Odor response characteristics of thalamic mediodorsal nucleus neurons in the rabbit

Extracellular responses were recorded from neurons in the thalamic mediodorsal nucleus (MD) of the lightly anesthetized rabbit. Eighty-seven neurons responded to electrical stimulation of the lateral olfactory tract (LOT shocks). They were located in the medial portion of the MD. In the same portion, negative field potentials with a short latency were evoked by the electrical stimulation of the olfactory projection area in the neocortex (OPA shocks). Fifty-nine MD neurons responded both to LOT and to OPA shocks. Among them, 17 thalamocortical relay neurons (which responded antidromically to OPA shocks) were found to respond transsynaptically to LOT shocks. Of the 87 LOT-responsive MD neurons, 48 responded to the odors applied. Eight odor-sensitive neurons were found to be the thalamocortical relay neurons. Thus, it was proven for the first time that a portion of the olfactory input to the OPA is mediated via relay neurons in the MD. Characteristics of response of MD neurons to odor stimulation were compared with those of OPA neurons. MD neurons did not show a selectivity of response to odors of urine, feces, or dry food pellets, to which OPA neurons responded exclusively. These results were discussed in relation to the functional role of the MD-OPA projection system in the discrimination of specific odors.     

Feeding related lateral hypothalamic neuron responses to odors depend on food deprivation in rats

It has been investigated feeding related LHA neuronal activity and responses to odor stimulation in rats at various levels of satiation. Extracellular responses of 168 neurons to three odors, isoamylacetate (AA), cineole (CL), and isovaleric acid (VA), were recorded from 168 LHA neurons of Wistar-SPF male rats. Of 168 units, 107 (63.7%) responded to from one to three odors, but not to light or phonic stimulation. Of the responding units, 94.4% (101/107) were excited, and 5.6% were inhibited. In response to a single electrical stimulation (0.5 msec, 1-10 V) of the OB, 61 units were excited with latencies of 6-43 msec (19.8 +/- 12.0 msec, mean +/- S.D.) indicating compound OB-LHA relations--mono- and polysynaptic through myelinated and nonmyelinated fibers. The results suggest predominantly excitatory effects of both electrical stimulation of the OB and odor stimulation on the LHA. Firing frequency in response to AA or VA was significantly (p less than 0.05) greater for the long fasting group (38 hr, LF, n = 8) than for the NF (nonfasting, n = 12) group; differences between the LF and MF (24 hr, n = 6) groups were not significant. Glucose-sensitive neurons (GSN, n = 19) responded more to odors than non-GSNs (n = 86), and discharge frequency increase depended markedly on food deprivation. Food deprivation results suggest that responsiveness of feeding related LHA neurons to odors depends on the degree of satiation. In conclusion, it was confirmed that olfactory functions are important in the responses of hypothalamic feeding related neurons.     

Olfactory input to the lateral hypothalamus of the old world monkey

Responses of lateral hypothalamic neurons to 8 odors were studied in chronic unanesthetized old world monkeys (Macaca irus). Many neurons (54.5%) responded to a single odor only, and the number of neurons responding to 2, 3 and 4 odors decreased successively. No neuron responded to as many as 5 odors. Thus, the presence of olfactory input and a highly discriminative ability for odors were found in the lateral hypothalamic area (LHA). Neuronal responses to the same odors were also studied in the septum (Spt). In anesthetized old world monkeys, evoked potentials were recorded in the LHA and in areas of the Spt and the nucleus accumbens (Acc) during stimulation of the olfactory bulb (OB). When the Spt (and probably the Acc with it) was subsequently destroyed, OB-evoked potentials in the LHA disappeared. Next, by injecting horseradish peroxidase (HRP) into the LHA, an olfactory pathway to the LHA was examined. Labeled neurons were found mainly in the Spt and the Acc, and only partly in other areas. However, labeled neurons were scarcely found in the prepyriform (PPF)-entorhinal (ER) area or in the olfactory tubercle (OT). The present study thus shows that an olfactory pathway to the LHA passes through the Spt and probably also the Acc, but not through the PPF-ER areas nor through the OT in the old world monkey.     

