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Titan's middle atmosphere is currently experiencing a rapid change of season after northern spring arrived in 2009 (refs 1, 2). A large cloud was observed for the first time above Titan's southern pole in May 2012, at an altitude of 300 kilometres. A temperature maximum was previously observed there, and condensation was not expected for any of Titan's atmospheric gases. Here we report that this cloud is composed of micrometre-sized particles of frozen hydrogen cyanide (HCN ice). The presence of HCN particles at this altitude, together with temperature determinations from mid-infrared observations, indicate a dramatic cooling of Titan's atmosphere inside the winter polar vortex in early 2012. Such cooling is in contrast to previously measured high-altitude warming in the polar vortex, and temperatures are a hundred degrees colder than predicted by circulation models. These results show that post-equinox cooling at the winter pole of Titan is much more efficient than previously thought.
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Ecological and laboratory studies have demonstrated that temperature modulates West Nile virus (WNV) transmission dynamics and spillover infection to humans. Here we explore whether inclusion of temperature forcing in a model depicting WNV transmission improves WNV forecast accuracy relative to a baseline model depicting WNV transmission without temperature forcing. Both models are optimized using a data assimilation method and two observed data streams: mosquito infection rates and reported human WNV cases. Each coupled model-inference framework is then used to generate retrospective ensemble forecasts of WNV for 110 outbreak years from among 12 geographically diverse United States counties. The temperature-forced model improves forecast accuracy for much of the outbreak season. From the end of July until the beginning of October, a timespan during which 70% of human cases are reported, the temperature-forced model generated forecasts of the total number of human cases over the next 3 weeks, total number of human cases over the season, the week with the highest percentage of infectious mosquitoes, and the peak percentage of infectious mosquitoes that on average increased absolute forecast accuracy 5%, 10%, 12%, and 6%, respectively, over the non-temperature forced baseline model. These results indicate that use of temperature forcing improves WNV forecast accuracy and provide further evidence that temperature influences rates of WNV transmission. The findings provide a foundation for implementation of a statistically rigorous system for real-time forecast of seasonal WNV outbreaks and their use as a quantitative decision support tool for public health officials and mosquito control programs.
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Predicción/métodos , Fiebre del Nilo Occidental/epidemiología , Fiebre del Nilo Occidental/transmisión , Animales , Culicidae , Brotes de Enfermedades , Humanos , Insectos Vectores/virología , Modelos Teóricos , Control de Mosquitos/tendencias , Salud Pública/tendencias , Estudios Retrospectivos , Estaciones del Año , Temperatura , Fiebre del Nilo Occidental/prevención & control , Virus del Nilo OccidentalRESUMEN
In October and November 2014, spectra covering the 1.436 - 1.863-µm wavelength range from the SINFONI Integral Field Unit Spectrometer on the Very Large Telescope showed the presence of a vast bright North polar cap on Uranus, extending northward from about 40°N and at all longitudes observed. The feature, first detected in August 2014 from Keck telescope images, has a morphology very similar to the southern polar cap that was seen to fade before the 2007 equinox. At strong methane-absorbing wavelengths (for which only the high troposphere or stratosphere is sampled) the feature is not visible, indicating that it is not a stratospheric phenomenon. We show that the observed northern bright polar cap results mainly from a decrease in the tropospheric methane mixing ratio, rather than from a possible latitudinal variation of the optical properties or abundance of aerosol, implying an increase in polar downwelling near the tropopause level.
