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• W4252051645 abstract "EPPO BulletinVolume 47, Issue 2 p. 174-197 Normes OEPP EPPO StandardsFree Access PM 7/40 (4) Globodera rostochiensis and Globodera pallida First published: 25 July 2017 https://doi.org/10.1111/epp.12391Citations: 40 Use of brand names of chemicals or equipment in these EPPO Standards implies no approval of them to the exclusion of others that may also be suitable. http://www.eppo.int/QUARANTINE/diag_activities/EPPO_TD_1056_Glossary.pdf AboutSectionsPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat Abstract Specific scope This Standard describes a diagnostic protocol for Globodera rostochiensis and Globodera pallida.1 Terms used are those in the EPPO Pictorial Glossary of Morphological Terms in Nematology.2 This Standard should be used in conjunction with PM 7/76 Use of EPPO diagnostic protocols. Specific approval and amendment Approved as an EPPO Standard in 2003-09. Revisions approved in 2009-09, 2012-09 and 2017-02. 1 Introduction Globodera rostochiensis and Globodera pallida (potato cyst nematodes, PCNs) cause major losses in Solanum tuberosum (potato) crops (van Riel & Mulder, 1998). The infective second-stage juveniles only move a maximum of about 1 m in the soil. Most movement to new localities is by passive transport. The main routes of spread are infested seed potatoes and movement of soil (e.g. on farm machinery) from infested land to other areas. Infestation occurs when the second-stage juvenile hatches from the egg and enters the root near the growing tip by puncturing the epidermal cell walls, and then internal cell walls, with its stylet. Eventually it begins feeding on cells in the pericycle, cortex or endodermis. The nematode induces enlargement of the root cells and breakdown of their walls to form a large, syncytial transfer cell. This syncytium provides nutrients for the nematode. Infested potato plants have a reduced root system and, because of the decreased water uptake, death of the plant can eventually occur. In this Diagnostic Protocol different tests for detection and identification are presented which can be used depending on the circumstances. In some EPPO countries official control is in place and routine testing is required. For such routine testing in the country itself molecular techniques can be very useful. In other situations, such as the testing of imported material for potential quarantine or damaging nematodes or new infestations, identification by morphological methods performed by experienced nematologists is more suitable (PM 7/76 Use of EPPO diagnostic protocols). A flow diagram describing the diagnostic procedure for G. rostochiensis and G. pallida is presented in Fig. 1. Figure 1Open in figure viewerPowerPoint Flow-diagram for identification of G. rostochiensis and G. pallida.Note that identification of G. rostochiensis and G. pallida should preferably combine morphological and molecular methods, especially when new introductions are suspected. 2 Identity Name: Globodera rostochiensis (Wollenweber, 1923), Skarbilovich, 1959. Synonyms: Heterodera rostochiensis, Wollenweber, 1923; Heterodera schachtii solani Zimmerman, 1927; Heterodera schachtii rostochiensis (Wollenweber) Kemner, 1929; Taxonomic position: Nematoda, Tylenchida1, Heteroderidae EPPO Code: HETDRO Phytosanitary categorization: EPPO A2 List no. 125, EU Annex designation I/A2 Name: Globodera pallida (Stone, 1973). Synonyms: Heterodera pallida Stone, 1973 Taxonomic position: Nematoda, Tylenchida1, Heteroderidae EPPO Code: HETDPA Phytosanitary categorization: EPPO A2 List no. 124, EU Annex designation I/A2 3 Detection 3.1 Symptoms Above-ground symptoms due to PCNs are not specific and often go undetected. General symptoms include patches of poor growth in the crop, with plants sometimes showing yellowing, wilting or death of foliage; tuber size is reduced and roots are extensively branched with soil stuck to them. However, there are many other causes of these symptoms. Plants should therefore be lifted for a visual check for the presence of cysts and young females on the roots, or a soil sample should be taken for