Same same but different: a stunning analogy between tracheal and vascular supply in the CNS of different arachnids

Described herein is an as yet unprecedented structural and functional analogy of both the tracheal supply of the prosomal ganglion in opilionids and the arterial supply of the prosomal ganglion in pulmonate arachnids. Within Arachnida, two different modes of respiration can be observed: the so-called book lungs, and the tube-like tracheae. These different respiratory modes always correlate with a specific setup concerning the complexity of the circulatory system. This fact has a particular influence on the supply of certain organ systems, such as the central nervous system. It has recently been shown that pulmonate arachnids possess a highly complex pattern of intraganglionic arteries. Here, we show that Opiliones (harvestmen) possess a complex tracheal system (which supplies the different organ systems with oxygen) and only a relatively simple vascular system, comprising a short heart and an anterior aorta that runs directly to the prosomal ganglion. Using a variety of modern and classical morphological methods, we studied the vascular, tracheal and nervous systems of different representatives from all higher taxa of Opiliones. We show that the prosomal ganglion is extensively supplied with intraganglionic tracheae. What is especially surprising is the high degree of correspondence between the pattern of these ganglionic tracheae in harvestmen and the pattern of arteries in the prosomal ganglion of pulmonate arachnids. We aim to provide mechanistic causal explanations of these analogous patterns by applying the concepts of role analogy and constructional analogy. We also aim to establish the circulatory system as a model organ system and hope that this may, in turn, provide a starting point for future research programmes.     

Evolution of locomotion in arachnida: The hydraulic pressure pump of the giant whipscorpion,Mastigoproctus Giganteus (Uropygi)

Many arachnids lack extensor muscles at the femoropatellar (knee) joint of their legs and extend this joint with hydraulic pressure during locomotion. Pressure is generated through compression of the prosoma, but there is disagreement about which muscles are involved in this process. Many arachhnologists consider contraction of the musculi laterales, a group of modified extrinsic leg muscles, as the cause of high prosomal pressure and regard hydraulic extension as a derived feature. However, integration of results from phylogenetic and comparative anatomical studies supports the view that hydraulic extension is primitive in Arachnida and that fluid pressure is generated by contraction of endosternal suspensor muscles. The functional predictions of the musculi laterales and endosternite hypotheses were tested by measuring muscle activity and prosomal pressure during unrestrained locomotion in a primitively “extensorless” arachnid, the giant whipscorpion. The results corroborate the endosternite model and refute the musculi laterales model. Changes in the prosomal pressure baseline were correlated with changes in endosternal muscle activity, while the musculi laterales fired in a step-coupled pattern of discrete bursts that appeared to be incapable of generating the pressure observed during locomotion. Step-coupled fluctuations in prosomal pressure were observed but were apparently caused by rapid flexing of the femoropatellar joints of the fourth leg pair rather than contraction of the musculi laterales.     

Comparing the effects of GM and non‐GM soybean varieties on non‐target arthropods

In order to guarantee the safety of genetically modified (GM) soybean crops, it is important to assess the potential toxicity of their expressed insecticidal proteins to non‐target organisms. In the present study, the effects of the GM soybean Insulin‐like Growth Factor (IGF), which is tolerant to the herbicide glufosinate, on plant‐dwelling non‐target insects and arachnids were evaluated in soybean agroecosystems. For comparison, the non‐GM parental cultivar of soybean Gwangan‐kong was used as a control. Data were collected in 2016 and 2017 via surveying at Ochang and Jeonju, Korea. In total, 13,031 individual insects and arachnids, representing 64 families in 11 orders, were captured during the study. Firstly, the results indicate that the GM soybean IGF did not negatively affect plant‐dwelling non‐target insects and arachnids. However, the numbers of captured individuals on both IGF and Gwangan‐kong were higher at Ochang in 2017. The occurrence of insect pests, natural enemies, and other insects differed significantly according to region, region and survey year, and survey year, respectively. In addition, the dominance, diversity, evenness, and richness indices for the collected insects varied significantly among the regions and survey years regardless of soybean variety. The score from PROXSCAL multidimensional scaling using combined data showed that insects and arachnids in different natural environments were separated by their cultivation regions and years irrespective of soybean cultivars.     

