Ecology (from Greek: οἶκος, "house"; -λογία, "study of") is the scientific study of the relation of living organisms to each

other and their surroundings.[1] Ecology includes the study of plant and animal populations, plant and animal communities and ecosystems. Ecologists study a range of living phenomena from the role of bacteria in nutrient recycling to the effects of tropical rain forest on the Earth's atmosphere. The word "ecology" ("oekologie") was coined by[citation needed] the German scientist Ernst Haeckel 1834–1919. Haeckel was a zoologist, artist, writer, and later in life a professor of comparative anatomy. Ecology is a sub-discipline of biology, which is the study of life, branching out from the natural sciences in the late 19th century. Ecology is not synonymous with environment, environmentalism, natural history or environmental science.[1] [2][3] Ecology is closely related to the biological disciplines of physiology, evolution, genetics and behavior.[4][5] Ecology seeks to explain:

schisms, and opposing camps, as ecologists took different approaches and often failed to meet on common ground. Scope Ecology is usually considered a branch of biology, the general science that studies living and once-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 multicellular systems (in physiology and anatomy, to individuals (in botany, zoology, and other similar disciplines), and finally at the level of populations, communities, and ecosystems, and to the biosphere as a whole. These latter strata, from populations to the biosphere, are the primary subjects of ecological inquiries.

The complexity of the subject of ecological studies can be see in this river in South America, which is part of the world's largest wetland, the Pantanal. 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, chemistry, and physics. Thus, ecology is said to be a holistic science, one that overarches older disciplines, such as biology, which in this view become sub-disciplines contributing to ecological knowledge. 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." The term ecology is sometimes confused with the term environmentalism. Environmentalism is a social movement aimed at the goal of protecting natural resources or the environment, and which may involve political lobbying, activism, education, and so forth. Ecology is the science that studies living organisms and their interactions with the environment. As such, ecology involves scientific methodology and does not dictate what is "right" or "wrong." However, findings in ecology may be used to support or counter various goals, assertions, or actions of environmentalists. Consider the ways an ecologist might approach studying the life of honeybees:

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life processes and adaptations distribution and abundance of organisms the movement of materials and energy through living communities the successional development of ecosystems,[4] and the abundance and distribution of biodiversity in context of the environment.[1][2][3][4]

There are many practical applications of ecology in conservation biology, wetland management, natural resource management (agriculture, forestry, fisheries), city planning (urban ecology), community health, economics, basic & applied science and it provides a conceptual framework for understanding and researching human social interaction (human ecology).[4][6][7][8][9] cology or ecological science, is the scientific study of the distribution and abundance of living organisms and how these properties are affected by interactions between the organisms and their environment. The environment of an organism includes both the physical properties, which can be described as the sum of local abiotic factors like climate and geology, as well as the other organisms that share its habitat. Ecology may be more simply defined as the relationship between living organisms and their abiotic and biotic environment or as "the study of the structure and function of nature" (Odum 1971). In this later case, structure includes the distribution patterns and abundance of organisms, and function includes the interactions of populations, including competition, predation, symbiosis, and nutrient and energy cycles. The term ecology (oekologie) was coined in 1866 by the German biologist Ernst Haeckel. The word is derived from the Greek oikos ("household," "home," or "place to live") and logos ("study")—therefore, "ecology" means the "study of the household of nature." The name is derived from the same root word as economics (management of the household), and thus ecology is sometimes considered the economics of nature, or, as expressed by Ernst Haeckel, "the body of knowledge concerning the economy of nature" (Smith 1996). The interactions between living organisms and their abiotic and biotic environments, the focus of ecology, generally convey an overall sense of unity and harmony in nature. See for instance, species interactions. On the other hand, the history of the science itself has often revealed conflicts,

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. 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 science which can be subdivided into major and minor sub-disciplines. The major sub-disciplines include:

Ecological units For modern ecologists, ecology can be studied at several levels: population level (individuals of the same species), biocenosis level (or community of species), ecosystem level, biome 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. Most life exists on or within a few meters of the Earth's surface. Relative to the volume of the Earth, the biosphere is only the very thin surface layer that 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 (the area of water exposed to sufficient sunlight for photosynthesis). Multicellular organisms then appeared and colonized benthic zones. 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, utilizing the chemistry of the hot volcanic vents, as the base of its food chain. The biosphere contains great quantities of elements such as carbon, nitrogen, 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 their mineral and organic states. A biome is a homogeneous ecological formation that exists over a vast region, such 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 the physical environment (for example, oceans or mountain ranges) and on 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 a more complicated scheme, latitude and altitude approximate a good representation of the distribution of biodiversity within the biosphere. Very generally, biodiversity is greater near the equator (as in Brazil) and decreases as one approaches the poles. The biosphere may also be divided into ecozones, which are biogeographical and ecological land classifications, such as Neartic, Neotropic, and Oceanic. Biozones are very well defined today and primarily follow the continental borders.

Physiological ecology (or ecophysiology), which studies the influence of the biotic and abiotic environment on the physiology of the individual, and the adaptation of the individual to its environment; Behavioral ecology, which studies the ecological and evolutionary basis for animal behavior, and the roles of behavior in enabling animals to adapt to their ecological niches; Population ecology (or autecology), which deals with the dynamics of populations within species and the interactions of these populations with environmental factors; Community ecology (or synecology) which studies the interactions between species within an ecological community; Ecosystem ecology, which studies the flows of energy and matter through ecosystems; Medical ecology, which studies issues of human health in which environmental disturbances play a role Landscape ecology, which studies the interactions between discrete elements of a landscape and spatial patterns, including the role of disturbance and human impacts; Global ecology, which looks at ecological questions at the global level, often asking macroecological questions; Evolutionary ecology, which either can be considered the evolutionary histories of species and the interactions between them, or approaches the study of evolution by including elements of species interaction; And ecolinguistics, which looks at the relation between ecology and language.

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Ecology can also be sub-divided on the basis of target groups:

Animal ecology, plant ecology, insect ecology, human ecology, and so forth.

Ecology can, in addition, be sub-divided from the perspective of the studied biomes:

Arctic ecology (or polar ecology), tropical ecology, desert ecology, aquatic ecology, terrestrial ecology, wetland ecology, and temperate zone ecology.

Ecology can also be sub-divided on whether or not the emphasis is on application to human activities, such as resource management, environmental conservation, and restoration:

Theoretical ecology and applied ecology (including such subfields as landscape ecology, conservation biology, and restoration ecology).

Basic concepts in ecology Ecology is a very broad-ranging and complex topic, and even its definition lacks consensus. Thus, there are numerous concepts that fit within this discipline, and diverse manners in which the content can be arranged and studied. Several of the basic concepts of ecology include ecological units, the ecosystem, energy flow, nutrient cycles, species interaction, productivity, and ecological challenges.

Ecological factors that can affect dynamic change in a population or species in a given ecology or environment are usually divided into two groups: biotic and abiotic. Biotic factors relate to living organisms and their interactions. A biotic community is an assemblage of plant, animal, and other living organisms. 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:

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How could the colonization of a barren area be carried out? What are the ecosystem's dynamics and changes? How does an ecosystem interact at local, regional, and global scale? Is the current state stable? What is the value of an ecosystem? How does the interaction of ecological systems provide benefit to humans, especially in the provision of healthy water?

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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 a 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; and Natural disasters can also be considered abiotic.

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. Energy flow One focus of ecologists is to study the flow of energy, a major process linking the abiotic and biotic constituents of ecosystems. 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 that 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. See ecosystem for a more extensive explanation of energy flow in ecosystems. Nutrient cycles Ecologists also study the flow of nutrients in ecosystems. Whereas energy is not cycled, nutrients are cycled. Living organisms are composed mainly of carbon, oxygen, hydrogen, and nitrogen, and these four elements are cycled through the biotic communities and the geological world. These permanent recyclings of the elements are called biogeochemical cycles. Three fundamental biogeochemical cycles are the nitrogen cycle, the water cycle, and the carbonoxygen cycle. Another key cycle is the phosphorus cycle. Water is also exchanged between the hydrosphere, lithosphere, atmosphere, and biosphere. The oceans are large tanks that store water; they ensure thermal and climatic stability, as well as the transport of chemical elements thanks to large oceanic currents. Species interactions

