Specialized ommatidia for polarization vision in the compound eye of cockchafers,Melolontha melolontha (Coleoptera,Scarabaeidae)

Summary The superposition eye of the cockchafer, Melolontha melolontha, exhibits the typical features of many nocturnal and crepuscular scarabaeid beetles: the dioptric apparatus of each ommatidium consists of a thick corneal lens with a strong inner convexity attached to a crystalline cone, that is surrounded by two primary and 9–11 secondary pigment cells. The clear zone contains the unpigmented extensions of the secondary pigment cells, which surround the cell bodies of seven retinula (receptor) cells per ommatidium and a retinular tract formed by them. The seven-lobed fused rhabdoms are composed by the rhabdomeres of the receptor cells 1–7. The rhabdoms are optically separated from each other by a tracheal sheath around the retinulae. The orientation of the microvilli diverges in a fan-like fashion within each rhabdomere. The proximally situated retinula cell 8 does not form a rhabdomere. This standard form of ommatidium stands in contrast to another type of ommatidium found in the dorsal rim area of the eye. The dorsal rim ommatidia are characterized by the following anatomical specializations: (1) The corneal lenses are not clear but contain light-scattering, bubble-like inclusions. (2) The rhabdom length is increased approximately by a factor of two. (3) The rhabdoms have unlobed shapes. (4) Within each rhabdomere the microvilli are parallel to each other. The microvilli of receptor 1 are oriented 90° to those of receptors 2–7. (5) The tracheal sheaths around the retinulae are missing. These findings indicate that the photoreceptors of the dorsal rim area are strongly polarization sensitive and have large visual fields. In the dorsal rim ommatidia of other insects, functionally similar anatomical specializations have been found. In these species, the dorsal rim area of the eye was demonstrated to be the eye region that is responsible for the detection of polarized light. We suggest that the dorsal rim area of the cockchafer eye subserves the same function and that the beetles use the polarization pattern of the sky for orientation during their migrations.     

Structural changes in light- and dark-adapted compound eyes of the australian earwig Labidura riparia truncata (dermaptera)

The apposition acone eye of Labidura is relatively small—550–600 facets—with a thick corneal lens and shallow retina. The retinula cell columns are each formed of six peripheral cells plus two central cells, a partially fused rhabdom, and dense pigment in two or three cell types. Upon adaptation from light to dark, the most striking photomechanical response is a proximal broadening of the cone cells, which results in a 38-fold increase in cross-sectional area of the aperture. While longitudinal rhabdom movement is small, microvillar diameters swell in response to light and contract in the dark. Irregularities of facet pattern and shape, and in ommatidial alignment were found, particularly towards eye margins. Three types of interommatidial sense organs on the eye surface are described, one of which has not been previously reported. An argument is presented to explain how the field of view and sensitivity are both apparently decreased in the acone eye by exposure to light.     

Do we need harvest regulations for European crayfish?

Summary Management of noble crayfish fisheries varies considerably between countries. Minimum legal harvest sizes range from 70 to 120 mm TL. National, regional and local crayfish harvesting regulations exist. The variations in regulations are strongly influenced by traditions for crayfish harvest and consumption. The current harvest of noble crayfish in Europe is approximately 220 tonnes; this is less than 10% of the pre-plague historic record. Improved fishery regulations may potentially increase the annual harvest. Because of allometric growth, noble crayfish increase by some 40–50% in weight by increasing in total length from 90 to 100 mm. Stock structure and stock characteristics vary between localities. Female noble crayfish mature at a size of 62–85 mm TL and the number of attached eggs is low, i.e. less than 200. Mean size of females in trap catches may be lower than mean size at maturity, indicating vulnerability to recruitment overfishing. There is a market for crayfish smaller than 90 mm TL, both for consumption and for stocking, and these often constitute more than 75% of trap catches. Regulations for catching crayfish are thus needed in order to prevent recruitment overfishing and to sustain high and stable yields. The authors therefore recommend a national minimum harvest size of 90–95 mm TL. The catching season should start after the first moult and after the brood have left their mother, and terminate when the mating period approaches in September. In localities with high exploitation a short harvest season (2–3 weeks) is recommended in the early August intermoult period. However, crayfish harvest regulations should also account for the variation that occurs between local stocks, and if necessary exemptions from the national regulations should be given where appropriate.     

