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M S Brainard

Publications and source records attributed to M S Brainard.

14 recordsLinked to original sources
PubMed2001

Postlearning consolidation of birdsong: stabilizing effects of age and anterior forebrain lesions.

Birdsong is a learned, sequenced motor skill. For the zebra finch, learned song normally remains unchanging beyond early adulthood. However, stable adult song will gradually deteriorate after deafening (Nordeen and Nordeen, 1992), indicating an ongoing influence of auditory feedback on learned song. This plasticity of adult song in response to deafening gradually declines with age (Lombardino and Nottebohm, 2000), suggesting that, after song learning, there continue to be changes in the brain that progressively stabilize the song motor program. A qualitatively similar stabilization of learned song can be precipitated artificially by lesions of a basal ganglia circuit in the songbird anterior forebrain (Brainard and Doupe, 2000), raising the question of whether and how these two forms of song stabilization are related. We investigated this issue by characterizing the deterioration of song that occurs after deafening in young adult birds and the degree to which that deterioration is reduced by age or by lesions of the anterior forebrain that were directed at the lateral portion of the magnocellular nucleus of the anterior neostriatum (LMAN). In most respects, LMAN lesions stabilized song to a significantly greater extent than did aging; whereas old-deafened birds eventually exhibited significant deterioration of song, lesioned-deafened birds generally did not differ from controls. The one exception was for song tempo, which was significantly stabilized by age, but not by LMAN lesions. The results indicate that LMAN lesions do not simply mimic a normal aging process, and likewise suggest that the anterior forebrain pathway continues to play a role even in the residual song plasticity that is observed after the age-dependent stabilization of song.

Aging↗
PubMed2000

Song selectivity and sensorimotor signals in vocal learning and production.

Bird song, like human speech, is a learned vocal behavior that requires auditory feedback. Both as juveniles, while they learn to sing, and as adults, songbirds use auditory feedback to compare their own vocalizations with an internal model of a target song. Here we describe experiments that explore a role for the songbird anterior forebrain pathway (AFP), a basal ganglia-forebrain circuit, in evaluating song feedback and modifying vocal output. First, neural recordings in anesthetized, juvenile birds show that single AFP neurons are specialized to process the song stimuli that are compared during sensorimotor learning. AFP neurons are tuned to both the bird's own song and the tutor song, even when these stimuli are manipulated to be very different from each other. Second, behavioral experiments in adult birds demonstrate that lesions to the AFP block the deterioration of song that normally follows deafening. This observation suggests that deafening results in an instructive signal, indicating a mismatch between feedback and the internal song model, and that the AFP is involved in generating or transmitting this instructive signal. Finally, neural recordings from behaving birds reveal robust singing-related activity in the AFP. This activity is likely to originate from premotor areas and could be modulated by auditory feedback of the bird's own voice. One possibility is that this activity represents an efference copy, predicting the sensory consequences of motor commands. Overall, these studies illustrate that sensory and motor processes are highly interrelated in this circuit devoted to vocal learning, as is true for brain areas involved in speech.

Animal Communication↗
PubMed2000

Interruption of a basal ganglia-forebrain circuit prevents plasticity of learned vocalizations.

Birdsong, like speech, is a learned vocal behaviour that relies greatly on hearing; in both songbirds and humans the removal of auditory feedback by deafening leads to a gradual deterioration of adult vocal production. Here we investigate the neural mechanisms that contribute to the processing of auditory feedback during the maintenance of song in adult zebra finches. We show that the deleterious effects on song production that normally follow deafening can be prevented by a second insult to the nervous system--the lesion of a basal ganglia-forebrain circuit. The results suggest that the removal of auditory feedback leads to the generation of an instructive signal that actively drives non-adaptive changes in song; they also suggest that this instructive signal is generated within (or conveyed through) the basal ganglia-forebrain pathway. Our findings provide evidence that cortical-basal ganglia circuits may participate in the evaluation of sensory feedback during calibration of motor performance, and demonstrate that damage to such circuits can have little effect on previously learned behaviour while conspicuously disrupting the capacity to adaptively modify that behaviour.

