Dorsal Light Reflex Research Articles

Visually induced vertical vergence as a motion processing biomarker associated with postural instability

The present study explored visually induced vertical vergence (VIVV) as non-specific motion processing response. Healthy participants (7 male, mean age 28.57 ± 2.30; 9 female, mean age 27.67 ± 3.65) were exposed to optokinetic stimuli in an HTC VIVE virtual reality headset while VIVV, pupil-size, and postural sway was recorded. The methodology was shown to produce VIVV in the roll plane at 30 deg/s. Subsequent trials consisted of 40 s optokinetic motion in yaw, pitch, and roll directions at 60 deg/s, and radial optic flow; optokinetic directions were inverted after 20 s of motion. Median VIVV amplitude changes were normalized to the clockwise roll rotation, analysed, and correlated with changes in pupil-size and body sway. VIVV, pupil-size, and body sway were all affected by changes in optokinetic direction. Post-hoc analyses showed significant VIVV responses during optokinetic yaw and pitch rotations, as well as during radial optic flow stimulations. VIVV magnitudes were universally correlated with pupil-size and body sway. In conclusion, VIVV was expressed in all tested dimensions and may consequently serve as a visual motion processing biomarker. Failing to support binocularity while responding to optokinetic directionality, VIVV may reflect an eye-movement response associated with increased postural instability and stress, similar to a dorsal light reflex.

Visually Driven Behavior Assays to Study Functionality and Recovery after Damage of the Aged African Turquoise Killifish Visual System.

Loss of vision is a prominent feature of aging and vision is considered by many to be the most valuable sense to be lost. In our graying society, we are increasingly challenged by age-related deterioration of the central nervous system (CNS), as well as by age-associated neurodegenerative diseases and brain injuries, all often affecting the visual system and thus its performance. Here, we describe two visually driven behavior assays to evaluate visual performance upon aging or CNS damage in the fast-aging killifish. The first test, the optokinetic response (OKR), measures the reflexive eye movement triggered by motion in the visual field and allows assessment of visual acuity. The second assay, the dorsal light reflex (DLR), evaluates the swimming angle based on input of light coming from above. The OKR can be used to study the effect of aging on visual acuity as well as visual improvement and recovery after rejuvenation therapy or visual system injury or disease, whereas the DLR is best used to assess functional repair after a unilateral optic nerve crush.

Analysis of Visual Recovery After Optic Nerve Crush in Adult Zebrafish.

Zebrafish can successfully regenerate axons after optic nerve crush (ONC). Here, we describe two different behavioral tests to map visual recovery: the dorsal light reflex (DLR) test and the optokinetic response (OKR) test. The DLR is based on the tendency of fish to orient their back to a light source, and it can be tested by rotating a flashlight around the dorsolateral axis of the animal or by measuring the angle between the left/right body axis and the horizon. The OKR, in contrast, consists of reflexive eye movements triggered by motion in the visual field of the subject and is measured by placing the fish in a drum on which rotating black-and-white stripes are projected.

Time-dependent bidirectional effects of chronic caffeine on functional recovery of the dorsal light reflex after hemilabyrinthectomy in the goldfish Carassius auratus

Caffeine works through a variety of complex mechanisms to exert an often bidirectional set of functional and structural neurological changes in vertebrates. We investigated the effects of chronic caffeine exposure on functional recovery of the dorsal light reflex (DLR) in hemilabyrinthectomized common goldfish, Carassius auratus. In this lesion model, the unilateral removal of the vestibular organs results in a temporary loss of gravitationally modulated postural control which is quantifiable via the DLR. We compared the functional recovery over 24days of post-surgery goldfish continuously held in a caffeine solution of 2.5mg/L (n=10), 5.0mg/L (n=10), 10.0mg/L (n=11), or 0.0mg/L control (n=9). Comparison to a sham surgery group (n=11) indicated statistically significant changes in the DLR of all hemilabyrinthectomized fish on day 1. The control group recovered over the study period and approached, but did not reach sham surgery DLR. Although the caffeine-treated fishes appeared to initiate some postural recovery within the first 2weeks, beginning on day 10, all caffeine groups diverged from the control group with a deterioration of postural control. All three caffeine groups were significantly deficient in comparison with the control on days 10–24. These results suggest that caffeine exposure can at first be benign, but that high dosage or prolonged exposure hinders functional recovery.

