Dorsal Nerve Cord Research Articles

Neuroanatomy of a Dactylogyrid monogenean, from gold fish Carassius auratus, Nilsson, from Meerut (U. P.), India

Chemical named 5-bromo indoxyl acetate has been used to describe the nervous system of anoviparous Dactylogyridmonogenean PellucidhaptorPrice and Mizelle (1964), a gill parasite of Carassius auratus. Central nervous system consists of paired cerebral ganglia from which anterior and posterior neuronal pathways arise. These neuronal pathways are interlinked by cross connectives and commissures. Paired dorsal, ventral and lateral nerve cords emanate from the cerebral ganglia, connected at intervals by transverse connectives. Huge arrangement of dorsal, ventral and lateral nerve cords and their innervations have been examined. Peripheral nervous system (PNS) includes innervations of the alimentary tract, reproductive organs and attachment organs (anterior adhesive areas and haptor). Both the CNS and PNS are bilaterally symmetrical, and better developed ventrally than laterally and dorsally.

Keeping amphioxus in the laboratory: an update on available husbandry methods

Cephalochordates, commonly known as amphioxus or lancelets, are small, marine animals that can be found in coastal habitats of temperate, subtropical, and tropical waters. Together with vertebrates and tunicates, the cephalochordates belong to the chordate phylum, whose members are characterized by a number of conserved morphological features, such as a dorsal nerve cord, a notochord, a pharynx, a segmented musculature as well as a post-anal tail. Due to their basal position within the phylum, cephalochordates have become essential models for studying the evolutionary origin and diversification of vertebrates. Here, we present the currently available methods for maintaining and rearing cephalochordates in a laboratory environment, focusing on five species: the European amphioxus (Branchiostoma lanceolatum), the Florida amphioxus (Branchiostoma floridae), the Chinese amphioxus (Branchiostoma belcheri), the Japanese amphioxus (Branchiostoma japonicum), and the Bahamas lancelet (Asymmetron lucayanum). In addition to reviewing the protocols for capture, transport, aquaculture, and feeding of adults, we discuss methods for controlling gonad development and spawning, as well as for growing embryos and larvae. This information is complemented by observations from our animal facility on the European amphioxus (Branchiostoma lanceolatum). In sum, this work summarizes the latest advances in cephalochordate animal husbandry and highlights challenges for improving the use of these animals as laboratory model systems.

T-Box Genes and Developmental Gene Regulatory Networks in Ascidians.

Ascidians are invertebrate chordates with a biphasic life cycle characterized by a dual body plan that displays simplified versions of chordate structures, such as a premetamorphic 40-cell notochord topped by a dorsal nerve cord and postmetamorphic pharyngeal slits. These relatively simple chordates are characterized by rapid development, compact genomes and ease of transgenesis, and thus provide the opportunity to rapidly characterize the genomic organization, developmental function, and transcriptional regulation of evolutionarily conserved gene families. This review summarizes the current knowledge on members of the T-box family of transcription factors in Ciona and other ascidians. In both chordate and nonchordate animals, these genes control a variety of morphogenetic processes, and their mutations are responsible for malformations and developmental defects in organisms ranging from flies to humans. In ascidians, T-box transcription factors are required for the formation and specialization of essential structures, including notochord, muscle, heart, and differentiated neurons. In recent years, the experimental advantages offered by ascidian embryos have allowed the rapid accumulation of a wealth of information on the molecular mechanisms that regulate the expression of T-box genes. These studies have also elucidated the strategies employed by these transcription factors to orchestrate the appropriate spatial and temporal deployment of the numerous target genes that they control.

