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In the new era of time-domain surveys, Type Ia supernovae are being caught sooner after explosion, which has exposed significant variation in their early light curves. Two driving factors for early-time evolution are the distribution of 56Ni in the ejecta and the presence of flux excesses of various causes. We perform an analysis of the largest young SN Ia sample to date. We compare 115 SN Ia light curves from the Zwicky Transient Facility to the turtls model grid containing light curves of Chandrasekhar mass explosions with a range of 56Ni masses, 56Ni distributions, and explosion energies. We find that the majority of our observed light curves are well reproduced by Chandrasekhar mass explosion models with a preference for highly extended 56Ni distributions. We identify six SNe Ia with an early-time flux excess in our gr -band data (four 'blue' and two 'red' flux excesses). We find an intrinsic rate of 18 ± 11 per cent of early flux excesses in SNe Ia at z < 0.07, based on three detected flux excesses out of 30 (10 per cent) observed SNe Ia with a simulated efficiency of 57 per cent. This is comparable to rates of flux excesses in the literature but also accounts for detection efficiencies. Two of these events are mostly consistent with circumstellar material interaction, while the other four have longer lifetimes in agreement with companion interaction and 56Ni-clump models. We find a higher frequency of flux excesses in 91T/99aa-like events (44 ± 13 per cent). [ABSTRACT FROM AUTHOR]

Type Ia supernovae (SNe Ia) in the nearby Hubble flow are excellent distance indicators in cosmology. The Zwicky Transient Facility (ZTF) has observed a large sample of SNe from an untargeted, rolling survey, reaching 20.8, 20.6, and 20.3 mag in g r , and i band, respectively. With an FoV of 47 deg2, ZTF discovered > 3000 SNe Ia in a little over 2.5 yr. Here, we report on the sample of 761 spectroscopically classified SNe Ia from the first year of operations (DR1). The sample has a median redshift |$\bar{z} =$| 0.057, nearly a factor of 2 higher than the current low- z sample. Our sample has a total of 934 spectra, of which 632 were obtained with the robotic SEDm on Palomar P60. We assess the potential for precision cosmology for a total of 305 SNe with redshifts from host galaxy spectra. The sample is already comparable in size to the entire combined literature low- z anchor sample. The median first detection is 13.5 d before maximum light, about 10 d earlier than the median in the literature. Furthermore, six SNe from our sample are at D L < 80 Mpc, for which host galaxy distances can be obtained in the JAMES WEBB SPACE TELESCOPE era, such that we have calibrator and Hubble flow SNe observed with the same instrument. In the entire duration of ZTF-I, we have observed nearly 50 SNe for which we can obtain calibrator distances, key for per cent level distance scale measurements. [ABSTRACT FROM AUTHOR]

As part of an on-going effort to identify, understand and correct for astrophysics biases in the standardization of Type Ia supernovae (SN Ia) for cosmology, we have statistically classified a large sample of nearby SNe Ia into those that are located in predominantly younger or older environments. This classification is based on the specific star formation rate measured within a projected distance of 1 kpc from each SN location (LsSFR). This is an important refinement compared to using the local star formation rate directly, as it provides a normalization for relative numbers of available SN progenitors and is more robust against extinction by dust. We find that the SNe Ia in predominantly younger environments are ΔY = 0.163 ± 0.029 mag (5.7σ) fainter than those in predominantly older environments after conventional light-curve standardization. This is the strongest standardized SN Ia brightness systematic connected to the host-galaxy environment measured to date. The well-established step in standardized brightnesses between SNe Ia in hosts with lower or higher total stellar masses is smaller, at ΔM = 0.119 ± 0.032 mag (4.5σ), for the same set of SNe Ia. When fit simultaneously, the environment-age offset remains very significant, with ΔY = 0.129 ± 0.032 mag (4.0σ), while the global stellar mass step is reduced to ΔM = 0.064  ±  0.029 mag (2.2σ). Thus, approximately 70% of the variance from the stellar mass step is due to an underlying dependence on environment-based progenitor age. Also, we verify that using the local star formation rate alone is not as powerful as LsSFR at sorting SNe Ia into brighter and fainter subsets. Standardization that only uses the SNe Ia in younger environments reduces the total dispersion from 0.142  ±  0.008 mag to 0.120  ±  0.010 mag. We show that as environment-ages evolve with redshift, a strong bias, especially on the measurement of the derivative of the dark energy equation of state, can develop. Fortunately, data that measure and correct for this effect using our local specific star formation rate indicator, are likely to be available for many next-generation SN Ia cosmology experiments. [ABSTRACT FROM AUTHOR]

