Effects of stimulus duration on neuronal response properties in the somatosensory cortex of the star-nosed mole

Star-nosed moles have a series of mechanosensory appendages surrounding each nostril. Each appendage is covered with sensory organs (Eimer's organs) containing both rapidly adapting and slowly adapting mechanoreceptors and each appendage is represented in primary somatosensory cortex (S1) by a single cortical module. When the skin surface of an appendage is depressed, neurons in the corresponding module in S1 respond in either a transient or sustained fashion. The aim of this study was to characterize and compare the responses of these two classes of neurons to both short (5 or 20 ms) and long (500 ms) mechanosensory stimulation. Activity from neurons in the representation of appendage 11, the somatosensory fovea, was recorded while delivering mechanosensory stimuli to the corresponding skin surface. Transient and sustained neurons had different levels of spontaneous activity and different responses to both short and long mechanosensory stimulation. Neurons with sustained responses had a significantly higher spontaneous firing rate than neurons with transient responses. Transient neurons responded to a 5 ms stimulus with excitation followed by suppression of discharge whereas sustained neurons did not exhibit post-excitatory suppression. Rather, responses of sustained neurons to 5 ms stimuli lasted several hundred milliseconds. Consequently sustained responses contained significantly more spikes than transient responses. These experiments suggest contact to the appendages causes two distinct firing patterns in cortex regardless of the duration of the stimulus. The sustained and transient responses could reflect either the activity of fundamentally different classes of neurons or activity in distinct subcortical and cortical networks.     

Orientation-Cue Invariant Population Responses to Contrast-Modulated and Phase-Reversed Contour Stimuli in Macaque V1 and V2

Visual scenes can be readily decomposed into a variety of oriented components, the processing of which is vital for object segregation and recognition. In primate V1 and V2, most neurons have small spatio-temporal receptive fields responding selectively to oriented luminance contours (first order), while only a subgroup of neurons signal non-luminance defined contours (second order). So how is the orientation of second-order contours represented at the population level in macaque V1 and V2? Here we compared the population responses in macaque V1 and V2 to two types of second-order contour stimuli generated either by modulation of contrast or phase reversal with those to first-order contour stimuli. Using intrinsic signal optical imaging, we found that the orientation of second-order contour stimuli was represented invariantly in the orientation columns of both macaque V1 and V2. A physiologically constrained spatio-temporal energy model of V1 and V2 neuronal populations could reproduce all the recorded population responses. These findings suggest that, at the population level, the primate early visual system processes the orientation of second-order contours initially through a linear spatio-temporal filter mechanism. Our results of population responses to different second-order contour stimuli support the idea that the orientation maps in primate V1 and V2 can be described as a spatial-temporal energy map.     

Separate modulations of human V1 associated with spatial attention and task structure

Functional magnetic resonance imaging (fMRI) was used while normal human volunteers engaged in simple detection and discrimination tasks, revealing separable modulations of early visual cortex associated with spatial attention and task structure. Both modulations occur even when there is no change in sensory stimulation. The modulation due to spatial attention is present throughout the early visual areas V1, V2, V3, and VP, and varies with the attended location. The task structure activations are strongest in V1 and are greater in regions that represent more peripheral parts of the visual field. Control experiments demonstrate that the task structure activations cannot be attributed to visual, auditory, or somatosensory processing, the motor response for the detection/discrimination judgment, or oculomotor responses such as blinks or saccades. These findings demonstrate that early visual areas are modulated by at least two types of endogenous signals, each with distinct cortical distributions.     

Transition between two excitabilities in mesencephalic V neurons

Neurons can make different responses to identical inputs. According to the emerging frequency of repetitive firing, neurons are classified into two types: type 1 and type 2 excitability. Though in mathematical simulations, minor modifications of parameters describing ionic currents can result in transitions between these two excitabilities, empirical evidence to support these theoretical possibilities is scarce. Here we report a joint theoretical and experimental study to test the hypothesis that changes in parameters describing ionic currents cause predictable transitions between the two excitabilities in mesencephalic V (Mes V) neurons. We developed a simple mathematical model of Mes V neurons. Using bifurcation analysis and model simulation, we then predicted that changes in conductance of two low-threshold currents would result in transitions between type 1 and type 2. Finally, by applying specific channel blockers, we observed the transition between two excitabilities forecast by the mathematical model.     

