Serum lipid fatty acids and temporal processing acuity in children with oral clefts.

We investigated the relation between a biological factor (fatty acids, FA) and a cognitive processing speed factor (temporal processing acuity, TPA) that are both suggested to relate to neuronal and cognitive functioning. Blood samples of 49 ten-year-old children with oral clefts were collected for FA analysis in serum triglycerides, cholesteryl esters, and phospholipids on the same day as they performed behavioral TPA tasks (simultaneity/nonsimultaneity judgments) in several perceptual modalities (visual, auditory, tactile, audiotactile, visuotactile, and audiovisual). This population has larger than expected variation in the relevant cognitive measures (TPA, learning ability, and intelligence). Sequential regression analyses (adjusted for age, gender, and cleft type) showed that saturated FAs were not generally associated with TPA. Monounsaturated erucic and nervonic acids were inversely related with TPA. Of the polyunsaturated fatty acids, eicosapentaenoic and docosahexaenoic acids were positively associated with TPA, whereas gamma-linolenic acid was inversely related to TPA. In summary, we found significant relations between a biological (certain FAs) and a cognitive factor (TPA).

Serum lipid fatty acids, phonological processing, and reading in children with oral clefts.

Reading skill is suggested to be related to phonological processing ability and polyunsaturated fatty acids (PUFAs). Here we investigated whether fatty acids (FAs) are related to phonological processing, whether the relations between PUFAs and reading generalize to other FAs, whether these relations are mediated by phonological processing, and whether relations of FAs are specific for language-related functions. Blood samples of 49 ten-year-old children with oral clefts were collected for FA proportion analysis in serum cholesteryl esters and phospholipids. On the same day, they performed tasks of phonological processing, reading, and both verbal and nonverbal intelligence. Sequential regression analyses (adjusted for age, gender, and cleft type) showed that phonological processing was inversely related to myristic acid in phospholipids and positively related to eicosapentaenoic acid in cholesteryl esters. Reading was inversely related to palmitoleic and gammalinolenic acids in phospholipids. The relations between FAs and reading were not mediated by phonological processing and FAs related only to language-related functions.

Temporal order and processing acuity of visual, auditory, and tactile perception in developmentally dyslexic young adults.

We studied the temporal acuity of 16 developmentally dyslexic young adults in three perceptual modalities. The control group consisted of 16 age- and IQ-matched normal readers. Two methods were used. In the temporal order judgment (TOJ) method, the stimuli were spatially separate fingertip indentations in the tactile system, tone bursts of different pitches in audition, and light flashes in vision. Participants indicated which one of two stimuli appeared first. To test temporal processing acuity (TPA), the same 8-msec nonspeech stimuli were presented as two parallel sequences of three stimulus pulses. Participants indicated, without order judgments, whether the pulses of the two sequences were simultaneous or nonsimultaneous. The dyslexic readers were somewhat inferior to the normal readers in all six temporal acuity tasks on average. Thus, our results agreed with the existence of a pansensory temporal processing deficit associated with dyslexia in a language with shallow orthography (Finnish) and in well-educated adults. The dyslexic and normal readers' temporal acuities overlapped so much, however, that acuity deficits alone would not allow dyslexia diagnoses. It was irrelevant whether or not the acuity task required order judgments. The groups did not differ in the nontemporal aspects of our experiments. Correlations between temporal acuity and reading-related tasks suggested that temporal acuity is associated with phonological awareness.

Rate of information segregation in developmentally dyslexic children.

Slowed processing of sequential perceptual information is related to developmental dyslexia. We investigated this unimodally and crossmodally in developmentally dyslexic children and controls ages 8-12 years. The participants judged whether two spatially separate trains of brief stimuli, presented at various stimulus onset asynchronies (SOA) in one or two senses, were synchronous or not. The stimulus trains consisted of light flashes in vision, clicks in audition, and indentations of the skin in the tactile sense. The dyslexic readers required longer SOAs than controls for successful performance in all six comparisons. The crossmodal spatiotemporal resolution of the groups differed more than unimodal performance. The dyslexic readers' segregation performance was also less differentiated than that of the controls. Our results show that not only sensory but also polysensory nonverbal information processing is temporally impaired in dyslexic children.

Mu rhythm modulation during changes of visual percepts.