Metamorphosis and definitive organogenesis in the holothurian Apostichopus japonicus

The structure of the late doliolaria, pentactula and 1-month-old juvenile of the holothurian Apostichopus japonicus was studied using light microscopy and 3D reconstruction methods. It was shown that metamorphosis in this species consists in the reorganization of the shape of the body and the destruction of provisional organs. The late doliolaria has a spindle-like form, ciliary rings and hyaline spheres shifted relative to the anterior-posterior axis of the body. Some provisional organs (ciliary rings, hyaline spheres) are destroyed during settlement, and others (hydropore and hydroporic canal) remain after metamorphosis. Definitive organogenesis in A. japonicus begins long before metamorphosis. The late doliolaria already has well-developed water-vascular and digestive systems, and the ectoneural part of the nervous system. Muscle and hemal systems begin to form in the pentactula. Moreover, the calcareous ring and connective tissue part of the body wall develop at this stage. The pentactula has anlages of the hyponeural part of the nervous system, which form in the mid-ventral and dorsal nerve cords. The hemal ring of the pentactula is located on the inner wall of the water-vascular ring. It remains unclosed in the left ventral radius. One-month-old juveniles have all the major organ systems except respiratory and reproductive systems. The hemal vessels of the intestine are well developed and begin to form the rete mirabile. Differentiation of the intestine into regions due to differential specialization of the enterocytes begins in 1-month-old juveniles. Obviously, emergence of new types of enterocytes enables the animal to consume a wider range of food items and indicates its increased feeding activity.

Presynaptic and postsynaptic regulation of muscle contractions in the ascarid nematodeAscaris suum: a target for drug action

The aim of this study was to determine the role in contractions of postsynaptic nicotinic acetylcholine (nACh) and γ-aminobutyric acid (GABA) receptors, in the bag region of Ascaris suum muscle cells, as well as the role of synaptic receptors between interneurons and motor neurons in the dorsal and ventral nerve cord. We have measured the isometric contractions of isolated segments of A. suum, with or without the nerve cord (dorsal or ventral). Contractions were caused by increasing concentrations of ACh or by electrical field stimulation (EFS). Based on our results, the presence of the nerve cord is essential for the contractile effects of ACh. The EC50 value of ACh for innervated muscle strips was 10.88μm. Unlike intact (innervated) preparations, there was no contraction of the muscle flaps when the nerve cord was mechanically removed. Furthermore, continuous EFS produced stable contractions of innervated muscle strips, but they are not sensitive to mecamylamine (100μm). However, GABA (30μm) significantly inhibited the EFS-induced contractions. EFS with the same characteristics did not cause muscle contractions of denervated muscle strips, but EFS with a wider pulse induced the increasing of tone and irregular contractions. These contractions were completely insensitive to the effect of GABA. The EC50 for ACh did not differ between the dorsal and ventral segments (9.83μm and 9.45μm), while GABA exhibited features of competitive and non-competitive antagonists, regardless of whether it acted on the dorsal or ventral segments of A. suum. It is obvious that drugs will be more effective if they act on both the synaptic and extrasynaptic nACh and GABA receptors.

Transcriptional Control of Synaptic Remodeling through Regulated Expression of an Immunoglobulin Superfamily Protein

Neural circuits are actively remodeled during brain development, but the molecular mechanisms that trigger circuit refinement are poorly understood. Here, we describe a transcriptional program in C.elegans that regulates expression of an Ig domain protein, OIG-1, to control the timing of synaptic remodeling. DD GABAergic neurons reverse polarity during larval development by exchanging the locations of pre- and postsynaptic components. In newly born larvae, DDs receive cholinergic inputs in the dorsal nerve cord. These inputs are switched to the ventral side by the end of the first larval (L1) stage. VD class GABAergic neurons are generated in the late L1 and are postsynaptic to cholinergic neurons in the dorsal nerve cord but do not remodel. We investigated remodeling of the postsynaptic apparatus in DD and VD neurons using targeted expression of the acetylcholine receptor (AChR) subunit, ACR-12::GFP. We determined that OIG-1 antagonizes the relocation of ACR-12 from the dorsal side in L1 DD neurons. During the L1/L2 transition, OIG-1 is downregulated in DD neurons by the transcription factor IRX-1/Iroquois, allowing the repositioning of synaptic inputs to the ventral side. In VD class neurons, which normally do not remodel, the transcription factor UNC-55/COUP-TF turns off IRX-1, thus maintaining high levels of OIG-1 to block the removal of dorsally located ACR-12 receptors. OIG-1 is secreted from GABA neurons, but its anti-plasticity function is cell autonomous and may not require secretion. Our study provides a novel mechanism by which synaptic remodeling is set in motion through regulated expression of an Ig domain protein.

Spatiotemporal control of a novel synaptic organizer molecule.

