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Picea Abies (L.) Karst .) in Managed Forest Stands of Latvia

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Picea Abies (L.) Karst .) in Managed Forest Stands of Latvia UNIVERSITY OF LATVIA, FACULTY OF BIOLOGY LATVIAN STATE FOREST RESEARCH INSTITUTE “SILAVA” DĀRTA KĻAVI ŅA ECTOMYCORRHIZAS OF NORWAY SPRUCE (PICEA ABIES (L.) KARST .) IN MANAGED FOREST STANDS OF LATVIA DOCTORAL THESIS Submitted for the degree of Doctor of Biology Subfield: Ecology Supervisors: Dr. silv. T ālis Gaitnieks (Latvian State Forest Research Institute „Silava”); Prof. Dr. habil. biol. Indri ķis Muižnieks (University of Latvia); Dr. phil. Audrius Menkis (Swedish University of Agricultural Sciences). Riga, 2015 Abbreviations AIC Akaike's information criterion ANOVA Analysis of variance CCA Canonical Correspondence Analysis DGGE Denaturing gradient gel electrophoresis DNA Deoxyribonucleic acid ECM Ectomycorrhizae / Ectomycorrhizal ECMf Ectomycorrhizal fungi GLMM Generalized linear mixed model ITS Internal transcribed spacer JSC Joint stock company LLC Limited Liability Company LSFRI Silava Latvian State Forest Research Institute “Silava” PCA Principal Component Analysis PCR Polymerase chain reaction rRNA Ribosomal ribonucleic acid 2 Content Summary ................................................................................................................................................. 4 Kopsavilkums.......................................................................................................................................... 6 1. Introduction......................................................................................................................................... 8 2. Background ....................................................................................................................................... 10 2.1. Ectomycorrhizal Symbiosis ....................................................................................................... 10 2.1.1. General Description............................................................................................................ 10 2.1.2. Role of Ectomycorrhizas .................................................................................................... 10 2.1.3. Description Methods of ECM Communities...................................................................... 11 2.2. Norway Spruce Ecology and Mycorrhization............................................................................ 13 2.3. Norway Spruce Forest Management.......................................................................................... 15 2.4. Some Problematic Aspects of Spruce Forest Management in Latvia........................................ 16 2.5. Specific Characteristics of Conifer Mycorrhization in Forest Nurseries and after Outplanting 18 2.6. Specific Characteristics of Spruce Mycorrhization in Mature and Managed Forest Stands...... 19 3. Material and Methods........................................................................................................................ 21 3.1. Studied Sites and Experimental Design..................................................................................... 21 3.1.1. Foliar Damage Study.......................................................................................................... 22 3.1.2. Root-Rot Study................................................................................................................... 22 3.1.3. Fertilization Study .............................................................................................................. 23 3.1.4. Stump Removal Study........................................................................................................ 24 3.1.5. Seed Origin Study............................................................................................................... 25 3.1.6. Cultivation System Study................................................................................................... 25 3.2. Root Analysis............................................................................................................................. 26 3.2.1. Root Sampling.................................................................................................................... 26 3.2.3. Analysis of Root Morphology ............................................................................................ 27 3.2.4. Ectomycorrhizal Morphotyping ......................................................................................... 28 3.2.5. Molecular Identification of ECM Fungi............................................................................. 29 3.3. Background Data: Seedling Growth Evaluation, Needle and Soil Analysis.............................. 30 3.3.1. Tree Growth Evaluation, Needle Chemical Analysis......................................................... 30 3.3.2. Soil Analysis....................................................................................................................... 31 3.4. Data Analysis............................................................................................................................. 32 4. Results and Discussion...................................................................................................................... 36 4.1. Foliar Damage Study ................................................................................................................. 36 4.1.1. Soil Parameters and Root Growth Stages........................................................................... 36 4.1.2. Fine Root Fungal Community ............................................................................................ 39 4.2. Root-Rot Study .......................................................................................................................... 43 4.2.1. Soil Parameters and Fine Root Growth .............................................................................. 43 4.2.2. Fine Root Fungal Community ............................................................................................ 44 4.3. Fertilization Study...................................................................................................................... 48 4.3.1. Soil Parameters and Growth of Fine Roots ........................................................................ 48 4.3.2. Fine Root Fungal Community ............................................................................................ 50 4.4. Stump Removal Study ............................................................................................................... 54 4.4.1. Seedling Growth................................................................................................................. 54 4.4.2. Seedling Mycorrhization and ECM Community................................................................ 55 4.5. Seed Origin Study...................................................................................................................... 59 4.6. Cultivation System Study .......................................................................................................... 63 4.6.1. Seedling Growth................................................................................................................. 63 4.6.2. Seedling Mycorrhization and ECM Community................................................................ 66 4.7. Overall Results and Discussion on ECM Community and Species Ecology............................. 68 5. Conclusions....................................................................................................................................... 71 Approbation of results........................................................................................................................... 73 Acknowledgements ............................................................................................................................... 75 Reference List ....................................................................................................................................... 77 Appendix............................................................................................................................................... 92 3 Summary This thesis assembles six separate studies on ectomycorrhizal fungi (ECMf) of young and mature Norway spruce (Picea abies (L.) Karst.) under different management practices. The studies provide insight in ectomycorrhizal (ECM) species composition in relation to tree growth rate, soil parameters and forest management practices applied. The main method used in this work was fine root analysis. Morphotyping was carried out in combination with sequencing of the fungal internal transcribed spacer (ITS) region of rRNA genes to identify fungal species. Stand or seedling growth rate and health status as well as soil parameters were taken into account for data analysis and interpretation. In total, 148 fungal species were identified from sequences of root tips. Most of the species represented basidiomycetes, 105 species, mainly from the orders of ECMf Atheliales , Agaricales , Russulales and Thelephorales . Ascomycetes were represented by 43 species, mainly from the Helotiales and Pezizales orders. Many species recorded were new for Latvia as only few of these taxonomic groups had been studied before. The results showed that fine root parameters were related to Norway spruce heath status and could serve as indicators of tree vitality. In stands with severe foliar damage symptoms, the proportion of living fine roots was reduced and greater abundance of saprotrophic species
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