Search PubMedSearch

Biomedical subjects

C Blakemore

Publications and source records attributed to C Blakemore.

At least 19 recordsLinked to original sources

EPSPs in rat neocortical pyramidal neurones in vitro are prolonged by NMDA receptor-mediated currents.

We investigated the influence of 2-amino-5-phosphonovalerate (APV), a selective antagonist of the N-methyl-D-aspartate (NMDA) receptor, on the time course of small excitatory postsynaptic potentials (EPSPs) in pyramidal neurones in layer 2/3 of adult rat visual cortex in vitro. Time constants of the voltage decay following the EPSPs (T(s)) and after brief (2 ms) pulses of current injected at the soma (T(p)) were determined from semilogarithmic plots of averages of 100-250 trials. The mean T(s)/T(p) ratio decreased from 1.53 +/- 0.29 (S.D.) to 1.10 +/- 0.08 on addition of 50 microM APV to the bathing medium (P less than 0.001; n = 23), but there was no significant change in EPSP peak amplitude or rise-time. These results suggest that the time course of many small EPSPs, even at negative membrane potentials and in the presence of Mg2+, can be prolonged by NMDA receptor-mediated currents.

2-Amino-5-phosphonovalerate

Lack of regional specificity for connections formed between thalamus and cortex in coculture.

The mammalian cerebral cortex consists of many structurally and functionally specialized areas, with characteristic input from particular nuclei of the thalamus. Some localized external influence, such as the arrival of fibres from the appropriate thalamic nucleus before or around the time of birth, could trigger the emergence of committed cortical fields from an undifferentiated 'protocortex. The guidance of axons from each thalamic nucleus to its appropriate target area in the cortex could, then, be crucial in the regulation of cortical differentiation. Recently, Yamamoto et al. and Bolz et al. have demonstrated that cocultured explants of rat lateral geniculate nucleus and visual cortex can form layer-specific interconnections. We have now tested the possibility that each cortical area exerts a selective trophic influence on axons from its appropriate thalamic nucleus, and vice versa, by coculturing explants of different regions of the thalamus and cortex taken at various stages of development. Although thalamo-cortical and cortico-thalamic connections formed in vitro can be remarkably normal in many respects, they lack regional specificity.

Animals

Sensitive and vulnerable periods in the development of the visual system.

In advanced mammals the visual system consists of a number of parallel channels for the efficient processing of different aspects of the visual scene. Much of the basic anatomical structure of the visual pathway is constructed before birth. A wave of maturation sweeps through the system, from the eye to the visual cortex, the correct formation of connections depending on precisely timed interactions between axons and their targets. Competition between growing axons (apparently dependent on spontaneous impulse activity in those axons), cell death (partly influenced by competition between those cells' axons), axon withdrawal, trophic interactions--these and other mechanisms play a part in constructing the visual pathway and laying down basic 'maps' of the visual field before birth. Disturbances in such processes might underlie disorders of the genesis of the nervous system. At the level of the visual cortex, synaptic plasticity continues after birth and may permit cortical neurons to refine their processing capacities on the basis of information provided by the visual environment. This makes the young animal vulnerable to disturbances of visual experience early in life, which can cause virtually irreversible deficits in stereoscopic vision, visual resolution and sensitivity to contrast (amblyopia) in adult life.

Animals

Development of stimulus selectivity and functional organization in the suprasylvian visual cortex of the cat.

