31/3/2018 Downy mildew of grape

Home | Log In
APS > Education > Introductory > Plant Disease Lessons > Fungi and Fungus- Share
like Organisms > Oomycetes
Ascomycetes/Imperfect
Fungi Downy mildew of grape
Basidiomycetes Ash, G. 2000. Downy mildew of grape. 2000. The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1112-01
Updated 2017.

Oomycetes
DISEASE: Downy mildew
Other
PATHOGEN: Plasmopara viticola

HOSTS: All cultivars of grapes in the species Vitis vinifera and many Vitis
intraspecific hybrid cultivars. Susceptibility within the North American
species Vitis labrusca, varies from highly susceptible to resistant.

Authors
Gavin Ash
Charles Sturt University,
Wagga Wagga, New South Wales, Australia

A typical method of growing grapevines on a trellis system.

(Courtesy G. Ash)

Downy mildew is a highly destructive disease of grapevines in all

grape-growing areas of the world where there is spring and summer
rainfall at temperatures above 10ºC (50ºF). In 1885, P.M.A. Millardet
first used Bordeaux mixture (copper sulfate and lime) to control
downy mildew in the vineyards of France, beginning the chemical era
of disease control.

Symptoms and Signs

Although all green parts of the grapevine are susceptible, the first symptoms of
downy mildew of grapes, caused by Plasmopara viticola, are usually seen on the
leaves as soon as 5 to 7 days after infection. Foliar symptoms appear as yellow
circular spots with an oily appearance (oilspots) (Figure 2). Young oilspots on young
leaves are surrounded by a brownish-yellow halo. This halo fades as the oilspot
matures. The spots are yellow in white grape varieties and red in some red grape
varieties (e.g., Ruby Red). Under favorable weather conditions, large numbers of
oilspots may develop and coalesce to cover most of the leaf surface (Figure 3).
After suitably warm, humid nights, a white downy fungal growth (sporangia) will

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 1/9
31/3/2018 Downy mildew of grape

appear on the underside of the leaves and other infected plant parts (Figure 4). The
disease gets its name "downy mildew" from the presence of this downy growth.

Figure 2 Figure 3 Figure 4

In late summer and early fall, the diseased leaves take on a tapestry-like
appearance when the growth of the pathogen is restricted by the veinlets (Figure 5).
Confirmation of active downy mildew is made by the "bag test." To do this test,
seal suspect diseased leaves and/or fruit bunches in a moistened (not wet)
plastic bag and incubate in a warm (13-28ºC/ 55-82ºF), dark place overnight.
Look for fresh, white downy sporulation beneath suspect oilspots or on shoots or
fruit bunches (Figure 4). Note that mature berries, although they may be
symptomatic and harbor the pathogen, may not support sporulation even when
provided with ideal conditions. Infected parts of young fruit bunches turn brown,
wither, and die rapidly. If infections occur on the young bunch stalk, the entire
inflorescence may die (Figure 6). Developing young berries will either die or, if
between 3 and 5 mm in diameter, become discolored (Figure 7). Berries become
resistant to infection within 2-3 week after bloom, although all parts of the rachis
may remain susceptible 2 months after bloom.

Figure 5 Figure 6 Figure 7

Pathogen Biology
The pathogen, Plasmopara viticola, produces asexual, biflagellate zoospores
and sexual oospores. Its mycelium is aseptate. It is an oomycete in the order
Peronosporales. Other important downy mildew pathogens that belong to this
group include species within the genera Bremia, Peronospora and Sclerospora.
Most taxonomists no longer include oomycetes within phylogenetic groupings
containing ascomycetes or basidiomycetes, and have placed them instead in the
kingdom Chromista. However, phylogenetic considerations aside, oomycetes
share many biological, ecological, and epidemiological characteristics with fungal
plant pathogens.

Plasmopara viticola is an obligate parasite, and it absorbs its nutrients from the
living host tissue via globose haustoria. The hyphae are largely internal in the
host. Sexual reproduction occurs through the fusion of antheridia and oogonia
within the host tissue. Plasmopara viticola has only recently been shown to be
heterothallic. The resulting sexual spore is an oospore, which is the survival and
resting stage of the pathogen.

