Cortical response field dynamics in cat visual cortex

Little is known about the "inverse" of the receptive field--the region of cortical space whose spatiotemporal pattern of electrical activity is influenced by a given sensory stimulus. We refer to this activated area as the cortical response field, the properties of which remain unexplored. Here, the dynamics of cortical response fields evoked in visual cortex by small, local drifting-oriented gratings were explored using voltage-sensitive dyes. We found that the cortical response field was often characterized by a plateau of activity, beyond the rim of which activity diminished quickly. Plateau rim location was largely independent of stimulus orientation. However, approximately 20 ms following plateau onset, 1-3 peaks emerged on it and were amplified for 25 ms. Spiking was limited to the peak zones, whose location strongly depended on stimulus orientation. Thus, alongside selective amplification of a spatially restricted suprathreshold response, wider activation to just below threshold encompasses all orientation domains within a well-defined retinotopic vicinity of the current stimulus, priming the cortex for processing of subsequent stimuli.     

Estimates of the net excitatory currents evoked by visual stimulation of identified neurons in cat visual cortex

The action potential discharge response of single neurons to both visual stimulation and injections of current were obtained during intracellular recordings in cat visual cortex in order to estimate the net excitatory current arriving at the soma during visual stimulation. Of 45 neurons recorded intracellularly, 19 pyramidal neurons and one basket cell were labelled with horseradish peroxidase. The discharge of all neurons adapted to constant current. For 40 neurons, a single exponential provided a good fit to the adapting discharge (r2 = 0.73 +/- 0.03) for all current intensities. Superficial layer neurons were significantly faster adapting [P < 0.001, mean (+/- SEM) time constant of adaptation = 11.5 +/- 1.3 ms; n = 20] than deep layer neurons (mean time constant of adaptation = 51.4 +/- 6.4 ms; n = 10). The percentage adaptation of the spike frequency, %(peak - adapted rate)/peak, was determined from the fitted exponential. Superficial layer neurons adapted significantly more strongly (P < 0.01, mean = 67 +/- 3%) than deep layer neurons (mean = 51 +/- 5%). The mean firing frequency in response to a current step of 320 ms duration had a linear relationship to the amplitude of the injected current (slope 66 spikes/s/nA; origin zero, mean r2 = 0.94; n = 33). This relationship provided a means of estimating the net peak excitatory current generated by visual stimuli. The estimated mean peak somatic current during the passage of a bar across the receptive field was 1.1 nA and the average current for the duration of the visually evoked discharge was 0.64 nA (n = 17). The transfer response of real and model neurons was obtained by differentiating the discharge response to a step input current and was then used to predict the output of the neuron following an arbitrary input. When these transfer responses were convolved with known input signals in model neurons, the predicted output was close to the simulated response of the model neuron to the same input waveforms. The transfer response was calculated for eight real neurons. Estimates of the net excitatory current arriving at the soma during visual stimulation was obtained by deconvolution. The mean peak somatic current for these neurons was 0.62 nA.      

Artificial vision for the blind by electrical stimulation of the visual cortex

Artificial vision for the blind may be feasible by interfacing a television camera with electronics stimulating the visual cortex. The status of a major collaborative effort involving the College of Physicians and Surgeons of Columbia University, the University of Utah, and the University of Western Ontario is reviewed. Results have been very encouraging, although much work remains to be done.     

在体膜片钳记录前扣带皮层神经元对伤害性电刺激的反应

目的 在麻醉状态下,采用在体膜片钳记录前扣带皮层(ACC)神经元对外周伤害性电刺激的反应.方法 健康成年雄性SD大鼠6只,经腹腔注射2.5％的戊巴比妥钠麻醉后,立即行气管插管术,随后固定于实验台,进行在体膜片钳记录,细胞记录稳定后,给予1.5mA、时长2s的电流刺激大鼠的前后肢,记录大鼠ACC神经元对电刺激的反应.结果 戊巴比妥钠麻醉状态下,给予大鼠电刺激后,其ACC神经元表现出明显的兴奋性反应,且其对电刺激的反应幅度为(4.02±1.32)mV,反应持续时间为(131.33±58.08)ms,反应潜伏期为(57.83±10.82)ms.结论 戊巴比妥钠麻醉状态下,伤害性电刺激可以诱导ACC神经元出现显著的兴奋性反应.     