testing. Young females and cysts are just visible to the naked eye as tiny white, yellow or brown pin-heads on the root surface (Figs 2 and 3). Detection by lifting plants is only possible for a short time as females mature into cysts and then can easily be lost at lifting, and it is time-consuming. Soil testing is therefore the best way to determine the presence of PCNs. Figure 2Open in figure viewerPowerPoint Potato roots infected by G. rostochiensis. (Courtesy: Plant Protection Service, NL.) Figure 3Open in figure viewerPowerPoint Broken cyst with eggs of G. pallida. (Courtesy: Plant Protection Service, NL.) 3.2 Statutory sampling procedures Recommendations on sampling can be found in Council Directive 2007/33/EC of 11 June 2007 on the control of PCN and Repealing Directive 69/465/EEC (EU, 2007). 3.3 Extraction procedures There are various processes for extracting cysts from the soil. Simple methods based on flotation can be as good as elutriation. Extraction methods are described in PM 7/119 Nematode extraction. Globodera cysts are generally round, which distinguishes them from most other types of nematode cysts. Prior to identification, cysts need to be removed from the floats. This process usually requires examination of the float by staff trained in separating nematode cysts from similar globular bodies in the soil. It can be time-consuming, depending upon the efficiency of extraction and whether any further clean-up has been used, e.g. acetone flotation. This process is critical to the efficiency of the diagnosis because false negative results may result if any Globodera cysts are missed at this stage. The distinction between PCNs and other cysts based on morphology can only be reliably performed by trained experts. When moist soil samples are not immediately processed and viability tests are envisaged, they should be stored above zero and below 5°C as temperature influences hatching behaviour (Muhammad, 1996; Sharma & Sharma, 1998). Soil samples should not be dried at a temperature higher than approximately 35°C as this might also influence the viability. 3.4 Bioassay Another procedure for detecting the nematodes is bioassay (Appendix 1, test A). 3.5 Direct testing of soil extracts A test for the direct detection and identification of G. rostochiensis and G. pallida in soil extracts/floats (cysts and debris) using real-time PCR has been developed by Reid et al. (2015) and is described in Appendix 7. 4 Identification Identification of G. rostochiensis and G. pallida should preferably combine morphological and molecular methods, especially when new introductions are suspected. For identification based on morphology, second-stage juveniles and cysts should be obtained from soil, plant roots or tubers. The colour of the female at the appropriate stage of development can be used as an indication of species: a female which changes during maturation from white to yellow then into a brown cyst is G. rostochiensis, while one which changes from white directly to brown is G. pallida. Identification of cysts and other stages is in general based on a combination of morphological and morphometric characters and/or molecular methods. In well-defined circumstances molecular tests alone can be applied. Differential interference contrast is highly recommended for identifying specimens mounted on microscope slides. 4.1 Identification on the basis of morphological features 4.1.1 Identification of cyst and juveniles to genus level 4.1.1.1 Cysts Identification of Heteroderidae cysts to genus level is based on the form of the cysts and the characteristics of the vulval–anal region (Table 1 and Figs 4-7). Further information is provided by keys of Brzeski (1998), Baldwin & Mundo-Ocampo (1991), Wouts & Baldwin (1998), Siddiqi (2000) and Subbotin et al. (2010). Table 1. Dichotomous key to Globodera species. (After Subbotin et al. (2010).) 