Search

Tree shrews are small squirrel-like mammals that are the closest living relative to primates available for detailed neurobiological study. In a recent study (Remple et al. [2006] J. Comp. Neurol. 497:133-154), we provided anatomical and electrophysiological evidence that the frontoparietal cortex of tree shrews has two motor fields (M1 and M2) and five somatosensory fields (3a, 3b, S2, somatosensory caudal area [SC], and parietal ventral area [PV]). In the present study, we injected anatomical tracers into M1, M2, 3a, 3b, SC, and posterior parietal cortex to establish the ipsilateral cortical connections of these areas. The results provide evidence for a number of new cortical areas including medial motor and somatosensory areas (MMA and MSA), three posterior parietal areas (PPd, PPv, and PPc), and an area ventral to temporal inferior cortex (TIV). Ml receives topographic projections from M2, MMA, 3a, and PPv, and nontopographic connections from the temporal anterior and dorsal areas (TA and TD), PPc, TIV, and MSA. The connections of M2 are similar to those of M1, except that M2 receives denser projections from TIV, PPc, and dorsal frontal cortex and sparser input from M1. Areas 3a, 3b, and SC receive dense topographic projections from each other, S2, and PV and sparser connections from PPd and PPv. Area 3a receives additional input from posterior parietal and temporal regions and from M1 and MMA. Overall, the frontoparietal connections of tree shrew cortex are most similar to those of prosimian primates and quite different from those of more distant relatives such as rats.

Dorsal visual cortical areas are thought to be dominated by input from the magnocellular (M) visual pathway, with little or no parvocellular (P) contribution. These relationships are supported by a close correlation between the functional properties of these areas and the M pathway, and by a lack of anatomical evidence for P input. Here we use rabies virus as a retrograde transynaptic tracer to show that the dorsal area MT receives strong input, via a single relay, from both M and P cells of the lateral geniculate nucleus. This surprising P input, likely relayed via layer 6 Meynert cells in primary visual cortex, can provide MT with sensitivity to a more complete range of spatial, temporal and chromatic cues than the M pathway alone. These observations provide the first definitive evidence for P pathway input to MT and show that convergence of parallel visual pathways occurs in the dorsal stream.

Tree shrews are highly visual mammals closely related to primates. They have a large visual pulvinar complex, but its organization and relation to visual cortex is only partly known. We processed brain sections through the pulvinar with seven different procedures in an effort to reveal histologically distinct compartments. The results revealed three major subdivisions. A dorsal subdivision, Pd, stains darkly for acetylcholinesterase (AChE) and occupies the dorsoposterior one-third of the pulvinar complex. A ventral subdivision, Pv, stains darkly when processed with the Cat-301 antibody and occupies the ventroanterior fifth of the pulvinar complex along the brachium of the superior colliculus. Unexpectedly, part of Pv is ventral to the brachium. A large central subdivision, Pc, stains moderately dark for AChE and cytochrome oxidase (CO), and very light for Cat-301. Pc includes about half of the pulvinar complex, with parts on both sides of the brachium of the superior colliculus. These architectonic results demonstrate that the pulvinar complex of tree shrews is larger and has more subdivisions than previously described. The complex resembles the pulvinar of primates by having a portion ventral to the brachium and by having histochemically distinct nuclei; the number of nuclei is less than in primates, however.

