1. |
- Heald, G. H., et al.
(author)
-
The LOFAR Multifrequency Snapshot Sky Survey (MSSS) : I. Survey description and first results
- 2015
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 582, s. 1-22
-
Journal article (peer-reviewed)abstract
- We present the Multifrequency Snapshot Sky Survey (MSSS), the first northern-sky Low Frequency Array (LOFAR) imaging survey. In this introductory paper, we first describe in detail the motivation and design of the survey. Compared to previous radio surveys, MSSS is exceptional due to its intrinsic multifrequency nature providing information about the spectral properties of the detected sources over more than two octaves (from 30 to 160 MHz). The broadband frequency coverage, together with the fast survey speed generated by LOFAR’s multibeaming capabilities, make MSSS the first survey of the sort anticipated to be carried out with the forthcoming Square Kilometre Array (SKA). Two of the sixteen frequency bands included in the survey were chosen to exactly overlap the frequency coverage of large-area Very Large Array (VLA) and Giant Metrewave Radio Telescope (GMRT) surveys at 74 MHz and 151 MHz respectively. The survey performance is illustrated within the MSSS Verification Field (MVF), a region of 100 square degrees centered at (α,δ)J2000 = (15h,69°). The MSSS results from the MVF are compared with previous radio survey catalogs. We assess the flux and astrometric uncertainties in the catalog, as well as the completeness and reliability considering our source finding strategy. We determine the 90% completeness levels within the MVF to be 100 mJy at 135 MHz with 108″ resolution, and 550 mJy at 50 MHz with 166″ resolution. Images and catalogs for the full survey, expected to contain 150 000–200 000 sources, will be released to a public web server. We outline the plans for the ongoing production of the final survey products, and the ultimate public release of images and source catalogs.
|
|
2. |
- Buitink, S., et al.
(author)
-
A large light-mass component of cosmic rays at 1017–1017.5 electronvolts from radio observations
- 2016
-
In: Nature. - : Springer Science and Business Media LLC. - 0028-0836 .- 1476-4687. ; 531:7592, s. 70-73
-
Journal article (peer-reviewed)abstract
- Cosmic rays are the highest-energy particles found in nature. Measurements of the mass composition of cosmic rays with energies of 1017–1018 electronvolts are essential to understanding whether they have galactic or extragalactic sources. It has also been proposed that the astrophysical neutrino signal1 comes from accelerators capable of producing cosmic rays of these energies2. Cosmic rays initiate air showers—cascades of secondary particles in the atmosphere—and their masses can be inferred from measurements of the atmospheric depth of the shower maximum3 (Xmax; the depth of the air shower when it contains the most particles) or of the composition of shower particles reaching the ground4. Current measurements5 have either high uncertainty, or a low duty cycle and a high energy threshold. Radio detection of cosmic rays6, 7, 8 is a rapidly developing technique9 for determining Xmax (refs 10, 11) with a duty cycle of, in principle, nearly 100 per cent. The radiation is generated by the separation of relativistic electrons and positrons in the geomagnetic field and a negative charge excess in the shower front6, 12. Here we report radio measurements of Xmax with a mean uncertainty of 16 grams per square centimetre for air showers initiated by cosmic rays with energies of 1017–1017.5 electronvolts. This high resolution in Xmax enables us to determine the mass spectrum of the cosmic rays: we find a mixed composition, with a light-mass fraction (protons and helium nuclei) of about 80 per cent. Unless, contrary to current expectations, the extragalactic component of cosmic rays contributes substantially to the total flux below 1017.5 electronvolts, our measurements indicate the existence of an additional galactic component, to account for the light composition that we measured in the 1017–1017.5 electronvolt range.
|
|
3. |
- Shulevski, A., et al.
(author)
-
The peculiar radio galaxy 4C 35.06 : a case for recurrent AGN activity?