1 Cuticle of cyst thin, transparent G. mali Cuticle of cyst thick, dark in colour 2 2 Mean length of J2 stylet ≤26 μm 3 Mean length of J2 stylet ≥27 μm G. zelandica 3 Mean length of J2 stylet <19 μm G. leptonepia Mean length of J2 stylet ≥19 μm 4 4 Hyaline region of J2 > 31 μm G. bravoae Hyaline region of J2 ≤ 31 μm 5 5 Mean Granek's ratio usually >2, mostly parasites of Solanaceae 6 Mean Granek's ratio ≤2, mostly parasites of Asteraceae 11 6 Combination of: mean J2 DGO ≥5.5 μm; mean Granek's ratio <3; J2 lip region with 4–6 annules, stylet knobs rounded to slightly anteriorly projected 7 Not with the above combination of all characters; mean J2 DGO <5.5 μm 8 7 Cyst wall lacking a network-like pattern, ridges close; mean number of cuticular ridges = 13 (10–18); ♂ spicules with a pointed, thorn-like tip G. ellingtonae Cyst wall exhibiting network-like or maze-like patterns; mean number of cuticular ridges = 7–8 (5–15); ♂ spicules with a finely rounded tip G. tabacum 8 Cysts with prominent bullae in the terminal region of most specimens; J2 lip region with 3 annules, mean hyaline region >28 μm G. capensis Cyst abullate, at most with small vulval bodies in some specimens; J2 lip region with 4–6 annules, mean hyaline region <28 μm 9 9 J2 stylet knobs distinctly anteriorly directed to flattened anteriorly; mean J2 stylet length >23 μm; Granek's ratio <3 10 J2 stylet knobs rounded to flattened anteriorly; mean J2 stylet length <23 μm; Granek's ratio ≥3 G. rostochiensis 10 Mean Granek's ratio = 2.1–2.5 G. pallida Mean Granek's ratio = 2.8 G. mexicana 11 J2 lip region with 5–6 annules 12 J2 lip region with 3 annules G. capensis 12 Mean stylet ≥25 μm in J2, ♂ gubernaculum = 11.2–12.9 μm G. millefolii Mean stylet <25 μm in J2, ♂ gubernaculum = 6.0–9.9 μm G. artemisiae The morphological key to Globodera species presented above has used the mean average of morphometric characters to assist with differentiation, due to the large overlap of ranges. If diagnosis of a population is carried out using morphological examination only, it is recommended to compare specimens with recent taxonomic descriptions and with the information provided in Table 2. Figure 4Open in figure viewerPowerPoint Form of cysts and characteristics of the vulval-anal region. (After Baldwin & Mundo-Ocampo (1991).) Figure 5Open in figure viewerPowerPoint The perineal region of a Globodera cyst (Hesling, 1978). Figure 6Open in figure viewerPowerPoint Heteroderidae cysts. Scale bar = 350 μm. (Courtesy NAK, NL.) Figure 7Open in figure viewerPowerPoint Perineal region. Green arrows indicate the vulva, black arrows the anus. Globodera spp. vulval fenestra/anal region non-fenestrate. Punctodera spp. vulval fenestra/anal region fenestrate. (Courtesy NAK, NL.) A dichotomous key to genus of Heteroderidae cysts is presented below: 1 Lemon-shaped cyst Not Globodera Round cyst 2 2 Two large separated fenestrae of equal size Not Globodera One large vulval fenestra Globodera Globodera cysts should present the following characteristics: Cysts of Globodera are smoothly rounded with a small projecting neck, no terminal cone, diameter ± 450 μm, and with a tanned brown skin (Fig. 6A). The cuticle surface has a zigzag pattern of ridges; a distinct D-layer is present. The perineal area (Figs 5 and 7A) consists of a single circumfenestration around the vulval slit, with tubercules on crescents near the vulva. The anus is subterminal without fenestra, the vulva is in a vulval basin; underbridge and bullae are rarely present (Fleming & Powers, 1998), and in particular not present in G. rostochiensis and G. pallida. Eggs retained in cyst, no egg-mass present. 4.1.1.2 Juveniles In addition to the juveniles in cysts, juveniles of cyst nematodes may be found incidentally in soil extracts after extraction for the detection of non-sedentary stages of nematodes. Distinction between juveniles of Globodera and other Heteroderidae is difficult; in such cases it is strongly advised to perform a cyst extraction where possible or to perform a molecular test on the juveniles (see section 4.2) and to proceed with this Diagnostic Protocol. Some information, however, is provided below. Globodera juveniles should present the following characteristics: The mobile second-stage juveniles of Globodera are vermiform, annulated, and taper at head and tail regions. Within the genus Globodera, body length ranges from 445 to 510 μm, stylet length is 18–29 