Maureen K. Powers

Development of the visual system

Many aspects of the retinal structure and visual function of fishes are typical of all vertebrates. Colour vision, based on the presence of multiple primary cone pigments (Marks, 1965) and on the colour-opponent organization of retinal ganglion cells (Wagner et al., 1960), is perhaps the best known example. While research on such typical properties has contributed greatly to our understanding of the neural basis of vertebrate vision, other properties deserve equal interest because they are not typical. The development of the visual system is an example of this kind of property.

MIGRATION OF LIMULUS FOR MATING: RELATION TO LUNAR PHASE, TIDE HEIGHT, AND SUNLIGHT

In the spring, horseshoe crabs (Limulus polyphemus) along the eastern coast of North America migrate toward shore to build nests close to the waters edge. In 1984 the mating season near Woods Hole, Massachusetts, extended from 14 May to 7 July. Mating activity during this period fluctuated with the phase of the moon, the height of the tide, and diurnal changes in daylight. As the moon approached new and full phases, large numbers of animals migrated into the intertidal zone to mate and build nests. They appeared 1-2 h before high tide, and returned to deep water about 2 h after high tide. No mating activity occurred during low tides. The two daily high tides in this region are unequal in height. The inequality is greatest during new and full moons. At these times most animals migrated toward shore on the higher tide, which occurred in the late afternoon and throughout the night. As the moon passed through quadrature the tidal inequality diminished and reversed. Mating activity changed accordingly: shore ...

Daily fluctuations in the detectability of dim lights by humans

Several aspects of retinal physiology are known to vary on daily or circadian cycles, suggesting that certain aspects of visual function might also vary. We measured absolute threshold--the dimmest visual stimulus that can be reliably detected--in seven human observers at mid-day (1200-1400) and mid-night (0000-0200), by means of standard psychophysical techniques. Five of the subjects were slightly more sensitive at night: they could detect stimuli that were on the average 20% dimmer at night than during the day, and this difference was not due to changes in the criterion for detection. Three of four subjects whose thresholds were measured hourly while they remained otherwise in total darkness for greater than 24 hours showed changes in absolute visual threshold that correlated with hourly changes in body temperature. These results suggest that the ability to detect dim lights varies systematically with time of day in many human observers.

MECHANISMS OF LIGHT ADAPTATION IN RAT RETINA

Green, Dowling, Siegel and Ripps (1975) J. gen. Physiol. 65, 483-502 found that both receptors and post-receptoral elements regulate the process of light adaptation in skate retina. As a test of the generality of this conclusion, we repeated their experiments on the retina of the intact albino rat. Increment threshold and intensity-response functions of aspartate-isolated receptor potentials, ERG b-waves and single retinal ganglion cell axons were measured, and all were found to be similar to those described by Green et al. (1975). In particular, the adaptive properties of the b-wave and ganglion cells were similar to each other, but different from those of the receptor potential. We conclude that the retinal mechanisms of light adaptation are similar in rat and skate.

Visual function in regenerating teleost retina following cytotoxic lesioning.

Teleost fish retinas can regenerate in vivo in adulthood. Retinal and visual function was assessed in adult goldfish following comprehensive retinal destruction by intraocular injection of ouabain. Electroretinograms (ERGs) and the dorsal light reflex (DLR) were used to evaluate the return of visual function. ERGs were detectable in regenerating eyes 50 to 70 days following ouabain injection. Amplitudes of both a- and b-waves increased steadily through day 210 following ouabain treatment, at which time a-wave amplitude was 90% and b-wave amplitude approached 50% of the contralateral control eye. The progressive gain observed in the a-wave was attributed to photoreceptor regeneration. The increase in b-wave amplitude was attributed to an increase in the number of inner nuclear layer cells and the number and efficacy of neuronal connections to or within the inner retina. The photopic spectral sensitivity of the b-wave in regenerating retina closely matched the intrafish control retina, suggesting that the relative numbers of cone photoreceptors was normal in regeneration. The recovery of the DLR (indicated by improved postural balance during regeneration) paralleled electrophysiological gains during retinal regeneration. Fish displayed a marked longitudinal body imbalance toward the control eye following retinal destruction. Improvement in equilibrium was correlated with increasing b-wave amplitudes. When the b-wave reached 50% of control amplitude (30 weeks), normal posture was restored. The return of the ERG indicates that photoreceptors and their synaptic connections must be functional in regenerating retina. Failure of the retina to regenerate produced an abnormal DLR that persisted through 30 weeks and ERGs were not measurable. The return of normal equilibrium indicates that the regenerating retina can establish central connections to the brain, and that the regenerated connections can mediate functional visual behavior.

