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Rhodopsin-based voltage imaging tools for use in muscles and neurons of Caenorhabditis elegans
(2019)
Genetically encoded voltage indicators (GEVIs) based on microbial rhodopsins utilize the voltage-sensitive fluorescence of all-trans retinal (ATR), while in electrochromic FRET (eFRET) sensors, donor fluorescence drops when the rhodopsin acts as depolarization-sensitive acceptor. In recent years, such tools have become widely used in mammalian cells but are less commonly used in invertebrate systems, mostly due to low fluorescence yields. We systematically assessed Arch(D95N), Archon, QuasAr, and the eFRET sensors MacQ-mCitrine and QuasAr-mOrange, in the nematode Caenorhabditis elegans ATR-bearing rhodopsins reported on voltage changes in body wall muscles (BWMs), in the pharynx, the feeding organ [where Arch(D95N) showed approximately 128% ΔF/F increase per 100 mV], and in neurons, integrating circuit activity. ATR fluorescence is very dim, yet, using the retinal analog dimethylaminoretinal, it was boosted 250-fold. eFRET sensors provided sensitivities of 45 to 78% ΔF/F per 100 mV, induced by BWM action potentials, and in pharyngeal muscle, measured in simultaneous optical and sharp electrode recordings, MacQ-mCitrine showed approximately 20% ΔF/F per 100 mV. All sensors reported differences in muscle depolarization induced by a voltage-gated Ca2+-channel mutant. Optogenetically evoked de- or hyperpolarization of motor neurons increased or eliminated action potential activity and caused a rise or drop in BWM sensor fluorescence. Finally, we analyzed voltage dynamics across the entire pharynx, showing uniform depolarization but compartmentalized repolarization of anterior and posterior parts. Our work establishes all-optical, noninvasive electrophysiology in live, intact C. elegans.
Results on the production of 4He and Image 1 nuclei in Pb–Pb collisions at √sNN=2.76TeV in the rapidity range |y|<1, using the ALICE detector, are presented in this paper. The rapidity densities corresponding to 0–10% central events are found to be dN/dyHe4=(0.8±0.4(stat)±0.3(syst))×10−6 and Image 2, respectively. This is in agreement with the statistical thermal model expectation assuming the same chemical freeze-out temperature (Tchem=156MeV) as for light hadrons. The measured ratio of Image 3 is 1.4±0.8(stat)±0.5(syst).
The elliptic flow (v2) of (anti-)3He is measured in Pb–Pb collisions at √sNN=5.02TeV in the transverse-momentum (pT) range of 2–6 GeV/c for the centrality classes 0–20%, 20–40%, and 40–60% using the event-plane method. This measurement is compared to that of pions, kaons, and protons at the same center-of-mass energy. A clear mass ordering is observed at low pT, as expected from relativistic hydrodynamics. The violation of the scaling of v2 with the number of constituent quarks at low pT, already observed for identified hadrons and deuterons at LHC energies, is confirmed also for (anti-)3He. The elliptic flow of (anti-)3He is underestimated by the Blast-Wave model and overestimated by a simple coalescence approach based on nucleon scaling. The elliptic flow of (anti-)3He measured in the centrality classes 0–20% and 20–40% is well described by a more sophisticated coalescence model where the phase-space distributions of protons and neutrons are generated using the iEBE-VISHNU hybrid model with AMPT initial conditions.
The production of prompt Λc+ baryons at midrapidity (|y|<0.5) was measured in central (0–10%) and mid-central (30–50%) Pb–Pb collisions at the center-of-mass energy per nucleon–nucleon pair √sNN=5.02 TeV with the ALICE detector. The results are more precise, more differential in centrality, and reach much lower transverse momentum (pT=1 GeV/c) with respect to previous measurements performed by the ALICE, STAR, and CMS Collaborations in nucleus–nucleus collisions, allowing for an extrapolation down to pT=0. The pT-differential Λc+/D0 ratio is enhanced with respect to the pp measurement for 4<pT<8 GeV/c by 3.7 standard deviations (σ), while the pT-integrated ratios are compatible within 1σ. The observed trend is similar to that observed in the strange sector for the Λ/KS0 ratio. Model calculations including coalescence or statistical hadronization for charm-hadron formation are compared with the data.
