Management of irregular astigmatism.

Using a liberal definition of corneal irregularity, modern videokeratoscopy may define approximately 40% of normal corneas with a toric refractive error as possessing primary irregular astigmatism. The causes of secondary forms of irregular astigmatism include corneal surgery, trauma, dystrophies, and infections. Internal refractive surface and media irregularity or noncorneal astigmatism (ocular residual astigmatism) contribute to irregular astigmatism of the entire refractive path of which crystaline lenticular astigmatism is usually the principal contributing component. Treatment options have increased in recent years, particularly, though not exclusively, through the advent of tailored corneal excimer laser ablations. However, discussion continues concerning the systematic approach necessary to enable treatment to achieve an optimal optical surface for the eye. Discussion also continues as to what constitutes the optimal corneal shape. Some refractive procedures may increase higher order aberrations in the attempt to neutralize refractive astigmatism. The way to further refinement of the commonly performed refractive techniques will ultimately lie in the integrated inclusion of a trio of technologies: topographic analysis of the corneal surface, wavefront analysis of ocular refractive aberrations, and vector planning to enable the appropriate balance in emphasis between these two diagnostic modalities. For the uncommon, irregularly roughened corneas, the ablatable polymer techniques show some promise.

Monocular focal retinal lesions induce short-term topographic plasticity in adult cat visual cortex.

Electrophysiological recording in primary visual cortex (VI) was performed both prior to and in the hours immediately following the creation of a discrete retinal lesion in one eye with an argon laser. Lesion projection zones (LPZs; 21-64 mm2) were defined in the visual cortex by mapping the extent of the lesion onto the topographic representation in cortex. There was no effect on neuronal responses to the unlesioned eye or on its topographic representation. However, within hours of producing the retinal lesion, receptive fields obtained from stimulation of the lesioned eye were displaced onto areas surrounding the scotoma and were enlarged compared with the corresponding field obtained through the normal eye. The proportion of such responsive recording sites increased during the experiment such that 8-11 hours post-lesion, 56% of recording sites displayed neurons responsive to the lesioned eye. This is an equivalent proportion to that previously reported with long-term recovery (three weeks to three months). Responsive neurons were evident as far as 2.5 mm inside the border of the LPZ. The reorganization of the lesioned eye representation produced binocular disparities as great as 15 degrees, suggesting interactions between sites in VI up to 5.5 mm apart.

Visuotopic reorganization in the primary visual cortex of adult cats following monocular and binocular retinal lesions.

The effect of discrete monocular retinal lesions on the representation of the visual field in the primary visual area (V1) was investigated in adult cats. Lesions were created using argon lasers, 8 d to 4(1/2) months prior to electrophysiological recording. This produced lesion projection zones (LPZs) in V1, 1.6-9.5 mm wide, that were deprived of their normal input from one eye, but that received a normal input from the other eye. Nevertheless, at the majority of recording sites within these zones neuronal responses were elicited by stimulation of the lesioned eye, with receptive fields being displaced onto regions of retina surrounding the lesion, while receptive fields determined through stimulation of the normal eye followed the normal visuotopic organization of V1. However, neuronal responses to stimulation of the lesioned eye within the LPZs were characterized by rapid habituation and unusually low firing rates in comparison with responses to stimulation of the normal eye. Stimulation of the normal eye temporarily marked the responsiveness of neurons within the LPZ to stimulation of the lesioned eye. The proportion of neurons responsive to stimulation of the lesioned eye was higher just inside the borders of the LPZs than at the centers of these zones. However, neurons responsive to stimulation of the test eye were found up to 3.6 mm from the perimeter of the LPZs, and therefore the shifts in the visuotopic map caused by retinal lesions cannot be explained solely on the basis of the normal scatter of receptive fields and point-image size in V1. The proportion of cells responsive to stimulation of the lesioned eye was highest in the infragranular layers, and lowest in the supragranular layers. By combining a restricted lesion of one eye with laser photocoagulation of the optic disc of the other eye, the effects of the normal eye on the lesion-induced visuotopic reorganization were also investigated. Neither chronic nor acute deactivation produced any discernible further changes in visuotopy or in the characteristics of neuronal responses to stimulation of the eye with the discrete lesions. Our findings show that the representations of the two eyes in adult visual cortex are capable of independent reorganization. These findings parallel those of work in auditory cortex, suggesting that topographic reorganization in primary sensory areas of adult cortex may be governed by similar mechanisms.

Visual areas in the dorsal and medial extrastriate cortices of the marmoset.

