Median Ocellus Research Articles

Drepanolejeunea laciniata(Lejeuneaceae), a New Species from Northern Thailand

Abstract Drepanolejeunea laciniata Qiong He et R.L. Zhu sp. nov., is described and illustrated from Doi Inthanon National Park, Chom Thong District, Chiang Mai Province, northern Thailand. It is similar to Drepanolejeunea pulla (Mitt.) Grolle and D. erecta (Steph.) Grolle, but differs mainly in its large leaf lobule that is 1/2–2/3 as long as the leaf lobe, the free lateral lobular margin proximal to the notch bordered by 9–13 rectangular cells, the obovate perianth with dense apical laciniae, and usual presence of median ocelli in the leaf lobe.

Wiring a Periscope – Ocelli, Retinula Axons, Visual Neuropils and the Ancestrality of Sea Spiders

The Pycnogonida or sea spiders are cryptic, eight-legged arthropods with four median ocelli in a ‘periscope’ or eye tubercle. In older attempts at reconstructing phylogeny they were Arthropoda incertae sedis, but recent molecular trees placed them as the sister group either to all other euchelicerates or even to all euarthropods. Thus, pycnogonids are among the oldest extant arthropods and hold a key position for the understanding of arthropod evolution. This has stimulated studies of new sets of characters conductive to cladistic analyses, e.g. of the chelifores and of the hox gene expression pattern. In contrast knowledge of the architecture of the visual system is cursory. A few studies have analysed the ocelli and the uncommon “pseudoinverted” retinula cells. Moreover, analyses of visual neuropils are still at the stage of Hanström's early comprehensive works. We have therefore used various techniques to analyse the visual fibre pathways and the structure of their interrelated neuropils in several species. We found that pycnogonid ocelli are innervated to first and second visual neuropils in close vicinity to an unpaired midline neuropil, i.e. possibly the arcuate body, in a way very similar to ancestral euarthropods like Euperipatoides rowelli (Onychophora) and Limulus polyphemus (Xiphosura). This supports the ancestrality of pycnogonids and sheds light on what eyes in the pycnogonid ground plan might have ‘looked’ like. Recently it was suggested that arthropod eyes originated from simple ocelli similar to larval eyes. Hence, pycnogonid eyes would be one of the early offshoots among the wealth of more sophisticated arthropod eyes.

The organization of honeybee ocelli: Regional specializations and rhabdom arrangements

We have re-investigated the organization of ocelli in honeybee workers and drones. Ocellar lenses are divided into a dorsal and a ventral part by a cusp-shaped indentation. The retina is also divided, with a ventral retina looking skywards and a dorsal retina looking at the horizon. The focal plane of lenses lies behind the retina in lateral ocelli, but within the dorsal retina in the median ocellus of both workers and drones. Ventral retinula cells are ca. 25 μm long with dense screening pigments. Dorsal retinula cells are ca. 60 μm long with sparse pigmentation mainly restricted to their proximal parts. Pairs of retinula cells form flat, non-twisting rhabdom sheets with elongated, straight, rectangular cross-sections, on average 8.7 μm long and 1 μm wide. Honeybee ocellar rhabdoms have shorter and straighter cross-sections than those recently described in the night-active bee Megalopta genalis. Across the retina, rhabdoms form a fan-shaped pattern of orientations. In each ocellus, ventral and dorsal retinula cell axons project into two separate neuropils, converging on few large neurons in the dorsal, and on many small neurons in the ventral neuropil. The divided nature of the ocelli, together with the particular construction and arrangement of rhabdoms, suggest that ocelli are not only involved in attitude control, but might also provide skylight polarization compass information.

