The venom and telopodal defence systems of the centipede Lithobius forficatus are functionally convergent serial homologues.

BACKGROUND: Evolution of novelty is a central theme in evolutionary biology, yet studying the origins of traits with an apparently discontinuous origin remains a major challenge. Venom systems are a well-suited model for the study of this phenomenon because they capture several aspects of novelty across multiple levels of biological complexity. However, while there is some knowledge on the evolution of individual toxins, not much is known about the evolution of venom systems as a whole. One way of shedding light on the evolution of new traits is to investigate less specialised serial homologues, i.e. repeated traits in an organism that share a developmental origin. This approach can be particularly informative in animals with repetitive body segments, such as centipedes. RESULTS: Here, we investigate morphological and biochemical aspects of the defensive telopodal glandular organs borne on the posterior legs of venomous stone centipedes (Lithobiomorpha), using a multimethod approach, including behavioural observations, comparative morphology, proteomics, comparative transcriptomics and molecular phylogenetics. We show that the anterior venom system and posterior telopodal defence system are functionally convergent serial homologues, where one (telopodal defence) represents a model for the putative early evolutionary state of the other (venom). Venom glands and telopodal glandular organs appear to have evolved from the same type of epidermal gland (four-cell recto-canal type) and while the telopodal defensive secretion shares a great degree of compositional overlap with centipede venoms in general, these similarities arose predominantly through convergent recruitment of distantly related toxin-like components. Both systems are composed of elements predisposed to functional innovation across levels of biological complexity that range from proteins to glands, demonstrating clear parallels between molecular and morphological traits in the properties that facilitate the evolution of novelty. CONCLUSIONS: The evolution of the lithobiomorph telopodal defence system provides indirect empirical support for the plausibility of the hypothesised evolutionary origin of the centipede venom system, which occurred through functional innovation and gradual specialisation of existing epidermal glands. Our results thus exemplify how continuous transformation and functional innovation can drive the apparent discontinuous emergence of novelties on higher levels of biological complexity.

Production, composition, and mode of action of the painful defensive venom produced by a limacodid caterpillar, Doratifera vulnerans.

Venoms have evolved independently several times in Lepidoptera. Limacodidae is a family with worldwide distribution, many of which are venomous in the larval stage, but the composition and mode of action of their venom is unknown. Here, we use imaging technologies, transcriptomics, proteomics, and functional assays to provide a holistic picture of the venom system of a limacodid caterpillar, Doratifera vulnerans Contrary to dogma that defensive venoms are simple in composition, D. vulnerans produces a complex venom containing 151 proteinaceous toxins spanning 59 families, most of which are peptides <10 kDa. Three of the most abundant families of venom peptides (vulnericins) are 1) analogs of the adipokinetic hormone/corazonin-related neuropeptide, some of which are picomolar agonists of the endogenous insect receptor; 2) linear cationic peptides derived from cecropin, an insect innate immune peptide that kills bacteria and parasites by disrupting cell membranes; and 3) disulfide-rich knottins similar to those that dominate spider venoms. Using venom fractionation and a suite of synthetic venom peptides, we demonstrate that the cecropin-like peptides are responsible for the dominant pain effect observed in mammalian in vitro and in vivo nociception assays and therefore are likely to cause pain after natural envenomations by D. vulnerans Our data reveal convergent molecular evolution between limacodids, hymenopterans, and arachnids and demonstrate that lepidopteran venoms are an untapped source of novel bioactive peptides.

A reversal in sensory processing accompanies ongoing ecological divergence and speciation in Rhagoletis pomonella.

Changes in behaviour often drive rapid adaptive evolution and speciation. However, the mechanistic basis for behavioural shifts is largely unknown. The tephritid fruit fly Rhagoletis pomonella is an example of ecological specialization and speciation in action via a recent host plant shift from hawthorn to apple. These flies primarily use specific odours to locate fruit, and because they mate only on or near host fruit, changes in odour preference for apples versus hawthorns translate directly to prezygotic reproductive isolation, initiating speciation. Using a variety of techniques, we found a reversal between apple and hawthorn flies in the sensory processing of key odours associated with host fruit preference at the first olfactory synapse, linking changes in the antennal lobe of the brain with ongoing ecological divergence. Indeed, changes to specific neural pathways of any sensory modality may be a broad mechanism for changes in animal behaviour, catalysing the genesis of new biodiversity.