Oospores represent the primary inoculum, and may overwinter in leaf litter or
may be released into the soil as leaves decay or are buried by detritivores. They
generally begin to germinate in significant numbers shortly after bud break of
grape, and populations of oospores may continue to germinate for the entire

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 2/9
31/3/2018 Downy mildew of grape

growing season in some growing regions. Oospores form a single germ tube
terminating in a sporangium. Zoospores form within the sporangia and are then
released. Zoospores germinate and penetrate the plant only through functioning
stomata, i.e., only on green host tissue. Sporangia and zoospores are easily
desiccated. They die within 2 to 3 hr of exposure to low humidity and sunlight, so
most infection occurs soon after their release. However, they may survive on leaf
surfaces for more than 24 hrs under cool humid conditions.

Sporangia (Figure 8) also serve as a means of secondary spread of the pathogen.

Treelike sporangiophores, bearing white, lemon-shaped sporangia, are produced
from a mycelial mat within the host tissue and emerge through stomata.
Sporangia are disseminated by wind and rain splash. Zoospores released from
the sporangia swim in free water (www.youtube.com/watch?v=BKs8WM01iIk) on
the grapevine surface, and encyst near a stoma. Zoospores then germinate and
penetrate through a stoma by the means of a germ tube. A time lapse video
showing sporulation, releasing of zoospores and germinated zoospore can be
found at www.youtube.com/watch?v=qnE91MbZrsg. Oospores are produced in
infected host material towards the end of the season. The resulting oospores are
thick-walled and serve not only as survival spores during the intercrop period, but
are also a source of genetic variation.

Figure 8

Disease Cycle and Epidemiology

Disease Cycle
The pathogen survives the winter period as oospores embedded in dead leaves
and other host tissue on the vineyard floor. Oospores may be released from the
decaying plant material onto the soil surface.

Oospores typically produce sporangia. These sporangia, in turn, produce

zoospores. Sporangia and zoospores are splashed by rain or carried by wind to
the lower leaves and tissues of the grapevines.

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 3/9
31/3/2018 Downy mildew of grape

The conditions necessary for oospore germination are wet soils with
temperatures above 10ºC (50ºF). An additional rule of thumb in some regions
includes rainfall of at least 10 mm (0.04 in.) in a 24-hr period to satisfy the
requirement of soil wetness. Once zoospores form and are dispersed to host
tissue, infection will occur given a minimum of 45 degree-hours of leaf wetness
[“degree hours” is the summation of hourly temperatures (C) until the specified
value is achieved]. Zoospores encyst and then germinate and penetrate through
stomates. Sporangia are usually inactivated after 5-7 hours in sunlight (Kennelly
et al., 2007), so most infections occur in the morning. After infection, the
pathogen grows intercellularly, producing haustoria.

Sporangia for secondary infections are produced on sporangiophores that

emerge through stomata of infected leaves and other grapevine tissues (e.g.
bunches). Production of sporangiophores and sporangia requires 95 to 100%
relative humidity and at least 4 hr of darkness at temperatures initially exceeding
13ºC (55ºF). The longevity and productivity of foliar lesions is related with how
often they were induced to sporulate. In the absence of an opportunity to
sporulate or extremely high temperatures, lesions typically maintained their
maximum potential to produce sporangia for at least 22-24 days or even as long
as 2-3 months (Kennelly et al., 2007). Caffi et al. (2013) reported the sporangia
are produced at a high rate for the first 4 days and then at a lower rate until
sporulation stopped. Sporangia are dispersed to new infection sites by rain
splash and/or wind; the latter occurs when sporangia are released into the air as
the humidity decreases. Zoospores from these sporangia are released in water
and are further spread by wind or rain splash.