Animals↗
PubMed2000

Auditory feedback in learning and maintenance of vocal behaviour.

Songbirds are one of the best-studied examples of vocal learners. Learning of both human speech and birdsong depends on hearing. Once learned, adult song in many species remains unchanging, suggesting a reduced influence of sensory experience. Recent studies have revealed, however, that adult song is not always stable, extending our understanding of the mechanisms involved in song maintenance, and their similarity to those active during song learning. Here we review some of the processes that contribute to song learning and production, with an emphasis on the role of auditory feedback. We then consider some of the possible neural substrates involved in these processes, particularly basal ganglia circuitry. Although a thorough treatment of human speech is beyond the scope of this article, we point out similarities between speech and song learning, and ways in which studies of these disparate behaviours complement each other in developing an understanding of general principles that contribute to learning and maintenance of vocal behaviour.

Acoustic Stimulation↗
PubMed1998

Sensitive periods for visual calibration of the auditory space map in the barn owl optic tectum.

Previous studies have identified sensitive periods for the developing barn owl during which visual experience has a powerful influence on the calibration of sound localization behavior. Here we investigated neural correlates of these sensitive periods by assessing developmental changes in the capacity of visual experience to alter the map of auditory space in the optic tectum of the barn owl. We used two manipulations. (1) We equipped owls with prismatic spectacles that optically displaced the visual field by 23 degrees to the left or right, and (2) we restored normal vision to prism-reared owls that had been raised wearing prisms. In agreement with previous behavioral experiments, we found that the capacity of abnormal visual experience to shift the tectal auditory space map was restricted to an early sensitive period. However, this period extended until later in life (approximately 200 d) than described previously in behavioral studies (approximately 70 d). Furthermore, unlike the previous behavioral studies that found that the capacity to recover normal sound localization after restoration of normal vision was lost at approximately 200 d of age, we found that the capacity to recover a normal auditory space map was never lost. Finally, we were able to reconcile the behaviorally and neurophysiologically defined sensitive periods by taking into account differences in the richness of the environment in the two sets of experiments. We repeated the behavioral experiments and found that when owls were housed in a rich environment, the capacity to adjust sound localization away from normal extended to later in life, whereas the capacity to recover to normal was never lost. Conversely, when owls were housed in an impoverished environment, the capacity to recover a normal auditory space map was restricted to a period ending at approximately 200 d of age. The results demonstrate that the timing and even the existence of sensitive periods for plasticity of a neural circuit and associated behavior can depend on multiple factors, including (1) the nature of the adjustment demanded of the system and (2) the richness of the sensory and social environment in which the plasticity is studied.

Aging↗
PubMed1996

Newly learned auditory responses mediated by NMDA receptors in the owl inferior colliculus.

Alignment of auditory and visual receptive fields in the optic tectum of the barn owl (Tyto alba) is maintained through experience-dependent modification of auditory responses in the external nucleus of the inferior colliculus (ICX), which provides auditory input to the tectum. Newly learned tectal auditory responses, induced by altered visual experience, were found to be pharmacologically distinct from normal responses expressed at the same tectal sites. N-methyl-D-aspartate (NMDA) receptor antagonists administered systemically or applied locally in the ICX reduced learned responses more than normal responses. This differential blockade was not observed with non-NMDA or broad-spectrum antagonists. Thus, NMDA receptors preferentially mediate the expression of novel neuronal responses induced by experience during development.

2-Amino-5-phosphonovalerate↗
PubMed1995

Dynamics of visually guided auditory plasticity in the optic tectum of the barn owl.

