Ecology (from Greek: οίκος, oikos, "household"; and λόγος, logos, "knowledge") is the scientific study of the distribution and abundance of life and the interactions between organisms and their environment. The environment of an organism includes physical properties, which can be described as the sum of local abiotic factors such as insolation (sunlight), climate, and geology, and biotic factors, which are other organisms that share its habitat.

The word "ecology" is often used more loosely in such terms as social ecology and deep ecology and in common parlance as a synonym for the natural environment or environmentalism. Likewise "ecologic" or "ecological" is often taken in the sense of environmentally friendly.

The term ecology or oekologie was coined by the German biologist Ernst Haeckel in 1866, when he defined it as "the comprehensive science of the relationship of the organism to the environment." Haeckel did not elaborate on the concept, and the first significant textbook on the subject (together with the first university course) was written by the Danish botanist, Eugenius Warming. For this early work, Warming is often identified as the founder of ecology.

Scope

Ecology is usually considered a branch of biology, the general science that studies living organisms. Organisms can be studied at many different levels, from proteins and nucleic acids (in biochemistry and molecular biology), to cells (in cellular biology), to individuals (in botany, zoology, and other similar disciplines), and finally at the level of populations, communities, and ecosystems, to the biosphere as a whole; these latter strata are the primary subjects of ecological inquiry. Ecology is a multi-disciplinary science. Because of its focus on the higher levels of the organization of life on earth and on the interrelations between organisms and their environment, ecology draws heavily on many other branches of science, especially geology and geography, meteorology, pedology, genetics, chemistry, and physics. Thus, ecology is considered by some to be a holistic science, one that over-arches older disciplines such as biology which in this view become sub-disciplines contributing to ecological knowledge. In support of viewing ecology as a subject in its own right as opposed to a sub-discipline of biology, Robert Ulanowicz stated that "The emerging picture of ecosystem behavior does not resemble the worldview imparted by an extrapolation of conceptual trends established in other sciences."

Agriculture, fisheries, forestry, medicine and urban development are among human activities that would fall within Krebs' (1972: 4) explanation of his definition of ecology: where organisms are found, how many occur there, and why.

Ecological knowledge such as the quantification of biodiversity and population dynamics have provided a scientific basis for expressing the aims of environmentalism and evaluating its goals and policies. Additionally, a holistic view of nature is stressed in both ecology and environmentalism.

Consider the ways an ecologist might approach studying the life of honeybees:

• The behavioral relationship between individuals of a species is behavioral ecology — for example, the study of the queen bee, and how she relates to the worker bees and the drones.

• The organized activity of a species is community ecology; for example, the activity of bees assures the pollination of flowering plants. Bee hives additionally produce honey which is consumed by still other species, such as bears.
• The relationship between the environment and a species is environmental ecology — for example, the consequences of environmental change on bee activity. Bees may die out due to environmental changes (see pollinator decline). The environment simultaneously affects and is a consequence of this activity and is thus intertwined with the survival of the species.

Disciplines of ecology

Ecology is a broad discipline comprising many sub-disciplines. A common, broad classification, moving from lowest to highest complexity, where complexity is defined as the number of entities and processes in the system under study, is:

• Ecophysiology examines how the physiological functions of organisms influence the way they interact with the environment, both biotic and abiotic
• Behavioral ecology examines the roles of behavior in enabling an animal to adapt to its environment
• Population ecology studies the dynamics of populations of a single species.
• Community ecology (or synecology) focuses on the interactions between species within an ecological community.
• Ecosystem ecology studies the flows of energy and matter through the biotic and abiotic components of ecosystems.
• Systems ecology is an interdisciplinary field focusing on the study, development, and organization of ecological systems from a holistic perspective.
• Landscape ecology examines processes and relationship across multiple ecosystems or very large geographic areas.
• Evolutionary ecology studies ecology in a way that explicitly considers the evolutionary histories of species and their interactions
• Political ecology connects politics and economy to problems of environmental control and ecological change

Ecology can also be sub-divided according to the species of interest into fields such as animal ecology, plant ecology, insect ecology, and so on. Another frequent method of subdivision is by biome studied, e.g., Arctic ecology (or polar ecology), tropical ecology, desert ecology, etc. The primary technique used for investigation is often used to subdivide the discipline into groups such as chemical ecology, genetic ecology, field ecology, statistical ecology, theoretical ecology, and so forth. These fields are not mutually exclusive.