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Saturn's moon Titan has a nitrogen atmosphere comparable to Earth's, with a surface pressure of 1.4 bar. Numerical models reproduce the tropospheric conditions very well but have trouble explaining the observed middle-atmosphere temperatures, composition and winds. The top of the middle-atmosphere circulation has been thought to lie at an altitude of 450 to 500 kilometres, where there is a layer of haze that appears to be separated from the main haze deck. This 'detached' haze was previously explained as being due to the co-location of peak haze production and the limit of dynamical transport by the circulation's upper branch. Here we report a build-up of trace gases over the south pole approximately two years after observing the 2009 post-equinox circulation reversal, from which we conclude that middle-atmosphere circulation must extend to an altitude of at least 600 kilometres. The primary drivers of this circulation are summer-hemisphere heating of haze by absorption of solar radiation and winter-hemisphere cooling due to infrared emission by haze and trace gases; our results therefore imply that these effects are important well into the thermosphere (altitudes higher than 500 kilometres). This requires both active upper-atmosphere chemistry, consistent with the detection of high-complexity molecules and ions at altitudes greater than 950 kilometres, and an alternative explanation for the detached haze, such as a transition in haze particle growth from monomers to fractal structures.
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BACKGROUND: Although many studies have been conducted regarding the effectiveness of medical nutrition therapy (MNT) for type 2 diabetes management, less is known about the effectiveness of MNT for low-income adults. This study evaluated the contribution of MNT in improving A1C and blood pressure in a population of low-income adults with type 2 diabetes. METHODS: This was a population-based, propensity score-matched cohort study using provincial health data from Altoona, Blair County, Pa. Patients who had been diagnosed with type 2 diabetes for at least 6 months before March 2014 were selected from two separate clinics that serve low-income populations. Patients who received MNT (n = 81) from a registered dietitian were compared to a matched group of patients who received primary care alone (n = 143). Outcome measures were A1C and systolic and diastolic blood pressure. The follow-up period was 1 year. RESULTS: Improvements in A1C and systolic and diastolic blood pressure were statistically significant for patients who received MNT at uniform 3-month intervals through 1 year. At the 1-year follow-up, A1C reduction was -0.8% (P <0.01), systolic blood pressure reduction was -8.2 mmHg (P <0.01), and diastolic blood pressure reduction was -4.3 mmHg (P <0.05). CONCLUSION: Although low-income individuals encounter a variety of barriers that reduce their capacity for success with and adherence to MNT, provision of nutrition therapy services by a registered dietitian experienced in addressing these barriers can be an effective addition to the existing medical components of type 2 diabetes care.
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Over the course of three years, 200 ft to 0.75 mi (60 m to 1.2 km) sections of 3 larger (>6 ft [1.8 m] diam) belowground storm sewer conveyance pipes in the northwestern Chicago suburbs were inspected for the presence of adult mosquitoes. Culex mosquitoes were by far the most common (555 of 556 [99.8%] total mosquitoes) collected within pipes during all four meteorological seasons (i.e. during months of October, January, May, August). These observations support prior work elsewhere, suggesting storm sewer pipes are consistent sites of refuge for adult Culex mosquitoes.
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Culex , Culicidae , Animales , Chicago , Estaciones del AñoRESUMEN
Over the course of three years, 200 ft to 0.75 mi (60 m to 1.2 km) sections of 3 larger (>6 ft [1.8 m] diam) belowground storm sewer conveyance pipes in the northwestern Chicago suburbs were inspected for the presence of adult mosquitoes. Culex mosquitoes were by far the most common (555 of 556 [99.8%] total mosquitoes) collected within pipes during all four meteorological seasons (i.e. during months of October, January, May, August). These observations support prior work elsewhere, suggesting storm sewer pipes are consistent sites of refuge for adult Culex mosquitoes.
