Contingency rules for pathogen competition and antagonism in a genetically based,plant defense hierarchy

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HISTONE DEACETYLASE 6 represses pathogen defence responses in Arabidopsis thaliana

Plant defence mechanisms are suppressed in the absence of pathogen attack to prevent wasted energy and growth inhibition. However, how defence responses are repressed is not well understood. Histone deacetylase 6 (HDA6) is a negative regulator of gene expression, and its role in pathogen defence response in plants is not known. In this study, a novel allele of hda6 (designated as shi5) with spontaneous defence response was isolated from a forward genetics screening in Arabidopsis. The shi5 mutant exhibited increased resistance to hemibiotrophic bacterial pathogen Pst DC3000, constitutively activated expression of pathogen‐responsive genes including PR1, PR2, etc. and increased histone acetylation levels at the promoters of most tested genes that were upregulated in shi5. In both wild type and shi5 plants, the expression and histone acetylation of these genes were upregulated by pathogen infection. HDA6 was found to bind to the promoters of these genes under both normal growth conditions and pathogen infection. Our research suggests that HDA6 is a general repressor of pathogen defence response and plays important roles in inhibiting and modulating the expression of pathogen‐responsive genes in Arabidopsis.     

Dressed to impress: impact of environmental adaptation on the Candida albicans cell wall

Candida albicans is an opportunistic fungal pathogen of humans causing superficial mucosal infections and life‐threatening systemic disease. The fungal cell wall is the first point of contact between the invading pathogen and the host innate immune system. As a result, the polysaccharides that comprise the cell wall act as pathogen associated molecular patterns, which govern the host–pathogen interaction. The cell wall is dynamic and responsive to changes in the external environment. Therefore, the host environment plays a critical role in regulating the host–pathogen interaction through modulation of the fungal cell wall. This review focuses on how environmental adaptation modulates the cell wall structure and composition, and the subsequent impact this has on the innate immune recognition of C. albicans.     

Evolution and maintenance of microbe‐mediated protection under occasional pathogen infection

Every host is colonized by a variety of microbes, some of which can protect their hosts from pathogen infection. However, pathogen presence naturally varies over time in nature, such as in the case of seasonal epidemics. We experimentally coevolved populations of Caenorhabditis elegans worm hosts with bacteria possessing protective traits (Enterococcus faecalis), in treatments varying the infection frequency with pathogenic Staphylococcus aureus every host generation, alternating host generations, every fifth host generation, or never. We additionally investigated the effect of initial pathogen presence at the formation of the defensive symbiosis. Our results show that enhanced microbe‐mediated protection evolved during host‐protective microbe coevolution when faced with rare infections by a pathogen. Initial pathogen presence had no effect on the evolutionary outcome of microbe‐mediated protection. We also found that protection was only effective at preventing mortality during the time of pathogen infection. Overall, our results suggest that resident microbes can be a form of transgenerational immunity against rare pathogen infection.     

Arbuscular mycorrhiza‐mediated resistance in tomato against Cladosporium fulvum‐induced mould disease

Root colonization with arbuscular mycorrhizal fungi (AMF) enhances plant resistance particularly against soil‐borne pathogenic fungi. In this study, mycorrhizal inoculation with Glomus mosseae (Gm) significantly alleviated tomato mould disease caused by the air‐borne fungal pathogen, Cladosporium fulvum (Cf). The disease index (DI) in local leaves (receiving pathogen inoculation) and systemic leaves (just above the local leaf without pathogen inoculation) was 36.4% and 11.7% in mycorrhizal plants, respectively, whereas DI was 59.6% and 36.4% in the corresponding leaves of AMF non‐inoculated plants, after 50 days of Gm inoculation, corresponding to 15 days after Cf inoculation by leaf infiltration. Foliar spray inoculation with Cf also revealed that AMF pre‐inoculated plants had a higher resistance against subsequent pathogen infection, where the DI was 41.3% in mycorrhizal plants vs. 64.4% in AMF non‐inoculated plants. AMF‐inoculated plants showed significantly higher fresh and dry weight than non‐inoculated plants under both control (without pathogen) and pathogen treatments. AMF‐inoculated plants exhibited significant increases in activities of superoxide dismutase and peroxidase, along with decreases in levels of H₂O₂ and malondialdehyde, compared with non‐inoculated plants after pathogen inoculation. AMF inoculation led to increases in total chlorophyll contents and net photosynthesis rate as compared with non‐inoculated plants under control and pathogen infection. Pathogen infection on AMF non‐inoculated plants led to decreases in chlorophyll fluorescence parameters. However, pathogen infection did not affect these parameters in mycorrhizal plants. Taken together, these results indicate that AMF colonization may play an important role in plant resistance against air‐borne pathogen infection by maintaining redox poise and photosynthetic activity.     

