Olfactory neuronal responses in the primate orbitofrontal cortex: analysis in an olfactory discrimination task

1. The primate orbitofrontal cortex receives inputs from the primary olfactory (pyriform) cortex and also from the primary taste cortex. To investigate how olfactory information is encoded in the orbitofrontal cortex, the responses of single neurons in the orbitofrontal cortex and surrounding areas were recorded during the performance of an olfactory discrimination task. In the task, the delivery of one of eight different odors indicated that the monkey could lick to obtain a taste of sucrose. If one of two other odors was delivered from the olfactometer, the monkey had to refrain from licking, otherwise he received a taste of saline. 2. Of the 1,580 neurons recorded in the orbitofrontal cortex, 3.1% (48) had olfactory responses and 34 (2.2%) responded differently to the different odors in the task. The neurons responded with a typical latency of 180 ms from the onset of odorant delivery. 3. Of the olfactory neurons with differential responses in the task, 35% responded solely on the basis of the taste reward association of the odorants. Such neurons responded either to all the rewarded stimuli, and none of the saline-associated stimuli, or vice versa. 4. The remaining 65% of these neurons showed differential selectivity for the stimuli based on the odor quality and not on the taste reward association of the odor. 5. The findings show that the olfactory representation within the orbitofrontal cortex reflects for some neurons (65%) which odor is present independently of its association with taste reward, and that for other neurons (35%), the olfactory response reflects (and encodes) the taste association of the odor. The additional finding that some of the odor-responsive neurons were also responsive to taste stimuli supports the hypothesis that odor-taste association learning at the level of single neurons in the orbitofrontal cortex enables such cells to show olfactory responses that reflect the taste association of the odor.     

Satiety-responsive neurons in the medial orbitofrontal cortex of the macaque.

Feeding-related gustatory, olfactory, and visual activation of the orbitofrontal cortex (OFC) decreases following satiety. Previous neurophysiological studies have concentrated on the caudolateral OFC (clOFC). We describe satiety-induced modulation of 23 gustatory, 5 water, and 15 control neurons in the medial OFC (mOFC), where gustatory neurons represent a much larger percentage of the population. For 15 of the 23 gustatory neurons (65%), every significant taste response evoked during pre-satiety testing decreased following satiety (X=70%). Responses evoked by the ineffective taste stimuli during pre-satiety testing were unchanged following satiety. The graded response decrements of the mOFC gustatory neurons stand in marked contrast to the clOFC responses, which are almost completely suppressed by satiety. Two other novel findings are reported here. First, all significant pre-satiety taste responses of four gustatory neurons increased following satiety (X=51%). Second, post-satiety emergent taste responses were observed in 7 of 15 neurons (47%) classified as non-responsive during pre-satiety testing. The presence of increased responsiveness and emergent gustatory neurons in the mOFC suggests that meal termination may require active processes as well as the passive loss of hedonic value. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sensory-specific satiety: comparison of taste and texture effects

The respective contributions of taste (saltiness and sweetness) and texture (the hardness dimension) to sensory-specific satiety (SSS) were compared. Sixteen male and 16 female, young, normal-weight adults rated the pleasantness of taste, pleasantness of texture and desire to eat on visual analog scales for eight test foods, were then given one of the foods to eat ad libitum for lunch, and re-rated the same parameters for the eight foods 2 and 20 min after the end of the meal. The experimental sets of eight test foods and four lunch foods were balanced for taste quality (salty vs. sweet) and texture quality (hard vs. soft). Lunch foods were the hard and soft versions of a salty food (ham and cheese sandwich on baguette vs. white bread) or of a sweet food (apples vs. applesauce). Sensory-specific satiety was observed for both saltiness and sweetness (e.g. pleasantness of the taste of, and desire to eat sweet test foods decreased significantly after eating a sweet lunch food and similarly for salty foods), and to a lesser extent for texture (e.g. pleasantness of the texture of, and desire to eat hard test foods decreased after eating a hard lunch food and similarly for one of the soft foods). The authors conclude texture-specific satiety may be a significant component of satiety.     

