Frequency Tuning Curves Research Articles

Lateral Sharpening of Cortical Frequency Tuning by Approximately Balanced Inhibition

Cortical inhibition plays an important role in shaping neuronal processing. The underlying synaptic mechanisms remain controversial. Here, in vivo whole-cell recordings from neurons in the rat primary auditory cortex revealed that the frequency tuning curve of inhibitory input was broader than that of excitatory input. This results in relatively stronger inhibition in frequency domains flanking the preferred frequencies of the cell and a significant sharpening of the frequency tuning of membrane responses. The less selective inhibition can be attributed to a broader bandwidth and lower threshold of spike tonal receptive field of fast-spike inhibitory neurons than nearby excitatory neurons, although both types of neurons receive similar ranges of excitatory input and are organized into the same tonotopic map. Thus, the balance between excitation and inhibition is only approximate, and intracortical inhibition with high sensitivity and low selectivity can laterally sharpen the frequency tuning of neurons, ensuring their highly selective representation.

Nonlinearities and Contextual Influences in Auditory Cortical Responses Modeled with Multilinear Spectrotemporal Methods

The relationship between a sound and its neural representation in the auditory cortex remains elusive. Simple measures such as the frequency response area or frequency tuning curve provide little insight into the function of the auditory cortex in complex sound environments. Spectrotemporal receptive field (STRF) models, despite their descriptive potential, perform poorly when used to predict auditory cortical responses, showing that nonlinear features of cortical response functions, which are not captured by STRFs, are functionally important. We introduce a new approach to the description of auditory cortical responses, using multilinear modeling methods. These descriptions simultaneously account for several nonlinearities in the stimulus-response functions of auditory cortical neurons, including adaptation, spectral interactions, and nonlinear sensitivity to sound level. The models reveal multiple inseparabilities in cortical processing of time lag, frequency, and sound level, and suggest functional mechanisms by which auditory cortical neurons are sensitive to stimulus context. By explicitly modeling these contextual influences, the models are able to predict auditory cortical responses more accurately than are STRF models. In addition, they can explain some forms of stimulus dependence in STRFs that were previously poorly understood.

Adaptation to Natural Binocular Disparities in Primate V1 Explained by a Generalized Energy Model

Sensory processing in the brain is thought to have evolved to encode naturally occurring stimuli efficiently. We report an adaptation in binocular cortical neurons that reflects the tight constraints imposed by the geometry of 3D vision. We show that the widely used binocular energy model predicts that neurons dedicate part of their dynamic range to impossible combinations of left and right images. Approximately 42% of the neurons we record from V1 of awake monkeys behave in this way (a powerful confirmation of the model), while about 58% deviate from the model in a manner that concentrates more of their dynamic range on stimuli that obey the constraints of binocular geometry. We propose a simple extension of the energy model, using multiple subunits, that explains the adaptation we observe, as well as other properties of binocular neurons that have been hard to account for, such as the response to anti-correlated stereograms.

Tonotopic Control of Auditory Thalamus Frequency Tuning by Reticular Thalamic Neurons

GABAergic cells of the thalamic reticular nucleus (TRN) can potentially exert strong control over transmission of information through thalamus to the cerebral cortex. Anatomical studies have shown that the reticulo-thalamic connections are spatially organized in the visual, somatosensory, and auditory systems. However, the issue of how inhibitory input from TRN controls the functional properties of thalamic relay cells and whether this control follows topographic rules remains largely unknown. Here we assessed the consequences of increasing or decreasing the activity of small ensembles of TRN neurons on the receptive field properties of medial geniculate (MG) neurons. For each MG cell, the frequency tuning curve and the rate-level function were tested before, during, and after microiontophoretic applications of GABA, or of glutamate, in the auditory sector of the TRN. For 66 MG cells tested during potent pharmacological control of TRN activity, group data did not reveal any significant effects. However, for a population of 20/66 cells (all but 1 recorded in the ventral, tonotopic, division), the breadth of tuning, the frequency selectivity and the acoustic threshold were significantly modified in the directions expected from removing, or reinforcing, a dominant inhibitory input onto MG cells. Such effects occurred only when the distance between the characteristic frequency of the recorded ventral MG cell and that of the TRN cells at the ejection site was <0.25 octaves; they never occurred for larger distances. This relationship indicates that the functional interactions between TRN cells and ventral MG cells rely on precise topographic connections.

