The etiology of hop chlorotic disease

Hop chlorotic disease was first described in England in 1930, but it has since been seldom seen and its etiology has remained unknown. In 1983 a patch of plants with the disease occurred in a large area of hops (Humulus lupulus) cv. Bramling Cross planted at Yalding, Kent in 1967. All plants in a rectangular area enclosing the disease outbreak were infected with hop mosaic, hop latent and prunus necrotic ringspot viruses; the diseased plants were additionally infected with arabis mosaic virus (AMV). The disease was also associated with seed-transmitted AMV, and was induced in hop seedlings inoculated with partially purified preparations of AMV originating from chlorotic disease-affected hops prepared from Chenopodium quinoa. The disease appears to be caused by AMV, but AMV isolates from hops with chlorotic disease were serologically indistinguishable from AMV isolates from hops with symptoms of bare-bine and/or nettlehead and showed similar pathogenicity in diagnostic hosts. The basis of the difference between isolates in their pathogenicity in hop remains unknown.     

Arabis mosaic and Prunus necrotic ringspot viruses in hop (Humulus lupulus L.)

Purified virus preparations made from nettlehead-diseased hop plants, or from Chenopodium quinoa, to which the virus was transmitted by inoculation of sap, contained polyhedral virus particles of 30 mμ diameter which were identified serologically as arabis mosaic virus (AMV). There were serological differences between AMV isolates from hop and from strawberry, and also differences in host range and in symptoms caused in C. quinoa and C. amaranticolor. AMV was always associated with nettlehead disease. The nematode Xiphinema diversicaudatum occurred in small numbers in most hop gardens, but was numerous where nettlehead disease was spreading rapidly. Preparations from nettlehead-affected hops also contained a second virus, serologically related to Prunus necrotic ringspot virus (NRSV), in mild and virulent forms which infected the same range of test plants but showed some serological differences. Mild isolates did not protect C. quinoa plants against infection by virulent isolates. Hop seedlings inoculated with virulent isolates of NRSV developed symptoms indistinguishable from those of split leaf blotch disease. Latent infection with NRSV was prevalent in symptomless hop plants. Nettlehead disease is apparently associated with dual infection of AMV and virulent isolates of NRSV. An unnamed virus with rod-shaped particles 650 mμ long was common in hop and was transmitted by inoculation of sap to herbaceous plants. Cucumber mosaic virus was obtained from a single plant of Humulus scandens Merr.     

Transmission of the hop strain of arabis mosaic virus by Xiphinema diversicaudatum*

Hop plants became infected with the hop strain of arabis mosaic virus (AMV(H)) when grown in hopfield and woodland soil in which infected plants had been growing. Infection occurred in soil infested with the dagger nematode Xiphinema diversicaudatum, but neither in uninfested soil nor in soil previously heated to kill nematodes. X. diversicaudatum transferred direct from hop soils transmitted AMV(H) to young herbaceous plants and to hop seedlings; some of the hop seedlings developed nettlehead disease. A larger proportion of plants was infected using X. diversicaudatum obtained from a woodland soil and then given access to the roots of hop or herbaceous plants infected with AMV(H). AMV(H) was transmitted by adults and by larvae, in which the virus persisted for at least 36 and 29 wk, respectively. Difficulties were encountered in detecting AMV(H) in infected hop plants, due partly to the delay in virus movement from roots to shoots. Infection of hop shoots was seldom detected until the year after the roots were infested and sometimes nettlehead symptoms did not appear until the third year. Isolates of arabis mosiac virus from strawberry did not infect hop. The results are discussed in relation to the etiology and control of nettlehead and related diseases of hop.     

Hop line-pattern virus in relation to the etiology and distribution of nettlehead disease

Hop line-pattern virus (HLPV) was transmissible by mechanical inoculation to hop plants; it induced characteristic severe symptoms in Humulus lupulus L. var. neo-mexicanus Nels. & Cockerell and the commercial derivatives College Cluster and Keyworth's Midseason, but none in the traditional English varieties of H. lupulus (e.g. Fuggle).
Mechanical transmission of hop nettlehead virus (HNV) was facilitated by the presence of HLPV in the test plants; hop seedlings and clonal plants escaped infection by sap inoculum that infected plants of two varieties already infected with HLPV. HNV was also transferred by stem contact and by knife cuts to plants carrying HLPV.
Infection with HLPV was latent in twelve nettlehead-diseased Fuggle plants from different fields, and in diseased and symptomless plants in a nettlehead outbreak in W.G.V., a variety that previously had escaped infection. It is suggested either that HLPV predisposes hop plants to infection with HNV or that nettlehead disease is caused by dual infection with both viruses.
Localized and scattered patterns of nettlehead spread were observed in hop plantations; these two types are usually attributed to different modes of spread which would be compatible with a complex etiology of the disease.     

