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H P Zeigler

Publications and source records attributed to H P Zeigler.

At least 19 recordsLinked to original sources

Topography of whisking II: interaction of whisker and pad.

The peripheral effector system mediating rodent whisking produces protraction/retraction movements of the whiskers and translation movements of the collagenous mystacial pad. To examine the interaction of these movements during whisking in air we used high-resolution, optoelectronic methods for two-dimensional monitoring of whisker and pad movements in head-fixed rats. Under these testing conditions (1) whisker movements on the same side of the face are synchronous and of similar amplitude; (2) pad movements exhibit the characteristic 'exploratory' rhythm (6-12 Hz) of whisking but their movements often have a low frequency (1-2 Hz) component; (3) Pad movements occur in both antero-posterior and dorso-ventral planes but there are considerable variations in the amplitude and topography of movement parameters in the two planes. We conclude that (a) both whisker and pad receive input from a common central rhythm generator; (b) differences in whisker and pad amplitude and topography probably reflect differences in the biomechanical properties of the structures receiving that input; (c) pad movements make a significant contribution to the kinematics of whisking behavior and (d) the two-dimensional nature of pad translation movements significantly increases the rat's flexible control of its mobile sensor.

Animals↗

Whisking as a "voluntary" response: operant control of whisking parameters and effects of whisker denervation.

The rat's ability to vary its whisking "strategies" to meet the functional demands of a discriminative task suggests that whisking may be characterized as a "voluntary" behavior--an operant--and like other operants, should be modifiable by appropriate manipulations of response-reinforcer contingencies. To test this hypothesis we have used high-resolution, optoelectronic "real-time" recording procedures to monitor the movements of individual whiskers and reinforce specific movement parameters (amplitude, frequency). In one operant paradigm (N = 9) whisks with protractions above a specified amplitude were reinforced (Variable Interval 30 s) in the presence of a tone, but extinguished (EXT) in its absence. In a second paradigm (N = 3), rats were reinforced on two different VI schedules (VI-20s/VI-120s) signaled, respectively, by the presence or absence of the tone. Selective reinforcement of whisking movements maintained the behavior over many weeks of testing and brought it under stimulus and schedule control. Subjects in the first paradigm learned to increase responding in the presence of the tone and inhibit responding in its absence. In the second paradigm, subjects whisked at significantly different rates in the two stimulus conditions. Bilateral deafferentation of the whisker pad did not impair conditioned whisking or disrupt discrimination behavior. Our results confirm the hypothesis that rodent whisking has many of the properties of an operant response. The ability to bring whisking movement parameters under operant control should facilitate electrophysiological and lesion/behavioral studies of this widely used "model" sensorimotor system.

Animals↗

Whisker deafferentation and rodent whisking patterns: behavioral evidence for a central pattern generator.

Even in the absence of explicit stimulation, rats emit patterns of rhythmic whisking movements. Because of their stereotyped nature and their persistence after sensory denervation and cortical ablation, whisking movements have been assumed to reflect the output of a central pattern generator (CPG). However, identification of a movement pattern as the product of a CPG requires evidence that its generation, patterning, and coordination are independent of sensory input. To provide such evidence, we used optoelectronic instrumentation to obtain high-resolution records of the movement trajectories of individual whiskers in rats whose heads were fixed to isolate their exploratory whisking from exafferent inputs. Unconditioned whisking patterns were quantitatively characterized by a biometric analysis of the kinematics, rhythmicity, and coordination of bilaterally homologous vibrissa movements. Unilateral and bilateral sectioning of the infraorbital nerve, which innervates the whiskers, was then performed to block reafferent inputs generated by the animal's own whisking movements. Unilateral sectioning of the nerve has no effect on whisking kinematics but is followed by a significant but relatively transient bilateral increase in whisking frequency. However, bilateral deafferentation, when performed in a single-stage procedure, does not disrupt the generation, patterning, or bilateral coordination of whisking patterns in the rat. These findings provide strong behavioral evidence for a whisking CPG and are discussed in relation to its possible location and properties.

