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Ingestive behaviors in mice are dependent on orosensory cues transmitted via the trigeminal nerve, as confirmed by transection studies. However, these studies cannot differentiate between deficits caused by the loss of the lemniscal pathway vs. the parallel paralemniscal pathway. The paired-like homeodomain protein Prrxl1 is expressed widely in the brain and spinal cord, including the trigeminal system. A knockout of Prrxl1 abolishes somatotopic barrellette patterning in the lemniscal brainstem nucleus, but not in the parallel paralemniscal nucleus. Null animals are significantly smaller than littermates by postnatal day 5, but reach developmental landmarks at appropriate times, and survive to adulthood on liquid diet. A careful analysis of infant and adult ingestive behavior reveals subtle impairments in suckling, increases in time spent feeding and the duration of feeding bouts, feeding during inappropriate times of the day, and difficulties in the mechanics of feeding. During liquid diet feeding, null mice display abnormal behaviors including extensive use of the paws to move food into the mouth, submerging the snout in the diet, changes in licking, and also have difficulty consuming solid chow pellets. We suggest that our Prrxl1(-/-) animal is a valuable model system for examining the genetic assembly and functional role of trigeminal lemniscal circuits in the normal control of eating in mammals and for understanding feeding abnormalities in humans resulting from the abnormal development of these circuits.

Rats generate stereotyped exploratory (5–12 Hz) vibrissa movements when navigating through their environment. Like other rhythmic behaviors, the production of whisking relies on a subcortical pattern generator. However, the relatively large vibrissae representation in motor cortex (vMCx) suggests that cortex also contributes to the control of whisker movements. The goal of this study was to examine the relationship between neuronal activity in vMCx and the kinematics of vibrissae movements. We recorded multiunit activity (MUA) and single units in the rhythmic region of vMCx while measuring vibrissa position in awake, head-restrained rats. The rats were engaged in one of two behavioral tasks where they were rewarded for either 1) producing noncontact whisking epochs that met specified criteria (epochs ≥4 Hz, whisks >5 mm) or 2) whisking to contact an object. There was significant coherence between the frequency of MUA and vibrissae movements during free-air whisking but not when animals were using their vibrissae to contact an object. Spike rate in vMCx was most frequently correlated with the amplitude of vibrissa movements; correlations with movement frequency did not exceed chance levels. These findings suggest that the specific parameter under cortical control may be the amplitude of whisker movements.

Objectives/Hypothesis: Rodent whisker movement has been used as a tool, after facial nerve manipulation, to quantify functional recovery. We have recently established a method to study functional correlates of aberrant regeneration of the facial nerve. Our objective was to establish normative parameters for both spontaneous and induced whisking and blinking behavior in a large group of normal rats. Study Design: Prospective animal study. Methods: Eighty animals underwent quantitative facial movement testing to measure simultaneous vibrissal movement and ocular closure for each side independently. Right and left C-1 whisker positions were continuously recorded for 5-minute sessions, and changes in infrared detection corresponding to eye closure were continuously recorded. Whisking and blinking were elicited by delivery of olfactory stimuli (10 s scented airflows) and corneal air puffs. Whisks were counted and analyzed, and eye closures were counted. Results: Whisking amplitude, velocity, and acceleration were consistent with literature values. Air puff delivery elicited an ipsilateral blink 99% of the time, a contralateral blink 18% of the time, and changes in or initiation of bilateral whisking 70% of the time. Olfactory stimulus delivery prompted a change in whisking behavior 83% of the time, and eye closure 20% of the time. Conclusions: This study establishes normative data for assessing cranial nerve VII-controlled facial movement in four separate facial regions. We demonstrate the capability and tendency of animals to move their orbicularis oculi muscles independently of and simultaneously with their midfacial muscles. This model provides an excellent tool for the study of aberrant regeneration after facial nerve injury in the rodent.

The occurrence of inappropriate co-contraction of facially innervated muscles in humans (synkinesis) is a common sequela of facial nerve injury and recovery. We have developed a system for studying facial nerve function and synkinesis in restrained rats using non-contact opto-electronic techniques that enable simultaneous bilateral monitoring of eyelid and whisker movements. Whisking is monitored in high spatio-temporal resolution using laser micrometers, and eyelid movements are detected using infrared diode and phototransistor pairs that respond to the increased reflection when the eyelids cover the cornea. To validate the system, eight rats were tested with multiple five-minute sessions that included corneal air puffs to elicit blink and scented air flows to elicit robust whisking. Four rats then received unilateral facial nerve section and were tested at weeks 3–6. Whisking and eye blink behavior occurred both spontaneously and under stimulus control, with no detectable difference from published whisking data. Proximal facial nerve section caused an immediate ipsilateral loss of whisking and eye blink response, but some ocular closures emerged due to retractor bulbi muscle function. The independence observed between whisker and eyelid control indicates that this system may provide a powerful tool for identifying abnormal co-activation of facial zones resulting from aberrant axonal regeneration.