1. In the optic tectum of normal barn owls, bimodal (auditory-visual) neurons are tuned to the values of interaural time difference (ITD) that are produced by sounds at the locations of their visual receptive fields (VRFs). The auditory tuning of tectal neurons is actively guided by visual experience during development: in the tectum of adult owls reared with an optically displaced visual field, neurons are tuned to abnormal values of ITD that are close to the values produced by sounds at the locations of their optically displaced VRFs. In this study we investigated the dynamics of this experience-dependent plasticity. 2. Owls were raised from shortly after eye-opening (14-22 days of age) with prismatic spectacles that displaced the visual field to the right or left. Starting at approximately 60 days of age, multiunit recordings were made to assess the tuning of tectal neurons to ITD presented via earphones. In the earliest recording sessions (ages 60-80 days), ITD tuning was often close to normal, even though the majority of the owls' previous experience was with an altered correspondence between ITD values and VRF locations. Subsequently, over a period of weeks, responses to the normal range of ITDs were gradually eliminated while responses to values of ITD corresponding with the optically displaced VRF were acquired. 3. At intermediate stages in this process, the ITD tuning at many sites became abnormally broad, so that responses were simultaneously present to both normal values of ITD and to values corresponding with the optically displaced VRF. At this stage the latencies and durations of newly acquired responses systematically exceeded the latencies and durations of the responses to normal values of ITD. 4. Dynamic changes in ITD tuning similar to those recorded in the optic tectum also occurred in the external nucleus of the inferior colliculus (ICX), which provides the major source of ascending auditory input to the tectum. 5. These results suggest the hypothesis that the neural selectivity for ITD in the barn owl's tectum is first established by vision-independent mechanisms and only subsequently calibrated by visual experience. This calibration involves both the elimination of responses to normal values of ITD and the visually guided acquisition of responses to novel values and can be accounted for by plasticity at the level of the ICX.

Animals↗
PubMed1994

Neural substrates of sound localization.

Recent behavioral experiments suggest that parallel pathways in the forebrain and midbrain are capable of mediating sound localization. Neurophysiological studies have advanced our understanding of how binaural and monaural localization cues contribute to the representations of auditory space within these pathways. Experience-dependent modifications of auditory spatial tuning in both the forebrain and midbrain pathways indicate that sensory input during development customizes an individual's neural processing of spatial information.

Animals↗
PubMed1993

Experience-dependent plasticity in the inferior colliculus: a site for visual calibration of the neural representation of auditory space in the barn owl.

The optic tectum (homolog of the superior colliculus) contains mutually aligned neural maps of auditory and visual space. During development, the organization of the auditory map is guided by spatial information provided by vision: barn owls raised wearing prismatic spectacles, which optically shift the visual field and the visual map in the optic tectum, develop an auditory map that is shifted by an approximately equivalent amount, such that alignment between the two maps is preserved (Knudsen and Brainard, 1991). In this study we investigated whether this shift in the auditory map is intrinsic to the optic tectum or whether it reflects plasticity at an earlier stage in the auditory pathway. Owls were raised wearing prismatic spectacles that displaced the visual field by 23 degrees to the left or right. This manipulation alters the normal correspondence between locations in the visual field and interaural time difference (ITD), the primary cue for the azimuth of a sound source. In normal owls and in owls with at least 150 d of prism experience, extracellular unit recordings were used to assess the representations of ITD at anatomically and physiologically defined sites in the optic tectum and in the two prior stages of the auditory pathway, the external and central nuclei of the inferior colliculus (ICx and ICc). In the optic tectum of normal owls, the values of ITD to which units responded most strongly (best ITDs) varied systematically with the azimuths of unit visual receptive fields (VRFs). In the prism-reared owls, best ITDs were shifted from normal toward the values of ITD produced by sounds at the locations of the units' optically displaced VRFs. In the ICx of prism-reared owls, the representation of ITD also was shifted from normal, by an amount and in a direction that could completely account for the shift in ITD measured in the optic tectum. At some sites in the ICx, the shift in ITD tuning was apparent within the first 7-8 msec of the response; shifted tuning at such short latencies argues that the altered representation of ITD in the ICx reflects plasticity in the ascending auditory pathway, and is not the result of descending activity from higher auditory centers. In the ICc, which immediately precedes the ICx in the ascending pathway, the representation of ITD was normal. The results indicate that the visual instruction of auditory spatial tuning of neurons in the optic tectum reflects plasticity at the level of the ICx, the site where the auditory map of space is first synthesized.

Acoustic Stimulation↗
PubMed1992

Neural derivation of sound source location: resolution of spatial ambiguities in binaural cues.