Synapse formation is a process tightly controlled in space and time. How gene regulatory mechanisms specify spatial and temporal aspects of synapse formation is not well understood. In the nematode C.elegans, two subtypes of the D-type inhibitory motor neuron (MN) classes, the dorsal D (DD) and ventral D (VD) neurons, extend axons along both the dorsal and ventral nerve cords 1. The embryonically generated DD MNs initially innervate ventral muscles in the first (L1) larval stage and receive their synaptic input from cholinergic MNs in the dorsal cord. They rewire by the end of the L1 molt to innervate dorsal muscles and to be innervated by newly formed ventral cholinergic MNs 1. VD MNs develop after the L1 molt; they take over the innervation of ventral muscles and receive their synaptic input from dorsal cholinergic MNs. We show here that the spatiotemporal control of synaptic wiring of the D-type neurons is controlled by an intersectional transcriptional strategy in which the UNC-30 Pitx-type homeodomain transcription factor acts together in embryonic and early larval stages with the temporally controlled LIN-14 transcription factor to prevent premature synapse rewiring of the DD MNs and, together with the UNC-55 nuclear hormone receptor, to prevent aberrant VD synaptic wiring in later larval and adult stages. A key effector of this intersectional transcription factor combination is a novel synaptic organizer molecule, the single immunoglobulin domain protein OIG-1. OIG-1 is perisynaptically localized along the synaptic outputs of the D-type MNs in a temporally controlled manner and is required for appropriate selection of both pre- and post-synaptic partners.

Hybrids between the Florida amphioxus (Branchiostoma floridae) and the Bahamas lancelet (Asymmetron lucayanum): developmental morphology and chromosome counts.

The cephalochordate genera Branchiostoma and Asymmetron diverged during the Mesozoic Era. In spite of the long separation of the parental clades, eggs of the Florida amphioxus, B. floridae, when fertilized with sperm of the Bahamas lancelet, A. lucayanum (and vice versa), develop through embryonic and larval stages. The larvae reach the chordate phylotypic stage (i.e., the pharyngula), characterized by a dorsal nerve cord, notochord, perforate pharynx, and segmented trunk musculature. After about 2 weeks of larval development, the hybrids die, as do the A. lucayanum purebreds, although all were eating the same algal diet that sustains B. floridae purebreds through adulthood in the laboratory; it is thus unclear whether death of the hybrids results from incompatible parental genomes or an inadequate diet. The diploid chromosome count in A. lucayanum and B. floridae purebreds is, respectively, 34 and 38, whereas it is 36 in hybrids in either direction. The hybrid larvae exhibit several morphological characters intermediate between those of the parents, including the size of the preoral ciliated pit and the angles of deflection of the gill slits and anus from the ventral midline. Based on the time since the two parent clades diverged (120 or 160 million years, respectively, by nuclear and mitochondrial gene analysis), the cross between Branchiostoma and Asymmetron is the most extreme example of hybridization that has ever been unequivocally demonstrated among multicellular animals.

The zinc finger RNA binding protein, ZFR, contributes to axon guidance in Caenorhabditis elegans

ZFR is an ancient and highly conserved chromosome-associated protein from nematodes to mammals, embryologically expressed in most species, with the exception of the nematode Caenorhabditis elegans. The ZFR encodes zinc and RNA binding protein, and in rat, the nuclear-cytoplasmic shuttling ZFR has been found with transport and translation-associated RNA granule-like structures in the somatodendritic compartments of hippocampal neurons. The majority of axons cross the midline before projecting to their contralateral synaptic target and this crossing decision is under tight control. Molecular factors contributing to these processes have been identified, although the mechanisms are not fully understood. In this study, we tested the role of ceZFR in axon guidance using ceZfr RNAi-treated animals to analyse axon midline crossing, axon fasciculation and cord commissures. In adult stages, RNAi-induced depletion of the ceZfr transcript leads to several phenotypes related to axon guidance. A midline crossing defect was observed in the ventral nerve cord (VNC) in axon type D, DD/VD motoneuron axons and axon type 1, interneuron axons. We further detected a dorsal nerve cord (DNC) axon fasciculation. Some ceZfr RNAi-treated animals revealed that cord commissures fail to reach their synaptic target. We provide evidence that ceZFR has a role in axon guidance. When Zfr was depleted by RNAi, the phenotypes are characterized by defects in axon midline crossing, axon defasciculation and cord commissures. Our results thus support the hypothesis that ZFR has essential roles during neurogenesis, and could support early steps of RNA transport and localization through RNA granule formation in the nucleus and/or to their nucleo-cytoplasmic shuttling.

