New insights into the morphology and evolution of the ventral pharynx and jaws in Histriobdellidae (Eunicida, Annelida)

The jaw apparatus in several annelid families represents a powerful tool for systematic approaches and evolutionary investigations. Nevertheless, for several taxa, this character complex has scarcely been investigated, and complete comparative analyses of all annelid jaws are lacking. In our comprehensive study, we described the fine structure of the jaw apparatus and the ventral pharyngeal organ (VPO) in Histriobdella homari – a minute ectocommensal of lobsters putatively belonging to the Eunicida – using different comparative morphological approaches, including SEM, TEM, CLSM and subsequent 3D reconstruction. The H. homari jaw apparatus is composed of ventral paired mandibles and dorsal symmetrical maxillae consisting of numerous dental plates, ventral carriers and an unpaired dorsal rod, and the general assemblage and arrangement of the different parts are highly comparable to those of other eunicid families. The jaw ultrastructure of histriobdellids resembles that of the families Dorvilleidae and (juvenile) Onuphidae. Furthermore, our data reveal that in the process of development of the jaw apparatus, the mandibles, maxillae II and unpaired dorsal rod are formed first, and the remaining maxillae and ventral carriers appear later. Notably, the muscular apparatus differs from that in Dorvilleidae and Onuphidae in terms of the number and arrangement of muscle fibers encompassing the jaws – not only because of the very small size of Histriobdella but also because histriobdellid maxillary protraction occurs due to straightening of the dorsal rod and thus requires a different muscular scaffold. Based on our investigations, the arrangement of the muscular apparatus of the jaws, the presence of paired ventral carriers and the dorsal rod, and the morphology of the ventral pharyngeal organ represent a histriobdellid autapomorphy. Our datasets form a basis for further comparative analyses to elucidate the evolution of Eunicida and jaw-bearing Annelida.publishedVersio

Comparative analyses of morphological characters in Sphaerodoridae and allies (Annelida) revealed by an integrative microscopical approach

Sphaerodoridae is a group of benthic marine worms (Annelida) characterized by the presence of spherical tubercles covering their whole surface. They are commonly considered as belonging to Phyllodocida although sistergroup relationships are still far from being understood. Primary homology assessments of their morphological features are lacking, hindering the appraisal of evolutionary relationships between taxa. Therefore, our detailed morphological investigation focuses on different Sphaerodoridae as well as on other members of Phyllodocida using an integrative approach combining scanning electron microscopy (SEM) as well as immunohistochemistry with standard neuronal (anti-5-HT) and muscular (phalloidin-rhodamine) markers and subsequent CLSM analysis of whole mounts and sections. Furthermore, we provide histological (HES) and light microscopical data to shed light on the structures and hypothetical function of sphaerodorid key morphological features. We provide fundamental details into the sphaerodorid morphology supporting a Phyllodocida ancestry of these enigmatic worms. However, the muscular arrangement and the presence of an axial muscular pharynx is similar to conditions observed in other members of the Errantia too. Furthermore, nervous system and muscle staining as well as SEM and histological observations of different types of tubercles indicate a homology of the so called microtubercles, present in the long-bodied sphaerodorids, to the dorsal cirri of other Errantia. The macrotubercles seem to represent a sphaerodorid autapomorphy based on our investigations. Therefore, our results allow comparisons concerning morphological patterns between Sphaerodoridae and other Phyllodocida and constitute a starting point for further comparative investigations to reveal the evolution of the remarkable Sphaerodoridae

The development of early pioneer neurons in the annelid Malacoceros fuliginosus

Background Nervous system development is an interplay of many processes: the formation of individual neurons, which depends on whole-body and local patterning processes, and the coordinated growth of neurites and synapse formation. While knowledge of neural patterning in several animal groups is increasing, data on pioneer neurons that create the early axonal scaffold are scarce. Here we studied the first steps of nervous system development in the annelid Malacoceros fuliginosus. Results We performed a dense expression profiling of a broad set of neural genes. We found that SoxB expression begins at 4 h postfertilization, and shortly later, the neuronal progenitors can be identified at the anterior and the posterior pole by the transient and dynamic expression of proneural genes. At 9 hpf, the first neuronal cells start differentiating, and we provide a detailed description of axonal outgrowth of the pioneer neurons that create the primary neuronal scaffold. Tracing back the clonal origin of the ventral nerve cord pioneer neuron revealed that it is a descendant of the blastomere 2d (2d221), which after 7 cleavages starts expressing Neurogenin, Acheate-Scute and NeuroD. Conclusions We propose that an anterior and posterior origin of the nervous system is ancestral in annelids. We suggest that closer examination of the first pioneer neurons will be valuable in better understanding of nervous system development in spirally cleaving animals, to determine the potential role of cell-intrinsic properties in neuronal specification and to resolve the evolution of nervous systems.publishedVersio

