The mechanisms for compression and reflection of cortical waves

Waves are common in cortical networks and may be important for carrying information about a stimulus from one local circuit to another. In a recent study of visually evoked waves in rat cortex, compression and reflection of waves are observed as the activation passes from visual areas V1 to V2. The authors of this study apply bicuculline (BMI) and demonstrate that the reflection disappears. They conclude that inhibition plays a major role in compression and reflection. We present several models for propagating waves in heterogeneous media and show that the velocity and thus compression depends weakly on inhibition. We propose that the main site of action of BMI with respect to wave propagation is on the threshold for firing which we suggest is related to action on potassium channels. We combine numerical and analytic methods to explore both compression and reflection in an excitable system with synaptic coupling.     

Reverberation of recent visual experience in spontaneous cortical waves

Spontaneous waves of activity propagating across large cortical areas may play important roles in sensory processing and circuit refinement. However, whether these waves are in turn shaped by sensory experience remains unclear. Here we report that visually evoked cortical activity reverberates in subsequent spontaneous waves. Voltage-sensitive dye imaging in rat visual cortex shows that following repetitive presentation of a given visual stimulus, spatiotemporal activity patterns resembling the evoked response appear more frequently in the spontaneous waves. This effect is specific to the response pattern evoked by the repeated stimulus, and it persists for several minutes without further visual stimulation. Such wave-mediated reverberation could contribute to short-term memory and help to consolidate the transient effects of recent sensory experience into long-lasting cortical modifications.     

Stimulus-entrained oscillatory activity propagates as waves from area 18 to 17 in cat visual cortex

Previous studies in cat visual cortex reported that area 18 can actively drive neurons in area 17 through cortico-cortical projections. However, the dynamics of such cortico-cortical interaction remains unclear. Here we used multielectrode arrays to examine the spatiotemporal pattern of neuronal activity in cat visual cortex across the 17/18 border. We found that full-field contrast reversal gratings evoked oscillatory wave activity propagating from area 18 to 17. The wave direction was independent of the grating orientation, and could not be accounted for by the spatial distribution of receptive field latencies, suggesting that the waves are largely mediated by intrinsic connections in the cortex. Different from the evoked waves, spontaneous waves propagated along both directions across the 17/18 border. Together, our results suggest that visual stimulation may enhance the flow of information from area 18 to 17.     

Temporal Sequence of Visuo-Auditory Interaction in Multiple Areas of the Guinea Pig Visual Cortex

Recent studies in humans and monkeys have reported that acoustic stimulation influences visual responses in the primary visual cortex (V1). Such influences can be generated in V1, either by direct auditory projections or by feedback projections from extrastriate cortices. To test these hypotheses, cortical activities were recorded using optical imaging at a high spatiotemporal resolution from multiple areas of the guinea pig visual cortex, to visual and/or acoustic stimulations. Visuo-auditory interactions were evaluated according to differences between responses evoked by combined auditory and visual stimulation, and the sum of responses evoked by separate visual and auditory stimulations. Simultaneous presentation of visual and acoustic stimulations resulted in significant interactions in V1, which occurred earlier than in other visual areas. When acoustic stimulation preceded visual stimulation, significant visuo-auditory interactions were detected only in V1. These results suggest that V1 is a cortical origin of visuo-auditory interaction.     

No evidence for early modulation of evoked responses in primary visual cortex to irrelevant probe stimuli presented during the attentional blink

Background

During rapid serial visual presentation (RSVP), observers often miss the second of two targets if it appears within 500 ms of the first. This phenomenon, called the attentional blink (AB), is widely held to reflect a bottleneck in the processing of rapidly sequential stimuli that arises after initial sensory registration is complete (i.e., at a relatively late, post-perceptual stage of processing). Contrary to this view, recent fMRI studies have found that activity in the primary visual area (V1), which represents the earliest cortical stage of visual processing, is attenuated during the AB. Here we asked whether such changes in V1 activity during the AB arise in the initial feedforward sweep of stimulus input, or instead reflect the influence of feedback signals from higher cortical areas.