The incubation period (the time from infection to appearance of new symptoms)
varies from 5 to 21 days depending on temperature, target organ of the host, and
on ontogenic resistance (Rossi et al., 2013). The incubation period is shortest
(~5 days) at mean temperatures from 20-25ºC (68-77ºF). At mean temperatures
of 12ºC (54ºF) or lower, the incubation period is 14 days or longer. The latent
period on leaves (the time from infection to first production of sporangia) is one
day less than the incubation period, given favorable conditions, i.e. sporulation
may occur one day prior to the appearance of oilspots.

Sexual reproduction occurs towards the end of the season. The resulting
oospores are thick-walled and serve as survival spores.

Disease Management
Cultural practices
When establishing vineyards the location, drainage, type of irrigation and
trellising system should all be selected to reduce the risk of disease. Because
moisture favors the development of downy mildew, grapevines should be
established in well-drained sites with good air movement. Grapevines should be
planted in rows that take advantage of natural patterns of air movement to help
minimize leaf wetness. Few cultural management options are available to control
downy mildew in established vineyards. Trellising systems and pruning can be
used to manage the leaf canopy to minimize leaf wetness. Avoid increasing
humidity and leaf wetness at night to mitigate secondary infection. Where
possible, the use of overhead irrigation should be avoided or scheduled so that
leaves will dry quickly. Reducing leaf litter and pruning may reduce the amount of
overwintering inoculum.

Genetic resistance
All cultivars of Vitis vinifera (the Eurasian species) are considered susceptible to
downy mildew, although cultivars such as Chardonnay, Pinot Noir, and Sultana

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 4/9
31/3/2018 Downy mildew of grape

are considered more susceptible than Cabernet Sauvignon and Semillon.

Several North American species show resistance to downy mildew (e.g. V.
labrusca and V. rotundifolia), although the V. labrusca cultivars Niagara and
Catawba are highly susceptible. Interspecific hybrids of V. vinifera and the North
American species have yielded cultivars with good wine-grape qualities and
greater resistance to downy mildew. However, wine consumers prefer the known
varietal wines over the less-known and sometimes fruitier hybrids. There are
several active research programs to genetically modify V. vinifera cultivars to
include disease resistance.

Chemical control
Both pre-infection (protective) and post-infection (systemic or penetrant)
fungicides are widely used for the control of downy mildew (Figure 9). Pre-infection
chemicals are applied prior to, but as close as possible, to an infection event. If
used in a regular spray schedule, their best use is in the period of greatest host
susceptibility, between shoot length of 10 cm (4 in.) and pea-sized berries.
Alternatively, pre-infection chemicals may be used irregularly by spraying as
close as possible prior to forecast weather events favorable for P. viticola
infection. The frequency of the sprays will be determined by the prevailing
weather conditions. Pre-infection fungicides include the copper-based fungicides,
such as Bordeaux mixture (see section on Significance) and the
dithiocarbamates. There have been no reports of resistance to these types of
fungicides in the pathogen.

Figure 9

Post-infection fungicides are more costly than pre-infection fungicides and are
best used sparingly. For optimal use, they should be applied as soon as possible
after an infection event and prior to the appearance of oilspots. These fungicides
are best used in conjunction with a forecasting program, which assesses the
likelihood of infection from canopy micro-climate data (Figure 10). A good number of
prediction models have been established, such as Australian D-Model (Magarey
et al., 1991), DMCast model in the US (Park et al.,1997), EPI model (Stryzik,
1983) and POM model in France (Tran Manh Sung et al., 1990), and complex
mechanistic UCSC model in Italy (Rossi et al., 2008). However, only limited
models have been tested independently by their developers or have been
integrated into more complex advisory systems (Gessler et al., 2011). Some
post-infection fungicides are less effective when applied to oilspots, although
these fungicides may have the capacity either to kill the pathogen active in
oilspots or to significantly reduce its sporulation potential. Currently used post-
infection fungicides include phosphonate (e.g. fosetyl-aluminum), phenylamides
(e.g. melalaxyl), QoI (e.g. azoxystrobin), and Carboxylic acid amides (CAA; e.g.
mandipropamid). Depending on the regulation of each state, some of these
chemistries are recommended for controlling grape downy mildew in the US.
However, fungicide resistance to some of above mentioned chemicals, such as
QoI or CAA fungicides, have also been reported in Australia, Japan, France, and
the US (FRAC, 2013). Although some new chemistries are available now, care
still need to be taken to prevent the development of fungicide resistance.

