IMILAST: A community effort to intercompare extratropical cyclone detection and tracking algorithms: assessing method-related uncertainties

The variability of results from different automated methods of detection and tracking of extratropical cyclones is assessed in order to identify uncertainties related to the choice of method. Fifteen international teams applied their own algorithms to the same dataset - the period 1989-2009 of interim European Centre for Medium-Range Weather Forecasts (ECMWF) Re-Analysis (ERAInterim) data. This experiment is part of the community project Intercomparison of Mid Latitude Storm Diagnostics (IMILAST; see www.proclim.ch/imilast/index.html). The spread of results for cyclone frequency, intensity, life cycle, and track location is presented to illustrate the impact of using different methods. Globally, methods agree well for geographical distribution in large oceanic regions, interannual variability of cyclone numbers, geographical patterns of strong trends, and distribution shape for many life cycle characteristics. In contrast, the largest disparities exist for the total numbers of cyclones, the detection of weak cyclones, and distribution in some densely populated regions. Consistency between methods is better for strong cyclones than for shallow ones. Two case studies of relatively large, intense cyclones reveal that the identification of the most intense part of the life cycle of these events is robust between methods, but considerable differences exist during the development and the dissolution phases.

First middle-atmospheric zonal wind profile measurements with a new ground-based microwave Doppler-spectro-radiometer

We report on the wind radiometer WIRA, a new ground-based microwave Doppler-spectro-radiometer specifically designed for the measurement of middle-atmospheric horizontal wind by observing ozone emission spectra at 142.17504 GHz. Currently, wind speeds in five levels between 30 and 79 km can be retrieved which makes WIRA the first instrument able to continuously measure horizontal wind in this altitude range. For an integration time of one day the measurement error on each level lies at around 25 m s−1. With a planned upgrade this value is expected to be reduced by a factor of 2 in the near future. On the altitude levels where our measurement can be compared to wind data from the European Centre for Medium-Range Weather Forecasts (ECMWF) very good agreement in the long-term statistics as well as in short time structures with a duration of a few days has been found. WIRA uses a passive double sideband heterodyne receiver together with a digital Fourier transform spectrometer for the data acquisition. A big advantage of the radiometric approach is that such instruments can also operate under adverse weather conditions and thus provide a continuous time series for the given location. The optics enables the instrument to scan a wide range of azimuth angles including the directions east, west, north, and south for zonal and meridional wind measurements. The design of the radiometer is fairly compact and its calibration does not rely on liquid nitrogen which makes it transportable and suitable for campaign use. WIRA is conceived in a way that it can be operated remotely and does hardly require any maintenance. In the present paper, a description of the instrument is given, and the techniques used for the wind retrieval based on the determination of the Doppler shift of the measured atmospheric ozone emission spectra are outlined. Their reliability was tested using Monte Carlo simulations. Finally, a time series of 11 months of zonal wind measurements over Bern (46°57′ N, 7°26′ E) is presented and compared to ECMWF wind data.

Polycyclic aromatic compounds (PAHs and oxygenated PAHs) and trace metals in fish species from Ghana (West Africa): Bioaccumulation and health risk assessment

We report the concentrations of 28 PAHs, 15 oxygenated PAHs (OPAHs) and 11 trace metals/metalloids (As, Cd, Co, Cr, Cu, Fe, Mn, Ni, Pb, Se, and Zn) in muscle and gut + gill tissues of demersal fishes (Drapane africana, Cynoglossus senegalensis and Pomadasys peroteti) from three locations along the coast of the Gulf of Guinea (Ghana). The concentrations of ∑ 28PAHs in muscle tissues averaged 192 ng g− 1 dw (range: 71–481 ng g− 1 dw) and were not statistically different between locations. The concentrations of ∑ 28 PAHs were higher in guts + gills than in muscles. The PAH composition pattern was dominated by low molecular weight compounds (naphthalene, alkyl-naphthalenes and phenanthrene). All fish tissues had benzo[a]pyrene concentrations lower than the EU limit for food safety. Excess cancer risk from consumption of some fish was higher than the guideline value of 1 × 10− 6. The concentrations of ∑ 15 OPAHs in fish muscles averaged 422 ng g− 1 dw (range: 28–1715 ng g− 1dw). The ∑ 15 OPAHs/∑ 16 US-EPA PAHs concentration ratio was > 1 in 68% of the fish muscles and 100% of guts + gills. The log-transformed concentrations of PAHs and OPAHs in muscles, guts + gills were significantly (p < 0.05) correlated with their octanol–water partitioning coefficients, strongly suggesting that equilibrium partitioning from water/sediment into fish tissue was the main mechanism of bioaccumulation. The trace metal concentrations in the fish tissues were in the medium range when compared to fish from other parts of the world. The concentrations of some trace metals (Cd, Cu, Fe, Mn, Zn) were higher in guts + gills than in muscle tissues. The target hazard quotients for metals were < 1 and did not indicate a danger to the local population. We conclude that the health risk arising from the consumption of the studied fish (due to their PAHs and trace metals content) is minimal.