Optical imaging and electrophysiology of rat barrel cortex. II. Responses to paired-vibrissa deflections

A study was undertaken to investigate the response of the rodent somatosensory barrel cortex to paired-whisker stimuli. Cortical responses to controlled whisker deflections were recorded by (i) conventional multi-unit extracellular recording within the cytochrome oxidase rich barrels centers, and (ii) intrinsic signal optical imaging, a technique that measures an optical correlate of neuronal activity thought to be related to the deoxygenation of hemoglobin in activated regions. Stimuli were applied to two whiskers in sequence, at temporal separations ranging from 0 to 60 ms. Over intervals of 10-40 ms, the primary effect of paired-whisker stimulation was suppressive. We suggest that paired-whisker inhibition results from the activation of layer IV fast-spike units within the principle whisker's barrel, by excitatory input arriving from a surround-whisker. Paired-whisker stimulation produces inhibition in intrinsic images, because it results in a net reduction in layer II/III and/or layer IV metabolism. Intra- cortical inhibition may serve to convert the sequence of inputs from the whisker array into a barrel cortex magnitude code that can be read by higher cortical areas.      

Optical imaging and electrophysiology of rat barrel cortex. I. Responses to small single-vibrissa deflections

A study was undertaken to investigate the response of the rodent somatosensory barrel cortex to single-whisker, near-threshold vibrissal stimuli. Cortical responses to controlled whisker deflections were recorded by (i) conventional multi-unit extracellular recording within the cytochrome oxidase rich barrels centers and the interbarrel septa, and (ii) intrinsic signal optical imaging, a technique that provides a spatial view of cortical activation thought to be related to the deoxygenation of hemoglobin in activated areas. Barrel cortex neurons responded weakly to whisker deflections of 0.04 degrees. Their response to a series of small stimuli of increasing amplitude was well-fitted by a logarithmic function. Responses to larger stimuli declined monotonically with distance from the center of the barrel column, and were characterized by greater onset and offset firing rates, by greater post-excitatory reduction of firing to below spontaneous levels, and by shorter response latency. In comparison to measurements taken previously from primary vibrissal afferent fibers, we conclude that cortical cells can respond to activity in a very small fraction of first-order sensory neurons.      

Vestibular inputs to human motion-sensitive visual cortex

Two crucial sources of information available to an organism when moving through an environment are visual and vestibular stimuli. Macaque cortical area MSTd processes visual motion, including cues to self-motion arising from optic flow and also receives information about self-motion from the vestibular system. In humans, whether human MST (hMST) receives vestibular afferents is unknown. We have combined 2 techniques, galvanic vestibular stimulation and functional MRI (fMRI), to show that hMST is strongly activated by vestibular stimulation in darkness, whereas adjacent area MT is unaffected. The activity cannot be explained in terms of somatosensory stimulation at the electrode site. Vestibular input appears to be confined to the anterior portion of hMST, suggesting that hMST as conventionally defined may contain 2 subregions. Vestibular activity was also seen in another area previously implicated in processing visual cues to self-motion, namely the cingulate sulcus visual area (CSv), but not in visual area V6. The results suggest that cross-modal convergence of cues to self-motion occurs in both hMST and CSv.     

Domain specificity in visual cortex

We investigated the prevalence and specificity of category-selective regions in human visual cortex. In the broadest survey to date of category selectivity in visual cortex, 12 participants were scanned with functional magnetic resonance imaging while viewing scenes and 19 different object categories in a blocked-design experiment. As expected, we found selectivity for faces in the fusiform face area (FFA), for scenes in the parahippocampal place area (PPA), and for bodies in the extrastriate body area (EBA). In addition, we describe 3 main new findings. First, evidence for the selectivity of the FFA, PPA, and EBA was strengthened by the finding that each area responded significantly more strongly to its preferred category than to the next most effective of the remaining 19 stimulus categories tested. Second, a region in the middle temporal gyrus that has been reported to respond significantly more strongly to tools than to animals did not respond significantly more strongly to tools than to other nontool categories (such as fruits and vegetables), casting doubt on the characterization of this region as tool selective. Finally, we did not find any new regions in the occipitotemporal pathway that were strongly selective for other categories. Taken together, these results demonstrate both the strong selectivity of a small number of regions and the scarcity of such regions in visual cortex.     

Very slow activity fluctuations in monkey visual cortex: implications for functional brain imaging

We examined fluctuations in band-limited power (BLP) of local field potential (LFP) signals recorded from multiple electrodes in visual cortex of the monkey during different behavioral states. We asked whether such signals demonstrated coherent fluctuations over time-scales of seconds and minutes, and would thus serve as good candidates for direct comparison with data obtained from functional magnetic resonance imaging (fMRI). We obtained the following results. (i) The BLP of the local field displayed fluctuations at many time-scales, with particularly large amplitude at very low frequencies (<0.1 Hz). (ii) These fluctuations exhibited high coherence between electrode pairs, particularly for BLP signals derived from the gamma frequency range. (iii) Coherence in the BLP, unlike that in the raw LFP, did not fall off sharply as a function of cortical distance. (iv) The structure and coherence of BLP changes were highly similar under distinctly different behavioral states. These results demonstrate the existence of widespread coherent activity fluctuations in the brain of the awake monkey over very long time-scales. We propose that such signals may make a significant contribution to the high variability observed in the time course of physiological signals, including those measured with functional imaging techniques. The results are discussed in the context of combined fMRI/electrophysiological recordings.     

