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Biomedical subjects

E I Knudsen

Publications and source records attributed to E I Knudsen.

At least 19 recordsLinked to original sources
PubMed1992

Anatomical pathways from the optic tectum to the spinal cord subserving orienting movements in the barn owl.

Electrical stimulation of the optic tectum in many vertebrate species elicits eye, head or body orienting movements in the direction of the receptive field location recorded at the site of stimulation; in the barn owl, tectal stimulation produces short latency saccadic head movements (du Lac and Knudsen 1990). However, the barn owl, like other avians, lacks a direct projection from the tectum to the spinal cord, implying that less direct connections underlie tectally mediated head movements. In order to determine the pathways by which the tectum gains access to spinal cord circuitry, we searched for overlap regions between tectal efferent projections and the locations of cells afferent to the spinal cord. Tectal efferent pathways and terminal fields were revealed by anterograde labeling using horseradish peroxidase (HRP) or tritiated amino acids injected into the optic tectum. Cells afferent to the spinal cord were identified by means of retrograde labeling using HRP, rhodamine, or rhodamine-coupled latex beads injected into the cervical spinal cord. A comparison of results from the anterograde and retrograde labeling experiments demonstrated several areas of overlap. All of the cell groups that both received heavy tectal input and contained a high proportion of cells projecting to the spinal cord were located in the medial half of the midbrain and rhombencephalic tegmentum, and included the red nucleus, the interstitial nucleus of Cajal, the medial reticular formation, the nucleus reticularis pontis giganto-cellularis, and the nucleus reticularis pontis oralis. All of these cell groups receive their tectal input from the medial efferent pathway, one of three major output pathways from the tectum. The other two output pathways (the rostral and the caudal) project to regions containing no more than a few scattered cells that are afferent to the spinal cord. Based on these data and on the functions of homologous cell groups in other vertebrates, we hypothesize that the medial efferent pathway and its brainstem target nuclei are primarily responsible for tectally mediated orienting head movements in the barn owl.

Animals↗
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↗
PubMed1992

Adaptive adjustment of unit tuning to sound localization cues in response to monaural occlusion in developing owl optic tectum.

Bimodal units in the barn owl's optic tectum are tuned to the location of auditory and visual stimuli, and are systematically organized according to their spatial tuning to form mutually aligned maps of auditory and visual space. Map alignment results from the fact that, normally, units are tuned to the values of interaural level difference (ILD) and interaural time difference (ITD) produced by a sound source at the location of their visual receptive fields (VRFs). Monaural occlusion alters the correspondence of ILD and ITD values with locations in space. We investigated the effect that raising owls with a chronic monaural occlusion has on the tuning of tectal units to ILD and ITD. Owls were monaurally occluded beginning at 1 month of age. The effects of monaural occlusion were assessed 2-4 months later by comparing the ILD and ITD tuning of units in monaurally occluded owls with the ILD and ITD tuning of units with equivalent VRFs in normal owls. ILD and ITD tuning was shifted substantially and in the direction of the unoccluded ear (the adaptive direction) in owls raised with a monaural occlusion. In most tecta, the mapped representations of ILD and ITD were shifted systematically. In addition, in some tecta, monaural occlusion induced a change in the topography of the ILD map such that ILD tuning remained essentially constant at values near 0 dB over abnormally large portions of the tectum. Across all recording sites, the average shift in ILD tuning was 9 dB (n = 396) and the average shift in ITD tuning was 40 microseconds (n = 414). In four of five animals, the magnitude of the effect was not equivalent on the two sides of the brain, the adjustments being significantly larger and more systematic on the side ipsilateral to the occlusion. Such differences in the altered ILD and ITD maps on the two sides of the brain in individual animals indicate that, although a component of the adaptive adjustment might be due to regulation of the gain and phase response of the monaural signals early in the auditory pathway, a major component of the adjustment must occur at or beyond the level where the encoding of ILDs and ITDs for left and right space separates.

Adaptation, Physiological↗
PubMed1992

Vision-independent adjustment of unit tuning to sound localization cues in response to monaural occlusion in developing owl optic tectum.

