Ectothermic vertebrates exhibit a diverse array of adult pigment patterns. A common element of these patterns is alternating dark and light stripes each comprising different classes of neural crest-derived pigment cells. In the zebrafish, Danio rerio, alternating horizontal stripes of black melanophores and yellow xanthophores are a prominent feature of the adult pigment pattern. In fms mutant zebrafish, however, xanthophores fail to develop and melanophore stripes are severely disrupted. fmsencodes a type III receptor tyrosine kinase expressed by xanthophores and their precursors and is the closest known homologue of kit, which has long been studied for roles in pigment pattern development in amniotes. In this study we assess the cellular and temporal requirements for Fms activity in promoting adult pigment pattern development. By transplanting cells betweenfms mutants and either wild-type or nacre mutant zebrafish,we show that fms acts autonomously to the xanthophore lineage in promoting the striped arrangement of adult melanophores. To identify critical periods for fms activity, we isolated temperature sensitive alleles of fms and performed reciprocal temperature shift experiments at a range of stages from embryo to adult. These analyses demonstrate that Fms is essential for maintaining cells of the xanthophore lineage as well as maintaining the organization of melanophore stripes throughout development. Finally, we show that restoring Fms activity even at late larval stages allows essentially complete recovery of xanthophores and the development of a normal melanophore stripe pattern. Our findings suggest that fms is not required for establishing a population of precursor cells during embryogenesis but is required for recruiting pigment cell precursors to xanthophore fates,with concomitant effects on melanophore organization.
Motivated by TeV-scale neutrino mass models, we propose a systematic treatment of heavy neutrino (N ) production at hadron colliders. Our simple and efficient modeling of the vector boson fusion (VBF) W ± γ → N ℓ ± and N ℓ ± + nj signal definitions resolve collinear and soft divergences that have plagued past studies, and is applicable to other color-singlet processes, e.g., associated Higgs (W ± h), sparticle (l ±ν ℓ ), and charged Higgs (h ±± h ∓ ) production. We present, for the first time, a comparison of all leading N production modes, including both gluon fusion (GF)ν ℓ and VBF. We obtain fully differential results up to next-to-leading order (NLO) in QCD accuracy using a Monte Carlo tool chain linking FeynRules, NLOCT, and MadGraph5 aMC@NLO. Associated model files are publicly available. At the 14 TeV LHC, the leading order GF rate is small and comparable to the NLO N ℓ ± + 1j rate; at a future 100 TeV Very Large Hadron Collider, GF dominates for mN = 300 − 1500 GeV, beyond which VBF takes lead.
The genetic and developmental bases for trait expression and variation in adults are largely unknown. One system in which genes and cell behaviors underlying adult traits can be elucidated is the larval-to-adult transformation of zebrafish, Danio rerio. Metamorphosis in this and many other teleost fishes resembles amphibian metamorphosis, as a variety of larval traits (e.g., fins, skin, digestive tract, sensory systems) are remodeled in a coordinated manner to generate the adult form. Among these traits is the pigment pattern, which comprises several neural crest-derived pigment cell classes, including black melanophores, yellow xanthophores, and iridescent iridophores. D. rerio embryos and early larvae exhibit a relatively simple pattern of melanophore stripes, but this pattern is transformed during metamorphosis into the more complex pattern of the adult, consisting of alternating dark (melanophore, iridophore) and light (xanthophore, iridophore) horizontal stripes. While it is clear that some pigment cells differentiate de novo during pigment pattern metamorphosis, the extent to which larval and adult pigment patterns are developmentally independent has not been known. In this study, we show that a subset of embryonic/early larval melanophores persists into adult stages in wild-type fish; thus, larval and adult pigment patterns are not completely independent in this species. We also analyze puma mutant zebrafish, derived from a forward genetic screen to isolate mutations affecting postembryonic development. In puma mutants, a wild-type embryonic/early larval pigment pattern forms, but supernumerary early larval melanophores persist in ectopic locations through juvenile and adult stages. We then show that, although puma mutants undergo a somatic metamorphosis at the same time as wild-type fish, metamorphic melanophores that normally appear during these stages are absent. The puma mutation thus decouples metamorphosis of the pigment pattern from the metamorphosis of many other traits. Nevertheless, puma mutants ultimately recover large numbers of melanophores and exhibit extensive pattern regulation during juvenile development, when the wild-type pigment pattern already would be completed. Finally, we demonstrate that the puma mutant is both temperature-sensitive and growth-sensitive: extremely severe pigment pattern defects result at a high temperature, a high growth rate, or both; whereas a wild-type pigment pattern can be rescued at a low temperature and a low growth rate. Taken together, these results provide new insights into zebrafish pigment pattern metamorphosis and the capacity for pattern regulation when normal patterning mechanisms go awry.
