龟纹瓢虫成虫的复眼形态及其显微结构

利用光镜、组织切片法观察了龟纹瓢虫Propylaea japonica(Thunberg)成虫的复眼形态及其显微结构。结果如下:(1)头正前方观,复眼外形似半球,且后方稍向内合拢。每个复眼约包括630个小眼。(2)每个小眼是由1套屈光器(1个角膜和1个晶锥)、6至8个小网膜细胞及其特化产生的视杆和基细胞等几部分组成。晶体周围及小网膜色素细胞内均含有丰富的色素颗粒。(3)小眼整体纵切显示,其上、下段色素颗粒分布相对较多,中段分布较少。(4)明、暗适应状态对小眼的色素颗粒分布有影响,性别对其分布无明显影响。明适应状态下,其色素颗粒较均匀地分布于视杆两侧上下,暗适应状态时色素颗粒则主要分布在视杆部位的上侧,显示其具有一定的重叠眼性质;而在相同的明、暗适应状态下其雌、雄成虫复眼的色素颗粒分布间无明显差异。     

Optical parameters of euphausiid eyes as a function of habitat depth

Summary The relationships between habitat depth, eye diameter relative to body length, and the dimensions of rhabdoms and crystalline cones have been examined for 13 species of three oceanic euphausiid genera with habitats ranging from near-surface waters to the deep-sea. Rate of eye growth decreases with depth. Longer rhabdoms may increase the visual sensitivity to point and extended light sources by an eye of a particular size with depth. Larger interommatidial angles suggest that visual acuity decreases at depth. Depth-related changes in euphausiid eyes are considered with respect to the probable roles of vision and bioluminescence in the deep-sea. Unusual features of the eyes of several species are described.     

亲土苔蛾成虫复眼超微结构

【目的】重叠型眼在昆虫复眼演化中起着重要作用。本研究旨在阐明夜出型亲土苔蛾Manulea affineola复眼类型及结构特征,以期填补灯蛾亚科昆虫复眼研究的空白,扩充夜出型昆虫复眼的特征数据,为探讨重叠型眼的变异趋势及复眼演化提供依据。【方法】运用光学和透射电子显微技术观察亲土苔蛾成虫复眼的超微结构。【结果】亲土苔蛾成虫复眼具有一个透明区,由6个次级色素细胞的透明胞质构成。小眼具8个视网膜细胞,其中1个视网膜细胞较短,仅位于小眼基部。在透明区内,7个视网膜细胞聚集成一束,其远端与晶体束末端相接,但并不形成视杆。在透明区下方,这7个视网膜细胞形成一个中心融合的视杆。在复眼背缘区的小眼的视杆具有近似矩形的横截面,而其余小眼的视杆具多分支状截面。【结论】亲土苔蛾成虫复眼属于重叠型眼;复眼背缘区的矩形视杆很可能与昆虫的偏振敏感性有关。     

Ultrastructural study of the naupliar eye of the ostracodeVargula graminicola (Crustacea,Ostracoda)

Summary Ostracodes, like other crustaceans, have a simple naupliar eye that is built upon a theme of three eye cups surrounded by a layer of screening pigments. The single naupliar eye of the ostracodeVargula graminicola is situated medially on the dorsal-anterior side of the body and has three fused eye cups, two dorso-lateral and one ventral. Each eye cup has the following components: (1) pigment cells between the eye cups, (2) tapetal cells, (3) retinular cells with (4) microvillar rhabdomeres, and (5) axons extending into the protocerebrum. Typically two retinular cells contribute lateral microvilli to each rhabdom. The two dorso-lateral eye cups have about 40 retinular cells (20 rhabdoms) and the ventral eye cup has about 30 retinular cells (15 rhabdoms). Typical of myodocopid naupliar eyes (as reported from light microscopic studies), no lens cells or cuticular lenses were observed. The presence of tapetal cells identifies theVargula eye as a maxillopod-ostracode type crustacean naupliar eye. It is unlikely that the naupliar eye ofV. graminicola functions in image formation, rather it probably functions in the mediation of simple taxis towards and away from light.     

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.     