Afferent Pathways↗

Discriminative whisking in the head-fixed rat: optoelectronic monitoring during tactile detection and discrimination tasks.

We compared whisking movement patterns during acquisition of tactile detection and object discrimination under conditions in which (a) head movements are excluded and (b) exposure to tactile discriminanda is confined to the large, moveable vibrissae (macrovibrissae). We used optoelectronic instrumentation to track the movements of an individual whisker with high spatio-temporal resolution and a testing paradigm, which allowed us to dissociate performance on an "indicator" response (lever pressing) from the rat's "observing" responses (discriminative whisking). We analyzed the relation between discrimination performance and whisking movement patterns in order to clarify the process by which the indicator response comes under the stimulus control of information acquired by the rat's whisking behavior. Whisking patterns over the course of task acquisition differed with task demands. Acquisition of the Detection task was correlated with modulation of only one whisking movement parameter-total number of whisks emitted, and more whisking was seen on trials in which the discriminandum was absent. Discrimination between a sphere and cube differing in size and texture was correlated with a reduction in whisk duration and protraction amplitude and with a shift towards higher whisking frequencies. Our findings confirm previous reports that acquisition of tactile discriminations involves modulation by the animal of both the amount and the type of whisking. In contrast with a previous report (Brecht et al., 1997), they indicate that rats can solve tactile object detection and discrimination tasks (a) using only the large, motile mystacial vibrissae (macrovibrissae) and (b) without engaging in head movements. We conclude that the functional contribution of the macrovibrissae will vary with the nature of the task and the conditions of testing.

Animals↗

Cortical barrel field ablation and unconditioned whisking kinematics.

The effects of "barrel cortex" ablation upon the biometrics of "exploratory" whisking were examined in three head-fixed rats which had previously sustained unilateral ablation of the left cortical "barrel field" under electrophysiological control. Unconditioned movements of a pair of bilaterally homologous whiskers (C-1, Right, Left) were monitored, optoelectronically, with other whiskers present. Whisking movements on the intact and ablated side were analyzed with respect to kinematics (protraction amplitude and velocity) whisking frequency and phase relationships between whisking movement on the two sides of the face. Histological analysis confirmed complete removal of S-1 "barrel cortex". In normal animals whisking movements have a characteristic rhythm (6-9 Hz), and protractions on the two sides of the face tend to be both synchronous and of very similar amplitudes. In the lesioned animals, whisking frequency was unchanged and whisking movements remained bilaterally synchronous. However, there was a significant difference between the amplitude of Right and Left whisker movements which was evident many months postoperatively. Our results suggest that the deficits in vibrissa-mediated tactile discrimination reported after "barrel" field ablation may reflect an impairment in the animal's ability to modulate whisking parameters on the two sides of the face to meet the functional requirements of a discriminative whisking task. The effects upon whisking amplitude seen after unilateral barrel field ablation are consistent with a model in which the activity of a whisking Central Pattern Generator is modulated by descending inputs to achieve sensorimotor control of whisking movement parameters.

Animals↗

"Real-time" monitoring of vibrissa contacts during rodent whisking.

Rodent whisking behavior provides active touch as input into a widely studied model system of information processing and behavior. We previously developed a simple optoelectronic system to monitor whisker movements in "real time" in head held rats at rest or performing various tasks such as tactile discrimination. We now describe a simple piezioelectic film device for detecting initial whisker contacts during whisking also in real time. In some applications this is as effective as high-speed videos and can be configured to isolate the contacts from different whiskers. The construction of this simple device is detailed. In addition to providing information during recordings from awake animals, the device could be used, for example, as an operant "manipulandum" for contingent reinforcement of object detection with a whisker.

Animals↗

Trigeminal deafferentation and conditioned pecking in pigeons.

