Das Doppelauge vonEntomobrya muscorum Nicolet (Insecta,Collembola)

Zusammenfassung Der bodenlebende CollembolaEntomobrya muscorum hat an jeder Kopfseite ein loses Aggregat von acht Ommatidien, von denen zwei einen deutlich kleineren Linsendurchmesser aufweisen. Die sechs großen Ommatidien werden als Haupt-, die zwei kleinen als Nebenaugen bezeichnet. Beide Ommatidientypen haben verschieden gebaute Rhabdome, so daß der Komplex ein Doppelauge darstellt. Es wird der Bau der Hauptaugen geschildert. Ihr sensibler Teil setzt sich aus acht Retinulazellen zusammen, die ein offenes Rhabdom in zwei Lagen bilden. Eine zentrale sechste Zelle in der distalen Schicht weicht in ihrer Feinstruktur deutlich von den übrigen ab. Die Rhabdome beider Ommatidientypen werden miteinander verglichen, und ihre funktionelle Bedeutung vor allem in Hinblick auf eine mögliche Arbeitsteilung für eine Polarisationswahrnehmung wird erörtert. Es wird kurz die phylogenetische Bedeutung dieser Augen für die Insekten und die Collembolen selbst angeschnitten.

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The double eye ofEntomobrya muscorum nicolet (Insecta, Collembola)

Summary The ground living CollembolaEntomobrya muscorum has on each side of the head eight circular ommatidia in an arrangement characteristic of the species. Of these two have a distinctly smaller lens. The six big ommatidia are called primary eyes, the two small ones accessory eyes. The two types of ommatidia have different rhabdom structures, so justifying the term double eye. The structure of the primary eyes is described. The photosensitive part is constructed of eight retinula cells which form an open rhabdom in two layers. The central sixth cell in the distal layer is quite different from the others. The rhabdoms of both types are compared. The possible function of the accessory eyes in perception of the e-vector is discussed. The phylogenetic significance of these eyes for the Insecta and the Collembola is considered.

     

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.     

The Spectral Sensitivities of Single Receptor Cells in the Lateral, Median, and Ventral Eyes of Normal and White-Eyed Limulus

Spectral sensitivity curves can be distorted by screening pigments. We have determined whether this is true for Limulus polyphemus by determining, from receptor potentials recorded using intracellular microelectrodes, spectral sensitivity curves for normal animals and for white-eyed animals (which lack screening pigment). Our results show: (a) In median ocelli, the curve for UV-sensitive receptor cells peaks at 360 nm and does not depend on the presence of screening pigment, (b) The curve for ventral eye photoreceptors is identical to that for retinular cells from the lateral eyes of white-eyed animals and peaks at 520–525 nm. (c) In normal lateral eyes, when the stimulating light passes through screening pigment, the curve indicates relatively more sensitivity in the red region of the spectrum than does the curve for white-eyed animals. Therefore, the screening pigment is probably red-transmitting, (d) In median ocelli, the curve for visible-sensitive cells peaks at 525 nm and is approximately the same whether the ocelli are from normal or white-eyed animals. However, the curve is significantly broader than that for ventral eyes and for lateral eyes from white-eyed animals.     

Morphologische regionale differenzierungen der retina und des facettenrasters vom komplexauge der pipunculidae (Diptera : Cyclorrhapha): ein neuartiges retinales muster der fliegen (brachycera)

