Variability in mitochondria of zebrafish photoreceptor ellipsoids.

Ultrastructural examination of photoreceptor inner segment ellipsoids in larval (4, 8, and 15 days postfertilization; dpf) and adult zebrafish identified morphologically different types of mitochondria. All photoreceptors had mitochondria of different sizes (large and small). At 4 dpf, rods had small, moderately stained electron-dense mitochondria (E-DM), and two cone types could be distinguished: (1) those with electron-lucent mitochondria (E-LM) and (2) those with mitochondria of moderate electron density. These distinctions were also apparent at later ages (8 and 15 dpf). Rods from adult fish had fewer mitochondria than their corresponding cones. The ellipsoids of some fully differentiated single and double cones contained large E-DM with few cristae; these were surrounded by small E-LM with typical internal morphology. The mitochondria within the ellipsoids of other single cones showed similar electron density. Microspectrophotometry of cone ellipsoids from adult fish indicated that the large E-DM had a small absorbance peak (∼0.03 OD units) and did not contain cytochrome-c, but crocetin, a carotenoid found in old world monkeys. Crocetin functions to prevent oxidative damage to photoreceptors, suggesting that the ellipsoid mitochondria in adult zebrafish cones protect against apoptosis and function metabolically, rather than as a light filter.

Communication using eye roll reflective signalling.

Body reflections in the ultraviolet (UV) are a common occurrence in nature. Despite the abundance of such signals and the presence of UV cones in the retinas of many vertebrates, the function of UV cones in the majority of taxa remains unclear. Here, we report on an unusual communication system in the razorback sucker, Xyrauchen texanus, that involves flash signals produced by quick eye rolls. Behavioural experiments and field observations indicate that this form of communication is used to signal territorial presence between males. The flash signal shows highest contrast in the UV region of the visual spectrum (lambdamax approximately 380 nm), corresponding to the maximum wavelength of absorption of the UV cone mechanism in suckers. Furthermore, these cones are restricted to the dorsal retina of the animal and the upwelling light background is such that their relative sensitivity would be enhanced by chromatic adaptation of the other cone mechanisms. Thus, the UV cones in the sucker have optimal characteristics (both in terms of absorbance and retinal topography) to constitute the main detectors of the flash signal. Our findings provide the first ecological evidence for restricted distribution of UV cones in the retina of a vertebrate.

Ontogenetic changes in visual sensitivity of the parasitic salmon louse Lepeophtheirus salmonis.

The salmon louse, Lepeophtheirus salmonis, is an ectoparasitic copepod of salmonid fishes whose life cycle involves two broadly defined, free-living larval stages, the nauplius and the copepodid. After settling on a host, the copepodid goes through various transformations to become a mobile adult. We recorded swimming responses of free-swimming salmon lice at the naupliar, copepodid and adult stages to the onset (ON) and offset (OFF) of lights of varying spectral irradiance and polarization. Nauplii showed a prominent swim-up OFF response across the spectrum 352-652 nm, but no ON response. Copepodids exhibited a swim-up ON response and a passive (sinking) OFF response across the same spectral range. Adults showed active swim-up responses to both ON and OFF stimuli, although the OFF response was proportionately stronger. The spectral range of the adult ON and OFF responses was the same as that of the copepodids and slightly greater than that of the nauplii, which did not exhibit responses at 652 nm. The absolute sensitivity of copepodids under white light (approx. 10(-13) photons m(2) s(1)) was higher than that of nauplii (approx. 10(-17) photons(-1) m(2 )s, OFF response) and that of adult female lice (approx. 10(-14) photons(-1)m(2)s). This suggests that the naupliar visual system is best suited for detection of shadows (e.g. the host) under a bright light field (daylight hours), while copepodids and adults may be more specialized for host detection at crepuscular periods and during the night, when light levels are low. None of the developmental stages responded to the rotation of the plane of polarized light or exhibited any difference in directed response when polarized light was used in place of diffuse light.

The ontogeny of ultraviolet sensitivity, cone disappearance and regeneration in the sockeye salmon Oncorhynchus nerka.

This study examines the spectral sensitivity and cone topography of the sockeye salmon Oncorhynchus nerka throughout its life history with special emphasis on ultraviolet sensitivity. Electrophysiological recordings from the optic nerve show that ultraviolet sensitivity is greatly diminished at the smolt stage but reappears in adult fish weighing about 201 g. Concomitantly, light microscopy observations of the retina show that ultraviolet cones disappear from the dorsal and temporal retina at the smolt stage but reappear at the adult stage. These changes occur for sockeye salmon raised in fresh water or salt water after smoltification. In contrast to this ultraviolet cycle, the other cone mechanisms (short-, middle- and long-wavelength-sensitive) and the rod mechanism remain present throughout ontogeny. The natural appearance and disappearance of ultraviolet cones in salmonid retinas follows surges in blood thyroxine at critical developmental periods. Their presence coincides with times of prominent feeding on zooplankton and/or small fish that may be more visible under ultraviolet light. It is proposed that the primary function of ultraviolet cones in salmonids is to improve prey contrast.

