Food web

A food web (or food cycle) is the natural interconnection of food chains and a graphical representation (usually an image) 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 of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs. To maintain their bodies, grow, develop, and to reproduce, autotrophs produce organic matter from inorganic substances, including both minerals and gases such as carbon dioxide. These chemical reactions require energy, which mainly comes from the Sun and largely by photosynthesis, although a very small amount comes from hydrothermal vents and hot springs. A gradient exists between trophic levels running from complete autotrophs that obtain their sole source of carbon from the atmosphere, to mixotrophs (such as carnivorous plants) that are autotrophic organisms that partially obtain organic matter from sources other than the atmosphere, and complete heterotrophs that must feed to obtain organic matter. The linkages in a food web illustrate the feeding pathways, such as where heterotrophs obtain organic matter by feeding on autotrophs and other heterotrophs. The food web is a simplified illustration of the various methods of feeding that links an ecosystem into a unified system of exchange. There are different kinds of feeding relations that can be roughly divided into herbivory, carnivory, scavenging and parasitism. Some of the organic matter eaten by heterotrophs, such as sugars, provides energy. Autotrophs and heterotrophs come in all sizes, from microscopic to many tonnes - from cyanobacteria to giant redwoods, and from viruses and bdellovibrio to blue whales.

Charles Elton pioneered the concept of food cycles, food chains, and food size in his classical 1927 book "Animal Ecology"; Elton's 'food cycle' was replaced by 'food web' in a subsequent ecological text. Elton organized species into functional groups, which was the basis for Raymond Lindeman's classic and landmark paper in 1942 on trophic dynamics. Lindeman emphasized the important role of decomposer organisms in a trophic system of classification. The notion of a food web has a historical foothold in the writings of Charles Darwin and his terminology, including an "entangled bank", "web of life", "web of complex relations", and in reference to the decomposition actions of earthworms he talked about "the continued movement of the particles of earth". Even earlier, in 1768 John Bruckner described nature as "one continued web of life".

Food webs are limited representations of real ecosystems as they necessarily aggregate many species into trophic species, which are functional groups of species that have the same predators and prey in a food web. Ecologists use these simplifications in quantitative (or mathematical representation) models of trophic or consumer-resource systems dynamics. Using these models they can measure and test for generalized patterns in the structure of real food web networks. Ecologists have identified non-random properties in the topographic structure of food webs. Published examples that are used in meta analysis are of variable quality with omissions. However, the number of empirical studies on community webs is on the rise and the mathematical treatment of food webs using network theory had identified patterns that are common to all. Scaling laws, for example, predict a relationship between the topology of food web predator-prey linkages and levels of species richness.

Taxonomy of a food web

FoodWebSimple
A simplified food web illustrating a three trophic food chain (producers-herbivores-carnivores) linked to decomposers. The movement of mineral nutrients is cyclic, whereas the movement of energy is unidirectional and noncyclic. Trophic species are encircled as nodes and arrows depict the links.[1][2]

Links in food webs map the feeding connections (who eats whom) in an ecological community. Food cycle is an obsolete term that is synonymous with food web. Ecologists can broadly group all life forms into one of two trophic layers, the autotrophs and the heterotrophs. Autotrophs produce more biomass energy, either chemically without the sun's energy or by capturing the sun's energy in photosynthesis, than they use during metabolic respiration. Heterotrophs consume rather than produce biomass energy as they metabolize, grow, and add to levels of secondary production. A food web depicts a collection of polyphagous heterotrophic consumers that network and cycle the flow of energy and nutrients from a productive base of self-feeding autotrophs.[3][4][5]

The base or basal species in a food web are those species without prey and can include autotrophs or saprophytic detritivores (i.e., the community of decomposers in soil, biofilms, and periphyton). Feeding connections in the web are called trophic links. The number of trophic links per consumer is a measure of food web connectance. Food chains are nested within the trophic links of food webs. Food chains are linear (noncyclic) feeding pathways that trace monophagous consumers from a base species up to the top consumer, which is usually a larger predatory carnivore.[6][7][8]

Linkages connect to nodes in a food web, which are aggregates of biological taxa called trophic species. Trophic species are functional groups that have the same predators and prey in a food web. Common examples of an aggregated node in a food web might include parasites, microbes, decomposers, saprotrophs, consumers, or predators, each containing many species in a web that can otherwise be connected to other trophic species.[9][10]

Trophic levels

TrophicWeb
A trophic pyramid (a) and a simplified community food web (b) illustrating ecological relations among creatures that are typical of a northern Boreal terrestrial ecosystem. The trophic pyramid roughly represents the biomass (usually measured as total dry-weight) at each level. Plants generally have the greatest biomass. Names of trophic categories are shown to the right of the pyramid. Some ecosystems, such as many wetlands, do not organize as a strict pyramid, because aquatic plants are not as productive as long-lived terrestrial plants such as trees. Ecological trophic pyramids are typically one of three kinds: 1) pyramid of numbers, 2) pyramid of biomass, or 3) pyramid of energy.[4]

Food webs have trophic levels and positions. Basal species, such as plants, form the first level and are the resource limited species that feed on no other living creature in the web. Basal species can be autotrophs or detritivores, including "decomposing organic material and its associated microorganisms which we defined as detritus, micro-inorganic material and associated microorganisms (MIP), and vascular plant material."[11]:94 Most autotrophs capture the sun's energy in chlorophyll, but some autotrophs (the chemolithotrophs) obtain energy by the chemical oxidation of inorganic compounds and can grow in dark environments, such as the sulfur bacterium Thiobacillus, which lives in hot sulfur springs. The top level has top (or apex) predators which no other species kills directly for its food resource needs. The intermediate levels are filled with omnivores that feed on more than one trophic level and cause energy to flow through a number of food pathways starting from a basal species.[12]

