Arcuate Body Research Articles

The visual pathway in sea spiders (Pycnogonida) displays a simple serial layout with similarities to the median eye pathway in horseshoe crabs

BackgroundPhylogenomic studies over the past two decades have consolidated the major branches of the arthropod tree of life. However, especially within the Chelicerata (spiders, scorpions, and kin), interrelationships of the constituent taxa remain controversial. While sea spiders (Pycnogonida) are firmly established as sister group of all other extant representatives (Euchelicerata), euchelicerate phylogeny itself is still contested. One key issue concerns the marine horseshoe crabs (Xiphosura), which recent studies recover either as sister group of terrestrial Arachnida or nested within the latter, with significant impact on postulated terrestrialization scenarios and long-standing paradigms of ancestral chelicerate traits. In potential support of a nested placement, previous neuroanatomical studies highlighted similarities in the visual pathway of xiphosurans and some arachnopulmonates (scorpions, whip scorpions, whip spiders). However, contradictory descriptions of the pycnogonid visual system hamper outgroup comparison and thus character polarization.ResultsTo advance the understanding of the pycnogonid brain and its sense organs with the aim of elucidating chelicerate visual system evolution, a wide range of families were studied using a combination of micro-computed X-ray tomography, histology, dye tracing, and immunolabeling of tubulin, the neuropil marker synapsin, and several neuroactive substances (including histamine, serotonin, tyrosine hydroxylase, and orcokinin). Contrary to previous descriptions, the visual system displays a serial layout with only one first-order visual neuropil connected to a bilayered arcuate body by catecholaminergic interneurons. Fluorescent dye tracing reveals a previously reported second visual neuropil as the target of axons from the lateral sense organ instead of the eyes.ConclusionsGround pattern reconstruction reveals remarkable neuroanatomical stasis in the pycnogonid visual system since the Ordovician or even earlier. Its conserved layout exhibits similarities to the median eye pathway in euchelicerates, especially in xiphosurans, with which pycnogonids share two median eye pairs that differentiate consecutively during development and target one visual neuropil upstream of the arcuate body. Given multiple losses of median and/or lateral eyes in chelicerates, and the tightly linked reduction of visual processing centers, interconnections between median and lateral visual neuropils in xiphosurans and arachnopulmonates are critically discussed, representing a plausible ancestral condition of taxa that have retained both eye types.

Anatomy of the Nervous System in Chelifer cancroides (Arachnida: Pseudoscorpiones) with a Distinct Sensory Pathway Associated with the Pedipalps.

Simple SummaryMost arthropods (uniting animals such as the chelicerates, e.g., spiders and their kin, as well as millipedes, centipedes, crustaceans, and insects) have distinct sensory appendages at the second head segment, the so-called antennae. The Arachnida (e.g., spiders and scorpions) do not possess antennae, but have evolved highly specialized sensory organs on different body regions. However, very limited information is available concerning pseudoscorpions (false scorpions). These animals do not seem to possess such specialized structures, but show dominant, multifunctional appendages prior to the first walking leg, called pedipalps. Here, we investigate the neuronal pathway of these structures as well as general aspects of the nervous system. We describe new details of typical arthropod brain compartments, such as the arcuate body and a comparatively small mushroom body. Neurons associated with the pedipalps terminate in two regions in the central nervous system of characteristic arrangement: a glomerular and a layered center, which we interpret as a chemo- and a mechanosensory center, respectively. The centers, which fulfill the same function in other animals, show a similar arrangement. These similarities in the sensory systems of different evolutionary origin have to be interpreted as functional prerequisites. Identifying these similarities helps to understand the general functionality of sensory systems, not only within arthropods.Many arachnid taxa have evolved unique, highly specialized sensory structures such as antenniform legs in Amblypygi (whip spiders), for instance, or mesosomal pectines in scorpions. Knowledge of the neuroanatomy as well as functional aspects of these sensory organs is rather scarce, especially in comparison to other arthropod clades. In pseudoscorpions, no special sensory structures have been discovered so far. Nevertheless, these animals possess dominant, multifunctional pedipalps, which are good candidates for being the primary sensory appendages. However, only little is known about the anatomy of the nervous system and the projection pattern of pedipalpal afferents in this taxon. By using immunofluorescent labeling of neuronal structures as well as lipophilic dye labeling of pedipalpal pathways, we identified the arcuate body, as well as a comparatively small mushroom body, the latter showing some similarities to that of Solifugae (sun spiders and camel spiders). Furthermore, afferents from the pedipalps terminate in a glomerular and a layered neuropil. Due to the innervation pattern and structural appearance, we conclude that these neuropils are the first integration centers of the chemosensory and mechanosensory afferents. Within Arthropoda, but also other invertebrates or even vertebrates, sensory structures show rather similar neuronal arrangement. Thus, these similarities in the sensory systems of different evolutionary origin have to be interpreted as functional prerequisites of the respective modality.