Which way is up? Asymmetric spectral input along the dorsal-ventral axis influences postural responses in an amphibious annelid.

Medicinal leeches are predatory annelids that exhibit countershading and reside in aquatic environments where light levels might be variable. They also leave the water and must contend with terrestrial environments. Yet, leeches generally maintain a dorsal upward position despite lacking statocysts. Leeches respond visually to both green and near-ultraviolet (UV) light. I used LEDs to test the hypothesis that ventral, but not dorsal UV would evoke compensatory movements to orient the body. Untethered leeches were tested using LEDs emitting at red (632 nm), green (513 nm), blue (455 nm) and UV (372 nm). UV light evoked responses in 100 % of trials and the leeches often rotated the ventral surface away from it. Visible light evoked no or modest responses (12-15 % of trials) and no body rotation. Electrophysiological recordings showed that ventral sensilla responded best to UV, dorsal sensilla to green. Additionally, a higher order interneuron that is engaged in a variety of parallel networks responded vigorously to UV presented ventrally, and both the visible and UV responses exhibited pronounced light adaptation. These results strongly support the suggestion that a dorsal light reflex in the leech uses spectral comparisons across the dorsal-ventral axis rather than, or in addition to, luminance.

The Halpern Syndrome of Monocular Visual Vertigo: A Rare Cause of Monocular Eye Closure

Background: Halpern syndrome is a vestibular disorder characterized by vertigo, a tendency to tilt to one side, and spontaneous closure of the ipsilateral eye. Symptoms are exacerbated during monocular viewing with the ipsilateral eye and alleviated during monocular viewing with the contralateral eye. It has long been attributed to a mismatch between visual and vestibular input.Methods: Reexamination of Halpern syndrome in light of new concepts of visuo-vestibular disease.Results: The visually dependent postural disequilibrium in Halpern syndrome recapitulates the dorsal light reflex in fish, in which unequal luminance input to the two eyes evokes a tilt of the body toward the brighter side.Conclusion: Patients with Halpern syndrome close one eye to restore vertical postural orientation. Halpern syndrome may result from reactivation the dorsal light reflex in the setting of central vestibular disease.

Responses to Light In Two Eyeless Cave Dwelling Amphipods (Niphargus ictus and Niphargus frasassianus)

Abstract Two amphipod species, Niphargus ictus and N. frasassianus, which are endemic to Frasassi Caves, Italy, possess morphological features typical of most troglobitic species. Most notably they lack eyes. Although cave waters flow directly into the adjoining Sentino River and N. frasassianus is found within two meters of the resurgence, neither species is present at or further outside the mouth of the cave, which raises the question of how they avoid leaving the cave. It was hypothesized that these animals might be able to detect light, and could use light cues to remain inside the cave. Individuals of both species exhibited greater activity levels in the presence of bright vs. low light levels. Neither species exhibited the dorsal light reflex, but both showed weak negative phototaxis when exposed to bright light directed at them from above and below. N. frasassianus, tested in an apparatus that permitted them to travel freely between bright or low lighted areas, demonstrated negative phototaxis. The...

Visual function in regenerating teleost retina following surgical lesioning

Regeneration of the teleost retina following surgical extirpation of 25% to 100% of the neural retina was investigated in goldfish (Carrasius auratus) and sunfish (Lepomis cyanellus). The retina will regenerate following removal of up to 95% of the neural retina, however complete extirpation prevented regeneration. Visual sensitivity was assessed by examining components of the electroretinogram (ERG) and the dorsal light reflex (DLR) during regeneration. B-wave amplitudes in the experimental eyes increased throughout the study and central connections were reestablished as indicated by the progressive improvement in the dorsal light reflex. The recovery of visual function was closely correlated with retinal regeneration. Visual recovery progressed more slowly than following complete cytotoxic destruction of the mature retina (Mensinger & Powers, 1999) because the surgery removed a large number of the pluripotent cell population and restricted the number and distribution of regenerating foci.