In quantum scattering processes between two particles, aspects characterizing the strong and Coulomb forces can be observed in kinematic distributions of the particle pairs. The sensitivity to the interaction potential reaches a maximum at low relative momentum and vanishing distance between the two particles. Ultrarelativistic heavy-ion collisions at the LHC provide an abundant source of many hadron species and can be employed as a measurement method of scattering parameters that is complementary to scattering experiments. This study confirms that momentum correlations of particles produced in Pb–Pb collisions at the LHC provide an accurate measurement of kaon–proton scattering parameters at low relative momentum, allowing precise access to the K−p→K−p process. This work also validates the femtoscopic measurement in ultrarelativistic heavy-ion collisions as an alternative to scattering experiments and a complementary tool to the study of exotic atoms with comparable precision. In this work, the first femtoscopic measurement of momentum correlations of K−p(K+p‾) and K+p(K−p‾) pairs in Pb–Pb collisions at centre-of-mass energy per nucleon pair of sNN=5.02 TeV registered by the ALICE experiment is reported. The components of the K−p complex scattering length are extracted and found to be ℜf0=−0.91±0.03(stat)−0.03+0.17(syst) and ℑf0=0.92±0.05(stat)−0.33+0.12(syst). The results are compared with chiral effective field theory predictions as well as with existing data from dedicated scattering and exotic kaonic atom experiments.
The measurement of the production of f0(980) in inelastic pp collisions at √s=5.02 TeV is presented. This is the first reported measurement of inclusive f0(980) yield at LHC energies. The production is measured at midrapidity, |y|<0.5, in a wide transverse momentum range, 0<pT<16 GeV/c, by reconstructing the resonance in the f0(980) →π+π− hadronic decay channel using the ALICE detector. The pT-differential yields are compared to those of pions, protons and ϕ mesons as well as to predictions from the HERWIG 7.2 QCD-inspired Monte Carlo event generator and calculations from a coalescence model that uses the AMPT model as an input. The ratio of the pT-integrated yield of f0(980) relative to pions is compared to measurements in e+e− and pp collisions at lower energies and predictions from statistical hadronisation models and HERWIG 7.2. A mild collision energy dependence of the f0(980) to pion production is observed in pp collisions from SPS to LHC energies. All considered models underpredict the pT-integrated 2f0(980)/(π+ +π−) ratio. The prediction from the canonical statistical hadronisation model assuming a zero total strangeness content of f0(980) is consistent with the data within 1.9σ and is the closest to the data. The results provide an essential reference for future measurements of the particle yield and nuclear modification in p–Pb and Pb–Pb collisions, which have been proposed to be instrumental to probe the elusive nature and quark composition of the f0(980) scalar meson.
In ultrarelativistic heavy-ion collisions, the event-by-event variation of the elliptic flow v2 reflects fluctuations in the shape of the initial state of the system. This allows to select events with the same centrality but different initial geometry. This selection technique, Event Shape Engineering, has been used in the analysis of charge-dependent two- and three-particle correlations in Pb–Pb collisions at √sNN=2.76 TeV. The two-particle correlator 〈cos(φα−φβ)〉, calculated for different combinations of charges α and β, is almost independent of v2 (for a given centrality), while the three-particle correlator 〈cos(φα+φβ−2Ψ2)〉 scales almost linearly both with the event v2 and charged-particle pseudorapidity density. The charge dependence of the three-particle correlator is often interpreted as evidence for the Chiral Magnetic Effect (CME), a parity violating effect of the strong interaction. However, its measured dependence on v2 points to a large non-CME contribution to the correlator. Comparing the results with Monte Carlo calculations including a magnetic field due to the spectators, the upper limit of the CME signal contribution to the three-particle correlator in the 10–50% centrality interval is found to be 26–33% at 95% confidence level.