Orbital cortex neuronal responses during an odor-based conditioned associative task in rats

Neuronal activity in the rat orbital cortex during discrimination of various odors [five volatile organic compounds (acetophenone, isoamyl acetate, cyclohexanone, p-cymene and 1,8-cineole), and food- and cosmetic-related odorants (black pepper, cheese, rose and perfume)] and other conditioned sensory stimuli (tones, light and air puff) was recorded and compared with behavioral responses to the same odors (black pepper, cheese, rose and perfume). In a neurophysiological study, the rats were trained to lick a spout that protruded close to its mouth to obtain sucrose or intracranial self-stimulation reward after presentation of conditioned stimuli. Of 150 orbital cortex neurons recorded during the task, 65 responded to one or more types of sensory stimuli. Of these, 73.8% (48/65) responded during presentation of an odor. Although the mean breadth of responsiveness (entropy) of the olfactory neurons based on the responses to five volatile organic compounds and air (control) was rather high (0.795), these stimuli were well discriminated in an odor space resulting from multidimensional scaling using Pearson's correlation coefficients between the stimuli. In a behavioral study, a rat was housed in an equilateral octagonal cage, with free access to food and choice among eight levers, four of which elicited only water (no odor, controls), and four of which elicited both water and one of four odors (black pepper, cheese, rose or perfume). Lever presses for each odor and control were counted. Distributions of these five stimuli (four odors and air) in an odor space derived from the multidimensional scaling using Pearson's correlation coefficients based on behavioral responses were very similar to those based on neuronal responses to the same five stimuli. Furthermore, Pearson's correlation coefficients between the same five stimuli based on the neuronal responses and those based on behavioral responses were significantly correlated. The results demonstrated a pivotal role of the rat orbital cortex in olfactory sensory processing and suggest that the orbital cortex is important in the manifestation of various motivated behaviors of the animals, including odor-guided motivational behaviors (odor preference).     

Orbital cortex neuronal responses during an odor-based conditioned associative task in rats

Neuronal activity in the rat orbital cortex during discrimination of various odors [five volatile organic compounds (acetophenone, isoamyl acetate, cyclohexanone, p-cymene and 1,8-cineole), and food- and cosmetic-related odorants (black pepper, cheese, rose and perfume)] and other conditioned sensory stimuli (tones, light and air puff) was recorded and compared with behavioral responses to the same odors (black pepper, cheese, rose and perfume). In a neurophysiological study, the rats were trained to lick a spout that protruded close to its mouth to obtain sucrose or intracranial self-stimulation reward after presentation of conditioned stimuli. Of 150 orbital cortex neurons recorded during the task, 65 responded to one or more types of sensory stimuli. Of these, 73.8% (48/65) responded during presentation of an odor. Although the mean breadth of responsiveness (entropy) of the olfactory neurons based on the responses to five volatile organic compounds and air (control) was rather high (0.795), these stimuli were well discriminated in an odor space resulting from multidimensional scaling using Pearson's correlation coefficients between the stimuli. In a behavioral study, a rat was housed in an equilateral octagonal cage, with free access to food and choice among eight levers, four of which elicited only water (no odor, controls), and four of which elicited both water and one of four odors (black pepper, cheese, rose or perfume). Lever presses for each odor and control were counted. Distributions of these five stimuli (four odors and air) in an odor space derived from the multidimensional scaling using Pearson's correlation coefficients based on behavioral responses were very similar to those based on neuronal responses to the same five stimuli. Furthermore, Pearson's correlation coefficients between the same five stimuli based on the neuronal responses and those based on behavioral responses were significantly correlated.The results demonstrated a pivotal role of the rat orbital cortex in olfactory sensory processing and suggest that the orbital cortex is important in the manifestation of various motivated behaviors of the animals, including odor-guided motivational behaviors (odor preference).     

