Modeling the effect of facial topographies upon the visual field of humans and primates.

PURPOSE: To establish a virtual device that can predict the effect of facial features on the visual field of humans and primates. METHODS: Virtual masks were obtained from human subjects, and macaque, chimpanzee and baboon taxidermic specimens, and aligned with upright head orientation at the center of a virtual perimeter-like dome (radius = 50 m) developed with Cinema 4D. Virtual searchlights positioned at the masks' pupils were then allowed to 'paint' facial elements obstructing their path, and demarcate the unobstructed rays at the perimetric surface and on a virtual ground floor related to eye level. Searchlight positions along the human mask's pupillary axis were identified by maximum congruence to Goldmann visual field limits. RESULTS: The human contours largely concur with large-stimulus isopters displaying the limiting role of the nasal ridge, and the relatively extended ventral and temporal limits. In contrast, the facial design of chimpanzees and baboons obstructs significant portions of the ventral foreground (>2 m cf < 0.5 m in human), while there appear to be larger binocular overlaps (125 degrees in chimp cf 90 degrees in human). CONCLUSIONS: The model provides information on anatomical constraints for monocular and binocular visual field extensions including projection of the ventral field on a virtual floor.

Photoreceptor cell types in the retina of the tuatara (Sphenodon punctatus) have cone characteristics.

The tuatara Sphenodon punctatus, restricted to a few New Zealand offshore islands and now strictly protected, belongs to the Rhynchocephalia, the smallest order of extant reptiles. Earlier light microscopical studies on the retina of this species described photoreceptors with both rod- and cone-like features and the presence of a fovea. A limited amount of retinal material from S. punctatus has now allowed us to prepare the first-ever electron microscopic observations on the eye of this reptile. We were able to distinguish three types of photoreceptor, all with fine structural features characteristic of cone cells. Large single cones as well as double cones had open discs in their outer segments and straight axons with pedicle-type terminals. An additional cone type, characterized by somewhat more slender inner and outer segments, vitreally-displaced cell bodies and oblique axons, resembled short-wavelength cones known from other sauropsids. No cells with rod characteristics could be confirmed in the samples, although they might occur in retinal regions not available for this study. We conclude that the tuatara has cone-like photoreceptors, which-as in other crepuscular or nocturnal reptiles-have acquired rod-like features. The phenotypic adaptations notwithstanding, the set of photoreceptor types is quite typical of the reptilian eye and in some respects reminiscent of those seen in lizards and turtles.

Horizontal cells in the retina of a diurnal rodent, the agouti ( Dasyprocta aguti ).

The morphology and distribution of normally placed and displaced A horizontal cells were studied in the retina of a diurnal hystricomorph rodent, the agouti Dasyprocta aguti. Cells were labeled with anti-calbindin immunocytochemistry. Dendritic-field size reaches a minimum in the visual streak, of about 9,000 microm(2), and increases toward the retinal periphery both in the dorsal and ventral regions. There is a dorsoventral asymmetry, with dorsal cells being larger than ventral cells at equal distances from the streak. The peak value for cell density of 281 +/- 28 cells/mm(2) occurs in the center of the visual streak, decreasing toward the dorsal and ventral retinal periphery, paralleling the increase in dendritic-field size. Along the visual streak, the decline in cell density is less pronounced, remaining between 100-200 cells/mm(2) in the temporal and nasal periphery. Displaced horizontal cells are rare and occur in the retinal periphery. They tend to be smaller than normally placed horizontal cells in the ventral region, whilst no systematic difference was observed between the two cell groups in the dorsal region. Mosaic regularity was studied using nearest-neighbor analysis and the Ripley function. When mosaic regularity was determined removing the displaced horizontal cells, there was a slight increase in the conformity ratio, but the bivariate Ripley function indicated some repulsive dependence between the two mosaics. Both results were near the level of significance. A similar analysis performed in the capybara retina, a closely related hystricomorph rodent bearing a higher density of displaced horizontal cells than found in the agouti, suggested spatial independence between the two mosaics, normally placed versus displaced horizontal cells.

Advances in broad bandwidth light sources for ultrahigh resolution optical coherence tomography.

