Post-embryonic photoreceptor development and dark/light adaptation in the spittle bug Philaenus spumarius (L.) (Homoptera, Cercopidae)

The aims of this paper have been to describe (1) the general structure of the compound eye of the spittle bug Philaenus spumarius, (2) the eye's post-embryonic development, (3) photomechanical changes upon dark/light adaptation in the eye, and (4) how leaving the semi-aquatic foam bubble and turning into an adult affects the organization of the eye. Spittle bugs, irrespective of size or sex, possess apposition type compound eyes. The eye's major components (i.e. facet, cornea, cone and rhabdom) grow rather isometrically from the smallest nymph to the adult. Photomechanical changes can occur during both nymphal and adult phases and manifest themselves through pigment granules and mitochondria migrating to and away from the rhabdom, and rhabdom diameters varying with time of day and ambient light level. When a nymph transforms into an adult, its compound eyes’ dorsoventral axes widen, facet diameters increase, facet shapes turn from circular to pentagonal and hexagonal, the cornea thickens and the rhabdoms become thinner. The agile adults, free from the foam that surrounds the nymphs, can be expected to need their vision more than the nymphs, and the changes in eye structure do, indeed, indicate that the adults have superior visual acuity. A thicker cornea in the adults reduces water loss and protects the compound eye from mechanical and light-induced damage: protection given to the nymphs by their foam bubbles.     

Daily changes of structure,function and rhodopsin content in the compound eye of the crabHemigrapsus sanguineus

The compound eye of the crab hemigrapsus sanguineus undergoes daily changes in morphology as determined by light and electron microscopy, both in the quantity of chromophore substances studied by HPLC and in visual sensitivity as shown by electrophysiological techniques. 1. At a temperature of 20 degrees C, the rhabdom occupation ratio (ROR) of an ommatidial retinula was 11.6% (maximum) at midnight, 8.0 times larger than the minimum value at midday (1.4%). 2. Observations by freeze-fracture revealed that the densities of intra-membranous particles (9-11 nm in diameter) of rhabdomeric membrane were ca. 2000/microns 2 and ca. 3000/microns 2 for night and daytime compound eyes, respectively. 3. Screening pigment granules migrated longitudinally and aggregated at night, but dispersed during the day. Reflecting pigment granules migrate transversally in the proximal half of the reticula layer i.e. cytoplasmic extensions containing reflecting pigment granules squeeze between neighbouring retinula cells causing optical isolation (Fig. 4). Thus the screening pigment granules within the retinula cells show longitudinal migration and radial movement so that the daytime rhabdoms are closely surrounded by the pigment granules. 4. At 20 degrees C, the total amount of chromophore of the visual pigment (11-cis and all-trans-retinal) was 1.4 times larger at night than during the day i.e. 46.6 pmol/eye at midnight and 33.2 pmol/eye at midday. Calculations of the total surface area of rhabdomeric membrane, total number of intra-membranous particles in rhabdomeric membrane and the total number of chromophore molecules in a compound eye, indicate that a considerable amount of chromophore-protein complex exists outside the rhabdom during the day. 5. The change in rhabdom size and quantity of chromophore were highly dependent on temperature. At 10 degrees C both rhabdom size and amount of chromophore stayed close to daytime levels throughout the 24 hours. 6. The intracellularly determined relative sensitivity of the dark adapted night eye to a point source of light was about twice as high as the dark-adapted day eye. Most of the increase in the sensitivity is attributed primarily to the effect of reflecting pigment migration around the basement membrane and, secondarily, to the changes in the amount and properties of the photoreceptive membrane. The results form the basis of a detailed discussion as to how an apposition eye can function possibly as a night-eye.     