Behavioral Attributes of Social Groups Determine the Strength and Direction of Selection on Neural Investment

The evolution of social systems can place novel selective forces on investment in expensive neural tissue by changing cognitive demands. Previous hypotheses about the impact of sociality on neural investment have received equivocal support when tested across diverse taxonomic groups and social structures. We suggest previous models for social behavior-brain relationships have overlooked important variation in social groups. Social groups vary significantly in structure and function, and the specific attributes of a social group may be more relevant to setting cognitive demands than sociality in general. We have identified intragroup competition, relationship differentiation, information sharing, dominance hierarchies, and task specialization and redundancy as attributes of social behavior which may impact selection for neural investment, and outline how variation in these attributes can result in increased or decreased neural investment with transitions to sociality in different taxa. Finally, we test some of the predictions generated using this framework in a phylogenetic comparison of neural tissue investment in Anelosimus social spiders. Social Anelosimus spiders engage in cooperative prey capture and brood care, which allows for individual redundancy in the completion of these tasks. We hypothesized that in social spider species, the presence of redundancy would reduce selection for individual neural investment relative to subsocial species. We found that social species had significantly decreased investment in the arcuate body, the cognitive center of the spider brain, supporting our predictions. Future comparative tests of brain evolution in social species should account for the special behavioral characteristics that accompany social groups in the subject taxa.

Outsourcing a visual neuropil – The central visual system of the median eyes of Galeodes granti Pocock, 1903 (Arachnida: Solifugae)

Only a few studies have examined the central visual system of Solifugae until now. To get new insights suitable for phylogenetic analysis we studied the R-cell (or retinula cell) projections and visual neuropils of Galeodes granti using various methods.G. granti possesses large median eyes and rudimentary lateral eyes. In this study, only the R-cells and neuropils of the median eyes were successfully stained. The R-cells terminate in two distinct visual neuropils. The first neuropil is located externally to the protocerebrum directly below the retina, the second neuropil lies in the cell body rind of the protocerebrum, and immediately adjacent is the arcuate body.This layout of the median eye visual system differs from Arachnopulmonata (Scorpiones + Tetrapulmonata). However, there are several similarities with Opiliones. In both, (1) the R-cells are connected to a first and second visual neuropil and not to any other region of the brain, (2) the first neuropil is not embedded in the cell body rind of the protocerebrum, it is rather external to the protocerebrum, (3) the second visual neuropil is embedded in the cell body rind, and (4) the second neuropil abuts the arcuate body. These findings may provide important new characters for the discussion on arachnid phylogeny.

Visual pathways in the brain of the jumping spider Marpissa muscosa.