Proteolytic Processing of the Extracellular Scaffolding Protein LEV-9 Is Required for Clustering Acetylcholine Receptors

Correct positioning of neurotransmitter-gated receptors at postsynapses is essential for synaptic transmission. At Caenorhabditis elegans neuromuscular junctions, clustering of levamisole-sensitive acetylcholine receptors (L-AChRs) requires the muscle-secreted scaffolding protein LEV-9, a multidomain factor containing complement control protein (CCP) modules. Here we show that LEV-9 needs to be cleaved at its C terminus to exert its function. LEV-9 cleavage is not required for trafficking nor secretion but directly controls scaffolding activity. The cleavage site is evolutionarily conserved, and post-translational cleavage ensures the structural and functional decoupling between different isoforms encoded by the lev-9 gene. Data mining indicates that most human CCP-containing factors are likely cleaved C-terminally from CCP tandems, suggesting that not only domain architectures but also cleavage location can be conserved in distant architecturally related proteins.

Genetic Interactions Between UNC-17/VAChT and a Novel Transmembrane Protein in Caenorhabditis elegans

The unc-17 gene encodes the vesicular acetylcholine transporter (VAChT) in Caenorhabditis elegans. unc-17 reduction-of-function mutants are small, slow growing, and uncoordinated. Several independent unc-17 alleles are associated with a glycine-to-arginine substitution (G347R), which introduces a positive charge in the ninth transmembrane domain (TMD) of UNC-17. To identify proteins that interact with UNC-17/VAChT, we screened for mutations that suppress the uncoordinated phenotype of UNC-17(G347R) mutants. We identified several dominant allele-specific suppressors, including mutations in the sup-1 locus. The sup-1 gene encodes a single-pass transmembrane protein that is expressed in a subset of neurons and in body muscles. Two independent suppressor alleles of sup-1 are associated with a glycine-to-glutamic acid substitution (G84E), resulting in a negative charge in the SUP-1 TMD. A sup-1 null mutant has no obvious deficits in cholinergic neurotransmission and does not suppress unc-17 mutant phenotypes. Bimolecular fluorescence complementation (BiFC) analysis demonstrated close association of SUP-1 and UNC-17 in synapse-rich regions of the cholinergic nervous system, including the nerve ring and dorsal nerve cords. These observations suggest that UNC-17 and SUP-1 are in close proximity at synapses. We propose that electrostatic interactions between the UNC-17(G347R) and SUP-1(G84E) TMDs alter the conformation of the mutant UNC-17 protein, thereby restoring UNC-17 function; this is similar to the interaction between UNC-17/VAChT and synaptobrevin.

Hagfish, Genome Duplications, and RFamide Neuropeptide Evolution

As a phylum, the chordates are remarkably similar in terms of body plan design compared with many of the other major phyla of metazoans. The minimal features for placement within the phylum are a dorsal nerve cord and a notochord and pharyngeal gills slits at some point in the developmental life history of the organism (1). As shown in Fig. 1A, chordates can be initially segregated based on whether the organism lacks an internal cartilaginous or bony skeleton (protochordates; such as Amphioxus or the tunicates) or have an internal skeleton (subphylum Vertebrata). The vertebrates can be further separated into those organisms that lack a hinged jaw (superclass Agnatha; such as the hagfishes or lampreys) and those organisms that have a hinged jaw (superclass Gnathostoma; such as the cartilaginous fishes, bony fishes, amphibians, reptiles, birds, and mammals). As indicated from the fossil record, the ancestral protochordates most likely emerged during the Cambrian Period, some 550 million years ago (MYA) (2), and were followed in relatively quick succession by the ancestral agnathan vertebrates (late Ordovician Period; 450 MYA), and later the ancestral gnathostomes (Silurian Period 420 MYA) (3). Although the external and internal morphological design of the vertebrates (agnathans and gnathostomes) is based on common themes, there are features of the chordates that clearly have undergone considerable change during the evolution of the phylum. Two of these features, the diversity of gene families, and the interactions between the brain and pituitary, are considered in the article by Osugi et al. (4) in this issue of Endocrinology on the evolution, characterization, and localization of RFamide peptides in the central nervous system (CNS) of a hagfish. Why study hagfish, and what information on neuropeptide gene family evolution and brain/pituitary interactions can be gleaned from these organisms?

