Globodera rostochiensis (yellow potato cyst nematode)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Latitude/Altitude Ranges
- Air Temperature
- Natural enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Globodera rostochiensis (Wollenweber, 1923) Skarbilovich, 1959
Preferred Common Name
- yellow potato cyst nematode
Other Scientific Names
- Globodera rostochiensis (Wollenweber, 1923) Behrens, 1975
- Heterodera (Globodera) rostochiensis Wollenweber, 1923 (Skarbilovich, 1959)
- Heterodera rostochiensis Wollenweber, 1923
- Heterodera schachtii rostochiensis Wollenweber, 1923
- Heterodera schachtii solani Zimmerman, 1927
International Common Names
- English: eelworm, golden; eelworm, potato root; golden eelworm; golden nematode; golden nematode eelworm; golden nematode of potato; golden potato cyst nematode; nematode of potato, golden; potato cyst nematode; potato golden nematode; potato root eelworm
- Spanish: nematodo dorado; nematodo dorado de la papa
- French: anguillule a kyste de la pomme de terre; anguillule des racines de la pomme de terre; nématode doré; nématode doré de la pomme de terre
Local Common Names
- Denmark: kartoffelal; kartottelcystenematod
- Finland: peruna-ankeroinen
- Germany: Aelchen, Goldfarbenes Kartoffelzysten-; Aelchen, Kartoffel-; Nematode, Kartoffel-
- Iran: nematode sibsamini
- Italy: Anguillula della patata
- Netherlands: Aardappelcystenaaltje
- Norway: potetcystenematode
- Sweden: potatiscystnematod
- HETDRO (Globodera rostochiensis)
Summary of InvasivenessTop of page
G. rostochiensis is a world wide pest of temperate areas, including both temperate countries and temperate regions of tropical countries, for example India’s Nigrilis region. Distribution is linked to that of the potato crop. Potato cyst nematode is considered to have originated from the Andes region of South America, from where it spread to Europe with potatoes. The ease with which it has been transported across continents proves what a resilient pest it is. The cyst form which adheres to host roots, stolons and tubers and to soil particles during transportation gives rise to new infestations where climate and food source are both available and favourable.
Secondary means of dispersal is through the movement of contaminated farm machinery, farming implements and contaminated footwear. Cysts are also successfully spread by wind dispersal, during winter storms or sand storms where top soil is redistributed. Rain which causes flooding and water to run off fields into trenches or irrigation channels also redistributes cysts into adjoining areas.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Nematoda
- Class: Secernentea
- Order: Tylenchida
- Family: Heteroderidae
- Genus: Globodera
- Species: Globodera rostochiensis
Notes on Taxonomy and NomenclatureTop of page
G. rostochiensis was first described by Wollenweber in 1923 as Heterodera rostochiensis. The type locality is Tessin, Mecklenburg, near Rostock, Germany (Golden and Ellington, 1972) and the host neotype is Solanum tuberosum. The species should not be confused with the very rare Globodera leptonepia (Cobb and Taylor, 1953). The origin of G. rostochiensis is considered to be the Andes mountains in South America (Krall and Krall, 1978), from where it was introduced into Europe in contaminated soil adhering to potato tubers. In 1959, Skarbilovich erected the subgenus Globodera to accommodate the round cyst nematode species of which there are several, including the potato cyst nematodes. Behrens (1975) raised Globodera to generic level, before Mulvey and Stone (1976).
In 1973, Stone described a second species of potato cyst nematode, Heterodera pallida, and it is worthy of note that information concerning H. rostochiensis before this date would have involved both species.
DescriptionTop of page
The eggs of G. rostochiensis are always retained within the cyst body and no egg sacs are produced. The eggshell surface is smooth and no microvilli are present.
Length=101-104 µm; width="46"-48 µm; L/W ratio=2.1-2.5
The females emerge from the root cortex about one month to six weeks after invasion by the second-stage juveniles. They are pure white initially, turning golden yellow on maturation. Mature females are approximately 500 µm in circumference without a cone. The cuticle of the female sometimes has a thin subcrystalline layer.
Stylet length=23 µm ± 1 µm; stylet base to dorsal oesophageal gland duct=6 µm ± 1 µm; head width at the base=5.2 µm ± 0.7 µm; head tip to median bulb=73 µm ± 14.6 µm; median bulb valve to excretory pore=65 µm ± 2.0 µm; head tip to excretory pore=145 µm ± 17 µm; mean diameter of the median bulb=30µm ± 3.0 µm; mean diameter of the vulval basin=22 µm ± 2.8 µm; vulval slit length=9.7 µm ± 2.0 µm; anus to vulval basin=60 µm ± 10 µm; number of cuticular ridges between the anus and vulva=21 ± 3.0.
The female head bears one to two annules and the neck region has numerous tubercules, which can be seen using a scanning electron microscope. The head skeleton is hexaradiate and weak. The stylet is divided equally in length between the conus and the shaft. An important diagnostic feature is the backward slope of the stylet knobs. The median bulb is large and circular and well developed. The large paired ovaries often displace the oesophageal glands. The excretory pore is well defined at the base of the neck. The posterior of the female, at the opposite pole to the neck and head, is referred to as the vulval basin and is contained within a rounded depression. The vulval slit is located in the centre of this region flanked on either side by papillae, which usually fill the translucent areas of cuticle in crescentic shape, from the slit to the edge of the fenestra. The anus is distinct and is often seen at the point in the cuticle where the 'V' shape tapers to an end. The number of cuticular ridges found in the area between the anus and the edge of the fenestra is counted as an aid to identification of Globodera species. The entire cuticle is covered in small subsurface punctations.
Length without neck=445 µm ± 50 µm; width="382" µm ± 60 µm; neck length=104 µm ± 19 µm; mean fenestral diameter=19.0 µm ± 2.0 µm; anus to fenestra=66.5 µm ± 10.3 µm; Granek's ratio=3.6 ± 0.8.
Cysts contain the eggs, the progeny for the next generation, and are formed from the hardened dead cuticle of the female. Newly produced cysts may still show an intact vulval basin but older cysts, particularly those which have been in the soil for many seasons, will have lost all signs of their genitalia with only a hole in the cuticle to show the position of the fenestral basin.
Length=0.89 -1.27 mm; width at excretory pore=28 µm ± 1.7 µm; head width at base=11.8 µm ± 0.6 µm; head length=7.0 µm ± 0.3 µm; stylet length=26 µm ± 1.0 µm; stylet base to dorsal oesophageal gland duct=5.3 µm ± 1.0 µm; head tip to median bulb valve=98.5 µm ± 7.4 µm; median bulb valve to excretory pore=74 µm ± 9µm; head tip to excretory pore=172 µm ± 12.0 µm; tail length=5.4 µm ± 1.0 µm; tail width at anus=13.5 µm ± 0.4 µm; spicule length=35.0 µm ± 3.0 µm; gubernaculum length=10.3 µm ± 1.5µm.
The male is vermiform in shape with a short tail and no bursa. On fixation, the body assumes a curved shape with the posterior region twisted at a 90 degree angle to the remainder of the body. There are four incisures in the mid-body i.e. three bands which terminate on the tail. The rounded head is offset and bears 6-7 annules. The head is strongly developed having a hexaradiate skeleton. The cephalids are located at body annules 2-4 and 6-9, respectively. The stylet is strong and has backward sloping knobs. The median bulb is well developed and has a large crescentic valve. The nerve ring is located around the oesophagus between the median bulb and the intestine. The hemizonid is found 2-3 annules anterior to the excretory pore and is itself two annules in length. The hemizonion is approximately nine body annules posterior to the excretory pore and is one annule in length. The single testis fills half the body cavity. The paired spicules are arcuate and end with single tips. The gubernaculum is around 10 µm in length and 2 µm in thickness and lies in a position dorsal to the spicules.
Body length=468 µm ± 100 µm; width at excretory pore=18 µm ± 0.6 µm; head length=4.6µm ± 0.6 µm; stylet length=22 µm ± 0.7 µm; head tip to median bulb valve=69 µm ± 2.0 µm; median bulb valve to excretory pore=31 µm ± 2.0 µm; head tip to excretory pore=100 µm ± 2.0 µm; tail length=44 µm ± 12 µm; tail width at anus=11.4 µm ± 0.6 µm; hyaline tail length=26.5 µm ± 2.0 µm.
The second-stage juvenile hatches from the egg, the first moult taking place within the egg. The juvenile, like the male, is vermiform with a rounded head and finely tapered tail. The hyaline portion of the tail represents about two thirds of its length. The lateral field has four incisures in the mid-body region reducing to three at the tail terminus and anterior end. The head is slightly offset and bears four to six annules. The head skeleton is well developed and hexaradiate in form. The cephalids are located at body annules 2-3 and 6-8, respectively. The stylet is strong, the conus being about 45% of the total length. The stylet knobs are an important diagnostic feature and typically slope backwards. The median bulb is well developed and elliptical in shape, having a large central valve. The nerve ring encircles the oesophagus between the median valve and the intestine. The hemizonion is about one body annule in width and is located five body annules posterior from the excretory pore. The hemizonid is around two body annules in width and is found just anterior to the excretory pore. The gonad primordium is four-celled and located at around 60% of the body length.
Other measurements can be found in Granek (1955), Spears (1968), Green (1971), Greet (1972), Golden and Ellington (1972), Hesling (1973, 1974), Mulvey (1973), Behrens (1975), Mulvey and Golden (1983), Othman et al. (1988) and Baldwin and Mundo-Ocampo (1991).
DistributionTop of page
G. rostochiensis is widely distributed in potato-growing regions, including the temperate regions of tropical countries.
See also CABI/EPPO (1998, No. 162).