The dorsal eye of the dragonfly Sympetrum: specializations for prey detection against the blue sky

Dragonflies of the genus Sympetrum have compound eyes conspicuously divided into dorsal and ventral regions. Using anatomical, optical, electrophysiological, in-vivo photochemical and microspectrophotometrical methods, we have investigated the design and physiology of the dorsal part which is characterized by a pale yellow-orange screening pigment and extremely large facets. The upper part of the yellow dorsal region is a pronounced fovea with interommatidial angles approaching 0.3°, contrasting to the much larger values of 1.5°–2° in the rest of the eye. The dorsal eye part is exclusively sensitive to short wavelengths (below 520 nm). It contains predominantly blue-receptors with a sensitivity maximum at 420 nm, and a smaller amount of UV-receptors. The metarhodopsin of the blue-receptors absorbs maximally at 535 nm. The yellow screening pigment transmits longwavelength light (cut-on 580 nm), which increases the conversion rate from metarhodopsin to rhodopsin (see Fig. 11a). We demonstrate that because of the yellow pigment screen nearly all of the photopigment is in the rhodopsin state under natural conditions, thus maximizing sensitivity. Theoretical considerations show that the extremely long rhabdoms (1.1 mm) in the dorsal fovea are motivated for absorption reasons alone. A surprising consequence of the long rhabdoms is that the sensitivity gain, caused by pumping photopigment into the rhodopsin state, is small. To explain this puzzling fact we present arguments for a mechanism producing a gradient of rhodopsin concentration along the rhabdom, which would minimize saturation of transduction units, and hence improve the signal-to-noise ratio at high intensities. The latter is of special importance for the short integration time and high contrast sensitivity these animals need for spotting small prey at long distances.Abbreviations ERG electroretinogram - R rhodopsin - M metarhodopsin     

Functional morphology of the divided compound eye of the honeybee drone (Apis mellifera)

Using different approaches, the functional morphology of the compound eye of the honeybee drone was examined. The drone exhibits an extended acute zone in the dorsal part of its eye. The following specializations were found here: enlarged facet diameters; smaller interommatidial angles; red-leaky screening pigment; enlarged rhabdom diameters; photopigment composition different from the drone's ventral eye region and the worker bee's eye. Thus, similar to other male insects, the drone compound eye is divided into a male-specific dorsal part and a ventral part resembling the worker bee's eye. The functional significance of the sex-specific acute zone is discussed with respect to mating behaviour.     

The ultrastructure of the uterine epithelium of the pig during the estrous cycle and early pregnancy

Summary In the compound eye of the moth Antheraea polyphemus, three types of visual pigments were found in extracts from the retina and by microspectrophotometry in situ. The absorption maxima of the receptor pigment P and the metarhodopsin M are at (1) P 520–530 nm, M 480–490 nm; (2) P 460–480 nm, M 530–540 nm; (3) P 330–340 nm, M 460–470 nm. Their localization was investigated by electron microscopy on eyes illuminated with different monochromatic lights. Within the tiered rhabdom, constituted of the rhabdomeres of nine visual cells, the basal cell contains a blue-and the six medial cells have a greenabsorbing pigment. The two distal cells of most ommatidia also have the blue pigment; only in the dorsal region of the eye, these cells contain a UV-absorbing pigment, which constitutes a portion of only 5% of the visual pigment content within the entire retina. The functional significance of this distribution is discussed.     

