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 concept of trophic level was developed by Raymond Lindeman (1942), based on the terminology of August Thienemann (1926): "producers", "consumers" and "reducers" (modified to "decomposers" by Lindeman).
The three basic ways in which organisms get food are as producers, consumers and decomposers.
Trophic levels can be represented by numbers, starting at level 1 with plants. Further trophic levels are numbered subsequently according to how far the organism is along the food chain.
In real world ecosystems, there is more than one food chain for most organisms, since most organisms eat more than one kind of food or are eaten by more than one type of predator. A diagram that sets out the intricate network of intersecting and overlapping food chains for an ecosystem is called its food web. Decomposers are often left off food webs, but if included, they mark the end of a food chain. Thus food chains start with primary producers and end with decay and decomposers. Since decomposers recycle nutrients, leaving them so they can be reused by primary producers, they are sometimes regarded as occupying their own trophic level.
The trophic level of a species may vary, if it has a choice of diet. Virtually all plants and phytoplankton are purely phototrophic and are at exactly level 1.0. Many worms are at around 2.1; insects 2.2; jellyfish 3.0; birds 3.6. A 2013 study estimates the average trophic level of human beings at 2.21, similar to pigs or anchovies. This is only an average, and plainly both modern and ancient human eating habits are complex and vary greatly. For example, a traditional Eskimo living on a diet consisting primarily of seals would have a trophic level of nearly 5.
In general, each trophic level relates to the one below it by absorbing some of the energy it consumes, and in this way can be regarded as resting on, or supported by, the next lower trophic level. Food chains can be diagrammed to illustrate the amount of energy that moves from one feeding level to the next in a food chain. This is called an energy pyramid. The energy transferred between levels can also be thought of as approximating to a transfer in biomass, so energy pyramids can also be viewed as biomass pyramids, picturing the amount of biomass that results at higher levels from biomass consumed at lower levels. However, when primary producers grow rapidly and are consumed rapidly, the biomass at any one moment may be low; for example, phytoplankton (producer) biomass can be low compared to the zooplankton (consumer) biomass in the same area of ocean.
The efficiency with which energy or biomass is transferred from one trophic level to the next is called the ecological efficiency. Consumers at each level convert on average only about 10% of the chemical energy in their food to their own organic tissue (the ten-percent law). For this reason, food chains rarely extend for more than 5 or 6 levels. At the lowest trophic level (the bottom of the food chain), plants convert about 1% of the sunlight they receive into chemical energy. It follows from this that the total energy originally present in the incident sunlight that is finally embodied in a tertiary consumer is about 0.001%
Both the number of trophic levels and the complexity of relationships between them evolve as life diversifies through time, the exception being intermittent mass extinction events.
Food webs largely define ecosystems, and the trophic levels define the position of organisms within the webs. But these trophic levels are not always simple integers, because organisms often feed at more than one trophic level. For example, some carnivores also eat plants, and some plants are carnivores. A large carnivore may eat both smaller carnivores and herbivores; the bobcat eats rabbits, but the mountain lion eats both bobcats and rabbits. Animals can also eat each other; the bullfrog eats crayfish and crayfish eat young bullfrogs. The feeding habits of a juvenile animal, and, as a consequence, its trophic level, can change as it grows up.
The fisheries scientist Daniel Pauly sets the values of trophic levels to one in plants and detritus, two in herbivores and detritivores (primary consumers), three in secondary consumers, and so on. The definition of the trophic level, TL, for any consumer species is:
where is the fractional trophic level of the prey j, and represents the fraction of j in the diet of i.
In the case of marine ecosystems, the trophic level of most fish and other marine consumers takes value between 2.0 and 5.0. The upper value, 5.0, is unusual, even for large fish, though it occurs in apex predators of marine mammals, such as polar bears and killer whales.
In addition to observational studies of animal behavior, and quantification of animal stomach contents, trophic level can be quantified through stable isotope analysis of animal tissues such as muscle, skin, hair, bone collagen. This is because there is a consistent increase in the nitrogen isotopic composition at each trophic level caused by fractionations that occur with the synthesis of biomolecules; the magnitude of this increase in nitrogen isotopic composition is approximately 3–4‰.
In fisheries, the mean trophic level for the fisheries catch across an entire area or ecosystem is calculated for year y as:
where is the catch of the species or group i in year y, and is the trophic level for species i as defined above.