Structure, Stability, and Spectra of Lateral Modes of a Broad-Area Semiconductor Laser

We present a theoretical analysis of the lateral modes of a broad-area semiconductor laser. The structure of the modes are classified into four categories and the modes are traced in the frequency versus pump rate diagram. It is shown how the branches of the frequency tuning curves for the different types of modes are interconnected and how the intensity profiles develop along the branches. The main result of the paper is the presentation of a small-signal stability analysis which identifies the saddle-node and Hopf bifurcation points on the mode tuning curves. For stable modes we derive expressions for small-signal noise and modulation spectra and present numerical examples of the spectra.

Processing of sounds by population spikes in a model of primary auditory cortex

We propose a model of the primary auditory cortex (A1), in which each iso-frequency column is represented by a recurrent neural network with short-term synaptic depression. Such networks can emit Population Spikes, in which most of the neurons fire synchronously for a short time period. Different columns are interconnected in a way that reflects the tonotopic map in A1, and population spikes can propagate along the map from one column to the next, in a temporally precise manner that depends on the specific input presented to the network. The network, therefore, processes incoming sounds by precise sequences of population spikes that are embedded in a continuous asynchronous activity, with both of these response components carrying information about the inputs and interacting with each other. With these basic characteristics, the model can account for a wide range of experimental findings. We reproduce neuronal frequency tuning curves, whose width depends on the strength of the intracortical inhibitory and excitatory connections. Non-simultaneous two-tone stimuli show forward masking depending on their temporal separation, as well as on the duration of the first stimulus. The model also exhibits non-linear suppressive interactions between sub-threshold tones and broad-band noise inputs, similar to the hypersensitive locking suppression recently demonstrated in auditory cortex. We derive several predictions from the model. In particular, we predict that spontaneous activity in primary auditory cortex gates the temporally locked responses of A1 neurons to auditory stimuli. Spontaneous activity could, therefore, be a mechanism for rapid and reversible modulation of cortical processing.

Responses to Social Vocalizations in the Inferior Colliculus of the Mustached Bat Are Influenced by Secondary Tuning Curves

Neurons in the inferior colliculus (IC) of the mustached bat integrate input from multiple frequency bands in a complex fashion. These neurons are important for encoding the bat's echolocation and social vocalizations. The purpose of this study was to quantify the contribution of complex frequency interactions on the responses of IC neurons to social vocalizations. Neural responses to single tones, two-tone pairs, and social vocalizations were recorded in the IC of the mustached bat. Three types of data driven stimulus-response models were designed for each neuron from single tone and tone pair stimuli to predict the responses of individual neurons to social vocalizations. The first model was generated only using the neuron's primary frequency tuning curve, whereas the second model incorporated the entire hearing range of the animal. The extended model often predicted responses to many social vocalizations more accurately for multiply tuned neurons. One class of multiply tuned neuron that likely encodes echolocation information also responded to many of the social vocalizations, suggesting that some neurons in the mustached bat IC have dual functions. The third model included two-tone frequency tunings of the neurons. The responses to vocalizations were better predicted by the two-tone models when the neuron had inhibitory frequency tuning curves that were not near the neuron's primary tuning curve. Our results suggest that complex frequency interactions in the IC determine neural responses to social vocalizations and some neurons in IC have dual functions that encode both echolocation and social vocalization signals.

The Representation of Complex Images in Spatial Frequency Domains of Primary Visual Cortex

The organization of cat primary visual cortex has been well mapped using simple stimuli such as sinusoidal gratings, revealing superimposed maps of orientation and spatial frequency preferences. However, it is not yet understood how complex images are represented across these maps. In this study, we ask whether a linear filter model can explain how cortical spatial frequency domains are activated by complex images. The model assumes that the response to a stimulus at any point on the cortical surface can be predicted by its individual orientation, spatial frequency, and temporal frequency tuning curves. To test this model, we imaged the pattern of activity within cat area 17 in response to stimuli composed of multiple spatial frequencies. Consistent with the predictions of the model, the stimuli activated low and high spatial frequency domains differently: at low stimulus drift speeds, both domains were strongly activated, but activity fell off in high spatial frequency domains as drift speed increased. To determine whether the filter model quantitatively predicted the activity patterns, we measured the spatiotemporal tuning properties of the functional domains in vivo and calculated expected response amplitudes from the model. The model accurately predicted cortical response patterns for two types of complex stimuli drifting at a variety of speeds. These results suggest that the distributed activity of primary visual cortex can be predicted from cortical maps like those of orientation and SF preference generated using simple, sinusoidal stimuli, and that dynamic visual acuity is degraded at or before the level of area 17.

