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.
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.
Recently, emission of volatile organic compounds (VOCs) has emerged as a mode of communication between bacteria and plants. Although some bacterial VOCs that promote plant growth have been identified, their underlying mechanism of action is unknown. Here we demonstrate that indole, which was identified using a screen for Arabidopsis growth promotion by VOCs from soil-borne bacteria, is a potent plant-growth modulator. Its prominent role in increasing the plant secondary root network is mediated by interfering with the auxin-signalling machinery. Using auxin reporter lines and classic auxin physiological and transport assays we show that the indole signal invades the plant body, reaches zones of auxin activity and acts in a polar auxin transport-dependent bimodal mechanism to trigger differential cellular auxin responses. Our results suggest that indole, beyond its importance as a bacterial signal molecule, can serve as a remote messenger to manipulate plant growth and development.
Many phytopathogenic type III secretion effectors (T3Es) have been shown to target and suppress plant immune signaling, but perturbation of the plant immune system by T3Es can also elicit a plant response. XopX is a “core” Xanthomonas T3E that contributes to growth and symptom development during Xanthomonas euvesicatoria (Xe) infection of tomato, but its functional role is undefined. We tested the effect of XopX on several aspects of plant immune signaling. XopX promoted ethylene production and plant cell death (PCD) during Xe infection of susceptible tomato and in transient expression assays in Nicotiana benthamiana, which is consistent with its requirement for the development of Xe-induced disease symptoms. Additionally, although XopX suppressed flagellin-induced reactive oxygen species, it promoted the accumulation of pattern-triggered immunity (PTI) gene transcripts. Surprisingly, XopX co-expression with other PCD elicitors resulted in delayed PCD, suggesting antagonism between XopX-dependent PCD and other PCD pathways. However, we found no evidence that XopX contributed to the suppression of effector-triggered immunity during Xe-tomato interactions, suggesting that XopX’s primary virulence role is to modulate PTI. These results highlight the dual role of a core Xanthomonas T3E in simultaneously suppressing and activating plant defense responses.
Plants have evolved disease resistance (R) genes encoding for nucleotide-binding site (NB) and leucine-rich repeat (LRR) proteins with N-terminals represented by either Toll/Interleukin-1 receptor (TIR) or coiled-coil (CC) domains. Here, a genome-wide study of presence and diversification of CC-NB-LRR and TIR-NB-LRR encoding genes, and shorter domain combinations in 19 Arabidopsis thaliana accessions and Arabidopsis lyrata, Capsella rubella, Brassica rapa and Eutrema salsugineum are presented.ResultsOut of 528 R genes analyzed, 12 CC-NB-LRR and 17 TIR-NB-LRR genes were conserved among the 19 A. thaliana genotypes, while only two CC-NB-LRRs, including ZAR1, and three TIR-NB-LRRs were conserved when comparing the five species. The RESISTANCE TO LEPTOSPHAERIA MACULANS 1 (RLM1) locus confers resistance to the Brassica pathogen L. maculans the causal agent of blackleg disease and has undergone conservation and diversification events particularly in B. rapa. On the contrary, the RLM3 locus important in the immune response towards Botrytis cinerea and Alternaria spp. has recently evolved in the Arabidopsis genus.ConclusionOur genome-wide analysis of the R gene repertoire revealed a large sequence variation in the 23 cruciferous genomes. The data provides further insights into evolutionary processes impacting this important gene family.
Plants posses an intricate innate immune system that enables them to fight off most invading pathogens. Around the world, agriculture relies on robust disease resistance to ensure adequate food and feed production. Researchers and breeders are constantly generating new resistant crop varieties mostly employing the lengthy process of conventional breeding. Nonetheless, crop losses due to plant pathogens are estimated to be over 15% every year - the main cause of such losses is rapid evolution of new virulent races. In order to keep up with emerging pathogens, we need to gain a deeper and more systematic understanding of the immune system of our crops. During the past two decades, molecular understanding of plant innate immune signaling has been greatly expanded using dicotyledonous model systems such as Arabidopsis thaliana. Now, it is time to connect this volume of knowledge with the immune system of the crop species.
