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Datasheet

Plasmopara viticola
(grapevine downy mildew)

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Datasheet

Plasmopara viticola (grapevine downy mildew)

Summary

  • Last modified
  • 22 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Plasmopara viticola
  • Preferred Common Name
  • grapevine downy mildew
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Chromista
  •     Phylum: Oomycota
  •       Class: Oomycetes
  •         Order: Peronosporales

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Pictures

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PictureTitleCaptionCopyright
Young, translucent oilspots with brown margins.
TitleSymptoms
CaptionYoung, translucent oilspots with brown margins.
CopyrightMegan M. Kennelly
Young, translucent oilspots with brown margins.
SymptomsYoung, translucent oilspots with brown margins. Megan M. Kennelly
Multiple oilspots can coalesce and cover much of the leaf surface.
TitleSymptoms
CaptionMultiple oilspots can coalesce and cover much of the leaf surface.
CopyrightMegan M. Kennelly
Multiple oilspots can coalesce and cover much of the leaf surface.
SymptomsMultiple oilspots can coalesce and cover much of the leaf surface.Megan M. Kennelly
Oilspots become dry and necrotic in the centre, with a yellow margin which retains the ability to sporulate.
TitleSymptoms
CaptionOilspots become dry and necrotic in the centre, with a yellow margin which retains the ability to sporulate.
CopyrightMegan M. Kennelly
Oilspots become dry and necrotic in the centre, with a yellow margin which retains the ability to sporulate.
SymptomsOilspots become dry and necrotic in the centre, with a yellow margin which retains the ability to sporulate.Megan M. Kennelly
Oilspots eventually become almost completely necrotic.
TitleSymptoms
CaptionOilspots eventually become almost completely necrotic.
CopyrightMegan M. Kennelly
Oilspots eventually become almost completely necrotic.
SymptomsOilspots eventually become almost completely necrotic.Megan M. Kennelly
On older leaves oilspots are restricted by veins and form yellow-to-brown small, angular spots in a patchwork pattern.
TitleSymptoms
CaptionOn older leaves oilspots are restricted by veins and form yellow-to-brown small, angular spots in a patchwork pattern.
CopyrightMegan M. Kennelly
On older leaves oilspots are restricted by veins and form yellow-to-brown small, angular spots in a patchwork pattern.
SymptomsOn older leaves oilspots are restricted by veins and form yellow-to-brown small, angular spots in a patchwork pattern.Megan M. Kennelly
After a warm, humid night sporangiophores and sporangia appear as a bright white, fluffy growth on the undersides of leaves.
TitleSymptoms
CaptionAfter a warm, humid night sporangiophores and sporangia appear as a bright white, fluffy growth on the undersides of leaves.
CopyrightMegan M. Kennelly
After a warm, humid night sporangiophores and sporangia appear as a bright white, fluffy growth on the undersides of leaves.
SymptomsAfter a warm, humid night sporangiophores and sporangia appear as a bright white, fluffy growth on the undersides of leaves.Megan M. Kennelly
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
TitleSymptoms
CaptionPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
Copyright©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
SymptomsPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
TitleSymptoms
CaptionPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
Copyright©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
SymptomsPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
TitleSymptoms
CaptionPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
Copyright©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.
SymptomsPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf prior to incubation in a dark polythene bag overnight to encourage sporulation. Surrey, England. September 2020.©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
TitleSymptoms
CaptionPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
Copyright©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.
SymptomsPlasmopara viticola (grapevine downy mildew); Underside of grapevine leaf after incubation in a dark polythene bag overnight resulting in sporulation, the sporangiophores and sporangia appear as a white fluffly growth. Surrey, England. September 2020.©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Oilspots on the upper surface of a grapevine leaf. Surrey, England. September 2020.
TitleSymptoms
CaptionPlasmopara viticola (grapevine downy mildew); Oilspots on the upper surface of a grapevine leaf. Surrey, England. September 2020.
Copyright©Dr Philip Taylor (CABI)
Plasmopara viticola (grapevine downy mildew); Oilspots on the upper surface of a grapevine leaf. Surrey, England. September 2020.
SymptomsPlasmopara viticola (grapevine downy mildew); Oilspots on the upper surface of a grapevine leaf. Surrey, England. September 2020.©Dr Philip Taylor (CABI)
Downy mildew symptoms  on grapevine leaf.
TitleSymptoms
CaptionDowny mildew symptoms on grapevine leaf.
CopyrightThorsten Kraska, University of Bonn, Germany
Downy mildew symptoms  on grapevine leaf.
SymptomsDowny mildew symptoms on grapevine leaf. Thorsten Kraska, University of Bonn, Germany
P. viticola leaf sporulation symptoms (downy mildew) on grapevine leaf.
TitleSymptoms
CaptionP. viticola leaf sporulation symptoms (downy mildew) on grapevine leaf.
CopyrightThorsten Kraska, University of Bonn, Germany
P. viticola leaf sporulation symptoms (downy mildew) on grapevine leaf.
SymptomsP. viticola leaf sporulation symptoms (downy mildew) on grapevine leaf.Thorsten Kraska, University of Bonn, Germany
Infected inflorescences and young fruit become covered with white, fluffy spores.
TitleSymptoms
CaptionInfected inflorescences and young fruit become covered with white, fluffy spores.
CopyrightMegan M. Kennelly
Infected inflorescences and young fruit become covered with white, fluffy spores.
SymptomsInfected inflorescences and young fruit become covered with white, fluffy spores.Megan M. Kennelly
Close-up of infected young berry with sporulation on berry surface and pedicel.
TitleSymptoms
CaptionClose-up of infected young berry with sporulation on berry surface and pedicel.
CopyrightMegan M. Kennelly
Close-up of infected young berry with sporulation on berry surface and pedicel.
SymptomsClose-up of infected young berry with sporulation on berry surface and pedicel.Megan M. Kennelly
Berries infected after 2-3 weeks postbloom become discoloured and shrivel, but do not support sporulation.
TitleSymptoms
CaptionBerries infected after 2-3 weeks postbloom become discoloured and shrivel, but do not support sporulation.
CopyrightMegan M. Kennelly
Berries infected after 2-3 weeks postbloom become discoloured and shrivel, but do not support sporulation.
SymptomsBerries infected after 2-3 weeks postbloom become discoloured and shrivel, but do not support sporulation.Megan M. Kennelly
Infected areas usually turn brown, dry out, and fall off of the cluster
TitleSymptoms
CaptionInfected areas usually turn brown, dry out, and fall off of the cluster
CopyrightMegan M. Kennelly
Infected areas usually turn brown, dry out, and fall off of the cluster
SymptomsInfected areas usually turn brown, dry out, and fall off of the clusterMegan M. Kennelly
Young inflorescences may distort or curl (arrowed) - note healthy inflorescence behind.
TitleSymptoms
CaptionYoung inflorescences may distort or curl (arrowed) - note healthy inflorescence behind.
CopyrightMegan M. Kennelly
Young inflorescences may distort or curl (arrowed) - note healthy inflorescence behind.
SymptomsYoung inflorescences may distort or curl (arrowed) - note healthy inflorescence behind.Megan M. Kennelly
(1) Sporangium detached from sporangiophore; (2) A sporangium which has released its zoospores; (3) Encysted zoospore beginning to germinate.
TitleSymptoms
Caption(1) Sporangium detached from sporangiophore; (2) A sporangium which has released its zoospores; (3) Encysted zoospore beginning to germinate.
CopyrightMegan M. Kennelly
(1) Sporangium detached from sporangiophore; (2) A sporangium which has released its zoospores; (3) Encysted zoospore beginning to germinate.
Symptoms(1) Sporangium detached from sporangiophore; (2) A sporangium which has released its zoospores; (3) Encysted zoospore beginning to germinate.Megan M. Kennelly