灰茶尺蠖成虫复眼的超微结构及其明暗适应中的变化

【目的】明确灰茶尺蠖Ectropis grisescens成虫复眼的超微结构及其明暗适应中的变化,探究其调光机制。【方法】采用超景深显微镜测定了灰茶尺蠖成虫复眼的小眼数量、间角、直径和曲率半径等外部参数,并通过组织切片、光学显微镜和透射电子显微镜等技术观察了复眼的内部超微结构;通过光学显微镜观察了灰茶尺蠖成虫复眼在明暗环境中分别适应2 h后晶锥结构及色素颗粒的位置变化。【结果】灰茶尺蠖成虫复眼呈半球形,雌、雄虫单个复眼分别有2 502±105和3 123±78个小眼。小眼自远端至近端由角膜、晶锥、透明区构成的屈光层和由15个视网膜细胞构成的感光层组成。2个初级色素细胞包裹着晶锥,自角膜近端延伸至视网膜细胞核区的远端;每个小眼外围由6个次级色素细胞围绕,自角膜近端延伸至基膜;在透明区内14个视网膜细胞聚集成束(非感杆束),远端与晶锥束末端连接,在感光层内形成闭合型感杆束,延伸至第15个视网膜细胞(基部视网膜细胞)。在明暗适应时,灰茶尺蠖复眼的晶锥细胞间出现开闭,色素颗粒进行纵向位移,以适应外界的光强度的变化。【结论】灰茶尺蠖成虫复眼属于重叠像眼,感杆束为“14+1”模式;屏蔽色素颗粒的移动是其复眼适应外界光强度变化的重要机制。     

The Development of the Rhabdom and the Appearance of the Electrical Response in the Insect Eye

Electron microscopic studies on the development of the rhabdom in the compound eye of the silkworm moth and pupa (Bombyx mori) were carried out in parallel with the recording of the electrical response to photic stimulation. No electrical response to photic stimulation was recorded from the pupal compound eye which had no trace of differentiation of the rhabdom. With the differentiation of development of the rhabdom in the pupal compound eye, electrical responses could be recorded, and the amplitude of such electrical responses increased with the progress of development of the rhabdom. These observations suggest that the rhabdom is probably the site of the photochemical reaction which leads to the generation of the slow retinal action potentials.     

飞蝗复眼生理和结构上的节律变化

采用细胞内记录和光镜方法研究了飞蝗(Locusta migratoria)夜间和日间在暗适应和明适应状态下小网膜细胞角敏感度以及晶锥和小网膜细胞之间区域结构上的变化.结果表明小网膜细胞角敏感度的变化不仅仅由于晶锥周围主色素细胞色素颗粒的移动,而且也由于小眼感杆束结构上的节律变化.     

Circadian rhythmicity in cockroaches: effects of early post-embryonic development and ageing

ABSTRACT. Effects of early post-embryonic development and ageing on the circadian rhythm of locomotor activity in the cockroach, Leucophaea maderae , were investigated. Cockroach nymphs (first to fourth instars) were found to exhibit a circadian rhythm of activity generally similar to that of adults, but there appeared to be a significant change in pacemaker period (τ) early in nymphal development, and nymphs exhibited regular fluctuations in activity level which may be related to the moulting cycle. Mean τ of adults was remarkably stable with age – no significant change was found in either males or females throughout the life span of the adult. There was, however, a small but significant difference between the average period (τ) of adult males (τ= 23.72 ± 0.12h) and adult females (τ= 23.84 ± 0.13 h). Lighting conditions during post-embryonic development were found to have major effects on τ of adults. Males raised in constant darkness had a significantly shorter period (τ= 23.52 ± 0.11 h) than males raised in LD 12:12, and adult males and females which had been raised in non-24-h light cycles (T = 22 h or 26 h) exhibited major differences in τ from animals raised in LD 12:12 which persisted for several months. Animals exposed as adults to non-24-h light cycles also showed 'after-effects' on τ, but the magnitude of the effect was much less than that exhibited by animals exposed as nymphs.     

