Host status and reaction of <Emphasis Type="Italic">Medicago truncatula</Emphasis> accessions to infection by three major pathogens of pea (<Emphasis Type="Italic">Pisum sativum</Emphasis>) and alfalfa (<Emphasis Type="Italic">Medicago sativa</Emphasis>)

Ditylenchus dipsaci, the stem nematode of alfalfa (Medicago sativa), Mycosphaerella pinodes, cause of Ascochyta blight in pea (Pisum sativum) and Aphanomyces euteiches, cause of pea root rot, result in major yield losses in French alfalfa and pea crops. These diseases are difficult to control and the partial resistances currently available are not effective enough. Medicago truncatula, the barrel medic, is the legume model for genetic studies, which should lead to the identification and characterization of new resistance genes for pathogens. We evaluated a collection of 34 accessions of M. truncatula and nine accessions from three other species (two from M. italica, six from M. littoralis and one from M. polymorpha) for resistance to these three major diseases. We developed screening tests, including standard host references, for each pathogen. Most of the accessions tested were resistant to D. dipsaci, with only three accessions classified as susceptible. A very high level of resistance to M. pinodes was observed among the accessions, none of which was susceptible to this pathogen. Conversely, a high level of variation, from resistant to susceptible accessions, was identified in response to infection by A. euteiches.     

Cloning and characterization of pea apyrases: involvement of <Emphasis Type="Italic">PsAPY1 </Emphasis>in response to signal molecules from the pea pathogen <Emphasis Type="Italic">Mycosphaerella pinodes</Emphasis>

 Two nucleoside triphosphatase (NTPase) cDNA clones were isolated from a cDNA library of Pisum sativum L., cv. Midoriusui. The genes encoding the cDNAs were designated PsAPY1 and PsAPY2. PsAPY1 included the N-terminal amino acid sequence of an NTPase bound to pea cell wall. The phylogenic analysis indicated that PsAPY1 belongs to an NTPase subfamily responsive to environmental stimuli and that PsAPY2 belongs to a discrete subfamily, the physiological role of which is almost unknown. The adenosine triphosphatase activity of recombinant PsAPY1 was regulated by an elicitor and a suppressor from the pea pathogen Mycosphaerella pinodes. Based on these findings, we discuss the role of NTPases in response to biological stresses. Received: May 27, 2002 / Accepted: July 31, 2002     

Mechanisms of resistance in<Emphasis Type="Italic">Lycopersicon</Emphasis> germplasm to the whitefly<Emphasis Type="Italic">Bemisia argentifolii</Emphasis>

The silverleaf whiteflyBemisia argentifolii Bellows & Perring (Homoptera: Aleyrodidae) [also known as strain B of the sweetpotato whiteflyB. tabaci (Gennadius)] is a major pest of tomatoes due to both feeding damage and transmission of plant viruses. Certain wild species ofLycopersicon have demonstrated high levels of resistance to the pest. Greenhouse studies were undertaken to quantify the effects on whitefly behavior and mortality of individual, resistant plants selected from three accessions ofL. pennellii (Corr.) D’Arcy (LA 1340, LA 1674 and LA 2560), five accessions ofL. hirsutum f.typicum Humb. & Bonpl. (LA 386, LA 1353, LA 1777, PI 127826 and PI 127827) and one accession ofL. hirsutum f.glabratum C.H. Mull. (PI 126449). In no-choice experiments, fewer adults settled on leaflets of the wild species and deposited 75–100% fewer eggs compared to the cultivated tomato,L. esculentum Mill. Adult mortality ranged from 77–100% on wild accessions but was only 1% onL. esculentum. Most dead adults were trapped in glandular trichome exudates. The effects of these resistant accessions onB. argentifolii were mechanically transferable by appressing the trichome exudates onto the leaves of the susceptible tomato, indicating an association between the factors mediating the resistance and the glandular trichomes. Laboratory studies evaluated the repellent, fumigant and residual toxic effects of representative constituents of trichome exudates onB. argentifolii adults by using selected concentrations and probit analyses. RC₅₀ values (estimated concentration to repel 50% of the adults) and LC₅₀ values for fumigant and residual toxicity indicated that 2-tridecanone had low levels of repellent and residual toxicity activity; that 2-undecanone had high levels of repellent and fumigant activity; and that ginger oil (composed, in part, of sesquiterpene hydrocarbons) had high levels of repellent and residual toxicity activity. These studies suggest that multi-factor resistance exists in wild tomato germplasm. By combining genetically the observed chemical constituents of resistance into a single germplasm, the resulting resistance may be more difficult forB. argentifolii to overcome. http://www.phytoparasitica.org     