Rats characteristically generate stereotyped exploratory (5–12 Hz) whisker movements, which can also be adaptively modulated. Here we tested the hypothesis that the vibrissal representation in motor cortex (vMCx) initiates and modulates whisking by acting on a subcortical whisking central pattern generator (CPG). We recorded local field potentials (LFPs) in vMCx of behaving Sprague-Dawley rats while monitoring whisking behavior through mystacial electromyograms (EMGs). Recordings were made during free exploration, under body restraint, or in a head-fixed animal. LFP activity increased significantly prior to the onset of a whisking epoch and ended prior to the epoch's termination. In addition, shifts in whisking kinematics within a whisk epoch were often reflected in changes in LFP activity. These data support the hypothesis that vMCx may initiate and modulate whisking behavior through its action on a subcortical CPG.

To examine the contribution of whisker inputs to the initial emergence and subsequent refinement of the rodent whisking pattern we combined surgical treatments producing varying degrees of postnatal whisker deafferentation with observations and video analysis of whisking across the first month of life. Whisking emerges during the second postnatal week, preceding eye opening by a few days. In contrast to the absence of deafferentation effects in adults, whisker deafferentation in pups, if carried out between the second and third postnatal week, delays (but does not prevent) the emergence of whisker movements and disrupts the development of normal whisking kinematics and coordination. The extent of the delay varies directly with the reduction in whisker input. When regeneration of the nerve is prevented by a cyanoacrylate block emergence of the normal pattern may be delayed indefinitely. Moreover, section of the whisker motor nerve contralateral to the deafferented side, substantially potentiates the effects of the initial deafferentation. These results confirm and extend an earlier description of the development of whisking in normal rat pups (Welker, Behaviour 12:223-244, 1964), fix the time of its initial emergence more precisely at P (postnatal day) 11-13, and suggest a critical role for trigeminal afference in the development of the normal whisking pattern. They are discussed in relation to the development of pattern generating mechanisms in the rodent whisker system.

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.

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.

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.

The following review and appreciation of the pioneering work and character of Wallace I. (Wally) Welker provides a historical perspective on Welker's life-long quest for answers to fundamental questions on the relationships among brain, behavior, and evolution, and evaluates his impact upon comparative behavioral neuroscience.

Previous studies have shown (1) that the form of the pigeon’s conditioned keypecking response resembles that of its ingestive pecking response, (2) that both ingestive and conditioned pecking in the pigeon are compound responses, including both transport (neck-movement) and gape (jaw-movement) components, and (3) that during operant conditioning or autoshaping of pecking behavior, the gape component comes under the associative control of the CS. In the present study, the gape component was experimentally isolated and a classical conditioning paradigm (water US) was used to bring jaw movements under the control of a CS (light). The results indicate that the topography of the jaw-movement CR is very similar to, though more variable than, that of the UR. They are consistent with the hypothesis that reported similarities in the form of ingestive and conditioned pecking responses reflect, in part, classical conditioning of the gape component.

Wally Welker died in Madison on 28 November 2007 after a long and innovative career at the University of Wisconsin–Madison. We were both influenced profoundly by his science and his approach to his...

In this paper we describe the effects of manipulating two kinds of sensory input in neonatal rats upon the development of the macrovibrissae--that movable subset of the rodent mystacial vibrissae. In an initial study of normal whisker development, data on whisker size were obtained from neonatal, perinatal, and adult rats. Data on whisker size were also obtained from rats sustaining either neonatal sensory or motor denervation of the whiskers and from both rats and mice bilaterally enucleated as neonates (BEN). In normally reared rats, most whiskers attain their final size over the first three postnatal weeks but development of rows 6 and 7 are not completed until after the first month. In normal animals we found a significant correlation both between body weight and whisker size and between the size of a whisker and the size of its corresponding cortical barrel. Rats sustaining neonatal denervation of the whiskers have shorter and thinner whiskers as adults than normally reared animals. In both rats and mice bilaterally enucleated as neonates a subset of the macrovibrissae are significantly larger than those of normal controls but no such effect is seen if the enucleation is carried out in adults. Moreover, BEN rats exposed to a novel stimulus environment whisk at a significantly higher frequency than normally reared animals. Mechanisms which might mediate these effects are discussed.

