Multiunit Electrophysiological Recordings Research Articles

Assessing Goodness-of-Fit in Marked Point Process Models of Neural Population Coding via Time and Rate Rescaling.

Marked point process models have recently been used to capture the coding properties of neural populations from multiunit electrophysiological recordings without spike sorting. These clusterless models have been shown in some instances to better describe the firing properties of neural populations than collections of receptive field models for sorted neurons and to lead to better decoding results. To assess their quality, we previously proposed a goodness-of-fit technique for marked point process models based on time rescaling, which for a correct model produces a set of uniform samples over a random region of space. However, assessing uniformity over such a region can be challenging, especially in high dimensions. Here, we propose a set of new transformations in both time and the space of spike waveform features, which generate events that are uniformly distributed in the new mark and time spaces. These transformations are scalable to multidimensional mark spaces and provide uniformly distributed samples in hypercubes, which are well suited for uniformity tests. We discuss the properties of these transformations and demonstrate aspects of model fit captured by each transformation. We also compare multiple uniformity tests to determine their power to identify lack-of-fit in the rescaled data. We demonstrate an application of these transformations and uniformity tests in a simulation study. Proofs for each transformation are provided in the appendix.

The Organization and Connections of Second Somatosensory Cortex in the Agouti.

In order to understand how the mammalian sensory cortex has been structured during evolution, it is necessary to compare data from different species across distinct mammalian lineages. Here, we investigated the organization of the secondary somatosensory area (S2) in the agouti (Dasyprocta aguti), a medium-sized Amazonian rodent, using microelectrode mapping techniques and neurotracer injections. The topographic map obtained from multiunit electrophysiological recordings were correlated with both cytochrome oxidase (CO) histochemistry and with patterns of corticocortical connections in tangential sections. The electrophysiological mapping of the lateral strip of parietal cortex adjacent to the primary somatosensory area (S1) revealed that S2 displays a mirror-reversed topographical representation of S1, but with a smaller cortical magnification factor. The caudal border of S2 is surrounded by sensory fields which also respond to auditory stimulation. BDA injections into the forelimb representation of S2 revealed a dense homotopic ipsilateral projection to S1, supplemented by a less dense projection to the caudolateral cortex located near the rhinal sulcus (parietal rhinal area) and to a frontal region probably associated with the motor cortex. Our findings were similar to those described in other mammalian species, reinforcing the existence of a common plan of organization for S2 in the mammalian parietal cortex.

Behavioral and neural auditory thresholds in a frog.

Vocalizations play a critical role in mate recognition and mate choice in a number of taxa, especially, but not limited to, orthopterans, frogs, and birds. But receivers can only recognize and prefer sounds that they can hear. Thus a fundamental question linking neurobiology and sexual selection asks—what is the threshold for detecting acoustic sexual displays? In this study, we use 3 methods to assess such thresholds in túngara frogs: behavioral responses, auditory brainstem responses, and multiunit electrophysiological recordings from the midbrain. We show that thresholds are lowest for multiunit recordings (ca. 45 dB SPL), and then for behavioral responses (ca. 61 dB SPL), with auditory brainstem responses exhibiting the highest thresholds (ca. 71 dB SPL). We discuss why these estimates differ and why, as with other studies, it is unlikely that they should be the same. Although all of these studies estimate thresholds they are not measuring the same thresholds; behavioral thresholds are based on signal salience whereas the 2 neural assays estimate physiological thresholds. All 3 estimates, however, make it clear that to have an appreciation for detection and salience of acoustic signals we must listen to those signals through the ears of the receivers.

Congenital foot deformation alters the topographic organization in the primate somatosensory system.

Limbs may fail to grow properly during fetal development, but the extent to which such growth alters the nervous system has not been extensively explored. Here we describe the organization of the somatosensory system in a 6-year-old monkey (Macaca radiata) born with a deformed left foot in comparison to the results from a normal monkey (Macaca fascicularis). Toes 1, 3, and 5 were missing, but the proximal parts of toes 2 and 4 were present. We used anatomical tracers to characterize the patterns of peripheral input to the spinal cord and brainstem, as well as between thalamus and cortex. We also determined the somatotopic organization of primary somatosensory area 3b of both hemispheres using multiunit electrophysiological recording. Tracers were subcutaneously injected into matching locations of each foot to reveal their representations within the lumbar spinal cord, and the gracile nucleus (GrN) of the brainstem. Tracers injected into the representations of the toes and plantar pads of cortical area 3b labeled neurons in the ventroposterior lateral nucleus (VPL) of the thalamus. Contrary to the orderly arrangement of the foot representation throughout the lemniscal pathway in the normal monkey, the plantar representation of the deformed foot was significantly expanded and intruded into the expected representations of toes in the spinal cord, GrN, VPL, and area 3b. We also observed abnormal representation of the intact foot in the ipsilateral spinal cord and contralateral area 3b. Thus, congenital malformation influences the somatotopic representation of the deformed as well as the intact foot.