When SEM becomes a deceptive tool of analysis: the unexpected discovery of epidermal glands with stalked ducts on the ultimate legs of geophilomorph centipedes.

BACKGROUND: The jointed appendage is a key novelty in arthropod evolution and arthropod legs are known to vary enormously in relation to function. Among centipedes, the ultimate legs always are distinctly different from locomotory legs, and different centipede taxa evolved different structural and functional modifications. In Geophilomorpha (soil centipedes), ultimate legs do not participate in locomotion and were interpret to serve a sensory function. They can be sexually dimorphic and in some species, male ultimate legs notably appear "hairy". It can be assumed that the high abundance of sensilla indicates a pronounced sensory function. This study seeks for assessing the sensory diversity, however, documents the surprising and unique case of an extensive glandular epithelium in the ultimate legs of three phylogenetically distant species. RESULTS: The tightly aggregated epidermal glands with stalked ducts - mistakenly thought to be sensilla - were scrutinized using a multimodal microscopic approach comprising histology as well as scanning and transmission electron microscopy in Haplophilus subterraneus. Hence, this is the first detailed account on centipede ultimate legs demonstrating an evolutionary transformation into a "secretory leg". Additionally, we investigated sensory structures as well as anatomical features using microCT analysis. Contrary to its nomination as a tarsus, tarsus 1 possesses intrinsic musculature, which is an indication that this podomere might be a derivate of the tibia. DISCUSSION: The presence and identity of ultimate leg associated epidermal glands with stalked ducts is a new discovery for myriapods. A pronounced secretory as well as moderate sensory function in Haplophilus subterraneus can be concluded. The set of characters will improve future taxonomic studies, to test the hypotheses whether the presence of these specialized glands is a common feature in Geophilomorpha, and that tarsus 1 may be a derivate of the tibia. As the number of epidermal glands with stalked ducts is sexually dimorphic, their function might be connected to reproduction or a sex-specific defensive role. Our results, in particular the unexpected discovery of 'glandular hairs', may account for a striking example for how deceptive morphological descriptions of epidermal organs may be, if based on non-invasive techniques alone.

The antennal scape organ of Scutigera coleoptrata (Myriapoda) and a new type of arthropod tip-pore sensilla integrating scolopidial components.

BACKGROUND: Centipedes are terrestrial, predatory arthropods with specialized sensory organs. However, many aspects of their sensory biology are still unknown. This also concerns hygroreception, which is especially important for centipedes, as their epicuticle is thin and they lose water rapidly at low humidity. Thus, the detection of humid places is vital but to date no definite hygroreceptor was found in centipedes. House centipedes (Scutigeromorpha) possess a peculiar opening at the base of their antenna, termed 'scape organ', that houses up to 15 cone-shaped sensilla in a cavity. Lacking wall and tip-pores, these socket-less sensilla may be hypothesized to function as hygroreceptors similar to those found in hexapods. RESULTS: The cone-shaped sensilla in the scape organ as well as nearby peg-shaped sensilla are composed of three biciliated receptor cells and three sheath cells. A tip-pore is present but plugged by a highly electron-dense secretion, which also overlays the entire inner surface of the cavity. Several solitary recto-canal epidermal glands produce the secretion. Receptor cell type 1 (two cells in cone-shaped sensilla, one cell in peg-shaped sensilla) possesses two long dendritic outer segments that project to the terminal pore. Receptor cell type 2 (one cell in both sensilla) possesses two shorter dendritic outer segments connected to the first (proximal) sheath cell that establishes a scolopale-like structure, documented for the first time in detail in a myriapod sensillum. CONCLUSIONS: The nearly identical configuration of receptor cells 1 with their long dendritic outer segments in both sensilla is similar to hexapod hygroreceptors. In Scutigera coleoptrata, however, the mechanism of stimulus transduction is different. Water vapor may lead to swelling and subsequent elongation of the plug pin that enters the terminal pore, thus causing stimulation of the elongated dendritic outer segments. The interconnection of receptor cell 2 with short outer dendritic segments to a scolopale-like structure potentially suits both sensilla for vibration or strain detection. Thus, both sensilla located at the antennal base of scutigeromorph centipedes fulfill a dual function.