Novel ultra-broad bandwidth light sources enabling unprecedented sub-2 microm axial resolution over the 400 nm-1700 nm wavelength range have been developed and evaluated with respect to their feasibility for clinical ultrahigh resolution optical coherence tomography (UHR OCT) applications. The state-of-the-art light sources described here include a compact Kerr lens mode locked Ti:sapphire laser (lambdaC = 785 nm, delta lambda = 260 nm, P(out) = 50 mW) and different nonlinear fibre-based light sources with spectral bandwidths (at full width at half maximum) up to 350 nm at lambdaC = 1130 nm and 470 nm at lambdaC = 1375 nm. In vitro UHR OCT imaging is demonstrated at multiple wavelengths in human cancer cells, animal ganglion cells as well as in neuropathologic and ophthalmic biopsies in order to compare and optimize UHR OCT image contrast, resolution and penetration depth.

Compact, low-cost Ti:Al2O3 laser for in vivo ultrahigh-resolution optical coherence tomography.

A compact, low-cost, prismless Ti:Al2O3 laser with 176-nm bandwidth (FWHM) and 20-mW output power was developed. Ultrahigh-resolution ophthalmic optical coherence tomography (OCT) ex vivo imaging in an animal model with approximately 1.2-microm axial resolution and in vivo imaging in patients with macular pathologies with approximately 3-microm axial resolution were demonstrated. Owing to the pump laser, this light source significantly reduces the cost of broadband OCT systems. Furthermore, the source has great potential for clinical application of spectroscopic and ultrahigh-resolution OCT because of its small footprint (500 mm x 180 mm including the pump laser), user friendliness, stability, and reproducibility.

The mammalian photoreceptor mosaic-adaptive design.

Unlike in birds and cold-blooded vertebrates' retinas, the photoreceptors of mammalian retinas were long supposed to be morphologically uniform and difficult to distinguish into subtypes. A number of new techniques have now begun to overcome the previous limitations. A hitherto unexpected variability of spectral and morphological subtypes and topographic patterns of distribution in the various retinas are being revealed. We begin to understand the design of the photoreceptor mosaics, the constraints of evolutionary history and the ecological specialization of these mosaics in all the mammalian subgroups. The review discusses current cytological identification of mammalian photoreceptor types and speculates on the likely "bottleneck-scenario" for the origin of the basic design of the mammalian retina. It then provides a brief synopsis of current data on the photoreceptors in the various mammalian orders and derives some trends for phenomena such as rod/cone dualism, spectral range, preservation or loss of double cones and oil droplets, photopigment co-expression and mono- and tri-chromacy. Finally, we attempt to demonstrate that, building on the limits of an ancient rod dominant (probably dichromatic) model, mammalian retinas have developed considerable radiation. Comparing the nonprimate models with the intensively studied primate model should provide us with a deeper understanding of the basic design of the mammalian retina.

Irregular S-cone mosaics in felid retinas. Spatial interaction with axonless horizontal cells, revealed by cross correlation.

In most mammals short-wavelength-sensitive (S) cones are arranged in irregular patterns with widely variable intercell distances. Consequently, mosaics of connected interneurons either may show some type of correlation to photoreceptor placement or may establish an independent lattice with compensatory dendritic organization. Since axonless horizontal cells (A-HC's) are supposed to direct all dendrites to overlying cones, we studied their spatial interaction with chromatic cone subclasses. In the cheetah, the bobcat, and the leopard, anti-S-opsin antibodies have consistently colabeled the A-HC's in addition to the S cones. We investigated the interaction between the two cell mosaics, using autocorrelation and cross-correlation procedures, including a Voronoi-based density probe. Comparisons with simulations of random mosaics show significantly lower densities of S cones above the cell bodies and primary dendrites of A-HC's. The pattern results in different long-wavelength-sensitive-L- and S-cone ratios in the central versus the peripheral zones of A-HC dendritic fields. The existence of a related pattern at the synaptic level and its potential significance for color processing may be investigated in further studies.

The photoreceptor mosaic.

The organisation of the human photoreceptor mosaic reflects evolutionary strategies for optimising visual information under a wide range of stimulus conditions: (1) The rod population dominates (max. 170,000/mm2 at c. 30 degrees sup.) except for the central 2 degrees and along the ora serrata. (2) Density of cone inner/outer segments reaches up to 300,000 mm2 in the fovea. A bundle of c. 300-500 foveolar cones are further distinguished by having their synaptic terminals located within the capillary-free zone. Radial displacement (> 350 microns) of foveal cone terminals may result in the lesion of two sets of cone pathways by perifoveal laser treatment. Along the ora serrata peripheral cone density (c. 4000) rises within a small rim (1 degree) to up to 20,000, but may be considerably decreased by cystoid degenerations. For the L- and M-cone subpopulations ratios of 2:1 to 1:1 and random arrangement are suggested. (3) Blue-sensitive (S-) cones constitute a regular and independent submosaic of c. 7% across the periphery. An annular maximum (1000-5000/mm2) at c. 1 degree surrounds the foveola. There density decreases and irregular zones lacking S-cones result in tritan deficiencies.