Wavelength-dependent polarization orientation in Daphnia.

The ability to detect and use the polarization of light for orientation is widespread among invertebrates. Among terrestrial insects, the retinula cells that are responsible for polarization detection contain a single visual pigment, either ultraviolet or short (blue) wavelength sensitive. With the exception of a few aquatic insects, the visual pigments underlying polarization sensitivity in aquatic invertebrates have yet to be determined. Here we report that polarotaxis in Daphnia pulex, a freshwater crustacean, is wavelength dependent and most likely mediated by two visual pigments with absorbance maxima in the middle (green) and long wavelength (red) parts of the spectrum. This contrasts with the response of a closely related species, D. magna, in which polarotaxis is wavelength independent and based on a single middle wavelength visual pigment. The visual systems in Daphnia are the first among crustaceans shown to utilize a middle wavelength pigment for polarization detection and, in the case of D. pulex, the first shown to use more than one visual pigment for such a purpose.

Double-cone internal reflection as a basis for polarization detection in fish.

Some species of fish are able to discriminate, in addition to intensity and wavelength (color), the direction of polarization of visible light. Optical experiments on axially oriented retinal cones from trout and sunfish with use of two types of polarization microscope indicate anisotropic light transmission through paired cones. The measured linear birefringence of paired cone ellipsoids is consistent with the presence of membranous partitions. It is proposed that the partition between the two members of a paired cone, which often appears extensive and flat, functions as a dielectric mirror and that polarization-dependent reflection and refraction at this partition constitutes the underlying mechanism in the transduction of polarization into intensity variation at the photoreceptor's outer segments. We support this hypothesis with linear birefringence and linear dichroism measurements, histological evidence, large-scale optical model measurements, and theoretical calculations based on Fresnel's formulas.

Cone photoreceptor topography in the retina of sexually mature Pacific salmonid fishes.

We examined the retinal cone topography in sexually mature individuals from four species of Pacific salmonid fishes by using semithin plastic sections. We identified variations in cone density and cone arrangements and noted the presence of putative ultraviolet (UV) cones. Putative UV cones were found over an area extending dorsotemporally from the center of the retina. Because most of the putative UV cones are believed to disappear in early ontogeny, their presence over a large proportion (15-20%) of the surface area of the adult retina suggests that they may be reincorporated prior to or at sexual maturity, at least in rainbow trout. Cone density varied across the retina, with highest values at the peripheral margin. Relatively high densities were observed ventrotemporally (in all specimens) and, to a lesser extent, dorsonasally (7 of 11 specimens). The higher cone density in the ventrotemporal retina may represent a retinal specialization in the part of the visual field located above and in front of the animal. Lowest cone densities were found dorsocentrally and coincided approximately with the distribution of putative UV cones, raising the possibility that these cones may not be used in visual tasks requiring the higher visual acuity normally associated with higher cone densities. We also report a novel cone arrangement that consists of rows of double cones inserted between rows composed of single-double cone pairs alternating in position.

No evidence of polarization sensitivity in freshwater sunfish from multi-unit optic nerve recordings.

The sensitivities of two species of sunfish (Lepomis gibbosus and Lepomis cyanellus) to the electric field (E-vector) of polarized light were assessed by compound action potential recordings from the optic nerve of live fish. Under white light and long wavelength adapting backgrounds, two cone mechanisms were found with maximum sensitivities in the long wavelength (lambda max approximately 620 nm) and middle wavelength (lambda max approximately 530 nm) regions of the spectrum. In contrast to previous findings (Cameron & Pugh, 1991), no evidence of polarization sensitivity was observed for either species. We conclude from these results that post-larval sunfish do not exhibit polarization sensitivity.

Is the use of underwater polarized light by fish restricted to crepuscular time periods?

We measured the spectral distributions of the underwater total and polarized light fields in the upper photic zone of meso-eutrophic waters (i.e., blue-green waters containing medium to high chlorophyll a concentrations). Per cent polarization levels during the day were always lower than 40%, but at crepuscular times these values could increase to 67%. A corresponding change occurred in the spectral distribution, with proportionately more shorter wavelength photons contributing to the total spectrum during crepuscular periods. Electrophysiological recordings from the optic nerve of rainbow trout subjected to light stimuli of varying polarization percentages show that the animal's threshold for detecting polarized light is between 63 and 72%. These physiological findings suggest that the use of water-induced polarized light cues by rainbow trout and similar percomorph fish should be restricted to crepuscular time periods.