History of ecology

Fundamental principles of ecology

Levels of ecological organization

Ecology can be studied at a wide range of levels, from a large to small scale. These levels of ecological organization, as well as an example of a question ecologists would ask at each level, include:

• Biosphere " What role does concentration of atmospheric Carbon Dioxide play in the regulation of global temperature?"
• Region "How has geological history influenced regional diversity within certain groups of organisms?"
• Landscape "How do vegetated corridors affect the rate of movement by mammals among isolated fragments?"
• Ecosystem "How does fire affect nutrient availability in grassland ecosystems?"
• Community "How does disturbance influence the number of mammal species in African grasslands?"
• Interactions "What evolutionary benefit do zebras gain by allowing birds to remove parasites?"
• Population "What factors control zebra populations?"
• Individual "How do zebras regulate internal water balance?"
  • These levels range from broadest to most specific

Biosphere

For modern ecologists, ecology can be studied at several levels: population level (individuals of the same species in the same or similar environment), biocoenosis level (or community of species), ecosystem level, and biosphere level.

The outer layer of the planet Earth can be divided into several compartments: the hydrosphere (or sphere of water), the lithosphere (or sphere of soils and rocks), and the atmosphere (or sphere of the air). The biosphere (or sphere of life), sometimes described as "the fourth envelope", is all living matter on the planet or that portion of the planet occupied by life. It reaches well into the other three spheres, although there are no permanent inhabitants of the atmosphere. Relative to the volume of the Earth, the biosphere is only the very thin surface layer which extends from 11,000 meters below sea level to 15,000 meters above.

It is thought that life first developed in the hydrosphere, at shallow depths, in the photic zone. (Recently, though, a competing theory has emerged, that life originated around hydrothermal vents in the deeper ocean. See Origin of life.) Multicellular organisms then appeared and colonized benthic zones. Photosynthetic organisms gradually produced the chemically unstable oxygen-rich atmosphere that characterizes our planet. Terrestrial life developed later, after the ozone layer protecting living beings from UV rays formed. Diversification of terrestrial species is thought to be increased by the continents drifting apart, or alternately, colliding. Biodiversity is expressed at the ecological level (ecosystem), population level (intraspecific diversity), species level (specific diversity), and genetic level. Recently technology has allowed the discovery of the deep ocean vent communities. This remarkable ecological system is not dependent on sunlight but bacteria, utilising the chemistry of the hot volcanic vents, are at the base of its food chain.

The biosphere contains great quantities of elements such as carbon, nitrogen, hydrogen and oxygen. Other elements, such as phosphorus, calcium, and potassium, are also essential to life, yet are present in smaller amounts. At the ecosystem and biosphere levels, there is a continual recycling of all these elements, which alternate between the mineral and organic states.

While there is a slight input of geothermal energy, the bulk of the functioning of the ecosystem is based on the input of solar energy. Plants and photosynthetic microorganisms convert light into chemical energy by the process of photosynthesis, which creates glucose (a simple sugar) and releases free oxygen. Glucose thus becomes the secondary energy source which drives the ecosystem. Some of this glucose is used directly by other organisms for energy. Other sugar molecules can be converted to other molecules such as amino acids. Plants use some of this sugar, concentrated in nectar to entice pollinators to aid them in reproduction.

Cellular respiration is the process by which organisms (like mammals) break the glucose back down into its constituents, water and carbon dioxide, thus regaining the stored energy the sun originally gave to the plants. The proportion of photosynthetic activity of plants and other photosynthesizers to the respiration of other organisms determines the specific composition of the Earth's atmosphere, particularly its oxygen level. Global air currents mix the atmosphere and maintain nearly the same balance of elements in areas of intense biological activity and areas of slight biological activity.

Water is also exchanged between the hydrosphere, lithosphere, atmosphere and biosphere in regular cycles. The oceans are large tanks, which store water, ensure thermal and climatic stability, as well as the transport of chemical elements thanks to large oceanic currents.

For a better understanding of how the biosphere works, and various dysfunctions related to human activity, American scientists simulated the biosphere in a small-scale model, called Biosphere II.

The ecosystem concept

The first principle of ecology is that each living organism has an ongoing and continual relationship with every other element that makes up its environment. An ecosystem can be defined as any situation where there is interaction between organisms and their environment.