Studies of sensorimotor systems such as the whisking system of rodents have suggested that associations between body movements and their sensory consequences during development may make an important contribution to the functional organization of the system. In the present study we have explored the possible utility of Botulinum toxin for developmental studies of whisking. Botox selectively blocks whisking-generated afference leaving other sources of whisker afference intact. We describe appropriate modes of injection, define dosage levels, and assess the effects of prolonged whisking paralysis during development upon the basic motor competency of the adult rat. Our findings indicate that: (a) Botulinum toxin may be used to block whisking behavior in adult and developing rats, (b) that the duration of the whisking paralysis produced by Botox treatment blockade is dose dependent in both developing and adult animals, (c) that the blockade is functionally reversible and (d) that Botox treatment during development does not impair either the kinematics or the rhythmic patterning of adult whisking behavior. Botox may be a useful tool for studying the role of experiential factors in the development of "active touch" in rodents.

During 'active touch' the rodent whiskers scan the environment in a series of repetitive movements ('whisks') generating afferent signals which transform the spatial properties of objects into spatio-temporal patterns of neural activity. Based upon analyses carried out in a single movement plane, it has been generally assumed that these trajectories are essentially uni-dimensional, although more complex movements have been described in some rodents. The present study was designed to examine this assumption and to more precisely characterize whisking topography by monitoring whisking trajectories along both the antero-posterior and dorso-ventral axes. Using optoelectronic monitoring techniques with high-spatio-temporal resolution, movement data were obtained from a population of vibrissae sampled at different locations on the mystacial pad in head-fixed rats isolated from the perturbing effects of contact. For a substantial proportion of the population of whisking movements sampled, the trajectories generated by a single whisker is most accurately described as occupying an expended two-dimensional space in which the A-P component predominates. However, the whisker system exhibits a considerable range of trajectory types, suggesting a high degree of movement flexibility. For each vibrissa position, it was possible to delineate a 'trajectory' domain-that portion of the animal's whisking space which is scanned by the movements of that vibrissa during whisking. Since the 'domains' of adjacent whiskers in the same row tend to overlap, synchronized movements of a subset of whiskers could generate a set of overlapping somatosensory fields analogous to overlapping retinal receptive fields. The organization of such trajectory domains within the rats' whisking space could provide the spatial component of the spatio-temporal integration process required to extract information about environmental features from the inputs generated by its recursive whisking movements.

The effects of a barrel cortexo ablation upon the biometrics of a exploratoryo whisking were examined in three head-fixed rats which had previously sustained unilateral ablation of the left cortical a barrel fieldo 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 a barrel cortexo. In normal animals whisking movements have a characteristic rhythm (6± 9Hz), 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 a barrelo 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.

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.

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.

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.

Section of peripheral sensory branches of the pigeon's trigeminal nerve abolishes or decreases feeding responses made in an operant conditioning situation without directly impairing performance of an operant key-pecking response reinforced with food. The results are discussed in relation to previous studies of trigeminal function in the pigeon and suggest the possibility of dissociating neural mechanisms underlying feeding behavior and operant conditioning in this species.

Pigeons offered a choice among several different types of seed presented separately in substantial quantities, show individual patterns of seed selection behavior which persist over prolonged periods. The relations among several different measures of selection were examined under a variety of presentation conditions and following experimental manipulations (food deprivation or trigeminal deafferentation) designed, respectively, to increase and decrease hunger. Different measures of selection were differentially affected by the manipulations, suggesting that they reflect the operation of different causal mechanisms. The results indicate that the pigeon's selection behavior is controlled by many of the same processes that normally control its quantitative intake.

Control of beak opening (gape) and peck location was examined in pigeons. Feeding pecks showed accurate guidance that positioned the seed between the beaks. At the moment of contact with the seed, gape was proportional to seed diameter, although pecks with gape less than seed diameter were more frequent following an increase in seed size during a meal. There were no substantial differences between pigeons trained to keypeck with autoshaping and those trained with operant conditioning procedures. With either procedure, water reinforcement produced keypecks with the beak closed; seed reinforcers of different sizes produced means for gape proportional to the seed diameters. Black or white circular stimuli of different sizes projected as conditioning signals had little influence upon gape, but a greater percentage of responses was directed to white stimuli. These results indicate that visual stimuli elicit and orient the peck, whereas the adjustment of gape also involves the somatosensory stimuli provided during previous experience with a particular reinforcer or food type.

The role of oral sensations in the control of hunger and thirst in the pigeon was studied in birds in which the trigeminal sensory nerves had been sectioned bilaterally. Trigeminal deafferentation, although it does not impair drinking, disrupts both the neurosensory and the motivational control of feeding behavior. These two types of deficit may be experimentally dissociated.

