The hop downy mildew pathogen Pseudoperonospora humuli

Pseudoperonospora humuli is an obligate biotrophic oomycete that causes downy mildew, one of the most devastating diseases of cultivated hop, Humulus lupulus. Downy mildew occurs in all production areas of the crop in the Northern Hemisphere and Argentina. The pathogen overwinters in hop crowns and roots, and causes considerable crop loss. Downy mildew is managed by sanitation practices, planting of resistant cultivars, and fungicide applications. However, the scarcity of sources of host resistance and fungicide resistance in pathogen populations complicates disease management. This review summarizes the current knowledge on the symptoms of the disease, life cycle, virulence factors, and management of hop downy mildew, including various forecasting systems available in the world. Additionally, recent developments in genomics and effector discovery, and the future prospects of using such resources in successful disease management are also discussed.TaxonomyClass: Oomycota; Order: Peronosporales; Family: Peronosporaceae; Genus: Pseudoperonospora; Species: Pseudoperonospora humuli.Disease symptomsThe disease is characterized by systemically infected chlorotic shoots called “spikes". Leaf symptoms and signs include angular chlorotic lesions and profuse sporulation on the abaxial side of the leaf. Under severe disease pressure, dark brown discolouration or lesions are observed on cones. Infected crowns have brown to black streaks when cut open. Cultivars highly susceptible to crown rot may die at this phase of the disease cycle without producing shoots. However, foliar symptoms may not be present on plants with systemically infected root systems.Infection processPathogen mycelium overwinters in buds and crowns, and emerges on infected shoots in spring. Profuse sporulation occurs on infected tissues and sporangia are released and dispersed by air currents. Under favourable conditions, sporangia germinate and produce biflagellate zoospores that infect healthy tissue, thus perpetuating the infection cycle. Though oospores are produced in infected tissues, their role in the infection cycle is not defined.ControlDowny mildew on hop is managed by a combination of sanitation practices and timely fungicide applications. Forecasting systems are used to time fungicide applications for successful management of the disease.Useful Websites https://content.ces.ncsu.edu/hop‐downy‐mildew (North Carolina State University disease factsheet), https://www.canr.msu.edu/resources/michigan‐hop‐management‐guide (Michigan Hop Management Guide), http://uspest.org/risk/models (Oregon State University Integrated Plant Protection Center degree‐day model for hop downy mildew), https://www.usahops.org/cabinet/data/Field‐Guide.pdf (Field Guide for Integrated Pest Management in Hops).     

Epidemiology and control of powdery mildew (Sphaerotheca pannosa) on roses

In examinations between September 1966 and December 1968 of 741 specimens of rose species and cultivars, cleistocarps of Sphaerotheca pannosa were found on thirty-two cultivars, mostly ramblers and old shrub roses, and on nine of these they were found in two or three successive seasons. On stem pieces placed on soil in each of the winters 1966-7, 1967-8 and 1968-9 the number of cleistocarps with asci and ascospores decreased during November and December and rose slightly in January, but none showed dehiscence. Cleistocarps on rose bushes examined during the winters of 1967-8 and 1968-9 showed a progressive degeneration of ascospores, and by December none was found. Perennation of S. pannosa in buds was demonstrated by field observations, by inducing bursting of dormant buds on surface-sterilized shoots and by dissection of dormant apical buds. In field studies in 1968 of the development of mildew, infected buds were noted on 22 March but secondary infections did not appear until 17 April, though viable conidia and susceptible leaf tissue were present during this period. Low temperatures appeared to be partly responsible for this lag. On detached leaves in the laboratory the fungus developed from germination to sporulation in 4 days at 20°, 7 days at 15°, 11 days at 10° and 28 days at 3°. Keeping inoculated detached leaves at 0° for 10 days apparently did not affect the viability of the conidia. In both 1967 and 1968 there were two host growth periods, each culminating in flowering, between May and September; mildew did not develop on the shoots until the second growth phase, then the disease increased logarithmically on shoots and blooms during August and September. The disease on the shoots was effectively controlled in the field during 1968 by applications of ‘Benlate’ (benomyl) or dinocap, but not by methy-rimol; these fungicides were less effective in controlling mildew on pedicels. Laboratory tests showed that ‘Benlate’ inhibited sporulation of S. pannosa by deforming the conidiophores.     