The ALICE Collaboration reports the measurement of semi-inclusive distributions of charged-particle jets recoiling from a high-transverse momentum trigger hadron in p–Pb collisions at √sNN=5.02 TeV. Jets are reconstructed from charged-particle tracks using the anti-kT algorithm with resolution parameter R=0.2 and 0.4. A data-driven statistical approach is used to correct the uncorrelated background jet yield. Recoil jet distributions are reported for jet transverse momentum 15<pT,jetch<50GeV/c and are compared in various intervals of p–Pb event activity, based on charged-particle multiplicity and zero-degree neutral energy in the forward (Pb-going) direction. The semi-inclusive observable is self-normalized and such comparisons do not require the interpretation of p–Pb event activity in terms of collision geometry, in contrast to inclusive jet observables. These measurements provide new constraints on the magnitude of jet quenching in small systems at the LHC. In p–Pb collisions with high event activity, the average medium-induced out-of-cone energy transport for jets with R=0.4 and 15<pT,jetch<50GeV/c is measured to be less than 0.4 GeV/c at 90% confidence, which is over an order of magnitude smaller than a similar measurement for central Pb–Pb collisions at √sNN=2.76TeV. Comparison is made to theoretical calculations of jet quenching in small systems, and to inclusive jet measurements in p–Pb collisions selected by event activity at the LHC and in d–Au collisions at RHIC.
In ultraperipheral collisions (UPCs) of relativistic nuclei without overlap of nuclear densities, the two nuclei are excited by the Lorentz-contracted Coulomb fields of their collision partners. In these UPCs, the typical nuclear excitation energy is below a few tens of MeV, and a small number of nucleons are emitted in electromagnetic dissociation (EMD) of primary nuclei, in contrast to complete nuclear fragmentation in hadronic interactions. The cross sections of emission of given numbers of neutrons in UPCs of 208Pb nuclei at sNN−−−√=5.02 TeV were measured with the neutron zero degree calorimeters (ZDCs) of the ALICE detector at the LHC, exploiting a similar technique to that used in previous studies performed at sNN−−−√=2.76 TeV. In addition, the cross sections for the exclusive emission of one, two, three, four, and five forward neutrons in the EMD, not accompanied by the emission of forward protons, and thus mostly corresponding to the production of 207,206,205,204,203Pb, respectively, were measured for the first time. The predictions from the available models describe the measured cross sections well. These cross sections can be used for evaluating the impact of secondary nuclei on the LHC components, in particular, on superconducting magnets, and also provide useful input for the design of the Future Circular Collider (FCC-hh).
The production of prompt D0, Ds+, and Λc+ hadrons, and their ratios, Ds+/D0 and Λc+/D0, are measured in proton–proton collisions at √s=13 TeV at midrapidity (|y|<0.5) with the ALICE detector at the LHC. The measurements are performed as a function of the charm-hadron transverse momentum (pT) in intervals of charged-particle multiplicity, measured with two multiplicity estimators covering different pseudorapidity regions. While the strange to non-strange Ds+/D0 ratio indicates no significant multiplicity dependence, the baryon-to-meson pT-differential Λc+/D0 ratio shows a multiplicity-dependent enhancement, with a significance of 5.3σ for 1<pT<12 GeV/c, comparing the highest multiplicity interval with respect to the lowest one. The measurements are compared with a theoretical model that explains the multiplicity dependence by a canonical treatment of quantum charges in the statistical hadronisation approach, and with predictions from event generators that implement colour reconnection mechanisms beyond the leading colour approximation to model the hadronisation process. The Λc+/D0 ratios as a function of pT present a similar shape and magnitude as the Λ/KS0 ratios in comparable multiplicity intervals, suggesting a potential common mechanism for light- and charm-hadron formation, with analogous multiplicity dependence. The pT-integrated ratios, extrapolated down to pT=0, do not show a significant dependence on multiplicity within the uncertainties.