Superposition optics and the time of flight in onitine dung beetles

Dung beetles fly to fresh dung, with vision essential for flight navigation. The daily period of flight varies among different species: some beetles fly only in sunlight, others only when ambient light levels change rapidly during dusk or dawn and others in the constant dark of night. Measurements of the optical properties of the lenses, eye geometry and photoreceptor dimensions were used in a computer ray-tracing model to determine the optical performance of the superposition eyes of nine species of onitine dung beetles. Eye sensitivity to light is determined mainly by body size, by the refractive-index parameters and size of the crystalline cones, and by the photoreceptor dimensions. Based on the optics of the ommatidial lenses and absorption of light in the retina, the most sensitive eyes, found in the crepuscular-nocturnal beetles, are 85 times or nearly two log units more sensitive than the eyes of the diurnal beetles. Three possible criteria are considered to determine the best position for the retina: maximum amount of light absorbed in the target rhabdom; maximum amount of light falling on the target rhabdom (best focus); and maximum resolution. The structure and physiological optics of the superposition compound eyes of an onitine dung beetle are matched to the range of light intensities at which it flies. Accepted: 4 February 1998     

Screening pigment,aperture and sensitivity in the dung beetle superposition eye

Summary Sensitivity to light was investigated in the refracting superposition eye of the dung beetle Onitis alexis using electrophysiological measurements and optical modelling. Intracellular recordings were made from single retinula cells over 24-h periods, with cells light and dark adapted, in order to measure the response/intensity (V-LogI) functions. The combined effects of a circadian rhythm and light adaptation allow the determination of the relative contributions of screening-pigment migration and transduction gain to changes in sensitivity in the eye. Between the extremes of dark adaptation at night and light adaptation during the day, the maximum sensitivity change possible is at least 4 log units, of which approximately 2 log units can be accounted for by changes in the transduction gain and at least 2 log units by screening-pigment migration. The role of the superposition aperture (the number of facets that contribute light to one rhabdom) in 3 species of dung beetle was investigated with an optical ray-tracing model of the eye. The facets of the superposition aperture do not contribute light equally to the target rhabdom; except in one species, the greatest contribution comes from facets located away from both the centre and periphery of the aperture. Light adaptation increases the optical density of the superposition aperture and decreases its size.     

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 visual ecology of pupillary action in superposition eyes

The two most common mechanisms of pupillary screening-pigment migration in arthropod superposition eyes are the cone and longitudinal pigment migration mechanisms. The dynamics of each were investigated by optical modelling and by determining experimentally the relationship between eye glow brightness and screening pigment position within the eyes of two representative insect species: the noctuid moth Agrotis infusa and the dung beetle Copris elphenor. During dark adaptation, in both mechanisms, the screening pigment is contracted distally to expose the proximal half of each crystalline cone. During light adaptation the pigment migrates proximally and reduces light flux in the retina. In the longitudinal mechanism, pigment migrates into the clear zone of the eye. In the cone mechanism, pigment never enters the clear zone and is instead restricted to the proximal half of each crystalline cone: a migrating sleeve of pigment creates a small aperture at the end of the crystalline cone, the area of which depends on the degree of light adaptation. According to the model, the cone mechanism provides a limited range of light attenuation (ca. 0.6 log units) for which both good spatial resolution and accuracy of control are maintained, and within this range attenuation is controlled very finely. Beyond this range, whilst attenuation is still possible, diffraction at the pigment aperture and increasing coarseness of control worsen visual performance significantly. In contrast, the longitudinal mechanism provides a much larger useful range of light attenuation (up to several log units) and maintains reasonable fineness of attenuation control over the entire range (although not as fine as the cone mechanism). The experimental results support the model. An extensive survey of arthropods with superposition eyes reveals that the cone mechanism is almost exclusively possessed by those animals experiencing a narrow range of light intensities, and the longitudinal mechanism by those experiencing a wide range.Dedicated to Professor Rolf Elofsson on the occasion of his retirement from the Chair of Zoology in Lund     

