Host Barriers and Responses to Uncinula necator in Developing Grape Berries

ABSTRACT Grape berries are highly susceptible to powdery mildew 1 week after bloom but acquire ontogenic resistance 2 to 3 weeks later. We recently demonstrated that germinating conidia of the grape powdery mildew pathogen (Uncinula necator) cease development before penetration of the cuticle on older resistant berries. The mechanism that halts U. necator at that particular stage was not known. Several previous studies investigated potential host barriers or cell responses to powdery mildew in berries and leaves, but none included observation of the direct effect of these factors on pathogen development. We found that cuticle thickness increased with berry age, but that ingress by the pathogen halted before formation of a visible penetration pore. Cell wall thickness remained unchanged over the first 4 weeks after bloom, the time during which berries progressed from highly susceptible to nearly immune. Autofluorescent polyphenolic compounds accumulated at a higher frequency beneath appressoria on highly susceptible berries than on highly resistant berries; and oxidation of the above phenolics, indicated by cell discoloration, developed at a significantly higher frequency on susceptible berries. Beneath the first-formed appressoria of all germinated conidia, papillae occurred at a significantly higher frequency on 2- to 5-day-old berries than on 30- to 31-day-old fruit. The relatively few papillae observed on older berries were, in most cases (82.8 to 97.3%), found beneath appressoria of conidia that had failed to produce secondary hyphae. This contrasted with the more abundantly produced papillae on younger berries, where only 35.4 to 41.0% were located beneath appressoria of conidia that had failed to produce secondary hyphae. A pathogenesis-related gene (VvPR-1) was much more highly induced in susceptible berries than in resistant berries after inoculation with U. necator. In contrast, a germin-like protein (VvGLP3) was expressed within 16 h of inoculation in resistant, but not in susceptible berries. Our results suggest that several putative barriers to infection, i.e., cuticle and cell wall thickness, antimicrobial phenolics, and two previously described pathogenesis-related proteins, are not principal causes in halting pathogen ingress on ontogenically resistant berries, but rather that infection is halted by one or more of the following: (i) a preformed physical or biochemical barrier near the cuticle surface, or (ii) the rapid synthesis of an antifungal compound in older berries during the first few hours of the infection process.     

Detection of Erysiphe necator in Air Samples Using the Polymerase Chain Reaction and Species-Specific Primers

ABSTRACT A polymerase chain reaction (PCR) assay employing species-specific primers was developed to differentiate Erysiphe necator from other powdery mildews common in the northwest United States. DNA was extracted from mycelia, conidia, and/or chasmothecia that were collected from grape leaves with a Burkard cyclonic surface sampler. To differentiate E. necator from other erysiphaeceous fungi, primer pairs Uncin144 and Uncin511 were developed to select unique sequences of the internal transcribed spacer regions of E. necator. Using these primers in PCR amplifications, a 367-bp amplicon specific to E. necator was generated, but no amplicons were generated from other erysiphaceous species collected from 48 disparate hosts representing 26 vascular plant families. The PCR limit of detection was one to five conidia of E. necator placed directly into reaction mixtures or 100 to 250 conidia placed on glass rods coated with silicon grease. During field studies, this PCR assay facilitated the detection of E. necator inoculum in air samples within hours of sample rod collection and prior to disease onset. Amplification of E. necator DNA did not occur when the PCR assay was conducted on vineyard air samples collected while grapes were dormant or during periods when vine growth occurred but E. necator remained dormant. The initial PCR detection of E. necator of the season occurred during seasonal ascospore releases caused by precipitation events between bud burst and the prebloom period during the 3 years of the study. Detection ceased for 7 to 11 days following ascospore release and then resumed several days prior to the observance of microscopic symptoms and signs of powdery mildew in the field. Results of this study represent the initial step toward the goal of incorporating an inoculum availability component into current and future grapevine powdery mildew risk assessment models.     