In the simplest scheme, the first trophic level (level 1) is plants, then herbivores (level 2), and then carnivores (level 3). The trophic level is equal to one more than the chain length, which is the number of links connecting to the base. The base of the food chain (primary producers or detritivores) is set at zero.[3][13] Ecologists identify feeding relations and organize species into trophic species through extensive gut content analysis of different species. The technique has been improved through the use of stable isotopes to better trace energy flow through the web.[14] It was once thought that omnivory was rare, but recent evidence suggests otherwise. This realization has made trophic classifications more complex.[15]

Trophic dynamics

The trophic level concept was introduced in a historical landmark paper on trophic dynamics in 1942 by Raymond L. Lindeman. The basis of trophic dynamics is the transfer of energy from one part of the ecosystem to another.[13][16] The trophic dynamic concept has served as a useful quantitative heuristic, but it has several major limitations including the precision by which an organism can be allocated to a specific trophic level. Omnivores, for example, are not restricted to any single level. Nonetheless, recent research has found that discrete trophic levels do exist, but "above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores."[15]

A central question in the trophic dynamic literature is the nature of control and regulation over resources and production. Ecologists use simplified one trophic position food chain models (producer, carnivore, decomposer). Using these models, ecologists have tested various types of ecological control mechanisms. For example, herbivores generally have an abundance of vegetative resources, which meant that their populations were largely controlled or regulated by predators. This is known as the top-down hypothesis or 'green-world' hypothesis. Alternatively to the top-down hypothesis, not all plant material is edible and the nutritional quality or antiherbivore defenses of plants (structural and chemical) suggests a bottom-up form of regulation or control.[17][18][19] Recent studies have concluded that both "top-down" and "bottom-up" forces can influence community structure and the strength of the influence is environmentally context dependent.[20][21] These complex multitrophic interactions involve more than two trophic levels in a food web.[22]

Another example of a multi-trophic interaction is a trophic cascade, in which predators help to increase plant growth and prevent overgrazing by suppressing herbivores. Links in a food-web illustrate direct trophic relations among species, but there are also indirect effects that can alter the abundance, distribution, or biomass in the trophic levels. For example, predators eating herbivores indirectly influence the control and regulation of primary production in plants. Although the predators do not eat the plants directly, they regulate the population of herbivores that are directly linked to plant trophism. The net effect of direct and indirect relations is called trophic cascades. Trophic cascades are separated into species-level cascades, where only a subset of the food-web dynamic is impacted by a change in population numbers, and community-level cascades, where a change in population numbers has a dramatic effect on the entire food-web, such as the distribution of plant biomass.[23]

Energy flow and biomass

EnergyFlowFrog
EnergyFlowTransformity

EnergyFlowFrog
EnergyFlowTransformity

Food webs depict energy flow via trophic linkages. Energy flow is directional, which contrasts against the cyclic flows of material through the food web systems.[26] Energy flow "typically includes production, consumption, assimilation, non-assimilation losses (feces), and respiration (maintenance costs)."[5]:5 In a very general sense, energy flow (E) can be defined as the sum of metabolic production (P) and respiration (R), such that E=P+R.

Biomass represents stored energy. However, concentration and quality of nutrients and energy is variable. Many plant fibers, for example, are indigestible to many herbivores leaving grazer community food webs more nutrient limited than detrital food webs where bacteria are able to access and release the nutrient and energy stores.[27][28] "Organisms usually extract energy in the form of carbohydrates, lipids, and proteins. These polymers have a dual role as supplies of energy as well as building blocks; the part that functions as energy supply results in the production of nutrients (and carbon dioxide, water, and heat). Excretion of nutrients is, therefore, basic to metabolism."[28]:1230–1231 The units in energy flow webs are typically a measure mass or energy per m2 per unit time. Different consumers are going to have different metabolic assimilation efficiencies in their diets. Each trophic level transforms energy into biomass. Energy flow diagrams illustrate the rates and efficiency of transfer from one trophic level into another and up through the hierarchy.[29][30]

It is the case that the biomass of each trophic level decreases from the base of the chain to the top. This is because energy is lost to the environment with each transfer as entropy increases. About eighty to ninety percent of the energy is expended for the organism’s life processes or is lost as heat or waste. Only about ten to twenty percent of the organism’s energy is generally passed to the next organism.[31] The amount can be less than one percent in animals consuming less digestible plants, and it can be as high as forty percent in zooplankton consuming phytoplankton.[32] Graphic representations of the biomass or productivity at each tropic level are called ecological pyramids or trophic pyramids. The transfer of energy from primary producers to top consumers can also be characterized by energy flow diagrams.[33]

Food chain

A common metric used to quantify food web trophic structure is food chain length. Food chain length is another way of describing food webs as a measure of the number of species encountered as energy or nutrients move from the plants to top predators.[34]:269 There are different ways of calculating food chain length depending on what parameters of the food web dynamic are being considered: connectance, energy, or interaction.[34] In its simplest form, the length of a chain is the number of links between a trophic consumer and the base of the web. The mean chain length of an entire web is the arithmetic average of the lengths of all chains in a food web.[35][12]

In a simple predator-prey example, a deer is one step removed from the plants it eats (chain length = 1) and a wolf that eats the deer is two steps removed from the plants (chain length = 2). The relative amount or strength of influence that these parameters have on the food web address questions about:

  • the identity or existence of a few dominant species (called strong interactors or keystone species)
  • the total number of species and food-chain length (including many weak interactors) and
  • how community structure, function and stability is determined.[36][37]

