In most natural listening environments, noise occludes objects of interest, and it would be benef... more In most natural listening environments, noise occludes objects of interest, and it would be beneficial for an organism to correctly identify those objects. When a sound of interest (“foreground” sound) is interrupted by a loud noise, subjects perceive the entire sound, even if the noise was intense enough to completely mask a part of it. This phenomenon can be exploited to create an illusion: when a silent gap is introduced into the foreground and high-intensity noise is superimposed into the gap, subjects report the foreground as continuing through the noise although that portion of the foreground was deleted. This phenomenon, referred to as auditory induction or amodal completion, is conceptually similar to visual induction, fill-in, illusory motion, and illusory contours. Two rhesus macaque monkeys performed a task designed to assess auditory induction. They were trained to discriminate complete stimuli from those containing a silent gap in the presence of two types of noise. Int...
The Journal of the Acoustical Society of America, 2015
The ability to segregate simultaneous sound sources based on their spatial locations is an import... more The ability to segregate simultaneous sound sources based on their spatial locations is an important aspect of auditory scene analysis. While the role of sound azimuth in segregation is well studied, the contribution of sound elevation remains unknown. Although previous studies in humans suggest that elevation cues alone are not sufficient to segregate simultaneous broadband sources, the current study demonstrates they can suffice. Listeners segregating a temporally modulated noise target from a simultaneous unmodulated noise distracter differing in elevation fall into two statistically distinct groups: one that identifies target direction accurately across a wide range of modulation frequencies (MF) and one that cannot identify target direction accurately and, on average, reports the opposite direction of the target for low MF. A non-spiking model of inferior colliculus neurons that process single-source elevation cues suggests that the performance of both listener groups at the po...
We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animal... more We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animals discriminated amplitude-modulated (AM) sounds and also while they sat passively. Engagement in AM discrimination improved ML and A1 neurons' ability to discriminate AM with both firing rate and phase-locking; however, task engagement affected neural AM discrimination differently in the two fields. The results suggest that these two areas utilize different AM coding schemes: a "single mode" in A1 that relies on increased activity for AM relative to unmodulated sounds and a "dual-polar…
When interfering objects occlude a scene, the visual system restores the occluded information. Si... more When interfering objects occlude a scene, the visual system restores the occluded information. Similarly, when a sound of interest (a ''foreground'' sound) is interrupted (occluded) by loud noise, the auditory system restores the occluded information. This process, called auditory induction, can be exploited to create a continuity illusion. When a segment of a foreground sound is deleted and loud noise fills the missing portion, listeners incorrectly report hearing the foreground continuing through the noise. Here we reveal the neurophysiological underpinnings of illusory continuity in single-neuron responses from awake macaque monkeys' primary auditory cortex (A1). A1 neurons represented the missing segment of occluded tonal foregrounds by responding to discontinuous foregrounds interrupted by intense noise as if they were responding to the complete foregrounds. By comparison, simulated peripheral responses represented only the noise and not the occluded foreground. The results reveal that many A1 single-neuron responses closely follow the illusory percept.
Despite the extensive physiological work performed on auditory cortex, our understanding of the b... more Despite the extensive physiological work performed on auditory cortex, our understanding of the basic functional properties of auditory cortical neurons is incomplete. For example, it remains unclear what stimulus features are most important for these cells. Determining these features is challenging given the considerable size of the relevant stimulus parameter space as well as the unpredictable nature of many neurons' responses to complex stimuli due to nonlinear integration across frequency. Here we used an adaptive stimulus optimization technique to obtain the preferred spectral input for neurons in macaque primary auditory cortex (AI). This method uses a neuron's response to progressively modify the frequency composition of a stimulus to determine the preferred spectrum. This technique has the advantage of being able to incorporate nonlinear stimulus interactions into a “best estimate” of a neuron's preferred spectrum. The resulting spectra displayed a consistent, re...
The focus of most research on auditory cortical neurons has concerned the effects of rather simpl... more The focus of most research on auditory cortical neurons has concerned the effects of rather simple stimuli, such as pure tones or broad-band noise, or the modulation of a single acoustic parameter. Extending these findings to feature coding in more complex stimuli such as natural sounds may be difficult, however. Generalizing results from the simple to more complex case may be complicated by non-linear interactions occurring between multiple, simultaneously varying acoustic parameters in complex sounds. To examine this issue in the frequency domain, we performed a parametric study of the effects of two global features, spectral pattern (here ripple frequency) and bandwidth, on primary auditory (A1) neurons in awake macaques. Most neurons were tuned for one or both variables and most also displayed an interaction between bandwidth and pattern implying that their effects were conditional or interdependent. A spectral linear filter model was able to qualitatively reproduce the basic effects and interactions, indicating that a simple neural mechanism may be able to account for these interdependencies. Our results suggest that the behavior of most A1 neurons is likely to depend on multiple parameters, and so most are unlikely to respond independently or invariantly to specific acoustic features.
