Functional and histological properties of caudal intraparietal area of macaque monkey

In our previous studies, we found that cells in the caudal intraparietal (CIP) area of the macaque monkey selectively responded to three-dimensional (3D) features, such as the axis and surface orientations, and we suggested that this area played a crucial role in 3D vision. In this study, we investigated (1) whether cells in CIP respond to other 3D features, such as curvature, and (2) whether CIP has any histological property to distinguish it from neighboring areas. Curvatures defined by a random-dot stereogram were presented on a display while the monkey performed a fixation task. The shape and amount of curvature were manipulated by two independent variables, shape index and curvedness, respectively. Two-way ANOVA showed that 19 out of 56 visually responsive cells (34.0%) showed the main effect of shape index. We tentatively designated these cells as 3D curvature-selective (3DCS). Of these, six 3DCS cells showed the main effects of shape index and curvedness, whereas 13 showed the main effect of shape index only. In both types of 3DCS cells, preferred shape indices calculated from tuning curves at two levels of curvedness matched well. These results indicate that the majority of 3DCS cells responded equally to a particular shape of curvatures with different curvedness levels. An immunohistochemical study showed that the recording sites of 3DCS cells were in a cortical region characterized by a dense SMI-32 immunoreactivity in the caudal portion of the lateral intraparietal sulcus (IPS), which suggests that this region is comparable to the lateral occipital parietal (LOP) designated in the caudal IPS previously. Further investigations showed that this region was separated from LIPv, the ventral subdivision of lateral intraparietal (LIP) located rostral to CIP/LOP. These results suggest that CIP is a cortical area distinct from LIP histologically as well as functionally.     

国人脑顶内沟的三维形态及其不对称性

目的 以MRI及三维可视化方法探索国人脑顶内沟的类型及形态特征,以期为端脑顶叶功能研究及相关疾病诊治提供解剖学依据。 方法 选取107例正常右利手中国人,应用3.0T GE磁共振扫描仪进行扫描,获取高分辨率脑MRI数据。利用BrainVISA软件建立顶内沟的三维解剖模型,并对顶内沟的形态类型进行观察分析。结果 顶内沟有3种分段模式：连续型、两段型和三段型。不同分段之间的分隔脑回或表面可见,或埋藏于脑沟内部。左脑以连续型为主（56.07%）,右脑以两段型为主（58.49%）,且左右脑顶内沟形态具有显著不对称性（P＜0.01）。男性大脑顶内沟以两段型为主(53.17%),女性大脑顶内沟以连续型为主(49.43%),但男女性顶内沟形态无明显性别差异（P＞0.05）。 结论 顶内沟形态复杂,具有3种形态类型,并有明显的侧别差异。     

A functional magnetic resonance imaging navigated repetitive transcranial magnetic stimulation study of the posterior parietal cortex in normal pain and hyperalgesia

Noxious stimuli activate a complex cerebral network. During central sensitization to pain, activity in most of these areas is changed. One of these areas is the posterior parietal cortex (PPC). The role of the PPC during processing of acute pain as well as hyperalgesia and tactile allodynia remains elusive. Therefore, we performed a functional magnetic resonance imaging (fMRI) based, neuro-navigated, repetitive transcranial magnetic stimulation (rTMS) study in 10 healthy volunteers. Firstly, pin-prick hyperalgesia was provoked on the right volar forearm, using the model of electrically-induced secondary mechanical hyperalgesia. fMRI was performed during pin-prick stimulation inside and outside the hyperalgesic areas. Secondly, on four different experimental sessions, the left and right individual intraparietal BOLD peak-activations were used as targets for a sham-controlled 1 Hz rTMS paradigm of 10 min duration. We measured psychophysically the (i) electrical pain stimulus intensity on an 11-point numeric pain rating scale (NRS, 0–10), the (ii) area of hyperalgesia, and the (iii) area of dynamic mechanical allodynia. Sham stimulation or rTMS was performed 16 min after induction of pin-prick hyperalgesia and tactile allodynia. Compared to sham stimulation, no significant effect of rTMS was observed on pain stimulus intensity and the area of allodynia. However, a reduction of the hyperalgesic area was observed for rTMS of the left PPC (P<0.05). We discuss the role of the PPC in central sensitization to pain, in spatial discrimination of pain stimuli and in spatial-attention to pain stimuli.     