The "amphi"-brains of amphipods: new insights from the neuroanatomy of Parhyale hawaiensis (Dana, 1853).

BACKGROUND: Over the last years, the amphipod crustacean Parhyale hawaiensis has developed into an attractive marine animal model for evolutionary developmental studies that offers several advantages over existing experimental organisms. It is easy to rear in laboratory conditions with embryos available year-round and amenable to numerous kinds of embryological and functional genetic manipulations. However, beyond these developmental and genetic analyses, research on the architecture of its nervous system is fragmentary. In order to provide a first neuroanatomical atlas of the brain, we investigated P. hawaiensis using immunohistochemical labelings combined with laser-scanning microscopy, X-ray microcomputed tomography, histological sectioning and 3D reconstructions. RESULTS: As in most amphipod crustaceans, the brain is dorsally bent out of the body axis with downward oriented lateral hemispheres of the protocerebrum. It comprises almost all prominent neuropils that are part of the suggested ground pattern of malacostracan crustaceans (except the lobula plate and projection neuron tract neuropil). Beyond a general uniformity of these neuropils, the brain of P. hawaiensis is characterized by an elaborated central complex and a modified lamina (first order visual neuropil), which displays a chambered appearance. In the light of a recent analysis on photoreceptor projections in P. hawaiensis, the observed architecture of the lamina corresponds to specialized photoreceptor terminals. Furthermore, in contrast to previous descriptions of amphipod brains, we suggest the presence of a poorly differentiated hemiellipsoid body and an inner chiasm and critically discuss these aspects. CONCLUSIONS: Despite a general uniformity of amphipod brains, there is also a certain degree of variability in architecture and size of different neuropils, reflecting various ecologies and life styles of different species. In contrast to other amphipods, the brain of P. hawaiensis does not display any striking modifications or bias towards processing one particular sensory modality. Thus, we conclude that this brain represents a common type of an amphipod brain. Considering various established protocols for analyzing and manipulating P. hawaiensis, this organism is a suitable model to gain deeper understanding of brain anatomy e.g. by using connectome approaches, and this study can serve as first solid basis for following studies.

An atlas of larval organogenesis in the European shore crab Carcinus maenas L. (Decapoda, Brachyura, Portunidae).

BACKGROUND: The life history stages of brachyuran crustaceans include pelagic larvae of the Zoea type which grow by a series of moults from one instar to the next. Zoeae actively feed and possess a wide range of organ systems necessary for autonomously developing in the plankton. They also display a rich behavioural repertoire that allows for responses to variations in environmental key factors such as light, hydrostatic pressure, tidal currents, and temperature. Brachyuran larvae have served as distinguished models in the field of Ecological Developmental Biology fostering our understanding of diverse ecophysiological aspects such as phenotypic plasticity, carry-over effects on life-history traits, and adaptive mechanisms that enhance tolerance to fluctuations in environmental abiotic factors. In order to link such studies to the level of tissues and organs, this report analyses the internal anatomy of laboratory-reared larvae of the European shore crab Carcinus maenas. This species has a native distribution extending across most European waters and has attracted attention because it has invaded five temperate geographic regions outside of its native range and therefore can serve as a model to analyse thermal tolerance of species affected by rising sea temperatures as an effect of climate change. RESULTS: Here, we used X-ray micro-computed tomography combined with 3D reconstruction to describe organogenesis in brachyuran larvae. We provide a detailed atlas of the larval internal organization to complement existing descriptions of its external morphology. In a multimethodological approach, we also used cuticular autofluorescence and classical histology to analyse the anatomy of selected organ systems. CONCLUSIONS: Much of our fascination for the anatomy of brachyuran larvae stems from the opportunity to observe a complex organism on a single microscopic slide and the realization that the entire decapod crustacean bauplan unfolds from organ anlagen compressed into a miniature organism in the sub-millimetre range. The combination of imaging techniques used in the present study provides novel insights into the bewildering diversity of organ systems that brachyuran larvae possess. Our analysis may serve as a basis for future studies bridging the fields of evolutionary developmental biology and ecological developmental biology.