The Zwicky Transient Facility (ZTF) is performing a three-day cadence survey of the visible northern sky (∼3π) with newly found transient candidates announced via public alerts. The ZTF Bright Transient Survey (BTS) is a large spectroscopic campaign to complement the photometric survey. BTS endeavors to spectroscopically classify all extragalactic transients with mpeak ≤ 18.5 mag in either the gZTF or rZTF filters, and publicly announce said classifications. BTS discoveries are predominantly supernovae (SNe), making this the largest flux-limited SN survey to date. Here we present a catalog of 761 SNe, classified during the first nine months of ZTF (2018 April 1–2018 December 31). We report BTS SN redshifts from SN template matching and spectroscopic host-galaxy redshifts when available. We analyze the redshift completeness of local galaxy catalogs, the redshift completeness fraction (RCF; the ratio of SN host galaxies with known spectroscopic redshift prior to SN discovery to the total number of SN hosts). Of the 512 host galaxies with SNe Ia, 227 had previously known spectroscopic redshifts, yielding an RCF estimate of 44% ± 4%. The RCF decreases with increasing distance and decreasing galaxy luminosity (for z < 0.05, or ∼200 Mpc, RCF ≈ 0.6). Prospects for dramatically increasing the RCF are limited to new multifiber spectroscopic instruments or wide-field narrowband surveys. Existing galaxy redshift catalogs are only ∼50% complete at r ≈ 16.9 mag. Pushing this limit several magnitudes deeper will pay huge dividends when searching for electromagnetic counterparts to gravitational wave events or sources of ultra-high-energy cosmic rays or neutrinos. [ABSTRACT FROM AUTHOR]

Context. Both multi-messenger astronomy and new high-throughput wide-field surveys require flexible tools for the selection and analysis of astrophysical transients. Aims. Here we introduce the alert management, photometry, and evaluation of light curves (AMPEL) system, an analysis framework designed for high-throughput surveys and suited for streamed data. AMPEL combines the functionality of an alert broker with a generic framework capable of hosting user-contributed code; it encourages provenance and keeps track of the varying information states that a transient displays. The latter concept includes information gathered over time and data policies such as access or calibration levels. Methods. We describe a novel ongoing real-time multi-messenger analysis using AMPEL to combine IceCube neutrino data with the alert streams of the Zwicky Transient Facility (ZTF). We also reprocess the first four months of ZTF public alerts, and compare the yields of more than 200 different transient selection functions to quantify efficiencies for selecting Type Ia supernovae that were reported to the Transient Name Server (TNS). Results. We highlight three channels suitable for (1) the collection of a complete sample of extragalactic transients, (2) immediate follow-up of nearby transients, and (3) follow-up campaigns targeting young, extragalactic transients. We confirm ZTF completeness in that all TNS supernovae positioned on active CCD regions were detected. Conclusions.AMPEL can assist in filtering transients in real time, running alert reaction simulations, the reprocessing of full datasets as well as in the final scientific analysis of transient data. This is made possible by a novel way of capturing transient information through sequences of evolving states, and interfaces that allow new code to be natively applied to a full stream of alerts. This text also introduces a method by which users can design their own channels for inclusion in the AMPEL live instance that parses the ZTF stream and the real-time submission of high-quality extragalactic supernova candidates to the TNS. [ABSTRACT FROM AUTHOR]

Current time domain facilities are discovering hundreds of new galactic and extra-galactic transients every week. Classifying the ever-increasing number of transients is challenging, yet crucial to furthering our understanding of their nature, discovering new classes, and ensuring sample purity, for instance, for Supernova Ia cosmology. The Zwicky Transient Facility is one example of such a survey. In addition, it has a dedicated very-low resolution spectrograph, the SEDMachine, operating on the Palomar 60-inch telescope. This spectrograph's primary aim is object classification. In practice most, if not all, transients of interest brighter than ∼19 mag are typed. This corresponds to approximately 10–15 targets a night. In this paper, we present a fully automated pipeline for the SEDMachine. This pipeline has been designed to be fast, robust, stable and extremely flexible. pysedm enables the fully automated spectral extraction of a targeted point source object in less than five minutes after the end of the exposure. The spectral color calibration is accurate at the few percent level. In the 19 weeks since pysedm entered production in early August of 2018, we have classified, among other objects, about 400 Type Ia supernovae and 140 Type II supernovae. We conclude that low resolution, fully automated spectrographs such as the "SEDMachine with pysedm" installed on 2-m class telescopes within the southern hemisphere could allow us to automatically and simultaneously type and obtain a redshift for most (if not all) bright transients detected by LSST within z <  0.2, notably potentially all Type Ia Supernovae. In comparison with the current SEDM design, this would require higher spectral resolution (R ≳ 1000) and slightly improved throughput. With this perspective in mind, pysedm is designed to easily be adaptable to any IFU-like spectrograph. [ABSTRACT FROM AUTHOR]

