Similar Orientation Preferences Research Articles

Development of local horizontal interactions in cat visual cortex studied by cross-correlation analysis

1. To explore the functional development of local horizontal interactions in the primary visual cortex, we carried out cross-correlation analysis of spike trains recorded simultaneously from a pair of neurons separated horizontally by < 1 mm, in kittens ranging in age postnatally from the second to ninth week. 2. Significantly correlated firings were found in 87 pairs of cells among 423 pairs analyzed, and 77 pairs of them were classified into three types on the basis of their functional implications: 1) excitatory interactions, 2) inhibitory interactions, and 3) common inputs to both neurons of the pair from other sources. 3. Common inputs and excitatory linkage were observed even in animals at the second postnatal week, whereas inhibitory linkage was not seen before the fourth week of age. The probability of observing common inputs and inhibitory linkage increased during development, whereas that of excitatory linkage tended to decrease after the sixth week of age. 4. Significant correlation was rarely seen in pairs with horizontal separation > 600 microns in the seventh to ninth week. In the fourth to sixth week of age, however, approximately 30% of the pairs with horizontal separation between 600 and 800 microns were significantly correlated. 5. Cells that were not sensitive for orientation or that lacked a visual response were observed mainly before the sixth week of age. These cells tended to receive excitatory effects from and share common inputs with other orientation-sensitive cells that were located within the horizontal distance of 400 microns. 6. All three types of correlations were observed mostly in cell pairs with preferred orientations that differed < 45 degrees at all ages studied. In the fourth to sixth week, however, the similarity of orientation preference was not strict, and correlated firings were observed even in a pair with orthogonal orientation preferences; whereas in the seventh to ninth week the tuning became sharper. 7. These results suggest that functional interactions between cortical neurons exist but are much less specific with respect to horizontal separation and orientation preference before the sixth week of age, and these interactions become more specific so as to operate between neurons with similar orientation preferences in more restricted region after the seventh to ninth week of age and thus in adulthood.

Functional specificity of a long-range horizontal connection in cat visual cortex: a cross-correlation study

Anatomical investigations of the visual cortex revealed a regular set of interlaminar connections and long-range horizontal connections. An important component of the intrinsic cortical circuit is the projection from layer 5 to layer 6 over long horizontal distances. Previous work has shown that when layer 5 is locally inactivated, layer 6 cells lose their response in a segment of their receptive fields corresponding to the blocked area in layer 5. It has therefore been proposed that the long receptive fields characteristic of layer 6 cells might be generated by concatenating layer 5 cells with similar orientation preferences. In the present study, we used cross-correlation analysis to examine both source and target cells of the interlaminar connection from layer 5 to layer 6. We found correlated firing between cells separated by up to 4.2 mm, the longest horizontal distance studied. The occurrence of correlated firing depended on the functional properties and on the topographic position of the cells in layer 5 and layer 6. Interactions were only observed if the cells had matching orientation tuning and similar eye preference and if the layer 5 cells lay within the summation area of the layer 6 cells. Both simple and complex cells in layer 6 fired in synchrony with layer 5 cells. In layer 5, mainly standard complex cells, but very few special complex cells, participated in correlated firing with layer 6 cells. These results suggest that cells in layer 6 receive their input from cooriented, coaxially aligned standard complex cells in layer 5. This topographic arrangement and the length summation properties of standard complex cells in layer 5 are well suited to generate long receptive fields in layer 6. Thus, our experiments provide evidence for the functional specialization of a particular component of cortical circuitry.

Stimulus-Dependent Neuronal Oscillations in Cat Visual Cortex: Inter-Columnar Interaction as Determined by Cross-Correlation Analysis.