In classical trace conditioning the acquisition of a conditioned response (CR) is possible even t... more In classical trace conditioning the acquisition of a conditioned response (CR) is possible even though an interval (the trace interval) elapses between the conditioned stimulus (CS) and unconditioned stimulus (US). This implies that some neural representation of the CS (the stimulus trace) is able to support association between the two stimuli. The medial geniculate nucleus (MGN), particularly the medial division (mMGN), has been identified as one site in the auditory pathway where associative related changes in neural activity occur. If neurons in the MGN are involved in such a sensory trace and in acquisition of a CR, then it is expected that activity following an acoustic CS should be related to both stimulus and response. This study examined the extracellular activity of single units in the MGN during differential auditory trace conditioning of the rabbit nictitating membrane response (NMR). Two 150-ms tones (600 Hz and 1200 Hz) served as CS+ and CS-, and the US was periorbital electrostimulation. Changes in activity during the stimulus and trace interval were largest in the medial and dorsal MGN divisions on CS+ trials and on trials in which a CR was made. Examination of probe stimuli of short (50 ms) and long (600 ms) duration suggested that both CR latency and activity changes in the trace interval were related to stimulus duration and time-locked to stimulus offset. Comparisons of neural activity on the basis of fast or slow CR responses revealed different patterns of response-activity on fast CR 'trials was generally greater and tended to occur earlier. These results suggest that MGN neurons are involved in the maintenance of a sensory memory trace and possibly play a part in CR generation and timing.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
It has been argued that dyslexics suffer from temporal sensory processing de®cits which affect th... more It has been argued that dyslexics suffer from temporal sensory processing de®cits which affect their ability to discriminate speech in quiet environments. The impact of auditory de®cits on non-language aspects of perception, however, is poorly understood. In almost every natural-listening environment, one must constantly construct scenes of the auditory world by grouping and analyzing sounds generated by multiple sources. We investigated whether dyslexics have dif®culties grouping sounds. The results demonstrate that dyslexics have an impairment in grouping auditory objects that depends both on the sounds' frequency and presentation rate (i.e. the spectrotemporal context of the sound). We conclude that dyslexics have dif®culty constructing scenes of the auditory world, and that these de®cits can contribute to learning impairments.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
It has become clear that spinal cord glia (microglia and astrocytes) importantly contribute to th... more It has become clear that spinal cord glia (microglia and astrocytes) importantly contribute to the creation of exaggerated pain responses. One model used to study this is peri-spinal (intrathecal, i.t.) administration of gp120, an envelope protein of HIV-1 known to activate glia. Previous studies demonstrated that i.t. gp120 produces pain facilitation via the release of glial proinflammatory cytokines. The present series of studies tested whether spinal nitric oxide (NO) contributes to i.t. gp120-induced mechanical allodynia and, if so, what effect NO has on spinal proinflammatory cytokines. gp120 stimulation of acutely isolated lumbar dorsal spinal cords released NO as well as proinflammatory cytokines (tumor necrosis factor-alpha, interleukin-1beta (IL1), interleukin-6 (IL6)), thus identifying NO as a candidate mediator of gp120-induced behavioral effects. Behaviorally, identical effects were observed when gp120-induced mechanical allodynia was challenged by i.t. pre-treatment with either a broad-spectrum nitric oxide synthase (NOS) inhibitor (L-NAME) or 7-NINA, a selective inhibitor of NOS type-I (nNOS). Both abolished gp120-induced mechanical allodynia. While the literature pre-dominantly documents that proinflammatory cytokines stimulate the production of NO rather than the reverse, here we show that gp120-induced NO increases proinflammatory cytokine mRNA levels (RT-PCR) and both protein expression and protein release (serial ELISA). Furthermore, gp120 increases mRNA for IL1 converting enzyme and matrix metalloproteinase-9, enzymes responsible for activation and release of proinflammatory cytokines.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
In most natural listening environments, noise occludes objects of interest, and it would be benef... more In most natural listening environments, noise occludes objects of interest, and it would be beneficial for an organism to correctly identify those objects. When a sound of interest (“foreground” sound) is interrupted by a loud noise, subjects perceive the entire sound, even if the noise was intense enough to completely mask a part of it. This phenomenon can be exploited to create an illusion: when a silent gap is introduced into the foreground and high-intensity noise is superimposed into the gap, subjects report the foreground as continuing through the noise although that portion of the foreground was deleted. This phenomenon, referred to as auditory induction or amodal completion, is conceptually similar to visual induction, fill-in, illusory motion, and illusory contours. Two rhesus macaque monkeys performed a task designed to assess auditory induction. They were trained to discriminate complete stimuli from those containing a silent gap in the presence of two types of noise. Int...