Methodology/Principal Findings

EEG signals were recorded while participants monitored a sequential stream of distractor letters for two target digits (T1 and T2). Neural responses associated with an irrelevant probe stimulus presented simultaneously with T2 were measured using an ERP marker – the C1 component – that reflects initial perceptual processing of visual information in V1. As expected, T2 accuracy was compromised when the inter-target interval was brief, reflecting an AB deficit. Critically, however, the magnitude of the early C1 component evoked by the probe was not reduced during the AB.

Conclusions/Significance

Our finding that early sensory processing of irrelevant probe stimuli is not suppressed during the AB is consistent with theoretical models that assume that the bottleneck underlying the AB arises at a post-perceptual stage of processing. This suggests that reduced neural activity in V1 during the AB is driven by re-entrant signals from extrastriate areas that regulate early cortical activity via feedback connections with V1.     

Beyond the aorta: partial transmission of reflected waves from aortic coarctation into supra-aortic branches modulates cerebral hemodynamics and left ventricular load

Wave reflection from the site of aortic coarctation produces a reflected backward compression wave (BCW) that raises left ventricular (LV) afterload. However, not all reflected wave power will propagate back to the LV. This study investigated the hypothesis that the BCW is partially transmitted into supra-aortic vessels as a forward wave and explored the consequences of this phenomenon for cerebral and LV haemodynamic load. In eight sheep, high fidelity pressure and flow were measured in the aortic trunk (AoT) and brachiocephalic trunk (BCT, the single supra-aortic vessel present in sheep) at baseline and during two levels of proximal descending aortic constriction. Wave power analysis showed that aortic constriction produced not only a BCW in the AoT, but also a second forward compression wave (\(\mathrm{FCW}_{2})\) in the BCT that augmented pressure and flow after the initial forward compression wave (\(\mathrm{FCW}_{1})\). Mathematical analysis and a one-dimensional model of the human systemic arteries and aortic coarctation suggested that the relative transmission of waves into supra-aortic vessels versus the aorta was determined by the relative admittances of these vessels. Reducing supra-aortic admittance (1) increased pressure and flow pulsatility in cerebral arteries, (2) produced carotid and middle cerebral arterial flow waveforms with an older adult phenotype, (3) promoted transmission of reflected wave power towards the LV and (4) substantially increased mid- to late-systolic myocardial stress, which may promote LV hypertrophy. These findings suggest that wave transmission into supra-aortic branches has an important impact on both cerebral hemodynamics and LV load in aortic coarctation.     

Retinotopic mapping of categorical and coordinate spatial relation processing in early visual cortex

Spatial relations are commonly divided in two global classes. Categorical relations concern abstract relations which define areas of spatial equivalence, whereas coordinate relations are metric and concern exact distances. Categorical and coordinate relation processing are thought to rely on at least partially separate neurocognitive mechanisms, as reflected by differential lateralization patterns, in particular in the parietal cortex. In this study we address this textbook principle from a new angle. We studied retinotopic activation in early visual cortex, as a reflection of attentional distribution, in a spatial working memory task with either a categorical or a coordinate instruction. Participants were asked to memorize a dot position, with regard to a central cross, and to indicate whether a subsequent dot position matched the first dot position, either categorically (opposite quadrant of the cross) or coordinately (same distance to the centre of the cross). BOLD responses across the retinotopic maps of V1, V2, and V3 indicate that the spatial distribution of cortical activity was different for categorical and coordinate instructions throughout the retention interval; a more local focus was found during categorical processing, whereas focus was more global for coordinate processing. This effect was strongest for V3, approached significance in V2 and was absent in V1. Furthermore, during stimulus presentation the two instructions led to different levels of activation in V3 during stimulus encoding; a stronger increase in activity was found for categorical processing. Together this is the first demonstration that instructions for specific types of spatial relations may yield distinct attentional patterns which are already reflected in activity early in the visual cortex.     

Calcium waves induced by large voltage pulses in fish keratocytes.