宁夏温室瓜菜白粉病菌鉴定及病害流行预测模型构建

为明确宁夏回族自治区温室瓜菜白粉病菌的分类地位,对采自该地区温室的南瓜、黄瓜和甜瓜上的白粉病菌基于ITS序列分析进行分子鉴定;利用孢子捕捉器对温室中甜瓜白粉病菌的孢子量进行监测,分析环境因子、孢子量和病情指数之间的关系,并采用逐步回归分析法构建温室甜瓜白粉病的流行预测模型。结果表明,基于ITS序列的分子鉴定结果,3种瓜菜白粉病的病原菌均为单囊壳白粉菌Podosphaera xanthii。发病期间,每日温室中甜瓜白粉病菌的孢子量在12:00—16:00时段最多,占24 h内总孢子量的34%~81%,20:00—08:00时段最少;白粉病菌孢子的释放与光照强度有关,相关系数为0.602。第t天的病情指数与标准累积温度、标准累积湿度、t-4 d前08:00—12:00时段的累积孢子量、第t-4天16:00—20:00时段的孢子量均具有显著的相关性,相关系数分别为0.935、0.938、0.956和0.921。以标准累积湿度和第t-4天16:00—20:00时段的孢子量为预测变量构建了温室甜瓜白粉病流行预测模型,决定系数为0.962,表明该模型具有较好的实际应用价值。     

Formation of overwintering structures of Uncinula necator and colonization of grapevine under field conditions

Developing shoots of grapevine ( Vitis vinifera cv. Kerner) were inoculated with conidia of the powdery mildew, Uncinula necator , at well-defined phenological stages of the host to provoke the development of flag shoots in the field and to investigate these shoots as early as possible in the following growing season for the presence of the pathogen. The disease progress was monitored and fungal growth and development on samples from a field trial were analysed by means of low-temperature scanning electron microscopy (LT–SEM). Mycelium was detected on the surface and in the interior parts of buds. The suitability of the field plot to analyse flag shoots was proven by the occurrence of such shoots in the following spring. Furthermore, early stages of cleistothecia development on berries were described for the first time. Establishment of U. necator in dormant buds of grapevine, giving rise to flag shoots in the following spring, is considered to play a significant role in overwintering of the pathogen in the vineyards of southern Germany.     

A single dominant locus, ren4, confers rapid non-race-specific resistance to grapevine powdery mildew

In the present study we screened the progeny of Vitis vinifera × V. romanetii populations segregating for resistance to powdery mildew and determined the presence of a single, dominant locus, Ren4, conferring rapid and extreme resistance to the grapevine powdery mildew fungus Erysiphe necator. In each of nine Ren4 pseudo-backcross 2 (pBC(2)) and pBC(3) populations (1,030 progeny), resistance fit a 1:1 segregation ratio and overall segregated as 543 resistant progeny to 487 susceptible. In full-sib progeny, microscopic observations revealed the reduction of penetration success rate (as indicated by the emergence of secondary hyphae) from 86% in susceptible progeny to below 10% in resistant progeny. Similarly, extreme differences were seen macroscopically. Ratings for Ren4 pBC(2) population 03-3004 screened using natural infection in a California vineyard and greenhouse and using artificial inoculation of an aggressive New York isolate were fully consistent among all three pathogen sources and environments. From 2006 to 2010, Ren4 pBC(2) and pBC(3) vines were continuously screened in California and New York (in the center of diversity for E. necator), and no sporulating colonies were observed. For population 03-3004, severity ratings on leaves, shoots, berries, and rachises were highly correlated (R(2) = 0.875 to 0.996) in the vineyard. Together, these data document a powdery mildew resistance mechanism not previously described in the Vitaceae or elsewhere, in which a dominantly inherited resistance prevents hyphal emergence and is non-race-specific and tissue-independent. In addition to its role in breeding for durable resistance, Ren4 may provide mechanistic insights into the early events that enable powdery mildew infection.     

Effects of humidity on the development of grapevine powdery mildew

ABSTRACT The effects of humidity on powdery mildew development on grape seedlings and the germination of Uncinula necator conidia in vitro were examined. Studies were conducted at an optimum temperature of 25 +/- 2 degrees C. Disease on foliage was markedly affected by humidity levels in the test range of 39 to 98% relative humidity (RH), corresponding to vapor pressure deficits (VPD) of 1,914 to 61 Pa. Incidence and severity increased with increasing humidity to an optimum near 85% RH, and then appeared to plateau or decrease marginally at higher values. Conidial density and chain length also were proportional to humidity, but were influenced less strongly. There was a strong, positive linear relationship between humidity level and frequency of conidium germination with RH treatments of 相似文献   

Conditions for development of powdery mildew of tomato caused by Oidium neolycopersici

