Effects of talker, rate, and amplitude variation on recognition memory for spoken words.
This study investigated the encoding of the surface form of spoken words using a continuous recognition memory task. The purpose was to compare and contrast three sources of stimulus variability--talker, speaking rate, and overall amplitude--to determine the extent to which each source of variability is retained in episodic memory. In Experiment 1, listeners judged whether each word in a list of spoken words was "old" (had occurred previously in the list) or "new." Listeners were more accurate at recognizing a word as old if it was repeated by the same talker and at the same speaking rate; however, there was no recognition advantage for words repeated at the same overall amplitude. In Experiment 2, listeners were first asked to judge whether each word was old or new, as before, and then they had to explicitly judge whether it was repeated by the same talker, at the same rate, or at the same amplitude. On the first task, listeners again showed an advantage in recognition memory for words repeated by the same talker and at same speaking rate, but no advantage occurred for the amplitude condition. However, in all three conditions, listeners were able to explicitly detect whether an old word was repeated by the same talker, at the same rate, or at the same amplitude. These data suggest that although information about all three properties of spoken words is encoded and retained in memory, each source of stimulus variation differs in the extent to which it affects episodic memory for spoken words. (+info
Distributed representation of spectral and temporal information in rat primary auditory cortex.
Modulations of amplitude and frequency are common features of natural sounds, and are prominent in behaviorally important communication sounds. The mammalian auditory cortex is known to contain representations of these important stimulus parameters. This study describes the distributed representations of tone frequency and modulation rate in the rat primary auditory cortex (A1). Detailed maps of auditory cortex responses to single tones and tone trains were constructed from recordings from 50-60 microelectrode penetrations introduced into each hemisphere. Recorded data demonstrated that the cortex uses a distributed coding strategy to represent both spectral and temporal information in the rat, as in other species. Just as spectral information is encoded in the firing patterns of neurons tuned to different frequencies, temporal information appears to be encoded using a set of filters covering a range of behaviorally important repetition rates. Although the average A1 repetition rate transfer function (RRTF) was low-pass with a sharp drop-off in evoked spikes per tone above 9 pulses per second (pps), individual RRTFs exhibited significant structure between 4 and 10 pps, including substantial facilitation or depression to tones presented at specific rates. No organized topography of these temporal filters could be determined. (+info
Temporal discharge patterns evoked by rapid sequences of wide- and narrowband clicks in the primary auditory cortex of cat.
The present study investigated neural responses to rapid, repetitive stimuli in the primary auditory cortex (A1) of cats. We focused on two important issues regarding cortical coding of sequences of stimuli: temporal discharge patterns of A1 neurons as a function of inter-stimulus interval and cortical mechanisms for representing successive stimulus events separated by very short intervals. These issues were studied using wide- and narrowband click trains with inter-click intervals (ICIs) ranging from 3 to 100 ms as a class of representative sequential stimuli. The main findings of this study are 1) A1 units displayed, in response to click train stimuli, three distinct temporal discharge patterns that we classify as regions I, II, and III. At long ICIs nearly all A1 units exhibited typical stimulus-synchronized response patterns (region I) consistent with previously reported observations. At intermediate ICIs, no clear temporal structures were visible in the responses of most A1 units (region II). At short ICIs, temporal discharge patterns are characterized by the presence of either intrinsic oscillations (at approximately 10 Hz) or a change in discharge rate that was a monotonically decreasing function of ICI (region III). In some A1 units, temporal discharge patterns corresponding to region III were absent. 2) The boundary between regions I and II (synchronization boundary) had a median value of 39.8 ms ICI ([25%, 75%] = [20.4, 58. 8] ms ICI; n = 131). The median boundary between regions II and III was estimated at 6.3 ms ([25%, 75%] = [5.2, 9.7] ms ICI; n = 47) for units showing rate changes (rate-change boundary). 3) The boundary values between different regions appeared to be relatively independent of stimulus intensity (at modest sound levels) or the bandwidth of the clicks used. 4) There is a weak correlation between a unit's synchronization boundary and its response latency. Units with shorter latencies appeared to also have smaller boundary values. And 5) based on these findings, we proposed a two-stage model for A1 neurons to represent a wide range of ICIs. In this model, A1 uses a temporal code for explicitly representing long ICIs and a rate code for implicitly representing short ICIs. (+info
Auditory nerve fiber responses to electric stimulation: modulated and unmodulated pulse trains.