New and resurgent insect pests on low trellis hops

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Simultaneous Detection of Three Viroid Species Infecting Hops in China by Multiplex RT‐PCR

We have developed a multiplex RT‐PCR protocol for the simultaneous detection of three viroids in three different genera that infect hops: Hop latent viroid (HLVd; Cocadviroid), Hop stunt viroid (HSVd; Hostuviroid) and Apple fruit crinkle viroid (AFCVd; Apscaviroid). The method was validated by testing 175 hop samples collected from the Xinjiang autonomous region of China. All samples were found to be positive for HLVd but negative for AFCVd, confirming the widespread or even ubiquitous infection of HLVd and the low incidence of AFCVd in hops in China. In addition, HSVd was detected in 22.86% of the samples tested. This rapid and reliable multiplex RT‐PCR assay provides an effective method for detection of three important viroid species in large‐scale surveys for disease management in hops.     

Cultivated Grapevines Represent a Symptomless Reservoir for the Transmission of Hop Stunt Viroid to Hop Crops: 15 Years of Evolutionary Analysis

Hop stunt was a mysterious disorder that first emerged in the 1940s in commercial hops in Japan. To investigate the origin of this disorder, we infected hops with natural Hop stunt viroid (HpSVd) isolates derived from four host species (hop, grapevine, plum and citrus), which except for hop represent possible sources of the ancestral viroid. These plants were maintained for 15 years, then analyzed the HpSVd variants present. Here we show that the variant originally found in cultivated grapevines gave rise to various combinations of mutations at positions 25, 26, 54, 193, and 281. However, upon prolonged infection, these variants underwent convergent evolution resulting in a limited number of adapted mutants. Some of them showed nucleotide sequences identical to those currently responsible for hop stunt epidemics in commercial hops in Japan, China, and the United States. Therefore, these results indicate that we have successfully reproduced the original process by which a natural HpSVd variant naturally introduced into cultivated hops was able to mutate into the HpSVd variants that are currently present in commercial hops. Furthermore, and importantly, we have identified cultivated grapevines as a symptomless reservoir in which HSVd can evolve and be transmitted to hop crops to cause epidemics.     

Hopped Beer: The Case For Cultivation

The history of hops, hopped beer, and hop cultivation is unclear and ambiguous. An assessment of the available literature reveals many contradictions, especially regarding the first use of hops in beer and the earliest incidence of hop cultivation. Historically, hops were used for a variety of purposes; now their primary use is as a preservative and flavoring in beer. Hop cultivation is poorly documented, but was certainly undertaken by the 10th century, most probably in response to the demand generated by beer-brewing. After comparing the literature and investigating source material, a chronology of hop use in beer and hop cultivation is proposed.     

Alfalfa mosaic virus isolates from lucerne in South Australia: biological variability and antigenic similarity

Alfalfa mosaic virus (AMV) was isolated from lucerne (Medicago sativa) plants with a variety of disease symptoms in eight of 13 sites in South Australia indicating that the virus is widespread in the state. The host ranges and symptomatology of the virus isolates varied considerably. Twelve selected local lesion isolates were shown to be distinct when mechanically inoculated to a range of plant species and cultivars. However, agar-gel diffusion and enzyme-linked immunoassay tests with polyclonal antisera prepared against glutaraldehyde-fixed virus preparations of the five most readily distinguishable AMV isolates, failed to reveal significant antigenic differences between the 12 virus isolates. This indicates that serological tests with polyclonal antisera can detect a wide range of AMV variants but would be unlikely to differentiate between strains. The wide host range and variability of AMV precluded the grouping of isolates into strains of the virus.     

Strains of Prunus necrotic ringspot virus in hop (Humulus lupulus L.)