A Perimotor Framework Reveals Functional Segmentation in the Motoneuronal Network Controlling Locomotion inCaenorhabditis elegans

The neuronal connectivity dataset of the nematode Caenorhabditis elegans attracts wide attention from computational neuroscientists and experimentalists. However, the dataset is incomplete. The ventral and dorsal nerve cords of a single nematode were reconstructed halfway along the body and the posterior data are missing, leaving 21 of 75 motoneurons of the locomotor network with partial or no connectivity data. Using a new framework for network analysis, the perimotor space, we identified rules of connectivity that allowed us to approximate the missing data by extrapolation. Motoneurons were mapped into perimotor space in which each motoneuron is located according to the muscle cells it innervates. In this framework, a pattern of iterative connections emerges which includes most (0.90) of the connections. We identified a repeating unit consisting of 12 motoneurons and 12 muscle cells. The cell bodies of the motoneurons of such a unit are not necessarily anatomical neighbors and there is no obvious anatomical segmentation. A connectivity model, composed of six repeating units, is a description of the network that is both simplified (modular and without noniterative connections) and more complete (includes the posterior part) than the original dataset. The perimotor framework of observed connectivity and the segmented connectivity model give insights and advance the study of the neuronal infrastructure underlying locomotion in C. elegans. Furthermore, we suggest that the tools used herein may be useful to interpret, simplify, and represent connectivity data of other motor systems.

MADD-4 Is a Secreted Cue Required for Midline-Oriented Guidance in Caenorhabditis elegans

The netrins and slits are two families of widely conserved cues that guide axons and cells along the dorsal-ventral (D-V) axis of animals. These cues typically emanate from the dorsal or ventral midlines and provide spatial information to migrating cells byforming gradients along the D-V axis. Some cell types, however, extend processes to both the dorsal and ventral midlines, suggesting the existence of additional guidance cues that are secreted from both midlines. Here, we report that a previously uncharacterized protein called MADD-4 is secreted by the dorsal and ventral nerve cords of the nematode C.elegans to attract sensory axons and musclemembrane extensions called muscle arms. MADD-4's activity is dependent on UNC-40/DCC, a netrin receptor, which functions cell-autonomously to direct membrane extension. The biological role of MADD-4 orthologs, including ADAMTSL1 and 3 in mammals, is unknown. MADD-4 may therefore represent the founding member of a family of guidance proteins.

Neuroanatomy of Cornudescoides kulkarnii n. sp., a gill parasite of Mystus vittatus in Meerut (UP), India

Chemical named 5-bromo indoxyl acetate has been used to describe the nervous system of a viviparous monogenean Cornudescoides Kulkarni (1969), a gill parasite of Mystus vittatus. Central nervous system consists of paired cerebral ganglia from which anterior and posterior neuronal pathways arise. These neuronal pathways are interlinked by cross connectives and commissures. Paired dorsal, ventral and lateral nerve cords emanate from the cerebral ganglia, connected at intervals by transverse connectives. Huge arrangement of dorsal, ventral and lateral nerve cords and their innervations have been examined. Peripheral nervous system (PNS) includes innervations of the alimentary tract, reproductive organs and attachment organs (anterior adhesive areas and haptor). Both the CNS and PNS are bilaterally symmetrical, and better developed ventrally than laterally and dorsally.