Many modern cochlear implants use sound processing strategies that stimulate the cochlea with modulated pulse trains. Rubinstein et al. [Hear. Res. 127, 108 (1999)] suggested that representation of the modulator in auditory nerve responses might be improved by the addition of a sustained, high-rate, desynchronizing pulse train (DPT). In addition, activity in response to the DPT may mimic the spontaneous activity (SA) in a healthy ear. The goals of this study were to compare responses of auditory nerve fibers in acutely deafened, anesthetized cats elicited by high-rate electric pulse trains delivered through an intracochlear electrode with SA, and to measure responses of these fibers to amplitude-modulated pulse trains superimposed upon a DPT. Responses to pulse trains showed variability from presentation to presentation, but differed from SA in the shape of the envelope of the interval histogram (IH) for pulse rates above 4.8 kpps (kilo pulses per second). These IHs had a prominent mode near 5 ms that was followed by a long tail. Responses to modulated biphasic pulse trains resembled responses to tones in intact ears for small (<10%) modulation depths, suggesting that acousticlike responses to sinusoidal stimuli might be obtained with a DPT. However, realistic responses were only observed over a narrow range of levels and modulation depths. Improved coding of complex stimulus waveforms may be achieved by signal processing strategies for cochlear implants that properly incorporate a DPT. (+info
From spectrum to space: the contribution of level difference cues to spatial receptive fields in the barn owl inferior colliculus.
Space-specific neurons in the owl's inferior colliculus have spatial receptive fields (RFs) computed from interaural time (ITD) and level (ILD) differences. Because of the shape of the owl's head, these cues vary with frequency in a manner specific for each location. We sought to determine the contribution of ILD to spatial selectivity. We measured the normal spatial receptive fields of space-specific neurons using virtual sound sources (i.e., noises filtered to simulate external sound sources, presented using headphones). The virtual-source filters were then altered so that ITD was fixed while frequency-specific ILDs varied according to location in the usual manner. The resulting "ILD-alone" RF typically revealed a horizontal band of excitation that included the normal RF. Above and below, the neurons were inhibited. Interestingly, the maxima of ILD-alone RFs were generally outside the normal RF, suggesting that space-specific neurons are not optimally tuned to the ILD spectrum occurring at the normal RF location. Congruously, frequency-specific ILD tuning, assessed with tones, better matched the ILDs at the peak of the ILD-alone RF than those at the peak of the normal RF. The firing evoked from the normal RF may thus reflect the balance of excitatory and inhibitory inputs needed to appropriately restrict the receptive field. Frequency-specific ILD tuning curves were combined with measured head-filtering characteristics to predict responses to the frequency-specific ILDs at each location. The predicted ILD-alone RFs, which are based on a simple sum of frequency-specific inputs, accounted for 56% of the variance in our measured ILD-alone RFs. (+info
Sound-level-dependent representation of frequency modulations in human auditory cortex: a low-noise fMRI study.
Recognition of sound patterns must be largely independent of level and of masking or jamming background sounds. Auditory patterns of relevance in numerous environmental sounds, species-specific vocalizations and speech are frequency modulations (FM). Level-dependent activation of the human auditory cortex (AC) in response to a large set of upward and downward FM tones was studied with low-noise (48 dB) functional magnetic resonance imaging at 3 Tesla. Separate analysis in four territories of AC was performed in each individual brain using a combination of anatomical landmarks and spatial activation criteria for their distinction. Activation of territory T1b (including primary AC) showed the most robust level dependence over the large range of 48-102 dB in terms of activated volume and blood oxygen level dependent contrast (BOLD) signal intensity. The left nonprimary territory T2 also showed a good correlation of level with activated volume but, in contrast to T1b, not with BOLD signal intensity. These findings are compatible with level coding mechanisms observed in animal AC. A systematic increase of activation with level was not observed for T1a (anterior of Heschl's gyrus) and T3 (on the planum temporale). Thus these areas might not be specifically involved in processing of the overall intensity of FM. The rostral territory T1a of the left hemisphere exhibited highest activation when the FM sound level fell 12 dB below scanner noise. This supports the previously suggested special involvement of this territory in foreground-background decomposition tasks. Overall, AC of the left hemisphere showed a stronger level-dependence of signal intensity and activated volume than the right hemisphere. But any side differences of signal intensity at given levels were lateralized to right AC. This might point to an involvement of the right hemisphere in more specific aspects of FM processing than level coding. (+info
Auditory cortical images of cochlear-implant stimuli: dependence on electrode configuration.