A Lagrangian Analysis of the Northern Hemisphere Subtropical Jet

This study presents a 5-yr climatology of 7-day back trajectories started from the Northern Hemisphere subtropical jet. These trajectories provide insight into the seasonally and regionally varying angular momentum and potential vorticity characteristics of the air parcels that end up in the subtropical jet. The trajectories reveal preferred pathways of the air parcels that reach the subtropical jet from the tropics and the extratropics and allow estimation of the tropical and extratropical forcing of the subtropical jet. The back trajectories were calculated 7 days back in time and started every 6 h from December 2005 to November 2010 using the Interim European Centre for Medium-Range Weather Forecasts (ECMWF) Re-Analysis (ERA-Interim) dataset as a basis. The trajectories were started from the 345-K isentrope in areas where the wind speed exceeded a seasonally varying threshold and where the wind shear was confined to upper levels. During winter, the South American continent, the Indian Ocean, and the Maritime Continent are preferred areas of ascent into the upper troposphere. From these areas, air parcels follow an anticyclonic pathway into the subtropical jet. During summer, the majority of air parcels ascend over the Himalayas and Southeast Asia. Angular momentum is overall well conserved for trajectories that reach the subtropical jet from the deep tropics. In winter and spring, the hemispheric-mean angular momentum loss amounts to approximately 6%; in summer, it amounts to approximately 18%; and in fall, it amounts to approximately 13%. This seasonal variability is confirmed using an independent potential vorticity–based method to estimate tropical and extratropical forcing of the subtropical jet.

A global radiosonde and tracked balloon archive on 16 pressure levels (GRASP) back to 1905 – Part 1: Merging and interpolation to 00:00 and 12:00 GMT

Many observed time series of the global radiosonde or PILOT networks exist as fragments distributed over different archives. Identifying and merging these fragments can enhance their value for studies on the three-dimensional spatial structure of climate change. The Comprehensive Historical Upper-Air Network (CHUAN version 1.7), which was substantially extended in 2013, and the Integrated Global Radiosonde Archive (IGRA) are the most important collections of upper-air measurements taken before 1958. CHUAN (tracked) balloon data start in 1900, with higher numbers from the late 1920s onward, whereas IGRA data start in 1937. However, a substantial fraction of those measurements have not been taken at synoptic times (preferably 00:00 or 12:00 GMT) and on altitude levels instead of standard pressure levels. To make them comparable with more recent data, the records have been brought to synoptic times and standard pressure levels using state-of-the-art interpolation techniques, employing geopotential information from the National Oceanic and Atmospheric Administration (NOAA) 20th Century Reanalysis (NOAA 20CR). From 1958 onward the European Re-Analysis archives (ERA-40 and ERA-Interim) available at the European Centre for Medium-Range Weather Forecasts (ECMWF) are the main data sources. These are easier to use, but pilot data still have to be interpolated to standard pressure levels. Fractions of the same records distributed over different archives have been merged, if necessary, taking care that the data remain traceable back to their original sources. If possible, station IDs assigned by the World Meteorological Organization (WMO) have been allocated to the station records. For some records which have never been identified by a WMO ID, a local ID above 100 000 has been assigned. The merged data set contains 37 wind records longer than 70 years and 139 temperature records longer than 60 years. It can be seen as a useful basis for further data processing steps, most notably homogenization and gridding, after which it should be a valuable resource for climatological studies. Homogeneity adjustments for wind using the NOAA-20CR as a reference are described in Ramella Pralungo and Haimberger (2014). Reliable homogeneity adjustments for temperature beyond 1958 using a surface-data-only reanalysis such as NOAA-20CR as a reference have yet to be created. All the archives and metadata files are available in ASCII and netCDF format in the PANGAEA archive

