Food web

{{quote box 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. 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. Trophic levels terrestrial ecosystem. The trophic pyramid roughly represents each level's biomass (usually measured as total dry weight). 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." 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 that no other species kills directly for their food resource needs. The intermediate levels are filled with omnivores that feed on more than one trophic level and cause energy to flow through several food pathways starting from a basal species. 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. It was once thought that omnivory was rare, but recent evidence suggests otherwise. This realization has made trophic classifications more complex. Like many wetlands, some ecosystems do not organize as a strict pyramid because aquatic plants are less productive than 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. Trophic dynamics and multitrophic interactions 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. 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." 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. These complex multitrophic interactions involve more than two trophic levels in a food web. For example, such interactions have been discovered in the context of arbuscular mycorrhizal fungi and aphid herbivores that utilize the same plant species. '' larvae sequester defensive compounds from specific types of plants they consume to protect themselves from bird predators Another example of a multitrophic 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. The field of chemical ecology has elucidated multitrophic interactions that entail the transfer of defensive compounds across multiple trophic levels. For example, certain plant species in the Castilleja and Plantago genera have been found to produce defensive compounds called iridoid glycosides that are sequestered in the tissues of the Taylor's checkerspot butterfly larvae that have developed a tolerance for these compounds and are able to consume the foliage of these plants. These sequestered iridoid glycosides then confer chemical protection against bird predators to the butterfly larvae. Energy flow and biomass in soil. This energy flow diagram illustrates how energy is lost as it fuels the metabolic process that transform the energy and nutrients into biomass. {{quote box 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. Energy flow "typically includes production, consumption, assimilation, non-assimilation losses (feces), and respiration (maintenance costs)." 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. "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." 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. 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. 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. 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. 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. 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. Ecological pyramids ecosystems exhibit inverted pyramids.Note: trophic levels are not drawn to scale and the pyramid of numbers excludes microorganisms and soil animals. Abbreviations: P=Producers, C1=Primary consumers, C2=Secondary consumers, C3=Tertiary consumers, S=Saprotrophs. The size of each level in the pyramid generally represents biomass, which can be measured as the dry weight of an organism. 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. 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. ==Material flux and recycling==