Oidium neolycopersici causes severe powdery mildew on all aerial parts of tomato, excluding the fruit. The objective of the present work was to examine factors that influence the development of O. neolycopersici on tomato and to identify potential methods for managing tomato powdery mildew. Under controlled conditions, the highest rates of conidial germination were observed at 25 degrees C, 99% relative humidity (RH) and minimal light, and the lowest on leaves adjacent to fruits. Optimal conditions for appressoria formation were 25 degrees C, RH ranging from 33 to 99%, and 1,750 lux light intensity. More conidia were formed at 20 degrees C, 70 to 85% RH, and 5,150 lux light intensity than at 16 and 26 degrees C, 99% RH, and 480 to 1,750 lux, respectively. Conidia survived and remained capable of germination for over four months when initially incubated at lower temperatures and higher RH, as compared with their fast decline under more extreme summer shade conditions. In growth chamber experiments, disease did not develop at 28 degrees C. Within the range of 70 to 99% RH, disease was less severe under the higher RH than the drier conditions. Disease was also less severe at lower light intensities. Data collected in three commercial-like greenhouse experiments involving various climate regimes were used to draw correlations regarding the effects of temperature and RH on the development of epidemics. Severity of powdery mildew was positively correlated with the duration of the range 15 to 25 degrees C, 1 to 4 weeks before disease evaluation (BDE), RH levels of 60 to 90% at 2 to 4 weeks BDE, and RH of 50 to 60% during the week BDE. Conversely, disease was negatively correlated with the duration of temperatures in the low and high ranges (5 to 15 degrees C and 35 to 40 degrees C) at 1 to 4 weeks BDE, with the duration of RH levels of 40% and below at 1 to 4 weeks BDE, and with 50 to 60% RH during the third week BDE. High (90 to 100%) RH was also negatively correlated with disease severity. These results suggest that the combination of high temperatures and low RH may help reduce O. neolycopersici powdery mildew severity in greenhouse tomatoes.     

Germinability of Venturia inaequalis Conidia Exposed to Sunlight

ABSTRACT Detached conidia of Venturia inaequalis were exposed simultaneously outdoors to direct sunlight or in an adjacent ventilated enclosure in darkness for periods ranging from 0.5 to 16.5 h. In addition, conidia were either exposed or not exposed to UV light (254 nm) from 1 to 64 min in the laboratory. After exposure, conidia were placed on water agar in closed petri dishes and allowed 24 h to germinate. Germinability of conidia in the laboratory was reduced up to 95% by doses of UV(254) of 8.1 to 10.8 kJ m(-2).Germinability of conidia in the field was reduced up to 95% by doses of UV-B of about 44 kJ m(-2). A significant percentage of conidia survived radiation doses equivalent to 12 h in full sunlight.     

Effects of Diffuse Colonization of Grape Berries by Uncinula necator on Bunch Rots, Berry Microflora, and Juice and Wine Quality

ABSTRACT Production of grape (principally cultivars of Vitis vinifera) for high-quality wines requires a high level of suppression of powdery mildew (Uncinula necator syn. Erysiphe necator). Severe infection of either fruit or foliage has well-documented and deleterious effects upon crop and wine quality. We found that berries nearly immune to infection by U. necator due to the development of ontogenic resistance may still support diffuse and inconspicuous mildew colonies when inoculated approximately 3 weeks post-bloom. Fruit with diffuse mildew colonies appear to be healthy and free of powdery mildew in late-season vineyard assessments with the naked eye. Nonetheless, presence of these colonies on berries was associated with (i) elevated populations of spoilage microorganisms; (ii) increased evolution of volatile ethyl acetate, acetic acid, and ethanol; (iii) increased infestation by insects known to be attracted to the aforementioned volatiles; (iv) increased rotting by Botrytis cinerea; and (v) increased frequency of perceived defects in wines prepared from fruit supporting diffuse powdery mildew colonies. Prevention of diffuse infection requires extending fungicidal protection until fruit are fully resistant to infection. Despite a perceived lack of improvement in disease control due to the insidious nature of diffuse powdery mildew, potential deleterious effects upon crop and wine quality thereby would be avoided.     

