Scanning behavior by larvae of the predacious diving beetle, <Emphasis Type="Italic">Thermonectus marmoratus</Emphasis> (Coleoptera: Dytiscidae) enlarges visual field prior to prey capture

Larvae of the predaceous diving beetle Thermonectus marmoratus bear six stemmata on each side of their head, two of which form relatively long tubes with linear retinas at their proximal ends. The physical organization of these eyes results in extremely narrow visual fields that extend only laterally in the horizontal body plane. There are other examples of animals possessing eyes with predominantly linear retinas, or with linear arrangements of specific receptor types. In these animals, the eyes, or parts of the eyes, are movable and perform scanning movements to increase the visual field. Based on anatomical data and observations of relatively transparent, immobilized young larvae, we report here that T. marmoratus larvae are incapable of moving their eyes or any part of their eyes within the head capsule. However, they do perform a series of bodily dorso-ventral pivots prior to prey capture, behaviorally extending the vertical visual field from 2° to up to 50°. Frame-by-frame analysis shows that such behavior is performed within a characteristic distance to the prey. These data provide first insights into the function of the very peculiar anatomical eye organization of T. marmoratus larvae.     

Fundamental differences in the optical structure of the eyes of nocturnal and diurnal mosquitoes

We have studied the anatomy and optics of the eyes of a range of mosquito species from the wholly dark-active blood-feeding Anopheles gambiae to the diurnal plant-feeder Toxorhynchites brevipalpis. Consistent with studies by Satô in the 1950s, we find that dark-active and crepuscular species have short fused rhabdoms with a conical construction. This maximises the amount of light the rhabdoms receive from the almost hemispherical wide-aperture lenses. Toxorhynchites, on the other hand, has long narrow rhabdomeres that are separated from each other over their entire length, and so resemble the open rhabdoms of advanced flies (Brachycera and Cyclorrhapha). These findings are confirmed by studies of the pseudopupil, whose form indicates the layout of the rhabdomere tips in the focal plane of each ommatidial lens. In anopheline species the pseudopupil is a single undivided ellipse, indicating a fused rhabdom structure, whereas in Toxorhynchites there is a ring of six outer elements surrounding a central one. This means that each rhabdomere views a separate direction in space, and our measurements indicate that, as in higher Diptera, adjacent rhabdomeres share their fields of view with one of the rhabdomeres in the immediately adjacent ommatidia. This in turn means that in the diurnal type of mosquito eye there is a basis for neural superposition, but the fused construction of anopheline rhabdoms precludes this. The Aedes species studied were similar to Anopheles but with lenses of less extreme aperture, and Sabethes cyaneus, a diurnal blood-feeder, was intermediate in structure, with fused conical rhabdoms in the centre of the eye and unfused rhabdomeres around the periphery.     

The eyes of a patrolling butterfly: visual field and eye structure in the Orange Sulphur, Colias eurytheme (Lepidoptera, Pieridae)

Sensory information plays a critical role in determining an animal's behavior on both proximate and evolutionary timescales. Butterflies, like many other insects, use vision extensively over their lifetimes, and yet relatively little work has been published to date on their visual capabilities. We describe the visual system of a pierid butterfly, Colias eurytheme, with the ultimate goal of better understanding its role in shaping the behavior of this animal. We made several measurements: visual field dimensions, eye surface area, interommatidial angle (Deltaphi), facet diameter (D), and eye parameter (p). C. eurytheme had a large visual field and considerable regional variation in visual acuity, as inferred by Deltaphi and D. When compared to females, males had larger eye surface areas, smaller Deltaphi, and larger D in all regions except ventrally. Both sexes had proportionally large eye surface areas compared to other butterflies. Minimum p in males was small, indicating that some regions of their eyes may operate close to the diffraction limit. Finally, we found that both eye surface area and D scaled positively, but with negative allometry to body size. We discuss the relevance of these visual characteristics to the biology and behavior of C. eurytheme.     