Preceding weak noise sharpens the frequency tuning and elevates the response threshold of the mouse inferior collicular neurons through GABAergic inhibition

In acoustic communication, animals must extract biologically relevant signals that are embedded in noisy environment. The present study examines how weak noise may affect the auditory sensitivity of neurons in the central nucleus of the mouse inferior colliculus (IC) which receives convergent excitatory and inhibitory inputs from both lower and higher auditory centers. Specifically, we studied the frequency sensitivity and minimum threshold of IC neurons using a pure tone probe and a weak white noise masker under forward masking paradigm. For most IC neurons, probe-elicited response was decreased by a weak white noise that was presented at a specific gap (i.e. time window). When presented within this time window, weak noise masking sharpened the frequency tuning curve and increased the minimum threshold of IC neurons. The degree of weak noise masking of these two measurements increased with noise duration. Sharpening of the frequency tuning curve and increasing of the minimum threshold of IC neurons during weak noise masking were mostly mediated through GABAergic inhibition. In addition, sharpening of frequency tuning curve by the weak noise masker was more effective at the high than at low frequency limb. These data indicate that in the real world the ambient noise may improve frequency sensitivity of IC neurons through GABAergic inhibition while inevitably decrease the frequency response range and sensitivity of IC neurons.

Nonlinear acoustic landmine detection: Comparison of tuning curve experiments profiling on and off the target

Recent experiments [Proc. SPIE 6217, 62170Y, Detection and Remediation Technologies for Mines and Minelike Targets XI; J.T. Broach, R.S. Harmon, J.H. Holloway, Jr.; Eds. (2006)] have shown that the soil vibrating over a drum-like acrylic landmine simulant is more nonlinear on the target than well off (due to soil alone). Here, a Laser Doppler Vibrometer (LDV) is configured to scan laterally across the buried simulant. The 10 cm diameter, 5 cm high rigid simulant has a flexible clamped 0.8 mm thick top plate and is buried 4 cm deep in dry, sifted masonry sand in a rigid soil container placed inside the anechoic chamber. At each location the sand is subjected to a series of increasing amplitude acoustic pressure level sweeps (across a 50–200 Hz bandwidth) that includes the resonant behavior. The soil surface particle velocity vs. frequency tuning curves, from locations ‘‘off target’’ show relatively small nonlinear effects, while those from locations ‘‘on target’’ exhibit clearly identifiable backbone (peak amplitude vs. corresponding resonant frequency) curves that exhibit softening. Profiles of the backbone slope vs. position indicate an extremely large contrast ratio between ‘‘on’’ and ‘‘off’’ locations compared to profiles of surface vibration obtained near resonance. [Work supported by ONR.]

Spatial Frequency-Specific Contrast Adaptation Originates in the Primary Visual Cortex

Adaptation to a high-contrast grating stimulus causes reduced sensitivity to subsequent presentation of a visual stimulus with similar spatial characteristics. This behavioral finding has been attributed by neurophysiological studies to processes within the visual cortex. However, some evidence indicates that contrast adaptation phenomena are also found in early visual pathways. Adaptation effects have been reported in retina and lateral geniculation nucleus (LGN). It is possible that these early pathways could be the physiological origin of the cortical adaptation effect. To study this, we recorded from single neurons in the cat's LGN. We find that contrast adaptation in the LGN, unlike that in the visual cortex, is not spatial frequency specific, i.e., adaptation effects apply to a broad range of spatial frequencies. In addition, aside from the amplitude attenuation, the shape of spatial frequency tuning curves of LGN cells is not affected by contrast adaptation. Again, these findings are unlike those found for cells in the visual cortex. Together, these results demonstrate that pattern specific contrast adaptation is a cortical process.

The auditory cortex

Recognizing other people, animals or objects by the sound they make is something that most of us take for granted. In fact, this ability relies on a series of rich and complex processes that begin when sounds are transduced into electrical signals by the exquisitely sensitive hair cell receptors that lie inside the cochlea of the inner ear. These messages are then encoded as volleys of action potentials by the axons of the vestibulo-cochlear nerve and transmitted via a complex chain of nuclei in the brainstem, midbrain and thalamus towards the auditory cortex (Figure 1A), where the interpretation and recognition of sounds is thought to take place.

Spatial frequency characteristics of nearby neurons in cats’ visual cortex

Various methods have allowed mapping of responses to several stimulus features on the cortical surface, particularly edge orientation and motion direction. The cortical mapping of spatial frequencies (SF), which is the basic property that leads to perception of spatial details of visual objects, is still controversial. We recorded simultaneously extracellular action potentials from neighboring cells in superficial layers of the area 17–18 border region of anesthetized cats. Responses of nearby cells to sine-wave gratings of varying SF were analyzed. Spatial frequency tuning curves were cross-correlated to establish the degree of similarity between the curves and optimal SFs were compared for each pair of neurons. The investigation showed that only about a half of nearby neurons exhibited close optimal SFs and similar tuning curves. The results suggest that SF channels do not show a clear clustering within a small pool of neurons. Such organization may contribute to the perception of spatial details at all orientations and motion directions.