The role of left and right intraparietal sulcus in the attentional blink: a transcranial magnetic stimulation study

Processing of one visual target (T1) makes it difficult to become aware of a second target (T2), when two targets, embedded in a stream of distractor stimuli, occur within about 500 ms. This phenomenon is known as attentional blink (AB) and reflects the temporal limitation in allocating visual attention. Although several studies suggest that parietal regions are concerned with the AB phenomenon, their functional relevance remains unclear. We investigated whether left and/or right intraparietal sulcus (IPS) contributed to the AB bottleneck using transcranial magnetic stimulation (TMS). The course of recovery from the AB deficit was facilitated when single pulse TMS induced a transient interruption of left or right IPS activity at a T1-TMS stimulus onset asynchrony of 350 ms, while there was no effect of TMS or sham stimulation delivered over Cz with the same timing. These results provide direct evidence that activation of left as well as right IPS is involved in the genesis of AB. This finding supports the idea that the IPS plays a critical role in the cortical network controlling the temporal dynamics of visual awareness.     

The functional neuroanatomy of classic delayed response tasks in humans and the limitations of cross-method convergence in prefrontal function

Three classic delay tasks: spatial delayed response, delayed spatial alternation and delayed object-alternation are prototypical experimental paradigms for mapping the functional neuroanatomy of prefrontal cortex in animals. These tasks have been applied in human lesion studies, yet there have been very few studies investigating their functional neuroanatomy in healthy human subjects. We used functional magnetic resonance imaging to investigate the functional neuroanatomy of these classic paradigms (and a fourth: object delayed response) in a single sample of healthy human participants. Consistent with previous animal, human lesion, and functional neuroimaging studies, activity was observed in prefrontal and posterior parietal cortices across all three delay tasks. Task-specific activations, however, were not entirely consistent with predictions drawn from animal lesion studies. For example, delayed object-alternation activated dorsolateral prefrontal cortex, a region not generally implicated in animal lesion reports. Spatial delayed response, classically associated with the dorsolateral prefrontal cortex, did not activate this region; it rather activated posterior premotor cortices involved in response preparation, as did spatial alternation. All three tasks activated the frontopolar cortex, a region not considered crucial in animal research but associated with manipulation of internally generated information in recent human research. While cross-method convergence may be attained for lower level perceptual or motor tasks, the results of this study caution against the assumption that lesion-specific effects in animals generalize to human prefrontal cortex function.     

The brain and the braincase: a spatial analysis on the midsagittal profile in adult humans

The spatial relationships between brain and braincase represent a major topic in surgery and evolutionary neuroanatomy. In paleoneurology, neurocranial landmarks are often used as references for brain areas. In this study, we analyze the variation and covariation of midsagittal brain and skull coordinates in a sample of adult modern humans in order to demonstrate spatial associations between hard and soft tissues. The correlation between parietal lobe size and parietal bone size is very low, and there is a marked individual variation. The distances between lobes and bones are partially influenced by the dimensions of the parietal lobes. The main pattern of morphological variability among individuals, associated with the size of the precuneus, apparently does not influence the position of the neurocranial sutures. Therefore, variations in precuneal size modify the distance between the paracentral lobule and bregma, and between the parietal lobe and lambda. Hence, the relative position of the cranial and cerebral landmarks can change as a function of the parietal dimensions. The slight correlation and covariation among these elements suggests a limited degree of spatial integration between soft and hard tissues. Therefore, although the brain influences the cranial size and shape during morphogenesis, the specific position of the cerebral components is sensitive to multiple effects and local factors, without a strict correspondence with the bone landmarks. This absence of correspondent change between brain and skull boundaries suggests caution when making inferences about the brain areas from the position of the cranial sutures. The fact that spatial relationships between cranial and brain areas may vary according to brain proportions must be considered in paleoneurology, when brain anatomy is inferred from cranial evidence.     

Relationship between the characteristics of visual short-term memory in monkeys and the spatial properties of images

Experiments were performed on Rhesus macaques to study the relationship between delayed visual differentiation processes and stimulus properties. These investigations showed that the processes of short-term storage of visual information in monkeys has significant features associated with differences in stimulus properties. These consisted of different durations of storage and motor response times. Because of these differences, stimuli (15 pairs) could be grouped into compact clusters on the bases of similarity between their delayed differentiation characteristics. These experiments characterized the processes of short-term information storage during the differentiation of stimuli differing in terms of spatial relationships between elements, as compared with stimuli differing in terms of other attributes (shape, color, etc.); spatial information was stored for shorter periods of time and motor response times were longer. It is suggested that visual short-term memory involves a set of mechanisms operating on attributes of different types and which, along with signs and working programs associated with the visual system, stores spatial discriminatory signs, in which the major role is played by visual-vestibular interactions. Cognitive Processes Modeling Group, Laboratory of Vestibular Apparatus Physiology, I. P. Pavlov Institute of Physiology, 6 Makarov Bank, St. Petersburg 199034, Russia. Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 83, No. 9, pp. 51–59, September, 1997.     

