Hop stunt viroid (HSVd) is an economically important pathogen that reduces growth and yield of hops. Visual symptoms of infected hop are highly dependent on cultivar. A study was conducted using six cultivars of hop to determine the impact on yield. Average dry cone yields of infected 'Glacier', 'Cascade', and 'Willamette' were reduced by 62, 14, and 34%, respectively, relative to noninoculated healthy plants. No significant yield reduction was observed for 'Nugget', 'Columbus', and 'Galena'. The alpha-acid and beta-acid contents showed a parallel pattern. Horticultural parameters of Willamette and Nugget were measured in the final year of the study. Internode length, shoot length, and side-arm length were reduced by 29, 26, and 73%, respectively, for infected Willamette bines relative to noninfected bines; no effects were observed resulting from infection of Nugget. To understand the current potential impact of HSVd, a survey was conducted to determine its distribution in central Washington. The survey revealed that 17% of hop plants tested are infected. Hop yield and hop plant longevity will be significantly affected by this level of infection.

Podosphaera macularis, the causal agent of hop powdery mildew, is a recurrent threat to hops in the Pacific Northwest because of the potential to reduce cone yield and quality. Early-season pruning is a common practice in hop production for horticultural reasons. Studies were conducted over a 3-year period in a commercial hop yard to quantify the effect of pruning method and timing on disease development, yield, and cone quality factors. A 4-week delay in pruning reduced the incidence of leaves with powdery mildew from 46 to 10% and cones from 9 to 1%, with the specific effect being season dependent. Pruning using chemical desiccants rather than by mechanical means had similar effects on disease levels on leaves. On cones, though, chemical pruning had a small but significant reduction in the incidence of powdery mildew compared with mechanical pruning. Cone yield, levels of bittering-acids, and color were not negatively affected in any individual year or cumulatively over three seasons when pruning treatments were applied repeatedly to the same plots during the study period. Delayed pruning may offer a low-cost means of reducing both the incidence of powdery mildew and early-season fungicide inputs in certain cultivars.

Host resistance, both quantitative and qualitative, is the preferred long-term approach for disease management in many pathosystems, including powdery mildew of hop (Podosphaera macularis). In 2012, an epidemic of powdery mildew occurred in Washington and Idaho on previously resistant cultivars whose resistance was putatively based on the gene designated R6. In 2013, isolates capable of causing severe disease on cultivars with R6-based resistance were confirmed in Oregon and became widespread during 2014. Surveys of commercial hop yards during 2012 to 2014 documented that powdery mildew is now widespread on cultivars possessing R6 resistance in Washington and Oregon, and the incidence of disease is progressively increasing. Pathogenic fitness, race, and mating type of R6-virulent isolates were compared with isolates of P. macularis lacking R6 virulence. All isolates were positive for the mating type idiomorph MAT1-1 and were able to overcome resistance genes Rb, R3, and R5 but not R1 or R2. In addition, R6-virulent isolates were shown to infect differential cultivars reported to possess the R6 gene and also the R4 gene, although R4 has not yet been broadly deployed in the United States. R6-virulent isolates were not detected from the eastern United States during 2012 to 2015. In growth chamber studies, R6-virulent isolates of P. macularis had a significantly longer latent period and produced fewer lesions on plants with R6 as compared with plants lacking R6, indicating a fitness cost to the fungus. R6-virulent isolates also produced fewer conidia when compared with isolates lacking R6 virulence, independent of whether the isolates were grown on a plant with or without R6. Thus, it is possible that the fitness cost of R6 virulence occurs regardless of host genotype. In field studies, powdery mildew was suppressed by at least 50% on plants possessing R6 as compared with those without R6 when coinoculated with R6-virulent and avirulent isolates. R6 virulence in P. macularis appears to be race specific and, at this time, imposes a measurable fitness penalty on the fungus. Resistance genes R1 and R2 appear to remain effective against R6-virulent isolates of P. macularis in the U.S. Pacific Northwest.