We have demonstrated previously that neurons in cat striate cortex, in response to their preferred stimuli, exhibit oscillatory responses in a frequency range of 40 - 60 Hz. Recently, we obtained evidence that such oscillatory responses can synchronize across columns. We have now performed an extensive analysis of this phenomenon for both unit and field potential responses. In addition, we studied the stimulus conditions leading to intercolumnar synchronization. We recorded both multi-unit activity and local field potentials from area 17 of adult cats with arrays of several electrodes. Interelectrode distances ranged from 0.4 to 12 mm. For all pairs of unit (n=200) and field potential (n=174) recordings, we computed auto- and cross-correlation functions. The modulation of the correlograms was quantified by fitting a damped sine wave (Gabor) function to the data. Cross-correlation analysis of the unit data revealed that in 90 out of 200 cases the recorded cells established a constant phase-relationship of their oscillatory responses. This occurred, on average, with no phase difference. If the receptive fields were nonoverlapping, we observed a synchronization primarily between cells with similar orientation preferences. Cells with overlapping receptive fields also showed a high incidence of synchronization if their orientation preferences were different. In this latter group, synchronization occurred even in cases where the stimulus was optimal for only one of the recording sites. Under conditions of monocular instead of binocular stimulation the oscillatory modulation of the responses was attenuated, but the cross-correlogram still indicated a significant interaction. Similar effects were seen with the application of stationary instead of moving stimuli. A synchronization of oscillatory field potential responses was observed in 136 out of 174 paired recordings. At all distances investigated, the probability of synchronization of field potential responses was independent of the orientation preferences of the cells. However, the strength of interaction decreased with increasing spatial separation. Control experiments showed that the synchronization of field potential responses was not due to volume conduction. The results demonstrate that oscillatory responses at separate cortical sites can transiently synchronize. The probability and strength of synchronization are dependent on the spatial separation of the recorded cells and their orientation preferences. In addition, the cross-columnar synchronization is influenced by features of the visual stimulus. It is suggested that this synchronization provides a mechanism for the formation of neuronal assemblies in the visual cortex.

Horizontal Interactions in Cat Striate Cortex: III. Ectopic Receptive Fields and Transient Exuberance of Tangential Interactions.

In this study the developmental changes of intracortical connectivity are related to changes of cortical receptor fields (RFs). The RFs of striate cortex neurons of 4- to 8.5-week-old kittens, reared under normal conditions (NR) or in a selective visual environment (SE), were analysed quantitatively and compared with adult cats. To unmask weak inputs from outside the conventional RF (CRF), cell excitability was raised by iontophoretic application of glutamate (GLU) and/or bicuculline methiodide (BIC) or by light stimulation of the CRF. Both the dominant discharge region (DDR) and the total RF (TRF) area were significantly larger in NR and SE kittens than in adult cats. Moreover, in kittens 18% of the cells had additional ectopic fields that were excitatory, had similar orientation preferences as the CRF, and ranged 4 degrees to 23 degrees from the centre of the CRF. In 74% of the cases the ectopic fields were direction-selective and 70% of them preferred stimuli moving toward the CRF. Ectopic fields occurred mainly in supragranular cells, were similarly frequent in simple and complex cells and slightly more frequent in SE (20.7%) than in NR (13.3%) kittens. In adult cats only one of 83 cells tested had an ectopic field. It is concluded that the age-dependent decrease in the RF size, the laminar distribution of cells having an ectopic RF, and the numerical reduction of these cells with age correlate well with the organization and postnatal pruning of tangential projections, suggesting that these contribute to the elaboration of specific response properties. Moreover, the authors infer from the early presence and from the selectivity of ectopic fields that the system of horizontal intrinsic connections mediates far-reaching, excitatory interactions between cortical neurons with similar functional properties and serves as a substrate for the processing of global aspects of visual patterns.

Organized arrangement of orientation-sensitive relay cells in the cat's dorsal lateral geniculate nucleus