Genetics of host-pathogen interactions in the <Emphasis Type="Italic">Pyrenophora teres</Emphasis> f. <Emphasis Type="Italic">teres</Emphasis> (net form) – barley (<Emphasis Type="Italic">Hordeum vulgare)</Emphasis> pathosystem

The genetics of host-pathogen interactions in the Hordeum vulgare – P. teres f. teres pathosystem was studied in twelve resistant barley accessions, i.e. CI 9825, CI 9819, Diamond, CI 4922, CI 5401, Harbin, c-8755, c-21849, c-8721 c-23874, c-19979, c-15811. F₂ analyses of crosses with susceptible genotypes employing various isolates (from Europe, USA, Canada, and Australia) revealed that resistance is mostly isolate-specific and controlled by one or two genes. Segregation in ascospore progeny from two crosses between isolates of different origin revealed that avirulence in P. teres is also determined by one or two genes. An epistatic effect of suppressor genes on avirulence genes is proposed for the genetics of virulence to Diamond, Harbin, CI 5401 and c-8721 in the fungal crosses D (181-6 × A80) and F (H-22 × 92-178/9). Segregation in F₂ of crosses of three new sources of resistance (c-23874, c-19979, c-15811) to the susceptible cv. Pirkka was studied in laboratory and greenhouse tests by using seven P. teres isolates, i.e. 181-6, d8-3, d8-4, d9-1, d9-4, F4 and F74. In addition, virulence to these barley accessions of ascospore progeny from crosses of the same isolates was studied. Based on these studies it was concluded that depending on the isolate used, resistance of c-23874 is determined at least by two genes and in c-19979 and c-15811 by three genes. The results of this parallel analyses of genetics of resistance and genetics of virulence allows the postulation of a gene–for–gene interaction in the P. teres – H. vulgare pathosystem.     

Susceptibility and resistance of <Emphasis Type="Italic">Beta vulgaris</Emphasis> subsp. <Emphasis Type="Italic">maritima</Emphasis> to foliar rub-inoculation with <Emphasis Type="Italic">Beet necrotic yellow vein virus</Emphasis>

Four lines (designated MR0, MR1, MR2, and M8) from 13 accessions of Beta vulgaris subsp. maritima were selected on the basis of phenotypes produced after foliar rub-inoculation with Beet necrotic yellow vein virus (BNYVV). The susceptible phenotype developed bright yellow local lesions, whereas the resistant phenotype had symptoms ranging from no visible lesions to necrotic lesions at the inoculation site. MR1 and MR2 lines had a resistant phenotype depending on the isolate and the MR0 line was susceptible to all isolates of BNYVV tested. The M8 line was highly susceptible; the virus spread systemically and caused severe stunting. These plant lines will be useful for distinguishing BNYVV isolates having different pathogenicities, especially those controlled by RNA3 and/or RNA5.     

Partial stem and leaf resistance against the fungal pathogen <Emphasis Type="Italic">Botrytis cinerea</Emphasis> in wild relatives of tomato

Tomato (Solanum lycopersicum) is one of many greenhouse crops that can be infected by the necrotrophic ascomycete Botrytis cinerea. Commercial cultivation of tomato is hampered by the lack of resistance. Quantitative resistance has been reported in wild tomato relatives, mostly based on leaf assays. We aimed to identify wild tomato relatives with resistance to B. cinerea based on quantitative assays both on leaves and stem segments, monitoring infection frequency and disease expansion rate as parameters. A quantitative tomato stem segment assay was developed. This stem assay and a previously described leaf assay were used to screen a collection of 22 Solanum accessions. Significant differences in disease parameters were observed among accessions. Resistance to B. cinerea was observed in a number of wild Solanum species, including accessions of S. chilense, S. habrochaites and S. neorickii, both in the leaf assay and the stem segment assay. A number of resistant and susceptible accessions were evaluated as adult plants under greenhouse conditions. The data obtained in greenhouse assays confirmed the leaf and stem disease data. The expression of several defence-related genes was studied in a subset of accessions. There was no apparent correlation between the expression levels of the genes tested and the quantitative resistance level to B. cinerea. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Assessment of early blight (<Emphasis Type="Italic">Alternaria solani</Emphasis>) resistance in tomato using a droplet inoculation method