Cues for sound localization are inherently spatially ambiguous. Nevertheless, most neurons in the barn owl's optic tectum (superior colliculus) have receptive fields for broadband noise stimuli that are restricted to a single region of space. This study characterizes the spatial ambiguities associated with two important sets of localization cues, interaural level differences (ILDs) and interaural phase differences (IPDs), and describes how information is integrated within and across frequencies to resolve these ambiguities. The auditory receptive fields of neurons in the optic tectum were measured with free-field sounds presented from a movable loudspeaker. In contrast to the single regions typical for broadband receptive fields, receptive fields for tonal stimuli usually included additional discrete regions of space (accessory fields). Based on acoustic measurements of ILD and IPD cues made in the external ear canals, it was shown that accessory fields corresponded to locations from which sound sources produced ILD and IPD values that were approximately the same as those arising from the broadband receptive field. In addition, accessory fields had inhibitory surrounds, corresponding to locations from which sound sources produced substantially different combinations of ILD and IPD values. Where an accessory field for one frequency overlapped with the inhibitory surround of a second frequency, an excitatory response to the first frequency could be reduced or eliminated by addition of the second frequency. Because tonal receptive fields for different frequencies always overlapped in the region of the broadband receptive field but tended not to overlap elsewhere, this integration of excitation and inhibition can account for the restriction of broadband receptive fields to a single region of space.

Animals↗
PubMed1991

Visual instruction of the neural map of auditory space in the developing optic tectum.

Neural maps of visual and auditory space are aligned in the adult optic tectum. In barn owls, this alignment of sensory maps was found to be controlled during ontogeny by visual instruction of the auditory spatial tuning of neurons. Large adaptive changes in auditory spatial tuning were induced by raising owls with displacing prisms mounted in spectacle frames in front of the eyes; neurons became tuned to sound source locations corresponding to their optically displaced, rather than their normal, visual receptive field locations. The results demonstrate that visual experience during development calibrates the tectal auditory space map in a site-specific manner, dictating its topography and alignment with the visual space map.

Animals↗
PubMed1988

Single-channel analysis of four distinct classes of potassium channels in Drosophila muscle.

A number of mutations have been shown to affect potassium channels in Drosophila muscle. Single-channel analysis of the effects of mutations will prove a powerful approach for studying the molecular mechanisms of ion channel gating. As an initial step towards studying the effects of mutations at the single-channel level, we have characterized wild-type potassium channels in cultured embryonic myotubes using whole-cell, cell-attached, inside-out, and outside-out configurations of the patch-clamp technique. The myotubes differentiate in vitro from primary cultures of late-gastrula stage embryos of Drosophila. The whole-cell outward currents develop in a characteristic sequence. At 8 hr after plating a small delayed outward current is present. Between 10 and 12 hr after plating an A-type outward current develops, followed, between 13 and 16 hr, by a large increase in the delayed current. The A-type current is absent at all developmental stages in myotubes homozygous for the mutant ShKS133. At least 4 different types of potassium channels contribute to the whole-cell outward currents: a fast transient 14 pS A-type potassium channel (A1), a slowly inactivating 14 pS potassium channel (KD), a 40 pS potassium channel that does not inactivate during voltage pulses up to 2.4 sec in duration (KO), and a 90 pS potassium channel that is strongly activated by membrane stretch (KST). Channels indistinguishable from the KD and KST channels were also observed in patch-clamp studies on larval body wall muscle fibers. A1 channels were also present in intact dorsal longitudinal flight muscles. The A1 channel underlies the rapidly inactivating component of the whole-cell current. It inactivates with a similar time course and voltage dependence to the A-current and is similarly blocked by 5 mM 4-aminopyridine. The KD channel underlies a large fraction of the delayed component of the whole-cell current. Ensemble averages of single KD channels inactivate with the same time course as the delayed current. The KO channel represents a smaller fraction of the whole-cell delayed outward current. Its increase in open probability with voltage is due primarily to a voltage dependence of its closed times. The KST channel is voltage and calcium independent and would therefore only contribute to the leak whole-cell current.

Animals↗