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Pathogenicity of Meloidogyne Chitwoodi1 J Journal of Nematology 31(4):386–392. 1999. © The Society of Nematologists 1999. A Pathotype System to Describe Intraspecific Variation in Pathogenicity of Meloidogyne chitwoodi1 J. G. Van der Beek,2 P. W. Th.Maas,2 G. J. W. Janssen,3 C. Zijlstra,2 and C. H. Van Silfhout2 Abstract: Tests of eight Dutch Meloidogyne chitwoodi isolates to the differential set for host races 1 and 2inM. chitwoodi provided no evidence for the existence of host race 2 in the Netherlands. The data showed deviations from expected reactions on the differential hosts, which raised doubts of the useful- ness of the host race classification in M. chitwoodi. The term ‘‘pathotype’’ is proposed for groups of isolates of one Meloidogyne sp. that exhibit the same level of pathogenicity on genotypes of one host species. We recommend that the pathotype classification be applied in pathogen-host relationships when several genotypes of a Meloidogyne sp. are tested on several genotypes of one host species. Three pathotypes of M. chitwoodi were identified on Solanum bulbocastanum, suggesting at least two different genetic factors for virulence and resistance in the pathogen and the host species, respectively. The occurrence of several virulence factors in M. chitwoodi will complicate the successful application of resistance factors from S. bulbocastanum for developing resistant potato cultivars. Key words: biotype, Daucus carota, host race, intraspecific variation, Medicago sativa, Meloidogyne chit- woodi, nematode, pathogen-by-host interaction, pathotype, resistance, root-knot nematode, Solanum bulbocastanum, virulence. For nematode control and risk avoidance, Sasser and Triantaphyllou, 1977; Taylor and crop rotation with unsuitable or resistant Sasser, 1978). Differential host tests are used host plants is one of the most effective envi- to detect mixed nematode populations con- ronmentally safe strategies. Practical appli- sisting of more than one species and to dis- cation, however, can be complicated by in- tinguish host races (Hartman and Sasser, traspecific variation in pathogenicity as 1985). The test differentiated among M. in- shown in some plant-parasitic nematodes. cognita (Kofoid & White) Chitwood host Pathogenicity refers to the ability of nema- races 1, 2, 3, and 4, M. javanica (Treub) todes to infect a plant species, reproduce on Chitwood (without host race specification), it, and cause disease symptoms. Different M. hapla Chitwood (without host race speci- types of variation are denoted in such terms fication), and M. arenaria (Neal) Chitwood as forma specialist, host race, race, biotype, host races 1 and 2, according to their repro- and pathotype. duction on specific cultivars of cotton, to- To describe intraspecific variation in bacco, pepper, watermelon, peanut, and to- pathogenicity in Meloidogyne spp., two con- mato. More recently, a differential set was cepts are used. The best known is the host proposed for M. chitwoodi Golden, race concept, which is based on host plant O’Bannon, Santo & Finley, to distinguish differentials (Hartman and Sasser, 1985; among host races 1, 2, and 3 with specific cultivars of carrot, alfalfa, and Solanum bul- bocastanum Dun. (Mojtahedi et al., 1988; Mo- Received for publication 30 March 1998. 1 Supported in part by EC grant No. FAIR1-CT95-0896. jtahedi and Santo, 1994). In nematology, 2 DLO-Research Institute for Plant Protection (IPO-DLO), the term ‘‘host race’’ is often related to in- P.O. Box 9060, NL-6700 GW Wageningen, The Netherlands. 3 DLO-Center for Plant Breeding and Reproduction Re- traspecific variation in nematodes to a set of search (CPRO-DLO), P.O. Box 16, NL-6700 AA Wageningen, plants from different host genera. The Netherlands. E-mail: [email protected] The second way of classifying intraspecific The authors thank L. M. Poleij for technical assistance, the variation, by biotypes, is based on responses Dutch Plant Protection Service (PD, Wageningen, The Nether- lands), Applied Research for Arable Farming and Field Produc- of populations of a Meloidogyne species to tion of Vegetables (PAV, Lelystad, The Netherlands), and more than one genotype of one host plant Washington State University (WSU, Prosser, USA) for kindly providing nematode isolates and WSU for providing in vitro species. Roberts (1995) applied and elabo- plants of S. bulbocastanum SB22 and seeds of carrot and alfalfa. rated this concept to the M. incognita-tomato The critical reading of the manuscript by R.F. Hoekstra is highly appreciated. relationship. This concept is comparable to This paper was edited by T. L. Niblack. the physiological race concept in fungi. 386 Pathotypes and Host Races in Meloidogyne: Van der Beek et al. 387 The aim of this research was to study the distinguished on Daucus carota L. cv. Red intraspecific variation in M. chitwoodi in The Cored Chantenay (susceptible to host race 1 Netherlands. In addition, we sought to de- isolates and resistant to host race 2) and termine the utility of host race and biotype Medicago sativa L. cv. Thor (resistant to host testing in describing variation in M. chit- race 1 isolates and susceptible to host race woodi. 