Identity

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Preferred Scientific Name

  • Plasmopara viticola (Berk. & M.A. Curtis) Berl. & de Toni

Preferred Common Name

  • grapevine downy mildew

Other Scientific Names

  • Botrytis viticola Berk. & M.A. Curtis
  • Peronospora viticola (Berk. & M.A. Curtis) de Bary
  • Plasmopara amurensis Prots.
  • Rhysotheca viticola (Berkeley & Curtis) G.W. Wilson

International Common Names

  • English: brown rot; downy mildew of grapevine; grey rot
  • Spanish: mildiu de la vid; mildiu velloso de la vid
  • French: mildiou de la grappe; mildiou de la vigne; rot-brun de la vigne; rot-gris de la vigne

Local Common Names

  • Germany: Falscher Mehltau: Weinrebe; Lederbeerenkrankheit: Weinrebe; Peronospora: Weinrebe
  • Italy: Peronospora della vite

EPPO code

  • PLASVI (Plasmopara viticola)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Chromista
  •         Phylum: Oomycota
  •             Class: Oomycetes
  •                 Order: Peronosporales
  •                     Family: Peronosporaceae
  •                         Genus: Plasmopara
  •                             Species: Plasmopara viticola

Notes on Taxonomy and Nomenclature

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As reviewed by Viala (1893) and Gregory (1915), P. viticola was first collected and studied in 1834 by Schweinitz, who named it Botrytis caca (Schweinitz, 1837). Berkeley and Curtis (1848) later described the organism as Botrytis viticola. De Bary (1863) transferred the pathogen to a new genus and described it as Peronospora viticola. Berlese and de Toni (1888) redescribed the pathogen as Plasmopara viticola after Schröter (1886) separated Peronospora into two genera, Peronospora and Plasmopara.