On the Co-Occurrence of Warm Conveyor Belt Outflows and PV Streamers*

The co-occurrence of warm conveyor belts (WCBs), strongly ascending moist airstreams in extratropical cyclones, and stratospheric potential vorticity (PV) streamers, indicators for breaking Rossby waves on the tropopause, is investigated for a 21-yr period in the Northern Hemisphere using Interim European Centre for Medium-Range Weather Forecasts (ECMWF) Re-Analysis (ERA-Interim) data. WCB outflows and PV streamers are respectively identified as two- and three-dimensional objects and tracked during their life cycle. PV streamers are more frequent than WCB outflows and nearly 15% of all PV streamers co-occur with WCBs during their life cycle, whereas about 60% of all WCB outflows co-occur with PV streamers. Co-occurrences are most frequent over the North Atlantic and North Pacific in spring and winter. WCB outflows are often located upstream of the PV streamers and form earlier, indicating the importance of diabatic processes for downstream Rossby wave breaking. Less frequently, PV streamers occur first, leading to the formation of new WCBs.

Land-atmosphere coupling over North America in CRCM5

Land-atmosphere coupling and its impact on extreme precipitation and temperature events over North America are studied using the fifth generation of the Canadian Regional Climate Model (CRCM5). To this effect, two 30 year long simulations, spanning the 1981–2010 period, with and without land-atmosphere coupling, have been performed with CRCM5, driven by the European Centre for Medium-Range Weather Forecasts reanalysis at the boundaries. In the coupled simulation, the soil moisture interacts freely with the atmosphere at each time step, while in the uncoupled simulation, soil moisture is replaced with its climatological value computed from the coupled simulation, thus suppressing the soil moisture-atmosphere interactions. Analyses of the coupled and uncoupled simulations, for the summer period, show strong soil moisture-temperature coupling over the Great Plains, consistent with previous studies. The maxima of soil moisture-precipitation coupling is more spread out and covers the semiarid regions of the western U.S. and parts of the Great Plains. However, the strength of soil moisture-precipitation coupling is found to be generally weaker than that of soil moisture-temperature coupling. The study clearly indicates that land-atmosphere coupling increases the interannual variability of the seasonal mean daily maximum temperature in the Great Plains. Land-atmosphere coupling is found to significantly modulate selected temperature extremes such as the number of hot days, frequency, and maximum duration of hot spells over the Great Plains. Results also suggest additional hot spots, where soil moisture modulates extreme events. These hot spots are located in the southeast U.S. for the hot days/hot spells and in the semiarid regions of the western U.S. for extreme wet spells. This study thus demonstrates that climatologically wet/dry regions can become hot spots of land-atmosphere coupling when the soil moisture decreases/increases to an intermediate transitional level where evapotranspiration becomes moisture sensitive and large enough to affect the climate.

Lake surface water temperatures of European Alpine lakes (1989–2013) based on the Advanced Very High Resolution Radiometer (AVHRR) 1 km data set

Lake water temperature (LWT) is an important driver of lake ecosystems and it has been identified as an indicator of climate change. Consequently, the Global Climate Observing System (GCOS) lists LWT as an essential climate variable. Although for some European lakes long in situ time series of LWT do exist, many lakes are not observed or only on a non-regular basis making these observations insufficient for climate monitoring. Satellite data can provide the information needed. However, only few satellite sensors offer the possibility to analyse time series which cover 25 years or more. The Advanced Very High Resolution Radiometer (AVHRR) is among these and has been flown as a heritage instrument for almost 35 years. It will be carried on for at least ten more years, offering a unique opportunity for satellite-based climate studies. Herein we present a satellite-based lake surface water temperature (LSWT) data set for European water bodies in or near the Alps based on the extensive AVHRR 1 km data record (1989–2013) of the Remote Sensing Research Group at the University of Bern. It has been compiled out of AVHRR/2 (NOAA-07, -09, -11, -14) and AVHRR/3 (NOAA-16, -17, -18, -19 and MetOp-A) data. The high accuracy needed for climate related studies requires careful pre-processing and consideration of the atmospheric state. The LSWT retrieval is based on a simulation-based scheme making use of the Radiative Transfer for TOVS (RTTOV) Version 10 together with ERA-interim reanalysis data from the European Centre for Medium-range Weather Forecasts. The resulting LSWTs were extensively compared with in situ measurements from lakes with various sizes between 14 and 580 km2 and the resulting biases and RMSEs were found to be within the range of −0.5 to 0.6 K and 1.0 to 1.6 K, respectively. The upper limits of the reported errors could be rather attributed to uncertainties in the data comparison between in situ and satellite observations than inaccuracies of the satellite retrieval. An inter-comparison with the standard Moderate-resolution Imaging Spectroradiometer (MODIS) Land Surface Temperature product exhibits RMSEs and biases in the range of 0.6 to 0.9 and −0.5 to 0.2 K, respectively. The cross-platform consistency of the retrieval was found to be within ~ 0.3 K. For one lake, the satellite-derived trend was compared with the trend of in situ measurements and both were found to be similar. Thus, orbital drift is not causing artificial temperature trends in the data set. A comparison with LSWT derived through global sea surface temperature (SST) algorithms shows lower RMSEs and biases for the simulation-based approach. A running project will apply the developed method to retrieve LSWT for all of Europe to derive the climate signal of the last 30 years. The data are available at doi:10.1594/PANGAEA.831007.

