Unthinned slow-growing ponderosa pine (Pinus ponderosa) trees contain muted isotopic signals in tree rings as compared to thinned trees

Trees - Structure and Function

Volumn 28, Issue 4, 2014, Pages 1035-1051

  Sohn, Julia A ^a   Brooks, J Renée ^b   Bauhus, Jürgen ^a   Kohler, Martin ^a   Kolb, Thomas E ^c   McDowell, Nathan G ^d  

^a UNIVERSITY OF FREIBURG   (Germany)

^b U S ENVIRONMENTAL PROTECTION AGENCY   (United States)

^c NORTHERN ARIZONA UNIVERSITY   (United States)

^d LOS ALAMOS NATIONAL LABORATORY   (United States)

Author keywords

Gas exchange; Oxygen isotopes; Pinus ponderosa (ponderosa pine); Sensitivity analysis; Thinning

Indexed keywords

CARBON; ISOTOPES; OXYGEN; PHYSIOLOGICAL MODELS; PHYSIOLOGY; PROCESS CONTROL; SENSITIVITY ANALYSIS; WOOD;

CARBON ISOTOPE DISCRIMINATION; ENVIRONMENTAL CONDITIONS; GAS EXCHANGE; INTERANNUAL VARIABILITY; OXYGEN ISOTOPES; PHYSIOLOGICAL PROCESS; PONDEROSA PINES; THINNING;

FORESTRY;

GAS; PHYSIOLOGY; PINUS PONDEROSA; SENSITIVITY; SIGNALS;

PINUS PONDEROSA;

EID: 84926686506     PISSN: 09311890     EISSN: None     Source Type: Journal    
DOI: 10.1007/s00468-014-1016-z     Document Type: Article

References (114)