2) provided by H. Mojtahedi and L. esculen- tum cv. Moneymaker as a susceptible con- Materials and Methods trol. In a second experiment, host race 3 Nematode isolates: A root-knot nematode and nematode × host genotype interaction isolate is a sample of a population, which is were studied on the same cultivars of carrot maintained in captivity. An isolate is not nec- and alfalfa, and S. bulbocastanum Dunnett essarily genetically homozygous or homoge- ‘SB22’ (provided by C.R. Brown), S. bulbo- neous but represents a group of conspecific castanum ‘93-60-2’, and S. tuberosum L. cv. individuals from a common geographical Nicola as a susceptible control. The host site. Twelve isolates of M. chitwoodi were used race 3 isolate of M. chitwoodi reproduces well in this study (Table 1). Four of the M. chit- on S. bulbocastanum ‘SB22’ and reacts simi- woodi isolates originated from the United larly to host race 2 isolates on the differen- States; all other isolates originated from vari- tial cultivars of carrot and alfalfa (Mojtahedi ous sites in The Netherlands. The four U.S. and Santo, 1994). Solanum bulbocastanum isolates represented the three host races: ‘93-60-2’ was used in previous experiments Cba (race 1), Cbd and Cbf (race 2), and in which it appeared to resist all tested iso- Cbh (race 3), according to their reproduc- lates of M. chitwoodi and M. fallax (Janssen et tion on specific cultivars of three genera. al., 1997), except to one M. chitwoodi isolate The isolates were maintained and propa- (Van der Beek et al., 1998). To obtain the gated on Lycopersicon esculentum Miller cv. proper number of in vitro plants of S. bulbo- Moneymaker. The specific identities of the castanum, shoots were cut every 2–3 weeks isolates were confirmed with isozyme pheno- and transferred into new tubes with MS me- types for esterase and malate dehydrogenase dium containing 30 g/liter sucrose (Mu- (Esbenshade and Triantaphyllou, 1990; rashige and Skoog, 1962). Three weeks after Karssen, 1995) and by analysis of internally the last cut, rooted cuttings were large transcribed spacer (ITS) regions of ribo- enough for transplantation into soil. somal DNA (Zijlstra et al., 1995). Nematode infestation and experimental design: Plant genotypes: Host races 1 and 2 were Carrot, alfalfa, and tomato seedlings, rooted TABLE 1. Isolates of Meloidogyne chitwoodi used for study. Year of Isolate Geographic origin Previous host isolation Sourcea Ca The Netherlands Maize 1989 PD Cb The Netherlands Wheat 1990 PD Ck The Netherlands Tomato 1989 PD Co The Netherlands Black salsify 1993 PAV Cx The Netherlands Potato 1993 PD Cy The Netherlands Potato 1993 PD Cz The Netherlands Potato 1993 PD Caq The Netherlands Potato 1993 PD Cba Oregon Potato Unknown WSU; ORMc12, race 1 Cbd Washington Potato Unknown WSU; WAMc30, race 2 Cbf Oregon Potato Unknown WSU; ORMc8, race 2 Cbh California Potato Unknown WSU; CAMc2, race 3 a PD: Plant Protection Service, Wageningen, The Netherlands; PAV: Applied Research for Arable Farming and Field Production of Vegetables, Lelystad, The Netherlands; WSU: Washington State University, Prosser, Washington, USA; ORMc: M. chitwoodi isolate from Oregon; WAMc: M. chitwoodi isolate from Washington; CAMc: M. chitwoodi isolate from California. 388 Journal of Nematology, Volume 31, No. 4, December 1999 cuttings of the two S. bulbocastanum geno- J2, and ␴2 is the dispersion parameter. types, and germinating potato tuber buds Treatment effects were tested by means of were transplanted to 350-cm3 clay pots, F-statistics. Results were considered signifi- which were filled with moist silver-sand con- cant at P < 0.05. The analyses were per- taining 0.2% slow-release fertilizer (Osmo- formed with the Genstat 5 program (Genstat cote Plus; NPK 15-11-13 + 2 MgO + micros) 5, Release 3, Clarendon Press, Oxford, UK). and 0.08% NPK 12-10-18, and placed in a greenhouse compartment at 20±2°Cand Results 70% to 80% relative humidity. The trans- Nine of the 10 M. chitwoodi isolates were planted cuttings were kept at approximately host race 1 in the test on carrot, alfalfa, and 100% humidity during the first 3 days under tomato (Table 2). All isolates were pure M. a plastic covering. chitwoodi isolates, and no contamination The nematode inoculum was collected with M. hapla or other species was detected from heavily galled tomato roots on trays in by rDNA analysis. Isolate Cbf, representing a mist cabinet at 20 °C with water tempera- host race 2, produced an average of only 3.0 ture of 25 °C (Seinhorst, 1988). Hatched egg masses/plant on alfalfa. In 4 of the 10 second-stage juveniles (J2) that settled on replicates of this isolate, no egg masses were the bottom of the tray were collected at least produced on alfalfa.
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