Infection periods in relation to the natural development of hop downy mildew (Pseudoperonospora humuli)

The relationships between temperature and surface wetness and subsequent infection of hop tissues by P. humuli were examined on potted plants and detached leaves kept in temperature-controlled growth rooms. Periods of wetness which would just allow leaf infection ranged from 1 1/2 h at 30d? to 24 h at 5d?. The corresponding ranges for shoots were: light infection, 3 h at 19–23d? to 6 h at 8–10d?; severe infection, 4 h at 19–23d? to 8 h at 12–13d?. These data were used to relate the development of downy mildew in an unsprayed hop garden during 1967 and 1968 to periods with temperature/surface wetness suitable for minimum (minor infection periods) and severe infection (major infection periods). In 1967 a sudden outbreak of infected basal shoots (spikes) was related to an isolated major infection period. By contrast, early in 1968, major shoot infection periods did not arise and spikes appeared gradually in response to a succession of minor infection periods. More spikes were formed in 196 than in 1967; this was not related to the incidence of infection periods but probably reflected the relatively higher concentrations of airborne sporangia early in 1968. In both years outbreaks of leaf and lateral shoot infection could be traced to major infection periods caused by rain; sudden disease increases again originated from isolated infection periods. There was a close similarity between the incubation period for each principal disease outbreak and that expected from growth-room experiments. Major infection periods occurred more frequently at the end of June 1968, resulting in a higher final concentration of diseased tissue than in 1967. Predicted major infection periods failed to induce large disease increases when dew alone provided wetness or when no airborne sporangia could be detected.     

Biology of the hop cyst nematode Heterodera humuli

In an intensively sampled English hop garden, cysts of Heterodera humuli were found in every sample taken. Most occurred in the top 15 cm of soil, decreasing in number by half with every 15 cm in depth. Numbers of second-stage juveniles fluctuated during the vegetative period in a pattern which indicated that more than one generation, possibly two to three, were produced. The generation time was about 1.5 months. In pot experiments under controlled conditions (20°C; 16 h light/day) the life cycle lasted 40 days. The optimal temperature was 20°C for the development of H. humuli in the roots of its host Humulus lupulus. Most H. humuli juveniles invaded hop roots at 15°C, but egg hatch was greatest at 20°C. In moist soil, second-stage juveniles survived for at least 54 days and thereafter invaded roots and reproduced.     

硅和白粉菌诱导接种对黄瓜幼苗白粉病抗性影响的研究

研究了硅酸盐和诱导接种白粉菌对黄瓜活性氧代谢、SiO2含量和抗病性的影响．结果表明,诱导接种能使叶片的超氧自由基(O2^-)产生速率、H2O2和丙二醛(MDA)含量升高,加硅接种处理的O2^-产生速率、H202和MDA含量明显低于不加硅接种处理．诱导接种能使叶片的过氧化氢酶(CAT)、过氧化物酶(POD)活性升高,超氧化物歧化酶(SOD)活性降低．加硅接种处理植株叶片的CAT、POD和SOD活性明显高于不加硅接种处理．诱导接种提高叶片的抗坏血酸(AsA)和还原型谷胱甘肽(GSH)含量,加硅处理的AsA含量明显低于不加硅处理,GSH含量高于不加硅处理．无论接种与否,加硅处理的SiO2含量显著高于不加硅处理,病情指数明显低于不加硅处理．     

Flight activity of the damson–hop aphid, Phorodon humuli.

