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New Phytol. 2015 Nov;208(3):736-49. doi: 10.1111/nph.13571. Epub 2015 Jul 22.

How does biomass distribution change with size and differ among species? An analysis for 1200 plant species from five continents.

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Plant Sciences (IBG-2), Forschungszentrum Jülich GmbH, D-52425, Jülich, Germany.
Polish Academy of Sciences, Institute of Dendrology, Parkowa 5, Kornik, PL-62-035, Poland.
Department of Game Management and Forest Protection, Faculty of Forestry, Poznan University of Life Sciences, Wojska Polskiego 71c, Poznan, PL-60-625, Poland.
Departamento de Selvicultura y Gestión de Sistemas Forestales, INIA-CIFOR, Avda. A Coruña, km 7.5., Madrid, 28040, Spain.
Sustainable Forest Management Research Institute, University of Valladolid-INIA, Madrid, Spain.
Jomo Kenyatta University of Agriculture and Technology (JKUAT), PO Box 62000, Nairobi, 00200, Kenya.
Department of Ecology, School of Horticulture and Plant Protection, Yangzhou University, 48 Wenhui East Road, Yangzhou, 225009, China.
State Key Laboratory of Urban and Regional Ecology, Research Centre for Eco-Environmental Sciences, Chinese Academy of Sciences, 18 Shuangqing Road, Haidian District, Beijing, 100085, China.
Department of Forest Resources, University of Minnesota, 1530 Cleveland Ave N, St Paul, MN, 55108, USA.
Ural State Forest Engineering University, Sibirskiy Trakt 37, Ekaterinburg, 620100, Russia.
Botanical Garden of Ural Branch of Russian Academy of Sciences, ul. Vos'mogo Marta 202a, Ekaterinburg, 620144, Russia.
IA Watson Grains Research Centre, Faculty of Agriculture and Environment, The University of Sydney, 12656 Newell Highway, Narrabri, NSW, Australia.
Hawkesbury Institute for the Environment, University of Western Sydney, Locked Bag 1797, Penrith, NSW, 2751, Australia.
Department of Ecology and Evolution, University of California Los Angeles, 621 Charles E. Young Drive South, Los Angeles, CA, 90095, USA.


We compiled a global database for leaf, stem and root biomass representing c. 11 000 records for c. 1200 herbaceous and woody species grown under either controlled or field conditions. We used this data set to analyse allometric relationships and fractional biomass distribution to leaves, stems and roots. We tested whether allometric scaling exponents are generally constant across plant sizes as predicted by metabolic scaling theory, or whether instead they change dynamically with plant size. We also quantified interspecific variation in biomass distribution among plant families and functional groups. Across all species combined, leaf vs stem and leaf vs root scaling exponents decreased from c. 1.00 for small plants to c. 0.60 for the largest trees considered. Evergreens had substantially higher leaf mass fractions (LMFs) than deciduous species, whereas graminoids maintained higher root mass fractions (RMFs) than eudicotyledonous herbs. These patterns do not support the hypothesis of fixed allometric exponents. Rather, continuous shifts in allometric exponents with plant size during ontogeny and evolution are the norm. Across seed plants, variation in biomass distribution among species is related more to function than phylogeny. We propose that the higher LMF of evergreens at least partly compensates for their relatively low leaf area : leaf mass ratio.


allometry; biomass allocation; biomass distribution; leaf mass fraction (LMF); leaf weight ratio; metabolic scaling theory; shoot : root ratio

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