Genetic transformation of Rangpur lime (Citrus limonia osbeck) with thebO (bacterio-opsin) genen and its initial evaluation forPhytophthora nicotianae resistance

Transgenic plants expressing the bacterio-opsin (bO) gene can spontaneously activate programmed cell death (ped) and may enhance broad-spectrum pathogen resistance by activating an intrinsic defense pathway in plant species such as tobacco and potato. In this work, we produced transgenic Rangpur lime plants with thebO gene, viaAgrobacterium tumefaciens-mediated transformation, and evaluated these plants forPhytophthora nicotianae resistance. Two transgenic lines were successfully regenerated and transformation was confirmed by GUS activity assay, PCR analysis, Southern, Northern and Western blot analyses, in addition to detecting the expressed bO protein by an immunological approach. Evaluation forPhytophthora nicotianae resistance was carried out by plant inoculations with the pathogen and quantification of the affected area. One of the two transgenic lines showed greater tolerance to the fungal pathogen as compared to the control, with significantly smaller stem lesions after pathogen challenge. This increase in pathogen tolerance is correlated with a significantly higher level of transgene expression in this line when compared with the other transgenic line. This is the first report of the introduction of a potentially important gene into Rangpur lime to provide novel pathogen tolerance.     

Incidence of “Candidatus Liberibacter asiaticus” in a Florida population of Asian citrus psyllid

The incidence of a bacterium “Candidatus Liberibacter asiaticus” was assessed in a Florida population of Asian citrus psyllid, Diaphorina citri. The bacterium is the presumed causal agent of Asiatic huanglongbing, a serious citrus disease. Adult D. citri were periodically collected between May 2010 and September 2012 in a block of diseased trees located in east‐central Florida. The psyllids were individually subjected to molecular analyses (quantitative polymerase chain reaction assays using HLBaspr primers) to determine whether the bacterium was present and, if so, the population level of the pathogen based on qPCR cycle threshold (CT) values. Significantly greater percentages of females tested positive for the pathogen than males, but there were no significant differences between females and males with respect to population levels of the pathogen within the psyllids. No significant differences were found among the three D. citri colour morphs with respect to percentages of adults testing positive for the pathogen. Among 47 sample dates, a mean of 17.5% of adults per sample date tested positive (CT < 36) for the pathogen with a mean CT value of 31.1. The incidence of the pathogen was generally higher during late fall or early winter and often lower during mid‐ to late summer. There was a significant negative correlation between percentages of D. citri testing positive and air temperature. Increases in the incidence of the pathogen may not necessarily correspond to increases in transmission, as a number of factors both internal and external to D. citri can influence transmission. Transmission rates may be highest during periods when D. citri infestation levels are large, a high percentage of adults carry a high population of the pathogen in their salivary glands, and citrus flush is abundant.     

中华蜜蜂的欧洲幼虫腐臭病病原研究

采用中华蜜蜂Apis cerana cerana F.临床自然发病的欧洲幼虫腐臭病(European Foulbrood, EFB)幼虫,对其病原体进行了分离鉴定。结果表明：中华蜜蜂EFB的病原系蜂房蜜蜂球菌Melissococcus pluton。该菌为革兰氏阳性的兼性厌氧菌,形态学及染色特性、致病性试验、血清学试验和细菌DNA G+C mol%试验均证明中华蜜蜂和西方蜜蜂的EFB病原属于同属的蜂房蜜蜂球菌。生理生化特性结果与国外的早期研究结果相近似。该试验为不同地理位置、不同种寄主间蜂房蜜蜂球菌的遗传差异的研究,以及中华蜜蜂EFB病的防治学研究打下基础。     

Galapagos ground finches balance investment in behavioural and immunological pathogen defences

The scientific community has long recognized the importance of individual behaviour in pathogen spread in wild animals. Furthermore, recent studies have highlighted the potential for behaviour to act as a pathogen defence mechanism. In line with these studies, the pathogen defence optimization hypothesis (PDOH) posits that individuals balance investment in costly behavioural and immunological defences against pathogen infection. Here, I test the PDOH using data from observations of wild Galapagos finches (Medium Ground Finches Geospiza fortis and Small Ground Finches Geospiza fuliginosa) at feeders, combined with immune data collected in the field. Both within and between species, those groups engaging to a greater extent in high transmission‐risk behaviours (not investing in behavioural defences against pathogen exposure) invest more heavily in immune pathogen defences. These data provide the first support for the PDOH in a wild population.     