We studied the physiological orientation biases of over 700 relay cells in the cat's dorsal lateral geniculate nucleus (LGNd). Relay cells were sampled at regular intervals along horizontally as well as vertically oriented electrode penetrations in a fashion analogous to that used previously in studies of visual cortex (Hubel and Wiesel, 1962). The strengths of the orientation biases and the distributions of the preferred orientations were determined for different classes of relay cells, relay cells in different layers of the LGNd, and relay cells subserving different parts of the visual field. We find that, at the population level, LGNd cells exhibit about the same degree of orientation bias as do the retinal ganglion cells providing their inputs (see also Soodak et al., 1987). Also, as in the retina (Levick and Thibos, 1982; Leventhal and Schall, 1983), most LGNd cells tend to prefer stimuli oriented radially, i.e., parallel to the line connecting their receptive fields to the area centralis projection. However, the radial bias in the LGNd is weaker than in the retina. Moreover, there is a relative overrepresentation of cells preferring tangentially oriented stimuli in the LGNd but not in the retina. As a result of the overrepresentation of cells preferring radial and tangential stimuli, the overall distribution of preferred orientations varies in regions of the LGNd subserving different parts of the visual field. Reconstructions of our electrode penetrations provide evidence that, unlike in the retina, cells having similar preferred orientations are clustered in the LGNd. This clustering is apparent for all cell types and in all parts of laminae A and A1. The tendency to cluster according to preferred orientation is evident for cells preferring radially, intermediately, and tangentially oriented stimuli and thus is not simply a reflection of the radial bias evident among retinal ganglion cells at the population level. It is already known that cells having inputs from different eyes, on-center, off-center, X-, Y-, W-type, and color-sensitive ganglion cells are distributed nonrandomly in the LGNd of cats and monkeys (for review, see Rodieck, 1979; Stone et al., 1979; Lennie, 1981; Stone, 1983). The finding that relay cells having similar preferred orientations are also distributed nonrandomly suggests that the initial sorting of virtually all properties segregated in visual cortex may begin in the LGNd.

Horizontal organization of orientation-sensitive cells in primate visual cortex

In the visual cortex of the monkey the horizontal organization of the preferred orientations of orientation-selective cells follows two opposing rules: (1) neighbors tend to have similar orientation preferences, and (2) many different orientations are observed in a local region. We have described a classification for orientation maps based on the types of topological singularities and the spacing of these singularities relative to the cytochrome oxidase blobs. Using the orientation drift rate as a measure we have compared simulated orientation maps to published records of horizontal electrode recordings.

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)

Structural deformation of the Leoville chondrite

Foliations defined by alignment of elongated chondrules have been noted previously in chondrites, but none displays this effect so well as Leoville (CV3). The shapes of Leoville chondrules were produced by deformation in situ, as indicated by inclusions and clasts with similar shapes and preferred orientations to those of chondrules. Similarities in the aspect ratios of apparent strain ellipses measured for chondrules alone (1.9 and 2.0 by several methods) and for the whole meteorite (2.0) indicate either that Leoville deformed homogeneously or that it deformed as a framework of touching chondrules. This amount of strain corresponds to approximately 33% uniaxial shortening, assuming constant volume. Because the strain ellipse was measured in only one orientation, this strain value is a minimum estimate for the meteorite. Lack of correlation between foliation and either shock or thermal effects argues that impact or metamorphism are unlikely to have produced this deformation. Compaction due to overburden from progressive accretion on the chondrite parent body is suggested to have been its cause. Estimates of maximum deviatoric stresses in the interiors of asteroid-sized bodies and constraints on maximum temperatures for CV3 chondrites are consistent with diffusional flow as the deformation mechanism for olivine in these chondrules. Diffusional flow is also suggested by the scarcity of observed lattice dislocations. Deformation of Leoville olivines by this mechanism at geologically reasonable strain rates appears to require higher temperatures than those believed to have been experienced by this meteorite (< 600°C). However, differences in olivine grain size, the presence of water, or a more complex deformation history might explain this discrepancy.

Long axons within the striate cortex: their distribution, orientation, and patterns of connection.

Rockland and Lung [Rockland, K. S. & Lung, J. S. (1982) Science 215, 1532-1534] have recently observed that an injection of horseradish peroxidase into the striate cortex of the tree shrew produces a patchy distribution of label adjacent to the injection site. They proposed that this pattern might be due to populations of neurons with long-range cortico-cortical connections that are interspersed with populations having no such connections. We suggest here an alternative explanation. We can account for the pattern by supposing that the label is carrier by a system of oriented axons. We suppose that these axons link cells with similar orientation preferences and make their connections within a narrow strip of cortex whose direction is related to the orientation of the cells in question. We suggest that such connections could be involved in generating complex receptive fields from simple ones. Other possibilities are that they are used to generate very elongated receptive fields, inhibitory flanks, or end-stopping. We suggest a number of experimental tests of these ideas.

Recovery from monocular deprivation in the monkey. I. Reversal of physiological effects in the visual cortex.