A 49 year hindcast of surface winds over the Iberian Peninsula

The use of hindcast climatic data is quite extended for multiple applications. However, this approach needs the support of a validation process to allow its drawbacks and, therefore, confidence levels to be assessed. In this work, the strategy relies on an hourly wind database resulting from a dynamical downscaling experiment, with a spatial resolution of 10 km, covering the Iberian Peninsula (IP), driven by the ERA40 reanalysis (1959–2001) extended by European Centre for Medium-Range Weather Forecast (ECMWF) analysis (2002–2007) and comprising two main steps. Initially, the skill of the simulation is evaluated comparing the quality-tested observational database (Lorente-Plazas et al., 2014) at local and regional scales. The results show that the model is able to portray the main features of the wind over the IP: annual cycles, wind roses, spatial and temporal variability, as well as the response to different circulation types. In addition, there is a significant added value of the simulation with respect to driving conditions, especially in regions with a complex orography. However, some problems are evident, the major drawback being the systematic overestimation of the wind speed, which is mainly attributed to a missrepresentation of frictional forces. The model skill is also lower along the Mediterranean coast and for the Pyrenees. In a second phase, the high spatio-temporal resolution of the pseudo-real wind database is used to explore the limitations of the observational database. It is shown that missing values do not affect the characterisation of the wind climate over the IP, while the length of the observational period (6 years) is sufficient for most regions, with only a few exceptions. The spatial distribution of the observational sampling schemes should be enhanced to improve the correct assessment of all IP wind regimes, particularly in some mountainous areas.

Flood triggering in Switzerland: the role of daily to monthly preceding precipitation

Determining the role of different precipitation periods for peak discharge generation is crucial for both projecting future changes in flood probability and for short- and medium-range flood forecasting. In this study, catchment-averaged daily precipitation time series are analyzed prior to annual peak discharge events (floods) in Switzerland. The high number of floods considered – more than 4000 events from 101 catchments have been analyzed – allows to derive significant information about the role of antecedent precipitation for peak discharge generation. Based on the analysis of precipitation times series, a new separation of flood-related precipitation periods is proposed: (i) the period 0 to 1 day before flood days, when the maximum flood-triggering precipitation rates are generally observed, (ii) the period 2 to 3 days before flood days, when longer-lasting synoptic situations generate "significantly higher than normal" precipitation amounts, and (iii) the period from 4 days to 1 month before flood days when previous wet episodes may have already preconditioned the catchment. The novelty of this study lies in the separation of antecedent precipitation into the precursor antecedent precipitation (4 days before floods or earlier, called PRE-AP) and the short range precipitation (0 to 3 days before floods, a period when precipitation is often driven by one persistent weather situation like e.g., a stationary low-pressure system). A precise separation of "antecedent" and "peak-triggering" precipitation is not attempted. Instead, the strict definition of antecedent precipitation periods permits a direct comparison of all catchments. The precipitation accumulating 0 to 3 days before an event is the most relevant for floods in Switzerland. PRE-AP precipitation has only a weak and region-specific influence on flood probability. Floods were significantly more frequent after wet PRE-AP periods only in the Jura Mountains, in the western and eastern Swiss plateau, and at the outlet of large lakes. As a general rule, wet PRE-AP periods enhance the flood probability in catchments with gentle topography, high infiltration rates, and large storage capacity (karstic cavities, deep soils, large reservoirs). In contrast, floods were significantly less frequent after wet PRE-AP periods in glacial catchments because of reduced melt. For the majority of catchments however, no significant correlation between precipitation amounts and flood occurrences is found when the last 3 days before floods are omitted in the precipitation amounts. Moreover, the PRE-AP was not higher for extreme floods than for annual floods with a high frequency and was very close to climatology for all floods. The fact that floods are not significantly more frequent nor more intense after wet PRE-AP is a clear indicator of a short discharge memory of Pre-Alpine, Alpine and South Alpine Swiss catchments. Our study poses the question whether the impact of long-term precursory precipitation for floods in such catchments is not overestimated in the general perception. The results suggest that the consideration of a 3–4 days precipitation period should be sufficient to represent (understand, reconstruct, model, project) Swiss Alpine floods.

