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PubMed2004

A dynamic system model-based technique for functional MRI data analysis.

Signals in functional magnetic resonance imaging (fMRI) are influenced by physiological fluctuations in addition to local brain activity. We have proposed a dynamic system model-based technique for separation of signal changes related to brain activation inputs from those related to physiological fluctuations. We applied this technique to a visual fMRI experiment to determine the validity and feasibility of this technique for fMRI data analyses. Gradient-echo echo planar images were obtained from 12 healthy volunteers with a Siemens ALLEGRA operating at 3 T, with a repetition time of 500 ms, echo time of 20 ms, field of view of 200-210 mm, matrix size of 64 x 64, and slice thickness of 5 mm. Twelve runs with two stimulation periods of varied duration (2-8 s) with 8-Hz flickering illumination were obtained for each subject. Local signal changes were modeled by an autoregressive model with two exogenous inputs, a visual stimulation input and a global reference signal. Local signal changes were appropriately predicted not only for stimulation periods but also resting periods. A significant linear relationship was found between model static gain based on the dynamic system modeling and beta coefficient based on a general linear model (GLM) analysis for active voxels in the primary visual cortex (analysis of covariance [ANCOVA], P < 0.001; estimated parameter, 0.967; 95% confidence interval, 0.734-1.201). This dynamic system model-based technique is sufficiently accurate and feasible for use in extracting signal changes related to brain activation inputs from measured signals with physiological fluctuations.

Adult↗
PubMed2004

Fos-tau-LacZ mice expose light-activated pathways in the visual system.

We have employed fos-tau-LacZ (FTL) transgenic mice to examine functional activation in the visual areas of the nervous system. The FTL mice express the marker gene lacZ in neurons and their processes following many different stimuli, and allow the imaging of activation from the level of the entire brain surface through individual neurons and their projections. Analysis of FTL expression in the retinas of mice following diurnal exposure to light shows that bipolar cells, specific classes of amacrine cells, ganglion cells, and a dense network of processes in the inner plexiform layer are functionally activated. In animals deprived of light, there is almost no activity in the retina. In the lateral geniculate nucleus (LGN), light exposure appears responsible for FTL expression in dorsal nuclei, but not for expression in the ventral nuclei or the intergeniculate leaflet. In the superficial layers of the superior colliculus, FTL expression is highly dependent on light exposure. Similarly, light exposure is required for FTL expression in primary visual cortex (area 17), but some expression remains in area 18 of dark-adapted animals. Finally, using mice with one or both eyes missing, we have determined which parts of the visual system are dependent on the presence of a functional connectivity from the eye. These data demonstrate the usefulness of the FTL mice to map functional activation within the entire visual system. Furthermore, we can capture visual activation in a conscious animal. Our findings give an insight into the architecture of activity within the retina and throughout the visual system.

Animals↗
PubMed2005

BOLD responses to visual stimulation in survivors of childhood cancer.

Children surviving certain cancers have a high incidence of cognitive deficits caused by central nervous system (CNS) disease or treatments directed at the CNS. To establish the feasibility of using blood oxygen level-dependent (BOLD) functional magnetic resonance imaging (fMRI) to study cognitive deficits in survivors of childhood cancer, we tested the hypothesis that this population has the same BOLD response to visual stimulation as healthy subjects. We used BOLD fMRI to measure spatial and temporal patterns of brain activity after brief visual stimulation in 16 survivors of childhood cancer, 11 age-similar healthy siblings of survivors, and 16 healthy adults. Functional data for the survivors were analyzed with two general linear models, one used a canonical hemodynamic response function (HRF) and the other used a Fourier set as basis functions. The measured BOLD signal and brain activation patterns were similar in the survivors with both models. The BOLD signal for survivors was qualitatively similar in timing and shape, but there were significant quantitative differences as compared with healthy subjects. The activation was normally located in the primary visual cortex in 13 survivors, but the activation volume was significantly smaller in brain tumor survivors than in other groups. These findings demonstrate the feasibility of using BOLD fMRI to investigate brain function in survivors of childhood cancer. However, fMRI studies in this population must take into account effects of quantitative differences in their BOLD responses as compared to healthy subjects.

Adolescent↗
PubMed2005

Identification of the neural sources of the pattern-reversal VEP.

This study aimed to characterize the neural generators of the early components of the visual-evoked potential (VEP) to pattern-reversal gratings. Multichannel scalp recordings of VEPs and dipole modeling techniques were combined with functional magnetic resonance imaging (fMRI) and retinotopic mapping in order to estimate the locations of the cortical sources giving rise to VEP components in the first 200 ms poststimulus. Dipole locations were seeded to visual cortical areas in which fMRI activations were elicited by the same stimuli. The results provide strong evidence that the first major component of the VEP elicited by a pattern-reversal stimulus (N75/P85) arises from surface-negative activity in the primary visual cortex (area V1). Subsequent waveform components could be accounted for by dipoles that were in close proximity to fMRI activations in the following cortical areas: P95 (area MT/V5), P125/N135 (area V1), N150 (transverse parietal sulcus, TPS), N160 (ventral occipital areas VP, V4v, and V4/V8), and N180 (dorsal occipital areas V3A/V7). These results provide a detailed spatiotemporal profile of the cortical origins of the pattern-reversal VEP, which should enhance its utility in both clinical and basic studies of visual-perceptual processing.

