Ecological stoichiometry (more broadly referred to as biological stoichiometry) considers how the balance of energy and elements influences living systems. Similar to chemical stoichiometry, ecological stoichiometry is founded on constraints of

mass balance In physics, a mass balance, also called a material balance, is an application of conservation of mass to the analysis of physical systems. By accounting for material entering and leaving a system, mass flows can be identified which might have b ...

as they apply to organisms and their interactions in

ecosystems An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...

.R. W. Sterner and J. J. Elser (2002) ''Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere''.

Princeton University Press Princeton University Press is an independent publisher with close connections to Princeton University. Its mission is to disseminate scholarship within academia and society at large. The press was founded by Whitney Darrow, with the financial ...

. pp.584. Specifically, how does the balance of energy and elements affect and how is this balance affected by organisms and their interactions. Concepts of ecological

stoichiometry Stoichiometry refers to the relationship between the quantities of reactants and products before, during, and following chemical reactions. Stoichiometry is founded on the law of conservation of mass where the total mass of the reactants equ ...

have a long history in

ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...

with early references to the constraints of mass balance made by

Liebig Justus Freiherr von Liebig (12 May 1803 – 20 April 1873) was a German scientist who made major contributions to agricultural and biological chemistry, and is considered one of the principal founders of organic chemistry. As a professor at t ...

Lotka Alfred James Lotka (March 2, 1880 – December 5, 1949) was a US mathematician, physical chemist, and statistician, famous for his work in population dynamics and energetics. An American biophysicist, Lotka is best known for his proposal ...

, and Redfield. These earlier concepts have been extended to explicitly link the elemental

physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemic ...

organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...

s to their

food web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one o ...

interactions and ecosystem function. Hvalba scenery

Most work in ecological stoichiometry focuses on the interface between an organism and its resources. This interface, whether it is between plants and their nutrient resources or large

herbivores A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

and

grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...

es, is often characterized by dramatic differences in the elemental composition of each part. The difference, or mismatch, between the elemental demands of organisms and the elemental composition of resources leads to an elemental imbalance. Consider

termites Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...

, which have a tissue

carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...

nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...

ratio (C:N) of about 5 yet consume

wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...

with a C:N

ratio In mathematics, a ratio shows how many times one number contains another. For example, if there are eight oranges and six lemons in a bowl of fruit, then the ratio of oranges to lemons is eight to six (that is, 8:6, which is equivalent to the ...

of 300–1000. Ecological stoichiometry primarily asks: # why do elemental imbalances arise in nature? # how is consumer

and life-history affected by elemental imbalances? and # what are the subsequent effects on ecosystem processes? Elemental imbalances arise for a number of physiological and

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

ary reasons related to the differences in the biological make up of organisms, such as differences in types and amounts of

macromolecule A macromolecule is a very large molecule important to biophysical processes, such as a protein or nucleic acid. It is composed of thousands of covalently bonded atoms. Many macromolecules are polymers of smaller molecules called monomers. The ...

organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...

s, and tissues. Organisms differ in the flexibility of their biological make up and therefore in the degree to which organisms can maintain a constant chemical composition in the face of variations in their resources. Variations in resources can be related to the types of needed resources, their relative availability in time and space, and how they are acquired. The ability to maintain internal chemical composition despite changes in the chemical composition and availability of resources is referred to as "stoichiometric homeostasis". Like the general biological notion of

homeostasis In biology, homeostasis ( British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...

, elemental homeostasis refers to the maintenance of elemental composition within some biologically ordered range. Photoautotrophic organisms, such as

algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...

and

vascular plants Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...

, can exhibit a very wide range of physiological plasticity in elemental composition and thus have relatively weak stoichiometric homeostasis. In contrast, other organisms, such as multicellular

animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...

s, have close to strict homeostasis and they can be thought of as having distinct chemical composition. For example, carbon to

phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ea ...

ratios in the suspended organic matter in

lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...

s (i.e.,

, bacteria, and

detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts comm ...

) can vary between 100 and 1000 whereas C:P ratios of ''

Daphnia ''Daphnia'' is a genus of small planktonic crustaceans, in length. ''Daphnia'' are members of the order Anomopoda, and are one of the several small aquatic crustaceans commonly called water fleas because their saltatory swimming style resembl ...

'', a

crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...

zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...

, remain nearly constant at 80:1. The general differences in stoichiometric homeostasis between plants and animals can lead to large and variable elemental imbalances between consumers and resources. Ecological stoichiometry seeks to discover how the chemical content of organisms shapes their ecology. Ecological stoichiometry has been applied to studies of nutrient recycling, resource competition, animal growth, and nutrient limitation patterns in whole ecosystems. The

Redfield ratio The Redfield ratio or Redfield stoichiometry is the consistent atomic ratio of carbon, nitrogen and phosphorus found in marine phytoplankton and throughout the deep oceans. The term is named for American oceanographer Alfred C. Redfield who in ...

of the world's oceans is one very famous application of stoichiometric principles to ecology. Ecological stoichiometry also considers phenomena at the sub-cellular level, such as the P-content of a

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to fo ...

, as well as phenomena at the whole biosphere level, such as the oxygen content of

Earth's atmosphere The atmosphere of Earth is the layer of gases, known collectively as air, retained by Earth's gravity that surrounds the planet and forms its planetary atmosphere. The atmosphere of Earth protects life on Earth by creating pressure allowing fo ...

. To date the research framework of ecological stoichiometry stimulated research in various fields of biology, ecology, biochemistry and human health, including

human microbiome The human microbiome is the aggregate of all microbiota that reside on or within human tissues and biofluids along with the corresponding anatomical sites in which they reside, including the skin, mammary glands, seminal fluid, uterus, ovarian ...

research, cancer research, food web interactions, population dynamics,

ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...

, productivity of agricultural crops and honeybee nutrition.

Consumer stoichiometry and food webs

The study of elemental ratios (i.e., C:N:P) within the tissues of organisms can be used to understand how organisms respond to changes in resource quality and quantity. For instance, in aquatic ecosystems, nitrogen and phosphorus pollution within streams, often due to agricultural activities, can increase the amount of N and P available to primary producers. This release in the limitation of N and P can impact the abundance, growth rates, and

biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...

of primary producers within the stream. This change in primary production can trickle through the food web via bottom-up processes and impact the stoichiometry of organisms, limiting elements, and biogeochemical cycling of streams. In addition, bottom-up changes in elemental availability can influence the morphology, phenology, and physiology of organisms that will be discussed below. The focus of this article is on aquatic systems; however, similar processes related to ecological stoichiometry can be applied in the terrestrial environment, as well.

Invertebrate stoichiometry

The demands for carbon, nitrogen and phosphorus at specific ratios by

invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...

can change at different life stages within invertebrate life history. The growth rate hypothesis (GRH) addresses this phenomenon and states that the demands for phosphorus increase during active growth phases to produce P-rich

nucleic acid Nucleic acids are biopolymers, macromolecules, essential to all known forms of life. They are composed of nucleotides, which are the monomers made of three components: a 5-carbon sugar, a phosphate group and a nitrogenous base. The two main ...

s in biomass production and are reflected in the P content of the consumer. During early growth stages, or earlier

instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...

s, invertebrates may have higher demands for N and P enriched resources to fuel the ribosomal production of proteins and RNA. At later stages, the demand for particular elements may shift as they are no longer actively growing as rapidly or generating protein rich biomass. Growth rates of invertebrate organisms can also be limited by the resources that are available to them.

References

{{modelling ecosystems, expanded=other Ecology Stoichiometry

Consumer stoichiometry and food webs

Invertebrate stoichiometry

See also

References