Differentially expressed profiles in the larval testes of Wolbachia infected and uninfected Drosophila

BACKGROUND: Wolbachia are endosymbiotic bacteria that are frequently found in arthropods and nematodes. These maternally inherited bacteria manipulate host reproduction by several mechanisms including cytoplasmic incompatibility (CI). CI is the most common phenotype induced by Wolbachia and results in the developmental arrest of embryos derived from crosses between Wolbachia-infected males and uninfected females. Although the molecular mechanisms of CI are currently unknown, several studies suggest that host sperm is modified by Wolbachia during spermatogenesis. RESULTS: We compared the gene expression of Drosophila melanogaster larval testes with and without the wMel strain of Wolbachia to identify candidate genes that could be involved in the interaction between Wolbachia and the insect host. Microarray, quantitative RT-PCR and in situ hybridization analyses were carried out on D. melanogaster larval testes to determine the effect of Wolbachia infection on host gene expression. A total of 296 genes were identified by microarray analysis to have at least a 1.5 fold change [q-value < 5%] in expression. When comparing Wolbachia-infected flies to uninfected flies, 167 genes were up-regulated and 129 genes down-regulated. Differential expression of genes related to metabolism, immunity, reproduction and other functions were observed. Quantitative RT-PCR (qRT-PCR) confirmed 12 genes are differentially expressed in the testes of the 3rd instar larvae of Wolbachia-infected and uninfected flies. In situ hybridization demonstrated that Wolbachia infection changes the expression of several genes putatively associated with spermatogenesis including JH induced protein-26 and Mst84Db, or involved in immune (kenny) or metabolism (CG4988-RA). CONCLUSIONS: Wolbachia change the gene expression of 296 genes in the larval testes of D. melanogaster including genes related to metabolism, immunity and reproduction. Interestingly, most of the genes putatively involved in immunity were up-regulated in the presence of Wolbachia. In contrast, most of the genes putatively associated with reproduction (especially spermatogenesis) were down-regulated in the presence of Wolbachia. These results suggest Wolbachia may activate the immune pathway but inhibit spermatogenesis. Our data provide a significant panel of candidate genes that may be involved in the interaction between Wolbachia and their insect hosts. This forms a basis to help elucidate the underlying mechanisms of Wolbachia-induced CI in Drosophila and the influence of Wolbachia on spermatogenesis

Life and Death of an Influential Passenger: Wolbachia and the Evolution of CI-Modifiers by Their Hosts

Wolbachia are intracellular bacteria widely distributed among arthropods and nematodes. In many insect species these bacteria induce a cytoplasmic incompatibility (CI) between sperm of infected males and eggs of uninfected females. From an evolutionary point of view, CI is puzzling: In order to induce this modification-rescue system, Wolbachia affect sperm of infected males even though Wolbachia are only transmitted maternally. Phylogenetic studies of Wolbachia and hosts show that the bacteria rarely cospeciate with their hosts, indicating that infections are lost in host species. However, the mechanisms leading to Wolbachia loss are not well understood.Using a population genetic model, we investigate the spread of host mutants that enhance or repress Wolbachia action by affecting either bacterial transmission or the level of CI. We show that host mutants that decrease CI-levels in males (e.g. by reducing Wolbachia-density during spermatogenesis) spread, even at cost to mutant males. Increase of these mutants can lead to loss of Wolbachia infections, either as a direct consequence of their increase or in a step-wise manner, and we derive analytically a threshold penetrance above which a mutation's spread leads to extinction of Wolbachia. Selection on host modifiers is sexually antagonistic in that, conversely, host mutants that enhance Wolbachia in females are favoured whereas suppressors are not.Our results indicate that Wolbachia is likely to be lost from host populations on long evolutionary time scales due to reduction of CI levels in males. This can occur either by evolution of single host modifiers with large effects or through accumulation of several modifier alleles with small effects on Wolbachia action, even at cost to mutant males and even if infected hosts do not incur fecundity costs. This possibility is consistent with recent findings and may help to explain the apparent short evolutionary persistence times of Wolbachia in many host systems

Bidirectional incompatibility among divergent Wolbachia and incompatibility level differences among closely related Wolbachia in Nasonia

