Fundam. appl. Nemalol., 1994, 17 (5),469-473

The relevance of races in dipsaci (Kiihn) Filipjev, the stem

GeertJ. W. JANSSEN DLO-Centre jor Plant Breeding and Reproduction Research (CPRO-DLO), P. o. Box 16, 6700 AA, Wageningen, the Netherlands. Accepted for publication 26 November 1993.

Summary - A review of reports regarding the (non-) host status of seventeen plant species towards eight host-races of Dilylenchus dipsaci, the stem nematode, is presented. Much variation in host renge was observed for the different host-races. The value of host ranges, host range tests and the designation of races is discussed. It is proposed to describe stem nematode populations with information about the place oforigin and host plant, with the exception of tetraploid stem nematode populations. Future research on molecular andlor biochemical characterization could lead to a new strategy of designating intraspecific variation.

Resume - Pertirnmce des races chez Ditylenchus dipsaci (Kuhn) Filipjev, le nbnatode des tiges et des bulbes - eet article passe en revue les signalisations concernant le statut de dix-sept plantes en tant qu'hote ou non-hote du nematode des tiges et des bulbes, Dilylenchus dipsaci. Une grande variabilite est notee dans la gamme d'hotes des differentes races d'hotes. La valeur de la gamme d'hotes et les tests permenant de la dHinir, ainsi que la designation des races sont discutes. I1 est propose de decrire les populations de D. dipsaci en precisant leur origine geographique et la plante hote anaquee, it I'exception toutefois des races tetraploldes. Dans le futur, une caracterisation moleculaire et (ou) biochirnique pourrait conduire it une nouvelle approche pour la definition de ces variations intraspecifiques.

Key-words: Dilylenchus dipsaci, host range, race, review, stem nematode.

A cosmopolitan distribution, including extreme cli­ Results matological areas (Viglierchio, 1971), and ability to parasitize some 500 plant species (Sturhan & Brzeski, In Table I seventeen plant species, i. e. lucerne, red 1991) have shown an almost unlimited adaptability of clover, white clover, tulip, hyacinth, , , stem , Ditylenchus dipsacl~ to various hosts oat, , sugar beet, potato, corn, , Phaseolus bean, and conditions. Ho\vever, as early as 1888 Ritzema Bos carrot, strawberry and teasel, are listed, that have been noticed different host preferences for stem nematode reported as either good, poor or non-host for eight host­ populations of different origins. Ever since, discussion races of D. dipsaci, i.e. lucerne-, red clover-, oat-, rye-, has continued regarding the extensive intraspecific var­ beet-, narcissus-, tulip-, and onion-race. Each letter iation in host range of D. dipsaci. Seinhorst (1957) be­ stands for a different reference to the literature. Owing lieved that these host preferences were constant and to a large variation in descriptions of (non-) hosts by the distinguished eleven biological races with nine plant spe­ various authors' observations are subdivided into three cies. Over the years, however, considerable variation in groups: " + " stands for a good host, on which nema­ host range of these biological - or host-races was no­ todes multiply moderately to well; "±" stands for a ticed, while the number of recorded races increased to poor host, on which nematodes multiply badly and" -" over 30 (Ladygina, 1982; Sturhan & Brzeski, 1991). stands for a non-host, where nematodes may enter, but The identification of races remained based on host do not multiply at all. Most reports are of original obser­ range observations. Although the usefulness of desig­ vations, but some literature references are reviews. This nating races has regularly been questioned (Hesling, might have caused double notations in some cases. 1966 b; Sturhan, 1969, 1971; Viglierchio, 1971; White­ head et al., 1987; Dropkin, 1988; Sturhan & Brzeski, Much variation is shown for the observed host ranges 1991), there has been no clear rejection of the race­ ofeight host-races. Observations on many plant species/ concept for D. dipsaci. host-race combinations are conflicting. Other reports, This review surveys a selected number of reports with which were not added to Table 1, could supply more information regarding the (non-) host status of different variation, like multiplication of the onion-race in tuJip crops for eight selected host-races. The reliability ofhost (Hesling, 1965) and of the red clover-race on rye (Kot­ ranges and the usefulness of designating races is evaluat­ thoff, 1950). For other races similar conflicting results ed. have been observed (e.g. Whitehead et al.) 1987). ISSN 1164-5571194105 S 4.001 © Gauthier-Villars - ORSTOM 469 G. J. W. Janssen

Table 1. Repol'led (non-) hosts of eight" hOS/-Trues " of Dirylenchus dipsaci. (+ : good host, nematodes multiply well; ± : POOT, host, nematodes multiply badly; -: non-hosl, nemalOdes don't multiply 0).