Purified preparations of Prunus necrotic ringspot virus (NRSV) from hop plants formed two light-scattering zones when centrifuged in sucrose density gradients; the upper and lower zones contained particles 25 mμ and 31 mμ in diameter respectively whose sedimentation coefficients were 79 S and 107 S. NSRV isolates from hop were of two distinct serological types: ‘A’ strains, serologically very closely related to NRSV isolates from apple; and ‘C’ strains more nearly related to NRSV from cherry. The variety Fuggle is tolerant to hop mosaic (not related to NRSV) and different selections of apparently healthy female plants usually contained A strains; but C strains were usually isolated from nettlehead-diseased plants. Either A or C strains occurred in male plants grown with the hop-mosaic tolerant varieties. In mosaic-sensitive varieties (Goldings and Bramlings) apparently healthy female plants tested were usually infected with C strains; either A or C types occurred in mosaic-sensitive male plants. NRSV was not detected in the seventy-four hop seedlings obtained from virus-infected plants. Some varieties developed nettlehead when infected with NRSV (A) or (C) + the hop form of arabis mosaic virus, but not with NRSV (A) or (C) alone. Others developed nettlehead when infected with arabis mosaic virus + NRSV (C) but not with arabis mosaic + NRSV (A). A and C strains can multiply together in the same hop plant. There is evidence of partial antagonism, however, and the fluctuating behaviour of the nettlehead syndrome probably reflects changes in the relative concentration of the two serotypes.     

A Survey of German Hops for the Presence of American Hop Latent Virus (AHLV)

The presence of rod-shaped viruses in German hops has been known for a long time and Hop Mosaic Virus (HMV) and Hop Latent Virus (HLV) were proved to be present by serological techniques. When another CARLA-Virus, the American Hop Latent Virus was detected in England an extensive survey was carried out in the German hop growing regions and among wild hops. No natural occurrence of the virus could be detected. Only recently introduced plants of the American clone USDA 21055 in an isolated breeding garden were infected. Spread of the virus to German seedlings could not be detected.     

Seed-borne alfalfa mosaic virus infecting annual medics (Medicago spp.) in Western Australia

Infection with alfalfa mosaic virus (AMV) was widespread in introduction, evaluation and seed increase plots of cultivars and numbered selections of annual medics (Medicago spp.) in Western Australia; the virus was detected in plots of seven species. When seed stocks from the West Australian annual medic collection harvested in 1984–1986 were sown and seedlings tested, seed-borne AMV was found in all 12 cultivars and in 44/50 numbered selections, belonging to 10 species. Seed transmission rates to seedlings ranged from 0.3–74% and exceeded 5% in 33 seed lots. By contrast, when seedlings of four species grown from seed harvested in 1971–1978 were tested, no AMV was detected; the oldest infected seed stock found was from 1980. In commercial seed stocks of two cultivars released in 1987, the levels of seedling infection with AMV found were 0–0.2% for M. polymorpha cv. Santiago and 526% for M. murex cv. Zodiac. In commercial 1986 seed of M. polymorpha cvs Serena and Circle Valley, AMV was detected in 3/13 and 6/9 stocks respectively; transmission rates to seedlings in infected stocks were 0.1–0.7%. In a survey of 47 annual medic pastures in medium and low rainfall zones of the Western Australian wheat belt in 1987, the virus was detected in leaf samples from only three sites. When inoculated mechanically, AMV systemically infected 11 cultivars and 12 selections belonging to 13 species, but did not infect one selection each of M. aculeata and M. orbicularis. Infected plants in ten species developed only faint mosaics or were symptomlessly infected, but M. littoralis, M. polymorpha and M. tornata developed distinct mottling, reduction in leaf size and, in some instances, leaf deformation and dwarfing. In pot tests, AMV infection decreased herbage and root production (dry wts) of M. polymorpha cvs Serena and Circle Valley by about 30% and 50–60% respectively, but did not decrease herbage production in M. murexcv. Zodiac. In spaced plants growing outside, AMV decreased herbage, root (dry wts) and seed production of M. polymorpha cvs Circle Valley and Santiago by about 60%.     