Ecological pyramids

Trophiclevels
TrophicEnergy

EcologicalPyramids

Trophiclevels
TrophicEnergy
EcologicalPyramids

In a pyramid of numbers, the number of consumers at each level decreases significantly, so that a single top consumer, (e.g., a polar bear or a human), will be supported by a much larger number of separate producers. There is usually a maximum of four or five links in a food chain, although food chains in aquatic ecosystems are more often longer than those on land. Eventually, all the energy in a food chain is dispersed as heat.[4]

Ecological pyramids place the primary producers at the base. They can depict different numerical properties of ecosystems, including numbers of individuals per unit of area, biomass (g/m2), and energy (k cal m−2 yr−1). The emergent pyramidal arrangement of trophic levels with amounts of energy transfer decreasing as species become further removed from the source of production is one of several patterns that is repeated amongst the planets ecosystems.[2]\[3][38] The size of each level in the pyramid generally represents biomass, which can be measured as the dry weight of an organism.[39] Autotrophs may have the highest global proportion of biomass, but they are closely rivaled or surpassed by microbes.[40][41]

Pyramid structure can vary across ecosystems and across time. In some instances biomass pyramids can be inverted. This pattern is often identified in aquatic and coral reef ecosystems. The pattern of biomass inversion is attributed to different sizes of producers. Aquatic communities are often dominated by producers that are smaller than the consumers that have high growth rates. Aquatic producers, such as planktonic algae or aquatic plants, lack the large accumulation of secondary growth as exists in the woody trees of terrestrial ecosystems. However, they are able to reproduce quickly enough to support a larger biomass of grazers. This inverts the pyramid. Primary consumers have longer lifespans and slower growth rates that accumulates more biomass than the producers they consume. Phytoplankton live just a few days, whereas the zooplankton eating the phytoplankton live for several weeks and the fish eating the zooplankton live for several consecutive years.[42] Aquatic predators also tend to have a lower death rate than the smaller consumers, which contributes to the inverted pyramidal pattern. Population structure, migration rates, and environmental refuge for prey are other possible causes for pyramids with biomass inverted. Energy pyramids, however, will always have an upright pyramid shape if all sources of food energy are included and this is dictated by the second law of thermodynamics.[4][43]

Material flux and recycling

Many of the Earth's elements and minerals (or mineral nutrients) are contained within the tissues and diets of organisms. Hence, mineral and nutrient cycles trace food web energy pathways. Ecologists employ stoichiometry to analyze the ratios of the main elements found in all organisms: carbon (C), nitrogen (N), phosphorus (P). There is a large transitional difference between many terrestrial and aquatic systems as C:P and C:N ratios are much higher in terrestrial systems while N:P ratios are equal between the two systems.[44][45][46] Mineral nutrients are the material resources that organisms need for growth, development, and vitality. Food webs depict the pathways of mineral nutrient cycling as they flow through organisms.[4][16] Most of the primary production in an ecosystem is not consumed, but is recycled by detritus back into useful nutrients.[47] Many of the Earth's microorganisms are involved in the formation of minerals in a process called biomineralization.[48][49][50] Bacteria that live in detrital sediments create and cycle nutrients and biominerals.[51] Food web models and nutrient cycles have traditionally been treated separately, but there is a strong functional connection between the two in terms of stability, flux, sources, sinks, and recycling of mineral nutrients.[52][53]

Kinds of food webs

Food webs are necessarily aggregated and only illustrate a tiny portion of the complexity of real ecosystems. For example, the number of species on the planet are likely in the general order of 107, over 95% of these species consist of microbes and invertebrates, and relatively few have been named or classified by taxonomists.[54][55][56] It is explicitly understood that natural systems are 'sloppy' and that food web trophic positions simplify the complexity of real systems that sometimes overemphasize many rare interactions. Most studies focus on the larger influences where the bulk of energy transfer occurs.[17] "These omissions and problems are causes for concern, but on present evidence do not present insurmountable difficulties."[3]:669

Food web and trophic level of the Chengjiang and Burgess Shale - journal.pbio.0060102.g001
Paleoecological studies can reconstruct fossil food-webs and trophic levels. Primary producers form the base (red spheres), predators at top (yellow spheres), the lines represent feeding links. Original food-webs (left) are simplified (right) by aggregating groups feeding on common prey into coarser grained trophic species.[57]

There are different kinds or categories of food webs:

  • Source web - one or more node(s), all of their predators, all the food these predators eat, and so on.
  • Sink web - one or more node(s), all of their prey, all the food that these prey eat, and so on.
  • Community (or connectedness) web - a group of nodes and all the connections of who eats whom.
  • Energy flow web - quantified fluxes of energy between nodes along links between a resource and a consumer.[3][39]
  • Paleoecological web - a web that reconstructs ecosystems from the fossil record.[57]
  • Functional web - emphasizes the functional significance of certain connections having strong interaction strength and greater bearing on community organization, more so than energy flow pathways. Functional webs have compartments, which are sub-groups in the larger network where there are different densities and strengths of interaction.[37][58] Functional webs emphasize that "the importance of each population in maintaining the integrity of a community is reflected in its influence on the growth rates of other populations."[39]:511

Within these categories, food webs can be further organized according to the different kinds of ecosystems being investigated. For example, human food webs, agricultural food webs, detrital food webs, marine food webs, aquatic food webs, soil food webs, Arctic (or polar) food webs, terrestrial food webs, and microbial food webs. These characterizations stem from the ecosystem concept, which assumes that the phenomena under investigation (interactions and feedback loops) are sufficient to explain patterns within boundaries, such as the edge of a forest, an island, a shoreline, or some other pronounced physical characteristic.[59][60][61]

Detrital web

In a detrital web, plant and animal matter is broken down by decomposers, e.g., bacteria and fungi, and moves to detritivores and then carnivores.[62] There are often relationships between the detrital web and the grazing web. Mushrooms produced by decomposers in the detrital web become a food source for deer, squirrels, and mice in the grazing web. Earthworms eaten by robins are detritivores consuming decaying leaves.[63]

Soil food webUSDA
An illustration of a soil food web.