Microstructural organization of the central nervous system in the harvestman Leiobunum japonicum (Arachnida: Opiliones)

Although the order Opiliones constitutes the third‐largest group of arachnids, this creature is still mysterious and has a rich unexplored field compared to what is known about insects and crustaceans. The order Opiliones is traditionally regarded as a close relative of mites, mainly because of morphological similarities in external body structure; however microstructural organization of the ganglionic neurons and nerves in the harvestman Leiobunum japonicum is quite similar to the central nervous system (CNS) in all extant arachnids. The CNS consists of a large neural cluster with paired appendicular nerves. The esophagus passes through the neural cluster and divides it into the upper supraesophageal ganglion (SpG) and the lower subesophageal ganglion (SbG). The dorsal part of the SpG has a quite condensed cell body compared with other parts of the CNS and has two main components, the protocerebrum and the cheliceral ganglion. The protocerebrum receives the optic nerves and has four main groups of neuropiles from the optic lobes, the superior central body, the lateral neuropils (corpora pedunculata) and the inferior neuropil. However, a pair of pedipalpal and four pairs of appendage nerves including several pairs of abdominal nerves arise from the nerve masses of the SbG.     

The affinities of mites and ticks: a review

Mites and ticks can be divided into two well-defined clades, Anactinotrichida and Actinotrichida, for which a recent work formalized a suite of putative autapomorphies and reciprocal differences. Whether they are sister-taxa – forming a monophyletic Acari – is more controversial. Earlier supporters of two independent origins for mites largely failed to demonstrate convincing synapomorphies between either of the two lineages and other arachnid orders; although recent work on reproductive biology revealed explicit characters of this nature. Furthermore, some of the characters proposed in support of a monophyletic Acari do not stand up to detailed scrutiny when compared with Arachnida in general. Effective morphological comparisons between mites and other arachnids are hindered by incompatible nomenclature and long-standing, mite-specific characters which are difficult to score for other arachnids. Furthermore, taxon-specific characters restricted to individual mite groups have sometimes been treated erroneously as 'typical' for all Acari. Here, previous hypotheses of mite affinities are reviewed. Historically, authors have debated whether mites are basal arachnids or highly derived. Excluding weakly supported early hypotheses, mites have been resolved – in whole or in part – as sister-group of all other Arachnida (based on tagmosis), closely related to Opiliones (based mostly on genital morphology), Palpigradi (based on controversial interpretations of limb morphology), Solifugae (based mostly on the mouthparts, but now perhaps also reproductive characters) and Ricinulei (based on hexapodal larvae and perhaps mouthparts). We cannot provide a final resolution here, but we aim to highlight important character sets which should be included in subsequent phylogenetic analyses, as well as useful areas for future investigations: particularly tagmosis and the nature of the gnathosoma.     

A trigonotarbid arachnid from the Lower Devonian of Tredomen, Wales

A new trigonotarbid (Arachnida: Trigonotarbida) Arianrhoda bennetti gen. et sp. nov. is described from the Lower Devonian (Lochkovian) of a quarry near Tredomen, Powys, mid Wales, UK. This relatively complete specimen is the first record of a pre-Carboniferous arachnid from Wales, one of only a handful of early Devonian arachnids, and the second oldest trigonotarbid recorded. Based on the rounded prosomal dorsal shield and the relatively narrow, elongate opisthosoma we refer this new fossil to the family Anthracosironidae. A distinct flange-like ornament on the leg 4 tibia in the new fossil is unique among trigonotarbids and is the primary autapomorphy for the new genus.     