A droplet inoculation method was used for evaluation of tomato resistance to early blight, a destructive foliar disease of tomato caused by Alternaria solani (Ellis and Martin) Sorauer. In this test method, leaflets are inoculated with small droplets of a spore suspension in either water or a 0.1% agar solution. Early blight resistance was evaluated based on lesion size. The droplet method better discriminated the level of resistance (P < 0.001) for a range of spore densities in comparison with the more commonly used spray inoculation method. Lesions generated by droplet inoculation at 7 days after inoculation ranged from small flecks to almost complete blight with an exponential-like distribution of lesion sizes. Significant correlations (r = 0.52, 0.58, and 0.63, P < 0.001) were observed across three glasshouse tests of 54 accessions including wild species using the droplet method. The most resistant accessions included wild species: one accession of Solanum arcanum, three accessions of Solanum peruvianum, one accession of Solanum neorickii, and one of Solanum chilense. Solanum pennellii and Solanum pimpinellifolium accessions were susceptible, whereas Solanum habrochaites and Solanum lycopersicum accessions ranged from susceptible to moderately resistant. The droplet test method is simple to apply, offers a fine discrimination of early blight resistance levels, and allows objective evaluation.     

Biofilm formation and resistance to bactericides of <Emphasis Type="Italic">Pseudomonas syringae</Emphasis> pv. <Emphasis Type="Italic">theae</Emphasis>

Biofilm-grown cells of Pseudomonas syringae pv. theae (P.s.theae) wild-type strain K9301 on abiotic surface had remarkable resistance to kasugamycin in comparison to planktonically grown cells; however, the biofilm-grown cells of K9301 had the same sensitivity to copper sulfate. Because both the lesser biofilm-forming strain K9301S3 and enhanced biofilm-forming strain K9301-6 also had remarkable biofilm resistance to kasugamycin just as K9301 did and because epigallocatechin gallate, which enhanced biofilm formation of P.s.theae, had no effect on biofilm resistance to kasugamaycin, the degree of biofilm formation was not correlated with the antibiotic susceptibilities. In addition, K9301 and K9301S3 had less sensitivity to kasugamycin but had high sensitivity to copper sulfate on nonwounded leaf surfaces. These results indicate a possibility that the mechanism of P.s.theae biofilm resistance to bactericide functions on both abiotic and nonwounded leaf surfaces.     

Indirect interactions between rust (<Emphasis Type="Italic">Melampsora epitea</Emphasis>) and leaf beetle (<Emphasis Type="Italic">Phratora vulgatissima</Emphasis>) damage on<Emphasis Type="Italic">salix</Emphasis>

Willows (Salix spp.) are beneficial as a potential source of renewable energy, riparian barriers and riverbank control, yet are considered invasive weeds when they clog watercourses and lead to erosion and flooding. Interactions between willow rustMelampsora epitea (Thüm.) (Uredinales: Melampsoraceae) and leaf beetlePhratora spp. (Coleoptera: Chrysomelidae) feeding damage have an impact on effective pest management and biological control. The present study investigated the effects of(a) prior mechanical leaf damage on rust development, and(b) rust infection on beetle feeding under laboratory conditions for different time intervals and levels of damage. Willow rust infection significantly reduced the amount of leaf area consumed by beetles. The result was similar when a compatible or an incompatible rust pathotype was sprayed ontoSalix viminalis (L.) ‘Mullatin’ plants. There were no overall significant effects of mechanical damage on rust development, although the lowest level of rust infection was found with the incremental damage treatment. There were, however, differences of significance for leaf position and damage status, with damaged leaves at all positions having fewer pustules and a smaller pustule area than the corresponding undamaged leaves. There was no detectable effect of possible volatile emissions from crushed willow leaves on rust infection and development, although the volatile compoundcis-3-hexenyl acetate significantly reduced pustule diameter and overall pustule area. The results are discussed in terms of the implications for pest management and biological control. Corresponding author http://www.phytoparasitica.org posting April 6, 2003.     

