The Rpi-mcq1 resistance gene family recognizes Avr2 of Phytophthora infestans but is distinct from R2

AbstractPotato late blight, which is caused by the destructive oomycete pathogen Phytophthora infestans, is a major threat to global food security. Several nucleotide binding, leucine-rich repeat (NLR) Resistance to P. infestans (Rpi) genes have been introgressed into potato cultivars from wild Solanum species that are native to Mexico, but these were quickly defeated. Positional cloning in Solanum mochiquense, combined with allele mining in Solanum huancabambense, were used to identify a new family of Rpi genes from Peruvian Solanum species. Rpi-mcq1, Rpi-hcb1.1 and Rpi-hcb1.2 confer race-specific resistance to a panel of P. infestans isolates. Effector assays showed that the Rpi-mcq1 family mediates a hypersensitive response upon recognition of the RXLR effector AVR2, which had previously been found to be exclusively recognized by the family of R2 resistance proteins. The Rpi-mcq1 and R2 genes are distinct and reside on chromosome IX and IV, respectively. This is the first report of two unrelated R protein families that recognize the same AVR protein. We anticipate that this likely is a consequence of a geographically separated dynamic co-evolution of R gene families of Solanum with an important effector gene of P. infestans.Author summaryPotato is the largest non-grain staple crop and essential for food security world-wide. However, potato plants are continuously threatened by the notorious oomycete pathogen Phytophthora infestans that causes late blight. This devastating disease has led to the Irish famine more then 150 years ago, and is still a major threat for potato. Resistance against P. infestans can be found in wild relatives of potato, which carry resistance genes that belong to the nucleotide binding site-leucine-rich repeat (NLR) class. Known NLR proteins typically recognize a matching effector from Phytophthora, which leads to a hypersensitive resistance response (HR). For example, R2 from Mexican Solanum species recognizes AVR2 from P. infestans. So far, these R genes exclusively match to one Avr gene. Here, we identified a new class of NLR proteins that are different from R2, but also recognize the same effector AVR2. This new family of NLR occurs in South American Solanum species, and we anticipate that it is likely a product of a geographically separated co-evolution with AVR2. This is the first report of two unrelated R protein families that recognize the same AVR protein.

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Rpi-amr3 confers resistance to multiple Phytophthora species by recognizing a conserved RXLR effector

10.1101/2021.06.10.447899 ◽

2021 ◽

Author(s):

Xiao Lin ◽

Andrea Olave-Achury ◽

Robert Heal ◽

Kamil Witek ◽

Hari S. Karki ◽

...

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Plant Pathogens ◽

Nucleotide Binding ◽

Phytophthora Species ◽

Crop Plants ◽

Leucine Rich Repeat ◽

Nucleotide Binding Domain ◽

Rxlr Effectors ◽

Rxlr Effector

Diverse pathogens from the genus Phytophthora cause disease and reduce yields in many crop plants. Although many Resistance to Phytophthora infestans (Rpi) genes effective against potato late blight have been cloned, few have been cloned against other Phytophthora species. Most Rpi genes encode nucleotide-binding domain, leucine-rich repeat- containing (NLR) proteins, that recognize RXLR effectors. However, whether NLR proteins can recognize RXLR effectors from multiple different Phytophthora pathogens has rarely been investigated. Here, we report the effector AVRamr3 from P. infestans that is recognized by Rpi-amr3 from S. americanum. We show here that AVRamr3 is broadly conserved in many different Phytophthora species, and that recognition of AVRamr3 homologs enables resistance against multiple Phytophthora pathogens, including P. parasitica and P. palmivora. Our findings suggest a novel path to identifying R genes against important plant pathogens.

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Rapid Detection of Phytophthora infestans in Late Blight-Infected Potato and Tomato Using PCR

Plant Disease ◽

10.1094/pdis.1997.81.9.1042 ◽

1997 ◽

Vol 81 (9) ◽

pp. 1042-1048 ◽

Cited By ~ 80

Author(s):