苦瓜白粉病病原菌和生理小种的鉴定及苦瓜对白粉病的抗性遗传分析

为明确海南省苦瓜白粉病的病原菌、生理小种及苦瓜对白粉病的抗性遗传规律,结合形态学鉴定和分子鉴定解析白粉病菌及生理小种种类,通过显微镜观察白粉病菌侵染过程,并应用主基因+多基因混合遗传模型分析法探讨苦瓜对白粉病的主要抗性遗传规律。结果表明:采集自海南省6个市(县)的苦瓜白粉病病原菌均为单囊壳白粉菌Sphaerotheca fuliginea,属生理小种2F,该菌在侵染苦瓜叶片时有4个关键时期:接种后4 h为分生孢子萌发高峰期,8 h为附着孢形成高峰期,16～24 h为次生菌丝形成高峰期,5 d为分生孢子梗形成高峰期。将其接种于苦瓜抗、感品系,对白粉病的抗性符合2对加性-显性-上位性主基因+加性-显性多基因模型,主基因和多基因共同控制苦瓜对白粉病的抗性,其中以主基因遗传为主,且会受到环境变异的影响。根据苦瓜抗性遗传规律,F2代主基因遗传率最高,受环境影响最小,在苦瓜的白粉病抗性育种中,以早期世代F2代作为有效选择世代。研究表明白粉病菌侵染叶片的前2 d是白粉病防治的最佳时期,所以在白粉病易发的物候期,可将防治时间提前1～2 d。     

Effects of light quality on conidiophore formation of the melon powdery mildew pathogen <Emphasis Type="Italic">Podosphaera xanthii</Emphasis>

The lengths of conidiophores in fungal colonies of the melon powdery mildew pathogen Podosphaera xanthii Pollacci KMP-6 N cultured under greenhouse (natural) conditions differed markedly from those cultured in a growth chamber. We hypothesized that light wavelength was responsible for the differences in conidiophore length. In this study, we examined the effects of light-emitting diode (LED) irradiation (purple, blue, green, orange, and red light) and white light on colony development and conidiophore formation in KMP-6 N using a stereomicroscope and a high-fidelity digital microscope. Colonies on leaves were flat under greenhouse conditions and under red LED light irradiation but were stacked under growth chamber conditions and under purple, blue, green, and orange LED light irradiation. In addition, KMP-6 N formed catenated conidia comprising six conidia per conidiophore under greenhouse conditions and red light but more than seven conidia per conidiophore under growth chamber conditions and purple, blue, green, and orange light. Furthermore, almost none of the conidia on top of the conidiophores grown under blue light were fully constricted. Therefore, these fungi could not scatter their conidia and spread infection. This is the first report of the effects of LED lights on conidiophore formation in the melon powdery mildew fungus P. xanthii. The results provide insight into the mechanisms underlying the responses of conidiophores to light of specific wavelengths and conidial scatter from conidiophores of melon powdery mildew fungi.     

Effect of Grapevine Training Systems on Development of Powdery Mildew

The effects of two training systems and row spacing on development of powdery mildew caused by Uncinula necator on clusters of 'Chardonnay' and 'Cabernet Sauvignon' grapevines were examined. Disease development was monitored in blocks with two different row spacing (2 and 3 m) in vertical shoot positioned vines (VSP) and in free-positioned, topped vines (FC) with no foliage support wires. The FC vines were hedged to about one meter shoot length. No fungicides were applied and disease powdery mildew level was recorded four to seven days after appearance of the first disease symptoms. During five consecutive years (1994–1998), disease incidence was higher in the VSP system than the FC vines. The difference was high when disease level was low (30% of the clusters in VSP vines infected, compared to 5% in the FC vines) and decreased when disease pressure was high (79% in VSP compared to 46% in FC vines). In the 'Cabernet Sauvignon', in four of the years, disease incidence was higher in the narrow spacing of 2 m between the rows than that in the wider 3 m spacing. Microclimate (temperature, relative humidity and light intensity) was monitored in the cluster zone near the spurs of 'Chardonnay' vines during three weeks in the 1998 season. In VSP vines light intensity was lower then that in FC vines both four and one week before disease symptoms appeared (72% and 18% respectively). The differences in temperature and relative humidity were less than 1°C and 3%, respectively, and most likely did not affect disease development. The results suggested that high light intensity is the primary factor, which limits powdery mildew growth development on field-grown grapevines in the Golan region of Israel. The use of the FC system might be useful in reducing the need of fungicides.     