Description of the Immature Stages of Bittacus planus Cheng (Mecoptera: Bittacidae) with Notes on Its Biology

The immature stages of Bittacus planus Cheng, 1949, including egg, four larval instars, and pupa, are described and illustrated for the first time. The egg is spherical, with two types of micropyles. The larva is eruciform and peripneustic, bearing eight pairs of prolegs on the abdomen. A pair of compound eyes and a median ocellus are present on the head. Each segment of the abdomen has rows of protuberances bearing setiferous clavate setae. The pupa is exarate and decticous. The species is univoltine, overwintering as diapausing eggs on the ground. Larval duration, including a prepupa of 10 days, lasts about 30 days. Pupation occurs in a soil cell, ranging from 10 to 15 days. The roles of larval compound eyes and the ocellus in the phylogeny of Holometabola are briefly discussed.

Directional Selectivity in the Simple Eye of an Insect

Among other sensory modalities, flight stabilization in insects is performed with the aid of visual feedback from three simple eyes (ocelli). It is thought that each ocellus acts as a single wide-field sensor that detects changes in light intensity. We challenge this notion by providing evidence that, when light-adapted, the large retinal L-neurons in the median ocellus of the dragonfly respond in a directional way to upward moving bars and gratings. This ability is pronounced under UV illumination but weak or nonexistent in green light and is optimal at angular velocities of approximately 750 degrees s(-1). Using a reverse-correlation technique, we analyze the functional organization of the receptive fields of the L-neurons. Our results reveal that L-neurons alter the structure of their linear spatiotemporal receptive fields with changes in the illuminating wavelength, becoming more inseparable and directional in UV light than in green. For moving bars and gratings, the strength of directionality predicted from the receptive fields is consistent with the measured values. Our results strongly suggest that, during the day, the retinal circuitry of the dragonfly median ocellus performs an early linear stage of motion processing. The likely advantage of this computation is to enhance pitch control.

Form vision in the insect dorsal ocelli: An anatomical and optical analysis of the dragonfly median ocellus

Previous work has suggested that dragonfly ocelli are specifically adapted to resolve horizontally extended features of the world, such as the horizon. We investigate the optical and anatomical properties of the median ocellus of Hemicordulia tau and Aeshna mixta to determine the extent to which the findings support this conclusion. Dragonfly median ocelli are shown to possess a number of remarkable properties: astigmatism arising from the elliptical shape of the lens is cancelled by the bilobed shape of the inner lens surface, interference microscopy reveals complex gradients of refractive index within the lens, the morphology of the retina results in zones of high acuity, and the eye has an exceedingly high sensitivity for a diurnal terrestrial invertebrate. It is concluded that dragonfly ocelli employ a number of simple, yet elegant, anatomical and optical strategies to ensure high sensitivity, fast transduction speed, wide fields of views and a modicum of spatial resolving power.

The mapping of visual space by dragonfly lateral ocelli

We study the extent to which the lateral ocelli of dragonflies are able to resolve and map spatial information, following the recent finding that the median ocellus is adapted for spatial resolution around the horizon. Physiological optics are investigated by the hanging-drop technique and related to morphology as determined by sectioning and three-dimensional reconstruction. L-neuron morphology and physiology are investigated by intracellular electrophysiology, white noise analysis and iontophoretic dye injection. The lateral ocellar lens consists of a strongly curved outer surface, and two distinct inner surfaces that separate the retina into dorsal and ventral components. The focal plane lies within the dorsal retina but proximal to the ventral retina. Three identified L-neurons innervate the dorsal retina and extend the one-dimensional mapping arrangement of median ocellar L-neurons, with fields of view that are directed at the horizon. One further L-neuron innervates the ventral retina and is adapted for wide-field intensity summation. In both median and lateral ocelli, a distinct subclass of descending L-neuron carries multi-sensory information via graded and regenerative potentials. Dragonfly ocelli are adapted for high sensitivity as well as a modicum of resolution, especially in elevation, suggesting a role for attitude stabilisation by localization of the horizon.