Neuron-specific proteotoxicity of mutant ataxin-3 in C. elegans : rescue by the DAF-16 and HSF-1 pathways

The risk of developing neurodegenerative diseases increases with age. Although many of the molecular pathways regulating proteotoxic stress and longevity are well characterized, their contribution to disease susceptibility remains unclear. In this study, we describe a new Caenorhabditis elegans model of Machado-Joseph disease pathogenesis. Pan-neuronal expression of mutant ATXN3 leads to a polyQ-length dependent, neuron subtype-specific aggregation and neuronal dysfunction. Analysis of different neurons revealed a pattern of dorsal nerve cord and sensory neuron susceptibility to mutant ataxin-3 that was distinct from the aggregation and toxicity profiles of polyQ-alone proteins. This reveals that the sequences flanking the polyQ-stretch in ATXN3 have a dominant influence on cell-intrinsic neuronal factors that modulate polyQ-mediated pathogenesis. Aging influences the ATXN3 phenotypes which can be suppressed by the downregulation of the insulin/insulin growth factor-1-like signaling pathway and activation of heat shock factor-1.

A specific antibody to neuropeptide AF1 (KNEFIRFamide) recognizes a small subset of neurons in Ascaris suum: differences from Caenorhabditis elegans

A monoclonal antibody, AF1-003, highly specific to the Ascaris suum neuropeptide AF1 (KNEFIRFamide), was generated. This antibody binds strongly to AF1 and extremely weakly to other peptides with C-terminal FIRFamide: AF5 (SGKPTFIRFamide), AF6 (FIRFamide), and AF7 (AGPRFIRFamide). It does not recognize 35 other AF (A. suum FMRFamide-like) peptides at the highest concentration tested, nor does it recognize FMRFamide. When crude peptide extracts of A. suum are fractionated by two-step HPLC, the only fractions recognized by AF1-003 are those comigrating with synthetic AF1. By immunocytochemistry, antibody AF1-003 recognizes a small subset of the 298 neurons of A. suum: these include the paired URX and RIP neurons, two pairs of lateral ganglion neurons in the head, and the unpaired PQR and PDA or -B tail neurons that send processes to the head along the dorsal and ventral nerve cords, respectively. AF1 immunoreactivity is also seen in three pairs of pharyngeal neurons. Mass spectroscopy (MS) shows the presence of AF1 in the head, pharynx, and dorsal and ventral nerve cords. In A. suum, the neurons that contain AF1 show little overlap with neurons that express green fluorescent protein constructs targeting the flp-8 gene, which encodes AF1 in Caenorhabditis elegans (Kim and Li [2004] J. Comp. Neurol. 475:540-550); the URX neurons express AF1 in both species, but, in C. elegans, flp-8 expression was not detected in RIP, PQR, and PDA or -B or in the pharynx. Other, less specific monoclonal antibodies recognize AF1, as well as other peptides to differing degrees; these antibodies are useful reagents for determination of neuronal morphology.

The postanal tail of the enteropneust Saccoglossus kowalevskii is a ciliary creeping organ without distinct similarities to the chordate tail