A single nucleotide polymorphism in the β-tubulin gene distinguishing two genotypes of Erysiphe necator expressing different symptoms on grapevine

The biotrophic fungus Erysiphe necator (formerly Uncinula necator ), for which two genetic groups have been described in European vineyards, is the causal agent of grapevine powdery mildew. By analysing the pathogen population with respect to polymorphism in the sequence of the β-tubulin gene, which distinguishes two groups of isolates, a new tool was developed for epidemiological and population studies and tested in the vineyard. As in many ascomycetes, the β-tubulin gene of E. necator ( Entub ) includes six introns and seven exons and encodes a 447-amino-acid protein. A single nucleotide polymorphism (SNP) in the intron-3 region of the Entub gene distinguished two genetic groups (A and B). This method was used to examine differences in the ratio of the two groups from a total of 289 grape powdery mildew samples collected at the beginning of the growing season from either flag shoots or leaves with sparse-spot symptoms in four different vineyards. The SNP in the intron-3 region of the β-tubulin gene, similar to SNPs in the CYP51 gene, was associated with genotypes A and B of E. necator and confirmed the existence of two sympatric populations of the pathogen in the French vineyards. Differences in the relative proportions of each group varied with the presence or absence of flag-shoot symptoms and with the region in which isolates had been collected.     

No indication of strict host associations in a widespread mycoparasite: grapevine powdery mildew (Erysiphe necator) is attacked by phylogenetically distant Ampelomyces strains in the field

Pycnidial fungi belonging to the genus Ampelomyces are common intracellular mycoparasites of powdery mildews worldwide. Some strains have already been developed as commercial biocontrol agents (BCAs) of Erysiphe necator and other powdery mildew species infecting important crops. One of the basic, and still debated, questions concerning the tritrophic relationships between host plants, powdery mildew fungi, and Ampelomyces mycoparasites is whether Ampelomyces strains isolated from certain species of the Erysiphales are narrowly specialized to their original mycohosts or are generalist mycoparasites of many powdery mildew fungi. This is also important for the use of Ampelomyces strains as BCAs. To understand this relationship, the nuclear ribosomal DNA internal transcribed spacer (ITS) and partial actin gene (act1) sequences of 55 Ampelomyces strains from E. necator were analyzed together with those of 47 strains isolated from other powdery mildew species. These phylogenetic analyses distinguished five major clades and strains from E. necator that were present in all but one clade. This work was supplemented with the selection of nine inter-simple sequence repeat (ISSR) markers for strain-specific identification of Ampelomyces mycoparasites to monitor the environmental fate of strains applied as BCAs. The genetic distances among strains calculated based on ISSR patterns have also highlighted the genetic diversity of Ampelomyces mycoparasites naturally occurring in grapevine powdery mildew. Overall, this work showed that Ampelomyces strains isolated from E. necator are genetically diverse and there is no indication of strict mycohost associations in these strains. However, these results cannot rule out a certain degree of quantitative association between at least some of the Ampelomyces lineages identified in this work and their original mycohosts.     

Effect of colored shade nets on pepper powdery mildew (<Emphasis Type="Italic">Leveillula taurica</Emphasis>)

The effect of colored shade nets with different shade intensities and qualities of irradiation transmittance on pepper powdery mildew was tested in mini-plots and field experiments. Leaf coverage byLeveillula taurica and leaf shedding due to the disease were more severe in the shade, by up to 275% and 70%, respectively, than in the open field. Leaf coverage byL. taurica symptoms and leaf shedding from plants grown under 25% shade black nets were higher, by up to 70% and 180%, respectively, than under 40% shade nets. The color of the shade nets affected the intensity of photosynthetically active (PAR), ultra-violet (UV), blue, red and far red radiations, the UV/blue light ratio, and percent PAR and UV transmitted. The various nets suppressed the disease differently. Black, blue-silver, green and red nets were associated with lower levels of disease in the field experiments. The red net was also superior in the mini-plots. The other results from the mini-plots were not similar to those from the field, probably reflecting more intensive epidemic development in the mini-plots. No interaction between net type and cultivar was found when two cultivars were grown under the nets. Yield was higher under nets than in the open; nevertheless, the yield from plants grown under the 40% shade black net was not higher than that of the plants under the 25% black net, despite the significantly lower levels of disease at the higher shade intensity. B-quality pepper yield was significantly higher in the plots covered by 25% shade. Yield differences between the different colored nets were also not well correlated with disease levels, probably due to factors negating the direct effect of the nets on the plants and their yield. Implementation of either ‘friendly’ (Ampelomyces quisqualis AQ10/Trichoderma harzianum T39/ sulfur/ neem seed extract) or chemical (sulfur/ pyrifenox/ Polyoxin AL/ myclobutanil/ azoxystrobin) spray regimes successfully reduced disease severity under the different nets. There was no interaction between net type and spray regime. Thus, growing sweet pepper under shade nets results in increased yields and also in higher powdery mildew severity. Disease is negatively associated with the rate of shading and is variably affected by the quality of light filtered through the different colored shade nets. http://www.phytoparasitica.org posting May 8, 2007.     