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The ecosystem concept Main article: Ecosystem Ecologists study ecosystems such as this section of the Salobra River in the Pantanal area of Brazil. Some consider the ecosystem (abbreviation for "ecological system") to be the basic unit in ecology. An ecosystem is an ecological unit consisting of a biotic community together with its environment. Examples include a swamp, a meadow, and a river. It is generally considered smaller than a biome ("major life zone"), which is a large, geographic region of the earth's surface with distinctive plant and animal communities. A biome is often viewed as a grouping of many ecosystems sharing similar features, but is sometimes defined as an extensive ecosystem spread over a wide geographic area. 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. The ecosystem is composed of two entities, the entirety of life (the community, or biocoenosis) and the medium that life exists in (the biotope). Within the ecosystem, species are connected and dependent upon one another in the food chain, and exchange energy and matter between themselves and with their environment. The concept of an ecosystem can apply to units of variable size, such as a pond, a field, or a piece of deadwood. A unit of smaller size is called a microecosystem. For example, an ecosystem can be a stone and all the life under it. A mesoecosystem could be a forest, and a macroecosystem a whole ecoregion, with its watershed. Some of the main questions when studying an ecosystem include:

Biocenose, or community, is a group of populations of plants, animals, and microorganisms. 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.

negative feedback controls, supports 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). Overall, the interactions of organisms convey a sense of unity and harmony (see Biology:Interactions). Plants, through photosynthesis, use carbon dioxide and provide oxygen, while animals use oxygen and give off carbon dioxide. On the level of the food web, plants capture the sun's energy and serve as food for herbivores, which serve as food for carnivores, and ultimately top carnivores. Decomposers (bacteria, fungi, etc.) break down organisms after they die into minerals that can be used by plants. The harmony of species’ interactions with other species and the environment, including the biogeochemical cycles, have proposed a theory by some that the entire planet acts as if one, giant, functioning organism (the Gaia theory). Lynn Margulis and Dorion Sagan in their book Microcosmos (1997) even propose that evolution is tied to cooperation and mutual dependence among organisms: "Life did not take over the globe by combat, but by networking." The observed harmony can be attributed to the concept of dual purpose: the view that every entity in the universe in its interactions simultaneously exhibits purposes for the whole and for the individual—and that these purposes are interdependent. "Individual purpose" refers to the individual's requirement to met basic needs of self-preservation, selfstrengthening, multiplication, and development. The "whole purpose" is that by which the individual contributes to the preservation, strengthening, and development of the larger entity of which it is a part. Thus, the cell of a multicellular body provides a useful function for the body of which it is part. This "whole purpose," which could be the secretion of an enzyme, harmonizes with the body's requirement of selfpreservation, development, self-strengthening, and reproduction. The body, on the other hand, supports the cell's "individual purpose" by providing essential nutrients and carrying away wastes, assisting the cell's self-preservation, self-strengthening, multiplication, and development. Likewise, each individual organism exhibits both an individual purpose and a purpose for the whole related to its place in the environment. The result is an extraordinary harmony evident in creation. Ecosystem productivity

Mutual symbiosis between clownfish that dwell among the tentacles of tropical sea anemones, protects the anemone from anemone-eating fish, and in turn the stinging tentacles of the anemone protects the clownfish from its predators. Biotic ecological factors influence biocenose viability; these factors are considered as either intraspecific or interspecific relations. Intraspecific relations are those which are established between individuals of the same species, forming a population. They are relations of cooperation or competition, with division of the territory, and sometimes organization in hierarchical societies. Interspecific relations—interactions between different species—are numerous, and are usually described according to their beneficial, detrimental, or neutral effect (for example, mutualism or competition). Symbiosis refers to an interaction between two organisms living together in more or less intimate association. A significant relation is 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. Other interspecific relations include parasitism, infectious disease, and competition for limiting resources, which can occur when two species share the same ecological niche. 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:

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Producers—plants which are capable of photosynthesis Consumers—animals, which can be primary consumers (herbivorous), or secondary or tertiary consumers (carnivorous). Decomposers—bacteria, mushrooms, which degrade organic matter of all categories, and restore minerals to the environment.

The concepts dealing with the movement of energy through an ecosystem (via producers, consumers, and decomposers) 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 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. 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. The interactions and biogeochemical cycles create 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

A meandering river. 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. Thus, from an energy point of view, it is more efficient for humans to be primary consumers (to get nourishment from grains and vegetables) than as secondary consumers (from herbivores

such as beef and veal), and more still than as tertiary consumers (from eating carnivores). The productivity of ecosystems is sometimes estimated by comparing three types of land-based ecosystems and the total of aquatic ecosystems:

The forests (one-third 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 (one-third 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—(one-third of the Earth's land area) have very sparse biomasses and low productivity Finally, the marine and fresh water ecosystems (three-fourths of Earth's surface) contain very sparse biomasses (apart from the coastal zones).

In the case of a global crisis, the consequences can be much more significant; some extinction events showed the disappearance of more than 90 percent 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 may benefit other species, genera, families, orders, or phyla of organisms. Sometimes, an ecological crisis can be a specific and reversible phenomenon at the ecosystem scale. But more generally, the crisis’ impact will last. Indeed, it rather is a connected series of events that occur until 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. 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 nondegradable wastes, threats on key species (great apes, pandas, whales) are also factors influencing the well-being of people. During the past decades, this increasing responsibility of humanity in some ecological crises has been clearly observed. 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 usually quoted examples as ecological crises are:

Humanity's actions over the last few centuries have 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 challenges Generally, an ecological crisis is what occurs when the environment of a species or a population evolves in a way unfavorable to that species' survival. It may be that 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 unfavorable for the survival of a species (or a population) due to an increased pressure of predation (e.g., overfishing). It may be that the situation becomes unfavorable to the quality of life of the species (or the population) due to a rise in the number of individuals (overpopulation). Although ecological crises are generally considered to be something that occurs in a short time span (days, weeks, or years), by definition, ecological crises can also be considered to occur over a very long time period, such as millions of years. They can also be of natural or anthropic origin. They may relate to one unique species or to many species (see the article on extinction).

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Permian-Triassic extinction event—250 million of years ago Cretaceous-Tertiary extinction event—65 million years ago Ozone layer hole issue Deforestation and desertification, with the disappearance of many species The nuclear meltdown at Chernobyl in 1986 that 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 because of the large amount of radiation generated by the meltdown.

History of ecology Ecology is generally spoken of as a new science, having only become prominent in the second half of the twentieth century. Nonetheless, ecological thinking at some level has been around for a long time, and the principles of ecology have developed gradually, closely intertwined with the development of other biological disciplines. There is no consensus on its beginnings, as it developed more like a multi-stemmed bush than a tree with a single trunk (Smith 1996). Thus, one of the first ecologists may have been Aristotle or perhaps his friend and associate, Theophrastus, both of whom had interest in many species of animals. Theophrastus described interrelationships between animals and between animals and their environment as early as the fourth century B.C.E. (Ramalay 1940).

People often use fires to clear land for agriculture or grazing livestock in the Pantanal. Lastly, an ecological crisis may be local (an oil spill, a fire, or eutrophication of a lake), widespread (the movement of glaciers during an ice age), or global (a rise in the sea level). 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. Of course, what is an ecological crisis to one species, or one group of species, may be beneficial or neutral with respect to other species, at least short-term.

In general, the modern movement to ecology through botanical geography (which led to plant ecology) developed earlier than animal ecology. Throughout the eighteenth and the beginning of the nineteenth century, the great maritime powers such as Britain, Spain, and Portugal launched many world exploratory expeditions. These expeditions were joined by many scientists, including botanists, such as the German explorer Alexander von Humboldt. Humboldt is often considered a father of ecology. He was the first to take on the study of the relationship between organisms and their environment. He exposed the existing relationships between observed plant species and climate, and described vegetation zones using latitude and altitude, a discipline now known as geobotany. With the publication of the work of Charles Darwin on The Origin of Species, ecology passed from a repetitive, mechanical model to a biological, organic, and hence evolutionary model. Alfred Russel Wallace, contemporary and competitor to Darwin, was first to propose a "geography" of animal species. Several authors recognized at the time that species were not independent of each other, and grouped them into plant species, animal species, and later into communities of living beings or "biocoenosis." This term, which comes from Greek, was coined in 1877 by marine biologist Karl Möbius, and essentially means "life having something in common." By the nineteenth century, ecology blossomed due to new discoveries in chemistry by Lavoisier and Horace-Bénédict de Saussure, notably the nitrogen cycle. After observing the fact that life developed only within strict limits of each compartment that makes up the atmosphere, hydrosphere, and lithosphere, the Austrian geologist Eduard Suess proposed the term biosphere in 1875. He used the name biosphere for the conditions promoting life, such as those found on Earth, which include flora, fauna, minerals, matter cycles, and so forth. In the 1920s, Vladimir Vernadsky, a Russian geologist who had defected to France, detailed the idea of the biosphere in his work The biosphere (1926), and described the fundamental principles of the biogeochemical cycles. Ecological damages were reported in the eighteenth century, as the multiplication of colonies impacted deforestation. Since the nineteenth century, with the Industrial Revolution, more and more pressing concerns have grown about the impact of human activity on the environment. The term ecologist has been in use since the end of the nineteenth century. Over the nineteenth century, botanical geography and zoogeography combined to form the basis of biogeography. This science, which deals with habitats of species, seeks to explain the reasons for the presence of certain species in a given location. Pioneers in animal ecology were early twentieth-century scientists R. Hesse and Charles Eton, Charles Adams, and Victor Shelford. It was in 1935 that Arthur Tansley, the British ecologist, coined the term ecosystem, the interactive system established between the biocoenosis (the group of living creatures), and their biotope (the environment in which they live). Ecology thus became the science of ecosystems. Tansley's concept of the ecosystem was adopted by the energetic and influential biology educator Eugene Odum. Along with his brother, Howard Odum, Eugene Odum wrote a textbook which (starting in 1953) educated multiple generations of biologists and ecologists in North America.