The Journal of the Acoustical Society of America, 2015
The ability to segregate simultaneous sound sources based on their spatial locations is an import... more The ability to segregate simultaneous sound sources based on their spatial locations is an important aspect of auditory scene analysis. While the role of sound azimuth in segregation is well studied, the contribution of sound elevation remains unknown. Although previous studies in humans suggest that elevation cues alone are not sufficient to segregate simultaneous broadband sources, the current study demonstrates they can suffice. Listeners segregating a temporally modulated noise target from a simultaneous unmodulated noise distracter differing in elevation fall into two statistically distinct groups: one that identifies target direction accurately across a wide range of modulation frequencies (MF) and one that cannot identify target direction accurately and, on average, reports the opposite direction of the target for low MF. A non-spiking model of inferior colliculus neurons that process single-source elevation cues suggests that the performance of both listener groups at the po...
We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animal... more We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animals discriminated amplitude-modulated (AM) sounds and also while they sat passively. Engagement in AM discrimination improved ML and A1 neurons' ability to discriminate AM with both firing rate and phase-locking; however, task engagement affected neural AM discrimination differently in the two fields. The results suggest that these two areas utilize different AM coding schemes: a "single mode" in A1 that relies on increased activity for AM relative to unmodulated sounds and a "dual-polar…
When interfering objects occlude a scene, the visual system restores the occluded information. Si... more When interfering objects occlude a scene, the visual system restores the occluded information. Similarly, when a sound of interest (a ''foreground'' sound) is interrupted (occluded) by loud noise, the auditory system restores the occluded information. This process, called auditory induction, can be exploited to create a continuity illusion. When a segment of a foreground sound is deleted and loud noise fills the missing portion, listeners incorrectly report hearing the foreground continuing through the noise. Here we reveal the neurophysiological underpinnings of illusory continuity in single-neuron responses from awake macaque monkeys' primary auditory cortex (A1). A1 neurons represented the missing segment of occluded tonal foregrounds by responding to discontinuous foregrounds interrupted by intense noise as if they were responding to the complete foregrounds. By comparison, simulated peripheral responses represented only the noise and not the occluded foreground. The results reveal that many A1 single-neuron responses closely follow the illusory percept.
Despite the extensive physiological work performed on auditory cortex, our understanding of the b... more Despite the extensive physiological work performed on auditory cortex, our understanding of the basic functional properties of auditory cortical neurons is incomplete. For example, it remains unclear what stimulus features are most important for these cells. Determining these features is challenging given the considerable size of the relevant stimulus parameter space as well as the unpredictable nature of many neurons' responses to complex stimuli due to nonlinear integration across frequency. Here we used an adaptive stimulus optimization technique to obtain the preferred spectral input for neurons in macaque primary auditory cortex (AI). This method uses a neuron's response to progressively modify the frequency composition of a stimulus to determine the preferred spectrum. This technique has the advantage of being able to incorporate nonlinear stimulus interactions into a “best estimate” of a neuron's preferred spectrum. The resulting spectra displayed a consistent, re...
The focus of most research on auditory cortical neurons has concerned the effects of rather simpl... more The focus of most research on auditory cortical neurons has concerned the effects of rather simple stimuli, such as pure tones or broad-band noise, or the modulation of a single acoustic parameter. Extending these findings to feature coding in more complex stimuli such as natural sounds may be difficult, however. Generalizing results from the simple to more complex case may be complicated by non-linear interactions occurring between multiple, simultaneously varying acoustic parameters in complex sounds. To examine this issue in the frequency domain, we performed a parametric study of the effects of two global features, spectral pattern (here ripple frequency) and bandwidth, on primary auditory (A1) neurons in awake macaques. Most neurons were tuned for one or both variables and most also displayed an interaction between bandwidth and pattern implying that their effects were conditional or interdependent. A spectral linear filter model was able to qualitatively reproduce the basic effects and interactions, indicating that a simple neural mechanism may be able to account for these interdependencies. Our results suggest that the behavior of most A1 neurons is likely to depend on multiple parameters, and so most are unlikely to respond independently or invariantly to specific acoustic features.