Context. Use of Type Ia supernovae (SNe Ia) as distance indicators has proven to be a powerful technique for measuring the darkenergy equation of state. However, recent studies have highlighted potential biases correlated with the global properties of their host galaxies, large enough to induce systematic errors into such cosmological measurements if not properly treated. Aims. We study the host galaxy regions in close proximity to SNe Ia in order to analyze relations between the properties of SN Ia events and environments where their progenitors most likely formed. In this paper we focus on local H± emission as an indicator of young progenitor environments. Methods. The Nearby Supernova Factory has obtained flux-calibrated spectral timeseries for SNe Ia using integral field spectroscopy. These observations enabled the simultaneous measurement of the SN and its immediate vicinity. For 89 SNe Ia we measured or set limits on H± emission, used as a tracer of ongoing star formation, within a 1 kpc radius around each SN. This constitutes the first direct study of the local environment for a large sample of SNe Ia with accurate luminosity, color, and stretch measurements. Results. Our local star formation measurements provide several critical new insights. We find that SNe Ia with local H± emission are redder by 0:036 ± 0:017 mag, and that the previously noted correlation between stretch and host mass is driven entirely by the SNe Ia coming from locally passive environments, in particular at the low-stretch end. There is no such trend for SNe Ia in locally star-forming environments. Our most important finding is that the mean standardized brightness for SNe Ia with local H± emission is 0:094 ± 0:031 mag fainter on average than for those without. This o±set arises from a bimodal structure in the Hubble residuals, with one mode being shared by SNe Ia in all environments and the other one exclusive to SNe Ia in locally passive environments. This structure also explains the previously known host-mass bias. We combine the star formation dependence of this bimodality with the cosmic star formation rate to predict changes with redshift in the mean SN Ia brightness and the host-mass bias. The strong change predicted is confirmed using high-redshift SNe Ia from the literature. Conclusions. The environmental dependences in SN Ia Hubble residuals and color found here point to remaining systematic errors in the standardization of SNe Ia. In particular, the observed brightness o±set associated with local H± emission is predicted to cause a significant bias in current measurements of the dark energy equation of state. Recognition of these e±ects o±ers new opportunities to improve SNe Ia as cosmological probes. For instance, we note that the SNe Ia associated with local H± emission are more homogeneous, resulting in a brightness dispersion of only 0:105 ± 0:012 mag. [ABSTRACT FROM AUTHOR]

Previously we used the Nearby Supernova Factory sample to show that Type Ia supernovae (SNe Ia) having locally star-forming environments are dimmer than SNe Ia having locally passive environments. Here we use the Constitution sample together with host galaxy data from GALEX to independently confirm that result. The effect is seen using both the SALT2 and MLCS2k2 lightcurve fitting and standardization methods, with brightness differences of 0.094 ± 0.037 mag for SALT2 and 0.155 ± 0.041 mag for MLCS2k2 with RV = 2.5. When combined with our previous measurement the effect is 0.094 ± 0.025 mag for SALT2. If the ratio of these local SN Ia environments changes with redshift or sample selection, this can lead to a bias in cosmological measurements. We explore this issue further, using as an example the direct measurement of H0. GALEX observations show that the SNe Ia having standardized absolute magnitudes calibrated via the Cepheid period-luminosity relation using the Hubble Space Telescope originate in predominately star-forming environments, whereas only ∼50% of the Hubble-flow comparison sample have locally star-forming environments. As a consequence, the H0 measurement using SNe Ia is currently overestimated. Correcting for this bias, we find a value of 70.6 ± 2.6 km s–1 Mpc–1 when using the LMC distance, Milky Way parallaxes, and the NGC 4258 megamaser as the Cepheid zero point, and 68.8 ± 3.3 km s–1 Mpc–1 when only using NGC 4258. Our correction brings the direct measurement of H0 within ∼1 σ of recent indirect measurements based on the cosmic microwave background power spectrum. [ABSTRACT FROM AUTHOR]