"Detritus can be broadly defined as any form of non-living organic matter, including different types of plant tissue (e.g. leaf litter, dead wood, aquatic macrophytes, algae), animal tissue (carrion), dead microbes, faeces (manure, dung, faecal pellets, guano, frass), as well as products secreted, excreted or exuded from organisms (e.g. extra-cellular polymers, nectar, root exudates and leachates, dissolved organic matter, extra-cellular matrix, mucilage). The relative importance of these forms of detritus, in terms of origin, size and chemical composition, varies across ecosystems."[47]:585

Quantitative food webs

Ecologists collect data on trophic levels and food webs to statistically model and mathematically calculate parameters, such as those used in other kinds of network analysis (e.g., graph theory), to study emergent patterns and properties shared among ecosystems. There are different ecological dimensions that can be mapped to create more complicated food webs, including: species composition (type of species), richness (number of species), biomass (the dry weight of plants and animals), productivity (rates of conversion of energy and nutrients into growth), and stability (food webs over time). A food web diagram illustrating species composition shows how change in a single species can directly and indirectly influence many others. Microcosm studies are used to simplify food web research into semi-isolated units such as small springs, decaying logs, and laboratory experiments using organisms that reproduce quickly, such as daphnia feeding on algae grown under controlled environments in jars of water.[36][64]

While the complexity of real food webs connections are difficult to decipher, ecologists have found mathematical models on networks an invaluable tool for gaining insight into the structure, stability, and laws of food web behaviours relative to observable outcomes. "Food web theory centers around the idea of connectance."[65]:1648 Quantitative formulas simplify the complexity of food web structure. The number of trophic links (tL), for example, is converted into a connectance value:

,

where, S(S-1)/2 is the maximum number of binary connections among S species.[65] "Connectance (C) is the fraction of all possible links that are realized (L/S2) and represents a standard measure of food web complexity..."[66]:12913 The distance (d) between every species pair in a web is averaged to compute the mean distance between all nodes in a web (D)[66] and multiplied by the total number of links (L) to obtain link-density (LD), which is influenced by scale dependent variables such as species richness. These formulas are the basis for comparing and investigating the nature of non-random patterns in the structure of food web networks among many different types of ecosystems.[66][67]

Scaling laws, complexity, choas, and patterned correlates are common features attributed to food web structure.[68][69]

Complexity and stability

Food webs are complex. Complexity is a measure of an increasing number of permutations and it is also a metaphorical term that conveys the mental intractability or limits concerning unlimited algorithmic possibilities. In food web terminology, complexity is a product of the number of species and connectance.[70][71][72] Connectance is "the fraction of all possible links that are realized in a network".[73]:12917 These concepts were derived and stimulated through the suggestion that complexity leads to stability in food webs, such as increasing the number of trophic levels in more species rich ecosystems. This hypothesis was challenged through mathematical models suggesting otherwise, but subsequent studies have shown that the premise holds in real systems.[70][74]

At different levels in the hierarchy of life, such as the stability of a food web, "the same overall structure is maintained in spite of an ongoing flow and change of components."[75]:476 The farther a living system (e.g., ecosystem) sways from equilibrium, the greater its complexity.[75] Complexity has multiple meanings in the life sciences and in the public sphere that confuse its application as a precise term for analytical purposes in science.[72][76] Complexity in the life sciences (or biocomplexity) is defined by the "properties emerging from the interplay of behavioral, biological, physical, and social interactions that affect, sustain, or are modified by living organisms, including humans".[77]:1018

Several concepts have emerged from the study of complexity in food webs. Complexity explains many principals pertaining to self-organization, non-linearity, interaction, cybernetic feedback, discontinuity, emergence, and stability in food webs. Nestedness, for example, is defined as "a pattern of interaction in which specialists interact with species that form perfect subsets of the species with which generalists interact",[78]:575 "—that is, the diet of the most specialized species is a subset of the diet of the next more generalized species, and its diet a subset of the next more generalized, and so on."[79] Until recently, it was thought that food webs had little nested structure, but empirical evidence shows that many published webs have nested subwebs in their assembly.[80]

Food webs are complex networks. As networks, they exhibit similar structural properties and mathematical laws that have been used to describe other complex systems, such as small world and scale free properties. The small world attribute refers to the many loosely connected nodes, non-random dense clustering of a few nodes (i.e., trophic or keystone species in ecology), and small path length compared to a regular lattice.[73][81] "Ecological networks, especially mutualistic networks, are generally very heterogeneous, consisting of areas with sparse links among species and distinct areas of tightly linked species. These regions of high link density are often referred to as cliques, hubs, compartments, cohesive sub-groups, or modules...Within food webs, especially in aquatic systems, nestedness appears to be related to body size because the diets of smaller predators tend to be nested subsets of those of larger predators (Woodward & Warren 2007; YvonDurocher et al. 2008), and phylogenetic constraints, whereby related taxa are nested based on their common evolutionary history, are also evident (Cattin et al. 2004)."[82]:257 "Compartments in food webs are subgroups of taxa in which many strong interactions occur within the subgroups and few weak interactions occur between the subgroups. Theoretically, compartments increase the stability in networks, such as food webs."[58]

Food webs are also complex in the way that they change in scale, seasonally, and geographically. The components of food webs, including organisms and mineral nutrients, cross the thresholds of ecosystem boundaries. This has led to the concept or area of study known as cross-boundary subsidy.[59][60] "This leads to anomalies, such as food web calculations determining that an ecosystem can support one half of a top carnivore, without specifying which end."[61] Nonetheless, real differences in structure and function have been identified when comparing different kinds of ecological food webs, such as terrestrial vs. aquatic food webs.[83]

History of food webs

EltonFW
Victor Summerhayes and Charles Elton's 1923 food web of Bear Island (Arrows point to an organism being consumed by another organism).

