Skip to main content Accessibility help

We use cookies to distinguish you from other users and to provide you with a better experience on our websites. Close this message to accept cookies or find out how to manage your cookie settings.

Close cookie message
×
Hostname: page-component-cd9895bd7-lnqnp Total loading time: 0 Render date: 2024-12-27T00:12:23.971Z Has data issue: false hasContentIssue false

2 - Toward a General Scaling Theory for Linking Traits, Stoichiometry, and Body Size to Ecosystem Function

from Part I - Connecting Ecosystem and Geoscience Processes

Published online by Cambridge University Press:  27 October 2016

By
Brian J. Enquist ,
Sean T. Michaletz  and
Andrew J. Kerkhoff
Edited by
Edward A. Johnson  and
Yvonne E. Martin
Brian J. Enquist
Affiliation:
University of Arizona
Sean T. Michaletz
Affiliation:
Los Alamos National Laboratory
Andrew J. Kerkhoff
Affiliation:
Departments of Biology and Mathematics
Edward A. Johnson
Affiliation:
University of Calgary
Yvonne E. Martin
Affiliation:
University of Calgary
Get access

Summary

Introduction

[W]ithout theory, we have only phenomenology and correlation, and we lose the opportunity to yoke the complexity of ecological systems using simple, quantitative principles.

Marquet et al. (2014)

Scaling has been heralded as one of the major challenges of ecology for more than two decades (Levin, 1992; Ehleringer and Field, 1993). Here, in the spirit of Marquet et al. (2014), we provide an overview of a general theory for scaling based on simple quantitative principles. We argue that a focus on scaling also presents some of the more powerful scientific tools available to ecologists facing problems that are unprecedented in both their scope and their stakes. Indeed, one of the central challenges of ecosystem science is to scale up from measurements on individual traits, organisms, and locations to predict the carbon and nutrient pools and fluxes of entire ecosystems.

In terrestrial ecosystems, this challenge requires us to integrate the physiological functioning of plants (e.g., leaf-level photosynthesis) across a collection of heterogeneous individuals (e.g., plants of different species) to understand the functioning of the entire ensemble (e.g., primary productivity) (Ehleringer and Field, 1993; Chapin, 2003). In order to better predict the future of plant communities and ecosystem functioning in response to rising CO2 and enhanced nitrogen (N) deposition with changes in climate (temperature and precipitation), this sort of understanding must be extended to connect simultaneous changes in multiple biogeochemical cycles.

Why a General Allometric and Metabolic Theory of Ecosystems Is Needed

Recent re-evaluations of global change models indicate that they could greatly benefit from incorporating allometry and ecosystem scaling. Specifically, global change models used to understand how ecosystems respond to climate change frequently do not produce realistic biomass and allometries, which suggests the need for better models of plant growth, nutrient uptake, and mortality (Wolf et al., 2011). Metabolic scaling provides a bridge between leaf-, plant-, and stand-level measurements and the biogeochemical and thermodynamic processes that drive global change models.

In this chapter, we focus on the powerful control that plant size and functional traits exert on ecosystem pattern and process. We use recent insights from the Metabolic Scaling Theory (MST) to scale up from individual plant metabolism, nutrient stoichiometry, and functional traits to ecosystem-level pools and fluxes.

Type
Chapter
Information
A Biogeoscience Approach to Ecosystems , pp. 9 - 46
Publisher: Cambridge University Press
Print publication year: 2016