Flight activity of Phorodon humuli was monitored using suction traps, laboratory studies and mark and recapture experiments. Emigrants were trapped as they flew from a Myrobalan (Prunus cerasifera) hedge and among dwarf hops (Humulus lupulus). Daily flight curves were bimodal with 69% and 38% of emigrants caught in the morning peak near Myrobalan and among hops, respectively. The median period of flight activity was from 2 h after sunrise until 30 min before sunset. The lower temperature for flight was 13.5°C in the field and 14.9°C for take off in the laboratory. Variations in wind speed had little effect on flight activity explaining <2.5% of the total variance among insect counts. The percentage of emigrants on hop declined exponentially with time. The relationship, y= 10.9(±2.0) + 64.3(±2.3) × 0.92(±0.01)^t where t = daylight hours (standard error in parentheses), explained 98.3% of the variance. Hence, 62% of new arrivals flew within 1 day of arrival and 79% within 2 days. Similar numbers arrived as departed at 08:30, 10:30 and 12:30 h, but at 14:30 h twice as many arrived than departed and at 16:30 h, the accumulation was threefold. Daily flight curves of return migrants and males leaving hop were bimodal with 70% and 80%, respectively, trapped in the earlier peak. In the field, the median lower temperature for flight was 13.2°C for return migrants and a nonsignificantly different 12.8°C for males. The mean temperature for take off by return migrants was 15.7°C in the laboratory.     

Overwintering of Sphaerotheca mors-uvae on black currant and gooseberry

The ability of Sphaerotheca mors-uvae to perennate as cleistocarps, and as mycelium in buds was examined during the winters of 1965-6, 1966-7 and 1967-8 in relation to its two principal hosts, gooseberry and black currant. Cleistocarps on black currant leaves were examined from August 1965 to April 1966 and from July 1966 to March 1967. In 1965 cleistocarps were first observed on the leaves on 5 August; in 1966 on 11 July. These continued to develop through August and September and by October approximately 70% contained well-defined ascospores. The ascospore content remained generally at this level until February 1966 and November 1966; then the numbers of cleistocarps with ascospores fell and by April 1966 and March 1967 few such cleistocarps remained. From 21 March 1966 and 15 February 1967, but not otherwise, discharge of ascospores from the overwintered cleistocarps was readily obtained in laboratory tests. The viability and infectivity of the ascospores was demonstrated by allowing them to discharge on to leaf discs of black currant in the laboratory and also on to leaf discs and plants in the field. Sporulating colonies of S. mors-uvae developed within 8 days. Cleistocarps from shoots of black currant were examined from 4 August 1966 to 9 March 1967, and from 27 July 1967 to 1 January 1968. They developed in a similar manner to those on black currant leaves and by September in both 1966 and 1967 over 60% contained ascospores. This level was not maintained; the number of cleistocarps with ascospores fell gradually and by 8 December 1966 and 1 January 1968 few remained. Only in one laboratory test (21 November 1967) were ascospores discharged from a sample of these cleistocarps. Cleistocarps from shoots of gooseberry were examined from July 1966 to March 1967, and from August 1967 to January 1968. The pattern of ascospore development and subsequent decline in number of cleistocarps with ascospores was similar to that observed for black currant shoots. No discharge of ascospores could be demonstrated in laboratory tests. Evidence that S. mors-uvae perennates in buds of gooseberry was obtained by dissecting buds and by inducing buds on surface-sterilized shoots to burst under conditions which precluded chance infection. Field observations also suggested that bud infection occurred on gooseberry. Similar experiments failed to demonstrate the fungus in buds of black currant, and there was no indication of bud infection of this host in the field.     