μm, tail length 37–55 μm and the hyaline tail part 21–31 μm. Juveniles of cyst nematodes can be distinguished from juveniles of root-knot nematodes (Meloidogyne spp.) by a more heavily sclerotized lip region, relatively strong stylet, shape of the tail and more robust appearance (Fig. 8). In such cases it is advised to perform a cyst extraction or to perform a molecular test on the juveniles. Figure 8Open in figure viewerPowerPoint Difference between Meloidogynidae and Heteroderidae juveniles. Comparison between Meloidogyne hapla and G. pallida. (Courtesy Fera.) 4.1.2 Identification to species level The identification of Globodera to species level based on morphology can be difficult because of the observed variability of key characteristics. Therefore the use of a combination of cyst and second-stage juvenile characteristics is recommended for reliable identification. First the nematodes should be identified with the key presented in Table 1. If the nematodes are identified as PCN species, species identification should be performed using the morphological and morphometric characters presented in Table 2. Table 2. Morphological and morphometric characters useful for identification of Globodera species, range and mean values in μm. (After Subbotin et al. (2010).) Species J2 body length J2 stylet Cyst measurements Knob width Knob shape Stylet length Number of cuticular ridges between anus and vulval basin Granek's ratio G. rostochiensis 468 (425–505) 3–4 Rounded to anteriorly flattened 21.8 (19–23) 16–31 (>14) 1.3–9.5 (>3) G. pallida 484 (440–525) 4–5 Distinct forward projections 23.8 (22–24) 8–20 (<14) 1.2–3.5 (<3) G. tabacum 477 (410–527) 4–5 Rounded to slightly anteriorly projected 24 (22–26) 5–15 (<10) 1–4.2 (<2.8) G. millefolii a 492 (472–515) 4–5 Rounded to anteriorly projected 25 (24–26) 4–11 (<10) 1.6 (1.3–1.9) G. artemisiae 413 (357–490) 3–5 Rounded to anteriorly flattened 23 (18–29) 5–16 (<10) 1.0 (0.8–1.7) a Krall (1978) considered G. millefolii (Kirjanova & Krall, 1965) Behrens, 1975 as species inquirenda, as the description was based on a single female. Brzeski (1998) reported on G. achilleae: ‘it may be conspecific with G. millefolii’. According to Subbotin et al., 2010, 2011 G. achilleae is a junior synonym of G. millefolii. So from this point onwards the species name G. achilleae will not be used and G. millefolii instead. Table 3. Sizes of RFLP fragments (Thiéry & Mugniéry, 1996) Species Bsh1236I RFLP pattern G. rostochiensis (European populations) 900, 190, 110 G. pallida (European populations) 500, 400, 350, 190, 110 G. ‘mexicana’ 500, 400, 190, 110 G. tabacum tabacum 445, 400, 190, 110 G. tabacum virginae 445, 400, 190, 110 G. tabacum solanacearum 445, 400, 190, 110 Globodera rostochiensis and G. pallida are morphologically and morphometrically closely related (Stone, 1973a,b). Figure 9 presents some drawings of different stages of G. rostochiensis (Fig. 9A) and G. pallida (Fig 9B). For cysts, the most important diagnostic differences are in the perineal area, i.e. the number of cuticular ridges between the vulva and anus and Granek's ratio (Fig. 10A,B). The second-stage juvenile characteristics are stylet length and stylet knob shape (Table 1, Fig. 10C). As the range of values for each of these characteristics can overlap between species care is needed. In such cases, confirmation with molecular techniques is recommended. It should also be noted that this data is for specific populations described in the publications and that natural deviations from the range may occur. Figure 9Open in figure viewerPowerPoint Illustrations on the left-hand side of plate (side labelled A in bold), G. rostochiensis: (A) entire juvenile; (B) head region of second-stage juvenile; (C) second-stage juvenile lateral field, mid-body; (D) pharyngeal region of second-stage juvenile; (E) pharyngeal region of male; (F) tail of male; (G) lateral field of male, mid-body; (H) entire cysts; (I) head and neck of female; (J) entire male. (After C.I.H. Descriptions of Plant-Parasitic Nematodes, Set 2, No. 16.) Illustrations on the right-hand side of plate (side labelled B in bold), G. pallida second-stage juvenile: (A) entire; (B) anterior; (C) head; (D) tail; (E) lateral field mid-body region; (F) lateral field tail; (G) head and face at level of lips; (H) head and face at level of base. (After Stone (1972).) Figure 10Open in figure viewerPowerPoint (A) Perineal measurements for Globodera identification. (B) Vulval-anal ridge patterns for four Globodera species. (C) Stylets from four species of Globodera. See footnote 6 (section 4.1.2) for G. achilleae. (After Fleming & Powers (1998).) When cysts without live content, meaning they do not contain viable eggs or second stage juveniles, are found, species identification is not possible2. The three other Globodera species which could cause confusion during identification of PCNs in Europe are Globodera millefolii (Kirjanova & Krall, 1965) Behrens, 19753, Globodera artemisiae (Eroshenko & Kazachenko, 1972) Behrens, 1975, and Globodera tabacum sensu lato. These first two species are not parasitic on potato but have been recorded on Achillea millefolium and Artemisia vulgaris, respectively, in comparable agricultural areas. The G. tabacum species complex (G. tabacum tabacum (Lownsbery & Lownsbery, 1954) Skarbilovich, 1959; G. tabacum solanacearum (Miller & Gray, 1972) Behrens, 1975, and G. tabacum virginiae (Miller & Gray, 1972) Behrens, 1975) is found in North and Central America. Globodera tabacum tabacum is also present in Southern Europe. It parasitizes Nicotiana tabacum (tobacco) and some other solanaceous plants (but not potato). Table 1 and Fig. 7 provide a morphometric and morphological comparison between the PCNs G. millefolii3, G. artemisiae and G. tabacum. See also Baldwin & Mundo-Ocampo (1991), Mota & Eisenback (1993), Brzeski (1998) Wouts & Baldwin (1998) and Subbotin et al. (2010) for more detailed information on other members of the Heteroderidae and identification keys. Two new Globodera species have recently been described, Globodera ellingtonae, detected on potato in Oregon, USA (Handoo et al., 2012) and in Argentina (Lax et al., 2014) and Globodera capensis detected in a potato field in South Africa (Knoetze et al., 2013). The differences between this species and PCN species are minute and only molecular methods can allow a reliable distinction. The species are only locally present in the USA, Argentina and South Africa and have not been detected in Europe so far. 4.2 Molecular methods As G. rostochiensis and G. pallida are morphologically closely related, several polymerase chain reaction (PCR)-based tests have been developed to separate the two PCN species. Recommended molecular tests are described in Appendices 3–8. It should be noted that many tests that were developed especially to distinguish G. rostochiensis from G. pallida have not been tested so far against species such as G. millefolii, G. tabacum or G. mexicana. This limitation should be noted. Tests that were developed after 2000 generally do not have these shortcomings. Specific identification of G. achilleae from G. rostochiensis and G. pallida is possible following the PCR restriction fragment length polymorphism (RFLP) test developed by Sirca et al. (2010). There are also differences between European and non-European populations of the two species, which might be made visible with sequencing techniques (Hockland et al., 2012). DNA barcoding can also be used to support identification. Identification of G. rostochiensis and G. pallida should preferably combine morphological and molecular methods, especially when new introductions are suspected. 4.2.1 PCR tests The following PCR tests are recommended for the identification of isolated cysts or individuals from G. rostochiensis and G. pallida: As performance characteristics of the different tests presented below vary (in particular with regard to their analytical specificity) the choice of test should be made according to the circumstances of use. Test Appendix Bulman & Marshall (1997): a multiplex PCR test using species-specific primers based on ribosomal 18S and ITS1 sequences 3 Thiéry & Mugniéry (1996): an internal transcribed spacer (ITS)-RFLP PCR test based on primers described by Vrain et al. (1992) 4 Real-time PCR tests for species specific identification as well as detection of G. rostochiensis, G. pallida and G. tabacum (based on LSU rDNA) available as an all-inclusive real-time PCR kit ( http://www. cleardetections.com) 5 A Taqman® real-time PCR targeting the internal transcribed spacer I (ITSI) gene (Fera) 6 High-throughput diagnosis of PCNs (Globodera spp.) in soil samples using real-time PCR (Reid et al., 2015) 7 Quantification of viable eggs of the PCNs (Globodera spp.) using either trehalose or RNA-specific real-time PCR 8 Appendix 2 describes nucleic acid isolation. 