Distribution TableTop of page
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Armenia||Present||Introduced||Invasive||Iskandaryan and Arutyunyan, 1990; CABI/EPPO, 2011; EPPO, 2014|
|India||Restricted distribution||Introduced||1961||Invasive||Prasad, 1996; CABI/EPPO, 2011; EPPO, 2014|
|-Kerala||Absent, reported but not confirmed||CABI/EPPO, 2011; EPPO, 2014|
|-Tamil Nadu||Present||Introduced||1977||Invasive||Prasad, 1996; CABI/EPPO, 2011; EPPO, 2014|
|Indonesia||Restricted distribution||2003||Indarti et al., 2004; CABI/EPPO, 2011; EPPO, 2014|
|-Java||Present||Introduced||2003||Invasive||Indarti et al., 2004; CABI/EPPO, 2011; EPPO, 2014|
|Iran||Present||Gitty and Maafi, 2010; CABI/EPPO, 2011; EPPO, 2014|
|Israel||Eradicated||1954||CABI/EPPO, 2000; CABI/EPPO, 2011; EPPO, 2014|
|Japan||Restricted distribution||Introduced||1972||Invasive||Aihara et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|-Hokkaido||Present||Introduced||1972||Invasive||Inagarki, 2004; CABI/EPPO, 2011; EPPO, 2014|
|-Kyushu||Present||Introduced||1992||Invasive||Aihara et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|Lebanon||Present||Introduced||Invasive||Ibrahim et al., 2000; CABI/EPPO, 2011; EPPO, 2014|
|Malaysia||Absent, unreliable record||EPPO, 2014|
|Oman||Present||Introduced||Invasive||Mani et al., 1993; CABI/EPPO, 2011; EPPO, 2014|
|Pakistan||Present||Introduced||1980||Invasive||Munir et al., 2004; CABI/EPPO, 2011; EPPO, 2014|
|Philippines||Present||Introduced||1983||Invasive||Davide and Zorilla, 1995; CABI/EPPO, 2011; EPPO, 2014|
|Sri Lanka||Restricted distribution||Introduced||1991||Invasive||Ekanayake, 1991; CABI/EPPO, 2011; EPPO, 2014|
|Tajikistan||Restricted distribution||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Turkey||Present, few occurrences||Introduced||Invasive||Enneli and Öztürk, 1995; CABI/EPPO, 2011; EPPO, 2014|
|Algeria||Restricted distribution||Introduced||1940s||Invasive||FRÉZAL, 1954; CABI/EPPO, 2011; EPPO, 2014|
|Egypt||Absent, unreliable record||Kleynhans, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Kenya||Present||Mwangi et al., 2015|
|Libya||Present||Introduced||1981||Invasive||Ben et al., 1981; CABI/EPPO, 2011; EPPO, 2014|
|Morocco||Absent, formerly present||CABI/EPPO, 2011; EPPO, 2014|
|Sierra Leone||Present||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|South Africa||Restricted distribution||Introduced||1950||Invasive||Kleynhans, 1998; CABI/EPPO, 2011; EPPO, 2014|
|-Canary Islands||Present||Introduced||1961||Gonzalez and Phillips, 1996; CABI/EPPO, 2011; EPPO, 2014|
|Tunisia||Restricted distribution||Introduced||1977||Invasive||Kleynhans, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Zimbabwe||Absent, intercepted only||CABI/EPPO, 2011; EPPO, 2014|
|Canada||Present, few occurrences||Introduced||1962||Invasive||IPPC, 2009; CABI/EPPO, 2011; EPPO, 2014|
|-Alberta||Present, few occurrences||CABI/EPPO, 2011; EPPO, 2014|
|-British Columbia||Present, few occurrences||Introduced||Invasive||Orchard, 1965; CABI/EPPO, 2011; EPPO, 2014|
|-Newfoundland and Labrador||Restricted distribution||Introduced||1962||Invasive||Proudfoot, 1977; CABI/EPPO, 2011; EPPO, 2014|
|-Quebec||Present, few occurrences||Introduced||Invasive||Mahran et al., 2010; CABI/EPPO, 2011; EPPO, 2014|
|Mexico||Restricted distribution||Desgarennes et al., 2009; CABI/EPPO, 2011; EPPO, 2014|
|USA||Restricted distribution||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-Delaware||Eradicated||CABI/EPPO, 2011; EPPO, 2014|
|-New York||Present, few occurrences||Introduced||1941||Invasive||NAPPO; CANNON, 1941; CABI/EPPO, 2011; EPPO, 2014|
Central America and Caribbean
|Costa Rica||Absent, formerly present||CABI/EPPO, 2011; EPPO, 2014|
|Cuba||Absent, intercepted only||CABI/EPPO, 2011|
|Panama||Present||Introduced||1968||Invasive||Brodie, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Argentina||Absent, formerly present||1956||Franco et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|Bolivia||Present||Native||Bendezu et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|Brazil||Absent, invalid record||CABI/EPPO, 2011; EPPO, 2014|
|Chile||Restricted distribution||Native||Cubillos and Fernández, 1990; CABI/EPPO, 2011; EPPO, 2014|
|Colombia||Restricted distribution||Native||BAEZA, 1972; CABI/EPPO, 2011; EPPO, 2014|
|Ecuador||Restricted distribution||Native||Franco et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|Peru||Present||Native||Picard et al., 2007; CABI/EPPO, 2011; EPPO, 2014|
|Venezuela||Present||Native||DAO et al., 1971; CABI/EPPO, 2011; EPPO, 2014|
|Albania||Restricted distribution||Introduced||1982||Invasive||Jovani, 1994; CABI/EPPO, 2011; EPPO, 2014|
|Austria||Widespread||Introduced||1940s||Invasive||Riel and Mulder, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Belarus||Present||Introduced||1957||Invasive||Ponin et al., 1978; CABI/EPPO, 2011; EPPO, 2014|
|Belgium||Restricted distribution||Introduced||1949||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Bosnia-Hercegovina||Present||Ostojic et al., 2011; EPPO, 2012; EPPO, 2014|
|Bulgaria||Restricted distribution||Introduced||1970s||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Croatia||Present, few occurrences||Introduced||Invasive||Grubisic et al., 2007; CABI/EPPO, 2011; Grubisic et al., 2013; EPPO, 2014|
|Cyprus||Restricted distribution||Introduced||1974||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Czech Republic||Restricted distribution||Introduced||1954||Invasive||Zouhar and Rysanek, 2002; CABI/EPPO, 2011; EPPO, 2014|
|Denmark||Widespread||Introduced||1928||Invasive||Hansen, 1988; CABI/EPPO, 2011; IPPC, 2013; EPPO, 2014|
|Estonia||Restricted distribution||Introduced||1953||Invasive||Koppel and Tsahkna, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Faroe Islands||Present||Introduced||1951||Invasive||Jakobsen, 1973; CABI/EPPO, 2011; EPPO, 2014|
|Finland||Restricted distribution||Introduced||1946||Invasive||Heikkilä and Tilikkala, 1992; CABI/EPPO, 2011; EPPO, 2014|
|France||Restricted distribution||Introduced||1948||Invasive||Riel and Mulder, 1998; CABI/EPPO, 2011; EPPO, 2014|
|-France (mainland)||Restricted distribution||CABI/EPPO, 2011|
|Germany||Widespread||Introduced||1913||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Greece||Restricted distribution||Introduced||1954||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-Crete||Present||Introduced||2004||Invasive||Tzortzakakis et al., 2004; CABI/EPPO, 2011; EPPO, 2014|
|-Greece (mainland)||Restricted distribution||Koliopanos, 1976; CABI/EPPO, 2011|
|Hungary||Restricted distribution||Introduced||1980||Invasive||Elekes-Kaminszky et al., 2004; CABI/EPPO, 2011; EPPO, 2014|
|Iceland||Widespread||Introduced||1953||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Ireland||Restricted distribution||Introduced||1922||Invasive||Riel and Mulder, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Italy||Widespread||Introduced||1961||Invasive||CABI/EPPO, 2011; EPPO, 2011; EPPO, 2014|
|-Italy (mainland)||Widespread||Greco et al., 1994; CABI/EPPO, 2011|
|-Sicily||Present||Lombardo et al., 2011|
|Latvia||Restricted distribution||Introduced||1949||Invasive||Eglitis, 1973; CABI/EPPO, 2011; EPPO, 2014|
|Liechtenstein||Widespread||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Lithuania||Restricted distribution||Introduced||1948||Invasive||Jogaite et al., 2007; CABI/EPPO, 2011; EPPO, 2014|
|Luxembourg||Restricted distribution||Introduced||1955||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Malta||Restricted distribution||Introduced||Invasive||Lamberti et al., 1987; CABI/EPPO, 2011; EPPO, 2014|
|Netherlands||Restricted distribution||Introduced||1941||Invasive||NPPO of the Netherlands, 2013; CABI/EPPO, 2011; EPPO, 2014|
|Norway||Widespread||Introduced||1955||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Poland||Restricted distribution||Introduced||1946||Invasive||Wolny, 1992; CABI/EPPO, 2011; Przetakiewicz, 2013; EPPO, 2014|
|Portugal||Restricted distribution||Introduced||1956||Invasive||Santos et al., 1995; CABI/EPPO, 2011; EPPO, 2014|
|-Azores||Absent, invalid record||CABI/EPPO, 2011; EPPO, 2014|
|-Madeira||Present||EPPO, 2009; CABI/EPPO, 2011; EPPO, 2014|
|-Portugal (mainland)||Restricted distribution||CABI/EPPO, 2011|
|Romania||Restricted distribution||Introduced||1986||Invasive||Rojancovski and Dehebanu, 1986; CABI/EPPO, 2011; EPPO, 2014|
|Russian Federation||Restricted distribution||Introduced||1945||Invasive||Subbotin et al., 1999; CABI/EPPO, 2011; EPPO, 2014|
|-Central Russia||Present||Introduced||Invasive||Gus'kova and Makovskaya, 1977; CABI/EPPO, 2011; EPPO, 2014|
|-Eastern Siberia||Present||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-Northern Russia||Present||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-Russian Far East||Present||Introduced||Invasive||Shvydkaya and Eroshenko, 1997; CABI/EPPO, 2011; EPPO, 2014|
|-Southern Russia||Present||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-Western Siberia||Present||Introduced||Invasive||Marks and Rojancovski, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Serbia||Present, few occurrences||Introduced||Invasive||Krnnjac et al., 2002; CABI/EPPO, 2011; EPPO, 2014|
|Slovakia||Restricted distribution||Introduced||Invasive||Marks and Rojancovski, 1998; CABI/EPPO, 2011; EPPO, 2014|
|Slovenia||Present, few occurrences||Introduced||Invasive||Sirca and Urek, 2004; Sirca and Urek, 2005; CABI/EPPO, 2011; EPPO, 2014|
|Spain||Restricted distribution||Introduced||Invasive||Martinez-Beringola et al., 1988; CABI/EPPO, 2011; EPPO, 2014|
|-Balearic Islands||Present, few occurrences||Introduced||Invasive||Andrés et al., 2006; CABI/EPPO, 2011; EPPO, 2014|
|-Spain (mainland)||Restricted distribution||CABI/EPPO, 2011|
|Sweden||Widespread||Introduced||1922||Invasive||Manduric and Andersson, 2003; CABI/EPPO, 2011; EPPO, 2014|
|Switzerland||Restricted distribution||Introduced||1958||Invasive||Riel and Mulder, 1998; CABI/EPPO, 2011; EPPO, 2014|
|UK||Restricted distribution||Introduced||Invasive||Minnis et al., 2002; CABI/EPPO, 2011; EPPO, 2014|
|-Channel Islands||Present||Introduced||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|-England and Wales||Restricted distribution||Minnis et al., 2002; CABI/EPPO, 2011; EPPO, 2014|
|-Northern Ireland||Restricted distribution||Introduced||1922||Invasive||Turner and Evans, 1998; CABI/EPPO, 2011; EPPO, 2014|
|-Scotland||Present||Introduced||1913||Invasive||CABI/EPPO, 2011; EPPO, 2014|
|Ukraine||Restricted distribution||Introduced||Invasive||Pylypenko et al., 2005; CABI/EPPO, 2011|
|Australia||Restricted distribution||Introduced||1986||Invasive||Stanton, 1986; IPPC, 2008; CABI/EPPO, 2011; EPPO, 2014|
|-Victoria||Restricted distribution||Introduced||1991||Invasive||Hinch et al., 1998; CABI/EPPO, 2011; EPPO, 2014|
|-Western Australia||Eradicated||Stanton, 1987; IPPC, 2010; CABI/EPPO, 2011; EPPO, 2011; EPPO, 2014; IPPC, 2015|
|New Zealand||Widespread||Introduced||1972||Invasive||CABI/EPPO, 2000; Dale, 1973; CABI/EPPO, 2011; EPPO, 2014|
|Norfolk Island||Present||Introduced||Invasive||Marshall, 1998; CABI/EPPO, 2011; EPPO, 2014|
History of Introduction and SpreadTop of page
The potato was first domesticated and cultivated by the Andean peoples as long as 8,000 years ago. When Peru was invaded by the Spanish in 1531, potatoes were already an important source of food for these peoples.
The potato was bought to Europe sometime in the sixteenth century and grown in the Seville region of Spain. An independent introduction of potato occurred in England around 1590, but not from the same source. From these introductions, potatoes spread across Europe and into other parts of the world. By the nineteenth century the potato had become more widely grown and had been dispersed over wider geographical areas by trade and the movement of populations.
With the advent of the hugely damaging potato blight (Phytophthora infestans) in the mid-nineteenth century, many people in Europe found starvation confronting them, particularly the Irish who by now were heavily dependant upon potatoes as a staple food. There are no records of potato cyst nematode damage from this time but whether it was not recognised or had not yet been introduced to areas where potato was grown is unclear. The potato now being a significant food source meant that Europeans began to search for varieties with resistance to blight from the area of origin of potatoes, South America. The new genetic material was also used to breed varieties of potato adapted to longer day lengths and with generally better adaptation to European growing conditions.
In all probability, potato cyst nematodes were introduced to Europe with the new genetic material around 1850. In 1881, Kuhn made the first record of cyst nematode damage to potato; although at the time it was described as the beet cyst nematode Heterodera schachtii. The thirty year period between 1881 and bringing in new potato material, most probably with adhering soil and cysts, was enough time for cyst nematodes to build up to damaging levels in the field and to cause crop loss.
In 1923, Wollenweber described the cysts he found on potato as the new species Heterodera rostochiensis, named after the type locality of Rostock in Germany. Later, H. pallida was described by Stone (1972) as a second species of PCN occupying an almost identical ecological niche, but having significantly different biological, morphological and biochemical characteristics.
In North America, G. rostochiensis was first discovered on Long Island, New York (Nassau County) in 1941 after a potato grower noticed isolated areas of poor plant growth (Cannon, 1941). The nematode may have come in on equipment returned from Europe after WWI - the field was an old military camp which became a potato field. The discovery led to establishment of a Federal Quarantine based on the Golden Nematode Act. Growers moved to Western New York following urbanization, and subsequent outbreaks appeared; required 8-10 years to reach a detectable level.
In the early 1970's, scientists in Mexico discovered an infestation of G. rostochiensis in the state of Guanajuato (Iverson, 1972). It was reported in South Africa in 1971 from an irrigated farm near Pretoria and then from small farms around Johannesburg and Bon Accord. Spread in Africa is detailed by Kleynhans (1998).
Two of the most recent outbreaks of Globodera rostochiensis have occurred in Quebec, Canada in 2006 (Sun et al., 2007) and in the Balearic Island of Mallorca, Spain 2006 (Andrés et al., 2006), so demonstrating the continuing need for good phytosanitary regulations and stringent quarantine measures.
IntroductionsTop of page
Risk of IntroductionTop of page
Potato cyst nematodes are A2 quarantine pests for EPPO (OEPP/EPPO, 1978, 1981, 2007). They are also of quarantine significance for APPPC and NAPPO. G. rostochiensis is also a quarantine pest for CPPC and IAPSC.
Virtually all areas within the EPPO region that grow potatoes are already contaminated with potato cyst nematodes. These areas are generally closely monitored. It is important to keep seed potato areas as free as possible of potato cyst nematodes. Domestic measures and import controls are justified as they help to reduce spread and introduction of new pathotypes into established areas. G. rostochiensis still seems to be the dominant species throughout Europe, with the exception of England, UK, where G. pallida is common.
Habitat ListTop of page
|Coastal areas||Present, no further details||Harmful (pest or invasive)|
|Soil||Present, no further details||Harmful (pest or invasive)|
|Stored products||Present, no further details||Harmful (pest or invasive)|
|Cultivated / agricultural land||Present, no further details||Harmful (pest or invasive)|
|Arid regions||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
The major hosts of G. rostochiensis are restricted to the Solanaceae, in particular potato, tomato and aubergine (Ellenby, 1945, 1954; Mai, 1951, 1952; Winslow, 1954; Stelter, 1957, 1959, 1987; Roberts and Stone, 1981; Sullivan et al., 2007). A number of weeds in the Solanaceae are also hosts.