Visual pigment spectra of the comma butterfly,<Emphasis Type="Italic"> Polygonia c-album</Emphasis>, derived from in vivo epi-illumination microspectrophotometry

The visual pigments in the compound eye of the comma butterfly, Polygonia c-album, were investigated in a specially designed epi-illumination microspectrophotometer. Absorption changes due to photochemical conversions of the visual pigments, or due to light-independent visual pigment decay and regeneration, were studied by measuring the eye shine, i.e., the light reflected from the tapetum located in each ommatidium proximal to the visual pigment-bearing rhabdom. The obtained absorbance difference spectra demonstrated the dominant presence of a green visual pigment. The rhodopsin and its metarhodopsin have absorption peak wavelengths at 532 nm and 492 nm, respectively. The metarhodopsin is removed from the rhabdom with a time constant of 15 min and the rhodopsin is regenerated with a time constant of 59 min (room temperature). A UV rhodopsin with metarhodopsin absorbing maximally at 467 nm was revealed, and evidence for a blue rhodopsin was obtained indirectly.     

How bees detect coloured targets using different regions of their compound eyes

Honeybees Apis mellifera detect coloured targets presented to the frontal region of their compound eyes using their colour vision system at larger visual angles (α > 15°), and an achromatic visual system based on the long-wave photoreceptor type at smaller visual angles (5° < α < 15°). Here we examine the capability of the dorsal, ventral and frontal regions of the eye for colour detection. The minimum visual angle α_min at which the bees detect a stimulus providing both chromatic contrast and receptor-specific contrasts to the three receptor types varies for the different regions of the eye: 7.1 ± 0.5° for the ventral region, 8.2 ± 0.6° for the dorsal region and 4.0 ± 0.5° for the frontal region. Flight trajectories show that when the target was presented in the horizontal plane, bees used only the ventral region of their eyes to make their choices. When the targets appeared dorsally, bees used the frontodorsal region. This finding suggests that pure dorsal detection of coloured targets is difficult in this context. Furthermore, α_min in the ventral plane depends on receptor-specific contrasts. The absence of S-receptor contrast does not affect the performance (α_min = 5.9 ± 0.5°), whilst the absence of M- and L-receptor contrast significantly impairs the detection task. Minimal visual angles of 10.3 ± 0.9° and 17.6 ± 3°, respectively, are obtained in these cases. Thus, as for many visual tasks, the compound eye of the honeybee shows a regionalisation of colour detection that might be related to peripheral or central specialisations. Accepted: 28 September 1999     

The unusual visual system of the Strepsiptera: external eye and neuropils

Adult males of the insect order Strepsiptera are characterized by an unusual visual system that may use design principles from compound as well as simple eyes. The lenses of this eye are unusually large and focus images onto extended retinae. The light-gathering ability of the lens is sufficient to resolve multiple points of an image in each optical unit. We regard each unit as an independent image-forming eye that contributes an inverted partial image. Each partial image is re-inverted by optic chiasmata between the retinae and the lamina, where the complete image could be assembled from the neighboring units. The lamina, medulla and lobula are present, but their organization into cartridges is not clearly discernable. Fluorescent fills, whole-tissue stains, and synaptotagmin immunohistochemistry show that the optic neuropils nevertheless are densely packed, and that several parallel channels within the medulla underlie each of the lenses. The size and shape of the rhabdoms, as well as a relatively slow flicker-fusion frequency could suggest that these eyes evolved through a nocturnal life stage.Abbreviations O object size - U object distance - I image size - f focal length - A lens aperture - D lens diameter - interommatidial angle - S light sensitivity of optical system     