According to Grunzel (1960) and Rafaila et al. (1968), several authors (Procenko, 1946; Savulescu and Savulescu, 1952; Golovina, 1955) have suggested distinguishing different varieties and special forms of P. viticola based on morphology or host range. Grunzel (1960) and Rafaila et al. (1968) concluded there was no evidence for such divisions.

Description

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After Hall (1989).

Mycelium composed of intercellular, colourless, aseptate hyphae 7-12 µm diameter, often irregularly shaped and swollen, bearing small, rounded vesicular haustoria, 4-10 µm diameter, formed predominantly in leaf tissues.

Sporangiophores hypophyllous, arborescent, 130-250(-700) x 11-14 µm, branching monopodially in the upper third at right angles to the main axis, and with a base tapering to a conical point; branches in a whorl of 4-5, 35-45 µm long, often with two opposite secondary branches 15-20 µm long, all having 3-4 conical tips 10 µm long 6 µm wide at base, diverging at right angles and tapering to a terminal swelling.

Sporangia ovoid, colourless, (17-)20(-25) x (10-)14(-16) µm, sometimes with a short pedicel, each producing 1-6 zoospores.

Zoospores reniform, laterally biflagellate, 6-8 x 4-5 µm, emerging from opposite the insertion point of the sporangium, via a papilla, or by direct penetration of the sporangium wall.

Oospores formed in leaf tissues, spherical, 28-40 µm diameter, containing 14-16 chromosomes, covered by two inner oospore membranes and an outer wrinkled oospore wall, germinating via a tube, 2-3 µm diameter, to give a pyriform sporangium, ca 25 x 35-40 µm, producing 8-20 zoospores.

Distribution

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P. viticola is native to the north-eastern USA. From the USA, P. viticola spread to Europe and worldwide. Today, P. viticola occurs in nearly all grape-growing regions worldwide. A few exceptionally dry grape-growing climates exist which support only minimal levels of the pathogen, such as parts of Argentina, Chile, California and Egypt (Weltzien, 1981; Lafon and Clerjeau, 1988; Emmett et al., 1992).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 25 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresent
AngolaPresent
Congo, Democratic Republic of thePresent
EgyptPresent
EthiopiaPresent
GhanaPresent
LibyaPresent
MadagascarPresent
MauritiusPresent
MoroccoPresent
MozambiquePresent
NigeriaPresent
Sierra LeonePresent
SomaliaPresent
South AfricaPresent
TanzaniaPresent
TunisiaPresent
UgandaPresent
ZambiaPresent
ZimbabwePresent

Asia

AzerbaijanPresent
CambodiaPresent
ChinaPresent
-LiaoningPresent
-NingxiaPresent
-XinjiangPresent
IndiaPresent
-Jammu and KashmirPresent
-KarnatakaPresent
-MaharashtraPresent
-Tamil NaduPresent
-Uttar PradeshPresent
IndonesiaPresent
IranPresent
IsraelPresent
JapanPresent
JordanPresent
KazakhstanPresent
LaosPresent
LebanonPresent
MalaysiaPresent
MyanmarPresent
PakistanPresent
PhilippinesPresent
Saudi ArabiaPresent
South KoreaPresent
Sri LankaPresent
SyriaPresent
TaiwanPresent
ThailandPresent
TurkeyPresent
VietnamPresent
YemenPresent

Europe

AustriaPresent
BulgariaPresent
CroatiaPresent
CyprusPresent
CzechiaPresent
CzechoslovakiaPresent
Union of Soviet Socialist RepublicsPresent
DenmarkPresent
FrancePresent
GermanyPresent
GreecePresent
HungaryPresent
ItalyPresent
-SardiniaPresent
MaltaPresent
MoldovaPresent
MontenegroPresent
NetherlandsPresent
North MacedoniaPresent
NorwayPresent
PolandPresent
PortugalPresent
-AzoresPresent
-MadeiraPresent
RomaniaPresent
RussiaPresentPresent based on regional distribution.
-Russian Far EastPresent
Serbia and MontenegroPresent
SlovakiaPresent
SloveniaPresent
SpainPresent
SwitzerlandPresent
UkrainePresent