Advancing mite phylogenomics: Designing ultraconserved elements for Acari phylogeny

Mites (Acari) are one of the most diverse groups of life on Earth; yet, their evolutionary relationships are poorly understood. Also, the resolution of broader arachnid phylogeny has been hindered by an underrepresentation of mite diversity in phylogenomic analyses. To further our understanding of Acari evolution, we design targeted ultraconserved genomic elements (UCEs) probes, intended for resolving the complex relationships between mite lineages and closely related arachnids. We then test our Acari UCE baits in‐silico by constructing a phylogeny using 13 existing Acari genomes, as well as 6 additional taxa from a variety of genomic sources. Our Acari‐specific probe kit improves the recovery of loci within mites over an existing general arachnid UCE probe set. Our initial phylogeny recovers the major mite lineages, yet finds mites to be non‐monophyletic overall, with Opiliones (harvestmen) and Ricinuleidae (hooded tickspiders) rendering Parasitiformes paraphyletic.     

On the central nervous system of Smeagolidae and Rhodopidae, two families questionably allied with the Gymnomorpha (Gastropoda: Euthyneura)

The central nervous systems ( CNS ) of Smeagol manneringi Climo, 1980 (Smeagolidae), of Rhodope veranü Kölliker, 1847, and of Rhodope transtrosa Salvini-Plawén, 1989 are redescribed in detail. The smeagolid CNS exhibits diagnostic pulmonate characters (procerebrum, cerebral gland, dorsal bodies). Sharing a distinctly structured abdominal ganglion and other peculiarities, the Smeagolidae are closely related to the Onchidüdae, and both families are united as Onchidioidea. The consequences of this close relationship for the origin and evolution of the Gymnomorpha are discussed and the value of the pulmonate CNS for phylogenetics is reconsidered. On the basis of neural characters, which are correlated with the mode of breathing organs, the Pulmonata are divided into Basommatophora, Systellommatophora (Otinidae and Gymnomorpha) and Eupulmonata ( ord. nov .: Trimusculidae, Ellobüdae, Stylommatophora).
In contrast, the CNSs of the rhodopid species lack the pulmonate attributes, but show diagnostic opisthobranch characters such as a rhinophoral nerve. Accordingly, the Rhodopidae are excluded from the Pulmonata and are classified as a separate order Rhodopemorpha among the Opisthobranchia.     

The distribution and relation to the cytoskeleton of specific prosomal proteins in the oogenesis of the clawed toad]

Using immunoblotting and immunofluorescent microscopy, we showed the presence in Xenopus laevis oocytes of two prosomal proteins (27 and 31-33 kDa) and studied their distribution during oogenesis. In the ooplasm, both proteins are detected in prosomal clusters of various size. During previtellogenesis, prosomal proteins are diffusely distributed in the nucleoplasm and form evenly distributed clusters in the cytoplasm. During oocyte growth, prosomal proteins disappear from the nucleus and form animal-vegetal and cortical gradients in the cytoplasm. In the course of oocyte maturation prosomal clusters become smaller. After artificial activation of the egg, the dorso-ventral gradient of distribution of prosomal proteins is observed. Double immunohistochemical labeling revealed morphological association between prosomal clusters and fibril-like structures of the oocyte containing actin and myosin. The latter are then replaced by diffusely distributed actin and myosin. Thus, correlation is observed between localization of the acto-myosin complex of the oocyte and that of prosomal proteins.     

Electron microscopy of the esophageal ganglion complex of the gastropod pulmonate Triodopsis divesta. I. Ultrastructure of the epineurium

The epineurium of the esophageal complex of the gastropod pulmonate Triodopsis divesta was examined by electron microscopy. The epineurium consists of two main regions: an inner dense fibrous region adjacent to the avascular neural tissue of the ganglion and an outer cellular region comprised of a variety of cell types embedded in a connective tissue matrix. The dense fibrous region contains smooth muscle cells and associated nerve processes and is invested on the neural side by thin processes of glial cells. The outer highly cellular region contains smooth muscle cells, nerve processes, wandering cells (amebocytes), globular cells, and myoepithelial cells comprising the walls of the vascular system. In addition, a cell type not previously identified in other gastropod epineuria is present. These cells resemble neurosecretory cells. The morphology and structural interrelationships of these various constituents are presented and the possible functions of individual cell types and the epineurium in general are discussed in relation to information available on other molluscs.     