Spatial and temporal frequency selectivity of neurons in the middle temporal visual area of new world monkeys (Callithrix jacchus)

Information about the responses of neurons to the spatial and temporal frequencies of visual stimuli is important for understanding the types of computations being performed in different visual areas. We characterized the spatiotemporal selectivity of neurons in the middle temporal area (MT), which is deemed central for the processing of direction and speed of motion. Recordings obtained in marmoset monkeys using high-contrast sine-wave gratings as stimuli revealed that the majority of neurons had bandpass spatial and temporal frequency tuning, and that the selectivity for these parameters was largely separable. Only in about one-third of the cells was inseparable spatiotemporal tuning detected, this typically being in the form of an increase in the optimal temporal frequency as a function of increasing grating spatial frequency. However, most of these interactions were weak, and only 10% of neurons showed spatial frequency-invariant representation of speed. Cells with inseparable spatiotemporal tuning were most commonly found in the infragranular layers, raising the possibility that they form part of the feedback from MT to caudal visual areas. While spatial frequency tuning curves were approximately scale-invariant on a logarithmic scale, temporal frequency tuning curves covering different portions of the spectrum showed marked and systematic changes. Thus, MT neurons can be reasonably described as similarly built spatial frequency filters, each covering a different dynamic range. The small proportion of speed-tuned neurons, together with the laminar position of these units, are compatible with the idea that an explicit neural representation of speed emerges from computations performed in MT.

Mild noise-induced hearing loss at young age affects temporal modulation transfer functions in adult cat primary auditory cortex

Kittens were exposed for 2 h to a 1/3rd octave band of noise centered at 5 kHz and at 120 dB SPL. After the exposure, they were kept in a quiet room for at least 4 weeks, and until they were mature. The noise-exposed cats showed on average 16.5 dB higher ABR thresholds and 13.2 dB higher thresholds at the characteristic frequency (CF) than the control cats for frequencies between 4 and 16 kHz. The frequency-tuning curve bandwidth at 20 dB above threshold was significantly increased compared to controls in the CF region of the hearing loss. In noise-exposed cats, temporal modulation-transfer functions (tMTFs) to amplitude-modulated (AM) noise, but not to periodic click trains, showed a marked increase for modulation frequencies (MFs) below 6 Hz. The vectorstrength in noise-exposed cats increased for all modulation frequencies below 32 Hz for neurons with a CF in the range of the hearing loss. The tMTFs for AMnoise in the noise-exposed group were less band-pass compared to the controls, and in that sense the mild hearing loss could be considered as effectively reducing the central activation in the same way as a reduced sound pressure level. Effects of reduced central inhibition are visible in the broadening of frequency-tuning curves, and in the increased limiting rates for AMnoise.

Hearing frequency selectivity in four species of toothed whales as revealed by the evoked-potential method

Frequency tuning curves were obtained using a tone-tone simultaneous masking paradigm in conjunction with the evoked potential recording. The masker was a continuous tone and the test was a sinusoidal amplitude-modulated (SAM) tonal signal, which evoked the envelope following response (EFR). The EFR was recorded in unanaesthetized animals from a head surface with the use of suction-cup electrodes. The obtained tuning curves featured very sharp tuning with Q(ERB) (quality estimated by the equivalent rectangular bandwidth) from 35 in Tursiops truncatus to nearly 50 in Delphinapterus leucas. This acuteness is several times better than in humans and many animals. The Q(ERB) dependence on probe frequency could be approximated by regression lines with a slope from 0.18 in Tursiops trucatus to 0.83–0.86 in Phocoena phocoena and Neophocoena phocoenoides. Thus, the frequency representation in the odontocete auditory system may be either near constant quality (in Tursiops) or near constant bandwidth (in porpoises). [Work supported by The Russian Foundation for Basic Research and Russian President Grant.]

