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 as they apply to organisms and their interactions in ecosystems. [1] 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 have a long history in ecology with early references to the constraints of mass balance made by Liebig, Lotka, and Redfield. These earlier concepts have been extended to explicitly link the elemental physiology of organisms to their food web interactions and ecosystem function. [2] [3]
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 and grasses, 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, which have a tissue carbon:nitrogen ratio (C:N) of about 5 yet consume wood with a C:N ratio of 300–1000. Ecological stoichiometry primarily asks:
Elemental imbalances arise for a number of physiological and evolutionary reasons related to the differences in the biological make up of organisms, such as differences in types and amounts of macromolecules, organelles, 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, elemental homeostasis refers to the maintenance of elemental composition within some biologically ordered range. Photoautotrophic organisms, such as algae and vascular plants, 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 animals, have close to strict homeostasis and they can be thought of as having distinct chemical composition. For example, carbon to phosphorus ratios in the suspended organic matter in lakes (i.e., algae, bacteria, and detritus) can vary between 100 and 1000 whereas C:P ratios of Daphnia , a crustacean zooplankton, 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 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, as well as phenomena at the whole biosphere level, such as the oxygen content of Earth's atmosphere.
To date the research framework of ecological stoichiometry stimulated research in various fields of biology, ecology, biochemistry and human health, including human microbiome research, [4] cancer research, [5] food web interactions, [6] population dynamics, [7] ecosystem services, [7] productivity of agricultural crops [7] and honeybee nutrition. [8]
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. [9] This release in the limitation of N and P can impact the abundance, growth rates, and biomass of primary producers within the stream. [10] 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.
The demands for carbon, nitrogen and phosphorus at specific ratios by invertebrates 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 acids in biomass production and are reflected in the P content of the consumer. [11] [12] During early growth stages, or earlier instars, 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.
An ecosystem 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 system through photosynthesis and is incorporated into plant tissue. By feeding on plants and on one another, animals play an important role in the movement of matter and energy through the system. They also influence the quantity of plant and microbial biomass present. By breaking down dead organic matter, decomposers release carbon back to the atmosphere and facilitate nutrient cycling by converting nutrients stored in dead biomass back to a form that can be readily used by plants and microbes.
Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.
Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytoplankton productivity". Water bodies with very low nutrient levels are termed oligotrophic and those with moderate nutrient levels are termed mesotrophic. Advanced eutrophication may also be referred to as dystrophic and hypertrophic conditions. Eutrophication can affect freshwater or salt water systems. In freshwater ecosystems it is almost always caused by excess phosphorus. In coastal waters on the other hand, the main contributing nutrient is more likely to be nitrogen, or nitrogen and phosphorus together. This depends on the location and other factors.
Isotope analysis is the identification of isotopic signature, abundance of certain stable isotopes of chemical elements within organic and inorganic compounds. Isotopic analysis can be used to understand the flow of energy through a food web, to reconstruct past environmental and climatic conditions, to investigate human and animal diets, for food authentification, and a variety of other physical, geological, palaeontological and chemical processes. Stable isotope ratios are measured using mass spectrometry, which separates the different isotopes of an element on the basis of their mass-to-charge ratio.
This glossary of ecology is a list of definitions of terms and concepts in ecology and related fields. For more specific definitions from other glossaries related to ecology, see Glossary of biology, Glossary of evolutionary biology, and Glossary of environmental science.
Energy flow is the flow of energy through living things within an ecosystem. All living organisms can be organized into producers and consumers, and those producers and consumers can further be organized into a food chain. Each of the levels within the food chain is a trophic level. In order to more efficiently show the quantity of organisms at each trophic level, these food chains are then organized into trophic pyramids. The arrows in the food chain show that the energy flow is unidirectional, with the head of an arrow indicating the direction of energy flow; energy is lost as heat at each step along the way.
The metabolic theory of ecology (MTE) is the ecological component of the more general Metabolic Scaling Theory and Kleiber's law. It posits that the metabolic rate of organisms is the fundamental biological rate that governs most observed patterns in ecology. MTE is part of a larger set of theory known as metabolic scaling theory that attempts to provide a unified theory for the importance of metabolism in driving pattern and process in biology from the level of cells all the way to the biosphere.