Pycnogonid affinities: a review

Early authors regarded Pycnogonida (sea spiders) either as aquatic arachnids, ‘degraded’ crustaceans or as some sort of intermediate form between the two. Subsequently, pycnogonids were either placed among the Chelicerata or considered as an isolated group, unrelated to other arthropods. The latter model is untenable under phylogenetic systematics and recent cladistic studies have supported one of two alternative hypotheses. The first is the traditional Chelicerata s.lat. concept, i.e. (Pycnogonida + Euchelicerata). This, however, has only one really convincing synapomorphy: chelate chelicerae. The second hypothesis recognizes (Pycnogonida + all other Euarthropoda) and has been recovered in various ‘total evidence’ studies. Morphologically some characters – the presence of gonopores on the trunk and absence of a labrum, nephridia and intersegmental tendons – support Cormogonida (Euarthropoda excluding pycnogonids). Advances in developmental biology have proposed clear interpretations of segmentation homologies. However, so far there is also a confrontation of the two hypotheses depending on whether the last walking leg segment is considered part of the prosoma. In this case pycnogonids have too many prosomal segments compared with Euchelicerata; perhaps implying they are not sister groups. Alternatively, if part of the postprosomal region, the last leg pair could correspond to the chilarial segment in euchelicerates and its uniramous state could be apomorphic with respect to other euarthropods. Molecular phylogenies need to be more rigorously analysed, better supported by data from different sources and technique‐sensitive aspects need to be explored. Chelicerata s.lat. may emerge as the more convincing model, yet even the putative autapomorphy of chelicerae needs to be treated with caution as there are fossil ‘great appendage’ arthropods in the early Palaeozoic which also have a robust, food‐gathering, pair of head limbs and which may lie on the chelicerate, or even the euarthropod, stem lineage.     

Prosomes and their multicatalytic proteinase activity.

Prosomes were first described as being mRNA-associated RNP (ribonucleoprotein) particles and subcomponents of repressed mRNPs (messenger ribonucleoprotein). We show here that prosomes isolated from translationally inactive mRNP have a protease activity identical to that described by others for the multicatalytic proteinase complex (MCP, 'proteasome'). By RNase or non-ionic detergent treatment, the MCP activity associated with repressed non-globin mRNP from avian erythroblasts, sedimenting at 35 S, could be quantitatively shifted on sucrose gradients to the 19-S sedimentation zone characteristic of prosomes, which were identified by monoclonal antibodies. The presence of small RNA in the enzymatic complex was shown by immunoprecipitation of the protease activity out of dissociated mRNP using a mixture of anti-prosome monoclonal antibodies; a set of small RNAs 80-120 nucleotides long was isolated from the immunoprecipitate. Furthermore, on CsCl gradients, colocalisation of the MCP activity with prosomal proteins and prosomal RNA was found, and no difference in the prosomal RNA pattern was observed whether the particles were fixed or not prior to centrifugation. These data indicate that the MCP activity is a property of prosomes, shown to be in part RNP and subcomplexes of in vivo untranslated mRNP. A hypothesis for the role of the prosome-MCP particles in maintaining homeostasis of specific protein levels is proposed.     

Automated tracking of stem cell lineages of Arabidopsis shoot apex using local graph matching

