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דוקטור לפילוסופיה Doctor of Philosophy

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דוקטור לפילוסופיה Doctor of Philosophy חבור לשם קבלת התואר Thesis for the degree דוקטור לפילוסופיה Doctor of Philosophy מאת By תמיר קליין Tamir Klein אקו - פיזיולוגיה של ניצול מים באורן ירושלים : מרמת העלה ועד רמת היער Eco-physiology of water use in Pinus halepensis : from leaf to forest scale מנחה : פרופ ' דן יקיר Advisor: Prof. Dan Yakir בשיתוף ד"ר שבתאי כהן ( מינהל Co-mentored by Dr. Shabtai המחקר החקלאי מכון וולקני) (Cohen (ARO Volcani תמוז תשע"ב November 2012 מוגש למועצה המדעית של Submitted to the Scientific Council of the מכון ויצמן למדע Weizmann Institute of Science רחובות , ישראל Rehovot, Israel This work has been conducted under the supervision of Prof. Dan Yakir Department of Environmental Sciences and Energy Research Weizmann Institute of Science Rehovot, Israel And Dr. Shabtai Cohen Institute of Soil, Water and Environmental Sciences Volcani Center ARO Beit Dagan, Israel 1 Acknowledgements I was blessed with two exceptional mentors, Prof. Dan Yakir and Dr. Shabtai Cohen, whose positive, passionate approach to scientific research and to any learning in general, has been an inspiration to me. Their enthusiasm about plant eco-physiology together with the highest level of professionalism has made a profound impact on my work. I wish to thank my advisory committee, Profs. Brian Berkowitz and Amram Eshel for their interest in the work and advice for improvements. I have had the privilege of illuminating discussions with Prof. Gabi Schiller, Dr. Jose Gruenzweig, Dr. Gilboa Arye, Dr. Ron Milo, Prof. Harvey Scher, Prof. Yohai Carmel and Dr. Yagil Osem, that broadened my knowledge and understanding. The members of Dan’s group with whom I have worked: Fyodor, David, Uri, Hemu, Naama, Keren, Gil and Amir; have been research collaborators and good friends. Special thanks go to Dr. Eyal Rotenberg, with whom I had many interesting discussions and good research ideas, and Yakir Preisler, a lasting companion to the field operations. Hagay, Manuela, Ruthi and Efrat are greatly acknowledged for their help at the lab. The work would not have been possible without the assistance of many ‘secret helpers’: the undergraduate students Yoav, Gil, Ehud, Shani and Mika, and the WIS security personnel. Around the department, Ishay, Ruthi, Dalia, Ofra, Avram, Hila and Dror have always been a source of help accompanied with a smile. I express my gratitude to a number of excellent, foreign scientists with whom I had the benefit to learn from and collaborate. This includes Prof. Katarina Strelcova, Dr. Jiri Kucera, Dr. Giovanni Di Matteo, Dr. Jerome Ogee, Dr. Lisa Wingate, Prof. Nate McDowell, Dr. Christopher Reyer, Dr. Sebastian Leuzinger, Dr. Guenter Hoch and Prof. Christian Koerner. These collaborations, facilitated by meetings with the support of various EU travel grants, have undoubtedly left a prominent footprint on my research. Finally, my wife Yasmin, who always encouraged and took an interest in my work, our little Tom (who was born amidst this term), and my parents Hanna and Yoel, I thank you for everything. 2 Declaration This Thesis summarizes my own research. Dr. Eyal Rotenberg, a research associate in our group, has contributed to the discussion of results from the provenance trial (chapter 5.2) and the study of soil and tree hydraulics (chapter 5.3). Yakir Preisler, an MSc student in our group, performed the deep soil survey in Yatir forest which facilitated the root distribution analysis in the study of soil and tree hydraulics (chapter 5.3). Yakir also established and maintained the stand density experiment in Yatir forest, enabling the study of stand density effects on leaf gas exchange (chapter 5.4.9-11). The study of soil and tree hydraulics (chapter 5.3) involved additional contributors. Ella Cohen-Hilaleh, an MSc student, participated in soil sampling and analysis. The soil moisture measurement system was established by Naama Raz-Yaseef, a former PhD student. Soil moisture simulations using the model MuSICA were performed together with Dr. Fyodor Tatarinov of our group, and the model developer, Prof. Jérôme Ogée of INRA, Bordeaux, France. Dr. Giovanni Di Matteo of CRA, Rome, Italy, has participated in tree coring as part of the provenance trial (chapter 5.2). Interpretation and analysis of the data is my own. All other work presented in this Thesis is also my own. Tamir Klein Rehovot, November 2012. 