The neurobiology of predation: sensory adaptations to carnivory (89.3)

Study objective: We investigated the central and peripheral nervous systems of northern grasshopper mice (Onychomys leucogaster), the most highly carnivorous rodents in North America, in order to explore potential neural correlates of the transition to carnivory.Methods: Multiunit electrophysiological recordings combined with analysis of flattened sections of neocortex processed for cytochrome oxidase were used to determine the topography of primary somatosensory cortex (S1) and the location and size of both the visual and auditory cortex. These findings were then related to flattened cortical sections of juvenile grasshopper mice labeled with the serotonin transporter (SERT) antibody. The number of myelinated axons in trigeminal, cochlear, and optic nerves were also quantified.Results: An enlarged forepaw representation was discovered within S1. Primary visual cortex (V1) also appeared to be larger in flattened cortex preparations compared to other rodents. Whereas the trigeminal and cochlear nerves were comparable to other rodents, the optic nerve contained a large number of axons.Conclusions: Grasshopper mice utilize a more specialized repertoire of attacks that rely on the forelimb, which perhaps corresponds to a magnification of this representation in S1 relative to other rodents. Enlargements of the optic nerve and V1 may also optimize processing of visual cues to enhance predatory success.Grant Funding Source: Supported by NIH grant # DE016061 and NSF grant # 0844743 to K.C.C.

Experience with Sugar Modifies Behavioral but not Taste-Evoked Medullary Responses to Sweeteners in Mice

Dietary exposure to sugars increases the preference for and intake of sugar solutions in mice. We used brief-access lick tests and multiunit electrophysiological recordings from the nucleus of the solitary tract (NST) to investigate the role of taste in diet-induced changes in sucrose responsiveness. We exposed C57BL/6J (B6) and 129X1/SvJ (129) mice to either a sucrose diet (chow, water, and a 500mM sucrose solution) or a control diet (chow and water) for 3 days. In B6 mice, exposure to the sucrose diet decreased the appetitive response (i.e., number of trials initiated) but had no effect on the consummatory response (i.e., rate of licking) to 500mM sucrose and 20mM saccharin. In 129 mice, exposure to the sucrose diet increased the appetitive response but had no effect on the consummatory response to the sweetener solutions. In the NST recordings, the B6 mice exhibited larger multiunit responses to sweeteners than 129 mice, but there was no effect of the sucrose diet in either strain. Our results indicate that sucrose exposure alters the appetitive response of B6 and 129 mice to sweeteners in diametrically opposed ways and that these changes are mediated by structures in the gustatory neuraxis above the NST (e.g., ventral forebrain).

A/J and C57BL/6J mice differ in chorda tympani responses to NaCl

The molecular mechanisms of sodium taste transduction are not completely understood, especially those responsible for the portion of NaCl's taste in rodents that is not blocked by amiloride. As a prelude to conducting genetic analyses of peripheral NaCl taste responsiveness, we performed multiunit electrophysiological recordings from the chorda tympani (CT) nerve in C57BL/6J (B6) and A/J mice. Mice were anesthetized, the CT was accessed, and taste solutions were flowed over the tongue in order to measure the integrated whole-nerve response. NaCl was delivered before and during application of 100μM amiloride. Pre-amiloride responses were significantly larger in A/J than B6 mice for 1–8mM NaCl. Responses to NaCl were suppressed significantly by amiloride in both strains and to similar degrees. However, the size of the amiloride-insensitive NaCl response component was significantly larger in A/J mice than in B6 mice for NaCl at 2–16mM. These data help to explain the prior observation that the strains differ in behavioral taste thresholds for NaCl. Specifically, the results suggest that perception of sodium-specific taste by mice depends on the ratio of amiloride-sensitive and -insensitive responses in the CT, rather than on the absolute level of the whole-nerve response to NaCl or on the size of the amiloride-sensitive component alone. Because the B6 and A/J mice differed in the size of their amiloride-insensitive components, they may prove useful in future genetic work designed to characterize the underlying transduction mechanisms.

