A new look at the ventral nerve centre of Sagitta: implications for the phylogenetic position of Chaetognatha (arrow worms) and the evolution of the bilaterian nervous system

<p>Abstract</p> <p>Background</p> <p>The Chaetognatha (arrow worms) are a group of marine carnivores whose phylogenetic relationships are still vigorously debated. Molecular studies have as yet failed to come up with a stable hypothesis on their phylogenetic position. In a wide range of metazoans, the nervous system has proven to provide a wealth of characters for analysing phylogenetic relationships (neurophylogeny). Therefore, in the present study we explored the structure of the ventral nerve centre ("ventral ganglion") in <it>Sagitta setosa </it>with a set of histochemical and immunohistochemical markers.</p> <p>Results</p> <p>In specimens that were immunolabeled for acetylated-alpha tubulin the ventral nerve centre appeared to be a condensed continuation of the peripheral intraepidermal nerve plexus. Yet, synapsin immunolocalization showed that the ventral nerve centre is organized into a highly ordered array of ca. 80 serially arranged microcompartments. Immunohistochemistry against RFamide revealed a set of serially arranged individually identifiable neurons in the ventral nerve centre that we charted in detail.</p> <p>Conclusion</p> <p>The new information on the structure of the chaetognath nervous system is compared to previous descriptions of the ventral nerve centre which are critically evaluated. Our findings are discussed with regard to the debate on nervous system organisation in the last common bilaterian ancestor and with regard to the phylogenetic affinities of this Chaetognatha. We suggest to place the Chaetognatha within the Protostomia and argue against hypotheses which propose a deuterostome affinity of Chaetognatha or a sister-group relationship to all other Bilateria.</p

Immunolocalization of serotonin in Onychophora argues against segmental ganglia being an ancestral feature of arthropods

<p>Abstract</p> <p>Background</p> <p>Onychophora (velvet worms) represent the most basal arthropod group and play a pivotal role in the current discussion on the evolution of nervous systems and segmentation in arthropods. Although there is a wealth of information on the immunolocalization of serotonin (5-hydroxytryptamine, 5-HT) in various euarthropods, as yet no comparable localization data are available for Onychophora. In order to understand how the onychophoran nervous system compares to that of other arthropods, we studied the distribution of serotonin-like immunoreactive neurons and histological characteristics of ventral nerve cords in <it>Metaperipatus blainvillei </it>(Onychophora, Peripatopsidae) and <it>Epiperipatus biolleyi </it>(Onychophora, Peripatidae).</p> <p>Results</p> <p>We demonstrate that paired leg nerves are the only segmental structures associated with the onychophoran nerve cord. Although the median commissures and peripheral nerves show a repeated pattern, their arrangement is independent from body segments characterized by the position of legs and associated structures. Moreover, the somata of serotonin-like immunoreactive neurons do not show any ordered arrangement in both species studied but are instead scattered throughout the entire length of each nerve cord. We observed neither a serially iterated nor a bilaterally symmetric pattern, which is in contrast to the strictly segmental arrangement of serotonergic neurons in other arthropods.</p> <p>Conclusion</p> <p>Our histological findings and immunolocalization experiments highlight the medullary organization of the onychophoran nerve cord and argue against segmental ganglia of the typical euarthropodan type being an ancestral feature of Onychophora. These results contradict <it>a priori </it>assumptions of segmental ganglia being an ancestral feature of arthropods and, thus, weaken the traditional Articulata hypothesis, which proposes a sistergroup relationship of Annelida and Arthropoda.</p

Mechanisms of eye development and evolution of the arthropod visual system: The lateral eyes of myriapoda are not modified insect ommatidia