Inheritance of resistance in <Emphasis Type="Italic">Solanum nigrum</Emphasis> to <Emphasis Type="Italic">Phytophthora infestans</Emphasis>

Solanum nigrum, black nightshade, is a wild non-tuber bearing hexaploid species with a high level of resistance to Phytophthora infestans (Colon et al. 1993), the causal agent of potato late blight, the most devastating disease in potato production. However, the genetic mode of resistance in S. nigrum is still poorly understood. In the present study, two S. nigrum accessions, 984750019 (N19) and #13, resistant (R) and susceptible (S), respectively, to three different isolates of P. infestans, were sexually crossed. The various kinds of progeny including F1, F2, F3, and backcross populations (BC₁; F1 × S), as well as two populations produced by self-pollinating the R parent and S parent, were each screened for susceptibility to P. infestans isolate MP 324 using detached leaf assays. Fifty seedling plant individuals of the F1 progeny were each resistant to this specific isolate, similarly to the seedling plants resulting from self-pollination of the resistant R parent. Thirty seedling plants obtained from self-pollination of the S parent were susceptible. Among a total of 180 F2 plants, the segregation ratio between resistant and susceptible plants was approximately 3: 1. Among the 66 seedling plants of the BC₁ progeny originating from crossing an F1 plant with the susceptible S parent, there were 26 susceptible and 40 resistant plants to P. infestans. The segregation patterns obtained indicated monogenic dominant inheritance of resistance to P. infestans isolate MP 324 in S. nigrum acc. 984750019. This gene, conferring resistance to P. infestans, may be useful for the transformation of potato cultivars susceptible to late blight.     

Characteristics of a <Emphasis Type="Italic">Plasmopara angustiterminalis</Emphasis> isolate from <Emphasis Type="Italic">Xanthium strumarium</Emphasis>

Leaves of Xanthium strumarium infected with downy mildew were collected in the vicinity of a sunflower field in southern Hungary in 2003. Based on phenotypic characteristics of sporangiophores, sporangia and oospores as well as host preference the pathogen was classified as Plasmopara angustiterminalis. Additional phenotypic characters were investigated such as the size of sporangia, the number of zoospores per sporangium and the time-course of their release. Infection studies revealed infectivity of the P. angustiterminalis isolate to both X. strumarium and Helianthus annuus. Inoculation of the sunflower inbred line, HA-335 with resistance to all known P. halstedii pathotypes, resulted in profuse sporulation on cotyledons and formation of oospores in the bases of hypocotyls. Infections of sunflower differential lines often led to damping-off. Molecular genetic analysis using simple sequence repeat primers and nuclear rDNA sequences revealed clear differences to Plasmopara halstedii, the downy mildew pathogen of sunflower.     

<Emphasis Type="Italic">Pythium</Emphasis> and <Emphasis Type="Italic">Phytophthora</Emphasis> species associated with root and stem rots of kalanchoe

Pythium and Phytophthora species were isolated from kalanchoe plants with root and stem rots. Phytophthora isolates were identified as Phytophthora nicotianae on the basis of morphological characteristics and restriction fragment length polymorphism (RFLP) analysis of the rDNA-internal transcribed spacer regions. Similarly, the Pythium isolates were identified as Pythium myriotylum and Pythium helicoides. In pathogenicity tests, isolates of the three species caused root and stem rots. Disease severity caused by the Pythium spp. and Ph. nicotianae was the greatest at 35°–40°C and 30°–40°C, respectively. Ph. nicotianae induced stem rot at two different relative humidities (60% and >95%) at 30°C. P. myriotylum and P. helicoides caused root and stem rots at high humidity (>95%), but only root rot at low humidity (60%).     

