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Null mutations in the human IQCB1/NPHP5 (nephrocystin-5) gene that encodes NPHP5 are the most frequent cause of Senior-LOken syndrome, a ciliopathy that is characterized by Leber congenital amaurosis and nephronophthisis. We generated germline Nphp5-knockout mice by placing a -Geo gene trap in intron 4, thereby truncating NPHP5 at Leu87 and removing all known functional domains. At eye opening, Nphp5(-/-) mice exhibited absence of scotopic and photopic electroretinogram responses, a phenotype that resembles Leber congenital amaurosis. Outer segment transmembrane protein accumulation in Nphp5(-/-) endoplasmic reticulum was evident as early as postnatal day (P)6. EGFP-CETN2, a centrosome and transition zone marker, identified basal bodies in Nphp5(-/-) photoreceptors, but without fully developed transition zones. Ultrastructure of P6 and 10 Nphp5(-/-) photoreceptors revealed aberrant transition zones of reduced diameter. Nphp5(-/-) photoreceptor degeneration was complete at 1 mo of age but was delayed significantly in Nphp5(-/-);Nrl(-/-) (cone only) retina. Nphp5(-/-) mouse embryonic fibroblast developed normal cilia, and Nphp5(-/-) kidney histology at 1 yr of age showed no significant pathology. Results establish that nephrocystin-5 is essential for photoreceptor outer segment formation but is dispensable for kidney and mouse embryonic fibroblast ciliary formation.
Paleomagnetic dating of the India-Asia collision hinges on determining the Paleogene latitude of the Lhasa terrane (southern Tibet). Reported latitudes range from 5 degrees N to 30 degrees N, however, leading to contrasting paleogeographic interpretations. Here we report new data from the Eocene Linzizong volcanic rocks in the Nanmulin Basin, which previously yielded data suggesting a low paleolatitude (similar to 10 degrees N). New zircon U-Pb dates indicate an age of similar to 52Ma. Negative fold tests, however, demonstrate that the isolated characteristic remanent magnetizations, with notably varying inclinations, are not primary. Rock magnetic analyses, end-member modeling of isothermal remanent magnetization acquisition curves, and petrographic observations are consistent with variable degrees of posttilting remagnetization due to low-temperature alteration of primary magmatic titanomagnetite and the formation of secondary pigmentary hematite that unblock simultaneously. Previously reported paleomagnetic data from the Nanmulin Basin implying low paleolatitude should thus not be used to estimate the time and latitude of the India-Asia collision. We show that the paleomagnetic inclinations vary linearly with the contribution of secondary hematite to saturation isothermal remanent magnetization. We tentatively propose a new method to recover a primary remanence with inclination of 38.1 degrees (35.7 degrees, 40.5 degrees) (95% significance) and a secondary remanence with inclination of 42.9 degrees (41.5 degrees,44.4 degrees) (95% significance). The paleolatitude defined by the modeled primary remanence21 degrees N (19.8 degrees N, 23.1 degrees N)is consistent with the regional compilation of published results from pristine volcanic rocks and sedimentary rocks of the upper Linzizong Group corrected for inclination shallowing. The start of the Tibetan Himalaya-Asia collision was situated at similar to 20 degrees N and took place by similar to 50Ma.
The Paleogene latitude of the Lhasa terrane (southern Tibet) can constrain the age of the onset of the India-Asia collision. Estimates for this latitude, however, vary from 5 degrees N to 30 degrees N, and thus, here, we reassess the geochronology and paleomagnetism of Paleogene volcanic rocks from the Linzizong Group in the Linzhou basin. The lower and upper parts of the section previously yielded particularly conflicting ages and paleolatitudes. We report consistent Ar-40/Ar-39 and U-Pb zircon dates of similar to 52Ma for the upper Linzizong, and Ar-40/Ar-39 dates (similar to 51Ma) from the lower Linzizong are significantly younger than U-Pb zircon dates (64-63Ma), suggesting that the lower Linzizong was thermally and/or chemically reset. Paleomagnetic results from 24 sites in lower Linzizong confirm a low apparent paleolatitude of similar to 5 degrees N, compared to the upper part (similar to 20 degrees N) and to underlying Cretaceous strata (similar to 20 degrees N). Detailed rock magnetic analyses, end-member modeling of magnetic components, and petrography from the lower and upper Linzizong indicate widespread secondary hematite in the lower Linzizong, whereas hematite is rare in upper Linzizong. Volcanic rocks of the lower Linzizong have been hydrothermally chemically remagnetized, whereas the upper Linzizong retains a primary remanence. We suggest that remagnetization was induced by acquisition of chemical and thermoviscous remanent magnetizations such that the shallow inclinations are an artifact of a tilt correction applied to a secondary remanence in lower Linzizong. We estimate that the Paleogene latitude of Lhasa terrane was 204 degrees N, consistent with previous results suggesting that India-Asia collision likely took place by similar to 52Ma at similar to 20 degrees N.