Optical imaging of somatosensory evoked potentials in the rat cerebral cortex after spinal cord injury

Optical imaging techniques have made it possible to monitor neural activity and to determine its spatiotemporal patterns. Traumatic spinal cord injury (SCI) results in both the death of gray matter neurons and the disruption of ascending and descending white matter tracts at the injury site, leading to the loss of motor and sensory functions. In this study, we monitored and compared cortical responses to the stimulation of sensory tracts in normal control and spinal-cord-injured rats using an optical imaging technique based on a voltage-sensitive dye (VSD). The sciatic nerve was stimulated with a platinum bipolar electrode, and the exposed cortical surface was stained with Di-2-ANEPEQ. Optical signals were recorded from the cerebral cortex using the MiCAM02 optical imaging system. Characteristic spatiotemporal patterns were observed in response to electrical stimulation of the sciatic nerve in normal control rats. In spinal-cord-injured rats, the optical signals were dramatically reduced compared to those of normal rats. Four weeks after SCI, however, the activation area increased in the vicinity of the focal sensory area compared to that of the rats 1 week after SCI. These results suggest that optical imaging with VSD may be useful to map functional changes after SCI.     

Functional and diffusion-weighted magnetic resonance imaging for visualization of the postthalamic visual fiber tracts and the visual cortex.

Diffusion-weighted magnetic resonance imaging (MRI) offers the possibility to study the course of the cerebral white matter tracts whereas functional MRI (fMRI) provides information about the specific functions of cortical areas. We evaluated the combination of fMRI and diffusion-weighted MRI to detect cortical visual areas with their corresponding visual fiber tracts in 15 healthy controls (age: 23 - 53 years, male : female = 8 : 7). We demonstrated activation within the primary visual cortex and white matter bundles connecting the lateral geniculate body and the striate cortex in all subjects investigated. Additional activation could be appreciated in some subjects within the lateral geniculate bodies (n = 2) and the motion-sensitive area V5 (n = 3). The combination of diffusion-weighted and functional imaging allows visualization of the origin, direction and functionality of large white matter tracts. This will prove helpful for imaging structural connectivity within the brain during functional imaging. Moreover, this technique might provide important information for neurosurgical patients presenting with space-occupying lesions close to the cortical and subcortical visual system since this technique can -- in contrast to diffusion tensor imaging -- easily be adopted into a neuronavigation system and can be performed on all MR scanners capable of diffusion-weighted imaging without specific post-processing programs.     

Localization of language-specific cortex by using magnetic source imaging and electrical stimulation mapping.

OBJECT: In this paper the authors demonstrate the concordance between magnetic source (MS) imaging and direct cortical stimulation for mapping receptive language cortex. METHODS: In 13 consecutive surgical patients, cortex specialized for receptive language functions was identified noninvasively by obtaining activation maps aided by MS imaging in the context of visual and auditory word-recognition tasks. Surgery was then performed for treatment of medically intractable seizure disorder (eight patients), and for resection of tumor (four), or angioma (one). Mapping of language areas with cortical stimulation was performed intraoperatively in 10 patients and extraoperatively in three. Cortical stimulation mapping verified the accuracy of the MS imaging-based localization in all cases. CONCLUSIONS: Information provided by MS imaging can be especially helpful in cases of atypical language representation, including bihemispheric representation, and location of language in areas other than those expected within the dominant hemisphere, such as the anterior portion of the superior temporal gyrus, the posteroinferior portion of the middle temporal gyrus, the basal temporal cortex, and the lateral temporooccipital cortex.     

Optical imaging of contrast response in Macaque monkey V1 and V2

Our studies on brightness information processing in Macaque monkey visual cortex suggest that the thin stripes in the secondary visual area (V2) are preferentially activated by brightness stimuli (such as full field luminance modulation and illusory edge-induced brightness modulation). To further examine this possibility, we used intrinsic signal optical imaging to examine contrast response of different functional domains in primary and secondary visual areas (V1 and V2). Color and orientation stimuli were used to map functional domains in V1 (color domains, orientation domains) and V2 (thin stripes, thick/pale stripes). To examine contrast response, sinusoidal gratings at different contrasts and spatial frequencies were presented. We find that, consistent with previous studies, the optical signal increased systematically with contrast level. Unlike single-unit responses, optical signals for both color domains and orientation domains in V1 exhibit linear contrast response functions, thereby providing a large dynamic range for V1 contrast response. In contrast to domains in V1, domains in V2 exhibit nonlinear responses, characterized by high gain at low contrasts, saturating at a mid-high contrast levels. At high contrasts, thin stripes exhibit increasing response, whereas thick/pale stripes saturate, consistent with a strong parvocellular input to thin stripes. These findings suggest that, with respect to contrast encoding, thin stripes have a larger dynamic range than thick/pale stripes and further support a role for thin stripes in processing of brightness information.     