The pigeon’s keypecking response includes both a head-transport (peck) and a jaw-movement (gape) component. Because the two components are mediated by different effector systems, they may potentially be viewed as orthogonal responses. A response differentiation procedure was used to bring gape amplitude under operant control. The procedure employed a conjunctive response requirement in which reinforcement was contingent upon both gaping and key contact. The key-contact requirement was held constant, while the gape contingency was systematically varied to reinforce either decreases or increases in gape amplitude with respect to baseline. The procedure was effective in shifting the gape distributions in both the upward and downward directions and in inducing new gape values that deviated from the baseline in the reinforced direction. These observations indicate that gape may be brought under operant control. However, subjects showed a bias in the differentiation of the gape response, such that larger gapes were more readily differentiated than smaller gapes. The results are discussed in relation to the methodological utility of the paradigm, the problem of biological constraints on learning, and the heuristic utility of a response components analysis.

In Experiment 1, the form of keypecks produced in an autoshaping procedure with food or water reinforcers was compared with that of eating and drinking responses. Because the responses involve a number of different effector systems, several elements of response form were measured, including peck force and duration, gape, and eye closure. Gape was the only measure to reliably distinguish between both ingestive responses and between conditioned keypecks reinforced with food or water. With either reinforcer, keypecks had greater force than did ingestive behaviors. In Experiment 2, a transition between two forms of keypeck was produced by manipulating deprivation and reinforcer conditions. Some measures appeared to vary in a dichotomous manner between two discrete response forms; gape showed a gradual and continuous change involving the production of intermediate forms of the response. It was concluded that the control of conditioned response form involves theconstruction of the response from movements produced by several effector systems, each with potentially different sources of control.

High-speed cinematography was used to analyze the eating behavior of the pigeon and to clarify the nature of its control by stimulus and motivational variables. Eating-response sequences may be functionally subdivided into a series of individual movement patterns whose duration and sequential organization exhibit a range of stereotypy comparable with that of movement patterns with an obvious communication function. Food deprivation does not affect the topography or spatiotemporal organization of the eating-response sequence but increases the rate at which such sequences are emitted. Stimuli coming from food elicit and orient many components of the eating response and serve as coordinating links in the causal chain that integrates individual movement patterns into an adaptive behavior for the efficient ingestion of food.

Electrolytic lesions were placed in the tractus septomesencephalicus (TSM) and tractus occipitomesencephalicus (TOM), efferent pathways originating in visual and somatosensorimotor areas of the avian telencephalon and distributing widely to brain stem and spinal nuclear regions. Lesion effects upon several aspects of ingestive behavior were examined using high speed cinematography, operant conditioning procedures and monitoring of intake and feeding responses. While there was no evidence for direct effects upon drinking, both TSM and TOM lesions were followed by periods of reduced food intake. Disruptions of feeding in TSM birds were mild and relatively transient but birds were hypophagic for prolonged periods. While there were no deficits in the efficiency or accuracy of their feeding responses or their performance in an operant situation, TSM birds displayed an inappropriate seed ''sorting'' behavior suggestive of lesion effects upon visually controlled food preferences. TOM lesions produced a significant reduction in responsiveness to food (aphagia, hypophagia), impairments in the control of grasping and peck accuracy and a disruption in operant key pecking reinforced by food. These ''sensorimotor'' and ''motivational'' deficits were similar to those seen after damage to central trigeminal structures in the pigeon and suggest that TOM is also a component of a putative ''feeding system'' in the pigeon.

An operant response in the pigeon, whose performance results in exposure to the discriminative stimuli, is described and suggested as an experimental analogue for “observing.” Such an operant response is then used to explore the relationship between observing responses and discrimination learning in a variety of discrimination situations, of progressively increasing complexity. In general, the results support the contention that the development and maintainence of observing responses is closely related to the degree of differential behaviour manifested toward the discriminative stimuli. Certain modifications are suggested in the theoretical formulation underlying the concept of “observing responses.”

An analysis of the deficits produced by lesions of quinto-frontal structures (PrV, QFT, NB) in the pigeon was carried out, using a variety of measures of hunger and thirst. The results may be summarized as follows: In pigeons maintained under ad-libitum conditions lesions of PrV, QFT and NB are followed by a period in which responsiveness to food is abolished or significantly reduced. Lesions of PrV and NB impair the performance of the consummatory response of eating by disrupting the neuro-sensory control of mandibulation. Lesions of PrV and NB abolish an operant key-pecking response reinforced with food for periods ranging from a few days to several weeks. No effects of quinto-frontal lesions upon either operant or consummatory measures of thirst were observed. We conclude that quinto-frontal structures mediate both sensorimotor and motivational processes involved in the control of feeding behavior in the pigeon and that these two functions may be experimentally dissociated. Our results are discussed in relation to previous studies of the sensory and neural control of hunger in birds and mammals.