C. L. Trout ◽

J. B. Ristaino ◽

M. Madritch ◽

T. Wangsomboondee

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Pcr Amplification ◽

Accurate Method ◽

Phytophthora Species ◽

Universal Primers ◽

Direct Pcr ◽

Field Samples ◽

Pcr Products ◽

Oomycete Pathogen

Late blight caused by the oomycete pathogen Phytophthora infestans is a devastating disease of potato and tomato worldwide. A rapid and accurate method for specific detection of P. infestans is necessary for determination of late blight in infected fruit, leaves, and tubers. Ribosomal DNA (rDNA) from four isolates of P. infestans representing the four genotypes US1, US6, US7, and US8 was amplified using polymerase chain reaction (PCR) and the universal primers internal transcribed spacer (ITS) 4 and ITS5. PCR products were sequenced using an automated sequencer. Sequences were aligned with published sequences from 5 other Phytophthora species, and a region specific to P. infestans was used to construct a PCR primer (PINF). Over 140 isolates representing 14 species of Phytophthora and at least 13 other genera of fungi and bacteria were used to screen the PINF primer. PCR amplification with primers PINF and ITS5 results in amplification of an approximately 600 base pair product with only isolates of P. infestans from potato and tomato, as well as isolates of P. mirabilis and P. cactorum. P. mirabilis and P. cactorum are not pathogens of potato; however, P. cactorum is a pathogen of tomato. P. infestans and P. cactorum were differentiated by restriction digests of the amplified product. The PINF primer was used with a rapid NaOH lysis technique for direct PCR of P. infestans from infected tomato and potato field samples. The PINF primer will provide a valuable tool for detection of P. infestans in potatoes and tomatoes.

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A Tomato Nucleotide Binding Sites−Leucine-Rich Repeat Gene Is Positively Involved in Plant Resistance to Phytophthora infestans

Phytopathology ◽

10.1094/phyto-12-17-0389-r ◽

2018 ◽

Vol 108 (8) ◽

pp. 980-987 ◽

Cited By ~ 8

Author(s):

Ning Jiang ◽

Jun Cui ◽

Jun Meng ◽

Yushi Luan

Keyword(s):

Phytophthora Infestans ◽

Binding Sites ◽

Nucleotide Binding ◽

Leucine Rich Repeat ◽

Tomato Plants ◽

Nucleotide Binding Sites ◽

Transgenic Tomato Plants ◽

Transgenic Lines ◽

Lrr Protein

The nucleotide binding sites−leucine-rich repeat (NBS-LRR) genes are key regulatory components of plant to pathogens. Phytophthora infestans-inducible coding sequence encoding an NBS-LRR (SpNBS-LRR) protein in tomato (Solanum pimpinellifolium L3708) was cloned and characterized based on our RNA-Seq data and tomato genome. After sequence analysis, SpNBS-LRR was identified as a hydrophilic protein with no transmembrane topological structure and no signal peptide. SpNBS-LRR had a close genetic relationship to RPS2 of Arabidopsis thaliana by phylogenetic analysis. In addition, SpNBS-LRR gene was mainly expressed in root, with low expression observed in leaf and stem. To further investigate the role of SpNBS-LRR in tomato−P. infestans interaction, SpNBS-LRR was introduced in susceptible tomatoes and three transgenic lines with higher expression level of SpNBS-LRR were selected. These transgenic tomato plants that overexpressed SpNBS-LRR displayed greater resistance than wild-type tomato plants after infection with P. infestans, as shown by decreased disease index, lesion diameters, number of necrotic cells, P. infestans abundance, and higher expression levels of the defense-related genes. This information provides insight into SpNBS-LRR involved in the resistance of tomato to P. infestans infection and candidate for breeding to enhance biotic stress-resistance in tomato.

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Metabolic Model of the Phytophthora infestans-Tomato Interaction Reveals Metabolic Switches during Host Colonization

mBio ◽

10.1128/mbio.00454-19 ◽

2019 ◽

Vol 10 (4) ◽

Cited By ~ 3

Author(s):

Sander Y. A. Rodenburg ◽

Michael F. Seidl ◽

Howard S. Judelson ◽

Andrea L. Vu ◽

Francine Govers ◽

...