Some optical features of the eyes of stomatopods

The optics of a variety of stomatopod eyes has been investigated using goniometric eye-mapping techniques and anatomical measurements. The species examined come from 3 of the 4 existing superfamilies: the Gonodactyloidea, Lysiosquilloidea and Squilloidea. This paper examines acuity, optical axes and general features of eye shape. Stomatopod eyes are divided into 3 clearly distinct zones; the mid-band and two hemispheres. Each hemisphere consists of an edge region, a visual streak and a near mid-band region. The optical axes of many ommatidia from both hemispheres are skewed inwards towards the centrally placed mid-band and are rarely normal to the corneal surface. The large skew angle enables each hemisphere to examine an area which extensively overlaps that of the other hemisphere. As a result monocular distance judgement is possible. Most of the ommatidia in each hemisphere are part of a horizontally aligned but vertically acute visual streak area. There is one visual streak per hemisphere and both look into the same 5–10° strip. This narrow strip is also the area in space the mid-band ommatidia examine. An acute zone is present in the eyes of lysiosquilloid and gonodactyloid stomatopods and includes ommatidia, from both the hemispheres and the mid-band. Here inter-ommatidial angles, especially those in the horizontal direction, are reduced. Acute zone facets are enlarged to increase sensitivity rather than aid spatial resolution.Abbreviations and definitions AZ Acute Zone - MB Mid-band - D Corneal facet diameter, as MB facets are asymmetrical, values for width and height of each facet are given - DH dorsal hemisphere - f Focal length of each ommatidium, estimated from the centre of the corneal lens to the tip of the rhabdom - NeMB Near mid-band ommatidia - R Resolving power=1/2_average - R_h Horizontal resolving power=1/2_h - R_v Vertical resolving power=1/2_v - VH ventral hemisphere - Geometrical acceptance angle (a/f×57.3) of each ommatidium - _h Horizontal inter-ommatidial angle, between facets along a row - _v Vertical inter-ommatidial angle, between rows; Superfamily Gonodactyloidea - g.c. Gonodactylus chiragra - O.s. Odontodactylus scyllarus - H.e. Hemisquilla ensigera; Superfamily Lysiosquilloidea - L.t. Lysiosquilla tredecimdentata - C.s. Coronis scolopendra; Superfamily Squilloidea - O.o. Oratosquilla solicitans     

Strategies for retinal design in arthropod eyes of low F-number

Summary The superposition eyes and simple eyes of many arthropods have apertures (A) with a diameter bigger than, or about the same size as, the focal length of the eye (f). That is, these eyes have low F-numbers (f/A). Many of the light rays focussed onto a photoreceptor will not be trapped by total internal reflection in the photoreceptor and will therefore pass through and be absorbed in photoreceptors other than that for which the light was intended. This spread of light in the retina leads to a broadening of the angular-sensitivity function and a consequent degrading of the image at the retinal level. A number of solutions to this problem are found in nature, with the most effective that of isolating the photoreceptors with a sheath of either light-absorbing pigment or reflecting tapetum. A ray-tracing model was used to assess the relative merits of the tapetal and pigment sheath designs in low F-number superposition eyes, and also to investigate the effect of changing the refractive index and absorption coefficient of the rhabdom. Which sheathing solution is best depends on the quality of the image on the retina, on the spacing of the rhabdoms in the retina and on the intensity of light normally experienced by the eye. In a retina with closely packed rhabdoms, the model predicts full sheathing to be the optimal solution if the image is well-focussed, partial sheathing if poorly focussed and no sheathing if moderately well-focussed. In a retina with rhabdoms spaced apart and a well-focussed image, the model predicts partial sheathing to be optimal. A pigment sheath is predicted to be useful in eyes which experience bright light and have no need for high sensitivity. A tapetal sheath is predicted to be useful at any intensity. A survey of arthropod eyes with low F-number supports the predictions of the model.     