The retinotopic organization of the middle temporal visual area (MT) was determined in six adult owl monkeys and one adult marmoset 69 d to 10 months after lesions of the dorsolateral primary visual cortex (V1). The lesions removed were limited to extensive parts of the representation of the lower visual quadrant in V1. Microelectrodes were used to record from neurons at numerous sites in MT to determine whether parts of MT normally devoted to the lower visual quadrant (1) were unresponsive to visual stimuli, (2) acquired responsiveness to inputs from intact portions of V1, or (3) became responsive to some other visually driven input such as a relay from the superior colliculus via the pulvinar to MT. All monkeys (n = 6) with moderate to moderately large lesions had unresponsive portions of MT even after 10 months of recovery. These unresponsive regions were retinotopically equivalent to the removed parts of V1 in normal animals. Thus, there was no evidence for an alternative source of activation. In addition, these results indicate that any retinotopic reorganization of MT based on inputs from intact portions of V1 was not extensive, yet neurons near the margins of responsive cortex may have acquired new receptive fields, and the smallest 5° lesion of V1 failed to produce an unresponsive zone. Deprived portions of MT were not remarkably changed in histological appearance in cytochrome oxidase, Nissl, andWisteria floribundaagglutinin preparations. Nevertheless, some reduction in myelin staining and other histological changes were suggested. We conclude that MT is highly dependent on V1 for activation in these monkeys, and alternative sources do not become effective over months when normal activation is absent. Additionally, remaining V1 inputs have only a limited capacity to expand their activation territory into deprived portions of MT.

Optical imaging of intrinsic responses to visual stimuli in extrastriate cortex of owl monkeys provided evidence for the dorsal half of the third visual area, V3. Visual stimuli were used to selectively activate locations in dorsolateral V2 and the rostrally adjoining presumptive V3. Consistent with the proposed retinotopies of dorsal V2 and dorsal V3, small bars of drifting gratings along the horizontal meridian of the contralateral hemifield activated cortex along the V2/V3 border, whereas such stimuli along the vertical meridian activated cortex along the rostral border of V3. Stimuli in limited locations in the lower visual quadrant revealed mirror reversals of retinotopy in dorsal V2 and V3, whereas stimuli in the upper visual quadrant failed to activate either region. Brain sections processed for cytochrome oxidase from the same cases provided architectonic borders of V2 that matched those indicated by the optical imaging. The results support the concept that a narrow dorsal V3 exists in monkeys. V3d borders dorsal V2 and contains a smaller, mirror-image representation of the lower visual quadrant.

Previously we described patterns of connections that support the concept of V3 in small New World marmoset monkeys, three species of larger New World monkeys, and two species of Old World macaque monkeys. Here we describe a pattern of V1 connections with extrastriate visual cortex in Galago garnetti (also known as Otolemur garnetti) that demonstrates the existence of a V3 in a strepsirhine (prosimian) primate. Injections of fluorochromes or cholera toxin subunit-B (CTB) in V1 labeled cells and terminals in retinotopically matched regions in V2, V3, DL (V4), and MT. Labeled axon terminations were more focused primarily in middle layers of cortex, likely representing 'feedforward' input from V1, whereas labeled cells were more widespread and found in both superficial and deeper cortical layers, indicative of feedback projections. Averaged across injections, V3 had the third largest percentage of labeled cells (11%), following only V2 (47%) and the middle temporal area (MT; 19%). The dorsolateral area (DL, or V4; 9%) also contained a relatively large number of retrogradely labeled cells. These results indicate that V2, V3, DL (V4), and MT are retinotopically connected with V1, and provide major sources of feedback. Other extrastriate areas were less densely connected to V1, and there was no clear indication of labeled terminals. Inferotemporal cortex (IT) provided nearly 7% of feedback connections, whereas the dorsomedial area (DM) contributed about 3%. The remaining areas that have been proposed for galago extrastriate cortex, MTc, MST, FST, LPP and VPP, each accounted for about 1% or less of the total number of labeled cells. Thus, six extrastriate areas, V2, MT, V3, DL (V4), IT, and DM provide over 96% of visual cortex projections to V1. These areas also provide most of the projections to V1 in New and Old World monkeys.