Some animals have evolved task differentiation among their eyes. A particular example is spiders, where most species have eight eyes, of which two (the principal eyes) are used for object discrimination, whereas the other three pairs (secondary eyes) detect movement. In the ctenid spider Cupiennius salei, these two eye types correspond to two visual pathways in the brain. Each eye is associated with its own first- and second-order visual neuropil. The second-order neuropils of the principal eyes are connected to the arcuate body, whereas the second-order neuropils of the secondary eyes are linked to the mushroom body. We explored the principal- and secondary eye visual pathways of the jumping spider Marpissa muscosa, in which size and visual fields of the two eye types are considerably different. We found that the connectivity of the principal eye pathway is the same as in C. salei, while there are differences in the secondary eye pathways. In M. muscosa, all secondary eyes are connected to their own first-order visual neuropils. The first-order visual neuropils of the anterior lateral and posterior lateral eyes are connected with a second-order visual neuropil each and an additional shared one (L2). In the posterior median eyes, the axons of their first-order visual neuropils project directly to the arcuate body, suggesting that the posterior median eyes do not detect movement. The L2 might function as an upstream integration center enabling faster movement decisions.

Tyrosine hydroxylase immunolabeling reveals the distribution of catecholaminergic neurons in the central nervous systems of the spiders Hogna lenta (Araneae: Lycosidae) and Phidippus regius (Araneae: Salticidae).

With over 48,000 species currently described, spiders (Arthropoda: Chelicerata: Araneae) comprise one of the most diverse groups of animals on our planet, and exhibit an equally wide array of fascinating behaviors. Studies of central nervous systems (CNSs) in spiders, however, are relatively sparse, and no reports have yet characterized catecholaminergic (dopamine [DA]- or norepinephrine-synthesizing) neurons in any spider species. Because these neuromodulators are especially important for sensory and motor processing across animal taxa, we embarked on a study to identify catecholaminergic neurons in the CNS of the wolf spider Hogna lenta (Lycosidae) and the jumping spider Phidippus regius (Salticidae). These neurons were most effectively labeled with an antiserum raised against tyrosine hydroxylase (TH), the rate-limiting enzyme in catecholamine synthesis. We found extensive catecholamine-rich neuronal fibers in the first- and second-order optic neuropils of the supraesophageal mass (brain), as well as in the arcuate body, a region of the brain thought to receive visual input and which may be involved in higher order sensorimotor integration. This structure likely shares evolutionary origins with the DA-enriched central complex of the Mandibulata. In the subesophageal mass, we detected an extensive filigree of TH-immunoreactive (TH-ir) arborizations in the appendage neuromeres, as well as three prominent plurisegmental fiber tracts. A vast abundance of TH-ir somata were located in the opisthosomal neuromeres, the largest of which appeared to project to the brain and decorate the appendage neuromeres. Our study underscores the important roles that the catecholamines likely play in modulating spider vision, higher order sensorimotor processing, and motor patterning.

The visual system of Thelyphonida (whip scorpions): Support for Arachnopulmonata

Only a few studies have examined the central visual system of Thelyphonida (whip scorpions) until now. To obtain new insights suitable for phylogenetic analysis we studied the axonal trajectories and neuropil architecture of the central visual systems of two whip scorpion species (Mastigoproctus giganteus and Typopeltis dalyi) with different neuroanatomical techniques (Cobalt fills, Wigglesworth stains, and μCT).The central visual system of whip scorpion comprises one pair of median eyes and one pair of lateral eye triplets. The R-cells (or retinula cells) of both eye types each terminate in a first and a second visual neuropil. Furthermore, a few R-cell fibres from the median eyes leave the second median eye visual neuropil and terminate in the second and the first lateral eye neuropil. This means R-cell terminals from the lateral eyes and the median eyes overlap here. Additionally, the arcuate body and mushroom bodies are described.A detailed comparison of our findings with previously studied chelicerate central visual systems seems to support a monophyly of Arachnopulmonata, i.e. a clade comprising Tetrapulmonata (Thelyphonida, Schizomida, Amblypygi, and Araneae) and Scorpions. Furthermore, the architecture of the central visual systems hints at a close evolutionary relationship of Arachnopulmonata and Xiphosura.