In this Research Topic we aim to collect manuscripts covering the current knowledge of the immune systems of major crop species. Specifically, we encourage the submission of manuscripts (Original Research, Hypothesis & Theory, Methods, Reviews, Mini Reviews, Perspective and Opinion) covering the following topics:
a. Manuscripts describing our current understanding of the plant immune system with a focus on crop species or comparative analyses between model systems and crops.
b. Manuscripts exploring how to best exploit our insight into genomes of plant pathogens and molecular understanding of effector function. c. Manuscripts debating (novel) strategies of how to generate more resistant crop varieties. These might include biotechnological, social and economical aspects of crop improvement.
We anticipate that this Research Topic will become an important resource for plant immunologists especially those interested in comparative studies of plant innate immune systems of model systems and crop species.
Benjamin Schwessinger UC Davis Davis, USA
Rebecca Bart Donald Danforth Plant Science Center St. Louis, USA
Gitta Coaker University of California, Davis Davis, USA
Ksenia V Krasileva University of California Davis Davis, USA
Plant immunity is often triggered by the specific recognition of pathogen effectors by intracellular nucleotide-binding, leucine-rich repeat (NLR) receptors. Plant NLRs contain an N-terminal signaling domain that is mostly represented by either a Toll-interleukin1 receptor (TIR) domain or a coiled coil (CC) domain. In many cases, single NLR proteins are sufficient for both effector recognition and signaling activation. However, many paired NLRs have now been identified where both proteins are required to confer resistance to pathogens. Recent detailed studies on the Arabidopsis thaliana TIR-NLR pair RRS1 and RPS4 and on the rice CC-NLR pair RGA4 and RGA5 have revealed for the first time how such protein pairs function together. In both cases, the paired partners interact physically to form a hetero-complex receptor in which each partner plays distinct roles in effector recognition or signaling activation, highlighting a conserved mode of action of NLR pairs across both monocotyledonous and dicotyledonous plants. We also discuss a new ‘integrated decoy’ effector recognition model to describe these receptor complexes that may be common to many other plant NLR pairs.
Xanthomonas axonopodis pathovar vasculorum strain NCPPB 900 was isolated from sugarcane on Reunion island in 1960. Consistent with its belonging to fatty acid type D, multi-locus sequence analysis confirmed that NCPPB 900 falls within the species X. axonopodis. This genome harbours sequences similar to plasmids pXCV183 from X. campestris pv. vesicatoria 85-10 and pPHB194 from Burkholderia pseudomallei. Its repertoire of predicted effectors includes homologues of XopAA, XopAD, XopAE, XopB, XopD, XopV, XopZ, XopC and XopI and transcriptional activator-like (TAL) effectors and it is predicted to encode a novel phosphonate natural product also encoded by the genome of the phylogenetically distant X. vasicola pv. vasculorum. Availability of this novel genome sequence may facilitate the study of interactions between xanthomonads and sugarcane, a host-pathogen system that appears to have evolved several times independently within the genus Xanthomonas and may also provide a source of target sequences for molecular detection and diagnostics.
Ca2+ is a ubiquitous second messenger for cellular signalling in various stresses and developmental processes. Here, we summarize current developments in the roles of Ca2+ during plant immunity responses. We discuss the early perception events preceding and necessary for triggering cellular Ca2+ fluxes, the potential Ca2+-permeable channels, the decoding of Ca2+ signals predominantly via Ca2+-dependent phosphorylation events and transcriptional reprogramming. To highlight the complexity of the cellular signal network, we briefly touch on the interplay between Ca2+-dependent signalling and selected major signalling mechanisms – with special emphasis on reactive oxygen species at local and systemic levels.
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.