host plant lucerne- red c1over- oat- rye· host plant lucerne- red clover· oat- rye- race race race race race race race race

lucerne + ** IV[ lucerne + 0 ± 0 AKP N C BCE ABDE ±ATV A ~:* -BTV AER BE ABHjlP r. clover + 0 1. clover + 0 ± AEGK AN - BCDP ± V AE ADS P BDE ACM - ABTV DEFR BEF ABCH]LP who clover + BK K wh. clover + T B ± 0 K EN M ± 0 - BCDEGP BCDEP BD C - BT ER BE BCH]LP tulip +AE AEN tulip + 0 E ** ± 0 ± 0 ARE -G ACE AC -T CF ACHL hyacinth + 0 AN hyacinth + 0 ACDF Q ± 0 E ± 0 A E - AEG ACE AC -T CEF ACHL narcissus + 0 narcissus + 0 ** ACDEF Q ± 0 E ± 0 A - AEG AC AE AC -T CHL onion + E E DEO C onion +TV CDEFRP CES ** ± 0 ±T DF -G CP oat + A ** ACM oat +ATIN PA BCP ± 0 E ±TV AE EL - CEG ACP -TV CFR F ABH]LP rye + 0 D ** rye + TIN Q ± G ± 0 -C CP -TV HjlP sugar beet + B B DO CM sugar beet + ** B BCP 0 ± ± 0 D -B BCP B B BJLP potato + 0 potato + ATV J ± 0 A AD C ± V CL - AG C AM -T A AHP corn + 0 0 C corn + V R D C ± 0 M ± 0 S -G C 0 -T H]lP pea + AG C ADO A pea + ATV RP DS ACHjlP ± G M ±T R S A C Phas. bean + G C DO C Phas. bean +TV R S CjL ± 0 ±T RD -T carrot + 0 C DO carrOl + U Cl ± 0 ± V DS L -G P C -TV R CHP strawberry + 0 A D srrawberry + 0 AP AF A ± 0 C ± 0 D D C -A Cl teasel + 0 D teasel + 0 S Q ± 0 AA ± 0 - AG C C A ACHLP

References: A =Metlirzky (1972); B =Whitehead et al. (1987); C = Seinhorst (unpubl.); D =lones & lones (1964); E = Webster (1967); F = Southey (1957); G = Barker & Sasser (1959); H = Sayre & Mountain (1962);1= Frirzsche (1967); K =Bingefors (1967); L =Edwards & Taylor (1963); M = Kradel (1957); N = StantOn et al. (1984); 0 = Hooper (1984); P = Kir'yanova & Krall (1971); Q = Thorne (1961); R = Hesling (1966 a); S =Hesling (1972); T = Srurhan (1965); U =Locher (1963); V =Goffart (1964) ,..:* : principle race host. 470 Fundam. appl. Nematol. Races o/Ditylenchus dipsaci