Adolescent↗
PubMed2005

The many faces of the gamma band response to complex visual stimuli.

While much is known about the functional architecture of the visual system, little is known about its large-scale dynamics during perception. This study describes this dynamics with a high spatial, temporal and spectral resolution. We recorded depth EEG of epileptic patients performing a face detection task and found that the stimuli induced strong modulations in the gamma band (40 Hz to 200 Hz) in selective occipital, parietal and temporal sites, in particular the fusiform gyrus, the lateral occipital gyrus and the intra-parietal sulcus. Occipito-temporal sites were the first to be activated, closely followed by the parietal sites, while portions of the primary visual cortex seemed to deactivate temporarily. Some of those effects were found to be correlated across distant sites, suggesting that a coordinated balance between regional gamma activations and deactivations could be involved during visual perception.

Adult↗
PubMed2005

Effects of EPI readout bandwidth on measured activation map and BOLD response in fMRI experiments.

The purpose of this study was to evaluate the effects of echo planar imaging (EPI) readout bandwidth and its interaction with data processing procedures on the measured blood oxygenation level dependent (BOLD) response and activation in fMRI experiments. Seventeen healthy subjects were scanned during a brief visual stimulation paradigm with two EPI pulse sequences having 'high' (1953 Hz/pixel) and 'low' (780 Hz/pixel) readout bandwidth. Functional data were analyzed with a general linear model including temporal filtering and a basic correlation model following (1) no preprocessing, (2) realignment, or (3) realignment and spatial smoothing. A range of statistical thresholds were used to generate activation maps. Despite slightly higher BOLD signal detected with the high bandwidth sequence from matched ROIs in the primary visual cortex, results showed that the low bandwidth pulse sequence was more sensitive under most conditions evaluated. That is, the low bandwidth sequence detected greater numbers of activated voxels with lower cluster average BOLD signal (e.g., low bandwidth detected 1.4 times more voxels, with average BOLD signal 30% lower compared to high bandwidth for P = 0.05 (corrected) with the 3rd preprocessing procedure using the general linear model). However, there was significant interaction between bandwidth and data preprocessing procedures. Of particular interest, the sensitivity advantage of the low bandwidth pulse sequence decreased for the smoothed data as the activation threshold became less conservative. For the frequently used threshold of P = 0.001 (uncorrected) and cluster size of at least 5 voxels, the bandwidth advantage became insignificant. These findings demonstrate that the effects of bandwidth should be considered carefully in the design, analysis, and interpretation of BOLD fMRI studies.

Adult↗
PubMed2005

When meaningless symbols become letters: neural activity change in learning new phonograms.

Left fusiform gyrus and left angular gyrus are considered to be respectively involved with visual form processing and associating visual and auditory (phonological) information in reading. However, there are a number of studies that fail to show the contribution of these regions in carrying out these aspects of reading. Considerable differences in the type of stimuli and tasks used in the various studies may account for the discrepancy in results. This functional magnetic resonance imaging (fMRI) study attempts to control aspects of experimental stimuli and tasks to specifically investigate brain regions involved with visual form processing and character-to-phonological (i.e., simple grapheme-to-phonological) conversion processing for single letters. Subjects performed a two-back identification task using known Japanese, and previously unknown Korean, and Thai phonograms before and after training on one of the unknown language orthographies. Japanese subjects learned either five Korean or five Thai phonograms. Brain regions related to visual form processing were assessed by comparing activity related to native (Japanese) phonograms with that of non-native (Korean and Thai) phonograms. There was no significant differential brain activity for visual form processing. Brain regions related to character-to-phonological conversion processing were assessed by comparing pre- and post-tests of trained non-native phonograms with that of native phonograms and non-trained non-native phonograms. Significant differential activation post-relative to pre-training exclusively for the trained non-native phonograms was found in left angular gyrus. In addition, psychophysiologic interaction (PPI) analysis revealed greater integration of left angular gyrus with primary visual cortex as well as with superior temporal gyrus for the trained phonograms post-relative to pre-training. The results suggest that left angular gyrus is involved with character-to-phonological conversion in letter perception.

Adult↗
PubMed2005

Optical imaging in cat area 18: strabismus does not enhance the segregation of ocular dominance domains.

