Frequency-modulated Sweeps Research Articles

Network models of frequency modulated sweep detection.

Frequency modulated (FM) sweeps are common in species-specific vocalizations, including human speech. Auditory neurons selective for the direction and rate of frequency change in FM sweeps are present across species, but the synaptic mechanisms underlying such selectivity are only beginning to be understood. Even less is known about mechanisms of experience-dependent changes in FM sweep selectivity. We present three network models of synaptic mechanisms of FM sweep direction and rate selectivity that explains experimental data: (1) The ‘facilitation’ model contains frequency selective cells operating as coincidence detectors, summing up multiple excitatory inputs with different time delays. (2) The ‘duration tuned’ model depends on interactions between delayed excitation and early inhibition. The strength of delayed excitation determines the preferred duration. Inhibitory rebound can reinforce the delayed excitation. (3) The ‘inhibitory sideband’ model uses frequency selective inputs to a network of excitatory and inhibitory cells. The strength and asymmetry of these connections results in neurons responsive to sweeps in a single direction of sufficient sweep rate. Variations of these properties, can explain the diversity of rate-dependent direction selectivity seen across species. We show that the inhibitory sideband model can be trained using spike timing dependent plasticity (STDP) to develop direction selectivity from a non-selective network. These models provide a means to compare the proposed synaptic and spectrotemporal mechanisms of FM sweep processing and can be utilized to explore cellular mechanisms underlying experience- or training-dependent changes in spectrotemporal processing across animal models. Given the analogy between FM sweeps and visual motion, these models can serve a broader function in studying stimulus movement across sensory epithelia.

Morphological, olfactory, and vocal development in big brown bats.

ABSTRACTUsing a within subjects design, we documented morphological, bioacoustical and behavioral developmental changes in big brown bats. Eptesicus fuscus pups are born naked and blind but assume an adult-like appearance by post-natal day (PND) 45 and flight by PND 30. Adult females use spatial memory, acoustic and olfactory cues to reunite with offspring, but it is unclear if pups can recognize maternal scents. We tested the olfactory discrimination abilities of young E. fuscus pups and found they exhibited no odor preferences. Pups also emit distinct vocalizations called isolation calls (i-calls) that facilitate mother-offspring reunions, but how pups shift their vocalizations from i-calls to downward frequency modulated (FM) sweeps used in echolocation remains unclear. Between PND 0–9, pups emitted mainly long duration, tonal i-calls rich in harmonics, but after they switched to short duration, downward FM sweeps with fewer harmonics. Call maximum frequency and repetition rate showed minor changes across development. Signal duration, bandwidth, and number of harmonics decreased, whereas the maximum, minimum and bandwidth of the fundamental, and peak spectral frequency all increased. We recorded vocalizations during prolonged maternal separation and found that isolated pups called longer and at a faster rate, presumably to signal for maternal assistance. To assess how PND 13 pups alter their signals during interactions with humans we compared spontaneous and provoked vocalizations and found that provoked calls were spectrally and temporally more similar to those of younger bats suggesting that pups in distress emit signals that sound like younger bats to promote maternal assistance.

Effect of Series of 1 to 2 kHz and 6 to 7 kHz Up-Sweeps and Down-Sweeps on the Behavior of a Harbor Porpoise (Phocoena phocoena)