Boring systematics: A genome skimmed phylogeny of ctenostome bryozoans and their endolithic family Penetrantiidae with the description of one new species.

Ctenostomes are a group of gymnolaemate bryozoans with an uncalcified chitinous body wall having few external, skeletal characters. Hence, species identification is challenging and their systematics remain poorly understood, even more so when they exhibit an endolithic (boring) lifestyle. Currently, there are four Recent families of endolithic bryozoans that live inside mineralized substrates like mollusk shells. In particular, Penetrantiidae Silén, 1946 has received considerable attention and its systematic affinity to either cheilostomes or ctenostomes has been debated. Species delimitation of penetrantiids remains difficult, owing to a high degree of colonial and zooidal plasticity. Consequently, an additional molecular approach is essential to unravel the systematics of penetrantiids, their phylogenetic placement and their species diversity. We therefore sequenced the mitochondrial (mt) genomes and two nuclear markers of 27 ctenostome species including nine penetrantiids. Our phylogeny supports the Penetrantiidae as a monophyletic group placed as sister taxon to the remaining ctenostomes alongside paludicellids, arachnidioids and terebriporids. The boring family Terebriporidae d'Orbigny, 1847 were previously considered to be among vesicularioids, but our results suggest an arachnidioid affinity instead. Ctenostome paraphyly is supported by our data, as the cheilostomes nest within them. A Multiporata clade is also well supported, including the former victorelloid genus Sundanella. Altogether, this study provides new insights into ctenostome systematics, assists with species delimitation and contributes to our understanding of the bryozoan tree of life.

Paraptotic Cell Death as an Unprecedented Mode of Action Observed for New Bipyridine-Silver(I) Compounds Bearing Phosphane Coligands.

In this work, we investigated the anticancer activity of several novel silver(I) 2,2'-bipyridine complexes containing either triphenylphosphane (PPh3) or 1,2-bis(diphenylphosphino)ethane (dppe) ligands. All compounds were characterized by diverse analytical methods including ESI-MS spectrometry; NMR, UV-vis, and FTIR spectroscopies; and elemental analysis. Moreover, several compounds were also studied by X-ray single-crystal diffraction. Subsequently, the compounds were investigated for their anticancer activity against drug-resistant and -sensitive cancer cells. Noteworthily, neither carboplatin and oxaliplatin resistance nor p53 deletion impacted on their anticancer efficacy. MES-OV cells displayed exceptional hypersensitivity to the dppe-containing drugs. This effect was not based on thioredoxin reductase inhibition, enhanced drug uptake, or apoptosis induction. In contrast, dppe silver drugs induced paraptosis, a novel recently described form of programmed cell death. Together with the good tumor specificity of this compound's class, this work suggests that dppe-containing silver complexes could be interesting drug candidates for the treatment of resistant ovarian cancer.

Comparative analysis of deutocerebral neuropils in Chilopoda (Myriapoda): implications for the evolution of the arthropod olfactory system and support for the Mandibulata concept.