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Ågren, G. I. (2004). The C: N: P stoichiometry of autotrophs–theory and observations. Ecology Letters, 7, 185–91.Google Scholar
Allegaert, K., Anderson, B. J., Cossey, V. and Holford, N. H. G. (2006). Limited predictability of amikacin clearance in extreme premature neonates at birth. British Journal of Clinical Pharmacology, 61, 39–48.Google Scholar
Allen, A. P. and Gillooly, J. F. (2009). Towards an integration of ecological stoichiometry and the metabolic theory of ecology to better understand nutrient cycling. Ecology Letters, 12, 369–84.Google Scholar
Allen, A. P., Gillooly, J. F. and Brown, J. H. (2005). Linking the global carbon cycle to individual metabolism. Functional Ecology, 19, 202–13.Google Scholar
Allen, C. D., Savage, M., Falk, D. A. et al. (2002). Ecological restoration of southwestern ponderosa pine ecosystems: a broad perspective. Ecological Applications, 12, 1418–33.Google Scholar
Anderson-Teixeira, K. J. and Vitousek, P. M. (2012). Ecosystems. In R. M. Sibly, J. H. Brown and A. Kodric-Brown, A. (eds.), Metabolic Ecology: A Scaling Approach, pp. 99–111.
Atkin, O. K. and Tjoelker, M. G. (2003). Thermal acclimation and the dynamic response of plant respiration to temperature. Trends in Plant Science, 8, 343–51.Google Scholar
Bentley, L. P., Stegen, J. C., Savage, V. M. et al. (2013). An empirical assessment of tree branching networks and implications for plant allometric scaling models. Ecology Letters, 16, 1069–78.Google Scholar
Blackman, V. H. (1919). The compound interest law and plant growth. Annals of Botany, 33, 353–60.Google Scholar
Brown, J. H., Gillooly, J. F., Allen, A. P., Savage, V. M. and West, G. B. (2004). Toward a metabolic theory of ecology. Ecology, 85, 1771–89.Google Scholar
Calder, W. A. (1984). Size, Function, and Life History. Cambridge, MA: Harvard University Press.
Chapin, F. S. (2003). Effects of plant traits on ecosystem and regional processes: a conceptual framework for predicting the consequences of global change. Annals of Botany, 91, 455–63.Google Scholar
Clark, J. S. (1990). Integration of ecological levels: individual plant-growth; population mortality and ecosystem processes. Journal of Ecology, 78, 275–99.Google Scholar
Clark, J. S. (1992). Density-independent mortality, density compensation, gap formation, and self-thinning in plant populations. Theoretical Population Biology, 42, 172–98.Google Scholar
Coomes, D. A. (2006). Challenges to the generality of WBE theory. Trends in Ecology & Evolution, 21, 593–6.Google Scholar
Coomes, D. A., Duncan, R. P., Allen, R. B. and Truscott, J. (2003). Disturbances prevent stem size-density distributions in natural forests from following scaling relationships. Ecology Letters, 6, 980–9.Google Scholar
Costanza, R., d'Arge, R., deGroot, R. et al. (1997). The value of the world's ecosystem services and natural capital. Nature, 387, 253–60.Google Scholar
Damuth, J. (1981). Population-density and body size in mammals. Nature, 290, 699–700.Google Scholar
Deng, J.-M., Wang, G.-X., Morris, E. C. et al. (2006). Plant mass–density relationship along a moisture gradient in north-west China. Journal of Ecology, 94, 953–8.Google Scholar
Duncanson, L. I., Dubayah, R. O. and Enquist, B. J. (2005). Assessing the general patterns of forest structure: Quantifying tree and forest allometrc scaling relationships in the United States. Global Ecology and Biogeography, 24(12), 1465–1475.Google Scholar
Economo, E. P., Kerkhoff, A. J. and Enquist, B. J. (2005). Allometric growth, life-history invariants and population energetics. Ecology Letters, 8, 353–60.Google Scholar
Ehleringer, J. R. and Field, C. B. (1993). Scaling Physiological Processes, Leaf to Globe. San Diego, CA: Academic Press.
Elser, J. J., Fagan, W. F., Kerkhoff, A. J., Swenson, N. G. and Enquist, B. J. (2010). Biological stoichiometry of plant production: metabolism, scaling and ecological response to global change. New Phytologist, 186, 593–608.Google Scholar
