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Resistance of Castanea Clones to Phytophthora Cinnamomi: Testing and Genetic Control

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Resistance of Castanea Clones to Phytophthora Cinnamomi: Testing and Genetic Control Miranda-Fontaina et. al.·Silvae Genetica (2007) 56-1, 11-21 Resistance of Castanea Clones to Phytophthora Cinnamomi: Testing and Genetic Control By M. E. MIRANDA-FONTAÍÑA1), J. FERNÁNDEZ-LÓPEZ1), A. M. VETTRAINO2) and A. VANNINI2) (Received 23th May 2005) Summary Chestnut hybrid clone forest reproduction material for The resistance of chestnut clones to Phytophthora cin- commercial use in Spain must be approved as belonging namomi was evaluated by a soil inoculation experiment to qualified or controlled categories, and its value must under controlled environmental conditions, as well as by be demonstrated according to the requirements outlined excised and intact stem tests. One-year-old plants of in Directive 1999/105/EC (EUROPEAN COUNCIL, 1999) and fifty different clones were inoculated with two isolates of RD 289/2003 (BOLETIN OFICIAL DEL ESTADO, 2003). For Phytophthora cinnamomi and evaluated fourteen weeks successful wood production in Atlantic areas, stem qual- after inoculation. There were significant differences ity and growth as well as a certain level of resistance to among clones for all root and collar rot variables. There Phytophthora sp. are used as selection criteria. While were significant differences for isolates of P. cinnamomi the first two traits can be assessed in field trials, evalu- but only for the collar rot variables. A total of 84% of ation of resistance in terms of survival is not straightfor- plants grown in infested soil showed symptoms of root rot but only 50% of the plants with root rot, showed also ward, as it is difficult to differentiate the different fac- had collar rot. The roots of resistant clones were able to tors related to survival in the field. confine the colonization, in roots and from roots to col- Different methods have been developed for screening lar. Percentage circumference of collar rot was the best the resistance of chestnut trees to Phytophthora sp. by indicator or descriptor of sensitivity, a 50% of clones using either intact seedlings (SANTINI et al., 2003; VET- were resistant or highly resistant clones, with respec- TRAINO et al., 2001a) or excised stems from seedlings or tively less than 20% and than 10% circumference of col- clones (FERNÁNDEZ-LÓPEZ et al., 2001; SALESSES et al., lar rot. Percentage of survival of plants is not sufficient 1993b; RAMOS GUEDES-LAFARGUE and SALESSES, 1998; to indicate resistance to the pathogen, as mortality may be affected by environmental conditions or by other VETTRAINO et al., 2001b). The variable used in intact or pathogens. The clonal heritability of collar rot variables excised stem inoculation tests is the length of lesion ranged between 0.54 and 0.71. The plants grown on after inoculation (SALESSES et al., 1993b; RAMOS GUEDES- inoculated soil showed a reduction in growth. The phe- LAFARGUE and SALESSES, 1998; FERNÁNDEZ-LÓPEZ et al., notypic and genotypic correlations between soil infesta- 2001; SANTINI et al., 2003; VETTRAINO et al., 2001a, tion characteristics and the length of necrosis in both 2001b). The variables used in the soil infestation test intact and excised stem tests were positive and highly are “health status of the plants (scale of 0–4)” and per- significant and indicated similar resistance rankings of centage of necrotized roots after carrying out a soil the clones. infestation test (VETTRAINO et al., 2001b; SANTINI et al., Key words: Chestnut, ink disease resistance, heritability, phe- 2003). Other qualitative and quantitative characters notypic and genotypic correlations, root collar rot, correlation were assessed by VETTRAINO et al. (2001a), such as stem test. growth in height, number of leaves, dry weight, length of the stem and root necroses, crown condition and mor- Introduction tality. Excised stem inoculation is the main method used to Phytophthora cinnamomi is an introduced pathogen test the resistance of chestnut clones to Phytophthora associated with the mortality in European chestnut cinnamomi, but this method is criticized because the trees (Castanea sativa Miller) growing on acid soils in results have not been shown to be clearly correlated the humid climate of northwestern Spain. This fungus with the results obtained in the soil inoculation test. causes ink disease, characterized by root and collar rot, Studies with species such as oak (ROBIN and DESPREZ- usually followed by death of the trees. Clonal forestry LOUSTAU, 1998, 2001) and Eucalyptus (TIPPET et al., for wood production has been promoted in the Atlantic 1985) have shown that the stem and root inoculation are coastal area of Galicia (NW Spain), with Euroasiatic closely correlated in terms of relative resistance rank- hybrid clones, mainly Castanea crenata x C. sativa. The ings. resistance of these hybrids to Phytophthora sp. is usual- ly higher than that of Castanea