The opossum photoreceptors--a model for evolutionary trends in early mammalian retina.

The topography and spectral characteristics of mammalian photoreceptors correlate with both, the present ecological demands and the evolutionary history. The South American Opossum is a marsupial mammal with unspecialized habitus and crepuscular lifestyle. A sparse population of cones (max. = 3000/mm2) can be differentiated into four subtypes by morphological, topographical and immunocytochemical criteria. In spite of this unusual diversity the cone types can be split into two functional groups: The population of single cones labeled by antibody OS-2 for short wavelength sensitive pigments was ubiquitous but at very low densities (200/mm2). The single cones labeled by antibody (COS-1) against long wavelength sensitive pigments constitute the dominant population in the area centralis (2300/mm2). These two single cone types correlate with the pair typically present in placental mammals. Discrimination of spatial and color contrast may be provided by this "modern" set. The COS-1 labeled double and single cones bearing an oil droplet, display a different pattern by being restricted to the inferior (non-tapetal) half of the retina (max = 800/mm2). This additional set of cones with oil droplets and long wavelength pigments is a conservative feature of the opossum retina and other marsupials. As an accessory cone system it is possibly providing enhanced sensitivity at mesopic conditions. During the early evolution of nocturnal mammals with its prominent expansion of rod vision these cone types were conserved but then were lost in placental mammals. Thus the unique features of mammalian retinas are the result of two evolutionary steps: first a reduction of cone based vision, followed by a secondary differentiation of photopic vision and behaviour relying on the remaining set of cones.

Photoreceptors in a primitive mammal, the South American opossum, Didelphis marsupialis aurita: characterization with anti-opsin immunolabeling.

The retinas of placental mammals appear to lack the large number and morphological diversity of cone subtypes found in diurnal reptiles. We have now studied the photoreceptor layer of a South American marsupial (Didelphis marsupialis aurita) by peanut agglutinin labeling of the cone sheath and by labeling of cone outer segments with monoclonal anti-visual pigment antibodies that have been proven to consistently label middle-to-long wavelength (COS-1) and short-wavelength (OS-2) cone subpopulations in placental mammals. Besides a dominant rod population (max. = 400,000/mm2) four subtypes of cones (max. = 3000/mm2) were identified. The outer segments of three cone subtypes were labeled by COS-1: a double cone with a principal cone containing a colorless oil droplet, a single cone with oil droplet, and another single cone. A second group of single cones lacking oil droplets was labeled by OS-2 antibody. The topography of these cone subtypes showed striking anisotropies. The COS-1 labeled single cones without oil droplets were found all over the retina and constituted the dominant population in the area centralis located in the temporal quadrant of the upper, tapetal hemisphere. The population of OS-2 labeled cones was also ubiquitous although slightly higher in the upper hemisphere (200/mm2). The COS-1 labeled cones bearing an oil droplet, including the principal member of double cones, were concentrated (800/mm2) in the inferior, non-tapetal half of the retina. The two spectral types of single cones resemble those of dichromatic photopic systems in most placental mammals. The additional set of COS-1 labeled cones is a distinct marsupial feature. The presence of oil droplets in this cone subpopulation, its absence in the area centralis, and the correlation with the non-tapetal inferior hemisphere suggest a functional specialization, possibly for mesopic conditions. Thus, sauropsid features have been retained but probably with a modified function.

A low cost computer aided design (CAD) system for 3D-reconstruction from serial sections.

This paper describes an approach to computer-assisted 3D-reconstruction of neuronal specimens based on a low cost yet powerful software package for a personal computer (Atari ST). It provides an easy to handle (mouse driven) object editor to create 3D models of medium complexity (15,000 vertices) from sections or from scratch. The models may be displayed in various modes including stereo viewing and complex animation sequences.

Iso-orientation areas in the foveal cone mosaic.