In addition to the main hosts listed, the following plants are hosts of G. rostochiensis:
Datura tatula, D. ferox, Hyoscyamus niger, Lycopersicon aureum, L. glandulosum, L. hirsutum, L. esculentum peruvianum, L. pimpinellifolium, L. pyriforme, L. racemigerum, Nicotiana acuminata, Physalis longifolia, P. philadelphica, Physochlaina orientalis, Salpiglossis sp., Saracha jaltomata, Solanum acaule, S. aethiopicum, S. ajanhuiri, S. ajuscoense, S. alandiae, S. alatum, S.americanum, S. anomalocalyx, S. antipoviczii, S. armatum, S. ascasabii, S. auriculatum, S. asperum, S. aviculare, S. berthaultii, S. blodgettii, S. boergeri, S. brevidens, S. brevimucronatum, S. bukasovii, S. bulbocastanum, S. calcense,S. calcense × S. cardenasii, S. caldasii, S. canasense, S. capsicibaccatum, S. capsicoides, S. cardiophyllum, S. carolinense, S. chacoense, S. chaucha, S. chenopodioides, S. chloropetalum, S. citrillifolium, S. coeruleiflorum, S. commersonii, S. curtilobum, S. curtipes, S. demissum, S. demissum × S. tuberosum, S. dulcamara, S. durum, S. elaeagnifolium, S. ehrenbergii, S. famatinae, S. fraxinifolium, S. fructo-tecto, S. garciae, S. gibberulosum, S. giganteum, S. gigantophyllum, S. gilo, S. glaucophyllum, S. goniocalyx, S. gourlayi, S. gracile, S. heterophyllum, S. heterodoxum, S. hirtum, S. hispidum, S. indicum, S. integrifolium, S. intrusum, S. jamesii, S. jujuyense, S. juzepczukii, S. kesselbrenneri, S. kurtzianum, S. lanciforme, S. lapazense, S. lechnoviczii, S. leptostygma, S. ligustrinum, S. longipedicellatum, S. luteum, S. macolae, S. macrocarpon, S. maglia, S. malinchense, S. mamilliferum, S. marginatum, S. mauritianum, S. melongena, S. miniatum, S. mochiquense, S. multidissectum, S. muricatum, S. neocardenasii, S. nigrum, S. nitidibaccatum, S. ochroleucum, S. okadae, S. oplocense, S. ottonis, S. pampasense, S. parodii, S. penelli, S. photeinocarpum, S. phureja, S. pinnatum, S. pinnatisectum, S. platense, S. platypterum, S. polyacanthos, S. polyadenium, S. prinophyllum, S. quitoense, S. radicans, S. raphanifolium, S. rostratum, S. rybinii S. salamanii, S. saltense, S. sambucinum, S. sanctae-rosae, S. sarrachoides, S. scabrum, S. schenkii, S. schickii, S. semidemissum, S. simplicifolium, S. sinaicum, S. sisymbrifolium, S. sodomaeum, S. soukupii, S. sparsipilum, S. spegazzinii, S. stenotomum, S. stoloniferum, S. suaveolens, S. subandigenum, S. sucrense, S. tarijense, S.tenuifilamentum , S. tlaxcalense, S. tomentosum, S. toralopanum, S. triflorum, S. tuberosum ssp. andigena, S. tuberosum ssp. tuberosum, S. tuberosum 'Aquila', S. tuberosum 'Xenia N', S. utile, S. vallis-mexicae, S. vernei, S. verrucosum, S. villosum, S. violaceimarmoratum, S. wittmackii, S. wittonense, S. xanti, S. yabari and S. zuccagnianum.
Note Oxalis tuberose (Oca), has been extensively tested in host range tests by Sullivan et al. (2007) and has been declared a non-host on this basis.
Host Plants and Other Plants AffectedTop of page
|Datura stramonium (jimsonweed)||Solanaceae||Other|
|Lycopersicon pimpinellifolium (currant tomato)||Solanaceae||Other|
|Solanum aviculare (kangaroo apple)||Solanaceae||Other|
|Solanum gilo (gilo)||Solanaceae||Other|
|Solanum lycopersicum (tomato)||Solanaceae||Main|
|Solanum marginatum (white-edged nightshade)||Solanaceae||Other|
|Solanum mauritianum (tobacco tree)||Solanaceae||Other|
|Solanum melongena (aubergine)||Solanaceae||Main|
|Solanum nigrum (black nightshade)||Solanaceae||Other|
|Solanum quitoense (naranjilla)||Solanaceae||Other|
|Solanum sarrachoides (green nightshade)||Other|
|Solanum tuberosum (potato)||Solanaceae||Main|
Growth StagesTop of page Pre-emergence, Seedling stage, Vegetative growing stage
SymptomsTop of page
Potato cyst nematodes, in common with other cyst nematodes, do not cause specific symptoms of infestation. Initially, crops will display patches of poor growth and plants in these patches may show chlorosis and wilting. When the tubers are harvested there will be a yield loss and tubers will be smaller. To be confident that these symptoms are caused by potato cyst nematodes and to give an indication of population density, soil samples must be taken or the females or cysts must be observed directly on the host roots. In heavily infested soils, plants have reduced root systems and often grow poorly due to nutrient deficiencies and to water stress. Plants may senesce prematurely as they are more susceptible to infection by fungi such as Verticillium spp. when heavily invaded by potato cyst nematodes.
Direct damage to roots and the yield of tubers
The infective second stage juveniles of both G. rostochiensis and G. pallida respond to environmental conditions when hatching. There is a short period of time for the second stage juvenile to locate a host root and begin the process of invasion, usually just behind the root tip. The juveniles then position themselves next to the stele within the root where, after a few hours, they will establish a feeding site (syncytium), which will become their nutrient source until their death. If a susceptible variety of potato is planted the plants will soon show signs of attack particularly when nematode density is high. In resistant plant varieties juveniles still hatch from the cyst and invade the plant roots, but they are unable successfully to establish a feeding site or syncytium. In this situation, males are more likely to be produced than females, as males have negligible nutrient requirements compared to females. Nevertheless, even resistant crops may show signs of attack.
The reduction in the yield of potato tubers, depending on the cultivar grown, is also related to or dependent on the plant's ability to tolerate the effects of nematode attack. The effects of potato cyst nematode on the plant include water stress and early senescence of the leaves. A heavily infested plant is unlikely to produce 100% ground cover with its reduced canopy of leaves. Many field studies have monitored the progression of ground cover by leaves and correlated the findings with yields (see Trudgill et al., 1998).
List of Symptoms/SignsTop of page
|Leaves / abnormal colours|
|Leaves / wilting|
|Roots / cysts on root surface|
|Roots / reduced root system|
|Vegetative organs / surface cracking|
|Whole plant / dwarfing|
|Whole plant / early senescence|
Biology and EcologyTop of page
Eggs contained within cysts are the persistent stage of the life cycle; new cysts contain around 500 eggs. Some eggs are able to survive within the cyst for as long as 30 years, although by this time very few are viable. Hatching in water is normal, especially within the 'hatching season' when most of the environmental factors are stable. Hatching is increased by the stimulus of a hatching factor, found in host root diffusate (Perry and Beane, 1988). This appears to affect the permeability of the eggshell lipoprotein membrane (Atkinson and Ballantyne, 1977a, b).
Many workers have studied the hydration of second-stage juveniles (Clarke and Perry, 1985; Perry, 1986; Perry and Feil, 1986; Holz et al., 1998) and pre-hatch behaviour within the egg (Doncaster and Shepherd, 1967; Doncaster and Seymour, 1973), as well as the responses of potato cyst nematodes to various hatching agents at different concentrations (Clarke and Hennessy, 1987). Studies have also been made of gene expression at hatching (Jones et al., 1997). Other factors affecting hatching are soil moisture, aeration and pH (Shepherd and Clarke, 1971). Soil temperature or the rate of heat accumulation (Jones and Parrot, 1969) are also important and Jones (1975) showed that the development of the nematode is governed by the rate of heat accumulation, except at extreme temperatures.
Nematode densities remain low at low temperatures (Mugniery, 1978; Franco, 1979; Hominick, 1979; Inagaki, 1984). The optimum temperature for the hatch of G. rostochiensis is about 15°C, with the largest proportion of adults in a population at 650-830 day degrees over a basal temperature of 4.4°C (Evans, 1968). Some populations seem to have adapted to climates with lower temperatures, as in the Scottish population from Ayrshire, which is able to mature even on early harvested potatoes (Ellenby and Smith, 1975; Hominick, 1979, 1982). The most northerly known point of survival for G. rostochiensis is the Finnish Arctic Circle after several years with long warm summers (Sarakoski, 1976).
The infective juveniles, as with most other cyst nematodes, penetrate the host root just behind the root tip. From this point the juveniles move up or down the root until they receive a specific signal, probably of a chemical nature, to set up a feeding site in the form of a syncytium. The ultrastructure of the feeding site and nematode interaction with the syncytium have been studied intensively (Wyss and Zunke, 1986; Golinowski et al., 1997; Endo, 1998). Infective second-stage juveniles that penetrate the pericycle cells of the plant are more likely to become males, whereas those that penetrate the procambial cells tend to become females (Golinowinski et al., 1997). Within a few hours of settling, the juvenile probes the selected cell, inserts its stylet into it while remaining motionless for several hours; the stylet is then withdrawn and re-inserted into the same cell. A secretory product from the oesophageal glands is injected via the stylet into the selected cell. The initial syncytial cell (ISC) is now altered to provide large amounts of nutrients to the developing nematode. The syncytium undergoes major changes, for example lysis of inner cell walls, formation of cell wall ingrowths next to plant conductive tissue, appearance of numerous lipid bodies and enlargement of now amoeboid nuclei. However, many events are still not fully understood. In resistant plants, the juvenile may try to form a syncytial feeding site but the walls of cells involved usually thicken and cells may die. This prevents the ready movement of nutrients to the juvenile (Rice et al., 1986; Robinson et al., 1988).
The life cycle takes approximately 45 days, during which time the males will moult and become vermiform, leave the host root and fertilize as many females as possible before dying about 10 days or so after first leaving the root (Evans, 1970). The females during this time have become saccate and their posterior ends have protruded through the root cortex, ready for mating.
Globodera spp. do not produce egg sacs or form a gelatinous mass into which eggs are deposited. However, it is known that pheromones are secreted which attract males (see data sheet on Globodera tabacum) (Green and Miller, 1969; Green and Plumb, 1970; Mugniery, 1979; Mugniery et al., 1992). Further studies into the nature of the pheromones (considered to be polar compounds by Green, 1980) have used electrophysiological techniques and behavioural assays (Riga et al., 1997). Extracellular recordings of electrical activity within the cephalic region of individual males were taken and correlated with exposure to extracts of females. The males exhibited a specific reaction to pheromones of females of G. rostochiensis, and it is also known that they will mate with females already fertilized. Trudgill (1967) noted that more males are present in a population when environmental conditions are poor, for example, lack of feeding sites due to overcrowding; the vermiform males are not known to feed (Evans, 1970).
Raising antibodies to nematodes permits investigations that were not previously possible (Fox and Atikinson, 1985b; Curtis, 1996), particularly studies on nematode secretions and their function and interaction within the host plant (Jones and Robertson, 1997; Jones and Harrower, 1998). The role of cuticular secretions in plant parasites is not yet fully understood (Forrest et al., 1989). Changes occur in the cuticle of second-stage juveniles when they begin to feed: the cuticle will alter due to natural growth patterns and, with the onset of moulting, changes occur that appear to be linked to interaction with the plant host (Jones and Robertson, 1997).
Recent observations show that one of the free radicals produced by plants is in the form of hydrogen peroxide, which is produced in response to nematode invasion and is in all probability broken down by an enzyme secreted by the nematode’s surface coat, thioredoxin peroxidase. Various other proteins have been identified from the surface coat and hypodermis of the nematode, such as the lipid binding protein GpFAR-1. This probably plays a role in the host defence signalling pathways as other plant parasitic nematodes, but not free-living types, also have similar compounds in their genetic makeup. Plant parasitic nematodes have probably evolved similar methods with which to protect and help to conceal themselves from the host plant.
Studies to explore the function of nematode genes, using the molecular tool RNAi, have given detailed insights into the host parasite relationship. Details of some of these findings regarding plant parasitic nematodes can be found in a review by Lilley et al., 2007. Genes related to the production of pectate lyases used for host penetration by G. rostochiensis (Kudla et al., 2007) provide an insight into the complexity of studies now being undertaken to elucidate aspects of the plant-host relationship.
Many studies are now focused on the interactions of chemical pathways, genetic function and metabolic concepts too numerous to cover here. Molecular aspects of plant-nematode interactions are reviewed by Gheysen and Jones (2006).
ClimateTop of page
|B - Dry (arid and semi-arid)||Preferred||< 860mm precipitation annually|
|BS - Steppe climate||Preferred||> 430mm and < 860mm annual precipitation|
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
|D - Continental/Microthermal climate||Tolerated||Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)|
|Ds - Continental climate with dry summer||Tolerated||Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)|
|ET - Tundra climate||Tolerated||Tundra climate (Average temp. of warmest month < 10°C and > 0°C)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Absolute minimum temperature (ºC)||-80|
|Mean annual temperature (ºC)||10||25|
|Mean maximum temperature of hottest month (ºC)||27|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Means of Movement and DispersalTop of page
Generally, cyst nematodes build up in numbers over time, usually in patches. When the areas of infestation are disturbed i.e. by agricultural machinery, implements or farm workers, the cysts often adhering to clods of soil are easily transferred to uninfested areas. This can occur at all levels, locally and internationally. Transportation of crop plants and tubers also requires extremely high levels of hygiene to prevent movement. Packaging materials can also harbour cysts, which will readily transfer to a new favourable environment.
Natural Dispersal (Non-Biotic)
Wind can have an impact by lifting and spreading cysts in dry soil and plant debris from one area to another. Run-off flood water from infested areas can pose a threat as dry cysts float and can be carried some distance from the original source to start new infestations in local situations.
