Phytopathogenic bacteria of the Xanthomonas genus cause severe diseases on hundreds host plants, including economically important crops, such as rice, wheat, cassava, banana, mango, tomato, citrus, cabbage, pepper, bean and cotton. Diseases occurring in nature comprise black rot, leaf/fruit spot, canker, wilt, leaf blight and streak. These bacteria are present worldwide where some phytopathogenic strains are emergent or re-emergent and, consequently, dramatically impact agriculture, economy and food safety.
Xanthomonas bacteria provide excellent models for genomic studies and hundreds of Xanthomonas genome sequences have been obtained since 2002 and many other are underway (www.xanthomonas.org/genomes.html). Comparative genomics between and/or within bacterial species and/or pathovars will be of a great help to decipher commonalities and particularities that underly host range definition.
Most of the Xanthomonas possesses a type III secretion system (T3SS) that is required for injection of various effectors inside plant cells, thus contributing to pathogenicity. Transcription Activator-Like (tal) genes, encode bacterial transcription factors which are injected through the T3SS by many Xanthomonas to promote pathogenicity. Some Ralstonia, Bulkholderia and marine bacteria also express TAL-like proteins which function and mode of action is starting to be deciphered. TALs are addressed to the plant nucleus where they activate plant gene expression by direct binding to the corresponding promoter sequences. Targeted genes essentially act as susceptibility genes. A few years after the cracking of the code allowing the TAL/Host promoter sequence recognition, combined to the ever-growing availability of plant genomes, many efforts have been done to identify TAL targets. These data collected for many Xanthomonas/host pathosystems will assuredly help breeders to breed resistance resistant in important crops.
In this Research Topic we aim to collect manuscripts covering the current knowledge of Xanthomonas genomics and effectomics. Specifically, we encourage the submission of manuscripts (Original Research, Hypothesis & Theory, Methods, Reviews, Mini Reviews, Perspective and Opinion) covering the following topics: 1. Manuscripts reporting genome sequencing of Xanthomonas strains. 2. Manuscripts describing functional and comparative genomics in Xanthomonas species/pathovars. 3. Manuscripts describing functional studies on Xanthomonas type III effectors. 3. Manuscripts describing discovery, evolution, bio-informatics and functional genomics of TAL effectors and their targets in plant genomes. 4. Manuscripts describing applications of TAL effector research for resistance breeding in crops.
We anticipate that this Research Topic will be of importance for plant pathologists and breeders.
The sensing of microbe-associated molecular patterns (MAMPs) triggers innate immunity in animals and plants. Lipopolysaccharide (LPS) from Gram-negative bacteria is a potent MAMP for mammals, with the lipid A moiety activating proinflammatory responses via Toll-like receptor 4 (TLR4). Here we found that the plant Arabidopsis thaliana specifically sensed LPS of Pseudomonas and Xanthomonas. We isolated LPS-insensitive mutants defective in the bulb-type lectin S-domain-1 receptor–like kinase LORE (SD1-29), which were hypersusceptible to infection with Pseudomonas syringae. Targeted chemical degradation of LPS from Pseudomonas species suggested that LORE detected mainly the lipid A moiety of LPS. LORE conferred sensitivity to LPS onto tobacco after transient expression, which demonstrated a key function in LPS sensing and indicated the possibility of engineering resistance to bacteria in crop species.
The modification of proteins by the attachment of fatty acids is a targeting tactic involved in mechanisms of both plant immunity and bacterial pathogenesis. The plant plasma membrane (PM) is a key battleground in the war against disease-causing microbes. This membrane is armed with an array of sensor proteins that function as a surveillance system to detect invading pathogens. Several of these sensor proteins are directed to the plasma membrane through the covalent addition of fatty acids, a process termed fatty acylation. Phytopathogens secrete effector proteins into the plant cell to subvert these surveillance mechanisms, rendering the host susceptible to infection. The targeting of effectors to specific locales within plant cells, particularly the internal face of the host PM, is critical for their virulence function. Several bacterial effectors hijack the host fatty acylation machinery to be modified and directed to this contested locale. To find and fight these fatty acylated effectors the plant leverages lipid-modified intracellular sensors. This review provides examples featuring how fatty acylation is a battle tactic used by both combatants in the molecular arms race between plants and pathogens. Also highlighted is the exploitation of a specific form of host-mediated fatty acid modification, which appears to be exclusively employed by phytopathogenic effector proteins.
