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Allelopathic Influence of Tall Hedge Mustard (Sisymbrium Loeseliil.) and Spotted Knapweed (Centaurea Maculosa Lam.) on Arbuscular Mycorrhizal Fungi

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Allelopathic Influence of Tall Hedge Mustard (Sisymbrium Loeseliil.) and Spotted Knapweed (Centaurea Maculosa Lam.) on Arbuscular Mycorrhizal Fungi ALLELOPATHIC INFLUENCE OF TALL HEDGE MUSTARD (SISYMBRIUM LOESELIIL.) AND SPOTTED KNAPWEED (CENTAUREA MACULOSA LAM.) ON ARBUSCULAR MYCORRHIZAL FUNGI by LUKE D. BAINARD B.Sc, Trinity Western University, 2003 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Plant Science) THE UNIVERSITY OF BRITISH COLUMBIA February 2007 © Luke D. Bainard, 2007 Abstract Many exotic weeds interfere with other species by releasing allelochemicals into the environment that have a negative effect on their growth and distribution. Allelochemicals can have a direct influence on plant growth and/or indirect influence by disrupting interactions between plants and soil organisms, such as arbuscular mycorrhizal (AM) fungi. The goal of this research was to explore the allelopathic influences of the exotic weeds tall hedge mustard (Sisymbrium loeselii L.) and spotted knapweed (Centaurea maculosa Lam.). The allelopathic potential of tall hedge mustard was assessed using aqueous root and shoot extracts in seed germination and seedling growth bioassays. Aqueous tall hedge mustard root and shoot extracts strongly inhibited germination and growth of bluebunch wheatgrass, Idaho fescue, and spotted knapweed. Glucosinolate (GSL) analysis of tall hedge mustard tissues revealed the presence of two major GSLs (isopropyl GSL and sec-butyl GSL) and four indolylic GSLs. The degradation products of the two major GSLs (isopropyl isothiocyanate [ITC] and sec-butyl ITC) were identified in tall hedge mustard aqueous root and shoot extracts. Commercially available isopropyl ITC and sec-butyl ITC inhibited seed germination and seedling growth, suggesting their role in the allelopathic influence of tall hedge mustard. Tall hedge mustard aqueous extracts and ITCs incorporated into an agar medium inhibited Glomus intraradices Shenck & Smith spore germination and hyphal growth. Tall hedge mustard aqueous extracts strongly inhibited spore germination and hyphal growth of G. intraradices. Isopropyl ITC and sec-butyl ITC inhibited spore germination and hyphal growth, with the former exhibiting a stronger effect. Tall hedge mustard infestations were also found to reduce the AM inoculum potential of soil. The AM colonization and total biomass was reduced in bluebunch wheatgrass and spotted knapweed plants growing in tall hedge mustard infested compared to noninfested soil. Spotted knapweed is known to produce two major allelochemicals, (±)-catechin and cnicin. Both allelochemicals inhibited Glomus intraradices spore germination, and cnicin also inhibited the hyphal growth, suggesting that these allelochemicals may be involved in the inhibitory effect spotted knapweed has on AM fungi. Results of this study show that tall hedge mustard and spotted knapweed both produce allelochemicals that have the potential to directly and/or indirectly inhibit the growth of neighboring species and their AM fungal associates. Table of Contents Abstract H Table of Contents iii List of Tables vi List of Figures vii Abbreviations , ix Acknowledgements x Co-Authorship Statement xi Chapter 1. General Introduction and Literature Review 1 General Introduction 1 Literature Review 5 1.1 Plant species used in this study. 5 1.1.1 Tall hedge mustard (Sisymbrium loeseliih.) 5 1.1.2 Spotted knapweed (Centaurea maculosa Lam.) 5 1.1.3 Bluebunch wheatgrass (Pseudoroegneria spicata [Pursh.] Love) 6 1.1.4 Idaho fescue (Festuca idahoensis Elmer.) 7 1.2 Allelopathy 7 1.3 Allelochemicals of the exotic weeds used in this study 8 1.3.1 Allellochemicals of Brassicaceae 8 1.3.2 Spotted knapweed allelochemicals 9 1.4 Arbuscular mycorrhizal fungi 10 1.4.1 Glomus intraradices Shenck & Smith 12 1.5 Literature Cited ..13 Chapter 2. Role of Glucosinolate Degradation Products in the Allelopathic Potential of Tall Hedge Mustard (Sisymbrium loeselii L.) .21 2.1 Introduction 21 iii 2.2 Materials & Methods. .... 25 2.2.1 Seed sources 25 2.2.2 Tall hedge mustard extracts and whole plant leachate 25 2.2.2.1 Plant material and aqueous extract and leachate preparation 25 2.2.2.2 Seed germination bioassay , 26 2.2.2.3 Seedling growth bioassay 26 2.2.2.4 Soil bioassay 27 2.2.3 Glucosinolate analysis 28 2.2.4 Glucosinolate degradation product analysis 29 2.2.5 Phytotoxicity of glucosinolate degradation products 31 2.2.5.1 Isothiocyanates - seed germination bioassay 31 2.2.5.2 Isothiocyanates - seedling growth bioassay 32 2.2.6 Data analysis 32 2.3 Results , , 33 2.3.1 Effect of aqueous extracts and leachate on seed germination 33 2.3.2 Effect of aqueous extracts and leachate on seedling growth 35 2.3.3 Effect of aqueous extracts and leachate in soil 38 2.3.4 Glucosinolate analysis 41 2.3.5 Glucosinolate degradation product analysis. 