To compare the effect of naval sonar up-sweeps and down-sweeps on the behavior of harbor porpoises, a harbor porpoise in a large pool was exposed to simulated low- and mid-frequency active sonar signals (series of 1-s duration frequency-modulated sweeps). Three sweep pairs were tested: (1) a 1 to 2 kHz up-sweep was compared with a 2 to 1 kHz down-sweep (both without harmonics) at a mean received sound pressure level (mean received SPL) of 114 dB re 1 µPa; (2) a 1 to 2 kHz up-sweep was compared with a 2 to 1 kHz downsweep (both with harmonics; mean received SPL: 123 dB re 1 µPa); and (3) a 6 to 7 kHz up-sweep was compared with a 7 to 6 kHz down-sweep (both without harmonics; mean SPL: 107 dB re 1 µPa). For each sweep pair, the level was chosen during a pretest session with the intention that the harbor porpoise would respond to the sounds by moving away from the projector and surfacing more often (i.e., he would show a change in behavior). The study consists of three separate parts, so only a comparison within sweep pairs could be made and not between sweep pairs. For the 1 to 2 kHz sweeps with harmonics, the harbor porpoise swam further away from the sound source in response to the up-sweeps than to the downsweeps. For the other two sweep pairs, sweep type (up-sweep or down-sweep) caused no significant difference in the harbor porpoise’s response. Thus, to allow the evaluation of potential effects of sonar sounds on harbor porpoises, sonar signal measurements should include the harmonics. For simulated naval sonar sounds with fundamental frequencies in the 1 to 2 kHz range containing harmonics, using down-sweeps appears to affect harbor porpoise behavior less than using up-sweeps.

An improved processing sequence for uncorrelated Chirp sonar data

Chirp sonar systems can be used to obtain high resolution seismic reflection images of the sub-seafloor during marine surveys. The exact knowledge of the Chirp signature allows the use of deterministic algorithms to process the data, similarly to that applied to Vibroseis data on land. Here, it is described an innovative processing sequence to be applied to uncorrelated Chirp data, which can improve vertical and lateral resolution compared to conventional methods. It includes application of a Wiener filter to transform a frequency-modulated sweep into a minimum-phase pulse sequence. In this way, the data become causal and can undergo predictive deconvolution to reduce ringing and enhance vertical resolution. Afterwards, FX-deconvolution and Stolt migration can be applied to obtain an improved imaging of the subsurface. The result of this procedure is a seismic reflection image with higher resolution than traditional ones, which are normally represented using the envelope function of the signal. This technique can be particularly useful for engineering-geotechnical surveys and archaeological investigations that require a fine detail imaging of the uppermost meters of the sub-seafloor.

Spatial hearing in Cope’s gray treefrog: II. Frequency-dependent directionality in the amplitude and phase of tympanum vibrations

Anuran ears function as pressure difference receivers, and the amplitude and phase of tympanum vibrations are inherently directional, varying with sound incident angle. We quantified the nature of this directionality for Cope's gray treefrog, Hyla chrysoscelis. We presented subjects with pure tones, advertisement calls, and frequency-modulated sweeps to examine the influence of frequency, signal level, lung inflation, and sex on ear directionality. Interaural differences in the amplitude of tympanum vibrations were 1-4dB greater than sound pressure differences adjacent to the two tympana, while interaural differences in the phase of tympanum vibration were similar to or smaller than those in sound phase. Directionality in the amplitude and phase of tympanum vibration were highly dependent on sound frequency, and directionality in amplitude varied slightly with signal level. Directionality in the amplitude and phase of tone- and call-evoked responses did not differ between sexes. Lung inflation strongly affected tympanum directionality over a narrow frequency range that, in females, included call frequencies. This study provides a foundation for further work on the biomechanics and neural mechanisms of spatial hearing in H. chrysoscelis, and lends valuable perspective to behavioral studies on the use of spatial information by this species and other frogs.

