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Vincent Bulone - One of the best experts on this subject based on the ideXlab platform.
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Identification of Growth Inhibitors of the Fish Pathogen Saprolegnia parasitica Using in silico Subtractive Proteomics, Computational Modeling, and Biochemical Validation
Frontiers in microbiology, 2020Co-Authors: Sanjiv Kumar, Vincent Bulone, Rahul Shubhra Mandal, Vaibhav SrivastavaAbstract:Many Stramenopile species belonging to oomycetes from the genus Saprolegnia infects fish, amphibians and crustaceans in aquaculture farms and natural ecosystems. Saprolegnia parasitica is one of the most severe fish pathogens, responsible for high losses in the aquaculture industry worldwide. Most of the molecules reported to date for the control of Saprolegnia infections either are inefficient or have negative impacts on the health of the fish hosts or the environment resulting in substantial economic losses. Until now, the whole proteome of S. parasitica has not been explored for a systematic screening of novel inhibitors against the pathogen. The present study was designed to develop a consensus computational framework for the identification of potential target proteins, their inhibitors and subsequent experimental validation of selected compounds. Comparative analysis between the proteomes of Saprolegnia , humans and fish species identified proteins that are specific and essential for the survival of the pathogen. The DrugBank database was exploited to select FDA-approved inhibitors whose high binding affinity to their respective protein targets was confirmed by computational modelling. At least six of the identified compounds significantly inhibited the growth of Saprolegnia parasitica in-vitro. Triclosan was found to be most effective with a minimum inhibitory concentration of 4 μg/mL. Optical microscopy showed that the inhibitors affect the morphology of hyphal cells, with hyper-branching being commonly observed. The inhibitory effects of the compounds identified in this study on Saprolegnia ’s mycelial growth indicate that they are potentially usable for disease control against this class of oomycete pathogens. Similar approach can be easily adopted for the identification of potential inhibitors against other plant and animal pathogenic oomycete infections.
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Identification and Characterization of the Chitin Synthase Genes From the Fish Pathogen Saprolegnia parasitica.
Frontiers in microbiology, 2019Co-Authors: Elzbieta Rzeszutek, Sara M. Díaz-moreno, Vincent BuloneAbstract:Saprolegnia parasitica is a pathogenic oomycete responsible for severe fish infections. Despite its low abundance in the cell wall of S. parasitica, chitin is essential for hyphal growth as the inhibition of its biosynthesis leads to highly reduced growth. Here we identified and characterized chitin synthases (CHS) from S. parasitica as potential targets for anti-oomycete drugs. Bioinformatics analyses allowed the identification of six different putative Chs genes in the genome of the pathogen. The total number of genes was confirmed by Southern blot analysis and their expression levels were determined by quantitative PCR. Four of the six Chs genes were expressed in the mycelium, while the two others exhibited undetectable levels of expression. The mycelium was highly sensitive to the addition of nikkomycin Z (NZ) in the culture medium, which led to a decreased amount of chitin in the cell wall by up to 40% in the conditions tested, and to the formation of abnormal branching structures in the hyphae. The presence of NZ increased the expression level of one of the genes, Chs3, suggesting that the corresponding product is compensating the disruption of chitin biosynthesis in the hyphae. In addition, the activity of isolated CHS was strongly inhibited by NZ in vitro. Altogether our data indicate the importance of CHS for the vegetative growth of S. parasitica and demonstrate that these enzymes represent promising targets for the control of diseases caused by oomycetes.
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Novel insights into chitin biosynthesis through heterologous expression and biochemical characterization of chitin synthase 5 from the pathogenic oomycete Saprolegnia parasitica
2019Co-Authors: Elzbieta Rzeszutek, Vaibhav Srivastava, Sara M. Díaz-moreno, Osei Y. Ampomah, Annie Inman, Qi Zhou, Vincent BuloneAbstract:Novel insights into chitin biosynthesis through heterologous expression and biochemical characterization of chitin synthase 5 from the pathogenic oomycete Saprolegnia parasitica
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Analysis of the cellulose synthase genes in the oomycete fish pathogen Saprolegnia parasitica and effect of cellulose biosynthesis inhibitors on enzyme activity and microbial growth
2019Co-Authors: Elzbieta Rzeszutek, Sara M. Díaz-moreno, Stefan Klinter, Vincent BuloneAbstract:Analysis of the cellulose synthase genes in the oomycete fish pathogen Saprolegnia parasitica and effect of cellulose biosynthesis inhibitors on enzyme activity and microbial growth
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Quantitative proteomic analysis of four developmental stages of Saprolegnia parasitica
Frontiers in microbiology, 2018Co-Authors: Vaibhav Srivastava, Svetlana Rezinciuc, Vincent BuloneAbstract:Several water mold species from the Saprolegnia genus infect fish, amphibians, and crustaceans in natural ecosystems and aquaculture farms. Saprolegnia parasitica is one of the most severe fish pat ...