41 2.3.6 Effect of isothiocyanates on seed germination 41 2.3.7 Effect of isothiocyanates on seedling growth 47 2.4 Discussion ..47 2.5 Literature Cited 56 Chapter 3. Inhibitory Effects of Tall Hedge Mustard (Sisymbrium loeselii L.) Allelochemicals on Arbuscular Mycorrhizal Fungi 60 3.1 Introduction 60 3.2 Materials & Methods 63 3.2.1 Seed and spore sources 63 3.2.2 Tall hedge mustard extracts and whole plant leachate preparation 63 3.2.3 Spore germination and hyphal growth bioassays 64 iv 3.2.3.1 Aqueous extracts and leachate. 64 3.2.3.2 Isothiocyanates ....65 3.2.4 Arbuscular mycorrhizal inoculum potential of tall hedge mustard infested soil 67 3.2.5 Data analysis 69 3.3 Results 70 3.3.1 Effect of aqueous extracts and leachate on spore germination. 70 3.3.2 Effect of isothiocyanates on spore germination,... 70 3.3.3 Effect of aqueous extracts and leachate on hyphal growth 70 3.3.4 Effect of isothiocyanates on hyphal growth 74 3.3.5 Arbuscular mycorrhizal inoculum potential of tall hedge mustard infested soil 74 3.4 Discussion : 74 3.5 Literature Cited , .80 CHAPTER 4. Effect of (±)-Catechin and Cnicin, Two Possible Allelochemicals of Spotted Knapweed (Centaurea maculosa), on Spore Germination and Hyphal Growth of Glomus intraradices 84 4.1 Introduction 84 4.2 Materials & Methods .....87 4.2.1 Allelochemicals ; 87 4.2.3 Spore germination and hyphal growth bioassay 87 4.2.4 Data analysis 88 4.3 Results ..! 88 4.3.1 Spore germination ..88 4.3.2 Hyphal growth '.. 90 4.4 Discussion 90 4.5 Literature Cited. .., 97 Chapter 5. General Discussion 101 5.1 Literature Cited 104 List of Tables Table 2.1a. Effect of tall hedge mustard root extract, shoot extract, and whole plant leachate on seedling growth in experiment #1 36 Table 2.1b. Effect of tall hedge mustard root extract, shoot extract, and. whole plant leachate seedling growth in,experiment #2 37 Table 2.2. Effect of tall hedge mustard root extract, shoot extract, and whole plant leachate on seed germination in soil 39 Table 2.3a. Effect of tall hedge mustard root extract, shoot extract, and whole plant leachate on seedling growth in soil in experiment #1 40 Table 2.3b. Effect of tall hedge mustard root extract, shoot extract, and whole plant leachate on seedling growth in soil in experiment #2 40 Table 2.4. Glucosinolate content of tall hedge mustard root and shoot tissues 43 Table 3.1. Characteristics of soils used in AM inoculum potential experiment 68 Table 3.2. Effect of tall hedge mustard aqueous root extract, shoot extract, and whole plant leachate on spore germination of Glomus intraradices 71 Table 3.3. Effect of tall hedge mustard aqueous root extract on hyphal growth of Glomus intraradices 73 vi List of Figures Figure 2.1. Dense stand of tall hedge mustard in southern British Columbia 24 Figure 2.2. UV absorption spectra of desulfo-GSLs extracted from tall hedge mustard root and shoot tissues 30 Figure 2.3. Effect Of tall hedge mustard root extracts, shoot extracts, and plant leachate on germination of (A) bluebunch wheatgrass, (B) Idaho fescue, (C) spotted knapweed, and (D) tall hedge mustard 34 Figure 2.4. HPLC chromatogram of desulfo-GSLs extracted from tall hedge mustard shoot tissues collected from the field 42 Figure 2.5. HPLC chromatogram of desulfo-GSLs extracted from tall hedge mustard root tissues collected from the field 42 Figure 2.6. GC-MS chromatogram of dichloromethane fraction of tall hedge mustard aqueous shoot extract 44 Figure 2.7. GC-MS chromatogram of dichloromethane fraction of tall hedge mustard aqueous root extract 44 Figure 2.8. Effect of isopropyl ITC on seed germination of (a) bluebunch wheatgrass, (b) Idaho fescue, (c) spotted knapweed, and (d) tall hedge mustard 45 Figure 2.9. Effect of sec-butyl ITC on seed germination of (a) bluebunch wheatgrass, (b) Idaho fescue, (c) spotted knapweed, and (d) tall hedge mustard 46 Figure 2.10. Effect of isopropyl ITC on radicle and coleoptile growth of bluebunch wheatgrass 48 Figure 2.11. Effect of isopropyl ITC on radicle and coleoptile growth of Idaho fescue 48 Figure 2.12. Effect of isopropyl ITC on radicle and coleoptile growth of spotted knapweed 48 Figure 2.13. Effect of isopropyl ITC on radicle and coleoptile growth of tall hedge mustard 48 Figure 2.14. Effect of sec-butyl ITC on radicle and coleoptile growth of bluebunch wheatgrass 49 Figure 2.15. Effect of sec-butyl ITC on radicle and coleoptile growth of Idaho fescue 49 Figure 2.16. Effect of sec-butyl ITC on radicle and coleoptile growth of spotted knapweed. ....49 Figure 2.17. Effect of sec-butyl ITC on radicle and coleoptile growth of tall hedge mustard '.. 49 Figure 3.1. (A) Ungerminated Glomus intraradices spore, and (B) germinated Glomus intraradices spore 66 Figure 3.2. Effect of (A) isopropyl ITC and (B) sec-butyl ITC on spore germination of Glomus intraradices 72 Figure 3.3. Effect of (A) isopropyl ITC and (B) sec-butyl ITC on hyphal growth of Glomus intraradices 75 Figure 3.4.
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