Intracellular responses of identified rat olfactory bulb interneurons to electrical and odor stimulation

1. Intracellular recordings were made from 28 granule cells and 6 periglomerular cells of the rat olfactory bulb during odor stimulation and electrical stimulation of the olfactory nerve layer (ONL) and lateral olfactory tract (LOT). Neurons were identified by injection of horseradish peroxidase (HRP) or biocytin and/or intracellular response characteristics. Odorants were presented in a cyclic sniff paradigm, as reported previously. 2. All interneurons could be activated from a wide number of stimulation sites on the ONL, with distances exceeding their known dendritic spreads and the dispersion of nerve fibers within the ONL, indicating that multisynaptic pathways must also exist at the glomerular region. All types of interneurons also responded to odorant stimulation, showing a variety of responses. 3. Granule cells responded to electrical stimulation of the LOT and ONL as reported previously. However, intracellular potential, excitability, and conductance analysis suggested that the mitral cell-mediated excitatory postsynaptic potential (EPSP) is followed by a long inhibitory postsynaptic potential (IPSP). An early negative potential, before the EPSP, was also observed in every granule cell and correlated with component I of the extracellular LOT-induced field potential. We have interpreted this negativity as a "field effect," that may be diagnostic of granule cells. 4. Most granule cells exhibited excitatory responses to odorant stimulation. Odors could produce spiking responses that were either nonhabituating (response to every sniff) or rapidly habituating (response to first sniff only). Other granule cells, while spiking to electrical stimulation, showed depolarizations that did not evoke spikes to odor stimulation. These depolarizations were transient with each sniff or sustained across a series of sniffs. These physiological differences to odor stimulation correlated with granule cell position beneath the mitral cell layer for 12 cells, suggesting that morphological subtypes of granule cells may show physiological differences. Some features of the granule cell odor responses seem to correlate with some of the features we have observed in mitral/tufted cell intracellular recordings. Only one cell showed inhibition to odors. 5. Periglomerular (PG) cells showed a response to ONL stimulation that was unlike that found in other olfactory bulb neurons. There was a long-duration hyperpolarization after a spike and large depolarization or burst of spikes (20-30 ms in duration). Odor stimulation produced simple bursts of action potentials, Odor stimulation produced simple bursts of action potentials, suggesting that PG cells may simply follow input from the olfactory nerve.(ABSTRACT TRUNCATED AT 400 WORDS)     

Electrophysiological evidence for the existence of a posterior cortical-prefrontal-basal forebrain circuitry in modulating sensory responses in visual and somatosensory rat cortical areas

The prefrontal cortex (PFC) receives input from sensory neocortical regions and sends projections to the basal forebrain (BF). The present study tested the possibility that pathways from sensory cortical regions via the PFC-BF and from the BF back to specific sensory cortical areas could modulate sensory responses. Two prefrontal areas that responded to stimulation of the primary somatosensory and visual cortices were delineated: an area encompassing the rostral part of the cingulate cortex that responded to visual cortex stimulation, and a region dorso-lateral to the first in the precentral-motor association area that reacted to somatosensory cortex stimulation. Moreover, BF neurons responded to PFC electrical stimulation. They were located in the ventral pallidum, substantia innominata and the horizontal limb of the diagonal-band areas. Of the responsive BF neurons 42% reacted only to stimulation of 'visually-responsive,' 33% responded only to the 'somatosensory-responsive' prefrontal sites and the remaining neurons reacted to both prefrontal cortical areas. The effect of BF and PFC stimulations on somatosensory and visual-evoked potentials was tested. BF stimulation increased the amplitude of both sensory-evoked potentials. However, stimulation of the 'somatosensory-responsive' prefrontal area increased only somatosensory-evoked potentials while 'visually-responsive' prefrontal-area stimulation increased only visual-evoked potentials. Atropine blocked both facilitatory effects.The proposed cortico-prefronto-basalo-cortical circuitry may have an important role in cortical plasticity and selective attention.     