Comparing injection, feeding and topical application methods for treatment of honeybees with octopamine

Entomologists have used a range of techniques to treat insects with neuroactive compounds, but it is not always clear whether different treatment methods are equally effective in delivering a compound to a target organ. Here, we used five different techniques to treat honeybees with 3H-octopamine ( 3H-OA), and analysed the distribution of the 3H radiolabelled compound within different tissues and how it changed over time. All treatment methods, including injection of the median ocellus, resulted in 3H-OA detection in all parts of the honeybee. Injection through the median ocellus was the most effective method for delivering 3H-OA to the brain. Topical application of 3H-OA dissolved in dimethylformamide (dMF) to the thorax was as effective as thoracic injections of 3H-OA in delivering 3H-OA to the brain, but topical applications to the abdomen were less so. Most of the 3H-OA applied topically remained associated with the cuticle and the tissues of the body segment to which it had been applied. For all treatment methods, 3H-OA was rapidly lost from the brain and head capsule, and accumulated in the abdomen. Our findings demonstrate the value of thoracic topical treatment with compounds dissolved in dMF as an effective non-invasive method for short-term, systemic pharmacological treatments.

Structure and development of onychophoran eyes: What is the ancestral visual organ in arthropods?

Scarce and controversial information on visual organs and their innervation in Onychophora currently do not allow a thorough comparison with Euarthropoda. Therefore, this study sets out to provide additional data on the architecture and morphogenesis of the onychophoran visual system and to explore similarities and differences between the visual organs of onychophorans and other arthropods. Based on the new data for Epiperipatus biolleyi (Peripatidae) and Metaperipatus blainvillei (Peripatopsidae), it is suggested that the compound eyes represent an autapomorphy of Euarthropoda since similarities with the onychophoran eyes are weak or absent. Instead, the innervation from a central rather than lateral part of the brain, the presence of only one (paired or unpaired) visual center, and a similar ontogenetic origin from an ectodermal groove rather than a proliferation zone suggest homology between the onychophoran eyes and the median ocelli of euarthropods. In conclusion, I suggest that the last common ancestor of arthropods bore only one pair of ocellus-like visual organs that were modified in several arthropod lineages. This hypothesis is supported by recent paleontological data.

The mapping of visual space by identified large second-order neurons in the dragonfly median ocellus

In adult dragonflies, the compound eyes are augmented by three simple eyes known as the dorsal ocelli. The outputs of ocellar photoreceptors converge on relatively few second-order neurons with large axonal diameters (L-neurons). We determine L-neuron morphology by iontophoretic dye injection combined with three-dimensional reconstructions. Using intracellular recording and white noise analysis, we also determine the physiological receptive fields of the L-neurons, in order to identify the extent to which they preserve spatial information. We find a total of 11 median ocellar L-neurons, consisting of five symmetrical pairs and one unpaired neuron. L-neurons are distinguishable by the extent and location of their terminations within the ocellar plexus and brain. In the horizontal dimension, L-neurons project to different regions of the ocellar plexus, in close correlation with their receptive fields. In the vertical dimension, dendritic arborizations overlap widely, paralleled by receptive fields that are narrow and do not differ between different neurons. These results provide the first evidence for the preservation of spatial information by the second-order neurons of any dorsal ocellus. The system essentially forms a one-dimensional image of the equator over a wide azimuthal area, possibly forming an internal representation of the horizon. Potential behavioural roles for the system are discussed.

Involvement of α-bungarotoxin-sensitive nicotinic receptors in long-term memory formation in the honeybee ( Apis mellifera)

In the honeybee Apis mellifera, multiple-trial olfactory conditioning of the proboscis extension response specifically leads to long-term memory (LTM) which can be retrieved more than 24 h after learning. We studied the involvement of nicotinic acetylcholine receptors in the establishment of LTM by injecting the nicotinic antagonists mecamylamine (1 mM), α-bungarotoxin (α-BGT, 0.1 mM) or methyllycaconitine (MLA, 0.1 mM) into the brain through the median ocellus 20 min before or 20 min after multiple-trial learning. The retention tests were performed 1, 3, and 24 h after learning. Pre-training injections of mecamylamine induced a lower performance during conditioning but had no effect on LTM formation. Post-training injections of mecamylamine did not affect honeybees’ performances. Pre-training injections of MLA or post-training injection of α-BGT specifically induced LTM impairment whereas acquisition as well as memory retrieval tested 1 or 3 h after learning was normal. This indicates that brain injections of α-BGT and MLA did not interfere with learning or medium-term memory. Rather, these blockers affect the LTM. To explain these results, we advance the hypothesis that honeybee α-BGT-sensitive acetylcholine receptors are also sensitive to MLA. These receptors could be essential for triggering intracellular mechanisms involved in LTM. By contrast, medium-term memory is not dependent upon these receptors but is affected by mecamylamine.