Potential and actual ascospore release of Erysiphe necator chasmothecia in Austria

Journal of Plant Diseases and Protection - Ascospores of grape powdery mildew (Erysiphe necator Schw.) play a crucial role in the disease onset in spring in many vine-growing areas. We investigated...     

Morphological and molecular characterization for a Japanese isolate of tomato powdery mildew Oidium neolycopersici and its host range

 A single conidium of tomato powdery mildew was isolated from heavily infected leaves of tomato (cv. Moneymaker) grown in the greenhouse of Kinki University, Nara Prefecture, Japan. It was successively multiplied so the morphological and taxonomic characteristics of the pathogen and its host range under high humidity conditions could be analyzed. The isolate KTP-01 of the tomato powdery mildew optimally developed infection structures at 25°C under continuous illumination of 3500 lx. More than 90% of the conidia germinated and developed moderately lobed appressoria. After forming haustoria, the pathogen elongated secondary hyphae from both appressoria and conidia. The hyphae attached to leaf surfaces by several pairs of appressoria and produced conidiophores with noncatenated conidia. In addition to its morphological similarity to Oidium neolycopersici, the phylogenetic analysis (based on the sequence of internal transcribed spacer regions of rDNA) revealed that KTP-01 could be classified into the same cluster group as O. neolycopersici. In host range studies, KTP-01 produced abundant conidia on the foliage of all tomato cultivars tested and tobacco (Nicotiana tabacum), and it developed faint colonies accompanied by necrosis on leaves of potato (Solanum tuberosum), red pepper (Capsicum annuum), petunia (Petunia × hybrida), and eggplant (S. melongena). The pathogen did not infect other plant species including Cucurubitaceae plants, which have been reported to be susceptible to some foreign isolates. Thus, the present isolate of the tomato powdery mildew was assigned as O. neolycopersici, a pathotype different from foreign isolates of the pathogen. Received: December 5, 2002 / Accepted: December 26, 2002 Acknowledgments This work was supported in part by a Grant-in-Aid (12660050) from the Ministry of Education, Culture, Sports, Science, and Technology of Japan. We express our deepest thanks to professor Dr. Y. Sato, Toyama Prefectural University, for his kind and valuable suggestion on taxonomic analysis of the powdery mildew pathogen described in the present study.     

Ontogenic resistance and plant disease management: a case study of grape powdery mildew

ABSTRACT A fundamental principle of integrated pest management is that actions taken to manage disease should be commensurate with the risk of infection and loss. One of the less-studied factors that determines this risk is ontogenic, or age-related resistance of the host. Ontogenic resistance may operate at the whole plant level or in specific organs or tissues. Until recently, grape berries were thought to remain susceptible to powdery mildew (Uncinula necator) until late in their development. However, the development of ontogenic resistance is actually quite rapid in berries, and fruit become nearly immune to infection within 4 weeks after fruit set. Our objective was to determine how and at what stage the pathogen was halted in the infection process on ontogenically resistant berries. Adhesion of conidia, germination, and appressorium formation were not impeded on older berries. However, once berries were approximately 3 weeks old and older, few germlings were able to form secondary hyphae. Ontogenically resistant berries responded rapidly to infection by synthesis of a germin-like protein that had been previously shown to play a role in host defense against barley powdery mildew. On susceptible berries, cell discoloration around penetration sites indicated the oxidation of phenolic compounds; a process that was followed by localized cell death. However, the pathogen was still able to infect such cells prior to their death, continue secondary growth, and thereby colonize young berries. Formation of papillae was not involved in the differential resistance mechanism of older berries. In susceptible berries, papillae formed frequently at infection sites but did not always contain the pathogen, whereas in resistant berries, the pathogen was always halted prior to the formation of papillae. The host defense, which conditions ontogenic resistance, operates in the earliest stages of the infection process, in the absence of gross anatomical barriers, prior to the formation of a functional haustorium and prior to the development of a conspicuous penetration pore. We also found that diffuse powdery mildew colonies that were not visible in the field predisposed berries to bunch rot by Botrytis cinerea, increased the levels of infestation by spoilage microorganisms, and substantially degraded wine quality. Our improved understanding of the nature, causes, and stability of ontogenic resistance in the grapevine/ powdery mildew system has supported substantial changes in how fungicides are used to control the disease. Present applications are more focused on the period of maximum fruit susceptibility instead of following a calendar-based schedule. This has improved control, reduced losses, and in many cases reduced the number of fungicide applications required to suppress the disease. Particularly where fungicides are deployed in a programmatic fashion and ontogenic resistance is dynamic, there may be equivalent improvements to be made in other hostpathogen systems through studies of how host susceptibility changes through time.     