The functional and anatomical organization of marsupial neocortex: evidence for parallel evolution across mammals

Marsupials are a diverse group of mammals that occupy a large range of habitats and have evolved a wide array of unique adaptations. Although they are as diverse as placental mammals, our understanding of marsupial brain organization is more limited. Like placental mammals, marsupials have striking similarities in neocortical organization, such as a constellation of cortical fields including S1, S2, V1, V2, and A1, that are functionally, architectonically, and connectionally distinct. In this review, we describe the general lifestyle and morphological characteristics of all marsupials and the organization of somatosensory, motor, visual, and auditory cortex. For each sensory system, we compare the functional organization and the corticocortical and thalamocortical connections of the neocortex across species. Differences between placental and marsupial species are discussed and the theories on neocortical evolution that have been derived from studying marsupials, particularly the idea of a sensorimotor amalgam, are evaluated. Overall, marsupials inhabit a variety of niches and assume many different lifestyles. For example, marsupials occupy terrestrial, arboreal, burrowing, and aquatic environments; some animals are highly social while others are solitary; different species are carnivorous, herbivorous, or omnivorous. For each of these adaptations, marsupials have evolved an array of morphological, behavioral, and cortical specializations that are strikingly similar to those observed in placental mammals occupying similar habitats, which indicate that there are constraints imposed on evolving nervous systems that result in recurrent solutions to similar environmental challenges.     

Thalamocortical and the dual pattern of corticothalamic projections of the posterior parietal cortex in macaque monkeys

The corticothalamic projection includes a main, modulatory projection from cortical layer VI terminating with small endings whereas a less numerous, driving projection from layer V forms giant endings. Such dual pattern of corticothalamic projections is well established in rodents and cats for many cortical areas. In non-human primates (monkeys), it has been reported for the primary sensory cortices (A1, V1, S1), the motor and premotor cortical areas and, in the parietal lobe, also for area 7. The present study aimed first at refining the cytoarchitecture parcellation of area 5 into the sub-areas PE and PEa and, second, establishing whether area 5 also exhibits this dual pattern of corticothalamic projection and what is its precise topography. To this aim, the tracer biotinylated dextran amine (BDA) was injected in area PE in one monkey and in area PEa in a second monkey. Area PE sends a major projection terminating with small endings to the thalamic lateral posterior nucleus (LP), ventral posterior lateral nucleus (VPL), medial pulvinar (PuM) and, but fewer, to ventral lateral posterior nucleus, dorsal division (VLpd), central lateral nucleus (CL) and center median nucleus (CM), whereas giant endings formed restricted terminal fields in LP, VPL and PuM. For area PEa, the corticothalamic projection formed by small endings was found mainly in LP, VPL, anterior pulvinar (PuA), lateral pulvinar (PuL), PuM and, to a lesser extent, in ventral posterior inferior nucleus (VPI), CL, mediodorsal nucleus (MD) and CM. Giant endings originating from area PEa formed restricted terminal fields in LP, VPL, PuA, PuM, MD and PuL. Furthermore, the origin of the thalamocortical projections to areas PE and PEa was established, exhibiting clusters of neurons in the same thalamic nuclei as above, in other words predominantly in the caudal thalamus. Via the giant endings CT projection, areas PE and PEa may send feedforward, transthalamic projections to remote cortical areas in the parietal, temporal and frontal lobes contributing to polysensory and sensorimotor integration, relevant for visual guidance of reaching movements for instance.     

The functional changes of the circadian system organization in aging

The circadian clock drives periodic oscillations at different levels of an organism from genes to behavior. This timing system is highly conserved across species from insects to mammals and human beings. The question of how the circadian clock is involved in the aging process continues to attract more attention. We aim to characterize the detrimental impact of aging on the circadian clock organization. We review studies on different components of the circadian clock at the central and periperal levels, and their changes in aged rodents and humans, and the fruit fly Drosophila. Intracellular signaling, cellular activity and intercellular coupling in the central pacemaker have been found to decline with advancing age. Evidence of degradation of the molecular clockwork reflected by clock gene expression in both central and peripheral oscillators due to aging is inadequate. The findings on age-associated molecular and functional changes of peripheral clocks are mixed. We conclude that aging can affect the circadian clock organization at various levels, and the impairment of the central network may be a fundamental mechanism of circadian disruption seen in aged species.     