Enquist, B. J. (2002). Universal scaling in tree and vascular plant allometry: toward a general quantitative theory linking plant form and function from cells to ecosystems. Tree Physiology, 22, 1045–64.Google Scholar
Enquist, B. J. and Bentley, L. P. (2012). Land plants: new theoretical directions and empirical prospects. In R. M. Sibly, J. H. Brown and A. Kodric-Brown, A. (eds.), Metabolic Ecology: A Scaling Approach, 164–87.
Enquist, B. J. and Niklas, K. J. (2001). Invariant scaling relations across tree-dominated communities. Nature, 410, 655–60.Google Scholar
Enquist, B. J. and Niklas, K. J. (2002). Global allocation rules for patterns of biomass partitioning in seed plants. Science, 295, 1517–20.Google Scholar
Enquist, B. J., Brown, J. H. and West, G. B. (1998). Allometric scaling of plant energetics and population density. Nature, 395, 163–5.Google Scholar
Enquist, B. J., West, G. B. and Brown, J. H. (2009). Extensions and evaluations of a general quantitative theory of forest structure and dynamics. Proceedings of the National Academy of Sciences of the United States of America, 106, 7046–51.Google Scholar
Enquist, B. J., Kerkhokff, A. J., Huxman, T. E. and Economo, E. P. (2007) Adaptive differences in plant physiology and ecosystem paradoxes: insights from Metabolic Scaling Theory. Global Change Biology, 13, 591–609.Google Scholar
Enquist, B. J., West, G. B., Charnov, E. L. and Brown, J. H. (1999). Allometric scaling of production and life-history variation in vascular plants. Nature, 401, 907–11.Google Scholar
Enquist, B. J., Allen, A. P., Brown, J. H. et al. (2007b). Biological scaling: does the exception prove the rule? Nature, 445, E9–E10.Google Scholar
Enquist, B. J., Economo, E. P., Huxman, T. E. et al. (2003). Scaling metabolism from organisms to ecosystems. Nature, 423, 639–42.Google Scholar
Enquist, B. J., Kerkhoff, A. J., Stark, S. C. et al. (2007a). A general integrative model for scaling plant growth, carbon flux, and functional trait spectra. Nature, 449, 218–22.Google Scholar
Enquist, B. J., Norberg, J., Bonser, S. P. et al. (2015). Scaling from traits to ecosystems: Developing a general trait driver theory via integrating trait-based and metabolic scaling theories. Advances in Ecological Research, 52, 249–318.Google Scholar
Folke, C., Carpenter, S., Walker, B. et al. (2004). Regime shifts, resilience, and biodiversity in ecosystem management. Annual Review of Ecology, Evolution, and Systematics, 35, 557–81.Google Scholar
Gifford, R. M. (2003). Plant respiration in productivity models: conceptualisation, representation and issues for global terrestrial carbon-cycle research. Functional Plant Biology, 30, 171–86.Google Scholar
Gillooly, J. F., Brown, J. H., West, G. B., Savage, V. M. and Charnov, E. L. (2001). Effects of size and temperature on metabolic rate. Science, 293, 2248–51.Google Scholar
Gillooly, J. F., Allen, A. P., Brown, J. H. et al. (2005). The metabolic basis of whole-organism RNA and phosphorus content. Proceedings of the National Academy of Sciences of the United States of America, 102, 11923–7.Google Scholar
Gillooly, J. F., Charnov, E. L., West, G. B., Savage, V. M. and Brown, J. H. (2002). Effects of size and temperature on development time. Nature, 417, 70–3.Google Scholar
Harte, J. (2002). Towards a synthesis of the Newtonian and Darwinian world views. Physics Today, 55, 29–37.Google Scholar
Hubbell, S. P. and Foster, R. B. (1990) Structure, dynamics, and equilibrium status of old-growth forest on Barro Colorado Island. In Gentry, A. H. (ed.), Four Neotropical Rainforests. New Haven, CT: Yale University Press, 522–41.
Hunt, R. (1978). Plant Growth Analysis. London: Edward Arnold Limited.
Kellner, J. R. and Asner, G. P. (2009) Convergent structural responses of tropical forests to diverse disturbance regimes. Ecology Letters, 12, 887–97.Google Scholar
