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.

Energy transfer

Energy transfer
A diagram of energy transfer between trophic levels

Primary production occurs in autotrophic organisms of an ecosystem. Photoautotrophs such as vascular plants and algae convert energy from the sun into energy stored as carbon compounds. Photosynthesis is carried out in the chlorophyll of green plants. The energy converted through photosynthesis is carried through the trophic levels of an ecosystem as organisms consume members of lower trophic levels.

Primary production can be broken down into gross and net primary production. Gross primary production is a measure of the energy that a photoautotroph harvests from the sun. Take, for example, a blade of grass that takes in x joules of energy from the sun. The fraction of that energy that is converted into glucose reflects the gross productivity of the blade of grass. The energy remaining after respiration is considered the net primary production. In general, gross production refers to the energy contained within an organism before respiration and net production the energy after respiration. The terms can be used to describe energy transfer in both autotrophs and heterotrophs.

Energy transfer between trophic levels is generally inefficient, such that net production at one trophic level is generally only 10% of the net production at the preceding trophic level (the Ten percent law). Due to non-predatory death, egestion, and cellular respiration, a significant amount of energy is lost to the environment instead of being absorbed for production by consumers. The figure approximates the fraction of energy available after each stage of energy loss in a typical ecosystem, although these fractions vary greatly from ecosystem to ecosystem and from trophic level to trophic level. The loss of energy by a factor of one half from each of the steps of non-predatory death, defecation, and respiration is typical of many living systems. Thus, the net production at one trophic level is or approximately ten percent that of the trophic level before it.

For example, assume 500 units of energy are produced by trophic level 1. One half of that is lost to non-predatory death, while the other half (250 units) is ingested by trophic level 2. One half of the amount ingested is expelled through defecation, leaving the other half (125 units) to be assimilated by the organism. Finally one half of the remaining energy is lost through respiration while the rest (63 units) is used for growth and reproduction. This energy expended for growth and reproduction constitutes to the net production of trophic level 1, which is equal to units.

Quantifying ecological efficiency

Ecological efficiency is a combination of several related efficiencies that describe resource utilization and the extent to which resources are converted into biomass.[1]

  • Exploitation efficiency is the amount of food ingested divided by the amount of prey production ()
  • Assimilation efficiency is the amount of assimilation divided by the amount of food ingestion ()
  • Net Production efficiency is the amount of consumer production divided by the amount of assimilation ()
  • Gross Production efficiency is the assimilation efficiency multiplied by the net production efficiency, which is equivalent to the amount of consumer production divided by amount of ingestion ()
  • Ecological efficiency is the exploitation efficiency multiplied by the assimilation efficiency multiplied by the net production efficiency, which is equivalent to the amount of consumer production divided by the amount of prey production ()

Theoretically, it is easy to calculate ecological efficiency using the mathematical relationships above. It is often difficult, however, to obtain accurate measurements of the values involved in the calculation. Assessing ingestion, for example, requires knowledge of the gross amount of food consumed in an ecosystem as well as its caloric content. Such a measurement is rarely better than an educated estimate, particularly with relation to ecosystems that are largely inaccessible to ecologists and tools of measurement. The ecological efficiency of an ecosystem is as a result often no better than an approximation. On the other hand, an approximation may be enough for most ecosystems, where it is important not to get an exact measure of efficiency, but rather a general idea of how energy is moving through its trophic levels.

Applications

In agricultural environments, maximizing energy transfer from producer (food) to consumer (livestock) can yield economic benefits. A sub-field of agricultural science has emerged that explores methods of monitoring and improving ecological and related efficiencies.

In comparing the net efficiency of energy utilization by cattle, breeds historically kept for beef production, such as the Hereford, outperformed those kept for dairy production, such as the Holstein, in converting energy from feed into stored energy as tissue.[2] This is a result of the beef cattle storing more body fat than the dairy cattle, as energy storage as protein was at the same level for both breeds. This implies that cultivation of cattle for slaughter is a more efficient use of feed than is cultivation for milk production.

While it is possible to improve the efficiency of energy use by livestock, it is vital to the world food question to also consider the differences between animal husbandry and plant agriculture. Caloric concentration in fat tissues are higher than in plant tissues, causing high-fat organisms to be most energetically-concentrated; however, the energy required to cultivate feed for livestock is only partially converted into fat cells. The rest of the energy input into cultivating feed is respired or egested by the livestock and unable to be used by humans.