Neurons in the developing optic tectum adjust their tuning to auditory localization cues in response to chronic monaural occlusion so that auditory spatial fields align with visual receptive fields (VRFs). We tested whether this adaptive adjustment of auditory tuning requires visual instruction. Both eyelids were sutured closed at the same time that one ear was occluded in two barn owls that were 1 month old. After 70 and 100 d, respectively, the tuning of units to interaural level difference (ILD) and to interaural time difference (ITD) was measured. These data were compared with equivalent data from 15 normal owls. Unit tuning to ITD was shifted from normal in both of the monaurally occluded owls. In one owl, ILD tuning was also clearly shifted. In the other owl, the map of ILD was flipped upside down and adaptive adjustments in ILD tuning could not be assessed. Instead, adjustments in ILD tuning were observed following removal of the earplug with the eyelids kept closed. Unit tuning was monitored at several sites in the tectum for 1 month after earplug removal using chronically implanted electrodes. Then, ILD tuning was resampled across the entire tectum. Both measures indicated shifts in ILD tuning in response to removal of the earplug in the second blind owl. In both animals, the magnitude of the shifts in ILD tuning and ITD tuning was smaller than has been observed previously in monaurally occluded but sighted owls. The results demonstrate that the brain can make adaptive adjustments in ILD and ITD tuning in response to early monaural occlusion even without the guiding influence of vision.

Adaptation, Physiological↗
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↗
PubMed1991

Early visual deprivation results in a degraded motor map in the optic tectum of barn owls.

The optic tectum contains a precise map of orienting movements: the size and direction of movements of the eyes, head, and/or body vary systematically with the locus of neural activation within the tectum. In adult animals, this motor map aligns closely with the tectal map of visual space. This study addressed the question of whether the motor map develops entirely independently of visual experience. We found that in barn owls (Tyto alba) raised without vision, although a tectal map of head movement develops, its topography and alignment with the map of visual (and auditory) space are abnormal. The results demonstrate that during early life vision is necessary either to maintain or to guide the development of a normal tectal motor map.

Animals↗
PubMed1991

Dynamic space codes in the superior colliculus.

Space coding in the superior colliculus has traditionally been viewed as a static representation by multiple, aligned, sensory and motor maps. Recent evidence has revealed that the maps are dynamic, shaped by sensory experience in developing animals, and by eye and head position signals in adults. The superior colliculus thus provides an ideal model for studying the neural mechanisms underlying developmental and real-time modifications of information representation in the brain.

Animals↗
PubMed1991

Stretched and upside-down maps of auditory space in the optic tectum of blind-reared owls; acoustic basis and behavioral correlates.

Vision during early life plays an important role in calibrating sound localization behavior. This study investigates the effects of visual deprivation on sound localization and on the neural representation of auditory space. Nine barn owls were raised with eyelids sutured closed; one owl was congenitally anophthalmic. Data from these birds were compared with data from owls raised with normal visual experience. Sound localization behavior was significantly less precise in blind-reared owls than in normal owls. The scatter of localization errors was particularly large in elevation, though it was abnormally large in both dimensions. However, there was no systematic bias to the localization errors measured over a range of source locations. This indicates that the representation of auditory space is degraded in some way for blind-reared owls, but on average is properly calibrated. The spatial tuning of auditory neurons in the optic tectum was studied in seven of the blind-reared owls to assess the effects of early visual deprivation on the neural representation of auditory space. In normal owls, units in the optic tectum are sharply tuned for sound source location and are organized systematically according to the locations of their receptive fields to form a map of auditory space. In blind-reared owls, the following auditory properties were abnormal: (1) auditory tuning for source elevation was abnormally broad, (2) the progression of the azimuths and elevations of auditory receptive fields across the tectum was erratic, and (3) in five of the seven owls, the auditory representation of elevation was systematically stretched, and in the two others large portions of the representation of elevation were flipped upside down. The following unit properties were apparently unaffected by blind rearing: (1) the sharpness of tuning for sound source azimuth, (2) the orientation of the auditory representation of azimuth, and (3) the mutual alignment of the auditory and visual receptive fields in the region of the tectum representing the area of space directly in front of the animal. The data demonstrate that the brain is capable of generating an auditory map of space without vision, but that the normal precision and topography of the map depend on visual experience. The space map results from the tuning of tectal units for interaural intensity differences (IIDs) and interaural time differences (ITDs; Olsen et al., 1989).(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1990

Horizontal and vertical components of head movement are controlled by distinct neural circuits in the barn owl.