Kerkhoff, A. J. and Enquist, B. J. (2006). Ecosystem allometry: the scaling of nutrient stocks and primary productivity across plant communities. Ecology Letters, 9, 419–27.Google Scholar
Kerkhoff, A. J. and Enquist, B. J. (2007). Implications of scaling approaches for understanding resilience and reorganization in ecosystems. Bioscience, 57, 489–99.Google Scholar
Kerkhoff, A. J., Enquist, B. J., Elser, J. J. and Fagan, W. F. (2005). Plant allometry, stoichiometry and the temperature-dependence of primary productivity. Global Ecology and Biogeography, 14, 585–98.Google Scholar
Kobe, R. K. (1999). Light gradient partitioning among tropical tree species through differential seedling mortality and growth. Ecology, 80, 187–201.Google Scholar
Koyama, K. and Kikuzawa, K. (2009.) Is whole-plant photosynthetic rate proportional to leaf area? A test of scalings and a logistic equation by leaf demography census. American Naturalist, 173, 640–9.Google Scholar
Lai, J., Coomes, D. A., Du, X. et al. (2013). A general combined model to describe tree diameter distributions within subtropical and temperate forest communities. Oikos, 122, 1636–42.Google Scholar
Lambers, H., Freijsen, N., Poorter, H., Hirose, T. and van der Werff, H. (1989). Analyses of growth based on net assimilation rate and nitrogen productivity: their physiological background. In Variation in Growth Rate and Productivity of Higher Plants, ed. Lambers, H., Cambridge, M. L., Konings, H. and Pons, T. L.. The Hague: SPB Academic Publishing, pp. 1–17.
Lemaire, G., Khaity, M., Onillon, B., Allirand, J. M., Chartier, M. and Gosse, G. (1992). Dynamics of accumulation and partitioning of N in leaves, stems and roots of Lucerne (Medicago-Sativa L) in a dense canopy. Annals of Botany, 70, 429–35.Google Scholar
Levin, S. (1992) The problem of pattern and scale in ecology. Ecology, 73, 1943–67.Google Scholar
Lieberman, D., Hartshorn, G. S., Lieberman, M. and Peralta, R. (1990) Forest dynamics at La Selva Biological Station, 1969–1985. In: Gentry, A. H. (ed.), Four Neotropical Rainforests. New Haven, CT: Yale University Press, pp. 509–21.
Marba, N., Duarte, C. M. and Agustí, S. (2007). Allometric scaling of plant life history. Proceedings of the National Academy of Sciences of the United States of America, 104, 15777–80.Google Scholar
Marquet, P. A., Allen, A. P., Brown, J. H. et al. (2014). On theory in ecology. BioScience, 64, 701–10.Google Scholar
Martin, J. G., Kloeppel, B. D., Schaefer, T. L., Kimbler, D. L. and McNulty, S. G. (1998). Aboveground biomass and nitrogen allocation of ten deciduous southern Appalachian tree species. Canadian Journal of Forest Research, 28, 1648–59.Google Scholar
Matzek, V. and Vitousek, P. M. (2009). N : P stoichiometry and protein : RNA ratios in vascular plants: an evaluation of the growth-rate hypothesis. Ecology Letters, 12, 765–71.Google Scholar
Michaletz, S. T., Weiser, M. D., Zhou, J., et al. (2015). Plant thermoregulation: energetics, Trait-environment interactions, and carbon economics. Trends in Ecology and Evolution, 30, 714–24. doi:10.1016/j.tree.2015.09.006.Google Scholar
Moses, M. E., Hou, C., Woodruff, W. H. et al. (2008). Revisiting a model of ontogenetic growth: estimating model parameters from theory and data. The American Naturalist, 171, 632–45.Google Scholar
Muller-Landau, H. C., Condit, R. S., Harms, K. E. et al. (2006a). Comparing tropical forest tree size distributions with the predictions of metabolic ecology and equilibrium models. Ecology Letters, 9, 589–602.Google Scholar
Muller-Landau, H. C., Condit, R. S., Chave, J. et al. (2006b). Testing metabolic ecology theory for allometric scaling of tree size, growth, and mortality in tropical forests. Ecology Letters, 9, 575–88.Google Scholar
Murray, C. D. (1927). A relationship between circumference and weight in trees and its bearing on branching angles. The Journal of General Physiology, 10, 725–39.Google Scholar