This is a study of the effects of monocular deprivation, reverse suturing (opening the deprived eye with closure of the other) and reopening of the deprived eye alone (without closing the other) on the physiological organization of the primary visual cortex in monkeys (Erythrocebus patas). All animals were initially monocularly deprived by suture of the lids of the right eye from soon after birth until about 4 weeks of age (24-29 days). In a monocularly deprived animal, recordings were taken from area 17 at 24 days. Already most neurons recorded outside layer IVc, were strongly or completely dominated by functional input from the left eye. The Non-oriented cells of layer IVc, where the bulk of the afferent input terminates, were also mainly dominated by the left eye. Although segregation of input from the two eyes was not complete, large areas of layer IVc were already monocularly dominated by the left eye. Four animals were reverse-sutured at about 4 weeks and recorded 3, 6, 15 and 126 days later. In each animal the pattern of ocular dominance was fairly similar within and outside layer IVc. Even with only 3 days of forced usage of the initially deprived right eye, about half of all cells recorded had become dominated by it, and the process of "recapture' of cortical cells by the initially deprived eye was apparently complete within 15 days. In layer IVc, the recovery took the form of an expansion of zones dominated by the deprived eye, as if the originally shrunken stripes of afferent termination had become enlarged. Binocularly driven neurons were rare at all stages, in all layers, but when present and orientation-selective, they had similar preferred orientations in the two eyes. Likewise the "columnar' sequences of preferred orientation continued without obvious disruption on shifting from regions dominated by one eye to those dominated by the other. Simply reopening the deprived eye at about 4 weeks, for 15 to 96 days caused no detectable change in the overall ocular dominance of cortical cells and, on average, no expansion of right-eye dominance columns in layer IVc. Therefore the recovery seen after reverse suturing depends not just on the restoration of normal activity to axons carrying information from the right eye, but on the establishment of a competitive advantage, through the right eye being made more active than the left.

Textural effects in splat-quenched aluminum, tin and AlSn alloys

A preferred orientation of grains has been reported recently in splat-quenched foils of high purity cadmium, zinc and bismuth. A similar preferred orientation was observed in splat-cooled alumium and tin and is reported in the present paper. However, when AlSn alloys were splat quenched under identical conditions, the aluminium and tin phases in the splat foil showed a random orientation.

The physiological effects of monocular deprivation and their reversal in the monkey's visual cortex.

1. 1127 single units were recorded during oblique penetrations in area 17 of one normal, three monocularly deprived and four reverse sutured monkeys. 2. In all animals most cells outside layer IV c were orientation-selective, and preferred orientation usually shifted from cell to cell in a regular progressive sequence. 3. The presence in layer IV c of non-oriented, monocularly driven units, organized in alternating right-eye and left-eye 'stripes' (LeVay, Hubel & Wiesel, 1975) was confirmed. 4. Early monocular deprivation (2--5 1/2 weeks) caused a strong shift of ocular dominance towards the non-deprived eye. However, even outside layer IV c, neural background and some isolated cells could still be driven from the deprived eye in regularly spaced, narrow columnar regions. In layer IV c the non-deprived eye's stripes were almost three times wider, on average, than the deprived. 5. Later monocular deprivation (11--16 months) had no detectable influence on layer IV c but seemed to cause a small shift in ocular dominance outside IV c. Deprivation for 6 1/4 months in an adult had no such effect. 6. After early reverse suturing (at 5 1/2 weeks) the originally deprived eye gained dominance over cells outside layer IV c just as complete as that originally exercised by the eye that was first non-deprived. 7. The later reverse suturing was delayed, the less effective was recapture by the originally deprived eye. Reversal at 8 weeks led to roughly equal numbers of cells being dominated by each eye; fewer cells became dominated by the newly open eye after reverse suturing at 9 weeks and most of them were non-oriented; reversal at 38 1/2 weeks had no effect. 8. Binocular cells, though rare in reverse sutured animals, always had very similar preferred orientations in the two eyes. The columnar sequences of preferred orientation were not interrupted at the borders of ocular dominance columns. 9. Even within layer IV c there was evidence for re-expansion of physiologically determined ocular dominance stripes. After early reverse suture, stripes for the two eyes became roughly equal in width. Possible mechanisms for these changes are discussed.

Quantitative study of cortical orientation selectivity in visually inexperienced kitten.