In classical trace conditioning the acquisition of a conditioned response (CR) is possible even t... more In classical trace conditioning the acquisition of a conditioned response (CR) is possible even though an interval (the trace interval) elapses between the conditioned stimulus (CS) and unconditioned stimulus (US). This implies that some neural representation of the CS (the stimulus trace) is able to support association between the two stimuli. The medial geniculate nucleus (MGN), particularly the medial division (mMGN), has been identified as one site in the auditory pathway where associative related changes in neural activity occur. If neurons in the MGN are involved in such a sensory trace and in acquisition of a CR, then it is expected that activity following an acoustic CS should be related to both stimulus and response. This study examined the extracellular activity of single units in the MGN during differential auditory trace conditioning of the rabbit nictitating membrane response (NMR). Two 150-ms tones (600 Hz and 1200 Hz) served as CS+ and CS-, and the US was periorbital electrostimulation. Changes in activity during the stimulus and trace interval were largest in the medial and dorsal MGN divisions on CS+ trials and on trials in which a CR was made. Examination of probe stimuli of short (50 ms) and long (600 ms) duration suggested that both CR latency and activity changes in the trace interval were related to stimulus duration and time-locked to stimulus offset. Comparisons of neural activity on the basis of fast or slow CR responses revealed different patterns of response-activity on fast CR 'trials was generally greater and tended to occur earlier. These results suggest that MGN neurons are involved in the maintenance of a sensory memory trace and possibly play a part in CR generation and timing.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
It has been argued that dyslexics suffer from temporal sensory processing de®cits which affect th... more It has been argued that dyslexics suffer from temporal sensory processing de®cits which affect their ability to discriminate speech in quiet environments. The impact of auditory de®cits on non-language aspects of perception, however, is poorly understood. In almost every natural-listening environment, one must constantly construct scenes of the auditory world by grouping and analyzing sounds generated by multiple sources. We investigated whether dyslexics have dif®culties grouping sounds. The results demonstrate that dyslexics have an impairment in grouping auditory objects that depends both on the sounds' frequency and presentation rate (i.e. the spectrotemporal context of the sound). We conclude that dyslexics have dif®culty constructing scenes of the auditory world, and that these de®cits can contribute to learning impairments.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
It has become clear that spinal cord glia (microglia and astrocytes) importantly contribute to th... more It has become clear that spinal cord glia (microglia and astrocytes) importantly contribute to the creation of exaggerated pain responses. One model used to study this is peri-spinal (intrathecal, i.t.) administration of gp120, an envelope protein of HIV-1 known to activate glia. Previous studies demonstrated that i.t. gp120 produces pain facilitation via the release of glial proinflammatory cytokines. The present series of studies tested whether spinal nitric oxide (NO) contributes to i.t. gp120-induced mechanical allodynia and, if so, what effect NO has on spinal proinflammatory cytokines. gp120 stimulation of acutely isolated lumbar dorsal spinal cords released NO as well as proinflammatory cytokines (tumor necrosis factor-alpha, interleukin-1beta (IL1), interleukin-6 (IL6)), thus identifying NO as a candidate mediator of gp120-induced behavioral effects. Behaviorally, identical effects were observed when gp120-induced mechanical allodynia was challenged by i.t. pre-treatment with either a broad-spectrum nitric oxide synthase (NOS) inhibitor (L-NAME) or 7-NINA, a selective inhibitor of NOS type-I (nNOS). Both abolished gp120-induced mechanical allodynia. While the literature pre-dominantly documents that proinflammatory cytokines stimulate the production of NO rather than the reverse, here we show that gp120-induced NO increases proinflammatory cytokine mRNA levels (RT-PCR) and both protein expression and protein release (serial ELISA). Furthermore, gp120 increases mRNA for IL1 converting enzyme and matrix metalloproteinase-9, enzymes responsible for activation and release of proinflammatory cytokines.
Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remai... more Despite dyslexia affecting a large number of people, the mechanisms underlying the disorder remain undetermined. There are numerous theories about the origins of dyslexia. Many of these relate dyslexia to low-level, sensory temporal processing deficits. Another group of theories attributes dyslexia to language-specific impairments. Here, we show that dyslexics perform worse than controls on an auditory perceptual grouping task. The results show differences in performance between the groups that depend on sound frequency and not solely on parameters related to temporal processing. Performance on this task suggests that dyslexics' deficits may result from impaired attentional control mechanisms. Such deficits are neither modality nor language-specific and may help to reconcile differences between theories of dyslexia.
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