Eye Optics in Semiaquatic Mammals for Aerial and Aquatic Vision.

Based on anatomical measurements of refractive structures in the eye, the positions of focused images were computed for several groups of semiaquatic mammals: rodents, a nonpinniped semiaquatic carnivore (the sea otter), and pinniped carnivores (seals, sea lions, and the walrus). In semiaquatic rodents, eye optics enable emmetropia in the air but cause substantial hypermetropia in the water. In semiaquatic carnivores, there are several mechanisms for amphibious vision that focus images on the retina in both air and water. These mechanisms include the potential for a substantial change in the lens shape of sea otters and the presence of the corneal emmetropic window in pinnipeds. The results suggest that several groups of mammals that independently adapted to aquatic environments vary in how their visual systems adapted to aquatic vision.

Retinal Ganglion Cell Topography and Retinal Resolution in the Baikal Seal (Pusa sibirica).

The total number, size, topographic distribution, and cell density of ganglion cells were studied in retinal wholemounts of Baikal seals (Pusa sibirica). The ganglion cell size varied from 10 to 38 µm. A distinct cell group consisted of large ganglion cells of more than 30 µm in diameter. The topographic distribution of ganglion cells showed a definite area of high cell density similar to the area centralis of terrestrial carnivores. This area was located approximately 6-7 mm dorsotemporally of the geometric center of the wholemount. In this area, the peak cell densities in two wholemounts were 3,800 and 3,400 cells/mm2 (mean 3,600 cells/mm2). With a posterior nodal distance of 24 mm (underwater), this density corresponds to 631 cells/square degree. These values predict a retinal resolution of 2.4' in water and 3.0' in air. The topographic distribution of large cells featured the highest density in the same location as the total ganglion cell population.

Ocular anatomy, ganglion cell distribution and retinal resolution of a killer whale (Orcinus orca).

Retinal topography, cell density and sizes of ganglion cells in the killer whale (Orcinus orca) were analyzed in retinal whole mounts stained with cresyl violet. A distinctive feature of the killer whale's retina is the large size of ganglion cells and low cell density compared to terrestrial mammals. The ganglion cell diameter ranged from 8 to 100 µm, with the majority of cells within a range of 20-40 µm. The topographic distribution of ganglion cells displayed two spots of high cell density located in the temporal and nasal quadrants, 20 mm from the optic disk. The high-density areas were connected by a horizontal belt-like area passing below the optic disk of the retina. Peak cell densities in these areas were evaluated. Mean peak cell densities were 334 and 288 cells/mm(2) in the temporal and nasal high-density areas, respectively. With a posterior nodal distance of 19.5 mm, these high-density data predict a retinal resolution of 9.6' (3.1 cycles/deg.) and 12.6' (2.4 cycles/deg.) in the temporal and nasal areas, respectively, in water.

Ganglion cell distribution and retinal resolution in the Florida manatee, Trichechus manatus latirostris.

The topographic organization of retinal ganglion cells was examined in the Florida manatee (Trichechus manatus latirostris) to assess ganglion cell size and distribution and to estimate retinal resolution. The ganglion cell layer of the manatee's retina was comprised primarily of large neurons with broad intercellular spaces. Cell sizes varied from 10 to 60 µm in diameter (mean 24.3 µm). The retinal wholemounts from adult animals measured 446-501 mm(2) in area with total ganglion cell counts of 62,000-81,800 (mean 70,200). The cell density changed across the retina, with the maximum in the area below the optic disc and decreasing toward the retinal edges and in the immediate vicinity of the optic disc. The maximum cell density ranged from 235 to 337 cells per millimeter square in the adult retinae. Two wholemounts obtained from juvenile animals were 271 and 282 mm(2) in area with total cell numbers of 70,900 and 68,700, respectively (mean 69,800), that is, nearly equivalent to those of adults, but juvenile retinae consequently had maximum cell densities that were higher than those of adults: 478 and 491 cells per millimeter square. Calculations indicate a retinal resolution of ∼19' (1.6 cycles per degree) in both adult and juvenile retinae.