1. Adams HD, Luce CH, Breshears DD, Allen CD, Weiler M, Hale VC, Smith AMS, Huxman TE (2012) Ecohydrological consequences of drought- and infestation- triggered tree die-off: insights and hypotheses. Ecohydrology 5:145–159
2. Allen CD, Breshears DD (1998) Drought-induced shift of a forest-woodland ecotone: rapid landscape response to climate variation. Pro Natl Acad Sci USA 95:14839–14842
3. Allen CD, Macalady AK, Chenchouni H, Bachelet D, McDowell N et al (2010) A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. For Ecol Manag 259:660–684
4. Aussenac G, Granier A (1988) Effects of thinning on water stress and growth in Douglas-fir. Can J For Res 18:100–105. doi:10.1139/x88-015
5. Ayub G, Smith RA, Tissue DT, Atkin OK (2011) Impacts of drought on leaf respiration in darkness and light in Eucalyptus saligna exposed to industrial-age atmospheric CO2 and growth temperature. New Phytol 190:1003–1018
6. Barbour MM (2007) Stable oxygen isotope composition of plant tissue: a review. Funct Plant Biol 34:83–94. doi:10.1071/fp06228
7. Barbour MM, Farquhar GD (2000) Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves. Plant Cell Environ 23:473–485
8. Barbour MM, Schurr U, Henry BK et al (2000) Variation in the oxygen isotope ratio of phloem sap sucrose from castor bean. Evidence in support of the Peclet effect. Plant Physiol 123:671–679
9. Barbour MM, Andrews TJ, Farquhar GD (2001) Correlations between oxygen isotope ratios of wood constituents of Quercus and Pinus samples from around the world. Funct Plant Biol 28:335–348
10. Barbour MM, Walcroft AS, Farquhar GD (2002) Seasonal variation in delta C-13 and delta O-18 of cellulose from growth rings of Pinus radiata. Plant Cell Environ 25:1483–1499
11. Barnard HR, Brooks JR, Bond BJ (2012) Applying the dual-isotope conceptual model to interpret physiological trends under uncontrolled conditions. Tree Physiol 32:1183–1198. doi:10.1093/treephys/tps078
12. Biondi F (1996) Decadal-scale dynamics at the Gus Pearson Natural Areas: evidence for inverse (a)symmetric competition? Can J For Res 26:1397–1406. doi:10.1139/x26-156
13. Brandes E, Kodama N, Whittaker K et al (2006) Short-term variation in the isotopic composition of organic matter allocated from the leaves to the stem of Pinus sylvestris: effects of photosynthetic and postphotosynthetic carbon isotope fractionation. Glob Change Biol 12:1922–1939. doi:10.1111/j.1365-2486.2006.01205.x
14. Breda N, Granier A, Aussenac G (1995) Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus-Petraea (Matt) Liebl). Tree Physiol 15:295–306
15. Breshears DD, Allen CD (2002) The importance of rapid, disturbance-induced losses in carbon management and sequestration: rapid, disturbance-induced C losses. Glob Ecol Biogeogr 11:1–5. doi:10.1046/j.1466-822X.2002.00274.x
16. Breshears DD, Cobb NS, Rich PM et al (2005) Regional vegetation die-off in response to global-change-type drought. Proc Natl Acad Sci 102:15144–15148. doi:10.1073/pnas.0505734102
17. Brooks JR, Coulombe R (2009) Physiological responses to fertilization recorded in tree rings: isotopic lessons from a long-term fertilization trial. Ecol Appl 19:1044–1060
18. Brooks JR, Mitchell AK (2011) Interpreting tree responses to thinning and fertilization using tree-ring stable isotopes. New Phytol 190:770–782. doi:10.1111/j.1469-8137.2010.03627.x
19. Burke EJ, Brown SJ, Christidis N (2006) Modeling the recent evolution of global drought and projections for the twenty-first century with the Hadley centre climate model. J Hydrometeorol 7:1113–1125. doi:10.1175/JHM544.1
20. Cannell MG, Dewar RC (1994) Carbon allocation in trees: a review of concepts for modelling. Adv Ecol Res 25:59–104
21. Cescatti A, Piutti E (1998) Silvicultural alternatives, competition regime and sensitivity to climate in a European beech forest. For Ecol Manag 102:213–223
22. Ciais P, Reichstein M, Viovy N et al (2005) Europe-wide reduction in primary productivity caused by the heat and drought in 2003. Nature 437:529–533. doi:10.1038/nature03972
23. Covington WW, Fule PZ, Moore MM et al (1997) Restoring ecosystem health in ponderosa pine forests of the southwest. J Forest 95:23–29
24. Craig H, Gordon LI (1965) Deuterium and oxygen 18 variations in the ocean and the marine atmosphere. In: Tongiorgi E (ed) Proceedings of a Conference on Stable Isotopes in Oceano-graphic Studies and Palaeotemperatures. Lischi and Figli, Pisa, Italy, pp 9–130