Also looking like Limulus? \u2013 retinula axons and visual neuropils of Amblypygi (whip spiders)

BackgroundOnly a few studies have examined the visual systems of Amblypygi (whip spiders) until now. To get new insights suitable for phylogenetic analysis we studied the axonal trajectories and neuropil architecture of the visual systems of several whip spider species (Heterophrynus elaphus, Damon medius, Phrynus pseudoparvulus, and P. marginemaculatus) with different neuroanatomical techniques. The R-cell axon terminals were identified with Cobalt fills. To describe the morphology of the visual neuropils and of the protocerebrum generally we used Wigglesworth stains and μCT.ResultsThe visual system of whip spiders comprises one pair of median and three pairs of lateral eyes. The R-cells of both eye types terminate each in a first and a second visual neuropil. Furthermore, a few R-cell fibres from the median eyes leave the second median eye visual neuropil and terminate in the second lateral eye neuropil. This means R-cell terminals from the lateral eyes and the median eyes overlap. Additionally, the arcuate body and the mushroom bodies are described.ConclusionsA detailed comparison of our findings with previously studied chelicerate visual systems (i.e., Xiphosura, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae) seem to support the idea of close evolutionary relationships between Xiphosura, Scorpiones, and Amblypygi.

The allometry of the arcuate body in the postembryonic development of the giant house spider Eratigena atrica

The brain of arachnids contains a special neuropil area called the arcuate body (AB), whose function has been widely discussed. Its growth and proportion in the brain volume during postembryogenesis have been investigated only in several spider species. Our allometric study is aimed at determining to what extent the development of the AB in Eratigena atrica, a spider with unique biology and behaviour, is similar to the development of this body in other species. We put forward a hypothesis of allometric growth of this body in relation to the volume of the central nervous system (CNS) and its neuropil as well as in relation to the volume of the brain and its neuropil. The analysis of paraffin embedded, H + E stained histological preparations confirmed our hypothesis. The AB developed more slowly than the CNS and the neuropil of both the brain and the CNS. In contrast, it exhibited positive allometry in relation to the volume of the brain. This body increased more than nine times within the postembryonic development. Its proportion in the brain volume varied; the lowest was recorded in larvae and nymphs I; then, it increased in nymphs VI and decreased to 2.93% in nymphs X. We conclude that in Eratigena atrica, the AB develops differently that in orb-weaver and wandering spiders. There is no universal model of the AB development, although in adult spiders, regardless of their behaviour, the proportion of this area in the brain volume is similar.

A tiny visual system — retinula axons and visual neuropils of Neobisium carcinoides (Hermann, 1804) (Chelicerata, Arachnida, Pseudoscorpiones)

Only a few studies have examined the visual system of Pseudoscorpiones until now. To fill this knowledge gap we analysed the axonal trajectories and neuropil architecture of the visual system of the pseudoscorpion Neobisium carcinoides (Hermann, 1804) with different neuroanatomical techniques. The R-cell axon terminals were identified with Cobalt fills and the morphology of the visual neuropils and the protocerebrum generally is described by means of the osmium-ethyl gallate procedure and TEM. N. carcinoides has two lateral eyes on each side of the prosoma. The R-cells of the eyes per hemisphere are linked to a first and a second visual neuropil, located in the dorsolateral protocerebrum. The first visual neuropil is subdivided into two hemineuropils, one for each lateral eye, while the second neuropil is not. Furthermore, the two-layered arcuate body is found − isolated from the visual neuropils − in the midline of the protocerebrum at a dorsoposterior position. These findings allow a detailed comparison of the pseudoscorpion visual system with that of other previously studied taxa and shed new light on lateral eye evolution in Chelicerata.