Acquired aversions as the basis for varied diets of ruminants foraging on rangelands

Ruminants eat an array of plant species that vary in nutrients and toxins. This selection makes intuitive sense, but no theories adequately explain this diversity. Some maintain it reduces the likelihood of overingesting toxins, whereas others contend it meets nutritional needs. Nevertheless, herbivores seek variety even when toxins are not a concern and nutritional needs are met. I offer another explanation for this behavior, one which encompasses the avoidance of toxins and the acquisition of nutrients. A key concept in this theory is aversion, the decrease in preference for food just eaten as a result of sensory input (a food's taste, odor, texture, i.e., its flavor) and postingestive effects (effects of nutrients and toxins on chemo-, osmo-, and mechano-receptors) unique to each food. Aversions are pronounced when foods contain toxins or high levels of rapidly digestible nutrients; they also occur when foods are deficient in specific nutrients. Aversions occur even when animals eat nutritionally adequate foods because satiety (satisfied to the full) and surfeit (filled to nauseating excess) represent points along a continuum, and there is a fine line between satiety and aversion. Thus, eating any food is likely to cause a mild aversion, and eating a food too frequently or in excess is likely to cause a strong aversion. Aversions are involuntary and are not the result of conscious decisions by an animal. Aversions yield benefits (e.g., obtain a balanced diet, reduce ingestion of toxic foods, optimize foraging and rumination times, sample foods, maintain a diverse microflora in the rumen) that are often mistaken as the cause of varied diets. In this article, I discuss the subtle ways in which aversions diminish preference and cause animals to eat a variety of foods.     

Perceptual significance of somatosensory cortical reorganization following peripheral denervation

The functional significance of reorganization in somatosensory cortex following peripheral denervation has not been thoroughly addressed. In this paper, two distinct hypotheses dealing with this issue are discussed. The first is the hypothesis of functional respecification. This influential view suggests that sets of partially deafferented cortical neurons, which respond to new peripheral inputs and acquire new receptive fields, undergo corresponding changes in perceptual meaning. Excitation of these neurons by stimulation of their novel receptive fields is thought to result in a change in referral of sensation from the original (now denervated) skin fields to the newly acquired skin fields. The second hypothesis is that of functional conservation. This equally plausible alternative is that sets of partially deprived neurons, although they respond to novel peripheral inputs, retain their original perceptual meaning. Excitation of these neurons by stimulation of their new receptive fields is thought to evoke sensation formerly mediated by those neurons, and hence is still projected to the original, now denervated skin regions or phantom. Behavioral evidence strongly suggests that cortical reorganization after peripheral denervation does not result in major functional respecification, but that the original perceptual function mediated by those neurons is preserved.     

A role for the right anterior temporal lobe in taste quality recognition

We conducted two experiments to examine central processing of the taste of citric acid. In the first experiment, elevated citric acid recognition thresholds, but normal detection thresholds, were observed in a group of patients who had undergone a right anterior temporal lobectomy for the treatment of epilepsy, compared with a control group and a group of patients who had undergone the same operation in the left hemisphere. In the second study, using positron emission tomography, we compared regional cerebral blood flow (rCBF) in a condition in which citric acid was presented with one in which water was presented (with similar somatosensory stimulation across both conditions). We observed increased rCBF bilaterally in the caudolateral orbitofrontal cortex, in the right anteromedial temporal lobe, and in the right caudomedial orbitofrontal cortex. The elevated recognition thresholds exhibited in patients with resection of the right anteromedial temporal lobe may be accounted for by damage in an area corresponding to that of the rCBF increase. These results suggest that although taste sensation may be computed in the primary taste cortex, recognition requires further processing by structures located in the anteromedial temporal lobe. Furthermore, they point to preferential processing of this higher-order gustatory function by the right cerebral hemisphere.     

Dietary variety, energy regulation, and obesity.

Increased variety in the food supply may contribute to the development and maintenance of obesity. Thirty-nine studies examining dietary variety, energy intake, and body composition are reviewed. Animal and human studies show that food consumption increases when there is more variety in a meal or diet and that greater dietary variety is associated with increased body weight and fat. A hypothesized mechanism for these findings is sensory-specific satiety, a phenomenon demonstrating greater reductions in hedonic ratings or intake of foods consumed compared with foods not consumed. Nineteen studies documenting change in preference, intake, and hedonic ratings of food after a food has been eaten to satiation in animals and humans are reviewed, and the theory of sensory-specific satiety is examined. The review concludes with the relevance of oral habituation theory as a unifying construct for the effects of variety of sensory-specific satiety, clinical implications of dietary variety and sensory-specific satiety on energy regulation, and suggestions for future research. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Olfactory memory and maternal behaviour-induced changes in c-fos and zif/268 mRNA expression in the sheep brain