Contribution of amygdalar and lateral hypothalamic neurons to visual information processing of food and nonfood in monkey

Visual information processing was investigated in the inferotemporal cortical (ITCx)-amygdalar (AM)-lateral hypothalamic (LHA) axis which contributes to food-nonfood discrimination. Neuronal activity was recorded from monkey AM and LHA during discrimination of sensory stimuli including sight of food or nonfood. The task had four phases: control, visual, bar press, and ingestion. Of 710 AM neurons tested, 220 (31.0%) responded during visual phase: 48 to only visual stimulation, 13 (1.9%) to visual plus oral sensory stimulation, 142 (20.0%) to multimodal stimulation and 17 (2.4%) to one affectively significant item. Of 669 LHA neurons tested, 106 (15.8%) responded in the visual phase. Of 80 visual-related neurons tested systematically, 33 (41.2%) responded selectively to the sight of any object predicting the availability of reward, and 47 (58.8%) responded nondifferentially to both food and nonfood. Many of AM neuron responses were graded according to the degree of affective significance of sensory stimuli (sensory-affective association), but responses of LHA food responsive neurons did not depend on the kind of reward indicated by the sensory stimuli (stimulus-reinforcement association). Some AM and LHA food responses were modulated by extinction or reversal. Dynamic information processing in ITCx-AM-LHA axis was investigated by reversible deficits of bilateral ITCx or AM by cooling. ITCx cooling suppressed discrimination by vision responding AM neurons (8/17). AM cooling suppressed LHA responses to food (9/22). We suggest deep AM-LHA involvement in food-nonfood discrimination based on AM sensory-affective association and LHA stimulus-reinforcement association.     

Non-specific olfactory aversion induced by intrabulbar infusion of the GABA(A) receptor antagonist bicuculline in young rats

On postnatal day 12, young rats show an aversion to an odor to which they had been exposed along with presentations of foot shock on postnatal day 11. The acquisition of this aversive learning involves and requires disinhibition of the mitral/tufted cells induced by centrifugal noradrenergic activation during somatosensory stimulation. This olfactory learning is established only for the odor to which the rat has been exposed during conditioning. Infusion of the GABA(A) receptor antagonist bicuculline at a high dose (2.0 nmol/each olfactory bulb) into the olfactory bulb in the presence of an odor is capable of developing olfactory aversive responses without somatosensory stimulation in young rats. The purpose of this study is to characterize the properties of bicuculline-induced aversive responses. In contrast to the odor specificity of aversive learning produced by odor-shock conditioning, bicuculline-induced aversive responses lack odor specificity. Namely, bicuculline infusion in the presence of a citral odor results, in a dose-dependent manner, in subsequent aversive responses to strange odors (benzaldehyde and vanillin) that have never been presented. Moreover, bicuculline infusion alone is sufficient to produce dose-dependent aversive responses to strange odors (citral, benzaldehyde and geraniol).From these results we suggest that disinhibition of mitral/tufted cells from granule cells by bicuculline infusion makes young rats aversive to strange odors non-specifically, as if the rats had learned the odor aversion as a result of odor exposure paired with foot shock. Different mechanisms of disinhibition of the mitral/tufted cells may underlie both the pharmacological manipulation and noradrenergic activation by somatosensory stimulation.     

Projection of a cutaneous nerve to the spinal cord of the pigeon II. Responses of dorsal horn neurons

Summary The responses of dorsal horn neurons to both electrical stimulation of a cutaneous nerve and natural stimulation of skin receptors have been studied in an avian species, the pigeon. Neurons located in either lamina I or lamina IV were recorded. Most lamina IV neurons had short-latency responses to electrical stimulation of a cutaneous nerve and were activated by stimulation of sensitive mechanoreceptors. This points to an input from mechanoreceptors innervated by large afferent fibers. Lamina I neurons which were usually located near the entrance zone of small fibers had longer latency responses and had often an input from several groups of afferent fibers including C-fibers. Many lamina I neurons were activated specifically by noxious stimulation. Some had an input from sensitive mechanoreceptors but possibly through an additional synapse. A few lamina I neurons responded specifically to activation of cold receptors. Some dorsal horn neurons showed segmental inhibition. Altogether, the characteristics of dorsal horn neurons in the pigeon studied so far were similar to those in mammalian species.     