Intrinsic Cortical Signal Gain

The role of the cortex in receiving incoming sensory information provided by thalamic inputs is not completely understood. Is the cortex act a passive receiving system, performing subsequent manipulations on a similar copy of the incoming data, or does it actively transform thalamic input to improve performance or detection? In a recent study, Li et al (Intracortical multiplication of thalamocortical signals in mouse auditory cortex. Nat Neurosci. 2013;16(9):1179-1181) used an optogenetics technique to support the latter interpretation. The authors dampened intracortical circuitry by activating parvalbumin (PV+) expressing inhibitory neurons locally, and found that in the uninhibited state, primary auditory cortex amplifies thalamocortical responses while leaving the spectral tuning curves preserved. This process appears to increase the duration and gain of thalamic input signals and serves to improve the signal-to-noise properties of the transmitted information. The authors performed this study in anesthetized mice, examining specifically primary auditory cortex (A1). Using blue LED illumination, they activated the PV+ inhibitory cellular component which had been modified (Cre-loxP recombination using an adeno-associated viral vector) to express channelrhodopsin-2 (ChR2). The PV+ neurons increased their firing rates considerably with illumination. Extracellular multi-unit recordings were performed in A1 for tonotopic mapping, with tone inputs applied to the contralateral ear. Similar recordings were also performed stereotactically in the ventral medial geniculate body (MGBv) in a tonotopic fashion, and loose-patch and whole-cell voltage clamp recordings were made from the Layer IV region of A1 (predominantly pyramidal cells). During optogenetic silencing, the authors found that the cortical response produced by equivalent thalamic inputs (for the same frequency tone stimulus) was reduced by a factor of 2.4 as compared to the normal non-silenced state. Additionally, the time duration of the cortical response was reduced during optogenetic silencing. The response amplification produced by the surrounding cortical input appeared additionally to preserve the preferred cortical response to upward going frequency-modulated sweeps, but the overall amplitude change brought on by optogenetic silencing was independent of direction of frequency sweeps. The onset latencies to sweep inputs remained unchanged between the silenced and non-silenced state, implying that the spectral range of the cortical response is determined by the thalamocortical input component, and not modulated by surrounding cortical input. In summary, Li et al describe an optogenetic method for silencing surrounding cortical influences on sensory input transmission to the primary auditory cortex of mice. The authors found amplification of the thalamic auditory signal input brought on by surrounding cortical input, which preserves the cortical frequency tuning response and appears to increase the signal-to-noise properties of the incoming MGBv relayed signal. This paper has relevance to the neurosurgeon or engineer interested in neuroprosthetic development. It may be possible to mimic this type of cortical pre-processing to amplify and augment incoming sensory information in future prosthetic systems that rely on sensory input as a feedback mechanism.Figure: A and C, 2 neurons exhibiting frequency Sdependent tone evoked responses, both during the case of optogenetic silencing of the cortical network (lower traces) and during the non-silenced case (upper traces). B and D, linear nature of pink amplitude of thalamic input vs the total excitation evoked by the same tone stimulus (left subplot). Onset latencies of the thalamocortical (blue) and total cortical (red) excitatory responses at different frequencies (right subplot). (Reprinted by permission from Macmillan Publishers Ltd: Li LY, Li YT, Zhou M, Tao HW, Zhang LI. Intracortical multiplication of thalamocortical signals in mouse auditory cortex. Nat Neurosci. 2013;16(9):1179-1181).

Footprints of inhibition in the response of cortical delay-tuned neurons of bats

Responses of echo-delay-tuned neurons that encode target distance were investigated in the dorsal auditory cortex of anesthetized short-tailed fruit bats (Carollia perspicillata). This species echolocates using short downward frequency-modulated (FM) biosonar signals. In response to FM sweeps of increasing level, 60 out of 131 studied neurons (47%) displayed a "paradoxical latency shift," i.e., longer response latency to loud sounds and shorter latency to faint sounds. In addition, a disproportionately large number of neurons (80%) displayed nonmonotonic responses, i.e., weaker responses to loud sounds and stronger responses to faint sounds. We speculate that the observed paradoxical latency shift and nonmonotonic responses are extracellular footprints of inhibitory processes evoked by loud sounds and that they could represent a specialization for the processing of the emitted loud biosonar pulse. Supporting this idea is the fact that all studied neurons displayed strong response suppression when an artificial loud pulse and a faint echo were presented together at a nonoptimal delay. In 24 neurons, iontophoresis of bicuculline (an antagonist of A-type γ-aminobutyric acid receptors) did not remove inhibitory footprints but did increase the overall spike output, and in some cases it also modified the response bandwidth and shifted the neuron's "best delay." We suggest that inhibition could play a dual role in shaping delay tuning in different auditory stations. Below the cortex it participates in delay-tuning implementation and leaves a footprint that is measurable in cortical responses, while in the cortex it provides a substrate for an in situ control of neuronal selectivity.

