Open Access
Issue
Published:
21 August 2020
Effects of climate, biotic factors, and phylogeny on allometric relationships: testing the metabolic scaling theory in plantations and natural forests across China
(
),
Dayong Fan2(
)
Download citation
397
Views
2
Downloads
6
Crossref
N/A
WoS
5
Scopus
0
CSCD
Metabolic scaling theory (MST) is still in debate because observed allometric exponents often deviate from MST predictions, and can change significantly depending on environment, phylogeny, and disturbance. We assembled published scaling exponents from literatures for three allometric relationships linked to biomass allocation: leaf biomass-diameter (L-D), stem biomass-diameter (S-D), and root biomass-diameter (R-D). We used data from natural forests and plantations across China to test the following hypotheses: 1) the allometric relationships of trees support the predictions of MST on a broad scale; 2) the observed deviations from MST predictions are caused by climate, biotic factors, and/or phylogeny; 3) abiotic and biotic factors influence allometric relationships in plantations and natural forests differently, and different allometric relationships (i.e. L-D, S-D, and R-D) are affected differently. We related these scaling exponents to geographic climate gradient, successional stage, stand density, leaf form and phenology, and phylogeny. We used mixed-effect models to examine the major factors affecting tree allometries.
In natural forests, S-D and R-D scaling exponents were consistent with MST predictions in primary forests, but were significantly lower in secondary forests. Both S-D and R-D scaling exponents in plantations had a medium value that fell between those of the secondary and primary forests, despite plantations being similar in species characteristics and age to secondary forests. The S-D and R-D exponents were significantly affected by factors that are not yet considered in MST, including winter coldness which explained 2.76% – 3.24% of variations, successional stage (7.91% – 8.20% of variations), density (a surrogate for competition, 5.86% – 8.54% of variations), and especially phylogeny (45.86% – 56.64% of variations explained). However, the L-D scaling exponents conformed to MST predictions in primary, secondary, and plantation forests, and was not strongly explained by most factors.
MST is only applicable to primary (steady-state) forests, and climate, biotic factors and phylogeny are causes of the observed deviations of allometric relationships from MST predictions. Forest management practices in plantations have a strong influence on tree allometries. L-D allometry is more strongly controlled by biophysical constraints than S-D and R-D allometries, however, the mechanisms behind this difference still need further examinations.
menu
Effects of climate, biotic factors, and phylogeny on allometric relationships: testing the metabolic scaling theory in plantations and natural forests across China
(
), Dayong Fan2(
)
Abstract
Metabolic scaling theory (MST) is still in debate because observed allometric exponents often deviate from MST predictions, and can change significantly depending on environment, phylogeny, and disturbance. We assembled published scaling exponents from literatures for three allometric relationships linked to biomass allocation: leaf biomass-diameter (L-D), stem biomass-diameter (S-D), and root biomass-diameter (R-D). We used data from natural forests and plantations across China to test the following hypotheses: 1) the allometric relationships of trees support the predictions of MST on a broad scale; 2) the observed deviations from MST predictions are caused by climate, biotic factors, and/or phylogeny; 3) abiotic and biotic factors influence allometric relationships in plantations and natural forests differently, and different allometric relationships (i.e. L-D, S-D, and R-D) are affected differently. We related these scaling exponents to geographic climate gradient, successional stage, stand density, leaf form and phenology, and phylogeny. We used mixed-effect models to examine the major factors affecting tree allometries.
In natural forests, S-D and R-D scaling exponents were consistent with MST predictions in primary forests, but were significantly lower in secondary forests. Both S-D and R-D scaling exponents in plantations had a medium value that fell between those of the secondary and primary forests, despite plantations being similar in species characteristics and age to secondary forests. The S-D and R-D exponents were significantly affected by factors that are not yet considered in MST, including winter coldness which explained 2.76% – 3.24% of variations, successional stage (7.91% – 8.20% of variations), density (a surrogate for competition, 5.86% – 8.54% of variations), and especially phylogeny (45.86% – 56.64% of variations explained). However, the L-D scaling exponents conformed to MST predictions in primary, secondary, and plantation forests, and was not strongly explained by most factors.
MST is only applicable to primary (steady-state) forests, and climate, biotic factors and phylogeny are causes of the observed deviations of allometric relationships from MST predictions. Forest management practices in plantations have a strong influence on tree allometries. L-D allometry is more strongly controlled by biophysical constraints than S-D and R-D allometries, however, the mechanisms behind this difference still need further examinations.