To generate behaviour, the brain must transform sensory information into signals that are appropriate to control movement. Sensory and motor coordinate frames are fundamentally different, however: sensory coordinates are based on the spatiotemporal patterns of activity arising from the various sense organs, whereas motor coordinates are based on the pulling directions of muscles or groups of muscles. Results from psychophysical experiments suggest that in the process of transforming sensory information into motor control signals, the brain encodes movements in abstract or extrinsic coordinate frames, that is ones not closely related to the geometry of the sensory apparatus or of the skeletomusculature. Here we show that an abstract code underlies movements of the head by the barn owl. Specifically, the data show that subsequent to the retinotopic code for space in the optic tectum yet before the motor neuron code for muscle tensions there exists a code for head movement in which upward, downward, leftward and rightward components of movement are controlled by four functionally distinct neural circuits. Such independent coding of orthogonal components of movement may be a common intermediate step in the transformation of sensation into behaviour.

Animals↗
PubMed1990

Neural maps of head movement vector and speed in the optic tectum of the barn owl.

1. This study investigates the contribution of the optic tectum in encoding the metric and kinetic properties of saccadic head movements. We describe the dependence of head movement components (size, direction, and speed) on parameters of focal electrical stimulation of the barn owl's optic tectum. The results demonstrate that both the site and the amount of activity can influence head saccade metrics and kinetics. 2. Electrical stimulation of the owl's optic tectum elicited rapid head movements that closely resembled natural head movements made in response to auditory and visual stimuli. The kinetics of these movements were similar to those of saccadic eye movements in primates. 3. The metrics and kinetics of head movements evoked from any given site depended strongly on stimulus parameters. Movement duration increased with stimulus duration, as did movement size. Both the size and the maximum speed of the movement increased to a plateau value with current strength and pulse rate. Movement direction was independent of stimulus parameters. 4. The initial position of the head influenced the size, direction, and speed of movements evoked from any given site: when the owl initially faced away from the direction of the induced saccade, the movement was larger and faster than when the owl initially faced toward the direction of the induced movement. 5. A characteristic movement of particular size, direction, and speed could be defined for each site by the use of stimulation parameters that elicited plateau movements with normal kinetic profiles and by having the head initially centered on the body. The size, direction, and speed of these characteristic movements varied systematically with the site of stimulation across the tectum. The map of head movement vector (size and direction) was aligned with the sensory representations of visual and auditory space, such that the movement elicited from a given site when the owl initially faced straight ahead brought the owl to face that region of space represented by the sensory responses of the neurons at the site of stimulation. 6. The results imply that both the site and the amount of neural activity in the optic tectum contribute to encoding the metrics and kinetics of saccadic movements. A comparison of the present findings with previous studies on saccadic eye movements in primates and combined eye and head movements in cats suggests striking similarities in the ways in which tectal activity specifies a redirection in gaze to such dissimilar motor effectors as the eyes and head.

Animals↗
PubMed1990

Sensitive and critical periods for visual calibration of sound localization by barn owls.

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)

Adaptation, Physiological↗
PubMed1989

Fused binocular vision is required for development of proper eye alignment in barn owls.

The eyes of adult barn owls (Tyto alba) are virtually fixed in the head in positions that are highly consistent from one individual to the next. However, early in development the eyes are exodeviated; the eyes achieve their adult positions during the owl's second month of life. Disruption of binocular vision in baby owls leads to permanent, highly abnormal eye positions and interocular alignment. Of three owls raised with both eyelids sutured closed, two developed exotropic strabismus and one developed esotropic strabismus. Two owls reared with monocular vision developed esotropic strabismus, whereas three owls reared with fused, but optically deviated binocular vision developed normal eye positions. Thus, the alignment of the eyes in adults results from an active process that depends on fused binocular vision during early life. Extracellular microelectrode recordings from the optic tecta of strabismic owls reveal that many units retain binocular inputs from corresponding points of the two eyes: the left-eye and right-eye receptive fields of individual units are misaligned by an amount predicted by the direction and magnitude of the strabismus. These results indicate that an innately determined pattern of connections in the brain anticipates the eye positions necessary to achieve binocular fusion. The hypothesis is put forth that the powerful activation of such binocular neurons by strong, synchronous inputs from the two eyes is the signal required by the optimotor system that proper eye alignment has been attained.

Animals↗
PubMed1989

Neural maps of interaural time and intensity differences in the optic tectum of the barn owl.