While early-onset strabismus leads to clearly segregated domains of the left and the right eye in cat primary visual cortex (area 17), far less is known about experience-dependent plasticity of ocular dominance in area 18. We therefore used optical imaging of intrinsic signals to analyze the influence of strabismus on cortical maps in cat area 18. Monocular visual stimulation of the left and right eye with moving square wave gratings of four different orientations induced patchy activity maps. Unlike our previous observations in cat area 17, the monocular activity maps in area 18 of strabismic cats were rather similar so that functional ocular dominance domains were not clearly segregated. Imaging of the 17/18 border region confirmed this observation and revealed a sudden change in the segregation of the left and right eye domains across the border. Our results demonstrate that modified visual input can have different consequences for different visual areas: while the decorrelation of activity between the two eyes (as induced by strabismus) clearly enhances the segregation of ocular dominance domains in cat area 17, area 18 does not show this effect although electrophysiological studies have confirmed that the percentage of binocularly driven neurons is as reduced as in area 17.

Animals↗
PubMed2006

Orientation selectivity in the common marmoset (Callithrix jacchus): the periodicity of orientation columns in V1 and V2.

Orientation selectivity is a ubiquitous property of the primary visual cortex of mammals. Within the primate, orientation selectivity is arranged into vertical columns that are organized into a regular patchy pattern. Previous studies, in old world primates, have noted an anisotropy in this arrangement that appears to be due to the presence of ocular dominance columns within the same tissue. In addition, orientation selective responses appear to be arranged into bands of activity within the adjoining extrastriate region V2. Little is known about the precise arrangement of orientation columns within V2. In this study, we examined the layout of orientation columns within both V1 and V2 of a new world primate, the common marmoset, using optical imaging. New world primates have the advantage that, unlike the macaque, V2 exists on the cortical surface, a requirement for this form of optical mapping. We found the arrangement of orientation columns to be isotropic within marmoset V1 with an average repeat distance of around 575 mum, smaller than the repeat distance previously reported for the macaque. We found no evidence of ocular dominance within the animals tested supporting the claim that ocular dominance columns when present distort the mapping of orientation in V1. In V2 we found that orientation columns were larger and as in other primates were represented in discrete bands throughout V2. Orientation columns were spaced on average around 1 mm apart. This suggests that, at least in the marmoset, the visual system maps orientation at a different scale within V1 and V2.

Animals↗
PubMed2006

Causal visual interactions as revealed by an information theoretic measure and fMRI.

In the present study, we evaluated the direction of the effective connectivity between fMRI activations in neural structures mediating preserved visual function in a patient with homonymous hemianopsia due to a posterior cerebral artery stroke. Although the lesion affected the primary visual cortex, the visual abilities of this patient included above-chance verbal reports of movement and color change as well as the discrimination of movement direction in his hemianopic field. These abilities were coupled with awareness (Riddoch syndrome). The strength and the direction of the interactions between visual regions were assessed by applying directed transinformation (T), a nonparametric information theoretic causal measure sensitive to linear as well as to nonlinear interactions. In the healthy hemisphere, T identified a strong flow of information from visual area V1 to V5 during stimulation by visual movement and from V1 to V4/V8 during stimulation by color change. In addition, during color change stimulation, a bi-directional flow was observed between V4/V8 and V5, suggesting crosstalk between these regions. In the lesioned hemisphere, the color change stimulation evoked a stronger flow from V5 to V4/V8 and a flow from V4/V8 to V2. These observations provide support for the hypothesis that visual information is mediated via subcortical pathways that bypass V1 and project first to higher-tier visual areas V5 and V4/V8 then subsequently to lower-tier area V2.

Adult↗
PubMed2006

Reduction in V1 activation associated with decreased visibility of a visual target.

The perception of a brief visual target stimulus can be affected by another visual mask stimulus immediately preceding or following the target. The link of this visual masking illusion, with visual cortical activation, offers insights into the neural mechanisms for visual perception. The present study investigated the association of the visibility of a target with cortical activation in humans using psychophysical testing and functional magnetic resonance imaging (fMRI). A visual masking protocol that was suitable for an fMRI study was developed. The event-related fMRI was used to measure activation in primary visual cortex (V1) during visual masking and unmasking stimulation. We found that the visibility of the target stimulus was reduced in the masking condition, due to the presence of mask stimuli, but not in the unmasking condition. We also found that the activation in V1 was modulated by the temporal separation of the mask stimuli from the target and was associated with the visibility of the target that was recorded during psychophysical testing and fMRI. These findings are consistent with what has been observed in the primate visual cortex of monkeys, i.e., the transient on-response and after-discharge of V1 neurons to the target stimulus were suppressed by forward and backward mask stimuli, respectively.

Data Interpretation, Statistical↗
PubMed2004

Dendritic spine dynamics are regulated by monocular deprivation and extracellular matrix degradation.