The ecosystem is of two entities, the entirety of life, the biocoenosis, and the medium that life exists in, the biotope. Within the ecosystem, species are connected by food chains or food webs. Energy from the sun, captured by primary producers via photosynthesis, flows upward through the chain to primary consumers (herbivores), and then to secondary and tertiary consumers (carnivores and omnivores), before ultimately being lost to the system as waste heat. In the process, matter is incorporated into living organisms, which return their nutrients to the system via decomposition, forming biogeochemical cycles such as the carbon and nitrogen cycles.

The concept of an ecosystem can apply to units of variable size, such as a pond, a field, or a piece of dead wood. An ecosystem within another ecosystem is called a micro ecosystem. For example, an ecosystem can be a stone and all the life under it. A meso ecosystem could be a forest, and a macro ecosystem'' a whole eco region, with its drainage basin.

The main questions when studying an ecosystem are:

• Whether the colonization of a barren area could be carried out
• Investigation the ecosystem's dynamics and changes
• The methods of which an ecosystem interacts at local, regional and global scale
• Whether the current state is stable
• Investigating the value of an ecosystem and the ways and means that interaction of ecological systems provides benefits to humans, especially in the provision of healthy water.

Ecosystems are often classified by reference to the biotopes concerned. The following ecosystems may be defined:

• As continental ecosystems, such as forest ecosystems, meadow ecosystems such as steppes or savannas, or agro-ecosystems
• As ecosystems of inland waters, such as lentic ecosystems such as lakes or ponds; or lotic ecosystems such as rivers
• As oceanic ecosystems.

Another classification can be done by reference to its communities, such as in the case of an human ecosystem.

Dynamics and stability

Ecological factors which affect dynamic change in a population or species in a given ecology or environment are usually divided into two groups: abiotic and biotic.

Abiotic factors are geological, geographical, hydrological and climatological parameters. A biotope is an environmentally uniform region characterized by a particular set of abiotic ecological factors. Specific abiotic factors include:

• Water, which is at the same time an essential element to life and a milieu
• Air, which provides oxygen, nitrogen, and carbon dioxide to living species and allows the dissemination of pollen and spores
• Soil, at the same time source of nutriment and physical support
  • Soil pH, salinity, nitrogen and phosphorus content, ability to retain water, and density are all influential
• Temperature, which should not exceed certain extremes, even if tolerance to heat is significant for some species
• Light, which provides energy to the ecosystem through photosynthesis
• Natural disasters can also be considered abiotic

Biocenose, or community, is a group of populations of plants, animals, micro-organisms. Each population is the result of procreations between individuals of same species and cohabitation in a given place and for a given time. When a population consists of an insufficient number of individuals, that population is threatened with extinction; the extinction of a species can approach when all biocenoses composed of individuals of the species are in decline. In small populations, consanguinity (inbreeding) can result in reduced genetic diversity that can further weaken the biocenose.

Biotic ecological factors also influence biocenose viability; these factors are considered as either intraspecific and interspecific relations.

Intraspecific relations are those which are established between individuals of the same species, forming a population. They are relations of co-operation or competition, with division of the territory, and sometimes organization in hierarchical societies.

Interspecific relations—interactions between different species—are numerous, and usually described according to their beneficial, detrimental or neutral effect (for example, mutualism (relation ++) or competition (relation --). The most significant relation is the relation of predation (to eat or to be eaten), which leads to the essential concepts in ecology of food chains (for example, the grass is consumed by the herbivore, itself consumed by a carnivore, itself consumed by a carnivore of larger size). A high predator to prey ratio can have a negative influence on both the predator and prey biocenoses in that low availability of food and high death rate prior to sexual maturity can decrease (or prevent the increase of) populations of each, respectively. Selective hunting of species by humans which leads to population decline is one example of a high predator to prey ratio in action. Other interspecific relations include parasitism, infectious disease and competition for limiting resources, which can occur when two species share the same ecological niche.

The existing interactions between the various living beings go along with a permanent mixing of mineral and organic substances, absorbed by organisms for their growth, their maintenance and their reproduction, to be finally rejected as waste. These permanent recyclings of the elements (in particular carbon, oxygen and nitrogen) as well as the water are called biogeochemical cycles. They guarantee a durable stability of the biosphere (at least when unchecked human influence and extreme weather or geological phenomena are left aside). This self-regulation, supported by negative feedback controls, ensures the perenniality of the ecosystems. It is shown by the very stable concentrations of most elements of each compartment. This is referred to as homeostasis. The ecosystem also tends to evolve to a state of ideal balance, reached after a succession of events, the climax (for example a pond can become a peat bog).