To define the number and limits of the visual areas in the primate extrastriate cortex, the visuotopy of the dorsal convexity and medial wall was studied by electrophysiological recordings in five marmosets anaesthetised with sufentanil and nitrous oxide and paralysed with pancuronium bromide. We identified five visuotopic representations in and around the densely myelinated zone between visual area 2 (V2) and the posterior parietal cortex. Most of the densely myelinated zone is formed by the homologue of the owl monkey's dorsomedial area (DM); thus, we also termed this area DM in the marmoset. Within DM, the lower quadrant representation is continuous, with central vision represented laterally, peripheral vision medially, the horizontal meridian caudally, and the vertical meridian rostrally. In contrast, the upper quadrant representation is split, with the central portion represented at the lateral edge of DM on the dorsal surface, and the periphery along the midline. Two other visual field representations, corresponding to the dorsointermediate area (DI) and to a new subdivision termed the dorsoanterior area (DA), are also densely myelinated but can be distinguished from DM based on the separation of the bands of Baillarger and visual topography. In addition, a homologue of the medial visual area (M) was identified. Our results reveal a highly complex visuotopy in primate cortex, with local discontinuities in representation and borders between areas that are often not coincident with either the horizontal or the vertical meridian. The topography of the dorsal extrastriate cortex in the marmoset strongly suggests that both visual area 3 (V3) and the parietooccipital area (PO) of other primates are portions of a single visuotopic representation, DM, and calls into question the existence of visual areas with partial or quadrantic representations of the visual field.

Retinal detachment induces massive immediate reorganization in visual cortex.

Large inactive regions of the retina of adult cats were produced by the novel method of inducing monocular retinal detachment. Within a few hours, neurones throughout the detachment projection zone in primary visual cortex (55-136 mm2), including some > 4.5 mm from its boundary, were found to have large receptive fields displaced onto intact retina. The new receptive fields of some neurones represented shifts of up to 9 mm across the retinotopic representation. For these rapid changes to occur pre-existing viable circuits must provide a cortical locus with inputs from a wide extent of the retina. Receptive fields, and the retinotopic map, for stimulation of the other eye were unchanged.

Responsiveness of cat area 17 after monocular inactivation: limitation of topographic plasticity in adult cortex.

1. Recordings were made from neurones in the splenial sulcus of normal adult cats and adult cats which had one eye inactivated by enucleation or photocoagulation of the optic disc. Two visually responsive regions were observed, corresponding to the peripheral representation of visual area 1 (V1) and the splenial visual area. In normal animals, responses to the ipsilateral eye in V1 were restricted to the medial half of the splenial sulcus, up to 45-50 deg eccentricity. Thus, by inactivating the eye contralateral to the experimental hemisphere, we created a region in V1, 1-2 mm wide, that lacked normal inputs. 2. In contrast to results from previous experiments where lesions were placed in the central retina, neurones in the deprived peripheral representation remained unresponsive to light stimuli for up to 12 h after deactivation of the contralateral eye. 3. In animals that were allowed to recover from the monocular deactivation for periods of 2 days to 16 months, there was rearrangement of the retinotopic maps. Receptive fields in regions of cortex that normally represented the monocular crescent were displaced to the temporal border of the binocular field of vision. However, most neurones in the deprived peripheral representation remained unresponsive to visual stimuli even more than 1 year after treatment. This is also in marked contrast with the extensive reorganization that is observed in the central representation of V1 after restricted retinal lesions. Analysis of the cortical magnification factor demonstrates that the change in visual topography is local, and does not involve an overall centro-peripheral shift of the retinotopic map. 4. Among the neurones that did show displaced receptive fields, the response properties were clearly abnormal. They showed a notable lack of spontaneous activity, low firing rates and rapid habituation to repeated stimulation. 5. The low potential for reorganization of the monocular sector of V1 demonstrates that the capacity for plasticity of mature sensory representations varies with location in cortex. Even relatively small pieces of cortex, such as the monocular crescent representations, may not reorganize completely if certain conditions are not met. These results suggest the existence of natural boundaries that may limit the process of reorganization of sensory representations.

Magnification factors, receptive field images and point-image size in the superior colliculus of flying foxes: comparison with the primary visual cortex.

The magnification factor (MF) of the stratum griseum superficiale (SGS) of the superior colliculus (SC) was calculated based on visual receptive fields recorded from anaesthetised and paralysed flying foxes (Pteropus spp.). In areal terms, the MF at the representation of central vision was 4-6 times larger than that in the peripheral representation. This variation is less marked than that observed in the primary visual area (V1), but is roughly that expected if the retinotopic map in the SC was defined by the distribution of ganglion cells in the retina. Two measures of the functional spread of activity in the SC, the receptive field images and the point-image size, were calculated. Receptive field images are remarkably similar throughout the SC. As in V1, the point-image size in the SGS of flying foxes is 0.5-0.6 mm and varies little with eccentricity. Bilateral ablation of the visual cortex results in a reduction of the mean receptive field size of neurones in the SGS, and the point-image size is reduced by half. However, the shape of the point-image function is not affected. These results demonstrate that the spread of activity in the SC is nearly constant throughout the retinotopic map and that this is primarily a result of the direct retinal projection. Although the visual cortex has an expanded central representation in comparison with the SC, the corticotectal pathway does not exert a preferential influence on the central representation of the SC.