Plant age and ambient temperature: significant drivers for powdery mildew (Erysiphe cruciferarum) epidemics on oilseed rape (Brassica napus)

Powdery mildew (Erysiphe cruciferarum) is an important disease in oilseed rape crops worldwide, but of sporadic importance in most southern Australian crops. Six Brassica napus cultivars were exposed to E. cruciferarum simultaneously in four plant age cohorts. First symptoms of powdery mildew appeared 9 days after inoculation (dai) on the oldest plants [42 days after seeding (das)], but 44 dai in the youngest plants that were exposed to inoculum from sowing, although final disease severity did not differ with the plant age at exposure. The maximum level of pod peduncle infestation was unaffected by plant age (P = 0.37) or cultivar (P = 0.28). The effect of temperature was also investigated. The development of disease on plants was slower and final severity reduced at a day/night temperature 14/10 °C compared with 22/17 °C. In vitro, maximum growth of germ tubes from conidia of E. cruciferarum was at 15–20 °C and survival of conidia reduced by temperatures >30 °C. The results explain the sporadic nature of powdery mildew outbreaks in winter‐grown oilseed rape in Australia, where slow rates of infection occur when seasonal colder prevailing winter conditions coincide with the presence of younger plants, together curtailing rapid disease development until temperatures increase in late winter/early spring. These results explain why epidemics are most severe in the two warmer cropping regions, viz. the northern agricultural region of Western Australia and New South Wales. This study suggests that with increases in winter temperatures under future climate scenarios, earlier and more severe powdery mildew outbreaks in Australia will be favoured.     

Coincidence of maximum severity of powdery mildew on grape leaves and the carbohydrate sink‐to‐source transition

Grapevine leaves infected with powdery mildew are a source of inoculum for fruit infection. Leaves emerging on a single primary shoot of Vitis vinifera cv. Cabernet Sauvignon were exposed to average glasshouse temperatures of 18°C (0·23 leaves emerging/day) or 25°C (0·54 leaves emerging/day). All leaves on 8–10 shoots with approximately 20 leaves each were inoculated with Erysiphe necator conidia to assess disease severity after 14 days in the 25°C glasshouse. Two photosynthetic ‘source’ leaves per shoot on the remaining 8–10 shoots were treated with ¹⁴CO₂ to identify, by autoradiography, the leaf position completing the carbohydrate sink‐to‐source transition. There was a clear association between the mean modal leaf position for maximum severity of powdery mildew (position 3·7 for 18°C; position 4·4 for 25°C) and the mean position of the leaf completing the sink‐to‐source transition (position 3·8 for 18°C; position 4·7 for 25°C). The mean modal leaf position for the maximum percentage of conidia germinating to form secondary hyphae was 4·2 for additional plants grown in the 25°C glasshouse. A higher rate of leaf emergence resulted in a greater proportion of diseased leaves per shoot. A Bayesian model, consisting of component models for disease severity and leaf ontogenic resistance, had parameters representing the rate and magnitude of pathogen colonization that differed for shoots developing in different preinoculation environments. The results support the hypothesis that the population of leaves in a vineyard capable of supporting substantial pathogen colonization will vary according to conditions for shoot development.