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Studies on the Storage Rot of Sweet Potato

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Studies on the Storage Rot of Sweet Potato STUDIES ON THE STORAGE ROT OF SWEET POTATO (IPOMOEA BATATAS L & LAM) BY BOTRYODIPLODIA THEOBROMAE PAT. AND OTHER FUNGI By Anthony Elue Arinze B.Sc., M.Sc. (Lagos) a A thesis submitted in part fulfilment of it) the requirements for the Degree of Doctor of Philosophy of the University of London. Department of Botany and Plant Technology Imperial College of Science and Technology Field Station Silwood Park Ascot Berkshire U.K. AUGUST, 1978 - 2 - ABSTRACT The storage rot of sweet potato (s.p.) (Ipomoea batatas) tuberous roots by Botryodiplodia theobromae (B.t.), Botrytis cinerea (B.c.) and Cladosporium cucumerinum (C.c.) was studied. The tuber was susceptible to rot by B. theobromae but was coloni,ed to a limited extent by B. cinerea and C. cucumerinum. The role of pectic enzymes in the successful rotting of s.p. by B.t. was investigated. B.t. produced four PG isoenzymes in vitro one of which was recovered from rotted sweet potato tissue. The properties of these isoenzymes were studied. The possible interaction between the host's metabolites (phenols and oxidative • enzymes) and the pectic enzymes of B.t. was discussed in relation to the successful rotting of the tuber by the fungus. Comparatively little pectic enzyme (PG) was recovered from tissues inoculated with B.c. and no pectic enzyme was found in tissues inoculated with C.c. Low temperature treatment (0-7°C) of the tuber induced chilling injury rendering the tissues more susceptible to rot by the fungi. The accumulation of antifungal compounds by s.p. inoculated with B.t., B.c. and C.c. was examined. All the fungi induced the formation of four terpenoid phytoalexins. The terpenes were present at lower concentrations in spreading B.t. lesions than • in the very restricted lesions formed by the other fungi. B.t. was insensitive to the phytoalexins at concentrations at which they occur in the tissue whereas B.c. and C.c. were sensitive in varying degrees to these compounds. In culture, B.t. degraded the compounds with(r24h while B.c. and C.c. degraded them little or 0 not at all. The roles of pectic enzymes and phytoalexins in the susceptibility and resistance of s.p. to rot is discussed. • - 3 - ABBREVIATIONS B.c. Botrytis cinerea B.t. Botryodiplodia theobromae oC degree(s) Centigrade • C.c. Cladosporium cucumerinum cm centimeter(s) Fig figure E.s. Fusarium solani g gram(s) hour(s) 1 litre(s) LANS Long Ashton Nutrient Solution LSA Leonian Solution Agar I NaPP Sodium polypectate NaDSS Sodium dioctylsulphosuccinate N Normal nm nanometer(s) No. number(s) mg milligram(s) ml millilitre(s) mm millimetre(s) • min minute(s) M Molar P., PP page, pages r • 4 P Pectin P.e. Phoma exigua P.G. polygalacturonase p .p .m. parts per million p.s.i. pounds per square inch 6 PTE Pectate trans-leliminase PVP Polyvinylpyrollidone R.H. relative humidity s second(s) SCA Sucrose-casamino acid TLC Thin layer chromatography T.B.A. Thiobarbituric acid uv ultra violet v/v volume/volume w/v weight/volume Pg microgram(s) pl microlitre(s) 16 p micron(s) percent • • -5 - CONTENTS Page TITLE PAGE 1 ABSTRACT 2 0 ABBREVIATIONS 3 CONTENTS 5 INTRODUCTION 13 LITERATURE REVIEW 15 1. ENZYME STUDIES 15 I The Pathogen 15 II Storage rots of sweet potato 16 III The production of pectic enzymes by plant pathogens 17 (A) Historical background 17 (B) Classification of pectic enzymes 18 (C) In vitro production of pectic enzymes by 20 plant pathogens (D) In vivo production of pectic enzymes by 22 plant pathogens (0 Pectic enzymes and tissue maceration 23 (F) Pectic enzymes and cell death 26 (G) Inhibitors of pectic enzymes 28 2. DEFENCE MECHANISMS IN PLANTS 32 r I Chemical basis for resistance 33 (A) Substances inhibitory to pathogens 33 (B) The phytoalexins 34 (C) Distribution of phytoalexins 35 6 Page II Phytoalexins of sweet potato 36 III Biological activity of phytoalexins 36 (A) Sensitivity of pathogens to phytoalexins 38 • (B) Phytoalexin degradation by pathogens 39 MATERIALS AND METHODS 42 1. ENZYME STUDIES 42 • I The culture and maintenance of fungi 42 II Growth media 42 (A) Stationary cultures 42 (B) Shake cultures 45 (C) Preparation of spore suspensions 45 (D) Culture filtrates for enzyme studies 46 III Chemical reagents 47 IV Pectic enzyme substrates 47 V Buffers 48 w VI Sterilisation 49 VII Measurement of pH 49 VIII Spectrophotometry 50 IX Preparation of rotted tissue for enzyme studies 50 • (A) Tissue inoculation 50 (B) Extraction of rotted tissue for enzyme assays 50 X Extraction of plant cell wall 51 s r - 7 - Page XI Enzyme purification 52 (A) Ultra filtration 52 (B) (NH4)2 SO4 precipitation 53 • (C) Sephadex gel filtration 53 (i) Preparation