昆虫复眼瞳孔调节的一种新机制

本文采用细胞内记录方法,研究了蝗虫和螽斯复眼侧区小眼在不同时间(日间和夜间)和不同适应状态(暗适应和明适应)下,小网膜细胞角灵敏度的变化规律.结果表明:小网膜细胞角灵敏度的变化不仅与适应状态有关,而且伴随时间的变化而变化.小眼感杆束直径相应变化用光镜方法得到证实,我们认为在昆虫复眼中存在一种新的瞳孔调节机制.     

Organisation of the compound eye of a tipulid fly during the day and night

Summary The ultrastructure of the compound eye of the Australian tipulid fly,Ptilogyna spectabilis, is described. The ommatidia are of the acone type. The rhabdom corresponds to the basic dipteran pattern with six outer rhabdomeres from retinular cells 1–6 (R1-6) that surround two tiered central rhabdomeres from R7 and 8. Distally, for about 8 m, the rhabdom is closed. For the remainder, where the rhabdomere of R8 replaces that of R7, the rhabdom is open, and the rhabdomeres lie in a large central ommatidial extracellular space. In the proximal two thirds of the rhabdom, the central space is partitioned by processes from the retinular cells so that the individual rhabdomeres are contained in pockets.At night the rhabdom abuts the cone cells, but during the day it migrates some 20 m proximally and is connected to a narrow (1–2 m) cone cell tract. This tract is surrounded by two primary pigment cells, which occupy a more lateral position at night and thus act like an iris. Pigment in secondary pigment cells also migrates so as to screen orthodromic light above the rhabdom during the day. Between midday and midnight, the rhabdom changes in length and cross-sectional area as a result of asynchrony of the shedding and synthetic phases of photoreceptor membrane turnover. The effects of these daily adaptive changes on photon capture ability are discussed with regard to the sensitivity of the eye.     

Larval and adult eye of the Western Rock Lobster (Panulirus longipes)

A number of differences exists between the compound eyes of larval and adult rock lobsters, Panulirus longipes. The larval eye more closely resembles the apposition type of compound eye, in which retinula cells and rhabdom lie immediately below the cone cells. The adult eye, on the other hand, is a typical clear-zone photoreceptor in which cones and retinula cell layers are separated by a wide transparent region. The rhabdom of the larval eye, if cut longitudinally, exhibits a "banded" structure over its entire length; in the adult the banded part is confined to the distal end, and the rhabdom is tiered. Both eyes have in common an eighth, distally-located retinula cell, which possesses orthogonally-oriented microvilli, and a peculiar lens-shaped "crystal", which appears to focus light onto the narrow column of the distal rhabdom. Migration of screening pigment on dark-light adaptation is accompanied by changes in sensitivity and resolution of the eye. Retinula cells belonging to one ommatidium do not arrange into one single bundle of axons, but interweave with axons of four neighbouring facets in an extraordinarily regular fashion.     

Postembryonic changes in circadian photo-responsiveness rhythms of optic lobe interneurons in the cricket Gryllus bimaculatus

The photo-responsiveness of 2 groups of interneurons responding to light in the protocerebrum was investigated at 2 developmental stages, the last instar nymphs and adults, in the cricket Gryllus bimaculatus. The cricket is diurnally active during the nymphal stage but becomes nocturnal as an adult. In both adults and nymphs, light-induced responses of optic lobe light-responding interneurons that conduct light information from the optic medulla to the lobula and the cerebral lobe showed a circadian rhythm peaking during the subjective night. Amplitudes of the rhythms were not significantly different between adults and nymphs, but adults showed more stable day and night states than did nymphs. The medulla bilateral neurons that interconnect the bilateral medulla areas of the optic lobe also showed circadian rhythms in their light-induced responses in both adults and nymphs. The rhythm had a clear peak and a trough in adults, and its amplitude was significantly greater than that of nymphs. These results suggest that the 2 classes of interneurons are differentially controlled by the circadian clock. The difference might be related to their functional roles in the animal's circadian behavioral organization.     