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.

Altered cortical spectrotemporal processing with age-related hearing loss

Presbycusis (age-related hearing loss) is a prevalent disability associated with aging that impairs spectrotemporal processing, but the mechanisms of such changes remain unclear. The goal of this study was to quantify cortical responses to frequency-modulated (FM) sweeps in a mouse model of presbycusis. Previous studies showed that cortical neurons in young mice are selective for the rate of frequency change in FM sweeps. Here single-unit data on cortical selectivity and response variability to FM sweeps of either direction and different rates (0.08-20 kHz/ms) were compared across young (1-3 mo), middle-aged (6-8 mo), and old (14-20 mo) groups. Three main findings are reported. First, there is a reduction in FM rate selectivity in the old group. Second, there is a slowing of the sweep rates at which neurons likely provide best detection and discrimination of sweep rates. Third, there is an increase in trial-to-trial variability in the magnitude and timing of spikes in response to sweeps. These changes were only observed in neurons that were selective for the fast or intermediate range of sweep rates and not in neurons that preferred slow sweeps or were nonselective. Increased variability of response magnitude, but not changes in temporal fidelity or selectivity, was seen even in the middle-aged group. The results show that spectrotemporal processing becomes slow and noisy with presbycusis in specific types of neurons, suggesting receptive field mechanisms that are altered. These data suggest neural correlates of presbycusis-related reduction in the ability of humans to process rapid spectrotemporal changes.

Regularity encoding and deviance detection of frequency modulated sweeps: Human middle‐ and long‐latency auditory evoked potentials

Fast encoding of frequency modulated (FM) sweeps is crucial for communication. In humans, FM sweeps deviating from the acoustic regularity elicit the mismatch negativity (MMN) evoked potential. Yet, direction sensitivity to FM sweeps is found in animals' primary auditory cortex, upstream of MMN sources found in humans. Here, we were interested in whether direction deviants of complex FM sweeps modulated brain responses earlier than MMN. We used a controlled oddball paradigm, and measured the middle latency responses (MLRs) and the MMN. Our results showed a repetition enhancement to the standards at the Pa component of the MLR and a genuine MMN in the later response range. These results show that, early in the cortical hierarchy, the system is sensitive to the physical characteristics of the repetitive stimuli, but a higher-order mechanism is needed to detect violations of the acoustic regularity.

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.

High Duty Cycle to Low Duty Cycle: Echolocation Behaviour of the Hipposiderid Bat Coelops frithii

Laryngeally echolocating bats avoid self-deafening (forward masking) by separating pulse and echo either in time using low duty cycle (LDC) echolocation, or in frequency using high duty cycle (HDC) echolocation. HDC echolocators are specialized to detect fluttering targets in cluttered environments. HDC echolocation is found only in the families Rhinolophidae and Hipposideridae in the Old World and in the New World mormoopid, Pteronotus parnellii. Here we report that the hipposiderid Coelops frithii, ostensibly an HDC bat, consistently uses an LDC echolocation strategy whether roosting, flying, or approaching a fluttering target rotating at 50 to 80 Hz. We recorded the echolocation calls of free-flying C. frithii in the field in various situations, including presenting bats with a mechanical fluttering target. The echolocation calls of C. frithii consisted of an initial narrowband component (0.5±0.3 ms, 90.6±2.0 kHz) followed immediately by a frequency modulated (FM) sweep (194 to 113 kHz). This species emitted echolocation calls at duty cycles averaging 7.7±2.8% (n = 87 sequences). Coelops frithii approached fluttering targets more frequently than did LDC bats (C.frithii, approach frequency = 40.4%, n = 80; Myotis spp., approach frequency = 0%, n = 13), and at the same frequency as sympatrically feeding HDC species (Hipposideros armiger, approach rate = 53.3%, n = 15; Rhinolophus monoceros, approach rate = 56.7%, n = 97). We propose that the LDC echolocation strategy used by C. frithii is derived from HDC ancestors, that this species adjusts the harmonic contents of its echolocation calls, and that it may use both the narrowband component and the FM sweep of echolocations calls to detect fluttering targets.