AbstractThe lateral eyes of Crustacea and Insecta consist of many single optical units, the ommatidia, that are composed of a small, strictly determined and evolutionarily conserved set of cells. In contrast, the eyes of Myriapoda (millipedes and centipedes) are fields of optical units, the lateral ocelli, each of which is composed of up to several hundreds of cells. For many years these striking differences between the lateral eyes of Crustacea/Insecta versus Myriapoda have puzzled evolutionary biologists, as the Myriapoda are traditionally considered to be closely related to the Insecta. The prevailing hypothesis to explain this paradox has been that the myriapod fields of lateral ocelli derive from insect compound eyes by disintegration of the latter into single ommatidia and subsequent fusion of several ommatidia to form multicellular ocelli. To provide a fresh view on this problem, we counted and mapped the arrangement of ocelli during postembryonic development of a diplopod. Furthermore, the arrangement of proliferating cells in the eyes of another diplopod and two chilopods was monitored by labelling with the mitosis marker bromodeoxyuridine. Our results confirm that during eye growth in Myriapoda new elements are added to the side of the eye field, which extend the rows of earlier-generated optical units. This pattern closely resembles that in horseshoe crabs (Chelicerata) and Trilobita. We conclude that the trilobite, xiphosuran, diplopod and chilopod mechanism of eye growth represents the ancestral euarthropod mode of visual-system formation, which raises the possibility that the eyes of Diplopoda and Chilopoda may not be secondarily reconstructed insect eyes

Physiological basis of interactive responses to temperature and salinity in coastal marine invertebrate: Implications for responses to warming

Developing physiological mechanistic models to predict species’ responses to climate‐driven environmental variables remains a key endeavor in ecology. Such approaches are challenging, because they require linking physiological processes with fitness and contraction or expansion in species’ distributions. We explore those links for coastal marine species, occurring in regions of freshwater influence (ROFIs) and exposed to changes in temperature and salinity. First, we evaluated the effect of temperature on hemolymph osmolality and on the expression of genes relevant for osmoregulation in larvae of the shore crab Carcinus maenas. We then discuss and develop a hypothetical model linking osmoregulation, fitness, and species expansion/contraction toward or away from ROFIs. In C. maenas, high temperature led to a threefold increase in the capacity to osmoregulate in the first and last larval stages (i.e., those more likely to experience low salinities). This result matched the known pattern of survival for larval stages where the negative effect of low salinity on survival is mitigated at high temperatures (abbreviated as TMLS). Because gene expression levels did not change at low salinity nor at high temperatures, we hypothesize that the increase in osmoregulatory capacity (OC) at high temperature should involve post‐translational processes. Further analysis of data suggested that TMLS occurs in C. maenas larvae due to the combination of increased osmoregulation (a physiological mechanism) and a reduced developmental period (a phenological mechanisms) when exposed to high temperatures. Based on information from the literature, we propose a model for C. maenas and other coastal species showing the contribution of osmoregulation and phenological mechanisms toward changes in range distribution under coastal warming. In species where the OC increases with temperature (e.g., C. maenas larvae), osmoregulation should contribute toward expansion if temperature increases; by contrast in those species where osmoregulation is weaker at high temperature, the contribution should be toward range contraction

Serotonin immunoreactive interneurons in the brain of the Remipedia: new insights into the phylogenetic affinities of an enigmatic crustacean taxon