Early environmental conditions affect the volume of higher‐order brain centers in a jumping spider

AbstractThe central nervous system is known to be plastic in volume and structure depending on the stimuli the organism is subjected to. We tested in the jumping spider Marpissa muscosa (Clerck, 1757), whether rearing environments affect the volume of two target higher‐order brain centers: the mushroom body (MB) and the arcuate body (AB). We reared female M. muscosa (N = 39) in three environments: solitarily (D: deprived), solitarily but in a physically enriched environment (P: physically enriched) and together with several siblings (G: group). We additionally investigated spiders caught from the field (W: wild). Volumes of MB and AB were compared using microCT analysis. We hypothesized that spiders reared in treatments P and G should have larger MB and AB than the spiders from treatment D, as the enriched environments are presumably cognitively more demanding than the deprived environment. Spiders from treatment P had significantly larger absolute brain volumes than spiders from treatment D, whereas brain volumes of treatment G lay in between. The relative volume of the MB was not significantly different between the treatments, whereas relative AB volumes were significantly larger in treatment P than in D, supporting the hypothesis that the AB is a center of locomotor control. W spiders had smaller absolute brain volumes and relatively smaller AB than spiders from laboratory treatments, which suggests developmental constraints under natural, possibly food‐limited conditions. Additionally, differences in the relative volume of MB substructures were found. Overall, our study demonstrates that brains of jumping spiders respond plastically to environmental conditions in that absolute brain volume, as well as the relative volume of higher‐order brain centers, is affected.

The synganglion of the jumping spider Marpissa muscosa (Arachnida: Salticidae): Insights from histology, immunohistochemistry and microCT analysis

Jumping spiders are known for their extraordinary cognitive abilities. The underlying nervous system structures, however, are largely unknown. Here, we explore and describe the anatomy of the brain in the jumping spider Marpissa muscosa (Clerck, 1757) by means of paraffin histology, X-ray microCT analysis and immunohistochemistry as well as three-dimensional reconstruction. In the prosoma, the CNS is a clearly demarcated mass that surrounds the esophagus. The anteriormost neuromere, the protocerebrum, comprises nine bilaterally paired neuropils, including the mushroom bodies and one unpaired midline neuropil, the arcuate body. Further ventrally, the synganglion comprises the cheliceral (deutocerebrum) and pedipalpal neuropils (tritocerebrum). Synapsin-immunoreactivity in all neuropils is generally strong, while allatostatin-immunoreactivity is mostly present in association with the arcuate body and the stomodeal bridge. The most prominent neuropils in the spider brain, the mushroom bodies and the arcuate body, were suggested to be higher integrating centers of the arthropod brain. The mushroom body in M. muscosa is connected to first and second order visual neuropils of the lateral eyes, and the arcuate body to the second order neuropils of the anterior median eyes (primary eyes) through a visual tract. The connection of both, visual neuropils and eyes and arcuate body, as well as their large size corroborates the hypothesis that these neuropils play an important role in cognition and locomotion control of jumping spiders. In addition, we show that the architecture of the brain of M. muscosa and some previously investigated salticids differs significantly from that of the wandering spider Cupiennius salei, especially with regard to structure and arrangement of visual neuropils and mushroom body. Thus, we need to explore the anatomical conformities and specificities of the brains of different spider taxa in order to understand evolutionary transformations of the arthropod brain.

The visual system of harvestmen (Opiliones, Arachnida, Chelicerata) - a re-examination.

