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Western Gall Rust (End0cr0nartium Harknessii (Moore)

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Western Gall Rust (End0cr0nartium Harknessii (Moore) WESTERN GALL RUST (END0CR0NARTIUM HARKNESSII (MOORE) HIRAT.) ON LODGEPOLE PINE (PINUS CONTORTA DOUGL.) IN BRITISH COLUMBIA - A STUDY OF VARIATION AND INHERITANCE OF RESISTANCE IN A NATURAL PATHOSYSTEM. By Harrison Ochieng Kojwang B. Sc. (Forestry) University of Nairobi, 1980 M. Sc. (Forestry) University of Helsinki, 1983 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES FOREST SCIENCES We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA 1989 © Harrison Ochieng Kojwang, 1989 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Forest Sciences The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date: 7a/,, 21, I9tq ABSTRACT Clones, open-pollinated, and full-sib families of lodgepole pine were inoculated with various spore collections of western gall rust to assess and describe variation in early symptom development, host resistance, and rust virulence and to determine the mode of inheritance of resistance. In addition, studies of the cytology of immature, mature and germinating aeciospores and one-dimensional SDS-PAGE (with silver stain) of total spore protein were undertaken. The frequency of some early symptoms varied significantly between open-pollinated families under some inoculation conditions, but was not related to the susceptibility of these families. In addition, the proportion of symptomatic seedlings that became galled was only slightly greater than that of asymptomatic seedlings. Early symptoms were not reliable indicators of successful infection. The frequency of uninucleate cells (58%) did not vary between the youngest and the oldest cells in immature spore chains. In mature spores, 57.5, 41.0 and 1.5 percent were uni-, bi- and trinucleate respectively. The number of nuclei in spores and germtubes increased gradually following germination up to an average of 5.6 (range of 2-9) at 34 hours. At no stage during the development and germination of aeciospores was there evidence of karyogamy in the form of a reduction in the number of nuclei per spore. Karyogamy and meiosis do not occur at spore germination in the coastal rust population sampled. Silver stained SDS-PAGE gels showed some variation among single gall spore sources. The approach has potential as a technique for distinguishing among spore sources. Sixteen grafted clones inoculated with four single-gall spore sources showed a sig• nificant interaction between clone and spore source. There were also large differences in relative susceptibility among pine clones and smaller differences among spore sources with respect to the average infection levels of pine clones. The infection levels of clones was considered to provide a better measure of the genetically determined resistance of parent trees than the degree of infection of those trees in the field. Forty open-pollinated pine families inoculated with coastal and interior spore collec• tions showed significant spore-family interactions attributable to six pine families that showed equal susceptibility to both spore sources. The coastal spore source caused much higher infection than the interior source on the other families. Estimates of narrow sense heritability 2 were as follows; 2 0.21 ± 0.10, 2 0.51 ± 0.16. Regressions h h Indiv = h Family = of the infection levels of offspring on those of their female parents were not significant. Hence selection of superior individuals requires progeny testing. A 4 by 4 diallel showed significant GCA effects and barely detectable SCA effects. The SCA component was about one third of the GCA component, indicating that inheritance of resistance is largely additive. Reciprocal and maternal effects were not significant. Stability in the pathosystem was attributed to the wide variation in host resistance and some degree of differential interactions between pine and spore genotypes. The highly variable host populations interact with much less variable pathogen populations; the latter possibly caused by the lack of sexual reproduction. As a result, the rate of selection for greater virulence may be matched by the rate of selection for resistance in spite of the much shorter life cycle of the pathogen. m Table of Contents List of Tables x List of Figures xix Acknowledgements xxii 1 GENERAL INTRODUCTION 1 2 LITERATURE REVIEW: NATURAL PATHOSYSTEMS 5 2.1 General 5 2.2 Disease In Some Natural Pathosystems 8 2.3 The Evolution of Stable Natural Pathosystems 11 2.3.1 General Theory 11 2.3.2 Stability Strategies of Vertical and Horizontal Pathosystems ... 