Mixed infections alter transmission potential in a fungal plant pathogen

Infections by more than one strain of a pathogen predominate under natural conditions. Mixed infections can have significant, though often unpredictable, consequences for overall virulence, pathogen transmission and evolution. However, effects of mixed infection on disease development in plants often remain unclear and the critical factors that determine the outcome of mixed infections remain unknown. The fungus Zymoseptoria tritici forms genetically diverse infections in wheat fields. Here, for a range of pathogen traits, we experimentally decompose the infection process to determine how the outcomes and consequences of mixed infections are mechanistically realized. Different strains of Z. tritici grow in close proximity and compete in the wheat apoplast, resulting in reductions in growth of individual strains and in pathogen reproduction. We observed different outcomes of competition at different stages of the infection. Overall, more virulent strains had higher competitive ability during host colonization, and less virulent strains had higher transmission potential. We showed that within-host competition can have a major effect on infection dynamics and pathogen population structure in a pathogen and host genotype-specific manner. Consequently, mixed infections likely have a major effect on the development of septoria tritici blotch epidemics and the evolution of virulence in Z. tritici.     

Biological Control of Pseudomonas syringae pv. glycinea by Epiphytic Bacteria under Field Conditions

The efficacy of a bacterial strain as a biocontrol agent in the field may be related to the ecological similarity between the biocontrol agent and the target pathogen. Therefore, a number of different Pseudomonas syringae strains were evaluated for their antagonistic activities in vitro (agar-diffusion assay) and in planta (greenhouse assay) against the target pathogen, Pseudomonas syringae pv. glycinea. Six strains of five different pathovars were found to be antagonistic in vitro as well as in planta. The epiphytic fitness of the antagonistic Pseudomonas syringae strain 22d/93 and its two antibiotic-resistant mutants were examined on soybean plants in the fields. After adaptation the parental strain and its mutants had the ability to establish and maintain large epiphytic populations (about 10⁶ cfu/g FW) over the whole growing season after a single spray inoculation. The epiphytic behaviors of the mutants and the parent were not significantly different. The introduced bacteria did not influence the total bacterial population size. When the antagonist was coinoculated with the pathogen, the development of the pathogen was significantly reduced during the whole growing season. When the antagonistic strain was inoculated 4 weeks in advance of the pathogen, this antagonistic effect could be markedly enhanced. The final population size of the pathogen reached just 10⁴ cfu/g FW and was significantly reduced to 0.12% compared to the pathogen alone. This study demonstrates that biological control of foliar pathogens through colonization of the host plants with near isogenic or ecologically similar antagonistical strains seems to be a realistic goal.     

Interaction between sunflower plants and five isolates of Plasmopara halstedii on the level of pathogenicity

The interaction between sunflower plants showing a high level of quantitative resistance and five Plasmopara halstedii (the causal agent of downy mildew) isolates of several races were studied using five single zoosporangium isolates per pathogen isolate. Aggressiveness criteria were analyzed for 25 P. halstedii single zoosporangium isolates. Based on the reaction for the P. halstedii isolates to four sunflower hybrids H1–H4 varying only in their downy mildew resistance genes, there were differences in virulence spectrum in pathogen isolates. Analysis of five single zoosporangium isolates for P. halstedii isolates showed significant variability within pathogen isolate for all aggressiveness criteria but not for all pathogen isolates. The hypothesis explaining the interaction between P. halstedii and its host plant was discussed on the level of pathogenicity.     