The ultrastructure of the compound eye of Munida rugosa (Crustacea: Anomura) and pigment migration during light and dark adaptation

The galatheid squat lobster, Munida rugosa, has compound eyes of the reflecting superposition type in which a distal cone cell layer and a proximal rhabdom layer are separated by an extensive clear zone. The eye is shown to have certain unique features. In all other reflecting superposition eyes, the clear zone is traversed by crystalline tracts formed by the cone cells. In M. rugosa a thin distal rhabdom thread, formed by the eighth retinula cell, connects the cones to the proximal fusiform rhabdoms. The cytoplasm of the other retinula cells also crosses the clear zone in a complex pattern. Fully light-adapted ommatidia are optically isolated by limited migrations of distal shielding pigments. A reflecting pigment multilayer lines each cone to facilitate the formation of a superposition image. This also shows a light-induced change which may limit the acceptance angle of the eye during light adaptation.     

Localization of the pupil trigger in insect superposition eyes

Summary In the superposition eyes of the sphingid moth Deilephila and the neuropteran Ascalaphus, adjustment to different intensities is subserved by longitudinal migrations of screening pigment in specialized pigment cells. Using ophthalmoscopic techniques we have localized the light-sensitive trigger that controls pigment position.In both species, local illumination of a small spot anywhere within the eye glow of a dark-adapted eye evokes local light adaptation in the ommatidia whose facets receive the light. Details of the response pattern demonstrate that a distal light-sensitive trigger is located axially in the ommatidium, just beneath the crystalline cone, and extends with less sensitivity deep into the clear zone. The distal trigger in Deilephila was shown to be predominantly UV sensitive, and a UV-absorbing structure, presumably the distal trigger, was observed near the proximal tip of the crystalline cone.In Ascalaphus we also found another trigger located more proximally, which causes local pigment reaction in the ommatidia whose rhabdoms are illuminated (the centre of the eye glow). The light-sensitive trigger for this response appears to be the rhabdom itself.     

Visual field structure in the Empress Leilia, Asterocampa leilia (Lepidoptera, Nymphalidae): dimensions and regional variation in acuity

Male Empress Leilia butterflies ( Asterocampa leilia) use a sit-and-wait tactic to locate mates. To see how vision might influence male behavior, we studied the morphology, optics, and receptor physiology of their eyes and found the following. (1) Each eye's visual field is approximately hemispherical with at most a 10 degrees overlap in the fields of the eyes. There are no large sexual differences in visual field dimensions. (2) In both sexes, rhabdoms in the frontal and dorsal ommatidia are longer than those in other eye regions. (3) Interommatidial angles are smallest frontally and around the equator of the eye. Minimum interommatidial angles are 0.9-1 degrees in males and 1.3-1.4 degrees in females. (4) Acceptance angles of ommatidia closely match interommatidial angles in the frontal region of the eye. We conclude that vision in these butterflies is mostly monocular and that males have more acute vision than females, especially in the frontal region (large facets, small interommatidial angles, small acceptance angles, long rhabdoms, and a close match between interommatidial angles and acceptance angles). This study also suggests that perched males direct their most acute vision where females are likely to appear but show no eye modifications that appear clearly related to a mate-locating tactic.     

The Ultrastructure of the Compound Eye of Two Species of Marine Ostracodes (Crustacea: Ostracoda: Cypridinidae)

Ultrastructurally, the compound eyes of the luminescent marine ostracodes Vargula graminkola and V. tsujii are similar. These ostracodes have two lateral compound eyes, with relatively few ommatidia (13 and 20 respectively). They exhibit apposition type compound eyes as seen in many other arthropods. Each ommatidium includes: a flat, ectodermal cuticular covering, corneagen cells, two long cone cells that give rise to a large conspicuous crystalline cone, retinular cells, pigment cells, a microvillar rhabdom and proximal axonal neurons. The axons merge to form an optic nerve that extends into the brain through a short, muscular stalk that is surrounded externally by a cuticle. The number of retinular cells is typically six per ommatidium in V. graminicola and eight per ommatidium in V. tsujii. Screening pigment cells surround each ommatidium forming a layer that is about 5–15 pigment granules thick. In addition to pigment cells, the cytoplasm of the retinular cells includes numerous screening pigment granules. In light/dark adaptation, there are no obvious morphological differences in the orientation of the rhabdom or in the organization of the screening pigments. Both Vargula species studied are nocturnally active and bioluminescent suggesting that these eyes are capable receptors of the bright conspecific luminescence.     