Pieter Van West - One of the best experts on this subject based on the ideXlab platform.
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Specialized attachment structure of the fish pathogenic oomycete Saprolegnia parasitica
PloS one, 2018Co-Authors: Svetlana Rezinciuc, Jose Vladimir Sandoval-sierra, Yolanda Ruiz-león, Pieter Van West, Javier Diéguez-uribeondoAbstract:The secondary cysts of the fish pathogen oomycete Saprolegnia parasitica possess bundles of long hooked hairs that are characteristic to this economically important pathogenic species. Few studies have been carried out on elucidating their specific role in the S. parasitica life cycle and the role they may have in the infection process. We show here their function by employing several strategies that focus on descriptive, developmental and predictive approaches. The strength of attachment of the secondary cysts of this pathogen was compared to other closely related species where bundles of long hooked hairs are absent. We found that the attachment of the S. parasitica cysts was around three times stronger than that of other species. The time sequence and influence of selected factors on morphology and the number of the bundles of long hooked hairs conducted by scanning electron microscopy study revealed that these are dynamic structures. They are deployed early after encystment, i.e., within 30 sec of zoospore encystment, and the length, but not the number, of the bundles steadily increased over the encystment period. We also observed that the number and length of the bundles was influenced by the type of substrate and encystment treatment applied, suggesting that these structures can adapt to different substrates (glass or fish scales) and can be modulated by different signals (i.e., protein media, 50 mM CaCl2 concentrations, carbon particles). Immunolocalization studies evidenced the presence of an adhesive extracellular matrix. The bioinformatic analyses of the S. parasitica secreted proteins showed that there is a high expression of genes encoding domains of putative proteins related to the attachment process and cell adhesion (fibronectin and thrombospondin) coinciding with the deployment stage of the bundles of long hooked hairs formation. This suggests that the bundles are structures that might contribute to the adhesion of the cysts to the host because they are composed of these adhesive proteins and/or by increasing the surface of attachment of this extracellular matrix.
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List of selected secreted proteins of Saprolegnia parasitica with their description and expression levels in different life stages.
2018Co-Authors: Svetlana Rezinciuc, Jose Vladimir Sandoval-sierra, Yolanda Ruiz-león, Pieter Van West, Javier Diéguez-uribeondoAbstract:List of selected secreted proteins of Saprolegnia parasitica with their description and expression levels in different life stages.
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A putative serine protease, SpSsp1, from Saprolegnia parasitica is recognised by sera of rainbow trout, Oncorhynchus mykiss
Fungal biology, 2014Co-Authors: Kirsty L. Minor, Albert Hendrik Van Den Berg, Stephan Wawra, Victoria L. Anderson, Christopher J. Secombes, Lars Löbach, Katie S. Davis, James S. Christie, Ali Reza Faruk, Pieter Van WestAbstract:Saprolegniosis, the disease caused by Saprolegnia sp., results in considerable economic losses in aquaculture. Current control methods are inadequate, as they are either largely ineffective or present environmental and fish health concerns. Vaccination of fish presents an attractive alternative to these control methods. Therefore we set out to identify suitable antigens that could help generate a fish vaccine against Saprolegnia parasitica. Unexpectedly, antibodies against S. parasitica were found in serum from healthy rainbow trout, Oncorhynchus mykiss. The antibodies detected a single band in secreted proteins that were run on a one-dimensional SDS-polyacrylamide gel, which corresponded to two protein spots on a two-dimensional gel. The proteins were analysed by liquid chromatography tandem mass spectrometry. Mascot and bioinformatic analysis resulted in the identification of a single secreted protein, SpSsp1, of 481 amino acid residues, containing a subtilisin domain. Expression analysis demonstrated that SpSsp1 is highly expressed in all tested mycelial stages of S. parasitica. Investigation of other non-infected trout from several fish farms in the United Kingdom showed similar activity in their sera towards SpSsp1. Several fish that had no visible saprolegniosis showed an antibody response towards SpSsp1 suggesting that SpSsp1 might be a useful candidate for future vaccination trial experiments.