Detritivores are heterotrophs that obtain nutrients by consuming detritus. There are many kinds of invertebrates, vertebrates, and plants that carry out coprophagy. By doing so, all these detritivores contribute to decomposition and the nutrient cycles. Detritivores should be distinguished from other decomposers, such as many species of bacteria, fungi and protists, which are unable to ingest discrete lumps of matter. Instead, these other decomposers live by absorbing and metabolizing on a molecular scale. The terms detritivore and decomposer are often used interchangeably, but they describe different organisms. Detritivores are usually arthropods and help in the process of remineralization. Detritivores perform the first stage of remineralization, by fragmenting the dead plant matter, allowing decomposers to perform the second stage of remineralization.
An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem engineers are important for maintaining the health and stability of the environment they are living in. Since all organisms impact the environment they live in one way or another, it has been proposed that the term "ecosystem engineers" be used only for keystone species whose behavior very strongly affects other organisms.
Coarse woody debris (CWD) or coarse woody habitat (CWH) refers to fallen dead trees and the remains of large branches on the ground in forests and in rivers or wetlands. A dead standing tree – known as a snag – provides many of the same functions as coarse woody debris. The minimum size required for woody debris to be defined as "coarse" varies by author, ranging from 2.5–20 cm (1–8 in) in diameter.
An arbuscular mycorrhiza (AM) is a type of mycorrhiza in which the symbiont fungus penetrates the cortical cells of the roots of a vascular plant forming arbuscules. Arbuscular mycorrhiza is a type of endomycorrhiza along with ericoid mycorrhiza and orchid mycorrhiza .They are characterized by the formation of unique tree-like structures, the arbuscules. In addition, globular storage structures called vesicles are often encountered.
A limiting factor is a variable of a system that causes a noticeable change in output or another measure of a type of system. The limiting factor is in a pyramid shape of organisms going up from the producers to consumers and so on. A factor not limiting over a certain domain of starting conditions may yet be limiting over another domain of starting conditions, including that of the factor.
Hydrobiology is the science of life and life processes in water. Much of modern hydrobiology can be viewed as a sub-discipline of ecology but the sphere of hydrobiology includes taxonomy, economic and industrial biology, morphology, and physiology. The one distinguishing aspect is that all fields relate to aquatic organisms. Most work is related to limnology and can be divided into lotic system ecology and lentic system ecology.
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 dynamic energy budget (DEB) theory is a formal metabolic theory which provides a single quantitative framework to dynamically describe the aspects of metabolism of all living organisms at the individual level, based on assumptions about energy uptake, storage, and utilization of various substances. The DEB theory adheres to stringent thermodynamic principles, is motivated by universally observed patterns, is non-species specific, and links different levels of biological organization as prescribed by the implications of energetics. Models based on the DEB theory have been successfully applied to over a 1000 species with real-life applications ranging from conservation, aquaculture, general ecology, and ecotoxicology. The theory is contributing to the theoretical underpinning of the emerging field of metabolic ecology.
The phosphorus cycle is the biogeochemical cycle that describes the movement of phosphorus through the lithosphere, hydrosphere, and biosphere. Unlike many other biogeochemical cycles, the atmosphere does not play a significant role in the movement of phosphorus, because phosphorus and phosphorus-based compounds are usually solids at the typical ranges of temperature and pressure found on Earth. The production of phosphine gas occurs in only specialized, local conditions. Therefore, the phosphorus cycle should be viewed from whole Earth system and then specifically focused on the cycle in terrestrial and aquatic systems.
Plant ecology is a subdiscipline of ecology that studies the distribution and abundance of plants, the effects of environmental factors upon the abundance of plants, and the interactions among plants and between plants and other organisms. Examples of these are the distribution of temperate deciduous forests in North America, the effects of drought or flooding upon plant survival, and competition among desert plants for water, or effects of herds of grazing animals upon the composition of grasslands.
A nutrient cycle is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cyclic. Mineral cycles include the carbon cycle, sulfur cycle, nitrogen cycle, water cycle, phosphorus cycle, oxygen cycle, among others that continually recycle along with other mineral nutrients into productive ecological nutrition.
Marine primary production is the chemical synthesis in the ocean of organic compounds from atmospheric or dissolved carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through chemosynthesis, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all life on Earth relies directly or indirectly on primary production. The organisms responsible for primary production are called primary producers or autotrophs.
James Elser is an American ecologist and limnologist. He is Director & Bierman Professor of Ecology, Flathead Lake Biological Station, University of Montana and research professor, School of Life Sciences, Arizona State University. He is known for his work in ecological stoichiometry. In 2019, he was elected to the National Academy of Sciences.