Functional organization of the insula in men and women with obstructive sleep apnea during Valsalva

Study ObjectivesObstructive sleep apnea (OSA) patients show impaired autonomic regulation, perhaps related to functional reorganization of the insula, which in healthy individuals shows sex-specific anterior and right dominance during sympathetic activation. We examined insular organization of responses to a Valsalva maneuver in OSA with functional magnetic resonance imaging (fMRI).MethodsWe studied 43 newly diagnosed OSA (age mean ± SD: 46.8 ± 8.7 years; apnea-hypopnea index (AHI) ± SD: 32.1 ± 20.1 events/hour; 34 males) and 63 healthy (47.2 ± 8.8 years; 40 males) participants. Participants performed four 18-second Valsalva maneuvers (1-minute intervals, pressure ≥ 30 mmHg) during scanning. fMRI time trends from five insular gyri—anterior short (ASG); mid short (MSG); posterior short (PSG); anterior long (ALG); and posterior long (PLG)—were assessed for within-group responses and between-group differences with repeated measures ANOVA (p < 0.05); age and resting heart rate (HR) influences were also assessed.ResultsRight and anterior fMRI signal dominance appeared in OSA and controls, with no between-group differences. Separation by sex revealed group differences. Left ASG anterior signal dominance was lower in OSA versus control males. Left ASG and ALG anterior dominance was higher in OSA versus control females. In all right gyri, only OSA females showed greater anterior dominance than controls. Right dominance was apparent in PSG and ALG in all groups; females showed right dominance in MSG and PLG. OSA males did not show PLG right dominance. Responses were influenced substantially by HR but modestly by age.ConclusionsAnterior and right insular fMRI dominance appears similar in OSA versus control participants during the sympathetic phase of the Valsalva maneuver. OSA and control similarities were present in just males, but not necessarily females, which may reflect sex-specific neural injury.     

Directional asymmetry in vertical smooth-pursuit and cancellation of the vertical vestibulo-ocular reflex in juvenile monkeys

Young primates exhibit asymmetric eye movements during vertical smooth-pursuit across a textured background such that upward pursuit has low velocity and requires many catch-up saccades. The asymmetric eye movements cannot be explained by the un-suppressed optokinetic reflex resulting from background visual motion across the retina during pursuit, suggesting that the asymmetry reflects most probably, a low gain in upward eye commands (Kasahara et al. in Exp Brain Res 171:306–321, 2006). In this study, we examined (1) whether there are intrinsic differences in the upward and downward pursuit capabilities and (2) how the difficulty in upward pursuit is correlated with the ability of vertical VOR cancellation. Three juvenile macaques that had initially been trained only for horizontal (but not vertical) pursuit were trained for sinusoidal pursuit in the absence of a textured background. In 2 of the 3 macaques, there was a clear asymmetry between upward and downward pursuit gains and in the time course of initial gain increase. In the third macaque, downward pursuit gain was also low. It did not show consistent asymmetry during the initial 2 weeks of training. However, it also exhibited a significant asymmetry after 4 months of training, similar to the other two monkeys. After 6 months of training, these two monkeys (but not the third) still exhibited asymmetry. As target frequency increased in these two monkeys, mean upward eye velocity saturated at ∼15°/s, whereas horizontal and downward eye velocity increased up to ∼40°/s. During cancellation of the VOR induced by upward whole body rotation, downward eye velocity of the residual VOR increased as the stimulus frequency increased. Gain of the residual VOR during upward rotation was significantly higher than that during horizontal and downward rotation. The time course of residual VOR induced by vertical whole body step-rotation during VOR cancellation was predicted by addition of eye velocity during pursuit and VOR x1. These results support our view that the directional asymmetry reflects the difference in the organization of the cerebellar floccular region for upward and downward directions and the preeminent role of pursuit in VOR cancellation.     