Post emergence maturation of the eye of the adult black carpet beetle,Attagenus megatoma (Fab.): An electron microscope study

The black carpet beetle, Attagenus megatoma (Fab.), has been reported to exhibit negative phototaxis immediately after emergence. In later adult life, after the period during which most oviposition has occurred, the beetles are found to be photopositive. The compound eyes of one-day-old (Post-emergence) and nine-day-old (Post-ovipositional) female beetles were studied by electron microscopy and a number of strong differences were found between eyes at the two ages. The corneal facets of one-day eyes had the form of convex-concave lenses, while those of nine-day beetles were double-convex lenses. The primary and secondary pigment cells of young eyes were large and contained much endoplasmic reticulum and little accessory pigment. In the older eyes the pigment cells were reduced and contained much pigment, the proportion of endoplasmic reticulum being greatly reduced by comparison with the one-day eye. The cross-sectional area of the rhabdom was greater in the older eye. The possible relationships between age-related changes in eye morphology and behavioral changes during the same period are discussed.     

The structures of dorsal and ventral regions of a dragonfly retina

Summary The apposition eyes of the corduliid dragonfly Hemicordulia tau are each divided by pigment colour, facet size and facet arrangement into three regions: dorsal, ventral, and a posterior larval strip. Each ommatidium has two primary pigment cells, twenty-five secondary pigment cells, and eight receptor cells, all surrounded by tracheae which probably prevent light passing between ommatidia, and reduce the weight of the eye. Electron microscopy reveals that the receptor cells are of two types: small vestigial cells making virtually no contribution to the rhabdom, and full-size typical cells. The ventral ommatidia have a distal typical cell (oriented either horizontally or vertically), four medial typical cells, two proximal typical cells and one full-length vestigial cell. The dorsal ommatidia have only four full-length typical cells, and one distal and three vestigial full-length cells. The cross-section of dorsal rhabdoms is small and circular distally, but expands to a large three-pointed star medially and proximally. The tiered receptor arrangement in the ventral ommatidia is typical of other Odonata but the dorsal structure has not been fully described in other species. Specialised dorsal eye regions are typical of insects that detect others against the sky.     

THE EFFECT OF TEMPERATURE UPON FACET NUMBER IN THE BAR-EYED MUTANT OF DROSOPHILA : PART III.

Three strains of the bar-eyed mutant of Drosophila melanogaster Meig have been reared at constant temperatures over a range of 15–31°C. The mean facet number in the bar-eyed mutant varies inversely with the temperature at which the larvæ develop. The temperature coefficient (Q₁₀) is of the same order as that for chemical reactions. The facet-temperature relations may be plotted as an exponential curve for temperatures from 15–31°. The rate of development of the immature stages gives a straight line temperature curve between 15 and 29°. Beyond 29° the rate decreases again with a further rise in temperature. The facet curve may be readily superimposed on the development curve between 15 and 27°. The straight line feature of the development curve is probably due to the flattening out of an exponential curve by secondary factors. Since both the straight line and the exponential curve appear simultaneously in the same living material, it is impractical to locate the secondary factors in enzyme destruction, differences in viscosity, or in the physical state of colloids. Differential temperature coefficients for the various separate processes involved in development furnish the best basis for an explanation of the straight line feature of the curve representing the effect of temperature on the rate of physiological processes. Facet number in the full-eyed wild stock is not affected by temperature to a marked degree. The mean facet number for fifteen full-eyed females raised at 27° is 859.06. The mean facet number for the Low Selected Bar females at 27° is 55.13; for the Ultra-bar females at 27° it is 21.27. A consistent sexual difference appears in all the bar stocks, the females having fewer facets. This relation may be expressed by the sex coefficient, the average value of which is 0.791. The average observed difference in mean facet number for a difference of 1°C. in the environment in which the flies developed is 3.09 for the Ultra-bar stock and 14.01 for the Low Selected stock. The average proportional differences in the mean for a difference of 1°C. are 9.22 per cent for Ultra-bar, and 14.51 for Low Selected. The differences in the number of facets per °C. are greatest at the low and least at the high temperatures. The difference in the number of facets per °C. varies with the mean. The proportional differences in the mean per °C. are greatest at the lower (15–17.5°) and higher (29–31°) temperatures and least at the intermediate temperatures. Temperature is a factor in determining facet number only during a relatively short period in larval development. This effective period, at 27°, comes between the end of the 3rd and the end of the 4th day. At 15°, this period is initiated at the end of 8 days following a 1st day at 27°. At 27° this period is approximately 18 hours long. At 15° it is approximately 72 hours long. The number of facets and the length of the immature stage (egg-larval-pupal) appear related when the whole of development is passed at one temperature. That the number of facets is not dependent upon the length of the immature stage is shown by experiments in which only a part of development was passed at one temperature and the remainder at another. Temperature affects the reaction determining the number of facets in approximately the same way that it affects the other developmental reactions, hence the apparent correlation between facet number and the length of the immature stage. Variability as expressed by the coefficient of variability has a tendency to increase with temperature. Standard deviation, on the other hand, appears to decrease with rise in temperature. Neither inheritance nor induction effects are exhibited by this material. This study shows that environment may markedly affect the somatic expression of one Mendelian factor (bar eye), while it has no visible influence on another (white eye).     