AbstractStach, T. and Kaul, S. 2011. The postanal tail of the enteropneust Saccoglossuskowalevskii is a ciliary creeping organ without distinct similarities to the chordatetail. — Acta Zoologica (Stockholm) 92: 150–160.The postanal tail of chordates is one of the key characters in chordate evolutionand it has been suggested to be homologous to the postanal tail of harrimaniidenteropneusts. We present electron microscopic data of the ontogeny of thepostanal tail in the enteropneust Saccoglossus kowalevskii. The postanal tail devel-ops as a ventral posterior allometric outgrowth with a ventral extension of thetelotroch. Transmission electron microscopy of serial sections reveals the epider-mal organization of the postanal tail with the exception of short, bilaterally sym-metric extensions of the paired metacoels. The epidermis cells are connected byapical junctions, rest basally on the ext racellular matrix surrounding the meso-derm, and possess a basiepidermal nerve net. The ventral cells in the postanal tailare multiciliated and used for creeping. Dorsal cells are monociliated withnumerous microvilli. Two types of glandular cells are present among the epider-mis cells. The mesoderm cells contain myofilaments. We were unable to detectanatomical structures similar to the ones present in the postanal locomotory tailof chordates, such as notochord, neural tube, or endodermal strand. Thus,results of our anatomical study do not support homology of the postanal chor-date tail and the postanal tail of harrimaniid enteropneusts.Thomas Stach, Department of Zoology, Systematics and EvolutionaryResearch, Faculty of Biology, Chemistry, and Pharmacy, Freie Universita¨tBerlin, Ko¨nigin-Luise-Strasse 1-3, 14195 Berlin, Germany. E-mail: tstach@zoosyst-berlin.deIntroductionWith a fish-like organization, chordates stand apart from therest of the millions of diverse invertebrate species. Morpholog-ical similarities among chordates abound and include a dorsalhollow nerve cord with a central canal that contains Reissner’sfiber (Olsson 1993), a pharynx perforated by gill slits sup-ported by chemically similar skeletal elements (Rychel et al.2006), a ventral glandular structure that binds iodine andsecretes a mucus net (Olsson 1963) and a postanal tail. Thispostanal tail functions in the peculiar undulatory movementcommon to all chordate animals (McHenry 2001). To servethis function, the postanal tail of chordates consists of severalsubstructures. A central skeletal element the notochord isflanked on both lateral sides by musculature (Kowalevsky1866). Dorsal to the notochord a hollow neural tube runsparallel to the notochord and on the ventral side a strand ofendodermal tissue is situated (e.g. Stach 2007). When Bur-don-Jones (1952) published the results of his detailed investi-gations into the embryology of the enteropneust Saccoglossushorsti, he described the presence of a postanal tail as a ventralextension of the posterior end of juvenile animals alreadyobserved in the juveniles of Saccoglossus kowalevskii (Bateson1885). While the homology of this postanal tail in harrimaniidenteropneusts to the stalk in pterobranchs and to the chordatelocomotory tail had been suggested (e.g. Eaton 1970), thishypothesis did not become widespread until recently. Thereason for the early reservation confronting this hypothesiswas the realization that both the postanal tail in enteropneustsand the stalk of pterobranchs are ventral extensions relative tothe position of the anus, whereas the position of the postanallocomotory tail in chordates is essentially dorsal. With the

A single immunoglobulin-domain protein required for clustering acetylcholine receptors inC. elegans

At Caenorhabditis elegans neuromuscular junctions (NMJs), synaptic clustering of the levamisole-sensitive acetylcholine receptors (L-AChRs) relies on an extracellular scaffold assembled in the synaptic cleft. It involves the secreted protein LEV-9 and the ectodomain of the transmembrane protein LEV-10, which are both expressed by muscle cells. L-AChRs, LEV-9 and LEV-10 are part of a physical complex, which localizes at NMJs, yet none of its components localizes independently at synapses. In a screen for mutants partially resistant to the cholinergic agonist levamisole, we identified oig-4, which encodes a small protein containing a single immunoglobulin domain. The OIG-4 protein is secreted by muscle cells and physically interacts with the L-AChR/LEV-9/LEV-10 complex. Removal of OIG-4 destabilizes the complex and causes a loss of L-AChR clusters at the synapse. Interestingly, OIG-4 partially localizes at NMJs independently of LEV-9 and LEV-10, thus providing a potential link between the L-AChR-associated scaffold and local synaptic cues. These results add a novel paradigm for the immunoglobulin super-family as OIG-4 is a secreted protein required for clustering ionotropic receptors independently of synapse formation.

Evolution of invertebrate nervous systems: the Chaetognatha as a case study

AbstractHarzsch, S. and Wanninger, A. 2010. Evolution of invertebrate nervous systems: the Chaetognatha as a case study. —Acta Zoologica(Stockholm)91: 35–43Although recent molecular studies indicate that Chaetognatha may be one of the earliest Bilaterian offshoots, the phylogenetic position of this taxon still is a matter of ongoing debate. In this contribution, we review recent attempts to contribute phylogenetic information on the Chaetognatha by analysing structure and development of their nervous system (neurophylogeny). Analysing this group of organisms also has a major impact on our understanding of nervous system evolution in Bilateria. We review recent evidence from this field and suggest that Urbilateria already was equipped with the genetic toolkit required to build a complex, concentrated central nervous system (CNS), although this was not expressed phenotypically so that Urbilateria was equipped with a nerve plexus and not a CNS. This implies that in the deep metazoan nodes, concentration of the ancestral plexus occurred twice independently, namely once after the protostome–deuterostome split on the branch leading to the protostomes (resulting in a ventrally positioned nerve cord) and once along the chordate line (with a dorsal nerve cord).