12 2.3.3 Theoretical Models 13 2.3.4 Resistance Frequency Distributions among Families Derived from Wild Populations 15 2.4 Genetic diversity and host-pathogen interactions in some stable natural Pathosystems 17 2.5 Genetic Diversity and Stability in Crop Multilines 19 3 CYTOLOGY OF DEVELOPING AND GERMINATING SPORES 22 3.1 Introduction 22 3.2 Materials and Methods 25 iv 3.2.1 Cytology of Spore Chains 25 3.2.2 Cytology of Dormant and Germinating Spores 26 3.2.3 Cytology of Germtubes 27 3.3 Results 28 3.4 Discussion 34 4 EARLY SYMPTOMS ON SEEDLINGS AND GALL FORMATION 38 4.1 Introduction 38 4.2 Materials and Methods 40 4.2.1 Materials 40 4.2.2 The Inoculation Chamber 41 4.2.3 Observations 43 4.2.4 Data analysis . 44 4.3 Results . 47 4.3.1 Description of Early Symptoms 47 4.3.2 Frequency of symptoms among families 52 4.3.3 Symptoms and gall formation at the family level 54 4.3.4 Early symptoms and infection of individual seedlings 58 4.3.5 Variation in Gall Formation 60 4.4 Discussion 64 4.4.1 Symptoms 64 4.4.2 Variation in Gall Formation . 65 5 PINE CLONES AND SINGLE-GALL SPORE SOURCES 68 5.1 Introduction and Literature Review 68 5.2 Materials and Methods 69 5.2.1 ' Sources and Preparation of the Clones . 69 v 5.2.2 Spore collection . 70 5.2.3 Inoculation Technique 70 5.2.4 Collection of Data and Analyses 71 5.2.5 Electrophoretic Variation among Single Galls 72 5.3 Results 72 5.4 Discussion 77 6 VARIATION AMONG OPEN-POLLINATED PINE FAMILIES 80 6.1 Introduction 80 6.2 Literature Review 81 6.3 Materials and Methods 82 6.3.1 Preparation of Seedlings 82 6.3.2 Collection of Spores 83 6.3.3 Inoculation of Seedlings 83 6.3.4 Observations and Analysis of Data 84 6.3.5 Estimates of Narrow Sense Heritability (h2) 85 6.4 Results 87 6.5 Discussion . 94 6.5.1 The Inoculation Technique 94 6.5.2 Variation among the 40 Open-pollinated Families 94 7 INHERITANCE OF RESISTANCE : A DIALLEL CROSS 97 7.1 Introduction 97 7.2 Materials and Methods 99 7.2.1 Preparation of Seedlings 99 7.2.2 Spore Collection and Inoculation 99 7.2.3 Analysis of Data 101 vi 7.3 Results 103 7.4 Discussion 109 8 CONCLUSIONS AND GENERAL DISCUSSION 111 8.1 General Conclusions Ill 8.2 Discussion . 112 8.2.1 Methodology '. 112 8.2.2 The Pathogen - Nuclear Cycle and Genetic Variability 114 8.2.3 The Host 115 8.2.4 The Mode of Inheritance - The core of stability 116 8.2.5 Pathosystem' Stability 118 Bibliography 119 APPENDICES 135 A Experimental Materials used in the Study 135 A.l Stand A 135 A.2 Stand B 137 A.3 Stand C 139 B Analyses of Variance of the Frequencies of Early Symptoms 140 C Resistance Frequency Distributions of 10 Pine Families 147 D A Diallel 150 E Electrophoretic Analysis of Rust Collections 154 E.l Results ." 154 vii Discussion and Conclusions ix List of Tables 4.1 Expected mean square table for ANOVA of early symptom frequency among 10 open-pollinated lodgepole pine families inoculated with west• ern gall rust at two spore loads and two times of inoculation . 46 4.2 Summary of ANOVA of the frequencies of three symptom types produced on lodgepole pine seedlings following inoculations with western gall rust. Analyses were done separately for each symptom type at 2, 4 and 8 weeks following inoculation 53 4.3 Results of analysis of arcsine-square root of % of seedlings of open-pollinated lodgepole pine families showing red flecks, 4 weeks following inoculations with two spore loads of western gall rust 53 4.4 Infection Levels on 1-year-old lodgepole pine seedlings from 10 open-pollinated families inoculated with western gall rust spores. This table presents data from the first stage of seedling maturity and high spore load (1.0g/250 seedlings) 55 4.5 Infection Levels on 1-year-old lodgepole pine seedlings from 10 open-pollinated families inoculated with western gall rust spores. This table presents data from the second stage of seedling maturity and low spore load (0.1g/250 seedlings) 55 x 4.6 Infection Levels on 1-year-old lodgepole pine seedlings from 10 open-pollinated families inoculated with western gall rust spores. This table presents data from the second stage of seedling maturity and high spore load (1.0g/250 seedlings) 56 4.7 Infection Levels on 1-year-old lodgepole pine seedlings from 10 open-pollinated families inoculated with western gall rust spores. This table presents data from the second stage of seedling maturity and low spore load (0.1g/250 seedlings) 56 4.8 Correlations between the occurrence of individual symptom types and gall formation (percent infected per family) on lodgepole pine seedlings fol• lowing inoculations with western gall rust.
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