North America

BarbadosPresent
CanadaPresent
-British ColumbiaPresent
-ManitobaPresent
-Nova ScotiaPresent
-OntarioPresent
-QuebecPresent
Costa RicaPresent
CubaPresent
Dominican RepublicPresent
El SalvadorPresent
GuatemalaPresent
HaitiPresent
JamaicaPresent
MexicoPresent
PanamaPresent
Puerto RicoPresent
United StatesPresent
-KentuckyPresent
-MarylandPresent
-MichiganPresent
-North CarolinaPresent
-PennsylvaniaPresent
-VirginiaPresent

Oceania

AustraliaPresent
-New South WalesPresent
-QueenslandPresent
-South AustraliaPresent
-TasmaniaPresent
-VictoriaPresent
-Western AustraliaPresent
New CaledoniaPresent
New ZealandPresent
Papua New GuineaPresent

South America

ArgentinaPresent
BoliviaPresent
BrazilPresent
-Minas GeraisPresent
-Rio Grande do SulPresent
-Santa CatarinaPresent
-Sao PauloPresent
ColombiaPresent
EcuadorPresent
UruguayPresent
VenezuelaPresent

Hosts/Species Affected

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P. viticola is most important as a pathogen of Vitis vinifera, V. labrusca and V. vinifera hybrids. P. viticola also attacks a number of Vitis species and related genera in the family Vitaceae, but wild hosts are not significant to disease development on the cultivated varieties (Barrett, 1939; Renfro and Bhat, 1981).

Wild Vitis host species include V. aestivalis,V. arizonica, V. berlandieri [V. cinerea var. helleri], V. californica, V. coignetiae, V. cordifolia, V. girdiana, V. monticola, V. pagnucci [V. piasezkii var. pagnuccii], V. riparia, V. romaneti, V. rotundifolia, V. rupestris and V. treleasei (Lafon and Bulit, 1981; Hall, 1989). Additionally, under laboratory conditions, Staudt and Kassemeyer (1995) observed downy mildew on V. acerifolia, V. amurensis, V. candicans [V. mustangensis], V. champinii, V. cinerea, V. doaniana, V. lanata, V. palmata, V. piasezkii, V. rubra, V. shuttleworthii, V. solonis, V. tiliifolia and V. vulpina.

Other wild host genera in the Vitaceae include Ampelopsis, Cissus, Cordifolia, Cinerea, Parthenocissus and Solonis (Renfro and Bhat, 1981; Hall, 1989).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
AmpelopsisVitaceaeOther
    CissusVitaceaeOther
      Vitis arizonica (canyon grape (USA))VitaceaeOther
        Vitis ficifoliaVitaceaeWild host
          Vitis labrusca (fox grape)VitaceaeMain
            Vitis rupestris (sand-grape)VitaceaeOther
              Vitis vinifera (grapevine)VitaceaeMain

                Growth Stages

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                Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

                Symptoms

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                P. viticola infects all green parts of the host plant that bear stomata. It generally causes yellow discoloration, necrosis and distortion.

                On young leaves, lesions appear as yellow, translucent 'oilspots' with a chocolate-brown halo (see Pictures). On cultivar Ruby Cabernet, oilspots are reddish instead of yellow (Nicholas et al., 1994). Multiple oilspots can coalesce to cover much of the leaf surface. Oilspots become dry and necrotic as they age, first in the centre and later throughout the entire lesion. On older leaves, the lesions are restricted by veins to form small, angular, yellow to reddish-brown spots which combine to form a patchwork or mosaic-like pattern.

                Sporulation only occurs on the lower leaf surface, where the stomata reside. The sporangiophores and sporangia appear as a white, downy, cottony growth. Under highly favourable conditions, sporulation may appear on the undersides of leaves before the yellow oilspot becomes visible on the upper leaf surface. On older oilspots, sporulation occurs primarily on the margins of the lesion.

                Infected shoot tips and rachises of young inflorescences distort into a curl or corkscrew.

                Infected inflorescences and young berries appear yellow or grey and may be covered with cottony spores under favourable conditions. Sporulation occurs on pedicels and berries. Clusters infected at an early stage can result in individual berries, sections of the cluster, or even entire clusters turning brown, drying and falling off the vine.

                Berries infected later in the season (after 2-3 weeks post-bloom) become discoloured and shrivel but do not support sporulation. This stage is sometimes referred to as the 'brown rot' phase. Berry stems continue to sporulate after sporulation ceases on the berries.