Ocellar optics in nocturnal and diurnal bees and wasps

Nocturnal bees, wasps and ants have considerably larger ocelli than their diurnal relatives, suggesting an active role in vision at night. In a first step to understanding what this role might be, the morphology and physiological optics of ocelli were investigated in three tropical rainforest species – the nocturnal sweat bee Megalopta genalis, the nocturnal paper wasp Apoica pallens and the diurnal paper wasp Polistes occidentalis – using hanging-drop techniques and standard histological methods. Ocellar image quality, in addition to lens focal length and back focal distance, was determined in all three species. During flight, the ocellar receptive fields of both nocturnal species are centred very dorsally, possibly in order to maximise sensitivity to the narrow dorsal field of light that enters through gaps in the rainforest canopy. Since all ocelli investigated had a slightly oval shape, images were found to be astigmatic: images formed by the major axis of the ocellus were located further from the proximal surface of the lens than images formed by the minor axis. Despite being astigmatic, images formed at either focal plane were reasonably sharp in all ocelli investigated. When compared to the position of the retina below the lens, measurements of back focal distance reveal that the ocelli of Megalopta are highly underfocused and unable to resolve spatial detail. This together with their very large and tightly packed rhabdoms suggests a role in making sensitive measurements of ambient light intensity. In contrast, the ocelli of the two wasps form images near the proximal boundary of the retina, suggesting the potential for modest resolving power. In light of these results, possible roles for ocelli in nocturnal bees and wasps are discussed, including the hypothesis that they might be involved in nocturnal homing and navigation, using two main cues: the spatial pattern of bright patches of daylight visible through the rainforest canopy, and compass information obtained from polarised skylight (from the setting sun or the moon) that penetrates these patches.     

Differences in temporal utilization patterns of dung pats among three scarabaeid dung beetles

1. Differences in the temporal utilization patterns of dung pats were investigated among three dung beetle species, Aphodius haroldianus, Onthophagus lenzii and Liatongus phanaeoides, and density effects of three species on emigration from a dung pat were compared.
2. A. haroldianus preferred fresh dung pats, immigrated to fresh pats and then emigrated from those pats quickly (mean duration of residence 48.4 h), whereas L. phanaeoides immigrated to older dung pats (over 7 days after deposition) as well as fresh ones and remained in the pats for the longest time (165.6 h).O. lenzii showed intermediate behavior in this respect (63.3 h).
3. In A. haroldianus, the presence of conspecifics induced faster emigration than presence of the other two species. L. phanaeoides showed the opposite relationship. In O. lenzii, the response to conspecifics was intermediate to those of the other two species.

     

Ontogeny and optics of the eyes of the common prawn Palaemon (Palaemon) serratus (Pennant, 1777)

A brief review of the work on crustacean compound eyes is given. Two main types of eye have been recognized: apposition and superposition. The ontogeny of the eyes of the common prawn Palaemon serratus is examined using a variety of methods: photography of live specimens, histological sections, SEM and TEM. In common with other decapod larvae, the common prawn hatches with apposition eyes having circular lenses packed hexagonally. After metamorphosis the gradual squaring of the eye facets, begun during the larval phase, is completed. This is an essential prerequisite for the functioning of the facultative superposition reflecting optics found in long-bodied decapods (e.g. shrimps, prawns and lobsters) and some Anomura. The possible phylogenetical significance of superposition, reflecting optics is also discussed.     