The Nymphal Eyes of Parabuthus transvaalicus Purcell, 1899 (Buthidae): An Accessory Lateral Eye in a Scorpion

In P. transvaalicus nymphs, 5 pairs of lateral ocelli each composed of a corneal lens, R-cell units forming a latticed rhabdom, arhabdomeric cells and pigment cells are present. In addition, we found a pair of unpigmented accessory sense organs situated ventroposteriorly to the lateral ocelli in prenymphs as well as in first nymphs. They are composed of primary, rhabdomeric sensory cells, and we infer that they represent a second type of lateral eye. They also comprise sensory units, but lenses and screening pigment are lacking. Their position and cellular architecture corresponds well with that of the “rudimentary” lateral eye of the xiphosuran, Limulus. The occurrence of a bipartite lateral visual system in Chelicerata and Arthropoda is discussed.     

Localization of myosin and actin in ocular nonmuscle cells

Summary Myosin and actin were localized by indirect immunofluorescence microscopy using specific antibodies prepared in rabbits against highly purified gizzard myosin and actin. A strong fluorescence staining with both antibodies was observed in rat corneal epithelial cells, anterior lens epithelial cells, rod inner segments, and in rat and frog pigment epithelial cells. The immunohistochemical localization of myosin in corneal epithelial cells was further supported by the electrophoretic and immunological identification of smooth muscle type myosin heavy chain in pure corneal epithelial abrasions. Electron-microscopic observations revealed a clear correlation between staining with actin antibodies and the presence of numerous thin cytoplasmic filaments (50–80 Å in diameter). The functional and biochemical nature of 90–110 Å filaments occurring in corneal and lens epithelial cells, as well as the ultrastructural localization of myosin in ocular nonmuscle cells under study remains obscure.     

Geometrical optics of a galatheid compound eye

The eyes of galatheid squat lobsters (Munida rugosa) are shown to be of the reflecting superposition type. In the dark-adapted state corneal lenses focus light at the level of the rhabdoms and light from more than 1000 facets is redirected to the superposition focus by the reflecting surfaces of the crystalline cones. When the eye is light adapted, apposition optics are used. In this state paraxial light is focused by the corneal lens and the parabolic proximal end of the cone onto the distal end of a rhabdomeric lightguide. The latter transmits light across the clear zone to the rhabdom layer. In the dorsal part of the eye the individual ommatidia become progressively shorter until the cones and rhabdoms are no longer separated by a clear zone. Although formerly considered to be developing ommatidia, they are shown to be retained specifically for scanning the downwelling irradiance.Abbreviations RI refractive index - SEM scanning electron microscope     