The retinae of the compound eyes of several species of Pipunculidae (Diptera : Cyclorrhapha), belonging to the subfamilies Chalarinae and Pipunculinae, were investigated in semithin sections in both sexes of the representative species. Whereas the boundary of the dorsal and ventral retinular cells in a mirror-image configuration is at the equator in the male, in the female it is situated in the anterior region of the eye at the level of the upper frons, and is located above the equator only in the lateral region. In the frontal view, it constitutes a concave arch to frontofacial region. The facets of the corneal lenses are strikingly enlarged in the anterior region of the eyes, compared with those in the remainder of the eye in the female. This area with the large-faceted ommatidia, was determined in more detail in total views as well as in histological preparations and compared with the eye of the male. In the frontal region, the mirror-image boundary in the retina of the female coincides exactly with the boundary between the large-faceted central and small-faceted peripheral ommatidia.By examining the dichoptic eyes of the Chalarus male, it has been demonstrated that the arcuate mirror-image boundary in the retinae of females is not associated with their dichoptic eye position. This is an example of sexual dimorphism. The retinal pattern of the female described in this paper was not found in other 37 families of the flies investigated till now. This new type of the retina of the suborder Brachycera (including Cyclorrhapha) is to be subsumed under the synapomorphic ground plans of the Pipunculidae. At the same time, it proves to be an autapomorphic characteristic of the family.     

The nauplius eye complex in 'conchostracans'(Crustacea, Branchiopoda: Laevicaudata, Spinicaudata, Cyclestherida) and its phylogenetic implications

The nauplius eye in Cyclestherida, Laevicaudata and Spinicaudata (previously collectively termed Conchostraca) consists of four cups of inverse sensory cells separated by a pigment layer and a tapetum layer. There are two lateral and two medial cups, a ventral medial cup and a posterior medial cup. The pigment and tapetum layers contain two different kinds of pigment granules, the inner pigment layer relatively large, dark (and electron dense) granules, and the outer tapetum layer light, reflective pigment granules. The presence of four cups and two different kinds of pigment granules are interpreted as autapomorphies of Phyllopoda. The position and shape of the nauplius eye in Spinicaudata is very distinct and herein interpreted as an autapomorphy of this taxon.Additional frontal eyes might be present dorsally or ventrally in varying proximity to the nauplius eye, but they have separate nerves from their sensory cells to the nauplius eye centre in the protocerebrum. Rhabdomeric structures are present in all these frontal eyes, evidencing their light sensitivity. In Lynceus biformis and L. tatei (Laevicaudata), two pairs of frontal eyes were found. In Cyclestheria hislopi (Cyclestherida), an unpaired ventral frontal eye is present. We did not find additional frontal eyes in Limnadopsis parvispinus and Caenestheriella sp. (Spinicaudata).     

Post-larval development of compound eyes and stemmata of Chaoborus crystallinus (De Geer, 1776) (Diptera : Chaoboridae): Stage-specific reconstructions within individual organs of vision

During metamorphosis, the dioptric apparatus of the larval compound eye of Chaoborus crystallinus (Diptera : Nematocera) is radically reconstructed. The thin larval cornea of the ommatidia is replaced by strongly curved corneal lenses, and the eucone larval cone is replaced by an imaginal cone of the acone type. Curvature of the future lens is already apparent in very young pupae, in which the cornea consists only of a thin epicuticle with corneal nipples. Fibrillary cuticle is secreted by cone and primary pigment cells throughout pupal development. Lens formation is accompanied by movement of the nuclei of the accessory pigment cells. The larval cone disintegrates unexpectedly late in young, images. During late pupal development, 7 cone cell projections emerge. In contrast to the dioptric apparatus, the retinula cells and rhabdom remain almost unchanged during metamorphosis. The main refractive element of the larval ommatidium appears to be the cone, while that of the imaginal ommatidium is the corneal lens. In addition to the compound eyes, the pairs of stemmata are retained during the whole post-larval development. Pupal stemmata show no structural differences from the larval stemmata. The stemmata are still present in 2-day-old images (“retained stemmata”), but the primary stemma loses its dioptric apparatus and is proximally relocated to the basal region of the compound eye. The reconstructions in the visual system of Chaoborus, which occur during ontogeny, are probably connected with the change from aquatic living larvae to aerial adults, and appear to fulfill stage-specific needs of vision.     

