Pseudoperonospora humuli is a plant pathogen that causes downy mildew on hops .
Hosts and Symptoms:
Downy mildew on hops is caused by the pathogen Pseudoperonospora humuli , an oomycete protist. P. humuli is an obligate biotrophic pathogen, meaning that it can only live and grow in living host tissue. P. humuli, like most mildews, is highly host-specific and thus only infected hop ( Humulus lupulus ) and also Japanese hop ( Humulus japonicas )  .
The most characteristic symptom of hop downy mildew are the “basal spikes” that form on the plant. These structures result from systemically infected shoots and are “stunted and brittle-have, downward-curled leaves from qui masses of purple to black and sporangia sporangia are Produced.”  These spikes can exist as three separate categories:
1) “Primary spikes”, which arises from infected hop crowns
2) “Secondary spikes”, which comes from infected apical meristems
3) “Aerial spikes”
Primary and secondary spikes result in stunting shoots and brittle leaves. Primary spikes also cause shortened internodes. Aerial spikes adversely affect the development of the plant, resulting in bines (the long, flexible stem the plant uses to climb). Infected leaves and cones show symptoms as well. Infected leaves will change color, ranging from purple to gray to black downy growth. Infected cones become hard and brown and display disrupted development  .
Downy mildew on hops has a polycyclic disease cycle.  Pathogen overwinters as mycelium in hop crowns. The pathogen infects crown buds, resulting in the emergence of infected shoots and primary basal spikes in the spring under correct conditions. This initiates the disease cycle.  Infected crowns can also produce uninfected shoots. The cycle becomes polycyclic as sporangiophores with sporangia emerge on the underside of infected leaves. Mature sporangia are dispersed via wind and release zoospores to infected leaves, cones, and shoots. This secondary cycle or sporulation and infection persists throughout the season. Mycelia grow systemically throughout the plant, leading to the infection of the crown and buds in which the pathogen will overwinter. 
A sexual stage also exists for P. humuli , in which antheridium fertilizes an oogonium to produce a recombinant oospore. While oospores are classically thought to be the chief survival structure of oomycetes, their role in primary infection in mildew of hops is uncertain. 
The environmental conditions favored by P. humuli are consistent with other oomycetes and fungal-like pathogens. High levels of moisture and warming temperatures facilitate the germination of many pathogens. 
The development and severity of the disease is dependent on a number of environmental factors. Studies have found that hours of relative humidity> 80%, degree-hours of wetness, and average temperature are paramount in
predicting model plants’ susceptibility to infection.  The pathogen favors extended periods of wetness, high humidity, and mildly warm temperatures ranging from 15.5 – 21 ° C (60 – 70 ° F).  Leaf infection may occur at temperatures as low as 5 ° C, where wetness persists for 24 hours or longer, indicating the primary role of moisture level in infection.  
- Fungorum Index
- USDA ARS Fungal Database
- ^ Jump up to:a b c “Downy Hop Mildew | NC State Extension Publications” . content.ces.ncsu.edu . Retrieved 2017-10-24 .
- ^ Jump up to:a b Gent, David; Ocamb, Cynthia (10 June 2009). “Predicting Infection Risk of Hop by Pseudoperonspora humuli”. Phytopathology . 99 : 1190-1198.
- ^ Jump up to:a b c d Lizotte, Erin (15 April 2015). “Managing hop downy mildew early in the season is critical” .
- Jump up^ Coley-Smith, JR 1962. Overwintering of hop downy mildewPseudoperonospora humuli(Miy and Tak.) Wilson. Ann. App. Biol. 50: 235-243.
- Jump up^ Johson, DA, et al. 2009. Downy mildew. Pages 18 to 22 in: Compendium of Hop Diseases and Arthropod Pests. American Phytopathological Society, St. Paul, MN.
- Jump up^ Burgess, Lester; Knight, Timothy (2008). “Diagnosis manual for plant diseases in Vietnam” . Archived from the original (PDF) on
|archive-date=( help ) .