We analyze Higgs condensate bubble expansion during a first-order electroweak phase transition in the early Universe. The interaction of particles with the bubble wall can be accompanied by the emission of multiple soft gauge bosons. When computed at fixed order in perturbation theory, this process exhibits large logarithmic enhancements which must be resummed to all orders when the wall velocity is large. We perform this resummation both analytically and numerically at leading logarithmic accuracy. The numerical simulation is achieved by means of a particle shower in the broken phase of the electroweak theory. The two approaches agree to the 10% level. For fast-moving walls, we find the scaling of the thermal pressure exerted against the wall to be P~γ2T4, independent of the particle masses, implying a significantly slower terminal velocity than previously suggested.
Latent precursors or stem cells of neural crest origin are present in a variety of post-embryonic tissues. Although these cells are of biomedical interest for roles in human health and disease, their potential evolutionary significance has been underappreciated. As a first step towards elucidating the contributions of such cells to the evolution of vertebrate form, we investigated the relative roles of neural crest cells and post-embryonic latent precursors during the evolutionary diversification of adult pigment patterns in Danio fishes. These pigment patterns result from the numbers and arrangements of embryonic melanophores that are derived from embryonic neural crest cells, as well as from post-embryonic metamorphic melanophores that are derived from latent precursors of presumptive neural crest origin. In the zebrafish D. rerio, a pattern of melanophore stripes arises during the larval-to-adult transformation by the recruitment of metamorphic melanophores from latent precursors. Using a comparative approach in the context of new phylogenetic data, we show that adult pigment patterns in five additional species also arise from metamorphic melanophores, identifying this as an ancestral mode of adult pigment pattern development. By contrast, superficially similar adult stripes of D. nigrofasciatus (a sister species to D. rerio) arise by the reorganization of melanophores that differentiated at embryonic stages, with a diminished contribution from metamorphic melanophores. Genetic mosaic and molecular marker analyses reveal evolutionary changes that are extrinsic to D. nigrofasciatus melanophore lineages, including a dramatic reduction of metamorphic melanophore precursors. Finally, interspecific complementation tests identify a candidate genetic pathway for contributing to the evolutionary reduction in metamorphic melanophores and the increased contribution of early larval melanophores to D. nigrofasciatus adult pigment pattern development. These results demonstrate an important role for latent precursors in the diversification of pigment patterns across danios. More generally, differences in the deployment of post-embryonic neural crest-derived stem cells or their specified progeny may contribute substantially to the evolutionary diversification of adult form in vertebrates, particularly in species that undergo a metamorphosis.
This report summarizes the present status of neutrino non-standard interactions (NSI). After a brief overview, several aspects of NSIs are discussed, including connection to neutrino mass models, model-building and phenomenology of large NSI with both light and heavy mediators, NSI phenomenology in both short- and long-baseline neutrino oscillation experiments, neutrino cross-sections, complementarity of NSI with other low- and high-energy experiments, fits with neutrino oscillation and scattering data, DUNE sensitivity to NSI, effective field theory of NSI, as well as the relevance of NSI to dark matter and cosmology. We also discuss the open questions and interesting future directions that can be pursued by the community at large. This report is based on talks and discussions during the Neutrino Theory Network NSI workshop held at Washington University in St.~Louis from May 29-31, 2019
The Deep Underground Neutrino Experiment (DUNE) will be a world-class neutrino observatory and nucleon decay detector designed to answer fundamental questions about the nature of elementary particles and their role in the universe.
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