4.2.2 DNA barcoding A protocol for DNA barcoding based on COI, 18S rDNA and 28S rDNA is described in Appendix 5 of PM 7/129 DNA barcoding as an identification tool for a number of regulated pests: DNA barcoding nematodes (EPPO, 2016) and can support the identification of G. pallida and G. rostocheinsis. Sequences are available in databases including Q-bank ( http://www.q-bank.eu/Nematodes/). 4.3 Pathotypes The term ‘pathotype’ is used by the International PCN Pathotype Scheme proposed by Kort et al. (1977) but is now considered too general. Many PCN populations cannot conclusively be assigned to the pathotypes described in this scheme. There are differences in virulence between the two PCN species, in particular between populations of G. pallida, and they are of utmost importance in populations from South America, but identification at this level is not adequate at the moment and it is time-consuming, expensive and requires specific analysis (Hockland et al., 2012). Any population showing signs of a new or unusual virulence (i.e. overcoming the resistance currently available in potato cultivars in Europe) should be tested as soon as possible. In practice, the virulence of populations can be tested on a set of cultivars used in each country. An EPPO Standard, PM 3/68 Testing of potato varieties to assess resistance to Globodera rostochiensis and Globodera pallida was adopted in 2006. 4.4 Testing the viability of eggs and juveniles Testing of the viability of the eggs and juveniles may be required for regulatory purposes. This can be done by different methods. Visual morphological determination of viability (a table with descriptions and figures is provided in Appendix 9). These observations require trained personnel. Determination of viability with a bioassay. Two tests are described in Appendix 1. Such tests require more time to perform than visual morphological determination of viability and generally more time than determination of viability by hatching tests. Dormancy might play a role and should be lifted. An additional aspect of bioassays is the possibility of false negative results due to a very low cyst content. Determination of viability by hatching tests. Three tests are described in Appendix 10. Such tests require more time to perform than visual morphological determination of viability. When determining the viability with a hatching test, it should be noted that cysts which have formed recently may be dormant (e.g. when sampling is performed in the autumn after potato harvest). To break the dormancy cysts should be exposed to +4°C for at least 4 months. Determination of the viability of eggs using trehalose. The test is described in Appendix 11, based on the publications by van den Elsen et al. (2012) and Ebrahimi et al. (2015) Determination of viability and identification on the basis of RNA. The test is described in Appendix 8, based on the publication by Beniers et al. (2014). Morphological determination of viability of eggs by staining with Meldola's Blue is also possible but the chemical is not easily available so this technique is not described in this Protocol. 5 Reference material Reference material can be obtained from: National Plant Protection Organization, National Reference Laboratory, PO Box 9102, 6700 HC Wageningen (NL). Food and Environmental Research Agency (Fera), Sand Hutton, York YO41 1LZ (GB). Julius Kühn-Institut (JKI), Federal Research Centre for Cultivated Plants, Messeweg 11–12, 38104 Braunschweig (DE). French National Institute for Agricultural Research (INRA) Biology of Organisms and Populations for Plant Protection Domaine de la Motte, BP 35327, 35653 Le Rheu Cedex (FR). 