The central nervous system of Oweniidae (Annelida) and its implications for the structure of the ancestral annelid brain

Figure S1: Histology Orrhage’s Owenia fusiformis. A: slide showing sections of Owenia fusiformis. B: Intermediate filaments (if) cross the neuropil of the brain (br). The ecm of the epidermis is less prominent where the neuropil layer is above it. C: Posterior part of the brain (br). if: intermediate filaments. (JPG 10649 kb

原著

摘出ラット上頸交感神経節を用いて,刺激頻度と薬物作用との関係を検討し,次の結果を得た。1.Hexamethonium,d-Tubocurarine,PhysostigmineおよびParaoxon作用下では,神経節伝達抑制作用が低頻度刺激よりも高頻度刺激により増強された。このような薬物効果をType Aとした。2.Adrenaline(Adr.),Nor-Adrenaline(Nor-Adr),MgCl_2,MnCl_2ならびにCa^-deficit Ringer(0.1〜0.4mM CaCl_2)では,低頻度刺激時にみられた抑制効果が高頻度刺激下で著しく回復され,この効果をType Bとした。3.ProcaineおよびPropranolol作用下の抑制効果は,刺激頻度の多少にほとんど影響されなかった。この効果をType Cとした。4.Adr.,Nor-Adr.のType B効果はPhentolamineにより拮抗されたが,ほとんどPropranololの影響をうけなかった。MgCl_2,MnCl_2およびCa^#-deficit効果は,α-,およびβ-効果遮断薬により影響されなかった。5.Type B作用群薬物の効果は,high Ca^-Ringer(9mM CaCl_2)で消失され,low Ca^-Ringer液中で増強された。6.Ringer液中のNa^+およびK^+イオンの増減ならびにOuabain(10μM)はType B効果に影響を与えなかった。7.Sucrose-gap法による実験で,Type B作用群薬物は単発刺激による神経節のspike potentialを消失し,比較的大きなsynaptic potentialを生じた。このsynaptic potentialは,300msec以下の刺激間隔で与えた2発刺激により容易にspike potentialに発展した。また高頻度刺激によっても漸次増大するspike potentialが容易に発生した。8.Type B作用群薬物の作用機構についてとくに神経終末からの伝達物質遊離抑制面から考察した。The relationship between the rate of stimulation and effects of drugs on the superior cervical ganglion was investigated. Under the effect of hexamethonium, d-tubocu-rarine, physostigmine or paraoxon, ganglionic transmission was blocked more effectively with higher rate of stimulation than lower rate of stimulation (Type A drugs). Under the effect of adrenaline, noradrenaline, MgCl_2, MnCl_2 or Ca^-deficit Ringer (0.1-0.4mM CaCl_2,) solution, ganglionic transmission was depressed with lower rate of stimulation, but it was restored with higher rate of stimulation (Type B drugs). The inhibitory effect of procaine and propranolol on the ganglionic transmission was not affected by frequency change in transmission (Type C drugs). The effect of adrenaline or noradrenaline, those belong to the type B group, was blocked by phentolamine but not by propranolol. Phentolamine and propranolol had no effect on the action of MgCl_2, MnCl_2 and Ca^-deficit Ringer solution. The effect of the type B group drugs was decreased in Ringer solution containing higher concentration of Ca^ and augmented in Ringer solution containing lower concentration of Ca^. The effect of the type B group drugs was not affected by increasing or decreasing of the concentration of Na^+ or K^+ in Ringer solution. Ouabain (10μM) had no effect on the action of the type B drugs. By means of the sucrose-gap method, the spike potentials evoked by applying single stimulation were transferred to the synaptic potentials under the influence of the type B drugs. When the synaptic potential was evoked by the doubleshock stimuli of an interval shorter than 300 msec, the synaptic potential easily devoloped to the spike potential. In this work, the mechanism of action of the type B drugs was discussed on the standpoint of the depressed transmitter release

Early divergence, broad distribution, and high diversity of animal chitin synthases

Even though chitin is one of themost abundant biopolymers in nature, current knowledge on chitin formation is largely based only on data from fungi and insects. This study reveals unanticipated broad taxonomic distribution and extensive diversification of chitin synthases (CSs) in Metazoa, shedding newlight on the relevance of chitin in animals and suggesting unforeseen complexity of chitin synthesis in many groups. We uncovered robust orthologs to insect type CSs in several representatives of deuterostomes, which generally are not thought to possess chitin. This suggests a broader distribution and function of chitin in this branch of the animal kingdom. We characterize a new CS type present not only in basal metazoans such as sponges and cnidarians but also in several bilaterian representatives. Themost extensive diversification of CSs took place during emergence of lophotrochozoans, the third large group of protostomes next to arthropods and nematodes, resulting in coexistence of up to ten CS paralogs inmolluscs. Independent fusion to different kinds of myosinmotor domains in fungi and lophotrochozoans points toward high relevance of CS interaction with the cytoskeleton for fine-tuned chitin secretion. Given the fundamental role that chitin plays in themorphology of many animals, the here presented CS diversification revealsmany evolutionary complexities. Our findings strongly suggest a very broad andmultifarious occurrence of chitin and question an ancestral role as cuticular component. The molecular mechanisms underlying regulation of animal chitin synthesis are most likely far more complex and diverse than existing data from insects suggest