Food webs serve as a framework to help ecologists organize the complex network of interactions among species observed in nature and around the world. One of the earliest descriptions of a food chain was described by a medieval Afro-Arab scholar named Al-Jahiz: "All animals, in short, cannot exist without food, neither can the hunting animal escape being hunted in his turn."[84]:143 The earliest graphical depiction of a food web was by Lorenzo Camerano in 1880, followed independently by those of Pierce and colleagues in 1912 and Victor Shelford in 1913.[85][86] Two food webs about herring were produced by Victor Summerhayes and Charles Elton[87] and Alister Hardy[88] in 1923 and 1924. Charles Elton subsequently pioneered the concept of food cycles, food chains, and food size in his classical 1927 book "Animal Ecology"; Elton's 'food cycle' was replaced by 'food web' in a subsequent ecological text.[89] After Charles Elton's use of food webs in his 1927 synthesis,[90] they became a central concept in the field of ecology. Elton[89] organized species into functional groups, which formed the basis for the trophic system of classification in Raymond Lindeman's classic and landmark paper in 1942 on trophic dynamics.[16][37][91] The notion of a food web has a historical foothold in the writings of Charles Darwin and his terminology, including an "entangled bank", "web of life", "web of complex relations", and in reference to the decomposition actions of earthworms he talked about "the continued movement of the particles of earth". Even earlier, in 1768 John Bruckner described nature as "one continued web of life".[3][92][93][94]

Interest in food webs increased after Robert Paine's experimental and descriptive study of intertidal shores[95] suggesting that food web complexity was key to maintaining species diversity and ecological stability. Many theoretical ecologists, including Sir Robert May[96] and Stuart Pimm,[97] were prompted by this discovery and others to examine the mathematical properties of food webs.

See also

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Further reading

  • Cohen, Joel E. (1978). Food webs and niche space. Monographs in Population Biology. 11. Princeton, NJ: Princeton University Press. pp. xv+1–190. ISBN 978-0-691-08202-8.
  • "Aquatic Food Webs". NOAA Education Resources. National Oceanic and Atmospheric Administration.
Cascade effect (ecology)

An ecological cascade effect is a series of secondary extinctions that is triggered by the primary extinction of a key species in an ecosystem. Secondary extinctions are likely to occur when the threatened species are: dependent on a few specific food sources, mutualistic (dependent on the key species in some way), or forced to coexist with an invasive species that is introduced to the ecosystem. Species introductions to a foreign ecosystem can often devastate entire communities, and even entire ecosystems. These exotic species monopolize the ecosystem's resources, and since they have no natural predators to decrease their growth, they are able to increase indefinitely. Olsen et al. showed that exotic species have caused lake and estuary ecosystems to go through cascade effects due to loss of algae, crayfish, mollusks, fish, amphibians, and birds. However, the principal cause of cascade effects is the loss of top predators as the key species. As a result of this loss, a dramatic increase (ecological release) of prey species occurs. The prey is then able to overexploit its own food resources, until the population numbers decrease in abundance, which can lead to extinction. When the prey's food resources disappear, they starve and may go extinct as well. If the prey species is herbivorous, then their initial release and exploitation of the plants may result in a loss of plant biodiversity in the area. If other organisms in the ecosystem also depend upon these plants as food resources, then these species may go extinct as well. An example of the cascade effect caused by the loss of a top predator is apparent in tropical forests. When hunters cause local extinctions of top predators, the predators' prey's population numbers increase, causing an overexploitation of a food resource and a cascade effect of species loss. Recent studies have been performed on approaches to mitigate extinction cascades in food-web networks.

Cross-boundary subsidy

Cross-boundary subsidies are caused by organisms or materials that cross or traverse habitat patch boundaries, subsidizing the resident populations. The transferred organisms and materials may provide additional predators, prey, or nutrients to resident species, which can affect community and food web structure. Cross-boundary subsidies of materials and organisms occur in landscapes composed of different habitat patch types, and so depend on characteristics of those patches and on the boundaries in between them. Human alteration of the landscape, primarily through fragmentation, has the potential to alter important cross-boundary subsidies to increasingly isolated habitat patches. Understanding how processes that occur outside of habitat patches can affect populations within them may be important to habitat management.

Ecological collapse

Ecological collapse refers to a situation where an ecosystem suffers a drastic, possibly permanent, reduction in carrying capacity for all organisms, often resulting in mass extinction. Usually, an ecological collapse is precipitated by a disastrous event occurring on a short time scale. Ecological collapse can be considered as a consequence of ecosystem collapse on the biotic elements that depended on the original ecosystem.Ecosystems have the ability to rebound from a disruptive agent. The difference between collapse or a gentle rebound is determined by two factors—the toxicity of the introduced element and the resiliency of the original ecosystem.Through natural selection the planet's species have continuously adapted to change through variation in their biological composition and distribution. Mathematically it can be demonstrated that greater numbers of different biological factors tend to dampen fluctuations in each of the individual factors.Scientists can predict tipping points for ecological collapse. The most frequently used model for predicting food web collapse is called R50, which is a reliable measurement model for food web robustness.