Infectibility of some new raspberry cultivars with arabismosaic and raspberry ringspot viruses and further evidence for variation in British isolates of these two nepoviruses

In field trials at sites of an outbreak of arabis mosaic nepovirus (AMV) in England and of raspberry ringspot nepovirus (RRV) in Scotland, the results of exposure of some new raspberry cultivars to natural infection with these viruses showed discrepancies from those obtained in graft inoculation tests using AMV-Lib and RRV-S, the Scottish type isolates. In particular, cv. Glen Prosen, which is immune to AMV-Lib and RRV-S, was infected with AMV and RRV in the field trials. Studies on these and other field isolates of AMV and RRV showed that they differed from the type isolates in Rubus host range and in symptomatology in herbaceous hosts. However, whereas four isolates of RRV found infecting Rubus were distinguishable by spur formation in gel double-diffusion serological tests, six AMV isolates were indistinguishable by this method. Immunoelectrophoresis of virus particles did not distinguish the six AMV isolates, but isolates RRV-MX and RRV-T were distinguishable from RRV-S and the English type isolate, RRV-E. Like the two RRV type isolates, RRV-MX contained a single electrophoretic component, but it migrated must faster whereas RRV-T contained two components, one with a migration rate similar to that of RRV-MX and the other similar to that of the type isolates. Polyacrylamide gel electrophoresis of protein preparations from highly purified virus particles of RRV isolates E, S and MX detected a single polypeptide of estimated mol. wt 54 × 10³, 54 × 10³ and 50 × 10³ respectively but that of isolate T contained two polypeptides of estimated mol. wt 54 × 10³ and 50 × 10³. These data suggest that RRV-T is a mixture of two isolates. In laboratory tests the nematode Xiphinema diversicaudatum transmitted four isolates of AMV efficiently whereas two populations of the nematode Longidorus elongatus were less efficient vectors of four RRV isolates. Neither vector species transmitted virus to any of nine raspberry cultivars. The results are discussed in relation to the control of nepoviruses in raspberry and to the biology of these viruses.     

The relationship between the Andean strain of potato virus S and pepino latent virus

The titres obtained in microprecipitin tests with purified preparations of pepino latent virus (PepLV) and the Andean strain of potato virus S (PVS^A) using PepLV antiserum and two antisera to the ordinary strain of PVS (PVS°) indicated a close serological relationship between PepLV and PVS^A. Using antiserum to PVS°, both viruses were detected by ELISA when infective Chenopodium quinoa sap was diluted to 10^-5but not to 10^-6. Particles of both viruses were decorated equally well by antibodies to PVS^o, PVS^Aand PepLV in all virus-antiserum combinations. When PepLV was inoculated to C. quinoa, C. amaranticolor and potato plants, the symptoms induced closely resembled those of PVS^Ain these hosts. It is concluded that PepLV is an isolate of PVS^Afrom pepino.     

The isolation and identification of rhubarb viruses occurring in Britain

Virus-like symptoms were common in British crops of rhubarb. All plants tested of the three main varieties, ‘Timperley Early’, ‘Prince Albert’ and ‘Victoria’, were virus-infected. Turnip mosaic virus and a severe isolate of arabis mosaic virus (AMV) were obtained from ‘Timperley Early’; and ‘Prince Albert’ contained turnip mosaic virus, cherry leaf roll virus (CLRV), a mild isolate of AMV and, infrequently, cucumber mosaic virus (CMV). The main commercial variety ‘Victoria’ contained turnip mosaic virus, CLRV, a mild isolate of AMV and, infrequently, strawberry latent ringspot virus (SLRV). All the viruses were identified serologically. The rhubarb isolates did not differ markedly from other isolates of these viruses in herbaceous host reactions, properties in vitro or particle size and shape. A rhubarb isolate of CLRV was distinguished serologically from a cherry isolate of the virus. Turnip mosaic virus, CLRV and SLRV, were transmitted with difficulty, but AMV isolates were readily transmitted by mechanical inoculation. Turnip mosaic virus was also transmitted to rhubarb by Myzus persicae and Aphis fabae. CLRV was transmitted in 6–8% of the seed of infected ‘Prince Albert’ and ‘Victoria’ rhubarb and in 72% of the seed of infected Chenopodium amaranticolor. Mild isolates of AMV were also transmitted in 10–24% of the seed of infected ‘Prince Albert’ and ‘Victoria’ plants.     

The Influence of Starch and Fibre on In Vitro Protein Digestibility of Dry Fractionated Quinoa Seed (Riobamba Variety)

The in vitro gastric digestibility of the quinoa variety Riobamba was investigated, especially the influence of the quinoa matrix. Dry-fractionated quinoa protein concentrate, which is just milled and sieved, was much better digestible than the same concentrate that was reconstituted from wet fractionated quinoa protein isolate, quinoa starch isolate, and quinoa fibre isolate. In the reconstituted concentrate, the presence of quinoa starch and fibre next to quinoa protein reduces its in vitro gastric digestibility significantly. However, the effect of starch is partially counteracted if the fibre is also present. While the effects of starch and fibre separately can be understood from the decrease of the accessibility for pepsin to hydrolyse proteins, due to the hydrated starch and fibre, we suspect that the synergistic effect of starch and fibre may be due to a relative reduction of the hydration of starch due to the presence of the also strongly hydrating fibre. We concluded that the presence of starch and fibre decreases the protein in vitro gastric digestibility. Therefore, the presence of fibre partially countered the decreased of the protein digestibility of starch. Heating of the matrices to 120 ^°C generally resulted in much lower digestion rates, due to extensive aggregation of the protein.