In eukaryotes, regulated protein turnover is required during many cellular processes, including defense against pathogens. Ubiquitination and degradation of ubiquitinated proteins via the ubiquitin – proteasome system (UPS) is the main pathway for the turnover of intracellular proteins in eukaryotes. The extensive utilization of the UPS in host cells makes it an ideal pivot for the manipulation of cellular processes by pathogens. Like many other Gram-negative bacteria, Xanthomonas species secrete a suite of type-III effector proteins (T3Es) into their host cells to promote virulence. Some of these T3Es exploit the plant UPS to interfere with immunity. This review summarizes T3E examples from the genus Xanthomonas with a proven or suggested interaction with the host UPS or UPS-like systems and also discusses the apparent paradox that arises from the presence of T3Es that inhibit the UPS in general while others rely on its activity for their function.
S. Üstün and F. Börnke
Nicolas Denancé's insight:
Mini-review published in the Research topic on Genomics and Effectomics of the crop killer Xanthomonas.
Recognition of extracellular peptides by plasma membrane‐localized receptor proteins is commonly used in signal transduction. In plants, very little is known about how extracellular peptides are processed and activated in order to allow recognition by receptors. Here, we show that induction of cell death in planta by a secreted plant protein GRIM REAPER (GRI) is dependent on the activity of the type II metacaspase METACASPASE‐9. GRI is cleaved by METACASPASE‐9 in vitro resulting in the release of an 11 amino acid peptide. This peptide bound in vivo to the extracellular domain of the plasma membrane‐localized, atypical leucine‐rich repeat receptor‐like kinase POLLEN‐SPECIFIC RECEPTOR‐LIKE KINASE 5 (PRK5) and was sufficient to induce oxidative stress/ROS‐dependent cell death. This shows a signaling pathway in plants from processing and activation of an extracellular protein to recognition by its receptor.
Identifying the factors that influence the outcome of host–microbial interactions is critical to protecting biodiversity, minimizing agricultural losses and improving human health. A few genes that determine symbiosis or resistance to infectious disease have been identified in model species, but a comprehensive examination of how a host genotype influences the structure of its microbial community is lacking. Here we report the results of a field experiment with the model plantArabidopsis thaliana to identify the fungi and bacteria that colonize its leaves and the host loci that influence the microbe numbers. The composition of this community differs among accessions of A. thaliana. Genome-wide association studies (GWAS) suggest that plant loci responsible for defense and cell wall integrity affect variation in this community. Furthermore, species richness in the bacterial community is shaped by host genetic variation, notably at loci that also influence the reproduction of viruses, trichome branching and morphogenesis.
Plant nucleotide-binding leucine-rich repeat (NB-LRR) disease resistance (R) proteins recognize specific “avirulent” pathogen effectors and activate immune responses. NB-LRR proteins structurally and functionally resemble mammalian Nod-like receptors (NLRs). How NB-LRR and NLR proteins activate defense is poorly understood. The divergently transcribed Arabidopsis R genes, RPS4 (resistance to Pseudomonas syringae 4) and RRS1 (resistance to Ralstonia solanacearum 1), function together to confer recognition of PseudomonasAvrRps4 and Ralstonia PopP2. RRS1 is the only known recessive NB-LRR R gene and encodes a WRKY DNA binding domain, prompting suggestions that it acts downstream of RPS4 for transcriptional activation of defense genes. We define here the early RRS1-dependent transcriptional changes upon delivery of PopP2 via Pseudomonas type III secretion. The Arabidopsis slh1 (sensitive to low humidity 1) mutant encodes an RRS1 allele (RRS1SLH1) with a single amino acid (leucine) insertion in the WRKY DNA-binding domain. Its poor growth due to constitutive defense activation is rescued at higher temperature. Transcription profiling data indicate that RRS1SLH1-mediated defense activation overlaps substantially with AvrRps4- and PopP2-regulated responses. To better understand the genetic basis of RPS4/RRS1-dependent immunity, we performed a genetic screen to identify suppressor of slh1 immunity (sushi) mutants. We show that many sushi mutants carry mutations in RPS4, suggesting that RPS4 acts downstream or in a complex with RRS1. Interestingly, several mutations were identified in a domain C-terminal to the RPS4 LRR domain. Using an Agrobacterium-mediated transient assay system, we demonstrate that the P-loop motif of RPS4 but not of RRS1SLH1 is required for RRS1SLH1 function. We also recapitulate the dominant suppression of RRS1SLH1 defense activation by wild type RRS1 and show this suppression requires an intact RRS1 P-loop. These analyses of RRS1SLH1shed new light on mechanisms by which NB-LRR protein pairs activate defense signaling, or are held inactive in the absence of a pathogen effector.