25. DeNiro MJ, Epstein S (1981) Isotopic composition of cellulose from aquatic organisms. Geochim Cosmochim Acta 45:1885–1894. doi:10.1016/0016-7037(81)90018-1
26. Dore S, Montes-Helu M, Hart SC et al (2012) Recovery of ponderosa pine ecosystem carbon and water fluxes from thinning and stand-replacing fire. Glob Change Biol 18:3171–3185. doi:10.1111/j.1365-2486.2012.02775.x
27. English NB, McDowell NG, Allen CD, Mora C (2011) The effects of a-cellulose extraction and blue-stain fungus on retrospective studies of carbon and oxygen isotope variation in live and dead trees: stable isotopes in tree-ring wood and cellulose of live and dead trees. Rapid Commun Mass Spectrom 25:3083–3090. doi:10.1002/rcm.5192
28. Farquhar GD, Lloyd J (1993) Carbon and oxygen isotope effects in the exchange of carbon dioxide between terrestrial plants and the atmosphere. In: Stable Isot. Plant Carbon-Water Relat. Elsevier, Amsterdam, pp 47–70
29. Farquhar GD, Ehleringer JR, Hubick KT (1989) Carbon isotope discrimination and photosynthesis. Annu Rev Plant Physiol Plant Mol Biol 40:503–537
30. Farquhar GD, Barbour MM, Henry BK (1998) Interpretation of oxygen isotope composition of leaf material. In: Griffiths H (ed) Stable isotopes: integration of biological, ecological and geo-chemical processes. BIOS Scientific Publishers, Oxford, pp 27–61
31. Feeney SR, Kolb TE, Covington WW, Wagner MR (1998) Influence of thinning and burning restoration treatments on presettlement ponderosa pines at the Gus Pearson Natural Area. Can J For Res Rev Can Rech For 28:1295–1306
32. Fettig CJ, Klepzig KD, Billings RF et al (2007) The effectiveness of vegetation management practices for prevention and control of bark beetle infestations in coniferous forests of the western and southern United States. For Ecol Manag 238:24–53. doi:10.1016/j.foreco.2006.10.011
33. Fiedler CE, Arno SF, Harrington MG (1996) Flexible silvicultural and prescribed burning approaches for improving health of ponder-osa pine forests. US For Serv Gen Tec. Rep RMRS, 278, pp 69–74
34. Flanagan LB, Ehleringer JR (1991) Stable isotope composition of stem and leaf water: applications to the study of plant water use. Funct Ecol 5:270–277
35. Francey RJ, Farquhar GD (1982) An explanation of 13C/12C variations in tree rings. Nature 297:28–31. doi:10.1038/297028a0
36. Fritts HC (1976) Tree rings and climate. Academic Press, London, New York, San Francisco
37. Galvez DA, Landhausser SM, Tyree MT (2011) Root carbon reserve dynamics in aspen seedlings: does simulated drought induce reserve limitation? Tree Physiol 31:250–257. doi:10.1093/treephys/tpr012
38. Gaylord ML, Kolb TE, Wallin KF, Wagner MR (2007) Seasonal dynamics of tree growth, physiology, and resin defenses in a northern Arizona ponderosa pine forest. Can J For Res 37:1173–1183. doi:10.1139/X06-309
39. Gessler A, Brandes E, Buchmann N et al (2009) Tracing carbon and oxygen isotope signals from newly assimilated sugars in the leaves to the tree-ring archive. Plant Cell Environ 32:780–795. doi:10.1111/j.1365-3040.2009.01957.x
40. Gessler A, Brandes E, Keitel C et al (2013) The oxygen isotope enrichment of leaf-exported assimilates—does it always reflect lamina leaf water enrichment? New Phytol 200:144–157. doi:10.1111/nph.12359
41. Gleixner G, Scrimgeour C, Schmidt H-L, Viola R (1998) Stable isotope distribution in the major metabolites of source and sink organs of Solanum tuberosum L.: a powerful tool in the study of metabolic partitioning in intact plants. Planta 207:241–245. doi:10.1007/s004250050479
42. Helle G, Schleser GH (2004) Beyond CO2-fixation by Rubisco—an interpretation of 13C/12C variations in tree rings from novel intra-seasonal studies on broad-leaf trees. Plant Cell Environ 27:367–380
43. Hill SA, Waterhouse JS, Field EM et al (1995) Rapid recycling of triose phosphates in oak stem tissue. Plant Cell Environ 18:931–936
44. Hogg EH, Brandt JP, Michaelian M (2008) Impacts of a regional drought on the productivity, dieback, and biomass of western Canadian aspen forests. Can J For Res 38:1373–1384. doi:10.1139/X08-001
45. Hsiao TC, Acevedo E (1974) Plant responses to water deficits, water-use efficiency, and drought resistance. Agric Meteorol 14:59–84
46. Huntington TG (2006) Evidence for intensification of the global water cycle: review and synthesis. J Hydrol 319:83–95. doi:10.1016/j.jhydrol.2005.07.003