Relationship between learning characteristics and the properties of visual objects in rhesus macaques with bilateral removal of parietal cortex field 7

Behavioral experiments were used in rhesus macaques with bilateral excision of field 7 of the lower parietal cortex to study the relationship between visual differentiation learning processes and a variety of stimulus properties. All animals showed significant differences associated with stimulus properties, which produced different types of learning curves. For each monkey, visual stimuli were divided into compact groups in terms of the “similarity” of their learning characteristics. Removal of field 7 had no effect on the process of learning visual image discrimination when this was based on properties such as color and geometrical shape, but worsened the learning characteristics when visual differentiation was based on spatial information, when the learning process became unstable, with increases in the numbers of peaks and troughs on the learning curve and a significant increase in the duration of the learning period. The time to a stable motor response also became significantly greater than for visual images distinguished by shape and color. It is suggested that during the process of learning visual discrimination, processing and extraction of image signs by the visual system for objects characterized by spatial relationships is accompanied by the formation of spatial distinguishing signs, this process involving neuronal structures in field 7 of the lower parietal cortex, which appears to be the main area determining visual-vestibular interactions. Increases in oscillations and in the difficulty of the learning process for differentiation on the basis of spatial information aftr removal of field 7 might be due to a transfer from one strategy to another, resulting from disruption of the mechanisms which evaluate body image and egocentric orientation on the basis of visual-vestibular interactions. Translated from Rossiiskii Fiziologicheskii Zhurmal imeni I. M. Sechenova, Vol. 84, No. 3 pp. 145–156, March, 1998     

Synchronization processes in the mechanisms of short-term memory in monkeys: The involvement of cholinergic and glutaminergic cortical structures

Behavioral experiments were carried out in which monkeys had to solve a task involving delayed visual discrimination, and activity was simultaneously recorded from several neurons of the visual, prefrontal, and lower temporal regions of the cortex before and after modification of cholinergic (by systemic infusion of the M-cholinoceptor blocker amizil) and glutaminergic (by intracortical perfusion with glutaminergic agonists and antagonists, i.e., NMDA, aminophosphonovalerianic acid (APV) and aminophosphonobutyric acid (APB)) systems. Amizil and APV reduced the duration of short-term information retention and increased the delay before the motor response was made. Worsening of these parameters was accompanied by a significant level of desynchronization of activity in the groups of neurons studied. NMDA and APB improved short-term memory and increased neuron synchronization. The role of synchronization of information processes in the mechanisms of short-term memory and the involvement of the cholinergic and glutaminergic systems are discussed. Laboratory for the Regulation of Brain Neuron Function (M. O. Samoilov, Director), I. P. Pavlov Institute of Physiology, Russian Academy of Sciences, St. Petersburg. Translated from Fiziologicheskii Zhurnal im. I. M. Sechenova, Vol. 81, No. 8, pp. 128–134, August, 1995.     

Characteristics of visual seeking and evoked potentials in the extrastriate areas of the cortex in humans

Studies in 11 young, healthy subjects addressed the characteristics of visual seeking (time taken, errors) on changes in the parameters of the target element to be sought (shape, color, and location) in an environment containing heterogeneous white distractors. Evoked potentials (EP) were recorded in six cortical leads (P3, P4, T3, T4, T5, T6) and the late endogenous components of EP were studied, i.e., the N2 and P3 components (standard terminology), as these components are known to change when the type of search changes, in the zone of so-called late selection. When the search difficulty increased (increased similarity between target and distractors), an increase in seeking time was accompanied by a delay in the P3 component and a decrease in its amplitude. Location of the target in a defined position resulted in a decrease in search time and a reduction in the latent period of the P3 component as compared with the situation in which the target position was indeterminate. Changes in the color of the target stimulus led to elimination of the inhibitory action of the distractors: EP parameters were no different from those recorded on presentation of single stimuli. A high level of correlation was found between search parameters and measures of the P3 component. Changes in EP in different types of search were essentially identical (no statistical differences) in the parietal and temporal leads. This suggests that on seeking the target in the environment, the parietal and temporal areas of the cortex function as a single system.     

The primary visual cortex in the mouse: Receptive field properties and functional organization