This report describes the binaural basis of the auditory space map in the optic tectum of the barn owl (Tyto alba). Single units were recorded extracellularly in ketamine-anesthetized birds. Unit tuning for interaural differences in timing and intensity of wideband noise was measured using digitally synthesized sound presented through earphones. Spatial receptive fields of the same units were measured with a free field sound source. Auditory units in the optic tectum are sharply tuned for both the azimuth and the elevation of a free field sound source. To determine the binaural cues that could be responsible for this spatial tuning, we measured in the ear canals the amplitude and phase spectra produced by a free field noise source and calculated from these measurements the interaural differences in time and intensity associated with each of 178 locations throughout the frontal hemisphere. For all frequencies, interaural time differences (ITDs) varied systematically and most strongly with source azimuth. The pattern of variation of interaural intensity differences (IIDs) depended on frequency. For low frequencies (below 4 kHz) IID varied primarily with source azimuth, whereas for high frequencies (above 5 kHz) IID varied primarily with source elevation. Tectal units were tuned for interaural differences in both time and intensity of dichotic stimuli. Changing either parameter away from the best value for the unit decreased the unit's response. The tuning of units to either parameter was sharp: the width of ITD tuning curves, measured at 50% of the maximum response with IID held constant (50% tuning width), ranged from 18 to 82 microsecs. The 50% tuning widths of IID tuning curves, measured with ITD held constant, ranged from 8 to 37 dB. For most units, tuning for ITD was largely independent of IID, and vice versa. A few units exhibited systematic shifts of the best ITD with changes in IID (or shifts of the best IID with changes in ITD); for these units, a change in the value of one parameter to favor one ear shifted the best value of the other parameter in favor of the same ear, i.e., in the direction opposite to that expected from "time-intensity trading." Overall sound intensity had little or no effect on ITD tuning, but did increase the best IIDs of units tuned to nonzero IIDs. The tuning of units for ITD and IID changed systematically along different dimensions of the optic tectum to create coextensive, independent neurophysiological maps of ITD and IID.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1989

Visuomotor adaptation to displacing prisms by adult and baby barn owls.

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.

Adaptation, Physiological↗
PubMed1989

Vision calibrates sound localization in developing barn owls.

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.

Animals↗
PubMed1988

Early blindness results in a degraded auditory map of space in the optic tectum of the barn owl.

The optic tectum of the barn owl (Tyto alba) contains a neural map of auditory space consisting of neurons that are sharply tuned for sound source location and organized precisely according to their spatial tuning. The importance of vision for the development of this auditory map was investigated by comparing space maps measured in normal owls with those measured in owls raised with both eyelids sutured closed. The results demonstrate that owls raised without sight, but with normal hearing, develop auditory space maps with degraded precision and with aspects of topography that are abnormal.

Animals↗
PubMed1987

Computational maps in the brain.

The nervous system performs computations to process information that is biologically important. Some of these computations occur in maps--arrays of neurons in which the tuning of neighboring neurons for a particular parameter value varies systematically. Computational maps transform the representation of information into a place-coded probability distribution that represents the computed values of parameters by sites of maximum relative activity. Numerous computational maps have been discovered, including visual maps of line orientation and direction of motion, auditory maps of amplitude spectrum and time interval, and motor maps of orienting movements. The construction of the auditory map of space is the most thoroughly understood: information about interaural delays and interaural intensity differences is processed in parallel by separate computational maps, and the outputs of these maps feed into a higher order processor that integrates sets of cues corresponding to sound source locations and creates a map of auditory space. Computational maps represent ranges of parameter values that are relevant to the animal, and may differentially magnify the representation of values that are of particular importance. The tuning of individual neurons for values of a mapped parameter is broad relative to the range of the map. Consequently, neurons throughout a large portion of a computational map are activated by any given stimulus, and precise information about the mapped parameter is coded by the locations of peak activity. There are a number of advantages of performing computations in maps. First, information is processed rapidly because the computations are preset and are executed in parallel. Second, maps simplify the schemes of connectivity required for processing and utilizing the information. Third, a common, mapped representation of the results of different kinds of computations allows the nervous system to employ a single strategy for reading the information. Finally, maps enable several classes of neuronal mechanisms to sharpen tuning in a manner not possible for information that is represented in a non-topographic code.

Animals↗