Developmental studies provide new insights into the evolution of sense organs in Sabellariidae (Annelida)

[Background] Sabellarids, also known as honeycomb or sandcastle worms, when building their tubes, produce chemical signals (free fatty acids) that are responsible for larval settlement and the formation of three-dimensional aggregations. The larval palps and the dorsal hump (becoming the median organ in adults) are presumed to participate in such a substrate selection during settlement. Notably, the sabellariid median organ is an apparently unique organ among annelids that has been attributed with a sensory function and perhaps with some affinities to the nuchal organs of other polychaetes. Nevertheless, detailed investigations of this prominent character complex including ultrastructural examinations are lacking so far.[Results] Our comprehensive investigations provide data about the anterior sensory organs in Sabellariidae and inform about their transformation during pelagic larval development. We used a comparative approach including immunostaining with subsequent confocal laser scanning microscopy (clsm), histological sections as well as electron microscopy in a range of larval and adult stages of two sabellariid species. We find that the neuronal innervation as well as the ultrastructure of the sabellariid ciliary structures along the median organ are highly comparable with that of nuchal organs known from other polychaetes. Furthermore, the myoinhibitory protein (MIP) – a protein known to be also involved into chemo-sensation - was detected in the region of the larval median organ. Moreover, we reveal the presence of an unusual type of photoreceptor as part of the median organ in Idanthyrsus australiensis with a corrugated sensory membrane ultrastructure unlike those observed in the segmental ocelli of other polychaetes.[Conclusions] We are describing for the first time the nuchal organ-like structures in different developmental stages of two species of Sabellariidae. The external morphology, neuronal innervation, developmental fate and ultrastructure of the newly-discovered median organ-based ciliary pits are comparable with the characteristics known for annelid nuchal organs and therefore indicate a homology of both sensory complexes. The presence of myoinhibitory peptide (MIP) in the respective region supports such a hypothesis and exhibits the possibility of an involvement of the entire sabellariid median organ complex, and in particular the prominent ciliated pits, in chemo-sensation.CH was financed by a personal research fellowship (“Rückkehrerstipendium”) from the DFG (HE 7224/2–1). MC was financed by ForBio – the Research School in Biosystematics, funded by the Norwegian Taxonomy Initiative (70184215), the Research Council of Norway (248799) and the Ramón y Cajal program (RYC-2016-20799) funded by Spanish Ministerio de Economía, Industria y Competitividad, Agencia Estatal de Investigación, Comunidad Autónoma de las Islas Baleares and the European Social Fund. The European Commission Taxonomic Initiative SYNTHESYS (ES-TAF-7033) financed a visit to Museo Nacional de Ciencias Naturales, Madrid, Spain. MJB is supported by the Biotechnology and Biological Sciences Research Council (BBSRC, BB/P011357/1) and the Knut and Alice Wallenberg Foundation and the Swedish Research Council (grants awarded to Dan-Eric Nilsson).Peer reviewe

On the Diversity of Phyllodocida (Annelida: Errantia), with a Focus on Glyceridae, Goniadidae, Nephtyidae, Polynoidae, Sphaerodoridae, Syllidae, and the Holoplanktonic Families