Induced systemic resistance of Chinese cabbage to bacterial leaf spot and <Emphasis Type="Italic">Alternaria </Emphasis>leaf spot by the root endophytic fungus, <Emphasis Type="Italic">Heteroconium chaetospira</Emphasis>

 The root endophytic fungus Heteroconium chaetospira isolate OGR-3 was tested for its ability to induce systemic resistance in Chinese cabbage against bacterial leaf spot caused by Pseudomonas syringae pv. maculicola and Alternaria leaf spot caused by Alternaria brassicae of the foliar diseases. Chinese cabbage seedlings planted in soil infested with an isolate of H. chaetospira were incubated in a growth chamber for 32 days. The first to fourth true leaves of the seedlings were challenge-inoculated with P. syringae pv. maculicola or A. brassicae. Chinese cabbage planted in soil infested with H. chaetospira showed significant decreases in the number of lesions of bacterial leaf spot or Alternaria leaf spot when compared to the control plants not treated with H. chaetospira. The results indicated that colonization of roots by H. chaetospira could induce systemic resistance in Chinese cabbage and reduce the incidence of bacterial leaf spot and Alternaria leaf spot. Received: April 24, 2002 / Accepted: August 9, 2002     

Resistance to infection by stealth: <Emphasis Type="Italic">Brassica napus</Emphasis> (winter oilseed rape) and <Emphasis Type="Italic">Pyrenopeziza brassicae</Emphasis> (light leaf spot)

Light leaf spot (Pyrenopeziza brassicae) is an important disease on winter oilseed rape crops (Brassica napus) in northern Europe. In regions where economically damaging epidemics occur, resistance to P. brassicae in commercial cultivars is generally insufficient to control the disease without the use of fungicides. Two major genes for resistance have been identified in seedling experiments, which may operate by decreasing colonisation of B. napus leaf tissues and P. brassicae sporulation. Much of the resistance present in current commercial cultivars is thought to be minor gene-mediated and, in crops, disease escape and tolerance also operate. The subtle strategy of the pathogen means that early colonisation of host tissues is asymptomatic, so a range of techniques and molecular tools is required to investigate mechanisms of resistance. Whilst resistance of new cultivars needs to be assessed in field experiments where they are exposed to populations of P. brassicae under natural conditions, such experiments provide little insight into components of resistance. Genetic components are best assessed in controlled environment experiments with single spore (genetically fixed) P. brassicae isolates. Data for cultivars used in the UK Recommended List trials over several seasons demonstrate how the efficacy of cultivar resistance can be reduced when they are deployed on a widespread scale. There is a need to improve understanding of the components of resistance to P. brassicae to guide the development of breeding and deployment strategies for sustainable management of resistance to P. brassicae in Europe.     

Isolation of pathogenicity- and gregatin-deficient mutants of <Emphasis Type="Italic">Phialophora gregata</Emphasis> f. sp. <Emphasis Type="Italic">adzukicola</Emphasis> through <Emphasis Type="Italic">Agrobacterium tumefaciens</Emphasis>-mediated transformation

Phialophora gregata f. sp. adzukicola, a causal agent of brown stem rot in adzuki beans, produces phytotoxic compounds: gregatins A, B, C, D, and E. Gregatins A, C, and D cause wilting and vascular browning in adzuki beans, which resemble the disease symptoms. Thus, gregatins are considered to be involved in pathogenicity. However, molecular analyses have not been conducted, and little is known about other pathogenic factors. We sought to isolate nonpathogenic and gregatin-deficient mutants through Agrobacterium tumefaciens-mediated transformation (ATMT) for cloning of pathogenicity-related genes. The co-cultivation of P. gregata and A. tumefaciens for 48 h at 20°C with 200 μM acetosyringone resulted in approximately 80 transformants per 10⁶ conidia. The presence of acetosyringone in the A. tumefaciens pre-cultivation period led to an increase in T-DNA copy number per genome. Of 420 and 110 transformants tested for their pathogenicity and productivity of gregatins, one nonpathogenic and three gregatin-deficient mutants were obtained, respectively. The nonpathogenic mutant produced gregatins, whereas the gregatin-deficient mutants had pathogenicity comparable to the wild-type strain. This is the first report of ATMT of P. gregata. Further analysis of these mutants will help reveal the nature of the pathogenicity of this fungus including the role of gregatin in pathogenesis.     