Spatial relationships and subdivisions of land

Ecosystems are not isolated from each other, but are interrelated. For example, water may circulate between ecosystems by the means of a river or ocean current. Water itself, as a liquid medium, even defines ecosystems. Some species, such as salmon or freshwater eels move between marine systems and fresh-water systems. These relationships between the ecosystems lead to the concept of a biome.

A biome is a homogeneous ecological formation that exists over a large region as tundra or steppes. The biosphere comprises all of the Earth's biomes -- the entirety of places where life is possible -- from the highest mountains to the depths of the oceans.

Biomes correspond rather well to subdivisions distributed along the latitudes, from the equator towards the poles, with differences based on to the physical environment (for example, oceans or mountain ranges) and to the climate. Their variation is generally related to the distribution of species according to their ability to tolerate temperature and/or dryness. For example, one may find photosynthetic algae only in the photic part of the ocean (where light penetrates), while conifers are mostly found in mountains.

Though this is a simplification of more complicated scheme, latitude and altitude approximate a good representation of the distribution of biodiversity within the biosphere. Very generally, the richness of biodiversity (as well for animal than plant species) is decreasing most rapidly near the equator and less rapidly as one approaches the poles.

The biosphere may also be divided into ecozones, which are very well defined today and primarily follow the continental borders. The ecozones are themselves divided into ecoregions, though there is not agreement on their limits.

Ecosystem productivity

In an ecosystem, the connections between species are generally related to food and their role in the food chain. There are three categories of organisms:

• Producers -- usually plants which are capable of photosynthesis but could be other organisms such as bacteria around ocean vents that are capable of chemosynthesis.
• Consumers -- animals, which can be primary consumers (herbivorous), or secondary or tertiary consumers (carnivorous and omnivores).
• Decomposers -- bacteria, mushrooms which degrade organic matter of all categories, and restore minerals to the environment. And decomposers can also decompose decaying animals

These relations form sequences, in which each individual consumes the preceding one and is consumed by the one following, in what are called food chains or food networks. In a food network, there will be fewer organisms at each level as one follows the links of the network up the chain.

These concepts lead to the idea of biomass (the total living matter in a given place), of primary productivity (the increase in the mass of plants during a given time) and of secondary productivity (the living matter produced by consumers and the decomposers in a given time).

These two last ideas are key, since they make it possible to evaluate the load capacity -- the number of organisms which can be supported by a given ecosystem. In any food network, the energy contained in the level of the producers is not completely transferred to the consumers. And the higher one goes up the chain, the more energy and resources is lost and consumed. Thus, from an energy—and environmental—point of view, it is more efficient for humans to be primary consumers (to subsist from vegetables, grains, legumes, fruit, etc.) than as secondary consumers (from eating herbivores, omnivores, or their products, such as milk, chickens, cattle, sheep, etc.) and still more so than as a tertiary consumer (from consuming carnivores, omnivores, or their products, such as fur, pigs, snakes, alligators, etc.). An ecosystem(s) is unstable when the load capacity is overrun and is especially unstable when a population doesn't have an ecological niche and overconsumers.

The productivity of ecosystems is sometimes estimated by comparing three types of land-based ecosystems and the total of aquatic ecosystems:

• The forests (1/3 of the Earth's land area) contain dense biomasses and are very productive. The total production of the world's forests corresponds to half of the primary production.
• Savannas, meadows, and marshes (1/3 of the Earth's land area) contain less dense biomasses, but are productive. These ecosystems represent the major part of what humans depend on for food.
• Extreme ecosystems in the areas with more extreme climates -- deserts and semi-deserts, tundra, alpine meadows, and steppes -- (1/3 of the Earth's land area) have very sparse biomasses and low productivity
• Finally, the marine and fresh water ecosystems (3/4 of Earth's surface) contain very sparse biomasses (apart from the coastal zones).

Humanity's actions over the last few centuries have seriously reduced the amount of the Earth covered by forests (deforestation), and have increased agro-ecosystems (agriculture). In recent decades, an increase in the areas occupied by extreme ecosystems has occurred (desertification).