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 5/9
31/3/2018 Downy mildew of grape

Figure 10
The disease should be monitored if a post-infection application strategy is used.
To scout a vineyard for the presence of downy mildew (Figure 11), the scout should
walk slowly along the vines looking for oilspots on at least 200 vines. More than 2
oilspots per 50 vines would be considered a risk to the vineyard.

Figure 11

Significance
Grapevines are one of the most widely grown fruit crops in the world with
significant plantings in Europe, North and South America, South Africa and
Australasia. Grapes are used in the production of wine, brandy, or non-fermented
drinks and are eaten fresh or dried as raisins.

In an effort to control a root-feeding aphid called Phylloxera, resistant grape

rootstocks were introduced to Europe from North America in 1878. At the same
time, the downy mildew pathogen was introduced into France, probably as
oospores on the imported rootstocks, and spread widely throughout Europe. In
1885, P.M.A. Millardet (Figure 12) first used Bordeaux mixture to control downy
mildew. It is said that a farmer had applied this mixture of copper sulfate and lime
to produce a chemical residue on grapes along the roadside to discourage
pilfering by passersby. Millardet's extensive experimentation led to the
development of Bordeaux mixture as the first widely used chemical to protect
plants from fungal infections. His description of this work has been translated into
English as a Phytopathological Classic and is cited in the references. A statue of
Millardet was erected in a park in Bordeaux to express the gratitude of the
French people for his important discovery. Bordeaux mixture remains an
important fungicide even today.

Figure 12

Downy mildew is a highly destructive disease of grapevines in all grape-growing

areas of the world where there is spring and summer rainfall at temperatures
above 10º C (50º F). Crop losses in individual years can be 100% if the disease
is not controlled during favorable weather. Early infection of young bunches can
lead to significant crop loss, whereas, severe leaf infection affects the source-
sink relationship in the vine and may lead to defoliation and possible sunburn or
lack of fruit ripening. This destruction of leaf tissue may affect sugar

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 6/9
31/3/2018 Downy mildew of grape

accumulation and growth in the subsequent season (Figures 13 and 14). Currently,
there are no suitable sources of resistance in commercially acceptable varieties,
so fungicides are the primary means of disease control.

Figure 13 Figure 14

Selected References
Caffi, T., G. Gilardi, M. Monchiero, and V. Rossi. 2013. Production and release of
asexual sporangia in Plasmopara viticola. Phytopathology 103: 64-73.

Carefoot, G.L. and E.R. Sprott.1967. Famine on the Wind. Rand McNally and
Co., Chicago.

Coombe, B.G. and P.R. Dry (eds.). 1992. Viticulture Volume 2. Practices.
Winetitles. Adelaide, Australia.

FRAC. 2013. FRAC List of plant pathogenic organisms resistant to disease

control agents. http://www.frac.info/docs/default-source/publications/list-of-
resistant-plant-pathogens/list-of-resistant-plant-pathogenic-organisms---february-
2013.pdf?sfvrsn=4.

Genet, J.L., and O. Vincent.1999. Sensitivity of European Plasmopara viticola

populations to cymoxanil. Pest. Sci. 55:129-136.

Gessler, C., I. Pertot, and M. Perazzolli. 2011. Plasmopara viticola: a review of

knowledge on downy mildew of grapevine and effective disease management.
Phytopathologia Mediterranea 50: 3-44.