Out of a total of 96.8 10^15 BTU of energy used in the US in 1999, 10.5% was used in food production,[3] with the percentage accounting for food from both producer and primary consumer trophic levels. In comparing the cultivation of animals versus plants, there is a clear difference in magnitude of energy efficiency. Edible kilocalories produced from kilocalories of energy required for cultivation are: 18.1% for chicken, 6.7% for grass-fed beef, 5.7% for farmed salmon, and 0.9% for shrimp. In contrast, potatoes yield 123%, corn produce 250%, and soy results in 415% of input calories converted to calories able to be utilized by humans.[4] This disparity in efficiency reflects the reduction in production from moving up trophic levels. Thus, it is more energetically efficient to form a diet from lower trophic levels.

Ten percent law

The ten percent law of transfer of energy from one trophic level to the next can be attributed to Raymond Lindeman (1942),[5] although Lindeman did not call it a "law" and cited ecological efficiencies ranging from 0.1% to 37.5%. According to this law, during the transfer of organic food energy from one trophic level to the next higher level, only about ten percent of the transferred energy is stored as flesh. The remaining is lost during transfer, broken down in respiration, or lost to incomplete digestion by higher trophic level.

The food chain

When organisms are consumed, 10% of the energy in the food is fixed into their flesh and is available for next trophic level (carnivores or omnivores). When a carnivore or an omnivore consumes that animal, only about 10% of energy is fixed in its flesh for the higher level.

For example, the Sun releases 1000 J of energy, then plants take only 100 J of energy from sunlight; thereafter, a deer would take 10 J from the plant. A wolf eating the deer would only take 1 J. A human eating the wolf would take 0.1J, etc.

The ten percent law provides a basic understanding on the cycling of food chains. Furthermore, the ten percent law shows the inefficiency of energy capture at each successive trophic level. The rational conclusion is that energy efficiency is best preserved by sourcing food as close to the initial energy source as possible.

Formula

Energy at n(th) level

         = (energy given by sun)/(10)^(n+1), 

and,

Energy at n(th) level

         =  (energy given by plant)/(10)^(n-1).

{Remember to count only plant energy in both equation}

See also

  • Eco-efficiency - the economic efficiency with which human society uses ecological resources

References

  1. ^ [1]
  2. ^ Gareett, W.N. Energetic Efficiency of Beef and Dairy Steers. Journal of Animal Science.1971. 32:451-456
  3. ^ U.S. Department of Energy, 2004: Annual energy review 2003. Rep. DOE/EIA-0384(2003), Energy Information Administration, 390 pp
  4. ^ Eshel, Gidon and Martin, Pamela A. Diet, Energy, and Global Warming. Earth Interactions. 2005. 10: 1- 17
  5. ^ Lindeman, RL (1942). "The trophic-dynamic aspect of ecology". Ecology. 23: 399–418. doi:10.2307/1930126.
21st century

The 21st (twenty-first) century is the current century of the Anno Domini era or Common Era, in accordance with the Gregorian calendar. It began on January 1, 2001, and will end on December 31, 2100. It is the first century of the 3rd millennium. It is distinct from the century known as the 2000s which began on January 1, 2000 and will end on December 31, 2099.The first years of the 21st century have thus far been marked by the rise of a global economy and Third World consumerism, mistrust in government, deepening global concern over terrorism and an increase in the power of private enterprise. The Arab Spring of the early 2010s led to mixed outcomes in the Arab world. The Third Industrial Revolution which began around the 1980s also continues into the present, and is expected to transition into Industry 4.0 and the Fourth Industrial Revolution by as early as 2030. Due to the proliferation of mobile devices, at least 40% of the world's population are currently connected to the internet, allowing humans to be more intertwined than ever before. Sequencing cost continues to fall exponentially: the first human genome cost three billion dollars. Today, sequencing only one human genome only costs about a thousand. Millennials and Generation Z come of age and rise to prominence in this century. In 2016, the United Kingdom decided to leave the European Union, causing Brexit.

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.

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 on Lake Victoria

Lake Victoria supports Africa's largest inland fishery, with the majority of the catch being the invasive Nile perch, introduced in the Lake in the 1950s.

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.

Glossary of biology

Most of the terms listed in Wikipedia glossaries are already defined and explained within Wikipedia itself. However, glossaries like this one are useful for looking up, comparing and reviewing large numbers of terms together. You can help enhance this page by adding new terms or writing definitions for existing ones.

This glossary of biology terms is a list of definitions of fundamental terms and concepts of biology, its sub-disciplines, and related fields. For more specific definitions from other glossaries related to biology, see Glossary of ecology, Glossary of botany, Glossary of genetics, and Glossary of speciation.