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Since the introduction of West Nile virus (WNV) to the United States over 20 years ago, thousands of cases of human disease and death have been reported. Yearly seasonal outbreaks continue to persist, and the city and suburbs of Chicago, Illinois, is considered a "hot spot" for WNV activity. To interrupt WNV transmission, ground ultra-low volume (ULV) adulticide applications are regularly used to reduce Culex pipiens L. and Culex restuans Theobold (Diptera: Culicidae) abundance and infection. The real-world effectiveness of adulticide applications has not been comprehensively assessed, and prior studies, including our own investigation, have yielded inconclusive or conflicting results. Therefore, we expanded our prior work and evaluated the effects of 5 sequential weekly truck-mounted ULV adulticide applications in large residential areas in the northern suburbs of Chicago, Illinois, in 2019 and 2020. Each day, Cx. pipiens and Cx. restuans host-seeking and gravid mosquitoes were collected to assess abundance, age structure, and WNV infection rates. Adulticide applications resulted in significant reductions of both host-seeking and gravid abundance on the night of treatment. The reduction in host-seeking mosquitoes was followed by a reduction in gravid mosquitoes trapped 3 and 4 days after adulticide application and an increase in the proportion of nulliparous mosquitoes. WNV infection rates were significantly reduced in treatment sites as compared to untreated sites when infection rates were higher in 2020. This large-scale study provides comprehensive evidence that ground ULV adulticide applications are an effective tool in an integrated mosquito management program for combating WNV vectors and infection risk.
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Culex , Insecticidas , Control de Mosquitos , Fiebre del Nilo Occidental , Virus del Nilo Occidental , Animales , Culex/efectos de los fármacos , Fiebre del Nilo Occidental/transmisión , Fiebre del Nilo Occidental/prevención & control , Chicago , Insecticidas/farmacología , Femenino , Mosquitos Vectores/efectos de los fármacos , Illinois , MasculinoRESUMEN
Insect growth regulators, like S-methoprene, are heavily relied upon worldwide for larval mosquito chemical control due to their target specificity and long-lasting effects. In this study, susceptibility to S-methoprene was evaluated in Culex pipiens, a globally important vector species. Populations from 14 sites throughout the Chicago area with a long history of S-methoprene use and two sites with minimal use in Wisconsin were examined. Using a bioassay methodology and probit analyses, LC50 and LC90 values were calculated and compared to a susceptible laboratory strain to develop resistance ratios, then categorized for resistance intensity. The resistance ratios observed required the addition of another category, termed 'extreme' resistance, indicating resistance ratios greater than 100. 'Low' to 'extreme' levels of resistance to S-methoprene were detected throughout Illinois populations, with resistance ratios ranging from 2.33 to 1010.52. Resistance was not detected in populations where S-methoprene pressure has been very limited. These 'extreme' resistance ratios observed have never been documented in a wild vector species mosquito population. The relationships between historical S-methoprene use, resistance detected with laboratory bioassays, and the potential for field product failure remain unclear. However, the profound resistance detected here demonstrates a potential critical threat to protecting public health from mosquito-borne diseases.
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Culex , Metopreno , Animales , Culex/efectos de los fármacos , Chicago , Metopreno/farmacología , Resistencia a los Insecticidas , Mosquitos Vectores/efectos de los fármacos , Larva/efectos de los fármacos , Control de Mosquitos/métodosRESUMEN
West Nile virus (WNV) invaded the continental United States over 20 years ago and continues to cause yearly seasonal outbreaks of human and veterinary disease. In the suburbs of Chicago, Illinois, ultra-low volume (ULV) truck-mounted adulticide spraying frequently is performed to reduce populations of Culex restuans Theobald and Cx. pipiens L. mosquitoes (Diptera: Culicidae) in an effort to lower the risk of WNV transmission. The effectiveness of this control method has not been rigorously evaluated, and evidence for Culex population reduction after ULV adulticide spraying has been inconclusive. Therefore, we evaluated the results of 5 sequential weekly truck-mounted adulticide applications of Zenivex® E20 (etofenprox) in 2 paired sites located in Cook County, IL, during the summer of 2018. Mosquito population abundance, age structure, and WNV infection prevalence were monitored and compared between paired treatment and nearby control sites. Adulticide treatment did not result in consistent short-term or long-term reductions in target WNV vector Culex abundance. However, there was a significant increase in the proportion of nulliparous females in the treated sites compared to control sites and a decrease in Cx. pipiens WNV infection rates at one of the treated sites. This evidence that ULV adulticide spraying altered the age structure and WNV infection prevalence in a vector population has important implications for WNV transmission risk management. Our findings also underscore the importance of measuring these important indicators in addition to abundance metrics when evaluating the efficacy of control methods.