Ecological network

An ecological network is a representation of the biotic interactions in an ecosystem, in which species (nodes) are connected by pairwise interactions (links). These interactions can be trophic or symbiotic. Ecological networks are used to describe and compare the structures of real ecosystems, while network models are used to investigate the effects of network structure on properties such as ecosystem stability.

Ecological stoichiometry

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. 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.

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:

why do elemental imbalances arise in nature?

how is consumer physiology 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 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, cancer research, food web interactions, population dynamics, ecosystem services, productivity of agricultural crops and honeybee nutrition.

Ecology of the San Francisco Estuary

The San Francisco Estuary together with the Sacramento–San Joaquin River Delta represents a highly altered ecosystem. The region has been heavily re-engineered to accommodate the needs of water delivery, shipping, agriculture, and most recently, suburban development. These needs have wrought direct changes in the movement of water and the nature of the landscape, and indirect changes from the introduction of non-native species. New species have altered the architecture of the food web as surely as levees have altered the landscape of islands and channels that form the complex system known as the Delta.This article deals particularly with the ecology of the low salinity zone (LSZ) of the estuary. Reconstructing a historic food web for the LSZ is difficult for a number of reasons. First, there is no clear record of the species that historically have occupied the estuary. Second, the San Francisco Estuary and Delta have been in geologic and hydrologic transition for most of their 10,000 year history, and so describing the "natural" condition of the estuary is much like "hitting a moving target". Climate change, hydrologic engineering, shifting water needs, and newly introduced species will continue to alter the food web configuration of the estuary. This model provides a snapshot of the current state, with notes about recent changes or species introductions that have altered the configuration of the food web. Understanding the dynamics of the current food web may prove useful for restoration efforts to improve the functioning and species diversity of the estuary.

Fishing down the food web

Fishing down the food web is the process whereby fisheries in a given ecosystem, "having depleted the large predatory fish on top of the food web, turn to increasingly smaller species, finally ending up with previously spurned small fish and invertebrates".The process was first demonstrated by the fisheries scientist Daniel Pauly and others in an article published in the journal Science in 1998. Large predator fish with higher trophic levels have been depleted in wild fisheries. As a result, the fishing industry has been systematically "fishing down the food web", targeting fish species at progressively decreasing trophic levels.

The trophic level of a fish is the position it occupies on the food chain. The article establishes the importance of the mean trophic level of fisheries as a tool for measuring the health of ocean ecosystems. In 2000, the Convention on Biological Diversity selected the mean trophic level of fisheries catch, renamed the "Marine Trophic Index" (MTI), as one of eight indicators of ecosystem health. However, many of the world's most lucrative fisheries are crustacean and mollusk fisheries, which are at low trophic levels and thus result in lower MTI values.

Food chain

A food chain is a linear network of links in a food web starting from producer organisms (such as grass or trees which use radiation from the Sun to make their food) and ending at apex predator species (like grizzly bears or killer whales), detritivores (like earthworms or woodlice), or decomposer species (such as fungi or bacteria). A food chain also shows how the organisms are related with each other by the food they eat. Each level of a food chain represents a different trophic level. A food chain differs from a food web, because the complex network of different animals' feeding relations are aggregated and the chain only follows a direct, linear pathway of one animal at a time. Natural interconnections between food chains make it a food web.

A common metric used to the quantify food web trophic structure is food chain length. In its simplest form, the length of a chain is the number of links between a trophic consumer and the base of the web and the mean chain length of an entire web is the arithmetic average of the lengths of all chains in a food web.Many food webs have a keystone species. A keystone species is a species that has a large impact on the surrounding environment and can directly affect the food chain. If this keystone species dies off it can set the entire food chain off balance. Keystone species keep herbivores from depleting all of the foliage in their environment and preventing a mass extinction.Food chains were first introduced by the Arab scientist and philosopher Al-Jahiz in the 10th century and later popularized in a book published in 1927 by Charles Elton, which also introduced the food web concept.

Forage fish

Forage fish, also called prey fish or bait fish, are small pelagic fish which are preyed on by larger predators for food. Predators include other larger fish, seabirds and marine mammals. Typical ocean forage fish feed near the base of the food chain on plankton, often by filter feeding. They include particularly fishes of the family Clupeidae (herrings, sardines, shad, hilsa, menhaden, anchovies, and sprats), but also other small fish, including halfbeaks, silversides, smelt such as capelin, and the goldband fusiliers pictured on the right.

Forage fish compensate for their small size by forming schools. Some swim in synchronised grids with their mouths open so they can efficiently filter plankton. These schools can become immense shoals which move along coastlines and migrate across open oceans. The shoals are concentrated energy resources for the great marine predators. The predators are keenly focused on the shoals, acutely aware of their numbers and whereabouts, and make migrations themselves that can span thousands of miles to connect, or stay connected, with them.The ocean primary producers, mainly contained in plankton, produce food energy from the sun and are the raw fuel for the ocean food webs. Forage fish transfer this energy by eating the plankton and becoming food themselves for the top predators. In this way, forage fish occupy the central positions in ocean and lake food webs.The fishing industry catches forage fish primarily for feeding to farmed animals. Some fisheries scientists are expressing concern that this will affect the populations of predator fish that depend on them.