On-Surface Synthesis of BN-Substituted Heteroaromatic Networks

We report on the bottom-up fabrication of BN-substituted heteroaromatic networks achieved by surface-assisted polymerization and subsequent cyclodehydrogenation of specifically designed BN-substituted precursor monomers based on a borazine core structural element. To get insight into the cyclodehydrogenation pathway and the influence of molecular flexibility on network quality, two closely related precursor monomers with different degrees of internal cyclodehydrogenation have been employed. Scanning tunneling microscopy shows that, for both monomers, surface-assisted cyclodehydrogenation allows for complete monomer cyclization and the formation of covalently interlinked BN-substituted polyaromatic hydrocarbon networks on the Ag(111) surface. In agreement with experimental observations, density functional theory calculations reveal a significantly lower energy barrier for the cyclodehydrogenation of the conformationally more rigid precursor monomer, which is also reflected in a higher degree of long-range order of the obtained heteroaromatic network. Our proof-of-concept study will allow for the fabrication of atomically precise substitution patterns within BNC heterostructures.

Composition Space Analysis in the Development of Copper Molybdate Hybrids Decorated by a Bifunctional Pyrazolyl/1,2,4-Triazole Ligand

A bitopic ligand, 4-(3,5-dimethylpyrazol-4-yl)-1,2,4-triazole (Hpz-tr) (1), containing two different heterocyclic moieties was employed for the design of copper(II)–molybdate solids under hydrothermal conditions. In the multicomponent CuII/Hpz-tr/MoVI system, a diverse set of coordination hybrids, [Cu(Hpz-tr)2SO4]·3H2O (2), [Cu(Hpz-tr)Mo3O10] (3), [Cu4(OH)4(Hpz-tr)4Mo8O26]·6H2O (4), [Cu(Hpz-tr)2Mo4O13] (5), and [Mo2O6(Hpz-tr)]·H2O (6), was prepared and characterized. A systematic investigation of these systems in the form of a ternary crystallization diagram approach was utilized to show the influence of the molar ratios of starting reagents, the metal (CuII and MoVI) sources, the temperature, etc., on the reaction products outcome. Complexes 2–4 dominate throughout a wide crystallization range of the composition triangle, while the other two compounds 5 and 6 crystallize as minor phases in a narrow concentration range. In the crystal structures of 2–6, the organic ligand behaves as a short [N–N]-triazole linker between metal centers Cu···Cu in 2–4, Cu···Mo in 5, and Mo···Mo in 6, while the pyrazolyl function remains uncoordinated. This is the reason for the exceptional formation of low-dimensional coordination motifs: 1D for 2, 4, and 6 and 2D for 3 and 5. In all cases, the pyrazolyl group is involved in H bonding (H-donor/H-acceptor) and is responsible for π–π stacking, thus connecting the chain and layer structures in more complicated H-bonding architectures. These compounds possess moderate thermal stability up to 250–300 °C. The magnetic measurements were performed for 2–4, revealing in all three cases antiferromagnetic exchange interactions between neighboring CuII centers and long-range order with a net moment below Tc of 13 K for compound 4.