We have recorded from single neurons in the medial bank of the middle suprasylvian sulcus (PMLS) of anaesthetized and paralysed cats aged between nine days and eight weeks. Visual responses were assessed qualitatively, by using conventional projected stimuli, and quantitatively for drifting, high-contrast gratings of optimum spatial and temporal frequencies, but varying in orientation and direction of drift. At 9 days of age, some cells in the PMLS were spontaneously active but in three long penetrations only one visually responsive neuron was isolated. Between 9 and 15 days there was a rapid increase in the proportion of responsive units, which first appeared in small clusters in the lower layers (IV, V, VI). During the second and third postnatal weeks, spontaneous activity and the strength of visual responses increased to adult levels, and the proportion of cells showing rapid habituation to visual stimulation decreased. Even before two weeks of age, at least 85% of responsive cells in the PMLS were selective, by quantitative criteria, for image motion along one particular axis, and a majority of these were clearly direction-selective (responding to movement in one direction significantly more strongly than to that in the opposite). By the end of the third postnatal week the proportion of units with strong direction preference reached adult levels. The selective cells were initially more broadly 'tuned', on average, for the direction of motion of a grating (mean half-width in animals of 10-12 days was 32.6 degrees), but the sharpness of tuning improved to reach the adult level (ca. 23 degrees) during the third postnatal week. In animals younger than three weeks a slightly smaller proportion of cells than in adults (but always more than one third of all visually responsive cells) responded to stationary, contrast-modulated gratings. The majority of these cells showed clear selectivity for the orientation of a flashed grating. A few 'non-selective' cells were found in the youngest animals but by the end of the third postnatal week virtually all cells responsive to stationary gratings displayed orientation selectivity. There was always good agreement between the preferred orientations for stationary and drifting gratings. Even before two weeks of age, when responsive cells occurred only in small clusters, there was a clear tendency for neighbouring neurons to have similar or opposite preferred directions, just as in adult cats. By 2-3 weeks of age there were clear progressive shifts in stimulus preference along oblique or tangential tracks.(ABSTRACT TRUNCATED AT 400 WORDS)

Age Factors

Developmental plasticity in the visual and auditory representations in the mammalian superior colliculus.

Environmental factors play an important role in certain aspects of the development of sensory systems. But the way in which the maturation of different sensory modalities is coordinated is poorly understood. We have investigated this question neurophysiologically in the mammalian superior colliculus (SC), which contains topographically aligned maps of visual and auditory space. We report here that an essentially normal auditory map, in approximate register with the visual map, is found in the SC of adult ferrets reared with abnormal binaural localization cues. Also, if, early in life, one eye is deviated laterally, there is a compensatory shift in the auditory map, but early eye rotation totally disorders the auditory representation. These results imply that development of the auditory map is affected by visual activity or by information about eye position and that there is definite, but limited, capacity for the auditory map to reorganize so that it remains aligned with the visual map.

Acoustic Stimulation

The organization of corticocortical projections from area 17 to area 18 of the cat's visual cortex.

Retrogradely transported tracers were injected into area 18 of the visual cortex of the adult cat to study the organization of corticocortical projections from area 17 to area 18. All injections, whether very small or relatively large, and irrespective of their exact location in area 18, produced a discontinuous, clustered distribution of labelled cells, mainly in layers II, III and upper IV, in a topographically related region of area 17. The mean centre-centre distance between neighbouring patches was about 750 microns. We conclude that the overall population of cells projecting to area 18 is genuinely distributed in a patchy fashion and that they provide an efficient spatial sample of information from area 17. Comparison of the dimensions of each injection site and of the retrogradely labelled territory suggested that each region in area 18 receives a convergent input from a zone in area 17 whose visual field representation is about 0.8 M-1 deg larger in all directions (where M is the magnification factor in millimetres per degree at the termination site in area 18). Pairs of injection were made in area 18 by placing small volumes of two fluorescent tracers, fast blue and diamidino yellow, side-by-side in either a rostrocaudal or a mediolateral plane, with different distances between them. When the boundaries of the dense central cores of two injection sites were separated, at their closest points, by about 1.6 mm, the two corresponding distributions of labelled cells in area 17 were just non-overlapping, suggesting that each group of cells in area 17 sends a divergent projection in innervate a zone about 0.8 mm larger in all directions in area 18. More closely spaced injections led to overlap of the distributions of labelling by the two dyes, with shared clusters containing a mixture of labelled cells. The proportion of double-labelled cells in these shared clusters never exceeded 4.4% (but was 70% after sequential injection of the two dyes at a single point). We conclude that, although each cluster of cells sends a divergent projection to area 18, the majority of individual axons terminate more discretely, perhaps providing specific inter-connections between functionally corresponding 'columns' in the two areas.