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Functional characterization of a tyrosinase gene from the oomycete Saprolegnia parasitica by RNAi silencing.
Fungal biology, 2014Co-Authors: Marcia Saraiva, Vincent Bulone, Debbie Mclaggan, Irene De Bruijn, Laura J. Grenville-briggs, Ariane Willems, Pieter Van WestAbstract:Here we describe the first application of transient gene silencing in Saprolegnia parasitica, a pathogenic oomycete that infects a wide range of fish, amphibians, and crustaceans. A gene encoding a putative tyrosinase from S. parasitica, SpTyr, was selected to investigate the suitability of RNA-interference (RNAi) to functionally characterize genes of this economically important pathogen. Tyrosinase is a mono-oxygenase enzyme that catalyses the O-hydroxylation of monophenols and subsequent oxidation of O-diphenols to quinines. These enzymes are widely distributed in nature, and are involved in the melanin biosynthesis. Gene silencing was obtained by delivering in vitro synthesized SpTyr dsRNA into protoplasts. Expression analysis, tyrosinase activity measurements, and melanin content analysis confirmed silencing in individual lines. Silencing of SpTyr resulted in a decrease of tyrosinase activity between 38 % and 60 %, dependent on the level of SpTyr-expression achieved. The SpTyr-silenced lines displayed less pigmentation in developing sporangia and occasionally an altered morphology. Moreover, developing sporangia from individual silenced lines possessed a less electron dense cell wall when compared to control lines, treated with GFP-dsRNA. In conclusion, the tyrosinase gene of S. parasitica is required for melanin formation and transient gene silencing can be used to functionally characterize genes in S. parasitica.
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The impact of the water moulds Saprolegnia diclina and Saprolegnia parasitica on natural ecosystems and the aquaculture industry
Fungal Biology Reviews, 2013Co-Authors: Albert Hendrik Van Den Berg, Javier Diéguez-uribeondo, Debbie Mclaggan, Pieter Van WestAbstract:Abstract A very large number of diseases in agriculture and aquaculture are caused by fungal-like micro-organisms, classified as Oomycetes or water moulds. Collectively, oomycetes represent a huge threat to global food security. In addition, several oomycetes can cause environmental disasters, by wiping out native species of trees, crustacians and amphibians. The group representing the aquatic oomycetes are relatively understudied in contrast to their terrestrial counterparts that predominantly infect plants (e.g. Phytophthora, Pythium and Bremia spp.). This review focuses on the unique characteristics of two aquatic Oomycetes, Saprolegnia parasitica and Saprolegnia diclina with respect to their impact on aquaculture, animal health and the surrounding environment. The species characteristics, ecology, biology, infectivity and identification methods are described and the latest research insights are discussed.
Ida Skaar - One of the best experts on this subject based on the ideXlab platform.
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In vitro passages impact on virulence of Saprolegnia parasitica to Atlantic salmon, Salmo salar L. parr
Journal of fish diseases, 2013Co-Authors: M M Songe, E Thoen, Øystein Evensen, Ida SkaarAbstract:The effect of serial in vitro subculturing on three pathogenic strains of Saprolegnia parasitica was investigated. The isolates were passed through Atlantic salmon, Salmo salar L. parr, and then re-isolated as single spore colonies. All strains caused infection. The isolate obtained from diseased fish served as a virulent reference culture and was designated ‘AP’ (‘activated through passage’). Successive subculturing was made by obtaining an inoculum from AP to produce the 2nd subculture and then passaged to the 3rd subculture (from the 2nd), until the 15th passage was obtained. Spores used to produce storage cultures were collected at passages 5, 10 and 15. The different passages of each strain were used to artificially infect Atlantic salmon parr. Morphological characterization of growth patterns was performed to observe differences occurring due to serial in vitro subculturing. Two of the strains declined in virulence after 15 successive in vitro subcultures, whereas one did not. This study is the first to investigate attenuation of virulence in Saprolegnia and whether or not isolates of S. parasitica should be passed through the fish host prior to challenge experiments. It reveals that some strains degenerate more rapidly than others when subjected to successive in vitro subculturing on glucose–yeast extract.