The morphology of the eyes of Limulus

Summary The dioptric apparatus of the Limulus compound eye is composed of the corneal cuticle with its internally projecting cuticular cones and the specialized underlying epidermis. The latter is composed of three distinct cell types. The guanophores, located between cuticular cones, contain guanine as a reflecting pigment. The distal pigment cells, which clothe the sides of the cuticular cones and form a sheath around the underlying ommatidium, contain massive bundles of microtubules, abundant pigment droplets and a large Golgi system. The cone cells are positioned between the flattened tip of the cuticular cone and the apex of the ommatidium. They serve to anchor the retinula cells to the cuticle and, by virtue of long processes along the periphery of the rhabdome, perform a glial function with respect to the interaction of adjacent retinula cells. The geometry and fine structure of the dioptric apparatus provide supporting evidence for the wide angle of acceptance and lack of polarized light perception by the ommatidia.This study constitutes publication No. 288 from the Oregon Regional Primate Research Center, supported in part by Grants FR 00163 and NB 07717-01 from the National Institutes of Health and in part by a Bob Hope Fight For Sight Grant-in-Aid of the National Council to Combat Blindness, Inc. The author wishes to thank Mrs. Audrey Griffin for patient and excellent technical assistance.     

Colour in the eyes of insects

Many insect species have darkly coloured eyes, but distinct colours or patterns are frequently featured. A number of exemplary cases of flies and butterflies are discussed to illustrate our present knowledge of the physical basis of eye colours, their functional background, and the implications for insect colour vision. The screening pigments in the pigment cells commonly determine the eye colour. The red screening pigments of fly eyes and the dorsal eye regions of dragonflies allow stray light to photochemically restore photoconverted visual pigments. A similar role is played by yellow pigment granules inside the photoreceptor cells which function as a light-controlling pupil. Most insect eyes contain black screening pigments which prevent stray light to produce background noise in the photoreceptors. The eyes of tabanid flies are marked by strong metallic colours, due to multilayers in the corneal facet lenses. The corneal multilayers in the gold-green eyes of the deer fly Chrysops relictus reduce the lens transmission in the orange-green, thus narrowing the sensitivity spectrum of photoreceptors having a green absorbing rhodopsin. The tapetum in the eyes of butterflies probably enhances the spectral sensitivity of proximal long-wavelength photoreceptors. Pigment granules lining the rhabdom fine-tune the sensitivity spectra.     

Polarization and spectral sensitivity of single photoreceptors of the domestic cricket

The polarization and spectral sensitivity of single photoreceptors ofAcheta domesticus L. was measured. The morphological characteristics of the cricket rhabdome satisfy the conditions for a symmetrical model, for which the polarization sensitivity of a single photoreceptor is identically equal to the dichroism of a single microvillus. Characteristic curves of spectral sensitivity of all photoreceptors measured (24 cells) were similar and had two maxima: the principal at 500 nm and a secondary peak at 360 nm, characteristic of a pigment such as rhodopsin in the rods of the vertebrate retina. The mean value of polarization sensitivity measured was 2.28 ± 0.85 (mean ± standard deviation, 70 cells), suggesting the existence of slight preferential orientation of the dipole moments of the rhodopsin molecules along the axes of the microvilli.I. N. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 5, pp. 483–490, September–October, 1979.     

Neither apposition nor superposition: the compound eyes of the Chestnut Leafminer Cameraria ohridella

The two lemon-shaped compound eyes of the moth Cameraria ohridella measure in dorsal–ventral direction 263.0 μm in male and 238.9 μm in female individuals. In anterior–posterior direction no significant differences were found between the sexes, eye length being about 194.6 μm. The eyes of males consist of ca. 417 hexagonal facets, while those of females contain 367. In both sexes facet diameters are approximately 11.5 μm. Despite the size differences of the eyes in the two sexes, ultrastructurally they are identical and both possess ommatidia of 80 μm mean length. The ultrastructure of the eye is described and compared with that of other compound eyes of Lepidoptera. Anatomically the eyes represent a type intermediate between apposition and refractive superposition kind. A distal rhabdom is present in the space that in the eyes of larger moths with superposition optics is occupied by the so-called clear zone. A tracheal tapetum and longitudinal screening pigment migrations, typical of superposition but not apposition eyes are present despite the lack of a clear zone. Thus, our results support an earlier calculated minimal theoretical limit for superposition eyes.     