A Spatiotemporal White Noise Analysis of Photoreceptor Responses to UV and Green Light in the Dragonfly Median Ocellus

Adult dragonflies augment their compound eyes with three simple eyes known as the dorsal ocelli. While the ocellar system is known to mediate stabilizing head reflexes during flight, the ability of the ocellar retina to dynamically resolve the environment is unknown. For the first time, we directly measured the angular sensitivities of the photoreceptors of the dragonfly median (middle) ocellus. We performed a second-order Wiener Kernel analysis of intracellular recordings of light-adapted photoreceptors. These were stimulated with one-dimensional horizontal or vertical patterns of concurrent UV and green light with different contrast levels and at different ambient temperatures. The photoreceptors were found to have anisotropic receptive fields with vertical and horizontal acceptance angles of 15° and 28°, respectively. The first-order (linear) temporal kernels contained significant undershoots whose amplitudes are invariant under changes in the contrast of the stimulus but significantly reduced at higher temperatures. The second-order kernels showed evidence of two distinct nonlinear components: a fast acting self-facilitation, which is dominant in the UV, followed by delayed self- and cross-inhibition of UV and green light responses. No facilitatory interactions between the UV and green light were found, indicating that facilitation of the green and UV responses occurs in isolated compartments. Inhibition between UV and green stimuli was present, indicating that inhibition occurs at a common point in the UV and green response pathways. We present a nonlinear cascade model (NLN) with initial stages consisting of separate UV and green pathways. Each pathway contains a fast facilitating nonlinearity coupled to a linear response. The linear response is described by an extended log-normal model, accounting for the phasic component. The final nonlinearity is composed of self-inhibition in the UV and green pathways and inhibition between these pathways. The model can largely predict the response of the photoreceptors to UV and green light.

Antennal tactile learning in the honeybee: Effect of nicotinic antagonists on memory dynamics

Restrained worker honeybees ( Apis mellifera L.) are able to learn to associate antennal-scanning of a metal plate with a sucrose reinforcement delivered to the mouthparts. Learning occurs reliably in a single association of the two sensory stimuli. The involvement of nicotinic pathways in memory formation and retrieval processes was tested by injecting, into the whole brain through the median ocellus, either mecamylamine (0.6 μg per bee) or α-bungarotoxin (2.4 ng per bee). Saline served as a control. Mecamylamine injected 10 min before the retrieval test impairs the retention level tested 3 h and 24 h after single- or multi-trial learning. Retrieval tests performed at various times after the injection show that the blocking effect of mecamylamine lasts about 1 h. The drug has no effect on the reconsolidation or extinction processes. Mecamylamine injected 10 min before conditioning impairs single-trial learning but has no effect on five-trial learning and on the consolidation process. By contrast, α-bungarotoxin only impairs the formation of long-term memory (24 h) induced by the five-trial learning and has no effect on medium-term memory (3 h), on single-trial learning or on the retrieval process. Hence, owing to previous data, at least two kinds of nicotinic receptors seem to be involved in honeybee memory, an α-bungarotoxin-sensitive and an α-bungarotoxin-insensitive receptor. Our results extend to antennal mechanosensory conditioning the role of the cholinergic system that we had previously described for olfactory conditioning in the honeybee. Moreover, we describe here in this insect a pharmacological dissociation between α-bungarotoxin sensitive long-term memory and α-bungarotoxin insensitive medium-term memory, the last one being affected by mecamylamine.