Happy Planet Index

The Happy Planet Index (HPI) is an index of human well-being and environmental impact that was introduced by the New Economics Foundation (NEF) in July 2006. The index is weighted to give progressively higher scores to nations with lower ecological footprints.

The index is designed to challenge well-established indices of countries’ development, such as the gross domestic product (GDP) and the Human Development Index (HDI), which are seen as not taking sustainability into account. In particular, GDP is seen as inappropriate, as the usual ultimate aim of most people is not to be rich, but to be happy and healthy. Furthermore, it is believed that the notion of sustainable development requires a measure of the environmental costs of pursuing those goals.Out of the 178 countries surveyed in 2006, the best scoring countries were Vanuatu, Colombia, Costa Rica, Dominica, and Panama. In 2009, Costa Rica was the best scoring country among the 143 analyzed, followed by the Dominican Republic, Jamaica, Guatemala and Vietnam. Tanzania, Botswana and Zimbabwe were featured at the bottom of the list.For the 2012 ranking, 151 countries were compared, and the best scoring country for the second time in a row was Costa Rica, followed by Vietnam, Colombia, Belize and El Salvador. The lowest ranking countries in 2012 were Botswana, Chad and Qatar. In 2016, out of 140 countries, Costa Rica topped the index for the third time in a row. It was followed by Mexico, Colombia, Vanuatu and Vietnam. At the bottom were Chad, Luxembourg and Togo.

Invasive species

An invasive species is a species that is not native to a specific location (an introduced species), and that has a tendency to spread to a degree believed to cause damage to the environment, human economy or human health.The term as most often used applies to introduced species that adversely affect the habitats and bioregions they invade economically, environmentally, or ecologically. Such species may be either plants or animals and may disrupt by dominating a region, wilderness areas, particular habitats, or wildland–urban interface land from loss of natural controls (such as predators or herbivores). This includes plant species labeled as exotic pest plants and invasive exotics growing in native plant communities. The European Union defines "Invasive Alien Species" as those that are, firstly, outside their natural distribution area, and secondly, threaten biological diversity. The term is also used by land managers, botanists, researchers, horticulturalists, conservationists, and the public for noxious weeds.The term "invasive" is often poorly defined or very subjective and some broaden the term to include indigenous or "native" species, that have colonized natural areas - for example deer considered by some to be overpopulating their native zones and adjacent suburban gardens in the Northeastern and Pacific Coast regions of the United States.The definition of "native" is also sometimes controversial. For example, the ancestors of Equus ferus (modern horses) evolved in North America and radiated to Eurasia before becoming locally extinct. Upon returning to North America in 1493 during their hominid-assisted migration, it is debatable as to whether they were native or exotic to the continent of their evolutionary ancestors.Notable examples of invasive plant species include The kudzu vine, Andean pampas grass, and yellow starthistle. Animal examples include the New Zealand mud snail, feral pigs, European rabbits, grey squirrels, domestic cats, carp and ferrets.Invasion of long-established ecosystems by organisms from distant bio-regions is a natural phenomenon, but has been accelerated massively by humans, from their earliest migrations though to the age of discovery, and now international trade.

Lakhta Center

The Lakhta Center (Russian: Ла́хта це́нтр, tr. Lakhta tsentr) is an 87-story skyscraper built in the outskirts of Lakhta in Saint Petersburg, Russia. Standing 462 metres (1,516 ft) tall, the Lakhta Center is the tallest building in Russia, the tallest building in Europe, and the 13th-tallest building in the world. The Lakhta Center is also the second-tallest structure in Russia and Europe, behind Ostankino Tower in Moscow.Construction of Lakhta Center started on 30 October 2012; it was topped out on 29 January 2018. The Lakhta Center surpassed Vostok Tower of the Federation Towers in Moscow as the tallest building in Russia and Europe on 5 October 2017. The center is designed for large-scale mixed-use development, consisting of public facilities and offices. The building was designed by RMJM. The project was then continued by GORPROJECT (2011-2017) based on the RMJM Concept (2011) under the main contractor, Rönesans Holding. The Lakhta Center is intended to become the new headquarters of Russian energy company Gazprom.

On December 24, 2018, Lakhta Center was certified according to the criteria of ecological efficiency at LEED Platinum. The concrete pouring of the bottom slab of Lakhta Center's foundation was registered by Guinness World Records as the largest continuous concrete pour; 19,624 cubic meters of concrete were used, which is approximately 3,000 cubic meters more than in the previous similar record registered at Wilshire Grand Tower. The tower's curtain wall is also the world’s largest cold-bent facade by area.