Cutaneous and periodontal inputs to the cerebellum of the naked mole-rat (Heterocephalus glaber)

The naked mole-rat (Heterocephalus glaber) is a small fossorial rodent with specialized dentition that is reflected by the large cortical area dedicated to representation of the prominent incisors. Due to naked mole-rats’ behavioral reliance on the incisors for digging and for manipulating objects, as well as their ability to move the lower incisors independently, we hypothesized that expanded somatosensory representations of the incisors would be present within the cerebellum in order to accommodate a greater degree of proprioceptive, cutaneous, and periodontal input. Multiunit electrophysiological recordings targeting the ansiform lobule were used to investigate tactile inputs from receptive fields on the entire body with a focus on the incisors. Similar to other rodents, a fractured somatotopy appeared to be present with discrete representations of the same receptive fields repeated within each folium of the cerebellum. These findings confirm the presence of somatosensory inputs to a large area of the naked mole-rat cerebellum with particularly extensive representations of the lower incisors and mystacial vibrissae. We speculate that these extensive inputs facilitate processing of tactile cues as part of a sensorimotor integration network that optimizes how sensory stimuli are acquired through active exploration and in turn adjusts motor outputs (such as independent movement of the lower incisors). These results highlight the diverse sensory specializations and corresponding brain organizational schemes that have evolved in different mammals to facilitate exploration of and interaction with their environment.

Organization of somatosensory cortex in the Northern grasshopper mouse (Onychomys leucogaster), a predatory rodent.

Northern grasshopper mice (Onychomys leucogaster) are among the most highly carnivorous rodents in North America. Because predatory mammals may have specialization of senses used to detect prey, we investigated the organization of sensory areas within grasshopper mouse neocortex and quantified the number of myelinated axons in grasshopper mouse trigeminal, cochlear, and optic nerves. Multiunit electrophysiological recordings combined with analysis of flattened sections of neocortex processed for cytochrome oxidase were used to determine the topography of primary somatosensory cortex (S1) and the location and size of both the visual and auditory cortex in adult animals. These findings were then related to the distinctive chemoarchitecture of layer IV visible in flattened cortical sections of juvenile grasshopper mice labeled with the serotonin transporter (SERT) antibody, revealing a striking correspondence between electrophysiological maps and cortical anatomy.

Circadian oscillators in the epithalamus

The habenula complex is implicated in a range of cognitive, emotional and reproductive behaviors, and recently this epithalamic structure was suggested to be a component of the brain's circadian system. Circadian timekeeping is driven in cells by the cyclical activity of core clock genes and proteins such as per2/PER2. There are currently no reports of rhythmic clock gene/protein expression in the habenula and therefore the question of whether this structure has an intrinsic molecular clock remains unresolved. Here, using videomicroscopy imaging and photon-counting of a PER2::luciferase (LUC) fusion protein together with multiunit electrophysiological recordings, we tested the endogenous circadian properties of the mouse habenula in vitro. We show that a circadian oscillator is localized primarily to the medial portion of the lateral habenula. Rhythms in PER2:: LUC bioluminescence here are visualized in single cells and oscillations continue in the presence of the sodium channel blocker, tetrodotoxin, indicating that individual cells have intrinsic timekeeping properties. Ependymal cells lining the dorsal third ventricle also express circadian oscillations of PER2. These findings establish that neurons and non-neuronal cells in the epithalamus express rhythms in cellular and molecular activities, indicating a role for circadian oscillators in the temporal regulation of habenula controlled processes and behavior.

Functional evidence for internal feedback in the songbird brain nucleus HVC

The song control system of songbirds consists mainly of the 'motor pathway' and 'anterior forebrain pathway'. The medial magnocellular nucleus of anterior nidopallium (mMAN) projects to the song control nucleus HVC, which is the point of divergence of the two pathways. We made simultaneous multiunit electrophysiological recordings from the mMAN and HVC in anesthetized Bengalese finches. We confirmed that the mMAN neurons showed song-selective auditory responses, and found temporal correlations between song-related activities of the mMAN and HVC neurons. The temporal relationship between the neural activation of the HVC and mMAN suggests that these nuclei are parts of a closed loop, which could provide internal feedback to the HVC for sequential syllable control.