Altered auditory processing in a mouse model of fragile X syndrome

This study provides the first description of auditory cortical processing in a mouse model of Fragile X Syndrome (FXS). FXS is a genetic cause of intellectual impairment and is an autism spectrum disorder. Human studies with auditory evoked potentials indicate that FXS is associated with abnormal auditory processing. The Fmr1 knock-out (KO) mouse is a useful model for studying FXS. The KO mice show acoustic hypersensitivity and propensity for audiogenic seizures, suggesting altered auditory responses. However, the nature of changes at the neuronal level is not known. Here we conducted in vivo single unit extracellular electrophysiology in the auditory cortex of urethane/xylazine-anesthetized Fmr1 KO mice in response to tones and frequency modulated (FM) sweeps. Using tones as stimuli, we report expanded frequency tuning, enhanced response magnitude, and more variable first spike latencies in Fmr1 KO mice compared to wild-type controls. FM sweep stimuli revealed altered sensitivity to the rate of frequency change indicating abnormal spectrotemporal processing. There was no difference in FM sweep direction selectivity. Consistent with studies of the somatosensory cortex, these data point to hyper-responsiveness of auditory neurons as a key processing abnormality in FXS. Auditory neural responses can serve as outcome measures in preclinical trials of therapeutics for FXS as well as serve as physiological probes to study their mechanisms of action.

Response properties underlying selectivity for the rate of frequency modulated sweeps in the auditory cortex of the mouse

This study focused on the response properties underlying selectivity for the rate of frequency modulated (FM) sweeps in the auditory cortex of anesthetized C57bl/6 (C57) mice. Linear downward FM sweeps with rates between 0.08 and 20 kHz/ms were tested. We show that at least two different response properties predict FM rate selectivity: sideband inhibition and duration tuning. Sideband inhibition was determined using the two-tone inhibition paradigm in which excitatory and inhibitory tones were presented with different delays. Sideband inhibition was present in the majority (88%, n = 53) of neurons. The spectrotemporal properties of sideband inhibition predicted rate selectivity and exclusion of the sideband from the sweep reduced/eliminated rate tuning. The second property predictive of sweep rate selectivity was duration tuning for tones. Theoretically, if a neuron is selective for the duration that a sweep spends in the excitatory frequency tuning curve, then rate selectivity will ensue. Duration tuning for excitatory tones was present and predicted rate selectivity in ∼34% of neurons (n = 97). Both sideband inhibition and duration tuning predicted rate selectivity equally well, but sideband inhibition was present in a larger percentage of neurons suggesting that it is the dominant mechanism in the C57 mouse auditory cortex. Similar mechanisms shape sweep rate selectivity in the auditory system of bats and mice and movement-velocity selectivity in the visual system, suggesting similar solutions to analogous problems across sensory systems. This study provides baseline data on basic spectrotemporal processing in the C57 strain for elucidation of changes that occur in presbycusis.