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.

Product-service system

Product-service systems (PSS) are business models that provide for cohesive delivery of products and services. PSS models are emerging as a means to enable collaborative consumption of both products and services, with the aim of pro-environmental outcomes.

Productivity (ecology)

In ecology, productivity refers to the rate of generation of biomass in an ecosystem. It is usually expressed in units of mass per unit surface (or volume) per unit time, for instance grams per square metre per day (g m−2 d−1). The mass unit may relate to dry matter or to the mass of carbon generated. Productivity of autotrophs such as plants is called primary productivity, while that of heterotrophs such as animals is called secondary productivity.

Resource efficiency

Resource efficiency is the maximising of the supply of money, materials, staff, and other assets that can be drawn on by a person or organization in order to function effectively, with minimum wasted (natural) resource expenses. It means using the Earth's limited resources in a sustainable manner while minimising environmental impact.

Soil Use Efficiency

Soil Use Efficiency (SUE) is the use of individual and inter-related factors (inherent and dynamic) related to soil quality, soil nutrient availability and nutrient uptake potential as effective reference points for improvement of crop productivity in individual and varying soil types. Assessing SUE involves a site evaluation of the land and pit excavation to examine the soil profile. Site characterization identifies impairments to biomass productivity and the provisioning of ecological services. Inherent impairments are from limits inherent to the land and soil such as steep slope, and subsoil claypan. Dynamic impairments are the result of land degradation, such as soil acidification, loss of soil carbon, water erosion, and wind erosion. Understanding these relationships informs land management decisions needed to restore land productivity. The determination of water-use efficiency (WUE) and Nutrient Use Efficiency (NUE) in agricultural production systems is governed primarily by the boundary conditions of Soil Use Efficiency (SUE).

Sustainable gardening

Sustainable gardening includes the more specific sustainable landscapes, sustainable landscape design, sustainable landscaping, sustainable landscape architecture, resulting in sustainable sites. It comprises a disparate group of horticultural interests that can share the aims and objectives associated with the international post-1980s sustainable development and sustainability programs developed to address the fact that humans are now using natural biophysical resources faster than they can be replenished by nature.Included within this compass are those home gardeners, and members of the landscape and nursery industries, and municipal authorities, that integrate environmental, social, and economic factors to create a more sustainable future.

Organic gardening and the use of native plants are integral to sustainable gardening.

Top-down and bottom-up design

Top-down and bottom-up are both strategies of information processing and knowledge ordering, used in a variety of fields including software, humanistic and scientific theories (see systemics), and management and organization. In practice, they can be seen as a style of thinking, teaching, or leadership.

A top-down approach (also known as stepwise design and in some cases used as a synonym of decomposition) is essentially the breaking down of a system to gain insight into its compositional sub-systems in a reverse engineering fashion. In a top-down approach an overview of the system is formulated, specifying, but not detailing, any first-level subsystems. Each subsystem is then refined in yet greater detail, sometimes in many additional subsystem levels, until the entire specification is reduced to base elements. A top-down model is often specified with the assistance of "black boxes", which makes it easier to manipulate. However, black boxes may fail to clarify elementary mechanisms or be detailed enough to realistically validate the model. Top down approach starts with the big picture. It breaks down from there into smaller segments.A bottom-up approach is the piecing together of systems to give rise to more complex systems, thus making the original systems sub-systems of the emergent system. Bottom-up processing is a type of information processing based on incoming data from the environment to form a perception. From a cognitive psychology perspective, information enters the eyes in one direction (sensory input, or the "bottom"), and is then turned into an image by the brain that can be interpreted and recognized as a perception (output that is "built up" from processing to final cognition). In a bottom-up approach the individual base elements of the system are first specified in great detail. These elements are then linked together to form larger subsystems, which then in turn are linked, sometimes in many levels, until a complete top-level system is formed. This strategy often resembles a "seed" model, by which the beginnings are small but eventually grow in complexity and completeness. However, "organic strategies" may result in a tangle of elements and subsystems, developed in isolation and subject to local optimization as opposed to meeting a global purpose.

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.

Vacant niche

The issue of what exactly defines a vacant niche, also known as empty niche, and whether they exist in ecosystems is controversial. The subject is intimately tied into a much broader debate on whether ecosystems can reach equilibrium, where they could theoretically become maximally saturated with species. Given that saturation is a measure of the number of species per resource axis per ecosystem, the question becomes: is it useful to define unused resource clusters as niche 'vacancies'?

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