At the turn of the twentieth century, Henry Chandler Cowles was one of the founders of the emerging study of "dynamic ecology," through his study of ecological succession at the Indiana Dunes, sand dunes at the southern end of Lake Michigan. Here Cowles found evidence of ecological succession in the vegetation and the soil with relation to age. Ecological succession is the process by which a natural community moves from a simpler level of organization to a more complex community (e.g., from bare sand, to grass growing on the sand, to grass growing on dirt produced from dead grass, to trees growing in the dirt produced by the grass). Human ecology began in the 1920s, through the study of changes in vegetation succession in the city of Chicago, Illinois. It became a distinct field of study in the 1970s. This marked recognition that humans, who had colonized all of the Earth's continents, were a major ecological factor. Humans greatly modify the environment through the development of the habitat (in particular urban planning), by intensive activities such as logging and fishing, and as side effects of agriculture, mining, and industry. Besides ecology and biology, this discipline involved many other natural and social sciences, such as anthropology and ethnology, economics, demography, architecture and urban planning, medicine and psychology, and many more. The development of human ecology led to the increasing role of ecological science in the design and management of cities. The history of ecology has been one of conflicts and opposing camps. Smith (1996) notes that the first major split in ecology was between plant ecology and animal ecology, which even lead to a controversy over the term ecology, with botanists dropping the initial "o" from oecology, the spelling in use at the time, and zoologists refusing to use the term ecology at all, because of its perceived affiliation with botany. Other historical schisms were between organismal and individualist ecology, holism versus reductionism, and theoretical versus applied ecology.

Levels of organization and study

Ecosystems regenerate after a disturbance such as fire, forming mosaics of different age groups structured across a landscape. Pictured are different seral stages in forested ecosystems starting from pioneers colonizing a disturbed site and maturing in successional stages leading to old-growth forests. The processes that influence ecological phenomena vary through space and time.[10] It can take thousands of years for ecological processes to mature; the life-span of a tree, for example, can encompass different successional stages. The ecological process is extended even further through time as trees die, decay and provide habitat as nurse logs or coarse woody debris. The area of an ecosystem can vary greatly from tiny to vast. A single tree is of little consequence to the classification of a forest ecosystem, but it is far more significant to smaller organisms.[11] Several generations of an aphid population can exist over the lifespan of a single leaf. Each of those aphids, in turn, support diverse bacterial communities.[12] Tree growth is related to local site variables, such as soil type, moisture content, slope of the land, and forest canopy closure. More complex global factors, such as climate, must be considered for the classification and understanding of processes leading to larger patterns spanning across a forested landscape.[13] Global patterns of biological diversity are complex. This biocomplexity stems from the interplay among ecological

processes that operate and influence patterns that grade into each other, such as transitional areas or ecotones that stretch across different scales.[14] "Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral, and geometric."[15]:3 There are different views on what constitutes complexity. One perspective lumps things that we do not understand into this category by virtue of the computational effort it would require to piece together the numerous interacting parts. Alternatively, complexity in life sciences can be viewed as emergent self-organized systems with multiple possible outcomes directed by random accidents of history.[16] Small scale patterns do not necessarily explain large scale phenomena, otherwise captured in the expression 'the sum is greater than the parts'.[17][18] Ecologists have identified emergent and self-organizing phenomena that operate at different environmental scales of influence, ranging from molecular to planetary, and these require different sets of scientific explanation.[19][20] Long-term ecological studies provide important track records to better understand the complexity of ecosystems over longer temporal and broader spatial scales. The International Long Term Ecological Network[21] manages and exchanges scientific information among research sites. The longest experiment in existence is the Park Grass Experiment that was initiated in 1856.[22] Another example includes the Hubbard Brook study in operation since 1960.[23] To structure the study of ecology into a manageable framework of understanding, the biological world is conceptually organized as a nested hierarchy of organization, ranging in scale from genes, to cells, to tissues, to organs, to organisms, to species and up to the level of the biosphere.[24] Together these hierarchical scales of life form a panarchy.[16] Ecosystems are primarily researched at three key levels of organization—organisms, populations, and communities. Ecologists study ecosystems by sampling a certain number of individuals that are representative of a population. Ecosystems consist of communities interacting with each other and the environment. In ecology, communities are created by the interaction of the populations of different species in an area.[25][26] Biodiversity is an attribute of a site or area that consists of the variety within and among biotic communities, whether influenced by humans or not, at any spatial scale from microsites and habitat patches to the entire biosphere.[27] :745 Biodiversity (an abbreviation of biological diversity) describes the diversity of life from genes to ecosystems and spans every level of biological organization. There are many ways to index, measure, and represent biodiversity.[28] Biodiversity includes species diversity, ecosystem diversity, genetic diversity and the complex processes operating at and among these respective levels.[28][29][30] Biodiversity plays an important role in ecological health as much as it does for human health. [31][32] Preventing or prioritizing species extinctions is one way to preserve biodiversity, but populations, the genetic diversity within them and ecological processes, such as migration, are being threatened on global scales and disappearing rapidly as well. Conservation priorities and management techniques require different approaches and considerations to address the full ecological scope of biodiversity. Populations and species migration, for example, are more sensitive indicators of ecosystem services that sustain and contribute natural capital toward the well-being of humanity.[33][34][35][36] An understanding of biodiversity has practical application for ecosystem-based conservation planners as they make ecologically responsible decisions in management recommendations to consultant firms, governments and industry.[37] [edit] Ecological niche and habitat

Termite mounds with varied heights of chimneys regulate gas exchange, temperature and other environmental parameters that are needed to sustain the internal physiology of the entire colony.[38][39] There are many definitions of the niche dating back to 1917, [40] but G. Evelyn Hutchinson made conceptual advances in 1957[41][42] and introduced the most widely accepted definition: "The niche is the set of biotic and abiotic conditions in which a species is able to persist and maintain stable population sizes."[40]:519 The ecological niche is a central concept in the ecology of organisms and is sub-divided into the fundamental and the realized niche. The fundamental niche is the set of environmental conditions under which a species is able to persist. The realized niche is the set of environmental plus ecological conditions under which a species persists.[25][40][42] The habitat of a species is a related but distinct concept that describes the environment over which a species is known to occur and the type of community that is formed as a result.[43] More specifically, "habitats can be defined as regions in environmental space that are composed of multiple dimensions, each representing a biotic or abiotic environmental variable; that is, any component or characteristic of the environment related directly (e.g. forage biomass and quality) or indirectly (e.g. elevation) to the use of a location by the animal."[44]:745 For example, the habitat might refer to an aquatic or terrestrial environment that can be further categorized as montane or alpine ecosystems. Biogeographical patterns and range distributions are explained or predicted through knowledge and understanding of a species traits and niche requirements.[45] Species have functional traits that are uniquely adapted to the ecological niche. A trait is a measurable property of an organism that influences its performance.[46] Traits of each species are suited ar uniquely adapted to their ecological niche. This means that resident species are at an advantage and able to competitively exclude other similarly adapted species from having an overlapping geographic range. This is called the competitive exclusion principle.[47]