                List of Symptoms/Signs

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                SignLife StagesType
                Fruit / discoloration
                Fruit / extensive mould
                Fruit / mummification
                Fruit / premature drop
                Growing point / discoloration
                Growing point / distortion
                Growing point / mycelium present
                Inflorescence / blight; necrosis
                Inflorescence / discoloration (non-graminaceous plants)
                Inflorescence / distortion (non-graminaceous plants)
                Inflorescence / lesions; flecking; streaks (not Poaceae)
                Leaves / abnormal colours
                Leaves / abnormal forms
                Leaves / abnormal leaf fall
                Leaves / fungal growth
                Leaves / necrotic areas
                Stems / distortion
                Stems / mould growth on lesion
                Stems / mycelium present

                Biology and Ecology

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                P. viticola is an obligately biotrophic plant pathogen with a sexual overwintering phase and asexual multiplication cycles during the growing season.

                Primary Infection

                The pathogen usually overwinters as oospores in fallen leaves infected in the previous season. P. viticola is heterothallic with two mating types (Wong et al., 2001). An antheridium fertilizes an oogonium to form the sexual oospore (Kortekamp et al., 1998). In mild climates such as in California the pathogen has been observed overwintering as mycelium in buds and canes of wild grape species (Barrett, 1939).

                Oospores germinate in the spring when temperatures reach about 10°C and vineyard soils are wet. Oospores germinate and form a germ tube which terminates in a macrosporangium. Germination and formation of the macrosporangia can occur in as little as 24 hours of soil wetness (Gregory, 1915; Ronzon-Tran Manh Sung and Clerjeau, 1988). When wet, the macrosporangia release an average of 8-20 (Hall, 1989) and up to 60 (Ronzon-Tran Manh Sung and Clerjeau, 1988) zoospores. At 20°C zoospore release occurs after 30 or more minutes (Ronzon-Tran Manh Sung and Clerjeau, 1988). Zoospores are dispersed onto host tissue by rainsplash.

                Penetration and colonization of the host

                Zoospores require surface wetness to infect the host. Infection takes place only through the stomata (Gregory, 1914). Zoospores swim on the tissue surface, encyst near stomata in groups of 2 to 5, and each spore forms a single germ tube which penetrates the stomata. In the substomatal cavity the germ tube swells, forming a substomatal vesicle from which arises a single hypha. Hyphae grow intercellularly, filling the space between the host mesophyll cells and taking on an irregular shape. In as little as 3.5 hours the first haustorium forms where the pathogen contacts the host cells. Additional haustoria form as the pathogen ramifies through host tissue, parasitizing the mesophyll cells (Langcake and Lovell, 1980; Kortekamp et al., 1998). The pathogen can initiate sporulation upon reaching a non-parasitized substomatal cavity (Langcake and Lovell, 1980; Kortekamp et al., 1998). Vascular tissue restricts pathogen growth to interveinal areas in older leaves (Langcake and Lovell, 1980).

                Incubation

                The incubation time, the period between infection and the first appearance of symptoms, depends on temperature and ranges from 4 to 21 days, with an average of 7-10 days. The incubation period is shortest on young leaves and at temperatures of 19-24°C (Muller and Sleumer, 1934; Zachos, 1959; Rafaila et al., 1968).

                Sporulation

                The pathogen sporulates through stomata during warm, humid nights. To sporulate, P. viticola requires at least 95-98% RH, temperatures between 10 and 30°C (peak production occurs at 20°C) and at least 4 hours of darkness (Blaeser and Weltzien, 1978; Lalancette et al., 1988b; Hill, 1989). Light inhibits sporulation of P. viticola (Brooks, 1979; Magarey and Butler, 1998). The sporangia remain viable for 4-8 days at high relative humidity and temperatures below about 22°C, and 1-2 days in hotter, drier conditions (Zachos, 1959; Blaeser and Weltzien, 1978; Kast and Stark-Urnau, 1999).

                Individual lesions resporulate a number of times under favourable conditions, and can retain the potential to sporulate for 2-3 months (Zachos, 1959; Hill, 1989).

                Secondary Infection

                Secondary cycles of infection occur repeatedly throughout the growing season if weather conditions are favourable. To infect the plant, P. viticola requires surface wetness and temperatures of between 5-6°C and 30°C (Blaeser and Weltzien, 1977; Lalancette et al., 1988a). At optimal temperatures (approximately 20°C) infection can occur in 2 hours of surface wetness, with more hours of wetness required at non-optimal temperatures (Blaeser and Weltzien, 1977; Lalancette et al., 1988a).

                When wet, the sporangia detach from their sporangiophores and germinate, releasing zoospores which are spread by windblown rain to new host tissue (Blaeser and Weltzien, 1978). Zoospores are released 30 to 180 minutes after the sporangia become wet (Langcake and Lovell, 1980; Kast and Stark-Urnau, 1999). Zoospores aggregate near stomata in groups of up to 10, possibly due to chemotactic signals (Royle and Thomas, 1973).

                The penetration and colonization process is the same as described for the primary infection.