BACKGROUND: Originating from a marine ancestor, the myriapods most likely invaded land independently of the hexapods. As these two evolutionary lineages conquered land in parallel but separately, we are interested in comparing the myriapod chemosensory system to that of hexapods to gain insights into possible adaptations for olfaction in air. Our study connects to a previous analysis of the brain and behavior of the chilopod (centipede) Scutigera coleoptrata in which we demonstrated that these animals do respond to volatile substances and analyzed the structure of their central olfactory pathway. RESULTS: Here, we examined the architecture of the deutocerebral brain areas (which process input from the antennae) in seven additional representatives of the Chilopoda, covering all major subtaxa, by histology, confocal laser-scan microscopy, and 3D reconstruction. We found that in all species that we studied the majority of antennal afferents target two separate neuropils, the olfactory lobe (chemosensory, composed of glomerular neuropil compartments) and the corpus lamellosum (mechanosensory). The numbers of olfactory glomeruli in the different chilopod taxa ranged from ca. 35 up to ca. 90 and the shape of the glomeruli ranged from spheroid across ovoid or drop-shape to elongate. CONCLUSION: A split of the afferents from the (first) pair of antennae into separate chemosensory and mechanosensory components is also typical for Crustacea and Hexapoda, but this set of characters is absent in Chelicerata. We suggest that this character set strongly supports the Mandibulata hypothesis (Myriapoda + (Crustacea + Hexapoda)) as opposed to the Myriochelata concept (Myriapoda + Chelicerata). The evolutionary implications of our findings, particularly the plasticity of glomerular shape, are discussed.

Comparative brain architecture of the European shore crab Carcinus maenas (Brachyura) and the common hermit crab Pagurus bernhardus (Anomura) with notes on other marine hermit crabs.

The European shore crab Carcinus maenas and the common hermit crab Pagurus bernhardus are members of the sister taxa Brachyura and Anomura (together forming the taxon Meiura) respectively. Both species share similar coastal marine habitats and thus are confronted with similar environmental conditions. This study sets out to explore variations of general brain architecture of species that live in seemingly similar habitats but belong to different major malacostracan taxa and to understand possible differences of sensory systems and related brain compartments. We examined the brains of Carcinus maenas, Pagurus bernhardus, and three other hermit crab species with immunohistochemistry against tyrosinated tubulin, f-actin, synaptic proteins, RF-amides and allatostatin. Our comparison showed that their optic neuropils within the eyestalks display strong resemblance in gross morphology as well as in detailed organization, suggesting a rather similar potential of processing visual input. Besides the well-developed visual system, the olfactory neuropils are distinct components in the brain of both C. maenas and P. bernhardus as well as the other hermit crabs, suggesting that close integration of olfactory and visual information may be useful in turbid marine environments with low visibility, as is typical for many habitats such as, e.g., the Baltic and the North Sea. Comparing the shape of the olfactory glomeruli in the anomurans showed some variations, ranging from a wedge shape to an elongate morphology. Furthermore, the tritocerebrum and the organization of the second antennae associated with the tritocerebrum seem to differ markedly in C. maenas and P. bernhardus, indicating better mechanosensory abilities in the latter close to those of other Decapoda with long second antennae, such as Astacida, Homarida, or Achelata. This aspect may also represent an adaptation to the "hermit lifestyle" in which competition for shells is a major aspect of their life history. The shore crab C. maenas, on the other hand seems to rely much less on mechanosensory information mediated by the second antennae but in water, the visual and the olfactory senses seem to be the most important modalities.

Modern venomics-Current insights, novel methods, and future perspectives in biological and applied animal venom research.

Venoms have evolved >100 times in all major animal groups, and their components, known as toxins, have been fine-tuned over millions of years into highly effective biochemical weapons. There are many outstanding questions on the evolution of toxin arsenals, such as how venom genes originate, how venom contributes to the fitness of venomous species, and which modifications at the genomic, transcriptomic, and protein level drive their evolution. These questions have received particularly little attention outside of snakes, cone snails, spiders, and scorpions. Venom compounds have further become a source of inspiration for translational research using their diverse bioactivities for various applications. We highlight here recent advances and new strategies in modern venomics and discuss how recent technological innovations and multi-omic methods dramatically improve research on venomous animals. The study of genomes and their modifications through CRISPR and knockdown technologies will increase our understanding of how toxins evolve and which functions they have in the different ontogenetic stages during the development of venomous animals. Mass spectrometry imaging combined with spatial transcriptomics, in situ hybridization techniques, and modern computer tomography gives us further insights into the spatial distribution of toxins in the venom system and the function of the venom apparatus. All these evolutionary and biological insights contribute to more efficiently identify venom compounds, which can then be synthesized or produced in adapted expression systems to test their bioactivity. Finally, we critically discuss recent agrochemical, pharmaceutical, therapeutic, and diagnostic (so-called translational) aspects of venoms from which humans benefit.