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A simple in vitro screening method to determine the effects of drugs against growth of Saprolegnia parasitica
Mycological Progress, 2005Co-Authors: Svein Stueland, Berit Tafjord Heier, Ida SkaarAbstract:To screen the effect of possible antifungal chemicals against growth of fish pathogenic Saprolegnia spp., a simple and rapid in vitro screening method has been developed. Heat sterilized hemp seeds ( Cannabis sativa ) colonized by Saprolegnia parasitica are exposed to different concentrations of the test drugs diluted in water. One Saprolegnia colonized hemp seed is transferred into each well of a 48-well flat bottom tissue culture plate, after which 1 mL of each concentration of the test drugs is added per well. Subsequent to exposure and incubation, the plate is inspected and any Saprolegnia growth on the seeds is graded. The method described in the present paper proved to be simple, effective and reproducible in screening of fungistatic and fungicidal drugs against Saprolegnia growth.
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A simple in vitro screening method to determine the effects of drugs against growth of Saprolegnia parasitica
Mycological Progress, 2005Co-Authors: Svein Stueland, Berit T. Heier, Ida SkaarAbstract:To screen the effect of possible antifungal chemicals against growth of fish pathogenic Saprolegnia spp., a simple and rapid in vitro screening method has been developed. Heat sterilized hemp seeds (Cannabis sativa) colonized by Saprolegnia parasitica are exposed to different concentrations of the test drugs diluted in water. One Saprolegnia colonized hemp seed is transferred into each well of a 48-well flat bottom tissue culture plate, after which 1 mL of each concentration of the test drugs is added per well. Subsequent to exposure and incubation, the plate is inspected and any Saprolegnia growth on the seeds is graded.
Javier Diéguez-uribeondo - One of the best experts on this subject based on the ideXlab platform.
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Specialized attachment structure of the fish pathogenic oomycete Saprolegnia parasitica
PloS one, 2018Co-Authors: Svetlana Rezinciuc, Jose Vladimir Sandoval-sierra, Yolanda Ruiz-león, Pieter Van West, Javier Diéguez-uribeondoAbstract:The secondary cysts of the fish pathogen oomycete Saprolegnia parasitica possess bundles of long hooked hairs that are characteristic to this economically important pathogenic species. Few studies have been carried out on elucidating their specific role in the S. parasitica life cycle and the role they may have in the infection process. We show here their function by employing several strategies that focus on descriptive, developmental and predictive approaches. The strength of attachment of the secondary cysts of this pathogen was compared to other closely related species where bundles of long hooked hairs are absent. We found that the attachment of the S. parasitica cysts was around three times stronger than that of other species. The time sequence and influence of selected factors on morphology and the number of the bundles of long hooked hairs conducted by scanning electron microscopy study revealed that these are dynamic structures. They are deployed early after encystment, i.e., within 30 sec of zoospore encystment, and the length, but not the number, of the bundles steadily increased over the encystment period. We also observed that the number and length of the bundles was influenced by the type of substrate and encystment treatment applied, suggesting that these structures can adapt to different substrates (glass or fish scales) and can be modulated by different signals (i.e., protein media, 50 mM CaCl2 concentrations, carbon particles). Immunolocalization studies evidenced the presence of an adhesive extracellular matrix. The bioinformatic analyses of the S. parasitica secreted proteins showed that there is a high expression of genes encoding domains of putative proteins related to the attachment process and cell adhesion (fibronectin and thrombospondin) coinciding with the deployment stage of the bundles of long hooked hairs formation. This suggests that the bundles are structures that might contribute to the adhesion of the cysts to the host because they are composed of these adhesive proteins and/or by increasing the surface of attachment of this extracellular matrix.
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List of selected secreted proteins of Saprolegnia parasitica with their description and expression levels in different life stages.
2018Co-Authors: Svetlana Rezinciuc, Jose Vladimir Sandoval-sierra, Yolanda Ruiz-león, Pieter Van West, Javier Diéguez-uribeondoAbstract:List of selected secreted proteins of Saprolegnia parasitica with their description and expression levels in different life stages.