Intracellular calcium waves in fish keratocytes are induced by the application of electric field pulses with amplitudes between 55 and 120 V/cm and full width at half-maximum of 65-100 ms. Calcium concentrations were imaged using two-photon excited fluorescence microscopy (Denk et al., 1990 Science. 248:73-76; Williams et al. 1994 FASEB J. 8:804-813) and the ratiometric calcium indicator indo-1. The applied electric field pulses induced waves with fast calcium rise times and slow decays, which nucleated in the lamellipodium at the hyperpolarized side of the cells and, less frequently, at the depolarized side. The effectiveness of wave generation was determined by the change induced in the membrane potential, which is about half the field strength times the cell width in the direction of the field. Stimulation of waves began at voltage drops across the cell above 150 mV and saturated at voltage drops above 300 mV, where almost all cells exhibited a wave. Waves were not induced in low-calcium media and were blocked by the nonselective calcium channel blockers cobalt chloride and verapamil, but not by specific organic antagonists of voltage-sensitive calcium channel conductance. Thapsigargin stopped wave propagation in the cell body, indicating that calcium release from intracellular stores is necessary. Thus a voltage pulse stimulates Ca2+ influx through calcium channels in the plasma membrane, and if the intracellular calcium concentration reaches a threshold, release from intracellular stores is induced, creating a propagating wave. These observations and the measured parameters (average velocity approximately 66 micron/s and average rise time approximately 68 ms) are consistent with a wave amplification model in which[equation, see text] determines the effective diffusivity of the propagating molecules, D approximately 300 micron2/s (Meyer, 1991. Cell. 64:675-678).     

Implications on visual apperception: Energy, duration, structure and synchronization

Although primary visual cortex (V1 or striate) activity per se is not sufficient for visual apperception (normal conscious visual experiences and conscious functions such as detection, discrimination, and recognition), the same is also true for extrastriate visual areas (such as V2, V3, V4/V8/VO, V5/M5/MST, IT, and GF). In the lack of V1 area, visual signals can still reach several extrastriate parts but appear incapable of generating normal conscious visual experiences. It is scarcely emphasized in the scientific literature that conscious perceptions and representations must have also essential energetic conditions. These energetic conditions are achieved by spatiotemporal networks of dynamic mitochondrial distributions inside neurons. However, the highest density of neurons in neocortex (number of neurons per degree of visual angle) devoted to representing the visual field is found in retinotopic V1. It means that the highest mitochondrial (energetic) activity can be achieved in mitochondrial cytochrome oxidase-rich V1 areas. Thus, V1 bear the highest energy allocation for visual representation.In addition, the conscious perceptions also demand structural conditions, presence of adequate duration of information representation, and synchronized neural processes and/or ‘interactive hierarchical structuralism.’ For visual apperception, various visual areas are involved depending on context such as stimulus characteristics such as color, form/shape, motion, and other features. Here, we focus primarily on V1 where specific mitochondrial-rich retinotopic structures are found; we will concisely discuss V2 where smaller riches of these structures are found. We also point out that residual brain states are not fully reflected in active neural patterns after visual perception. Namely, after visual perception, subliminal residual states are not being reflected in passive neural recording techniques, but require active stimulation to be revealed.     

Contrast invariance in the human lateral occipital complex depends on attention

The human visual system has a remarkable ability to successfully operate under a variety of challenging viewing conditions. For example, our object-recognition capabilities are largely unaffected by low-contrast (e.g., foggy) environments. The basis for this ability appears to be reflected in the neural responses in higher cortical visual areas that have been characterized as being invariant to changes in luminance contrast: neurons in these areas respond nearly equally to low-contrast as compared to high-contrast stimuli. This response pattern is fundamentally different than that observed in earlier visual areas such as primary visual cortex (V1), which is highly dependent on contrast. How this invariance is achieved in higher visual areas is largely unknown. We hypothesized that directed spatial attention is an important prerequisite of the contrast-invariant responses in higher visual areas and tested this with functional MRI (fMRI) while subjects directed their attention either toward or away from contrast-varying shape stimuli. We found that in the lateral occipital complex (LOC), a visual area important for processing shape information, attention changes the form of the contrast response function (CRF). By directing attention away from the shape stimuli, the CRF in the LOC was similar to that measured in V1. We describe a number of mechanisms that could account for this important function of attention.     