BackgroundThe visual systems in chelicerates are poorly understood, even though they show strong variation in eye and visual neuropil architecture, thus may provide valuable insights for the understanding of chelicerate phylogeny and eye evolution. Comparable morphological characters are desperately sought for reconstructions of the phylogeny of Chelicerata, especially with respect to Arachnida. So far, reliable data exist only for Pycnogonida, Xiphosura, Scorpiones, and Araneae. The few earlier studies of the organisation of the visual system in harvestmen are contradictory concerning the number, morphology, and position of the visual neuropils.ResultsWe undertook a descriptive and comparative analysis of the neuroanatomy of the visual system in several phalangid harvestmen species. Various traditional and modern methods were used that allow comparisons with previous results (cobalt fills, DiI/DiO labelling, osmium ethyl gallate procedure, and TEM). The R-cells (photoreceptor and arhabdomeric cells) in the eyes of Opiliones are linked to a first and a second visual neuropil. The first visual neuropil receives input from all R-cell axons, in the second only few R-cells terminate in the distal part. Hence, the second visual neuropil is subdivided in a part with direct R-cell input and a part without. The arcuate body is located in a subsequent position with direct contact to the second visual neuropil.ConclusionsThis re-examination comes to conclusions different from those of all previous studies. The visual system of phalangid Opiliones occupies an intermediate position between Pycnogonida, Xiphosura, and Scorpiones on the one side, and Araneae on the other side. The projection of the R-cells is similar to that in the former grouping, the general neuropil arrangement to that in the latter taxon. However, more research on the visual systems in other chelicerate orders is needed in order to draw inferences on phylogeny or eye evolution.

Differential expression of retinal determination genes in the principal and secondary eyes of Cupiennius salei Keyserling (1877).

BackgroundTranscription factors that determine retinal development seem to be conserved in different phyla throughout the animal kingdom. In most representatives, however, only a few of the involved transcription factors have been sampled and many animal groups remain understudied. In order to fill in the gaps for the chelicerate group of arthropods, we tested the expression pattern of the candidate genes involved in the eye development in the embryo of the wandering spider Cupiennius salei. One main objective was to profile the molecular development of the eyes and to search for possible variation among eye subtype differentiation. A second aim was to form a basis for comparative studies in order to elucidate evolutionary pathways in eye development.ResultsWe screened the spider embryonic transcriptome for retina determination gene candidates and discovered that all except one of the retinal determination genes have been duplicated. Gene expression analysis shows that the two orthologs of all the genes have different expression patterns. The genes are mainly expressed in the developing optic neuropiles of the eyes (lateral furrow, mushroom body, arcuate body) in earlier stages of development (160 to 220 h after egg laying). Later in development (180 to 280 h after egg laying), there is differential expression of the genes in disparate eye vesicles; for example, Cs-otxa is expressed only in posterior-lateral eye vesicles, Cs-otxb, Cs-six1a, and Cs-six3b in all three secondary eye vesicles, Cs-pax6a only in principal eye vesicles, Cs-six1b in posterior-median, and posterior-lateral eye vesicles, and Cs-six3a in lateral and principal eye vesicles.ConclusionsPrinciple eye development shows pax6a (ey) expression, suggesting pax6 dependence, although secondary eyes develop independently of pax6 genes and show differential expression of several retinal determination genes. Comparing this with the other arthropods suggests that pax6-dependent median eye development is a ground pattern of eye development in this group and that the ocelli of insects, the median eyes of chelicerates, and nauplius eyes can be homologised. The expression pattern of the investigated genes makes it possible to distinguish between secondary eyes and principal eyes. Differences of gene expression among the different lateral eyes indicate disparate function combined with genetic drift.Electronic supplementary materialThe online version of this article (doi:10.1186/s13227-015-0010-x) contains supplementary material, which is available to authorized users.

Petrology and geochemistry of the ultramafic-mafic Mawpyut complex, Meghalaya: a Sylhet trap differentiation centre in northeastern India