Photic regulation of heme oxygenase activity in the golden hamster retina: involvement of dopamine

The photic regulation of heme oxygenase (HO) activity was examined in the golden hamster retina. This enzymatic activity was significantly higher at midday than at midnight. When the hamsters were placed under constant darkness for 48 h and killed at subjective day or at subjective night, the differences in HO activity disappeared. Western blot analysis showed no differences in HO levels among these time points. Dopamine significantly increased this activity in retinas excised at noon or at midnight, with a higher sensitivity at night. The effect of dopamine was reversed by SCH 23390 but not by spiperone and clozapine and it was not reproduced by quinpirole. In vitro, the increase in HO activity found in retinas incubated under light for 1 h was significantly reduced by SCH 23390. Two cAMP analogs increased HO activity and their effect, as well as the effect of dopamine was blocked by H-89, a protein kinase A (PKA) inhibitor. Tin protoporphyrin IX, an HO inhibitor, significantly decreased cGMP accumulation with maximal effects during the day. Low concentrations of bilirubin decreased retinal thiobarbituric acid substances levels (an index of lipid peroxidation) in basal conditions and after exposing retinal cells to H2O2. These results suggest that hamster retinal HO activity is regulated by the photic stimulus, probably through a dopamine/cAMP/PKA dependent pathway.     

Performance of nymphal and adult southern green stink bug on an overwintering host plant and impact of nymph to adult food-switch

Performance of nymphs and adults of the southern green stink bug,Nezara viridula (L.) (Heteroptera: Pentatomidae) feeding on immature fruits of radish,Raphanus raphanistrum L. (Brassicaceae) was poor compared with feeding on immature soybean [Glycine max (L.) Merrill] pods, but a change from radish to soybean from nymph to adult alleviated some of the detrimental effects of this food on adult performance. Over 50% of nymphs died on radish, while only 15% died on soybean. To complete nymph development (stadia 2 through 5), male and female nymphs required 9.5 and 13.3 days longer on radish compared to soybean. Fresh body weight of 1-d-old adult females and males was ≅65% greater for those reared as nymphs on soybean than on radish, with females being significantly heavier than males on both foods. Survivorship and longevity of females, however, was greater for those fed on radish as nymphs and as adults, as compared to those that fed on soybean or radish followed by soybean. Males lived longer when fed exclusively on soybean. In general, reproductive performance and percentage gain in adult body weight was greatest for insects fed on soybean only, and were better for those whose food was switched from nymph to adult, as compared to those insects fed on radish solely.     

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.     

Habitat use and phenology of the large insular endemic grasshopper Acrostira euphorbiae (Orthoptera: Pamphagidae)

The habitat use and the phenology of the large grasshopper Acrostira euphorbiae García and Oromí endemic to La Palma (Canary Islands) are studied. This grasshopper is entirely dependent on the Canarian endemic shrub Euphorbia lamarckii both for food and to avoid predation. Adults stay on subapical branches during the day, probably to reduce the risk of predation, and climb up to the apex at night to feed. While females seem to ensure the genetic diversity of offspring by waiting for visits by different males, the latter have to move to guarantee their reproductive success. Monophagy in this species may be related to the year-round presence of tender shoots, and to the predator-repellent toxic latex found in the host plant. Unlike related species from continental areas, adults and nymphs of A. euphorbiae are present almost all year round, probably in adaptation to the particular climate of the islands. Nymphs are more abundant in winter, when Euphorbia leaves are most available. However, adults are more abundant than nymphs in spring, summer and autumn. Males develop more quickly than females, an apparent reproductive strategy based on achieving sexual maturity to coincide with females undergoing imaginal moulting. Matings start immediately after adult females emerge. Densities oscillated between 73 and 193 individuals ha(-1), which can be considered a low value compared with other continental pamphagid species.     