To clarify the contribution of peripheral trigeminal input to the control of pecking behavior we examined head and jaw movement kinematics and peck localization in pigeons with surgical section of trigeminal nerves providing somatosensory input to the beak. Conditioning procedures were used to bring the pecking/grasping components of pecking under the control of a visual target. Conditioned head and jaw movements were monitored 'on-line' using movement transducers and terminal peck location was recorded using 'touch-screen' technology. The periodic delivery of a food reinforcer provided repeated opportunities to monitor the kinematics of ingestive pecks. Deafferentation produced deficits in mandibulation during ingestive pecking and in the coordination of head and jaw movements during conditioned pecking. These results are attributed to disruptions in trigeminal feedback and feedforward mechanisms, respectively. In contrast with previous studies, deafferentation did not impair the precision of peck localization. Possible reasons for the absence of localization deficits are presented. The results are discussed in relation to the role of peripheral inputs in the control of prehensile movements.

Animals↗

A "pre-satiety sequence" in rats drinking sucrose solutions.

Food-deprived rats were given intermittent access or, in a replication, continuous access to a 20% sucrose solution. Both drinking and nondrinking behaviors were recorded. During the ensuing drinking bout, latency of lapping after snout apposition, and duration of lapping bouts, did not change. Approaches to the sipper tube usually eventuated in lapping, though aborted approaches increased in frequency late in the session. Drinking was interrupted by nondrinking behaviors which appeared in a characteristic rostro-caudal sequence: partial head withdrawal from the drinking aperture, then full head withdrawal, then movements of the front paws, then movement of the hind paws with full-body locomotion. All these behavioral changes occurred before there was any appreciable reduction in rate of lapping.

Animals↗

Optoelectronic monitoring of individual whisker movements in rats.

We describe two systems for the real-time recording and display of individual vibrissa movements in head-fixed rats. Both systems utilize high-speed, linear image sensors, each composed of an array of light sensitive elements (CCDs). Uniform illumination of the array generates a constant baseline voltage in each element. The shadow produced by the movement of a whisker interposed between the light source and the sensors produces a voltage shift in a subset of elements. The successive position of the shift is linearly related to the momentary whisker position. Associated software/hardware scans the array at regular intervals to identify the successive positions of voltages above a preset threshold and outputs the data to a microprocessor for computation of the whisker movement trajectory. In both systems, movements of a single whisker may be monitored 'on-line' with high spatial and temporal resolution; in one case with, in the other without the presence of neighboring whiskers. Optoelectronic monitoring facilitates rapid and efficient (computer-assisted) acquisition and analysis of data on rodent whisking behavior.

Animals↗

Organization of quinto-frontal structures in hatchling ring doves (Streptopelia risoria).

Transganglionic and retrograde tracing procedures were applied to peripheral and central trigeminal structures in hatchling ring doves. The organization of the trigeminal ganglion, its somatotopic projections upon the principal sensory nucleus (PrV), and the projections of PrV upon the telencephalon are similar in adult and hatchling Columbiformes. The results suggest that development of feeding patterns in these species involves experiential differentiation of trigeminal sensorimotor circuits present at hatching.

Animals↗

Conditioned 'prehension' in the pigeon: kinematics, coordination and stimulus control of the pecking response.

Like human prehensile behavior, the pigeon's ingestive pecking response is elicited by visual stimuli conveying information about the location and size of the target. This information is used to generate localized ingestive pecks whose gapes are amplitude-scaled to seed size, prior to contact. We employed high-resolution, 'real-time' monitoring of head acceleration, jaw movements and terminal peck location to examine the kinematics, coordination and stimulus control of conditioned pecking. Conditioning procedures were used to bring pecking under the control of visual targets whose stimulus properties (size, location) were independently varied, while simultaneously monitoring pecking response parameters. Stimulus control of the transport component (peck localization) is extremely precise, even in the absence of a specific localization-dependent reinforcement contingency. Subjects also showed amplitude-scaling of gape size to the size of a visual target, but over a more restricted range than shown to food pellets of comparable sizes. Comparison of the kinematic profiles of conditioned and ingestive pecks suggests that conditioned pecking is functionally analogous to human 'pointing' rather than 'grasping' behavior.