Ecological crisis

Generally, an ecological crisis occurs with the loss of adaptive capacity when the resilience of an environment or of a species or a population evolves in a way unfavourable to coping with perturbations that interfere with that ecosystem, landscape or species survival. It may be that the environment quality degrades compared to the species needs, after a change in an abiotic ecological factor (for example, an increase of temperature, less significant rainfalls). It may be that the environment becomes unfavourable for the survival of a species (or a population) due to an increased pressure of predation (for example overfishing). Lastly, it may be that the situation becomes unfavourable to the quality of life of the species (or the population) due to a rise in the number of individuals (overpopulation).

Ecological crises vary in length and severity, occurring within a few months or taking as long as a few million years. They can also be of natural or anthropic origin. They may relate to one unique species or to many species, as in an Extinction event. Lastly, an ecological crisis may be local (as an oil spill) or global (a rise in the sea level due to global warming).

According to its degree of endemism, a local crisis will have more or less significant consequences, from the death of many individuals to the total extinction of a species. Whatever its origin, disappearance of one or several species often will involve a rupture in the food chain, further impacting the survival of other species.

In the case of a global crisis, the consequences can be much more significant; some extinction events showed the disappearance of more than 90% of existing species at that time. However, it should be noted that the disappearance of certain species, such as the dinosaurs, by freeing an ecological niche, allowed the development and the diversification of the mammals. An ecological crisis thus paradoxically favored biodiversity.

Sometimes, an ecological crisis can be a specific and reversible phenomenon at the ecosystem scale. But more generally, the crises impact will last. Indeed, it rather is a connected series of events, that occur till a final point. From this stage, no return to the previous stable state is possible, and a new stable state will be set up gradually (see homeorhesy).

Lastly, if an ecological crisis can cause extinction, it can also more simply reduce the quality of life of the remaining individuals. Thus, even if the diversity of the human population is sometimes considered threatened (see in particular indigenous people), few people envision human disappearance at short span. However, epidemic diseases, famines, impact on health of reduction of air quality, food crises, reduction of living space, accumulation of toxic or non degradable wastes, threats on keystone species (great apes, panda, whales) are also factors influencing the well-being of people.

Due to the increases in technology and a rapidly increasing population, humans have more influence on their own environment than any other ecosystem engineer.

Some common examples of ecological crises are:

• The Exxon Valdez oil spill off the coast of Alaska in 1989
• Permian-Triassic extinction event 250 million of years ago
• Cretaceous–Tertiary extinction event 65 million years ago
• Global warming related to the Greenhouse effect. Warming could involve flooding of the Asian deltas (see also eco refugees), multiplication of extreme weather phenomena and changes in the nature and quantity of the food resources (see Global warming and agriculture). See also international Kyoto Protocol.
• Ozone layer hole issue
• Deforestation and desertification, with disappearance of many species.
• Volcanic eruptions such as Mount St. Helens and the Tunguska and other impact events
• The nuclear meltdown at Chernobyl in 1986 caused the death of many people and animals from cancer, and caused mutations in a large number of animals and people. The area around the plant is now abandoned by humans because of the large amount of radiation generated by the meltdown. Twenty years after the accident, the animals have returned.

Bibliography

• Warming, E. (1909) Oecology of Plants - an introduction to the study of plant-communities. Clarendon Press, Oxford.
• Haeckel, E. (1866) General Morphology of Organisms; General Outlines of the Science of Organic Forms based on Mechanical Principles through the Theory of Descent as reformed by Charles Darwin. Berlin.

References

See also

Lists

• List of basic biology topics
• List of biology topics
• List of basic ecology topics
• List of ecology topics
• List of ecologists
• Important publications in ecology

Related topics

• Acoustic ecology
• Deep ecology
• Ecological economics
• Ecology movement
• Ecosystem
• Ecosystem model
• Ecohydrology
• ELDIS, a database on ecological aspects of economical development.
• Environmental art
• Environmental science
• Environmental technology
• Environmental communication
• Environmental psychology
• Forest farming
• Forest gardening
• Human ecology
• Knowledge ecology
• Permaculture
• Social ecology
• Systems ecology

External links

• What is Ecology?
• Fundamentals of Ecology Textbook-style investigation to the economy of nature, breaks down in 4 chapters from Population to Ecosystem.
• Ecology (Stanford Encyclopedia of Philosophy)