P F Knudsen

Stanford University, Stanford, California, United States

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Publications (15)134.31 Total impact

  • Source
    Ilana B Witten · Phyllis F Knudsen · Eric I Knudsen
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    ABSTRACT: Barn owls integrate spatial information across frequency channels to localize sounds in space. We presented barn owls with synchronous sounds that contained different bands of frequencies (3-5 kHz and 7-9 kHz) from different locations in space. When the owls were confronted with the conflicting localization cues from two synchronous sounds of equal level, their orienting responses were dominated by one of the sounds: they oriented toward the location of the low frequency sound when the sources were separated in azimuth; in contrast, they oriented toward the location of the high frequency sound when the sources were separated in elevation. We identified neural correlates of this behavioral effect in the optic tectum (OT, superior colliculus in mammals), which contains a map of auditory space and is involved in generating orienting movements to sounds. We found that low frequency cues dominate the representation of sound azimuth in the OT space map, whereas high frequency cues dominate the representation of sound elevation. Significance: We argue that the dominance hierarchy of localization cues reflects several factors: 1) the relative amplitude of the sound providing the cue, 2) the resolution with which the auditory system measures the value of a cue, and 3) the spatial ambiguity in interpreting the cue. These same factors may contribute to the relative weighting of sound localization cues in other species, including humans.
    Preview · Article · Apr 2010 · PLoS ONE
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    Kristin A Maczko · Phyllis F Knudsen · Eric I Knudsen
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    ABSTRACT: The nucleus isthmi pars parvocellularis (Ipc) is a midbrain cholinergic nucleus that shares reciprocal, topographic connections with the optic tectum (OT). Ipc neurons project to spatially restricted columns in the OT, contacting essentially all OT layers in a given column. Previous research characterizes the Ipc as a visual processor. We found that, in the barn owl, the Ipc responds to auditory as well as to visual stimuli. Auditory responses were tuned broadly for frequency, but sharply for spatial cues. We measured the tuning of Ipc units to binaural sound localization cues, including interaural timing differences (ITDs) and interaural level differences (ILDs). Units in the Ipc were tuned to specific values of both ITD and ILD and were organized systematically according to their ITD and ILD tuning, forming a map of space. The auditory space map aligned with the visual space map in the Ipc. These results demonstrate that the Ipc encodes the spatial location of objects, independent of stimulus modality. These findings, combined with the precise pattern of projections from the Ipc to the OT, suggest that the role of the Ipc is to regulate the sensitivity of OT neurons in a space-specific manner.
    Preview · Article · Jan 2007 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
  • Eric I. Knudsen · Phyllis F. Knudsen
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    ABSTRACT: Barn owls not only localize auditory stimuli with great accuracy, they also remember the locations of auditory stimuli and can use this remembered spatial information to guide their flight and strike. Although the mechanisms of sound localization have been studied extensively, the neurobiological basis of auditory spatial memory has not. Here we show that the ability of barn owls to orient their gaze towards and fly to the remembered location of auditory targets is lost during pharmacological inactivation of a small region in the forebrain, the anterior archistriatum. In contrast, archistriatal inactivation has no effect on stimulus-guided responses to auditory targets. The memory-dependent deficit is evident only for acoustic events that occur in the hemifield contralateral to the side that is inactivated. The data demonstrate that in the avian archistriatum, as in the mammalian frontal cortex, there exists a region that is essential for the expression of spatial working memory and that, in the barn owl, this region encodes auditory spatial memory.
    No preview · Article · Nov 1996 · Nature
  • Eric I. Knudsen · Phyllis F. Knudsen
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    ABSTRACT: A region in the barn owl forebrain, referred to as the archistriatal gaze fields (AGF), is shown to be involved in auditory orienting behavior. In a previous study, electrical microstimulation of the AGF was shown to produce saccadic movements of the eyes and head, and anatomical data revealed that neurons in the AGF region of the archistriatum project directly to brainstem tegmental nuclei that mediate gaze changes. In this study, we investigated the effects of AGF inactivation on the auditory orienting responses of trained barn owls. The AGF and/or the optic tectum (OT) were inactivated pharmacologically using the GABAA agonist muscimol. Inactivation of the AGF alone had no effect on the probability or accuracy of orienting responses to contralateral acoustic stimuli. Inactivation of the OT alone decreased the probability of responses to contralateral stimuli, but the animals were still capable of orienting accurately toward stimuli on about 60% of the trials. Inactivation of both the AGF and the OT drastically decreased the probability of responses to 16-21% and, on the few trials that the animals did respond, there was no relationship between the final direction of gaze and the location of the stimulus. Thus, with the AGF and OT both inactivated, the animals were no longer capable of orienting accurately toward acoustic stimuli located on the contralateral side. These data confirm that the AGF is involved in gaze control and that the AGF and the OT have parallel access to gaze control circuitry in the brainstem tegmentum. In these respects, the AGF in barn owls is functionally equivalent to the frontal eye fields in primates.