- 2015
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 579, s. 1-10
-
Journal article (peer-reviewed)abstract
- Using observations obtained with the LOw Fequency ARray (LOFAR), the Westerbork Synthesis Radio Telescope (WSRT) and archival Very Large Array (VLA) data, we have traced the radio emission to large scales in the complex source 4C 35.06 located in the core of the galaxy cluster Abell 407. At higher spatial resolution (~ 4″), the source was known to have two inner radio lobes spanning 31 kpc and a diffuse, low-brightness extension running parallel to them, offset by about 11 kpc (in projection). At 62 MHz, we detect the radio emission of this structure extending out to 210 kpc. At 1.4 GHz and intermediate spatial resolution (~ 30″), the structure appears to have a helical morphology. We have derived the characteristics of the radio spectral index across the source. We show that the source morphology is most likely the result of at least two episodes of AGN activity separated by a dormant period of around 35 Myr. The outermost regions of radio emission have a steep spectral index (α< − 1), indicative of old plasma. We connect the spectral index properties of the resolved source structure with the integrated fluxdensity spectral index of 4C 35.06 and suggest an explanation for its unusual integrated flux density spectral shape (a moderately steep power law with no discernible spectral break), possibly providing a proxy for future studies of more distant radio sources through inferring their detailed spectral index properties and activity history from their integrated spectral indices. The AGN is hosted by one of the galaxies located in the cluster core of Abell 407. We propose that it is intermittently active as it moves in the dense environment in the cluster core. In this scenario, the AGN turned on sometime in the past, and has produced the helical pattern of emission, possibly a sign of jet precession/merger during that episode of activity. Using LOFAR, we can trace the relic plasma from that episode of activity out to greater distances from the core than ever before. Using the the WSRT, we detect H I in absorption against the center of the radio source. The absorption profile is relatively broad (FWHM of 288 kms-1), similar to what is found in other clusters. The derived column density is NHI ~ 4 × 1020 cm-2 for a Tspin = 100 K. This detection supports the connection – already suggested for other restarted radio sources – between the presence of cold gas and restarting activity. The cold gas appears to be dominated by a blue-shifted component although the broad H I profile could also include gas with different kinematics. Understanding the duty cycle of the radio emission as well as the triggering mechanism for starting (or restarting) the radio-loud activity can provide important constraints to quantify the impact of AGN feedback on galaxy evolution. The study of these mechanisms at low frequencies using morphological and spectral information promises to bring new important insights in this field.
|
|
4. |
- Stewart, A. J., et al.
(author)
-
LOFAR MSSS : detection of a low-frequency radio transient in 400 h of monitoring of the North Celestial Pole
- 2016
-
In: Monthly notices of the Royal Astronomical Society. - : Oxford University Press (OUP). - 0035-8711 .- 1365-2966. ; 456:3, s. 2321-2342
-
Journal article (peer-reviewed)abstract
- We present the results of a four-month campaign searching for low-frequency radio transients near the North Celestial Pole with the Low-Frequency Array (LOFAR), as part of the Multifrequency Snapshot Sky Survey (MSSS). The data were recorded between 2011 December and 2012 April and comprised 2149 11-min snapshots, each covering 175 deg2. We have found one convincing candidate astrophysical transient, with a duration of a few minutes and a flux density at 60 MHz of 15–25 Jy. The transient does not repeat and has no obvious optical or high-energy counterpart, as a result of which its nature is unclear. The detection of this event implies a transient rate at 60 MHz of 3.9−3.7+14.7×10−4" style="position: relative;" tabindex="0" id="MathJax-Element-1-Frame" class="MathJax">3.9+14.7−3.7×10−4 d−1 deg−2, and a transient surface density of 1.5 × 10−5 deg−2, at a 7.9-Jy limiting flux density and ∼10-min time-scale. The campaign data were also searched for transients at a range of other time-scales, from 0.5 to 297 min, which allowed us to place a range of limits on transient rates at 60 MHz as a function of observation duration.
|
|
5. |
- Corstanje, A., et al.
(author)
-
The shape of the radio wavefront of extensive air showers as measured with LOFAR
- 2015
-
In: Astroparticle physics. - : Elsevier BV. - 0927-6505 .- 1873-2852. ; 61, s. 22-31
-
Journal article (peer-reviewed)abstract
- Extensive air showers, induced by high energy cosmic rays impinging on the Earth’s atmosphere, produce radio emission that is measured with the LOFAR radio telescope. As the emission comes from a finite distance of a few kilometers, the incident wavefront is non-planar. A spherical, conical or hyperbolic shape of the wavefront has been proposed, but measurements of individual air showers have been inconclusive so far. For a selected high-quality sample of 161 measured extensive air showers, we have reconstructed the wavefront by measuring pulse arrival times to sub-nanosecond precision in 200 to 350 individual antennas. For each measured air shower, we have fitted a conical, spherical, and hyperboloid shape to the arrival times. The fit quality and a likelihood analysis show that a hyperboloid is the best parameterization. Using a non-planar wavefront shape gives an improved angular resolution, when reconstructing the shower arrival direction. Furthermore, a dependence of the wavefront shape on the shower geometry can be seen. This suggests that it will be possible to use a wavefront shape analysis to get an additional handle on the atmospheric depth of the shower maximum, which is sensitive to the mass of the primary particle.
|
|
6. |
- Jackson, N., et al.