Niklas, K. J. (2006). Plant allometry, leaf nitrogen and phosphorus stoichiometry, and interspecific trends in annual growth rates. Annals of Botany, 97, 155–63.Google Scholar
Niklas, K. J. and Enquist, B. J. (2001). Invariant scaling relationships for interspecific plant biomass production rates and body size. Proceedings of the National Academy of Sciences of the United States of America, 98, 2922–7.Google Scholar
Niklas, K. J., Owens, T., Reich, P. B. and Cobb, E. D. (2005). Nitrogen/phosphorus leaf stoichiometry and the scaling of plant growth. Ecology Letters, 8, 636–42.Google Scholar
Odum, E. P. (1969). The strategy of ecosystem development. Science, 164, 262–70.Google Scholar
Pearsall, W. H. (1927). Growth studies. VI: on the relative sizes of growing plant organs. Annals of Botany, 41, 549–56.Google Scholar
Peng, Y., Niklas, K. J. and Sun, S. (2011). The relationship between relative growth rate and whole-plant C: N: P stoichiometry in plant seedlings grown under nutrient-enriched conditions. Journal of Plant Ecology, 4, 147–56.Google Scholar
Peters, R. H. (1980). Useful concepts for predictive ecology. Synthese, 43, 257–69.Google Scholar
Peters, R. H. (1983). The Ecological Implications of Body Size. Cambridge, UK: Cambridge University Press.
Peterson, G., Allen, C. R. and Holling, C. S. (1998). Ecological resilience, biodiversity, and scale. Ecosystems, 1, 6–18.Google Scholar
Poorter, H. (1989). Interspecific variation in relative growth rate: on ecological causes and physiological consequences. In Causes and Consequences of Variation in Growth Rate and Productivity in Higher Plants, ed. Lambers, H., Cambridge, M. L., Konings, H. and Pons, T. L.. The Hague: SPB Academic Publishing, pp. 45–68.
Price, C., Enquist, B. and Savage, V. (2007). A general model for allometric covariation in botanical form and function. Proceedings of the National Academy of Sciences, 104, 13204–9.Google Scholar
Price, C. A., Gilooly, J. F., Allen, A. P., Weitz, J. S. and Niklas, K. J. (2010). The metabolic theory of ecology: prospects and challenges for plant biology. New Phytologist, 188, 696–710.Google Scholar
Price, C. A., Wright, I. J., Ackerly, D. D. et al. (2014). Are leaf functional traits ‘invariant’ with plant size and what is ‘invariance’ anyway? Functional Ecology, 28, 1330–43.Google Scholar
Reams, G. A., Smith, W. D., Hanse, M. H., Bechtold, W. A., Roesch, F. A., et al. (2005). The forest inventory and analysis sampling frame. Gen. Tech. Rep. SRS-80. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station, 21–36.
Reich, P. B., Tjoelker, M. G., Machado, J.-L. and Oleksyn, J. (2006). Universal scaling of respiratory metabolism, size and nitrogen in plants. Nature, 439, 457–61.Google Scholar
Reich, P. B., Tjoelker, M. G., Walters, M. B., Vanderklein, D. W. and Bushena, C. (1998). Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedlings of nine boreal tree species grown in high and low light. Functional Ecology, 12, 327–38.Google Scholar
Robinson, D. (2004). Scaling the depths: below-ground allocation in plants, forests and biomes. Functional Ecology, 18, 290–5.Google Scholar
Robinson, D. (2007). Implications of a large global root biomass for carbon sink estimates and for soil carbon dynamics. Proceedings of the Royal Society B, 274, 2753–9.Google Scholar
Russo, S. E., Wiser, S. K. and Coomes, D. A. (2007). Growth-size scaling relationships of woody plant species differ from predictions of the Metabolic Ecology Model. Ecology Letters, 10, 889–901.Google Scholar
Savage, V. M., Deeds, E. J. and Fontana, W. (2008). Sizing up allometric scaling theory. PLoS Computational Biology, 4, e1000171, doi: 10.1371/journal.pcbi.1000171.Google Scholar
Savage, V. M., Bentley, L. P., Enquist, B. J. et al. (2010). Hydraulic tradeoffs and space filling enable better predictions of vascular structure and function in plants. Proceedings of the National Academy of Sciences of the USA, 107, 22722–7.Google Scholar