    No preview · Article · Mar 1996 · Experimental Brain Research
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    E I Knudsen · P F Knudsen · T Masino
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    ABSTRACT: The hypothesis that sound localization and gaze control are mediated in parallel in the midbrain and forebrain was tested in the barn owl. The midbrain pathway for gaze control was interrupted by reversible inactivation (muscimol injection) or lesion of the optic tectum. Auditory input to the forebrain was disrupted by reversible inactivation or lesion of the primary thalamic auditory nucleus, nucleus ovoidalis (homolog of the medial geniculate nucleus). Barn owls were trained to orient their gaze toward auditory or visual stimuli presented from random locations in a darkened sound chamber. Auditory and visual test stimuli were brief so that the stimulus was over before the orienting response was completed. The accuracy and kinetics of the orienting responses were measured with a search coil attached to the head. Unilateral inactivation of the optic tectum had immediate and long-lasting effects on auditory orienting behavior. The owls failed to respond on a high percentage of trials when the auditory test stimulus was located on the side contralateral to the inactivated tectum. When they did respond, the response was usually (but not always) short of the target, and the latency of the response was abnormally long. When the auditory stimulus was located on the side ipsilateral to the inactivated tectum, responses were reliable and accurate, and the latency of responses was shorter than normal. In a tectally lesioned animal, response probability and latency to contralateral sounds returned to normal within 2 weeks, but the increase in response error (due to undershooting) persisted for at least 12 weeks. Despite abnormalities in the response, all of the owls were capable of localizing and orienting to contralateral auditory stimuli on some trials with the optic tectum inactivated or lesioned. This was not true for contralateral visual stimuli. Immediately following tectal inactivation, the owls exhibited complete neglect for visual stimuli located more than 20 degrees to the contralateral side (i.e., beyond the edge of the visual field of the ipsilateral eye). In the tectally lesioned animal, this neglect diminished with time. Unilateral inactivation of nucleus ovoidalis had different effects in three owls. Response error to contralateral sound sources increased for one owl and decreased for two; response error to ipsilateral sources did not change significantly for any. The probability of response to ipsilateral (but not contralateral) stimuli decreased for one owl. The latency of response to ipsilateral (but not contralateral) stimuli increased for one and decreased for another. All of the owls, however, routinely localized and oriented toward ipsilateral and contralateral auditory stimuli with nucleus ovoidalis inactivated.(ABSTRACT TRUNCATED AT 400 WORDS)
    Preview · Article · Aug 1993 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen
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    ABSTRACT: This study describes developmental changes in the capacity of owls to adjust sound localization in response to chronic prismatic displacement of the visual field and to recover accurate sound localization following the restoration of normal vision. Matched, binocular displacing prisms were mounted over the eyes of 19 barn owls (Tyto alba) beginning at ages ranging from 10 to 272 d. In nearly all cases, the visual field was shifted 23 degrees to the right. Sound localization was assessed on the basis of head orientations to sound sources, measured in a darkened sound chamber with a search coil system. Chronic exposure to a displaced visual field caused the owls to alter sound localization in the direction of the visual field displacement, thereby inducing a sound-localization error. The size of the sound-localization error that resulted depended on the age of the animal when prism experience began. Maximal errors of about 20 degrees were induced only when prism experience began by 21 d of age. As prism experience began at later ages, the magnitude of induced errors decreased. A bird that wore prisms beginning at 102 d of age, altered sound localization by only 6 degrees. An adult owl, when exposed chronically to a displaced visual field, altered sound localization by about 3 degrees. We refer to the early period in life when displaced vision induces exceptionally large sound-localization errors (relative to those induced in the adult) as a sensitive period. The capacity to recover accurate sound localization following restoration of normal vision was tested in 7 owls that had been raised wearing prisms. Four owls that had prisms removed by 182 d of age recovered accurate localization rapidly (over a period of weeks), whereas 3 owls that were older when the prisms were removed did not recover accurate localization when tested for up to 7 months after prism removal. Adjustment of sound localization slowed greatly or ceased at about 200 days of age, referred to here as the critical period for visual calibration of sound localization. Three owls were subjected repetitively to displacement of the visual field. An owl that adjusted sound localization to the left of normal during the sensitive period retained the capacity to adjust again to the left, but not to the right of normal, later in the critical period. The converse was true for an owl that adjusted sound localization to the right of normal during the sensitive period.(ABSTRACT TRUNCATED AT 400 WORDS)
    Preview · Article · Feb 1990 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen
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    ABSTRACT: The capacity of barn owls to adapt visuomotor behavior in response to prism-induced displacement of the visual field was tested in babies and adults. Matched, binocular Fresnel prisms, which displaced the visual field 11 degrees, 23 degrees, or 34 degrees to the right, were placed on owls for periods of up to 99 d. Seven baby owls wore the prisms from the day the eyelids first opened; 2 owls wore them as adults. Prism adaptation was measured by the accuracy with which a target was approached and struck with the talons, a behavior similar to pointing behavior used commonly to assess prism adaptation in primates. Baby and adult owls exhibited a limited capacity to adapt this visuomotor behavior. Acquisition of adapted behavior was slow, taking place over a period of weeks, and was never complete even for owls that were raised viewing the world through relatively weak (11 degrees) displacing prisms. When the prisms were removed from adapted owls, they struck to the opposite side of the target. The recovery of strike accuracy following prism removal was rapid; 7 of 9 owls recovered normal accuracy within 30 min of prism removal, despite having worn the prisms for months. This limited capacity for adaptation contrasts dramatically with the extensive and rapid adaptation exhibited by adult primates exposed to comparable prismatic displacements. The mechanism of adaptation used by the owls was to alter the movements employed for approaching targets. Instead of moving straight ahead, the head and body moved diagonally relative to the orientation of the head. Thus, in contrast to prism adaptation by humans that can involve reinterpretation of eye, head, and limb position, prism adaptation by owls is based on changes in the motor commands that underlie approach behavior.
    Preview · Article · Oct 1989 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen
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    ABSTRACT: This study demonstrates that continuous exposure of baby barn owls to a displaced visual field causes a shift in sound localization in the direction of the visual displacement. This implies an innate dominance of vision over audition in the development and maintenance of sound localization. Twelve owls were raised from the first day of eye opening wearing binocular prisms that displaced the visual field to the right by 11 degrees, 23 degrees, or 34 degrees. The prisms were worn for periods of up to 7 months. Consistent with previous results (Knudsen and Knudsen, 1989a), owls reared with displacing prisms did not adjust head orientation to visual stimuli. While wearing prisms, owls consistently oriented the head to the right of visual targets, and, as soon as the prisms were removed, they oriented the head directly at visual targets, as do normal owls. In contrast, prism-reared owls did change head orientation to sound sources even though auditory cues were not altered significantly. Birds reared wearing 11 degrees or 23 degrees prisms oriented the head to the right of acoustic targets by an amount approximately equal to the optical displacement induced by the prisms. Birds raised wearing 34 degrees prisms adjusted sound localization by only about 50% of the optical displacement. Thus, visually guided adjustment of sound localization appears to be limited to about 20 degrees in azimuth. The data indicate that when confronted with consistently discordant localization information from the auditory and visual systems, developing owls use vision to calibrate associations of auditory localization cues with locations in space in an attempt to bring into alignment the perceived locations of auditory and visual stimuli emanating from a common source. Vision exerts this instructive influence on sound localization whether or not visual information is accurate.
    Preview · Article · Oct 1989 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen
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    ABSTRACT: Early in life, the barn owl passes through a sensitive period during which it can interpret and make use of abnormal auditory cues for accurate sound localization. This capacity is lost at about 8 weeks of age, just after the head and ears reach adult size (knudsen et al. 1984a). The end of the sensitive period could be triggered either by an age-dependent process or by the exposure of the auditory system to stable or adult-like cues. To distinguish between these alternatives, we subjected baby owls to constant abnormal cues (chronic monaural occlusion) or to frequently changing abnormal cues (alternating monaural occlusion) throughout the sensitive period. In the first group of animals (n = 2), one ear was plugged continuously until 73 or 79 d of age, respectively, and then the earplug was switched to the opposite ear. Although these animals adjusted sound localization accuracy during the initial chronic monaural occlusion, they could not localize sounds at all after the earplug was switched to the opposite ear, and they remained unable to localize sounds as long as the opposite ear remained occluded (7 and 27 weeks, respectively). When the second monaural occlusion was finally removed, both birds localized sounds with errors that were similar to the errors they exhibited immediately after removal of the first monaural occlusion. One bird that was 127-d-old at the time the second earplug was removed corrected its localization error; the other bird, 250-d-old when the second earplug was removed, did not.(ABSTRACT TRUNCATED AT 250 WORDS)
    Preview · Article · Aug 1986 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen
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    ABSTRACT: Barn owls raised with one ear plugged make systematic errors in auditory localization when the earplug is removed. Young owls correct their localization errors within a few weeks. However, such animals did not correct their auditory localization errors when deprived of vision. Moreover, when prisms were mounted in front of their eyes, they adjusted their auditory localization to match the visual error induced by the prisms, as long as the visual and auditory errors were within the same quadrant of directions. The results demonstrate that, during development, the visual system provides the spatial reference for fine-tuning auditory localization.
    Preview · Article · Dec 1985 · Science
  • E. I. Knudsen · P. F. Knudsen