Fungal cell walls play dynamic functions in interaction of fungi with their surroundings. In pathogenic fungi, the cell wall is the first structure to make physical contact with host cells. An important structural component of fungal cell walls is chitin, a well-known elicitor of immune responses in plants. Research into chitin perception has sparked since the chitin receptor from rice was cloned nearly a decade ago. Considering the widespread nature of chitin perception in plants, pathogens evidently evolved strategies to overcome detection, including alterations in the composition of cell walls, modification of their carbohydrate chains and secretion of effectors to provide cell wall protection or target host immune responses. Also non-pathogenic fungi contain chitin in their cell walls and are recipients of immune responses. Intriguingly, various mutualists employ chitin-derived signaling molecules to prepare their hosts for the mutualistic relationship. Research on the various types of interactions has revealed different molecular components that play crucial roles and, moreover, that various chitin-binding proteins contain dissimilar chitin-binding domains across species that differ in affinity and specificity. Considering the various strategies from microbes and hosts focused on chitin recognition, it is evident that this carbohydrate plays a central role in plant–fungus interactions.
Andrea Sánchez-Vallet , Jeroen R. Mesters , Bart P.H.J. Thomma
In plant innate immunity, individual cells have the capacity to sense and respond to pathogen attack. Intracellular recognition mechanisms have evolved to intercept perturbations by pathogen virulence factors (effectors) early in host infection and convert it to rapid defense. One key to resistance success is a polymorphic family of intracellular nucleotide-binding/leucine-richrepeat (NLR) receptors that detect effector interference in different parts of the cell. Effector-activated NLRs connect, in various ways, to a conserved basal resistance network in order to transcriptionally boost defense programs. Effector-triggered immunity displays remarkable robustness against pathogen disturbance, in part by employing compensatory mechanisms within the defense network. Also, the mobility of some NLRs and coordination of resistance pathways across cell compartments provides flexibility to fine-tune immune outputs. Furthermore, a number of NLRs function close to the nuclear chromatin by balancing actions of defense-repressing and defense-activating transcription factors to program cells dynamically for effective disease resistance.
Phytopathogens are a global threat to plant agriculture and biodiversity. The genomics era has lead to an exponential rise in comparative gene and genome studies of both economically significant and insignificant microorganisms. In this review we highlight some recent comparisons and discuss how they identify shared genes or genomic regions associated with host virulence. The two major mechanisms of rapid genome adaptation – horizontal gene transfer and hybridisation – are reviewed and we consider how intra-specific pan-genome sequences encode alternative host specificity. We also discuss the power that access to expansive gene databases provides in aiding the study of phytopathogen emergence. These databases can rapidly enable the identification of an unknown pathogen and its origin, as well as genomic adaptations required for emergence.
Elisha Thynne, Megan C. McDonald, Peter S. Solomon
Next to numerous abiotic stresses, plants are constantly exposed to a variety of pathogens within their environment. Thus, their ability to survive and prosper during the course of evolution was strongly dependent on adapting efficient strategies to perceive and to respond to such potential threats. It is therefore not surprising that modern plants have a highly sophisticated immune repertoire consisting of diverse signal perception and intracellular signaling pathways. This signaling network is intricate and deeply interconnected, probably reflecting the diverse lifestyles and infection strategies used by the multitude of invading phytopathogens. Moreover it allows signal communication between developmental and defense programs thereby ensuring that plant growth and fitness are not significantly retarded. How plants integrate and prioritize the incoming signals and how this information is transduced to enable appropriate immune responses is currently a major research area. An important finding has been that pathogen-triggered cellular responses involve massive transcriptional reprogramming within the host. Additional key observations emerging from such studies are that transcription factors (TFs) are often sites of signal convergence and that signal-regulated TFs act in concert with other context-specific TFs and transcriptional co-regulators to establish sensory transcription regulatory networks required for plant immunity.