Animals

Factors limiting the postnatal development of visual acuity in the monkey.

A major factor underlying the prolonged postnatal improvement of visual acuity in primates is an increase in the sampling limit imposed by the photoreceptor mosaic, which may be as much as 5-fold. Further factors operating peripheral to neurons of the lateral geniculate nucleus, which might include changes in neural connectivity, appear to contribute at most 1.5 octaves, between 3 weeks and 6 months of age. Neural factors at the geniculo-cortical synapse may add another half-octave between 10 weeks and 1 yr. Factors operating after the level of the striate cortex (changes in spatial degradation, attention, motivation, etc.) contribute up to another octave or more to the increase in behavioural acuity between birth and about 11 weeks of age. The foveal image is substantially undersampled in young animals and therefore aliasing could occur over a wide range of spatial frequencies.

Animals

The in vitro slice preparation for combined morphological and electrophysiological studies of rat visual cortex.

The morphological condition of slices of rat visual cortex, maintained in vitro in an interface-type recording chamber, was assessed. In addition, neurones in these slices were impaled with glass micropipettes for intracellular recording and horseradish peroxidase (HRP) injection. After fixation and embedding, slices were examined by light and electron microscopy. Slices sectioned orthogonally to the original plane of cutting showed a vertical zonation of tissue preservation. The upper zone contained dense and flattened neuronal somata, although the neuropil appeared normal. The central zone was well preserved, with the appearance of most somata, dendrites, axons and synapses comparing favourably with perfusion-fixed material. The lower zone contained many abnormal, vacuolated somata. The morphology of HRP-injected neurones was assessed by light microscopy. Dendrites could be visualised in great detail and spines were clearly visible. Local axon arbors were well represented. There was good correlation between electrophysiological and morphological criteria for the assessment of the condition of the slice. We conclude that, provided the extent of degeneration within the slice is monitored and appreciated, slices of visual cortex can provide both electrophysiological and morphological data of high quality.

Animals

Development of spatial and temporal selectivity in the suprasylvian visual cortex of the cat.

We have studied the development of the spatial and temporal properties of neurons in the medial bank of the suprasylvian visual cortex (PMLS) in kittens aged between 9 d and 8 weeks. Quantitative measurements were made of the responses to drifting high-contrast gratings of optimum orientation and direction of motion, but varying in spatial and temporal frequency. The spatial resolution ("acuity") of cells increased rapidly and was fully mature (over 2 cycles/deg for the best cells) at 3 weeks of age. The optimum spatial frequency also tended to improve and reached adult values (around 0.5 cycles/deg for the best cells) at about the end of the third week. In younger kittens, the spatial resolution of neurons was not obviously correlated with the eccentricity of their receptive fields, but in older animals acuity was clearly elevated for receptive fields in the central visual field. The proportion of "low-pass" cells (showing no obvious attenuation of response for gratings of low spatial frequency) decreased with age and simultaneously there was a slight increase in the mean spatial bandwidth of "bandpass" cells. Responses to drifting sinusoidal gratings were generally dominated by an unmodulated elevation of discharge at all ages. In tests with stationary, contrast-modulated gratings presented at different spatial positions, cells in the youngest kittens behaved nonlinearly and showed mainly an unmodulated increase in discharge, whereas in older kittens, as in adult cats, most neurons responded to contrast-modulated gratings with a small, phase-dependent response at the temporal frequency of modulation and a larger component at twice the fundamental frequency. None of the cells recorded at any age had a true "null position." As in adult PMLS, the widths of receptive fields in kittens were, on average, about twice the size of the preferred spatial period (4 times the preferred bar width). At all ages, therefore, neurons in PMLS resembled striate complex cells with respect to the nonlinearity of their responses and the spatial structure of their receptive fields. The preferred temporal frequency and high-temporal-frequency cutoff also improved, on average, during the first 3 weeks of life, and the range of temporal frequencies over which cells responded continued to increase until at least 8 weeks. Although the low-spatial-frequency inhibition that creates spatial bandpass characteristics probably depends on cortical mechanisms, the postnatal development of both temporal and spatial resolution might well be limited by maturation at the level of the retina.