Does Reorganization in the Cuneate Nucleus Following Neonatal Forelimb Amputation Influence Development of Anomalous Circuits Within the Somatosensory Cortex?

Neonatal forelimb amputation in rats produces sprouting of sciatic nerve afferent fibers into the cuneate nucleus (CN) and results in 40% of individual CN neurons expressing both forelimb-stump and hindlimb receptive fields. The forelimb-stump region of primary somatosensory cortex (S-I) of these rats contains neurons in layer IV that express both stump and hindlimb receptive fields. However, the source of the aberrant input is the S-I hindlimb region conveyed to the S-I forelimb-stump region via intracortical projections. Although the reorganization in S-I reflects changes in cortical circuitry, it is possible that these in turn are dependent on the CN reorganization. The present study was designed to directly test whether the sprouting of sciatic afferents into the CN is required for expression of the hindlimb inputs in the S-I forelimb-stump field. To inhibit sprouting, neurotrophin-3 (NT-3) was applied to the cut nerves following amputation. At P60 or older, NT-3-treated rats showed minimal sciatic nerve fibers in the CN. Multiunit electrophysiological recordings in the CN of NT-3-treated, amputated rats revealed 6.3% of sites were both stump/hindlimb responsive, compared with 30.5% in saline-treated amputated animals. Evaluation of the S-I following GABA receptor blockade, revealed that the percentage of hindlimb responsive sites in the stump representation of the NT-3-treated rats (34.2%) was not significantly different from that in saline-treated rats (31.5%). These results indicate that brain stem reorganization in the form of sprouting of sciatic afferents into the CN is not necessary for development of anomalous hindlimb receptive fields within the S-I forelimb/stump region.

Subthalamic nucleus stimulation in Parkinson's disease: Postoperative CT–MRI fusion images confirm accuracy of electrode placement using intraoperative multi-unit recording

Deep brain stimulation (DBS) of the subthalamic nucleus (STN) is increasingly used to treat advanced Parkinson's disease (PD). The optimal method for targeting the STN before implanting the definitive DBS electrode is still a matter of debates. Beside methods of direct visualization of the nucleus based on stereotactic magnetic resonance imaging (MRI), the most often used technique for targeting STN consists in recording single-cell activity along exploratory tracks of 10-15mm in length, centered on the theoretical or MRI-defined target coordinates. Single-unit recordings with a microelectrode present various drawbacks. They are time-consuming if correctly performed and a single-cell precision is probably superfluous, taking into account the size of the implanted electrode. In this study, we present an original method of recording and quantification of a multi-unit signal recorded intraoperatively with a semi-microelectrode for targeting the STN. Twelve patients with advanced PD have been included and assessed clinically before and one year after bilateral STN-DBS electrode implantation guided by multi-unit electrophysiological recordings. After one year of chronic stimulation, all patients showed a marked clinical improvement. The motor score of the unified Parkinson's disease rating scale decreased by more than 57% and the required levodopa-equivalent daily dose by 59.5% in on-stimulation off-medication condition compared to off-stimulation off-medication condition. The accuracy of STN-DBS lead placement was confirmed on postoperative computed tomography (CT) scans, which were fused to preoperative T2-weighted MRI. The boundaries of the STN were easily determined by an increase in multi-unit signal amplitude, which was observed on average from 0.492mm below the rostral border of the STN down to 0.325mm above its caudal border. Signal amplitude significantly increased at the both rostral and caudal STN margins (P<0.05) and the level of neuronal activity easily distinguished inside from outside the nucleus. This study showed that STN boundaries could be adequately determined on the basis of intraoperative multi-unit recording with a semi-microelectrode. The accuracy of our method used for positioning DBS electrodes into the STN was confirmed both on CT-MRI fusion images and on the rate of therapeutic efficacy.