盆神经和阴部神经传入在大鼠腰骶髓的相互作用

应用条件－检验刺激技术观察时间依赖性抑制现象是研究传入信息相互作用的方式之一。用１．５－３倍阈刺激强度的电脉冲交替刺激麻醉、麻痹的盆神经（Ｐｅ）和阴部神经（Ｐｕ）,以玻璃微电极在Ｌ６－Ｓ１节段脊髓背角会聚神经元上记录细胞外放电。条件输入可对深层（＞３００μｍ）单位的检验反应产生时间依赖性抑制效应,产生抑制的刺激间期为１－３６０ｍｓ,Ｐｅ为条件刺激时较长。浅层细胞（＜３００μｍ）发生抑制的间期为１－     

电刺激蝙蝠小脑对中脑上丘神经元听反应的影响

实验在２３只成年中华鼠耳蝠（Ｍｙｏｌｉｓｃｈｉｎｅｎｓｉｓ）上进行。使用常规电生理学方法,观察了电刺激小脑对上丘神经元听反应的影响。在所观察的１７１个上丘神经元中,有１１６个（占６７．８４％）神经元听反应受到影响,其中７２个（占４２．１１％）表现为抑制效应,４４个（占２５．７３％）为易化效应。刺激小脑对上丘神经听反应的影响是双侧的。抑制或易化程度与电刺激强度、声刺激强度以及声、电刺激间隔有关。结果提示,小脑可以对上丘神经元听反应进行调制,这种调制作用可能是小脑调控回声定位过程中听觉－运动的中枢机制之一。     

Visually Driven Activation in Macaque Areas V2 and V3 without Input from the Primary Visual Cortex

Creating focal lesions in primary visual cortex (V1) provides an opportunity to study the role of extra-geniculo-striate pathways for activating extrastriate visual cortex. Previous studies have shown that more than 95% of neurons in macaque area V2 and V3 stop firing after reversibly cooling V1 [1], [2], [3]. However, no studies on long term recovery in areas V2, V3 following permanent V1 lesions have been reported in the macaque. Here we use macaque fMRI to study area V2, V3 activity patterns from 1 to 22 months after lesioning area V1. We find that visually driven BOLD responses persist inside the V1-lesion projection zones (LPZ) of areas V2 and V3, but are reduced in strength by ∼70%, on average, compared to pre-lesion levels. Monitoring the LPZ activity over time starting one month following the V1 lesion did not reveal systematic changes in BOLD signal amplitude. Surprisingly, the retinotopic organization inside the LPZ of areas V2, V3 remained similar to that of the non-lesioned hemisphere, suggesting that LPZ activation in V2, V3 is not the result of input arising from nearby (non-lesioned) V1 cortex. Electrophysiology recordings of multi-unit activity corroborated the BOLD observations: visually driven multi-unit responses could be elicited inside the V2 LPZ, even when the visual stimulus was entirely contained within the scotoma induced by the V1 lesion. Restricting the stimulus to the intact visual hemi-field produced no significant BOLD modulation inside the V2, V3 LPZs. We conclude that the observed activity patterns are largely mediated by parallel, V1-bypassing, subcortical pathways that can activate areas V2 and V3 in the absence of V1 input. Such pathways may contribute to the behavioral phenomenon of blindsight.     

Responses of neurons in primary and inferior temporal visual cortices to natural scenes.