BACKGROUND: Remipedia, a group of homonomously segmented, cave-dwelling, eyeless arthropods have been regarded as basal crustaceans in most early morphological and taxonomic studies. However, molecular sequence information together with the discovery of a highly differentiated brain led to a reconsideration of their phylogenetic position. Various conflicting hypotheses have been proposed including the claim for a basal position of Remipedia up to a close relationship with Malacostraca or Hexapoda. To provide new morphological characters that may allow phylogenetic insights, we have analyzed the architecture of the remipede brain in more detail using immunocytochemistry (serotonin, acetylated α-tubulin, synapsin) combined with confocal laser-scanning microscopy and image reconstruction techniques. This approach allows for a comprehensive neuroanatomical comparison with other crustacean and hexapod taxa. RESULTS: The dominant structures of the brain are the deutocerebral olfactory neuropils, which are linked by the olfactory globular tracts to the protocerebral hemiellipsoid bodies. The olfactory globular tracts form a characteristic chiasm in the center of the brain. In Speleonectes tulumensis, each brain hemisphere contains about 120 serotonin immunoreactive neurons, which are distributed in distinct cell groups supplying fine, profusely branching neurites to 16 neuropilar domains. The olfactory neuropil comprises more than 300 spherical olfactory glomeruli arranged in sublobes. Eight serotonin immunoreactive neurons homogeneously innervate the olfactory glomeruli. In the protocerebrum, serotonin immunoreactivity revealed several structures, which, based on their position and connectivity resemble a central complex comprising a central body, a protocerebral bridge, W-, X-, Y-, Z-tracts, and lateral accessory lobes. CONCLUSIONS: The brain of Remipedia shows several plesiomorphic features shared with other Mandibulata, such as deutocerebral olfactory neuropils with a glomerular organization, innervations by serotonin immunoreactive interneurons, and connections to protocerebral neuropils. Also, we provided tentative evidence for W-, X-, Y-, Z-tracts in the remipedian central complex like in the brain of Malacostraca, and Hexapoda. Furthermore, Remipedia display several synapomorphies with Malacostraca supporting a sister group relationship between both taxa. These homologies include a chiasm of the olfactory globular tract, which connects the olfactory neuropils with the lateral protocerebrum and the presence of hemiellipsoid bodies. Even though a growing number of molecular investigations unites Remipedia and Cephalocarida, our neuroanatomical comparison does not provide support for such a sister group relationship

Contrasting offspring responses to variation in salinity and temperature among populations of a coastal crab: A maladaptive ecological surprise?

Current understanding of species capacities to respond to climate change is limited by the amount of information available about intraspecific variation in the responses. Therefore, we quantified between- and within- population variation in larval performance (survival, development, and growth to metamorphosis) of the shore crab Carcinus maenas in response to key environmental drivers (temperature, salinity) in 2 populations from regions with contrasting salinities (32-33 PSU: Helgoland, North Sea; 16-20 PSU: Kerteminde, Baltic Sea). We also accounted for the effect(s) of salinity experienced during embryogenesis, which differs between populations. We found contrasting patterns between populations and embryonic salinity conditions. In the Helgoland population, we observed a strong thermal mitigation of low salinity stress (TMLS) for all performance indicators, when embryos were kept in seawater. The negative effects of low salinity on survival were mitigated at increased temperatures; only at high temperatures were larvae exposed to low salinity able to sustain high growth rates and reduced developmental time, thereby metamorphosing with comparable levels of carbon and nitrogen to those reared in seawater. By contrast, larvae from the Kerteminde population showed a detrimental effect of low salinity, consistent with a maladaptive response and a weak TMLS. Low salinity experienced during embryogenesis pre-empted the development of TMLS in both populations, and reduced survival for the Kerteminde population, which is exposed to low salinity. Our study emphasises the importance of evaluating species responses to variation in temperature and salinity across populations; the existence of maladaptive responses and the importance of the maternal habitat should not be underestimated

Neurogenesis in the central olfactory pathway of adult decapod crustaceans: development of the neurogenic niche in the brains of procambarid crayfish

<p>Abstract</p> <p>Background</p> <p>In the decapod crustacean brain, neurogenesis persists throughout the animal's life. After embryogenesis, the central olfactory pathway integrates newborn olfactory local and projection interneurons that replace old neurons or expand the existing population. In crayfish, these neurons are the descendants of precursor cells residing in a neurogenic niche. In this paper, the development of the niche was documented by monitoring proliferating cells with S-phase-specific markers combined with immunohistochemical, dye-injection and pulse-chase experiments.</p> <p>Results</p> <p>Between the end of embryogenesis and throughout the first post-embryonic stage (POI), a defined transverse band of mitotically active cells (which we will term 'the deutocerebral proliferative system' (DPS) appears. Just prior to hatching and in parallel with the formation of the DPS, the anlagen of the niche appears, closely associated with the vasculature. When the hatchling molts to the second post-embryonic stage (POII), the DPS differentiates into the lateral (LPZ) and medial (MPZ) proliferative zones. The LPZ and MPZ are characterized by a high number of mitotically active cells from the beginning of post-embryonic life; in contrast, the developing niche contains only very few dividing cells, a characteristic that persists in the adult organism.</p> <p>Conclusions</p> <p>Our data suggest that the LPZ and MPZ are largely responsible for the production of new neurons in the early post-embryonic stages, and that the neurogenic niche in the beginning plays a subordinate role. However, as the neuroblasts in the proliferation zones disappear during early post-embryonic life, the neuronal precursors in the niche gradually become the dominant and only mechanism for the generation of new neurons in the adult brain.</p

