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Relationships Between Succession and Community Structure and Function of Alnus-Associated Ectomycorrhizal Fungi in Alaskan Boreal Forests

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Relationships Between Succession and Community Structure and Function of Alnus-Associated Ectomycorrhizal Fungi in Alaskan Boreal Forests Relationships between succession and community structure and function of Alnus-associated ectomycorrhizal fungi in Alaskan boreal forests Item Type Thesis Authors Swanson, Michaela M. Download date 01/10/2021 04:42:20 Link to Item http://hdl.handle.net/11122/6860 RELATIONSHIPS BETWEEN SUCCESSION AND COMMUNITY STRUCTURE AND FUNCTION OF y4ZM/S-ASSOCIATED ECTOMYCORRHIZAL FUNGI IN ALASKAN BOREAL FORESTS By Michaela M. Swanson RECOMMENDED: Dj^Jack JylcF^irland I Dr. D iljee Taylor Dr. Roger Ruess Advisory Committee Chair Dr. Diane Wagner Chair, Departmejilof Biology and Wildlife APPROVED: . John of the Graduate School Date y RELATIONSHIPS BETWEEN SUCCESSION AND COMMUNITY STRUCTURE AND FUNCTION OF ALNUS-ASSOCIATED ECTOMYCORRHIZAL FUNGI IN ALASKAN BOREAL FORESTS A THESIS Presented to the Faculty of the University of Alaska Fairbanks in Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE By Michaela M. Swanson, B.A. Fairbanks, Alaska August 2016 Abstract Rates of production and carbon cycling in northern ecosystems depend heavily on nitrogen (N) availability across the landscape. Since much of the available N enters these systems through biological N-fixation,Alnus , with its capacity to fix large amounts of N, plays a critical role in ecosystem response to environmental change. However, because of its high phosphorus (P) demands, the abundance, distribution, and N-fixing capacity of Alnus is tightly controlled by the availability of P and its ability to assimilate P by associating with ectomycorrhizal fungal (EMF) symbionts. We assessed the potential of A.tenuifolia-associated EMF to access organic P forms of varying complexities. More than half of the communityA. on tenuifolia were individuals from the genera Alnicola and Tomentella, indicating that the community of EMF onAlnus is a relatively distinct group of host-specific ectomycorrhizal fungi. However, the aggregated acid phosphatase, phosphodiesterase and phytase activities of the Alnus-EMF community were not dramatically different from other boreal plant hosts on the root tip level. We detected variability in the activities of the two Alnus dominants to mobilize acid phosphatase and phosphodiesterase. However, it appears that contrary to the hypothesis that nitrogen-fixing species would associate with EMF types well suited to P acquisition, the potential acid phosphatase activity ofAlnicola luteolofibrillosa was significantly below the community mean. Our finding that enzyme activities of Alnus-EMF are not substantially greater than those found on other plant hosts suggests that if host specific EMF onAlnus facilitates P mobilization and uptake, the steps between P hydrolysis and assimilation into plant tissue as well as other pathways of P acquisition may be of greater importance in determining P provisioning to Alnus by EMF. v vi Table of Contents Page Signature Page .............................................................................................................................................i Title Page ................................................................................................................................................. iii Abstract ....................................................................................................................................................... v Table of Contents.....................................................................................................................................vii List of Figures............................................................................................................................................ix Acknowledgements ..................................................................................................................................xi 1. Introduction............................................................................................................................................. 1 2. Methods................................................................................................................................................... 5 2.1 Site description.................................................................................................................................. 5 2.2 Root tip sampling............................................................................................................................. 6 2.3 Root tip selection..............................................................................................................................6 2.4 Enzyme assays..................................................................................................................................7 2.5 Molecular analysis........................................................................................................................... 9 2.6 Plant host identification ................................................................................................................10 2.7 Statistical analysis...........................................................................................................................11 3. Results ....................................................................................................................................................12 3.1 Stage and horizon effects on enzyme activities..........................................................................12 3.2 Enzyme activity differences across hosts....................................................................................18 3.3 Individualistic responses within hosts across stages..................................................................20 vii Page 3.4 Host by horizon effects..................................................................................................................22 3.5 Fungal community identification................................................................................................. 23 3.6 EMF specific activities within and among hosts....................................................................... 28 4. Discussion .............................................................................................................................................30 4.1 Successional patterns of root-tip P mobilizing enzymes.......................................................... 30 4.2 Differences in enzyme activities among host species................................................................32 4.3 Structure of Alnus ectomycorrhizal communities.................................................................... 33 4.4 Patterns of EMF enzyme activities..............................................................................................36 4.5 Phytase activity...............................................................................................................................38 5. Conclusions ...........................................................................................................................................39 Appendix .................................................................................................................................................. 41 Literature Cited ........................................................................................................................................ 48 viii List of Figures Page Figure 3.1 Potential activities of A) acid phosphatase, B) phosphodiesterase and C) phytase ....13 Figure 3.2 Potential activities of phosphorus mobilizing enzymes on the surfaces....................... 16 Figure 3.3 Potential activities of acid phosphatase (100^M), acid phosphatase (250^M).............19 Figure 3.4 Potential activities of A) acid phosphatase (100^M), B) acid phosphatase (250^M) ..21 Figure 3.5 Potential activities of acid phosphatase (100^M), acid phosphatase (250^M) ............ 24 Figure 3.6 Frequency of the 19 most abundant EMF species on four host plant species.............. 26 Figure 3.7 Non-metric multidimensional scaling (NMS) ordination of EMF species ..................27 Figure 3.8 Potential activities of acid phosphatase (100^M), acid phosphatase (250^M)............ 29 ix x Acknowledgements I would like to thank my advisor Roger Ruess and my committee Knut Kielland, Jack McFarland and Lee Taylor. Ivan Lein, Laurel Lynch, Mitchel Slife, Collin Stackhouse and Josh Tanner spent long hours in challenging conditions to help collect and process samples and assist with enzyme assays. Thank you to Karl Olson for help in the lab and in the field. This work was funded by two grants from the Center for Global Change, University of Alaska Fairbanks (UAF), and was part of a larger project funded by the National Science Foundation (NSF DEB-0641033) and the Bonanza Creek Long Term Ecological Research Site (BNZ LTER). Thank you to the Institute of Arctic Biology and the Biology and Wildlife Department for your help and support. Xi xii 1. Introduction Studies on nutrient controls over net primary production (NPP) in boreal forests have focused almost entirely on nitrogen (N). There is good reason for this, due to the close coupling between plant N content, photosynthesis and growth rates (Chapin III et al. 1986), and the slow rates of soil organic matter turnover in boreal forest soils (Flanagan and Van Cleve 1983, Rineau and Garbaye 2009). We know comparatively little about terrestrial phosphorus (P) cycling in boreal forests (but see Marion et al. 1993; Giesler et al. 2002) despite the tight stoichiometric
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