Plants and microbes are in a continuous arms race to maintain their predominance within their particular niche. Understanding the complexity of these plant–microbe interactions is of utmost importance as it can provide new insights into the mechanisms mediating disease processes and in turn inspire new plant breeding strategies. The International Society for Molecular Plant–Microbe Interactions (IS-MPMI) invited scientists from around the world to share their findings during the XVI International Congress on Molecular Plant–Microbe Interactions, which was held on the beautiful island of Rhodes in Greece. The congress was organized by the Agricultural University of Athens, the Hellenic Phytopathology Society, and the Hellenic Society of Phytiatry and provided over 1100 participants from 55 countries with the opportunity to present and discuss their current and future research. A great number of talks and posters were presented, however our aim within this report is to provide a snapshot of the discipline by focusing on just some of the exciting research and discussions which took place. The key topics discussed were virulence factors, epigenetic regulation, hormones, symbiosis factors, toxins, signaling pathways, microbe recognition, immunity, and pathogen diagnostics. Effector biology was also a recurrent theme in many plenary and concurrent sessions, indicating the importance of a topic that was also highlighted recently by a Virtual Special Issue in New Phytologist (see Kuhn & Panstruga, 2014). In addition to this, throughout the meeting next generation sequencing (NGS) techniques were described and shown to be shedding new light on long-standing issues in microbial ecology.
Genome editing opens up opportunities for the precise and rapid alteration of crops to boost yields, protect against pests and diseases and enhance nutrient content. The extent to which applied plant research and crop breeding benefit will depend on how the EU decides to regulate this fledgling technology.
Symposium aim - We aim to organize a cutting edge meeting focused on the application of cell biology approaches to understand the mechanisms that diverse microbes use to manipulate plant cells to benefit their life styles. The meeting will bring together researchers working on a broad spectrum of microbes across different taxa (bacteria, fungi, oomycetes) that form a variety of different interactions (pathogenic, symbiotic) with plant organs/tissues (leaves, roots). With the explosion in microbial/host genome sequences and the identification of genes/proteins involved in these interactions, the focus of the field is moving rapidly towards using cell and molecular biology techniques and new imaging technologies to understand the molecular dialogue between plants and their microbial pathogens/symbionts. The need for a conference on this topic, the first of its type, is evidenced by the growing prominence of cell biology in the literature. Students and scientists in this field face many challenges in the application and interpretation of cell biology data and would greatly benefit from a specialized conference on this topic. The symposium will bring together a broad representation of researchers focussing on different cell biology aspects and will allow researchers across the different disciplines to present and exchange their recent advances in this important topic of plant biology.
Symposium rationale and scope - Plant organs are subject to colonisation and manipulation by microbes, and this requires reprogramming of host cell biology to accommodate microbial structures within tissues/cells and to mediate responses for proper immunity or for symbiosis. Host cell biology changes during microbial invasion were first reported more than 100 years ago based on microscopy studies revealing that many microbes project structures (haustoria, arbuscules) into plant cells that are enveloped with a specialized plant-derived membrane and evidence now suggests an intimate molecular exchange takes place across these membrane interfaces. However, recent identification of some of the molecular players in these interactions is only now providing appropriate tools to analyse these events. The symposium will focus on advances in understanding the molecular interactions that occur between a microbe and its host at a cellular and subcellular level, such as:
how root and leaf cells accommodate microbial structures through biogenesis of specialized plant derived membranes, microbial invasion and spreading strategies (via stomata, roots, vasculature, plasmodesmata), the dynamic localization of cell surface and cytosolic receptors recognizing microbial signals the reprogramming of host membrane trafficking (focal accumulation, secretion), the delivery of microbial molecules from fungal and oomycete species into plant cells.
With recent advances in high resolution/throughput bioimaging we are gaining new insights into the cell biology mechanisms and pathways of plant cell interactions with diverse microbes. Therefore the symposium provides a timely and important opportunity to overview the application of these technologies to plant–microbe interactions, and to discuss recent discoveries emerging from diverse host–microbe interactions illustrating common underlying principles and differences of strategies used by the microbes to gain access to plant tissues/cells. The symposium will certainly trigger a wealth of discussions, exchange of findings and methodologies, and will promote new lines of research and ideas in this rapidly expanding field.
You will find the summer programs of the University of Angers in the field of health sciences and plant science, which will be held from June 29th to July 10th 2015. Our summer schools offer an unique opportunity for students from all over the world to enjoy Science in a beautiful environment. Each program entirely conducted in English includes conferences by international researchers, hands-on activities, visits of research facilities and biotech companies and an attractive social program. Come and meet in Angers international students who share your passion for Science!
Plenary conference by Prof. Gareth Williams, renowned professor.