The primary visual cortex (V1) is the first cortical area to receive visual input, and inferior temporal (IT) areas are among the last along the ventral visual pathway. We recorded, in area V1 of anaesthetized cats and area IT of awake macaque monkeys, responses of neurons to videos of natural scenes. Responses were analysed to test various hypotheses concerning the nature of neural coding in these two regions. A variety of spike-train statistics were measured including spike-count distributions, interspike interval distributions, coefficients of variation, power spectra, Fano factors and different sparseness measures. All statistics showed non-Poisson characteristics and several revealed self-similarity of the spike trains. Spike-count distributions were approximately exponential in both visual areas for eight different videos and for counting windows ranging from 50 ms to 5 seconds. The results suggest that the neurons maximize their information carrying capacity while maintaining a fixed long-term-average firing rate, or equivalently, minimize their average firing rate for a fixed information carrying capacity.     

Medullary taste responses are modulated by the bed nucleus of the stria terminalis

Previous studies have shown a modulatory influence of limbic forebrain areas, such as the central nucleus of the amygdala and lateral hypothalamus, on the activity of taste-responsive cells in the nucleus of the solitary tract (NST). The bed nucleus of the stria terminalis (BST), which receives gustatory afferent information, also sends descending axons to the NST. The present studies were designed to investigate the role of the BST in the modulation of NST gustatory activity. Extracellular action potentials were recorded from 101 taste-responsive cells in the NST of urethane-anesthetized hamsters and analyzed for a change in excitability following bilateral electrical stimulation of the BST. The response of NST taste cells to stimulation of the BST was predominately inhibitory. Orthodromic inhibitory responses were observed in 29 of 101 (28.7%) NST taste-responsive cells, with four cells inhibited bilaterally. An increase in excitability was observed in seven of the 101 (6.9%) NST taste cells. Of the 34 cells showing these responses, 25 were modulated by the ipsilateral BST and 15 by the contralateral; four were inhibited bilaterally and two inhibited ipsilaterally and excited contralaterally. The duration of inhibitory responses (mean = 177.9 ms) was significantly longer than that of excitatory responses (35.4 ms). Application of subthreshold electrical stimulation to the BST during taste trials inhibited or excited the taste responses of every BST-responsive NST cell tested with this protocol. NST neurons that were most responsive to sucrose, NaCl, citric acid or quinine hydrochloride were all affected by BST stimulation, although citric acid-best cells were significantly more often modulated and NaCl-best less often modulated than expected by chance. These results combine with excitatory and inhibitory modulation of NST neurons by the insular cortex, lateral hypothalamus and central nucleus of the amygdala to demonstrate extensive centrifugal modulation of brainstem gustatory neurons.     

电刺激蝙蝠中脑上丘对下丘听神经元电活动的影响

实验在24只鲁氏菊头蝠(Rhinolophus rouxi)上进行.使用玻璃微电极在中脑下丘中央核记录听神经元电反应.刺激点位于上丘核.共观察了294个对超声刺激产生反应的下丘听单位.当电刺激上丘时,有122个听单位的反应受到影响,占所观察总数的41.5%.其中96个单位表现为抑制性影响(占32.65%),26个单位表现为易代性效应(占8.84%).其余172个单位不受上丘刺激的影响(58.50%).实验中发现,上述抑制潜伏期一般在5毫秒以上,抑制时程较长.抑制程度与上丘刺激电流强度呈相关关系(r=0.99).实验中还发现,刺激上丘同样可抑制部分下丘神经元的自发放电活动,其抑制后效应相当长,有的可达120毫秒以上.     

Modulation by arginine vasopressin of glutamate excitation in the ventral septal area of the rat brain