1. Extracellular recordings were made from single units in the visual cortices of six kittens deprived of experience with pattern vision by binocular lid suture. 2. Selectivity for stimulus orientation was quantitatively assessed in 98 units; 90 responded selectively to the orientation of a moving bar stimulus, the remainder responding nonselectively or too poorly to classify. Cells in these visually inexperienced kittens were similar in their degree of selectivity for orientation to cells tested in adult cats. However, responses tended to be weaker and somewhat more erratic. 3. About half the cells in this simple responded to both directions of stimulus motion at the optimal orientation. Most of those responding to only one direction of motion were considered orientation rather than direction selective because they responded more strongly or more selectively to a moving bar than to a moving spot. 4. Cells appeared to be organized within the cortex in a pattern similar to that found in adult cats, with cells in one column selective for the same orientation, and adjacent column having similar preferred orientations. 5. It is concluded that selectivity for stimulus orientation in the cat's visual cortex is innately determined.

Pyrolytic carbons deposited in fluidized beds at 1200 to 1400°c from various hydrocarbons

Pyrolytic carbons deposited in fluidized beds at 1200 to 1400°C from various hydrocarbon-helium mixtures using otherwise identical process parameters were compared. The trends of microstructure or density with increasing deposition rate were very similar at a given temperature. For each hydrocarbon investigated at 1400°C there was a minimum in the density vs. deposition rate curve. Pyrolytic carbons deposited at 1200°C at similar initial rates near 10 μ/hr from seven hydrocarbon-helium mixtures (methane, ethane, ethylene, acetylene, propane, cyclohexane and benzene) had very similar microstructures, densities, preferred orientations and apparent crystallite sizes. The results support the view that pyrolysis of hydrocarbons in fluidized beds involves dissociation into small fragments which then take place in polymerization-dehydrogenation reactions and indicate that the general mechanisms of pyrolytic carbon deposition in fluidized beds are independent of the hydrocarbon. The various microstructures were explained in terms of the extent of incorporation of gas-borne droplets or soot particles into the deposits or the variations of coating environment within the fluidized beds. The systematic variations of the properties of the pyrolytic carbons deposited from methanehelium mixtures at 1400, 1300 and 1200°C were investigated in detail and found to be consistent with a published model of pyrolytic carbon deposition in fluidized beds.

Sedimentary structures in Huronian rocks of Ontario

A study of cross-bedding in the southern part of the Huronian outcrop belt indicates that the sediments were transported in a southerly direction (vector mean 160°). Symmetrical ripple marks show a similar preferred orientation (178°), and, together with dessication cracks, indicate shallow water conditions. Dimensional fabric of embedded clasts in massive polymictic conglomerates of the Gowganda Formation probably reflects glacial transport in a southerly direction. Clastic dykes in the Espanola and Serpent Formations are interpreted as injection structures from the underlying Bruce conglomerate. The dykes and ball and pillow structures may indicate tectonic instability during sedimentation. In the Cobalt-Gowganda area both directional sedimentary structures and preferred orientation of clasts in the Gowganda conglomerates indicate a southerly direction of transportation. A study of cross-bedding in the region around the Sudbury Basin suggests the presence of a positive area there in early Huronian times. In the Bruce Mines-Blind River area dimensional fabric of the Gowganda conglomerates suggests that the glaciers moved in a northeast-southwest direction in response to a regional palaeoslope. However, directional structures in associated waterlaid sediments indicate transport to the southeast, possibly reflecting a local southeasterly slope in the Huronian sedimentary basin. The presence of flute casts, groove casts, and graded bedding in the waterlaid sediments of the Gowganda Formation suggests the possibility of the activity of turbidity currents, but many other lines of evidence support a glacial origin for the conglomerates and laminated argillites.

Effect of orientation on the modulation sensitivity for interference fringes on the retina.

It is now well established that, for many test targets, vertical and horizontal orientations yield higher visual acuities than oblique orientations. In order to assess the role of the optics of the eye in this effect, focusing errors of the eye were bypassed by using as the measure of resolving capacity the modulation sensitivity for sinusoidal interference fringes formed on the retina. The modulation sensitivity for vertical and horizontal orientations of the fringes was greater than for oblique orientations for a wide range of spatial frequencies. A similar orientation preference was found for the cut-off spatial frequencies. Measurements of the modulation sensitivity at 15° orientation intervals indicated that maxima in sensitivity were spaced at 90° intervals. Since the effects of the optics of the eye have been eliminated, the origin of meridional variations in acuity must lie in the retina and/or higher visual pathways.