First record of the genus Amauronematus Konow (Hymenoptera: Symphyta: Tenthredinidae: Nematinae) with three new species from the Oriental region

With three new species, the genus Amauronematus Konow is reported for the first time from India and thus it has extended its faunistic limits to the Oriental region. Described and illustrated in this paper as new are A. dilatus, A. kulwantae and A. tesa. A key for separation of the Oriental species is provided. ABBREVIATIONS AWMT - Apical width of metatibia; EL - Eye length; IATS - Inner apical tibial spur; IDMO - Interoocular distance at level of median ocellus; LID - Lower interocular distance; MB - Metabasitarsus; OATS - Outer apical tibial spur; OCL - Ocellooccipital line; POL - Postocellar line; UCL - Oculooccipital line; UOL - Oculoocellar lineAfter being erected by Konow (1890) the genus Amauronematus was so far recorded from the Holarctic region only. It is based on Nematus fallax Lepeletier as its type species designated by Rohwer (1911). However, with the discovery of three new species from India, this genus has extended its faunistic limits to the Oriental region. The new species are described and illustrated on a uniform pattern.

Anisotropic imaging in the dragonfly median ocellus: a matched filter for horizon detection

It is suggested that the dragonfly median ocellus is specifically adapted to detect horizontally extended features rather than merely changes in overall intensity. Evidence is presented from the optics, tapetal reflections and retinal ultrastructure. The underfocused ocelli of adult insects are generally incapable of resolving images. However, in the dragonfly median ocellus the geometry of the lens indicates that some image detail is present at the retina in the vertical dimension. Details in the horizontal dimension are blurred by the strongly astigmatic lens. In the excised eye the image of a point source forms a horizontal streak at the level of the retina. Tapetal reflections from the intact eye show that the field of view is not circular as in most other insects but elliptical with the major axis horizontal, and that resolution in the vertical direction is better than in the horizontal. Measurements of tapetal reflections in locust ocelli confirm their visual fields are wide and circular and their optics strongly underfocused. The ultrastructure suggests adaptation for resolution, sensitivity and a high metabolic rate, with long, widely separated rhabdoms, retinulae cupped by reflecting pigment, abundant tracheoles and mitochondria, and convoluted, amplified retinula cell plasma membranes.

Phase-shifting and Entrainment of a Circadian Rhythm in Limulus polyphemus by Ocular and Extraocular Photoreceptors

Illumination of just the telson of the horseshoe crab Limulus polyphemus produced increasing phase-shifts in the circadian rhythm of the ERG amplitudes of the compound lateral eye with increasing length of exposure to light, up to 12 h for 10-h exposures, equal to the phase-shifts produced by illumination of the whole animal or of the anterior photoreceptors alone. Periodic illumination of the anterior photoreceptors or of just the telson entrained the lateral eye rhythm, which reliably re-entrained when the phase of the illumination was shifted by ± 6 h. Illumination for 10 h of one compound eye alone or of the median ocelli alone produced small phase-shifts. Periodic illumination of just one compound lateral eye could be arranged to entrain the rhythm of the other compound eye, but the rhythm did not always re-entrain following a ± 6 h phase-shift in illumination. The apparent redundancy in circadian photoreceptors may be useful to Limulus in dealing with the various photic environments which it inhabits.

Meliponini neotropicais: o gênero camargoia moure, 1989 (apinae, apidae, hymenoptera)

Camargoia nordestina, sp. n., is described from the dry valleys of Piaui state, NE Brazil. It differs from C. camargoi by the yellow maculations ofparaocular areas ending at the level of superior tangent of antennal sockets, basitarsi II and tibiae and basitarsi III blackish, microtrichiae of anterior wing uniformly fuscous and the median ocellus above the orbital tangent. C. camargoi has the yellow paraocular maculation reaching the middle of the front; the basitarsi and tibiae are yellowish, only slightly darkish on apex; the microtrichiae fuscous at the anterior wing apex and yellowish at basis, median ocellus reaching the level of orbital tangent. From C. pilicornis, it differs by the very shorter setae of the antennal scape with about 1/4 of its diameter. C. pilicornis has the setae longer or as long as the diameter of scape. The subterraneous nest of, C. nordestina is described in detail. Some considerations about possible convergency of architectonic attributes . among the nests of Camargoia, Geotrigona and Schwarziana are presented. The result of a cladistic analysis suggests Camargoia as sister-group of Ptilotrigona.

