A mosaic of conserved and novel modes of gene expression and morphogenesis in mesoderm and muscle formation of a larval bivalve

The mesoderm gives rise to several key morphological features of bilaterian animals including endoskeletal elements and the musculature. A number of regulatory genes involved in mesoderm and/or muscle formation (e.g., Brachyury (Bra), even-skipped (eve), Mox, myosin II heavy chain (mhc)) have been identified chiefly from chordates and the ecdysozoans Drosophila and Caenorhabditis elegans, but data for non-model protostomes, especially those belonging to the ecdysozoan sister clade, Lophotrochozoa (e.g., flatworms, annelids, mollusks), are only beginning to emerge. Within the lophotrochozoans, Mollusca constitutes the most speciose and diverse phylum. Interestingly, however, information on the morphological and molecular underpinnings of key ontogenetic processes such as mesoderm formation and myogenesis remains scarce even for prominent molluscan sublineages such as the bivalves. Here, we investigated myogenesis and developmental expression of Bra, eve, Mox, and mhc in the quagga mussel Dreissena rostriformis, an invasive freshwater bivalve and an emerging model in invertebrate evodevo. We found that all four genes are expressed during mesoderm formation, but some show additional, individual sites of expression during ontogeny. While Mox and mhc are involved in early myogenesis, eve is also expressed in the embryonic shell field and Bra is additionally present in the foregut. Comparative analysis suggests that Mox has an ancestral role in mesoderm and possibly muscle formation in bilaterians, while Bra and eve are conserved regulators of mesoderm development of nephrozoans (protostomes and deuterostomes). The fully developed Dreissena veliger larva shows a highly complex muscular architecture, supporting a muscular ground pattern of autobranch bivalve larvae that includes at least a velum muscle ring, three or four pairs of velum retractors, one or two pairs of larval retractors, two pairs of foot retractors, a pedal plexus, possibly two pairs of mantle retractors, and the muscles of the pallial line, as well as an anterior and a posterior adductor. As is typical for their molluscan kin, remodelling and loss of prominent larval features such as the velum musculature and various retractor systems appear to be also common in bivalves

Non-collinear Hox gene expression in bivalves and the evolution of morphological novelties in mollusks

Hox genes are key developmental regulators that are involved in establishing morphological features during animal ontogeny. They are commonly expressed along the anterior--posterior axis in a staggered, or collinear, fashion. In mollusks, the repertoire of body plans is widely diverse and current data suggest their involvement during development of landmark morphological traits in Conchifera, one of the two major lineages that comprises those taxa that originated from a uni-shelled ancestor (Monoplacophora, Gastropoda, Cephalopoda, Scaphopoda, Bivalvia). For most clades, and bivalves in particular, data on Hox gene expression throughout ontogeny are scarce. We thus investigated Hox expression during development of the quagga mussel, Dreissena rostriformis, to elucidate to which degree they might contribute to specific phenotypic traits as in other conchiferans. The Hox/ParaHox complement of Mollusca typically comprises 14 genes, 13 of which are present in bivalve genomes including Dreissena. We describe here expression of 9 Hox genes and the ParaHox gene Xlox during Dreissena development. Hox expression in Dreissena is first detected in the gastrula stage with widely overlapping expression domains of most genes. In the trochophore stage, Hox gene expression shifts towards more compact, largely mesodermal domains. Only few of these domains can be assigned to specific developing morphological structures such as Hox1 in the shell field and Xlox in the hindgut. We did not find traces of spatial or temporal staggered expression of Hox genes in Dreissena. Our data support the notion that Hox gene expression has been coopted independently, and to varying degrees, into lineage-specific structures in the respective conchiferan clades. The non-collinear mode of Hox expression in Dreissena might be a result of the low degree of body plan regionalization along the bivalve anterior--posterior axis as exemplified by the lack of key morphological traits such as a distinct head, cephalic tentacles, radula apparatus, and a simplified central nervous system

A model independent and rephase invariant parametrization of CP violation

The phenomenological description of the neutral B meson system is proposed in terms of the fundamental CP-violating observables and within a rephasing invariant formalism. This generic formalism can select the time-dependent and time-integrated asymmetries which provide the basic tools to discriminate the different kinds of possible CP-violating effects in dedicated experimental B-meson facilities.Comment: 19 pages, Plain Te

Comparative transcriptomics enlarges the toolkit of known developmental genes in mollusks

Data used for the phylogenetic analysis of Hox and ParaHox genes, including the respective GenBank accession numbers. (DOC 31 kb

Inelastic Final-State Interactions and Two-body Hadronic B decays into Single-Isospin channels

The role of inelastic final-state interactions in CP asymmetries and branching ratios is investigated in certain chosen single isospin two-body hadronic B decays. Treating final-state interactions through Pomeron and Regge exchanges, we demonstrate that inelastic final state interactions could lead to sizeable effects on the CP asymmetry.Comment: 23 pages, Latex, 1 eps-figur

Optimization of the design of OMNIS, the observatory of multiflavor neutrinos from supernovae

A Monte Carlo code has been developed to simulate the operation of the planned detectors in OMNIS, a supernova neutrino observatory. OMNIS will detect neutrinos originating from a core collapse supernova by the detection of spalled neutrons from Pb- or Fe-nuclei. This might be accomplished using Gd-loaded liquid scintillator. Results for the optimum configuration for such modules with respect to both neutron detection efficiency and cost efficiency are presented. Careful consideration has been given to the expected levels of radioactive backgrounds and their effects. The results show that the amount of data to be processed by a software trigger can be reduced to the <10kHz region and a neutron, once produced in the detector, can be detected and identified with an efficiency of >30%.Comment: Elsevier preprint; 29 pages, 23 figure

Embryonic chirality and the evolution of spiralian left-right asymmetries

This work has been funded by the Sars core budget to A.H