47. IPCC (2007) Climate change 2007: synthesis report. Contribution of working groups I, II and III to the fourth assessment report of the intergovernmental panel on climate change
48. Kagawa A, Sugimoto A, Maximov TC (2006) Seasonal course of translocation, storage and remobilization of 13C pulse-labeled photoassimilate in naturally growing Larix gmelinii saplings. New Phytol 171:793–804. doi:10.1111/j.1469-8137.2006.01780.x
49. Kahmen A, Simonin K, Tu K et al (2009) The influence of species and growing conditions on the 18-O enrichment of leaf water and its impact on ‘‘effective path length’’. New Phytol 184:619–630. doi:10.1111/j.1469-8137.2009.03008.x
50. Kahmen A, Sachse D, Arndt SK et al (2011) Cellulose 18O is an index of leaf-to-air vapor pressure difference (VPD) in tropical plants. Proc Natl Acad Sci 108:1981–1986. doi:10.1073/pnas.1018906108
51. Keel SG, Siegwolf RTW, Körner C (2006) Canopy CO2 enrichment permits tracing the fate of recently assimilated carbon in a mature deciduous forest. New Phytol 172:319–329. doi:10.1111/j.1469-8137.2006.01831.x
52. Kerhoulas LP, Kolb TE, Koch GW (2013) Tree size, stand density, and the source of water used across seasons by ponderosa pine in northern Arizona. For Ecol Manag 289:425–433. doi:10.1016/j.foreco.2012.10.036
53. Koepke DF, Kolb TE (2013) Species variation in water relations and xylem vulnerability to cavitation at a forest-woodland ecotone. Forensic Sci 59:524–535. doi:10.5849/forsci.12-053
54. Koepke DF, Kolb TE, Adams HD (2010) Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone. Oeco-logia 163:1079–1090. doi:10.1007/s00442-010-1671-8
55. Kolb TE, Stone JE (2000) Differences in leaf gas exchange and water relations among species and tree sizes in an Arizona pine-oak forest. Tree Physiol 20:1–12
56. Kolb TE, Holmberg KM, Wagner MR, Stone JE (1998) Regulation of ponderosa pine foliar physiology and insect resistance mechanisms by basal area treatments. Tree Physiol 18:375–381
57. Kolb TE, Agee JK, Fule PZ, McDowell NG, Pearson K, Sala A, Waring RH (2007) Perpetuating old ponderosa pine. For Ecol Manag 249:141–157
58. Körner C (2003) Carbon limitation in trees. J Ecol 91:4–17
59. Kriedemann PE (1986) Stomatal and photosynthetic limitations to leaf growth. Funct Plant Biol 13:15–31
60. Kuptz D, Fleischmann F, Matyssek R, Grams TEE (2011) Seasonal patterns of carbon allocation to respiratory pools in 60-yr-old deciduous (Fagus sylvatica) and evergreen (Picea abies) trees assessed via whole-tree stable carbon isotope labeling. New Phytol 191:160–172. doi:10.1111/j.1469-8137.2011.03676.x
61. Kurz WA, Dymond CC, Stinson G et al (2008) Mountain pine beetle and forest carbon feedback to climate change. Nature 452:987–990. doi:10.1038/nature06777
62. Latham P, Tappeiner J (2002) Response of old-growth conifers to reduction in stand density in western Oregon forests. Tree Physiol 22:137–146
63. Laurent M, Antoine N, Joel G (2003) Effects of different thinning intensities on drought response in Norway spruce (Picea abies (L.) Karst.). For Ecol Manag 183:47–60. doi:10.1016/s0378-1127(03)00098-7
64. Leavitt SW (1993) Seasonal 13 C/12 C changes in tree rings: species and site coherence, and a possible drought influence. Can J For Res 23:210–218. doi:10.1139/x93-028
65. Legoff N, Ottorini JM (1993) Thinning and climate effects on growth of beech (Fagus-sylvatica L) in experimental stands. For Ecol Manag 62:1–14
66. Ma S, Concilio A, Oakley B et al (2010) Spatial variability in microclimate in a mixed-conifer forest before and after thinning and burning treatments. For Ecol Manag 259:904–915. doi:10.1016/j.foreco.2009.11.030
67. Marshall JD, Monserud RA (1996) Homeostatic gas-exchange parameters inferred from 13C/12C in tree rings of conifers. Oecologia 105:13–21. doi:10.1007/BF00328786
68. Martin-Benito D, Del Rio M, Heinrich I et al (2010) Response of climate-growth relationships and water use efficiency to thinning in a Pinus nigra afforestation. For Ecol Manag 259:967–975. doi:10.1016/j.foreco.2009.12.001
69. Martinez-Vilalta J, Sala A, Pinol J (2004) The hydraulic architecture of Pinaceae—a review. Plant Ecol 171:3–13
70. McDowell N, Brooks JR, Fitzgerald SA, Bond BJ (2003) Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions. Plant Cell Environ 26:631–644
71. McDowell NG, Adams HD, Bailey JD et al (2006) Homeostatic maintenance of ponderosa pine gas exchange in response to stand density changes. Ecol Appl 16:1164–1182
72. McDowell NG, Adams HD, Bailey JD, Kolb TE (2007) The role of stand density on growth efficiency, leaf area index, and resin flow in southwestern ponderosa pine forests. Can J For Res Rev Can Rech For 37:343–355. doi:10.1139/x06-233
73. McDowell NG, White S, Pockman WT (2008) Transpiration and stomatal conductance across a steep climate gradient in the southern Rocky Mountains. Ecohydrology 1:193–204. doi:10.1002/eco.20
74. McDowell NG, Allen CD, Marshall L (2010) Growth, carbon-isotope discrimination, and drought-associated mortality across a Pinus ponderosa elevational transect. Glob Change Biol 16:399–415. doi:10.1111/j.1365-2486.2009.01994.x
75. McDowell NG, Bond BJ, Dickman LT et al (2011) Relationships between tree height and carbon isotope discrimination. In: Meinzer FC, Lachenbruch B, Dawson TE (eds) Size- Age-Relat. Chang. Tree Struct. Funct. Springer, Dordrecht, pp 255–286
76. Michaelian M, Hogg EH, Hall RJ, Arsenault E (2011) Massive mortality of aspen following severe drought along the southern edge of the Canadian boreal forest: aspen mortality following severe drought. Glob Change Biol 17:2084–2094. doi:10.1111/j.1365-2486.2010.02357.x
77. Mitchell PJ, O’Grady AP, Tissue DT, White DA, Ottenschlaeger ML, Pinkard EA (2013) Drought response strategies define the relative contributions of hydraulic dysfunction and carbohydrate depletion during tree mortality. New Phytol 197:862–872
78. Moore MM, Casey CA, Bakker JD et al (2006) Herbaceous vegetation responses (1992–2004) to restoration treatments in a ponderosa pine forest. Rangel Ecol Manag 59:135–144. doi:10.2111/05-051R2.1
79. Moreno-Gutiérrez C, Barberá GG, Nicolás E et al (2011) Leaf d18O of remaining trees is affected by thinning intensity in a semiarid pine forest: thinning intensity influences tree water status. Plant Cell Environ 34:1009–1019. doi:10.1111/j.1365-3040.2011.02300.x
80. Negron JF, McMillin JD, Anhold JA, Coulson D (2009) Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona, USA. For Ecol Manag 257:1353–1362
81. Offermann C, Ferrio JP, Holst J et al (2011) The long way down—are carbon and oxygen isotope signals in the tree ring uncoupled from canopy physiological processes? Tree Physiol 31:1088–1102. doi:10.1093/treephys/tpr093
82. Parsons DJ, DeBenedetti SH (1979) Impact of fire suppression on a mixed-conifer forest. For Ecol Manag 2:21–33. doi:10.1016/0378-1127(79)90034-3
83. Peterson DL, Johnson MC, Agee JK, Jain TB, McKenzie D, Reinhardt ED (2005) Forest structure and fire hazard in the western United States. USDA For Serv Gen Tech Re. PNW-GTR-628
84. Phillips OL, Aragao LEOC, Lewis SL et al (2009) Drought sensitivity of the Amazon rainforest. Science 323:1344–1347. doi:10.1126/science.1164033
85. Pinol J, Sala A (2000) Ecological implications of xylem cavitation for several Pinaceae in the Pacific Northern USA. Funct Ecol 14:538–545. doi:10.1046/j.1365-2435.2000.00451.x
86. Powers MD, Pregitzer KS, Palik BJ, Webster CR (2009) Wood delta C-13, delta O-18 and radial growth responses of residual red pine to variable retention harvesting. Tree Physiol 30:326–334. doi:10.1093/treephys/tpp119
87. Rambo TR, North MP (2009) Canopy microclimate response to pattern and density of thinning in a Sierra Nevada forest. For Ecol Manag 257:435–442. doi:10.1016/j.foreco.2008.09.029
88. Richardson AD, Carbone MS, Keenan TF et al (2013) Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees. New Phytol 197:850–861. doi:10.1111/nph.12042
89. Roden JS, Ehleringer JR (2000) Hydrogen and oxygen isotope ratios of tree ring cellulose for field-grown riparian trees. Oecologia 123:481–489
90. Roden JS, Farquhar GD (2012) A controlled test of the dual-isotope approach for the interpretation of stable carbon and oxygen isotope ratio variation in tree rings. Tree Physiol 32:490–503. doi:10.1093/treephys/tps019
91. Roden J, Siegwolf R (2012) Is the dual-isotope conceptual model fully operational? Tree Physiol 32:1179–1182. doi:10.1093/treephys/tps099
92. Roden JS, Lin G, Ehleringer JR (2000) A mechanistic model for interpretation of hydrogen and oxygen isotope ratios in tree-ring cellulose. Geochim Cosmochim Acta 64:21–35. doi:10.1016/S0016-7037(99)00195-7
93. Ronco F, Edmister CB, Trujillo DB (1985) Growth of ponderosa pine thinned to different stocking levels in northern Arizona. USDA For Serv Res Pap RM-262, p 15
94. Sala A, Hoch G (2009) Height-related growth declines in ponderosa pine are not due to carbon limitation. Plant Cell Environ 32:22–30. doi:10.1111/j.1365-3040.2008.01896.x
95. Saurer M, Aellen K, Siegwolf R (1997) Correlating delta C-13 and delta O-18 in cellulose of trees. Plant Cell Environ 20:1543–1550
96. Scheidegger Y, Saurer M, Bahn M, Siegwolf R (2000) Linking stable oxygen and carbon isotopes with stomatal conductance and photosynthetic capacity: a conceptual model. Oecologia 125:350–357
97. Schwalm CR, Williams CA, Schaefer K et al (2012) Reduction in carbon uptake during turn of the century drought in western North America. Nat Geosci 5:551–556. doi:10.1038/ngeo1529
98. Seibt U, Rajabi A, Griffiths H, Berry JA (2008) Carbon isotopes and water use efficiency: sense and sensitivity. Oecologia 155:441–454. doi:10.1007/s00442-007-0932-7
99. Skov KR, Kolb TE, Wallin KF (2004) Tree size and drought affect ponderosa pine physiological response to thinning and burning treatments. For Sci 50:81–91
100. Skov KR, Kolb TE, Wallin KF (2005) Difference in radial growth response to restoration thinning and burning treatments between young and old ponderosa pine in Arizona. Western J Appl For 20:36–43
101. Sohn JA, Kohler M, Gessler A, Bauhus J (2012) Interactions of thinning and stem height on the drought response of radial stem growth and isotopic composition of Norway spruce (Picea abies). Tree Physiol 32:1199–1213
102. Sohn JA, Gebhardt T, Ammer C et al (2013) Mitigation of drought by thinning: short-term and long-term effects on growth and physiological performance of Norway spruce (Picea abies). For Ecol Manag 308:188–197. doi:10.1016/j.foreco.2013.07.048
103. Song X, Barbour MM, Farquhar GD et al (2013) Transpiration rate relates to within- and across-species variations in effective path length in a leaf water model of oxygen isotope enrichment: transpiration-related variation in Péclet path length. Plant Cell Environ 36:1338–1351. doi:10.1111/pce.12063
104. Sternberg L, DeNiro M (1983) Biogeochemical implications of the isotopic equilibrium fractionation factor between oxygen atoms of acetone and water. Geochim et Cosmochim Acta 47: 2271–2274
105. Sternberg LDSL, Deniro MJ, Savidge RA (1986) Oxygen isotope exchange between metabolites and water during biochemical reactions leading to cellulose synthesis. Plant Physiol 82:423–427. doi:10.1104/pp.82.2.423
106. Tissue DT, Wright SJ (1995) Effect of seasonal water availability on phenology and the annual shoot carbohydrate cycle of tropical forest shrubs. Funct Ecol 9:518–527
107. Van Mantgem PJ, Stephenson NL, Byrne JC et al (2009) Widespread increase of tree mortality rates in the Western United States. Science 323:521–524. doi:10.1126/science.1165000
108. Walcroft AS, Silvester WB, Whitehead D, Kelliher FM (1997) Seasonal changes in stable carbon isotope ratios within annual rings of Pinus radiata reflect environmental regulation of growth processes. Aust J Plant Physiol 24:57–68
109. Williams MA, Baker WL (2012) Comparison of the higher-severity fire regime in historical (A.D. 1800s) and modern (A.D. 1984–2009) Montane forests across 624,156 ha of the Colorado front range. Ecosystems 15:832–847. doi:10.1007/s10021-012-9549-8
110. Wollum AG, Schubert GH (1975) Effect of thinning on the foliage and forest floor properties of ponderosa pine stands. Soil Sci Soc Am Proc 39:968–972
111. Woodruff DR, Meinzer FC (2011) Size-dependent changes in biophysical control of tree growth: the role of turgor. In: Size-and age-related changes in tree structure and function. Springer, Netherlands, pp 363–384
112. Würth MKR, Pelez-Riedl S, Wright SJ, Körner C (2005) Non-structural carbohydrate pools in a tropical forest. Oecologia 143:11–24. doi:10.1007/s00442-004-1773-2
113. Zausen GL, Kolb TE, Bailey JD, Wagner MR (2005) Long-term impacts of stand management on ponderosa pine physiology and bark beetle abundance in northern Arizona: a replicated landscape study. For Ecol Manag 218:291–305. doi:10.1016/j.foreco.2005.08.023
114. Zhang JW, Feng Z, Cregg BM, Schumann CM (1997) Carbon isotopic composition, gas exchange, and growth of three populations of ponderosa pine differing in drought tolerance. Tree Physiol 17:461–466