The eyes of a “nobody”,Anoplodactylus petiolatus (Pantopoda; Anoplodactylidae)

The fine structure of the four ocelli ofAnoplodactylus petiolatus was examined using serial longitudinal and transversal sections of the eye hill. Each pigment cup ocellus is composed of a (planconvex) cuticular lens, lens forming hypodermal cells, inverse retinula cells with latticed rhabdom and surrounding tapetum and pigment layers. Within the retinula cells a distal “vitreous” zone, a nucleus zone and a proximal rhabdomeric zone can be distinguished. Retinula cell axons originate proximally. The tapetum cells contain several layers of reflecting crystals. Distally, they have a common microvillous region. The intraretinal “vitreous” zone contains glycogen-like particles in the centre and rough ER in the periphery. Contrary to other Pantopoda vitreous cells, a praeretinal membrane and a vertical lens groove have not been observed inAnoplodactylus. While the presence of four (median) ocelli appears to be a primitive characteristic, the inverse orientation of the retinula cells in combination with a tapetum lucidum represents a highly derived characteristic among arthropod median eyes.

The morphology and physiology of a "mini-ommatidium" in the median optic nerve of Limulus polyphemus.

Examination of the Limulus median optic nerve with low-magnification light microscopy allows clear visualization of an ultraviolet-sensitive mini-ommatidium enshrouded by pigment cells, glial cells, and guanophores. Serial 1-micron sections of median optic nerves containing mini-ommatidia revealed the presence of a single, heavily pigmented photoreceptor (retinular) cell and a single, unpigmented arhabdomeric cell. Computer-assisted serial reconstructions from 1-micron sections confirmed the presence of two cells, each bearing a nucleus, and two axons leaving the mini-ommatidium. The retinular cell is morphologically similar to retinular cells from the median and lateral eyes. Its rhabdomere appears to be a continuous sheet of microvilli with much infolding. The structure of the arhabdomeric cell is nearly identical to those found in the median ocellus. As in other photoreceptors in Limulus, the retinular cell of the mini-ommatidium is innervated by efferent fibers from the brain. Each mini-ommatidium generates a single train of nerve impulses in response to light, presumably from the arhabdomeric cell. Measurement of the spectral sensitivity of the mini-ommatidium based upon a constant-response criterion indicated that the retinular cell is maximally sensitive to near ultraviolet light with lambda max = 380 nm. Comparison of intensity-response functions revealed that those of the mini-ommatidium are significantly steeper than those of the ocellus almost certainly as the result of neural processing in the ocellus which is absent in the mini-ommatidium.

Opsins from the lateral eyes and ocelli of the horseshoe crab, Limulus polyphemus.

cDNA clones encoding opsins from the lateral eyes and median ocelli of the horseshoe crab, Limulus polyphemus, were isolated from cDNA libraries. The opsin cDNAs obtained from the lateral eye and ocellar libraries code for deduced proteins with 376 amino acids. The two cDNAs are 96% identical at the nucleic acid level, differing primarily at the 3' untranslated region, and are apparently the products of two separate genes. The deduced opsin proteins are 99% identical to each other, differing at only 5 amino acids. The opsins encoded by these cDNAs are most likely the protein moiety of the visible-wavelength rhodopsins in this animal. In the lateral eye, expression of the opsin gene is restricted to the photoreceptor cells of the ommatidia. Comparisons with opsins of other species show that the Limulus opsin proteins are most similar (53% identity) to the opsin from the R1-6 photoreceptors of flies.