Shoot apical meristems (SAMs) of higher plants harbor stem‐cell niches. The cells of the stem‐cell niche are organized into spatial domains of distinct function and cell behaviors. A coordinated interplay between cell growth dynamics and changes in gene expression is critical to ensure stem‐cell homeostasis and organ differentiation. Exploring the causal relationships between cell growth patterns and gene expression dynamics requires quantitative methods to analyze cell behaviors from time‐lapse imagery. Although technical breakthroughs in live‐imaging methods have revealed spatio‐temporal dynamics of SAM‐cell growth patterns, robust computational methods for cell segmentation and automated tracking of cells have not been developed. Here we present a local graph matching‐based method for automated‐tracking of cells and cell divisions of SAMs of Arabidopsis thaliana. The cells of the SAM are tightly clustered in space which poses a unique challenge in computing spatio‐temporal correspondences of cells. The local graph‐matching principle efficiently exploits the geometric structure and topology of the relative positions of cells in obtaining spatio‐temporal correspondences. The tracker integrates information across multiple slices in which a cell may be properly imaged, thus providing robustness to cell tracking in noisy live‐imaging datasets. By relying on the local geometry and topology, the method is able to track cells in areas of high curvature such as regions of primordial outgrowth. The cell tracker not only computes the correspondences of cells across spatio‐temporal scale, but it also detects cell division events, and identifies daughter cells upon divisions, thus allowing automated estimation of cell lineages from images captured over a period of 72 h. The method presented here should enable quantitative analysis of cell growth patterns and thus facilitating the development of in silico models for SAM growth.     

In vitro formation of the Merkel cell‐neurite complex in embryonic mouse whiskers using organotypic co‐cultures

A Merkel cell‐neurite complex is a touch receptor composed of specialized epithelial cells named Merkel cells and peripheral sensory nerves in the skin. Merkel cells are found in touch‐sensitive skin components including whisker follicles. The nerve fibers that innervate Merkel cells of a whisker follicle extend from the maxillary branch of the trigeminal ganglion. Whiskers as a sensory organ attribute to the complicated architecture of the Merkel cell‐neurite complex, and therefore it is intriguing how the structure is formed. However, observing the dynamic process of the formation of a Merkel cell‐neurite complex in whiskers during embryonic development is still difficult. In this study, we tried to develop an organotypic co‐culture method of a whisker pad and a trigeminal ganglion explant to form the Merkel cell‐neurite complex in vitro. We initially developed two distinct culture methods of a single whisker row and a trigeminal ganglion explant, and then combined them. By dissecting and cultivating a single row from a whisker pad, the morphogenesis of whisker follicles could be observed under a microscope. After the co‐cultivation of the whisker row with a trigeminal ganglion explant, a Merkel cell‐neurite complex composed of Merkel cells, which were positive for both cytokeratin 8 and SOX2, Neurofilament‐H‐positive trigeminal nerve fibers and Schwann cells expressing Nestin, SOX2 and SOX10 was observed via immunohistochemical analyses. These results suggest that the process for the formation of a Merkel cell‐neurite complex can be observed under a microscope using our organotypic co‐culture method.     

Flashback and foreshadowing—a review of the taxon Opisthobranchia

Opisthobranchia have experienced an unsettled taxonomic history. At the moment their taxonomy is in state of dramatic flux as recent phylogenetic studies have revealed traditional Opisthobranchia to be paraphyletic or even polyphyletic, allocating some traditional opisthobranch taxa to other groups of Heterobranchia, e.g. Pulmonata. Here we review the history of Opisthobranchia and their subgroups, explain their traditionally proposed relationships, and outline the most recent phylogenetic analyses based on various methods (morphology, single gene and multiple gene analyses, as well as genomic data). We also present a phylogenetic hypothesis on Heterobranchia that, according to the latest results, represents a consensus and is the most probable one available to date. The proposed phylogeny supports the Acteonoidea outside of monophyletic Euthyneura, the basal euthyneuran split into Nudipleura (Nudibranchia plus Pleurobranchoidea) and the recently established taxon Tectipleura. The latter divides into the Euopisthobranchia, containing most of the major traditional opisthobranch clades, and the Panpulmonata, with a mix of the former opisthobranch, putative allogastropod and pulmonate taxa. This “new euthyneuran tree” rejects the traditional taxa Opisthobranchia and Pulmonata, and, in particular, has profound implications for preconceived textbook scenarios of opisthobranch and pulmonate evolution, which must now be reconsidered. In the absence of systematic barriers, research communities—which have traditionally investigated marine and non-marine heterobranchs separately—need to interact and finally merge for the sake of science.     