The 2015 Molecular Biology of Plant Pathogens (MBPP) conference will be held at the University of the West of England (UWE), Bristol on the 8th-9th April 2015. This will be the 23rd MBPP conference!
UWE is the largest university in the South West of England with over 30,000 students and approximately 3,500 staff. UWE has a long and interesting history starting life as a Merchant Venturer’s Navigation College in 1595 and undergoing many changes before gaining University status in 1992. Today UWE attracts students from all over the UK as well as a significant number of international students from 140 countries worldwide.
UWE has an active research community which makes a significant contribution to advances in industry, commerce, health and technology both nationally and internationally. The organisers of this years’ MBPP conference, Professor Dawn Arnold, Dr Carrie Brady and Dr Helen Neale work within the Centre for Research in Bioscience (CRIB) which leads world-class research in areas of strategic importance including plant science, agri-food, bio-sensing and biomedicine.
MBPP provides an excellent forum for networking between junior and senior scientists. The primary focus is on providing PhD students and post-doctoral scientists the opportunity to give oral presentations in front of a wide range of national and international researchers.
There will also be three keynote talks by internationally renowned scientists Professor Pietro Spanu (Imperial College), Dr Chris Ridout (John Innes Centre) and Professor Teresa Coutinho (Forestry and Agricultural Biotechnology Institute, University of Pretoria, South Africa). Please see our biographies tab for more information on these speakers.
Bananas (Musa spp.) belong to the most important global food commodities, and their cultivation represents the world's largest monoculture. Although the plant-associated microbiome has substantial influence on plant growth and health, there is a lack of knowledge of the banana microbiome and its influencing factors. We studied the impact of (i) biogeography, and (ii) agroforestry on the banana-associated gammaproteobacterial microbiome analyzing plants grown in smallholder farms in Nicaragua and Costa Rica. Profiles of 16S rRNA genes revealed high abundances of Pseudomonadales, Enterobacteriales, Xanthomonadales, and Legionellales. An extraordinary high diversity of the gammaproteobacterial microbiota was observed within the endophytic microenvironments (endorhiza and pseudostem), which was similar in both countries. Enterobacteria were identified as dominant group of above-ground plant parts (pseudostem and leaves). Neither biogeography nor agroforestry showed a statistically significant impact on the gammaproteobacterial banana microbiome in general. However, indicator species for each microenvironment and country, as well as for plants grown in Coffea intercropping systems with and without agri-silvicultural production of different Fabaceae trees (Inga spp. in Nicaragua and Erythrina poeppigiana in Costa Rica) could be identified. For example, banana plants grown in agroforestry systems were characterized by an increase of potential plant-beneficial bacteria, like Pseudomonas and Stenotrophomonas, and on the other side by a decrease of Erwinia. Hence, this study could show that as a result of legume-based agroforestry the indigenous banana-associated gammaproteobacterial community noticeably shifted.
Martina Köberl, Miguel Dita, Alfonso Martinuz, Charles Staver, and Gabriele Berg
En agriculture, les agents phytopathogènes (champignons, virus, bactéries) sont responsables de pertes considérables de rendements et de qualité. Ces pathologies combinées avec la croissance démographique mondiale et les changements climatiques constituent un risque majeur pour la sécurité alimentaire. Pour mettre en place des actions appropriées il est indispensable de détecter, d'identifier et de mieux connaitre ces organismes. Après le congres de Paris en 2012, la SFP continue son compagnonnage et, cette année, le colloque se tiendra en Alsace à Colmar du 2 au 5 juin 2015 au CREF (centre de rencontres, d'échanges et de formation) au 5 Rue des jardins.
Ce colloque, est ouvert aux phytopathologistes français et étrangers. C'est une opportunité d’aborder tous les domaines de la santé des végétaux et de faciliter le dialogue entre chercheurs à thématiques différentes. Pour présenter vos travaux, vous informer des recherches récentes dans ce domaine en France et au-delà, faire des rencontres et des échanges entre collègues, venez au 9éme colloque de la SFP, qui propose un programme passionnant, tant scientifique que touristique. C'est aussi l'occasion de découvrir la gastronomie alsacienne et les richesses touristiques de Colmar.
The last 20 years have provided a sophisticated understanding of how plants recognise relatively conserved microbial patterns to activate defence. In recent years DNA sequencing allowed genomes and transcriptomes of eukaryotic rusts and mildew pathogens to be studied and high-throughput imaging permit the study and visualisation of intracellular interactions during pathogenesis and defence.
