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Chapter 1 obtusisporum. According to Booth, however, 2–3-septate macroconidia of “C.” obtusisporum measure [34–50×6–7.5 µm]. “Cylindrocarpon” obtusisporum isolates obtained from California formed perithecia when cross-inoculated with “C.” macrodidymum, giving further evidence to support the misidentification theory. This was also confirmed by sequence comparisons (Halleen et al., 2006a). In 2005, Petit and Gubler confirmed the presence of “C.” macrodidymum in the USA, and concluded that black-foot disease in California is caused by “C.” macrodidymum and “C.” destructans (Petit and Gubler, 2005). Moreover, Halleen et al., (2004) established a new genus, Campylocarpon Halleen, Schroers & Crous, which is “Cylindrocarpon”-like in morphology, associated with black-foot disease of grapevines. Species of this genus and members of the former “Nectria” mammoidea group, are excluded from Neonectria/“Cylindrocarpon”, because phylogenetic analyses revealed that these species are phylogenetically not closely related to Neonectria/“Cylindrocarpon” genera (Halleen et al., 2004; Schroers et al., 2008). From this genus, two species were included as the causal agents of black-foot disease: Campylocarpon fasciculare Schroers, Halleen & Crous, which has been reported in South Africa (Halleen et al., 2004), Brazil (Correia et al., 2012), and Spain (Alaniz et al., 2011b) and Campyl. pseudofasciculare Halleen, Schroers & Crous, which has been reported in South Africa (Halleen et al., 2004), Uruguay (Abreo et al., 2010), Brazil (Correia et al., 2012) and Perú (Álvarez et al., 2012). As highlighted before, “C.” destructans was originally identified as the causal agent of black-foot disease (Maluta and Larignon, 1991), but the status of “C.” destructans as the causal agent of the disease has since then questioned. In fact, Halleen et al. (2006b), compared “C.” destructans strains isolated from diseased grapevines in France, New Zealand, Portugal and South Africa with “C.” destructans-like anamorphs obtained from various herbaceous or woody hosts. DNA analyses of their internal transcribed spacer (ITS) and partial β-tubulin genes (TUB) showed that these isolates were genetically identical with “C.” liriodendri J.D. MacDon. & E.E. Butler, which was first associated with root rot of tulip poplar (Liriodendron tulipifera L.) in California by MacDonald and Butler (1981). Thus, because these species had identical sequences, “C.” destructans isolates collected from asymptomatic or diseased grapevines affected by black-foot disease were renamed as “C.” liriodendri, associating “C.” destructans only with root rot on other herbaceous or woody hosts (Halleen et al., 12
General introduction 2006b). In addition, in order to clarify the taxonomy of “C.” destructans causing black- foot in California, Petit and Gubler (2007) also compared “C.” destructans isolates obtained from grapevines in California with “C.” liriodendri isolates from South Africa. All of them were identical, and consequently “C.” destructans isolates were also renamed as “C.” liriodendri. This species has been later reported as a black-foot pathogen of grapevine in Australia (Whitelaw-Weckert et al., 2007), Spain (Alaniz et al., 2007), Brazil (Russi et al., 2010), Iran (Mohammadi et al., 2009), Switzerland (Casieri et al., 2009), Uruguay (Abreo et al., 2010) and northeastern United States and southeastern Canada (Petit et al., 2011). The teleomorphs of “C.” liriodendri and “C.” macrodidymum were described as Neonectria liriodendri Halleen, Rego & Crous and N. macrodidyma Halleen, Schroers & Crous (Halleen et al., 2004, 2006b). In 2008, a new species associated with black-foot disease of grapevines, “C.” pauciseptatum Schroers & Crous, was described in New Zealand and Slovenia (Schroers et al., 2008). To date, this species has been isolated from affected grapevines in Uruguay (Abreo et al., 2010), Canada (O’Gorman and Haag, 2011) and Spain (Martin et al., 2011). Phylogenetic studies carried out in New Zealand and Slovenia by Schroers et al. (2008), indicated that “C.” pauciseptatum is the closest phylogenetic sister-taxon of “C.” macrodidymum and both species are closely related to the “C.” destructans-complex, which also includes “C.” liriodendri. In addition, other “Cylindrocarpon” species have been associated with black- foot disease of grapevine: “Cylindrocarpon” didymum (Harting) Wollenw. in Canada (Petit et al., 2011), “C.” olidum (Wollenw.) Wollenw. in Spain (De Francisco et al., 2009) and “C.” olidum var. crassum Gerlach in Uruguay (Abreo et al., 2010). Thus, at this moment, “C.” destructans, “C.” liriodendri, “C.” macrodidymum, “C.” obtusisporum, “C.” pauciseptatum, Campyl. fasciculare and Campyl. pseudofasciculare were considered as the main species associated with young vines showing symptoms of black-foot disease in most of grapevine producing areas worldwide. In 2011, Chaverri et al., performed a phylogenetic study of Neonectria, “Cylindrocarpon” and related genera with “Cylindrocarpon”-like anamorphs (Chaverri et al., 2011). Morphological and molecular phylogenetic analyses data accumulated over several years have indicated that Neonectria sensu stricto and “Cylindrocarpon” sensu stricto are phylogenetically congeneric, while Neonectria sensu lato and 13
Page 1: TESIS DOCTORAL ESCUELA TÉCNICA SUP
Page 5: A mis padres A Quique
Page 8 and 9: Agradecimientos A Arianna Rustignol
Page 10 and 11: Summary The soil of grapevine roots
Page 13 and 14: Resumen Desde principios de los añ
Page 15: Resumen Es importante destacar que
Page 18 and 19: Resum Mitjançant l’ús de quatre
Page 21 and 22: Content Index Figure index………
Page 23 and 24: Figure Index Figure 1.1. A, black d
Page 25: Table Index Table 1.1. Fungal speci
Page 29 and 30: 1.- GENERAL INTRODUCTION 1.1.- Youn
Page 31 and 32: General introduction associated wit