Binocular vision and accommodation in prey-catching chameleons

We have studied the role of accommodation and binocular convergence in the predatory behaviour of two chameleon species (Chamaeleocalyptratus, C. dilepis). Accommodation measurements support earlier observations that accommodation is the major distance cue. Specifically, accommodation speed (60 D s⁻¹), amplitude (45 D) and precision (no significant under-accommodation) were superior to those of other terrestrial vertebrates. Similar to other vertebrates, accommodation was accompanied by a prominent pupillary constriction (pupillary near response). Accommodation could be coupled or uncoupled in both eyes, depending on the experimental situation or the phase of the predatory behavioural sequence. Uncoupled accommodation occurred: 1. During scanning saccadic eye movements for prey detection. Only one eye accommodated appropriately, the other adopted a hyperopic resting refractive state. Attention switched from one eye to the other at approximately 1-s intervals. 2. During initial stages of distance estimation. Coupled accommodation only occurred shortly before the tongue shot. Coupling was demonstrated by either covering one eye with a lens or covering one eye with an infrared light transmitting cut-off filter which still permitted refraction to be measured. In both cases the amount of accommodation was identical in both eyes. Search-coil measurements showed that the angle of convergence of both eyes is too variable to permit triangulation or to provide the basic requirement for stereopsis (matching corresponding points). We conclude that coupling of accommodation serves to improve accommodation precision rather than to permit stereopsis. Accepted: 3 September 1997     

Binocular vision: the eyes add and subtract

Our two eyes' views of the outside world are slightly different, providing the basis for stereopsis. A new study has found evidence that the human visual system has separately adaptable channels for adding and subtracting the neural signals from the two eyes, supporting an unconventional view of the initial stages of stereopsis.     

Properties of 3D rotations and their relation to eye movement control

Rotations of the eye are generated by the torques that the eye muscles apply to the eye. The relationship between eye orientation and the direction of the torques generated by the extraocular muscles is therefore central to any understanding of the control of three-dimensional eye movements of any type. We review the geometrical properties that dictate the relationship between muscle pulling direction and 3D eye orientation. We then show how this relation can be used to test the validity of oculomotor control hypotheses. We test the common modeling assumption that the extraocular muscle pairs can be treated as single bidirectional muscles. Finally, we investigate the consequences of assuming fixed muscle pulley locations when modeling the control of eye movements.     

Relationship between melanin content and superoxide dismutase (SOD) activity in the liver of various species of animals

The scavenger effect of melanin and of superoxide dismutase (SOD) activity on superoxide anion has been shown. In this work we show the relationship between melanin content and SOD activity in livers containing different quantities of melanin which were taken from various species of animals. The mitochondrial SOD activity disappears when the melanin content in the liver is very high; moreover it increases, in the liver of various species of animals examined, proportionally to the decrease of melanin content. No significant variation of the SOD activity localized in the soluble fraction has been detected when related to the melanin content. We think that in the pigmented liver the antioxidant activity of the melanin could mimic part of the function of SOD. The loss of Mn SOD activity could be mediated by a low intracellular level of superoxide anion due to the scavenger effect of melanin on superoxide anion; in fact, it is well known that the biosynthesis of Mn SOD is induced by intracellular levels of superoxide anion.     

Pheromone-modulated optomotor response in male gypsy moths,Lymantria dispar L.: Directionally selective visual interneurons in the ventral nerve cord

In response female pheromone the male gypsy moth flies a zigzagging path upwind to locate the source of odor. He determines wind direction visually. To learn more about the mechanism underlying this behavior, we studied descending interneurons with dye-filled micro-electrodes. We studied the interneuronal responses to combinations of pheromone and visual stimuli.

The PM1 neurons,movement sensitive centrifugal visual brain neurons in the locust: anatomy,physiology, and modulation by identified octopaminergic neurons

The locust’s optic lobe contains a system of wide-field, multimodal, centrifugal neurons. Two of these cells, the protocerebrum-medulla-neurons PM4a and b, are octopaminergic. This paper describes a second pair of large centrifugal neurons (the protocerebrum-medulla-neurons PM1a and PM1b) from the brain of Locusta migratoria based on intracellular cobalt fills, electrophysiology, and immunocytochemistry. They originate and arborise in the central brain and send processes into the medulla of the optic lobe. Double intracellular recording from the same cell suggests input in the central brain and output in the optic lobe. The neurons show immunoreactivity to gamma-amino-butyric acid and its synthesising enzyme, glutamate decarboxylase. The PM1 cells are movement sensitive and show habituation to repeated visual stimulation. Bath application of octopamine causes the response to dishabituate. A very similar effect is produced by electrical stimulation of one of an octopaminergic PM4 neuron. This effect can be blocked by application of the octopamine antagonists, mianserin and phentolamine. This readily accessible system of four wide-field neurons provides a system suitable for the investigation of octopaminergic effects on the visual system at the cellular level.     