Cooperation between vision and somatomotor behavior, such as manual exploration of objects, suggests close functional coupling between the visual and sensorimotor systems. We observed this type of interaction in human volunteers during binocular rivalry while following the level of sensorimotor mu rhythm with a whole-scalp neuromagnetometer. The observers viewed a weak vertical grating in the lower visual field of one eye and a strong horizontal grating in the same spatial window of the other eye. When stationary, the weak grating was permanently invisible because of its low contrast and spatial frequency. A sudden brief drifting movement of the weak grating wiped out the dominant grating, and the weak grating became visible for less than the 3-s interval between the movements. The postcentral 8- to 15-Hz mu rhythm was found in six of nine observers, and its level increased transiently by 10-15%, starting about 450 ms after the beginning of the movement. The mu level was also enhanced by the actual disappearance of the stronger stimulus, when it occurred in random order with the rivalry stimuli. Identical visual motion, when not accompanied by a perceptual dominance change, produced only minor effects on the mu rhythm. Our results show that a change in visual percept, even with no real or imagined motor response, is associated with modified activity of the postcentral gyrus. This modification may reflect visuohaptic interactions and/or activity of the distributed cortical network implementing visually guided movements.

Stronger occipital cortical activation to lower than upper visual field stimuli. Neuromagnetic recordings.

We recorded whole-scalp magnetoencephalographic (MEG) responses to black-and-white checkerboards to study whether the human cortical responses are quantitatively similar to stimulation of the lower and upper visual field at small, 0-6 degrees, eccentricities. All stimuli evoked strong occipital responses peaking at 50-100 ms (mean 75 ms). The activation was modeled with a single equivalent current dipole in the contralateral occipital cortex, close to the calcarine fissure, agreeing with an activation of the V1/V2 cortex. The dipole was, on average, twice as strong to lower than to upper field stimuli. Responses to hemifield stimuli that extended to both lower and upper fields resembled the responses to lower field stimuli in source current direction and strength. These results agree with psychophysical data, which indicate lower visual field advantage in complex visual processing. Parieto-occipital responses in the putative V6 complex were similar to lower and upper field stimuli.

Activation of human V5 complex and rolandic regions in association with moving visual stimuli.

We recorded magnetoencephalographic responses from seven healthy humans during the presentation of stationary and rotating radial gratings. Rotations lasting 1 s evoked movement-specific sustained activity in the parieto-occipitotemporal border area, in agreement with the activation of the V5 complex specialized for the analysis of movement. The source areas of the movement-specific sustained fields were transiently active 100-130 ms after the onsets of both rotating and stationary stimuli, suggesting that movement-related cortical areas respond to any transient changes in the visual environment. Transients were evoked also in other brain areas 60-200 ms after onsets of both stimuli. Four subjects displayed additional motion-related sustained activity in the rolandic region. Sustained activity continued after the stimulus movement in several subjects during perception of the movement aftereffect. The transient activity may evoke visual attention while sustained activity of the V5 complex may be related to the conscious perception of movement.

Cortical magnification, scale invariance and visual ecology.

The visual world of an organism can be idealized as a sphere. Locomotion towards the pole causes translation of retinal images that is proportional to the sine of eccentricity of each object. In order to estimate the human striate cortical magnification factor M, we assumed that the cortical translations, caused by retinal translations due to the locomotion, were independent of eccentricity. This estimate of M agrees with previous data on magnifications, visual thresholds and acuities across the visual field. It also results in scale invariance in which the resolution of objects anywhere in the visual field outside the fixated point is about the same for any viewing distance. Locomotion seems to be a possible determinant in the evolution of the visual system and the brain.

Illusory perception of gratings stimulating a small number of neurones.

We studied pattern perceptions caused by drifting gratings presented monocularly in the nasal and temporal visual fields at various suprathreshold contrasts. The grating and its surround and background were matched in luminance. Small grating produced illusions and reduced perceptions. When grating area or contrast increased from a subthreshold value, the gratings were first seen as mere flashes. Then each grating was sometimes perceived as a single small bright spot or point. Next each grating was seen as a single dark or bright line. Finally the stimuli were perceived as gratings consisting of several bars. Orientation or direction of movement were perceived correctly, but velocity, colour and number of bars were often perceived as illusions. Thus, in spite of the illusions, some features of the stimuli could have allowed correct discriminations. The area and contrast limits of illusory perception depended on eccentricity. Irrespective of retinal size, the stimuli were not perceived correctly as gratings at any eccentricities when the gratings were smaller than about 1 x 1 mm in their calculated cortical area and stimulated a small constant number of retinal ganglion cells. Relations between the results and retinal aliasing, cortical columns and phase locking of neuronal oscillations are discussed.