Patterned response to odor in mammalian olfactory bulb: the influence of intensity

Responses of single neurons in the olfactory bulb of anesthetized hamsters were recorded extracellularly while odors of defined concentration and time course were delivered to the olfactory system at constant flow. Responses could be either excitatory or suppressive, as judged by the first distinguishable change in firing rate during odor delivery. However, when the time course of the response was examined in more detail, approximately one-third of all tests and one-half of the tests at high concentration resulted in complex temporal patterns of firing rate that involved both increases and decreases with respect to spontaneous activity. Approximately two-thirds of all tests produced responses where increased firing rate preceded any distinguishable suppression. Excitatory and suppressive responses were each classified into four groups according to their temporal patterns. Different patterns were not equally represented in the data and the proportions of patterns elicited by the same odor changed with stimulus intensity. Complex responses, where temporal patterns included periods of firing rate above and below spontaneous rate, were increasingly common and intensity was increased. Magnitude of response is difficult to define when a single response includes both increases and decreases of firing rate but more than half of the neurons that responded to more than one stimulus concentration clearly had nonmonotonic intensity-response functions. Forty-one out of 101 neurons were classified as output cells because they could be driven at short constant latency by lateral olfactory tract stimulation. Their responses were not clearly different from the remaining cells that could not be classified as output cells. The contribution of the inhibitory circuits of the olfactory bulb to the generation of patterned response and to changes in pattern with intensity are discussed. The lateral inhibitory circuits of the bulb appear to be sufficient to explain the data presented here.     

Discrimination among odorants by single neurons of the rat olfactory bulb

1. Intracellular and extracellular recordings were made from rat olfactory bulb mitral and tufted cells during odor stimulation and during electrical stimulation of the olfactory nerve. Neurons were identified by horseradish peroxidase injections and/or antidromic activation. The presentation of multiple concentrations of at least one odorant in a cyclic artificial sniff paradigm, as reported previously (10), allowed the study of odor responses. This approach was extended to multiple odorants to compare their concentration-response profiles. This procedure avoids the problems of interpretation resulting from nonequivalence of the effective concentrations of different odorants used as stimuli that have characterized previous studies of odor quality effects. Comparisons of intracellular events and responses to electrical stimulation with the odor-induced spike train activity allow us to begin to delineate the local circuitry involved in generating odor-induced responses. 2. The concentration-response profiles of the 72 cells in the present study are comparable to those previously reported for output neurons of the olfactory bulb, showing ordered changes in the temporal patterning of spike activity with step changes in odor concentration. However, eight of the neurons exhibited inhibitory responses to lower concentrations, but excitation, at similar latency, to higher concentrations of the same odorant. These data emphasize that to study pattern changes induced by changing odor quality the influence of stimulus intensity must also be carefully examined. The data also provide evidence that the temporal pattern evoked by an odorant is probably not in itself the code for odor quality recognition. 3. Complete concentration-response profiles, including subthreshold concentrations, to more than one odorant show that, although responses to the different odorant can evolve systematically with concentration, the responses to different odorants can evolve through very different patterns. For example, in some cells, the response patterns to different odors were complementary in form. These results demonstrate that the patterned responses of olfactory bulb neurons can reflect changes in odor quality as well as intensity. 4. Intracellular recording was employed to compare the temporal patterning of spikes during odor stimulation with membrane potential changes. In some cases, the spike pattern was closely correlated with apparent postsynaptic potentials. However, there were several clear exceptions. In five cells, a prominent hyperpolarization, seen in the first sniff of a series of 10 consecutive sniffs, was associated with pauses in spike activity. In the following     

Taste and olfactory modulation of feeding related neurons in behaving monkey.

Single neuron activity in the monkey lateral hypothalamus (LHA) was recorded by multibarreled electrode during a bar press feeding task. Activity of glucose-sensitive (GS) neurons decreased during bar press (BP) and reward (RW) periods. The inhibition was caused by activation of beta-adrenoceptors and opioid receptors respectively. Glucose-insensitive (GIS) neurons were excited during BP and RW, and at cue light (CL). Excitation at CL and BP was caused by activation of dopaminergic receptors. Among GS neurons, 66% responded to taste and 88% to odor. These responses were 39% and 52% in GIS neurons. GS neurons responded predominantly to two or more taste and odor stimuli while GIS neurons responded to only one stimulant. GS neurons have dense mutual connections with the prefrontal area, and GIS neurons are connected with the motor area. Gustatory and olfactory stimulation elicited responses in 67% of GS neurons and in only 21% of GIS neurons. Data suggest that GS and GIS neurons may have different functions in feeding: GS neurons process endogenous chemical information and integrated chemical sensations, and GIS neurons process external information processing, motor control and discriminative chemical sensations.     