Changes in circadian rhythms during puberty in Rattus norvegicus: developmental time course and gonadal dependency

During puberty, humans develop a later chronotype, exhibiting a phase-delayed daily rest/activity rhythm. The purpose of this study was to determine: 1) whether similar changes in chronotype occur during puberty in a laboratory rodent species, 2) whether these changes are due to pubertal hormones affecting the circadian timekeeping system. We tracked the phasing and distribution of wheel-running activity rhythms during post-weaning development in rats that were gonadectomized before puberty or left intact. We found that intact peripubertal rats had activity rhythms that were phase-delayed relative to adults. Young rats also exhibited a bimodal nocturnal activity distribution. As puberty progressed, bimodality diminished and late-night activity phase-advanced until it consolidated with early-night activity. By late puberty, intact rats showed a strong, unimodal rhythm that peaked at the beginning of the night. These pubertal changes in circadian phase were more pronounced in males than females. Increases in gonadal hormones during puberty partially accounted for these changes, as rats that were gonadectomized before puberty demonstrated smaller phase changes than intact rats and maintained ultradian rhythms into adulthood. We investigated the role of photic entrainment by comparing circadian development under constant and entrained conditions. We found that the period (τ) of free-running rhythms developed sex differences during puberty. These changes in τ did not account for pubertal changes in entrained circadian phase, as the consolidation of activity at the beginning of the subjective night persisted under constant conditions in both sexes. We conclude that the circadian system continues to develop in a hormone-sensitive manner during puberty.     

Day–night changes in the spatial distribution and habitat preferences of freshwater shrimps, Gammarus pulex, in a stony stream

1. As many invertebrates are nocturnal, their spatial distribution and habitat preferences may change from day to night. Both aspects are examined for Gammarus pulex by testing the hypotheses: (i) a power function was a suitable model for the spatial distribution of the shrimps in both day and night; (ii) diurnal and nocturnal spatial distributions were significantly different; (iii) diurnal and nocturnal habitat preferences were significantly different. Five different life‐stages were treated separately. To ensure that the conclusions were consistent, large samples were taken near midday and midnight in April, June and November over 4 years at two sites about 3 km apart in a stony stream: downstream (n = 30) and upstream (n = 50). 2. The first and second hypotheses were supported at both sites. A power function, relating spatial variance (s²) to mean (m), was an excellent fit in all analyses (P < 0.001, r² > 0.91), i.e. the spatial variance was density‐dependent. All five life‐stages were aggregated in the day. At night, the degree of aggregation increased for juveniles at higher densities but decreased for juveniles at lower densities, increased for immature females and males, but decreased slightly for mature females and especially mature males, the latter being close to a random distribution. There were no significant differences between sites, in spite of the lower numbers at the downstream site. 3. The third hypothesis was tested at only the upstream site and supported by comparisons between shrimp densities and 13 physical variables (distance from bank, water depth, water velocity, ten particle size‐classes), and three non‐physical variables (dry weights of bryophytes, leaf material, organic detritus). During the day, densities were strongly related to particle sizes with the following preferences: 0.5–8 mm for juveniles, 8–256 mm for the other life‐stages with a weaker relationship for males. There were no significant positive relationships with the other variables, apart from bryophytes for immature shrimps and adults. At night, densities were unrelated to particle size; juveniles and immature shrimps preferred low water velocities near the banks, often where leaf material and organic detritus accumulated, females often preferred medium water velocities slightly away from the banks, and males showed no habitat preferences. 4. Day samples do not provide a complete picture of habitat preferences and probably identify refuge habitats. Day–night changes in spatial distribution and habitat preferences are an essential part of the behavioural dynamics of the shrimps and should be investigated in other species.     