In sheep maternal behaviour and the formation of the selective olfactory, ewe/lamb bond are induced by feedback to the brain from stimulation of the vagina and cervix during parturition. In the present study, we have used in situ hybridization histochemistry to quantify changes in cellular expression of two immediately-early genes, c-fos and zif/268, in order to identify activated brain regions during the induction of maternal behaviour and olfactory bonding as well as regions where plastic changes are occurring during with the formation of the olfactory memory associated with bonding. Three different treatment groups were used. One group gave birth normally, became maternal and were allowed to interact with their lambs for 30 min. A second group received exogenous treatment with oestradiol and progesterone to induce lactation and then received a 5-min period of artificial stimulation of the vagina and cervix (VCS) which reliably induces maternal behaviour but could not interact with lambs. A final control group received exogenous hormone treatment but no VCS or interaction with lambs. Compared to the control group, post-partum animals and animals that had received VCS showed increased c-fos expression in a number of cortical regions (cingulate, entorhinal and somatosensory), the mediodorsal thalamic nucleus and the lateral habenula, the limbic system (bed nucleus of the stria terminalis, lateral septum, medial arnygdala, dentate gyrus and the CA3 region of the hippocampus) and the hypothalamus (medial preoptic area, mediobasal hypothalamus, paraventricular nucleus, supraoptic nucleus and periventricular complex). The group that gave birth and had contact with their lambs for 30 min had significantly enhanced c-fos mRNA expression in the cingulate cortex compared to those receiving VCS and additionally showed significantly increased c-fos mRNA expression in olfactory processing regions (olfactory bulb, piriform cortex and orbitofrontal cortex). Expression of zif/268 was significantly increased in the entorhinal cortex, orbitofrontal cortex and dentate gyrus of the parturition group compared to either the control or the VCS alone groups. These results show a clear differentiation between neural substrates controlling the expression of maternal behaviour and those involved in the olfactory memory process associated with selective recognition of offspring although at the level of the hippocampus and cingulate cortex there may be some degree of overlap. Alterations in zif/268 at tertiary processing sites for olfactory information (orbitofrontal cortex) and the entorhinal cortex and dentate gyrus may reflect plastic changes occurring during the early stages of olfactory memory formation.     

Effect of sensory perception of foods on appetite and food intake: a review of studies on humans

OBJECTIVE: How much do the sensory properties of food influence the way people select their food and how much they eat? The objective of this paper is to review results from studies investigating the link between the sensory perception of food and human appetite regulation. CONTENT OF THE REVIEW: The influence of palatability on appetite and food intake in humans has been investigated in several studies. All reviewed studies have shown increased intake as palatability increased, whereas assessments of the effect of palatability using measures of subjective appetite sensations have shown diverging results, for example, subjects either feel more hungry and less full after a palatable meal compared to a less palatable meal, or they feel the opposite, or there is no difference. Whether palatability has an effect on appetite in the period following consumption of a test meal is unclear. Several studies have investigated which sensory properties of food are involved in sensory-specific satiety. Taste, smell, texture and appearance-specific satieties have been identified, whereas studies on the role of macronutrients and the energy content of the food in sensory-specific satiety have given equivocal results. Different studies have shown that macronutrients and energy content play a role in sensory-specific satiety or that macronutrients and energy content are not a factor in sensory-specific satiety. Sensory-specific satiety may have an important influence on the amount of food eaten. Studies have shown that increasing the food variety can increase food and energy intake and in the short to medium term alter energy balance. Further knowledge about the importance of flavour in appetite regulation is needed, for example, which flavour combinations improve satiety most, the possible connection between flavour intensity and satiety, the effect of persistence of chemesthetic sensation on palatability and satiety, and to what extent genetic variation in taste sensitivity and perception influences dietary habits and weight control.     

Somatosensory and auditory convergence in the lateral nucleus of the amygdala.

Previous studies have shown that the lateral nucleus of the amygdala (AL) is essential in auditory fear conditioning and that neurons in the AL respond to auditory stimuli. The goals of the present study were to determine whether neurons in the AL are also responsive to somatosensory stimuli and, if so, whether single neurons in the AL respond to both auditory and somatosensory stimulation. Single-unit activity was recorded in the AL in anesthetized rats during the presentation of acoustic (clicks) and somatosensory (footshock) stimuli. Neurons in the dorsal subdivision of the AL responded to both somatosensory and auditory stimuli, whereas neurons in the ventrolateral AL responded only to somatosensory stimuli and neurons in the ventromedial AL did not respond to either stimuli. These findings indicate that the dorsal AL is a site of auditory and somatosensory convergence and may therefore be a focus of convergence of conditioned and unconditioned stimuli (CS and UCS) in auditory fear conditioning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Deafferentation causes apoptosis in cortical sensory neurons in the adult rat