Intertidal ecology

Intertidal ecology is the study of intertidal ecosystems, where organisms live between the low and high tide lines. At low tide, the intertidal is exposed whereas at high tide, the intertidal is underwater. Intertidal ecologists therefore study the interactions between intertidal organisms and their environment, as well as between different species of intertidal organisms within a particular intertidal community. The most important environmental and species interactions may vary based on the type of intertidal community being studied, the broadest of classifications being based on substrates—rocky shore and soft bottom communities.Organisms living in this zone have a highly variable and often hostile environment, and have evolved various adaptations to cope with and even exploit these conditions. One easily visible feature of intertidal communities is vertical zonation, where the community is divided into distinct vertical bands of specific species going up the shore. Species ability to cope with abiotic factors associated with emersion stress, such as desiccation determines their upper limits, while biotic interactions e.g.competition with other species sets their lower limits.Intertidal regions are utilized by humans for food and recreation, but anthropogenic actions also have major impacts, with overexploitation, invasive species and climate change being among the problems faced by intertidal communities. In some places Marine Protected Areas have been established to protect these areas and aid in scientific research.

Marine life

Marine life, or sea life or ocean life, is the plants, animals and other organisms that live in the salt water of the sea or ocean, or the brackish water of coastal estuaries. At a fundamental level, marine life affects the nature of the planet. Marine organisms produce oxygen. Shorelines are in part shaped and protected by marine life, and some marine organisms even help create new land.

Most life forms evolved initially in marine habitats. By volume, oceans provide about 90 percent of the living space on the planet. The earliest vertebrates appeared in the form of fish, which live exclusively in water. Some of these evolved into amphibians which spend portions of their lives in water and portions on land. Other fish evolved into land mammals and subsequently returned to the ocean as seals, dolphins or whales. Plant forms such as kelp and algae grow in the water and are the basis for some underwater ecosystems. Plankton, and particularly phytoplankton, are key primary producers forming the general foundation of the ocean food chain.

Marine invertebrates exhibit a wide range of modifications to survive in poorly oxygenated waters, including breathing tubes as in mollusc siphons. Fish have gills instead of lungs, although some species of fish, such as the lungfish, have both. Marine mammals, such as dolphins, whales, otters, and seals need to surface periodically to breathe air.

A total of 230,000 documented marine species exist, including about 20,000 species of marine fish, with some two million marine species yet to be documented. Marine species range in size from the microscopic, including plankton and phytoplankton which can be as small as 0.02 micrometres, to huge cetaceans (whales, dolphins and porpoises), including the blue whale – the largest known animal reaching up to 33 metres (108 ft) in length. Marine microorganisms, including bacteria and viruses, constitute about 70% of the total marine biomass.

Mesopredator

A mesopredator is a mid-ranking predator in the middle of a trophic level, which typically preys on smaller animals. Mesopredators often vary in ecosystems depending on the food web. It is also important to note that there is no specific size or weight restrictions to qualify as a mesopredator, as it depends on how large the apex predator is, and what the mesopredator's prey is. When new species are introduced into an ecosystem, the role of mesopredator often changes; the same happens if a species is removed.

Microalgae

Microalgae or microphytes are microscopic algae, typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist individually, or in chains or groups. Depending on the species, their sizes can range from a few micrometers (µm) to a few hundred micrometers. Unlike higher plants, microalgae do not have roots, stems, or leaves. They are specially adapted to an environment dominated by viscous forces. Microalgae, capable of performing photosynthesis, are important for life on earth; they produce approximately half of the atmospheric oxygen and use simultaneously the greenhouse gas carbon dioxide to grow photoautotrophically. Microalgae, together with bacteria, form the base of the food web and provide energy for all the trophic levels above them. Microalgae biomass is often measured with chlorophyll a concentrations and can provide a useful index of potential production. The standing stock of microphytes is closely related to that of its predators. Without grazing pressures the standing stock of microphytes dramatically decreases.The biodiversity of microalgae is enormous and they represent an almost untapped resource. It has been estimated that about 200,000-800,000 species in many different genera exist of which about 50,000 species are described. Over 15,000 novel compounds originating from algal biomass have been chemically determined. Most of these microalgae species produce unique products like carotenoids, antioxidants, fatty acids, enzymes, polymers, peptides, toxins and sterols.

Microbial food web

The microbial food web refers to the combined trophic interactions among microbes in aquatic environments. These microbes include viruses, bacteria, algae, heterotrophic protists (such as ciliates and flagellates).

In aquatic environments, microbes constitute the base of the food web. Single celled photosynthetic organisms such as diatoms and cyanobacteria are generally the most important primary producers in the open ocean. Many of these cells, especially cyanobacteria, are too small to be captured and consumed by small crustaceans and planktonic larvae. Instead, these cells are consumed by phagotrophic protists which are readily consumed by larger organisms. Viruses can infect and break open bacterial cells and (to a lesser extent), planktonic algae (a.k.a. phytoplankton). Therefore, viruses in the microbial food web act to reduce the population of bacteria and, by lysing bacterial cells, release particulate and dissolved organic carbon (DOC). DOC may also be released into the environment by algal cells. One of the reasons phytoplankton release DOC termed "unbalanced growth" is when essential nutrients (e.g. nitrogen and phosphorus) are limiting. Therefore, carbon produced during photosynthesis is not used for the synthesis of proteins (and subsequent cell growth), but is limited due of a lack of the nutrients necessary for macromolecules. Excess photosynthate, or DOC is then released, or exuded.

The microbial loop describes a pathway in the microbial food web where DOC is returned to higher trophic levels via the incorporation into bacterial biomass.

Microbial loop

The microbial loop describes a trophic pathway in the marine microbial food web where dissolved organic carbon (DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by phytoplankton-zooplankton-nekton. The term microbial loop was coined by Farooq Azam and Tom Fenchel et al. to include the role played by bacteria in the carbon and nutrient cycles of the marine environment.