The quality of the foveal cone mosaic in human and primate retinas is a basic parameter of spatial vision function. The present study uses digital-texture analysis procedures to analyze the crystalline order of inner segment sections containing the rod-free portions of foveal cone mosaics. Definition of the cone cross-sectional centers made possible by adequate preprocessing allows precise mapping of lattice vertices and differentiation of hexagonal positions by procedures for direct neighbor recognition. In a further step, the existence of subunits within the hexagonal areas is revealed by the determination of axial orientation. The lattice of the subunits is characterized by similar orientation and high positional correlation of its hexagonal units. The axial orientation of the areas differs from that of neighboring subunits by angular shifts of 10-15 deg and linear series of nonhexagonal irregularities demarcate the borders. Although larger patches with continuous hexagonal order occur in the surrounding rod-free regions, elevated degrees of disorder (30%) are found within the foveolar center (ca. 300 cones). Analysis of a mosaic showing labeled B cones (Szél et al., 1988) demonstrates that lattice disorder is in part associated with the blue cone subpopulation. The foveal mosaic from a glaucomatuous eye reveals severe lattice degradation throughout the rod-free zone, presumably due to extensive receptor loss. The low-frequency superstructure results in local sets of sampling grids (5'-8') with differing orientational bias. Besides a horizontal/vertical difference of mosaic compression (ca. 1:1.15), the present analysis gives no hints for the existence of systematic meridional anisotropies at the receptor mosaic level. The study reveals a discontinuous organization of the foveal mosaic and points to possible sources for the induction and location of lattice disorder.

Background-induced flicker enhancement in cat retinal horizontal cells. I. Temporal and spectral properties.

1. Dim backgrounds can enhance small-spot flicker responses of cat retinal horizontal cells by a factor of 2 or more. 2. Intracellular marking with horseradish peroxidase (HRP) reveals that this enhancement effect occurs in--but is not necessarily limited to--the cone-connected, A-type horizontal cell. 3. Flicker amplitudes decrease over a frequency range from 3 to 36 Hz of square-wave photic stimulation. There is little evidence of flicker-response enhancement at 3 Hz. Flicker-response enhancement is typically 2-6 times larger at 35 than at 6 Hz. 4. Inspection of flicker waveforms indicates both a scaling-up of response signals with backgrounds and a distortion composed of 2- to 5-ms-latency decrease, expressed primarily within a quick component of OFF-repolarization. 5. Flicker enhancement first increases as a function of background irradiance and then decreases. The increasing limb has the dynamic range and spectral sensitivity of cat rods (507-nm peak). Enhancement is maintained during rod after-effects. The decreasing limb of the background-versus-intensity function results from light adaptation of cat, long-wavelength (red) cones. 6. The flicker responses themselves peak spectrally at approximately 555 nm and reflect only the activity of cat long-wavelength (red) cones, without evidence of intermixing of other photoreceptor mechanisms. 7. Thus within the first synaptic layer of the cat visual system, rod signals interact with the flicker responses of red cones, both increasing cone-signal amplitudes and modifying cone-signal waveforms. 8. The results are closely analogous to "suppressive rod-cone interaction" (SRCI) as described in human psychophysics. 9. An outer-plexiform-layer circuit involving rods, horizontal cells and cones may mediate rod-induced enhancement of cone flicker. This being the case, notions of horizontal-cell feedback interactions with cones may have to be modified and extended. A specific feedback model is elaborated in the companion paper.

Identification of a subtype of cone photoreceptor, likely to be blue sensitive, in the human retina.

A minor population of cone photoreceptors (called B-cones) can be distinguished from the major population (called R-cones) on morphological criteria as seen by light microscopy in foveal and peripheral human retina. The B-cones are characterized by a longer inner segment projecting into subretinal space, a larger-diameter inner segment, an increased staining intensity of the inner segment, and a different distribution relative to the R-cones in the cone mosaic. B-cones occur even in the foveolar center (3-5%) and rise to a maximum (15%) in the foveolar slope. They can also be identified in peripheral retina where they form 7-10% of the total cone population. The B-cone population follows the distribution profile postulated for the blue-sensitive system from histochemical studies on monkeys and from psychophysical studies on humans. The B-cones also share many of the same morphological features of the putative blue cones of the ground squirrel and monkey retinas. For these reasons we suggest that our B-cone group is the blue cone population of the human retina.

Telodendrial contacts between foveolar cone pedicles in the human retina.

The synaptic pedicles of foveolar cones in the human retina contact each other by means of telodendrial processes. Thus direct lateral coupling of photoreceptor terminals exists even in the area of highest acuity function.

Characterization of the color related receptor mosaic in the ground squirrel retina.

Light microscopic and histochemical studies reveal that the retina of the European ground squirrel (Citellus citellus L.) contains a mosaic pattern of two cone types and a small population of rods. A minority (7%) of the cones can be characterized by their ellipsoids having larger diameters and increased staining density over the majority population. Exposure to green light selectively elicited intense NBT-diformazan labeling in the major population of cones while the larger diameter cone type was labeled after blue illumination. The two cone subpopulations are probably the blue and green cone types of ground squirrel protanopic color vision.