of gel slurry 53 (ii) Column set-up 54 (iii) Sample application 54 (D) Isoelectric focusing 55 XII Enzyme assays 57 (A) Polygalacturonase 57 (i) ,Viscometry 57 (ii) Cup-plate assay 57 (iii) Release of reducing group 58 (iv) T.B.A. reaction 60 (B) Polyphenol oxidase 61 (C) Peroxidase 62 XIII Activity of pectic enzymes on plant tissues 62 (A) Macerating activity 62 (B) Cell death 63 (C) Release of ions from tissues 64 w XIV Enzyme adsorption by tissues 65 XV Km determination 67 s - 8 - ♦ Page XVI Determination of total phenols 67 (A) Extraction of total phenols 67 (B) Phenol estimation 68 XVII Thin layer electrofocusing 69 (A) Preparation of polyacrylamide gel 69 (PAG) solution (B) Gel polymerization 70 4 (C) Sample application and run 71 (D) Protein location 72 XVIII Protein measurement 73 (A) Colorimetric method with Folin-Ciocalteu 73 reagent • (B) Ultra violet absorption method 74 2. THE PHYTOALEXINS OF SWEET POTATO 75 I Preparation of furanoterpenoids from sweet 75 porato tubers 75 (A) Inoculation of tubers (B) Extraction of terpenes from infected tissue 75 II Thin layer chromatography of furanoterpenoids 77 (A) Set up of TLC 77 (B) Visualization reagents 77 III Bioassay of compounds separated on chromatogram against C.c. 79 (A) Location of antifungal zones 79 (B) Elution of antifungal compounds 79 • 9 Page 4 IV Quantitative evaluation of terpenes 80 (A) Total terpenes 80 (B) Ipomeamarone and other components 81 V Sensitivity of fungi to phytoalexains of sweet 82 • potato (A) Spore germination assay 82 (B) Sporeling growth 83 VI Isolation of other phenol derivatives from 83 rotted tissue 4 VII Degradation of Zpomeamarone and total terpenes 85 by fungi (A) Preparation of mycelial suspension 85 (B) Degradation arrangement 86 • EXPERIMENTAL RESULTS 88 1. ENZYME STUDIES 88 I Preliminary experiments 88 1. Conditions for B. theobromae to produce 88 pectic enzymes in culture (A) Effect of various carbon sources on 88 pectic enzyme secretion (B) Effect of age of culture on enzyme 89 production 2. Pectic enzyme activities of crude culture 91 filtrates (A) Release of reducing groups 93 (B) Effect of dilution of culture filtrate 93 (C) Thiobarbituric acid (TBA) test of 96 culture filtrate s - 10 - Page. • 3. Extraction of rotted tissue from sweet potato 96 4. Pectic enzyme activities of crude rot extracts 98 (A) Viscosity-reducing activity 98 • (B) Release of reducing groups 100 (C) TBA test of rot extract 100 II Purification of PG from culture filtrate of B.t. 103 and sweet potato rot extract 1. Ammonium sulphate fractional precipitation 103 2. Gel filtration with Sephadex G75 superfine 106 3. Isoelectric focusing 106 (A) pH 3 - 10 focusing 108 (B) pH 3.5 - 6 focusing 108 r 4. Thin-layer isoelectric focusing 113 III Properties of purified culture filtrate PG 116 isoenzymes and nit extract PG 1. Effect on tissues 116 • (A) Maceration 116 (B) Cell death 117 (C) Release of electrolytes from tissue 119 2. Km 119 IV Binding of enzymes on tissues 121 V Enzyme inhibition studies 128 1. Effect of aqueous extracts of sweet potato 129 on activity of purified PG • Page 2. Sweet potato rot zones caused by B. theobromae 130 (A) Inhibition activity of rot zones 133 (B) Distribution of PG activity in the 133 rot zones N (C) Phenol oxidase and peroxidase activities 136 (D) Total phenol content 137 2. STORAGE PROBLEMS OF SWEET POTATO 139 I Effect of environmental conditions on rot development 139 II Inoculation with other fungi 148 III Chilling injury studies 156 1. Effect of pretreatment temperatures on 156 deterioration of tubers 2. Effect of pretreatment temperatures on 157 i susceptibility to infection by B.t., B.c. and C.c. 3. Effect of low temperature treatment of 160 tissues on maceration and cell death by purified rot extract enzyme (PG3) I. 3. PHYTOALEXIN STUDIES 165 Extraction of terpenoids of infected sweet potato 165 1. TLC on silica gel 165 2. Isolation of s.p. phytoalexins 165 (A)Bioassay of phytoalexins against C.c. 168 Ilt (B) Identification of phytoalexins 171 (C) Qualitative determination of the compounds 173 A, B, C, D, in different rot zones caused by B.t., B.c. and C.c. (D) Quantitative estimation of the phytoalexins 175 formed in the different rot zones (E) Time course analysis of the phytoalexins 177 formed by s.p. inoculated with B.t. and B.c. s • -12 - Page II Extraction of some other phenol derivatives 177 1. TLC of phenol derivatives 179 2. Bioassay of phenol derivatives against 181 C.c. spores s 3. Effect of combined phenol derivatives on 184 germination of C.c. spores III Sensitivity of B.t., B.c. and C.c. to s.p. 186 phytoalexins 1. Spore germination assays 186 2. Sporeling growth assays 192 3. Activity of total terpene extract from 198 different rot zones against sporeling growth of B.t., B.c.
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