Topography and extent of visual-field representation in the superior colliculus of the megachiropteran Pteropus.

It has been proposed that flying foxes (genus Pteropus) have a primate-like pattern of representation in the superficial layers of the superior colliculus (SC), whereby the visual representation in this structure is limited by the same decussation line that limits the retino-geniculo-cortical projection (Pettigrew, 1986). To test this hypothesis, visual receptive fields were plotted based on single- and multi-unit recordings in the SC of ten flying foxes. A complete representation of the contralateral hemifield was observed in the SC. Although the binocular hemifield of vision in Pteropus is 54 deg wide, receptive-field centers invaded the ipsilateral hemifield by only 8 deg, and the receptive-field borders by 13 deg. This invasion is similar to that observed at the border between visual areas V1 and V2 in the occipital cortex. The extent of the ipsilateral invasion was not affected by a lesion that completely ablated the occipital visual areas, thus suggesting that this invasion may be consequence of a zone of nasotemporal overlap in the retinal projections to the two colliculi. Neurones located in the superficial layers typically responded briskly to stimulation of both eyes, with a bias towards the contralateral eye. After cortical lesions the neuronal responses to the ipsilateral eye were depressed, and the ocular-dominance histograms shifted towards an even stronger dominance by the contralateral eye. However, cells located in the rostral pole of the SC remained responsive to the ipsilateral eye after cortical lesions. Responses in the stratum opticum and stratum griseum intermediale were more severely affected by cortical lesions than those in the stratum griseum superficiale. Our results demonstrate that the SC in flying foxes retain some generalized mammalian characteristics, such as the stronger direct projections of the contralateral eye and the location of the upper, lower, central, and peripheral representations in the SC. Nonetheless, the extent of visual representation in the SC demonstrates a specialized, primate-like pattern. These observations are consistent with the hypothesis that megachiropterans are members of a group that branched off early during the differentiation of primates from basal mammals.

Organization of the second visual area in the megachiropteran bat Pteropus.

The organization of peristriate cortex was studied in nine flying foxes (genus Pteropus). Based on receptive field mapping and architectonic data, we report on the organization of the second visual area (V2). V2 forms a continuous belt 2-4 mm wide bordering V1 anteriorly. In each hemisphere, V2 contains a precisely organized representation of the entire contralateral visual field. The vertical meridian of the visual field (VM), and a short strip of the ipsilateral hemifield are represented at the posterior border of V2, with V1. The area centralis is represented approximately at the center of the posterior border of V2. At each mediolateral level, progressively more peripheral portions of the visual field are represented as V2 is crossed from posterior to anterior. The representation of the upper quadrant is continuous, and confined to the lateral half of V2. In contrast, the representation of the lower quadrant is split along a line running from the temporal edge of the field of vision to the optic disk. As a result of this arrangement, the portions of the lower quadrant close to the VM are represented medially, and those away from the VM laterally in V2. The entire representation of the horizontal meridian is located in lateral V2, and is not split between medial and lateral V2 as in primates. The linear cortical magnification factor (CMF) decays by a factor of 3-5 from the central to the peripheral representation. The CMF is anisotropic, and equal distances in the visual field are magnified twice as much parallel to the V1/V2 border than perpendicular to this border. Moreover, points in the lower quadrant are magnified relative to symmetrical points in the upper quadrant. V2 is histologically distinct from all surrounding areas in both cytochrome oxidase- and Nissl-stained sections. These results suggest that V2 is an homologous area common to all archontans, and imply that much of the variability reported among mammals may be due to technical factors, rather than true species differences.

Retinotopic organization of the primary visual cortex of flying foxes (Pteropus poliocephalus and Pteropus scapulatus).