                Means of Movement and Dispersal

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                Natural dispersal (non-biotic):

                Blaeser and Weltzien (1978) observed sporangial dispersion only in wind-blown rain, not in the air. Zachos (1959) noted that high mortality of sporangia in dry conditions would prevent air dispersal. However, there is some evidence of long-distance (500-600 km) spore dispersal in regional air currents (Szoke et al., 1998).

                Vector transmission

                None.

                Seedborne spread

                None.

                Agricultural practices

                Historically, agricultural practices have led to catastrophic dispersal of P. viticola. P. viticola is native to the north-eastern USA, and it is generally accepted that the pathogen spread to Europe on American grapes imported for use as rootstocks resistant to phylloxera (Daktulosphaira vitifolii [Viteus vitifoliae]) (Viennot-Bourgin, 1981). Downy mildew was first observed in France in September 1878 (Viala, 1893).

                According to Viala (1893), the dangers of importing infected American
                plants had already been asserted by Cornu several years earlier.

                Strategies to prevent the spread of P. viticola on plant material include heat treatment of cuttings, maintaining disease-free tissue culture plantlets, and avoiding the spread of soil and leaf debris which may bear oospores (Emmett et al., 1992).

                Plant Trade

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                Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
                Flowers/Inflorescences/Cones/Calyx hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
                Fruits (inc. pods) hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
                Growing medium accompanying plants spores Pest or symptoms usually invisible
                Leaves hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
                Seedlings/Micropropagated plants hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
                Plant parts not known to carry the pest in trade/transport
                Bark
                Bulbs/Tubers/Corms/Rhizomes
                Roots
                Stems (above ground)/Shoots/Trunks/Branches
                True seeds (inc. grain)
                Wood

                Impact

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                P. viticola has caused significant impacts on grape production since the 1800s. During the early culture of European varieties in the USA, yield losses were commonly 75% (Viala, 1893). Catastrophic losses arose in Europe in the late 1800s when P. viticola was first introduced on American rootstocks. The disease caused severe losses in favourable seasons and growers abandoned the use of several highly susceptible varieties (Viala, 1893).

                Potential yield losses remain high, ranging from 50 to 100% under favourable conditions. The pathogen directly attacks the young inflorescences and fruit. Indirect damage occurs when severe foliar infections cause early defoliation, exposing the fruit to sunburn and reducing winter hardiness (Emmett et al., 1992). The estimated annual crop loss in an average year in Australia is $22.5 million (Australian dollars) with an additional $10 million spent on control measures. In wet years, direct crop losses may be as high as $64 million (Magarey and Butler, 1998).

                Diagnosis

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                Place leaves displaying oilspots and clusters displaying epinasty or oilspots in a lightly moistened plastic box or plastic bag and incubate overnight in the dark at 15-25°C. Fresh spores will be visible on infected tissue in the morning. Mature berries will not produce spores but their infected pedicels will sporulate.

                P. viticola is an obligate biotroph and thus cannot be grown in culture.

                Detection and Inspection

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                Oilspots are readily visible in the field and will be noticed on inspection of the leaves (Seem et al., 1985). Monitoring procedures have been described to detect primary infection at levels as low as 0.01% (Seem et al., 1985). The upper leaf surfaces should be inspected for oilspots by gently moving the shoots and leaves, spending about 30 seconds per vine for 200-300 vines. If a primary infection is observed, that spot should be monitored for the first secondary infections, which are usually within 1 m of the original primary oilspots (Nicholas et al., 1994). Vines should be inspected every 1-2 weeks, or as necessary based on weather conditions (Nicholas et al., 1994). Computer training programs have been developed for improved accuracy of disease assessments (Nutter et al., 1998).

                Similarities to Other Species/Conditions

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                Uncinula necator, causal agent of grape powdery mildew, also infects grape leaves and fruit clusters. Symptoms of early powdery mildew can resemble those of early downy mildew. Both diseases cause light-yellow patches on the upper leaf surface. However, powdery mildew is characterized by a flat, powdery, whitish-grey appearance on upper or lower leaf surfaces and on the fruit (Pearson, 1988; Nicholas et al., 1994). In contrast, downy mildew colonies are fluffy, a brighter white, and only found on the lower leaf surface.

                Botrytis cinerea, the cause of Botrytis bunch rot or grey mould, produces fluffy colonies on grape clusters. However, Botrytis sporulation is brownish-grey or olive, unlike the bright white appearance of P. viticola (Bulit and Dubos, 1988; Nicholas et al., 1994).

                Herbicides such as paraquat can cause yellowing of leaf tissue but these spots produce no spores (Nicholas et al., 1994).