Analysis of spatial structure in eccentric vision.

The analysis of spatial structure, ie, the encoding of relative positions between pattern elements, was studied in central and eccentric vision. In a two-alternative forced-choice task the observer had to discriminate between two patterns consisting of short line segments. At each trial the two patterns were flashed for 140 msec and the observer indicated whether the patterns were identical or mirror symmetric. Psychometric functions were measured by changing pattern size at each eccentricity in order to find the threshold size allowing 75% of correct responses. The scaling factor, required for discriminating between mirror symmetric and identical patterns independent of eccentricity, was found to be similar to the size-scaling proposed by Levi et al (Vision Res 25:963, 1985) for vernier acuity tasks.

Cortical magnification and peripheral vision.

In a generalized form, the cortical magnification theory of peripheral vision predicts that the thresholds of any visual stimuli are similar across the whole visual field if the cortical stimulus representations calculated by means of the cortical magnification factor are similar independently of eccentricity. Failures of the theory in spatial vision were analyzed, and the theory was tested with five visual acuity tasks and two hyperacuity tasks. Almost all increases in thresholds with eccentricity were explained by the theory in five of these tasks, which included the two-dot vernier hyperacuity test, the measurement of visual acuities with gratings, the Snellen E test, and two acuity tests that required either separation between dots or discrimination between two mirror-symmetric forms. The two-dot vernier thresholds could be explained as a special case of orientation discrimination, and orientation discrimination at different eccentricities was in agreement with the cortical magnification theory. The increase of thresholds in peripheral vision was larger than predicted by the theory in the Landolt visual acuity and bisection hyperacuity tests, possibly because of retinal undersampling.

Texture discrimination at different eccentricities.

Differences in preattentive texture discrimination between central vision and peripheral vision were studied with textures composed of random dots. The subject had to discriminate between two textures whose first-order statistics were kept identical but whose second-order statistics were different. For textures of constant retinal size the discrimination was easy in central vision, but the decrease of visual acuity with increasing eccentricity made the textures unresolvable in peripheral vision. When the textures were scaled by the cortical magnification factor derived from the frequency of retinal ganglion cells so that the calculated neural representations of the textures became similar at different eccentricities, texture discrimination became independent of visual field location. This indicates that preattentive texture discrimination based on differences in second-order statistics of random dots operates similarly in central vision and peripheral vision.

Mesopic spectral responses and the Purkinje shift of macaque lateral geniculate nucleus cells.

Spectral responsiveness of different classes of macaque LGN cells at eccentricities smaller than 12 degrees was studied at low light intensities. Cone thresholds of cells varied from 1 to 10 td. Rod inputs were found in all classes of cells, including inhibitory inputs in some cells. Rod inputs were not apparent above 10-40 td, giving a total mesopic range of about 1-40 td. Strong rod-mediated responses could be evoked in broadband phasic cells and in spectrally opponent cells excited by short wavelengths. Only weak if any excitatory responses could be evoked by short wavelengths at scotopic levels in spectrally opponent long-wavelength excited cells. Hence, rod inputs do not confound the spectral responsiveness of cells because no spectrally opponent cell excited by long-wavelength stimuli at photopic levels became significantly responsive to short wavelength stimuli at mesopic or scotopic intensities. The so-called "rod color" may be blue. An increase in the dominance of wide-band cell responses that may explain the Bezold-Brücke hue shift was observed at higher stimulus radiances at wavelengths near 450 and 650 nm.

Temporal integration and contrast sensitivity in foveal and peripheral vision.

Spatial contrast sensitivity functions and temporal integration functions for gratings with dark surrounds were measured at various eccentricities in photopic vision. Contrast sensitivity decreased with increasing eccentricity at all exposure durations and spatial frequencies tested. The decrease was faster at high than at low spatial frequencies, but similar at different exposure durations. When cortically similar stimulus conditions were produced at different eccentricities by M-scaling, contrast sensitivity became independent of visual field location at all exposure durations tested. The results support the view that in photopic vision spatiotemporal information processing is qualitatively similar across the visual field, and that quantitative differences result from retino-topical differences in ganglion cell sampling. For gratings of constant retinal area temporal integration (improvement of contrast sensitivity with increasing exposure duration) was more extensive at high than at low retinal spatial frequencies but independent of cortical spatial frequency and eccentricity. For M-scaled gratings temporal integration was more extensive at high than at low cortical spatial frequencies but independent of retinal spatial frequency and eccentricity. The results suggest that the primary determinant of temporal integration is not spatial frequency but grating value that is calculated as AF2 square cycles (cycle2), where A is grating area and F spatial frequency.