Integration of local inputs in visual cortex

In mammalian visual cortex, local connections are ubiquitous, extensively linking adjacent neurons of all types. In this study, optical maps of intrinsic signals and responses from single neurons were obtained from the same region of cat visual cortex while the effectiveness of the local cortical circuitry was altered by focally disinhibiting neurons within a column of known orientation preference. Maps of intrinsic signals indicated that local connections provide strong and functional subthreshold inputs to neighboring columns of other orientation preferences, altering the observed orientation preference to that of the disinhibited column. However, measuring the suprathreshold response using single-cell recordings revealed only mild changes of preferred orientation over the affected region. Because strongly tuned subthreshold inputs from cortex only marginally affect the tuning of a cortical cell's output, it is concluded that local cortical inputs are integrated weakly compared to geniculate inputs. Such circuitry potentially allows for the normalization of responses across a wide range of input activity through local averaging.      

Monocular visual deprivation suppresses excitability in adult human visual cortex

The adult visual cortex maintains a substantial potential for plasticity in response to a change in visual input. For instance, transcranial magnetic stimulation (TMS) studies have shown that binocular deprivation (BD) increases the cortical excitability for inducing phosphenes with TMS. Here, we employed TMS to trace plastic changes in adult visual cortex before, during, and after 48 h of monocular deprivation (MD) of the right dominant eye. In healthy adult volunteers, MD-induced changes in visual cortex excitability were probed with paired-pulse TMS applied to the left and right occipital cortex. Stimulus-response curves were constructed by recording the intensity of the reported phosphenes evoked in the contralateral visual field at range of TMS intensities. Phosphene measurements revealed that MD produced a rapid and robust decrease in cortical excitability relative to a control condition without MD. The cortical excitability returned to preinterventional baseline levels within 3 h after the end of MD. The results show that in contrast to the excitability increase in response to BD, MD acutely triggers a reversible decrease in visual cortical excitability. This shows that the pattern of visual deprivation has a substantial impact on experience-dependent plasticity of the human visual cortex.     

Optical imaging in oncology

Optical imaging is useful during preclinical drug development for monitoring biomarkers and molecular events in living tissue. In animal models, the technology has been used to monitor tumor growth and metastasis, determine biological effects of novel drugs, monitor tumor enzyme activity, and examine host-tumor interactions. Clinical application of optical imaging remains in the developmental stages. Fluorescence signals have a limited dept of penetration through tissue. Therefore, completely noninvasive application of optical imaging in humans is currently not possible. However, laparoscopy and endoscopy provide opportunities to apply optical imaging in patients     

The spatial pattern of response magnitude and selectivity for orientation and direction in cat visual cortex

Optical imaging studies of orientation and direction preference in visual cortex have typically used vector averaging to obtain angle and magnitude maps. This method has shown half-rotation orientation singularities (pinwheels) located within regions of low orientation vector magnitude. Direction preference is generally orthogonal to orientation preference, but often deviates from this, particularly in regions of low direction vector magnitude. Linear regions of rapid change in direction preference terminate in or near orientation singularities. The vector-averaging method is problematic however because it does not clearly disambiguate spatial variation in orientation tuning width from variation in height. It may also wrongly estimate preferred direction in regions where preference is weak. In this paper we analyze optical maps of cat visual cortex by fitting model tuning functions to the responses. This new method reveals features not previously evident. Orientation tuning height and width vary independently across the map: tuning height is always low near singularities, however regions of broad and narrow orientation tuning width can be found in regions of low tuning height, often alternating in a spoke-like fashion around singularities. Orientation and direction preference angles are always closely orthogonal. Reversals in direction preference form lines that originate precisely in orientation singularities.     

Nitrous oxide suppresses the electromyographic response evoked by electrical stimulation of the motor cortex

The influence of nitrous oxide on motor evoked potential (MEP) elicited in rats by cortical and midcervical electrical stimulation was studied and compared with early components of somatosensory evoked potential (SEP) following stimulation of the posterior tibial nerve in 6 rats. We found that nitrous oxide gradually suppresses MEP, depending on the concentration of this inhalation agent. At a concentration of 66 vol% of nitrous oxide, the MEP was completely abolished, whereas the initial component N1-P1 of the SEP was only slightly reduced. We conclude that the descending impulse elicited by electrical stimulation of the corticospinal tract is mainly inhibited at the level of the spinal neuronal or interneuronal system, since (1) neuromuscular transmission is not blocked by nitrous oxide, and (2) MEP suppression is the same following cortical and midcervical stimulation.