The Tibetan Himalaya represents the northernmost continental unit of the Indian plate that collided with Asia in the Cenozoic. Paleomagnetic studies on the Tibetan Himalaya can help constrain the dimension and paleogeography of "Greater India,' the Indian plate lithosphere that subducted and underthrusted below Asia after initial collision. Here we present a paleomagnetic investigation of a Jurassic (limestones) and Lower Cretaceous (volcaniclastic sandstones) section of the Tibetan Himalaya. The limestones yielded positive fold test, showing a prefolding origin of the isolated remanent magnetizations. Detailed paleomagnetic analyses, rock magnetic tests, end-member modeling of acquisition curves of isothermal remanent magnetization, and petrographic investigation reveal that the magnetic carrier of the Jurassic limestones is authigenic magnetite, whereas the dominant magnetic carrier of the Lower Cretaceous volcaniclastic sandstones is detrital magnetite. Our observations lead us to conclude that the Jurassic limestones record a prefolding remagnetization, whereas the Lower Cretaceous volcaniclastic sandstones retain a primary remanence. The volcaniclastic sandstones yield an Early Cretaceous paleolatitude of 55.5 degrees S [52.5 degrees S, 58.6 degrees S] for the Tibetan Himalaya, suggesting it was part of the Indian continent at that time. The size of "Greater India' during Jurassic time cannot be estimated from these limestones. Instead, a paleolatitude of the Tibetan Himalaya of 23.8 degrees S [21.8 degrees S, 26.1 degrees S] during the remagnetization process is suggested. It is likely that the remagnetization, caused by the oxidation of early diagenetic pyrite to magnetite, was induced during 103-83 or 77-67 Ma. The inferred paleolatitudes at these two time intervals imply very different tectonic consequences for the Tibetan Himalaya.
The formation of the Pamir is a key component of the India-Asia collision with major implications for lithospheric processes, plateau formation, land-sea configurations and associated climate changes. Although the formation of the Pamir is traditionally linked to Cenozoic processes associated with the India-Asia collision, the contribution of the Mesozoic tectonic evolution remains poorly understood. The Pamir was formed by the suturing of Gondwanan terranes to the south margin of Eurasia, however, the timing and tectonic mechanisms associated with this Mesozoic accretion remain poorly constrained. These processes are recorded by several igneous belts within these terranes, which are not well studied. Within the Southern Pamir, the Albian-Turonian volcanic rocks and comagmatic plutons of the Kyzylrabat Igneous Complex (KIC) provide an important and still unconstrained record of the Pamir evolution. Here we provide the age, origin and the geodynamic setting of the KIC volcanics by studying their petrology, zircon U-Pb geochronology, geochemistry and isotope composition.17 samples from the KIC volcanics yield U-Pb ages spanning from 92 to 110 Ma. The volcanics are intermediate to acidic in composition (SiO2 = 56-69 wt%) and exhibit high-K calc-alkaline and shoshonitic affinity (K2O/Na2O = 12.2 wt%). They show enrichment in LILE and LREE and depletion in HFSE and HREE with negative Ta, Ti and Nb anomalies, suggesting an arc-related tectonic setting for their formation. Low sNd(t) values (from 9.1 to 4.7), relatively high Sr-87/Sr-86(i) ratios (0.7069-0.7096) and broad range of zircon stif values (from 22.6 to 1.5) suggest a mixture of different magma sources. These features suggest that volcanics were derived by crustal under- or intraplating of an enriched subduction-related mantle shoshonitic magmas, by heating and partial melting of the lower crust, and by mixing of both magma components. Our results further imply that the KIC volcanics represent a shoshonitic suite typical of an evolution from active continental arc to post-collisional setting with a steepening of the Benioff zone and thickening of the crust toward the back-arc. This setting is best explained by the subduction- collision transition along the Shyok suture due to accretion of the Kohistan island arc to the Karakoram. This suggests that a significant part of the crustal shortening and thickening accommodated in the Pamir occurred in the Mesozoic before the India-Asia collision with implications for regional tectonic models. This further suggests the Pamir was already a major topographic feature with potentially important paleoclimate forcing such as the monsoonal circulation. (C) 2017 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.
Building and changing a microbiome at will and maintaining it over hundreds of generations has so far proven challenging. Despite best efforts, complex microbiomes appear to be susceptible to large stochastic fluctuations. Current capabilities to assemble and control stable complex microbiomes are limited. Here, we propose a looped mass transfer design that stabilizes microbiomes over long periods of time. Five local microbiomes were continuously grown in parallel for over 114 generations and connected by a loop to a regional pool. Mass transfer rates were altered and microbiome dynamics were monitored using quantitative high-throughput flow cytometry and taxonomic sequencing of whole communities and sorted subcommunities. Increased mass transfer rates reduced local and temporal variation in microbiome assembly, did not affect functions, and overcame stochasticity, with all microbiomes exhibiting high constancy and increasing resistance. Mass transfer synchronized the structures of the five local microbiomes and nestedness of certain cell types was eminent. Mass transfer increased cell number and thus decreased net growth rates mu'. Subsets of cells that did not show net growth mu'SCx were rescued by the regional pool R and thus remained part of the microbiome. The loop in mass transfer ensured the survival of cells that would otherwise go extinct, even if they did not grow in all local microbiomes or grew more slowly than the actual dilution rate D would allow. The rescue effect, known from metacommunity theory, was the main stabilizing mechanism leading to synchrony and survival of subcommunities, despite differences in cell physiological properties, including growth rates.