GROMOS-C, a novel ground-based microwave radiometer for ozone measurement campaigns

Stratospheric ozone is of major interest as it absorbs most harmful UV radiation from the sun, allowing life on Earth. Ground-based microwave remote sensing is the only method that allows for the measurement of ozone profiles up to the mesopause, over 24 hours and under different weather conditions with high time resolution. In this paper a novel ground-based microwave radiometer is presented. It is called GROMOS-C (GRound based Ozone MOnitoring System for Campaigns), and it has been designed to measure the vertical profile of ozone distribution in the middle atmosphere by observing ozone emission spectra at a frequency of 110.836 GHz. The instrument is designed in a compact way which makes it transportable and suitable for outdoor use in campaigns, an advantageous feature that is lacking in present day ozone radiometers. It is operated through remote control. GROMOS-C is a total power radiometer which uses a pre-amplified heterodyne receiver, and a digital fast Fourier transform spectrometer for the spectral analysis. Among its main new features, the incorporation of different calibration loads stands out; this includes a noise diode and a new type of blackbody target specifically designed for this instrument, based on Peltier elements. The calibration scheme does not depend on the use of liquid nitrogen; therefore GROMOS-C can be operated at remote places with no maintenance requirements. In addition, the instrument can be switched in frequency to observe the CO line at 115 GHz. A description of the main characteristics of GROMOS-C is included in this paper, as well as the results of a first campaign at the High Altitude Research Station at Jungfraujoch (HFSJ), Switzerland. The validation is performed by comparison of the retrieved profiles against equivalent profiles from MLS (Microwave Limb Sounding) satellite data, ECMWF (European Centre for Medium-Range Weather Forecast) model data, as well as our nearby NDACC (Network for the Detection of Atmospheric Composition Change) ozone radiometer measuring at Bern.

Comparison of co-located independent ground-based middle atmospheric wind and temperature measurements with numerical weather prediction models

High-resolution, ground-based and independent observations including co-located wind radiometer, lidar stations, and infrasound instruments are used to evaluate the accuracy of general circulation models and data-constrained assimilation systems in the middle atmosphere at northern hemisphere midlatitudes. Systematic comparisons between observations, the European Centre for Medium-Range Weather Forecasts (ECMWF) operational analyses including the recent Integrated Forecast System cycles 38r1 and 38r2, the NASA’s Modern-Era Retrospective Analysis for Research and Applications (MERRA) reanalyses, and the free-running climate Max Planck Institute–Earth System Model–Low Resolution (MPI-ESM-LR) are carried out in both temporal and spectral dom ains. We find that ECMWF and MERRA are broadly consistent with lidar and wind radiometer measurements up to ~40 km. For both temperature and horizontal wind components, deviations increase with altitude as the assimilated observations become sparser. Between 40 and 60 km altitude, the standard deviation of the mean difference exceeds 5 K for the temperature and 20 m/s for the zonal wind. The largest deviations are observed in winter when the variability from large-scale planetary waves dominates. Between lidar data and MPI-ESM-LR, there is an overall agreement in spectral amplitude down to 15–20 days. At shorter time scales, the variability is lacking in the model by ~10 dB. Infrasound observations indicate a general good agreement with ECWMF wind and temperature products. As such, this study demonstrates the potential of the infrastructure of the Atmospheric Dynamics Research Infrastructure in Europe project that integrates various measurements and provides a quantitative understanding of stratosphere-troposphere dynamical coupling for numerical weather prediction applications.

Glycoprotein D of bovine herpesvirus 5 (BoHV-5) confers an extended host range to BoHV-1 but does not contribute to invasion of the brain

Bovine herpesvirus 1 (BoHV-1) and BoHV-5 are closely related pathogens of cattle, but only BoHV-5 is considered a neuropathogen. We engineered intertypic gD exchange mutants with BoHV-1 and BoHV-5 backbones in order to address their in vitro and in vivo host ranges, with particular interest in invasion of the brain. The new viruses replicated in cell culture with similar dynamics and to titers comparable to those of their wild-type parents. However, gD of BoHV-5 (gD5) was able to interact with a surprisingly broad range of nectins. In vivo, gD5 provided a virulent phenotype to BoHV-1 in AR129 mice, featuring a high incidence of neurological symptoms and early onset of disease. However, only virus with the BoHV-5 backbone, independent of the gD type, was detected in the brain by immunohistology. Thus, gD of BoHV-5 confers an extended cellular host range to BoHV-1 and may be considered a virulence factor but does not contribute to the invasion of the brain.