Ontogeny of sexual dimorphism via tissue duplication in an ostracod (Crustacea)

SUMMARY The adaptive significance of specific sexual dimorphism is well studied. However, the evolutionary history and ontogenic origins of the dimorphism are often unknown. As dimorphism represents two phenotypes generated from relatively similar genotypes, it is of interest to understand both its evolutionary and developmental/genetic underpinnings. Here, we present the first ontogenetic examination of the eyes of philomedid ostracods (Crustacea), which exhibit extremely sexually dimorphic lateral eyes. Adult male philomedids have large compound lateral eyes, whereas females have rudimentary lateral eyes. First, we show that eye dimorphism is unlikely to be due to additional genes present on a male-specific chromosome because karyotype analysis suggests philomedids are XX/XO. We then examine the ontogeny of eye development and find that in at least two species of Euphilomedes , this dimorphism is not generated solely by differences in tissue growth rates, as has been commonly shown for sexually dimorphic characters of other species. Instead, the dimorphism appears to arise during development via tissue duplication, where a single tissue becomes two, perhaps with different developmental potentials. The second eye field is only observed in male Euphilomedes , producing most of the adult eye tissue. We point out that tissue duplication is a developmental process with evolutionary implications because novel characters could evolve via alternative modification of the duplicated fields, analogous to the origin of new genes by gene duplication and alternative modification. Depending on the evolutionary history of the duplicated field, it may have either facilitated or directly caused the observed sexual dimorphism of philomedid ostracods.     

Visual fields of orb web and single line web spiders of the family Uloboridae (Arachnida,Araneida)

Summary In the family Uloboridae, web reduction is associated with changes in web monitoring posture and prosomal features. A spider must extend its first pair of legs directly forward to monitor the signal line of a reduced web. This posture is facilitated by shifts in prosomal musculature that cause reduced web uloborids to have a narrower anterior prosoma, a reduced or absent anterior eye row, and prominent posterior lateral eye tubercles. The eye tubercles and larger posterior eyes of these uloborids suggest that web reduction may also be accompanied by ocular changes that compensate for reduction of the anterior eyes by expanding the visual fields of the posterior eyes. A comparison of the visual fields of the eight-eyed, orb web species Octonoba octonaria and a four-eyed, reduced web Miagrammopes species was made to determine if this is true. Physical and optical measurements determined the visual angles of each species' eyes and the pattern of each species' visual surveillance. Despite loss of the anterior four eyes, the Miagrammopes species has a visual coverage similar to that of O. octonaria. This is due to (1) an increase in the visual field of each of the four remaining Miagrammopes eyes, accruing from an extension of the retina and an increase in the lens' rear radius of curvature, and (2) a ventral shift of each visual axis, associated with the development of an eye tubercle and an asymmetrical expansion of the retina. Miagrammopes monitor their simple webs from twigs or moss where they are vulnerable to predation. Therefore, maintenance of visual cover may enable them to detect predators in time to assume or maintain their characteristic, cryptic posture. It may also allow them to observe approaching prey and permit them to adjust web tension or prepare to jerk their webs when prey strikes.     

Das Geruchsorgan des TiefseefischesAphanopus carbo (Percomorphi,Trichiuridae)

The paper deals with the structure of the olfactory organ, its accessory parts and the forebrain in the deep-sea fishAphanopus carbo. On each side of the head only one opening leads to the olfactory chamber. The olfactory folds are arranged in a rosette-like pattern, resembling the 360°-type. Secondary folds on the main folds may serve as an enlargement of the surface of the olfactory epithelium. Most of the surface area of the olfactory folds is covered by the olfactory epithelium, indicating that the receptive area is of optimal extension. The histological structure of the olfactory epithelium is similar to that in other teleost species. The number of olfactory receptors amounts to about 5×10⁶ to 10⁷ for the single organ. Numerous secretory cells of unknown function are located within the olfactory epithelium. The olfactory chamber is enlarged by three accessory sacs: Two ethmoidal sacs and one lacrymal sac (consisting of two parts). These sacs serve as ventilation aparatus which causes a permanent water current within the olfactory chamber and between the olfactory folds. The action of the accessory sacs is induced by the splanchnokinetic. The forebrain ofAphanopus carbo is well developed; its size ranges between that of forebrains in microsmatic and macrosmatic teleost species. A detailed investigation of the forebrain is in preparation. The diagnosis of the different parts of the olfactory apparatus ofAphanopus carbo demonstrates clearly that — in addition to the eye — this sense organ is well developed (relative to that in other teleosts). This fact suggests thatAphanopus carbo is related to a group of teleost species characterized by optical and olfactory orientation mechanisms of high performance.     