References(52)
Apol MEF, Etienne RS, Olff H (2008) Revisiting the evolutionary origin of allometric metabolic scaling in biology. Funct Ecol 22: 1070-1080
Binkley D, Stape JL, Ryan MG (2004) Thinking about efficiency of resource use in forests. Forest Ecol Manag 193: 5–16
Brockerhoff EG, Jactel H, Parrotta JA, Quine CP, Sayer J (2008) Plantation forests and biodiversity: oxymoron or opportunity? Biodivers Conserv 17: 925–951
Brown JH, Gillooly JF, Allen AP, Savage VM, West GB (2004) Toward a metabolic theory of ecology. Ecology 85: 1771–1789
Chave J, Andalo C, Brown S, Cairns MA, Chambers JQ, Eamus D, Fölster H, Fromard F, Higuchi N, Kira T, Lescure J-P, Nelson BW, Ogawa H, Puig H, Riéra B, Yamakura T (2005) Tree allometry and improved estimation of carbon stocks and balance in tropical forests. Oecologia 145: 87–99
Chave J, Réjou-Méchain M, Búrquez A, Chidumayo E, Colgan MS, Delitti WBC, Duque A, Eid T, Fearnside PM, Goodman RC, Henry M, Martínez-Yrízar A, Mugasha WA, Muller-Landau HC, Mencuccini M, Nelson BW, Ngomanda A, Nogueira EM, Ortiz-Malavassi E, Pélissier R, Ploton P, Ryan CM, Saldarriaga JG, Vieilledent G (2014) Improved allometric models to estimate the aboveground biomass of tropical trees. Glob Chang Biol 20: 3177–3190
Deng J, Wang G, Morris EC, Wei X, Li D, Chen B, Zhao C, Liu J, Wang Y (2006) Plant mass-density relationship along a moisture gradient in north-West China. J Ecol 94: 953–958
Duncanson LI, Dubayah RO, Enquist BJ (2015) Assessing the general patterns of forest structure: quantifying tree and forest allometric scaling relationships in the United States. Glob Ecol Biogeogr 24: 1465–1475
Enquist BJ, Niklas KJ (2001) Invariant scaling relations across tree-dominated communities. Nature 410: 655–660
Enquist BJ, West GB, Brown JH (2009) Extensions and evaluations of a general quantitative theory of forest structure and dynamics. PNAS 106: 7046–7051
Fang J, Brown S, Tang Y, Nabuurs GJ, Wang X, Shen H (2006) Overestimated biomass carbon pools of the northern mid- and high latitude forests. Clim Chang 74: 355–368
Fang J, Shen Z, Tang Z, Wang X, Wang Z, Feng J, Liu Y, Qiao X, Wu X, Zheng C (2012) Forest community survey and the structural characteristics of forests in China. Ecography 35: 1059–1071
Getzin S, Dean C, He F, Trofymow JA, Wiegand K, Wiegand T (2006) Spatial patterns and competition of tree species in a Douglas-fir chronosequence on Vancouver Island. Ecography 29: 671–682
He JS, Wang X, Flynn DFB, Wang L, Schmid B, Fang J (2009) Taxonomic, phylogenetic, and environmental trade-offs between leaf productivity and persistence. Ecology 90: 2779–2791
Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A (2005) Very high resolution interpolated climate surfaces for global land areas. Int J Climatol 25: 1965–1978
Hulshof CM, Swenson NG, Weiser MD (2015) Tree height–diameter allometry across the United States. Ecol Evol 5: 1193–1204
Jacobsen AL, Pratt RB, Tobin MF, Hacke UG, Ewers FW (2012) A global analysis of xylem vessel length in woody plants. Am J Bot 99: 1583–1591
Johnson JB, Omland KS (2004) Model selection in ecology and evolution. Trends Ecol Evol 19: 101–108
Lines ER, Zavala MA, Purves DW, Coomes DA (2012) Predictable changes in aboveground allometry of trees along gradients of temperature, aridity and competition. Glob Ecol Biogeogr 21: 1017–1028
Luo Y, Wang X, Lu F (2015) Comprehensive database of biomass regressions for China's tree species. China Forestry Publishing House, Beijing
Luo Y, Wang X, Zhang X, Booth TH, Lu F (2012) Root: shoot ratios across China's forests: forest type and climatic effects. Forest Ecol Manag 269: 19–25
Luo Y, Zhang X, Wang X, Ren Y (2014) Dissecting variation in biomass conversion factors across China's forests: implications for biomass and carbon accounting. PLoS One 9: e94777
McMahon TA, Kronauer RE (1976) Tree structures: deducing the principle of mechanical design. J Theor Biol 59: 443–466
Mokany K, Raison RJ, Prokushkin AS (2006) Critical analysis of root: shoot ratios in terrestrial biomes. Glob Chang Biol 12: 84–96
Muller-Landau HC, Condit RS, Chave J, Thomas SC, Bohlman SA, Bunyavejchewin S, Davies S, Foster R, Gunatilleke S, Gunatilleke N, Harms KE, Hart T, Hubbell SP, Itoh A, Kassim AR, LaFrankie JV, Lee HS, Losos E, Makana J-R, Ohkubo T, Sukumar R, Sun I-F, Supardi MNN, Tan S, Thompson J, Valencia R, Muñoz GV, Wills C, Yamakura T, Chuyong G, Dattaraja HS, Esufali S, Hall P, Hernandez C, Kenfack D, Kiratiprayoon S, Suresh HS, Thomas D, Vallejo MI, Ashton P (2006) Testing metabolic ecology theory for allometric scaling of tree size, growth and mortality in tropical forests. Ecol Lett 9: 575–588