                Erinose mites cause a white growth on the undersides of leaves which can resemble downy mildew infections. The growths are only in blister-like galls and are not accompanied by oilspots (Nicholas et al., 1994).

                Prevention and Control

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                Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

                Disease pressure varies significantly with weather conditions. Management must be rigorous in wet climates such as eastern North America and parts of Europe, and during unusually wet seasons in dry locations such as California or Australia.

                Chemical Control

                Chemical control has been an important control measure since the late 1800s, after the classic discovery of Bordeaux mix (copper sulfate plus lime) by Millardet in 1885 (Viala, 1893). Fungicides remain the most widely used management tool against P. viticola today (Lafon and Bulit, 1981; Lafon and Clerjeau, 1988; Emmett et al., 1992).

                There are multiple pre- and post-infection chemicals available. Currently used pre-infection chemicals, or protectants, include: copper-based compounds (e.g. copper hydroxide, copper oxychloride and cuprous oxide); cyclimide (e.g. captan); dithiocarbamates (e.g. mancozeb); chlorothalonil; and dithianon.
                Phenylamide compounds such as metalaxyl and benalaxyl have both pre- and post-infection activity (Nicholas et al., 1994) as do strobilurines such as azoxystrobin (Wong and Wilcox, 2001). Phosphonates provide post-infection activity (Nicholas et al., 1994).

                First applications are generally advised at 3-8 inches of shoot-growth, immediate pre-bloom, and post-bloom to protect the young inflorescences and fruit. For the remainder of the season sprays may be based on a routine schedule (usually every 10-14 days) to maintain continuous protection of the vines. Alternatively, sprays may be based on disease risk as determined by weather conditions and forecasting models.

                Resistance to phenylamide fungicides has been observed in Europe (Leroux and Clerjeau, 1985). The possibility of future resistance has triggered interest in monitoring and preventing this threat (Genet and Vincent, 1999; Wong and Wilcox, 2000). Gauthier and Amsden (2014) reported Qol-resistant downy mildew of grape in Kentucky.

                Warning Systems

                The complex interactions of P. viticola with the environment render grape downy mildew a candidate for disease forecasting and modelling. Spray timing models have been used in Europe since the early 1900s (Muller and Sleumer, 1934; Populer, 1981). In more recent years computer-based forecasting models and decision tools have been developed in Italy (Rosa et al., 1993), France (Fouassier et al., 1997), Austria (Denzer, 1998), Germany (Huber et al., 1998), Hungary and Slovakia (Szoke et al., 1998), the USA (Park et al., 1997; Madden et al., 2000) and Australia (Magarey et al., 1991; Wachtel and Magarey, 1997). Most downy mildew models incorporate temperature, rainfall, relative humidity and leaf wetness, and more complex simulators incorporate information on host growth stage and varietal susceptibility. Models can be integrated into pre- or post-infection treatment strategies.

                Resistant Crop Cultivars

                Most cultivars of V. vinifera are highly susceptible to downy mildew, as are many cultivars of V. labrusca and interspecific hybrids.

                Breeding for resistance to downy mildew has been ongoing since the 1930s (Matthews, 1981). Wild American Vitis species and related genera have been identified as possible sources of resistance. Putative resistance mechanisms include surface features such as leaf hairs or warty structures (Matthews, 1981; Staudt and Kassemeyer, 1995; Kortekamp and Zyprian, 1999), single-gene hypersensitive response (Matthews, 1981) and defence chemicals including peroxidase, viniferin, resveratrol and flavonoids (Langcake, 1981; Dai et al., 1995; Kortekamp et al., 1998).

                Attempts to genetically engineer grapevine began in the late 1980s with Agrobacterium-mediated transformation (Baribault et al., 1989) and more recently with biolistics, or the 'gene gun' (Kikkert et al., 1996). Research continues on the introduction of genes for resistance to downy mildew and other pathogens (Reisch et al., 1996).

                Cultural Control

                Cultural practices alone are unlikely to give sufficient control, especially under conditions favourable to downy mildew (Lafon and Clerjeau, 1988). However, cultural methods can influence disease development. High humidity and extended periods of leaf wetness enhance the development of downy mildew, thus techniques to promote air circulation and minimize surface wetness may reduce disease development (Palti and Rotem, 1981). Pruning and trellising methods which reduce canopy density decrease downy mildew levels (Wearing et al., 1999). Using furrow irrigation instead of overhead irrigation also reduces disease (Emmett et al., 1992). Primary infection may be prevented by ploughing to bury oospores in leaf litter, and avoidance of irrigating soil for long periods in order to prevent oospore germination (Palti and Rotem, 1981).