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Metabolic Model ◽

Host Cells ◽

Complex Nature ◽

Infection Cycle ◽

In Planta ◽

Content Type ◽

Metabolic Fluxes ◽

Oomycete Pathogen

ABSTRACT The oomycete pathogen Phytophthora infestans causes potato and tomato late blight, a disease that is a serious threat to agriculture. P. infestans is a hemibiotrophic pathogen, and during infection, it scavenges nutrients from living host cells for its own proliferation. To date, the nutrient flux from host to pathogen during infection has hardly been studied, and the interlinked metabolisms of the pathogen and host remain poorly understood. Here, we reconstructed an integrated metabolic model of P. infestans and tomato (Solanum lycopersicum) by integrating two previously published models for both species. We used this integrated model to simulate metabolic fluxes from host to pathogen and explored the topology of the model to study the dependencies of the metabolism of P. infestans on that of tomato. This showed, for example, that P. infestans, a thiamine auxotroph, depends on certain metabolic reactions of the tomato thiamine biosynthesis. We also exploited dual-transcriptome data of a time course of a full late blight infection cycle on tomato leaves and integrated the expression of metabolic enzymes in the model. This revealed profound changes in pathogen-host metabolism during infection. As infection progresses, P. infestans performs less de novo synthesis of metabolites and scavenges more metabolites from tomato. This integrated metabolic model for the P. infestans-tomato interaction provides a framework to integrate data and generate hypotheses about in planta nutrition of P. infestans throughout its infection cycle. IMPORTANCE Late blight disease caused by the oomycete pathogen Phytophthora infestans leads to extensive yield losses in tomato and potato cultivation worldwide. To effectively control this pathogen, a thorough understanding of the mechanisms shaping the interaction with its hosts is paramount. While considerable work has focused on exploring host defense mechanisms and identifying P. infestans proteins contributing to virulence and pathogenicity, the nutritional strategies of the pathogen are mostly unresolved. Genome-scale metabolic models (GEMs) can be used to simulate metabolic fluxes and help in unravelling the complex nature of metabolism. We integrated a GEM of tomato with a GEM of P. infestans to simulate the metabolic fluxes that occur during infection. This yields insights into the nutrients that P. infestans obtains during different phases of the infection cycle and helps in generating hypotheses about nutrition in planta.

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First Report of Late Blight Caused by Phytophthora infestans Clonal Lineage US-24 on Potato (Solanum tuberosum) in Wisconsin

Plant Disease ◽

10.1094/pdis-09-12-0825-pdn ◽

2013 ◽

Vol 97 (1) ◽

pp. 152-152 ◽

Cited By ~ 6

Author(s):

A. J. Gevens ◽

A. C. Seidl

Keyword(s):

Solanum Tuberosum ◽

Late Blight ◽

Phytophthora Infestans ◽

Weather Conditions ◽

Clonal Lineage ◽

Banding Patterns ◽

First Report ◽

The Us ◽

Epidemiological Characteristics ◽

The U.S

Potato (Solanum tuberosum) crops are grown on over 25,090 ha in Wisconsin annually. Late blight, caused by Phytophthora infestans (Mont.) deBary, is a potentially devastating disease that affects tomato and potato crops in Wisconsin every few years when inoculum is introduced and weather conditions favor disease. Incidence and severity of late blight are highly variable in these few years due to differences in pathogen clonal lineages, their timing and means of introduction, and weather conditions. Prevention of this disease through prophylactic fungicide application can cost producers millions of dollars annually in additional chemical, fuel, and labor expenses. Populations of P. infestans in the U.S. have recently undergone significant genetic change, resulting in isolates with unique clonal lineages and epidemiological characteristics (1). In 2010, late blight epidemics were of low severity in discrete portions of a few fields and were seen exclusively on potato in two counties of central Wisconsin. Symptoms included water-soaked to dark brown circular lesions with pale green haloes accompanied by white fuzzy pathogen sporulation typically on leaf undersides in high humidity conditions. Infected plants were collected by professional crop consultants and submitted to the authors at the University of Wisconsin Vegetable Pathology Laboratory in Madison, Wisconsin. Eight isolates of P. infestans were generated from individual leaf samples, representing separate fields, by removing sporangia from sporulating lesions and placing onto Rye A agar amended with rifampicin and ampicillin. Axenic, single zoospore-derived cultures of isolates were generated from parent cultures and maintained on Rye A agar for further characterization. Mycelium was coenocytic with hyphal diameter of 5 to 8 μm (n = 50). Sporangia were limoniform to ovoid, semi- to fully papillate, caducous, had short pedicels, and were 36.22 × 19.11 μm (height × width; n = 50). The average length-width ratio was 1.91. Allozyme banding patterns at the glucose-6-phosphate isomerase (Gpi) locus indicated a 100/100/111 profile, consistent with the US-24 clonal lineage (3,4). Mating type assays confirmed the isolates to be A1 and intermediate insensitivity to mefenoxam was observed in vitro (4). Genomic DNA was extracted with a phenol:chloroform:isoamyl alcohol solution and restriction fragment length polymorphism (RFLP) analysis was performed using the RG-57 probe on a representative isolate and resulted in banding patterns consistent with US-24 (2,3). Clonal lineages of P. infestans documented in Wisconsin in previous epidemics included US-8 in the mid-1990s and US-1 in the 1970s. The US-24 (A1) clonal lineage was very widespread in the U.S. in 2010 and its presence in Wisconsin in the same year as identification of US-22 (A2) posed great concern for potential sexual recombination, oospore production, and soil persistence. Fortunately, the opposite mating types were separated spatiotemporally. To the best of our knowledge, this is the first report of the P. infestans clonal lineage US-24 causing late blight on potato in Wisconsin. References: (1) K. Deahl. (Abstr.) Phytopathology 100:S161, 2010. (2) S. B. Goodwin et al. Curr. Genet. 22:107, 1992. (3) Hu et al. Plant Dis. 96:1323, 2012. (4) A. C. Seidl and A. J. Gevens. (Abstr.) Phytopathology 101:S162, 2011.