Este artículo contiene 65 páginas, 26 figuras, 1 tabla.Phyllodocida is a clade of errantiate annelids characterized by having ventral sensory palps, anterior enlarged cirri, axial muscular proboscis, compound chaetae (if present) with a single ligament, and of lacking dorsolateral folds. Members of most families date back to the Carboniferous, although the earliest fossil was dated from the Devonian. Phyllodocida holds 27 well-established and morphologically homogenous clades ranked as families, gathering more than 4600 currently accepted nominal species. Among them, Syllidae and Polynoidae are the most specious polychaete groups. Species of Phyllodocida are mainly found in the marine benthos, although a few inhabit freshwater, terrestrial and planktonic environments, and occur from intertidal to deep waters in all oceans. In this review, we (1) explore the current knowledge on species diversity trends (based on traditional species concept and molecular data), phylogeny, ecology, and geographic distribution for the whole group, (2) try to identify the main knowledge gaps, and (3) focus on selected families: Alciopidae, Goniadidae, Glyceridae, Iospilidae, Lopadorrhynchidae, Polynoidae, Pontodoridae, Nephtyidae, Sphaerodoridae, Syllidae, Tomopteridae, Typhloscolecidae, and Yndolaciidae. The highest species richness is concentrated in European, North American, and Australian continental shelves (reflecting a strong sampling bias). While most data come from shallow coastal and surface environments most world oceans are clearly under-studied. The overall trends indicate that new descriptions are constantly added through time and that less than 10% of the known species have molecular barcode information availableWe acknowledge support of the publication fees by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI) and the Open Access Publication Funds of the Georg-August-Universität Göttingen. This research was funded by the Spanish “Agencia Estatal de Investigación” (AEI) and the European Funds for Regional Development (FEDER), Research Project PopCOmics (CTM2017-88080) to DM; the Russian Scientific Foundation for Basic Research, grant no. RFBR 18-05-00459 to TAB; Fundação para a Ciência e a Tecnologia (FCT), contract foreseen in the Decree-Law 57/2016 (Nrs 4-6, art. 23), changed by Law 57/2017 to AR and FCT/MCTES to CESAM (UIDP/50017/2020+UIDB/50017/2020) through national funds; São Paulo Research Foundation (FAPESP), fellowship proc. 2007/53040-9 to MVF; Spanish MINECO, AEI, Comunidad Autónoma de las Islas Baleares, European Social Funds and Ramón y Cajal program, RYC-2016-20799 to MC; FCT and ESF (SFRH/BD/131527/2017) through a PhD grant to MALT.Peer reviewe

Perspectives and Integration in SOLAS Science

Why a chapter on Perspectives and Integration in SOLAS Science in this book? SOLAS science by its nature deals with interactions that occur: across a wide spectrum of time and space scales, involve gases and particles, between the ocean and the atmosphere, across many disciplines including chemistry, biology, optics, physics, mathematics, computing, socio-economics and consequently interactions between many different scientists and across scientific generations. This chapter provides a guide through the remarkable diversity of cross-cutting approaches and tools in the gigantic puzzle of the SOLAS realm. Here we overview the existing prime components of atmospheric and oceanic observing systems, with the acquisition of ocean–atmosphere observables either from in situ or from satellites, the rich hierarchy of models to test our knowledge of Earth System functioning, and the tremendous efforts accomplished over the last decade within the COST Action 735 and SOLAS Integration project frameworks to understand, as best we can, the current physical and biogeochemical state of the atmosphere and ocean commons. A few SOLAS integrative studies illustrate the full meaning of interactions, paving the way for even tighter connections between thematic fields. Ultimately, SOLAS research will also develop with an enhanced consideration of societal demand while preserving fundamental research coherency. The exchange of energy, gases and particles across the air-sea interface is controlled by a variety of biological, chemical and physical processes that operate across broad spatial and temporal scales. These processes influence the composition, biogeochemical and chemical properties of both the oceanic and atmospheric boundary layers and ultimately shape the Earth system response to climate and environmental change, as detailed in the previous four chapters. In this cross-cutting chapter we present some of the SOLAS achievements over the last decade in terms of integration, upscaling observational information from process-oriented studies and expeditionary research with key tools such as remote sensing and modelling. Here we do not pretend to encompass the entire legacy of SOLAS efforts but rather offer a selective view of some of the major integrative SOLAS studies that combined available pieces of the immense jigsaw puzzle. These include, for instance, COST efforts to build up global climatologies of SOLAS relevant parameters such as dimethyl sulphide, interconnection between volcanic ash and ecosystem response in the eastern subarctic North Pacific, optimal strategy to derive basin-scale CO2 uptake with good precision, or significant reduction of the uncertainties in sea-salt aerosol source functions. Predicting the future trajectory of Earth’s climate and habitability is the main task ahead. Some possible routes for the SOLAS scientific community to reach this overarching goal conclude the chapter

The James Webb Space Telescope Mission

Twenty-six years ago a small committee report, building on earlier studies, expounded a compelling and poetic vision for the future of astronomy, calling for an infrared-optimized space telescope with an aperture of at least $4m$. With the support of their governments in the US, Europe, and Canada, 20,000 people realized that vision as the $6.5m$ James Webb Space Telescope. A generation of astronomers will celebrate their accomplishments for the life of the mission, potentially as long as 20 years, and beyond. This report and the scientific discoveries that follow are extended thank-you notes to the 20,000 team members. The telescope is working perfectly, with much better image quality than expected. In this and accompanying papers, we give a brief history, describe the observatory, outline its objectives and current observing program, and discuss the inventions and people who made it possible. We cite detailed reports on the design and the measured performance on orbit.Comment: Accepted by PASP for the special issue on The James Webb Space Telescope Overview, 29 pages, 4 figure