Fish at higher trophic levels usually have a higher economic value, which can result in overfishing at the higher trophic levels. Earlier reports found precipitous declines in mean trophic level of fisheries catch, in a process known as fishing down the food web. However, more recent work finds no relation between economic value and trophic level; and that mean trophic levels in catches, surveys and stock assessments have not in fact declined, suggesting that fishing down the food web is not a global phenomenon. However Pauly et al. note that trophic levels peaked at 3.4 in 1970 in the northwest and west-central Atlantic, followed by a subsequent decline to 2.9 in 1994. They report a shift away from long-lived, piscivorous, high-trophic-level bottom fishes, such as cod and haddock, to short-lived, planktivorous, low-trophic-level invertebrates (e.g., shrimps) and small, pelagic fish (e.g., herrings). This shift from high-trophic-level fishes to low-trophic-level invertebrates and fishes is a response to changes in the relative abundance of the preferred catch. They argue this is part of the global fishery collapse.
Since biomass transfer efficiencies are only about 10%, it follows that the rate of biological production is much greater at lower trophic levels than it is at higher levels. Fisheries catches, at least to begin with, will tend to increase as the trophic level declines. At this point the fisheries will target species lower in the food web. In 2000, this led Pauly and others to construct a "Fisheries in Balance" index, usually called the FiB index. The FiB index is defined, for any year y, by
where is the catch at year y, is the mean trophic level of the catch at year y, is the catch, the mean trophic level of the catch at the start of the series being analyzed, and is the transfer efficiency of biomass or energy between trophic levels.
The FiB index is stable (zero) over periods of time when changes in trophic levels are matched by appropriate changes in the catch in the opposite direction. The index increases if catches increase for any reason, e.g. higher fish biomass, or geographic expansion. Such decreases explain the “backward-bending” plots of trophic level versus catch originally observed by Pauly and others in 1998.
One aspect of trophic levels is called tritrophic interaction. Ecologists often restrict their research to two trophic levels as a way of simplifying the analysis; however, this can be misleading if tritrophic interactions (such as plant–herbivore–predator) are not easily understood by simply adding pairwise interactions (plant–herbivore plus herbivore–predator, for example). Significant interactions can occur between the first trophic level (plant) and the third trophic level (a predator) in determining herbivore population growth, for example. Simple genetic changes may yield morphological variants in plants that then differ in their resistance to herbivores because of the effects of the plant architecture on enemies of the herbivore. Plants can also develop defenses against herbivores such as chemical defenses.
An apex predator, also known as an alpha predator or top predator, is a predator at the top of a food chain, with no natural predators.Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic levels. Food chains are often far shorter on land, usually limited to being secondary consumers – for example, wolves prey mostly upon large herbivores (primary consumers), which eat plants (primary producers). The apex predator concept is applied in wildlife management, conservation and ecotourism.
Apex predators have a long evolutionary history, dating at least to the Cambrian period when animals such as Anomalocaris dominated the seas.
Humans have for many centuries interacted with apex predators including the wolf, birds of prey and cormorants to hunt game animals, birds, and fish respectively. More recently, ecotourism such as with the tiger shark has become popular, and rewilding with predators such as the lynx has been proposed.Biomagnification
Biomagnification, also known as bioamplification or biological magnification, is the increasing concentration of a substance, such as a toxic chemical, in the tissues of tolerant organisms at successively higher levels in a food chain. This increase can occur as a result of:
Persistence – where the substance cannot be broken down by environmental processes
Food chain energetics – where the substance's concentration increases progressively as it moves up a food chain
Low or non-existent rate of internal degradation or excretion of the substance – often due to water-insolubility
Biological magnification often refers to the process whereby certain substances such as pesticides or heavy metals work their way into lakes, rivers and the ocean, and then move up the food chain in progressively greater concentrations as they are incorporated into the diet of aquatic organisms such as zooplankton, which in turn are eaten perhaps by fish, which then may be eaten by bigger fish, large birds, animals, or humans. The substances become increasingly concentrated in tissues or internal organs as they move up the chain. Bioaccumulants are substances that increase in concentration in living organisms as they take in contaminated air, water, or food because the substances are very slowly metabolized or excreted.Biomass (ecology)
The biomass is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms, plants or animals. The mass can be expressed as the average mass per unit area, or as the total mass in the community.
How biomass is measured depends on why it is being measured. Sometimes, the biomass is regarded as the natural mass of organisms in situ, just as they are. For example, in a salmon fishery, the salmon biomass might be regarded as the total wet weight the salmon would have if they were taken out of the water. In other contexts, biomass can be measured in terms of the dried organic mass, so perhaps only 30% of the actual weight might count, the rest being water. For other purposes, only biological tissues count, and teeth, bones and shells are excluded. In some applications, biomass is measured as the mass of organically bound carbon (C) that is present.