The ontogeny of facultative superposition optics in a shrimp eye: hatching through metamorphosis

Summary Compound eyes of larval and first postlarval grass shrimp (Palaemonetes pugio Holthuis) were studied with light and electron microscopy following adaptation to darkness or bright light. Larvae have well-developed apposition eyes, including 3 main types of accessory screening and reflecting pigments and a fourth class of putatively reflective granules recently described in adult shrimps. Rhabdoms contain orthogonally layered microvilli, and by the last larval stage, 8 retinular cells. Ocular accessory pigments in both light- and dark-adapted larvae are distributed much like those of light-adapted adults, but the distal mass of reflecting pigment is concentrated dorsally in larvae and ventrally in adults. Since larvae swim upside-down, reflecting pigment is oriented downward in all developmental stages and may function for countershading. Light and dark adaptational migrations of all 3 major accessory pigments commence abruptly at metamorphosis to the first postlarva. Upon dark adaptation in postlarvae, superposition optics remain impossible because (1) distal screening pigment migrates only slightly, (2) no clear zone has developed, and (3) the crystalline cones remain circular in cross section. Nevertheless, a slight improvement in photon catch is expected due to extensive redistributions of reflecting pigment and retinular cell screening pigment granules.

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The nucleolar fibrillar centres in various cell types in vivo or in vitro

Summary The compound eye of male (haploid) Xyleborus ferrugineus beetles was examined with scanning and transmission electron microscopy. The eye externally consists of ca. 19 to 33 facets. Each ommatidium is composed of a thickly biconvex lenslet with about 50 electron dense and rare layers, but at the junction area between two lenslets there are only about 35 to 37 layers that can be distinguished. A very short (3.4–4.0 m) acone type crystalline cone is located directly beneath the lenslet. Each ommatidium is surrounded by pigment cells, and pigment granules also appear throughout the cytoplasm of the retinular cells. Some pigment granules are even present below the basement membrane. There are 8 retinular cells. The rhabdomeres of 2 centrally situated photoreceptor cells fuse into a rhabdom which is enveloped by the rhabdomeres of 6 peripheral retinular cells. The rhabdomeres of the 6 peripheral retinular cells join laterally to form a rhabdomeric ring around the central rhabdom. No tracheation was observed among the retinular cells. Virus-like particles are evident near the nucleus in each Semper cell of the crystalline cone.This research was supported by the Director of the Research Division, C.A.L.S., University of Wisconsin, Madison; and in part by research grant No. RR-00779 from the Division of Research Resources, National Institutes of Health and by funds from the Schoenleber Foundation, Milwaukee, WI to D.M.N.     

Ultrastructure of the compound eye of the diploid female beetle,Xyleborus ferrugineus

Summary The compound eye of female (diploid) Xyleborus ferrugineus beetles was examined with scanning and transmission electron microscopy. The eye is emarginate, and externally consists of roughly 70–100 facets. Each ommatidium is composed of a thickly biconvex lenslet with about 50 electron dense and rare layers. The lens facet overlies a crystalline cone of the acone type which is roughly hourglass-shaped. Pigment cells envelop the entire ommatidium, and pigment granules also are abundant throughout the cytoplasm of the 8 retinular cells. The rhabdomeres of 2 centrally situated photoreceptor cells effectively fuse into a rhabdom that extends from the base of the crystalline cone deeply into the ommatidium. Six distal peripheral retinular cells encircle the 2 central cells, and their rhabdomeres join laterally to form a rhabdomeric ring around the central rhabdom. The rhabdom and rhabdomeric ring are effectively separated by the cytoplasm of the two central retinular cells which contains the usual organelles and an abundance of shielding pigment granules. Eight axons per ommatidium gather in a tracheae-less fascicle before exiting the eye through the fenestrate basement membrane. No tracheation was observed among the retinular cells. Each Semper cell of each observed crystalline cone contained an abundance of virus-like particles near the cell nucleus. The insect is laboratory reared, and the visual system seems very amenable to photoreceptor investigations.This research was supported by the Director of the Research Division, C.A.L.S., University of Wisconsin, Madison; and in part by research grant No. RR-00779 from the Division of Research Resources, National Institutes of Health and by funds from the Schoenleber Foundation, Milwaukee, WI to D.M.N.     