We will present many aspects of plant- microbe interactions including:
- gene discovery - genome analysis - intra-cellular interactions with high-throughput imaging technology - mechanistic understanding of cellular and molecular processes to translational activities
The focus on the dynamic and interactive practical sessions will naturally promote strong interactions between lecturers and participants.
Members of the genus Xanthomonas are among the most important phytopathogens. A key feature of Xanthomonas pathogenesis is the translocation of type III secretion system (T3SS) effector proteins (T3SEs) into the plant target cells via a T3SS. Several T3SEs and a murein lytic transglycosylase gene (mlt, required for citrus canker symptoms) are found associated with three transposition-related genes in Xanthomonas citri plasmid pXAC64. These are flanked by short inverted repeats (IRs). The region was identified as a transposon, TnXax1, with typical Tn3 family features, including a transposase and two recombination genes. Two 14-bp palindromic sequences within a 193-bp potential resolution site occur between the recombination genes. Additional derivatives carrying different T3SEs and other passenger genes occur in different Xanthomonas species. The T3SEs include transcription activator-like effectors (TALEs). Certain TALEs are flanked by the same IRs as found in TnXax1 to form mobile insertion cassettes (MICs), suggesting that they may be transmitted horizontally. A significant number of MICs carrying other passenger genes (including a number of TALE genes) were also identified, flanked by the same TnXax1 IRs and delimited by 5-bp target site duplications. We conclude that a large fraction of T3SEs, including individual TALEs and potential pathogenicity determinants, have spread by transposition and that TnXax1, which exhibits all of the essential characteristics of a functional transposon, may be involved in driving MIC transposition. We also propose that TALE genes may diversify by fork slippage during the replicative Tn3 family transposition. These mechanisms may play a crucial role in the emergence of Xanthomonas pathogenicity.
Plants host distinct microbial communities on and inside their tissues designated the plant microbiota. Microbial community profiling enabled the description of the phylogenetic structure of the plant microbiota to an unprecedented depth, whereas functional insights are largely derived from experiments using individual microorganisms. The binary interplay between isolated members of the plant microbiota and host plants ranges from mutualistic to commensalistic and pathogenic relationships. However, how entire microbial communities capable of executing both growth-promoting and growth-compromising activities interfere with plant fitness remains largely unknown. Ultimately, unravelling the net result of microbial activities encoded in the extended plant genome—the plant microbiome—will be key to understanding and exploiting the full yield potential of a crop plant. In this perspective, we summarize first achievements of plant-microbiome research, we discuss future research directions, and we provide ideas for the translation of basic science to application to capitalize on the plant microbiome at work.
We report the draft genome sequences of two Xanthomonas euvesicatoria strains from the Balkan Peninsula, which were isolated from symptomatic pepper plants. The availability of these genome sequences will facilitate the development of modern genotyping assays for X. euvesicatoria strains and to define targets for resistance breeding.
Comparison with the completely sequenced strain 85-10, which was isolated in Florida in 1985, revealed some unique effectors that are present or absent in one or the other strain (e.g., AvrBs1, AvrBs3, XopE3, XopG, XopH, XopAF, and XopAQ).
During plant immunity, surface-localized pattern recognition receptors (PRRs) recognize pathogen-associated molecular patterns (PAMPs). The transfer of PRRs between plant species is a promising strategy for engineering broad-spectrum disease resistance. Thus, there is a great interest in understanding the mechanisms of PRR-mediated resistance across different plant species. Two well-characterized plant PRRs are the leucine-rich repeat receptor kinases (LRR-RKs) EFR and XA21 from Arabidopsis thaliana (Arabidopsis) and rice, respectively. Interestingly, despite being evolutionary distant, EFR and XA21 are phylogenetically closely related and are both members of the sub-family XII of LRR-RKs that contains numerous potential PRRs. Here, we compared the ability of these related PRRs to engage immune signaling across the monocots-dicots taxonomic divide. Using chimera between Arabidopsis EFR and rice XA21, we show that the kinase domain of the rice XA21 is functional in triggering elf18-induced signaling and quantitative immunity to the bacteria Pseudomonas syringae pv. tomato (Pto) DC3000 and Agrobacterium tumefaciens in Arabidopsis. Furthermore, the EFR:XA21 chimera associates dynamically in a ligand-dependent manner with known components of the EFR complex. Conversely, EFR associates with Arabidopsis orthologues of rice XA21-interacting proteins, which appear to be involved in EFR-mediated signaling and immunity in Arabidopsis. Our work indicates the overall functional conservation of immune components acting downstream of distinct LRR-RK-type PRRs between monocots and dicots.