Biodiversity of a coral reef. Corals adapt and modify their environment by forming calcium carbonate skeletons that provide growing conditions for future generations and form habitat for many other species.[48] Organisms are subject to environmental pressures, but they are also modifiers of their habitats. The regulatory feedback between organisms and their environment can modify conditions from local (e.g., a pond) to global scales (e.g., Gaia), over time and even after death, such as decaying logs or silica skeleton deposits from marine organisms.[49] The

process and concept of ecosystem engineering has also been called niche construction. Ecosystem engineers are defined as: "...organisms that directly or indirectly modulate the availability of resources to other species, by causing physical state changes in biotic or abiotic materials. In so doing they modify, maintain and create habitats."[50]:373 The ecosystem engineering concept has stimulated a new appreciation for the degree of influence that organisms have on the ecosystem and evolutionary process. The terms niche construction are more often used in reference to the under appreciated feedback mechanism of natural selection imparting forces on the abiotic niche.[38][51] An example of natural selection through ecosystem engineering occurs in the nests of social insects, including ants, bees, wasps, and termites. There is an emergent homeostasis in the structure of the nest that regulates, maintains and defends the physiology of the entire colony. Termite mounds, for example, maintain a constant internal temperature through the design of air-conditioning chimneys. The structure of the nests themselves are subject to the forces of natural selection. Moreover, the nest can survive over successive generations, which means that ancestors inherit both genetic material and a legacy niche that was constructed before their time.[38][39][52] Diatoms in the Bay of Fundy, Canada, provide another example of an ecosystem engineer. Benthic diatoms living in estuarine sediments secrete carbohydrate exudates that bind the sand and stabilizes the environment. The diatoms cause a physical state change in the properties of the sand that allows other organisms to colonize the area. The concept of ecosystem engineering brings new conceptual implications for the discipline of conservation biology.[53] [edit] Population ecology Main article: Population ecology The population is the unit of analysis in population ecology. A population consists of individuals of the same species that live, interact and migrate through the same niche and habitat. [54] A primary law of population ecology is the Malthusian growth model.[55] This law states that: "...a population will grow (or decline) exponentially as long as the environment experienced by all individuals in the population remains constant."[55]:18 This Malthusian premise provides the basis for formulating predictive theories and tests that follow. Simplified population models usually start with four variables including death, birth, immigration, and emigration. Mathematical models are used to calculate changes in population demographics using a null model. A null model is used as a null hypothesis for statistical testing. The null hypothesis states that random processes create observed patterns. Alternatively the patterns differ significantly from the random model and require further explanation. Models can be mathematically complex where "...several competing hypotheses are simultaneously confronted with the data."[56] An example of an introductory population model describes a closed population, such as on an island, where immigration and emigration does not take place. In these island models the rate of population change is described by: , where N is the total number of individuals in the population, B is the number of births, D is the number of deaths, b and d are the per capita rates of birth and death respectively, and r is the per capita rate of population change. This formula can be read out as the rate of change in the population (dN/dT) is equal to births minus deaths (B – D).[55][57]

Using these modelling techniques, Malthus' population principle of growth was later transformed into a model known as the logistic equation:

, where N is the number of individuals measured as biomass density, a is the maximum per-capita rate of change, and K is the carrying capacity of the population. The formula can be read as follows: the rate of change in the population (dN/dT) is equal to growth (aN) that is limited by carrying capacity (1 – N/K). The discipline of population ecology builds upon these introductory models to further understand demographic processes in real study populations and conduct statistical tests. The field of population ecology often uses data on life history and matrix algebra to develop projection matrices on fecundity and survivorship. This information is used for managing wildlife stocks and setting harvest quotas.[57][58]

A list of terms that define various types of natural groupings of individuals that are used in population studies[59] Term Species population Definition All individuals of a species.

Metapopulatio A set of spatially disjunct populations, n among which there is some immigration. Population A group of conspecific individuals that is demographically, genetically, or spatially disjunct from other groups of individuals. A spatially clustered group of individuals. A group of individuals more genetically similar to each other than to other individuals, usually with some degree of spatial isolation as well. A group of individuals within an investigatordelimited area smaller than the geographic range of the species and often within a population (as defined above). A local population could be a disjunct population as well.

Aggregation Deme

Local population

Subpopulatio An arbitrary spatially delimited subset of n individuals from within a population (as defined above).

[edit] Metapopulation ecology Main article: Metapopulation Populations are also studied and modeled according to the metapopulation concept. The metapopulation concept was introduced in 1969:[60] "as a population of populations which go extinct locally and recolonize."[61]:105 Metapopulation ecology is another statistical approach that is often used in conservation research.[62] Metapopulation research simplifies the landscape into patches of varying levels of quality.[63]

Metapopulation models have been used to explain life-history evolution, such as the ecological stability of amphibian metamorphosis in small vernal ponds. Alternative ecological strategies have evolved. For example, some salamanders forgo metamorphosis and sexually mature as aquatic neotenes. The seasonal duration of wetlands and the migratory range of the species determines which ponds are connected and if they form a metapopulation. The duration of the life history stages of amphibians relative to the duration of the vernal pool before it dries up regulates the ecological development of metapopulations connecting aquatic patches to terrestrial patches.[64] In metapopulation terminology there are emigrants (individuals that leave a patch), immigrants (individuals that move into a patch) and sites are classed either as sources or sinks. A site is a generic term that refers to places where ecologists sample populations, such as ponds or defined sampling areas in a forest. Source patches are productive sites that generate a seasonal supply of juveniles that migrate to other patch locations. Sink patches are unproductive sites that only receive migrants and will go extinct unless rescued by an adjacent source patch or environmental conditions become more favorable. Metapopulation models examine patch dynamics over time to answer questions about spatial and demographic ecology. The ecology of metapopulations is a dynamic process of extinction and colonization. Small patches of lower quality (i.e., sinks) are maintained or rescued by a seasonal influx of new immigrants. A dynamic metapopulation structure evolves from year to year, where some patches are sinks in dry years and become sources when conditions are more favorable. Ecologists use a mixture of computer models and field studies to explain metapopulation structure.[65][66] [edit] Community ecology Main article: Community ecology Community ecology examines how interactions among species and their environment affect the abundance, distribution and diversity of species within communities. Johnson & Stinchcomb[67]:250 Community ecology is a subdiscipline of ecology which studies the distribution, abundance, demography, and interactions between coexisting populations. An example of a study in community ecology might measure primary production in a wetland in relation to decomposition and consumption rates. This requires an understanding of the community connections between plants (i.e., primary producers) and the decomposers (e.g., fungi and bacteria).[68] or the analysis of predator-prey dynamics affecting amphibian biomass.[69] Food webs and trophic levels are two widely employed conceptual models used to explain the linkages among species.[70][71] [edit] Food webs Main article: Food webs A food web is the archetypal ecological network. They are a type of concept map that illustrate pathways of energy flows in an ecological community, usually starting with solar energy being used by plants during photosynthesis. As plants grow, they accumulate carbohydrates and are eaten by grazing herbivores. Step by step lines or relations are drawn until a web of life is illustrated.[72][73][74][75]

Generalized food web of waterbirds from Chesapeake Bay There are different ecological dimensions that can be mapped to create more complicated food webs, including: species composition (type of species), richness (number of species), biomass (the dry weight of plants and animals), productivity (rates of conversion of energy and nutrients into growth), and stability (food webs over time). A food web diagram illustrating species composition shows how change in a single species can directly and indirectly influence many others. Microcosm studies are used to simplify food web research into semi-isolated units such as small springs, decaying logs, and laboratory experiments using organisms that reproduce quickly, such as daphnia feeding on algae grown under controlled environments in jars of water.[76][77] Principles gleaned from food web microcosm studies are used to extrapolate smaller dynamic concepts to larger systems.[77] Food webs are limited because they are generally restricted to a specific habitat, such as a cave or a pond. The food web illustration (right) only shows a small part of the complexity connecting the aquatic system to the adjacent terrestrial land. Many of these species migrate into other habitats to distribute their effects on a larger scale. In other words, food webs are incomplete, but are nonetheless a valuable tool in understanding community ecosystems.[78] 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.[79]:269 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.[79] 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 (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.[77]

[edit] Trophic dynamics Main article: Trophic dynamics The Greek root of the word troph, τροφή, trophē, means food or feeding. Links in food-webs primarily connect feeding relations or trophism among species. Biodiversity within ecosystems can be organized into vertical and horizontal dimensions. The vertical dimension represents feeding relations that become further removed from the base of the food chain up toward top predators. The horizontal dimension represents the abundance or biomass at each level.[80] When the relative abundance or biomass of each functional feeding group is stacked into their respective trophic levels they naturally sort into a ‘pyramid of numbers’.[81] Functional groups are broadly categorized as autotrophs (or primary producers), heterotrophs (or consumers), and detrivores (or decomposers). Heterotrophs can be further sub-divided into different functional groups, including: primary consumers (strict herbivores), secondary consumers (predators that feed exclusively on herbivores) and tertiary consumers (predators that feed on a mix of herbivores and predators).[82] Omnivores do not fit neatly into a functional category because they eat both plant and animal tissues. It has been suggested that omnivores have a greater functional influence as predators