The present article describes, for the first time, petrological and geochemical details of the Mawpyut differentiated complex which is related to the Sylhet trap located at Jaintia Hills district, Meghalaya, northeastern India. The Mawpyut complex occurs as an arcuate body that intrudes into the surrounding Shillong Group rocks. The complex in general contains ‘ultramafic’ and ‘mafic’ rocks, as well as minor syenitic veins that postdate the main units. The lithotypes correspond to cumulate and noncumulate units. The cumulate unit is represented by olivine clinopyroxenite, clinopyroxenite, plagioclase-bearing ultramafic, olivine gabbronorite, mela-gabbronorite, melagabbro, orthopyroxene gabbro, and gabbro, all with a pronounced cumulus texture. The noncumulate unit is marked by gabbro, monzonite, monzodiorite, and quartzsyenite. The use of several major and trace element variation diagrams suggests that magmatic differentiation led to the formation of cumulate and noncumulate units. In chondrite-normalized REE diagrams the cumulate rocks show flat LREE and MREE patterns and a moderate positive Eu anomaly (in plagioclase-bearing ultramafics) due to plagioclase cumulation. The rocks of the noncumulate unit show a strongly fractionated REE pattern and no Eu anomaly. The noncumulate mafic rocks are geochemically comparable to high-phosphorous/high-titanium basalts (HPT) indicative of low pressure fractional crystallization. In a primitive mantle-normalized multielement diagram some of the cumulate rocks show pronounced negative anomalies for K and P, indicating anorogenic mafic magmatism in a within-plate setting. The rocks of the noncumulate unit show a slight negative anomaly for Yb and a Nb–Ta trough, indicating a subduction-related signature that perhaps is inherited from subducted sedimentary rocks incorporated during crustal contamination of the derived magma (left after crystal cumulation) with country rocks. Various trace element ratios for the cumulate mafic rocks indicate parent EMI/EMII/HIMU sources with a very limited crustal signature. The noncumulate mafic rocks (corresponding to the derived evolved magma) indicate EMI/EMII/HIMU sources with a pronounced crustal contamination. The Sr–Nd isotopic compositions of the Mawpyut samples typically plot in the continental flood basalt field, with an affinity to the EMII source. The isotopic compositions of the noncumulate rocks also clearly indicate crustal contamination. We suggest that partial melting (involving garnet in the residue) of the enriched mantle source EMI/EMII/HIMU could have derived the parental melt; this melt, in turn, underwent assimilation and fractional crystallization to produce the variety of cumulate-noncumulate lithologies of the Mawpyut complex. Copyright © 2013 John Wiley & Sons, Ltd.

Looking like Limulus? - Retinula axons and visual neuropils of the median and lateral eyes of scorpions.

BackgroundDespite ongoing interest in the neurophysiology of visual systems in scorpions, aspects of their neuroanatomy have received little attention. Lately sets of neuroanatomical characters have contributed important arguments to the discussion of arthropod ground patterns and phylogeny. In various attempts to reconstruct phylogeny (from morphological, morphological + molecular, or molecular data) scorpions were placed either as basalmost Arachnida, or within Arachnida with changing sister-group relationships, or grouped with the extinct Eurypterida and Xiphosura inside the Merostomata. Thus, the position of scorpions is a key to understanding chelicerate evolution. To shed more light on this, the present study for the first time combines various techniques (Cobalt fills, DiI / DiO labelling, osmium-ethyl gallate procedure, and AMIRA 3D-reconstruction) to explore central projections and visual neuropils of median and lateral eyes in Euscorpius italicus (Herbst, 1800) and E. hadzii Di Caporiacco, 1950.ResultsScorpion median eye retinula cells are linked to a first and a second visual neuropil, while some fibres additionally connect the median eyes with the arcuate body. The lateral eye retinula cells are linked to a first and a second visual neuropil as well, with the second neuropil being partly shared by projections from both eyes.ConclusionsComparing these results to previous studies on the visual systems of scorpions and other chelicerates, we found striking similarities to the innervation pattern in Limulus polyphemus for both median and lateral eyes. This supports from a visual system point of view at least a phylogenetically basal position of Scorpiones in Arachnida, or even a close relationship to Xiphosura. In addition, we propose a ground pattern for the central projections of chelicerate median eyes.