Trichoderma longibrachiatum as an antagonist of Botrydiplodia theobromae

The antagonistic potential of Trichoderma longibrachiatum against the pathogen Botrydiplodia theobromae was examined in vitro. Both fungi were paired on 9 cm Petri plates of acidified potato dextrose agar (APDA). Three pairing methods were employed, viz., ‘inoculating antagonist before pathogen’, ‘inoculating pathogen before antagonist’ and ‘simultaneous inoculation of pathogen and antagonist’. Radial growth (cm) of both fungi were later analysed using the GLM Procedure of SAS. Growth inhibition of B. theobromae by T. longibrachiatum in all pairing methods was significantly different from control (P = 0.05, R ² = 0.89). Growth inhibition of the pathogen was best when using ‘inoculation of antagonist before pathogen’. Duration of pairing (DAP) and pairing method are both critical to significant antagonistic impact of T. longibrachiatum on B. theobromae (P > 0.0001). Inhibition mechanism includes competition for space and nutrients. T. longibrachiatum could thus be a promising antagonist of B. theobromae.     

Research advances in the Pyrenophora teres–barley interaction

Pyrenophora teres f. teres and P. teres f. maculata are significant pathogens that cause net blotch of barley. An increased number of loci involved in P. teres resistance or susceptibility responses of barley as well as interacting P. teres virulence effector loci have recently been identified through biparental and association mapping studies of both the pathogen and host. Characterization of the resistance/susceptibility loci in the host and the interacting effector loci in the pathogen will provide a path for targeted gene validation for better-informed release of resistant barley cultivars. This review assembles concise consensus maps for all loci published for both the host and pathogen, providing a useful resource for the community to be used in pathogen characterization and barley breeding for resistance to both forms of P. teres.     

Defining Plant Resistance to Phytophthora cinnamomi: A Standardized Approach to Assessment

Phytophthora cinnamomi is a soil‐borne plant pathogen that causes devastating disease in agricultural and natural systems worldwide. While a small number of species survive infection by the pathogen without producing disease symptoms, the nature of resistance, especially under controlled conditions, remains poorly understood. At present, there are no standardized criteria by which resistance or susceptibility to P. cinnamomi can be assessed, and we have used five parameters consisting of plant fresh weight, root growth, lesion length, relative chlorophyll content of leaves and pathogen colonization of roots to analyse responses to the pathogen. The parameters were tested using two plant species, Zea mays and Lupinus angustifolius, through a time course study of the interactions and resistance and susceptibility defined 7 days after inoculation. A scoring system was devised to enable differentiation of these responses. In the resistant interaction with Z. mays, there was no significant difference in fresh weight, root length and relative chlorophyll content in inoculated compared with control plants. Both lesion size and pathogen colonization of root tissues were limited to the site of inoculation. Following inoculation L. angustifolius showed a significant reduction in plant fresh weight and relative leaf chlorophyll content, cessation of root growth and increased lesion lengths and pathogen colonization. We propose that this technique provides a standardized method for plant–P. cinnamomi interactions that could be widely used to differentiate resistant from susceptible species.     

Geographic range, impact, and parasitism of lepidopteran species associated with the invasive weed Lantana camara in South Africa

Six isolates of Trichoderma were screened for antagonism to Armillaria in tea stem sections buried in the soil. The inability of Armillaria to invade Trichoderma-colonized stem sections and the reduction of its viability in the plant materials following invasion of these by Trichoderma were used as indicators of antagonism. Four isolates of the species Trichoderma harzianum significantly (P<0.001) reduced the incidence of the pathogen in the plant materials. Isolate T4 completely eliminated the pathogen from plant materials in sterile soil and also antagonized two different isolates of the pathogen in nonsterile soil. Application of this T. harzianum isolate to the soil as a wheat bran culture significantly (P<0.001) reduced viability of Armillaria in woody blocks of inoculum. Soil amendment with coffee pulp also reduced the inoculum viability but did not affect the incidence of Trichoderma in the blocks of inoculum. We conclude that the direct application of wheat bran-formulated T. harzianum into soil surrounding woody Armillaria inoculum sources can suppress the pathogen. Further, no organic amendment is needed to enhance development of the antagonist in the soil as a pre-requisite to suppressing the pathogen.     

