Detritus

For other uses, see Detritus (disambiguation).

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In biology, detritus (/dᵻˈtraɪtəs/) is dead particulate organic material (as opposed to dissolved organic material). It typically includes the bodies or fragments of dead organisms as well as fecal material. Detritus is typically colonized by communities of microorganisms which act to decompose (or remineralize) the material. In terrestrial ecosystems, it is encountered as leaf litter and other organic matter intermixed with soil, which is referred to as humus. Detritus of aquatic ecosystems is organic material suspended in water and piles up on seabed floors which is referred to as marine snow.

Theory

Giant sequoia (Sequoiadendron giganteum) cones and foliage, sugar pine and white fir foliage, and other plant litter cover the floor of Mariposa Grove in Yosemite National Park.

Dead plants or animals, material derived from animal tissues (such as skin cast off during moulting et cetera) gradually lose their form, due to both physical processes and the action of decomposers, including grazers, bacteria and fungi. Decomposition, the process through which organic matter is decomposed, takes place in many stages. Materials like proteins, lipids and sugars with low molecular weight are rapidly consumed and absorbed by micro-organisms and organisms that feed on dead matter. Other compounds, such as complex carbohydrates are broken down more slowly. The various micro-organisms involved in the decomposition break down the organic materials in order to gain the resources they require for their own survival and proliferation. Accordingly, at the same time that the materials of plants and animals are being broken down, the materials (biomass) making up the bodies of the micro-organisms are built up by a process of assimilation. When micro-organisms die, fine organic particles are produced, and if these are eaten by small animals which feed on micro-organisms, they will collect inside the intestine, and change shape into large pellets of dung. As a result of this process, most of the materials from dead organisms disappears from view and is not obviously present in any recognisable form, but is in fact present in the form of a combination of fine organic particles and the organisms using them as nutrients. This combination is detritus.

In ecosystems on land, detritus is deposited on the surface of the ground, taking forms such as the humic soil beneath a layer of fallen leaves. In aquatic ecosystems, most detritus is suspended in water, and gradually settles. In particular, many different types of material are collected together by currents, and much material settles in slowly flowing areas.

Much detritus is used as a source of nutrition for animals. In particular, many bottom feeding animals (benthos) living in mud flats feed in this way. In particular, since excreta are materials which other animals do not need, whatever energy value they might have, they are often unbalanced as a source of nutrients, and are not suitable as a source of nutrition on their own. However, there are many micro-organisms which multiply in natural environments. These micro-organisms do not simply absorb nutrients from these particles, but also shape their own bodies so that they can take the resources they lack from the area around them, and this allows them to make use of excreta as a source of nutrients. In practical terms, the most important constituents of detritus are complex carbohydrates, which are persistent (difficult to break down), and the micro-organisms which multiply using these absorb carbon from the detritus, and materials such as nitrogen and phosphorus from the water in their environment to synthesise the components of their own cells.

A characteristic type of food chain called the detritus cycle takes place involving detritus feeders (detritivores), detritus and the micro-organisms that multiply on it. For example, mud flats are inhabited by many univalves which are detritus feeders, such as moon shells. When these detritus feeders take in detritus with micro-organisms multiplying on it, they mainly break down and absorb the micro-organisms, which are rich in proteins, and excrete the detritus, which is mostly complex carbohydrates, having hardly broken it down at all. At first this dung is a poor source of nutrition, and so univalves pay no attention to it, but after several days, micro-organisms begin to multiply on it again, its nutritional balance improves, and so they eat it again. Through this process of eating the detritus many times over and harvesting the micro-organisms from it, the detritus thins out, becomes fractured and becomes easier for the micro-organisms to use, and so the complex carbohydrates are also steadily broken down and disappear over time.

What is left behind by the detritivores is then further broken down and recycled by decomposers, such as bacteria and fungi.

This detritus cycle plays a large part in the so-called purification process, whereby organic materials carried in by rivers is broken down and disappears, and an extremely important part in the breeding and growth of marine resources. In ecosystems on land, far more essential material is broken down as dead material passing through the detritus chain than is broken down by being eaten by animals in a living state. In both land and aquatic ecosystems, the role played by detritus is too large to ignore.

Aquatic ecosystems

In contrast to land ecosystems, dead materials and excreta in aquatic ecosystems do not settle immediately, and the finer the particles involved are, the longer they tend to take.