Climate change has adversely affected maize productivity. Thereby, a holistic understanding of metabolic crosstalk among its organs is important to address this issue. Thus, we reconstructed the first multi-organ maize metabolic model, i ZMA6517, and contextualized it with heat and cold stress transcriptomics data using expression distributed reaction flux measurement (EXTREAM) algorithm. Furthermore, implementing metabolic bottleneck analysis on contextualized models revealed differences between these stresses. While both stresses had reducing power bottlenecks, heat stress had additional energy generation bottlenecks. We also performed thermodynamic driving force analysis, revealing thermodynamics-reducing power-energy generation axis dictating the nature of temperature stress responses. Thus, a temperature-tolerant maize ideotype can be engineered by leveraging the proposed thermodynamics-reducing power-energy generation axis. We experimentally inoculated maize root with a beneficial mycorrhizal fungus, Rhizophagus irregularis , and as a proof-of-concept demonstrated its efficacy in alleviating temperature stress. Overall, this study will guide the engineering effort of temperature stress-tolerant maize ideotypes., Competing Interests: The authors declare no competing interests., (© 2023 The Authors.)

Detecting gravitationally lensed supernovae is among the biggest challenges in astronomy. It involves a combination of two very rare phenomena: catching the transient signal of a stellar explosion in a distant galaxy and observing it through a nearly perfectly aligned foreground galaxy that deflects light towards the observer. Here we describe how high-cadence optical observations with the Zwicky Transient Facility, with its unparalleled large field of view, led to the detection of a multiply imaged type Ia supernova, SN Zwicky, also known as SN 2022qmx. Magnified nearly 25-fold, the system was found thanks to the standard candle nature of type Ia supernovae. High-spatial-resolution imaging with the Keck telescope resolved four images of the supernova with very small angular separation, corresponding to an Einstein radius of only θ E  = 0.167″ and almost identical arrival times. The small θ E and faintness of the lensing galaxy are very unusual, highlighting the importance of supernovae to fully characterize the properties of galaxy-scale gravitational lenses, including the impact of galaxy substructures., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2023, corrected publication 2023.)

Tidal disruption events (TDEs) are bursts of electromagnetic energy that are released when supermassive black holes at the centres of galaxies violently disrupt a star that passes too close 1 . TDEs provide a window through which to study accretion onto supermassive black holes; in some rare cases, this accretion leads to launching of a relativistic jet 2-9 , but the necessary conditions are not fully understood. The best-studied jetted TDE so far is Swift J1644+57, which was discovered in γ-rays, but was too obscured by dust to be seen at optical wavelengths. Here we report the optical detection of AT2022cmc, a rapidly fading source at cosmological distance (redshift z = 1.19325) the unique light curve of which transitioned into a luminous plateau within days. Observations of a bright counterpart at other wavelengths, including X-ray, submillimetre and radio, supports the interpretation of AT2022cmc as a jetted TDE containing a synchrotron 'afterglow', probably launched by a supermassive black hole with spin greater than approximately 0.3. Using four years of Zwicky Transient Facility 10 survey data, we calculate a rate of [Formula: see text] per gigapascals cubed per year for on-axis jetted TDEs on the basis of the luminous, fast-fading red component, thus providing a measurement complementary to the rates derived from X-ray and radio observations 11 . Correcting for the beaming angle effects, this rate confirms that approximately 1 per cent of TDEs have relativistic jets. Optical surveys can use AT2022cmc as a prototype to unveil a population of jetted TDEs., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

The origins of the high-energy cosmic neutrino flux remain largely unknown. Recently, one high-energy neutrino was associated with a tidal disruption event (TDE). Here we present AT2019fdr, an exceptionally luminous TDE candidate, coincident with another high-energy neutrino. Our observations, including a bright dust echo and soft late-time x-ray emission, further support a TDE origin of this flare. The probability of finding two such bright events by chance is just 0.034%. We evaluate several models for neutrino production and show that AT2019fdr is capable of producing the observed high-energy neutrino, reinforcing the case for TDEs as neutrino sources.