视力与视觉诱发电位的相关分析

对104例病人的图形翻转VEP的瞬态波形各参数,以及9例正常或近视学生的稳态曲线功率谱与视力之间的关系进行了多元相关统计分析,旨在探讨VEP的哪些参数可客观地评估视力.结果表明,瞬态VEP的波形参数中以13’格诱发的N_1P_1、P_1N_2的峰峰值及P_(100)潜伏期与视力的相关系数最大,故认为,分析视力时以平均P_(100)波的波幅值和P_(100)波潜伏期作指标较为灵敏;而稳态、VEP能谱曲线则显示,视力与平均相叶能谱或刺激频率点的能谱相关性较大,与二次谐波的相关性则小.     

Two Cortical Circuits Control Propagating Waves in Visual Cortex

Visual stimuli produce waves of activity that propagate across the visual cortex of fresh water turtles. This study used a large-scale model of the cortex to examine the roles of specific types of cortical neurons in controlling the formation, speed and duration of these waves. The waves were divided into three components: initial depolarizations, primary propagating waves and secondary waves. The maximal conductances of each receptor type postsynaptic to each population of neurons in the model was systematically varied and the speed of primary waves, durations of primary waves and total wave durations were measured. The analyses indicate that wave formation and speed are controlled principally by feedforward excitation and inhibition, while wave duration is controlled principally by recurrent excitation and feedback inhibition.     

Calcium waves in mammalian heart: quantification of origin, magnitude, waveform, and velocity

A dual, digital, indo-1 fluorescence imaging system was used to obtain high-speed ratiometric images of [Ca2+]i waves in single voltage-clamped mammalian cardiac cells. The spatiotemporal origin of [Ca2+]i waves in depolarized cells was detected as the spontaneous appearance, over 100-300 ms, of domelike regions of elevated [Ca2+]i, approximately 20 microns in diameter and 300 nM at the center. Images of [Ca2+]i taken at 67-ms intervals during propagation of [Ca2+]i waves revealed that the [Ca2+]i wave front was 1) constant in shape, 2) spatially steep, typically rising from 500 to 1200 nM in about 10 microns, and 3) propagating at constant velocity, typically 100 microns/s at 22 degrees C. The observed spatial and temporal patterns of origin and propagation of [Ca2+]i waves are consistent with the hypothesis that [Ca2+]i waves arise from propagating Ca2(+)-induced release of Ca2+ mediated by diffusion of cytosolic Ca2+. The [Ca2+]i waves are smaller in peak magnitude and can occupy a larger fraction of the cell than thought previously on the basis of indirect observations.     

Role of tapering in aortic wave reflection: hydraulic and mathematical model study

Pressure and flow have been measured simultaneously at six locations along the aorta of an anatomically correct 1:1 scale hydraulic elastic tube model of the arterial tree. Our results suggest a discrete reflection point at the level of the renal arteries based on (i) the quarter-wavelength formula and (ii) the comparison of foot-to-foot (c(ff)) and apparent phase velocity (c(app)). However, separation of the pressure wave into an incident and reflected wave at all six locations indicates continuous reflection: a reflected wave is generated at each location as the forward wave passes by. We did a further analysis using a mathematical transmission line model with a simple tapering geometry (length 50 cm, 31 and 11 mm proximal and distal diameter, respectively) for a low (0.32 ml/mmHg), normal (1.6 ml mmHg) and high (8 ml/mmHg) value of total arterial compliance. Using the quarter-wavelength formula, a discrete reflection point is found at x = 33 cm, the level of the renal arteries, independent of the value of total compliance. However, local analysis comparing c(ff) and c(app) does not reveal a marked reflection site, and the analysis of incident and reflected waves merely suggests a continuous reflection. We therefore conclude that the measured in vivo aortic wave reflection indices are the result of at least two interacting phenomena: a continuous wave reflection due to tapering, and local reflections arising from branches at the level of the diaphragm. The continuous reflection is hidden in the input impedance pattern. Using the quarter-wavelength formula or the classical wave separation theory, it appears as a reflection coming from a single discrete site, confusingly also located at the level of the diaphragm. Therefore, the quarter-wavelength formula and the linear wave separation theory should be used with caution to identify wave reflection zones in the presence of tapering, i.e., in most mammalian arteries.     

Nonlinear periodic space-charge waves in plasma

A solution is obtained in the form of coupled nonlinear periodic space-charge waves propagating in a magnetoactive plasma. The wave spectrum in the vicinity of the critical point, where the number of harmonics increases substantially, is found to fall with harmonic number as ∝ s ^?1/3. Periodic space-charge waves are invoked to explain the zebra pattern in the radio emission from solar flares.     