Retinal ganglion cell layer of the Caspian seal Pusa caspica: topography and localization of the high-resolution area.

Retinal topography, cell density and sizes of ganglion cells in the Caspian seal (Pusa caspica) were analyzed in retinal whole mounts stained with cresyl-violet. The topographic distribution of ganglion cells displayed an area of high cell density located in the temporal quadrant of the retina and was similar to the area centralis of terrestrial carnivores. It extended nasally, above the optic disk, as a streak of increased cell density. In different whole mounts, the peak cell density in the high-density area ranged from 1,684 to 1,844 cells/mm² (mean 1,773 cells/mm²). The cell density data predict a retinal resolution of around 8.5 cycles/degree in water. A distinctive feature of the Caspian seal's retina is the large size of ganglion cells and the low cell density compared to terrestrial mammals. The ganglion cell diameter ranged from 10 to 58 µm. Cell size histograms featured bimodal patterns with groups of small and large ganglion cells. The large cells appeared similar to α-cells of terrestrial mammals and constituted 7% of the total ganglion cell population.

Adaptive features of aquatic mammals' eye.

The eye of aquatic mammals demonstrates several adaptations to both underwater and aerial vision. This study offers a review of eye anatomy in four groups of aquatic animals: cetaceans (toothed and baleen whales), pinnipeds (seals, sea lions, and walruses), sirenians (manatees and dugongs), and sea otters. Eye anatomy and optics, retinal laminar morphology, and topography of ganglion cell distribution are discussed with particular reference to aquatic specializations for underwater versus aerial vision. Aquatic mammals display emmetropia (i.e., refraction of light to focus on the retina) while submerged, and most have mechanisms to achieve emmetropia above water to counter the resulting aerial myopia. As underwater vision necessitates adjusting to wide variations in luminosity, iris muscle contractions create species-specific pupil shapes that regulate the amount of light entering the pupil and, in pinnipeds, work in conjunction with a reflective optic tapetum. The retina of aquatic mammals is similar to that of nocturnal terrestrial mammals in containing mainly rod photoreceptors and a minor number of cones (however, residual color vision may take place). A characteristic feature of the cetacean and pinniped retina is the large size of ganglion cells separated by wide intercellular spaces. Studies of topographic distribution of ganglion cells in the retina of cetaceans revealed two areas of ganglion cell concentration (the best-vision areas) located in the temporal and nasal quadrants; pinnipeds, sirenians, and sea otters have only one such area. In general, the visual system of marine mammals demonstrates a high degree of development and several specific features associated with adaptation for vision in both the aquatic and aerial environments.

Retinal topography of the harp seal Pagophilus groenlandicus.

The total number, size, topographic distribution, and cell density of ganglion cells were studied in retinal wholemounts of the harp seal Pagophilus groenlandicus. Ganglion cell size varied from 10 to 60 mum. A distinct group were large ganglion cells more than 30-35 mum in diameter which were similar to alpha-cells known in terrestrial mammals. The number of alpha-cells constituted 5.3-5.9% of the total ganglion cell population. The cell size distribution was bimodal, with the second mode composed of alpha-cells. The topographic distribution of ganglion cells showed a definite area of high cell density similar to area centralis of terrestrial carnivores. This area was located in the temporal retinal quadrant, 7-8 mm (16-18 degrees ) from the optic disk. In this area, the peak cell densities in two wholemounts were 2,500 and 1,650 (mean 2,075) cells/mm(2). With a posterior nodal distance of 25.5 mm (underwater), this density corresponded to 495 and 327 (mean 411) cells/deg(2). These values predict a retinal resolution of 2.7-3.3' (11.1-9.0 cycles/deg) in water and 3.6-4.4' (8.3-6.8 cycles/deg) in air. Topographic distributions of alpha-cells was qualitatively similar to that of the total ganglion cell population, but the density of alpha-cells constituted only a few percent (mostly 3-7.5%) of the total ganglion cell density.