Owing to the large genetic diversity of barley and its resilience under harsh environments, this crop is of great value for agroecological transition and the need for reduction of nitrogen (N) fertilizers inputs. In the present work, we investigated the diversity of a North African barley genotype collection in terms of growth under limiting N (LN) or ample N (HN) supply and in terms of physiological traits including amino acid content in young seedlings. We identified a Moroccan variety, Laanaceur, accumulating five times more lysine in its leaves than the others under both N nutritional regimes. Physiological characterization of the barley collection showed the genetic diversity of barley adaptation strategies to LN and highlighted a genotype x environment interaction. In all genotypes, N limitation resulted in global biomass reduction, an increase in C concentration, and a higher resource allocation to the roots, indicating that this organ undergoes important adaptive metabolic activity. The most important diversity concerned leaf nitrogen use efficiency (LNUE), root nitrogen use efficiency (RNUE), root nitrogen uptake efficiency (RNUpE), and leaf nitrogen uptake efficiency (LNUpE). Using LNUE as a target trait reflecting barley capacity to deal with N limitation, this trait was positively correlated with plant nitrogen uptake efficiency (PNUpE) and RNUpE. Based on the LNUE trait, we determined three classes showing high, moderate, or low tolerance to N limitation. The transcriptomic approach showed that signaling, ionic transport, immunity, and stress response were the major functions affected by N supply. A candidate gene encoding the HvNRT2.10 transporter was commonly up-regulated under LN in the three barley genotypes investigated. Genes encoding key enzymes required for lysine biosynthesis in plants, dihydrodipicolinate synthase (DHPS) and the catabolic enzyme, the bifunctional Lys-ketoglutarate reductase/saccharopine dehydrogenase are up-regulated in Laanaceur and likely account for a hyperaccumulation of lysine in this genotype. Our work provides key physiological markers of North African barley response to low N availability in the early developmental stages., Competing Interests: MA was employed by company TIMAC AGRO International SAS. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Decouard, Bailly, Rigault, Marmagne, Arkoun, Soulay, Caïus, Paysant-Le Roux, Louahlia, Jacquard, Esmaeel, Chardon, Masclaux-Daubresse and Dellagi.)

We showed previously that nitrogen (N) limitation decreases Arabidopsis resistance to Erwinia amylovora (Ea). We show that decreased resistance to bacteria in low N is correlated with lower apoplastic reactive oxygen species (ROS) accumulation and lower jasmonic acid (JA) pathway expression. Consistently, pretreatment with methyl jasmonate (Me-JA) increased the resistance of plants grown under low N. In parallel, we show that in planta titres of a nonvirulent type III secretion system (T3SS)-deficient Ea mutant were lower than those of wildtype Ea in low N, as expected, but surprisingly not in high N. This lack of difference in high N was consistent with the low expression of the T3SS-encoding hrp virulence genes by wildtype Ea in plants grown in high N compared to plants grown in low N. This suggests that expressing its virulence factors in planta could be a major limiting factor for Ea in the nonhost Arabidopsis. To test this hypothesis, we preincubated Ea in an inducing medium that triggers expression of hrp genes in vitro, prior to inoculation. This preincubation strongly enhanced Ea titres in planta, independently of the plant N status, and was correlated to a significant repression of JA-dependent genes. Finally, we identify two clusters of metabolites associated with resistance or with susceptibility to Ea. Altogether, our data showed that high susceptibility of Arabidopsis to Ea, under low N or following preincubation in hrp-inducing medium, is correlated with high expression of the Ea hrp genes in planta and low expression of the JA signalling pathway, and is correlated with the accumulation of specific metabolites., (© 2021 The Authors. Molecular Plant Pathology published by British Society for Plant Pathology and John Wiley & Sons Ltd.)

Autophagy is a ubiquitous vesicular process for protein and organelle recycling in eukaryotes. In plant, autophagy is reported to play pivotal roles in nutrient recycling, adaptation to biotic and abiotic stresses. The role of autophagy in plant immunity remains poorly understood. Several reports showed enhanced susceptibility of different Arabidopsis autophagy mutants (atg) to necrotrophic fungal pathogens. Interaction of necrotrophic bacterial pathogens with autophagy is overlooked. We then investigated such interaction by inoculating the necrotrophic enterobacterium Dickeya dadantii in leaves of the atg2 and atg5 mutants and an ATG8a overexpressing line. Overexpressing ATG8a enhances plant tolerance to D. dadantii. While atg5 mutant displayed similar susceptibility to the WT, the atg2 mutant exhibited accelerated leaf senescence and enhanced susceptibility upon infection. Both phenotypes were reversed when the sid2 mutation, abolishing SA signaling, was introduced in the atg2 mutant. High levels of SA signaling in atg2 mutant resulted in repression of the jasmonic acid (JA) defense pathway known to limit D. dadantii progression in A. thaliana. We provide evidence that in atg2 mutant, the disturbed hormonal balance leading to higher SA signaling is the main factor causing increased susceptibility to the D. dadantii necrotroph by repressing the JA pathway and accelerating developmental senescence.