The physical and morphological properties of the pigment screen in the compound eye of a shrimp (Crustacea)

Summary The pigment cells of the compound eye of the shrimps (Crangon crangon andC. allmani) were studied by electron microscopy (SEM and TEM) and microspectrophotometry. The compound eyes of these species contain light-absorbing and -reflecting pigments contained in granules, located in 5 different cells. The light absorbing pigment granules (light screen) are situated in (1) the distal pigment cells, (2) the retinular cells, (3) the basal pigment cells. The reflecting pigment granules are located in (4) the distal, and (5) the proximal reflecting pigment cells. Another innominate cell type investing the ommatidia contains vacuoles without pigment content. The innominate cell type, and the basal absorbing pigment cell (3) listed above, have not earlier been reported for a crustacean species. Measurements of the spectral absorption on sliced and squashed ommatidia show that all components of the light screen have an increased absorption in the wavelength regions 400–450 nm and 530–570 nm, probably due to xanthommatin and ommin. The spectral absorbancy of the reflecting pigment cells were not determined. Similar cells in other species are known to contain pteridines.We thank Prof. Dr. Langer, Bochum, Germany, for his kind help. The work was supported by funds from the Karolinska Institutet to Doc. G. Struwe, and grant NFR No. 2760-007 to Doc. R. Elofsson.     

Size and structure of crayfish (Astacus astacus) populations on different habitats in Finland

The many inland waters in Finland make crayfish production an important potential resource. The rational utilization and management of this resource requires knowledge of the size and structure of the crayfish populations. The difficulties often encountered in catching crayfish complicate population studies. Mark-recapture and electric fishing have been used in the studies. The number of adult crayfish measuring more than 70 mm in a 4-ha lake was estimated at 620, and the number in a 13-ha lake at 3 480. In the lakes, the density of adult crayfish was around 0.6–1.4 m^–2 and in one stream studied about 2.5 m^–2 rising to several individuals per m² in the best biotopes.     

Variation in compound eye structure: effects of diet and family

Studies of compound eyes have revealed that variation in eye structure can substantially affect visual performance. Here, we investigate the degree to which a stressful rearing environment, which decreases body size, affects the eye phenotype. Full siblings of the Orange Sulphur butterfly, Colias eurytheme, were collected from known parents and split within families among two diet treatments that varied in quality. In both sexes, individuals reared on the high-quality diet had larger eye height and anterior facet diameter, and therefore, by inference, superior vision. However, relative to their reduced body size, individuals reared on low-quality diet had proportionally larger eyes and facets than individuals reared on high-quality diet. We interpret this finding as evidence that butterflies encountering nutritional stress increased proportional investment in eye development to reduce loss of visual performance. We also found significant broad-sense genetic variation underlying eye structure in both males and females, and report novel heritability estimates for eye height and facet diameter. Surprisingly, there was greater genetic variation in eye height among males than among females, despite apparently stronger directional selection on male vision. We discuss the implications of these data for our understanding of eye development and evolution.     

Sexual Dimorphism in the Compound Eye of Rhagophthalmus ohbai (Coleoptera: Rhagophthalmidae): I. Morphology and Ultrastructure

The eyes of the winged males and larvi-form, wingless females of the firefly Rhagophthalmus ohbai differ from each other in several respects. Compared with the eyes of the males, those of the females contain fewer (35 versus ca. 3500) and smaller (20 μm versus 24-31 μm) facets and anatomically they are of the apposition type. Their main function appears to be to detect light intensity changes from day to nighttime; resolving power of the female eye must be poor and e-vector discrimination would be absent. The eyes of the males consist of a smaller, dorsal region of ca. 500 om-matidia of about 250 μm length and a larger, ventral region of ca. 2000 ommatidia of about 640 urn length. The microvilli of the dorsal eye region are somewhat wider than those of the ventral region (55 nm versus 45 nm) and are less regularly arranged. A tapetal reflecting layer is only present in the dorsal eye region. The small clear-zone between dioptric apparatus and retina in the dorsal eye region would not allow as good a superposition image to be produced as in the ventral eye region with its 5 times wider clear-zone. The regular orientations of the microvilli in the rhabdoms and the lack of a proper tapetum in the ventral eye region suggest that e-vector discrimination should be possible.