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Culex , Culicidae , Fiebre del Nilo Occidental , Virus del Nilo Occidental , Femenino , Animales , Humanos , Mosquitos VectoresRESUMEN
BACKGROUND: West Nile virus (WNV), primarily vectored by mosquitoes of the genus Culex, is the most important mosquito-borne pathogen in North America, having infected thousands of humans and countless wildlife since its arrival in the USA in 1999. In locations with dedicated mosquito control programs, surveillance methods often rely on frequent testing of mosquitoes collected in a network of gravid traps (GTs) and CO2-baited light traps (LTs). Traps specifically targeting oviposition-seeking (e.g. GTs) and host-seeking (e.g. LTs) mosquitoes are vulnerable to trap bias, and captured specimens are often damaged, making morphological identification difficult. METHODS: This study leverages an alternative mosquito collection method, the human landing catch (HLC), as a means to compare sampling of potential WNV vectors to traditional trapping methods. Human collectors exposed one limb for 15 min at crepuscular periods (5:00-8:30 am and 6:00-9:30 pm daily, the time when Culex species are most actively host-seeking) at each of 55 study sites in suburban Chicago, Illinois, for two summers (2018 and 2019). RESULTS: A total of 223 human-seeking mosquitoes were caught by HLC, of which 46 (20.6%) were mosquitoes of genus Culex. Of these 46 collected Culex specimens, 34 (73.9%) were Cx. salinarius, a potential WNV vector species not thought to be highly abundant in upper Midwest USA. Per trapping effort, GTs and LTs collected > 7.5-fold the number of individual Culex specimens than HLC efforts. CONCLUSIONS: The less commonly used HLC method provides important insight into the complement of human-biting mosquitoes in a region with consistent WNV epidemics. This study underscores the value of the HLC collection method as a complementary tool for surveillance to aid in WNV vector species characterization. However, given the added risk to the collector, novel mitigation methods or alternative approaches must be explored to incorporate HLC collections safely and strategically into control programs.
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Culex , Fiebre del Nilo Occidental , Virus del Nilo Occidental , Animales , Femenino , Humanos , Mosquitos Vectores , Animales Salvajes , Control de Mosquitos/métodosRESUMEN
ESA's Jupiter Icy Moons Explorer (JUICE) will provide a detailed investigation of the Jovian system in the 2030s, combining a suite of state-of-the-art instruments with an orbital tour tailored to maximise observing opportunities. We review the Jupiter science enabled by the JUICE mission, building on the legacy of discoveries from the Galileo, Cassini, and Juno missions, alongside ground- and space-based observatories. We focus on remote sensing of the climate, meteorology, and chemistry of the atmosphere and auroras from the cloud-forming weather layer, through the upper troposphere, into the stratosphere and ionosphere. The Jupiter orbital tour provides a wealth of opportunities for atmospheric and auroral science: global perspectives with its near-equatorial and inclined phases, sampling all phase angles from dayside to nightside, and investigating phenomena evolving on timescales from minutes to months. The remote sensing payload spans far-UV spectroscopy (50-210 nm), visible imaging (340-1080 nm), visible/near-infrared spectroscopy (0.49-5.56 µm), and sub-millimetre sounding (near 530-625 GHz and 1067-1275 GHz). This is coupled to radio, stellar, and solar occultation opportunities to explore the atmosphere at high vertical resolution; and radio and plasma wave measurements of electric discharges in the Jovian atmosphere and auroras. Cross-disciplinary scientific investigations enable JUICE to explore coupling processes in giant planet atmospheres, to show how the atmosphere is connected to (i) the deep circulation and composition of the hydrogen-dominated interior; and (ii) to the currents and charged particle environments of the external magnetosphere. JUICE will provide a comprehensive characterisation of the atmosphere and auroras of this archetypal giant planet.