Microelectrode mapping methods were used to define the parietal ventral somatosensory area (PV) on the upper bank of the lateral sulcus in five marmosets (Callithrix jacchus). In the same animals, neuroanatomical tracers were placed into electrophysiologically identified sites in PV and/or the second somatosensory area (S2). Foci of anterograde and retrograde label were related to electrophysiological maps of cortical areas and cortical and thalamic architecture. The results lead to the following conclusions: (1) Multiunit recordings from cortex on the upper bank of the lateral sulcus demonstrate that PV is somatotopically organized, with the face representation adjoining area 3b and the hindlimb and tail representations away from this border in cortex deep on the upper bank of the lateral sulcus. The forelimb representation is caudal in PV adjacent to the S2 forelimb representation. The body surface representation in PV approximates a mirror image of that in S2; (2) Areas PV and S2 are less myelinated and have less cytochrome oxidase enzyme activity than area 3b; (3) The ventroposterior inferior nucleus (VPI) of the thalamus provides the major somatosensory projections to PV. PV is reciprocally connected with VPI and anterior pulvinar; (4) PV has ipsilateral cortical connections with areas 3a, 3b, 1, and M1 and higher order somatosensory fields, and at least most of these connections are somatotopically matched; and (5) Callosal connections of PV are with S2 and PV of the other cerebral hemisphere. These results further establish PV as one of at least four somatosensory areas of the lateral sulcus of primates.

After years of experimentation and substantial progress, there is still only limited agreement on how visual cortex in primates is orcognized, and what features of this organization are variable or stable across lines of primate phylogeny. Only three visual areas, V1, V2, and MT, are widely recognized as common to all primates, although there are certainly more. Here we consider various concepts of how the cortex along the outer border of V2 is organized. An early proposal was that this region is occupied by a V3 that is as wide and as long as V2, and represents the visual hemifield as a mirror image of V2. We refer to this notion as the classical V3 or V3-C. Another proposal is that only the dorsal half of V3-C exists, the half representing the lower visual quadrant, and thus the representation is incomplete (V3-I) by half. A version of this proposal is that V3-I is discontinuous, extremely thin in places, and highly variable across individuals, much as a vestigial or degenerate structure might be (V3-IF-incomplete and fragmented). A fourth proposal is that there is no V3. Many results suggest that a series of visual areas border V2, none of which has the characteristics of V3. Alternatively, the possibility exists that primate taxa differ with regard to visual areas bordering V2. Currently, much of the supporting evidence for a classical V3 comes from fMRI studies in humans, much of the evidence for a series of bordering areas comes from New World Monkeys and prosimian galagos, and much of the evidence for a V3-I or V3-IF comes from macaque monkeys. Possibly all these interpretations of visual cortex organization are valid, but each for only one of the major groups of primate evolution. Here, we suggest that none of these interpretations is correct, and propose instead that a modified V3 (V3-M) exists in a similar form in all primates. This V3-M is smaller and thinner than V3-C, discontinuous in the middle, but with comparable dorsal and ventral halves representing the lower and upper visual hemifields, respectively. Because the evidence for V3-M is limited, and it stems in part from our ongoing but incomplete comparative studies of V1 connections in primates, this suggestion requires further experimental evaluation and it remains tentative.

SummaryThere has never been a wholesale way of identifying neurons that are monosynaptically connected either to some other cell group or, especially, to a single cell. The best available tools, transsynaptic tracers, are unable to distinguish weak direct connections from strong indirect ones. Furthermore, no tracer has proven potent enough to label any connected neurons whatsoever when starting from a single cell. Here we present a transsynaptic tracer that crosses only one synaptic step, unambiguously identifying cells directly presynaptic to the starting population. Based on rabies virus, it is genetically targetable, allows high-level expression of any gene of interest in the synaptically coupled neurons, and robustly labels connections made to single cells. This technology should enable a far more detailed understanding of neural connectivity than has previously been possible.

Eight patients with long standing intermittent abdominal pain were found to have solitary cysts in the head of the pancreas. These cysts were demonstrated by retrograde pancreatography but could not be either visualized or palpated at operation. In only two cases was there sufficient clinical evidence to make a diagnosis of pancreatitis; in the remaining six the demonstration of a cyst was the first evidence of any abnormality that could account for their symptoms. All patients were treated by sphincteroplasty. Biopsy of the ampulla revealed fibrosis or inflammation in four of the seven patients in whom biopsy was performed. There was one postoperative death, and there has been little discernible relief in the case associated with alcoholism. The remaining six patients have had symptomatic relief for one to five years.