Vector Transmission (Biotic)
It is known that cyst nematodes are capable of passing through the digestive tract of farm animals and are excreted intact in a viable condition ready to begin a new infestation. Soil debris transported on animal hoofs can move cysts from one area to another.
Probably the most likely cause of new introductions to different countries has been unintentional and has come with the advent of trade and travel.
Pathway CausesTop of page
|Breeding and propagation||Potato breeding material from South America to Ireland||Yes||Yes||Turner and Evans, 1998|
|Crop production||Peru to Europe||Yes||Yes||Turner and Evans, 1998|
|Digestion and excretion||Yes|
|Flooding and other natural disasters||Yes||Been and Schomaker, 2006|
|Food||Yes||Yes||Turner and Evans, 1998|
|Garden waste disposal||Yes|
|Military movements||Yes||Yes||Brodie, 1998|
|People sharing resources||Yes|
|Seed trade||Netherlands to South America||Yes||DAO et al., 1971|
Pathway VectorsTop of page
|Clothing, footwear and possessions||Cysts||Yes||Been and Schomaker, 2006|
|Containers and packaging - non-wood||Cysts||Yes||Inagarki, 2004|
|Containers and packaging - wood||Cysts as contamination||Yes|
|Land vehicles||Cysts as contamination, juveniles||Yes||Yes|
|Machinery and equipment||Cysts||Yes||Yes||Been and Schomaker, 2006|
|Plants or parts of plants||Cysts, juveniles, eggs||Yes||Yes|
|Soil, sand and gravel||Cysts in soil and dust storms, juveniles||Yes||Yes|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bulbs/Tubers/Corms/Rhizomes||cysts||Yes||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Growing medium accompanying plants||cysts||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Roots||cysts; eggs; juveniles||Yes||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Seedlings/Micropropagated plants||cysts; eggs; juveniles||Yes||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|True seeds (inc. grain)|
Impact SummaryTop of page
Economic ImpactTop of page
The actual cost of damage caused by G. rostochiensis is difficult to determine, but potato cyst nematodes (see also Globodera pallida) cause extensive damage, particularly in temperate areas and particularly when virulent pathotypes occur and any resistance has failed. The situation is worse with G. pallida, where commercial cultivars with good resistance are few and often have other undesirable properties. Damage is related to the number of eggs per unit of soil, and is reflected in the weight of tubers produced (see Seinhorst, 1986 for further details). Severe infestation with G. rostochiensis and/or G. pallida can result in yields that are smaller than the quantity of seed originally planted (Oerke et al., 1994).
South and Central America
In Chile, microplot studies were carried out to determine the effect of densities of G. rostochiensis (as eggs/g soil) on the yield of summer, winter and spring sown potatoes (Moreno et al., 1984; Greco and Moreno, 1992). For the summer crop, a damage threshold for tuber yield of 1.3 eggs/g soil was determined. At population densities greater than this, yields were greatly suppressed. At initial inoculum levels of 12, 32 and 128 eggs/g soil, reductions in yield of 20, 50 and 70%, respectively, were recorded. Winter-sown potatoes indicated a tolerance limit of 0.8 eggs/g soil. Spring-sown potatoes had a tolerance limit of 1.56 eggs/g soil. Yield losses of 20, 50 and 90% were obtained with population densities of 9, 28 and 128 eggs/g soil, respectively.
In Panama, Pinochet (1987) estimated that average annual crop losses due to G. rostochiensis, G. pallida and species of Meloidogyne were 10-30%.
In Canada, G. rostochiensis was found in Newfoundland in 1962, followed by G. pallida 15 years later. Since their discovery, ca $Can 800,000/year has been spent on control and research (Miller, 1986). In 2006, G. rostochiensis was discovered in a field in the Saint-Amable region of Quebec, Canada. (Sun et al., 2007).
Between 1967 and 1973 in the UK, Brown and Sykes (1983) investigated the losses due to G. rostochiensis and G. pallida in Staffordshire and Worcestershire. At densities of 0-24 eggs/g soil, losses of 6.25 t/ha per 20 eggs/g soil were recorded. At 40-160 eggs/g soil, losses of 1.67 t/ha were recorded. An average loss of 2.75 t/ha per 20 eggs/g soil was determined for all population densities. The maximum crop loss was 22 t/ha.
The economic threshold for G. rostochiensis in the UK is ca 15 eggs/g soil. Unless control measures are implemented above this threshold, crop losses can be as high as 80%. Dale (1988) found that, on average, the losses are 3-4 t/ha per 10 eggs/g soil at planting time. A loss figure of 9% nationwide does not take into account any indirect losses sustained as a result of control or quarantine measures, or for growing crops which give a lower return (Evans and Brodie, 1980; Brodie, 1984). The annual UK expenditure on nematicides is £9 million and these are applied over ca 40,000 ha, a cost of £210-250/ha; losses in spite of control measures are ca £6 million (Trudgill et al., 1987; Dale, 1988).
In Germany, Engel et al. (1982) estimated the marketable crop losses dependent on G. rostochiensis infestation density in long-term trials using regression analysis. They estimated crop losses of 11, 27 and 43% at nematode populations of 100, 1000 and 10,000 larvae/100 cm³, based on an attainable yield of 22.4 t/ha on nematode-free plots. As populations of 500 juveniles/100 cm³ are regularly reached, losses on affected areas were estimated at 10-15% (Kleinhempel, 1986).
Greco et al. (1982) reported that the loss threshold for G. rostochiensis and G. pallida at three Italian sites was between 1.4 and 2.1 eggs/g soil. In a field trial in 1982, a yield of 23.9 t/ha was achieved on nematicide treated areas. Without control, G. rostochiensis reduced the yield by 76%; the marketable yield of 13.75 t/ha on treated areas was reduced (by 85%) to 1.96 t/ha (Greco et al., 1984).
In Arcadia, Greece, application of nematicides to control G. rostochiensis increased the yield of potatoes grown on sandy soils to 31 t/ha. In untreated areas, the loss was 37% (Kalyviotis-Gazelas, 1982).
In Poland, monoculture has been used as a method of growing potatoes. This has led to a build-up in the population of nematodes, particularly G. rostochiensis. Zawislak et al. (1981) reported that after growing potatoes in the same ground in Balayny for 4 years, the number of viable cysts rose to 63/100 g soil, though yield was unaffected. In the fifth year, however, there were 180 cysts/100 g soil (up to 3000 cysts/plant) and yields fell by 72%.
G. rostochiensis is a principal potato pest in Byelorussia. At a population of 1000 juveniles/100 cm³ soil, crop losses of 17-20% were recorded in susceptible varieties. At population densities of 500, 10,000 and 25,000 juveniles/100 cm³ soil, losses of 31, 45 and 74%, respectively, were recorded. With repeated cropping (as in the Polish monoculture systems), the population of G. rostochiensis could reach 38,000/100 cm³ (Gladkaja and Korzhentsevskaya, 1985).
Risk and Impact FactorsTop of page Invasiveness
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Pioneering in disturbed areas
- Highly mobile locally
- Long lived
- Fast growing
- Has high reproductive potential
- Has propagules that can remain viable for more than one year
- Has high genetic variability
- Changed gene pool/ selective loss of genotypes
- Host damage
- Increases vulnerability to invasions
- Modification of nutrient regime
- Modification of successional patterns
- Negatively impacts agriculture
- Negatively impacts cultural/traditional practices
- Negatively impacts livelihoods
- Competition - monopolizing resources
- Pest and disease transmission
- Interaction with other invasive species
- Parasitism (incl. parasitoid)
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
Uses ListTop of page
- Laboratory use
- Research model
- Test organisms (for pests and diseases)
DiagnosisTop of page
Following the demonstration that isoelectric focusing could be used to identify G. rostochiensis and G. pallida because they display different protein profiles (Fleming and Marks, 1983), DNA based techniques of identification have been developed (Bakker and Gommers, 1986; Marshall and Crawford, 1987; Bossis and Mugniery, 1993; Thiery and Mugniery, 1996). More recently, these techniques have been improved upon and identification and the use of species-specific primers (Mulholland et al, 1996; Fullaondo et al, 1999), for RAPD PCR RFLP, and AFLP multiplex PCR are now recommended techniques for molecular identification. Micro array techniques are now also more commonly used for diagnostic work (Picard et al., 2007).
Molecular diagnostics are now the “norm” having advanced dramatically in the last decade. Their discriminatory powers can provide a deeper level of species separation and identification. Molecular diagnostics, although still comparatively expensive, require less skill then traditional methods and can give a greater throughput and standardisation of samples on a routine basis. It is also possible to determine the identity of known pathotypes of potato cyst nematode using a high performance capillary electrophoresis (CE) technique. This uses sodium dodecyl sulphate (SDS) capillary gel electrophoresis to display polypeptide profiles (Hinch et al., 1998). Other means of discriminating nematode species using biochemical techniques have led to the use of monoclonal antibodies as a diagnostic tool (Fox and Atkinson, 1985a; Curtis, 1996). Perry and Jones (1998) reviewed further in-depth biochemical and genetic studies, including the disruption of the nematode life cycle by the addition of resistance genes or the insertion of other genetic constructs into the plant genome by genetic engineering techniques (Burrows, 1996; Burrows and De Waele, 1997).
Detection and InspectionTop of page
Potato cyst nematodes, in common with other cyst nematodes, do not cause specific symptoms of infestation. Initially, crops will display patches of poor growth and these plants may show chlorosis and wilting. When the tubers are harvested there will be a yield loss and tubers will be smaller. To be confident that these symptoms are caused by potato cyst nematodes and to give an indication of population density, soil samples must be taken or the females or cysts must be observed directly on the host roots. Detection based on host plant symptoms and identification by morphological and molecular methods are detailed in EPPO (2009).
Surveys of the numbers and distribution of potato cyst nematode are prerequisites for making informed choices for their management. Samples taken within a field are either to check whether potato cyst nematode is present or not in the field for statutory purposes or to determine the extent of the infestation, which might include a determination as to what species is present.
At one time, it was considered that nematodes had a haphazard distribution in the field but this has been disproved. Aspects of the environment and ecological factors such as disease, predators and soil type favour aggregated distributions. Many models help to describe distributions, for example Taylor's Power Law (Taylor, 1961), Iwao's regression model (Allsopp, 1990) and others. However, geostatistical techniques may provide a more purposeful definition of the spatial distribution of nematodes. Although these techniques are young, 3-D maps can be generated to study nematode population levels more effectively. These methods have already proved useful in mapping other types of field related data (Chellemi et al., 1988) and have recently been applied to the distribution of potato cyst nematodes (Evans et al., 2003).
Similarities to Other Species/ConditionsTop of page
G. rostochiensis is very similar to the other members of the round cyst nematode group. Its morphology, phenotypic appearance and biology as well as host range can make difficult the initial diagnosis of the species from Globoderatabacum tabacum (Lownsbery and Lownsbery, 1954), Globoderatabacum solanacearum (Miller and Gray, 1972), Globoderatabacum virginiae (Miller and Gray, 1968) and Globoderapallida (Stone, 1972). G. rostochiensis, the G. tabacum 'complex' and Globodera spp. from Compositae all produce females that have a golden phase upon maturity just before death, after which the cyst cuticle of all becomes brown. The second-stage juveniles are similar in, for example, body and stylet length. Other studies have shown hybridisation of G. rostochiensis with the G. tabacum group, proving a strong genetic link. Their host ranges are also very similar (Roberts and Stone, 1981; Stelter, 1987). G. pallida is often found together with G. rostochiensis but has white or cream females, the second-stage juveniles are longer and have longer stylets and the cyst has a larger Granek's ratio and usually fewer ridges between the anus and fenestra. The two species can be differentiated using simple isoelectric focusing techniques. For details, see the Diagnosis section.
Prevention and ControlTop of page
To prevent further spread of potato cyst nematodes into uninfested areas, several methods are used. These include legislation on the movement of seed potatoes, nursery stock, flower bulbs and soil. These apply internationally and nationally.
The specific EPPO quarantine regulations (OEPP/EPPO, 1990; EPPO, 2007) for these nematodes require that fields in which seed potatoes or rooted plants for export are grown are inspected by taking soil samples according to an EPPO-recommended method (OEPP/EPPO, 1991; EPPO, 2007) and must be found free from viable cysts of both species of potato cyst nematode. The sampling must be performed after harvest and after removal of the previous potato crop.
Quarantine is a very necessary and often expensive way of attempting to limit the damage caused by disease organisms such as nematodes. Methods to limit or prevent the introduction of alien or existing pests by providing ways of very accurate identification plus sensible legislation (MAFF, 2000) are already used in many countries. Continuous records must be kept of previous land usage and crop rotations. Various organizations throughout the world help to back up the phytosanitary regulations.
The previous PCN Control Directive for the European Union was in place for several decades, over which time many practices within the potato industry have changed and a lot more has been learned from intensive studies on this pest. New European legislation was introduced on 1 July 2010. The revised Control Directive (2007/33/EC) was adopted by the Council of Ministers and has been published in the Official Journal of the European Communities in June 2007.
Physical barriers help to confine pests to their own locality, for example, seas, mountains and other natural phenomena. Trade is probably the keystone to the problem of spread (Parker, 2000). Trade is essential and, when it comes to the movement of soil and plants, is of unparalleled importance with regard to nematode quarantine. Plant parasitic nematodes occur worldwide on virtually every crop, and plant movement can probably be blamed for all potato cyst nematode occurrences outside South America.
1. Check that machinery is thoroughly clean and free from plant debris.
2. Do not return soil to fields as it may cause infestation of potato cyst nematode to spread.
3. Clean soil from potato tubers and have the soil tested to be sure of non-transference of potato cyst nematode.
4. Make sure that laboratories that test soil for potato cyst nematode are properly qualified and that they test 500 g of soil per sample.