Ocular morphology of the Baikal sculpin-oilfishes,Comephorus baicalensis and C. dybowskii (Comephoridae)

Synopsis Baikal sculpin-oilfishes (Comephorus baicalensis and C. dybowskii) were captured from Lake Baikal by mid-water trawl. External morphology was assessed using camera lucida drawings of whole heads. Retinal morphology was examined in material prepared for light and electron microscopy. A single rod-like photoreceptor is present in the retina, and this is at low density compared with marine species of similar depth distribution. C. baicalensis displays tubular eyes, with a dorsally displaced spherical lens and pupil, creating a dorsally directed visual axis. This corresponds to an area with a high density of photoreceptors in the ventral retina. Retinal structure is very similar in C. dybowskii with the exception that regional variation of cell densities within the retina is less marked than in C. baicalensis. In contrast to C. baicalensis, C. dybowskii has a caudal aphakic space and highest photoreceptor density in the rostral retina. Both species display changes in body attitude in association with diurnal vertical migration. We suggest that this may facilitate inspection of the water column from which predator strikes are most likely to occur. Comephorus in general display less specialization for vision in low light than marine counterparts with similar depth distributions.     

Spectral sensitivity of the photonegative reaction of the blood-sucking bug Triatoma infestans (Heteroptera: Reduviidae)

We studied the spectral sensitivity of the visual system of the blood-sucking bug Triatoma infestans, one of the main vectors of Chagas Disease in South America. We quantified the photonegative reaction of this insect in a rectangular arena, half of which was kept dark and the other half illuminated with various intensities of different monochromatic lights (or broadband stimuli for λ>665 nm). As a behavioral parameter of the photonegative response, we measured the time each insect spent in the dark half of the arena. We found that low intensity levels (under 0.06 μW/cm²) of monochromatic lights of 397, 458, 499, and 555 nm evoked a statistically significant (i.e., different from that of control groups) photonegative reaction. Insects were less sensitive to monochromatic lights of 357 nm (UV) and 621 nm (dark orange), and to broadband stimuli in the red part of the spectrum (665–695 nm). These findings indicate that the visual system of T. infestans is sensitive to broader regions of the spectrum than those previously reported.     

Acute effects of UVR on human eyes and skin

Solar UVR ( approximately 295-400 nm) has acute clinical effects on the eyes and the skin. The only effect on the eye is inflammation of the cornea (photokeratitis), which is caused by UVB (and non-solar UVC) and resolves without long-term consequences within 48 h. The effects on the skin are more extensive and include sunburn (inflammation), tanning and immunosuppression for which UVB is mainly responsible. Tanning is modestly photoprotective against further acute UVR damage. Skin colour is also transiently changed by UVA-dependent immediate pigment darkening, the function of which is unknown. Skin type determines sensitivity to the acute and chronic effects of UVR on the skin. Some of the photochemical events that initiate acute effects are also related to skin cancer. Solar UVB is also responsible for the synthesis of vitamin D.     

Prey size selection and distance estimation in foraging adult dragonflies

To determine whether perching dragonflies visually assess the distance to potential prey items, we presented artificial prey, glass beads suspended from fine wires, to perching dragonflies in the field. We videotaped the responses of freely foraging dragonflies (Libellula luctuosa and Sympetrum vicinum—Odonata, suborder Anisoptera) to beads ranging from 0.5 mm to 8 mm in diameter, recording whether or not the dragonflies took off after the beads, and if so, at what distance. Our results indicated that dragonflies were highly selective for bead size. Furthermore, the smaller Sympetrum preferred beads of smaller size and the larger Libellula preferred larger beads. Each species rejected beads as large or larger than their heads, even when the beads subtended the same visual angles as the smaller, attractive beads. Since bead size cannot be determined without reference to distance, we conclude that dragonflies are able to estimate the distance to potential prey items. The range over which they estimate distance is about 1 m for the larger Libellula and 70 cm for the smaller Sympetrum. The mechanism of distance estimation is unknown, but it probably includes both stereopsis and the motion parallax produced by head movements.