The daily onset of flight in the crepuscular dung beetle Onitis alexis

The role of light intensity and temperature in determining the onset of flight in the crepuscular dung beetle Onitis alexis Klug (Coleoptera: Scarabaeidae) was examined. Flight under natural light was highly synchronized, with two-thirds of the beetles that flew flying over a period of 10–12 min. In six flights on different days, the mean time of onset varied by up to 13 min, but mean onset occurred at fixed light intensity. Absolute light intensity therefore appears to be a vital cue in determining flight onset. Mean onset remained at this intensity when dusk was advanced artificially by up to about 8 min. However, when dusk was brought further forward, mean flight occurred at lower intensities and onset of flight took place over a longer period. This is interpreted as an overlapping of the period of light intensities suitable for flight with the circadian rhythm that brings the beetles to the surface. No beetles flew when kept under constant bright light or in the dark during the dusk period. Under conditions of constant dusk, the beetles appeared uncoordinated and, although the median time of onset of flight was only 3 min later than in the control flight, onset of flight was spread out over a much longer period. At soil temperatures of 20–22 °C, over 90% of the beetles flew. This percentage decreased with decreasing temperature, and less than 10% flew at temperatures of 16–17 °C.

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L'incitation quotidienne au vol chez e bousier crépusculaire Onitis alexis Klug (Col. Scarabaeidae)

Résumé L'examen a porté sur le rôle de l'intensité lumineuse et de la température sur l'incitation au vol chez Onitis alexis. Le vol en lumière naturelle est fortement synchronisé, avec les deux tiers des adultes qui s'envolent en 10–12 min. Sur six vols pour différents jours, le moment moyen d'envol variait de 13 min, mais l'envol moyen avait lieu à une intensité lumineuse déterminée. La valeur absolue de l'intensité lumineuse paraît ainsi un signal crucial dans la détermination et l'incitation au vol. Le moment d'envol se maintient à cette intensité quand le crépuscule est artificiellement avancé jusqu'à 8 min environ. Cependant, quand le crépuscule est rendu encore plus précoce, l'envol moyen se produit à des intensités plus basses et est étalé. Ceci peut être interprété comme un chevauchement de la période des intensités lumineuses favorables à l'envol avec le rythme circadien qui conduit les bousiers à la surface. Aucun adulte ne vole quand il y a maintien de lumière constante ou d'obscurité à l'heure du crépuscule. En présence de crépuscule constant, les réponses sont hétérogènes, et, bien que le moment médian d'envol ne soit retardé que de 3 min, par rapport aux témoins, l'incitation à l'envol est étalée sur une période plus longue. Avec des températures au sol de 20–22 °C, plus de 90% des adultes s'envolent. Le pourcentage diminue avec la température, et moins de 10% s'envolent à 16–17 °C.

     

Effect of population density on reproduction of two sympatric dung beetle species,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

The effect of population density on reproduction as well as the oviposition modes of two sympatric dung beetle species, Aphodius haroldianus (a low fecundity/high parental effort species) and A. elegans (a high fecundity/low parental effort species) was studied at a pasture in central Japan from 1982 to 1986.

1. The adult population density of A. haroldianus was high (>40 per dung pat). The density of A. elegans was low (<5 per dung pat). Oviposition of A. haroldianus was suppressed above the density of about 10 adults of conspecifics but not affected by the density of other species. The intra- and interspecific density effects on oviposition were not detected in A. elegans.
2. The number of A. haroldianus adults per pat decreased with dung age, while that of A. elegans increased until sixth day after deposition and then decreased. The number of eggs laid per pat was not different between 1-day-and 3-day-old pats for A. haroldianus. However, more eggs of A. elegans were found in 3-day-old pats than in 1-day-old ones.
3. In the both species, the amplitude of population fluctuations was not remarkable; the maximum/minimum ratio for five years being 2.3 for A. haroldianus and 2.9 for A. elegans. Different density dependent processes were suggested to function for the small fluctuation of population density between the two species, i.e. intraspecific density effect on oviposition for A. haroldianus and contest type competition for food among larvae of A. elegans.