Organization of deutocerebral neuropils and olfactory behavior in the centipede Scutigera coleoptrata (Linnaeus, 1758) (Myriapoda: Chilopoda).

Myriapods represent an arthropod lineage, that originating from a marine arthropod ancestor most likely conquered land independently from hexapods and crustaceans. Establishing aerial olfaction during a transition from the ocean to land requires molecules to be detected in gas phase instead of in water solution. Considering that the olfactory sense of myriapods has evolved independently from that in hexapods and crustaceans, the question arises if and how myriapods have solved the tasks of odor detection and odor information processing in air. Comparative studies between arthropod taxa that independently have established a terrestrial life style provide a powerful means of investigating the evolution of chemosensory adaptations in this environment and to understand how the arthropod nervous system evolved in response to new environmental and ecological challenges. In general, the neuroethology of myriapods and the architecture of their central nervous systems are insufficiently understood. In a set of experiments with the centipede Scutigera coleoptrata, we analyzed the central olfactory pathway with serial semi-thin sectioning combined with 3-dimensional reconstruction, antennal backfilling with neuronal tracers, and immunofluorescence combined with confocal laser-scanning microscopy. Furthermore, we conducted behavioral experiments to find out if these animals react to airborne stimuli. Our results show that the primary olfactory and mechanosensory centers are well developed in these organisms but that the shape of the olfactory neuropils in S. coleoptrata is strikingly different when compared with those of hexapods and malacostracan crustaceans. Nevertheless, the presence of distinct neuropils for chemosensory and mechanosensory qualities in S. coleoptrata, malacostracan Crustacea, and Hexapoda could indicate a common architectural principle within the Mandibulata. Furthermore, behavioral experiments indicate that S. coleoptrata is able to perceive airborne stimuli, both from live prey and from a chemical extract of the prey. These results are in line with the morphological findings concerning the well-developed olfactory centers in the deutocerebrum of this species.

A comparative analysis of the nervous system of cheilostome bryozoans.

BACKGROUND: Bryozoans are sessile aquatic suspension feeders in mainly marine, but also freshwater habitats. Most species belong to the marine and calcified Cheilostomata. Since this taxon remains mostly unstudied regarding its neuroanatomy, the focus of this study is on the characterization and ground pattern reconstruction of the autozooidal nervous system based on six representatives. RESULTS: A common neuronal innervation pattern is present in the investigated species: a cerebral ganglion is located at the base of the lophophore, from where neurite bundles embrace the mouth opening to form a circumoral nerve ring. Four neurite bundles project from the cerebral ganglion to innervate peripheral areas, such as the body wall and parietal muscles via the tentacle sheath. Five neurite bundles comprise the main innervation of the visceral tract. Four neurite bundles innervate each tentacle via the circumoral nerve ring. Mediofrontal tentacle neurite bundles emerge directly from the nerve ring. Two laterofrontal- and one abfrontal tentacle neurite bundles emanate from radial neurite bundles, which originate from the cerebral ganglion and circumoral nerve ring in between two adjacent tentacles. The radial neurite bundles terminate in intertentacular pits and give rise to one abfrontal neurite bundle at the oral side and two abfrontal neurite bundles at the anal side. Similar patterns are described in ctenostome bryozoans. CONCLUSIONS: The present results thus represent the gymnolaemate situation. Innervation of the tentacle sheath and visceral tract by fewer neurite bundles and tentacular innervation by four to six tentacle neurite bundles support cyclostomes as sister taxon to gymnolaemates. Phylactolaemates feature fewer distinct neurite bundles in visceral- and tentacle sheath innervation, which always split in nervous plexus, and their tentacles have six neurite bundles. Thus, this study supports phylactolaemates as sistergroup to myolaemates.