Canopy management is an important aspect of control of powdery mildew diseases and may influence the intensity of fungicide applications required to suppress disease. In hop, powdery mildew (caused by Podosphaera macularis) is most damaging to cones when infection occurs during bloom and the juvenile stages of cone development. Experiments were conducted over 3 years to evaluate whether fungicide applications could be ceased after the most susceptible stages of cone development (late July) without unduly affecting crop yield and quality when disease pressure was moderated with varying levels of basal foliage removal. In experimental plots of 'Galena' hop, the incidence of leaves with powdery mildew was similar whether fungicides were ceased in late July or made in late August. Disease levels on leaves were unaffected by the intensity of basal foliage removal, whereas the intensity of basal foliage removal interacted with the duration of fungicide applications to affect disease levels on cones. Similar experiments conducted in large plots of 'Tomahawk' hop in a commercial hop yard similarly found no significant impact on disease levels on leaves from either the duration of fungicide applications or intensity of basal foliage removal. In contrast, on cones, application of fungicides into August had a modest, suppressive effect on powdery mildew. There was also some evidence that the level of powdery mildew on cones associated with fungicide treatment was influenced by the intensity of basal foliage removal. When fungicide applications ceased in late July, there was a progressive decrease in the incidence of cones with powdery mildew with increasing intensity of basal foliage removal. Removing basal foliage two to three times allowed fungicide applications to be terminated in late July rather than late August without diminishing disease control on cones, yield, or cone quality factors. Thus, this study further establishes that fungicide applications made during the early stages of hop cone development have the strongest effect on suppression of powdery mildew on cones. The additive effect of fungicide applications targeted to the periods of greatest cone susceptibility and canopy management to reduce disease favorability may obviate the need for fungicide applications later in the season. This appears to be a viable strategy in mature hop yards of certain cultivars when disease pressure is not excessively high.

Results of 28 field trials conducted over a 12-year period investigating management of hop powdery mildew caused by Podosphaera macularis were quantitatively summarized by meta-analysis to compare product efficacy and use patterns by mode of action as defined by Fungicide Resistance Action Committee (FRAC) groups. Availability of original observations enabled individual participant data meta-analysis. Differences in control of powdery mildew on leaves and cones were apparent among fungicide FRAC groups when individual products were evaluated over the course of a growing season. FRAC groups 13, 3, and U13 provided the most efficacious control of powdery mildew on leaves. Percent disease control on cones was influenced by midseason foliar disease and fungicide mode-of-action. FRAC 13 provided significantly better disease control on cones than all other groups except U13, 3, and premixes of 7 with 11. Disease control on leaves was similar when a rotational program of fungicides was used, independent of the modes of action, but improved on cones if FRAC groups 13 and 3 were both included compared with programs consisting of FRAC groups 11 and 3, 11 and 5, or 3 and 5. Disease control on cones was improved from 32 to 52%, on average, when the fungicide quinoxyfen (FRAC 13) was applied at least once during the early stages of cone development, defined in this analysis as 20 July to 10 August, as compared with all other treatments. Efficacy of disease control on cones by quinoxyfen was moderated by and interacted with the incidence of leaves with powdery mildew. Disease control on cones was further improved if two applications of quinoxyfen were made during this period. Collectively, these findings suggest that disease control during juvenile stages of cone development largely influences the success of fungicide programs and point to the critical importance of focusing management efforts during this stage of development, independent of what actual management strategy is employed.