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The impact of the water moulds Saprolegnia diclina and Saprolegnia parasitica on natural ecosystems and the aquaculture industry
Fungal Biology Reviews, 2013Co-Authors: Albert Hendrik Van Den Berg, Javier Diéguez-uribeondo, Debbie Mclaggan, Pieter Van WestAbstract:Abstract A very large number of diseases in agriculture and aquaculture are caused by fungal-like micro-organisms, classified as Oomycetes or water moulds. Collectively, oomycetes represent a huge threat to global food security. In addition, several oomycetes can cause environmental disasters, by wiping out native species of trees, crustacians and amphibians. The group representing the aquatic oomycetes are relatively understudied in contrast to their terrestrial counterparts that predominantly infect plants (e.g. Phytophthora, Pythium and Bremia spp.). This review focuses on the unique characteristics of two aquatic Oomycetes, Saprolegnia parasitica and Saprolegnia diclina with respect to their impact on aquaculture, animal health and the surrounding environment. The species characteristics, ecology, biology, infectivity and identification methods are described and the latest research insights are discussed.
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Re-evaluation of the enigmatic species complex Saprolegnia diclina-Saprolegnia parasitica based on morphological, physiological and molecular data.
Fungal genetics and biology : FG & B, 2007Co-Authors: Javier Diéguez-uribeondo, Lage Cerenius, Kenneth Söderhäll, J.m. Fregeneda-grandes, J. M. Aller-gancedo, Elena Pérez-iniesta, M. Teresa Telleria, María P. MartínAbstract:The phylogenetic relationships among isolates of the Saprolegnia diclina-Saprolegnia parasitica complex were investigated based on ITS rDNA sequences, and correlated with morphological and physiological characters. The isolates studied belong to five phylogenetically separate clades. The majority of presumed parasitic isolates, mostly isolated from fish lesions, fell within a clade that comprises isolates which has been variously named as S. diclina Type 1, S. parasitica, Saprolegnia salmonis or just as unnamed Saprolegnia sp. Presence of bundles of long-hooked hairs on secondary cysts, high frequency of retracted germination, and oogonia production at 7 degrees C (when occurring) were characteristic of this clade. A single isolate identified as S. diclina Type 2 clustered in a clade along with Saprolegnia ferax isolates. The isolates identified as S. diclina s. str. (S. diclina Type 3) distributed in two clades and appeared closely related to Saprolegnia multispora and to a number of Chilean isolates identified as Saprolegnia australis. The ITS sequences of clade I were almost identical even though the isolates were of diverse geographical origins and showed physiological and morphological differences and variations in their pathogenicity. This suggest these species reproduces clonally even in apparently sexually competent isolates. Adaptation to parasitism in Saprolegnia might have occurred at spore level by the development of long-hooked hairs to facilitate host attachment and selection of a retracting germination. The use of the name S. parasitica should be assigned to isolates of clade I that contained isolates forming cysts with bundles of long-hooked hairs.
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Repeated zoospore emergence in Saprolegnia parasitica
Mycological Research, 1994Co-Authors: Javier Diéguez-uribeondo, Lage Cerenius, Kenneth SöderhällAbstract:A method of determining the number of phases of repeated zoospore emergence in vitro in the water mould Saprolegnia parasitica was developed. Using vortexing to encyst zoospores it was possible, in a synchronous manner, to achieve up to six consecutive generations of cysts and laterally biflagellated zoospores. It was found that the length of time the zoospores were allowed to swim influenced the number of zoospore generations which could be produced. After zoospore encystment the spores can be triggered to either germinate or release a new zoospore depending upon the addition of external nutrients. Germination of cysts could only be triggered within 45 min of mechanically induced encystment. Thus, it is now possible in this species to study in detail the two alternative developmental pathways which follow after encystment: either repeated zoospore emergence or germ tube formation.
Christopher J. Secombes - One of the best experts on this subject based on the ideXlab platform.