The representation of the visual field in the occipital cortex was studied by multiunit recordings in seven flying foxes (Pteropus spp.), anesthetized with thiopentone/N2O and immobilized with pancuronium bromide. On the basis of its visuotopic organization and architecture, the primary visual area (V1) was distinguished from neighboring areas. Area V1 occupies the dorsal surface of the occipital pole, as well as most of the tentorial surface of the cortex, the posterior third of the mesial surface of the brain, and the upper bank of the posterior portion of the splenial sulcus. In each hemisphere, it contains a precise, visuotopically organized representation of the entire extent of the contralateral visual hemifield. The representation of the vertical meridian, together with 8-15 degrees of ipsilateral hemifield, forms the anterior border of V1 with other visually responsive areas. The representation of the horizontal meridian runs anterolateral to posteromedial, dividing V1 so that the lower visual quadrant is represented medially, and the upper quadrant laterally. The total surface area of V1 is about 140 mm2 for P. poliocephalus, and 110 mm2 for P. scapulatus. The representation of the central visual field is greatly magnified relative to that of the periphery. The cortical magnification factor decreases with increasing eccentricity, following a negative power function. Conversely, receptive field sizes increase markedly with increasing eccentricity, and therefore the point-image size is approximately constant throughout V1. The emphasis in the representation of the area centralis in V1 is much larger than that expected on the basis of ganglion cell counts in flat-mounted retinas. Thus, a larger degree of convergence occurs at the peripheral representations in the retino-geniculo-cortical pathway, in comparison with the central representations. The marked emphasis in the representation of central vision, the wide extent of the binocular field of vision, and the relatively large surface area of V1 reflect the importance of vision in megachiropterans.

Connections of somatosensory cortex in megachiropteran bats: the evolution of cortical fields in mammals.

The cortical connections of the primary somatosensory area (SI or 3b), a caudal somatosensory field (area 1/2), the second somatosensory area (SII), the parietal ventral area (PV), the ventral somatosensory area (VS), and the lateral parietal area (LP) were investigated in grey headed flying foxes by injecting anatomical tracers into electrophysiologically identified locations in these fields. The receptive fields for clusters of neurons were mapped with sufficient density for injection sites to be related to the boundaries of fields, and to representations of specific body parts within the fields. In all cases, cortex was flattened and sectioned parallel to the cortical surface. Sections were stained for myelin and architectonic features of cortex were related to physiological mapping and connection patterns. We found patterns of topographic and nontopographic connections between 3b and adjacent anterior parietal fields 3a and 1/2, and fields caudolateral to 3b (SII and PV). Area 1/2 had both topographic and nontopographic connections with 3b, PP, and SII. Connections of SII and PV with areas 3b, 3a, and 1/2 were roughly topographic, although there was clear evidence for nontopographic connections between these fields. SII was most densely connected with area 1/2, while PV was most densely connected with 3b. SII had additional connections with fields in lateral parietal cortex and with subdivisions of motor cortex. Other connections of PV were with subdivisions of motor cortex and pyriform cortex. Laminar differences in connection patterns of SII and PV with surrounding cortex were also observed. Injections in the ventral somatosensory area revealed connections with SII, PV, area 1/2, auditory cortex, entorhinal cortex, and pyriform cortex. Finally, the lateral parietal field had very dense connections with posterior parietal cortex, caudal temporal cortex, and with subdivisions of motor cortex. Our results indicate that the 3b region is not homogeneous, but is composed of myelin dense and light regions, associated with 3b proper and invaginations of area 1/2, respectively. Connections of myelin dense 3b were different from invaginating portions of myelin light area 1/2. Our findings that 3b is densely interconnected with PV and moderately to lightly interconnected with SII supports the notion that SII and PV have been confused across mammals and across studies. Our connectional evidence provides further support for our hypothesis that area 1/2 is partially incorporated in 3b and has led to theories of the evolution of cortical fields in mammals.

Retinal topography in the koala (Phascolarctos cinereus).

Nissl-stained retinal wholemounts were used to investigate the topographical organization of the ganglion cell layer of the koala (Phascolarctos cinereus); the visual resolution limit of this animal was subsequently estimated from retinal ganglion cell density data. Two types of cells could be differentiated on the basis of their size and staining characteristics: a subpopulation of presumed ganglion cells, consisting of medium to large cells with Nissl substance in the cytoplasm and pale uniformly staining nuclei, and a further subpopulation of small, densely staining cells. The latter group were presumed to be neuroglia and displaced amacrine cells. Iso-density contour maps were prepared from total cell counts and also counts of presumed ganglion cells; in all cases, the density of cells was greatest in the inferior retina where there was an area of peak density occurring as a poorly developed, horizontal streak that extended across the inferior retina. The inferior position of the streak in the koala contrasts with reports of the superior position of streaks in other marsupials. Peak cell densities of 2370 cells/mm2 and 1480 cells/mm2 were recorded for the total cell population and the presumed ganglion cell subpopulation, respectively. The latter value is equivalent to a visual resolution of 2.4 cycles/degree, based on sampling theory and a square packing paradigm, placing the koala close in visual performance to two other marsupials, the Australian Northern native cat and the American Virginia opossum.