Hong, C.F. 2016. Release of Plasmopara viticola zoospore.

https://www.youtube.com/watch?v=BKs8WM01iIk

Hong, C.F. 2016. Time lapse video of sporulation and sporangia germination of
Plasmopara viticola. https://www.youtube.com/watch?v=qnE91MbZrsg

Hoppmann D. and K.P. Wittich. 1997. Epidemiology-related modelling of the leaf-

wetness duration as an alternative to measurements, taking Plasmopara viticola
as an example. Zeit. Pflanzenkrank. Pflanzenschutz (Journal Plant Dis. Prot.)
104:533-544.

Kennelly, M. M., D. M. Gadoury, W. F. Wilcox, P.A. Magarey, and R.C. Seem.

2005. Seasonal development of ontogenic resistance to downy mildew in grape
berries and rachises. Phytopathology 95:1445-1452.

Kennelly M.M., D. M. Gadoury, W. F. Wilcox, P. A. Magarey, and R.C. Seem.

2007. Primary Infection, Lesion Productivity, and Survival of Sporangia in the
Grapevine Downy Mildew Pathogen Plasmopara viticola. Phytopathology 97,
512-22.

Kortekamp A. 1997. Epicoccum nigrum link - a biological control agent of

Plasmopara viticola (Berk. Et Curt.) Berl. Et de-toni. Vitis 36:215-216.

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 7/9
31/3/2018 Downy mildew of grape

Lalancette, N., L.V. Madden, and M.A. Ellis. 1988. Development of an infection
efficiency model of Plasmopara viticola on American grape based on
temperature and duration of leaf wetness. Phytopathology 78:794-800.

Lalancette, N., L.V. Madden, and M.A. Ellis. 1988. A quantitative model for
describing the sporulation of Plasmopara viticola on grape leaves.
Phytopathology 78:1316-1321.

Langcake, P. and A. Lovell. 1980. Light and electron microscopical studies of the
infection of Vitis spp. by Plasmopara viticola, the downy mildew pathogen. Vitis
19:321-337.

Large, E.C. 1940. The Advance of the Fungi. Dover Publications, New York, NY.
Reprinted by American Phytopathological Society Press, St. Paul, MN.

Madden, L.V., M.A. Ellis, N. Lanlancette, and L. L. Wilson. 2000. Evaluation of a

disease warning system for downy mildew of grapes. Plant Dis. 84:549-554.

Magarey, P.A., M.F. Wachtel, P.C. Weir, and R.C. Seem.1991. A computer-based
simulator for rational management of grapevine downy mildew Plasmopara
viticola. Aust. Plant Prot. Q. 6:29-33.

Millardet, P.M.A. The discovery of Boreaux mixture, 1885. Phytopathological

Classic translated into English by F.J. Schneiderhan. American Phytopathological
Society Press, St. Paul, MN.

Park, E.W., R.C. Seem, D.M. Gadoury, and R.C. Pearson.1997. DMCAST: A
prediction model for grape downy mildew development. Vitic. Enol. Sci. 52:182-
189.

Pearson, R.C. and A.C. Goheen. 1994. Compendium of Grape Diseases.

American Phytopathological Society Press, St. Paul, MN.

Rossi V, T. Caffi, S. Giosuè, and R. Bugiani. 2008. A mechanistic model

simulating primary infections of downy mildew in grapevine. Ecological Modelling
212, 480-91.

Rossi V, T. Caffi, and D. Gobbin. 2013. Contribution of molecular studies to

botanical epidemiology and disease modelling: grapevine downy mildew as a
case-study. European Journal of Plant Pathology 135, 641-54.

Wong, F.P., H.N. Burr, and W. F. Wilcox. 2000. Heterothallism in Plasmopara

viticola (grapevine downy mildew). Phytopathology 90:S85.

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 8/9
31/3/2018 Downy mildew of grape

Get ALL the Latest Updates for ICPP2018: PLANT HEALTH IN A GLOBAL ECONOMY. © 2018 The American Phytopathological Society. All rights
Follow APS! reserved.

Contact Us - Report a Bad Link

https://www.apsnet.org/edcenter/intropp/lessons/fungi/Oomycetes/Pages/DownyMildewGrape.aspx 9/9