Plant defense stimulators, used in crop protection, are an attractive option to reduce the use of conventional crop protection products and optimize biocontrol strategies. These products are able to activate plant defenses and thus limit infection by pathogens. However, the effectiveness of these plant defense stimulators remains erratic and is potentially dependent on many agronomic and environmental parameters still unknown or poorly controlled. The developmental stage of the plant as well as its fertilization, and essentially nitrogen nutrition, play major roles in defense establishment in the presence of pathogens or plant defense stimulators. The major nitrogen source used by plants is nitrate. In this study, we investigated the impact of Arabidopsis thaliana plant developmental stage and nitrate nutrition on its capacity to mount immune reactions in response to two plant defense stimulators triggering two major defense pathways, the salicylic acid and the jasmonic acid pathways. We show that optimal nitrate nutrition is needed for effective defense activation and protection against the pathogenic bacteria Dickeya dadantii and Pseudomonas syringae pv. tomato . Using an npr1 defense signaling mutant, we showed that nitrate dependent protection against D. dadantii requires a functional NPR1 gene. Our results indicate that the efficacy of plant defense stimulators is strongly affected by nitrate nutrition and the developmental stage. The nitrate dependent efficacy of plant defense stimulators is not only due to a metabolic effect but also invloves NPR1 mediated defense signaling. Plant defense stimulators may have opposite effects on plant resistance to a pathogen. Together, our results indicate that agronomic use of plant defense stimulators must be optimized according to nitrate fertilization and developmental stage., (Copyright © 2020 Verly, Djoman, Rigault, Giraud, Rajjou, Saint-Macary and Dellagi.)

Few secreted proteins involved in plant infection common to necrotrophic bacteria, fungi and oomycetes have been identified except for plant cell wall-degrading enzymes. Here we study a family of iron-binding proteins that is present in Gram-negative and Gram-positive bacteria, fungi, oomycetes and some animals. Homolog proteins in the phytopathogenic bacterium Dickeya dadantii (IbpS) and the fungal necrotroph Botrytis cinerea (BcIbp) are involved in plant infection. IbpS is secreted, can bind iron and copper, and protects the bacteria against H 2 O 2 -induced death. Its 1.7 Å crystal structure reveals a classical Venus Fly trap fold that forms dimers in solution and in the crystal. We propose that secreted Ibp proteins binds exogenous metals and thus limit intracellular metal accumulation and ROS formation in the microorganisms.

In their natural environment, plants are generally confronted with multiple co-occurring stresses. However, the interaction between stresses is not well known and transcriptomic data in response to combined stresses remain scarce. This study aims at characterizing the interaction between transcriptomic responses to biotic stress and nitrogen (N) limitation. Plants were grown in low or full N, infected or not with Erwinia amylovora ( Ea ) and plant gene expression was analyzed through microarray and qRT-PCR. Most Ea -responsive genes had the same profile (induced/repressed) in response to Ea in low and full N. In response to stress combination, one third of modulated transcripts responded in a manner that could not be deduced from their response to each individual stress. Many defense-related genes showed a prioritization of their response to biotic stress over their response to N limitation, which was also observed using Pseudomonas syringae as a second pathosystem. Our results indicate an interaction between transcriptomic responses to N and biotic stress. A small fraction of transcripts was prioritized between antagonistic responses, reflecting a preservation of the plant defense program under N limitation. Furthermore, this interaction also led to a complex and specific response in terms of metabolism and cellular homeostasis-associated genes.

Among the most devastating bacterial diseases of plants, soft rot provoked by Dickeya spp. cause crop yield losses on a large range of species with potato being the most economically important. The use of antibiotics being prohibited in most countries in the field, identifying tolerance genes is expected to be one of the most effective alternate disease control approaches. A prerequisite for the identification of tolerance genes is to develop robust disease quantification methods and to identify tolerant plant genotypes. In this work, we investigate the feasibility of the exploitation of Arabidopsis thaliana natural variation to find tolerant genotypes and to develop robust quantification methods. We compared different quantification methods that score either symptom development or bacterial populations in planta . An easy to set up and reliable bacterial quantification method based on qPCR amplification of bacterial DNA was validated. This study demonstrates that it is possible to conduct a robust phenotyping of soft rot disease, and that Arabidopsis natural accessions are a relevant source of tolerance genes.