Species differences in auditory processing dynamics in songbird auditory telencephalon

The caudomedial nidopallium (NCM) of songbirds is a telencephalic area involved in the auditory processing and memorization of complex vocal communication signals. We used pure tone stimuli and multiunit electrophysiological recordings in awake birds to investigate whether the basic properties of song-responding circuits in NCM differ between canaries and zebra finches, two species whose songs are markedly different in their spectral and temporal organization. We found that the responses in zebra finch NCM are characterized by broad tuning and sustained responses that may facilitate the integration of zebra finch song syllables and call notes that are of long duration and have a broad harmonic structure. In contrast, we found that the responses in canary NCM show narrower tuning and less sustained responses over the time periods analyzed. These characteristics may contribute to enhanced processing of the narrow-band whistles, rapid trills, and steep frequency modulations that are prominent features of canary song. These species differences are much less pronounced in field L2, the direct thalamorecipient region that represents a preceding station in the central avian auditory pathway. NCM responses did not differ across sexes of either species, but field L2 did show wider tuning in zebra finch females relative to males. In sum, species differences in the response properties of NCM likely reflect selectivity for the acoustic elements of each species' vocal repertoire.

Organization of the somatosensory cortex in elephant shrews (E. edwardii)

The superorder Afrotheria consists of a diverse group of mammals, including elephants, hyraxes, dugongs, sea cows, aardvarks, tenrecs, golden moles, and elephant shrews. Recent studies suggest this clade diverged from other placental mammals 100 million years ago and thus may represent the sister group to the remaining placental mammals. Despite this important taxonomic position, relatively few studies have investigated cortical organization in these species. Here we present results of an investigation of the somatosensory cortex in the Cape elephant shrew (Elephantulus edwardii). Using multiunit electrophysiological recording techniques, we identified a topographic map of the elephant shrew's body in a location and orientation consistent with the primary somatosensory cortex (S1). The elephant shrew's elongated snout, extensive facial vibrissae, and long tongue accounted for a large portion of the somatosensory representation, located in a relatively rostral area of cortex. Evidence for an additional somatosensory area, presumed to be secondary somatosensory cortex (S2), was found just lateral to S1. Visual and auditory responsive areas were also identified and the extent of visual cortex appeared to be quite large in these highly visual mammals. Despite the elephant shrew's exceptionally well-developed eyes, ears, and vibrissae, there were no anatomical correlates to sensory areas, or body part representations (e.g., barrels), that could be identified in the flatted cortex.

The Organization and Connections of Anterior and Posterior Parietal Cortex in Titi Monkeys: Do New World Monkeys Have an Area 2?

We used multiunit electrophysiological recording techniques to examine the topographic organization of somatosensory area 3b and cortex posterior to area 3b, including area 1 and the presumptive area 5, in the New World titi monkey, Callicebus moloch. We also examined the ipsilateral and contralateral connections of these fields, as well as those in a region of cortex that appeared to be similar to both area 7b and the anterior intraparietal area (7b/AIP) described in macaque monkeys. All data were combined with architectonic analysis to generate comprehensive reconstructions. These studies led to several observations. First, area 1 in titi monkeys is not as precisely organized in terms of topographic order and receptive field size as is area 1 in macaque monkeys and a few New World monkeys. Second, cortex caudal to area 1 in titi monkeys is dominated by the representation of the hand and forelimb, and contains neurons that are often responsive to visual stimulation as well as somatic stimulation. This organization is more like area 5 described in macaque monkeys than like area 2. Third, ipsilateral and contralateral cortical connections become more broadly distributed away from area 3b towards the posterior parietal cortex. Specifically, area 3b has a relatively restricted pattern of connectivity with adjacent somatosensory fields 3a, 1, S2 and PV; area 1 has more broadly distributed connections than area 3b; and the presumptive areas 5 and 7b/AIP have highly diverse connections, including connections with motor and premotor cortex, extrastriate visual areas, auditory areas and somatosensory areas of the lateral sulcus. Fourth, the hand representation of the presumptive area 5 has dense callosal connections. Our results, together with previous studies in other primates, suggest that anterior parietal cortex has expanded in some primate lineages, perhaps in relation to manual abilities, and that the region of cortex we term area 5 is involved in integrating somatic inputs with the motor system and across hemispheres. Such connections could form the substrate for intentional reaching, grasping and intermanual transfer of information necessary for bilateral coordination of the hands.