Cell responses in dorsal layers of macaque lateral geniculate nucleus as a function of intensity and wavelength.

We studied the relationship between light intensity and cell response to various wavelengths and wavelength combinations in the dorsal, parvocellular layers of the macaque lateral geniculate nucleus. When response is plotted as a function of the logarithm of stimulus intensity, the slope and shape of curves depends on wavelength. For wavelengths near the crossover point between excitatory and suppressive responses, nonmonotonic curves are common. Consequently, the form of spectral-response functions depends on stimulus intensity. Responses to combined stimuli made up of wavelengths close together near one spectral extreme are approximately additive. If one wavelength is near the crossover point, responses are nonadditive so that a midspectral wavelength, only producing a weak excitatory response, is able to occlude more vigorous responses to wavelengths near the spectral ends. Responses of parvocellular layer cells are consistent with their being a result of linear interaction of opponent cone mechanisms, the response of each of which follows a modified hyperbolic tangent function (22). Responses to all wavelength combinations, even those showing strikingly nonadditive effects, could be predicted from the additive opponent model described above.

Light adaptation in cells of macaque lateral geniculate nucleus and its relation to human light adaptation.

Responses of macaque lateral geniculate nucleus (LGN) cells to stimuli of different incremental intensities and wavelength compositions were studied at different levels of light adaptation from scotopic to low photopic levels. Stimuli were large in comparison with receptive-field size. Human increment thresholds were measured for comparison. The strength of responses grew in many cells from threshold up to a saturation level as a logarithmic function of incremental intensity. More complex intensity-response functions were also obtained, particularly from parvocellular layer (PCL) cells, but the shape and slope of the intensity-response function changed as a function of adaptation level only with chromatic backgrounds. As a function of adaptation level, the intensity-response functions shifted along the logarithmic abscissa but not sufficiently for a complete contrast constancy. Thus responses to any constant contrast became smaller when adaptation level decreased. The change from cone to rod responses, when possible, took place without noticeable change in shape of intensity-response functions, and much of the adaptive shift of the functions could be attributed to the change-over between rods and cones. Differences between different cells in light adaptation and dark-adapted sensitivity were large, mostly because of variation in the strength of rod input. The strongest excitatory rod inputs were found in PCL cells activated by short-wavelength light, so that the highest sensitivity at low levels of illumination occurred in blue- and blue-green-sensitive cells. The lowest increment thresholds based on cones matched the thresholds of macaque cone late receptor potentials recorded by Boynton and Whitten (3). They were also similar to human cone thresholds measured psychophysically but only for small stimulus sizes that may approximate the size of the receptive-field centers. Human sensitivity was much higher when measured with large stimulus sizes, indicating integration at post-geniculate neural levels. Light adaptation, as evaluated with respect to contrast constancy and Weber law behavior, was similarly incomplete in monkey single cells and human perception. A few cat LGN cells were studied in a control experiment; results agreed with previous findings. The light adaptation of cat cels was more complete and sensitivity higher than those observed under comparable conditions in macaque single cells and human. The maintained activity level of cells was little affected by the intensity of steady backgrounds. Thus, the steady-state hyper-polarisation of receptors was not transmitted to LGN cells.

Linear signal transmission from prepotentials to cells in the macaque lateral geniculate nucleus.

Prepotentials preceding a neuronal action potential were recorded extracellularly in the lateral geniculate nucleus (LGN) of the macaque. Although prepotentials are found less frequently in the macaque than in the cat LGN, their electrical characteristics are similar, suggesting that they represent the arrival of impulses in a retinal afferent, as in the cat. The visual response properties of prepotentials and associated cells were similar under a variety of conditions, indicating that, apart from some response attenuation, little signal processing takes place in macaque LGN. A constant fraction of prepotentials above a threshold frequency gave rise to neuronal action potentials independent of the stimuli used, so that the frequency of cell action potentials was linearly related to the frequency of prepotentials. Since the maintained discharge rates of a cell and its prepotential always fell on the linear relation, the net responses of a cell and its prepotential to visual stimuli were approximately proportional to one another.