5. Grow susceptible and resistant varieties of potato alternately, thus reducing the possibility of selecting a highly virulent or new pathotypes.
For further details, see Seinhorst (1986).
Rotation is frequently used to reduce population densities of the potato cyst nematodes, G. rostochiensis and G. pallida. The major hosts of these two species are restricted to the plant family Solanaceae, with the main commercial crops being potato, tomato and aubergine. When these crops are grown in monoculture for several seasons in infested soil, nematode densities can increase to extremely high levels and crop yields become uneconomical. To reduce nematode population densities, non-host crops such as barley are grown between host crops. Magnusson (1987) recorded an 87% decline in G. rostochiensis population density using this type of rotation. Whitehead (1995) also found good control of G. rostochiensis when barley was grown in infested microplots. The annual decline rate of potato cyst nematode in soil is variable, depending on the non-hosts used, the initial population density, various soil-related factors and the population under study. If the reduction of population densities by rotation alone is too slow, then additional means of control may be necessary, such as the use of resistant cultivars, trap cropping or nematicides.
Trap cropping has been used successfully for the reduction of cyst nematode populations (Halford, et al., 1999). Potatoes are grown in order to cause the second stage juveniles to hatch. These are given sufficient time to penetrate the roots and develop into young adults. By monitoring soil temperature from the date of planting, fertilization and formation of new eggs can be avoided by destroying the crop some 6 or even 7 weeks after planting, before too many heat units have accumulated. If crop destruction is left too late, the nematode density will increase. Using this method, G. rostochiensis populations have been reduced by more than 80% (Halford et al., 1999).
The use of resistant potato clones as trap crops has been used in field trials in Northern Ireland. Ten clones had a strong hatch-inducing effect and resistance to currently known PCN pathotypes. Some of these showed potential for further development to reduce high population levels of PCN in the field and for the organic potato market (Turner et al., 2006) Another solanaceous host, S. sisymbriifolium, acts in a similar way, stimulating hatch without the nematode being able to complete its life-cycle, with the added benefit of then being used as a green manure. The seed line of S. sisymbriifolium is probably important as seed lines used for trap cropping are listed by Scholtze (2000) as totally resistant to PCN, whereas two lines (nos. 72 and 121) were recorded as poor hosts (with less then 5 cysts per plant in a host test performed by Stelter (1987). Where the nematode density is reduced, there will be a significant yield benefit for any subsequent potato crop.
Solarization is a good method of killing nematodes in very hot climates. The soil is covered with two layers of polyethylene, allowing the soil underneath to heat up quickly. In Oman, Mani et al. (1993) found that 62 days of solarization reduced G. rostochiensis population density by 95%. In New York State, USA, 97% of nematode eggs were inactivated in the top 10 cm of soil (LaMondia and Brodie, 1984.). Solarization in cooler climates and at depths greater than 10 cm is much less effective.
Natural parasites and biological control options are being studied intensively in the search for natural ways of controlling plant parasitic nematode populations without the use of nematicides, which are highly toxic and a burden on the environment. These biological control agents are part of a grander objective to manage nematodes more effectively using a variety of biological strategies that include trap cropping and rotation.
Work on biological control agents was started in the late 1930s (Linford et al., 1938) and still continues (Crump and Flynn, 1995; Segers et al., 1996; Crump, 2004). At present, there is still no commercial biological product available to control potato cyst nematodes. The majority of studies in the late 1990s have concentrated on the fungal control agents Pochonia, Hirsutella and Arthrobotrys and the bacterium Pasteuria.
Several workers (e.g. Roessner, 1986) have studied biological control of G. rostochiensis in pots and in vitro. Almost no field trial data are available. This is probably due in part to the logistics of such operations e.g. producing enough inoculum for field scale trials. Also, some tests do not produce the expected results for reasons as yet undefined, but are probably related in some way to the physiology or ecology of the nematode or to the host parasite relationship.
Pochoniachlamydosporia will infect young females in pots but is less effective when potatoes are grown in the presence of low nematode population densities.
Progress in the area of biological control requires a better understanding of the population dynamics of potato cyst nematodes and its parasites (Davies et al., 1991; Davies, 1998). A variety of factors, such as plant host, the action of root exudates, soil type and the mode of parasitism of the control organism, interact to determine success in the biological control of the nematode. Cyst nematodes may be more susceptible to infection at certain points in their life cycle. For example, the three major fungal parasites Pochonia chlamydosporia, Fusarium oxysporum and Cylindrocarpon destructans, have all been detected throughout the potato cyst nematode life cycle but the most active will vary at different times of the cycle (Crump, 1987).
At the molecular level, the high specificity known to occur between cyst nematodes and their plant hosts is important, in terms of intra- and inter-specific variation, particularly with regard to virulence and avirulence. Work by Segers et al. (1996) on the effect of a protease-like enzyme (designated VCPI), from the nematophagous fungus Pochonia chlamydosporia, showed that when this enzyme was used as a pre-treatment of Meloidogyne incognita eggs they became more susceptible to P. chlamydosporia. The same treatment applied to G. rostochiensis eggs gave no response.
In the last decade some products have come to market that have nematicidal effects, such as DiTera, a compound produced from the fermentation extracts of a bacterium. Paecilomyces chlamydosporia, a fungal biocontrol agent, is also available on the market. Most other potential biocontrol agents are still being tested or studied to overcome problems with delivery systems or application methods. Many other mutualistic bacterial and fungal endophytes probably exist in the agroecosystem that would greatly improve plant health while at the same time be detrimental to plant parasitic nematodes. However, many technologies are involved in discovering the most appropriate candidates for commercialisation (Sikora et al., 2007). With time, appropriate study of plant parasites, their molecular properties and modes of parasitism will improve biological control options and identify new ways forward.
Fumigant nematicides are toxic and expensive, but have been used to help reduce large densities of potato cyst nematode. Soil fumigants can kill large numbers of nematodes, especially in moist sandy soils under polythene sheeting. Soil fumigants can be injected into the soil and this is also very effective, but there may be some risks of ground water contamination in certain circumstances. Many chemicals previously used for nematode control have now been banned, or are in the process of being phased out, due to safety concerns or their effect on the ozone layer.
The nematicide 1, 3-Dichloropropene is available mixed in various quantities with other compounds, such as C3-chlorinated hydrocarbons and 1, 2-dichloropropane; Telone II consists of 94% 1, 3-D whereas D-D contains around 50%. Other types of fumigants release methyl isothiocyanate (MITC), e.g. dazomet, which is effective against potato cyst nematode (Whitehead, 1975).
Fumigant nematicides are usually applied several weeks before planting to avoid phytotoxicity. Conditions affecting the efficacy of any of this type of chemical include soil type, moisture content, drainage and temperature.
For further information, see Whitehead (1998), Whitehead and Turner (1998) and Haydock et al. (2006).
These compounds are used in smaller amounts and usually do not persist in the soil as long as fumigant nematicides. Organophosphates and oximecarbamates are very effective nematicides. Their effect on nematodes is to paralyse rather than kill, unless very large doses are used. Organophoshates offer good control of G. rostochiensis, but need to be incorporated into the soil using rotary cultivation. These types of chemical are better suited to light, silty soils and are not as effective on organic soils. Isazophos (CGA 12223) was used by Moss et al. (1975) on silty loams and gave partial control of G. rostochiensis and of a mixed population of G. rostochiensis and G. pallida.
Organophosphates are cumulative poisons and, as such, are not used as much as they once were. The pH of the soil is an important factor if used: at more than pH 8.0 the soil is too alkaline. Higher temperatures also cause degradation of carbamates at faster rates than at lower temperatures.
Non-fumigant chemicals are most effective when used in granular form and at 15 cm. below the soil surface incorporated by machines that won't damage the soil structure.
Ellenby (1954) was the first to report a gene for resistance to G. rostochiensis. The gene was found in Solanum tuberosum ssp. andigena and is a single dominant gene, referred to as gene H1, but it conveys resistance only to certain populations of G. rostochiensis. Some European cultivars of potato have resistance (often only partial) to European pathotypes of potato cyst nematode but some South American populations of potato cyst nematode are known to be more virulent and are able to override resistance in the European cultivars (Kort and Jaspers, 1973; Turner et al., 1995).
Intensive studies in South America, at the International Potato Centre (CIP) in Lima, Peru, over the past 20 years, have considered 3000 accessions of potato and have focused on pathotypes in the local regions. Some resistant cultivars reduce potato cyst nematode field populations by up to 50%, but may select new, more virulent pathotypes. The use of any resistant potato cultivar repeatedly carries the risk of selecting a virulent nematode population, perhaps even a type not previously recognized, especially where unknown mixtures of species and/or pathotypes occur in the same location. It is wise to grow a different cultivar (resistant or tolerant) each year to avoid this problem. In the UK, frequent growing of the cultivar Maris Piper, resistant to UK G. rostochiensis potato cyst nematode populations but susceptible to G. pallida, has caused UK populations to switch to G. pallida, which is much more difficult to control.
In Asia both G. pallida and G. rostochiensis are present in India, Pakistan and Sri Lanka but growing areas are restricted; so far only the R1A and P5A pathotypes are known to be present (Zaheer, 1998). In South Africa, the potato is a major crop with some 5600 ha grown, most of which are sold for consumption. In Morocco, 5 eggs/ml soil caused appreciable damage (Spears, 1968).
There is also a need for tolerant cultivars, which suffer less damage and growth of which may also prevent selection of virulent pathotypes by resistant cultivars in a field. Strict quarantine measures are important supporting measures. There are no potato cultivars with full resistance to G. pallida (Marshall, 1998). However, in New Zealand the Institute for Crop and Food Research has released a series of cultivars with high resistance to G. rostochiensis and G. pallida, e.g. Karaka (Anderson et al., 1993) and Gladiator (Genet et al., 1995). For other resistant, tolerant and partially resistant cultivars, see Whitehead (1998).
ReferencesTop of page
Aihara T, Sumiya T, Suzuki K, 1998. Studies on the pathotype of the potato cyst nematode (Globodera rostochiensis) in Nagasaki prefecture. Research Bulletin of the Plant Protection Service, Japan, No. 34:71-79.
Allsopp PG, 1990. Sequential sampling plans for nematodes affecting sugar cane in Queensland, Australia. Journal of Agricultural Science, 41:351 58.
Anderson JAD, Lewthwaite S L, Genet RA, Broam F, Gallagher D, 1993. " Karka": A new fresh market potato with high resistance to Globodera pallida and Globodera rostochiensis, New Zealand Journal of Crop and Horticultural Science 21, 95-97.
Anon, 1976. Known distribution of potato cyst nematodes (Heterodera rostochiensis Woll. and H.pallida Stone) in Norway from a twenty years' survey. As, Norway: Statens Plantevern Zoologisk Avdeling, 24 pp.
Anon, 1988. ===. FAO Plant Protection Bulletin, 36. 42.
APPPC, 1987. Insect pests of economic significance affecting major crops of the countries in Asia and the Pacific region. Technical Document No. 135. Bangkok, Thailand: Regional Office for Asia and the Pacific region (RAPA).
Bakker J, Gommers FJ, 1986. Genotyping European potato cyst nematode populations with two dimensional gel electrophoresis of total protein extracts. Revue de Nématologie, 9:287.
Baldwin JG, Mundo-Ocampo M, 1991. Heteroderinae, cyst and non-cyst forming nematodes. In: Nickle WR, ed. Manual of Agricultural Nematology. New York, USA: Marcel Dekker, 275-362.
Bazan de Segura C, 1953. More about the golden nematode in Peru. Plant Disease Reporter, 37:326.
Been TH, Schomaker CH, 2006. Distribution patterns and sampling. In: Plant nematology [ed. by Perry, R. N.\Moens, M.]. Wallingford, UK: CABI, 302-326. http://www.cabi.org/CABeBooks/default.aspx?site=107&page=45&LoadModule=PDFHier&BookID=292
Behrens E, 1975. Taxonomically useful characters for the differentiation of Heterodera species. Probleme der Phytonematologie. Vortrage anlasslich der 10 Tagung uber Probleme der Phytonematologie im Institut fur Pflanzenzuchtung Gross-Lusewitz der Deutschen Akademie der Landwirtschaftswissenschaften zu Berlin am 11 Juni 1971., 122-142.
Bello A, Gonzalez JA, 1994. Potato cyst nematodes in the Canary Islands: an epidemiologic model for the Mediterranean region. Bulletin OEPP, 24:429-438.
Belokursakaya VI, 1973. ===. In: Materialy vsesoyuznogo simpoziuma po bor'be s kartofel'noi nematodoi, Tartu, 3-5 iyulya 1973. Tartu, Estonia: Institut Zoolgii I Botaniki Akademii Nauk Estonskoi SSR, 29-30.
Ben Saad A, Khalil JA, Farag I, Abugnia A, Saleh A, 1981. Diseases and Pests of Crop Plants of Libya., Libya: University Al-Fateh.
Bendezu IF, Evans K, Burrows PR, Pomerai Dde, Canto-Saenz M, 1998. Inter and intra-specific genomic variability of the potato cyst nematodes Globodera pallida and G. rostochiensis from Europe and South America using RAPD-PCR. Nematologica, 44(1):49-61.
Bendezu IF, Russell MD, Evans K, 1998. Virulence of populations of potato cyst nematodes (Globodera spp.) from Europe and Bolivia towards differential potato clones frequently used for pathotype classification. Nematologica, 44(6):667-681.
Bossis M, MugniTry D, 1993. Specific status of six Globodera parasites of solanaceous plants studied by means of two-dimensional gel electrophoresis with a comparison of gel patterns by a computed system. Fundamental and Applied Nematology, 16(1):47-56; 35 ref.