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Arabidopsis late blight: Infection of a nonhost plant by Albugo laibachii enables full colonization by Phytophthora infestans

10.1101/035006 ◽

2015 ◽

Cited By ~ 2

Author(s):

Khaoula Belhaj ◽

Liliana M. Cano ◽

David C. Prince ◽

Ariane Kemen ◽

Kentaro Yoshida ◽

...

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Molecular Mechanisms ◽

Transcriptional Profiling ◽

Unexpected Finding ◽

Potato Plants ◽

Nonhost Plant ◽

Gene Expression Dynamics ◽

Oomycete Pathogen ◽

Arabidopsis Cells

AbstractThe oomycete pathogen Phytophthora infestans causes potato late blight, and as a potato and tomato specialist pathogen, is seemingly poorly adapted to infect plants outside the Solanaceae. Here, we report the unexpected finding that P. infestans can infect Arabidopsis thaliana when another oomycete pathogen, Albugo laibachii, has colonized the host plant. The behaviour and speed of P. infestans infection in Arabidopsis pre-infected with A. laibachii resemble P. infestans infection of susceptible potato plants. Transcriptional profiling of P. infestans genes during infection revealed a significant overlap in the sets of secreted-protein genes that are induced in P. infestans upon colonisation of potato and susceptible Arabidopsis, suggesting major similarities in P. infestans gene expression dynamics on the two plant species. Furthermore, we found haustoria of A. laibachii and P. infestans within the same Arabidopsis cells. This Arabidopsis - A. laibachii - P. infestans tripartite interaction opens up various possibilities to dissect the molecular mechanisms of P. infestans infection and the processes occurring in co-infected Arabidopsis cells.

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First Report of Late Blight Caused by Phytophthora infestans on Cape Gooseberry (Physalis peruviana) in Colombia

Plant Disease ◽

10.1094/pdis-91-4-0464b ◽

2007 ◽

Vol 91 (4) ◽

pp. 464-464 ◽

Cited By ~ 5

Author(s):

A. M. Vargas ◽

A. Correa ◽

D. C. Lozano ◽

A. González ◽

A. J. Bernal ◽

...

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Morphological Characteristics ◽