Interneurone bursts are spontaneously associated with muscle contractions only during early phases of mouse spinal network development: a study in organotypic cultures

For a short time during development immature circuits in the spinal cord and other parts of the central nervous system spontaneously generate synchronous patterns of rhythmic activity. In the case of the spinal cord, it is still unclear how strongly synchronized bursts generated by interneurones are associated with motoneurone firing and whether the progressive decline in spontaneous bursting during circuit maturation proceeds in parallel for motoneurone and interneurone networks. We used organotypic cocultures of spinal cord and skeletal muscle in order to investigate the ontogenic evolution of endogenous spinal network activity associated with the generation of coordinate muscle fibre contractions. A combination of multiunit electrophysiological recordings, videomicroscopy and optical flow computation allowed us to measure the correlation between interneurone firing and motoneurone outputs after 1, 2 and 3 weeks of in vitro development. We found that, in spinal organotypic slices, there is a developmental switch of spontaneous activity from stable bursting to random patterns after the first week in culture. Conversely, bursting recorded in the presence of strychnine and bicuculline became increasingly regular with time in vitro. The time course of spontaneous activity maturation in organotypic slices is similar to that previously reported for the spinal cord developing in utero. We also demonstrated that spontaneous bursts of interneurone action potentials strongly correlate with muscular contractions only during the first week in vitro and that this is due to the activation of motoneurones via AMPA-type glutamate receptors. These results indicate the occurrence in vitro of motor network development regulating bursting inputs from interneurones to motoneurones.

Organization of area 3a in macaque monkeys: contributions to the cortical phenotype.

The detailed organization of somatosensory area 3a was examined in macaque monkeys using multiunit electrophysiological recording techniques. By examining topographic relationships, changes in receptive field size, and the type of stimulus that neurons responded to, functional boundaries of area 3a were determined and related to architectonic boundaries. One striking observation was that the location of area 3a varied with respect to the central sulcus. In one-half of the cases area 3a was on the rostral bank and fundus of the central sulcus and in the other half of the cases it was on the caudal bank and fundus of the central sulcus. In terms of topographic organization, we found that area 3a contains a complete representation of deep receptors and musculature of the contralateral body, and that the general organization of body part representations mirrors that of the primary somatosensory area, 3b. These results as well as results from studies of area 3a in ours and other laboratories indicate that area 3a is part of a network involved in proprioception, postural control, and the generation of coordinated movements. Further, comparative analysis of area 3a in a variety of species suggests that its construction is based, to a large extent, on the use of a particular body part rather than on innervation density.

Characterisation of multiple physiological fields within the anatomical core of rat auditory cortex

The organisation and response properties of the rat auditory cortex were investigated with single and multi-unit electrophysiological recording. Two tonotopically organised ‘core’ fields, i.e. the primary (A1) and anterior (AAF) auditory fields, as well as three non-tonotopically organised ‘belt’ fields, i.e. the posterodorsal (PDB), dorsal (DB) and anterodorsal (ADB) belt fields, were identified. Compared to neurones in A1, units in AAF exhibited broader frequency tuning, as well as shorter minimum, modal and mean first spike latencies. In addition, units in AAF showed significantly higher thresholds and best SPLs, as well as broader dynamic ranges. Units in PDB, DB and ADB were characterised by strong responses to white noise and showed either poor or no responses to pure tones. The differences in response properties found between the core and belt fields may reflect a functional specificity in processing different features of auditory stimuli. The present study also combined microelectrode mapping with Nissl staining to determine if the physiological differences between A1 and AAF corresponded to cytoarchitectonically defined borders. Both A1 and AAF were located within temporal cortex 1 (Te1), with AAF occupying an anteroventral subdivision of Te1, indicating that the two neighbouring, physiologically distinct fields are cytoarchitectonically homogeneous.

Somatotopic reorganization in the brainstem and thalamus following peripheral nerve injury in adult primates

Injury-induced reorganization of central somatotopic maps is a phenomenon that has proven to be useful for elucidating the mechanisms and time course of neural plasticity. To date, the overwhelming majority of this line of research has focused on such plastic events in cortical areas, at the expense of subcortical structures. In this study, we used multi-unit electrophysiological recording techniques to assess the somatotopic organization of brainstem and thalamic areas following chronic survival from paired median and ulnar nerve section in adult squirrel monkeys. We report that the extent of cutaneously-driven reorganization in both the cuneate nucleus of the brainstem and the ventroposterior lateral nucleus of the thalamus is comparable to that previously documented for area 3b of cortex. These observations are consistent with those previously reported in thalamus, and are unique for brainstem.