Brande J van den, Kips RH, D'Herde J, Mol L van, 1952. Onderzoek van aardappelvarieten en van Amerikanense Solanum-soorten in verband met het aardappelcysten aaltje Heterodera rostochiensis Wollenweber. Meded. Landbouwhogesch. OpzoekStns Gent, 17:51-60.
Brodie BB, 1984. Nematode parasites of potato. In: Nickle WR, ed. Plant and Insect Nematodes. New York, USA: Marcel Dekker, 167-212.
Brodie BBJ, 1998. Potato cyst nematodes (Globodera species) in Central and North America. In: Potato cyst nematodes: biology and distribution and control [ed. by Marks Brodie, R. J. ;. B. B. J.]. Wallingford, UK: CAB International, 317-331.
Bruchner E, 1961. [English title not available]. (Primer hallazgo de Heterodera rostochiensis Woll. Sobre papas silvestres (Solanum sect.Tuberarium) en la República Argentina y su significación como plaga de la especies cultivadas.) Universidad Nacional De Cuyo,Revista de la Facultad de Ciencias Agrarias, 8:7-18.
Buhr H, 1954. Untersuchungen uber den Kartoffelnematoden. I. Die (Papierstrifen- Methode), ein vereinfachtes Verfahren zur Utersuchung von Bodemproben auf ihren Besatz mit Nematodzysten. NachrBl, Dt. PflschDienst, Berl., 8:45-48.
Burrows PR, 1996. Modifying resistance to plant parasitic nematodes. In: Pierpoint WS, Shewry P, eds. Genetic Engineering of Crop Plants for Resistance to Pests and Diseases. London, UK: British Crop Protection Council, 38-65.
Burrows PR, de Waele D, 1997. Engineering resistance against plant parasitic nematodes using anti-nematode genes. In: Fenoll C, Grundler FMW, Ohl SA, eds. Cellular and Molecular Aspects of Plant-Nematode Interactions. Dordrecht, Netherlands: Kluwer Academic Publishers, 217-236.
CEC, 1969. Council Directive 69/465/EEC of 8 December 1969 on control of potato golden nematode. Official Journal of the European Communities No. L 323, 3-4.
Clarke AJ, Hennessy J, 1987. Hatching agents as stimulants of movement of Globodera rostochiensis juveniles. Revue de Nématologie, 10:471-476.
Cobb GS, Taylor AL, 1953. Heterodera leptonepia n. sp., a cyst forming nematode found in soil with stored potatoes. Proceedings of the Helminthological Society of Washington, 20:13-15.
Crump DH, 1987. Effect of time sampling, method of isolation and age of nematode on the species of fungi isolated from females of Heterodera schachtii and H. avenae.. Revue de Nématologie, 10(3):369-373; 17 ref.
Crump DH, 2004. Biocontrol- a route to market. In: Proceedings of the Fourth International congress of Nematology, 8-13 June 2002, Tenerife, Spain [ed. by Cook, R. \Hunt, D. J.]. Leiden, Netherlands: Brill, 165-174. [Nematology Monographs and Perspectives 2.]
Cubillos A, Fernández C, 1990. Breeding for resistance to bacteria, nematodes and other pathogens in Chile. (Mejoramiento genético para resistencia a bacterias, nematodos y otros patógenos en Chile.) In: Avances en el mejoramiento genético de la papa en los países del cono sur [ed. by Hidalgo, O. A.\Rincón R., H.]. Lima, Peru: International Potato Center, 181-186.
Dale MFB, 1988. Breeding for tolerance to potato cyst nematode. Aspects of Applied Biology, 17(1):95-101; [^italic~In Environmental aspects of applied biology, York, UK, 19-21 September 1988^roman~]; 10 ref.
DAO D, F, GONZÂLEZ JA, 1971. The golden nematode of potato, Heterodera rostochiensis Woll. and its presence in the Venezuelan Andes. (El nematodo dorado de la papa, Heterodera rostochiensis Woll. y su presencia en los Andes Venezolanos.) Agronomia Tropical, 21(2):105-110.
Davide RG, Zorilla RA, 1995. Evaluation of three nematophagous fungi and some nematicides for the control of potato cyst nematode Globodera rostochiensis. (Evaluacion de tres hongos nematofagos yalgunos nematicidas para el control de nematodo quiste de la papa Globodera rostochiensis.) BIOCONTROL, 1(3):45-55.
Davies KG, 1998. Natural parasites and biological control. In: Sharma SB, ed. The cyst nematodes. London, UK: Chapman and Hall.
Davies KG, Laird V, Kerry BR, 1991. The motility, development and infection of Meloidogyne incognita encumbered with spores of the obligate hyperparasite Pasteuria penetrans. Revue de Ne^acute~matologie, 14(4):611-618; 24 ref.
Desgarennes D, Sánchez-Nava P, Peña-Santiago R, Carrión G, 2009. Nematode fauna associated with the rhizosphere of potato crop (Solanum tuberosum) grown in the region of Cofre de Perote, Veracruz, Mexico. (Nematofauna asociada a la rizosfera de papas (Solanum tuberosum) cultivadas en la zona productora del Cofre de Perote, Veracruz, México.) Revista Mexicana de Biodiversidad, 80(3):611-614. http://www.ibiologia.unam.mx
Doncaster CC, Seymour M, 1973. Exploration and selection of penetration site by Tylenchida. Nematologica, 19:137-145.
Doncaster CC, Shepherd AM, 1967. The behaviour of second-stage Heterodera rostochiensis larvae leading to their emergence from the egg. Nematologica, 13:476-478.
Eglitis V, 1973. ===. In: Materialy vsesoyuznogo simpoziuma po bor'be s Kartofel'noi nematode, Tartu, 3-5 iyulya 1973. Tartu, Estonia: Institut Zoolgii I Botaniki Akademii Nauk Estonskoi SSR, 26-28.
Ekanayake HMRK, 1991. Identification of Globodera rostochiensis pathotype in Sri Lanka. Quarterly newsletter, Asia and Pacific Plant Protection Commission, April-June. 17 pp.
Elekes-Kaminszky M, Feketé-Palkovics Â, Avar K, Baranyai-Tóth R, Bártfai J, Budai C, Cziklin M, Farkas I, Gál T, Gyo´´rffy-Molnár J, Gyulai P, Havasréti B, Herczig B, Hoffmann É, Jobbágy J, Kleineizel S, Komlósi É, Kováts Z, Lukács M, Mero´´ F, Simon A, Szendrey L, Szeo´´ke K, Tóth AC, Tóth B, To´´kés-Papp E(et al), 2004. Second nation-wide survey of the potato cyst nematodes (Globodera rostochiensis and G. pallida) between 1999-2002. (A burgonya cisztaképzo´´ fonálférgei (Globodera rostochiensis [Woll.] és G. pallida stone) elterjedésének: II. Országos felmérése (1999-2002).) Növényvédelem, 40(9):463-469.
Ellenby C, 1945. Susceptibility of South American tuber-forming species of Solanum to the potato-root eelworm Heterodera rostochiensis Wollenweber. Epn. J. exp. Agric., 13:158-168.
Ellenby C, 1954. Tuber forming species and varieties of the genus Solanum tested for resistance to the potato cyst eelworm Heterodera rostochiensis Wollenweber. Euphytica, 3:195-202.
Endo BY, 1998. Atlas on Ultrastructure of Infective Juveniles of the Soybean Cyst Nematode, Heterodera glycines. Agriculture Handbook No. 711. Washington, USA: USDA.
Enneli S, Öztürk G, 1995. The important plant parasitic nematodes harmful on potatoes in central Anatolia. (Orta anadolu bölgesinde patateslerde zarar yapan önemli bitki paraziti nematodlar.) Zirai Mücadele Arastirma Yilligi, No. 30:35-36.
EPPO, 1990. Specific quarantine requirements. EPPO Technical Documents, No. 1008. Paris, France: European and Mediterranean Plant Protection Organization.
EPPO, 2007. Basic requirements for quality management in plant pest diagnosis in laboratories. European and Mediterranean Plant Protection Organization Bulletin, 37:580-588.
EPPO, 2007. EPPO Standards, Diagnostics. European and Mediterranean Plant Protection Organization Bulletin, 37:499-502.
EPPO, 2011. EPPO Reporting Service. EPPO Reporting Service. Paris, France: EPPO. http://archives.eppo.org/EPPOReporting/Reporting_Archives.htm
EPPO, 2012. EPPO Reporting Service. EPPO Reporting Service. Paris, France: EPPO. http://archives.eppo.org/EPPOReporting/Reporting_Archives.htm
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
Evans K, 1968. The Influence of some factors on the Reproduction of Heterodera rostochiensis. Ph.D. Thesis. London, UK: London University.
Evans K, 1970. Longevity of males and fertilization of females of Heterodera rostochiensis. Nematologica, 16:369-374.
Forrest JMS, Robertson WM, Milne EW, 1989. Observations on the cuticle surface of second stage juveniles of Globodera rostochiensis and Meloidogne incognita. Revue de Ne^acute~matologie, 12(4):337-341; 23 ref.
Fox PC, Atkinson HJ, 1985. Enzyme variation in pathotypes of the potato cyst nematodes Globodera rostochiensis and G. pallida. Parasitology, 91:499-506.
Fox PC, Atkinson HJ, 1985. Immunochemical studies on pathotypes of the potato cyst nematodes Globodera rostochiensis and G. pallida. Parasitology, 90:417-483.
Franco J, Oros R, Main G, Ortun~o N, 1998. Potato cyst nematodes (Globodera species) in South America. In: Marks RJ, Brodie BB, eds. Potato cyst nematodes biology, distribution and control. Wallingford, UK: CAB International, 239-270.
Franklin MT, 1940. On the identification of strains of Heterodera schachtii. Journal of Helminthology, 18:63-84.
FRÉZAL P, 1954. [English title not available]. (Importance et repercussions dc la contamination de l'Algérie par le nématode doré (Heterodera rostochiensis Wooll. [Woll.].) Comptes Rendus des Seances de l'Académie d'Agriculture de France, 40(2):71-74.
Fullaondo A, Barrena E, Viribay M, Barrena I, Salazar A, Ritter E, 1999. Identification of potato cyst nematode species Globodera rostochiensis and G. pallida by PCR using specific primer combinations. Nematology, 1(2):157-163.
Genet RA, Braam WF, Gallagher DTP, Anderson JAD, Lewthwaite SL, 1995. 'Gladiator': a new potato cultivar with high resistance to potato cyst nematode and powdery scab suitable for french fries and fresh market. New Zealand Journal of Crop and Horticultural Science, 23(1):105-107; 5 ref.
Gheysen G, Jones JT, 2006. Molecular Aspects of Plant-Nematode Interactions. In: Plant Nematology [ed. by Perry, R. N. \Moens, M.]. Wallingford, UK: CABI, 234-152.
Gheysen G, Jones JT, 2006. Molecular aspects of plant-nematode interactions. In: Plant nematology [ed. by Perry, R. N.\Moens, M.]. Wallingford, UK: CABI, 234-254. http://www.cabi.org/CABeBooks/default.aspx?site=107&page=45&LoadModule=PDFHier&BookID=292
Gladkaya RM, Korzhentsevskaya NV, 1985. The potato nematode. Zashchita Rastenii, Moscow, No.8:39.
Golden AM, Ellington DMS, 1972. Re-description of Heterodera rostochiensis (Nematoda: Heteroderidae), with a key and notes on closely related species. Proceedings of the Helminthological Society of Washington, 39:64-78.
Golinowski W, Sobczak M, Kurek W, Grymaszewska, 1997. The structure of Syncytia. In: Fenoll C, Grundler FMW, Ohl SA, eds. Cellular and Molecular Aspects of Plant Nematode Interactions. Dordhecht, Netherlands: Kluwer Academic Publishers, 80-97.
Granek I, 1955. Additional morphological differences between the cysts of Heterodera rostochiensis and Heterodera tabacum. Plant Disease Reporter, 39:716 -718.
Greco N, Moreno I, 1992. Influence of Globodera rostochiensis on yield of summer, winter and spring potato in Chile. Nematropica, 22:165-173.
Green CD, 1971. The morphology of the terminal area of the round-cyst nematodes S.G. Heterodera rostochiensis and allied species. Nematologica, 17:34-46.
Green CD, Miller LI, 1969. Cyst nematodes: attraction of males to females. Report of Rothamsted Experimental Station for 1968. Part 1. Rothamsted, UK: Rothamsted Experimental Station, 153-158.
Green CD, Plumb SC, 1970. The interrelationships of some Heterodera species indicated by the specificity of the male attractants emitted by their females. Nematologica, 16:39-46.
Grubisic D, Ostrec L, Culjak TG, Blümel S, 2007. The occurrence and distribution of potato cyst nematodes in Croatia. Journal of Pest Science, 80(1):21-27. http://www.springerlink.com/content/u0884x7r10821475/?p=79709419b777459784f355b7b2fe8bbc&pi=3
Grubisic D, Zivkovic IP, Culjak TG, Brmez M, Benkovic-Lacic T, Mesic A, 2013. First molecular detection of Croatian potato cyst nematode (PCN) populations using the polymerase chain reaction (PCR). Entomologia Croatica, 17(1/4):35-40. http://hrcak.srce.hr/entomologia-croatica
Gus'kova LA, Makovskaya SA, 1977. A study of the composition of the pathotypes of the cyst nematode in the north-west of the Nechernozem zone of the RSFSR. Svobodnozhivushchie, pochvennye, entomopatogennye i fitonematody. (Sbornik nauchnykh rabot). Leningrad, USSR: Akademiya Nauk SSSR, Zoologicheskii Institut., 57-60.