Plant pathogens display impressive versatility in adapting to host immune systems. Pathogen effector proteins facilitate disease but can become avirulence (Avr) factors when the host acquires discrete recognition capabilities that trigger immunity. The mechanisms that lead to changes to pathogen Avr factors that enable escape from host immunity are diverse, and include epigenetic switches that allow for reuse or recycling of effectors. This perspective outlines possibilities of how epigenetic control of Avr effector gene expression may have arisen and persisted in plant pathogens, and how it presents special problems for diagnosis and detection of specific pathogen strains or pathotypes.
Rust diseases caused by fungi of the order Pucciniales afflict a wide range of plants, including cereals, legumes, ornamentals, and fruit trees, and pose a serious threat to cropping systems and global food security. The obligate parasitic lifestyle of these fungi and their complex life cycles, often involving alternate hosts for the sexual and asexual stages, also make this group of pathogens of great biological interest. One of the most remarkable adaptations of rust fungi is the specialized infection structure that underpins the sustained biotrophic association with hosts; the haustorium (Figure 1A and C). This organ forms after penetration of the wall of a live host cell, expanding on the inner side of the cell wall while invaginating the surrounding host plasma membrane (Figure 1C). Through haustoria, the pathogen derives nutrients from the host and secretes virulence proteins called effectors, which are believed to be the key players that manipulate the physiological and immune responses of host cells –. Analogous terminal feeding structures have independently evolved in other organisms such as the haustorium in powdery mildews (ascomycetes) and downy mildews (oomycetes, not true fungi), and the arbuscules in arbuscular mycorrhizae, suggesting that such architecture represents a successful adaptation of these organisms to interact with their respective host plants , .
Garnica DP, Nemri A, Upadhyaya NM, Rathjen JP, Dodds PN
While conceptual principles governing plant immunity are becoming clear, its systems-level organization and the evolutionary dynamic of the host-pathogen interface are still obscure. We generated a systematic protein-protein interaction network of virulence effectors from the ascomycete pathogen Golovinomyces orontii and Arabidopsis thaliana host proteins. We combined this data set with corresponding data for the eubacterial pathogen Pseudomonas syringae and the oomycete pathogen Hyaloperonospora arabidopsidis. The resulting network identifies host proteins onto which intraspecies and interspecies pathogen effectors converge. Phenotyping of 124 Arabidopsis effector-interactor mutants revealed a correlation between intraspecies and interspecies convergence and several altered immune response phenotypes. Several effectors and the most heavily targeted host protein colocalized in subnuclear foci. Products of adaptively selected Arabidopsis genes are enriched for interactions with effector targets. Our data suggest the existence of a molecular host-pathogen interface that is conserved across Arabidopsis accessions, while evolutionary adaptation occurs in the immediate network neighborhood of effector targets.
Ralf Weßling, Petra Epple, Stefan Altmann,Yijian He, Li Yang, Stefan R. Henz, Nathan McDonald, Kristin Wiley, Kai Christian Bader, Christine Glaßer, M. Shahid Mukhtar, Sabine Haigis, Lila Ghamsari, Amber E. Stephens, Joseph R. Ecker, Marc Vidal, Jonathan D.G. Jones,Klaus F.X. Mayer, Emiel Ver Loren van Themaat, Detlef Weigel, Paul Schulze-Lefert, Jeffery L. Dangl, Ralph Panstruga, and Pascal Braun