     

Limitation of reproduction by feeding condition in a carabid beetle,Carabus yaconinus

1. Effects of the amount of food consumed on reproduction of the carabid beetle, Carabus yaconinus B., were studied in the laboratory by rearing beetles at different food levels, and the feeding and oviposition rates in the field were estimated on the basis of the relationships between the amount of food consumed, body weight and egg production obtained in the experiment.
2. The maximum amount of food consumed was 150 mg of minced beef per day. The number of eggs laid per day and the mean body weight increased with an increase in the amount of food consumed. High mortality occurred only when the beetles consumed less than 25 mg of minced beef per day.
3. The ratio of current body weight to the minimum one just before death by starvation, W/W_min, was used for the estimation of the rates of food consumption and egg production. The relationships between mean W/W_min ratio, the amount of food consumed and the number of eggs laid per day were clarified.
4. The relationships between ovary states (ovary weight and the number of mature eggs in the ovary) and W/W_min ratio were examined for the females caught in the field. Females with higher values of W/W_min ratio had more mature eggs.
5. The amount of food consumed by females in the field during the reproductive period was estimated to be 50–70% of the maximum value attained in the experiment and the estimated rate of oviposition was 45–59% of the maximum rate attained in the experiment.

     

Long adult life spain and polyphagy of a carabid beetle,Leptocarabus kumagaii in relation to reproduction and survival

The reproductive patterns of two different age classes of Leptocarabus kumagaii adults, i.e. young beetles which were in the first year of adult stage and old ones which survived for more than two years, were studied in the field. The numbers of eggs laid were compared between young and old females and the effects of food quality and quantity were studied on egg production and maintenance in the laboratory.

1. Both of the young and old beetles reproduced in the field population. The number of eggs laid did not significantly differ between the two.
2. The number of eggs laid by female increased with an increase in the amount of miced beef eaten. Plant materials were ineffective for egg production but sufficient for the survival of both male and female beetles.
3. The two life historical aspects, i.e. repeated reproduction and polyphagy were discussed in relation to environmental fluctuation and limitation of food resources.

     

蛋白激酶C分子的成熟与活化

蛋白激酶C（Proteinkinase C,PKc）是细胞内一类重要的Ser／Thr激酶,调控多种生命活动的信号转导过程,目前已发现了至少11种亚型,其结构有一定的保守性而又有所差别,导致其功能和调控的多样性。新合成的PKC一般需要经历活化茎环（Acti．vation-loop,A—loop）、转角模体（Tummotif,T1V1）以及疏水模体（hydrophobic motif,HM）的程序性磷酸化过程才能成熟,获得进一步活化的功能。本文综述了近年来PKC的程序性磷酸化成熟以及活化的研究进展情况。     

Ultrastructure of pigmented eyes in Dorvilleidae (Annelida,Errantia, Eunicida) and their importance for understanding the evolution of eyes in polychaetes

Among polychaetes, the errant forms are the only group known so far possessing true multicellular eyes in adults which are preceded by bicellular larval eyes in many species. Most likely, two pairs of such eyes showing a specific structure belong to the ground pattern of Errantia = Aciculata. However, these eyes have primarily been investigated in only two subgroups of Errantia, but data on the third main taxon, Eunicida, are available for only two taxa. In the present investigation, the eyes in two additional species of Eunicida, the dorvilleids Protodorvillea kefersteini and Schistomeringos neglecta, were studied. In P. kefersteini, usually described as possessing one pair of small eyes, two pairs could be detected, whereas in S. neglecta only one pair was found. Each eye is made up of rhabdomeric photoreceptor cells, pigment cells and unpigmented supportive cells. Lenses or vitreous bodies are absent. From their structure most likely all eyes represent adult eyes and even the small anterior eyes in P. kefersteini structurally resemble miniaturized adult eyes. Neither persisting larval eyes nor unpigmented rhabdomeric ocelli were found in the two species. The observations in Dorvilleidae confirm the hypothesis of a common origin of adult eyes in Errantia.     