Siderophores are specific ferric iron chelators synthesized by virtually all microorganisms in response to iron deficiency. We have previously shown that they promote infection by the phytopathogenic enterobacteria Dickeya dadantii and Erwinia amylovora. Siderophores also have the ability to activate plant immunity. We have used complete Arabidopsis transcriptome microarrays to investigate the global transcriptional modifications in roots and leaves of Arabidopsis (Arabidopsis thaliana) plants after leaf treatment with the siderophore deferrioxamine (DFO). Physiological relevance of these transcriptional modifications was validated experimentally. Immunity and heavy-metal homeostasis were the major processes affected by DFO. These two physiological responses could be activated by a synthetic iron chelator ethylenediamine-di(o-hydroxyphenylacetic) acid, indicating that siderophores eliciting activities rely on their strong iron-chelating capacity. DFO was able to protect Arabidopsis against the pathogenic bacterium Pseudomonas syringae pv tomato DC3000. Siderophore treatment caused local modifications of iron distribution in leaf cells visible by ferrocyanide and diaminobenzidine-H₂O₂ staining. Metal quantifications showed that DFO causes a transient iron and zinc uptake at the root level, which is presumably mediated by the metal transporter iron regulated transporter1 (IRT1). Defense gene expression and callose deposition in response to DFO were compromised in an irt1 mutant. Consistently, plant susceptibility to D. dadantii was increased in the irt1 mutant. Our work shows that iron scavenging is a unique mechanism of immunity activation in plants. It highlights the strong relationship between heavy-metal homeostasis and immunity.

Iron is an essential element for most living organisms, and pathogens are likely to compete with their hosts for the acquisition of this element. The bacterial plant pathogen Dickeya dadantii has been shown to require its siderophore-mediated iron uptake system for systemic disease progression on several host plants, including Arabidopsis thaliana. In this study, we investigated the effect of the iron status of Arabidopsis on the severity of disease caused by D. dadantii. We showed that symptom severity, bacterial fitness and the expression of bacterial pectate lyase-encoding genes were reduced in iron-deficient plants. Reduced symptoms correlated with enhanced expression of the salicylic acid defence plant marker gene PR1. However, levels of the ferritin coding transcript AtFER1, callose deposition and production of reactive oxygen species were reduced in iron-deficient infected plants, ruling out the involvement of these defences in the limitation of disease caused by D. dadantii. Disease reduction in iron-starved plants was also observed with the necrotrophic fungus Botrytis cinerea. Our data demonstrate that the plant nutritional iron status can control the outcome of an infection by acting on both the pathogen's virulence and the host's defence. In addition, iron nutrition strongly affects the disease caused by two soft rot-causing plant pathogens with a large host range. Thus, it may be of interest to take into account the plant iron status when there is a need to control disease without compromising crop quality and yield in economically important plant species., (© 2012 THE AUTHORS. MOLECULAR PLANT PATHOLOGY © 2012 BSPP AND BLACKWELL PUBLISHING LTD.)

Siderophores (ferric ion chelators) are secreted by organisms in response to iron deficiency. The pathogenic enterobacterium Erwinia chrysanthemi produces two siderophores, achromobactin and chrysobactin (CB), which are required for systemic dissemination in host plants. Previous studies have shown that CB is produced in planta and can trigger the up-regulation of the plant ferritin gene AtFER1. To further investigate the function of CB during pathogenesis, we analyzed its effect in Arabidopsis (Arabidopsis thaliana) plants following leaf infiltration. CB activates the salicylic acid (SA)-mediated signaling pathway, while the CB ferric complex is ineffective, suggesting that the elicitor activity of this siderophore is due to its iron-binding property. We confirmed this hypothesis by testing the effect of siderophores structurally unrelated to CB, including deferrioxamine. There was no activation of SA-dependent defense in plants grown under iron deficiency before CB treatment. Transcriptional analysis of the genes encoding the root ferrous ion transporter and ferric chelate reductase, and determination of the activity of this enzyme in response to CB or deferrioxamine, showed that these compounds induce a leaf-to-root iron deficiency signal. This root response as well as ferritin gene up-regulation in the leaf were not compromised in a SA-deficient mutant line. Using the Arabidopsis-E. chrysanthemi pathosystem, we have shown that CB promotes bacterial growth in planta and can modulate plant defenses through an antagonistic mechanism between SA and jasmonic acid signaling cascades. Collectively, these data reveal a new link between two processes mediated by SA and iron in response to microbial siderophores.