Ommatidial structure in relation to turnover of photoreceptor membrane in the locust

Summary In the compound eye of the locust, Locusta, the cross-sectional area of the rhabdoms increases at dusk by 4.7-fold due to the rapid assembly of new microvillar membrane, and decreases at dawn by a corresponding amount as a result of pinocytotic shedding from the microvilli. The rhabdoms at night have more and longer photoreceptor microvilli than rhabdoms during the day. The orientations of the six rhabdomeres that comprise the distal rhabdom also change. The density of intramembrane particles on the P-face of the microvillar membrane, putatively representing mostly rhodopsin molecules, or aggregates thereof, does not change.An alteration in the size of the ommatidial field-stop, produced by the primary pigment cells, is concomitant with the change in rhabdom size. At night the increase in size of the field-stop must widen the angular acceptance of a rhabdom, increasing the capture of photons from an extended field. Conversely, during the day, when photons are more abundant, its decrease must narrow the acceptance angle, increasing angular resolution. Because of the presence of this field-stop, the optics of the ommatidium would not be greatly affected if the rhabdom were to remain always at its night size. It is argued, therefore, that the variable-size rhabdom must have resulted from some demand other than that of light/dark adaptation.Changes in size and organisation of the rhabdoms in response to various light regimes indicate that: (1) Rapid shedding of photoreceptor membrane is induced by the onset of light, but shedding also occurs slowly in darkness during the day. (2) Microvillar assembly is initiated by the onset of darkness, but also occurs at the normal time of dusk without a change in ambient lighting, provided there has been some light during the day. Therefore, both shedding and assembly of microvillar membrane are affected by the state of illumination, but also appear to be under some endogenous control.     

Mobility of mass-reared diapaused and nondiapaused Cydia pomonella (Lepidoptera: Tortricidae): effect of mating status and treatment with gamma radiation

Mass-reared insects may differ in their behavioral traits depending on whether they have undergone diapause. We studied the mobility of mass-reared diapaused and nondiapaused Cydia pomonella (L.) with a focus on understanding the effect of mating status and treatment with gamma radiation as these insects are destined for use in an areawide program that uses the sterile insect technique (SIT). Actograph-measured mobility was assessed one gender at a time for 4 h during which the photoperiod transitioned from day to night. We tested 20-30 individuals per treatment. For experiments on the effect of mating status, we used 24-48-h-old adults (diapause [D]-virgin, D-mated, standard [N]-virgin, and N-mated), which is the typical age class that is released in the SIT program. Diapaused females were significantly more mobile than females reared through standard production, whereas no differences were detected in male mobility because of rearing strategy. Mated females were significantly more mobile than virgin females, whereas no difference in mobility because of mating status was detected for males. Mated females were significantly more mobile than mated males. In contrast, virgin females were significantly less mobile than virgin males. For experiments on the effect of treatment with gamma radiation, adults from all rearing strategies and treatments (D-0Gy, D-100Gy, D-250Gy, N-0Gy, N-100Gy, and N-250Gy), were tested simultaneously. Adult males were tested at two different constant temperatures (25 and 20 degrees C), whereas adult females were tested only at 25 degrees C. For standard-reared adults, we found a significant linear decrease in mobility as the radiation dose increased from 0 to 250 Gy. In contrast, the relationship between mobility and dose of radiation was quadratic for diapaused males and absent for diapaused females.     

Interspecific variations in the morphology and ultrastructure of the rhabdoms of oplophorid shrimps

Interspecific variations in rhabdom structure between various oplophorid shrimps are described and the differences are related to the light environment at different depths within the mesopelagic zone. The ultrastructure of the distal rhabdom in these species is described for the first time. Quantitative measurements show that the proportion of the rhabdom layer occupied by the distal rhabdom varies from 3.5-25% in the dorsoventral plane of the eye of Systellaspis debilis. The distal rhabdom occupies less than 1% of the rhabdoms in the eye of Acanthephyra pelagica, where it can only be seen by using the electron microscope. It is suggested that the rhabdoms of those species that remain within the photic zone (such as S. debilis) are adapted to maximize contrast, whereas in those whose depth ranges extend into the aphotic zone (such as A. pelagica) they are adapted for maximum sensitivity.