(author)
-
LBCS: The LOFAR Long-Baseline Calibrator Survey
- 2016
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 595, s. Art no A86-
-
Journal article (peer-reviewed)abstract
- We outline the LOFAR Long-Baseline Calibrator Survey (LBCS), whose aim is to identify sources suitable for calibrating the highest-resolution observations made with the International LOFAR Telescope, which include baselines > 1000 km. Suitable sources must contain significant correlated flux density (greater than or similar to 50 - 100 mJy) at frequencies around 110-190 MHz on scales of a few hundred milliarcseconds. At least for the 200-300-km international baselines, we find around 1 suitable calibrator source per square degree over a large part of the northern sky, in agreement with previous work. This should allow a randomly selected target to be successfully phase calibrated on the international baselines in over 50% of cases. Products of the survey include calibrator source lists and fringe-rate and delay maps of wide areas-typically a few degrees-around each source. The density of sources with significant correlated flux declines noticeably with baseline length over the range 200-600 km, with good calibrators on the longest baselines appearing only at the rate of 0.5 per sq. deg. Coherence times decrease from 1-3 min on 200-km baselines to about 1 min on 600-km baselines, suggesting that ionospheric phase variations contain components with scales of a few hundred kilometres. The longest median coherence time, at just over 3 min, is seen on the DE609 baseline, which at 227 km is close to being the shortest. We see median coherence times of between 80 and 110 s on the four longest baselines (580-600 km), and about 2 min for the other baselines. The success of phase transfer from calibrator to target is shown to be influenced by distance, in a manner that suggests a coherence patch at 150-MHz of the order of 1 deg. Although source structures cannot be measured in these observations, we deduce that phase transfer is affected if the calibrator source structure is not known. We give suggestions for calibration strategies and choice of calibrator sources, and describe the access to the online catalogue and data products.
|
|
7. |
- Sobey, C., et al.
(author)
-
LOFAR discovery of a quiet emission mode in PSR B0823+26
- 2015
-
In: Monthly notices of the Royal Astronomical Society. - : Oxford University Press (OUP). - 0035-8711 .- 1365-2966. ; 451, s. 2493-2506
-
Journal article (peer-reviewed)abstract
- PSR B0823+26, a 0.53-s radio pulsar, displays a host of emission phenomena over time-scales of seconds to (at least) hours, including nulling, subpulse drifting, and mode-changing. Studying pulsars like PSR B0823+26 provides further insight into the relationship between these various emission phenomena and what they might teach us about pulsar magnetospheres. Here we report on the LOFAR (Low-Frequency Array) discovery that PSR B0823+26 has a weak and sporadically emitting ‘quiet’ (Q) emission mode that is over 100 times weaker (on average) and has a nulling fraction forty-times greater than that of the more regularly-emitting ‘bright’ (B) mode. Previously, the pulsar has been undetected in the Q mode, and was assumed to be nulling continuously. PSR B0823+26 shows a further decrease in average flux just before the transition into the B mode, and perhaps truly turns off completely at these times. Furthermore, simultaneous observations taken with the LOFAR, Westerbork, Lovell, and Effelsberg telescopes between 110 MHz and 2.7 GHz demonstrate that the transition between the Q mode and B mode occurs within one single rotation of the neutron star, and that it is concurrent across the range of frequencies observed.
|
|
8. |
- Orrù, E., et al.