Control Measures ◽

Distilled Water ◽

First Report ◽

Physalis Peruviana ◽

Potato Slice ◽

Solanaceae Family ◽

Cape Gooseberry

Late blight caused by Phytophthora infestans is the most limiting disease for several species of the Solanaceae family in Colombia. A potential host for P. infestans is Cape gooseberry (Physalis peruviana), a species belonging to the Solanaceae family. Its center of origin is the highlands of Peru and it is grown at approximately 1,500 to 3,000 m above sea level. Cape gooseberry has become an important export fruit in Colombia. Consequently, in the last few years, the area cultivated with Physalis peruviana has increased dramatically. P. infestans was isolated from this crop in the province of Cundinamarca, Colombia. Symptoms caused by this oomycete appeared initially on the leaf margins as small, irregular, necrotic spots that expanded and merged, increasing the necrotic area. These spots had a soft texture resulting from the degradation of plant tissue by the pathogen. On old lesions, white mycelia and sporangia were observed. Affected plants were rarely killed, but under favorable conditions, severe symptoms were observed in leaves and yield was reduced. Ten isolates were obtained from infected tissue by placing a lesion directly on a potato slice in a moist chamber (2). Mycelia grown on the potato slice were then transferred to rye agar. Identification of the pathogen was performed based on morphological characteristics, specifically, sporangiophores of P. infestans are compoundly branched and develop sympodially, with swellings at the points where sporangia were attached (1). Further confirmation was obtained by sequencing the internal transcribed spacer (ITS) regions (GenBank Accession Nos. EF173467-EF173476). Koch's postulates were completed in the laboratory by spray inoculating detached leaves of Cape gooseberry with a zoospore suspension obtained from each of the 10 isolates. Inoculum was prepared by flooding 10-day-old cultures with sterile distilled water to obtain a 104/ml sporangial suspension followed by zoospore induction at 4°C. Leaves were sprayed with this suspension, placed in moist chambers, and incubated at 20°C in the dark. Control leaves were sprayed with sterile distilled water. Two separate leaves were inoculated with each isolate. The pathogen was reisolated from leaf lesions in all cases. The period between infection and the appearance of symptoms ranged from 5 to 7 days. To our knowledge, this is the first report of P. infestans causing damage on Cape gooseberry in Colombia. Chemical control measures are to some extent successfully applied in most regions where solanaceous crops are grown in Colombia. Nevertheless, suitable disease management for Physalis peruviana has not been achieved and further studies on the epidemiology of the disease on this new host are needed. References: (1) D. C. Erwin and O. K. Ribeiro. Phytophthora Diseases Worldwide. The American Phytopathological Society. St. Paul, MN, 1996. (2) G. A. Forbes et al. Phytopathology 87:375, 1997.

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First Report of Phytophthora infestans on Petunia × hybrida in South Africa

Plant Disease ◽

10.1094/pd-90-1550b ◽

2006 ◽

Vol 90 (12) ◽

pp. 1550-1550

Author(s):

A. McLeod ◽

S. Coertze

Keyword(s):

South Africa ◽

Plant Pathology ◽

Late Blight ◽

Phytophthora Infestans ◽

Petunia Hybrida ◽

Plant Diseases ◽

Distilled Water ◽

Western Cape ◽

Western Cape Province ◽

First Report

Phytophthora infestans is known worldwide as the destructive, late blight pathogen of potatoes and tomatoes. However, erratic reports dating back to 1856 also have shown it to be pathogenic on petunia (Petunia × hybrida), although it has not been regarded as an important pathogen on this host (4). Recently, reports from North America showed that P. infestans is of commercial importance in greenhouse-grown petunias (2), and that late blight-infected petunias may serve as inoculum to tomatoes growing in the same greenhouse (1,2). In the Western Cape Province of South Africa, two petunia samples were received at the Stellenbosch University Plant Disease Clinic in 2005 that showed symptoms resembling P. infestans infections. The two samples were from nurseries where petunias were either grown under shading nets or in a greenhouse. In the greenhouse-grown petunias, the presumptive late blight infections resulted in substantial losses to the grower. Symptoms included gray, slightly sunken leaf lesions with white sporulation mainly on the adaxial side of the leaves. Leaflets of the petunias were incubated in moist chambers, and sporangia sporulating from lesions were identified morphologically as being P. infestans (3). Subsequently, one isolate was cultured onto synthetic media by carefully transferring sporangia from a lesion with the tip of a bended glass rod onto wheat medium (120 g/L of crushed wheat seeds, blended, boiled and filtered through cheesecloth, plus 15 g of sucrose and agar). The identity of the culture was further confirmed through sequence analyses of the internal transcribed spacer regions (GenBank Accession No. DQ479409). The isolated P. infestans strain (STE-U 6134) has been submitted to the Stellenbosch University culture collection. Inoculum for the pathogenicity tests was produced by first flooding 14-day-old cultures with sterile distilled water to obtain a 2 × 104/ml sporangial suspension, followed by zoospore induction at 4°C. A mixture of petunia cultivars (n = 24) were spray inoculated to runoff with the zoospore induced sporangial suspension. Control plants were sprayed with sterile distilled water. Inoculated plants were incubated at 22 to 25°C and high relative humidity (≥93%) within perspex humidity chambers (60 × 30 × 60 cm) lined with a wet sheet of chromatography paper. The experiment was repeated twice. The first late blight symptoms similar to those of the submitted samples appeared 5 to 7 days after inoculation, with some lesions containing profuse white mycelia and sporangiophores typical for P. infestans. The pathogen was reisolated from the leaf lesions, completing Koch's postulate. To our knowledge, this is the first report of P. infestans causing damage on petunias in South Africa. Future studies should be aimed at investigating whether late blight-infected petunias provide an important source of inoculum for potatoes and tomatoes, which are widely grown in the Western Cape Province of South Africa. References: (1) M. C. Becktell et al. Plant Dis. 89:1000, 2005. (2) K. L. Deahl and D. K. Farel. Plant Dis. 87:1004, 2003. (3) D. C. Erwin and O. K. Ribeiro, Phytophthora Diseases Worldwide. The American Phytopathological Society St. Paul, MN, 1996. (4) J. M. Hirst and W. C. Moore. Phytophthora infestans on petunia and datura. Page 76 in: Plant Pathology-A Record of Current Work on Plant Diseases and Pests. Vol. 6. Ministry of Agriculture, Fisheries and Food Plant Pathology Laboratory, Harpenden, England, 1957.