Restoration of visual function following optic nerve regeneration in bluegill (Lepomis macrochirus) × pumpkinseed (Lepomis gibbosus) hybrid sunfish

Simple (dorsal light reflex) and complex (predator-prey interactions) visually mediated behaviors were used concurrently with morphological examination to assess restoration of visual function following optic nerve crush in bluegill (Lepomis macrochirus) x pumpkinseed (Lepomis gibbosus) hybrid sunfish. Regenerating optic nerve axons projected into the stratum opticum-stratum fibrosum et griseum superficiale by week 2, the stratum griseum centrale by week 4, and stratum album centrale by week 6. Initial projections into the laminae were diffuse and less stratified compared to controls. By week 12, the projection pattern of regenerating nerve fibers closely resembled the innervation of normal tecta. Visual improvements were correlated with increasing projections into the tectum. The dorsal light reflex improved from a 45 degrees vertical deviation following nerve crush to 4.5 degrees by week 16. Initial predator-prey interactions were exclusively mediated by the control eye. As regeneration progressed, there was a gradual expansion of the visual field. The reaction distance and attack angles within the visual field of the experimental eye were initially less than controls, however, these differences disappeared by week 10. Improvements in visual function were closely correlated with an increase of regenerating ganglion cell axons into the optic tectum indicating sufficient synaptogenesis to mediate both simple and complex visual behavior.

Visual behavior of adult goldfish with regenerating retina

To determine whether regenerating neural pathways can support visual behavior, adult goldfish (Carassius auratus) were injected intraocularly with ouabain and tested for the presence of reflexive visual behaviors (dorsal light reflex and optokinetic nystagmus) and the ability to respond to visual stimuli in a classical conditioning paradigm. All visual behaviors were absent or greatly diminished until 8 to 10 weeks, when retinal layering had returned. At 10 weeks post-ouabain, reflexive behaviors to supra-threshold stimuli were near normal; however the ability to detect supra-threshold stimuli in the conditioning paradigm did not recover until 13 weeks. Absolute dark-adapted threshold and light-adapted spectral sensitivity measured at 13 to 17 weeks were abnormal: Dark-adapted threshold was elevated by 1.5 log units and light-adapted spectral sensitivity was markedly narrower than normal. No responses to 50% contrast sinusoidal gratings could be obtained through ouabain-treated eyes using the classical conditioning technique, even though responses through the untreated eye remained. Results demonstrate that: (a) visually mediated behaviors return in goldfish with ouabain-treated retinas; (b) the time course of recovery of reflexive responses in luminance and spatial domains parallels return of ERG function and of tectal activity; and (c) visual function that is mediated by regenerating retina appears not to be as sensitive as vision via normally developed retinal pathways.

Acuity and contrast sensitivity of the bluegill sunfish and how they change during optic nerve regeneration

Spatial vision was studied in the bluegill sunfish, Lepomis macrochirus (9.5-14 cm standard length) to assess the limitations imposed by the optics of the eye, the retinal receptor spacing and the retinotectal projection during regeneration. Examination of images formed by the dioptric elements of the eye showed that spatial frequencies up to 29 c/ degrees could be imaged on the retina. Cone spacing was measured in the retina of fresh, intact eyes. The spacing of rows of double cones predicted 3.4 c/degrees as the cutoff spatial frequency; the spacing between rows of single and double cones predicted 6.7 c/degrees. Contrast sensitivity functions were obtained psychophysically in normals and fish with one regenerating optic nerve. Fish were trained to orient to gratings (mean luminance = 25 cd/m2) presented to either eye. In normals, contrast sensitivity functions were similar in shape and bandwidth to those of other species, peaking at 0.4 c/degrees with a minimum contrast threshold of 0.03 and a cutoff at about 5 c/degrees, which was within the range predicted by cone spacing. Given that the optical cutoff frequency exceeds that predicted by cone spacing, it is possible that gratings could be detected by aliasing with the bluegill's regular cone mosaic. However, tests with high contrast gratings up to 15 c/ degrees found no evidence of such detection. After crushing one optic nerve in three trained sunfish, recovery of visual avoidance, dorsal light reflex and orienting to gratings, were monitored over 315 days. At 64-69 days postcrush, responses to gratings reappeared, and within 2-5 days contrast sensitivity at low (0.15 c/degrees) and medium (1.0 c/ degrees) spatial frequencies had returned to normal. At a high spatial frequency (2.93 c/degrees) recovery was much slower, and complete only in one fish.