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To increase the effectiveness of adult mosquito control, more frequent ultra-low volume (ULV) applications have been suggested. In the Chicago area, weather may be a limiting factor to more frequent applications. To investigate this, ULV adulticidal application dates from the Northwest Mosquito Abatement District during June 2011 to September 2020 were compared to historical weather data-wind and rain patterns to determine optimum conditions for conducting ULV treatments. During the 85 applications performed during 2011-20, there would have been an opportunity, based on weather conditions, for an additional consecutive night for about half (48.2%) of the time. These opportunities diminished with each additional application. The most intensive response possible would have been a spray of 5 consecutive nights, which occurred once in 3 of the 10 years investigated.
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Insecticidas , Chicago , Control de Mosquitos , Vehículos a Motor , Tiempo (Meteorología)RESUMEN
Culex pipiens complex is an important vector of epizootic and zoonotic pathogens, including West Nile virus. Chicago, Illinois and its suburbs have suffered high incidence of human West Nile virus infections in the past. This makes abatement programs in and around the Chicago area an essential service. The control of Cx. pipiens is often complicated by rapidly evolving resistance to pyrethroids, which are the most widely used chemical class in US mosquito abatement programs. The present study assessed Sumithrin® resistance in Cx. pipiens collected from five locations around Cook County, Illinois, neighboring the city limits of Chicago. According to CDC guidelines, samples from all five locations demonstrated some resistance to Sumithrin®. When assessed with Anvil®, a formulated product made of Sumithrin® synergized with piperonyl butoxide, susceptibility was rescued in mosquitoes from three out of the five locations, suggesting involvement of mixed-function oxidases and/or carboxylesterases in Sumithrin® resistance at these locations. Not all locations had susceptibility rescued by Anvil®, but these locations had relatively low knockdown resistance allele frequencies, suggesting that mechanisms other than knockdown resistance may be involved. Enzyme activities did not reveal any marked trends that could be related back to mortality in the bottle bioassays, which highlights the need for multiple types of assays to infer enzymatic involvement in resistance. Future directions in pyrethroid resistance management in Chicago area Cx. pipiens are discussed.
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Culex , Culicidae , Piretrinas , Virus del Nilo Occidental , Animales , Bioensayo , Chicago , Culex/genética , Humanos , Mosquitos Vectores , Piretrinas/farmacologíaRESUMEN
The Asian tiger mosquito, Aedes albopictus (Skuse), is a public health threat because it can potentially transmit multiple pathogenic arboviruses, exhibits aggressive diurnal biting, and is highly invasive. As Ae. albopictus moved northward into the United States, the limits of expansion were predicted as locations with a mean January temperature warmer than -2.5°C. We postulated that the range of Ae. albopictus could exceed these temperature limits if eggs in diapause overwinter in tires that provide an insulating effect from extreme temperatures. Fifteen tires with Ae. albopictus and Aedes triseriatus (Say) eggs, a native cold hardy species, were placed outside at five locations along a latitudinal gradient in Wisconsin and Illinois during the winter of 2018-2019; notably, in January 2019, a regional arctic air event brought the lowest temperatures recorded in over 20 yr. External and internal tire temperatures were recorded at 3 hr intervals, and egg survival was recorded after six months. Aedes albopictus eggs survived only from tires at northernmost locations. The mean internal January temperature of tires that supported survival was -1.8°C, while externally the mean temperature was -5.3°C, indicating that tires provided an average of +3.5°C of insulation. Tires that supported egg survival also had over 100 mm of snow cover during January. In the absence of snow cover, tires across the study area provided an average +0.79°C [95% CI 0.34-1.11] insulation. This work provides strong argument for the inclusion of microhabitats in models of dispersal and establishment of Ae. albopictus and other vector species.