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A putative serine protease, SpSsp1, from Saprolegnia parasitica is recognised by sera of rainbow trout, Oncorhynchus mykiss
Fungal biology, 2014Co-Authors: Kirsty L. Minor, Albert Hendrik Van Den Berg, Stephan Wawra, Victoria L. Anderson, Christopher J. Secombes, Lars Löbach, Katie S. Davis, James S. Christie, Ali Reza Faruk, Pieter Van WestAbstract:Saprolegniosis, the disease caused by Saprolegnia sp., results in considerable economic losses in aquaculture. Current control methods are inadequate, as they are either largely ineffective or present environmental and fish health concerns. Vaccination of fish presents an attractive alternative to these control methods. Therefore we set out to identify suitable antigens that could help generate a fish vaccine against Saprolegnia parasitica. Unexpectedly, antibodies against S. parasitica were found in serum from healthy rainbow trout, Oncorhynchus mykiss. The antibodies detected a single band in secreted proteins that were run on a one-dimensional SDS-polyacrylamide gel, which corresponded to two protein spots on a two-dimensional gel. The proteins were analysed by liquid chromatography tandem mass spectrometry. Mascot and bioinformatic analysis resulted in the identification of a single secreted protein, SpSsp1, of 481 amino acid residues, containing a subtilisin domain. Expression analysis demonstrated that SpSsp1 is highly expressed in all tested mycelial stages of S. parasitica. Investigation of other non-infected trout from several fish farms in the United Kingdom showed similar activity in their sera towards SpSsp1. Several fish that had no visible saprolegniosis showed an antibody response towards SpSsp1 suggesting that SpSsp1 might be a useful candidate for future vaccination trial experiments.
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Immune gene expression in trout cell lines infected with the fish pathogenic oomycete Saprolegnia parasitica
Developmental and comparative immunology, 2012Co-Authors: Irene De Bruijn, Pieter Van West, Marcia Saraiva, Rodrigo Belmonte, Vicky L. Anderson, Tiehui Wang, Christopher J. SecombesAbstract:The oomycete Saprolegnia parasitica causes significant losses in the aquaculture industry, mainly affecting salmon, trout and catfish. Since the ban of malachite green, effective control measures are currently not available prompting a re-evaluation of the potential for immunological intervention. In this study, the immune response of salmonid cells is investigated at the transcript level, by analysis of a large set of immune response genes in four different rainbow trout cell lines (RTG-2, RTGill, RTL and RTS11) upon infection with S. parasitica. Proinflammatory cytokine transcripts were induced in all four cell lines, including IL-1β1, IL-8, IL-11, TNF-α2, as well as other components of the innate defences, including COX-2, the acute phase protein serum amyloid A and C-type lectin CD209a and CD209b. However, differences between the four cell lines were found. For example, the fold change of induction was much higher in the epithelial RTL and macrophage-like RTS11 cell lines compared to the fibroblast cell lines RTG-2 and RTGill. Several antimicrobial peptides (AMPs) were also up-regulated in response to Saprolegnia infection, including hepcidin and cathelicidin 1 (rtCATH1) and 2 (rtCATH2). An rtCATH2 peptide was synthesised and tested for activity and whilst it showed no killing activity for zoospores, it was able to delay sporulation of S. parasitica. These results demonstrate that particular immune genes are up-regulated in response to S. parasitica infection and that AMPs may play a crucial role in the first line of defence against oomycetes in fish.
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The putative RxLR effector protein SpHtp1 from the fish pathogenic oomycete Saprolegnia parasitica is translocated into fish cells
FEMS microbiology letters, 2010Co-Authors: Pieter Van West, Irene De Bruijn, Kirsty L. Minor, Andrew J. Phillips, Emma J. Robertson, Stephan Wawra, Judith M. Bain, Victoria L. Anderson, Christopher J. SecombesAbstract:The fish pathogenic oomycete Saprolegnia parasitica causes the disease Saprolegniosis in salmonids and other freshwater fish, resulting in considerable economic losses in aquaculture. Very little is known about the molecular and cellular mechanisms underlying the infection process of fish pathogenic oomycetes. In order to investigate the interaction in detail, an in vitro infection assay using an Oncorhynchus mykiss (rainbow trout) cell line (RTG-2) was developed. In a zoospore/cyst cDNA library, we identified the ORF SpHtp1, which encodes a secreted protein containing an RxLR motif. Detailed expression analysis indicated that SpHtp1 is highly expressed in zoospores/cysts from S. parasitica and in the very early stages of infection on RTG-2 cells, when compared with in vitro-grown mycelium. Moreover, the protein, SpHtp1, was found to translocate into the RTG-2 trout cells, during the interaction with S. parasitica, and also when the RTG-2 cells were treated with recombinant SpHtp1 fused to a C-terminal His-tag. These findings suggest that protein translocation could play an important role in Saprolegniosis.