                References

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                Chavarria G, Santos H P dos, Sônego O R, Marodin G A B, Bergamaschi H, Cardoso L S, 2007. Incidence of diseases and needs of control in overhead covered grapes. (Incidência de doenças e necessidade de controle em cultivo protegido de videira.). Revista Brasileira de Fruticultura. 29 (3), 477-482. http://www.scielo.br/scielo.php?script=sci_issues&pid=0100-2945&lng=pt&nrm=iso DOI:10.1590/S0100-29452007000300014

                Chen WeiMin, Jiao ZiWei, Song HongMei, He Qiong, Li ZhenGui, Xu Yi, Chen RenNing, Sun QingHua, Tuersun, 2007. Occurrence of Red Global grape diseases and weeds, and their integrated management in Ili Valley, Xinjiang. Xinjiang Agricultural Sciences. 44 (1), 91-95.

                Choi Y J, Cho S E, Shin H D, 2017. First report of downy mildew caused by Plasmopara viticola on Vitis ficifolia var. sinuata in Korea. Plant Disease. 101 (11), 1958. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-04-17-0571-PDN

                Ferrari J T, Nogueira E M de C, Santos A J T, Louzeiro I M, 2003. Evaluation of fungicides in the control of grapevine downy mildew on grape. (Avaliação de fungicidas no controle do míldio em videira.). Arquivos do Instituto Biológico (São Paulo). 70 (Suplemento 3), 045. http://www.biologico.sp.gov.br/ARQUIVOS/V70_suplemento23/raib.pdf

                Gauthier N A W, Amsden B, 2014. First report of QoI-resistant downy mildew (Plasmopara viticola) of grape (Vitis vinifera cv. Vidal Blanc) in Kentucky. Plant Disease. 98 (2), 276-277. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-11-12-1020-PDN

                Hrubý R, 2003. Early warning system for grapevine in South Moravia (Czech Republic). Bulletin OEPP. 33 (3), 433-436. DOI:10.1111/j.1365-2338.2003.00669.x

                Khilare V C, Deokate A S, Gangawane L V, 2003. Occurrence of aluminium phosethyl (Allitte) resistance in Plasmopara viticola causing downy mildew of grapevine in Maharashtra. Journal of Phytological Research. 16 (2), 239-241.

                Kilimnik A N, Samoĭlov Yu K, 2000. Approaches to control downy mildew. Zashchita i Karantin Rasteniĭ. 29.

                Latinovic N, Maras V, 2005. Susceptibility of Kratosija variety's biotypes to causal agent of grapevine dead arm disease (Phomopsis viticola Sacc.). Agroznanje - Agroknowledge Journal. 6 (2), 25-29.

                Luskar S, Košir I J, Kač M, 2005. Deposit of copper ions on vine leaves and its effect on biological efficacy of different plant protection products against downy mildew: 2. Deposit of copper ions on vine leaves. (Depozit bakrovih ionov na listih vinske trte in njegov vpliv na biološko učinkovitost uporabljenih fitofarmacevtskih sredstev za zatiranje peronospore vinske trte: 2. Depozit bakrovih ionov na listih vinske trte.). Hmeljarski Bilten. 31-41.

                McKirdy S J, Riley I T, Cameron I J, Magarey P A, 1999. First report of grapevine downy mildew (Plasmopara viticola) in commercial viticulture in Western Australia. Plant Disease. 83 (3), 301. DOI:10.1094/PDIS.1999.83.3.301D

                Pereira V F, Resende M L V de, Monteiro A C A, Ribeiro Júnior P M, Regina M de A, Medeiros F C L, 2010. Alternative products for the protection of vine against downy mildew. (Produtos alternativos na proteção da videira contra o míldio.). Pesquisa Agropecuária Brasileira. 45 (1), 25-31. http://www.scielo.br/pab DOI:10.1590/S0100-204X2010000100004

                Peruch L A M, Bruna E D, 2008. Relation between doses of Bordeaux mixture and phosphites potassium on the intensity of Downy Mildew on grape cv. 'Goethe'. (Relação entre doses de calda bordalesa e de fosfito potássico na intensidade do míldio e na produtividade da videira cv. "Goethe".). Ciência Rural. 38 (9), 2413-2418. http://www.ufsm.br/ccr/revista DOI:10.1590/S0103-84782008000900001

                Prabhu H V, 2007. Biological efficacy of new combination products against downy mildew of grapes. Annals of Biology. 23 (1), 53-56.

                Prakash V R, Eswaran A, Sanjeevkumar K, Usharani S, 2007. Effectiveness of novel combination fungicide against downy mildew incidence, fruit quality, shelf life and post harvest pathogens of grapevine in India. Plant Archives. 7 (2), 775-780.

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