6 Reporting and documentation Guidance on reporting and documentation is given in EPPO Standard PM 7/77 Documentation and reporting on a diagnosis. 7 Performance criteria When performance criteria are available, these are provided with the description of the test. Validation data is also available in the EPPO Database on Diagnostic Expertise ( http://dc.eppo.int), and it is recommended to consult this database as additional information may be available there (e.g. more detailed information on analytical specificity, full validation reports, etc.). 8 Further information Further information on this organism can be obtained from: L den Nijs and G Karssen, National Plant Protection Organization, National Reference Laboratory, PO Box 9102, 6700 HC Wageningen (NL). E-mail: l.j.m.f.dennijs@nvwa.nl or g.karssen@nvwa.nl 9 Feedback on this Diagnostic Protocol If you have any feedback concerning this Diagnostic Protocol, or any of the tests included, or if you can provide additional validation data for tests included in this protocol that you wish to share please contact: diagnostics@eppo.int. 10 Protocol revision An annual review process is in place to identify the need for revision of diagnostic protocols. Protocols identified as needing revision are marked as such on the EPPO website. When errata and corrigenda are in press, this will also be marked on the website. Acknowledgements This Standard was originally developed under the EU DIAGPRO Project (SMT 4-CT98-2252) by a partnership of contractor laboratories and intercomparison laboratories in European countries. The first draft was prepared by: L den Nijs and G Karssen, National Plant Protection Organization, National Reference Laboratory, PO Box 9102, 6700 HC Wageningen (NL). It was revised by G Anthoine (Anses, Plant Health Laboratory, Angers, FR), L Kox, G Karssen, B van den Vossenberg and L den Nijs (National Plant Protection Organization, National Reference Laboratory, Wageningen, NL). Methods performed in Germany and Austria provided by D Kaemmerer, Bayerische Landesanstalt für Landwirtschaft. Methods performed in Norway provided by C Magnusson. Methods performed in Sweden provided by S Manduric. The Standard was reviewed by the Panel on Diagnostics in Nematology. Notes 1 Developments combining a classification based on morphological data and molecular analysis refer to ‘Tylenchomorpha’ (De Ley & Blaxter, 2004). 2 It should be noted that under European conditions, especially when cysts without live content have been detected in fields used for the production of potato in the past, it is highly probable that these cysts belong to either one of the PCN species G. rostochiensis or G. pallida. 3 Krall (1978) considered G. millefolii (Kirjanova & Krall, 1965) Behrens, 1975 as species inquirenda, as the description was based on a single female. Brzeski (1998) reported on Globodera achilleae: ‘it may be conspecific with G. millefolii’. According to Subbotin et al., 2010, 2011 G. achilleae is a junior synonym of G. millefolii. So from this point onwards the species name G. achilleae will not be used but G. millefolii will be used instead. 4 They can be obtained from Ritter GmbH, Schwabenstraße 50–54, D-86836 Untermeitingen (DE). 5 Based on a test performance study between laboratories in The Netherlands (L. den Nijs pers. comm.) Appendix 1: Bioassays Test A Bioassay (method performed in Germany and Austria) This method relies on the principle that if PCNs are present in a soil sample (even in very low numbers) they will multiply when given access to the roots of growing potato plantlets in a small container. The presence of developing cysts on the roots can then be observed through the transparent walls of the special containers used4 Depending on the size of the container about 100–200 mL of soil from the sample should be put into each container, ensuring that the s" @default.
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• W4252051645 title "PM 7/40 (4)<i>Globodera rostochiensis</i>and<i>Globodera pallida</i>" @default.
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