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Aedes , Animales , Frío , Mosquitos Vectores , Estaciones del Año , Nieve , Estados UnidosRESUMEN
Over the course of 2019 and 2020, 7 larvicide formulations for use in catch basins were evaluated using a standardized pass/fail protocol. A comparison between 1- and 2-pouch doses of VectoLex® water-soluble pouches (WSP; 10 g per pouch) and 20 g of VectoLex FG (loose granules) suggests that the WSP formulation may have a shorter duration than the FG formulation. Results also suggest that 20 g doses of each of 2 larvicides, Duplex™-G and Sumilarv® 0.5G, may have a minimum effective duration in basins for approximately 40 days. A 20 g dose of Altosid® XR-G and a single briquet Altosid XR each had an observed minimum effective duration of 20 days. Sustain MBG had an observed duration of only 7 wk posttreatment. These observations provide evidence that, in some locations, 20 g granular applications may have a longer effective duration than a single briquet. This work highlights the utility of applying a standardized protocol for routine quality control assessments of the thousands of catch basin larvicide applications performed seasonally by mosquito abatement districts.
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Culex , Insecticidas , Animales , Illinois , Larva , Control de MosquitosRESUMEN
Modeling vector-borne diseases is best conducted when heterogeneity among interacting biotic and abiotic processes is captured. However, the successful integration of these complex processes is difficult, hindered by a lack of understanding of how these relationships influence disease transmission across varying scales. West Nile virus (WNV) is the most important mosquito-borne disease in the United States. Vectored by Culex mosquitoes and maintained in the environment by avian hosts, the virus can spill over into humans and horses, sometimes causing severe neuroinvasive illness. Several modeling studies have evaluated drivers of WNV disease risk, but nearly all have done so at broad scales and have reported mixed results of the effects of common explanatory variables. As a result, fine-scale relationships with common explanatory variables, particularly climatic, socioeconomic, and human demographic, remain uncertain across varying spatial extents. Using an interdisciplinary approach and an ongoing 12-year study of the Chicago region, this study evaluated the factors explaining WNV disease risk at high spatiotemporal resolution, comparing the human WNV model and covariate performance across three increasing spatial extents: ultrafine, local, and county scales. Our results demonstrate that as spatial extent increased, model performance increased. In addition, only six of the 23 assessed covariates were included in best-fit models of at least two scales. These results suggest that the mechanisms driving WNV ecology are scale-dependent and covariate importance increases as extent decreases. These tools may be particularly helpful for public health, mosquito, and disease control personnel in predicting and preventing disease within local and fine-scale jurisdictions, before spillover occurs.
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Demografía , Modelos Biológicos , Fiebre del Nilo Occidental/epidemiología , Humanos , Illinois , Factores de RiesgoRESUMEN
West Nile virus (WNV) is a globally distributed mosquito-borne virus of great public health concern. The number of WNV human cases and mosquito infection patterns vary in space and time. Many statistical models have been developed to understand and predict WNV geographic and temporal dynamics. However, these modeling efforts have been disjointed with little model comparison and inconsistent validation. In this paper, we describe a framework to unify and standardize WNV modeling efforts nationwide. WNV risk, detection, or warning models for this review were solicited from active research groups working in different regions of the United States. A total of 13 models were selected and described. The spatial and temporal scales of each model were compared to guide the timing and the locations for mosquito and virus surveillance, to support mosquito vector control decisions, and to assist in conducting public health outreach campaigns at multiple scales of decision-making. Our overarching goal is to bridge the existing gap between model development, which is usually conducted as an academic exercise, and practical model applications, which occur at state, tribal, local, or territorial public health and mosquito control agency levels. The proposed model assessment and comparison framework helps clarify the value of individual models for decision-making and identifies the appropriate temporal and spatial scope of each model. This qualitative evaluation clearly identifies gaps in linking models to applied decisions and sets the stage for a quantitative comparison of models. Specifically, whereas many coarse-grained models (county resolution or greater) have been developed, the greatest need is for fine-grained, short-term planning models (m-km, days-weeks) that remain scarce. We further recommend quantifying the value of information for each decision to identify decisions that would benefit most from model input.