(author)
-
Wide-field LOFAR imaging of the field around the double-double radio galaxy B1834+620 : A fresh view on a restarted AGN and doubeltjes
- 2015
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 584, s. 1-12
-
Journal article (peer-reviewed)abstract
- Context. The existence of double-double radio galaxies (DDRGs) is evidence for recurrent jet activity in active galactic nuclei (AGN), as expected from standard accretion models. A detailed study of these rare sources provides new perspectives for investigating the AGN duty cycle, AGN-galaxy feedback, and accretion mechanisms. Large catalogues of radio sources, on the other hand, provide statistical information about the evolution of the radio-loud AGN population out to high redshifts.Aims. Using wide-field imaging with the LOFAR telescope, we study both a well-known DDRG as well as a large number of radio sources in the field of view.Methods. We present a high resolution image of the DDRG B1834+620 obtained at 144 MHz using LOFAR commissioning data. Our image covers about 100 square degrees and contains over 1000 sources.Results. The four components of the DDRG B1834+620 have been resolved for the first time at 144 MHz. Inner lobes were found to point towards the direction of the outer lobes, unlike standard FR II sources. Polarized emission was detected at +60 rad m-2 in the northern outer lobe. The high spatial resolution allows the identification of a large number of small double-lobed radio sources; roughly 10% of all sources in the field are doubles with a separation smaller than 1′.Conclusions. The spectral fit of the four components is consistent with a scenario in which the outer lobes are still active or the jets recently switched off, while emission of the inner lobes is the result of a mix-up of new and old jet activity. From the presence of the newly extended features in the inner lobes of the DDRG, we can infer that the mechanism responsible for their formation is the bow shock that is driven by the newly launched jet. We find that the density of the small doubles exceeds the density of FR II sources with similar properties at 1.4 GHz, but this difference becomes smaller for low flux densities. Finally, we show that the significant challenges of wide-field imaging (e.g., time and frequency variation of the beam, directional dependent calibration errors) can be solved using LOFAR commissioning data, thus demonstrating the potential of the full LOFAR telescope to discover millions of powerful AGN at redshift z ~ 1.
|
|
9. |
- Moldón, J., et al.
(author)
-
The LOFAR long baseline snapshot calibrator survey
- 2015
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 574
-
Journal article (peer-reviewed)abstract
- Aims. An efficient means of locating calibrator sources for international LOw Frequency ARray (LOFAR) is developed and used to determine the average density of usable calibrator sources on the sky for subarcsecond observations at 140 MHz.Methods. We used the multi-beaming capability of LOFAR to conduct a fast and computationally inexpensive survey with the full international LOFAR array. Sources were preselected on the basis of 325 MHz arcminute-scale flux density using existing catalogues. By observing 30 different sources in each of the 12 sets of pointings per hour, we were able to inspect 630 sources in two hours to determine if they possess a sufficiently bright compact component to be usable as LOFAR delay calibrators.Results. More than 40% of the observed sources are detected on multiple baselines between international stations and 86 are classified as satisfactory calibrators. We show that a flat low-frequency spectrum (from 74 to 325 MHz) is the best predictor of compactness at 140 MHz. We extrapolate from our sample to show that the sky density of calibrators that are sufficiently bright to calibrate dispersive and non-dispersive delays for the international LOFAR using existing methods is 1.0 per square degree.Conclusions. The observed density of satisfactory delay calibrator sources means that observations with international LOFAR should be possible at virtually any point in the sky provided that a fast and efficient search, using the methodology described here, is conducted prior to the observation to identify the best calibrator.
|
|
10. |
- Pilia, M., et al.
(author)
-
Wide-band, low-frequency pulse profiles of 100 radio pulsars with LOFAR
- 2016
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 586
-
Journal article (peer-reviewed)abstract
- Context. LOFAR offers the unique capability of observing pulsars across the 10−240 MHz frequency range with a fractional bandwidth of roughly 50%. This spectral range is well suited for studying the frequency evolution of pulse profile morphology caused by both intrinsic and extrinsic effects such as changing emission altitude in the pulsar magnetosphere or scatter broadening by the interstellar medium, respectively.Aims. The magnitude of most of these effects increases rapidly towards low frequencies. LOFAR can thus address a number of open questions about the nature of radio pulsar emission and its propagation through the interstellar medium.Methods. We present the average pulse profiles of 100 pulsars observed in the two LOFAR frequency bands: high band (120–167 MHz, 100 profiles) and low band (15–62 MHz, 26 profiles). We compare them with Westerbork Synthesis Radio Telescope (WSRT) and Lovell Telescope observations at higher frequencies (350 and 1400 MHz) to study the profile evolution. The profiles were aligned in absolute phase by folding with a new set of timing solutions from the Lovell Telescope, which we present along with precise dispersion measures obtained with LOFAR.Results. We find that the profile evolution with decreasing radio frequency does not follow a specific trend; depending on the geometry of the pulsar, new components can enter into or be hidden from view. Nonetheless, in general our observations confirm the widening of pulsar profiles at low frequencies, as expected from radius-to-frequency mapping or birefringence theories.
|
|
11. |
- Girard, J. N., et al.