3 Table of Contents 1. List of abbreviations 7 2. Abstract 8 3. Introduction 10 3.1. Forests under change 10 3.2. Water transport in trees 10 3.3. Xylem embolism and refilling 12 3.4. Forests and the hydrological cycle 13 3.5. Aleppo pine as a model forest tree 14 3.6. A forest in the dry timberline 15 3.7. Research Objectives 15 4. Materials and Methods 16 4.1. Forest sites and plant material 16 4.2. Experimental set-up and design of the greenhouse experiment 18 4.3. Experimental design of the thinning experiment 18 4.4. Tree water use 19 4.5. Xylem hydraulic conductivity and cavitation 20 4.6. Leaf gas exchange and water potential 21 4.7. Field Performance and growth phenology 21 4.8. Tree- ring δ13 C 22 4.9. Leaf chlorophyll concentration 23 4.10. Soil mechanical arrangement and water retention 23 4.11. Root depth distribution 24 4.12. Simulation of soil water content dynamics 25 4.13. Upscaling from leaf- to forest stand-scale transpiration 25 4.14. Statistical analysis 26 4 5. Results 27 5.1. Hydraulic adjustments underlying drought resistance of Pinus halepensis . Klein T, Cohen S, Yakir D. 2011. Hydraulic adjustments underlying drought resistance of Pinus halepensis. Tree Physiology 31 :637-48. 28 5.1.1. Seasonal shifts in xylem hydraulics on the diurnal scale 40 5.1.2. Comparison with previous reports of cavitation and refilling 45 5.1.3. Stomata, VPD, leaf water potential and cavitation 46 5.2. The role of trait plasticity in sustaining growth and survival of Pinus halepensis under climate change in the Mediterranean. 50 Klein T, Di Matteo G, Rotenberg E, Cohen S, Yakir D. 2012. Differential ecophysiological response of a major Mediterranean pine species across a climatic gradient. Tree Physiology , in press. 51 5.3. The role of soil and tree hydraulics in the success of a water-limited pine forest. 62 5.3.1. Soil water content dynamics and mechanical structure 62 5.3.2. Soil water retention properties 63 5.3.3. Transpirable soil water content and tree water use 65 5.3.4. Annual soil water budget is influenced by depth-dependent infiltration 69 5.3.5. Root depth distribution 70 5.3.6. Water δ18O of soil and tree sap 71 5.3.7. Relationship between soil texture and water content from ecosystem model simulations 72 5.3.8. Depth-dependent water availability 74 5.3.9. Inter-annual variations in transpirable soil water content 75 5.4. From leaf to forest scale: water use of P. halepensis and the coexisting, anisohydric Quercus calliprinos and the impact of a semi-arid pine forest on local water yield. 79 5.4.1. Stomatal regulation in P. halepensis and Q. calliprinos 79 5.4.2. Stomatal adjustments and transpiration 79 5.4.3. Seasonal gas exchange patterns of P. halepensis and Q. calliprinos 80 5.4.4. Tree-ring δ13 C 83 5 5.4.5. Stand-scale water use of P. halepensis and Q. calliprinos 84 5.4.6. Relationship between stomatal regulation and leaf gas exchange 85 5.4.7. Relationship between tree-ring δ13 C and leaf gas exchange 87 5.4.8. Water-use of Pinus halepensis and Quercus calliprinos and the forest water balance 87 5.4.9. Stand density effects on leaf-scale conditions and gas exchange rates 90 5.4.10. Upscale to stand-scale gas exchange rates using leaf area index (LAI) 90 5.4.11. The thinning bias 91 6. Discussion 93 6.1. Eco-physiology of water use in P. halepensis 93 6.2. A broader perspective 94 6.3 Implications for forest management under change 96 References 98 6 1. List of abbreviations Abbreviation Definition Units -2 -1 A Net carbon assimilation µmol CO 2 m s BAI Basal area increment cm DBH Diameter at breast height cm Es Evaporation from soil mm ET Evapotranspiration mm -2 -1 gs Leaf stomatal conductance µmol H 2O m s I Interception mm -1 -1 -1 Ks Specific hydraulic conductivity kg m MPa s 2 La Leaf area m P Annual precipitation mm PAR Photosynthetically active radiation µmol m -2 s-1 PDI Plant drought index (dimensionless) PLC Percent loss of conductivity % RH Relative humidity % SF Sap flow rate kg hr -1 SFD Sap flux density cm 3 d-1 cm -2 SWC Soil water content (volumetric) m3 m–3 -2 -1 T Leaf transpiration mmol H 2O m s Td Diurnal tree water use mm tSWC Transpirable soil water content mm Tt Tree water use mm Tu Understory water use mm VPD Vapor pressure deficit kPa -1 WUE i Intrinsic water-use efficiency µmol CO 2 mol H2O 13 13 δ C C fractionation ‰ 13 13 Δ C Discrimination against C ‰ Ψl Leaf water potential MPa Ψs Soil water potential MPa 7 2. Abstract Tree water use and transport are fundamental eco-physiological processes with major implications on phenomena of various scales. Among these are the biological mechanisms underlying tree drought resistance; the interactions between hydraulic and physiological traits; and the differences of water use with forest type, with important implications for water yield under increasing water limitations. In this study, these processes were measured in Pinus halepensis trees growing in the dry timberline, where they are regularly challenged by seasonal drought.
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