Arginine vasopressin is hypothesized to act as a neurotransmitter or neuromodulator in the ventral septal area of the rat brain. To examine this role of vasopressin further, it was applied by microiontophoresis or micropressure from multiple-barrelled micropipettes onto spontaneously active or glutamate-activated neurons. Applied in this manner, vasopressin reduced glutamate-evoked excitation in 32 of the 47 cells studied. Further, micropressure application of the vasopressin antagonist d(CH2)5Tyr(Me)AVP reversed the vasopressin effects. In contrast, administration of vasopressin had no effect on excitations evoked by acetylcholine iontophoresis or on the spontaneous activity of the majority of the ventral septal neurons studied. These observations suggest that vasopressin may be acting on a V1-like receptor on specific neurons in the ventral septal area as a modulator of glutamate actions. Evoked responses were also obtained in the same population of ventral septal cells following stimulation of a variety of limbic areas. Inhibitory input onto most of the vasopressin responsive neurons studied was obtained following electrical stimulation of the paraventricular nucleus and bed nucleus of the stria terminalis, two cell groupings that are potential sources of vasopressin to the ventral septal area. Thus, the similarity in action of exogenously applied vasopressin and the evoked responses following paraventricular nucleus and bed nucleus stimulation suggests that vasopressin may be a neurotransmitter in this pathway.     

Visual cortex: the continuing puzzle of area V2

Surprisingly little is known about the role of V2 in visual processing. A recent study found that the responses of V2 neurons to pairs of angled lines could be predicted from their responses to the individual line components. A simple analysis shows how these neurons may simply sum the responses from one or more orientation selective V1 neurons.     

Brain stem mechanisms underlying acupuncture modality-related modulation of cardiovascular responses in rats.

The present study was designed to investigate brain stem responses to manual acupuncture (MA) and electroacupuncture (EA) at different frequencies at pericardial P (5-6) acupoints located over the median nerve. Activity of premotor sympathetic cardiovascular neurons in the rostral ventral lateral medulla (rVLM) was recorded during stimulation of visceral and somatic afferents in ventilated anesthetized rats. We stimulated either the splanchnic nerve at 2 Hz (0.1-0.4 mA, 0.5 ms) or the median nerve for 30 s at 2, 10, 20, 40, or 100 Hz using EA (0.3-0.5 mA, 0.5 ms) or at approximately 2 Hz with MA. Twelve of 18 cells responsive to splanchnic and median nerve stimulation could be antidromically driven from the intermediolateral columns of the thoracic spinal cord, T2-T4, indicating that they were premotor sympathetic neurons. All 18 neurons received baroreceptor input, providing evidence of their cardiovascular sympathoexcitatory function. Evoked responses during stimulation of the splanchnic nerve were inhibited by 49 +/- 6% (n = 7) with EA and by 46 +/- 4% (n = 6) with MA, indicating that the extent of inhibitory effects of the two modalities were similar. Inhibition lasted for 20 min after termination of EA or MA. Cardiovascular premotor rVLM neurons responded to 2-Hz electrical stimulation at P 5-6 and to a lesser extent to 10-, 20-, 40-, and 100-Hz stimulation (53 +/- 10, 16 +/- 2, 8 +/- 2, 2 +/- 1, and 0 +/- 0 impulses/30 stimulations, n = 7). These results indicate that rVLM premotor sympathetic cardiovascular neurons that receive convergent input from the splanchnic and median nerves during low-frequency EA and MA are inhibited similarly for prolonged periods by low-frequency MA and EA.     

电刺激大鼠苍白球对黑质Ⅰ型神经元的影响

采用细胞外记录方法,分别观察了黑质（ＳＮ）Ⅰ型神经元对刺激苍白球（ＧＰ）内侧部及外侧部的反应。实验共记录了９６个Ⅰ型神经元。刺激ＧＰ的内侧部,有５７个（５９．３８％）神经元被顺行抑制。刺激ＧＰ的外侧部,有８６个（８９．５８％）神经元被顺行抑制;２个（２．０８％）被逆行激活,被逆行激活的神经元产生的诱发电位潜伏期恒定且短（分别为８．０和８．５ｍｓ）。被顺行抑制的神经元中,有的产生抑制、兴奋交替出现的振荡反应。本文讨论了ＧＰ与ｓＮⅠ型神经元的联系途径及ＳＮⅠ型神经元与ＧＰ的联系途径。     

Modulation of V1 Spike Response by Temporal Interval of Spatiotemporal Stimulus Sequence