Freshwater aquatic and terrestrial food-webs

because relative to herbivores they are comparatively inefficient at grazing.[83] Ecologist collect data on trophic levels and food webs to statistically model and mathematically calculate parameters, such as those used in other kinds of network analysis (e.g., graph theory), to study emergent patterns and properties shared among ecosystems. The emergent pyramidal arrangement of trophic levels with amounts of energy transfer decreasing as species become further removed from the source of production is one of several patterns that is repeated amongst the planets ecosystems.[75][84][85] The size of each level in the pyramid generally represents biomass, which can be measured as the dry weight of an organism.[86] Autotrophs may have the highest global proportion of biomass, but they are closely rivaled or surpassed by microbes.[87][88] The decomposition of dead organic matter, such as leaves falling on the forest floor, turns into soils that feed plant production. The total sum of the planet's soil ecosystems is called the pedosphere where a very large proportion of the Earth's biodiversity sorts into other trophic levels. Invertebrates that feed and shred larger leaves, for example, create smaller bits for smaller organisms in the feeding chain. Collectively, these are the detrivores that regulate soil formation.[89][90] Tree roots, fungi, bacteria, worms, ants, beetles, centipedes, spiders, mammals, birds, reptiles, amphibians and other less familiar creatures all work to create the trophic web of life in soil ecosystems. As organisms feed and migrate through soils they physically displace materials, which is an important ecological process called bioturbation. Biomass of soil microorganisms are influenced by and feed back into the trophic dynamics of the exposed solar surface ecology. Paleoecological studies of soils places the origin for bioturbation to a time before the Cambrian period. Other events, such as the evolution of trees and amphibians moving into land in the Devonian period played a significant role in the development of soils and ecological trophism.[69][90][91]

species into a trophic system gives a macroscopic image of the larger functional design.[94] 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 herbibores 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 foodweb, such as the distribution of plant biomass.[95] [edit] Keystone species A keystone species is a species that is disproportionately connected to more species in the food-web. Keystone species have lower levels of biomass in the trophic pyramid relative to the importance of their role. The many connections that a keystone species holds means that it maintains the organization and structure of entire communities. The loss of a keystone species results in a range of dramatic cascading effects that alters trophic dynamics, other food-web connections and can cause the extinction of other species in the community.[96][97] Sea otters (Enhydra lutris) are commonly cited as an example of a keystone species because they limit the density of sea urchins that feed on kelp. If sea otters are removed from the system, the urchins graze until the kelp beds disappear and this has a dramatic effect on community structure.[98] Hunting of sea otters, for example, is thought to have indirectly led to the extinction of the Steller's Sea Cow (Hydrodamalis gigas). [99] While the keystone species concept has been used extensively as a conservation tool, it has been criticized for being poorly defined from an operational stance. It is very difficult to experimentally determine in each different ecosystem what species may hold a keystone role. Furthermore, food-web theory suggests that keystone species may not be all that common. It is therefore unclear how generally the keystone species model can be applied.[95][98] [edit] Ecosystem ecology Main article: Ecosystem ecology These ecosystems, as we may call them, are of the most various kinds and sizes. They form one category of the multitudinous physical systems of the universe, which range from the universe as a whole down to the atom. Tansley[100]:299 The concept of the ecosystem was first introduced in 1935 to describe habitats within biomes that form an integrated whole and a dynamically responsive system having both physical and biological complexes. Within an ecosystem there are inseparable ties that link organisms to the physical and biological components of their environment to which they are adapted.[100] Ecosystems are complex adaptive systems where the interaction of life processes form self-organizing patterns across different scales of time and space.[101] This section introduces key areas of ecosystem ecology that are used to inquire, understand and explain observed patterns of biodiversity and ecosystem function across different scales of organization.

List of ecological functional groups, definitions and examples Functional group Producers or autotrophs Definition and examples Usually plants or cyanobacteria that are capable of photosynthesis but could be other organisms such as the bacteria near ocean vents that are capable of chemosynthesis.

Consumers or Animals, which can be primary consumers heterotrophs (herbivorous), or secondary or tertiary consumers (carnivorous and omnivores). Decomposers Bacteria, fungi, and insects which degrade or detritivores organic matter of all types and restore nutrients to the environment. The producers will then consume the nutrients, completing the cycle.

Functional trophic groups sort out hierarchically into pyramidic trophic levels because it requires specialized adaptations to become a photosynthesizer or a predator, so few organisms have the adaptations needed to combine both abilities. This explains why functional adaptations to trophism (feeding) organizes different species into emergent functional groups.[83] Trophic levels are part of the holistic or complex systems view of ecosystems.[92][93] Each trophic level contains unrelated species that grouped together because they share common ecological functions. Grouping functionally similar

[edit] The biome Main article: Biome Ecological units of organization are defined through reference to any magnitude of space and time on the planet. Communities of organisms, for example, are somewhat arbitrarily defined, but the processes of life integrate at different levels and organize into more complex wholes. Biomes, for example, are a larger unit of organization that categorize regions of the Earth's ecosystems mainly according to the structure and composition of vegetation.[102] Different researchers have applied different methods to define continental boundaries of biomes dominated by different functional types of vegetative communities that are limited in distribution by climate, precipitation, weather and other environmental variables. Examples of biome names include: tropical rainforest, temperate broadleaf and mixed forests, temperate deciduous forest, taiga, tundra, hot desert, and polar desert.[103] Other researchers have recently started to categorize other types of biomes, such as the human and oceanic microbiomes. To a microbe, the human body is a habitat and a landscape.[104] The microbiome has been largely discovered through advances in molecular genetics that have revealed a hidden richness of microbial diversity on the planet. The oceanic microbiome plays a significant role in the ecological biogeochemistry of the planet's oceans.[105] [edit] The biosphere Main article: Biosphere Ecological theory has been used to explain self-emergent regulatory phenomena at the planetary scale. The largest scale of ecological organization is the biosphere: the total sum of ecosystems on the planet. Ecological relations regulate the flux of energy, nutrients, and climate all the way up to the planetary scale. For example, the dynamic history of the planetary CO2 and O2 composition of the atmosphere has been largely determined by the biogenic flux of gases coming from respiration and photosynthesis, with levels fluctuating over time and in relation to the ecology and evolution of plants and animals.[106] When sub-component parts are organized into a whole there are oftentimes emergent properties that describe the nature of the system. This the Gaia hypothesis,[20] and is an example of holism applied in ecological theory.[107] The ecology of the planet acts as a single regulatory or holistic unit called Gaia. The Gaia hypothesis states that there is an emergent feedback loop generated by the metabolism of living organisms that maintains the temperature of the Earth and atmospheric conditions within a narrow self-regulating range of tolerance.
[20]

processes operating across different spatial or temporal scales of organization.[20][110][111] While the boundary between ecology and evolution is not always clear, it is understood that ecologists study the abiotic and biotic factors that influence the evolutionary process.[2][86] [edit] Behavioral ecology Main article: Behavioral ecology

Social display and color variation in differently adapted species of chameleons (Bradypodion spp.). Chameleons change their skin color to match their background as a behavioral defense mechanism and also use color to communicate with other members of their species, such as dominant (left) versus submissive (right) patterns shown in the three species (A-C) above.[112] All organisms are motile to some extent. Even plants express complex behavior, including memory and communication.[113] Behavioural ecology is the study of ethology and its ecological and evolutionary implications. Ethology is the study of observable movement or behaviour in nature. This could include investigations of motile sperm of plants, mobile phytoplankton, zooplankton swimming toward the female egg, the cultivation of fungi by weevils, the mating dance of a salamander, or social gatherings of amoeba.[114][115][116][117][118] Adaptation is the central unifying concept in behavioral ecology."International Society for Behavioral Ecology". http://www.behavecol.com/pages/society/welcome.html. Behaviors can be recorded as traits and inherited in much the same way that eye and hair color can. Behaviours evolve and become adapted to the ecosystem because they are subject to the forces of natural selection.[29] Hence, behaviors can be adaptive, meaning that they evolve functional utilities that increases reproductive success for the individuals that inherit such traits.[119] This is also the technical definition for fitness in biology, which is a measure of reproductive success over successive generations.[29] Predator-prey interactions are an introductory concept into food-web studies as well as behavioural ecology.[120] Prey species can exhibit different kinds of behavioural adaptations to predators, such as avoid, flee or defend. Many prey species are faced with multiple predators that differ in the degree of danger posed. To be adapted to their environment and face predatory threats, organisms must balance their energy budgets as they invest in different aspects of their life history, such as growth, feeding, mating, socializing, or modifying their habitat. Hypotheses posited in behavioural ecology are generally based on adaptive principals of conservation, optimization or efficiency.[1][2][3][121] For example, "The threat-sensitive predator avoidance hypothesis predicts that prey should assess the degree of threat posed by different predators and match their behavior according to current levels of risk."[122]