It is further noteworthy that barley (not listed), if han & Brzeski, 1991), and moreover, nominating bi­ observed, was a non-host for stem nematodes. Infesta­ ological of host-races, based on host range alone, seems tion of wheat has been reported as a rare event (Lbcher, neither possible nor justifiable. 1963; Hesling, 1966 b.; Kir'yanova & Krall, 1971). On Some authors have considered the species Ditylenchus the other hand and Phaseolus vulgaris beans dipsaci as wild type polyphagous with race deviations were often good hosts for D. dipsaci populations. under selective conditions (Hesling, 1966 b; Viglierchio, 1971; Green, 1981). In line with this hypothesis, it is proposed to describe stem nematode populations ac­ Discussion cording to the host plant from which it was recovered and additionally the place of origin. For example: D. There are several arguments to question the value and dipsaci, derived from lucerne, Wageningen (The Neth­ reliability of the described host ranges and its variation; erlands). Additional information from host range tests this survey contains a collection of resuJts from different can lead to further specification, but it should be recog­ researchers, experimental conditions and time. Several nised that the variability and genetic behaviour ofpatho­ possible explanations for errors have been suggested, genicity, as well as the resistance factors of the plant like using mixrures of stem nematode populations (e.g. species, will make classification a delicate job. Thorne, 1961; Ladygina, 1982), the existence ofseveral biological races from a (original) host (e.g. Srurhan, An exception on the use of race-designation can be 1965; Kir'yanova & Krall, 1971), seasonal variation in made for populations which are cytogenetically differ­ aggressiveness of nematode populations (Fritzsche, ent, like the tetraploid giant bean-race. Successful cross­ 1967), differences in viability of the used inoculates es with other cytogenetically different diploid popula­ (Barker & Sasser, 1959; Ladygina, 1982), false negative tions confirmed no complete genetical isolation of this results due to failure of (artificial) inoculation (e.g. group (Srurhan, 1969). Differentiation in terms of a Thorne, 1961; Whitehead et al., 1987), use of resistant sibling species has been posrulated (Srurhan, 1971), but cultivars of a plant species (e.g. Jones & Jones, 1964; also for Meloidogyne hapla diploid and tetraploid var­ Srurhan, 1969; Whitehead et al., 1987; Srurhan & iants have been designated as race A and B respectively Brzeski, 1991) and variation in the physiological state of (Triantaphyllou, 1985). ' the plants (Ladygina, 1982). Although several Limitations of host range tests have In addition it is commonly recognized that variation in been discussed, they will retain their usefulness in char­ host ranges and pathogenicity of local populations of a acterizing narurally occurring stem nematode popula­ race can be genetically determined (Srurhan & Brzeski, tions for rotation management. However, results will be 1991). Numerous crossing experiments with individual reliable only for limited time and restricted areas (e.g. nematodes confirmed not only the unity of the species, Hesling, 1966 b; Whitehead et al., 1987). For scientific but also proved pathogenicity to be genetically deter­ purposes, like resistance srudies, the use of genetically mined (e.g. Srurhan 1964; Webster, 1967; Eriksson, more uniform nematode populations will be a pre­ 1974; Ladygina, 1978). Although incompatibility fac­ requisite. tors and abnormal hybrids were sometimes observed, Present molecular techniques might give rise to a new interbreeding is possibly an important reason for the method for designating virulence groups within D. dip­ variation in host plantlbiological race interaction (Srur­ saci. Economical impact by means of development of han, 1969), as is shown in Table 1. Interbreeding can new strategies in rotation management or (= R) restric­ easily occur under narural conditions in plants being tion measures seems evident. Palmer et al. (1992) used penetrated by specimens of several races, even though monoclonal antibodies to determine intraspecific var­ the nematodes of one of these races are unable to mul­ iation of stem nematode populations, whereas Wendt el tiply (Srurhan, 1964; Hesling, 1966 bj Ladygina, 1982). al. (1993) were able to differentiate populations on ribo­ The variation in host range, presented in Table I, somal DNA using Southern blot analysis with the ribo­ implies that populations from crops like onion, beet and somal cistron as a probe. However, the consistency of oat can hardly be distinguished from each other and that these molecular characters for diagnostic use still needs a differentiation and designation of these host-races to be investigated. For Globodera spp. promising results seem to have lost their value. In the case of the more have been obtained by use of molecular techniques, in specialized races, like the lucerne- and red clover-race, order to classify (virulence) polymorphisms as a more not only has considerable variation in host range been reliable and stable alternative for the pathotype scheme shown, but also some multiplication on each other's (Bakker el al., 1993). Therefore a successful strategy in hosts has been observed and even good multiplication search for molecular and/or biochemical characteriza­ on white clover (Table 1). Therefore a workable host tion of stem nematode populations will be based on differential test for distinguishing host-races seems un­ markers for pathogenicity characters, and most likely likely to be attained (e.g. Whitehead el al., 1987; Srur- not on the present race-concept of Ditylenchus dipsaci.

Vol. 17, n° 5 - 1994 471 G, J. W, Janssen

Acknowledgments KAADEL, J, (1957), Untersuchungen zum Wirtspflanzenkreis The author gratefuUy thanks A, F, van der Wal and J, van einer HerkeJnft des Stock- und Stengelalchens (Dilylenchus Bezooijen for their assistance with preliminary work and dipsaci [Kuhn 1858] Filipjev, 1936) - 2, Mitteilung, D, Sturhan, R. Janssen and J, Hoogendoorn for their critical NachrBI, dl PjlSchUlzdiensl, 11 : 32-34, comments on the manuscript. LADYGINA, N, M, (1978), rrhe genetic and physiological References compatibility of different forms of stem nematodes. VI. Crossbreeding of Dilylenchus from cultivated plants and BAKKER, L FOLKERTS,'vIA, R. T" ROUPPE VAN DER VOORT, from weedsJ, ParaziLOlogiya, 12: 349-353, J, N, A, M" DE BOER, J, M, & G 01',,1 IV! ERS, F. J, (1993), Changing concepts and molecular approaches in the man­ L.\DYGINA N. M, (1982). [Biological races, karyotypes and agement of virulence genes in potato cySt nematodes. A. hybridization,J In,' Gubina, V, G. (Ed,). [Plam and soil ne­ Rev. 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472 Fundam, appl, NemaLOI, Races of Ditylenchus dipsaci

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