Author Posting. © The Author(s), 2007. This is the author's version of the work. It is posted here by permission of Nature Publishing Group for personal use, not for redistribution. The definitive version was published in Heredity 99 (2007): 278–287, doi:10.1038/sj.hdy.6800994.Most insect groups harbor obligate bacterial symbionts from the alphaproteobacterial genus Wolbachia. These bacteria alter insect reproduction in ways that enhance their cytoplasmic transmission. One of the most common alterations is cytoplasmic incompatibility (CI) - a post-fertilization modification of the paternal genome that renders embryos inviable or unable to complete diploid development in crosses between infected males and uninfected females or infected females harboring a different strain. The parasitic wasp species complex Nasonia (N. vitripennis, N. longicornis, and N. giraulti) harbor at least six different Wolbachia that cause cytoplasmic incompatibility. Each species have double infections with a representative from both the A and B Wolbachia subgroups. CI relationships of the A and B Wolbachia of N. longicornis with those of N. giraulti and N. vitripennis are investigated here. We demonstrate that all pairwise crosses between the divergent A strains are bidirectionally incompatible. We were unable to characterize incompatibility between the B Wolbachia, but we establish that the B strain of N. longicornis induces no or very weak CI in comparison to the closely related B strain in N. giraulti that expresses complete CI. Taken together with previous studies, we show that independent acquisition of divergent A Wolbachia has resulted in three mutually incompatible strains, while codivergence of B Wolbachia in N. longicornis and N. giraulti is associated with differences in CI level. Understanding the diversity and evolution of new incompatibility strains will contribute to a fuller understanding of Wolbachia invasion dynamics and Wolbachia-assisted speciation in certain groups of insects.This work was supported by grant EF-0328363 and DEB-9981634 from the National Science Foundation to J.H.W. and an Ernst Caspari Research Fellowship to S.R.B while he was at the University of Rochester. S.R.B. acknowledges support from the NASA Astrobiology Institute (NNA04CC04A)

Spin alignment measurements of the K∗0(892)K^{*0}(892) and ϕ(1020)\phi(1020) vector mesons at RHIC

We present the first spin alignment measurements for the $K^{*0}(892)$ and $\phi(1020)$ vector mesons produced at mid-rapidity with transverse momenta up to 5 GeV/c at $\sqrt{s_{NN}}$ = 200 GeV at RHIC. The diagonal spin density matrix elements with respect to the reaction plane in Au+Au collisions are $\rho_{00}$ = 0.32 $\pm$ 0.04 (stat) $\pm$ 0.09 (syst) for the $K^{*0}$ ($0.8<p_T<5.0$ GeV/c) and $\rho_{00}$ = 0.34 $\pm$ 0.02 (stat) $\pm$ 0.03 (syst) for the $\phi$ ($0.4<p_T<5.0$ GeV/c), and are constant with transverse momentum and collision centrality. The data are consistent with the unpolarized expectation of 1/3 and thus no evidence is found for the transfer of the orbital angular momentum of the colliding system to the vector meson spins. Spin alignments for $K^{*0}$ and $\phi$ in Au+Au collisions were also measured with respect to the particle's production plane. The $\phi$ result, $\rho_{00}$ = 0.41 $\pm$ 0.02 (stat) $\pm$ 0.04 (syst), is consistent with that in p+p collisions, $\rho_{00}$ = 0.39 $\pm$ 0.03 (stat) $\pm$ 0.06 (syst), also measured in this work. The measurements thus constrain the possible size of polarization phenomena in the production dynamics of vector mesons.Comment: 7 pages, 4 figures. fig.1 updated; one more reference added, one typo corrected, published in PRC.77.06190

Enhanced strange baryon production in Au+Au collisions compared to p+p at sqrts = 200 GeV

We report on the observed differences in production rates of strange and multi-strange baryons in Au+Au collisions at sqrts = 200 GeV compared to pp interactions at the same energy. The strange baryon yields in Au+Au collisions, then scaled down by the number of participating nucleons, are enhanced relative to those measured in pp reactions. The enhancement observed increases with the strangeness content of the baryon, and increases for all strange baryons with collision centrality. The enhancement is qualitatively similar to that observed at lower collision energy sqrts =17.3 GeV. The previous observations are for the bulk production, while at intermediate pT, 1 < pT< 4 GeV/c, the strange baryons even exceed binary scaling from pp yields.Comment: 7 pages, 4 figures. Printed in PR