Quantifying the portfolio of larval responses to salinity and temperature in a coastal-marine invertebrate: a cross population study along the European coast

portfolio available for a species to cope with and mitigate effects of climate change. Here, we quantified variation in larval survival and physiological rates of Carcinus maenas among populations occurring in distant or contrasting habitats (Cádiz: Spain, Helgoland: North Sea, Kerteminde: Baltic Sea). During the reproductive season, we reared larvae of these populations, in the laboratory, under a combination of several temperatures (15–24 °C) and salinities (25 and 32.5 PSU). In survival, all three populations showed a mitigating effect of high temperatures at lower salinity, with the strongest pattern for Helgoland. However, Cádiz and Kerteminde differed from Helgoland in that a strong thermal mitigation did not occur for growth and developmental rates. For all populations, oxygen consumption rates were driven only by temperature; hence, these could not explain the growth rate depression found at lower salinity. Larvae from Cádiz, reared in seawater, showed increased survival at the highest temperature, which differs from Helgoland (no clear survival pattern), and especially Kerteminde (decreased survival at high temperature). These responses from the Cádiz population correspond with the larval and parental habitat (i.e., high salinity and temperature) and may reflect local adaptation. Overall, along the European coast, C. maenas larvae showed a diversity of responses, which may enable specific populations to tolerate warming and subsidise more vulnerable populations. In such case, C. maenas would be able to cope with climate change through a spatial portfolio effect

Brain architecture of the largest living land arthropod, the Giant Robber Crab Birgus latro (Crustacea, Anomura, Coenobitidae): evidence for a prominent central olfactory pathway?

<p>Abstract</p> <p>Background</p> <p>Several lineages within the Crustacea conquered land independently during evolution, thereby requiring physiological adaptations for a semi-terrestrial or even a fully terrestrial lifestyle. <it>Birgus latro </it>Linnaeus, 1767, the giant robber crab or coconut crab (Anomura, Coenobitidae), is the largest land-living arthropod and inhabits Indo-Pacific islands such as Christmas Island. <it>B. latro </it>has served as a model in numerous studies of physiological aspects related to the conquest of land by crustaceans. From an olfactory point of view, a transition from sea to land means that molecules need to be detected in gas phase instead of in water solution. Previous studies have provided physiological evidence that terrestrial hermit crabs (Coenobitidae) such as <it>B. latro </it>have a sensitive and well differentiated sense of smell. Here we analyze the brain, in particular the olfactory processing areas of <it>B. latro</it>, by morphological analysis followed by 3 D reconstruction and immunocytochemical studies of synaptic proteins and a neuropeptide.</p> <p>Results</p> <p>The primary and secondary olfactory centers dominate the brain of <it>B. latro </it>and together account for ca. 40% of the neuropil volume in its brain. The paired olfactory neuropils are tripartite and composed of more than 1,000 columnar olfactory glomeruli, which are radially arranged around the periphery of the olfactory neuropils. The glomeruli are innervated ca. 90,000 local interneurons and ca. 160,000 projection neurons per side. The secondary olfactory centers, the paired hemiellipsoid neuropils, are targeted by the axons of these olfactory projection neurons. The projection neuron axonal branches make contact to ca. 250.000 interneurons (per side) associated with the hemiellipsoid neuropils. The hemiellipsoid body neuropil is organized into parallel neuropil lamellae, a design that is quite unusual for decapod crustaceans. The architecture of the optic neuropils and areas associated with antenna two suggest <it>that B. latro </it>has visual and mechanosensory skills that are comparable to those of marine Crustacea.</p> <p>Conclusions</p> <p>In parallel to previous behavioral findings that <it>B. latro </it>has aerial olfaction, our results indicate that their central olfactory pathway is indeed most prominent. Similar findings from the closely related terrestrial hermit crab <it>Coenobita clypeatus </it>suggest that in Coenobitidae, olfaction is a major sensory modality processed by the brain, and that for these animals, exploring the olfactory landscape is vital for survival in their terrestrial habitat. Future studies on terrestrial members of other crustacean taxa such as Isopoda, Amphipoda, Astacida, and Brachyura will shed light on how frequently the establishment of an aerial sense of olfaction evolved in Crustacea during the transition from sea to land. Amounting to ca. 1,000,000, the numbers of interneurons that analyse the olfactory input in <it>B. latro </it>brains surpasses that in other terrestrial arthropods, as e.g. the honeybee <it>Apis mellifera </it>or the moth <it>Manduca sexta</it>, by two orders of magnitude suggesting that <it>B. latro </it>in fact is a land-living arthropod that has devoted a substantial amount of nervous tissue to the sense of smell.</p