[edit] Relation to evolution Ecology and evolution are considered sister disciplines of the life sciences. Natural selection, life history, development, adaptation, populations, and inheritance are examples of concepts that thread equally into ecological and evolutionary theory. Morphological, behavioural and/or genetic traits, for example, can be mapped onto evolutionary trees to study the historical development of a species in relation to their functions and roles in different ecological circumstances. In this framework, the analytical tools of ecologists and evolutionists overlap as they organize, classify and investigate life through common systematic principals, such as phylogenetics or the Linnaean system of taxonomy.[108] The two disciplines often appear together, such as in the title of the journal Trends in Ecology and Evolution.[109] There is no sharp boundary separating ecology from evolution and they differ more in their areas of applied focus. Both disciplines discover and explain emergent and unique properties and

"The optimal flight initiation distance occurs where expected postencounter fitness is maximized, which depends on the prey’s initial fitness, benefits obtainable by not fleeing, energetic escape costs, and expected fitness loss due to predation risk."[123] The behaviour of long-toed salamanders (Ambystoma macrodactylum) presents an example in this context. When threatened, the long-toed salamander defends itself by waving its tail and secreting a white milky fluid.[124][125] The excreted fluid is distasteful, toxic and adhesive, but it is also used for nutrient and energy storage during hibernation. Hence, salamanders subjected to frequent predatory attack will be energetically compromised as they use up their energy stores.[126][127] Ecological interactions can be divided into host and associate relationships. A host is any entity that harbors another that is called the associate.[129] Host and associate relationships among species that are mutually or reciprocally beneficial are called mutualisms. If the host and associate are physically connected, the relationship is called symbiosis. Approximately 60% of all plants, for example, have a symbiotic relationship with arbuscular mycorrhizal fungi. Symbiotic plants and fungi exchange carbohydrates for mineral nutrients.[130] Symbiosis differs from indirect mutualisms where the organisms live apart. For example, tropical rainforests regulate the Earth's atmosphere. Trees living in the equatorial regions of the planet supply oxygen into the atmosphere that sustains species living in distant polar regions of the planet. This relationship is called commensalism because many other host species receive the benefits of clean air at no cost or harm to the associate tree species supplying the oxygen.[131] The host and associate relationship is called parasitism if one species benefits while the other suffers. Competition among species or among members of the same species is defined as reciprocal antagonism, such as grasses competing for growth space.[132]

poor years and all birds to survive when food is plentiful.[137]
[138]

Elaborate sexual displays and posturing are encountered in the behavioural ecology of animals. The birds of paradise, for example, display elaborate ornaments and song during courtship. These displays serve a dual purpose of signalling healthy or well-adapted individuals and good genes. The elaborate displays are driven by sexual selection as an advertisement of quality of traits among male suitors.[139] [edit] Biogeography Main article: Biogeography The word biogeography is an amalgamation of biology and geography. Biogeography is the comparative study of the geographic distribution of organisms and the corresponding evolution of their traits in space and time.[140] The Journal of Biogeography was established in 1974.[141] Biogeography and ecology share many of their disciplinary roots. For example, the theory of island biogeography, published by the mathematician Robert MacArthur and ecologist Edward O. Wilson in 1967[142] is considered one of the fundamentals of ecological theory.[143] Biogeography has a long history in the natural sciences where questions arise concerning the spatial distribution of plants and animals. Ecology and evolution provide the explanatory context for biogeographical studies.[140] Biogeographical patterns result from ecological processes that influence range distributions, such as migration and dispersal.[143] and from historical processes that split populations or species into different areas.[144] The biogeographic processes that result in the natural splitting of species explains much of the modern distribution of the Earth's biota. The splitting of lineages in a species is called vicariance biogeography and it is a subdiscipline of biogeography.[144][145][146] There are also practical applications in the field of biogeography concerning ecological systems and processes. For example, the range and distribution of biodiversity and invasive species responding to climate change is a serious concern and active area of research in context of global warming.[34][147] [edit] r/K-Selection theory Main article: r/K selection A population ecology concept (introduced in MacArthur and Wilson's (1967) book, The Theory of Island Biogeography) is r/K selection theory, one of the first predictive models in ecology used to explain life-history evolution. The premise behind the r/K selection model is that natural selection pressures change according to population density. For example, when an island is first colonized, density of individuals is low. The initial increase in population size is not limited by competition, leaving an abundance of available resources for rapid population growth. These early phases of population growth experience density-independent forces of natural selection, which is called r-selection. As the population becomes more crowded, it approaches the island's carrying capacity, thus forcing individuals to compete more heavily for fewer available resources. Under crowded conditions the population experiences density-dependent forces of natural selection, called K-selection.[148] In the r/K-selection model, the first variable r is the intrinsic rate of natural increase in population size and the second variable K is the carrying capacity of a population.[25] Different species evolve different life-history strategies spanning a continuum between these two selective forces. An r-selected

Popular ecological study systems for mutualism include, fungus-growing ants employing agricultural symbiosis, bacteria living in the guts of insects and other organisms, the fig wasp and yucca moth pollination complex, lichens with fungi and photosynthetic algae, and corals with photosynthetic algae.[133][134] Intraspecific behaviours are notable in the social insects, slime moulds, social spiders, human society, and naked mole rats where eusocialism has evolved. Social behaviours include reciprocally beneficial behaviours among kin and nest mates. [29][116][135] Social behaviours evolve from kin and group selection. Kin selection explains altruism through genetic relationships, whereby an altruistic behaviour leading to death is rewarded by the survival of genetic copies distributed among surviving relatives. The social insects, including ants, bees and wasps are most famously studied for this type of relationship because the male drones are clones that share the same genetic make-up as every other male in the colony. [29] In contrast, group selectionists find examples of altruism among non-genetic relatives and explain this through selection acting on the group, whereby it becomes selectively advantageous for groups if their members express altruistic behaviours to one another. Groups that are predominantely altruists beat groups that are predominantely selfish.[29][136] A often quoted behavioural ecology hypothesis is known as Lack's brood reduction hypothesis (named after David Lack). Lack's hypothesis posits an evolutionary and ecological explanation as to why birds lay a series of eggs with an asynchronous delay leading to nestlings of mixed age and weights. According to Lack, this brood behaviour is an ecological insurance that allows the larger birds to survive in

species is one that has high birth rates, low levels of parental investment, and high rates of mortality before individuals reach maturity. Evolution favors high rates of fecundity in rselected species. Many kinds of insects and invasive species exhibit r-selected characteristics. In contrast, a K-selected species has low rates of fecundity, high levels of parental investment in the young, and low rates of mortality as individuals mature. Humans and elephants are examples of species exhibiting K-selected characteristics, including longevity and efficiency in the conversion of more resources into fewer offspring.[142][149] [edit] Molecular ecology Main article: Molecular ecology The important relationship between ecology and genetic inheritance predates modern techniques for molecular analysis. Molecular ecological research became more feasible with the development of rapid and accessible genetic technologies, such as the polymerase chain reaction (PCR). The rise of molecular technologies and influx of research questions into this new ecological field resulted in the publication Molecular Ecology in 1992.[150] Molecular ecology uses various analytical techniques to study genes in an evolutionary and ecological context. In 1994, professor John Avise also played a leading role in this area of science with the publication of his book, Molecular Markers, Natural History and Evolution.[151] Newer technologies opened a wave of genetic analysis into organisms once difficult to study from an ecological or evolutionary standpoint, such as bacteria, fungi and nematodes. Molecular ecology engendered a new research paradigm to investigate ecological questions considered otherwise intractable. Molecular investigations revealed previously obscured details in the tiny intricacies of nature and improved resolution into probing questions about behavioral and biogeographical ecology. For example, molecular ecology revealed promiscuous sexual behavior and multiple male partners in tree swallows previously thought to be socially monogamous.[152] In a biogeographical context, the marriage between genetics, ecology and evolution resulted in a new sub-discipline called phylogeography.[153] [edit] Relation to the environment The environment is dynamically interlinked, imposed upon and constrains organisms at any time throughout their life cycle.[154] Like the term ecology, environment has different conceptual meanings and to many these terms also overlap with the concept of nature. Environment "...includes the physical world, the social world of human relations and the built world of human creation."[155]:62 The environment in ecosystems includes both physical parameters and biotic attributes. The physical environment is external to the level of biological organization under investigation, including abiotic factors such as temperature, radiation, light, chemistry, climate and geology. The biotic environment includes genes, cells, organisms, members of the same species (conspecifics) and other species that share a habitat.[156] The laws of thermodynamics applies to ecology by means of its physical state. Armed with an understanding of metabolic and thermodynamic principles a complete accounting of energy and material flow can be traced through an ecosystem.[157] Environmental and ecological relations are studied through reference to conceptually manageable and isolated parts. Once the effective environmental components are understood they conceptually link back together as a holocoenotic[158] system. In other words, the organism and the environment form a dynamic whole (or umwelt).[159]:252 Change in one