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Phytophthora infestans Produces Oospores in Fruits and Seeds of Tomato

Phytopathology ◽

10.1094/phyto.2001.91.11.1074 ◽

2001 ◽

Vol 91 (11) ◽

pp. 1074-1080 ◽

Cited By ~ 15

Author(s):

Evgenia Rubin ◽

Alexander Baider ◽

Yigal Cohen

Keyword(s):

Late Blight ◽

Phytophthora Infestans ◽

Seed Coat ◽

Growing Season ◽

Tomato Fruits ◽

First Report ◽

Tomato Seeds ◽

Saturated Atmosphere ◽

Causal Fungus ◽

Green Stage

Tomato fruits at the mature green stage coinoculated with A1 + A2 sporangia of Phytophthora infestans, the late blight causal fungus, showed abundant oospores in the vascular tissues, pericarp, columella, and placenta. Oospores were also formed on the surface of fruits kept in moisture-saturated atmosphere. Occasionally, oospores were enclosed between the epidermal hairs of the seed coat. In a few seeds, oospores were detected inside the embryo. The data suggest that blighted tomato fruits may carry a large number of oospores, thus making them a threatening source of blight inoculum. Such fruits may also release airborne oosporic inoculum that may introduce recombinant genotypes within a growing season. Although Phytophthora infestans is seedborne in tomato, to our knowledge, this is the first report on the occurrence of oospores in tomato seeds. Whether such tomato seeds produce blighted seedlings remains to be shown.

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Migration and Selection Enforced Multiple Phenotypic and Genotypic Changes in the Population of Phytophthora infestans in Israel during the Last 36-Year Period

10.20944/preprints202001.0321.v1 ◽

2020 ◽

Author(s):

Yigal Cohen ◽

Avia E. Rubin ◽

Mariana Galperin ◽

Esti Shamaba ◽

Uri Zig ◽

...

Keyword(s):

Population Structure ◽

Late Blight ◽

Phytophthora Infestans ◽

Phenylamide Fungicides ◽

Selection Processes ◽

Heterogeneous Nature ◽

Oomycete Pathogen ◽

Single Field ◽

One Year ◽

Genotypic Structure

Late blight caused by the oomycete pathogen Phytophthora infestans is a devastating disease of potato and tomato worldwide, including Israel. The population structure of this pathogen was monitored in potato and tomato fields in Israel during a 36-year period of 1983-2019. Isolates of the pathogen were tested for sensitivity to phenylamide fungicides, mating type, race structure, and genotype. The phenotypic and genotypic structure of the population from potato have changed greatly from one year to another, from one season to the next, within a season and within a single field. Major changes also occurred in the population collected from tomato crops. The mechanisms driving these multiple changes and the heterogeneous nature of the population in Israel are shown to derive from multiple migration events of the pathogen via seed tubers from Europe and from fitness-driven selection processes.

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