Anterograde and retrograde tracing experiments employing WGA-HRP were used to identify an auditory projection area within the frontal telencephalon of the pigeon. The projection originates in a nucleus of the lateral lemniscus, travels with the quintofrontal tract and terminates within nucleus basalis. The location of the projection area and the absence of a thalamic relay in its pathway are consistent with previous reports of short-latency auditory potentials evoked in the vicinity of the nucleus basalis in several avian species.

Trigeminal orosensory deafferentation produces a reduction in intake whose magnitude is a joint function of the extent of the deafferentation and the sensory properties of the diet. Intake deficits are attributed to depressed oromotor reflexes, disrupted ingestive response sequences, decreased responsiveness to food and water, and a reduced level of long-term body weight regulation.

A set of software utility programs is described which, in association with a microcomputer and a commercially available analog-to-digital converter may be used to acquire, store, manipulate and display large amounts of behavioral and/or physiological data in digitized form.

Section of trigeminal orosensory or oromotor nerves reduces food intake and disrupts performance of a food-reinforced lever pressing response but increases the rate of an identical lever pressing response reinforced by electrical stimulation of the lateral hypothalamus. Our findings suggest that trigeminal orosensation plays a critical role in the neural control of hunger in the rat. These results also fail to support a response-oriented theory of intracranial self-stimulation where the latter is thought to involve selective facilitation of sensorimotor circuits underlying species-typical consummatory behavior.

For the past thirty years neurobehavioral research on ingestive behavior has been characterized by a decline in the study of peripheral sensorimotor mechanisms and an increasing concentration upon the role of central neural structures in general and of the hypothalamus in particular. The consequence of this “hypothalamocentric” strategy has been a subtle shift in research goals and tactics. What had begun, at first, as a search for the neural basis of hunger and thirst gradually evolved into a search for the causal basis of the lateral hypothalamic (LH) syndrome (Anand and Brobeck, 1951; Epstein, 1971; Stricker and Zigmond, 1976; Teitelbaum and Epstein, 1962). Although some aspects of this research have paid handsome empirical and conceptual dividends (see, e.g., Teitelbaum, 1977), our continued preoccupation with this one behavioral syndrome and this one brain region may actually have impeded the neurobehavioral analysis of ingestive behavior mechanisms.

Electrophysiological, neurobehavioral, and cinematographic techniques were used to study the role of the trigeminal nerve in the neurosensory control of eating. Analaysis of single-unit data recorded in the pigeon's trigeminal ganglion indicates that the trigeminal nerve provides somatosensory inputs from the oral region that signal the location and movement of food and monitor the extent of mouth opening. Trigeminal deafferentation, although it does not affect pecking or swallowing, severly impairs the efficiency of eating. Cinematographic analysis shows that the impairment is due to disruption of the neurosensory control of mandibulation-the process by which grain is grasped at the beak tip and moved to the back of the mouth.

With the aid of autoradiographic and histochemical (WGA-HRP) tracing techniques, the projections of the nucleus of the nucleus of the tractus solitarius (nTS) in the pigeon have been delineated and related to the viscerotopic organization of the nucleus. As in mammals, nTS projects to both brainstem and forebrain structures. At medullary levels, projections were seen to nTS itself, to the dorsal motor nucleus of the vagus and to the subjacent and more ventral reticular formation. There is a substantial projection to the parabrachial nuclear complex with terminations in all its subnuclei and minor projections to locus coeruleus and several mesencephalic areas, including the ventral area of Tsai, the nucleus of the ascending brachium conjunctivum, and the compact portion of the tegmental pedunculopontine nucleus. At diencephalic levels, projections to the hypothalamus (magnocellular periventricular nucleus, stratum cellulare internum and externum) and dorsal thalamus were seen. Terminal fields within the basal telencephalon included the nucleus of the pallial commissure, the bed nucleus of the stria terminalis, and the nucleus accumbens. The organization of nTS projections in pigeons is correlated with the pattern of inputs to specific nTS subnuclei. Lateral tier subnuclei receiving cardiovascular and pulmonary inputs project upon the ventrolateral reticular formation and the ventrolateral parabrachial complex. Medial tier subnuclei receiving gustatory and gastrointestinal inputs project upon dorsal and medial parabrachial nuclei. Transparabrachial projections arise from nTS subnuclei receiving little or no primary input from the viscera.