Neural correlates of social odor recognition and the representation of individual distinctive social odors within entorhinal cortex and ventral subiculum

Recognition of individual conspecifics is important for social behavior and requires the formation of memories for individually distinctive social signals. Individual recognition is often mediated by olfactory cues in mammals, especially nocturnal rodents such as golden hamsters. In hamsters, this form of recognition requires main olfactory system input to the lateral entorhinal cortex (LEnt). Here, we tested whether neurons in LEnt and the nearby ventral subiculum (VS) would show cellular correlates of this natural form of recognition memory. Two hundred ninety single neurons were recorded from both superficial (SE) and deep layers of LEnt (DE) and VS while male hamsters investigated volatile odorants from female vaginal secretions. Many neurons encoded differences between female's odors with many discriminating between odors from different individual females but not between different odor samples from the same female. Other neurons discriminated between odor samples from one female and generalized across collections from other females. LEnt and VS neurons showed enhanced or suppressed cellular activity during investigation of previously presented odors and in response to novel odors. A majority of SE neurons decreased firing to odor repetition and increased activity to novel odors. In contrast, DE neurons often showed suppressed activity in response to novel odors. Thus, neurons in LEnt and VS of male hamsters encode information that is critical for the identification and recognition of individual females by odor cues. This study reveals cellular mechanisms in LEnt and VS that may mediate a natural form of recognition memory in hamsters. These neuronal responses were similar to those observed in rats and monkeys during performance in standard recognition memory tasks. Consequently, the present data extend our understanding of the cellular basis for recognition memory and suggest that individual recognition requires similar neural mechanisms as those employed in laboratory tests of recognition memory.     

Discrimination of odors from stressed rats by non-stressed rats

Comparison of earlier reports of rat stress odors is complicated by the many differences in experimental parameters and responses measured. To evaluate whether these stress odors provide a special signal, rats were subjected to different levels of stressful foot-shock in one half of a simple two-compartment test box whilst the other half was clean and unoccupied. The results show that whilst test subjects preferred the half containing odors from non-stressed rats, this preference was decreased by the presence of stress odors to an extent concordant with the level of stressor applied to the odor donors. There were no differences in plasma corticosterone among the odor donors indicating that this hormone is probably not the source of stress odors. Plasma corticosterone levels of the subjects were similar to each other and to the odor donors. Compared to odors from non-stressed rats, stress odors increased the activity of the subjects. The evidence strongly suggests a special signalling function for stress odors although responses to this signal are not stereotyped.     

Odor- and context-dependent modulation of mitral cell activity in behaving rats.

The projections and odor responses of mammalian olfactory receptor neurons, as well as the physiology of the bulb's principal neurons-the mitral cells (MCs)-are known from studies in slices and anesthetized animals. In behaving rats trained to discriminate between two odors associated with different reinforcers, we examined MC responses following alternated odor-reinforcer pairings. Whereas only 11% of the recorded MCs showed changes in odor-selective firing rate during the odor-sampling phase, 94% of MCs modulated activity during specific behaviors surrounding odor sampling. These cell- and odor-selective responses were not primary sensory responses; rather, they depended (reversibly) on the predictive value of each odor. MC activity thus depends critically on efferent influences linked to the animal's experience and behavior.     