The present study provides an experimental model of the apoptotic death of pyramidal neurons in rat olfactory cortex after total bulbectomy. Terminal transferase (TdT)-mediated deoxyuridine triphosphate (d-UTP)-biotin nick end labeling (TUNEL), DNA electrophoresis, and neuronal ultrastructure were used to provide evidence of apoptosis; neurons in olfactory cortex were counted by stereology. Maximal TUNEL staining occurred in the piriform cortex between 18 and 26 hr postbulbectomy. Within the survival times used in the present study (up to 48 hr postlesion), cell death was observed exclusively in the piriform cortex; there was no evidence of cell death in any other areas connected with the olfactory bulb. Neurons undergoing apoptosis were pyramidal cells receiving inputs from, but not projecting to, the olfactory bulb. The apical dendrites of these neurons were contacted by large numbers of degenerating axonal terminals. Gel electrophoresis of DNA purified from lesioned olfactory cortex showed a ladder pattern of fragmentation. Inflammatory cells or phagocytes were absent in the environment of degenerating neurons in the early stages of the apoptotic process. The present model suggests that deafferentation injury in sensory systems can cause apoptosis. In addition, olfactory bulbectomy can be used for investigating molecular mechanisms that underlie apoptosis in mature mammalian cortical neurons and for evaluating strategies to prevent the degeneration of cortical neurons.     

Patterns of Fos expression in the amygdala and ventral perirhinal cortex induced by training in an olfactory fear conditioning paradigm.

The activation of amygdaloid nuclei, the ventral perirhinal cortex (vPRh), and several other brain areas in the rat during the acquisition and expression of olfactory fear conditioning was assessed through Fos immunocytochemistry in 3 separate experiments. The results of Experiment 1 suggest that olfactory and somatosensory inputs may functionally converge in the anterior region of the medial nucleus (aMe). The results of Experiment 2 indicate that the aMe exhibited significantly greater Fos-like immunoreactivity (FLI) in subjects acquiring conditioned stimulus–unconditioned stimulus associations than in those presented with the same olfactory and somatosensory stimuli in a manner that precluded acquisition. The results of Experiment 3 indicate that the vPRh appeared to exhibit learning-related increases in FLI during the expression of previously acquired associations. Collectively, these data suggest that the aMe and vPRh may be critically involved in different aspects of olfactory fear conditioning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Destruction of olfactory inputs affects the morphogenesis of the telencephalon in rats

We unilaterally destroyed the nasal radix of rat embryos on day 15.5 of gestation (E15.5) in utero so as to block the olfactory inputs to the ipsilateral forebrain vesicle. The embryonic brains were examined after 6 days' survival (E21.5). In the deafferented half of the brain, LHRH neurons were significantly reduced in number, indicating the successful blocking of the olfactory input. On the deafferented side, the olfactory bulb failed to develop, and the telencephalic hemisphere, small in size, accompanied various histogenetic retardations in the primary olfactory cortex, in the cortical plate, and in the hippocampal formation. The striatum revealed remarkable structural differences between the ipsilateral and contralateral sides: on the ipsilateral side, the striatum was small in size and displayed numerical reductions of immunoreactive tyrosine hydroxylase (TH) fibers and substance P (SP) neurons in comparison with those in the contralateral one; in the substantia nigra, TH neurons and SP fibers were less numerous on the deafferented side. There were no remarkable differences in the distribution of TH neurons in the hypothalamus. In view of these sequential histogenetic alterations, it can be assumed that the olfactory inputs play a key role in the telencephalic morphogenesis.     

Selective loss of cholinergic neurons projecting to the olfactory system increases perceptual generalization between similar, but not dissimilar, odorants.

The neuromodulator acetylcholine is thought to modulate information processing in the olfactory system. The authors used 192 IgG-saporin, a lesioning agent selective for basal forebrain cholinergic neurons, to determine whether selective lesions of cholinergic neurons projecting to the olfactory bulb and cortex affect odor perception in rats. Lesioned and sham-operated rats were tested in an olfactory generalization paradigm with sets of chemically related odorants (n-aliphatic aldehydes, acids, and alcohols). Lesioned rats generalized more between chemically similar odorants but did not differ from controls in their response to chemically unrelated odorants or in acquisition of the conditioned odor. Results show that cholinergic inputs to the olfactory system influence perceptual qualities of odorants and confirm predictions made by computational models of this system. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