A new Pseudophoxinus (Teleostei,Cyprinidae) species from Asi River Drainage (Turkey)

Pseudophoxinus turani sp. n. is described from the İncesu Spring (Hassa-Hatay) drainage of Asi River, Turkey. It is distinguished from other Eastern Mediterranean Region Pseudophoxinus species by a combination of characters: lateral line incomplete, with 12–25 (commonly 16–21) perforated scales and 38–46+2-3 scales in lateral series (commonly 41–44+2-3); 10–11 scale rows between the lateral line and dorsal-fin origin; 3–4 scale rows between the lateral line and the pelvic–fin origin; dorsal fin with 7½ branched rays; anal fin commonly with 7½ branched rays; 8-11gill rakers on the first branchial arch; dorsal profile markedly convex with marked hump at the nape, ventral profile less convex than dorsal profile; a small, irregular, black blotch on the base of the caudal fin; mouth terminal, with slightly distinct chin, its corner not reaching vertical through anterior margin of eye; snout somewhat long, with rounded tip; and its length greater than eye diameter.     

On Levymanus,a remarkable new spider genus from?Israel,with notes on the Chediminae (Araneae,Palpimanidae)

Levymanus gershomi gen. n. et sp. n., is described from southern Israel. The eye arrangement and structure of the male palp indicate that this genus belongs to Chediminae Simon, 1893. Levymanus gen. n. differs from other chedimine genera by its unusually long and slender legs, an elongate body, a unique shape of the bipartite thoracic fovea, reduced leg scopulae, smaller spinnerets, and other characters, which are presumably apomorphic. We propose two taxonomic changes: 1) based on widely spaced lateral eyes the Western African genus Badia Roewer, 1961 is transferred from Chediminae to Palpimaninae, and 2) Fernandezina gyirongensis Hu & Li, 1987 from China, based on palpal morphology, is transferred to the Asian genus Steriphopus Simon, 1887 for a new combination Steriphopus gyirongensis (Hu & Li, 1987) comb. n.     

Fine structure of the compound eyes of the midwater amphipod Phronima in relation to behavior and habitat

Pelagic amphipods belonging to the genus Phronima have four compound eyes; two lateral eyes and two large transparent medial eyes which comprise the entire top of the head. The eyes are structurally similar but the crystalline cones of the medial eyes are more than twenty times as long as those of the lateral eyes, reaching 5 mm in a large animal. The dioptric system of each ommatidium consists of an unfaceted cornea, a layer of hypodermal cells, two rudimentary cone cells, two cells which surround and form the crystalline cone, and the cone itself. The cone and its surrounding cells penetrate the layer of accessory pigment cells which surrounds the retina. The fused rhabdom is formed by the five retinula cells but is separated from them by an extracellular palisade which is crossed by bridges. The retinula cell nuclei lie proximal to the basement membrane. Further proximally the bundle of retinula cell axons is crossed by a second basement membrane, which surrounds each axon with a collar. Medial and lateral eyes on each side of the head share a common lamina. The medial eyes of Phronima appear to be a solution to the problem of remaining inconspicuous to predators while still maintaining sensitivity and resolution.     

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.     