Encoding of ultrasonic vocalizations in the auditory cortex

One of the central tasks of the mammalian auditory system is to represent information about acoustic communicative signals, such as vocalizations. However, the neuronal computations underlying vocalization encoding in the central auditory system are poorly understood. To learn how the rat auditory cortex encodes information about conspecific vocalizations, we presented a library of natural and temporally transformed ultrasonic vocalizations (USVs) to awake rats while recording neural activity in the primary auditory cortex (A1) with chronically implanted multielectrode probes. Many neurons reliably and selectively responded to USVs. The response strength to USVs correlated strongly with the response strength to frequency-modulated (FM) sweeps and the FM rate tuning index, suggesting that related mechanisms generate responses to USVs as to FM sweeps. The response strength further correlated with the neuron's best frequency, with the strongest responses produced by neurons whose best frequency was in the ultrasonic frequency range. For responses of each neuron to each stimulus group, we fitted a novel predictive model: a reduced generalized linear-nonlinear model (GLNM) that takes the frequency modulation and single-tone amplitude as the only two input parameters. The GLNM accurately predicted neuronal responses to previously unheard USVs, and its prediction accuracy was higher than that of an analogous spectrogram-based linear-nonlinear model. The response strength of neurons and the model prediction accuracy were higher for original, rather than temporally transformed, vocalizations. These results indicate that A1 processes original USVs differentially than transformed USVs, indicating preference for temporal statistics of the original vocalizations.

Influence of Core Auditory Cortical Areas on Acoustically Evoked Activity in Contralateral Primary Auditory Cortex

In contrast to numerous studies of transcallosal communication in visual and somatosensory cortices, the functional properties of interhemispheric connections between auditory cortical fields have not been widely scrutinized. Therefore, the purpose of the present investigation was to measure the magnitude and type (inhibitory/excitatory) of modulatory properties of core auditory fields on contralateral primary auditory cortex (A1) activity. We combined single-unit neuronal recordings with reversible cooling deactivation techniques to measure variations in contralateral A1 response levels during A1, anterior auditory field (AAF), or simultaneous A1 and AAF neuronal discharge suppression epochs in cat auditory cortex. Cortical activity was evoked by presentation of pure tones, noise bursts, and frequency-modulated (FM) sweeps before, during, and after cortical deactivation periods. Comparisons of neuronal response changes before and during neuronal silencing revealed three major findings. First, deactivation of A1 and AAF-induced significant peak response reductions in contralateral A1 activity during simple (tonal) and complex (noise bursts and FM sweeps) acoustic exposure. Second, decreases in A1 neuronal activity appear to be in agreement with anatomical laminar termination patterns emanating from contralateral auditory cortex fields. Third, modulatory properties of core auditory areas lack hemispheric lateralization. These findings demonstrate that during periods of acoustic exposure, callosal projections emanating from core auditory areas modulate A1 neuronal activity via excitatory inputs.

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.

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.

The noseleaf of Rhinolophus formosae focuses the Frequency Modulated (FM) component of the calls

Bats of the family Rhinolophidae emit their echolocation calls through their nostrils and feature elaborate noseleaves shaping the directionality of the emissions. The calls of these bats consist of a long constant-frequency component preceded and/or followed by short frequency-modulated sweeps. While Rhinolophidae are known for their physiological specializations for processing the constant frequency part of the calls, previous evidence suggests that the noseleaves of these animals are tuned to the frequencies in the frequency modulated components of the calls. In this paper, we seek further support for this hypothesis by simulating the emission beam pattern of the bat Rhinolophus formosae. Filling the furrows of lancet and removing the basal lappets (i.e., two flaps on the noseleaf) we find that these conspicuous features of the noseleaf focus the emitted energy mostly for frequencies in the frequency-modulated components. Based on the assumption that this component of the call is used by the bats for ranging, we develop a qualitative model to assess the increase in performance due to the furrows and/or the lappets. The model confirms that both structures decrease the ambiguity in selecting relevant targets for ranging. The lappets and the furrows shape the emission beam for different spatial regions and frequency ranges. Therefore, we conclude that the presented evidence is in line with the hypothesis that different parts of the noseleaves of Rhinolophidae are tuned to different frequency ranges with at least some of the most conspicuous ones being tuned to the frequency modulated components of the calls—thus yielding strong evidence for the sensory importance of the component.

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.