Animals↗

Behavioral morphology of the pigeon's peck: ingestion, prehension and cognition.

In nature, pecking involves the transport of a prehensile effector organ (the beak) towards biologically significant targets, and the generation jaw movement patterns whose probability and topography are appropriate to the stimulus properties of the target (e.g., distance, direction, size) and the motivational state of the animals (hunger, thirst). Because conditioned pecking preserves essential topographic features of the ingestive peck, we have used the pigeon's conditioned pecking response as a "model system" for the study of motor control, response form modulation and cognitive processes. In the first study, conditioning procedures were used to bring pecking under the control of visual targets whose stimulus properties (size, location) were independently varied, while simultaneously monitoring head acceleration, jaw movements and terminal peck location. Comparison of the kinematic profiles of conditioned and ingestive pecks suggests that conditioned pecking is functionally analogous to human "pointing" rather than "grasping" behavior. In the second study, we manipulated reinforcer type and size to generate populations of "virtual" eating and drinking responses and used them to study the process of response form modulation. The data suggest that different response forms do not reflect "fixed" movement patterns but the modulation of movement patterns across a topographic continuum. In the third study we manipulated stimulus, response and deprivational variables to test a cognitive model for the study of "expectancy", "intention", and "choice" in pigeon.

Animals↗

Key-peck probability and topography in a concurrent variable-interval variable-interval schedule with food and water reinforcers.

The relation between variables that modulate the probability and the topography of key pecks was examined using a concurrent variable-interval variable-interval schedule with food and water reinforcers. Measures of response probability (response rates, time allocation) and topography (peck duration, gape amplitude) were obtained in 5 water- and food-deprived pigeons. Key color signaled reinforcer type. During baseline, response rates and time allocations were greater to the food key than to the water key, and food-key pecks had larger gapes and shorter durations. Relative probability measures (for the food key) were increased by prewatering and decreased by prefeeding. Deprivation effects upon topography measures were apparent only when food- and water-key pecks were analyzed separately. Food-key gape amplitudes increased with prewatering and decreased with prefeeding. The clearest effect occurred with prewatering. There were no consistent effects upon water-key gapes. The key color-reinforcer relation was reversed for 3 pigeons to determine how response topography was modulated during the transition from food- to water-key pecks. Reacquisition was faster for the probability than for the topography measures. Analysis of gape-amplitude distributions during reversal indicated that response-form modulation proceeded through the generation of intermediate gape sizes.

Animals↗

Central projections and somatotopic organisation of trigeminal primary afferents in pigeon (Columba livia).

Injections of cholera toxin B-chain conjugated to horseradish peroxidase into individual peripheral branches of the trigeminal nerve or into the trigeminal ganglion showed that an ascending trigeminal tract (TTA) terminated in distinct ventral and dorsal divisions of the principal sensory nucleus (PrVv and PrVd, respectively), and a descending tract (TTD) terminated within pars oralis, pars interpolaris, and pars caudalis divisions of the nucleus of TTD (nTTD) and within the dorsal horn of the first six cervical spinal segments. In PrVd, mandibular, ophthalmic, and maxillary projections were predominantly located dorsally, ventrally, and medially, respectively. In nTTD, mandibular projections lay dorsomedially, ophthalmic projections lay ventrolaterally, and maxillary projections lay in between. At caudal medullary and spinal levels, mandibular projections were situated medially, ophthalmic projections were situated laterally, and maxillary projections were situated centrally. The terminations within the dorsal horn were most dense in laminae III and IV and were least dense in lamina II, with laminae III-IV also receiving topographically organised contralateral projections. Extratrigeminal projections were mainly to the external cuneate nucleus by way of a lateral descending trigeminal tract (lTTD; Dubbeldam and Karten [1978] J. Comp. Neurol. 180:661-678) and to the region of the tract of Lissauer and lamina I of the dorsal horn. Other projections were to a region medial to the apex of pars interpolaris, to the nuclei ventrolateralis anterior (Vla) and presulcalis anterior (Pas) of the solitary complex, and sparsely to the lateral reticular formation (plexus of Horsley) ventral to TTD. No projections were seen to the trigeminal motor nuclei or to the cerebellum.