The mammalian primary visual cortex (V1) is especially susceptible to changes in visual input over a well-defined critical period, during which closing one eye leads to a loss of responsiveness of neurons to the deprived eye and a shift in response toward the open eye. This functional plasticity can occur rapidly, following even a single day of eye closure, although the structural bases of these changes are unknown. Here, we show that rapid structural changes at the level of dendritic spines occur following brief monocular deprivation. These changes are evident in the supra- and infragranular layers of the binocular zone and can be mimicked by degradation of the extracellular matrix with the tPA/plasmin proteolytic cascade. Further, monocular deprivation occludes a subsequent effect of matrix degradation, suggesting that this mechanism is active in vivo to permit structural remodeling during ocular dominance plasticity.

Animals↗
PubMed2005

Adaptation of oriented and unoriented color-selective neurons in human visual areas.

Primary visual cortex contains at least two distinct populations of color-selective cells: neurons in one have circularly symmetric receptive fields and respond best to reddish and greenish light, while neurons in another have oriented receptive fields and a variety of color preferences. The relative prevalence and perceptual roles of the two kinds of neurons remain controversial, however. We used fMRI and a selective adaptation technique to measure responses attributable to these two populations. The technique revealed evidence of adaptation in both populations and indicated that they each produced strong signals in V1 and other human visual areas. The activity of both sets of neurons was also reflected in color appearance measurements made with the same stimuli. Thus, both oriented and unoriented color-selective cells in V1 are important components of the neural pathways that underlie perception of color.

Adaptation, Physiological↗
PubMed2005

Spatiotemporal elements of macaque v1 receptive fields.

Neurons in primary visual cortex (V1) are commonly classified as simple or complex based upon their sensitivity to the sign of stimulus contrast. The responses of both cell types can be described by a general model in which the outputs of a set of linear filters are nonlinearly combined. We estimated the model for a population of V1 neurons by analyzing the mean and covariance of the spatiotemporal distribution of random bar stimuli that were associated with spikes. This analysis reveals an unsuspected richness of neuronal computation within V1. Specifically, simple and complex cell responses are best described using more linear filters than the one or two found in standard models. Many filters revealed by the model contribute suppressive signals that appear to have a predominantly divisive influence on neuronal firing. Suppressive signals are especially potent in direction-selective cells, where they reduce responses to stimuli moving in the nonpreferred direction.

Action Potentials↗
PubMed2005

Cortical maps: where theory meets experiments.

Primary visual cortex (V1) has remarkably systematic functional maps. One commonly used class of computational models proposes that such maps are generated by a mechanism that projects the multiple dimensions of neuronal responses smoothly onto the two dimensions of cortex. In this issue of Neuron, Mriganka Sur and colleagues find a close match between such model predictions and measurements from ferret V1.

Animals↗
PubMed2006

The parvocellular LGN provides a robust disynaptic input to the visual motion area MT.

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

Animals↗
PubMed2006

Envisioning the reward.

The primary visual cortex (area V1) is for vision. At least, that is what most researchers believe. However, in a recent issue of Science, Shuler and Bear demonstrate a correlate of reward timing in area V1. This surprising result indicates that brain circuits for reward processing are more extensive than expected and that area V1 has more functionality than previously thought.

Animals↗
PubMed2006

Attentional shifting and the role of the dorsal pathway in visual word recognition.

A substantial amount of evidence has been collected to propose an exclusive role for the dorsal visual pathway in the control of guided visual search mechanisms, specifically in the preattentive direction of spatial selection [Vidyasagar, T. R. (1999). A neuronal model of attentional spotlight: Parietal guiding the temporal. Brain Research and Reviews, 30, 66-76; Vidyasagar, T. R. (2001). From attentional gating in macaque primary visual cortex to dyslexia in humans. Progress in Brain Research, 134, 297-312]. Moreover, it has been suggested recently that the dorsal visual pathway is specifically involved in the spatial selection and sequencing required for orthographic processing in visual word recognition. In this experiment we manipulate the demands for spatial processing in a word recognition, lexical decision task by presenting target words in a normal spatial configuration, or where the constituent letters of each word are spatially shifted relative to each other. Accurate word recognition in the Shifted-words condition should demand higher spatial encoding requirements, thereby making greater demands on the dorsal visual stream. Magnetoencephalographic (MEG) neuroimaging revealed a high frequency (35-40Hz) right posterior parietal activation consistent with dorsal stream involvement occurring between 100 and 300ms post-stimulus onset, and then again at 200-400ms. Moreover, this signal was stronger in the shifted word condition, compared to the normal word condition. This result provides neurophysiological evidence that the dorsal visual stream may play an important role in visual word recognition and reading. These results further provide a plausible link between early stage theories of reading, and the magnocellular-deficit theory of dyslexia, which characterises many types of reading difficulty.

Attention↗