Persisting stemma neuropils in Chaoborus crystallinus (Diptera: Chaoboridae): Development and evolution of a bipartite visual system

Stemmata or “larval” eyes are of crucial importance for the understanding of the evolution and ontogeny of the hexapod's main visual organs, the compound eyes. Using classical neuroanatomical techniques, I showed that the persisting stemmata of Chaoborus imagos are connected to persisting stemma neuropils neighboring the first and second order neuropils of the compound eyes, and therefore also the imago possesses a stemma lamina and medulla closely associated with the architecture and the developmental pattern of those of the compound eyes. The findings are compared with other arthropods, e.g. accessory lateral eyes in Amandibulata and Myriapoda, suggesting some ancestral rather than derived character states. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Sperm morphology and phylogeny of lagriids (Coleoptera,Tenebrionidae)

The systematics of tenebrionids remain unclear, principally at the subfamily level, as is the case of the Lagriinae. Considering that sperm morphology has contributed to the various insect group phylogenies, in this work we describe the structure and ultrastructure of these cells in Lagria villosa. Sperm in this species exhibit a strong morphological similarity to those of Tenebrio molitor and Tribolium castaneum, the only two species of Tenebrionidae with previously described sperm. In tenebrionids, the flagellar components offer good diagnostic characters, e.g. the symmetry of mitochondrial derivatives in L. villosa differentiates sperm of this species from those of Te. molitor and Tr. castaneum. However, the lateral association of the nucleus with flagellar components, the form of accessory bodies, and the presence of material connecting the axoneme to the accessory bodies and mitochondrial derivatives indicate that the three species form a sister group. Therefore, the sperm morphology of L. villosa support lagriid beetles as a subfamily (Lagriinae) of Tenebrionidae.     

Fine structure of the eyes of orb-weavers,Argiope amoena L. Koch (Aranea: Argiopidae)

The anterolateral eye, the posterolateral eye and the posteromedial eye of the web-building spider, Argiope amoena have been examined by light and electron microscopy. The dioptric apparatus of all three eyes is similar in structure, and consists of a cornea, a lens and a vitreous body. The retina contains monopolar receptor cells, the cell bodies of which are present beneath the vitreous body in all three eyes. Proximal processes of the receptor cells form rhabdoms beneath the cell body layer and then extend toward the first optic glomerulus as an ocellar nerve. Two distinct patterns of retinal organization are present in the three eyes. In one type the rhabdomic layer of the retina is backed by a pigmented layer. In the other type the rhabdomic layer is backed by a tapetal reflecting layer. Rhabdomic structure and cytoplasmic inclusions of the receptor cells differ greatly between the two types. The anterolateral eye possesses a single type of retina with the rhabdoms backed by the tapetum. Both the posterolateral and the posteromedial eye are similar in structure, each possessing beneath the common dioptric apparatus retinae of both types.     

Sinocyclocheilus zhenfengensis,a new cyprinid species (Pisces: Teleostei) from Guizhou Province,Southwest China

Sinocyclocheilus zhenfengensis sp. nov. is a new species from an underground water environment in a karst cave located in Beipanjiang River of the Pearl River Basin in Shuangrufeng (Double Breasts Hill) Scenic, Etian Village, Zhexiang Township, Zhenfeng County, Guizhou Province, Southwest China. The new species has eyes, its body is completely scaled, an anteriorly directed bulge has formed on the back of its head, and a long pectoral fin exceeds the pelvic fin insertion. The body of the new species is apparently similar to that of S. bicornutus, but differs in scale rows above the lateral line, scale rows below the lateral line, snout length, and eye diameter. Phylogenetic reconstruction of the cytochrome b gene using the maximum parsimony (MP) and Bayesian inference (BI) methods indicates that S. zhenfengensis is closely related to S. bicornutus, S. tianeensis, S. jiuxuensis, and S. altishoulderus; however, it differs in terms of eye size, lateral line scales, scale rows above and below the lateral line.