sativa, but there is vari- The overall aim of the present study was to select ability among different inter-specific hybrid genotypes clones with high levels of resistance to Phytophthora (VIEITEZ, 1960; SCHAD et al., 1952). cinnamomi, which are able to survive on infected soils and therefore can be recommended as forest reproduc- tion materials. The specific aims were: 1) to find useful 1 ) Centro de Investigacións Forestais e Ambientais de Lourizán. indicator variables for screening resistant clones in a Departamento de Producción Forestal. Xunta de Galicia. Apdo 127, 36080. Pontevedra, Spain. E-mail: memiranda.cifal@ soil inoculation experiment; 2) to assess the resistance siam-cma.org of chestnut clones; 3) to compare the rankings of resis- 2) Department of Plant Protection, University of Studies of tance of chestnut clones resulting from the soil infesta- Tuscia, Via S. Camillo de Lellis, 01100 Viterbo, Italy. tion test and excised and intact stem tests. Silvae Genetica 56, 1 (2007) 11 DOI:10.1515/sg-2007-0002 edited by Thünen Institute of Forest Genetics Miranda-Fontaina et. al.·Silvae Genetica (2007) 56-1, 11-21 Materials and Methods Three plants per isolate were selected to reisolate P. Plant material cinnamomi at the end of experiment, as described by THEMANN and WERRES (1998, www.bba.de/phytoph/ Fifty chestnut clones were selected in field trials by diagn_r.htm). growth and crown form (FERNÁNDEZ-LÓPEZ, 1996) and their genealogical origin determined using diagnostic Experiment design isozyme loci of chestnut species (FERNÁNDEZ-LÓPEZ, 1996): 44 clones are hybrids between Castanea crenata The trial consisted of five blocks or plots (1–5) with Sieb. et Zucc. x C. sativa Mill., one clone is a hybrid three treatments per plot/block (control (Cont), P. cinna- between Castanea mollissima Blume x C. sativa Mill. momi Lourizán (PcL) and P. cinnamomi Sigras (PcS)). Each of the fifteen block-treatment combinations was a (URQUIJO, 1956; VIEITEZ, 1960) (clone 7521), four clones belong to the species Castanea sativa Mill. (clones 3205, complete randomized design including one plant for 130, 90.042 and MARA) and one is a Castanea crenata each one of 50 clones. Fifteen plants per clone were ini- Sieb. et Zucc. clone (2091). The French hybrid clone (CA- tially used for the test, so there were 5 replicates per 15) (SALESSES et al., 1993a, 1993b) and the Spanish clone per treatment. Some plants died during prepara- hybrid clones (111 and 2073) (FERNÁNDEZ-LÓPEZ et al., tion of the test, thus, the mean number of plants per 2001) were used as resistant controls. The Spanish clone and treatment was 4,56 and the design of the test hybrid clones (110 and 125) were used as less resistant was unbalanced. A total of 440 plants were finally tested controls (FERNÁNDEZ-LÓPEZ et al., 2001). for the presence of P. cinnamomi coming from inocula- tion with PcL and PcS isolations, at the end of the The chestnut clones were propagated from cuttings experiment. during June and July 2002 and planted in plastic pots (4L) in February 2003. The plants were grown in a In addition to the soil inoculation experiment, two greenhouse at 23 ± 3°C in a mixed substrate (50% per- tests were carried out: the first experiment consisted on lite, 25% peat and 25% composted sterilized pine bark) the inoculation on the apex of excised stem of fifty one fertilized with Osmocote plus. Plants of mean height chestnut clones (33 common clones to soil inoculation 41.03 ± 21.8 cm and mean diameter 5.69 ± 1.87 mm test), using three treatments (control (without inocu- were used for the test at the end of May 2003. lum) and two isolates of P. cinnamomi (PcL and PcS)), in controlled environmental conditions in greenhouse and Isolation, identification and culture of Phytophthora with stems collected in June (spring) and October cinnamomi (autumn) (MIRANDA-FONTAÍÑA et al., 2007). The second Soil samples containing diseased chestnut roots were trial consisted on the inoculation on the apex of intact collected at two chestnut sites on the coast of Galicia plant stem of thirty clones (20 common clones to soil (northwest Spain): the nursery of the Centro de Investi- inoculation test), using two treatments (control (without gacións Forestais e Ambientais de Lourizán (L) and the inoculum) and one isolate of P. cinnamomi (PcL), in Sigrás orchard (S). Two isolates of Phytophthora cin- June (MIRANDA-FONTAÍÑA and FERNÁNDEZ-LÓPEZ, 2005). namomi were obtained and identified as belonging to In these tests, the length of lesions produced after 14 mating type A2 (VETTRAINO et al., 2001, 2005). The iso- and 21 days was used as an indicator of P. cinnamomi lates designated P. cinnamomi Lourizán (PcL) and resistance; also survival after 14, 21 and 28 days was P. cinnamomi Sigrás (PcS), were maintained on carrot- recorded in the second trial. Analyses of variance were agar (CA) in Petri-plates at 20°C in darkness (BRASIER, applied to the results to study the effects of genotype, 1969), and subcultured at 4-week intervals (VETTRAINO the isolates of P.
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