In general, dissolved organic carbon (DOC) is introduced into the ocean environment from bacterial lysis, the leakage or exudation of fixed carbon from phytoplankton (e.g., mucilaginous exopolymer from diatoms), sudden cell senescence, sloppy feeding by zooplankton, the excretion of waste products by aquatic animals, or the breakdown or dissolution of organic particles from terrestrial plants and soils (Van den Meersche et al. 2004). Bacteria in the microbial loop decompose this particulate detritus to utilize this energy-rich matter for growth. Since more than 95% of organic matter in marine ecosystems consists of polymeric, high molecular weight (HMW) compounds (e.g., protein, polysaccharides, lipids), only a small portion of total dissolved organic matter (DOM) is readily utilizable to most marine organisms at higher trophic levels. This means that dissolved organic carbon is not available directly to most marine organisms; marine bacteria introduce this organic carbon into the food web, resulting in additional energy becoming available to higher trophic levels. Recently the term "microbial food web" has been substituted for the term "microbial loop".

Soil ecology

Soil ecology is the study of the interactions among soil biology, and between biotic and abiotic aspects of the soil environment. It is particularly concerned with the cycling of nutrients, formation and stabilization of the pore structure, the spread and vitality of pathogens, and the biodiversity of this rich biological community.

Soil food web

The soil food web is the community of organisms living all or part of their lives in the soil. It describes a complex living system in the soil and how it interacts with the environment, plants, and animals.

Food webs describe the transfer of energy between species in an ecosystem. While a food chain examines one, linear, energy pathway through an ecosystem, a food web is more complex and illustrates all of the potential pathways. Much of this transferred energy comes from the sun. Plants use the sun’s energy to convert inorganic compounds into energy-rich, organic compounds, turning carbon dioxide and minerals into plant material by photosynthesis. Plant flowers exude energy-rich nectar above ground and plant roots exude acids, sugars, and ectoenzymes into the rhizosphere, adjusting the pH and feeding the food web underground.Plants are called autotrophs because they make their own energy; they are also called producers because they produce energy available for other organisms to eat. Heterotrophs are consumers that cannot make their own food. In order to obtain energy they eat plants or other heterotrophs.

Trophic cascade

Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce the abundance, or alter the behavior, of their prey, thereby releasing the next lower trophic level from predation (or herbivory if the intermediate trophic level is a herbivore).

The trophic cascade is an ecological concept which has stimulated new research in many areas of ecology. For example, it can be important for understanding the knock-on effects of removing top predators from food webs, as humans have done in many places through hunting and fishing.

A top-down cascade is a trophic cascade where the top consumer/predator controls the primary consumer population. In turn, the primary producer population thrives. The removal of the top predator can alter the food web dynamics. In this case, the primary consumers would overpopulate and exploit the primary producers. Eventually there would not be enough primary producers to sustain the consumer population. Top-down food web stability depends on competition and predation in the higher trophic levels. Invasive species can also alter this cascade by removing or becoming a top predator. This interaction may not always be negative. Studies have shown that certain invasive species have begun to shift cascades; and as a consequence, ecosystem degradation has been repaired.For example, if the abundance of large piscivorous fish is increased in a lake, the abundance of their prey, smaller fish that eat zooplankton, should decrease. The resulting increase in zooplankton should, in turn, cause the biomass of its prey, phytoplankton, to decrease.

In a bottom-up cascade, the population of primary producers will always control the increase/decrease of the energy in the higher trophic levels. Primary producers are plants, phytoplankton and zooplankton that require photosynthesis. Although light is important, primary producer populations are altered by the amount of nutrients in the system. This food web relies on the availability and limitation of resources. All populations will experience growth if there is initially a large amount of nutrients.In a subsidy cascade, species populations at one trophic level can be supplemented by external food. For example, native animals can forage on resources that don't originate in their same habitat, such a native predators eating livestock. This may increase their local abundances thereby affecting other species in the ecosystem and causing an ecological cascade. For example, Luskin et al (2017) found that native animals living in protected primary rainforest in Malaysia found food subsidies in neighboring oil palm plantations. This subsidy allowed native animal populations to increase, which then triggered powerful secondary ‘cascading’ effects on forest tree community. Specifically, crop-raiding wild boar (Sus scrofa) built thousands of nests from the forest understory vegetation and this caused a 62% decline in forest tree sapling density over a 24-year study period. Such cross-boundary subsidy cascades may be widespread in both terrestrial and marine ecosystems and present significant conservation challenges.

These trophic interactions shape patterns of biodiversity globally. Humans and climate change have affected these cascades drastically. One example can be seen with sea otters (Enhydra lutris) on the Pacific coast of the United States of America. Over time, human interactions caused a removal of sea otters. One of their main prey, the pacific purple sea urchin (Strongylocentrotus purpuratus) eventually began to overpopulate. The overpopulation caused increased predation of giant kelp (Macrocystis pyrifera). As a result, there was extreme deterioration of the kelp forests along the California coast. This is why it is important for countries to regulate marine and terrestrial ecosystems.Predator-induced interactions could heavily influence the flux of atmospheric carbon if managed on a global scale. For example, a study was conducted to determine the cost of potential stored carbon in living kelp biomass in Sea Otter enhanced ecosystems. The study valued the potential storage between $205 million and $408 million dollars (US) on the European Carbon Exchange (2012).

Trophic level

The trophic level of an organism is the position it occupies in a food chain. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it is from the start of the chain. A food chain starts at trophic level 1 with primary producers such as plants, can move to herbivores at level 2, carnivores at level 3 or higher, and typically finish with apex predators at level 4 or 5. The path along the chain can form either a one-way flow or a food "web". Ecological communities with higher biodiversity form more complex trophic paths.

The word trophic derives from the Greek τροφή (trophē) referring to food or nourishment.

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