Visual performance of horseshoe crabs day and night

A circadian clock modulates the structure and function of the lateral eyes of Limulus polyphemus, greatly increasing their sensitivity at night. During the mating season, male Limulus are visually attracted both day and night to females and objects that resemble females. This paper asks how well Limulus can see day and night, and whether the circadian changes in retinal sensitivity might influence the ability of these animals to find mates. We recorded the visual behavior of male and female horseshoe crabs in the vicinity of an object--a cement hemisphere (29.5 cm diameter) similar in size and shape to a female horseshoe crab--placed in a mating area near Mashnee Dike, Bourne, Massachusetts. Males oriented toward this target from an average distance of 0.94 m during the day and 0.88 m at night; and females appeared to avoid the target. We conclude that males can see potential mates at night almost as well as they can during the day. Apparently the circadian changes in the retina help compensate for the daily changes in illumination in the animals normal environment. This study provides the first evidence for a role of visual circadian rhythms in an animals natural behavior.

Spectral sensitivity of ON and OFF responses from the optic nerve of goldfish.

The vertebrate retina processes visual information in parallel neural pathways known as the ON and OFF pathways. These pathways encode increments and decrements of light independently as excitatory responses. We examined the photopic spectral response of ON and OFF mechanisms in goldfish by measuring the sensitivity of optic nerve responses to the onset and termination of stimuli of various wavelengths. Using various adapting backgrounds, we found that the ON and OFF responses have different spectral sensitivities. The weighting of the cone inputs to the responses was estimated by an algebraic summation model. This model suggests that for the ON response, input from S-cones is stronger and more independent than for the OFF response, and M- and L-cones show stronger antagonism in the ON response than in the OFF response. The OFF response probably receives input from all cone types, but spectral antagonism is weak and its dominant input is from L-cones.

Visual detection by the rod system in goldfish of different sizes.

New rods are continually generated and inserted across the entire differentiated retina in juvenile and adult goldfish; no other retinal cells share this characteristic. How does the preferential addition of rods affect visual function? To examine the relation between continued rod addition and visual sensitivity, we measured absolute threshold in fish of different sizes. Twenty-nine fish were trained in a classical conditioning paradigm, and psychometric functions were obtained for each of them for detection of a 532 nm light 5 sec in duration, 140 deg in angular subtense, presented while the fish was fully dark adapted. We found that absolute threshold (expressed in terms of retinal photon density) was lower in larger fish, but by a very small amount; on average, large fish (15.4 +/- 0.5 cm standard body length) were 1.45 times more sensitive than small fish (4.3 +/- 0.3 cm). Morphometric analysis showed that the planimetric density of rods in goldfish retina increases at a similar rate between small and large fish, while the density of retinal ganglion cells declines between small and large fish (by a factor of 3.8). The ratio of rods to ganglion cells (a possible indicator of neural convergence) increased, but by a factor that is too large to reconcile with the psychophysical results (5.3 x). The results suggest that absolute visual threshold in the goldfish is closely related to the density of rods in the retina.

Psychosocial Findings in Radial Keratotomy Patients Two Years After Surgery

In a psychosocial study of patients who participated in an evaluation of the visual, refractive and keratometric results of radial keratotomy, respondents stated that their primary reasons for electing the operation related to anticipated changes in vision. Improving appearance was not reported as a primary reason for seeking radial keratotomy by these patients. When asked two years after surgery if their vision had improved, remained the same, or worsened, 94.3% reported improvement over preoperative vision. About 40% still wear corrective lens, 26% full time and 14% only part of the time. Most patients (84.1%) reported overall satisfaction with the surgical outcome. Patient satisfaction was strongly related to perceived improvement of vision after surgery, and not to patient self-esteem, to changes in appearance or lifestyle brought about by the operation, or to having a particular physician perform the surgery.

Spectral sensitivity of the electroretinogram b-wave in dark-adapted goldfish

The action spectrum of the ERG b-wave was measured under dark-adapted conditions in intact goldfish (Carassius auratus). It is substantially broader than the absorption spectrum of goldfish rod porphyropsin. Neither prolonged dark adaptation nor removal of possible efferent neural activity affected its shape. Moreover, a 682-nm background did not produce a selective loss of sensitivity to long wavelengths. The results imply that the spectral sensitivity of the b-wave in dark-adapted goldfish reflects the influence of at least two photoreceptor types which act as a single univariant mechanism near absolute threshold.