Animals

The organization and post-natal development of area 18 of the cat's visual cortex.

1. We made extracellular recordings from 1176 single units in area 18 of adult cats and kittens aged between 7 days and 10 weeks, and from 137 single units in area 17 in kittens aged between 12 days and 10 weeks. 2. All cells examined in area 18 of adult cats were visually responsive, 84% being orientation selective, 9% orientation biased and 7% non-oriented. Orientation columns and ocular dominance columns were identified. There was an over-all bias towards horizontal and vertical in the distribution of preferred orientations in the rostral part of area 18, where we were recording. 3. In area 18 of 7-day-old, visually inexperienced kittens the majority of cells (60%) were visually unresponsive; the remainder were either non-oriented (25%) or orientation biased (15%), and no orientation-selective units were found. Nevertheless there seemed to be a rudimentary columnar system, even in the youngest animals, in that orientation-biased cells tended to occur in clusters with neighboring neurones having similar orientation preferences. In normal animals of 3 weeks and younger we found that the distribution of preferred orientations of a sample of neurones in area 18 with receptive fields scattered over much of the left lower quadrant of the visual hemifield was biased to oblique orientations. 4. As in adult cats, most cells in area 18 in young kittens were binocularly driven, and periodic alternation of dominance along oblique penetrations, characteristic of ocular dominance columns, was sometimes seen, even at the earliest ages. 5. Many of the developmental changes that we observed in area 18 occurred during the first 4 post-natal weeks. Orientation selectivity, orientation tuning, directionality and responsiveness of neurones matured rapidly over this period. The proportions of simple and complex cells were similar in kittens aged 4 weeks or more to those in adult cats, whereas prior to this most neurones that displayed an orientation preference appeared to be immature simple cells. 6. A laminar analysis revealed that very few units in the superficial layers (I, II and III) in area 18 are visually responsive in kittens during their first and second weeks, but orientation selectivity rapidly develops during the third week. By contrast, even in very young, visually inexperienced kittens, the majority of neurones found in deeper laminae (IV, V and VI) are visually responsive and a few of them already show an orientation preference; however, the subsequent appearance of larger proportions of orientation-selective cells in these lower layers is a more prolonged process than in upper laminae.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Effects of dark-rearing on the development of area 18 of the cat's visual cortex.

1. We recorded extracellularly from 420 single units in area 18 in visually inexperienced kittens aged 7 days and dark-reared kittens aged between 3 and 12 weeks and from 60 single units in area 17 in dark-reared kittens aged 5 and 11 weeks. 2. Visual deprivation generally depressed the maturation of area 18, although some features were affected more than others and certain developmental improvements still occurred. The percentage of visually responsive units in area 18 increased from 40% in 7-day-old kittens to about 75% in dark-reared animals 10-12 weeks. At each age a proportion of cells was orientation biased (between 15 and 45%) and these neurones appeared to be arranged in a crude columnar fashion. However, dark-rearing, from birth, prevented the development of a significant proportion of orientation-selective cells in area 18; no more than 5% of neurones were orientation-selective at any age. We found no major bias in the over-all distribution of preferred orientations of cells in area 18 in dark-reared kittens. 3. Simple cells, which are found in area 18 even in very young, visually inexperienced kittens, persisted after dark-rearing, although most retained immature properties. Relatively few complex cells were found in area 18 in visually deprived animals. 4. The majority of neurones in area 18 of dark-reared kittens were binocularly driven, many equally well by either eye; evidence for regional variation in ocular dominance (indicative of a columnar pattern) was found in these deprived animals. 5. A laminar analysis in area 18 showed that percentages of non-oriented and orientation-biased cells changed little, if at all, in lower laminae (IV, V, and VI) but increased substantially in upper layers (above layer IV) in the absence of visual stimulation, over the first 12 post-natal weeks. 6. A comparison of the effects of dark-rearing on areas 17 and 18 indicates that the normal development of visual responsiveness and specific receptive field properties is suppressed in both areas during the first 12 post-natal weeks. It is possible that area 17 has a greater degree of orientation selectivity than area 18 in young visually deprived kittens and this may reflect a difference in the type of afferent inputs.