Niklas KJ, Spatz HC (2004) Growth and hydraulic (not mechanical) constraints govern the scaling of tree height and mass. PNAS 101: 15661–15663
Poorter H, Jagodzinski AM, Ruiz-Peinado R, Kuyah S, Luo Y, Oleksyn J, Usoltsev VA, Buckley TN, Reich PB, Sack L (2015) How does biomass distribution change with size and differ among species? An analysis for 1200 plant species from five continents. New Phytol 208: 736–749
Poorter H, Niklas KJ, Reich PB, Oleksyn J, Poot P, Mommer L (2012) Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. New Phytol 193: 30–50
Price CA, Enquist BJ, Savage VM (2007) A general model for allometric covariation in botanical form and function. PNAS 104: 13204–13209
Price CA, Ogle K, White EP, Weitz JS (2009) Evaluating scaling models in biology using hierarchical Bayesian approaches. Ecol Lett 12: 641–651
Quinn G, Keough M (2002) Experimental design and data analysis for biologists. Cambridge University Press, Cambridge
R Core Team (2014) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
Rubner M (1883) On the influence of body size on metabolism and energy exchange. Zeitschriftfür Biologie 19: 535–562
Rüger N, Condit R (2012) Testing metabolic theory with models of tree growth that include light competition. Funct Ecol 26: 759–765
Savage VM, Bentley LP, Enquist BJ, Sperry JS, Smith DD, Reich PB, von Allmen EI (2010) Hydraulic trade-offs and space filling enable better predictions of vascular structure and function in plants. PNAS 107: 22722–22727
Sun H, Wang X, Fan Y, Liu C, Wu P, Li Q, Yin W (2017b) Effects of biophysical constraints, climate and phylogeny on forest shrub allometries along an altitudinal gradient in Northeast China. Sci Rep 7: 43769
Sun H, Wang X, Wu P, Han W, Xu K, Liang P, Liu C, Yin W, Xia X (2017a) What causes greater deviations from predictions of metabolic scaling theory in earlier successional forests? Forest Ecol Manag 405: 101–111
Wang X, Fang J, Sanders NJ, White PS, Tang Z (2009) Relative importance of climate vs local factors in shaping the regional patterns of forest plant richness across Northeast China. Ecography 32: 133–142
Wang X, Fang J, Tang Z, Zhu B (2006) Climatic control of primary forest structure and DBH–height allometry in Northeast China. Forest Ecol Manag 234: 264–274
Wang X, Fang J, Zhu B (2008) Forest biomass and root-shoot allocation in Northeast China. Forest Ecol Manag 255: 4007–4020
Wang X, Ouyang S, Sun OJ, Fang J (2013) Forest biomass patterns across Northeast China are strongly shaped by forest height. Forest Ecol Manag 293: 149–160
Wang Y, Chu C, Zhu K, Shen Z (2011) Effects of inter-specific variability on biomass allocation: a hierarchical Bayesian approach. Ecol Inform 6: 341–344
Wang Y, Pederson N, Ellison AM, Buckley HL, Case BS, Liang E, Julio Camarero J (2016) Increased stem density and competition may diminish the positive effects of warming at alpine treeline. Ecology 97: 1668–1679
Warton DI, Wright IJ, Falster DS, Westoby M (2006) Bivariate line-fitting methods for allometry. Biol Rev 81: 259–291
West GB, Brown JH, Enquist BJ (1997) A general model for the origin of allometric scaling laws in biology. Science 276: 122–126
West GB, Brown JH, Enquist BJ (1999) A general model for the structure and allometry of plant vascular systems. Nature 400: 664–667
White CR, Cassey P, Blackburn TM (2007) Allometric exponents do not support a universal metabolic allometry. Ecology 88: 315–323
Wright IJ, Dong N, Maire V, Prentice IC, Westoby M, Díaz S, Gallagher RV, Jacobs BF, Kooyman R, Law EA, Leishman MR, Niinemets Ü, Reich PB, Sack L, Villar R, Wang H, Wilf P (2017) Global climatic drivers of leaf size. Science 357: 917–921
Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Groom PK, Hikosaka K, Lee W, Lusk CH, Niinemets Ü, Oleksyn J, Osada N, Poorter H, Warton DI, Westoby M (2005) Modulation of leaf economic traits and trait relationships by climate. Glob Ecol Biogeogr 14: 411–421
Zanne AE, Tank DC, Cornwell WK, Eastman JM, Smith SA, Fitzjohn RG, McGlinn DJ, O'Meara BC, Moles AT, Reich PB, Royer DL, Soltis DE, Stevens PF, Westoby M, Wright IJ, Aarssen L, Bertin RI, Calaminus A, Govaerts R, Hemmings F, Leishman MR, Jacek O, Soltis PS, Swenson NG, Warman L, Beaulieu JM (2014) Three keys to the radiation of angiosperms into freezing environments. Nature 506: 89–92
Zhang H, Song T, Wang K, Wang G, Liao J, Xu G, Zeng F (2015) Biogeographical patterns of forest biomass allocation vary by climate, soil and forest characteristics in China. Environ Res Lett 10: 44014
Publication history
Copyright
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
FEEDBACK 
TOP