The spike activity of single neurons of the primary visual cortex (V1) becomes more selective and reliable in response to wide-field natural scenes compared to smaller stimuli confined to the classical receptive field (RF). However, it is largely unknown what aspects of natural scenes increase the selectivity of V1 neurons. One hypothesis is that modulation by surround interaction is highly sensitive to small changes in spatiotemporal aspects of RF surround. Such a fine-tuned modulation would enable single neurons to hold information about spatiotemporal sequences of oriented stimuli, which extends the role of V1 neurons as a simple spatiotemporal filter confined to the RF. In the current study, we examined the hypothesis in the V1 of awake behaving monkeys, by testing whether the spike response of single V1 neurons is modulated by temporal interval of spatiotemporal stimulus sequence encompassing inside and outside the RF. We used two identical Gabor stimuli that were sequentially presented with a variable stimulus onset asynchrony (SOA): the preceding one (S1) outside the RF and the following one (S2) in the RF. This stimulus configuration enabled us to examine the spatiotemporal selectivity of response modulation from a focal surround region. Although S1 alone did not evoke spike responses, visual response to S2 was modulated for SOA in the range of tens of milliseconds. These results suggest that V1 neurons participate in processing spatiotemporal sequences of oriented stimuli extending outside the RF.     

强电针穴位对背角神经元镇痛效应广泛性的中枢机制

实验用雄性大鼠,玻璃微电极细胞外记录Ｔ１２－Ｌ１脊髓背角会聚神经元对后爪伤害性刺激的反应,观察到低强度（２Ｖ）电针作用于与痛源接近的“足三里”穴对背角神经元的伤害性反应有明显的抑制作用,而远隔穴位“下关”穴则无效。而当采用超过Ｃ类纤维阈值１８Ｖ电针时,则远隔穴位“下关”也有明显的镇痛作用。表现为强电针穴位镇痛作用的广泛性。而损毁ＮＲＭ后,强电针（１８Ｖ）远节段“下关”穴的镇痛作用消失,而近节段“足     

Respiratory resetting induced by spinal cord stimulation in the cat

Electrical stimulation (50-150 microA, 0.5-ms duration, 3-300 Hz) was performed within three different regions (lateral, ventrolateral, and ventral) of the C2-C3 spinal cord of decerebrate, vagotomized, paralyzed, and artificially ventilated cats. Spinal cord stimulation sites were located by inserting monopolar or bipolar stimulating electrodes either at the dorsolateral sulcus or at least 1 mm medial or lateral to the sulcus. With stimulation at each site, alterations in respiratory rhythm, orthodromic phrenic nerve responses, and antidromic activation of medullary respiratory-modulated neurons were examined. Phrenic nerve responses to cervical spinal cord stimulation consisted of an early excitation (2-4 ms) and/or a late excitation (4-8 ms). Stimulation of the lateral region evoked the greatest amplitude early response and stimulation of the ventrolateral region produced the greatest late excitation. All three stimulus sites elicited antidromic activation of some respiratory-modulated neurons in the dorsal (DRG) and ventral respiratory groups (VRG). The lateral region was the least effective resetting site, and it had the highest incidence of antidromic activation of both DRG and VRG neurons. The ventrolateral region of the cervical spinal cord was the most effective resetting site, but it had the lowest incidence of antidromic activation of DRG respiratory-modulated neurons. In addition, resetting responses were observed with spinal cord stimulation at similar sites in the thoracic and lumbar spinal cord regions thought to be devoid of inspiratory bulbospinal axons.(ABSTRACT TRUNCATED AT 250 WORDS)     

Vasopressin V1 receptors contribute to hemodynamic and sympathoinhibitory responses evoked by stimulation of adenosine A2a receptors in NTS