The total live biomass on Earth is about 550–560 billion tonnes C, and the total annual primary production of biomass is just over 100 billion tonnes C/yr. The total live biomass of bacteria may be as much as that of plants and animals or may be much less. The total number of DNA base pairs on Earth, as a possible approximation of global biodiversity, is estimated at (5.3±3.6)×1037, and weighs 50 billion tonnes. In comparison, the total mass of the biosphere has been estimated to be as much as 4×1012 tonnes of carbon.Calanoida
Calanoida is an order of copepods, a kind of zooplankton. They include around 46 families with about 1800 species of both marine and freshwater copepods. Calanoid copepods are dominant in the plankton in many parts of the world's oceans, making up 55%–95% of plankton samples. They are therefore important in many food webs, taking in energy from phytoplankton and algae and 'repackaging' it for consumption by higher trophic level predators. Many commercial fish are dependent on calanoid copepods for diet in either their larval or adult forms. Baleen whales such as bowhead whales, sei whales, right whales and fin whales eat calanoid copepods.Calanoids can be distinguished from other planktonic copepods by having first antennae at least half the length of the body and biramous second antennae. Their key defining feature anatomically, however, is the presence of a joint between the fifth and sixth body segments. The largest specimens reach 18 millimetres (0.71 in) long, but most are 0.5–2.0 mm (0.02–0.08 in) long.Clupeidae
Clupeidae is a family of ray-finned fishes, comprising, for instance, the herrings, shads, sardines, ilish, and menhadens. The clupeids include many of the most important food fishes in the world, and are also commonly caught for production of fish oil and fish meal. Many members of the family have a body protected with shiny cycloid scales (very smooth and uniform scales), a single dorsal fin, with a fusiform body for quick, evasive swimming and pursuit of prey composed of small planktonic animals. Due to their small size, and position in the lower trophic level of many marine food webs, the levels of methylmercury they bioaccumulate are very low, reducing the risk of mercury poisoning when consumed.Ecological efficiency
Ecological efficiency describes the efficiency with which energy is transferred from one trophic level to the next. It is determined by a combination of efficiencies relating to organismic resource acquisition and assimilation in an ecosystem.Ecological pyramid
An ecological pyramid (also trophic pyramid, eltonian pyramid, energy pyramid, or sometimes food pyramid) is a graphical representation designed to show the biomass or bio productivity at each trophic level in a given ecosystem.
Biomass pyramids show how much biomass (the amount of living or organic matter present in an organism) is present in the organisms at each trophic level, while productivity pyramids show the procreation or turnover in biomass. There is also pyramid of numbers which represent the number of organisms in each trophic level. They may be upright (e.g. Grassland ecosystem), inverted (parasitic ecosystem) or dumbbell shaped (forest ecosystem).
Energy pyramids begin with producers on the bottom (such as plants) and proceed through the various trophic levels (such as herbivores that eat plants, then carnivores that eat flesh, then omnivores that eat both plants and flesh, and so on). The highest level is the top of the food chain.Energy flow (ecology)
In ecology, energy flow, also called the calorific flow, refers to the flow of energy through a food chain, and is the focus of study in ecological energetics. In an ecosystem, ecologists seek to quantify the relative importance of different component species and feeding relationships.
A general energy flow scenario follows:
Solar energy is fixed by the photoautotrophs, called primary producers, like green plants. Primary consumers absorb most of the stored energy in the plant through digestion, and transform it into the form of energy they need, such as adenosine triphosphate (ATP), through respiration. A part of the energy received by primary consumers, herbivores, is converted to body heat (an effect of respiration), which is radiated away and lost from the system. The loss of energy through body heat is far greater in warm-blooded animals, which must eat much more frequently than those that are cold-blooded. Energy loss also occurs in the expulsion of undigested food (egesta) by excretion or regurgitation.
Secondary consumers, carnivores, then consume the primary consumers, although omnivores also consume primary producers. Energy that had been used by the primary consumers for growth and storage is thus absorbed into the secondary consumers through the process of digestion. As with primary consumers, secondary consumers convert this energy into a more suitable form (ATP) during respiration. Again, some energy is lost from the system, since energy which the primary consumers had used for respiration and regulation of body temperature cannot be utilized by the secondary consumers.
Tertiary consumers, which may or may not be apex predators, then consume the secondary consumers, with some energy passed on and some lost, as with the lower levels of the food chain.
