Biological Stoichiometry

Abstract

Biological stoichiometry is the study of the balance of energy and multiple chemical elements in living systems. It compares elemental requirements of organisms for growth, reproduction and maintenance with that provided by their nutritional resources. It considers the physiological, cellular and biochemical underpinnings of stoichiometric differences as well as their evolutionary basis. Primary producers generally exhibit greater flexibility in elemental composition compared to consumers, which leads to elemental imbalances between adjacent trophic levels. For individual organisms, the relatively low supply of an element can alter metabolic and physiological processes involving the acquisition, incorporation and release of multiple chemical elements. When sustained, elemental imbalances slow growth and limit reproduction of organisms, particularly those with relatively high elemental requirements. Elemental imbalances have been documented in diverse ecosystems and at multiple trophic levels and affect key ecological and evolutionary processes underlying population dynamics, life‐history evolution, community structure, trophic interactions and ecosystem function.

Key Concepts

  • Biological stoichiometry studies the balance of energy and multiple chemical elements in living systems.
  • Biological stoichiometry compares the elemental compositions of resources with the elemental requirements of organisms. It also considers the environmental and evolutionary origins of elemental imbalances between producers and consumers.
  • Biological stoichiometry approaches processes such as organism growth, population dynamics and trophic interactions as if such processes were composite chemical reactions that must simultaneously meet the law of mass conservation for multiple chemical elements and the rules of exact proportions in chemical reactions.
  • Biological stoichiometry uses its elemental perspective on biochemical and physiological processes to understand intra‐ and interspecific interactions that involve the transfer or transformation of matter in food webs.
  • Biological stoichiometry also provides a mechanistic framework for how animal species mediate ecosystem processes such as nutrient recycling. The stoichiometric approach can also be used to study trophic interactions as well as decomposition and microbial release of elements.
  • Biological stoichiometry considers the molecular and evolutionary basis of major differences in the C:N:P ratios of living things. Understanding how evolution affects these ratios provides considerable insight into processes that link all levels of organisation in biology.

Keywords: metabolism; biochemistry; physiology; ecology; evolution; colimitation; biodiversity

Figure 1. Regressions between consumer and resource N:P for (a) algae, (b) bacteria, (c) zooplankton, (d) aquatic macroinvertebrates and (e) terrestrial insects. Black regression lines indicate least square regressions with p < 0.1 (plastic), and grey lines indicate regressions with p > 0.1 (strictly homeostatic). Species with insignificant (p > 0.1) regression slopes were considered as strictly homeostatic and their slope was displayed as zero (1/HN:P = 0). The length of the regression lines displayed corresponds to the respective data range. The dotted diagonal line shows the 1:1 relation. Reproduced with permission from Persson et al. 2010 © John Wiley & Sons Ltd.
Figure 2. (a) The components of the ‘growth rate hypothesis’ which link evolution of important life‐history traits associated with growth and development rate to ecological and ecosystem and ecological impacts because of their effects on cellular and biochemical allocations and biomass C:N:P stoichiometry. (b) Variation in cellular or body P content (% total P of dry mass; y‐axis) is strongly correlated with P content derived from RNA (x‐axis) both intraspecifically (various grey lines for each organism studied) and interspecifically (green line). In each case, high P/high RNA data are associated with fast growth rates. On average, RNA contributed approximately 50% to total organism P across the entire data set. Note that the slope of the green line fit to the entire data set is 0.97 (not significantly different than 1), indicating that not only is RNA correlated with P, it is quantitatively explanatory of the observed variation. Reproduced with permission from Elser et al. 2003 © John Wiley & Sons Ltd.
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Further Reading

Andersen T, Elser JJ and Hessen DO (2004) Stoichiometry and population dynamics. Ecology Letters 7: 884–900.

Cardinale BJ, Hillebrand H, Harpole WS, Gross K and Ptacnik R (2009) Separating the influence of resource “availability” from resource “imbalance” on productivity‐diversity relationships. Ecology Letters 12: 475–487. DOI: 10.1111/j.1461-0248.2009.01317.x.

Cross W, Wallace J, Rosemond A and Eggert S (2006) Whole‐system nutrient enrichment increases secondary production in a detritus‐based ecosystem. Ecology 87: 1556–1565.

Danger M, Daufresne T, Lucas F, Pissard S and Lacroix G (2008) Does Liebig's law of the minimum scale up from species to communities? Oikos 117: 1741–1751. DOI: 10.1111/j.1600-0706.2008.16793.x.

Elser JJ, Loladze I, Peace AL and Kuang Y (2012) Lotka re‐loaded: modeling trophic interactions under stoichiometric constraints. Ecological Modelling 245: 3–11.

Elser JJ and Urabe J (1999) The stoichiometry of consumer‐driven nutrient recycling: theory, observations, and consequences. Ecology 80: 735–751.

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Hillebrand H, Borer ET, Bracken MES, et al. (2009) Herbivore metabolism and stoichiometry each constrain herbivory at different organizational scales across ecosystems. Ecology Letters 12: 516–527. DOI: 10.1111/j.1461-0248.2009.01304.x.

Sperfeld E, Martin‐Creuzburg D and Wacker A (2012) Multiple resource limitation theory applied to herbivorous consumers: Liebig's minimum rule vs. interactive co‐limitation. Ecology Letters 15: 142–150. DOI: 10.1111/j.1461-0248.2011.01719.x.

Sterner RW and Elser JJ (2002) Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere. Princeton, NJ: Princeton University Press.

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Striebel, Maren, Frost, Paul C, and Elser, James J(Aug 2017) Biological Stoichiometry. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0021229.pub2]