Halford PH, Russell MR, Evans K, 1999. Trap cropping for cyst nematode management. Annals of Applied Biology, 134:321-327.
Hansen LM, 1988. Use of stylet measurements to distinguish between two species of potato nematode Globodera rostochiensis and G. pallida. (Avendelse af stiletmalinger til at skelne mellem kartoffelnematodens to arter Globodera rostochiensis og G. pallida.) In: Växtskyddsrapporter, Jordbruk, No. 53. S-750 77 Uppsala, Sweden: Sveriges Lantbruksuniversitet, 29-31.
Haydock PPJ, Woods SR, Grove IG, Hare MC, 2006. Chemical control of nematodes. In: Plant nematology [ed. by Perry, R. N.\Moens, M.]. Wallingford, UK: CABI, 392-410. http://www.cabi.org/CABeBooks/default.aspx?site=107&page=45&LoadModule=PDFHier&BookID=292
Heikkilä J, Tilikkala K, 1992. Globodera rostochiensis (Woll.) Behrens (Tylenchida, Heteroderidae), the only potato cyst nematode species found in Finland. Agricultural Science in Finland, 1(5):519-524.
Hinch JM, Alberdi F, Smith SC, Woodward JR, Evans K, 1998. Discrimination of European and Australian Globodera rostochiensis and G. pallida pathotypes by high performance capillary electrophoresis. Fundamental and Applied Nematology, 21(2):123-128; 14 ref.
Holz RA, Wright DJ, Perry RN, 1998. Changes in the lipid content and fatty acid composition of 2nd-stage juveniles of Globodera rostochiensis after rehydration, exposure to the hatching stimulus and hatch. Parasitology, 116(2):183-190; 45 ref.
Hooper DJ, 1990. Extraction and processing of plant and soil nematodes. In: Luc M, Sikora RA, Bridge J, eds. Plant Parasitic Nematodes in Subtropical and Tropical Agriculture. Wallingford, UK: CAB International, 45-68.
Huijsman CA, 1959. Nature and inheritance of the resistance to the potato-eelworm Heterodera rostochiensis W. in Solanum kurtzianum. Meded. LandbHoogesch. OpzoekStns. Gent, 24:611-613.
Inagarki H, 2004. Distribution means of the potato cyst nematode to Japan, its ecology and control. Japanese Journal of Tropical Agriculture, 48(5):324-329.
Indarti S, Bambang RTP, Triman M, Triman B, 2004. First record of potato cyst nematode Globodera rostochiensis in Indonesia. Australasian Plant Pathology, 33:325-326.
IPPC, 2009. Golden Nematode (Globodera rostochiensis) - Update on the Canadian Situation (2009). IPPC Official Pest Report, CAN-03/2. Rome, Italy: FAO. https://www.ippc.int/index.php?id=1110520&no_cache=1&type=pestreport&L=0
IPPC, 2013. Globodera rostochiensis subject to official control in Denmark. IPPC Official Pest Report, No. DNK-13/1. Rome, Italy: FAO. https://www.ippc.int/
IPPC, 2015. Eradication of Potato Cyst Nematode (PCN) from Western Australia. IPPC Official Pest Report, No. AUS-39/2. Rome, Italy: FAO. https://www.ippc.int/
Iverson LGK, 1972. Golden nematode-infestation found in Mexico. Plant Disease Reporter, 49:281.
Jogaite V, Cepulyte R, Stanelis A, Bu¯da V, 2007. Monitoring of Globodera spp. in Lithuania using diagnostic morphometric analysis and polymerase chain reaction. Acta Zoologica Lituanica, 17(2):184-186. http://www.ekoi.lt
Jones FGW, Parrot DM, 1969. Population fluctuations of Heterodera rostochiensis Woll. when susceptible potato varieties are grown continuously. Annals of Applied Biology, 63:175-181.
Jones FGW, Pawelska K, 1963. The behaviour of populations of potato-root eelworm (Heterodera rostochiensis Woll.) towards some resistant tuberous and other Solanum species. Annals of Applied Biology, 51:277-294.
Jones JT, Harrower BE, 1998. A comparison of the efficiency of differential display and cDNA-AFLPs as tools for the isolation of differentially expressed parasite genes. Fundamental and Applied Nematology, 21(1):81-88; 14 ref.
Jones JT, Perry RN, Johnston MRL, 1993. Changes in the ultrastructure of the cuticle of the potato cyst nematode, Globodera rostochiensis, during development and infection. Fundamental and Applied Nematology, 16(5):433-445; 23 ref.
Jones JT, Robertson L, Perry RN, Robertson WM, 1997. Changes in gene expression during stimulation and hatching of the potato cyst nematode Globodera rostochiensis. Parasitology, 114(3):309-315; 13 ref.
Jones JT, Robertson WM, 1997. Nematode secretions In: Fenoll C, Grundler FMW, Ohl SA, eds. Cellular and Molecular Aspects of Plant Nematode Interactions. Dordrecht, Netherlands: Kluwer Academic Publishers, 98-106.
Kemner NA, 1929. Potatisnematoden eller potatosalen (Heterodera schachtii subsp. rostochiensis Woll.) och dess framtradande i Sverige. Medd. CentAnst. Forsoksv. Jordbr., Stockh., No.355 (Lantbruksent. Avd., No. 56).
Kleynhans KPN, 1998. Potato cyst nematodes (Globodera species) in Africa. In: Potato cyst nematodes, biology, distribution and control [ed. by Marks, R. J.\Brodie, B. B.]. Wallingford, UK: CAB INTERNATIONAL, 347-351.
Koppel M, Tsahkna A, 1998. Potato cyst nematode (Globodera rostochiensis) resistance breeding in Estonia. In: Beiträge zur Züchtungsforschung - Bundesanstalt für Züchtungsforschung an Kulturpflanzen, 4(2) [ed. by Peter, K.]. 92-93.
Krall E, Krall H, 1978. Reconstruction of phylogeny and systematics of nematodes of the Heteroderidae on the basis of host specialisation and co-evolution with their hosts. Fitogel'mintol. Issledovania Moscow, SSR: " Nauk" Acad. Sci. U.S.S.R. Acad. Sciences, 39-56.
Krnnjac DJ, Oro V, Gladovic S, Trkulja N, Scekic D, Kecovic V, 2002. Zastita bilja ( Serbia and Montenegro). Plant Protection, 53(4):147-156.
Kudla U, Milac AL, Qin Ling, Overmars H, Roze E, Holterman M, Petrescu AJ, Goverse A, Bakker J, Helder J, Smant G, 2007. Structural and functional characterization of a novel, host penetration-related pectate lyase from the potato cyst nematode Globodera rostochiensis. Molecular Plant Pathology, 8(3):293-305. http://www.blackwell-synergy.com/doi/pdf/10.1111/j.1364-3703.2007.00394.x
Kühn, 1881. Die Ergebnisse der Versuche zur Ermittelung der Ursache der Rübenmüdigkeit und zur Erforschung der Natur der Nematoden. Ber. physiol. Lab. landw. Inst. Univ. Halle, 3:1-153.
Lilley CJ, Bakhetia M, Charlton WL, Urwin PE, 2007. Recent progress in the development of RNA interference for plant parasitic nematodes. Molecular Plant Pathology, 8(5):701-711. http://www.blackwell-synergy.com/loi/mpp
Linford MB, Yap F, Olivera JM, 1938. Reduction of soil populations of the root knot nematode during decomposition of organic matter. Soil Science, 45:127-141.
Lombardo S, Colombo A, Rapisarda C, 2011. Cyst nematodes of the genus Heterodera and Globodera in Sicily. Redia [X Congress of the Italian Society of Nematology (SIN), Naples, Italy, 28-30 October 2010.], 94:137-141. http://email@example.com
Lownsbery BF, Lownsbery JW, 1954. Heterodera tabacum new species a parasite of solanaceous plants in Connecticut. Proceedings of the Helminthological Society of Washington, 21:42-47.
MAFF, 2000. Potato Cyst nematode: a management guide. London, UK: MAFF.
Magnusson ML, 1987. Effects of some non-host crops on population density of Globodera rostochiensis. VSxtskyddsnotiser, 12:17-25.
Mahran A, Turner S, Martin T, Yu Q, Miller S, Sun F, 2010. The golden potato cyst nematode Globodera rostochiensis pathotype Ro1 in the Saint-Amable Regulated Area in Quebec, Canada. Plant Disease, 94(12):1510. http://apsjournals.apsnet.org/loi/pdis
Mai WF, 1951. Solanum xanti Gray and Solanum integrifolium Poir., new hosts of he golden nematode, Heterodera rostochiensis Wollenweber. American Potato Journal, 28:578-579.
Mai WF, 1952. Susceptibilityof Lycopersicon species to the golden nematode. Phytopathology, 42:461.
Mai WF, Spears JF, 1954. The golden nematode in the United States. American Potato Journal, 31:387-394.
Male-Kayiwa BS, 1983. Screening wild potatoes for resistance to potato cyst nematode, Globodera pallida (Stone) Mulvey and Stone. MSC. Thesis, Department of Plant Biology University of Birmingham, September 1983, 9-14.
Manduric S, Andersson S, 2003. Potato cyst nematodes (Globodera rostochiensis and G. pallida) from Swedish potato cultivation - an AFLP study of their genetic diversity and relationships to other European populations. Nematology, 5(6):851-858.
Marks RJ, Rojancovski E, 1998. Potato cyst nematodes (Globodera species) in central Europe, the Balkans and the Baltic states. In: Potato cyst nematodes, biology, distribution and control [ed. by Marks, R. J.\Brodie, B. B.]. Wallingford, UK: CAB INTERNATIONAL, 299-315.
Marshall JW, 1998. Potato cyst nematodes (Globodera) species New Zealand and Australia. In: Marks RJ, Brodie BB. Potato cyst nematodes biology, distribution and control. Wallingford, UK: CAB International, 359-394.
Marshall JW, Crawford AM, 1987. A cloned DNA fragment that can be used as a sensitive probe to distinguish Globodera pallida from G. rostochiensis and other common cyst forming nematodes (abstract). Journal of Nematology, 19:541.
Martinez-Beringola ML, Franco L, Paz LM, Gutierrez MP, 1988. Cyst forming nematodes of potato in Spain. (Los nematodos formadores de quistes de la patata en España.) Boletin de Sanidad Vegetal, Plagas, Spain, 14(3):405-414.
Miller LI, 1986. Economic importance of cyst nematodes in North America. In: Lamberti F, Taylor CE, eds. Cyst Nematodes. New York, USA: Plenum Press, 373-385.
Miller LI, Gray BJ, 1968. Horsenettle cyst nematode, Heterodera virginiae n. sp., a parasite of solanaceous plants. Nematologica, 14:535-543.
Miller LI, Gray BJ, 1972. Heterodera solanacearum n. sp., a parasite of solanaceous plants. Nematologica, 18:404-413.
Moreno I, Vovlas N, Lamberti F, 1984. Species of potato cyst nematodes from Chile. Nematologia Mediterranea, 12:247-252.
Morgan DO, 1925. Investigations on eelworm in potatoes in South Lincolnshire. Journal of Helminthology, 3:185-192.
Moss SR, Crump D, Whitehead AG, 1975. Control of potato cyst-nematodes, Heterodera rostochiensis and H. pallida, in sandy, peaty and silt loam soils by oximecarbamate and organophosphate nematicides. Annals of Applied Biology, 81(3):359-365
MugniTry D, Bossis M, Pierre JS, 1992. Hybridization between Globodera rostochiensis (Wollenweber), G. pallida (Stone), G. virginiae (Miller & Gray), G. solanacearum (Miller & Gray) and Globodera "mexicana" (Campos-Vela). Description and future of the hybrids. Fundamental and Applied Nematology, 15(4):375-382; 19 ref.
Mulholland V, Carde L, O'Donnell KJ, Fleming CC, Powers TO, 1996. Use of the polymerase chain reaction to discriminate potato cyst nematode at the species level. In: Diagnostics in Crop Production, BCPC Symp. Proceedings, 65. 247-252.
Mulvey RH, 1973. Morphology of the terminal areas of white females and cysts of the genus Heterodera (e.g. Globodera) Journal of Nematology, 5:303-311.
Mulvey RH, Golden AM, 1983. An illustrated key to the cyst-forming genera and species of Heteroderidae in the Western Hemisphere with species morphometrics and distribution. Journal of Nematology, 15(1):1-59.
Mulvey RH, Stone AR, 1976. Description of Punctodera matadorensis n.gen., n.sp. (Nematoda: Heteroderidae) from Saskatchewan with lists of species and generic diagnoses of Globodera (n. rank), Heterodera, and Sarisodera. Canadian Journal of Zoology, 54(5):772-785
OEPP/EPPO, 1991. Quarantine procedure No. 30, Globodera pallida, G. rostochiensis, Soil sampling methods. Bulletin OEPP/EPPO Bulletin, 21:233-240.
OEPP/EPPO, 2004. Diagnostic protocols for regulated pests Globodera rostochiensis and Globodera pallida. European and Mediterranean Plant Protection Organization Bulletin, 34(2):309-314.
Oerke EC, Dehne HW, Schönbeck F, Weber A, 1994. Crop production and crop protection: estimated losses in major food and cash crops. Crop production and crop protection: estimated losses in major food and cash crops., xxii + 808 pp.; [ref. at ends of chapters, available from publishers at PO Box 1991, Amsterdam, Netherlands or PO Box 945, Madison Square Station, NY 10160-0757, USA].