The tracheal system of scutigeromorph centipedes and the evolution of respiratory systems of myriapods.

The tracheal system of scutigeromorph centipedes (Chilopoda) is special, as it consists of dorsally arranged unpaired spiracles. In this study, we investigate the tracheal systems of five different scutigeromorph species. They are strikingly similar to each other but depict unique characters compared to the tracheal systems of pleurostigmophoran centipedes, which has engendered an ongoing debate over a single versus independent origin of tracheal systems in Chilopoda. Up to now, only the respiratory system of Scutigera coleoptrata was investigated intensively using LM-, TEM-, and SEM-techniques. We supplement this with data for species from all three families of Scutigeromorpha. These reveal interspecific differences in atrial width and the shape and branching pattern of the tracheal tubules. Further, we investigated the tracheal system of Scutigera coleoptrata with three additional techniques: light sheet microscopy, microCT and synchrotron radiation based microCT analysis. This set of techniques allows a comparison between fresh versus fixed and dried material. The question of a unique vs. multiple origin of tracheal systems in centipedes and in Myriapoda as a whole is discussed with regard to their structural similarities and differences and the presence of hemocyanin as an oxygen carrier. We used morphological and molecular data and the fossil record to evaluate the alternative hypotheses.

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.

Primary processing neuropils associated with the malleoli of camel spiders (Arachnida, Solifugae): a re-evaluation of axonal pathways.

BACKGROUND: Arachnids possess highly specialized and unorthodox sense organs, such as the unique pectines of Scorpiones and the malleoli of Solifugae. While the external morphology, numbers, and shapes of sensory organs are widely used in taxonomic studies, little is known about the internal anatomy of these organs and their associated processing neuropils in the central nervous system. Camel spiders (Solifugae) possess pedipalps and first walking legs heavily endowed with sensory structures, as well as conspicuous malleoli located ventrally on the proximal fourth walking legs. Malleoli are fan-shaped organs that contain tens of thousands of presumptive chemoreceptor neurons, but mechanoreceptive structures are absent. RESULTS: Here, we examine the organization of the synganglion based on microCT analysis, 3D reconstruction of serial paraffin sections, and backfill preparations to trace the malleolar pathway. The projection area of malleolar afferents is intriguingly located in the most anterior ventral nerve cord, located in between the pedipalpal neuromere hemispheres. However, malleolar axon bundles are separated by a thin soma layer that points to an anteriad projection of the fourth walking leg neuromere. A conspicuous projection neuron tract that may receive additional input from pedipalpal sensory organs connects the malleolar neuropil with the mushroom bodies in the protocerebrum. CONCLUSION: Arthropod chemosensory appendages or organs and primary processing neuropils are typically located in the same segment, which also holds true in Solifugae, although the malleolar neuropil is partially shifted towards the pedipalpal neuromere. A comparison of the malleoli in Solifugae and the pectines in Scorpiones, and of their primary processing neuropils, reveals certain similarities, while striking differences are also evident. Similarities include the ventral arrangement of peg-shaped sensory structures on the respective segmental appendage, exposing dense arrays of chemoreceptive sensilla, and projections to a primary processing neuropil with glomerular subdivision. Differences are, e.g., the lack of mechanoreceptive afferents and an associated processing neuropil.

Comparative morphology of ultimate and walking legs in the centipede Lithobius forficatus (Myriapoda) with functional implications.