Persönlichkeit und Entspannung vs. Ablenkung beim Zahnarzt

Die vorliegende Studie untersuchte den Einfluß von hoch und niedrig ausgeprägter Zahnbehandlungsangst und den Einfluß der Intervention durch ein Entspannungs- und Ablenkungsverfahren (und Kontrolle) bei der Antizipation einer Zahnbehandlung. Die Teilnehmer waren 49 Patienten einer Giessener Zahnarztpraxis, die parallelisiert nach Alter und Geschlecht auf die Bedingungen aufgeteilt wurden. Anders als in anderen Studien wurden die verschiedenen Meßaspekte in umfassender Weise betrachtet: es wurden peripher-physiologische, immunologische, subjektiv-verbale und behaviorale Indikatoren herangezogen. Außerdem wurde der Einfluß der Ausprägung der Zahnbehandlungsangst und der Art der Experimentalbedingung auf das aktuelle Coping-verhalten untersucht. Es zeigte sich, dass peripher-physiologische und immunologische Indikatoren nicht von der Ausprägung der Zahnbehandlungsangst beeinflußt waren. Allerdings zeigten sich subjektiv-verbale Unterschiede: Hochängstliche schätzten ihre körperliche Erregung höher ein als Niedrigängstliche, was auf eine Tendenz der Hochängstlichen zurückgeführt wurde, körperliche Reaktionen stärker als Niedrigängstliche zu beachten. Weiterhin zeigten sich behaviorale Unterschiede: Hochängstliche wirkten starrer und wurden als ängstlicher, weniger entspannt und weniger kooperativ eingeschätzt. Allerdings hat sich das aktuelle Copingverhalten von Hoch- und Niedrigängstlichen nicht unterschieden. Die Ergebnisse sprechen jedoch für eine Wirksamkeit des Entspannungsverfahrens, vor allem bei Hochängstlichen. Zwar haben sich die Experimentalgruppen, wie auch die Persönlichkeitsgruppen, nicht in peripher-physiologischen und immunologischen Indikatoren unterschieden, allerdings verbesserte das Entspannungsverfahren die subjektive Befindlichkeit besonders bei den Hochängstlichen. Auch behaviorale Unterschiede weisen auf die Wirksamkeit des Entspannungsverfahrens hin, dass sich von der Wirkung des Ablenkungsverfahrens und der Kontrollbedingung unterscheidet. Die Ergebnisse zur Wirksamkeit des Ablenkungsverfahrens sind weniger eindeutig. Das Ablenkungsverfahren unterscheidet sich in einigen Punkten vom Entspannungsverfahren, allerdings kaum von der Kontrollbedingung. Dies wird der verwendeten Technik (Audio-Präsentation) angelastet, die vergleichsweise wenig Ablenkungskraft hat. Allerdings beeinflußte das Ablenkungsverfahren das aktuelle Copingverhalten während der Zahnbehandlung