GABAergic neurons in barrel cortex show strong, whisker-dependent metabolic activation during normal behavior

Electrophysiological data from the rodent whisker/barrel cortex indicate that GABAergic, presumed inhibitory, neurons respond more vigorously to stimulation than glutamatergic, presumed excitatory, cells. However, these data represent very small neuronal samples in restrained, anesthetized, or narcotized animals or in cortical slices. Histochemical data from primate visual cortex, stained for the mitochondrial enzyme cytochrome oxidase (CO) and for GABA, show that GABAergic neurons are more highly reactive for CO than glutamatergic cells, indicating that inhibitory neurons are chronically more active than excitatory neurons but leaving doubt about the short-term stimulus dependence of this activation. Taken together, these results suggest that highly active inhibitory neurons powerfully influence relatively inactive excitatory cells but do not demonstrate directly the relative activities of excitatory and inhibitory neurons in the cortex during normal behavior. We used a novel double-labeling technique to approach the issue of excitatory and inhibitory neuronal activation during behavior. Our technique combines high-resolution 2-deoxyglucose (2DG), immunohistochemical staining for neurotransmitter-specific antibodies, and automated image analysis to collect the data. We find that putative inhibitory neurons in barrel cortex of behaving animals are, on average, much more heavily 2DG-labeled than presumed excitatory cells, a pattern not seen in animals anesthetized at the time of 2DG injection. This metabolic activation is dependent specifically on sensory inputs from the whiskers, because acute trimming of most whiskers greatly reduces 2DG labeling in both cell classes in columns corresponding to trimmed whiskers. Our results provide confirmation of the active GABAergic cell hypothesis suggested by CO and single-unit data. We conclude that strong activation of inhibitory cortical neurons must confer selective advantages that compensate for its inherent energy inefficiency.     

Effects of intake level of a mixed diet on chewing activity and on particle size of ruminated boli, ruminal digesta fractions and faeces of steers

In the rat cortical taste area (CTA), we recorded 31 pairs of taste neurons and seven pairs of taste and non-taste neurons, with single or double electrodes. By using a cross-correlogram (CCG) in a stationary state, we examined the functional interaction between neurons of the pairs while activating them by taste stimulation. Though only 14.3% of the taste and non-taste neuron pairs were correlated, 54.8% of the taste neuron pairs showed correlated activities, 41.9% of them showing common inputs, including one with an additional excitatory connection. The remainder (12.9%) showed excitatory connections with a time lag of 1-3 ms. When pairs were recorded using single or double electrodes with an intertip distance of < 50 microns in a dorsoventral direction, a larger fraction had correlated activities than when the intertip distance was > 50 microns. Whereas pairs of neurons showed correlated activities in area DI whatever the vertical intertip distance was, most of the pairs having correlated activities in area GI were found within 50 microns of the vertical intertip distance. The taste profiles of common inputs to the pair were estimated on the basis of peak at time 0 in CCGs for various taste stimuli. The efficacy contribution of the source to target neurons tended to be larger when both had the same best stimulus. This tendency held true for pairs showing excitatory connections. Interlayer excitatory connections were also evident. It is concluded that a functional column with a diameter of 50 microns may present in the CTA in rats, and that information flow is larger between pairs of neurons with the same best stimulus.     

Neural mechanisms for synthesizing sensory information and producing adaptive behaviors

The ability to integrate information from different sensory systems is a fundamental characteristic of the brain. Because different bits of information are derived from different sensory channels, their synthesis markedly enhances the detection and identification of external stimuli. The neural substrate for such "multisensory" integration is provided by neurons that receive convergent input from two or more sensory modalities. Many such multisensory neurons are found in the superior colliculus (SC), a midbrain structure that plays a significant role in overt attentive and orientation behaviors. The various principles governing the integration of visual, auditory, and somatosensory inputs in SC neurons have been explored in several species. Thus far, the evidence suggests a remarkable conservation of integrative features during vertebrate evolution. One of the most robust of these principles is based on spatial relationships: a striking enhancement in activity is induced in a multisensory neuron when two different sensory stimuli (e.g., visual and auditory) are in spatial concordance, whereas a profound response depression can be induced when these cues are spatially discordant. The most extensive physiological observations have been made in cat, and in this species the same principles that have been shown to govern multisensory integration at the level of the individual SC neuron have also been shown to govern overt attentive and orientation responses to multisensory stimuli. Most surprising, however, is the critical role played by association (i.e. anterior ectosylvian) cortex in facilitating these midbrain processes. In the absence of the modulating corticotectal influences, multisensory SC neurons in cat are unable to integrate the different sensory cues converging upon them in an adult-like fashion, and are unable to mediate overt multisensory behaviors. This situation appears quite similar to that observed during early postnatal life. When multisensory SC neurons first appear, they are able to respond to multiple sensory inputs but are unable to synthesize these inputs to significantly enhance or degrade their responses. During ontogeny, individual multisensory neurons develop this capacity abruptly, but at very different ages, until the mature population condition is reached after several postnatal months. It appears likely that the abrupt onset of this capacity in any individual SC neuron reflects the maturation of inputs from anterior ectosylvian cortex. Presumably, the functional coupling of cortex with an individual SC neuron is essential to initiate and maintain that neuron's capability for multisensory integration throughout its life.     