Action Potentials

Stimulus selectivity and functional organization in the lateral suprasylvian visual cortex of the cat.

1. We have examined the responses of cells in the postero-medial and postero-lateral lateral suprasylvian areas (p.m.l.s./p.l.l.s.) in the medial and lateral banks of the middle suprasylvian sulcus of the anaesthetized, paralysed cat. 2. Visual responses were assessed qualitatively (for projected spot and bar stimuli) and quantitatively (for drifting, high-contrast gratings of optimum spatial and temporal frequencies, but varying in orientation and direction of drift). There was excellent agreement between qualitative and quantitative estimates of preferred direction of motion. 3. Comparison of responses to motion in the preferred direction and the opposite direction confirmed that the percentage of units with strong directional preference is higher in this region of cortex than in other cortical and subcortical visual structures so far investigated. 4. Cells were comparatively well 'tuned' for the direction of motion of a grating: on average the half-width at half-amplitude for the variation in response around the principal preferred direction was 23.2 deg for p.m.l.s. and 25.3 deg for p.l.l.s., thus falling within the range found for complex cells in area 17. In this and other aspects of their direction selectivity, neurones in p.m.l.s. and p.l.l.s. were very similar to each other. 5. Quantitative analysis of the direction-response functions revealed them frequently to be more complex than previously described. For many cells there were subsidiary response peaks in addition to the main peak at the principal preferred direction. Often there was an accessory peak 180 deg from the optimum direction (i.e. directional preference was incomplete), but there was also a clear tendency for responses to be specifically elevated for directions orthogonal to the principal direction. 6. Tests with stationary, contrast-modulated gratings of the optimum spatial frequency, but differing in orientation, revealed that neurones responsive to such stimuli (though limited to about half the population) were orientation selective, their preferred orientations for stationary and drifting gratings being very similar. 7. The functional architecture of the lateral suprasylvian cortex was studied by means of single and multiple penetrations at different angles to the cortical surface: in most penetrations in p.m.l.s. the preferred directions of cells tended to shift progressively in small steps across the cortex, at a maximum rate of about 360 deg mm-1. However, there were occasional 180 deg differences between neighbouring recording sites or between cells recorded at different depths within a single radial column.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Spatial and temporal selectivity in the suprasylvian visual cortex of the cat.