Ultrastructure of the eyes of the grass shrimp,Palaemonetes pugio

Summary The cone cells and corneagenous cells possess extensive networks of smooth tubular endoplasmic reticulum that may be involved in optical reflectance and light-adaptational responses, respectively. The extracellular basal lamina of the basement membrane is confluent with glial cell capillary walls and may prove to be a viaduct for the transmission of hemolymph-borne substances to the retina or of retinal degradation products to the hemolymph. In addition to dense pigment granules, the distal pigment cells are shown for the first time to contain migratory reflecting platelets that are usually polymorphic in light-adapted eyes but are rectangular in dark-adapted eyes. In the latter these plates become aligned against the crystalline cones and presumably contribute to the reflection superposition optics of the grass shrimp. Dark-adapted retinular cells possess well-developed perirhabdomal cisternae, oblong or ovoid mitochondria, generally vesicular rough endoplasmic reticulum, and occasional, spherical, calcium-like intrarhabdomal inclusions. Light-adapted retinular cells possess poorly developed perirhabdomal cisternae, lamelliform rough endoplasmic reticulum, and condensed mitochondria frequently associated with lipid droplets and pigment granules. The cytoplasmic boundaries of the reflecting pigment cells expand into the extracellular spaces between individual ommatidial retinular cells during dark adaptation and recede to the interommatidial extracellular spaces during light adaptation. Cytoplasmic microfilament bundles found only at the bases of partially light-adapted rhabdomeric microvilli may be involved in microvillar shortening.     

Evolution of eye size and shape in primates

Strepsirrhine and haplorhine primates exhibit highly derived features of the visual system that distinguish them from most other mammals. Comparative data link the evolution of these visual specializations to the sequential acquisition of nocturnal visual predation in the primate stem lineage and diurnal visual predation in the anthropoid stem lineage. However, it is unclear to what extent these shifts in primate visual ecology were accompanied by changes in eye size and shape. Here we investigate the evolution of primate eye morphology using a comparative study of a large sample of mammalian eyes. Our analysis shows that primates differ from other mammals in having large eyes relative to body size and that anthropoids exhibit unusually small corneas relative to eye size and body size. The large eyes of basal primates probably evolved to improve visual acuity while maintaining high sensitivity in a nocturnal context. The reduced corneal sizes of anthropoids reflect reductions in the size of the dioptric apparatus as a means of increasing posterior nodal distance to improve visual acuity. These data support the conclusion that the origin of anthropoids was associated with a change in eye shape to improve visual acuity in the context of a diurnal predatory habitus.     

Visual behaviour and the structure of dark and light-adapted larval and adult eyes of the New Zealand glowworm Arachnocampa luminosa (Mycetophilidae: Diptera)

Both larval and adult New Zealand cave glowworms exhibit reactions to light; their photoreceptors must, therefore, be regarded as functional. The two principal stemmata of the larva possess large biconvex lenses and voluminous rhabdoms. Approximately 12 retinula cells are present. In light-adapted larvae the diameter of the rhabdom is 8 μm and that of an individual microvillus is 49.5 nm. Dark-adapted eyes have rhabdoms that measure 14 μm in cross section and microvilli with an average diameter of 54 nm. The compound eye of the adult comprises approximately 750 ommatidia, each with a facet diameter of 27–28 μm. A facet is surrounded by 1–6 interommatidial hairs which are up to 30 μm long. The interommatidial angle is 5.5°. Cones, consisting of 4 crystalline cone cells, are of the ‘acone’ type. Pigment granules in the primary pigment cells are twice as large as those of the retinula cells which measure 0.6–0.75 μm in diameter. The rhabdom is basically of the dipteran type, i.e. six open peripheral rhabdomeres surround 2 central rhabdomers arranged in a tandem position. The microvilli of cells 1–6 and cell 8 have diameters ranging from 68 to 73 nm, but those of the distally-located central rhabdomere 7 are 20% larger. This is irrespective of whether the eye is dark or light-adapted. In the latter the cones are long and narrow, the screening pigment granules closely surround the rhabdomeres, and the rhabdom is less voluminous than that of the dark-adapted eye.     