Insecticide resistance in damson-hop aphid, Phorodon humuli in commercial hop gardens in Kent

From 1966 to 1976, samples of Phorodon humuli were collected annually from five commerical hop gardens in Kent and from other hop gardens where problems in control occurred. A susceptible stock was obtained from wild hop growing in northern England in 1969. The samples were cultured in isolation on potted hops and bioassayed against insecticides in common use. The level of resistance to demeton-S-methyl was c. 10X in 1966 after 10 yr use, and more than doubled from 1968–1974 apparently due to the spread of a more resistant type; there was a further increase to c. 50X in 1975–1976. There was also resistance of C. 20–30X to omethoate, 2–7X to methidathion and 4–8X to methomyl. Assays and field results show an increased resistance to methidathion and less certainly to methomyl after 5 yr use. There was no clear change in response to endosulfan. The LC₅₀'s estimated from a single dose and a mean probit slope were found to agree satisfactorily with the LC₅₀'s calculated from serial doses and so should be adequate for monitoring trends in resistance.     

Evaluation of the activity of a number of procaine derivatives and analogous compounds against powdery mildew (Sphaerotheca fuliginea) on cucumber*

About fifty derivatives and analogues of procaine have been prepared and tested in three different tests for activity against powdery mildew of cucumber. Most of the compounds were active but few equalled procaine. The results obtained seem to afford little evidence for simple structure-activity relationship based on physico-chemical or electro-chemical properties.     

Use of potassium bicarbonate (Armicarb) on the control of powdery mildew (Sphaerotheca mors-uvae) of gooseberry (Ribes uva-crispa)

Powdery mildew (Sphaerotheca mors-uvae) severely infects young shoots, stems and fruits of gooseberry (Ribes uva-crispa). Environmental friendly and biological control measures are being sought throughout the world. Especially in organic gooseberry growing effective control measures are needed, because powdery mildew infections may result in a total loss of the crop. In organic currant growing the number of adequate control methods is very limited. Sulphur as a fungicide against powdery mildew in e.g. gooseberry or table grape growing is not recommendable due to possible bleaching of berries and scorching of tender shoots. Various bicarbonate salts are suggested as a good option to control powdery mildew. In a field trial the effect of potassium bicarbonate (Armicarb) on the on the control of powdery mildew of gooseberry was evaluated. Four treatments; i.e. two preventive strategies and two curative strategies, were applied. PLants were sprayed until runoff. The percent of infected fruits and disease severity were assessed. In the unsprayed control plots very high disease incidences were observed; on average more than 90% of the berries were infected with powdery mildew. The weekly (preventive) spray applications with potassium carbonate reduced the disease incidences on fruits significantly. On average approximately 10% of the fruits were affected by powdery mildew. However, the number of spray applications was high. In conclusion, our results indicate that applications of potassium bicarbonate (as Armicarb) are effective in reducing the incidence and severity of American powdery mildew in gooseberry. Early spray applications are necessary to protect berries against powdery mildew infections. Future research will focus on reducing the number of applications, e.g. warning models based on powdery mildew of rose (Sphaerotheca pannosa).     

Observations on the effect of black root rot (Phomopsis sclerotioides) on the susceptibility of cucumber to powdery mildew (Sphaerotheca fuliginea)

Powdery mildew (Sphaerotheca fuliginea) infection of cucumber leaves was earlier and increased more rapidly on plants with black root rot (Phomopsis sclerotioides) than on plants with healthy root systems in the same glasshouse.     

Field experiments on sugar-beet powdery mildew, Erysiphe betae

Sugar-beet powdery mildew, Erysiphe betae appeared in East Anglia in late July 1976 and became wide-spread in eastern England during August and September but was scarce in crops in the north and west of England. Fentinhydroxide, sulphur, benomyl and ethirimol controlled the disease but benomyl applied once only and ethirimol were less effective than the other materials. In heavily infected crops two sulphur sprays, the first applied at the onset of the mildew attack, increased sugar yield by 13%. A single early sulphur spray increased yield on average by 9% giving a return of six to seven times the cost of treatment. When the first spray was delayed, mildew control was less effective.