     

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.     

Proteomic Approach for Characterization of Hop-Inducible Proteins in Lactobacillus brevis

Resistance to hops is a prerequisite for the capability of lactic acid bacteria to grow in beer and thus cause beer spoilage. Bactericidal hop compounds, mainly iso-α-acids, are described as ionophores which exchange H⁺ for cellular divalent cations, e.g., Mn²⁺, and thus dissipate ion gradients across the cytoplasmic membrane. The acid stress response of Lactobacillus brevis TMW 1.465 and hop adaptation in its variant L. brevis TMW 1.465A caused changes at the level of metabolism, membrane physiology, and cell wall composition. To identify the basis for these changes, a proteomic approach was taken. The experimental design allowed the discrimination of acid stress and hop stress. A strategy for improved protein identification enabled the identification of 84% of the proteins investigated despite the lack of genome sequence data for this strain. Hop resistance in L. brevis TMW 1.465A implies mechanisms to cope with intracellular acidification, mechanisms for energy generation and economy, genetic information fidelity, and enzyme functionality. Interestingly, the majority of hop-regulated enzymes are described as manganese or divalent cation dependent. Regulation of the manganese level allows fine-tuning of the metabolism, which enables a rapid response to environmental (stress) conditions. The hop stress response indicates adaptations shifting the metabolism into an energy-saving mode by effective substrate conversion and prevention of exhaustive protein de novo synthesis. The findings further demonstrate that hop stress in bacteria not only is associated with proton motive force depletion but obviously implies divalent cation limitation.     

A damaging outbreak of arabis mosaic nepovirus in blackcurrant, the occurrence of other nepoviruses in Ribes species, and the demonstration that alfalfa mosaic virus is the cause of interveinal white mosaic in blackcurrant

In a crop of blackcurrant (Ribes nigrum), cv. Baldwin in Eire, chlorotic mottling and ringspot symptoms in leaves on plants and severe crop loss was associated with infection with arabis mosaic nepovirus (ArMV) and the presence in the soil of its nematode vector, Xiphinema diversicaudatum. This is only the second report of ArMV damaging a crop of blackcurrant. Tomato black ring (TBRV) and raspberry ringspot nepoviruses were detected in single plants of redcurrant (R. rubrum) in England and flowering currant (R. sanguineum) in Scotland respectively; each of these infected plants showed foliar chlorotic line-pattern symptoms. This is the first record of TBRV in redcurrant. A single blackcurrant plant in New Zealand showing symptoms typical of those described for interveinal white mosaic disease, contained alfalfa mosaic virus (AMV). When AMV particles were purified and concentrated from herbaceous test plants and mechanically inoculated to young blackcurrant plants, several became infected with AMV and most infected plants developed systemic symptoms typical of the original disease. This provides the strongest evidence to date that AMV is the causal agent of interveinal white mosaic disease.     

Evaluation of a damage threshold for two-spotted spider mites, Tetranychus urticae Koch (Acari: Tetranychidae), in hop culture

Damage caused by two‐spotted spider mites (Tetranychus urticae) at harvest to yield, quality (measured in percentage α‐acids content) and cone infestation was assessed on hop cvs Hallertauer Magnum, Hallertauer Tradition and Perle. Acaricide‐untreated hop plants with known levels of T. urticae infestation were compared with neighbouring acaricide‐treated plants. Although in 24 of the 36 experimental harvests the untreated hop plants had spider mite infestations of > 100 mites leaf^?1, yields and α‐acids content from the untreated plants were significantly lower than the treated plants in only four instances. However, although mite infestation of cones from untreated hops were significantly higher than acaricide‐treated plants in 27 of the 36 cases, in only one instance did that cause economic loss. Spider mite infestation levels of c. 90 mites leaf^?1 are tolerable at harvest time with little or no risk of causing economic loss to hop growers.