Synaptic relationships between the chorda tympani and tyrosine hydroxylase-immunoreactive dendritic processes in the gustatory zone of the nucleus of the solitary tract in the hamster

The toxic lectin ricin was applied to the hamster chorda tympani (CT), producing anterograde degeneration of its terminal boutons within the gustatory zone of the nucleus of the solitary tract (NST). Immunocytochemistry was subsequently performed with antiserum against tyrosine hydroxylase (TH), and the synaptic relationships between degenerating CT terminal boutons and either TH-immunoreactive or unlabeled dendritic processes were examined at the electron microscopic level. Degenerating CT terminal boutons formed asymmetric axodendritic synapses and contained small, clear, spherical synaptic vesicles that were densely packed and evenly distributed throughout the ending, with no accumulation at the active synaptic. The degenerating CT terminated on the dendrites of TH-immunoreactive neurons in 36% (35/97) of the cases. The most frequent termination pattern involved the CT and two or three other inputs in synaptic contact with a single immunoreactive dendrite, resulting in a glomerular-like structure that was enclosed by glial processes. In 64% (62/97) of the cases, the degenerating CT was in synaptic contact with unlabeled dendrites, often forming a calyx-like synaptic profile that surrounded much of the perimeter of a single unlabeled dendrite. These results indicate that the TH-immunoreactive neurons of the gustatory NST receive direct input from the CT and taste receptors of the anterior tongue and that the termination patterns of the CT vary with its target neuron in the gustatory NST. The glomerular-like structure that characterizes many of the terminations of the CT provides an opportunity for the convergence of several functionally distinct inputs (both gustatory and somatosensory) onto putative dopaminergic neurons that may shape their responsiveness to the stimulation of the oral cavity.     

A study of a highly specific pyroglutamyl aminopeptidase type-II from the membrane fraction of bovine brain

The purpose of these experiments was to define the topography of cuneate and spinal projections to the forelimb representation in the rostral dorsal accessory olive (rDAO). We were interested in determining whether the spinal and cuneate inputs constitute a homogeneous afferent source, and whether there is evidence that they serve different functional roles. We were also interested in determining whether the somatotopy of rDAO is the result of a point-to-point projection from its afferent sources, or whether the projection suggests a reorganization of afferents at the olive. Single unit recording was used to identify specific regions of rDAO, and the topography of inputs to the identified regions was determined by using wheat germ agglutinin-horseradish peroxidase (WGA-HRP) as a tracer. The results from retrograde tracing were confirmed by using WGA-HRP as an anterograde tracer from input sources. The cuneate and spinal neurons providing input to rDAO constitute two distinct neural populations. One consists of cells in the caudal cuneate nucleus and lamina VI of the rostral two cervical segments, the other consists of cells in the rostral cuneate nucleus. The cells in the caudal cuneate nucleus and the rostral cervical segments are large, multipolar neurons that form a single column of rDAO input cells. The column of cells projects to the contralateral rDAO in a topographic fashion with rostral regions of the column projecting to rostral rDAO, which contains cells that respond to somatosensory stimulation of the contralateral shoulder, trunk, and proximal forelimb. Caudal regions of the column project to caudal rDAO, which contains cells that respond to stimulation of the distal forelimb. Despite this topography, there is a large degree of overlap in the terminations from neighboring regions of the input column, indicating that a major reorganization occurs at the rDAO. The projection from the rostral cuneate nucleus arises from small neurons that project bilaterally to rDAO, and the input from the rostral cuneate nucleus lacks a clear topography. We propose that input from the cell column is responsible for the somatosensory sensitivity of rDAO neurons, whereas input from rostral cuneate is most likely modulatory, probably inhibitory, in nature.