The micromorphology of the nauplius eye of the estuarine calanoid copepod, Sulcanus conflictus Nicholls (Crustacea)

The nauplius eye consists of one median and two lateral ocelli, each within a pigment cup. The three pigment cups are made up from two multi-nucleate pigment cells: each cell forming one lateral cup and half of the median cup. The three cups are lined on the insides by tapetal cells which contain layers of reflectile crystals. Each of the ocelli contains six sensory cells which protrude from the rims of the pigment cups and the protruding parts are sheathed by the conjunctiva cells. The whole eye is enveloped by a thin membrane which also sheaths the proximal parts of the five nerve bundles that leave the eye. All the sensory cells of the lateral ocelli are similar and have rhabdomeric microvilli on the terminal end, and contain phaosomes and a multitude of other organelles and cytoplasmic inclusions. The complex median ocellus contains a superior group of three retinular cells, linked by interdigitating processes, and an inferior group consisting of a large central cell enclosed in two cup-shaped peripheral retinular cells. A two-tiered rhabdome arrangement exists, with a rather complex inferior rhabdome set made up of a central rhabdomere and two hemi-annulate rhabdomeres. The cytoplasm of the retinular cells of the median ocellus lack phaosomes but instead contain double-walled tubular elements, possibly formed by the inpushings of microvilli into adjacent cells. The possible functional significance of the unique arrangement seen in the median ocellus is discussed. The retinular cells are of the inverse type. There are no efferent nerve fibres from the brain nor any nervous connection between the lateral and the median ocelli.     

Structure and functional aspects of the scotopic compound eye of the sugarcane borer moth

Morphology and functional aspects of the scotopic compound eye of the moth Diatraea saccharalis, studied using light and electron microscopy, is presented. An ommatidium is composed of a laminate corneal lens, four Semper cells, a refractive cone, two primary pigment cells, six screening pigment cells, a crystalline tract that functions as an optical waveguide, and six to eight sensory retinular cells. Accessory light regulators consist of screening pigment cells that, in the dark-adapted position, increase receptor sensitivity by permitting light rays to cross over to adjacent ommatidia and specialized tracheal regions that enhance sensitivity by reflecting light back toward sensory receptors.     

Visual pigments: trading noise for fast recovery

Spontaneous activation of rhodopsin without light absorption occurs at a much lower rate in rod photoreceptors and insect rhabdoms than in cones. The difference lies in the pigment molecules themselves, and has implications for the design of visual photoreceptors.     

黑翅土白蚁有翅成虫复眼的形态结构

采用组织切片法光镜下观察黑翅土白蚁Odontotermes formosanus(Shiraki)有翅成虫的复眼形态结构及光、暗适应条件下色素颗粒移动的规律。结果如下:(1)头正前方观,复眼外部形态略呈圆形。(2)有翅成虫复眼类型属于并列像眼,每只复眼约由360个小眼组成。(3)每个小眼是由1套屈光器(1个角膜和1个晶锥)、小网膜色素细胞、视杆和基细胞等几部分组成。小网膜色素细胞内均含有丰富的色素颗粒。(4)在光适应条件状态下,屈光器及视杆周围的色素颗粒主要分布在视杆部位的上侧,暗度适应条件状态时则较均匀地分布于视杆两侧上下;性别对色素颗粒分布无明显影响。     

Comparative studies of chromophore contents inside and outside the rhabdoms of arthropod compound eyes

Summary The intracellular distribution of visual pigments in the retinular cells of compound eyes of a butterfly (Papilio), a moth (Actias), a water scorpion (Ranatra), a dragonfly (Pantala), a crayfish (Procambarus), and a crab (Hemigrapsus) were investigated.The experiments measured the total surface area of rhabdom membrane in single compound eyes by light and electron microscopy, the densities of intramembrane particles in rhabdoms of compound eyes by freeze-fracture methods, and the quantities of chromophore molecules in compound eyes by high pressure liquid chromatography.Four species (moth, water scorpion, dragonfly, and crayfish) contain more than 75% of total chromophore probably as visual pigments in their rhabdoms, but the butterfly and the crab contain only 28.6% and 39.2%, respectively.The remainder of the chromophore molecules (Papilio 71.4%,Actias 5.7%,Ranatra 18.9%,Pantala 23.2%,Procambarus 24.3%, andHemigrapsus 60.8%) are supposed to exist mostly in the cytoplasm of the retinular cells. The localizing sites of such extrarhabdomeric chromophores are discussed in relation to the cytoplasmic membrane systems such as endoplasmic reticulum and lysosomal elements.In view of the results of the present experiment, it seems clear that the butterfly,Papilio, possesses a very large extra-rhabdomeric pool of chromophore that is available for rapid regeneration of visual pigment.Abbreviations HPLC high pressure liquid chromatography - ROR ratio of rhabdom to retinulae volume