The largest uncertainty in projections of future sea-level change results from the potentially changing dynamical ice discharge from Antarctica. Basal ice-shelf melting induced by a warming ocean has been identified as a major cause for additional ice flow across the grounding line. Here we attempt to estimate the uncertainty range of future ice discharge from Antarctica by combining uncertainty in the climatic forcing, the oceanic response and the ice-sheet model response. The uncertainty in the global mean temperature increase is obtained from historically constrained emulations with the MAGICC-6.0 (Model for the Assessment of Greenhouse gas Induced Climate Change) model. The oceanic forcing is derived from scaling of the subsurface with the atmospheric warming from 19 comprehensive climate models of the Coupled Model Intercomparison Project (CMIP-5) and two ocean models from the EU-project Ice2Sea. The dynamic ice-sheet response is derived from linear response functions for basal ice-shelf melting for four different Antarctic drainage regions using experiments from the Sea-level Response to Ice Sheet Evolution (SeaRISE) intercomparison project with five different Antarctic ice-sheet models. The resulting uncertainty range for the historic Antarctic contribution to global sea-level rise from 1992 to 2011 agrees with the observed contribution for this period if we use the three ice-sheet models with an explicit representation of ice-shelf dynamics and account for the time-delayed warming of the oceanic subsurface compared to the surface air temperature. The median of the additional ice loss for the 21st century is computed to 0.07 m (66% range: 0.02-0.14 m; 90% range: 0.0-0.23 m) of global sea-level equivalent for the low-emission RCP-2.6 (Representative Concentration Pathway) scenario and 0.09 m (66% range: 0.04-0.21 m; 90% range: 0.01-0.37 m) for the strongest RCP-8.5. Assuming no time delay between the atmospheric warming and the oceanic subsurface, these values increase to 0.09 m (66% range: 0.04-0.17 m; 90% range: 0.02-0.25 m) for RCP-2.6 and 0.15 m (66% range: 0.07-0.28 m; 90% range: 0.04-0.43 m) for RCP-8.5. All probability distributions are highly skewed towards high values. The applied ice-sheet models are coarse resolution with limitations in the representation of grounding-line motion. Within the constraints of the applied methods, the uncertainty induced from different ice-sheet models is smaller than that induced by the external forcing to the ice sheets.
We report on both high-precision photometry from the Microvariability and Oscillations of Stars (MOST) space telescope and ground-based spectroscopy of the triple system delta Ori A, consisting of a binary O9.5II+early-B (Aa1 and Aa2) with P = 5.7 days, and a more distant tertiary (O9 IV P > 400 years). This data was collected in concert with X-ray spectroscopy from the Chandra X-ray Observatory. Thanks to continuous coverage for three weeks, the MOST light curve reveals clear eclipses between Aa1 and Aa2 for the first time in non-phased data. From the spectroscopy, we have a well-constrained radial velocity (RV) curve of Aa1. While we are unable to recover RV variations of the secondary star, we are able to constrain several fundamental parameters of this system and determine an approximate mass of the primary using apsidal motion. We also detected second order modulations at 12 separate frequencies with spacings indicative of tidally influenced oscillations. These spacings have never been seen in a massive binary, making this system one of only a handful of such binaries that show evidence for tidally induced pulsations.
In this study, rapid loss of relativistic radiation belt electrons at low L* values (2.4-3.2) during a strong geomagnetic storm on 22 June 2015 is investigated along with five possible loss mechanisms. Both the particle and wave data are obtained from the Van Allen Probes. Duskside H+ band electromagnetic ion cyclotron (EMIC) waves were observed during a rapid decrease of relativistic electrons with energy above 5.2 MeV occurring outside the plasma sphere during extreme magnetopause compression. Lower He+ composition and enriched O+ composition are found compared to typical values assumed in other studies of cyclotron resonant scattering of relativistic electrons by EMIC waves. Quantitative analysis demonstrates that even with the existence of He+ band EMIC waves, it is the H+ band EMIC waves that are likely to cause the depletion at small pitch angles and strong gradients in pitch angle distributions of relativistic electrons with energy above 5.2 MeV at low L values for this event. Very low frequency wave activity at other magnetic local time can be favorable for the loss of relativistic electrons at higher pitch angles. An illustrative calculation that combines the nominal pitch angle scattering rate due to whistler mode chorus at high pitch angles with the H+ band EMIC wave loss rate at low pitch angles produces loss on time scale observed at L = 2.4-3.2. At high L values and lower energies, radial loss to the magnetopause is a viable explanation.