System-Size Independence of Directed Flow Measured at the BNL Relativistic Heavy-Ion Collider

We measure directed flow (ν_1) for charged particles in Au+Au and Cu+Cu collisions at √S_(NN)=200 and 62.4 GeV, as a function of pseudorapidity (η), transverse momentum (p_t), and collision centrality, based on data from the STAR experiment. We find that the directed flow depends on the incident energy but, contrary to all available model implementations, not on the size of the colliding system at a given centrality. We extend the validity of the limiting fragmentation concept to ν_1 in different collision systems, and investigate possible explanations for the observed sign change in ν_1(p_t)

Longitudinal double-spin asymmetry for inclusive jet production in p+p collisions at sqrt(s)=200 GeV

We report a new STAR measurement of the longitudinal double-spin asymmetry A_LL for inclusive jet production at mid-rapidity in polarized p+p collisions at a center-of-mass energy of sqrt(s) = 200 GeV. The data, which cover jet transverse momenta 5 < p_T < 30 GeV/c, are substantially more precise than previous measurements. They provide significant new constraints on the gluon spin contribution to the nucleon spin through the comparison to predictions derived from one global fit of polarized deep-inelastic scattering measurements.Comment: 7 pages, 4 figures + 1 tabl

Forward Neutral Pion Transverse Single Spin Asymmetries in p+p Collisions at \sqrt{s}=200 GeV

We report precision measurements of the Feynman-x dependence, and first measurements of the transverse momentum dependence, of transverse single spin asymmetries for the production of \pi^0 mesons from polarized proton collisions at \sqrt{s}=200 GeV. The x_F dependence of the results is in fair agreement with perturbative QCD model calculations that identify orbital motion of quarks and gluons within the proton as the origin of the spin effects. Results for the p_T dependence at fixed x_F are not consistent with pQCD-based calculations.Comment: 6 pages, 4 figure

Measurements of ϕ\phi meson production in relativistic heavy-ion collisions at RHIC

We present results for the measurement of $\phi$ meson production via its charged kaon decay channel $\phi \to K^+K^-$ in Au+Au collisions at $\sqrt{s_{_{NN}}}=62.4$, 130, and 200 GeV, and in $p+p$ and $d$+Au collisions at $\sqrt{s_{_{NN}}}=200$ GeV from the STAR experiment at the BNL Relativistic Heavy Ion Collider (RHIC). The midrapidity ($|y|<0.5$) $\phi$ meson transverse momentum ($p_{T}$) spectra in central Au+Au collisions are found to be well described by a single exponential distribution. On the other hand, the $p_{T}$ spectra from $p+p$, $d$+Au and peripheral Au+Au collisions show power-law tails at intermediate and high $p_{T}$ and are described better by Levy distributions. The constant $\phi/K^-$ yield ratio vs beam species, collision centrality and colliding energy is in contradiction with expectations from models having kaon coalescence as the dominant mechanism for $\phi$ production at RHIC. The $\Omega/\phi$ yield ratio as a function of $p_{T}$ is consistent with a model based on the recombination of thermal $s$ quarks up to $p_{T}\sim 4$ GeV/$c$, but disagrees at higher transverse momenta. The measured nuclear modification factor, $R_{dAu}$, for the $\phi$ meson increases above unity at intermediate $p_{T}$, similar to that for pions and protons, while $R_{AA}$ is suppressed due to the energy loss effect in central Au+Au collisions. Number of constituent quark scaling of both $R_{cp}$ and $v_{2}$ for the $\phi$ meson with respect to other hadrons in Au+Au collisions at $\sqrt{s_{_{NN}}}$=200 GeV at intermediate $p_{T}$ is observed. These observations support quark coalescence as being the dominant mechanism of hadronization in the intermediate $p_{T}$ region at RHIC.Comment: 22 pages, 21 figures, 4 table