Page 33 and 34: General introduction perennial host
Page 35 and 36: General introduction chlorosis and
Page 37: General introduction the species an
Page 41 and 42: General introduction Portugal (Cabr
Page 43 and 44: General introduction produced from
Page 45 and 46: General introduction They are gener
Page 47 and 48: General introduction covering graft
Page 49 and 50: General introduction foot pathogens
Page 51 and 52: General introduction a semi-commerc
Page 53 and 54: General introduction obtained by Bl
Page 55 and 56: General introduction Aroca, A., Gar
Page 57 and 58: General introduction Chaverri, P.,
Page 59 and 60: General introduction Gramaje, D., A
Page 61 and 62: General introduction Larignon, P. 1
Page 63 and 64: General introduction Petit, E. and
Page 65 and 66: General introduction Serra, S., Man
Page 67: Chapter 2
Page 70 and 71: Chapter 2 5. Study the effects of t
Page 73: Chapter 3 Detection of black-foot d
Page 76 and 77: Chapter 3 grapevines as symptomatic
Page 78 and 79: Chapter 3 been recently used to det
Page 80 and 81: Chapter 3 of denaturation at 94°C
Page 82 and 83: Chapter 3 cut grafting machines and
Page 84 and 85: Chapter 3 macrodidyma-complex in bo
Page 86 and 87: Chapter 3 obtained by Halleen et al
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Chapter 3 Alaniz, S., Abad-Campos,
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Chapter 3 canes from trunks of infe
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Chapter 3 control with combined hot
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Chapter 4 Detection of black-foot a
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Chapter 4 External symptoms of this
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Chapter 4 Greenhouse experiments So
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Chapter 4 The identification of Ily
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Chapter 4 Table 4.1. Incidence and
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Chapter 4 isolation ranged from 1.0
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Chapter 4 al., 2006), the positive
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Chapter 4 Aroca A, Raposo R (2007)
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Chapter 4 Gramaje D, Mostert L, Arm
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Chapter 4 Whiteman SA, Jaspers MV,
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Chapter 5 Detection and quantificat
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Chapter 5 Based on current data, bl
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Chapter 5 Materials and methods Fie
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Chapter 5 Sensitivity of the qPCR a
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Chapter 5 Table 5.2. Fungal identif
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Chapter 5 In general, qPCR analyses
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Chapter 5 the qPCR techniques have
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Chapter 5 Filion M, St-Arnaud M, Ja
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Chapter 6
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Abstract Weeds as potential hosts W
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Weeds as potential hosts undetected
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A C E F Weeds as potential hosts Fi
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Weeds as potential hosts the E.Z.N.
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Weeds as potential hosts Table 6.1.
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Weeds as potential hosts very varia
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Weeds as potential hosts 256 (mean
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Weeds as potential hosts in the C.
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Weeds as potential hosts 10. Black,
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Weeds as potential hosts disease in
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Weeds as potential hosts 54. Wisler
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Chapter 7 Effects of temperature, p
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Chapter 7 discoloration and necrosi
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Chapter 7 Table 7.1. Sources of iso
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Chapter 7 presence/absence of chlam
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Chapter 7 isolate Cy59 (C. lirioden
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Radial growth (cm day -1 ) Chapter
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Radial growth (as % of control) Cha
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Chapter 7 nonamended PDA (-0.3 MPa)
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Chapter 7 liriodendri isolates and
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Chapter 7 dsRNA on growth rate and
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Chapter 7 Martin M.T., L. Martin, M
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Chapter 8
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Chapter 8 The main objective of thi
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Chapter 8 be investigated is the ro
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Chapter 8 difficult isolation of th
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Chapter 8 isolates were able to gro
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Chapter 8 2010a). Thus, an integrat
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Chapter 8 Dubrowski, S. and Fabriti
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Chapter 8 Pascoe, I. G., and Cottra
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CONCLUSIONS Chapter 9 1.- Inoculum
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