Terrestrial ecosystems

Detritus occurs in a variety of terrestrial habitats including forest, chaparral and grassland. In forests the detritus is typically dominated by leaf, twig, and bacteria litter as measured by biomass dominance. This plant litter provides important cover for seedling protection as well as cover for a variety of arthropods, reptiles^[1] and amphibians. Some insect larvae feed on the detritus.^[2] Fungi and bacteria continue the decomposition process^[3] after grazers have consumed larger elements of the organic materials, and animal trampling has assisted in mechanically breaking down organic matter. At the later stages of decomposition, mesophilic micro-organisms decompose residual detritus, generating heat from exothermic processes; such heat generation is associated with the well known phenomenon of the elevated temperature of composting.

Consumers

There is an extremely large number of detritus feeders in water. After all, a large quantity of material is carried in by water currents. Even if an organism stays in a fixed position, as long as it has a system for filtering water, it will be able to obtain enough food to get by. Many rooted organisms survive in this way, using developed gills or tentacles to filter the water to take in food, a process known as filter feeding.

Another more widely used method of feeding, which also incorporates filter feeding, is a system where an organism secretes mucus to catch the detritus in lumps, and then carries these to its mouth using an area of cilia. This is called mucus feeding.

Many organisms, including sea slugs and serpent's starfish, scoop up the detritus which has settled on the water bed. Bivalves which live inside the water bed do not simply suck in water through their tubes, but also extend them to fish for detritus on the surface of the bed.

Producers

In contrast, from the point of view of organisms using photosynthesis such as plants and plankton, detritus reduces the transparency of the water and gets in the way of this process. Given that these organisms also require a supply of nutrient salts—in other words fertilizer—for photosynthesis, their relationship with detritus is a complex one.

In land ecosystems, the waste products of plants and animals collect mainly on the ground (or on the surfaces of trees), and as decomposition proceeds, plants are supplied with fertiliser in the form of inorganic salts. In water ecosystems, relatively little waste collects on the water bed, and so the progress of decomposition in water takes a more important role. Investigating the level of inorganic salts in sea ecosystems shows that unless there is an especially large supply, the quantity increases from winter to spring—but is normally extremely low in summer. As such, the quantity of seaweed present reaches a peak in early summer and then decreases. The thinking is that organisms like plants grow quickly in warm periods and thus the quantity of inorganic salts is not enough to keep up with the demand. In other words, during winter, plant-like organisms are inactive and collect fertiliser, but if the temperature rises to some extent they will use this up in a very short period.

It is not entirely true that their productivity falls during the warmest periods. Organisms such as dinoflagellate have mobility, the ability to take in solid food, and the ability to photosynthesise. This type of micro-organism can take in substances such as detritus to grow, without waiting for it to be broken down into fertiliser.

Aquariums

In recent years, the word detritus has also come to be used in relation to aquariums (the word "aquarium" is a general term for any installation for keeping aquatic animals).

When animals such as fish are kept in an aquarium, substances such as excreta, mucus and dead skin cast off during moulting are produced by the animals and, naturally, generate detritus, and are continually broken down by micro-organisms.

Modern sealife aquariums often use the Berlin Method, which employs a piece of equipment called a protein skimmer, which produces air bubbles which the detritus adheres to, and forces it outside the tank before it decomposes, and also a highly porous type of natural rock called live rock where many bentos and bacteria live (hermatype which has been dead for some time is often used), which causes the detritus-feeding bentos and micro-organisms to undergo a detritus cycle. The Monaco system, where an anaerobic layer is created in the tank, to denitrify the organic compounds in the tank, and also the other nitrogen compounds, so that the decomposition process continues until the stage where water, carbon dioxide and nitrogen are produced, has also been implemented.

Initially, the filtration systems in water tanks often worked as the name suggests, using a physical filter to remove foreign substances in the water. Following this, the standard method for maintaining the water quality was to convert ammonium or nitrates in excreta, which have a high degree of neurotoxicity, but the combination of detritus feeders, detritus and micro-organisms has now brought aquarium technology to a still higher level.