<Emphasis Type="Italic">Physalis ixocarpa</Emphasis> and <Emphasis Type="Italic">P. peruviana</Emphasis>, new natural hosts of <Emphasis Type="Italic">Tomato chlorosis virus</Emphasis>

Tomato chlorosis virus causes yellow leaf disorder epidemics in many countries worldwide. Plants of Physalis ixocarpa showing abnormal interveinal yellowing and plants of Physalis peruviana showing mild yellowing collected in the vicinity of tomato crops in Portugal were found naturally infected with ToCV. Physalis ixocarpa and P. peruviana were tested for susceptibility to ToCV by inoculation with Bemisia tabaci, Q biotype. Results confirmed that ToCV is readily transmissible to both species. The infection was expressed in P. ixocarpa by conspicuous interveinal yellow areas on leaves that developed into red or brown necrotic flecks, while P. peruviana test plants remained asymptomatic. Infected plants of both P. ixocarpa and P. peruviana served as ToCV sources for tomato infection via B. tabaci transmission. This is the first report of P. ixocarpa and P. peruviana as natural hosts of ToCV.     

Role of host specificity in the speciation of <Emphasis Type="Italic">Ascochyta</Emphasis> pathogens of cool season food legumes

Ascochyta/legume interactions are attractive systems for addressing evolutionary questions about the role of host specificity in fungal speciation because many wild and cultivated cool season food legumes are infected by Ascochyta spp. and most of these fungi have described teleomorphs (Didymella spp.) that can be induced in the laboratory. Recent multilocus phylogenetic analyses of a worldwide sample of Ascochyta fungi causing ascochyta blights of chickpea (Cicer arietinum), faba bean (Vicia faba), lentil (Lens culinaris), and pea (Pisum sativum) have revealed that fungi causing disease on each host formed a monophyletic group. Host inoculations of these fungi demonstrated that they were host-specific, causing disease only on the host species from which they were isolated. In contrast to the strict association between monophyletic group and host observed for pathogens of cultivated legumes, Ascochyta fungi causing disease on wild bigflower vetch (Vicia grandiflora) were polyphyletic. Genetic crosses between several pairs of closely related, host-specific, and phylogenetically distinct Ascochyta fungi were fully sexually compatible. Progeny from these crosses had normal cultural morphology and segregation of molecular markers indicating a lack of intrinsic, post-zygotic mating barriers between the parental taxa. However, when progeny from a cross between a faba bean-adapted isolate (A. fabae) and a pea-adapted isolate (A. pisi) were assessed for their pathogenicity to the parental hosts, almost all progeny were non-pathogenic to either faba bean or pea. These results suggest that although these fungi have retained the ability to mate and produce progeny with normal saprophytic fitness, progeny are severely compromised in parasitic fitness. The host specificity of these fungi, coupled with the inability of hybrid progeny to colonize and reproduce on a host, may constitute strong extrinsic, pre-zygotic and post-zygotic mating barriers in these fungi and promote the genetic isolation and speciation of host-specific taxa. A phylogeny of the host plants is also being developed, and with more extensive sampling of pathogens and hosts from sympatric populations in the centre of origin, the hypothesis of cospeciation of pathogens and hosts will be tested. The objectives of this review are: (1) to summarize recent phylogenetic, host specificity and speciation studies of Ascochyta fungi, and (2) to suggest how current and future research using these pathosystems may lead to a better understanding of the role of host specificity in the speciation of plant-pathogenic fungi and the cospeciation of pathogens and their hosts.     

Identification of a CAPS marker tightly linked to the Tomato yellow leaf curl disease resistance gene <Emphasis Type="Italic">Ty-1</Emphasis> in tomato

During the process of breeding programmes, several resistance genes have been introgressed into tomato (Solanum lycopersicum) cultivars from different wild tomato relatives. A number of these resistance genes have been mapped to chromosome 6. Among them, Ty-1 and Mi, which confer resistance to Tomato yellow leaf curl disease and to Meloidogyne spp., respectively, are in most cases incorporated in commercial hybrids. Several molecular markers tightly linked to Mi have been identified. This study was conducted in order to find an informative molecular marker linked to Ty-1. Six markers mapped in the same region as Ty-1 were analysed in plant material carrying different combinations of Ty-1 and Mi alleles. Three of the six markers revealed polymorphism among the assayed accessions. One allele of JB-1 marker showed association with Ty-1. Furthermore, the presence of Mi did not interfere with the results. The analysis of several accessions of wild tomato relatives with the three polymorphic markers allowed the establishment of the origin of the alleles found in cultivated plant material, showing that introgressions from S. lycopersicum, S. pimpinellifolium and S. habrochaites will not interfere with the results of this marker which tags Ty-1. Furthermore this analysis enabled the location of CT21, the RFLP marker from which JB-1 was designed.