Powdery mildew of hop ( P odosphaera macularis ) may cause economic loss due to reductions in cone yield and quality. Quantitative estimates of crop damage from powdery mildew remain poorly characterized, especially the effect of late season disease management on crop yield and quality. Field studies in Washington State evaluated cone yield, bittering acid content and quality factors when fungicide applications were ceased at different stages of cone development. The incidence of cones with powdery mildew was linearly correlated with yield of cones, bittering acids and accelerated cone maturation. In cultivar Galena, the cumulative effect of every 1% increase in cones powdery mildew incidence was to reduce alpha‐acid yield by 0·33%, which was due to direct effects on cone yield but also indirect effects mediated by dry matter. In the more susceptible cultivar Zeus, alpha‐acid yield was increased 20% by controlling powdery mildew through the transition of bloom to early cone development compared to ceasing fungicide applications at bloom: additional applications provided only modest improvements in alpha‐acid yield. In both cultivars, the impact of powdery mildew on aroma characteristics and bittering acid content were less substantial than cone yield. The damage caused by powdery mildew to cone colour and alpha‐acid yield, as well as the effectiveness of fungicide applications made to manage the disease, appears inseparably linked to dry matter content of cones at harvest. Realising achievable yield potential in these cultivars requires control of the disease through early stages of cone development and harvest before maturity exceeds c . 25% dry matter.

Downy mildew (caused by Pseudoperonospora humuli) and powdery mildew (caused by Podosphaera macularis) are important diseases of hop in the Pacific Northwest United States, and cultural practices may affect the severity of both diseases. The association of spring pruning quality and timing with severity of downy mildew and powdery mildew was assessed through analysis of survey data collected from commercial hop yards in Oregon and Washington. Among 149 hop yards surveyed, the most common pruning method was chemical desiccation (48% of yards), mechanical pruning (23%), or a combination of these practices (15%). The quality of pruning was assessed using a three-category ordinal scale (“excellent”, “moderate”, or “poor”) based on the amount of foliage remaining on plants following pruning. Excellent pruning quality was attained more often in yards pruned twice (74.6 to 82.1% of yards) versus once (33.8% of yards), independent of pruning method. Seasonal severity of downy mildew in Oregon increased approximately twofold with reduction in pruning quality from excellent to moderate to poor. Pruning quality was not significantly related to levels of powdery mildew on leaves or cones in Oregon. Under more severe disease pressure in Washington, however, seasonal severity of powdery mildew on leaves and the incidence of cones with powdery mildew were significantly greater in yards that had poor pruning compared with excellent pruning. Moreover, yards that had excellent pruning quality received, on average, 1.1 to 1.5 fewer fungicide applications per season for downy mildew or powdery mildew compared with yards that had moderate or poor pruning quality. This savings was associated with delayed initiation of the first application by 7.5 to 14.2 days in yards with excellent pruning quality. Replicated experiments in commercial yards in Oregon quantified the effect of delaying pruning timing 5 to 21 days compared with growers' standard practices on the diseases and yield. Downy mildew suppression by delayed pruning was dependent on cultivar and year of sampling, being significantly reduced fivefold only in ‘Willamette’ in 2007. Severity of powdery mildew and cone yield was similar between plots that received the delayed or standard pruning timing treatments. Collectively, these studies emphasize that early spring sanitation measures are associated with reduced primary inoculum and are critically important for managing both downy mildew and powdery mildew. A savings of at least one fungicide application per year appears achievable when spring pruning is conducted thoroughly and slightly delayed compared with growers' current practices.

Downy mildew, caused by Pseudoperonospora humuli, is an important disease of hop in most production regions in the northern hemisphere. Insensitivity to phenylamide fungicides was detected in isolates of P. humuli in production regions in Oregon and Idaho in 1992, and these fungicides since have been used on a limited basis. In this study, the prevalence of phenylamide insensitivity among isolates of P. humuli collected from 2005 to 2007 in the northwestern United States was quantified using a leaf disk assay with a discriminating dose (25 μg/ml) of either metalaxyl or mefenoxam with inoculum derived from 201 systemically infected diseased shoots (basal spikes) collected from 6, 10, and 11 hop yards in Idaho, Washington, and Oregon, respectively. A subset of 47 basal spike isolates and 42 monosporic isolates collected from two yards in Idaho and nine yards in Oregon during 2006 and 2007 were assayed using a dilution series of metalaxyl to determine the effective dose that inhibited 50% of the incidence of sporulation (ED50). Insensitivity to mefenoxam was detected in 31 of 74 (41.9%) basal spikes collected from 13 hop yards. Insensitivity to the related compound metalaxyl was detected in 52 of 80 (65%) spikes collected from nine hop yards, including four hop yards in Washington. Log ED50 values ranged from -2.25 to 2.67 for basal spike isolates and -2.27 to 2.98 for monosporic isolates and had a similar distribution. Log ED50 values for monosporic isolates and entire basal spike isolates were significantly associated. However, the slope of the regression line was less than 1, indicating that the log ED50 values obtained from entire basal spike isolates were greater than the corresponding log ED50 values obtained from a monosporic isolate obtained from that spike. This research suggests that insensitivity to phenylamide fungicides is a stable phenotype in P. humuli. Management of downy mildew should rely on measures other than these fungicides in yards or regions where phenylamide insensitivity is prevalent, which is now known to include certain hop yards in Idaho, Oregon, and Washington.