(author)
-
Imaging Jupiter’s radiation belts down to 127 MHz with LOFAR
- 2016
-
In: Astronomy and Astrophysics. - : EDP Sciences. - 0004-6361 .- 1432-0746. ; 587
-
Journal article (peer-reviewed)abstract
- Context. With the limited amount of in situ particle data available for the innermost region of Jupiter’s magnetosphere, Earth-based observations of the giant planets synchrotron emission remain the sole method today of scrutinizing the distribution and dynamical behavior of the ultra energetic electrons magnetically trapped around the planet. Radio observations ultimately provide key information about the origin and control parameters of the harsh radiation environment.Aims. We perform the first resolved and low-frequency imaging of the synchrotron emission with LOFAR. At a frequency as low as 127 MHz, the radiation from electrons with energies of ~1–30 MeV are expected, for the first time, to be measured and mapped over a broad region of Jupiter’s inner magnetosphere.Methods. Measurements consist of interferometric visibilities taken during a single 10-hour rotation of the Jovian system. These visibilities were processed in a custom pipeline developed for planetary observations, combining flagging, calibration, wide-field imaging, direction-dependent calibration, and specific visibility correction for planetary targets. We produced spectral image cubes of Jupiter’s radiation belts at the various angular, temporal, and spectral resolutions from which flux densities were measured.Results. The first resolved images of Jupiter’s radiation belts at 127–172 MHz are obtained with a noise level ~20–25 mJy/beam, along with total integrated flux densities. They are compared with previous observations at higher frequencies. A greater extent of the synchrotron emission source (≥4 RJ) is measured in the LOFAR range, which is the signature – as at higher frequencies – of the superposition of a “pancake” and an isotropic electron distribution. Asymmetry of east-west emission peaks is measured, as well as the longitudinal dependence of the radial distance of the belts, and the presence of a hot spot at λIII = 230° ± 25°. Spectral flux density measurements are on the low side of previous (unresolved) ones, suggesting a low-frequency turnover and/or time variations of the Jovian synchrotron spectrum.Conclusions. LOFAR proves to be a powerful and flexible planetary imager. In the case of Jupiter, observations at 127 MHz depict the distribution of ~1–30 MeV energy electrons up to ~4–5 planetary radii. The similarities of the observations at 127 MHz with those at higher frequencies reinforce the conclusion that the magnetic field morphology primarily shapes the brightness distribution features of Jupiter’s synchrotron emission, as well as how the radiating electrons are likely radially and latitudinally distributed inside about 2 planetary radii. Nonetheless, the detection of an emission region that extends to larger distances than at higher frequencies, combined with the overall lower flux density, yields new information on Jupiter’s electron distribution, and this information may ultimately shed light on the origin and mode of transport of these particles.
|
|
12. |
- Nelles, A., et al.
(author)
-
Calibrating the absolute amplitude scale for air showers measured at LOFAR
- 2015
-
In: Journal of Instrumentation. - 1748-0221. ; 10
-
Journal article (peer-reviewed)abstract
- Air showers induced by cosmic rays create nanosecond pulses detectable at radio frequencies. These pulses have been measured successfully in the past few years at the LOw-Frequency ARray (LOFAR) and are used to study the properties of cosmic rays. For a complete understanding of this phenomenon and the underlying physical processes, an absolute calibration of the detecting antenna system is needed. We present three approaches that were used to check and improve the antenna model of LOFAR and to provide an absolute calibration of the whole system for air shower measurements. Two methods are based on calibrated reference sources and one on a calibration approach using the diffuse radio emission of the Galaxy, optimized for short data-sets. An accuracy of 19% in amplitude is reached. The absolute calibration is also compared to predictions from air shower simulations. These results are used to set an absolute energy scale for air shower measurements and can be used as a basis for an absolute scale for the measurement of astronomical transients with LOFAR.
|
|
13. |
- Nelles, A., et al.
(author)
-
Measuring a Cherenkov ring in the radio emission from air showers at 110-190 MHz with LOFAR
- 2015
-
In: Astroparticle physics. - : Elsevier BV. - 0927-6505 .- 1873-2852. ; 65, s. 11-21
-
Journal article (peer-reviewed)abstract
- Measuring radio emission from air showers offers a novel way to determine properties of the primary cosmic rays such as their mass and energy. Theory predicts that relativistic time compression effects lead to a ring of amplified emission which starts to dominate the emission pattern for frequencies above ∼100∼100 MHz. In this article we present the first detailed measurements of this structure. Ring structures in the radio emission of air showers are measured with the LOFAR radio telescope in the frequency range of 110–190 MHz. These data are well described by CoREAS simulations. They clearly confirm the importance of including the index of refraction of air as a function of height. Furthermore, the presence of the Cherenkov ring offers the possibility for a geometrical measurement of the depth of shower maximum, which in turn depends on the mass of the primary particle.
|
|
14. |
- de Vries, Paul S., et al.