Orientation by polarized light in the crayfish dorsal light reflex: behavioral and neurophysiological studies

In decapod crustaceans, the dorsal light reflex rotates the eyestalk so that the dorsal retina faces the brightest segment of dorsal visual space. Stepwise displacements of white stripes elicit eyestalk rotations in the same direction as that of the stripe. Conversely, stepwise displacements of black stripes on a white background elicit eyestalk rotations in the opposite direction as that of the stripe. The reversal of the response with contrast inversion distinguishes the dorsal light reflex from an optokinetic reflex. When the visual scene is composed of polarized light, segmented by variations in e-vector orientation, displacement of segments containing near vertical e-vectors elicit responses similar to those elicited by a white stripe. Displacement of polarized stripes containing near horizontal e-vectors elicit eyestalk rotations similar to those elicited by a black stripe. The results are consistent with the use of polarized light in orientation. The stimulus conditions described above were also applied to visual interneurons (sustaining fibers) and oculomotor neurons and the results were generally in accord with the behavior. In the neural studies, it was possible to show that responses to polarized stripe displacements are predictable from the receptive field location and the neuron's polarization tuning function.

Physiologic Hyperphoria and the Dorsal Light Reflex in Humans with Normal Binocular Vision

We read with interest the study by Wang and Bedell (2005) that examined the influence of asymmetrical illumination on physiologic hyperphoria (an ocular movement qualitatively similar to dissociated vertical divergence) in normal humans. Of 30 subjects tested with a Maddox rod to subjectively assess diplopia, eight showed evidence of bilateral hyperphoria of the occluded eye when a dark occluder was used. Interestingly, four of these subjects displayed a physiologic hyperphoria that converted to a hypophoria of the occluded eye when an illuminated occluder was used, consistent with a human dorsal light reflex (Brodsky, 1999). We are concerned that the methodology of this study may have led the authors to the erroneous conclusion that “little or no vestige of the dorsal light reflex occurs in normal individuals.” First, diplopia was used as an endpoint rather than eye position. Physiologic hyperphoria is a tiny response, usually

Physiologic Hyperphoria and the Dorsal Light Reflex in Humans with Normal Binocular Vision: Reply to Brodsky & Van Rijn

Physiologic Hyperphoria and the Dorsal Light Reflex in Humans with Normal Binocular Vision: Reply to Brodsky & Van Rijn

Asymmetrical Vertical Phorias in Normal Subjects: The Influence of Unbalanced Illumination

Purpose: One goal of this investigation was to determine whether asymmetrical vertical hetereophorias, which are qualitatively similar to a dissociated vertical deviation (DVD), occur commonly in normal people. A second goal was to establish whether the magnitude or direction of vertical phoria depends on the illumination of the occluded eye, as would be expected if vertical eye position were influenced by a dorsal light reflex. Methods: Vertical phorias were estimated in 30 normal subjects using a Maddox rod in conjunction with a dark and an illuminated occluder. Results: On average, subjects exhibited a hypophoria of less than 1 min arc with both the dark and the illuminated occluder. Asymmetrical vertical phorias that were consistent with previously published criteria for a minute DVD were found in eight subjects when a dark occluder was used and in four subjects when an illuminated occluder was used. The proportions of subjects whose vertical phorias were consistent with a DVD did not differ significantly under the two conditions of occlusion. Conclusions: Asymmetrical vertical phorias that mimic a minute DVD do not occur commonly in normal people. Our finding that vertical phorias do not change systematically when an illuminated instead of a dark occluder is used suggests that normal subjects exhibit little or no vestige of the dorsal light reflex.

The response of squid and fish to changes in the angular distribution of light

This paper describes the responses of a squid (Alloteuthis subulata) and a fish (Trachurus trachurus) to changes in the angular distribution of light. An apparatus was made that simulated the angular distribution of daylight in the sea. The apparatus enabled the direction of the brightest light to be changed and the positions of the animals in response to these changes were observed. Squid viewed head-on were observed to roll by a maximum of 20° when the incident light source was at angles between 20° and 90° (where 0° is vertically downwards). When viewed laterally, i.e. in the pitch plane, the squid were observed to position themselves more closely with respect to the angle of the light source, they swam in a near vertical plane when the incident light source was at an angle of 90°. Swimming movements in the roll and pitch plane became more horizontal with positions of the light source between 90 and 180°. Horse mackerel, in contrast, inclined their dorsal surfaces to almost perfectly match the angle of the incident light source, even swimming upside-down when light came from below. These experiments also revealed that squid display a counter-shading chromatophore pattern (‘Flexible Countershading’) in response to light coming from the sides, which involves darkening the side of the body facing the brightest light. The use of chromatophores in this way may explain why the dorsal light reflex in squid is so weak compared to that of fish.