Viroid status was investigated in commercial hop (Humulus lupulus) plantings in three regions of intensive hop production in Washington State. Hop stunt viroid (HSVd), the causal agent of hop stunt disease, was detected for the first time in hop plants in North America. HSVd was detected in samples from 10 of the 33 hop gardens sampled and in 19 of 126 plant samples. Infection with HSVd was associated with chlorosis and reduced plant vigor. Reverse transcription-polymerase chain reaction amplification, sequence identity, and pathogenicity of hop extracts on cucumber seedlings confirmed the presence of HSVd. Apple fruit crinkle viroid (AFCVd), recently reported in hops in Japan, was not detected by RT-PCR in any samples. Hop latent viroid (HLVd), which is frequently associated with hop germplasm world-wide, was detected with high frequency (98 of 126 hop plants surveyed). Accepted for publication 23 September 2007. Published 27 November 2007.

Pseudoperonospora humuli populations from Oregon and Washington were analyzed for genetic variation using random amplified polymorphic DNA (RAPD) and DNA amplification fingerprinting (DAF) markers. The genetic structure of the Oregon and Washington populations differed considerably. There was little genetic diversity in Washington, with only five RAPD and six DAF groups detected among 40 isolates tested. One genotype was predominant in Washing-ton. In contrast, 18 RAPD and 34 DAF groups were found among the 40 isolates tested from Oregon. No unique band profile associated with host cultivar was observed. It is suggested that the distinct difference in population structure between the two geographic regions might be due to climatic differences resulting in a higher frequency of sexual reproduction of P. humuli in Oregon than in Washington.

Failures of fosetyl-Al (Aliette) to control hop (Humulus lupulus) downy mildew, caused by Pseudoperonospora humuli, have recently occurred in northern Idaho and Oregon. To determine if resistance of the pathogen to the fungicide has developed, leaf disk assays were conducted to compare sensitivity of P. humuli isolates from the different U.S. hop-growing regions to isolates from a research yard where exposure to fosetyl-Al had not occurred for at least 10 years. Dose response curves of transformed data were linear. The fosetyl-Al concentration effective against 50% of the P. humuli isolates (ED 50 ) from each location was estimated from the linearized data. The ED 50 values indicate that fosetyl-Al was about one-third as effective against P. humuli isolates from commercial hop yards in northern Idaho and Oregon and about one-half as effective against isolates from southern Idaho compared to isolates from the research yard. Commercial yards in Washington were similar to the research yard. Accepted for publication 3 August 2004. Published 11 August 2004.

Powdery mildew of hop (Humulus lupus L.), which is caused by Podosphaera macularis (formerly Sphaerotheca macularis) was found in the Yakima Valley, WA in 1996 and subsequently spread to the growing regions in Oregon and northern and southern Idaho. To rapidly assist growers in reducing the cost associated with the preventive fungicide program, the Gubler/Thomas grape powdery mildew risk infection model was adapted for hops. In addition, field surveys were utilized to identify other management practices that impacted disease development. Weather networks were established and utilized to deliver daily regional maps indicating the risk index. These maps were posted to the web for daily access. Lessons learned from this experience will be useful in addressing future pathogen introductions. Accepted for publication 28 March 2003. Published 13 November 2003.