Activation of adenosine A2a receptors in the nucleus of the solitary tract (NTS) decreases mean arterial pressure (MAP), heart rate (HR), and renal sympathetic nerve activity (RSNA), whereas increases in preganglionic adrenal sympathetic nerve activity (pre-ASNA) occur, a pattern similar to that observed during hypotensive hemorrhage. Central vasopressin V1 receptors may contribute to posthemorrhagic hypotension and bradycardia. Both V1 and A2a receptors are densely expressed in the NTS, and both of these receptors are involved in cardiovascular control; thus they may interact. The responses elicited by NTS A2a receptors are mediated mostly via nonglutamatergic mechanisms, possibly via release of vasopressin. Therefore, we investigated whether blockade of NTS V1 receptors alters the autonomic response patterns evoked by stimulation of NTS A2a receptors (CGS-21680, 20 pmol/50 nl) in alpha-chloralose-urethane anesthetized male Sprague-Dawley rats. In addition, we compared the regional sympathetic responses to microinjections of vasopressin (0.1-100 ng/50 nl) into the NTS. Blockade of V1 receptors reversed the normal decreases in MAP into increases (-95.6 +/- 28.3 vs. 51.4 +/- 15.7 integralDelta%), virtually abolished the decreases in HR (-258.3 +/- 54.0 vs. 18.9 +/- 57.8 integralDeltabeats/min) and RSNA (-239.3 +/- 47.4 vs. 15.9 +/- 36.1 integralDelta%), and did not affect the increases in pre-ASNA (279.7 +/- 48.3 vs. 233.1 +/- 54.1 integralDelta%) evoked by A2a receptor stimulation. The responses partially returned toward normal values approximately 90 min after the blockade. Microinjections of vasopressin into the NTS evoked dose-dependent decreases in HR and RSNA and variable MAP and pre-ASNA responses with a tendency toward increases. We conclude that the decreases in MAP, HR, and RSNA in response to NTS A2a receptor stimulation may be mediated via release of vasopressin from neural terminals in the NTS. The differential effects of NTS V1 and A2a receptors on RSNA versus pre-ASNA support the hypothesis that these receptor subtypes are differentially located/expressed on NTS neurons/neural terminals controlling different sympathetic outputs.     

Electrophysiological relations between the superior colliculus and the red nucleus in the cat.

The electrical activity of single units located in the parvicellular part of the red nucleus (pRP) was recorded extracellularly in nitrous oxide anesthetized and C1-transected adult cats. In this area, neurons were found to respond to electrical stimulation applied to intermediate and deep layers of the right superior colliculus (SC). Forty neurons located in the pRN of both sides were studied. Three neurons out of 18 (16.6%) located in the contralateral pRN and six neurons out of 22 (27.3%) located in the ipsilateral pRN were driven by the right SC stimulation. The pRN neurons were separated into four groups according to the latency response to the SC stimulation: 1) 0.6-1.9 ms, 2) 2-4 ms, 3) 4-6 ms, 4) variable latency responses. Each of these four groups of neurons showed a particular pattern of discharge, even though their discharge frequency showed a strong consistency. Four pRN neurons, which responded to SC stimulation, showed a significant correlation with spontaneous horizontal eye movements of saccadic type. It is known that the SC represents one of the main outputs of the striato-nigral motor system. The relation between the SC and the pRN described in the present study suggest that connections exist between the cortico-rubral and the striato-nigral systems, since both have the SC as a common output structure. It is likely, therefore, that the cortico-rubral-SC system is involved in the control of oculomotor functions, and that the SC may serve to establish interactions between systems concerned with eye movements.     

Limited plasticity of the rabbit visual cortex and hippocampal neurons in the oddball test

The activity of 41 visual cortex and 20 hippocampal neurons from field CA1 was registered in experiments using oddball-stimulation with different color stimuli varied in intensity. 34% cortical and 37% hippocampal neurons demonstrated plasticity reactions. The significant increase of latest phases of neuronal activity (200-500 and 200-1000 ms after stimulation for cortical neurons and 300-550 ms for hippocampal neurons) was shown in responses to rare deviant stimuli, which had a less intensity than frequently standards. The quantity of the earliest neuronal phase of activity (40-120 ms after stimulation) was stabilized in responses to deviants and standards during the experiment. We propose that such increase of the latest phases of neuronal activity (the limited plasticity) may reflect the mechanisms of orienting reaction.