Dissociated vertical divergence: perceptual correlates of the human dorsal light reflex.

Dissociated vertical divergence (DVD) has been attributed to a human dorsal light reflex that emerges when single binocular vision is precluded in infancy. If this is the case, then DVD should be associated with a subjective sensation of tilt. Prospective examination of 9 patients with DVD and 9 control subjects to determine whether monocular occlusion and alternate occlusion induces a subjective sensation of visual tilt or body tilt. Alternate occlusion disclosed a tilt in the subjective visual vertical in 8 of the 9 patients with DVD and in none of 9 control subjects. On occlusion of the fixating eye, a vertical pencil positioned in the sagittal plane was perceived as instantaneously tilted, with its upper pole tipped toward the side of the covered eye. This visual tilt was quickly followed by a perceived rotation back to vertical, which coincided with the dorsally directed drift of the covered eye. In patients with DVD, monocular occlusion is associated with a subjective visual tilt that is annulled by a cyclovertical divergence movement of the eyes. This observation supports the notion that DVD is a human dorsal light reflex, which functions to restore vertical visual orientation when unequal binocular visual input evokes a subjective sensation of visual tilt.

D2-dopamine receptor blockade impairs motion detection in goldfish.

Under photopic illumination conditions, motion detection in goldfish is dominated by the long-wavelength-sensitive cone type (L-cone), and under scotopic conditions motion it is determined by rods (Schaerer & Neumeyer, 1996). The switch from rod-dominated to cone-dominated motion detection occurs during light adaptation. It has been suggested that dopamine acts as a neuronal light-adaptative signal. It is known that dopamine affects wavelength discrimination through D1-dopamine receptors (Mora-Ferrer & Neumeyer, 1996), and the dorsal light reflex through D1- and D2-dopamine receptors (Lin & Yazulla, 1994a). The purpose of this study was to determine whether dopamine influenced movement detection by goldfish, and if so, which dopamine receptor was involved. The D2-dopamine receptor antagonist sulpiride reduced the animal's sensitivity to the moving stimulus, whereas SCH 23390, a D1-dopamine receptor antagonist, did not have any effect. The effect of sulpiride is discussed in relation to known sulpiride effects on retinal neurons and the retinal pigment epithelium.

ACTH4-10, substance P, and dizolcipine (MK-801) accelerate functional recovery after hemilabyrinthectomy in goldfish.

In this study, we evaluated the goldfish model of hemilabyrinthectomy for investigating potential recovery-promoting drugs. In this lesion model, the unilateral removal of the labyrinth induces a postural imbalance in response to light (Dorsal Light Reflex), from which the animals can recover over time. The behavioral effects of two neuropeptides were tested–namely, of substance P and ACTH4-10, both of which are known to promote functional recovery in several other lesion models. Furthermore, the effect of MK- 801, an antagonist of the glutamatergic NMDAreceptor subtype, was tested because this substance has also been shown to exert a neuroprotective effect. After lesion of the right labyrinth, the animals (n=12) were treated intraperitoneally daily either with vehicle (n=12), substance P (n=11), ACTH4-10 (n=12), or MK- 801 (n=12). Another group (n=11), which served as a non-lesion control, did not receive hemilabyrinthectomy or systemic injections. The lesion group, treated post-operatively with vehicle, did not recover from the postural deviation over the 24-d testing period. In contrast, all three test substances accelerated the functional recovery after unilateral labyrinthectomy. The decrease of the dorsal light reflex persisted even after cessation of drug treatment after 20d. The results indicate that using the dorsal light reflex in the model of hemilabyrinthectomy in goldfish provides a useful approach to studying the ability of potential new neurotrophic or neuroprotective drugs to promote functional recovery.

Dissociated vertical divergence: a righting reflex gone wrong.

Dissociated vertical divergence (DVD) is an ocular motor disorder characterized by a slow, upward drift of 1 eye when the other eye is fixating a target. I propose that DVD is a dorsal light reflex in which asymmetrical visual input to the 2 eyes evokes a vertical divergence movement of the eyes. This primitive visuo-vestibular reflex functions as a righting response to restore vertical orientation in lower lateral-eyed animals by equalizing binocular visual input. The dorsal light reflex is suppressed in humans but can manifest as DVD when early-onset strabismus precludes normal binocular development.