Nicholas Holton, Vladimir Nekrasov, Pamela C. Ronald, Cyril Zipfel
"Our multi-species genome-wide analysis reveals: i) AUX, CK and SL signaling pathways originated in charophyte lineages; ii) ABA, JA, and SA signaling pathways arose in the last common ancestor of land plants; iii) the GA signaling evolved after the divergence of bryophytes from land plants; iv) the canonical BR signaling originated before the emergence of angiosperms but likely after the split of gymnosperms and angiosperms; v) the origin of the canonical ETH signaling pathway postdates shortly the emergence of angiosperms. Our findings might have important implications in understanding the molecular mechanisms underlying the emergence of land plants"
Stable and efficient knockdown of multiple gene targets is highly desirable for dissection of molecular pathways. Because it allows sequence-specific DNA binding, transcription activator-like effector (TALE) offers a new genetic perturbation technique that allows for gene-specific repression. Here, we constructed a multicolor lentiviral TALE-Kruppel-associated box (KRAB) expression vector platform that enables knockdown of multiple gene targets. This platform is fully compatible with the Golden Gate TALEN and TAL Effector Kit 2.0, a widely used and efficient method for TALE assembly. We showed that this multicolor TALE-KRAB vector system when combined together with bone marrow transplantation could quickly knock down c-kit and PU.1 genes in hematopoietic stem and progenitor cells of recipient mice. Furthermore, our data demonstrated that this platform simultaneously knocked down both c-Kit and PU.1 genes in the same primary cell populations. Together, our results suggest that this multicolor TALE-KRAB vector platform is a promising and versatile tool for knockdown of multiple gene targets and could greatly facilitate dissection of molecular pathways.
Understanding how plants balance between enabling microbial symbionts and fending off pathogens has direct implications both for basic plant biology and optimal use of crop plants in agriculture. The degree to which the processes associated with these two types of interactions overlap is poorly known. Recent studies revealed that symbiotic and pathogenic filamentous microbes require common plant genetic elements to establish colonization (Wang et al., 2012; Rey et al., 2013), supporting the long-held view that plants have evolved the ability to accommodate microbes (Parniske, 2000) and that pathogens have exploited these pathways. However, the extent to which plant genes implicated in fungal or bacterial symbioses are involved in interactions with biotrophic pathogens is unknown and research has been hampered by the lack of suitable common host experimental systems.
In this study, we took advantage of a newly established quantitative Phytophthora palmivora–Medicago truncatula system to assess the extent to which mutants perturbed in colonization by arbuscular mycorrhiza fungi (AM fungi) and/or bacterial root nodule symbiosis are affected in the early/biotrophic stages of oomycete pathogenesis (Supporting Information Table S1). We devised and implemented a high throughput seedling infection assay and applied it to 19 M. truncatula lines mutated in 14 genes (for details see Methods S1; for explanation of gene abbreviations see Table S1). We measured both the overall root length and disease development, then plotted them as a ratio (Figs 1a,c, S1; Table S2). Of the 14 genes tested, seven (nine alleles) showed an altered response to P. palmivora inoculation compared with the wild-type Jemalong A17. Mutants in RAM2 and NIP/LATD showed enhanced resistance whereas mutants in five genes, NFP, LYK3, ERN, EFD, and LIN, all of which are impaired in the interaction with nitrogen fixing rhizobia displayed enhanced susceptibility. Expression levels of two defence response genes in M. truncatula mutants with altered disease symptoms were not overall significantly different from levels observed during infection of wild-type A17 seedlings (Fig. S2). This suggests that observed differences in disease extent are not attributable to altered defence responses in these mutants. These findings reveal a significant overlap between processes that define symbiosis and disease in M. truncatula roots. However, the remaining M. truncatula mutants unaltered in P. palmivora disease development include the common symbiotic signalling pathway (CSSP) mutants dmi1, dmi2 and dmi3, suggesting that the CSSP is not a major modulator of susceptibility to P. palmivora in M. truncatula.
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