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Downloaded for Personal Non-Commercial Research Or Study, Without Prior Permission Or Charge https://theses.gla.ac.uk/ Theses Digitisation: https://www.gla.ac.uk/myglasgow/research/enlighten/theses/digitisation/ This is a digitised version of the original print thesis. Copyright and moral rights for this work are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This work cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Enlighten: Theses https://theses.gla.ac.uk/ [email protected] University of Glasgow A study of the interactions between Alternaria linicola and linseed Neal Evans BSc. (Hons.) (Liverpool Polytechnic/CNAA) Department of Plant Science S.A.C., Auchincruive Ayr UK. A thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy, at the Faculty of Science, University of Glasgow. © Neal Evans, April 1996. ProQuest Number: 10391484 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 10391484 Published by ProQuest LLO (2017). C o pyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLO. ProQuest LLO. 789 East Eisenhower Parkway P.Q. Box 1346 Ann Arbor, Ml 4 8 1 0 6 - 1346 SA C Auchincruive 032459 . l O ù O - L GLASGOW UNrvERsnr LIBRARY s I•v! * I 'I Dedication This work is dedicated to my wife Lisa, ' with all the love in the world.......... I !■ :# 111 A bstract The principal aim of the study was to further the knowledge of the interaction between Alternaria linicola and the host plant linseed {Linum iisitatissimim). A novel detached cotyledon in viti'o bioassay was developed to allow the quantification of disease resistance of Linum material to A. linicola. Differences were apparent between the disease response scores of four linseed varieties when tested with seven isolates of the pathogen which differed in aggressiveness. However, there was no significant difference between the disease response scores of the varieties and no change in the ranking of varieties over three experiments. This indicated that the varieties behaved in a predictable manner to each isolate during each test. Accordingly, in a subsequent study, 102 Linum accessions were challenged with an aggressive and a non-aggressive isolate. About 75 % of the accessions gave a moderate response, although there was a continuous distribution of resistance from high susceptibility to resistant. Accessions at both extremes of the disease response consisted of breeding material, currently grown varieties and near relatives of the host species. For example, one of the more resistant accessions tested was Linum angustifolium. A sub-set o f nine Linum accessions was chosen (a range of susceptible, moderately-resistant and resistant material) and the resistance response of the material to an aggressive and a non- aggressive A. linicola isolate was investigated using a whole seedling inoculation technique. A comparison of the response of the material during the seedling test with that of the detached in vitro assay indicated that the latter test systematically, but marginally, overestimated the disease response. The in vitro bioassay scores and the seedling test scores were positively correlated following inoculation with the more aggressive of the two isolates. It was suggested that the resistance response of material could be accurately predicted by the in vitro bioassay but that a certain level of isolate aggressiveness was necessary to differentiate between responses of the accessions. Since large isolate-line interactions with respect to resistance scores were not observed, the results implied that resistance was polygenically determined. These results indicate that the bioassay for disease resistance produces an accurate measure of resistance and provides plant breeders with a useful tool which can be utilised during breeding programs. A microscopic analysis was carried out to examine differences in pathogen behaviour and host response in interactions between Alternaria linicola and three genotypes of Linum usitatissimum (previously identified as susceptible, moderately resistant and resistant to the IV . pathogen). Cotyledons of whole seedlings were inoculated with a conidial suspension of the pathogen and seedlings were incubated in a controlled environment cabinet. Observations of . the infection process were carried out at 18, 24 and 40 h post-inoculation by UV fluorescence microscopy. Data of 15 variables of the interaction was analysed using multivariate analysis of variance (MANOVA) and canonical variâtes analysis (CVA). Significant differences in pathogen development were found among the Limim accessions at 18, 24, and 40 hours after inoculation. At 18 hours after inoculation, attempted penetration by the pathogen was relatively rare on all three accessions. Canonical variâtes analysis revealed that overall differences among accessions resulted from the successful penetration of the most susceptible accession. Beyond 18 h post-inoculation attempted penetration was more common in the moderate and resistant accessions also. Overall differences among accessions were found to result from differences in the rate of colonisation of the host tissue. Pathogen development was obseiwed to occur at a quicker rate on susceptible material. Subsequent studies demonstrated that linicola growing in vitro produced a wide range of secondary metabolites. Extracts of culture filtrate were taken by reverse phase chromatography. Crude extracts produced disease-like symptoms on linseed cultivars and a range of non-host species indicating the presence of phytotoxic components in the extract. Characterised via thin layer chromatography, these included the non-host specific phytotoxins tenuazonic acid, alternariol monomethyl ether, tentoxin and two destruxin-type compounds (which closely resembled destruxin A and destruxin B). The identity of four of the compounds was confirmed by two dimensional thin layer chromatography and proton nuclear magnetic resonance spectroscopy. Linum leaf material infected with conidia of A. linicola and blastospores of Melampsora Uni were extracted using a facilitated diffusion extraction technique. The resultant extracts contained a number of compounds which were fungitoxic to Cladosporiiim cladosporioides and, to a lesser extent, A. brassicicola. One such compound corresponded to the phytoalexin conifeiyl alcohol which had previously reported to have been produced by flax seedlings inoculated with M Uni. Quantitative differences in the amount of the fungitoxic compounds produced between the inoculated and uninoculated resistant and susceptible host genotype combinations suggested that the production of fungitoxic compounds was greater in response to attempted colonisation and that as such, phytoalexin production was a component of the resistance reaction. : The main conclusions from the study were that due to the absence of qualitative differences in resistance response, resistance was polygenically determined. This observation, and the lack of a specific inhibitory response by a single host factor during the infection process indicated that the mechanism of resistance was multicomponent in nature. It is suggested that one of the major components in the interaction would appear to be the ability of the pathogen to produce non-host specific phytotoxins (some of which may be host-selective). At the cellular level, changes are induced in the host which result in the modification of the cell wall. Changes in the physical and/or chemical construction of the cell wall slow down the rate of pathogen ingress. Concurrent production of phytoalexin compounds by the host cells also has the effect of slowing or preventing pathogen ingress. Susceptible accessions apparently are induced too late or produce less of a resistance response in comparison to resistant accessions. The possibilities of an improvement in the resistance of linseed to A. linicola are discussed and it is suggested that such improvement would be both possible and beneficial. VI Table of Contents Title Page i Declaration ii Dedication hi Abstract iv Table of Contents vii List of Figures xiv List of Tables xvii List of Plates xix Publications xxi Acknowledgements xxii 1.0 Chapter 1: Introduction 1.1 The linseed host 1.1.1 The history and importance of linseed {Linum usitatissimum L.) The Linaceae Ancient History o f linseed and flax cultivation Recent histoiy o f linseed and flax cultivation Aspects o f linseed production 1.1.2 The cytogenetics of the Linum genus. 1.2 The pathogen 6 1.2.1 The genus Alternaria and the pathogen Alternaria linicola Groves and 6 Skolko. The systematics o f the genus Alternaria 6 Alternaria species pathogenic on linseed/flax 6 Characteristics and epidemiology of A. linicola 7 Seed infection 9 Climatic factors affecting disease development in the field 10 1.2.2 CouXvoX of Alternaria
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