ecological or environmental factor can concurrently affect the dynamic state of an entire ecosystem.[160][161] Ecological studies are necessarily holistic as opposed to reductionistic.[19][162] Holism has three scientific meanings or uses that identify with: 1) the mechanistic complexity of ecosystems, 2) the practical description of patterns in quantitative reductionist terms where correlations may be identified but nothing is understood about the causal relations without reference to the whole system, which leads to 3) a metaphysical hierarchy whereby the causal relations of larger systems are understood without reference to the smaller parts. An example of the metaphysical aspect to holism is the trend of increased exterior thickness in shells of different species. The reason for a thickness increase can be understood through reference to principals of natural selection via predation without any reference to the biomolecular properties of the exterior shells.[163] [edit] Metabolism and the early atmosphere Metabolism – the rate at which energy and material resources are taken up from the environment, transformed within an organism, and allocated to maintenance, growth and reproduction – is a fundamental physiological trait. Ernst et al.[164]:991 The Earth formed approximately 4.5 billion years ago[165] and environmental conditions were too extreme for life to form for the first 500 million years. During this early Hadean period, the Earth started to cool, allowing a crust and oceans to form. Environmental conditions were unsuitable for the origins of life for the first billion years after the Earth formed. The Earth's atmosphere transformed from being dominated by hydrogen, to one composed mostly of methane and ammonia. Over the next billion years the metabolic activity of life transformed the atmosphere to higher concentrations of carbon dioxide, nitrogen, and water vapor. These gases changed the way that light from the sun hit the Earth's surface and greenhouse effects trapped heat. There were untapped sources of free energy within the mixture of reducing and oxidizing gasses that set the stage for primitive ecosystems to evolve and, in turn, the atmosphere also evolved.[166]

The leaf is the primary site of photosynthesis in most plants. Throughout history, the Earth's atmosphere and biogeochemical cycles have been in a dynamic equilibrium with planetary ecosystems. The history is characterized by periods of significant transformation followed by millions of years of stability.[167] The evolution of the earliest organisms, likely anaerobic methanogen microbes, started the process by converting atmospheric hydrogen into methane (4H2 + CO2 → CH4 + 2H2O). Anoxygenic photosynthesis converting hydrogen sulfide into other sulfur compounds or water (2H2S + CO2 → hv → CH2O → H2O → + 2S or 2H2 + CO2 + hv → CH2O + H2O), as occurs in deep sea hydrothermal vents today, reduced hydrogen concentrations and increased atmospheric methane. Early forms of fermentation also increased levels of atmospheric methane. The transition to an oxygen dominant atmosphere (the Great Oxidation) did not begin until approximately 2.4-2.3 billion years ago, but photosynthetic processes started 0.3 to 1 billion years prior.[167][168] [edit] Radiation: heat, temperature and light The biology of life operates within a certain range of temperatures. Heat is a form of energy that regulates temperature. Heat affects growth rates, activity, behavior and

primary production. Temperature is largely dependent on the incidence of solar radiation. The latitudinal and longitudinal spatial variation of temperature greatly affects climates and consequently the distribution of biodiversity and levels of primary production in different ecosystems or biomes across the planet. Heat and temperature relate importantly to metabolic activity. Poikilotherms, for example, have a body temperature that is largely regulated and dependent on the temperature of the external environment. In contrast, homeotherms regulate their internal body temperature by expending metabolic energy.[2][86][157] There is a relationship between light, primary production, and ecological energy budgets. Sunlight is the primary input of energy into the planet's ecosystems. Light is composed of electromagnetic energy of different wavelengths. Radiant energy from the sun generates heat, provides photons of light measured as active energy in the chemical reactions of life, and also acts as a catalyst for genetic mutation.[2][86][157] Plants, algae, and some bacteria absorb light and assimilate the energy through photosynthesis. Organisms capable of assimilating energy by photosynthesis or through inorganic fixation of H2S are autotrophs. Autotrophs—responsible for primary production—assimilate light energy that becomes metabolically stored as potential energy in the form of biochemical enthalpic bonds.[2][86][157] [edit] Physical environments [edit] Water Main article: Aquatic ecosystem

shape and movement of tectonic plates as well as having an influence on geomorphic processes such as orogeny and erosion. These forces govern many of the geophysical properties and distributions of ecological biomes across the Earth. On a organism scale, gravitational forces provide directional cues for plant and fungal growth (gravitropism), orientation cues for animal migrations, and influence the biomechanics and size of animals.[2] Ecological traits, such as allocation of biomass in trees during growth are subject to mechanical failure as gravitational forces influence the position and structure of branches and leaves.[171] The cardiovascular systems of all animals are functionally adapted to overcome pressure and gravitational forces that change according to the features of organisms (e.g., height, size, shape), their behavior (e.g., diving, running, flying), and the habitat occupied (e.g., water, hot deserts, cold tundra).[172] [edit] Pressure Climatic and osmotic pressure places physiological constraints on organisms, such as flight and respiration at high altitudes, or diving to deep ocean depths. These constraints influence vertical limits of ecosystems in the biosphere as organisms are physiologically sensitive and adapted to atmospheric and osmotic water pressure differences.[2] Oxygen levels, for example, decrease with increasing pressure and are a limiting factor for life at higher altitudes.[173] Water transportation through trees is another important ecophysiological parameter dependent upon pressure.[174][175] Water pressure in the depths of oceans requires adaptations to deal with the different living conditions. Mammals, such as whales, dolphins and seals are adapted to deal with changes in sound due Environmental science

Wetland conditions such as shallow water, high plant productivity, and anaerobic substrates provide a suitable environment for important physical, biological, and chemical processes. Because of these processes, wetlands play a vital role in global nutrient and element cycles.:29[169] The rate of diffusion of carbon dioxide and oxygen is approximately 10,000 times slower in water than it is in air. When soils become flooded, they quickly lose oxygen from low-concentration (hypoxic) to an (anoxic) environment where anaerobic bacteria thrive among the roots. Water also influences the spectral properties of light that becomes more diffuse as it is reflected off the water surface and submerged particles.[169] Aquatic plants exhibit a wide variety of morphological and physiological adaptations that allow them to survive, compete and diversify these environments. For example, the roots and stems develop large cellular air spaces to allow for the efficient transportation gases (for example, CO2 and O2) used in respiration and photosynthesis. In drained soil, microorganisms use oxygen during respiration. In aquatic environments, anaerobic soil microorganisms use nitrate, manganic ions, ferric ions, sulfate, carbon dioxide and some organic compounds. The activity of soil microorganisms and the chemistry of the water reduces the oxidationreduction potentials of the water. Carbon dioxide, for example, is reduced to methane (CH4) by methanogenic bacteria. Salt water also requires special physiological adaptations to deal with water loss. Salt water plants (or halophytes) are able to osmo-regulate their internal salt (NaCl) concentrations or develop special organs for shedding salt away.[169] The physiology of fish is also specially adapted to deal with high levels of salt through osmoregulation. Their gills form electrochemical gradients that mediate salt excrusion in salt water and uptake in fresh water.[170] [edit] Gravity The shape and energy of the land is affected to a large degree by gravitational forces. On a larger scale, the distribution of gravitational forces on the earth are uneven and influence the