A final link in the food chain are decomposers which break down the organic matter of the tertiary consumers (or whichever consumer is at the top of the chain) and release nutrients into the soil. They also break down plants, herbivores and carnivores that were not eaten by organisms higher on the food chain, as well as the undigested food that is excreted by herbivores and carnivores. Saprotrophic bacteria and fungi are decomposers, and play a pivotal role in the nitrogen and carbon cycles.The energy is passed on from trophic level to trophic level and each time about 90% of the energy is lost, with some being lost as heat into the environment (an effect of respiration) and some being lost as incompletely digested food (egesta). Therefore, primary consumers get about 10% of the energy produced by autotrophs, while secondary consumers get 1% and tertiary consumers get 0.1%. This means the top consumer of a food chain receives the least energy, as a lot of the food chain's energy has been lost between trophic levels. This loss of energy at each level limits typical food chains to only four to six links.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 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.Food chains were first introduced by the African-Arab scientist and philosopher Al-Jahiz in the 9th century and later popularized in a book published in 1927 by Charles Elton, which also introduced the food web concept.Foundation species
In ecology, the term foundation species is used to refer to a species that has a strong role in structuring a community. A foundation species can occupy any trophic level in a food web (i.e., they can be primary producers, herbivores or predators). The term was coined by Paul K. Dayton in 1972, who applied it to certain members of marine invertebrate and algae communities. It was clear from studies in several locations that there were a small handful of species whose activities had a disproportionate effect on the rest of the marine community and they were therefore key to the resilience of the community. Dayton’s view was that focusing on foundation species would allow for a simplified approach to more rapidly understand how a community as a whole would react to disturbances, such as pollution, instead of attempting the extremely difficult task of tracking the responses of all community members simultaneously. The term has since been applied to range of organisms in ecosystems around the world, in both aquatic and terrestrial environments. Aaron Ellison et al. introduced the term to terrestrial ecology by applying the term foundation species to tree species that define and structure certain forest ecosystems through their influences on associated organisms and modulation of ecosystem processes.Lake Waikare
Lake Waikare is the largest of several shallow lakes in the upper floodplain of the Waikato River in New Zealand's North Island. It is a riverine lake, located to the east of Te Kauwhata and 40 kilometres north of Hamilton. It covers 34 km².
Due to its shallow nature (its depth is never more than two metres) and the heavy use of fertiliser in the surrounding farming district, the waters of the lake are in poor condition. A 2010 report showed that of all the measured lakes it had the highest trophic level index, a measurement of the amount of pollutants.Lakes of New Zealand
In New Zealand there are 3,820 lakes with a surface area larger than one hectare. The lakes are of varying types and origins. Many of the lakes in the central North Island area are volcanic crater lakes, while the majority of the lakes near the Southern Alps were carved by glaciers. Hydroelectric reservoirs are common in South Canterbury, Central Otago and along the Waikato River.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 be qualified 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.Mesopredator release hypothesis
The mesopredator release hypothesis is an ecological theory used to describe the interrelated population dynamics between apex predators and mesopredators within an ecosystem, such that a collapsing population of the former results in dramatically-increased populations of the latter. This hypothesis describes the phenomenon of trophic cascade in specific terrestrial communities.
A mesopredator is a medium-sized, middle trophic level predator, which both preys and is preyed upon. Examples are raccoons, skunks, snakes, cownose rays, and small sharks.Primary producers
secondary producers convert an abiotic source of energy (e.g. light) into energy stored in organic compounds, which can be used by other organisms (e.g. heterotrophs). The primary producers can convert the energy in the light (phototroph and photoautotroph) or the energy in inorganic chemical compounds (chemolithotrophs) to build organic molecules, which is usually accumulated in the form of biomass and will be used as carbon and energy source by other organisms (e.g. heterotrophs and mixotrophs). The photoautotrophs are the main primary producers, converting the energy of the light into chemical energy through photosynthesis, ultimately building organic molecules from carbon dioxide, an inorganic carbon source. Examples of chemolithotrophs are some archaea and bacteria (unicellular organisms) that produce biomass from the oxidation of inorganic chemical compounds, these organisms are called chemoautotrophs, and are frequently found in hydrothermal vents in the deep ocean. Primary producers ares at the lowest trophic level, and are the reasons why Earth is sustainable for life to this day .Trophic
Trophic, from Ancient Greek τροφικός (trophikos) "pertaining to food or nourishment", may refer to:
Trophic factor receptor
Trophic level index
Trophic state index
Trophic webTrophic 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 may 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 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. Specically, crop-raiding wild boar (Sus scofa) built thousands of nests from the forest understory vegatation 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.Trophic level index
The trophic level index (TLI) is used in New Zealand as a measure of nutrient status of lakes. It is similar to the trophic state index but was proposed as alternative that suited New Zealand.The system uses four criteria, phosphorus and nitrogen concentrations, as well as visual clarity and algal biomass weighted equally.