Different Serotonin Receptor Agonists Have Distinct Effects on Sound-Evoked Responses in Inferior Colliculus

The neuromodulator serotonin has a complex set of effects on the auditory responses of neurons within the inferior colliculus (IC), a midbrain auditory nucleus that integrates a wide range of inputs from auditory and nonauditory sources. To determine whether activation of different types of serotonin receptors is a source of the variability in serotonergic effects, four selective agonists of serotonin receptors in the serotonin (5-HT) 1 and 5-HT2 families were iontophoretically applied to IC neurons, which were monitored for changes in their responses to auditory stimuli. Different agonists had different effects on neural responses. The 5-HT1A agonist had mixed facilitatory and depressive effects, whereas 5-HT1B and 5-HT2C agonists were both largely facilitatory. Different agonists changed threshold and frequency tuning in ways that reflected their effects on spike count. When pairs of agonists were applied sequentially to the same neurons, selective agonists sometimes affected neurons in ways that were similar to serotonin, but not to other selective agonists tested. Different agonists also differentially affected groups of neurons classified by the shapes of their frequency-tuning curves, with serotonin and the 5-HT1 receptors affecting proportionally more non-V-type neurons relative to the other agonists tested. In all, evidence suggests that the diversity of serotonin receptor subtypes in the IC is likely to account for at least some of the variability of the effects of serotonin and that receptor subtypes fulfill specialized roles in auditory processing.

Amplitude modulation and apodization of quasi-phase-matched interactions

We propose several techniques to modulate the local amplitude of quasi-phase-matched (QPM) interactions in periodically poled lithium niobate waveguides and demonstrate apodization by using each of these techniques. When the hard edges are removed in the spatial profile of the nonlinear coupling, the sidelobes of the frequency tuning curves are suppressed by 13 dB or more, compared with a uniform grating, consistent with theoretical predictions. The sidelobe-suppressed gratings are useful for frequency conversion devices in optical communication systems to minimize interchannel cross talk, while the amplitude modulation techniques in general have potential uses in applications that require altering the tuning curve shapes.

Frequency tuning of long-wavelength VCSELs

Tuning properties of long-wavelength VCSELs have been studied experimentally, for the first time to our knowledge. Injection current and temperature tuning rates of two VCSELs operating near 1512 and 1577 nm have been measured using a Fabry–Perot etalon with free spectral range 0.056 cm −1. A 100-Hz saw-tooth modulation with depths of modulation of ∼10% or less was superimposed on a direct injection current (dc bias) to tune lasers in narrow spectral intervals (0.3–1.2 cm −1) around a central frequency set by the dc bias. The lasers have been found to be capable of being tuned faster at higher levels of dc bias. The enhancement factors were up to ∼2 and ∼3 for the 1512- and 1577-nm lasers, respectively, as compared with their tuning rates measured at the levels of the dc bias close to the threshold of lasing. A linear dependence between injection current tuning rates and the levels of dc bias has been observed. Temperature tuning coefficients have been proved to be independent of the laser heat sink temperature and of the dc bias. Frequency tuning curves were approximated with a second-order polynomial. The frequencies of more than 40 absorption lines of CO, CO 2, H 2O and NH 3 known from spectral databases were compared with the calculated frequencies. The accuracy of the approximation was found to be within 0.2 cm −1 for spectral intervals up to 38 cm −1. The dependence of current tuning rates of the VCSELs on dc bias was shown to be taken into account for accurate analysis of absorption line profiles. The results obtained can be used for precise spectroscopic measurements with long-wavelength VCSELs.

Responses of Neurons in the Rat's Ventral Nucleus of the Lateral Lemniscus to Monaural and Binaural Tone Bursts

Responses to monaural and binaural tone bursts were recorded from neurons in the rat's ventral nucleus of the lateral lemniscus (VNLL). Most of the neurons (55%) had V- or U-shaped frequency-tuning curves with a single clearly defined characteristic frequency (CF). However, many neurons had more complex, multipeaked tuning curves (37%), or other patterns (8%). Temporal firing patterns included both onset and sustained responses to contralateral tone bursts. Onset and sustained responses were distributed along the dorsoventral length of VNLL with no indication of segregation into different regions. Onset neurons had shorter average first-spike latencies than neurons with sustained responses (means, 8.3 vs. 14.8 ms). They also had less jitter, as reflected in the SD of first-spike latencies, than neurons with sustained responses (means, 0.59 and 4.2 ms, respectively). The extent of jitter decreased with an increase in stimulus intensity for neurons with sustained responses, but remained unchanged for onset neurons tested over the same range. Many neurons had binaural responses, primarily of the excitatory/inhibitory (EI) type, widely distributed along the dorsoventral extent of VNLL. Local application of the AMPA receptor antagonist NBQX reduced excitatory responses, indicating that responses were dependent on synaptic activity and not recorded from passing fibers. The results show that many neurons in VNLL have a precision of timing that is well suited for processing auditory temporal information. In the rat, these neurons are intermingled among cells with less precise temporal response features and include cells with binaural as well as monaural responses.