蛋白激酶C分子的成熟与活化

蛋白激酶C(Protein kinase C,PKC)是细胞内一类重要的Ser/Thr激酶,调控多种生命活动的信号转导过程,目前已发现了至少11种亚型,其结构有一定的保守性而又有所差别,导致其功能和调控的多样性。新合成的PKC一般需要经历活化茎环(Activation-loop,A-loop)、转角模体(Turn motif,TM)以及疏水模体(hydrophobic motif,HM)的程序性磷酸化过程才能成熟,获得进一步活化的功能。本文综述了近年来PKC的程序性磷酸化成熟以及活化的研究进展情况。     

On the visual perception of the eyes in adult Hyalomma truncatum ticks (Acari: Ixodidae)

Adult Hyalomma truncatum ticks with uncovered and foil-covered eyes were exposed to an upright-positioned rectangle as a target giving a luminance contrast ratio of 5:1 at a sun-simulating radiation. The transmission rate of the foil was less than 0.003%. Significantly (p0.05) more locomotorally active ticks with uncovered eyes (36.6%) responded to the target than ticks with foil-covered eyes (7.3%). When the rectangle was illuminated by monochromatic light at wavelengths ranging between 420 and 648 nm, the target induced a positive scototaxis in ticks with uncovered eyes regardless of the wavelength range. In contrast, ticks with covered eyes did not exhibita positive scototaxis at wavelength ranges of 553–585 and 608–648 nm and very few ticks responded only to other wavelength ranges. The results indicate that the eyes are the only or at least the most essential sense organs in the visual system of adult H. truncatum ticks.     

粪食性金龟的行为及其适应演化

粪食性金龟的行为简单分为3类:粪居型(endocoprid或dwelling behaviour)、直接掘洞型(paracoprid或tunnelling behaviour)和滚粪球型(telocoprid或rolling behaviour)。文章详细阐述各类型的特点,探讨粪食性金龟行为的进化趋势和过程,并进一步阐述粪食性金龟的适应演化。     

Effect of dung burial by the dung beetle Bubas bison on numbers and viability of Cryptosporidium oocysts in cattle dung

Cryptosporidium oocysts were inoculated into fresh dung (∼1.2 × 10⁴ oocysts per gram wet weight) and fed to dung beetles to assess the effect of dung burial by the dung beetle Bubas bison on the distribution of the oocysts in small cores of soil in the laboratory. The experiment consisted of five replicates of each of two treatments; controls (dung but no dung beetles) and the experimental treatment (inoculated dung and seven pairs of dung beetles). After 5 days, when approximately 90% of the dung was buried, the surface and buried dung was recovered and subsampled. The oocysts in the subsamples were recovered and enumerated using qPCR. Oocyst viability was evaluated using an assay based on the exclusion or inclusion of two fluorogenic vital dyes, 4′,6-diamidino-2-phenylindole (DAPI) and propidium iodide (PI). Results revealed that overall 13.7% of oocysts remained on the surface and 86.3% of oocysts were buried. The viability of oocysts in buried dung was only 10% compared to oocysts the surface dung (58%). Therefore, widespread dung burial by B. bison during the winter months could substantially reduce the numbers of Cryptosporidium oocysts available to be washed into waterways following winter rains.     

Maturation of Potassium-Stimulated Respiration in Rat Cerebral Cortical Slices

Abstract: Maximal dinitrophenol-stimulated respiration and K⁺-stimulated respiration were measured polarographically in cerebral cortical slices taken from rats aged 2–60 days. Increasing K⁺ concentrations produced an increase in respiration in slices from animals aged 15 days and older, but not in slices from animals aged 10 days and younger. Dinitrophenol-stimulated respiration, or the maximal respiratory capacity of the tissue, showed a similar increase between 10 and 15 days of age. At each age the maximal respiratory capacity was 6–8 ng at 0/mg protein min greater than the maximal K⁺-stimulated respiration.