    No preview · Article · Jan 1985
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    E I Knudsen · S D Esterly · P F Knudsen
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    ABSTRACT: Sound localization was disrupted in young barn owls by chronically plugging one ear. Owls that were younger than 8 weeks of age at the time of ear plugging recovered normal localization accuracy while plugged, whereas those that were older than 8 weeks at the time of ear plugging did not. The end of the sensitive period for the adjustment of sound localization accuracy coincides with the maturation of the head and ears, suggesting that the exposure of the auditory system to stable, adult-like acoustic cues could play a role in bringing the sensitive period to a close. The results demonstrate that, early in development, associations between auditory cues and locations in space can be altered by experience.
    Preview · Article · May 1984 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
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    E I Knudsen · P F Knudsen · S D Esterly
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    ABSTRACT: We studied the ability of barn owls to recover accurate sound localization after being raised with one ear occluded. Most of the owls had ear plugs inserted before they reached adult size, and therefore they never experienced normal adult localization cues until their ear plugs were removed. Upon removal of their ear plugs, these owls exhibited large systematic sound localization errors. The rate at which they recovered accurate localization decreased with the age of the bird at the time of plug removal, and recovery essentially ceased when owls reached 38 to 42 weeks of age. We interpret this age as the end of a critical period for the consolidation of associations between auditory cues and locations in space. Owls that had experienced adult localization cues for a short period of time before ear plugging recovered normal accuracy rapidly, even if they remained plugged well past the end of the critical period. This suggests that a brief exposure to normal adult cues early in the critical period is sufficient to enable the recovery of localization accuracy much later in life.
    Preview · Article · May 1984 · The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
  • Eric I. Knudsen · Phyllis F. Knudsen
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    ABSTRACT: The optic tectum of the owl contains a topographic representation of auditory space. We have investigated the source of this space-mapped auditory activity by using retrograde tracing with horseradish peroxidase. The major source of auditory input to the optic tectum is the ipsilateral external nucleus of the inferior colliculus (ICX), which is known to contain a map of auditory space also. Additional minor projections originate in the superficial nucleus of the inferior colliculus and the nucleus isthmi parvocellularis. There is no apparent projection to the optic tectum from the contralateral ICX or any other brainstem auditory nucleus. The projection from the ICX to the optic tectum is point-to-point: rostral ICX projects to rostral tectum, caudal ICX to caudal tectum, dorsal ICX to dorsomedial tectum, and ventral ICX to ventral tectum. Thus, the space-mapped organization that exists in the ICX is passed on by topographic projections to the optic tectum.
    No preview · Article · Aug 1983 · The Journal of Comparative Neurology
  • Eric I. Knudsen · Phyllis F. Knudsen · Steven D. Esterly
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    ABSTRACT: The auditory system localizes sounds by comparing the timing and intensity of sounds arriving at the two ears, and associating specific binaural differences with directions of sounds in space. Thus, location for the auditory system, like stereoscopic depth for the visual system, is a percept created in the brain by a comparison of inputs from two receivers. We considered whether the neural mechanism underlying sound localization is determined entirely genetically, or if it is modified and regulated by auditory experience. The influence of experience on sound localization has been examined previously in various species, including man, with contradictory results1–9. For our study we used the barn owl (Tyto alba) as an experimental model because it localizes sounds with extreme accuracy10,11. By placing a plug in one ear, we disrupted the owl's binaural localization cues and induced large errors in sound localization. We report here that young owls adjusted to the altered cues and regained normal localization accuracy over a period of weeks. However, as the owls aged, their rate of adjustment slowed, and beyond 6–7 months of age, their capacity to adjust was lost or greatly reduced. This plasticity of the sound localization mechanism early in life thus enables the auditory system to establish precise correlations between binaural cues (which vary between individuals) and directions of sounds in space.
    No preview · Article · Jan 1982

Publication Stats

809 Citations
134.31 Total Impact Points

Institutions

  • 1996-2010
    • Stanford University
      • Department of Neurobiology
      Stanford, California, United States
  • 1983-1989
    • Stanford Medicine
      • Department of Neurobiology
      Stanford, California, United States