AtNRAMP3 and AtNRAMP4 are two Arabidopsis metal transporters sharing about 50% sequence identity with mouse NRAMP1. The NRAMP1/Slc11A1 metal ion transporter plays a crucial role in the innate immunity of animal macrophages targeted by intracellular bacterial pathogens. AtNRAMP3 and AtNRAMP4 localize to the vacuolar membrane. We found that AtNRAMP3 is upregulated in leaves challenged with the bacterial pathogens Pseudomonas syringae and Erwinia chrysanthemi, whereas AtNRAMP4 expression is not modified. Using single and double nramp3 and nramp4 mutants, as well as lines ectopically expressing either of these genes, we show that AtNRAMP3 and, to a lesser extent, AtNRAMP4 are involved in Arabidopsis thaliana resistance against the bacterial pathogen E. chrysanthemi. The susceptibility of the double nramp3 nramp4 mutant is associated with the reduced accumulation of reactive oxygen species and ferritin (AtFER1), an iron storage protein known to participate in A. thaliana defense. Interestingly, roots from infected plants accumulated transcripts of AtNRAMP3 as well as the iron-deficiency markers IRT1 and FRO2. This finding suggests the existence of a shoot-to-root signal reminiscent of an iron-deficiency signal activated by pathogen infection. Our data indicate that the functions of NRAMP proteins in innate immunity have been conserved between animals and plants.

Many taxonomically diverse plant species are attacked by Erwinia chrysanthemi, a member of the causal agents of soft-rotting diseases. Symptom development is due to the collective action of pectin-degrading enzymes secreted by the bacterium through a type II secretion system (T2SS). Using Arabidopsis thaliana as a susceptible host, we show that plants respond to E. chrysanthemi 3937 by expressing cell-wall reactions, production of an oxidative burst, and activation of salicylic acid (SA) and jasmonic acid (JA) or ethylene (ET) signaling pathways. We found that the oxidative burst is mainly generated via the expression of the AtrbohD gene, constitutes a barrier of resistance to bacterial attack, and acts independently of the SA-mediated response. To determine the importance of T2SS-secreted proteins in elicitation of these defenses, we used a T2SS deficient mutant and purified enzymatic preparations of representative members of strain 3937 pectate lyase activity. The T2SS-secreted proteins were responsible only partially for the activation of SA and JA or ET signaling pathways observed after infection with the wild-type bacterium and were not involved in the expression of other identified defense reactions. Our study shows the differential role played by pectate lyases isoenzymes in this process and highlights the complexity of the host immune network, which is finely controlled by the bacterium.

Ferritins are multimeric iron storage proteins encoded by a four-member gene family in Arabidopsis (AtFer1-4). To investigate whether iron sequestration in ferritins is a part of an iron-withholding defense system induced in response to bacterial invasion, we used Arabidopsis thaliana as a susceptible host for the pathogenic bacterium Erwinia chrysanthemi. In this study, we used a T-DNA insertion mutant line to show that the lack of a functional AtFer1 gene resulted in an enhanced susceptibility of Arabidopsis plants to E. chrysanthemi. We found that the AtFer1 gene is upregulated during infection, with a biphasic accumulation of the transcript at critical time points 0.5 and 24 h post-infection (p.i.). The activation of AtFer1 expression observed at 24 h p.i. was independent of the iron-dependent regulatory sequence (IDRS) known to mediate the transcriptional response of the AtFer1 gene to iron excess and to nitric oxide. Upregulation of AtFer1 gene expression was compromised after inoculation with an E. chrysanthemi siderophore null mutant. Infiltration of the purified siderophores chrysobactin and desferrioxamine strongly increased AtFer1 transcript abundance and it did not occur with the iron-loaded forms of these siderophores. We found that neither oxidative stress nor nitric oxide was involved in the plant response to chrysobactin. Our data show that ferritin accumulation during infection of Arabidopsis by E. chrysanthemi is a basal defense mechanism which is mainly activated by bacterial siderophores. The potential role of siderophores in this process is discussed.

Starting from November 1990, an international External Quality Assessment Scheme (EQAS) for immmunoassays of tumor markers has been organized. Presently, 238 laboratories from France, Germany, Italy, Japan and Spain participate in the scheme. In this report the main features of the EQAS and data processing are outlined. Results collected during the 1992-cycle allow evaluation of the state of the art of AFP, CEA, CA 19-9, CA 15-3, CA 125 and PSA immunoassays. According to their analytical performances, the 6 tumor marker immunoassays can be classified into several groups, the first including AFP and CA 15-3 for which both total variability and within-kit agreement are good. For CEA assay, performance can be considered as satisfactory even though further improvements of between-lab agreement would be welcome. For the 3 other tumor markers, the higher total variability indicates an urgent need for a better standardization by improvement of either both within-kit and between-kit agreements (CA 19-9) or between-kit agreement mainly (PSA, CA 125).