Afferent Pathways↗

Effects of food-pellet size on rate, latency, and topography of autoshaped key pecks and gapes in pigeons.

Four pigeons responded under autoshaping contingencies in which different conditional stimuli (red or green keylights) were associated with unconditional stimuli of different magnitudes (large or small food pellets) over successive trials within a session. Both topography (beak opening or gape) and strength (rates and latencies of key pecks and gapes) of responding during the conditional stimuli depended on the magnitude of the correlated unconditional stimulus. Key-peck and gape rates were higher and latencies were shorter in large-pellet trials than in small-pellet trials. Gape amplitudes varied directly with pellet size, although conditional and unconditional gapes were larger than either pellet. These findings were replicated when the key colors were presented either on one or two keys and after reversals of the color-size correlations. Because the unconditional stimulus was varied through pellet size, magnitude was not confounded with food-access duration or quality. These results demonstrate the effects of the magnitude of the unconditional stimulus, in that rates and latencies of both key pecks (which are directed movements toward the key) and gapes (which are independent of the bird's position and key properties) varied with pellet size. Gape measures were unique in that two dimensions (response strength and topography) of a single response class varied simultaneously with magnitude.

Animals↗

Conditioned whisking in the rat.

The rat's mystacial vibrissae are active during exploratory and discriminative behaviors, with individual vibrissae serving as elements in a receptive array scanned across object surfaces. To facilitate neurobehavioral analysis of this sensorimotor system, we have developed an experimental paradigm that confines vibrissa movements to a defined physical location, makes possible on-line monitoring of "whisking" activity, and brings such activity under associative control using operant conditioning procedures. Rats were secured, and movements of an identified bilaterally homologous pair of vibrissae (right and left gamma straddlers) were detected by laser-based photodetectors. Subjects were maintained on a water deprivation schedule, and whisker movements were monitored during adaptation to the test situation and after the clipping of other vibrissae on both sides of the snout. Rats were reinforced with water delivery for emitting vibrissa movements in the presence of a conditioned stimulus (tone) whose presentation was made contingent upon a prior period of nonwhisking. The rate and temporal distribution of vibrissa movements were brought under experimental control by means of interval and ratio reinforcement schedules. Although the procedures provide minimal information about the kinematics or topography of conditioned vibrissa movements, they permit the investigator to manipulate response parameters normally under the voluntary control of the animal in a preparation amenable to neurophysiological analysis.

Animals↗

Water deprivation and subfornical organ activity in the pigeon a [14C]2-deoxyglucose study.

Following varying degrees of water deprivation (0, 24 and 72 h), functional activity in the subfornical organ (SFO) of pigeons was measured using the [14C]2-deoxyglucose method. Increasing levels of water deprivation produced a significant increase in glucose uptake in SFO. The magnitude of the effect was systematically correlated with morphologically defined SFO subdivisions.

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Jaw muscle (EMG) activity and amplitude scaling of jaw movements during eating in pigeon (Columba livia)

During each phase of the pigeon's eating sequence, jaw opening amplitude (gape) is adjusted to the size of the food object; first prior to contact (Grasping), again in positioning the food (Stationing), and finally, during its movement through the oral cavity (Intraoral Transport). Part I of this study examined jaw movement kinematics during ingestion of different size food pellets to determine the relative contribution of velocity and rise time variables. Part II specified the muscle activity patterns mediating each phase of the eating sequence, and determined how these patterns are modulated to produce adjustments of gape size. The relative contribution of velocity and rise time variables to the control of gape differs in each phase of the eating sequence. However, for any pellet size, variations in opening rise time may function in a compensatory manner to minimize gape "undershooting". Each phase of the eating sequence is mediated by a characteristic muscle activity pattern. The adjustment of gape size to pellet size involves systematic modulation of this pattern, and the parameters modulated differ in the different phases in a manner which may reflect the functional requirements of each phase.

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