BACKGROUND: In the context of evolutionary arthopodial transformations, centipede ultimate legs exhibit a plethora of morphological modifications and behavioral adaptations. Many species possess significantly elongated, thickened, or pincer-like ultimate legs. They are frequently sexually dimorphic, indicating a role in courtship and mating. In addition, glandular pores occur more commonly on ultimate legs than on walking legs, indicating a role in secretion, chemical communication, or predator avoidance. In this framework, this study characterizes the evolutionarily transformed ultimate legs in Lithobius forficatus in comparison with regular walking legs. RESULTS: A comparative analysis using macro-photography, SEM, µCT, autofluorescence, backfilling, and 3D-reconstruction illustrates that ultimate legs largely resemble walking legs, but also feature a series of distinctions. Substantial differences are found with regard to aspects of the configuration of specific podomeres, musculature, abundance of epidermal glands, typology and distribution of epidermal sensilla, and architecture of associated nervous system structures. CONCLUSION: In consideration of morphological and behavioral characteristics, ultimate legs in L. forficatus primarily serve a defensive, but also a sensory function. Moreover, morphologically coherent characteristics in the organization of the ultimate leg versus the antenna-associated neuromere point to constructional constraints in the evolution of primary processing neuropils.

The assassin bug Pristhesancus plagipennis produces two distinct venoms in separate gland lumens.

The assassin bug venom system plays diverse roles in prey capture, defence and extra-oral digestion, but it is poorly characterised, partly due to its anatomical complexity. Here we demonstrate that this complexity results from numerous adaptations that enable assassin bugs to modulate the composition of their venom in a context-dependent manner. Gland reconstructions from multimodal imaging reveal three distinct venom gland lumens: the anterior main gland (AMG); posterior main gland (PMG); and accessory gland (AG). Transcriptomic and proteomic experiments demonstrate that the AMG and PMG produce and accumulate distinct sets of venom proteins and peptides. PMG venom, which can be elicited by electrostimulation, potently paralyses and kills prey insects. In contrast, AMG venom elicited by harassment does not paralyse prey insects, suggesting a defensive role. Our data suggest that assassin bugs produce offensive and defensive venoms in anatomically distinct glands, an evolutionary adaptation that, to our knowledge, has not been described for any other venomous animal.

Serotonergic neurons in the ventral nerve cord of Chilopoda - a mandibulate pattern of individually identifiable neurons.

BACKGROUND: Given the numerous hypotheses concerning arthropod phylogeny, independent data are needed to supplement knowledge based on traditional external morphology and modern molecular sequence information. One promising approach involves comparisons of the structure and development of the nervous system. Along these lines, the morphology of serotonin-immunoreactive neurons in the ventral nerve cord has been investigated in numerous tetraconate taxa (Crustacea and Hexapoda). It has been shown that these neurons can be identified individually due to their comparably low number, characteristic soma position, and neurite morphology, thus making it possible to establish homologies at the single cell level. Within Chilopoda (centipedes), detailed analyses of major branching patterns of serotonin-immunoreactive neurons are missing, but are crucial for developing meaningful conclusions on the homology of single cells. RESULTS: In the present study, we re-investigated the distribution and projection patterns of serotonin-immunoreactive neurons in the ventral nerve cord of three centipede species: Scutigera coleoptrata, Lithobius forficatus, and Scolopendra oraniensis. The centipede serotonergic system in the ventral nerve cord contains defined groups of individually identifiable neurons. An anterior and two posterior immunoreactive neurons per hemiganglion with contralateral projections, a pair of ipsilateral projecting lateral neurons (an autapomorphic character for Chilopoda), as well as a postero-lateral group of an unclear number of cells are present in the ground pattern of Chilopoda. CONCLUSIONS: Comparisons to the patterns of serotonin-immunoreactive neurons of tetraconate taxa support the homology of anterior and posterior neurons. Our results thus support a sister group relationship of Myriapoda and Tetraconata and, further, a mandibulate ground pattern of individually identifiable serotonin-immunoreactive neurons in the ventral nerve cord. Medial neurons are not considered to be part of the tetraconate ground pattern, but could favor the 'Miracrustacea hypothesis', uniting Remipedia, Cephalocarida, and Hexapoda.