Alteration of the Enantioselective Toxicity of Diclofop Acid by Nonylphenol: Effect on Ascorbate‐Glutathione Cycle in Microcystis Aeruginosa

The enantioselective effects of chiral compounds have been the subject of extensive studies in recent years due to their important implications for contaminant behavior and risk as well as the design of drug and pesticide formulations. The potential alterations of enantioselectivity, however, still remain elusive from the available data suggesting the effects of numerous environmental factors and the coexisting achiral and chiral compounds. Herein we studied the effect of nonylphenol (NP), a ubiquitous contaminant and ingredient in pesticide formulation, on the enantioselectivity of diclofop acid (DC) through ascorbate‐glutathione (AsA‐GSH) cycle in Microcystis aeruginosa. The enantioselectivity of DC in the AsA and GSH antioxidant defense system of M. aeruginosa was affected significantly by the addition of NP. Specifically, although R‐ DC and S‐DC were added with an equal toxic concentration (at their EC₅₀ values), NP addition to the DC treatments altered the enantiomeric ratios of the activities of monodehydroascorbate reductase (MDHAR) and glutathione reductase (GR), key enzymes in the regeneration of AsA and GSH, respectively. NP also modified the enantiomeric ratios of AsA and GSH levels in both the AsA and GSH antioxidant defense systems of M. aeruginosa. Overall, the oxidative damage induced by R‐DC was further deteriorated, whereas that induced by S‐DC was alleviated after NP addition. These altered enantioselectivities indicate a need to reexamine the risks and biological effects of chiral compounds in the complex environmental matrices containing a multitude of other chemicals, including commercial chiral agricultural chemicals. Chirality 28:475–481, 2016. © 2016 Wiley Periodicals, Inc.     

Slit sense organs on the scorpion leg (Androctonus australis L., Buthidae)

As in other arthropods the exoskeleton of arachnids is subjected to loads generated by external stimuli and behavioral activities. Far from being mere by-products of various activities such loads act as signals for mechanoreceptors capable of detecting minute displacements caused by them in the cuticle. In arachnids the slit sense organs serve in this capacity. Spiders have the most elaborate system of slit sense organs. Our previous studies clearly pointed to a functional significance of their specific location and orientation, as well as degree and type of aggregation (isolated, grouped, compound or lyriform) on respective body parts. The present study extends our work to the slit sense organs of scorpions. It gives a detailed account of the topography of the organs on the walking legs. In general slits are less orderly arranged on the legs of scorpions than on those of spiders. In the scorpion they never aggregate to form lyriform organs. Instead there are groups at comparable locations forming much more irregular, but still specific patterns. Isolated slits are more numerous on the scorpion leg, but are also less regularly distrubuted there. A common feature of the majority of slits on both the spider and the scorpion leg is their position on the lateral surfaces and their orientation roughly parallel to the long axis of the leg.     

Beyond modules and hubs: the potential of gene coexpression networks for investigating molecular mechanisms of complex brain disorders

In a research environment dominated by reductionist approaches to brain disease mechanisms, gene network analysis provides a complementary framework in which to tackle the complex dysregulations that occur in neuropsychiatric and other neurological disorders. Gene–gene expression correlations are a common source of molecular networks because they can be extracted from high‐dimensional disease data and encapsulate the activity of multiple regulatory systems. However, the analysis of gene coexpression patterns is often treated as a mechanistic black box, in which looming ‘hub genes’ direct cellular networks, and where other features are obscured. By examining the biophysical bases of coexpression and gene regulatory changes that occur in disease, recent studies suggest it is possible to use coexpression networks as a multi‐omic screening procedure to generate novel hypotheses for disease mechanisms. Because technical processing steps can affect the outcome and interpretation of coexpression networks, we examine the assumptions and alternatives to common patterns of coexpression analysis and discuss additional topics such as acceptable datasets for coexpression analysis, the robust identification of modules, disease‐related prioritization of genes and molecular systems and network meta‐analysis. To accelerate coexpression research beyond modules and hubs, we highlight some emerging directions for coexpression network research that are especially relevant to complex brain disease, including the centrality–lethality relationship, integration with machine learning approaches and network pharmacology .