(author)
-
Comparison of HapMap and 1000 Genomes Reference Panels in a Large-Scale Genome-Wide Association Study
- 2017
-
In: PLOS ONE. - : Public Library of Science (PLoS). - 1932-6203. ; 12:1
-
Journal article (peer-reviewed)abstract
- An increasing number of genome-wide association (GWA) studies are now using the higher resolution 1000 Genomes Project reference panel (1000G) for imputation, with the expectation that 1000G imputation will lead to the discovery of additional associated loci when compared to HapMap imputation. In order to assess the improvement of 1000G over HapMap imputation in identifying associated loci, we compared the results of GWA studies of circulating fibrinogen based on the two reference panels. Using both HapMap and 1000G imputation we performed a meta-analysis of 22 studies comprising the same 91,953 individuals. We identified six additional signals using 1000G imputation, while 29 loci were associated using both HapMap and 1000G imputation. One locus identified using HapMap imputation was not significant using 1000G imputation. The genome-wide significance threshold of 5x10(-8) is based on the number of independent statistical tests using HapMap imputation, and 1000G imputation may lead to further independent tests that should be corrected for. When using a stricter Bonferroni correction for the 1000G GWA study (P-value < 2.5x10(-8)), the number of loci significant only using HapMap imputation increased to 4 while the number of loci significant only using 1000G decreased to 5. In conclusion, 1000G imputation enabled the identification of 20% more loci than HapMap imputation, although the advantage of 1000G imputation became less clear when a stricter Bonferroni correction was used. More generally, our results provide insights that are applicable to the implementation of other dense reference panels that are under development.
|
|
15. |
|
|
16. |
- Huffman, JE, et al.
(author)
-
Rare and low-frequency variants and their association with plasma levels of fibrinogen, FVII, FVIII, and vWF
- 2015
-
In: Blood. - : American Society of Hematology. - 1528-0020 .- 0006-4971. ; 126:11, s. E19-E29
-
Journal article (peer-reviewed)abstract
- Twelve independent, novel, low-frequency (n = 2) and rare (n = 10) genetic variants were associated with fibrinogen, FVII, FVIII, or vWF. Nine were within previously associated genes, and 3 novel candidate genes (KCNT1, HID1, and KATNB1) were confined to cohorts of African ancestry.
|
|
17. |
|
|
18. |
|
|
19. |
|
|
20. |
|
|
21. |
|
|
22. |
- van Hensbergen, Vincent P., et al.
(author)
-
Streptococcal Lancefield polysaccharides are critical cell wall determinants for human Group IIA secreted phospholipase A2 to exert its bactericidal effects
- 2018
-
In: PLoS Pathogens. - : Public Library of Science (PLoS). - 1553-7374. ; 14:10, s. 1007348-1007348
-
Journal article (peer-reviewed)abstract
- Human Group IIA secreted phospholipase A2 (hGIIA) is an acute phase protein with bactericidal activity against Gram-positive bacteria. Infection models in hGIIA transgenic mice have suggested the importance of hGIIA as an innate defense mechanism against the human pathogens Group A Streptococcus (GAS) and Group B Streptococcus (GBS). Compared to other Gram-positive bacteria, GAS is remarkably resistant to hGIIA activity. To identify GAS resistance mechanisms, we exposed a highly saturated GAS M1 transposon library to recombinant hGIIA and compared relative mutant abundance with library input through transposon-sequencing (Tn-seq). Based on transposon prevalence in the output library, we identified nine genes, including dltA and lytR, conferring increased hGIIA susceptibility. In addition, seven genes conferred increased hGIIA resistance, which included two genes, gacH and gacI that are located within the Group A Carbohydrate (GAC) gene cluster. Using GAS 5448 wild-type and the isogenic gacI mutant and gacI-complemented strains, we demonstrate that loss of the GAC N-acetylglucosamine (GlcNAc) side chain in the ΔgacI mutant increases hGIIA resistance approximately 10-fold, a phenotype that is conserved across different GAS serotypes. Increased resistance is associated with delayed penetration of hGIIA through the cell wall. Correspondingly, loss of the Lancefield Group B Carbohydrate (GBC) rendered GBS significantly more resistant to hGIIA-mediated killing. This suggests that the streptococcal Lancefield antigens, which are critical determinants for streptococcal physiology and virulence, are required for the bactericidal enzyme hGIIA to exert its bactericidal function.
|
|