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Toma de Decisiones , Modelos Biológicos , Administración en Salud Pública , Fiebre del Nilo Occidental/prevención & control , HumanosRESUMEN
The occurrence of multiple pathogens in mosquitoes and birds could affect the dynamics of disease transmission. We collected adult Culex pipiens and Cx. restuans (Cx. pipiens/restuans hereafter) from sites in Wisconsin and tested them for West Nile virus (WNV) and for avian malaria (Plasmodium). Gravid Cx. pipiens/restuans were tested for WNV using a commercial immunoassay, the RAMP WNV test, and positive results were verified by reverse transcriptase-polymerase chain reaction. There were 2 WNV-positive pools of Cx. pipiens/restuans in 2006 and 1 in 2007. Using a bias-corrected maximum likelihood estimation, the WNV infection rate for Cx. pipiens/restuans was 5.48/1,000 mosquitoes in 2006 and 1.08/1,000 mosquitoes in 2007. Gravid Cx. pipiens or Cx. restuans were tested individually for avian Plasmodium by a restriction enzyme-based assay. Twelve mosquitoes were positive for avian Plasmodium (10.0%), 2 were positive for Haemoproteus, and 3 were positive for Leucocytozoon. There were 4 mixed infections, with mosquitoes positive for > 1 of the hemosporidian parasites. This work documents a high rate of hemosporidian infection in Culex spp. and illustrates the potential for co-infections with other arboviruses in bird-feeding mosquitoes and their avian hosts. In addition, hemosporidian infection rates may be a useful tool for investigating the ecological dynamics of Culex/avian interactions.
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Culex/parasitología , Culex/virología , Malaria Aviar/transmisión , Plasmodium/aislamiento & purificación , Virus del Nilo Occidental/aislamiento & purificación , Animales , Aves , Insectos Vectores/parasitología , Insectos Vectores/virología , WisconsinRESUMEN
Atmospheric circulation patterns derived from multi-spectral remote sensing can serve as a guide for choosing a suitable entry location for a future in situ probe mission to the Ice Giants. Since the Voyager-2 flybys in the 1980s, three decades of observations from ground- and space-based observatories have generated a picture of Ice Giant circulation that is complex, perplexing, and altogether unlike that seen on the Gas Giants. This review seeks to reconcile the various competing circulation patterns from an observational perspective, accounting for spatially-resolved measurements of: zonal albedo contrasts and banded appearances; cloud-tracked zonal winds; temperature and para-H2 measurements above the condensate clouds; and equator-to-pole contrasts in condensable volatiles (methane, ammonia, and hydrogen sulphide) in the deeper troposphere. These observations identify three distinct latitude domains: an equatorial domain of deep upwelling and upper-tropospheric subsidence, potentially bounded by peaks in the retrograde zonal jet and analogous to Jovian cyclonic belts; a mid-latitude transitional domain of upper-tropospheric upwelling, vigorous cloud activity, analogous to Jovian anticyclonic zones; and a polar domain of strong subsidence, volatile depletion, and small-scale (and potentially seasonally-variable) convective activity. Taken together, the multi-wavelength observations suggest a tiered structure of stacked circulation cells (at least two in the troposphere and one in the stratosphere), potentially separated in the vertical by (i) strong molecular weight gradients associated with cloud condensation, and by (ii) transitions from a thermally-direct circulation regime at depth to a wave- and radiative-driven circulation regime at high altitude. The inferred circulation can be tested in the coming decade by 3D numerical simulations of the atmosphere, and by observations from future world-class facilities. The carrier spacecraft for any probe entry mission must ultimately carry a suite of remote-sensing instruments capable of fully constraining the atmospheric motions at the probe descent location.