Summary Receptive field (RF) characteristics of cells in primary visual cortex of the mouse (C57B16 strain) were studied by single unit recording. We have studied the functional organization of area 17 along both the radial and tangential dimensions of the cortex. Eighty seven percent of the visual neurons could be classified according to their responses to oriented stimuli and to moving stimuli. Cells which preferred a flashed or moving bar of a particular orientation and responded less well to bars of other orientations or to spots, were classified as orientation selective (simple RF 23%, complex RF 18%). The majority of them were, moreover, unidirectional (24%). All orientations were roughly equally represented. Cells with oriented RFs were recorded mostly in the upper part of cortical layers II–III, where they appeared to be clustered according to their preferred orientation. Neurons that responded equally well to spots and bars of all orientations (46%) were classified as non-oriented; among these neurons there were several subcategories. Cells which responded equally well to spots and bars but preferred stimuli moving along one or both directions of a particular axis were classified as non oriented asymmetric cells (unidirectional 14%, bidirectional 4%). They were recorded mainly in supra- and infra-granular layers. Cells unaffected by stimulus shape and orientation which responded equally well to all directions of movement were classified as symmetric units. They had receptive field classified as ON (11%), OFF (1%), ON/ OFF (11%), or were unresponsive to stationary stimuli (5%). These cells were mostly found in layer IV, in which they constituted the majority of recorded cells. There was no apparent correlation between the functional type and size of RFs. However, the greatest proportion of small RFs was found in layer IV. In the binocular segment of the mouse striate cortex, the influence of the contralateral eye predominated. Ninety five percent of cells in this segment were driven through the contralateral eye. However, 70% of cells were binocularly activated, showing that considerable binocular integration occured in this cortical segment. Ocular dominance varied less along the radial than along the tangential dimension of the cortex.     

染铅大鼠胎盘一氧化氮、基质金属蛋白酶-9表达与胎盘组织超微结构的相关性

目的观察NO和基质金属蛋白酶-9（MMP-9）在孕期铅暴露大鼠胎盘组织中的变化特征,及其与胎盘组织超微结构的相关性。方法108只大鼠分为4组（A、B、C组和对照组）,对照组孕期饮服蒸馏水,A、B和C组分别于孕期不同阶段饮服0.025%醋酸铅溶液。原子吸收光谱法测定血铅水平。硝酸还原酶法测定胎盘组织NO含量。免疫组化法测定胎盘滋养层细胞MMP-9表达阳性率。电镜观察胎盘组织超微结构。结果①A和B组孕鼠胎盘组织NO含量均高于对照组,差异有显著统计学意义（P＆lt;0.01）;②MMP-9评分4组间总体差异有显著统计学意义（H=39.226,P＆lt;0.01）,以B组的MMP-9阳性表达率最高,C组最低;③各组孕鼠胎盘组织超微结构示：对照组合体滋养层细胞微绒毛致密,胞质内细胞器丰富,内质网致密,有较多的线粒体,细胞间连接和基底膜清晰完整;A组胎盘微血管内皮细胞肿胀,基底膜增厚,微绒毛突起;B组滋养层巨细胞增生,空泡化细胞岛增多;C组滋养层细胞坏死,微绒毛稀疏,线粒体数目减少和粗面内质网池扩张。结论铅暴露孕鼠胎盘组织中NO含量和MMP-9表达阳性率与胎盘组织超微结构的病理改变密切相关。     

Perivascular nerves with immunoreactivity to vasoactive intestinal polypeptide in cephalic arteries of the cat: distribution, possible origins and functional implications

The distribution of nerves containing vasoactive intestinal polypeptide(VIP)-immunoreactive material was examined in the cephalic arteries and cranial nerves of cats using an indirect immunofluorescence procedure on whole mounts. Perivascular VIP-immunoreactive nerves were widely distributed in arteries and arterioles supplying glands, muscles and mucous membranes of the face. Within the cerebral circulation, perivascular VIP-immunoreactive nerves were most abundant in the circle of Willis and the proximal portions of the major cerebral arteries and their proximal branches supplying the rostral brainstem and ventral areas of the cerebral cortex. Nerves containing VIP-immunoreactive material were absent from distal portions of arteries supplying the posterior brainstem, cerebellum and dorsal cerebral cortex. Cerebral perivascular VIP-immunoreactive nerves had extracerebral origins probably from VIP-immunoreactive perikarya within microganglia in the cavernous plexus and external rete. Extracerebral perivascular VIP-immunoreactive nerves probably arose from VIP-immunoreactive perikarya in microganglia associated with the tympanic plexus, chorda tympani, lingual nerve and Vidian nerve as well as from cells in the otic, sphenopalatine, submandibular and sublingual ganglia. Therefore, it seems likely that each major segment of the cephalic circulation is supplied by local VIP-immunoreactive neurons. If the VIP-immunoreactive nerves cause vasodilation, they are well placed to allow redistribution of arterial blood flow within the head. During heat stress, neurogenic vasodilation of the appropriate beds would permit efficient cooling of cerebral blood, particularly that supplying the rostral brainstem and surrounding areas of the cerebral cortex.