic Mechanisms of Direction Selectivity in Primary Auditory Cortex

Frequency modulation (FM) is a prominent feature in animal vocalization and human speech. Although many neurons in the auditory cortex are known to be selective for FM direction, the synaptic mechanisms underlying this selectivity are not well understood. Previous studies of both visual and auditory neurons have suggested two general mechanisms for direction selectivity: (1) differential delays of excitatory inputs across the spatial/spectral receptive field and (2) spatial/spectral offset between excitatory and inhibitory inputs. In this study, we have examined the contributions of both mechanisms to FM direction selectivity in rat primary auditory cortex. The excitatory and inhibitory synaptic inputs to each cortical neuron were measured by in vivo whole-cell recording. The spectrotemporal receptive field of each type of inputs was mapped with random tone pips and compared with direction selectivity of the neuron measured with FM stimuli. We found that both the differential delay of the excitatory input and the spectral offset between excitation and inhibition are positively correlated with direction selectivity of the neuron. Thus, both synaptic mechanisms are likely to contribute to FM direction selectivity in the auditory cortex. Finally, direction selectivity measured from the spiking output is significantly stronger than that based on the subthreshold membrane potentials, indicating that the selectivity is further sharpened by the spike generation mechanism.