Schmidt-Nielsen, K. (1984). Scaling: Why is Animal Size so Important? Cambridge, UK: Cambridge University Press.
Shaver, G. R. and Chapin, F. S. (1991). Production–biomass relationships and element cycling in contrasting Arctic vegetation types. Ecological Monographs, 61, 1–31.Google Scholar
Shinozaki, K., Yoda, K., Hozumi, K. and Kira, T. (1964). A quantitative analysis of plant form – the pipe model theory: I. Basic analysis. Japanese Journal of Ecology, 14, 97–105.Google Scholar
Standish, R. J., Hobbs, R. J., Mayfield, M. M. et al. (2014). Resilience in ecology: abstraction, distraction, or where the action is? Biological Conservation, 177, 43–51.Google Scholar
Stephenson, N. L., Das, A. J., Condit, R. et al. (2014). Rate of tree carbon accumulation increases continuously with tree size. Nature, 507, 90–3.Google Scholar
Tilman, D., HilleRisLambers, J., Harpole, S., et al. (2004). Does metabolic theory apply to community ecology? It's a matter of scale. Ecology, 85, 1797–9.Google Scholar
Vrede, T., Dobberfuhl, D. R., Kooijman, S. and Elser, J. J. (2004). Fundamental connections among organism C: N: P stoichiometry, macromolecular composition, and growth. Ecology, 85, 1217–29.Google Scholar
West, G. B., Brown, J. H. and Enquist, B. J. (1997). A general model for the origin of allometric scaling laws in biology. Science, 276, 122–6.Google Scholar
West, G. B., Brown, J. H. and Enquist, B. J. (1999a). The fourth dimension of life: fractal geometry and allometric scaling of organisms. Science, 284, 1677–9.Google Scholar
West, G. B., Brown, J. H. and Enquist, B. J. (1999b). A general model for the structure and allometry of plant vascular systems. Nature, 400, 664–7.Google Scholar
West, G. B., Brown, J. H. and Enquist, B. J. (2001). A general model for ontogenetic growth. Nature, 413, 628–31.Google Scholar
West, G. B., Enquist, B. J. and Brown, J. H. (2009). A general quantitative theory of forest structure and dynamics. Proceedings of the National Academy of Science of the United States of America, 106, 7040–5.Google Scholar
White, E. P., Enquist, B. J. and Green, J. L. (2008). On estimating the exponent of power law frequency distributions. Ecology, 89, 905–12.Google Scholar
White, E. P., Ernest, S. K. M., Kerkhoff, A. J. and Enquist, B. J. (2007). Relationships between body size and abundance in ecology. Trends in Ecology & Evolution, 22, 323–30.Google Scholar
Wolf, A., Ciais, P., Bellassen, V. et al. (2011). Forest biomass allometry in global land surface models. Global Biogeochemical Cycles, 25, GB3015, doi:10.1029/2010GB003917.Google Scholar
Yoda, K., Kira, T., Ogawa, H. and Hozumi, K. (1963). Self-thinning in overcrowded pure stands under cultivated and natural conditions. Journal of Biology Osaka City University, 14, 107–29.Google Scholar
Yvon-Durocher, G., Caffrey, J. M., Cescatti, A. et al. (2012). Reconciling the temperature dependence of respiration across timescales and ecosystem types. Nature, 487, 472–6.Google Scholar

Save book to Kindle

To save this book to your Kindle, first ensure no-reply@cambridge.org is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about saving to your Kindle.

Note you can select to save to either the @free.kindle.com or @kindle.com variations. ‘@free.kindle.com’ emails are free but can only be saved to your device when it is connected to wi-fi. ‘@kindle.com’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

Find out more about the Kindle Personal Document Service.

Available formats
×

Save book to Dropbox

To save content items to your account, please confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your account. Find out more about saving content to Dropbox.

Available formats
×

Save book to Google Drive

To save content items to your account, please confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your account. Find out more about saving content to Google Drive.

Available formats
×