We recorded from single units in the medial and lateral banks of the posterolateral suprasylvian visual cortex (PMLS/PLLS) of the cat. The responses to drifting high-contrast gratings of optimum orientation and direction of motion, but varying in spatial and temporal frequency, were examined quantitatively for a sample of cells, whose receptive fields covered a wide range of eccentricities. The optimum spatial frequencies (average about 0.2 cycles/deg) were low compared to the values reported for striate cortex but similar to those for area 18. The mean spatial bandwidth (about 2 octaves) was slightly broader than that of cells in other cortical visual areas. The cut-off spatial frequencies ("acuities") covered a wide range, from 0.05 to 2.1 cycles/deg, similar to those of cells in area 18. Responses to drifting sinusoidal gratings were usually dominated by an unmodulated elevation of discharge, although some modulation occurred at the temporal frequency of drift, especially at low spatial frequencies. Modulated responses were relatively stronger in PMLS than in PLLS. For those cells that responded to flashed stimuli, stationary, contrast-modulated gratings presented at different spatial positions typically evoked small responses at the fundamental frequency (dependent on spatial phase) and a larger component at the second harmonic of temporal frequency, with no overall "null-position." The optimum spatial frequency was usually higher than would be predicted by simple summation within the dimensions of the receptive field. Thus, neurons in PMLS and PLLS, like complex cells in areas 17 and 18, behave nonlinearly and their spatial selectivity is determined by "subunits" smaller than their receptive fields. The range of preferred temporal frequencies ranged from less than 2.5 Hz to more than 10 Hz. In their temporal selectivity neurons in PMLS resembled cells in area 17, with little attenuation at low temporal frequencies, whereas there was a tendency for cells in PLLS to prefer higher temporal frequencies, as is common in area 18.

Animals

Differences of visual field representation in the medial and lateral banks of the suprasylvian cortex (PMLS/PLLS) of the cat.

We have studied the orderliness of representation of visual space in the medial and lateral banks of the middle suprasylvian sulcus. Penetrations were made either parallel to the sulcus, in one bank or the other, or vertical, thus crossing the sulcus between the postero-medial (PMLS) and posterolateral (PLLS) divisions of this area. In some cases we found clear evidence for topographical order in the representation of the visual field with a tendency (greater in PMLS than in PLLS) for the receptive fields of cells recorded deeper in the walls of the sulcus to lie closer to the area centralis, but along many penetrations the receptive fields were so large and so scattered that no retinotopic arrangement could be discerned. In PMLS the receptive fields of the majority of units we studied were centered below and close to the horizontal meridian, whereas in PLLS they were distributed over both the upper and lower visual fields with an over-representation of the upper field. Receptive fields were significantly larger in PLLS (mean field area = 442.2 deg2) than in PMLS (mean area = 154.4 deg2); there was also less clear correlation between receptive field size and eccentricity in PLLS (correlation coefficient = +0.25) than in PMLS (corr. coeff. = +0.72). Analysis of the distance between the receptive field centres of consecutively recorded units demonstrated that the mean scatter in both PMLS and PLLS amounts to about half the average receptive field diameter. In summary the topographical representation of visual space is less orderly in PLLS, and may involve a wider area of the visual field. These findings may relate to the segregated visual cortical and extrageniculate thalamic connections that the medial and lateral banks of the LS receive.

Animals

Organization of the visual pathways in the newborn kitten.

We have used retrograde and anterograde transport to examine the major visual pathways in newborn kittens. Retinal projections from both eyes to the dorsal lateral geniculate nucleus (dLGN) and superior colliculus (SC) are present and topographically organized. The dLGN projects topographically to areas 17 and 18 and receives reciprocal projections from cells in layer VI of areas 17, 18, 19 and suprasylvian cortex on the same side of the brain. Area 19 also has a sparse thalamic input but probably not from the dLGN. The laminar distribution of [3H]proline transported from dLGN to area 17 was quantified: label was spread through all layers, with a minimum at the border of layers I/II. Layer I was always labelled less heavily than IV. These results are critically compared with those based on other tracing techniques. Cells of layer V in areas 17, 18, 19 and the suprasylvian cortex project topographically to the superficial layers of the ipsilateral SC. Area 19 and the lateral suprasylvian cortex also send a crossed projection to restricted parts of the opposite SC. Thus these visual projections are not only present and topographically ordered on the day of birth, but, unlike certain highly exuberant interhemispheric and cortico-cortical projections, they are qualitatively remarkably mature, some days before the onset of visual activity. The major subcortical projections to and from the visual cortex appear to be constructed without the benefit of visual experience and much of the activity-dependent plasticity of cortical cells may well involve only local modulation of synaptic input.

Animals