Petrology, geochemistry of hornblende gabbro and associated dolerite dyke of Paharpur, Puruliya, West Bengal: Implication for petrogenetic process and tectonic setting

Paharpur gabbroic intrusive is an arcuate body running east–west paralleling the foliation of Chhotanagpur Granite Gneiss which acts as country rock. The main gabbroic body is intruded by a number of dolerite dykes running north–south. It is composed of clinopyroxene (Wo48En40Fs12–Wo51En40Fs09, mg no. 72–82), plagioclase (An52–An90), hornblende (magnesian hornblende to ferro-tschermackite), orthopyroxene (En76–En79) and ilmenite. Hornblende occurs as large poikilitic grain and constitutes around 60% of the rock. Both gabbro and associated dolerite dykes, show relatively primitive character (mg no. 65–73). Primitive mantle-normalized and MORB-normalized spider diagrams indicate enrichment in Rb, Ba, Th, La, Sr and depletion in Nb, Zr, Y, Ti and Nd. The LILE enrichment and Nb, Ti, Zr, Y depletion suggest arc like geochemical signature for the gabbroic and doleritic rocks of Paharpur. Flat to slightly LREE fractionated pattern and variable degree of REE enrichment is observed. An early stage fractionation of clinopyroxene, plagioclase, orthopyroxene, ilmenite and late stage reaction of cumulate pile and evolved melt/hydrous fluid is suggested for magmatic evolution of gabbro. Associated dolerite dykes, which are geochemically similar to the gabbro, have tholeiitic with boninitic character. The mineralogical and chemical compositions of intrusive rocks also have some similarity with mafic rocks of ophiolite complex of subduction zone.

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.

Neuroarchitecture of the arcuate body in the brain of the spider Cupiennius salei (Araneae, Chelicerata) revealed by allatostatin-, proctolin-, and CCAP-immunocytochemistry and its evolutionary implications

Here we describe the neuronal organization of the arcuate body in the brain of the wandering spider Cupiennius salei. The internal anatomy of this major brain center is analyzed in detail based on allatostatin-, proctolin-, and crustacean cardioactive peptide (CCAP)-immunohistochemistry. Prominent neuronal features are demonstrated in graphic reconstructions. The stainings revealed that the neuroarchitecture of the arcuate body is characterized by several distinct layers some of which comprise nerve terminals that are organized in columnar, palisade-like arrays. The anatomy of the spider’s arcuate body exhibits similarities as well as differences when compared to the central complex in the protocerebrum of the Tetraconata. Arguments for and against a possible homology of the arcuate body of the Chelicerata and the central complex of the Tetraconata and their consequences for the understanding of arthropod brain evolution are discussed.

Compartmentalization of the precheliceral neuroectoderm in the spider Cupiennius salei: Development of the arcuate body, optic ganglia, and mushroom body

Similarly to vertebrates, arthropod brains are compartmentalized into centers with specific neurological functions such as cognition, behavior, and memory. The centers can be further subdivided into smaller functional units. This raises the question of how these compartments are formed during development and how they are integrated into brain centers. We show here for the first time how the precheliceral neuroectoderm of the spider Cupiennius salei is compartmentalized to form the distinct brain centers of the visual system: the optic ganglia, the mushroom bodies, and the arcuate body. The areas of the visual brain centers are defined by the formation of grooves and vesicles and express the proneural gene CsASH1, followed by expression of the neural differentiation marker Prospero. Furthermore, the transcription factor dachshund, which is strongly enriched in the mushroom bodies and the outer optic ganglion of Drosophila, is expressed in the optic anlagen and the mushroom bodies of the spider. The developing brain centers are further subdivided into single neural precursor groups, which become incorporated into the grooves and vesicles but remain distinguishable throughout development, suggesting that they encode spatial information for neural subtype identity. Several molecular and morphological aspects of the development of the optic ganglia and the mushroom bodies are similar in the spider and in insects. Furthermore, we show that the primary engrailed head spot contributes neurons to the optic ganglia of the median eyes, whereas the secondary head spot, which has been associated with the optic ganglia in insects and crustaceans, is absent.