The evoked cortical activity of the cerebral hemispheres in man during the active and passive perception of facial expression]

Twenty-two right-handed subjects were asked either to identify sad, neutral, or laughing faces presented on a computer monitor or to view passively the same pictures without classification. Visual evoked potentials were recorded from F3/4, C3/4, P3/4, O1/2, and T5/6 derivations. In comparison with the passive viewing, the emotion recognition was characterized by the higher level of cortical activity reflected in the higher N1, N2, N3 amplitudes and shortened latencies of N1, P2, and N2 waves. In contrast, the latencies of the later P3 and N3 waves were longer. During emotion recognition, the dynamical brain mapping technique revealed symmetrical activation of the frontocentral areas and only the right-side activation during the passive perception. Factor analysis demonstrated a complication of N2 structure in the task of face emotion recognition. A principal component corresponding to the descending part of the N2 wave was revealed, which probably reflected the stage of image classification.     

Personality-related differences in recognition of facial emotional expression and cortical evoked potentials

A correlation between some characteristics of the visual evoked potentials and individual personality traits (by the Kettell scale) was revealed in 40 healthy subjects when they recognized facial expressions of anger and fear. As compared to emotionally stable subjects, emotionally unstable subjects had shorter latencies of evoked potentials and suppressed late negativity in the occipital and temporal areas. In contrast, amplitude of these waves in the frontal areas was increased. In emotionally stable group of subjects differences in the evoked potentials related to emotional expressions were evident throughout the whole signal processing beginning from the early sensory stage (P1 wave). In emotionally unstable group differences in the evoked potentials related to recognized emotional expressions developed later. Sensitivity of the evoked potentials to emotional salience of faces was also more pronounced in the emotionally stable group. The involvement of the frontal cortex, amygdala, and the anterior cingulate cortex in the development of individual features of recognition of facial expressions of anger and fear is discussed.     

Abnormal attentional modulation of retinotopic cortex in parietal patients with spatial neglect

Brain regions beyond visual cortex are thought to be responsible for attention-related modulation of visual processing [1, 2], but most evidence is indirect. Here, we applied functional magnetic resonance imaging (fMRI), including retinotopic mapping of visual areas, to patients with focal right-parietal lesions and left spatial neglect [3, 4]. When attentional load at fixation was minimal, retinotopic areas in right visual cortex showed preserved responses to task-irrelevant checkerboards in the contralateral left hemifield, analogously to left visual cortex for right-hemifield checkerboards, indicating a "symmetric" pattern in both hemispheres with respect to contralateral stimulation under these conditions. But when attentional load at fixation was increased, a functional asymmetry emerged for visual cortex, with contralateral responses in right visual areas being pathologically reduced (even eliminated for right V4/TEO), whereas left visual areas showed no such reduction in their contralateral response. These results reveal attention-dependent abnormalities in visual cortex after lesions in distant (parietal) regions. This may explain otherwise puzzling aspects of neglect [5, 6], as confirmed here by additional behavioral testing.     

Comparison of evoked potentials to tones and clicks recorded simulataneously by multiple electrodes from the auditory cortex in unanesthetized cats

Evoked potentials to tones and clicks were recorded simultaneously from seven points of the auditory cortex and one or two points of the somatosensory cortex in unanesthetized cats. Comparison of evoked potentials to tones of equal loudness in the 250–7000 Hz band showed no common pattern of cortical tonotopic distribution. However, an individual dependence of the components of the evoked potential on pitch and on localization of the recording point exists for each animal. With a change in stimulus intensity the absolute and relative values of these components of the evoked potential vary. The initial positive waves are the most variable; besides the two waves already known a third, intermediate wave, particulary sensitive to loudness, was discovered. The negative wave of the primary response increases proportionally to loudness. Evoked potentials to clicks are more uniform over the auditory cortex and more stable than those to tones. Responses appeared in the somatosensory cortex to loud stimuli, more regularly to clicks than to tones. It is concluded that the parameter of pitch is reflected in the cat cortex as a complex spatially-individual distribution of the amplitude and time parameters of the evoked potentials.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 7, No. 2, pp. 115–125, March–April, 1975.