Citations

↑ C.M. Hogan, 2008
↑ D.A. Grimaldi, 2005
↑ B.C. Patten, 1975

Sources

Bernard C. Patten (1975) Systems Analysis and Simulation in Ecology, Academic Press, 607 pages ISBN 0-12-547203-X
C. Michael Hogan (2008) "Western fence lizard (Sceloporus occidentalis)", Globaltwitcher, ed. Nicklas Stromberg
David Author Grimaldi and Michael S. Author (2005) Engelevolution of the insects, Cambridge University Press ISBN 0-521-82149-5
Some of this article was translated from the equivalent article in the Japanese-language Wikipedia, as it was on September 1, 2006.

Modelling ecosystems – trophic components

General	Abiotic component Abiotic stress Behaviour Biogeochemical cycle Biomass Biotic component Biotic stress Carrying capacity Competition Ecosystem Ecosystem ecology Ecosystem model Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Resource

Producers	Autotrophs Chemosynthesis Chemotrophs Foundation species Mixotrophs Myco-heterotrophy Mycotroph Organotrophs Photoheterotrophs Photosynthesis Photosynthetic efficiency Phototrophs Primary nutritional groups Primary production

Consumers	Apex predator Bacterivore Carnivores Chemoorganotroph Foraging Generalist and specialist species Intraguild predation Herbivores Heterotroph Heterotrophic nutrition Insectivore Mesopredator release hypothesis Omnivores Optimal foraging theory Predation Prey switching

Decomposers	Chemoorganoheterotrophy Decomposition Detritivores Detritus

Microorganisms	Archaea Bacteriophage Environmental microbiology Lithoautotroph Lithotrophy Microbial cooperation Microbial ecology Microbial food web Microbial intelligence Microbial loop Microbial mat Microbial metabolism Phage ecology

Food webs	Biomagnification Ecological efficiency Ecological pyramid Energy flow Food chain Trophic level

Example webs	Cold seeps Hydrothermal vents Intertidal Kelp forests Lakes North Pacific Subtropical Gyre Rivers San Francisco Estuary Soil Tide pool

Processes	Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer-resource systems Copiotrophs Dominance Ecological network Ecological succession Energy quality Energy Systems Language f-ratio Feed conversion ratio Feeding frenzy Mesotrophic soil Nutrient cycle Oligotroph Paradox of the plankton Trophic cascade Trophic mutualism Trophic state index

Defense/counter	Animal coloration Antipredator adaptations Camouflage Deimatic behaviour Herbivore adaptations to plant defense Mimicry Plant defense against herbivory Predator avoidance in schooling fish

Modelling ecosystems – other components

Population ecology	Abundance Allee effect Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation in wild animals Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey (Lotka–Volterra) equations Recruitment Resilience Small population size Stability

Species	Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species Latitudinal gradients in species diversity Minimum viable population Neutral theory Occupancy-abundance relationship Population viability analysis Priority effect Rapoport's rule Relative abundance distribution Relative species abundance Species diversity Species homogeneity Species richness Species distribution Species-area curve Umbrella species

Species interaction	Antibiosis Biological interaction Commensalism Community ecology Ecological facilitation Interspecific competition Mutualism Storage effect

Spatial ecology	Biogeography Cross-boundary subsidy Ecocline Ecotone Ecotype Disturbance Edge effects Foster's rule Habitat fragmentation Ideal free distribution Intermediate Disturbance Hypothesis Island biogeography Landscape ecology Landscape epidemiology Landscape limnology Metapopulation Patch dynamics r/K selection theory Source–sink dynamics

Niche	Ecological niche Ecological trap Ecosystem engineer Environmental niche modelling Guild Habitat Marine habitats Limiting similarity Niche apportionment models Niche construction Niche differentiation

Other networks	Assembly rules Bateman's principle Bioluminescence Ecological collapse Ecological debt Ecological deficit Ecological energetics Ecological indicator Ecological threshold Ecosystem diversity Emergence Extinction debt Kleiber's law Liebig's law of the minimum Marginal value theorem Thorson's rule Xerosere

Other	Allometry Alternative stable state Balance of nature Biological data visualization Constructal theory Ecocline Ecological economics Ecological footprint Ecological forecasting Ecological humanities Ecological stoichiometry Ecopath Ecosystem based fisheries Endolith Evolutionary ecology Functional ecology Industrial ecology Macroecology Microecosystem Natural environment Regime shift Systems ecology Theoretical ecology

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