From Wikipedia, the free encyclopedia Jump to: navigation, search

Blue Marble composite images generated by NASA in 2001 (left) and 2002 (right). Environmental science is an interdisciplinary academic field that integrates physical and biological sciences, (including physics, chemistry, biology, soil science, geology, and geography) to the study of the environment, and the solution of environmental problems. Environmental science provides an integrated, quantitative, and interdisciplinary approach to the study of environmental systems.[1] Related areas of study include environmental studies and environmental engineering. Environmental studies incorporates more of the social sciences for understanding human relationships, perceptions and policies towards the environment. Environmental engineering focuses on design and technology for improving environmental quality. Environmental scientists work on subjects like the understanding of earth processes, evaluating alternative energy systems, pollution control and mitigation, natural resource management, and the effects of global climate change. Environmental issues almost always include an interaction of physical, chemical, and biological processes. Environmental scientists bring a systems approach to the

analysis of environmental problems. Key elements of an effective environmental scientist include the ability to relate space, and time relationships as well as quantitative analysis. Environmental science came alive as a substantive, active field of scientific investigation in the 1960s and 1970s driven by (a) the need for a multi-disciplinary approach to analyze complex environmental problems, (b) the arrival of substantive environmental laws requiring specific environmental protocols of investigation and (c) the growing public awareness of a need for action in addressing environmental problems. Events that spurred this development included the publication of Rachael Carson's landmark environmental book Silent Spring[2] along with major environmental issues becoming very public, such as the 1969 Santa Barbara oil spill, and the Cuyahoga River of Cleveland, Ohio, "catching fire" (also in 1969), and helped increase the visibility of environmental issues and create this new field of study. Environmental science is the study of interactions among the physical, chemical, and biological components of the environment. It provides an integrated, quantitative, and interdisciplinary approach to the study of environmental systems.[1] It includes such diverse areas as geology, agronomy, meteorology, atmospheric chemistry, soil chemistry, water chemistry, systems modeling, and biological responses of systems to anthropogenic influences. Environmental scientists monitor the quality of the environment (air, water, and soil), interpret the impact of human activities on terrestrial and aquatic ecosystems, and develop strategies for restoring ecosystems. In addition, environmental scientists help planners develop and construct buildings, transportation corridors, and utilities in ways that protect water resources and reflect efficient and beneficial land use.[2] Given the interdisciplinary nature of environmental science, teams of professionals commonly work together to conduct research or produce environmental impact statements, as required by governmental laws and regulations. In addition, various professional organizations engender work in environmental science and aid in interdisciplinary communications. [edit] Components Atmospheric sciences focuses on the Earth's atmosphere, with an emphasis upon its interrelation to other systems. Atmospheric sciences can include studies of meteorology, greenhouse gas phenomena, atmospheric dispersion modeling of airborne contaminants,[3][4] sound propagation phenomena related to noise pollution, and even light pollution. Taking the example of the global warming phenomena, physicists create computer models of atmospheric circulation and infra-red radiation transmission, chemists examine the inventory of atmospheric chemicals and their reactions, biologists analyze the plant and animal contributions to carbon dioxide fluxes, and specialists such as meteorologists and oceanographers add additional breadth in understanding the atmospheric dynamics. Ecology. An interdisciplinary analysis of an ecological system which is being impacted by one or more stressors might include several related environmental science fields. For example, one might examine an estuarine setting where a proposed industrial development could impact certain species by water and air pollution. For this study, biologists would describe the flora and fauna, chemists would analyze the transport of water pollutants to the marsh, physicists would calculate air pollution emissions and geologists would assist in understanding the marsh soils and bay muds.

Environmental chemistry is the study of chemical alterations in the environment. Principal areas of study include soil contamination and water pollution. The topics of analysis include chemical degradation in the environment, multi-phase transport of chemicals (for example, evaporation of a solvent containing lake to yield solvent as an air pollutant), and chemical effects upon biota. As an example study, consider the case of a leaking solvent tank which has entered the habitat soil of an endangered species of amphibian. As a method to resolve or understand the extent of soil contamination and subsurface transport of solvent, a computer model would be implemented. Chemists would then characterize the molecular bonding of the solvent to the specific soil type, and biologists would study the impacts upon soil arthropods, plants, and ultimately ponddwelling organisms that are the food of the endangered amphibian. Geosciences include environmental geology, environmental soil science, volcanic phenomena and evolution of the Earth's crust. In some classification systems this can also include hydrology, including oceanography. As an example study of soils erosion, calculations would be made of surface runoff by soil scientists. Fluvial geomorphologists would assist in examining sediment transport in overland flow. Physicists would contribute by assessing the changes in light transmission in the receiving waters. Biologists would analyze subsequent impacts to aquatic flora and fauna from increases in water turbidity. [edit] Regulations driving the studies

Environmental science examines the effects of humans on nature (Glen Canyon Dam in the U.S.) In the U.S. the National Environmental Policy Act (NEPA) of 1969 set forth requirements for analysis of major projects in terms of specific environmental criteria. Numerous state laws have echoed these mandates, applying the principles to localscale actions. The upshot has been an explosion of documentation and study of environmental consequences before the fact of development actions. One can examine the specifics of environmental science by reading examples of Environmental Impact Statements prepared under NEPA such as: Wastewater treatment expansion options discharging into the San Diego/Tijuana Estuary, Expansion of the San Francisco International Airport, Development of the Houston, Metro Transportation system, Expansion of the metropolitan Boston MBTA transit system, and Construction of Interstate 66 through Arlington, Virginia. In England and Wales the Environment Agency (EA),[5] formed in 1996, is a public body for protecting and improving the environment and enforces the regulations listed on the communities and local government site.[6] (formerly the office of the deputy prime minister). The agency was set up under the Environment Act 1995 as an independent body and works closely with UK Government to enforce the regulations. [edit] Terminology In common usage, "environmental science" and "ecology" are often used interchangeably, but technically, ecology refers only to the study of organisms and their interactions with each other and their environment. Ecology could be considered a subset of environmental science, which also could involve purely chemical or public health issues (for

example) ecologists would be unlikely to study. In practice, there is considerable overlap between the work of ecologists and other environmental scientists. The National Center for Education Statistics in the United States defines an academic program in environmental science as follows: A program that focuses on the application of biological, chemical, and physical principles to the study of the physical environment and the solution of environmental problems, including subjects such as abating or controlling environmental pollution and degradation; the interaction between human society and the natural environment; and natural resources management. Includes instruction in biology, chemistry, physics, geosciences, climatology, statistics, and mathematical Timeline of ecologists

Warming

1924

Geography[17]

Ellen Swallow 1842– Richards 1911

Pioneer and educator who linked urban ecology to human health[18]

Stephen Forbes

1844– 1930

Early founder of entomology and ecological concepts in 1887 [5][19][20]

Vito Volterra

1860– 1940

Independently pioneered mathematical populations models around the same time as Alfred J. Lotka.[21][22]

Vladimir Vernadsky A list of founders, innovators and their significant contributions to ecology, from Romanticism onward. Henry C. Cowles Major contribution & citation

1869– 1939

Founded the biosphere concept

Notable figure

Lifespa n

1869– 1939

Pioneering studies and conceptual development in studies of ecological succession[23]

Antoni van 1632– Leeuwenhoek 1723

First to develop concept of food chains

Jan Christian Smuts

1870– 1950

Coined the term holism in a 1926 book Holism and Evolution.[6]

1707– Carl Linnaeus 1778

Influential naturalist, inventor of science on the economy of nature[10][11]

Arthur G. Tansley

1871– 1955

First to coin the term ecosystem in 1936 and notable researcher[15][24][25]

Alexander Humboldt

1769– 1859

First to describe ecological gradient of latitudinal biodiversity increase toward the tropics [4] in 1807

Charles Christopher Adams

1873– 1955

Animal ecologist, biogeographer, author of first American book on animal ecology in 1913, founded ecological energetics[26][27]

Charles Darwin

1809– 1882

Founder of evolution by means of natural selection, founder of ecological studies of soils[12]

Friedrich Ratzel

1844– 1904

German geographer who first coined the term biogeography in 1891.

Herbert Spencer

1820– 1903

Early founder of social ecology, coined the phrase 'survival of the fittest'[11][13]

Frederic Clements

1874– 1945

Authored the first influential American ecology book in 1905[28]

Karl Möbius

1825– 1908

First to develop concept of ecological community, biocenosis, or living community[14][15][16]

Victor Ernest Shelford

1877– 1968

Founded physiological ecology, pioneered food-web and biome concepts, founded The Nature Conservancy[29][30]

Ernst Haeckel

1834– 1919

Invented the term ecology, popularized research links between ecology and evolution

Alfred J. Lotka

1880– 1949

First to pioneer mathematical populations models explaining trophic (predator-prey) interactions using logistic equation[31]

Victor Hensen

1835– 1924

Invented term plankton, developed quantitative and statistical measures of productivity in the seas

Henry Gleason

1882– 1975

Early ecology pioneer, quantitative theorist, author, and founder of the individualistic concept of ecology[28][32]

Eugenius

1841–

Early founder of Ecological Plant

Charles S. Elton

1900– 1991

'Father' of animal ecology, pioneered food-web & niche concepts and authored influential Animal Ecology text[29][33]

1. 2. 2.

Lentic, the ecosystem of a lake, pond or swamp. Lotic, the ecosystem of a river, stream or spring.

Artificial, ecosystems created by humans.

G. Evelyn Hutchinson

1903– 1991

Limnologist and conceptually advanced the niche concept[34][35][36]

Eugene P. Odum

1913– 2002

Co-founder of ecosystem ecology and ecological thermodynamic concepts[25]
[29][37][38]

Central to the ecosystem concept is the idea that living organisms interact with every other element in their local environment. Eugene Odum, a founder of ecology, stated: "Any unit that includes all of the organisms (ie: the "community") in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (i.e.: exchange of materials between living and nonliving parts) within the system is an ecosystem."[3] [edit] