Evidence supporting hydraulic limitation hypothesis was found using foliar δ13C in combination with nitrogen content per unit leaf area and statistical partitioning for three conifer species.
One theory behind the productivity decline of mature forests is the hydraulic limitation hypothesis (HLH); leaf-level gas exchange is reduced with increasing forest canopy height via increased hydraulic resistance in the xylem pathway, which in turn limits photosynthesis via stomatal regulation. Foliar \(\delta\)13C can be used to assess the HLH as it reflects the history of leaf-level gas exchange. However, this method should be used with caution as co-varying factors, including light levels and foliar nutrient status, can also influence foliar \(\delta\)13C. We explore the potential use of foliar \(\delta\)13C to assess leaf-level hydraulic limitation using three coniferous species across three height classes (short, intermediate and tall) in northern Idaho, USA. Foliar samples were collected from multiple canopy locations varying in height from each height class to measure \(\delta\)13C of bulk foliar materials as well as sugar and starch extracted from the samples. We also quantified nitrogen content per unit leaf area (Narea) as an integrated measure of nutrient status and light environment of a given foliar sample, which can partly account for various non-stomatal limitations for photosynthesis, and thus affect foliar \(\delta\)13C. Using sequential ANOVAs, we tested the hypothesis that foliar \(\delta\)13C variation was attributable to foliar sample height changes after accounting for Narea. The hypothesis was supported by analyses using foliage samples from the top canopy location across the three height classes for each conifer species, especially for bulk foliage and extracted sugar. In conclusion, we found evidence supporting the HLH using foliar \(\delta\)13C from three conifer species.
This is a preview of subscription content, access via your institution.
Buy single article
Instant access to the full article PDF.
Tax calculation will be finalised during checkout.
Subscribe to journal
Immediate online access to all issues from 2019. Subscription will auto renew annually.
Tax calculation will be finalised during checkout.
Availability of data and material
All the data used in this study can be found as an electronic supplemental material.
Adams HD, Germino MJ, Breshears DD, Barron-Gafford GA, Guardiola-Claramonte M, Zou CB, Huxman TE (2013) Nonstructural leaf carbohydrate dynamics of Pinus edulis during drought-induced tree mortality reveal role for carbon metabolism in mortality mechanism. New Phytol 197:1142–1151
Ambrose AR, Sillett SC, Dawson TE (2009) Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods. Plant Cell Environ 32:743–757
Bachofen C, D’Odorico P, Buchmann N (2020) Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir. Oecologia 192:323–339
Badeck FW, Tcherkez G, Nogues S, Piel C, Ghashghaie J (2005) Post-photosynthetic fractionation of stable carbon isotopes between plant organs—a widespread phenomenon. Rapid Commun Mass Spectrom 19:1381–1391
Baker KV, Tai X, Miller ML, Johnson DM (2019) Six co-occurring conifer species in northern Idaho exhibit a continuum of hydraulic strategies during an extreme drought year. AoB Plants 11:plz056
Baret M, Pepin S, Ward C, Pothier D (2015) Long-term changes in belowground and aboveground resource allocation of boreal forest stands. For Ecol Manag 350:62–69
Barnard HR, Ryan M (2003) A test of the hydraulic limitation hypothesis in fast-growing Eucalyptus saligna. Plant Cell Environ 26:1235–1245
Bates D, Maechler M, Bolker B, Walker S (2014) lme4: Linear mixed-effects models using Eigen and S4. R package version. 1:1-23. Accessed 2 May 2020
Berry SC, Varney GT, Flanagan LB (1997) Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients. Oecologia 109:499–506
Blessing CH, Werner RA, Siegwolf R, Buchmann N (2015) Allocation dynamics of recently fixed carbon in beech saplings in response to increased temperatures and drought. Tree Physiol 35:585–598
Bown HE, Watt MS, Mason EG, Clinton PW, Whitehead D (2009) The influence of nitrogen and phosphorus supply and genotype on mesophyll conductance limitations to photosynthesis in Pinus radiata. Tree Physiol 29:1143–1151
Brienen R, Gloor E, Clerici S, Newton R, Arppe L, Boom A, Bottrell S, Callaghan M, Heaton T, Helama S (2017) Tree height strongly affects estimates of water-use efficiency responses to climate and CO2 using isotopes. Nat Commun 8:1–10
Brugnoli E, Hubick KT, von Caemmerer S, Wong SC, Farquhar GD (1988) Correlation between the carbon isotope discrimination in leaf starch and sugars of C3 plants and the ratio of intercellular and atmospheric partial pressures of carbon dioxide. Plant Physiol 88:1418–1424
Brugnoli E, Lauteri M, Guido M (1994) Carbon isotope discrimination and photosynthesis: response and adaptation to environmental stress. In: de Kouchkovsky Y, Larher F (eds) Plant sciences. Second General Colloquium on Plant Sciences Universite de Renners, Renners, pp 269–272
Brugnoli E, Scartazza A, Lauteri M, Monteverdi M, Máguas C (1998) Carbon isotope discrimination in structural and non-structural carbohydrates in relation to productivity and adaptation to unfavourable conditions. In: Friffiths H (ed) Stable isotopes. Integration of biological, ecological geochemical processes. BIOS Scientific Publishers, Oxford, pp 133–144
Buchmann N, Kao W-Y, Ehleringer J (1997) Influence of stand structure on carbon-13 of vegetation, soils, and canopy air within deciduous and evergreen forests in Utah, United States. Oecologia 110: 109–119Buckley TN (2005) The control of stomata by water balance. New Phytol 168:275–292
Buckley TN (2005) The control of stomata by water balance. New Phytologist. 168:275-292.
Chatterton NJ, Silvius JE (1981) Photosynthate partitioning into starch in soybean leaves: II. Irradiance level and daily photosynthetic period duration effects. Plant Physiol 67:257–260
Chen HY (1997) Interspecific responses of planted seedlings to light availability in interior British Columbia: survival, growth, allometric patterns, and specific leaf area. Can J For Res 27:1383–1393
Chen HY, Klinka K, Kayahara GJ (1996) Effects of light on growth, crown architecture, and specific leaf area for naturally established Pinus contorta var. latifolia and Pseudotsuga menziesii var. glauca saplings. Can J For Res 26:1149–1157
Coble AP, Fogel ML, Parker GG (2017) Canopy gradients in leaf functional traits for species that differ in growth strategies and shade tolerance. Tree Physiol 37:1415–1425
Damesin C, Lelarge C (2003) Carbon isotope composition of current-year shoots from Fagus sylvatica in relation to growth, respiration and use of reserves. Plant Cell Environ 26:207–219
Dawson TE (1996) Determining water use by trees and forests from isotopic, energy balance and transpiration analyses: the roles of tree size and hydraulic lift. Tree Physiol 16:263–272
Dewar R, Mauranen A, Mäkelä A, Hölttä T, Medlyn B, Vesala T (2018) New insights into the covariation of stomatal, mesophyll and hydraulic conductances from optimization models incorporating nonstomatal limitations to photosynthesis. New Phytol 217:571–585
Dietze MC, Sala A, Carbone MS, Czimczik CI, Mantooth JA, Richardson AD, Vargas R (2014) Nonstructural carbon in woody plants. Annu Rev Plant Biol 65:667–687
Douthe C, Dreyer E, Brendel O, Warren CR (2012) Is mesophyll conductance to CO2 in leaves of three Eucalyptus species sensitive to short-term changes of irradiance under ambient as well as low O2? Funct Plant Biol 39:435–448
Duursma R, Marshall J (2006) Vertical canopy gradients in δ13C correspond with leaf nitrogen content in a mixed-species conifer forest. Trees 20:496–506
Ellsworth D, Reich PB (1993) Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest. Oecologia 96:169–178
Finklin AI (1983) Climate of Priest River experimental forest, northern Idaho. US Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station, General Technical Report INT-159
Flanagan LB, Brooks JR, Varney GT, Berry SC, Ehleringer JR (1996) Carbon isotope discrimination during photosynthesis and the isotope ratio of respired CO2 in boreal forest ecosystems. Global Biogeochem Cy 10:629–640
Flexas J, Medrano H (2002) Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited. Ann Bot 89:183–189
Flexas J, Ribas-Carbo M, Diaz-Espejo A, GalmES J, Medrano H (2008) Mesophyll conductance to CO2: current knowledge and future prospects. Plant Cell Environ 31:602–621
Frank DC, Poulter B, Saurer M, Esper J, Huntingford C, Helle G, Treydte K, Zimmermann NE, Schleser GH, Ahlström A, Ciais P, Friedlingstein P, Levis S, Lomas M, Sitch S, Viovy N, Andreu-Hayles L, Bednarz Z, Berninger F, Boettger T, D‘Alessandro CM, Daux V, Filot M, Grabner M, Gutierrez E, Haupt M, Hilasvuori E, Jungner H, Kalela-Brundin M, Krapiec M, Leuenberger M, Loader NJ, Marah H, Masson-Delmotte V, Pazdur A, Pawelczyk S, Pierre M, Planells O, Pukiene R, Reynolds-Henne CE, Rinne KT, Saracino A, Sonninen E, Stievenard M, Switsur VR, Szczepanek M, Szychowska-Krapiec E, Todaro L, Waterhouse JS, Weigl M, (2015) Water-use efficiency and transpiration across European forests during the Anthropocene. Nat Clim Change 5:579–583
Galiano L, Timofeeva G, Saurer M, Siegwolf R, Martínez-Vilalta J, Hommel R, Gessler A (2017) The fate of recently fixed carbon after drought release: towards unravelling C storage regulation in Tilia platyphyllos and Pinus sylvestris. Plant Cell Environ 40:1711–1724
Gessler A, Ferrio JP, Hommel R, Treydte K, Werner RA, Monson RK (2014) Stable isotopes in tree rings: towards a mechanistic understanding of isotope fractionation and mixing processes from the leaves to the wood. Tree Physiol 34:796–818
Gessler A, Keitel C, Kodama N, Weston C, Winters AJ, Keith H, Grice K, Leuning R, Farquhar GD (2007) δ13C of organic matter transported from the leaves to the roots in Eucalyptus delegatensis: short-term variations and relation to respired CO2. Funct Plant Biol 34:692–706
Gleixner G, Schmidt H-L (1997) Carbon isotope effects on the fructose-1, 6-bisphosphate aldolase reaction, origin for non-statistical 13C distributions in carbohydrates. J Biol Chem 272:5382–5387
Göttlicher S, Knohl A, Wanek W, Buchmann N, Richter A (2006) Short-term changes in carbon isotope composition of soluble carbohydrates and starch: from canopy leaves to the root system. Rapid Commun Mass Spectrom 20:653–660
Grassi G, Magnani F (2005) Stomatal, mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees. Plant Cell Environ 28:834–849
Grossiord C, Buckley TN, Cernusak LA, Novick KA, Poulter B, Siegwolf RT, Sperry JS, McDowell NG (2020) Plant responses to rising vapor pressure deficit. New Phytol 226:1550–1566
Hanba YT, Mori S, Lei TT, Koike T, Wada E (1997) Variations in leaf δ13C along a vertical profile of irradiance in a temperate Japanese forest. Oecologia 110:253–261
Harwood KG, Gillon JS, Roberts A, Griffiths H (1999) Determinants of isotopic coupling of CO2 and wter vapour within a Quercus petraea forest canopy. Oecologia 119:109–119
Hultine K, Marshall J (2000) Altitude trends in conifer leaf morphology and stable carbon isotope composition. Oecologia 123:32–40
Irvine J, Law B, Anthoni P, Meinzer F (2002) Water limitations to carbon exchange in old-growth and young ponderosa pine stands. Tree Physiol 22:189–196
Irvine J, Law B, Kurpius M, Anthoni P, Moore D, Schwarz P (2004) Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine. Tree Physiol 24:753–763
Jansen E, Overpeck J, Briffa KR, Duplessy J-C, Joos F, Masson-Delmotte V, Olago D, Otto-Bliesner B, Peltier WT, Rahmstorf S, Ramesh R, Raynaud D, Rind D, Solomina O, Villalba R, Zhang D (2007) Palaeoclimate. In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt KB, Tignor M, Miller HL (eds) Climate Change 2007: the physical science basis. Contribution of Working Group I to the fourth assessment report of the intergovernmental panel on climate change. Cambridge University Press, Cambridge
Koch GW, Sillett SC, Jennings GM, Davis SD (2004) The limits to tree height. Nature 428:851–854
Kumagai T, Ichie T, Yoshimura M, Yamashita M, Kenzo T, Saitoh TM, Ohashi M, Suzuki M, Koike T, Komatsu H (2006) Modeling CO2 exchange over a Bornean tropical rain forest using measured vertical and horizontal variations in leaf‐level physiological parameters and leaf area densities. J. Geophys. Res., 111, D10107, https://doi.org/10.1029/2005JD006676.
Lavergne A, Graven H, De Kauwe MG, Keenan TF, Medlyn BE, Prentice IC (2019) Observed and modelled historical trends in the water-use efficiency of plants and ecosystems. Global Change Biol 25:2242–2257
Lawlor DW, Cornic G (2002) Photosynthetic carbon assimilation and associated metabolism in relation to water deficits in higher plants. Plant Cell Environ 25:275–294
Lawlor DW, Tezara W (2009) Causes of decreased photosynthetic rate and metabolic capacity in water-deficient leaf cells: A critical evaluation of mechanisms and integration of processes. Ann Bot 103:561–579
Le Roux X, Bariac T, Sinoquet H, Genty B, Piel C, Mariotti A, Girardin C, Richard P (2001) Spatial distribution of leaf water-use efficiency and carbon isotope discrimination within an isolated tree crown. Plant Cell Environ 24:1021–1032
Li W, Hartmann H, Adams HD, Zhang H, Jin C, Zhao C, Guan D, Wang A, Yuan F, Wu J (2018) The sweet side of global change–dynamic responses of non-structural carbohydrates to drought, elevated CO2 and nitrogen fertilization in tree species. Tree Physiol 38:1706–1723
Livingston N, Whitehead D, Kelliher F, Wang YP, Grace J, Walcroft A, Byers J, McSeveny T, Millard P (1998) Nitrogen allocation and carbon isotope fractionation in relation to intercepted radiation and position in a young Pinus radiata D. Don tree Plant Cell Environ 21:795–803
Marchand W, Girardin MP, Hartmann H, Depardieu C, Isabel N, Gauthier S, Boucher É, Bergeron Y (2020) Strong overestimation of water-use efficiency responses to rising CO2 in tree-ring studies. Global Change Biol 26:4538–4558
Marshall JD, Linder S (2013) Mineral nutrition and elevated [CO2] interact to modify δ13C, an index of gas exchange, in Norway spruce. Tree Physiol 33:1132–1144
Marshall JD, Monserud RA (2003) Foliage height influences specific leaf area of three conifer species. Can J For Res 33:164–170
Martínez-Vilalta J, Garcia-Forner N (2017) Water potential regulation, stomatal behaviour and hydraulic transport under drought: deconstructing the iso/anisohydric concept. Plant Cell Environ 40:962–976
McDowell NG, Allen CD, Anderson-Teixeira K, Aukema BH, Bond-Lamberty B, Chini L, Clark JS, Dietze M, Grossiord C, Hanbury-Brown A, Hurtt GC, Jackson RB, Johnson DJ, Kueppers L, Lichstein JW, Ogle K, Poulter B, Pugh TAM, Seidl R, Turner MG, Uriarte M, Walker AP, Xu C (2020) Pervasive shifts in forest dynamics in a changing world. Science 368:eaaz9463
McDowell NG, Bond BJ, Dickman LT, Ryan MG, Whitehead D (2011) Relationships between tree height and carbon isotope discrimination. In: Meinzer F, Dawson T, Lachenbruch B (eds) Size-and age-related changes in tree structure and function. Springer, New York, pp 255–286
McDowell NG, Licata J, Bond BJ (2005) Environmental sensitivity of gas exchange in different-sized trees. Oecologia 145:9
McDowell NG, Phillips N, Lunch C, Bond BJ, Ryan MG (2002) An investigation of hydraulic limitation and compensation in large, old Douglas-fir trees. Tree Physiol 22:763–774
Mcdowell NG, Williams A, Xu C, Pockman W, Dickman L, Sevanto S, Pangle R, Limousin J, Plaut J, Mackay D (2016) Multi-scale predictions of massive conifer mortality due to chronic temperature rise. Nat Clim Change 6:295–300
Monti A, Brugnoli E, Scartazza A, Amaducci M (2006) The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L.). J Exp Bot 57:1253–1262
Morales F, Pavlovič A, Abadía A, Abadía J (2018) Photosynthesis in poor nutrient soils, in compacted soils, and under drought. In: Adams Iii WW, Terashima I (eds) The leaf: a platform for performing photosynthesis. Springer International Publishing, Berlin, pp 371–399
National Centers for Environmental Information (National Oceanic and Atmospheric Administration) (2020) Monthly precipitation in 2004. https://www.ncdc.noaa.gov/. Accessed 20 July 2020
Niinemets Ü, Díaz-Espejo A, Flexas J, Galmés J, Warren CR (2009) Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field. J Exp Bot 60:2249–2270
Pangle R, Kavanagh K, Duursma R (2015) Decline in canopy gas exchange with increasing tree height, atmospheric evaporative demand, and seasonal drought in co-occurring inland Pacific Northwest conifer species. Can J For Res 45:1086–1101
Peichl M, Arain MA (2007) Allometry and partitioning of above-and belowground tree biomass in an age-sequence of white pine forests. For Ecol Manag 253:68–80
Pflug EE, Buchmann N, Siegwolf RTW, Schaub M, Rigling A, Arend M (2018) Resilient leaf physiological response of European Beech (Fagus sylvatica L.) to Summer Drought and Drought Release. Front. Plant Sci. 9:187. https://doi.org/10.3389/fpls.2018.00187
Phillips N, Bond BJ, McDowell NG, Ryan MG (2002) Canopy and hydraulic conductance in young, mature and old Douglas-fir trees. Tree Physiol 22:205–211
R Core Team (2018) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/. Accessed 1 Aug 2018
Restaino CM, Peterson DL, Littell J (2016) Increased water deficit decreases Douglas fir growth throughout western US forests. Proc Natl Acad Sci 113:9557–9562
RStudio Team (2016) RStudio: integrated development for R. RStudio, Inc., Boston, MA URL http://www.rstudio.com/. Accessed 1 Dec 2016
Ryan MG, Phillips N, Bond BJ (2006) The hydraulic limitation hypothesis revisited. Plant Cell Environ 29:367–381
Ryan MG, Yoder BJ (1997) Hydraulic limits to tree height and tree growth. Bioscience 47:235–242
Saldarriaga JG, Luxmoore RJ (1991) Solar energy conversion efficiencies during succession of a tropical rain forest in Amazonia. J Trop Ecol 7:233–242
Sanginés de Cárcer P, Vitasse Y, Peñuelas J, Jassey VE, Buttler A, Signarbieux C (2018) Vapor–pressure deficit and extreme climatic variables limit tree growth. Glob Change Biol 24:1108–1122
Scartazza A, Lauteri M, Guido M, Brugnoli E (1998) Carbon isotope discrimination in leaf and stem sugars, water-use efficiency and mesophyll conductance during different developmental stages in rice subjected to drought. Funct Plant Biol 25:489–498
Scartazza A, Mata C, Matteucci G, Yakir D, Moscatello S, Brugnoli E (2004) Comparisons of δ13C of photosynthetic products and ecosystem respiratory CO2 and their responses to seasonal climate variability. Oecologia 140:340–351
Seibt U, Rajabi A, Griffiths H, Berry JA (2008) Carbon isotopes and water use efficiency: sense and sensitivity. Oecologia 155:441
Sendall KM, Reich PB (2013) Variation in leaf and twig CO2 flux as a function of plant size: a comparison of seedlings, saplings and trees. Tree Physiol 33:713–729
Simard M, Lecomte N, Bergeron Y, Bernier PY, Paré D (2007) Forest productivity decline caused by successional paludification of boreal soils. Ecol Appl 17:1619–1637
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
Smith DM, Larson BC, Kelty MJ, Ashton PMS (1997) The practice of silviculture: applied forest ecology. Wiley, New York
Swidrak I, Schuster R, Oberhuber W (2013) Comparing growth phenology of co-occurring deciduous and evergreen conifers exposed to drought. Flora 208:609–617
Tang J, Luyssaert S, Richardson AD, Kutsch W, Janssens IA (2014) Steeper declines in forest photosynthesis than respiration explain age-driven decreases in forest growth. Proc Natl Acad Sci USA 111:8856–8860
Ubierna N, Marshall JD (2011) Estimation of canopy average mesophyll conductance using δ13C of phloem contents. Plant Cell Environ 34:1521–1535
van de Weg MJ, Meir P, Grace J, Ramos GD (2012) Photosynthetic parameters, dark respiration and leaf traits in the canopy of a Peruvian tropical montane cloud forest. Oecologia 168:23–34
Vitousek PM, Field CB, Matson PA (1990) Variation in foliar δ13C in Hawaiian Metrosideros polymorpha: a case of internal resistance? Oecologia 84:362–370
Wanek W, Heintel S, Richter A (2001) Preparation of starch and other carbon fractions from higher plant leaves for stable carbon isotope analysis. Rapid Commun Mass Spectrom 15:1136–1140
Warren CR (2008a) Stand aside stomata, another actor deserves centre stage: the forgotten role of the internal conductance to CO2 transfer. J Exp Bot 59:1475–1487
Warren C (2008b) Does growth temperature affect the temperature responses of photosynthesis and internal conductance to CO2? A test with Eucalyptus regnans. Tree Physiol 28:11–19
Warren C, Ethier G, Livingston N, Grant N, Turpin D, Harrison D, Black T (2003) Transfer conductance in second growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) canopies. Plant Cell Environ 26:1215–1227
Werner C, Gessler A (2011) Diel variations in the carbon isotope composition of respired CO2 and associated carbon sources: a review of dynamics and mechanisms. Biogeosciences Discussions. 8(2):2437–2459
Western Regional Climate Center (2020) 30-year average monthly precipitation between 1971 and 2000. http://www.wrcc.dri.edu/index.html. Accessed 20 July 2020.
Woodruff D, Meinzer F, Lachenbruch B, Johnson D (2009) Coordination of leaf structure and gas exchange along a height gradient in a tall conifer. Tree Physiol 29:261–272
Yang J, Duursma RA, De Kauwe MG, Kumarathunge D, Jiang M, Mahmud K, Gimeno TE, Crous KY, Ellsworth DS, Peters J (2019) Incorporating non-stomatal limitation improves the performance of leaf and canopy models at high vapour pressure deficit. Tree Physiol 39:1961–1974
Zeeman SC, Kossmann J, Smith AM (2010) Starch: its metabolism, evolution, and biotechnological modification in plants. Annu Rev Plant Biol 61:209–234
We would like to thank the Priest River Experimental Forest for allowing us to establish our research sites and for use of their facilities. We are indebted to our sawyer Dana Townsend and to Benjamin Miller for assistance in processing foliar samples. We also thank Bob Brander and Idaho Stable Isotopes Laboratory, as well as Dr. Ray Lee and his lab at Washington State University for isotopic sample analysis. Thanks to John Marshall, Jodi Johnson-Maynard, and Alan Black for providing thoughtful comments on this manuscript, and three anonymous reviewers whose suggestions greatly improved the article.
This research was funded with a grant from McIntire-Stennis Cooperative Forestry Research Program (Grant number: 0199175: Seasonal changes in water use across forest stands of differing age and height at the Priest River Experimental Forest) awarded to KLK.
Conflict of interest
The authors declare that they have no conflict of interest.
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Communicated by A. Gessler.
About this article
Cite this article
Koyama, A., Schotzko, A.D., Schedlbauer, J.L. et al. Can variation in canopy \(\delta\)13C be attributed to changes in tree height? An investigation of three conifer species. Trees (2021). https://doi.org/10.1007/s00468-020-02069-5
- Hydraulic limitation theory
- Foliar \(\delta\) 13C
- N area
- Sequential ANOVAs