This study examines patterns of auditory cortical activity elicited by single-pulse cochlear implant stimuli that vary in electrode configuration, cochlear place of stimulation, and stimulus level. Recordings were made from the primary auditory cortex (area A1) of ketamine-anesthetized guinea pigs. The spatiotemporal pattern of neural spike activity was measured simultaneously across 16 cortical locations spanning approximately 2-3 octaves of the tonotopic axis. Such a pattern, averaged over 40 presentations of any particular stimulus, was defined as the "cortical image" of that stimulus. Acutely deafened guinea pigs were implanted with a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 4), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. Cortical images typically showed a focus of maximum spike probability and minimum latency. Spike probabilities tended to decrease, and latencies tended to increase, with increasing cortical distance from that focus. Cortical images of TP stimuli were the most spatially compact, followed by BP + N images, and then MP images, which were the broadest. Images of CG stimuli were rather variable across animals and stimulus channels. The locations of cortical images shifted systematically from caudal to rostral as the cochlear place of stimulation changed from basal to apical. At the most sensitive cortical site for each condition, the dynamic ranges over which spike rates increased with increased current level were restricted to about 1-2 dB, regardless of configuration. Dynamic ranges tended to increase with increasing cortical distance from the most sensitive site. Electrode configurations that produced compact cortical images (e.g., TP and BP + 0) showed the greatest range of thresholds within each cortical image and the largest dynamic range at cortical sites removed from the most sensitive site. (+info
Auditory cortical images of cochlear-implant stimuli: coding of stimulus channel and current level.
This study quantified the accuracy with which populations of neurons in the auditory cortex can represent aspects of electrical cochlear stimuli presented through a cochlear implant. We tested the accuracy of coding of the place of stimulation (i.e., identification of the active stimulation channel) and of the stimulus current level. Physiological data came from the companion study, which recorded spike activity of neurons simultaneously from 16 sites along the tonotopic axis of the guinea pig's auditory cortex. In that study, cochlear electrical stimuli were presented to acutely deafened animals through a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 3), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. In the present analysis, an artificial neural network was trained to recognize spatiotemporal patterns of cortical activity in response to single presentations of particular stimuli and, thereby, to identify those stimuli. The accuracy of pair-wise discrimination of stimulation channels or of current levels was represented by the discrimination index, d', where d' = 1 was taken as threshold. In many cases, the threshold for discrimination of place of cochlear stimulation was < 0.75 mm, and the threshold for discrimination of current levels was < 1 dB. Cochlear electrode configurations varied in the accuracy with which they signaled to the auditory cortex the place of cochlear stimulation. The BP + N and TP configurations provided considerably greater sensitivity to place of stimulation than did the MP configuration. The TP configuration maintained accurate signaling of place of stimulation up to the highest current levels, whereas sensitivity was degraded at high current levels in BP + N configurations. Electrode configurations also varied in the dynamic range over which they signaled stimulus current level. Dynamic ranges were widest for the BP + 0 configuration and narrowest for the TP configuration. That is, the configuration that showed the most accurate signaling of cochlear place of stimulation (TP) showed the most restricted dynamic range for signaling of current level. These results suggest that the choice of the optimal electrode configuration for use by human cochlear-prosthesis users would depend on the particular demands of the speech-processing strategy that is to be employed. (+info