Olsen OA, Mulvey RH, 1962. The discovery of the golden cyst nematode in Newfoundland. Canadian Plant Disease Survey, 42(4).
Oostenbrink M, 1950. Het Aardappelaaltje (Heterodera rostochiensis Wollenweber), een gevaarlijke parasiet voor de eenzijdige aardappel-cultuur. Verslagen en Mededelingen von den Plantenziekten-Kundige Dienst te Wageningen: H. Veenman and Zonen, 115:230pp.
Orchard WR, 1965. Occurrence of the golden nematode in Newfoundland. Canadian Plant disease Survey, 42(4).
Ostojic I, Grubisic D, Zovko M, Milicevic T, Culjak TG, 2011. First report of the golden potato cyst nematode, Globodera rostochiensis, in Bosnia and Herzegovina. Plant Disease, 95(7):883. http://apsjournals.apsnet.org/loi/pdis
Othman AA, Baldwin JG, Mundo-Ocampo M, 1988. Comparative morphology of Globodera, Cactodera, and Punctodera spp. (Heteroderidae) with scanning electron microscopy. Revue de Nematologie, 11:53-63.
Parker B, 2000. Experts seek out reliable advice on PCN. Potato Review: 36-38.
Perry RN, 1986. Physiology of hatching. In: Lamberti F, Taylor CE, eds. Cyst Nematodes. New York, USA: Plenum Press, 119-131.
Perry RN, Beane J, 1988. Effects of activated charcoal on hatching and infectivity of Globodera rostochiensis in pot tests. Revue de Nématologie, 11:229-233.
Perry RN, Feil JJ, 1986. Observations on a novel hatching bioassay for Globodera rostochiensis using fluorescence microscopy. Revue de Nématologie, 9:280-282.
Picard D, Sempere T, Plantard O, 2007. A northward colonisation of the Andes by the potato cyst nematode during geological times suggests multiple host-shifts from wild to cultivated potatoes. Molecular Phylogenetics and Evolution, 42(2):308-316. http://www.sciencedirect.com/science/journal/10557903
Pinochet J, 1987. Management of plant parasitic nematodes in central America: The Panama experience. In: Veetch JA, Dickson DW, eds. Vistas on Nematology. Hyattsville, Maryland, USA: Society of Nematologists, 105-113.
Potocek J, Gaar V, Hnízdil M, Novák F, 1991, publ. 1992. Protection against the spread of potato wart disease and potato root nematode. (Ochrana proti sirení rakoviny brambor a hádhacek~átka bramborového.) Metodiky pro Zavádení Výsledku Výzkumu do Zemedelské Praxe, No.18. 88 pp.
Pylypenko LA, Uehara T, Phillips MS, Sigareva DD, Blok VC, 2005. Identification of Globodera rostochiensis and G. pallida in the Ukraine by PCR. European Journal of Plant Pathology, 111(1):39-46. http://springerlink.metapress.com/link.asp?id=100265
Rice SL, Leadbeater BSA, Stone AR, 1986. Changes in roots of resistant potatoes parasitized by potato cyst-nematodes. I. Potatoes with resistance gene H1 derived from Solanum tuberosum ssp. andigena. Physiology and Plant Pathology, 27:219-234.
Riel HRvan, Mulder A, 1998. Potato cyst nematodes (Globodera species) in western Europe. In: Potato cyst nematodes, biology, distribution and control [ed. by Marks, R. J.\Brodie, B. B.]. Wallingford, UK: CAB INTERNATIONAL, 271-298.
Riga E, Holdsworth DR, Perry RN, Barrett J, Johnston MRL, 1997. Electrophysiological analysis of the response of males of the potato cyst nematode, Globodera rostochiensis, to fractions of their homospecific sex pheromone. Parasitology, 115(3):311-316; 10 ref.
Robinson MP, Atkinson HJ, Perry RN, 1988. The association and partial characterization of a fluorescent hypersensitive response of potato roots to the potato cyst nematode Globodera rostochiensis and G. pallida. Revue de Nématologie 11: 99-108.
Roessner J, 1986. Parasitism of Globodera rostochiensis by nematophagous fungi. Revue de Nématologie, 9:307-308.
Rojancovski E, Dehebanu A, 1986. Potato cyst nematode Globodera rostochiensis, a new pest detected in our country. Bulletinul de Protectia Plantelor, 2:43-50.
Rothacker D, 1957. Beitrage zur Resistenzzuchtung gegen den Kartoffelnematoden. 1. Prufung von Primitiv-und Wildkartoffeln auf dos Verhalten gegenüber dem Kartoffelnematoden. Zuchter, 27:124-132.
Santos MSNAde, Evans K, Abreu CA, Martins FF, Abrantes IMOde, 1995. A review of potato cyst nematodes in Portugal. Nematologia Medditerreanea, 23:35-42.
Santos MSNAde, Fernandes MFM, 1988. ===. Nematologia Medditerranea, 16:145.
Scholte K, 2000. Screening of non-tuber bearing Solanceae for resistance to and induction of juvenile hatch of potato cyst nematodes and their potential for trap cropping. Annals of Applied Biology, 136:239-246.
Segers R, Butt TM, Kerry BR, Beckett A, Pedberdy JF, 1996. The role of the proteinase VCP1 produced by the nematophagous Verticillium chlamydosporium in the infection process of nematode eggs. Mycological Research, 100(4):421-428; 27 ref.
Seinhorst JW, 1986. Effects of nematode attack on growth and yield of crop plants. In: Lamberti F, Taylor CE, eds. Cyst Nematodes. New York, USA: Plenum Press, 191-209.
Sembdner G, 1959. Uber das Eindringen der Kartoffelnematoden- Larven und ihre Weiterentwicklung in pflanzlichen Geweben. TagBer. Dt. Akad, LandwWiss. Berl. No. 20, 53-55.
Sembdner G, 1963. Anatomie der Heterodera-rostochiensis-gallen an Tomatenwurzeln. Nematologica, 9:55-64.
Sembdner G, Buhr H, Osske G, Schreiber K, 1960. Untersuchgen uber Wirt- Parasit-Beziehungen beim Kartoffelnematoden, Heterodera rostochiensis Woll. Wiss. PflSchKonf, Bpest, 339-342.
Shepherd AM, Clarke AJ, 1971. Moulting and hatching stimuli. In: Zuckerman BM, Mai WF, Rhode RA, eds. Plant Parasitic Nematodes, Volume II. New York, USA and London, UK: Academic Press, 267-287.
Sikora RA, Schäfer K, Dababat AA, 2007. Modes of action associated with microbially induced in planta suppression of plant-parasitic nematodes. Australasian Plant Pathology, 36(2):124-134. http://www.publish.csiro.au/nid/39.htm
Sirca S, Urek G, 2004. Morphometrical and ribosomal DNA sequence analysis of Globodera rostochiensis and Globodera achilleae from Slovenia. Russian Journal of Nematology, 12(2):161-168. http://www.russjnematology.com
Sirca S, Urek G, 2005. Results of the study of the yellow potato cyst nematode G. rostochiensis Woll. (Behrens) in Slovenia. (Rezultati preucevanja rumene krompirjeve ogorcice G. rostochiensis Woll. (Behrens) v Sloveniji.) In: Zbornik predavanj in referatov. 7. Slovensko posvetovanje o varstvu rastlin, 8.-10 marec, 2005, Zrece, Slovenija [ed. by Vajs, S.\Lesnik, M.]. Ljubljana, Slovenia: Drustvo za Varstvo Rastlin Slovenije, 349-352.
Skarbilovich TS, 1959. On the structure of the nematodes of the order Tylenchida Thorne, 1949. Acta Parasitologica Polonica, 15:117-132.
Spears JF, 1968. The golden nematode handbook, survey, laboratory, control and quarantine procedures. U.S. Department of Agriculture Handbook, No. 353.
Spears M, 1968. The golden nematode handbook, survey, laboratory, control and quarantine procedures. U.S. Department of Agriculture Handbook, No. 353.
Stanton JM, 1986. First record of potato cyst nematode Globodera rostochiensis in Australia. Australasian Plant Pathology, 15:87.
Stare BG, Sirca S, Urek G, 2013. Are we ready for the new nematode species of the Globodera genus? In: Zbornik Predavanj in Referatov, 11. Slovenskega Posvetovanja o Varstvu Rastlin Z Mednarodno Udelezbo (in okrogle mize o zmanjsanju tveganja zaradi rabe FFS v okviru projekta CropSustaIn), Bled, Slovenia, 5.-6. Marec 2013 [ed. by Trdan, S.\Macek, J.]. Ljubljana, Slovenia: Plant Protection Society of Slovenia, 144-150.
Stelter H, 1957. Untersuchungen über den Kartoffelnematoden Heterodera rostochiensis Wollenweber. III. Neue Wirtspflanzen des Kartoffelnematoden. Parasitica, 13:87-93.
Stelter H, 1959. Einige Beobachtungen an nicht-Knollentragenden Solanaceen in bezug auf den Kartoffelnematoden (Heterodera rostochiensis Wr). Nachr. Bl. Dt. PflschutzDienst, Berl., 13(7):135.
Stelter H, 1987. The host suitability of Solanum spp. for the three Globodera spp. Nematologica, 33:310-315.
Stone AR, 1973. Heterodera pallida n.sp. (Nematoda: Heteroderidae), a second species of potato cyst nematode. Nematologica, 18:591-606.
Subbotin SA, Halford PD, Perry RN, 1999. Identification of populations of potato cyst nematodes from Russia using protein electrophoresis, rDNA-RFLPs and RAPDs. Russian Journal of Nematology, 7(1):57-63.
Subbotin, S. A., Mundo-Ocampo, M., Baldwin, J. G., 2010. Systematics of cyst nematodes (Nematoda: Heteroderinae), Volume 8, Part A., In: Systematics of cyst nematodes (Nematoda: Heteroderinae), Volume 8, Part A. Brill Academic Publishers. xii + 352 pp..
Sun F, Miller S, Wood S, Côté MJ, 2007. Occurrence of potato cyst nematode, Globodera rostochiensis, on potato in the Saint-Amable region, Quebec, Canada. Plant Disease, 91(7):908. HTTP://www.apsnet.org
Taylor LR, 1961. Aggregation, variance and the mean. Nature (London) 189:732-735.
Thiery M, Mugniery D, 1996. Interspecific rDNA restriction fragment length polymorphism in Globodera species, Parasites of solanceous plants. Fundamental and Applied Nematology, 19:417-479.
Tiago JS, 1956. [English title not available]. (Un grave inimigo da cultura de batata.) Gazetta Agric Mozambique, 8:334.
Trudgill DL, 1967. The effect of environment on sex determination in Heterodera rostochiensis. Nematologica, 13:263-272.
Trudgill DL, Evans K, Phillips MS, 1998. Potato cyst nematodes: damage mechanisms and tolerance in the potato. In: Marks RJ, Brodie BB, eds. Potato cyst nematodes biology, distribution and control. Wallingford, UK: CAB International, 7-26.
Turner SJ, Evans K, 1998. The origins, global distribution and biology of potato cyst nematodes (Globodera rostochiensis (Woll.) and Globodera pallida Stone). In: Potato cyst nematodes, biology, distribution and control [ed. by Marks, R. J.\Brodie, B. B.]. Wallingford, UK: CAB INTERNATIONAL, 7-26.
Turner SJ, Fleming CC, Marks RJ, Watts PJ, 1994. A strategy for maintaining effective host-plant resistance to potato cyst nematodes. Proceedings - Brighton Crop Protection Conference, Pests and Diseases, 1994, vol. 2, No. 2:905-910; 10 ref.
Turner SJ, Martin TJG, McAleavey PBW, Fleming CC, 2006. The management of potato cyst nematodes using resistant Solanaceae potato clones as trap crops. Annals of Applied Biology, 149(3):271-280. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1744-7348.2006.00089.x
Tzortzakakis EA, Conceição ILPMda, Abrantes IMde O, Santos MSNde A, 2004. Characterisation and identification of potato cyst nematode populations from Crete, Greece, by isoelectric focusing of proteins. Nematology, 6(1):153-154. http://www.brill.nl
Whitehead AG, Turner SJ, 1998. Management and regulatory control strategies for potato cyst nematodes ( Globodera rostochiensis and Globodera pallida) In: Marks RJ, Brodie BB, eds. Potato cyst nematodes biology, distribution and control. CAB International, Wallingford, UK: CAB International.
Wille JE, Segura CB de, 1952. La angulilula dorado, Heterodera rostochiensis. Una Plaga del cultivo de las papas, recien discubierta en el Peru. Lima, Peru: Boln Cent. Nac. Invest. Exp. Agric.
Winslow RD, 1955. Provisional lists of host plants of some root eelworms (Heterodera species). Annals of Applied Biology, 41:591-605.
Wollenweber HW, 1923. Krankheiten und BeschSdigungen der Kartoffel. Arb. Forsch. Inst. Kartof. Berlin, Heft 7, 1-56.
Wyss U, Zunke U, 1986. Observations on the behaviour of second-stage juveniles of Heterodera schachtii inside host roots. Revue de Nématologie, 9:153-165.
Yameda E, Takakura S, Tezuka H, 1972. On the occurrence of potato cyst nematodes, Heterodera rostochiensis, Japan. Japanese Journal of Nematology, 2:12-15.
Zimmerman H, 1927. Versuche über die Kartoffelnematode (Heterodera schachtii, forma solani). Arb. ForschInst. Kartoff., Berl., No. 8, 151-154.
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18/03/2008 Updated by:
Janet Rowe, IACR-Rothamsted, Rothamsted Experimental Station, Harpenden, Hertfordshire, AL5 2JQ, UK
Distribution MapsTop of page
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