Age-associated loss of selectivity in human olfactory sensory neurons

We report a cross-sectional study of olfactory impairment with age based on both odorant-stimulated responses of human olfactory sensory neurons (OSNs) and tests of olfactory threshold sensitivity. A total of 621 OSNs from 440 subjects in 2 age groups of younger (≤ 45 years) and older (≥ 60 years) subjects were investigated using fluorescence intensity ratio fura-2 imaging. OSNs were tested for responses to 2 odorant mixtures, as well as to subsets of and individual odors in those mixtures. Whereas cells from younger donors were highly selective in the odorants to which they responded, cells from older donors were more likely to respond to multiple odor stimuli, despite a loss in these subjects' absolute olfactory sensitivity, suggesting a loss of specificity. This degradation in peripheral cellular specificity may impact odor discrimination and olfactory adaptation in the elderly. It is also possible that chronic adaptation as a result of reduced specificity contributes to observed declines in absolute sensitivity.     

Neuronal activity in the dorsolateral pontine nucleus of the alert monkey modulated by visual stimuli and eye movements

Summary The activity of neurons in the dorsolateral pontine nucleus (dlpn) was studied in two awake rhesus monkeys trained to participate in a variety of visual and oculomotor tests. The visual and eye movement related responses of 73 neurons encountered in the more caudal part of the dlpn were analyzed. Thirty eight of these could be assigned to one of the three following groups. Visual-only neurons (Type 1, n = 10) responded to movement of a broad range of visual stimuli in certain preferred directions. Their receptive fields were usually large, not restricted to the contralateral visual field and always included the fovea. Visual-tracking (VT) neurons (n = 28) discharged in relation to smooth pursuit of a small target in particular preferred directions. Nine of these (Type 2) did not respond to visual stimulation during stationary fixation. Nineteen VT-cells (Type 3) discharged in relation to both visual tracking and visual stimulation. In 9 of the Type 3 neurons, the preferred directions for visual stimulation and tracking were opposite, whereas they were the same in the other 10. Visual responses of Type 3 neurons were indistinguishable from those of Type 1 neurons. Testing of an additional 9 neurons driven by either visual-tracking or pattern movement was not sufficient to allow a definite assignment to one of the groups 1, 2 or 3. The distribution of preferred directions for both visual stimulation and visual tracking was widely scattered between 0 and 360 deg. Our results suggest that the dlpn is a constituent in a cerebro-cerebellar loop important for the generation of smooth pursuit eye movements.     

Olfactory bulb responses to odor stimulation: analysis of response pattern and intensity relationships

Extracellular recordings were made from mitral cells, tufted cells, and presumed glomerular layer and external plexiform layer interneurons of the olfactory bulb of anesthetized rats during odor stimulation. Intensity responses of these cells were studied by presenting a series of six or seven concentrations, spanning a range greater than two log units, in a cyclic artificial sniff paradigm, which produced repeated response measures at each concentration. Experiments focused on obtaining a complete intensity series, including interspersed unstimulated spontaneous activity records, for a single odorant (usually amyl acetate), but concentration responses to other odorants were tested when possible. Odor responses of 46 cells were studied with two approaches. Response form was examined in an attempt to define response classes based on qualitative characteristics of the temporal pattern of response. Assessment of response magnitude was attempted, in order to construct stimulus-response functions for each cell, independent of response form. As previously reported for olfactory bulb cells, the cells in our sample responded to odor stimulation with spike trains of a variety of temporal patterns, consisting of excitatory and inhibitory components that were frequently recognizable in the responses of a cell across a range of concentrations. However, response patterns usually changed significantly with concentration, such that response form across the concentration range could not be predicted from the response at any one concentration. Responses of different cells were sometimes similar to each other in form at one concentration and quite different from each other in the rest of their concentration-response profiles. Classification of response profiles into discrete types, based on consistency of response form throughout the profile, was therefore not feasible. In agreement with other reports, response of a single cell to different odorants sometimes showed similar forms and sometimes showed very different forms across the concentration-response profiles. Since the response form depends on the stimulus intensity as well as the stimulus quality, characterization of response magnitude and of the pattern of response to different odors require testing with a series of stimulus concentrations. Because odor responses consisted of temporally patterned spike trains, whose components changed in complex ways with stimulus intensity, it was not possible to quantify response magnitude by measuring characteristics of particular response components or counting mean frequency.(ABSTRACT TRUNCATED AT 400 WORDS)