Fluctuating temperatures alter environmental pathogen transmission in a Daphnia–pathogen system

Environmental conditions are rarely constant, but instead vary spatially and temporally. This variation influences ecological interactions and epidemiological dynamics, yet most experimental studies examine interactions under constant conditions. We examined the effects of variability in temperature on the host–pathogen relationship between an aquatic zooplankton host (Daphnia laevis) and an environmentally transmitted fungal pathogen (Metschnikowia bicuspidata). We manipulated temperature variability by exposing all populations to mean temperatures of 20°C for the length of the experiments, but introducing periods of 1, 2, and 4 hr each day where the populations were exposed to 28°C followed by periods of the same length (1, 2, and 4 hr, respectively) where the populations were exposed to 12°C. Three experiments were performed to assess the role of thermal variability on Daphnia–pathogen interactions, specifically with respect to: (1) host infection prevalence and intensity; (2) free‐living pathogen survival; and (3) host foraging ecology. We found that temperature variability affected host filtering rate, which is closely related to pathogen transmission in this system. Further, infection prevalence was reduced as a function of temperature variability, while infection intensity was not influenced, suggesting that pathogen transmission was influenced by temperature variability, but the growth of pathogen within infected hosts was not. Host survival was reduced by temperature variability, but environmental pathogen survival was unaffected, suggesting that zooplankton hosts were more sensitive than the fungal pathogen to variable temperatures. Together, these experiments suggest that temperature variability may influence host demography and host–pathogen interactions, providing a link between host foraging ecology and pathogen transmission.     

Interactions between the vesicular-arbuscular mycorrhizal fungus Glomus fascicuhtum and Aphanomyces euteiches root rot of peas

Interactions between Glomus fasciculatum and Aphanomyces euteiches root rot of peas (Pisum sativum), were studied in pot experiments using irradiated soil. Infections with the pathogen were suppressed by VAM when plants were challenge inoculated after two weeks. No reduction of the pathogen was detected when the plants were inoculated with both fungi at the same time. The suppression of the pathogen, obtained by preinoculation with G. fasciculatum, was not reduced when the inoculum level of the pathogen was increased thirty times. The induced resistance to A. euteiches in VAM plants was partially a systemic effect. When root systems were split into two halves, one with mycorrhiza and one with A. euteiches, the oospore production was reduced in both root systems. The infection with the pathogen was only suppressed when both fungi were present in the same pot. The background for the induced resistance is discussed.     

Oxidative enzymes activities of mycorrhizal inoculated pepper plant infected with phytophthora infestans

The activities of polyphenol oxidase, peroxidase and catalase were investigated in Glomus etunicatum an arbuscular mycorrhizal inoculated pepper plant infected with Phytophthora infestans. Pepper plant inoculated with pathogen alone, simultaneously inoculated with mycorrhizal and pathogen or dually inoculated with pathogen before mycorrhizal had increased level of the oxidative enzymes activities especially at 4 weeks after planting. Of the three oxidases investigated catalase had the highest level of activities in all the treatments while the level of peroxidase was the lowest. The results showed that pathogen alone, build up the level of the oxidases in the inoculated pepper plant while inoculation of the pepper plant with AM mycorrhizal either simultaneously or dually with pathogen lower the activity of the oxidases indicating a passive protective effect of mycorrhizal.     

Novel insights into the emergence of pathogens: the case of chestnut blight

Exotic, invasive pathogens have emerged repeatedly and continue to emerge to threaten the world’s forests. Ecosystem structure and function can be permanently changed when keystone tree species such as the American chestnut (Castanea dentata) are eliminated from a whole range by disease. The fungal ascomycete pathogen Cryphonectria parasitica is responsible for causing chestnut blight. Once the pathogen was introduced into the Eastern US, where chestnuts were predominant, chestnuts were all but eliminated. This pathogen is currently causing extensive damage in Europe. A study in this issue of Molecular Ecology sheds new light on the pattern and process of emergence of this devastating plant pathogen ( Dutech et al. 2012 ). The authors used microsatellite markers to investigate the evolutionary history of C. parasitica populations introduced into North America and Europe. To infer sources of migrants and the migration events, the authors included putative source populations endemic to China and Japan, inferred potentially unsampled populations and conducted a multivariate population genetic and complex ABC analysis. Cryphonectria parasitica emerges as an example of an introduced pathogen with limited genotypic diversity and some admixture in the invaded ranges, yet repeated invasions into different areas of Europe and the United States. This work sheds new light on the emergence of C. parasitica providing compelling evidence that this pathogen emerged by repeated migration and occasional admixture.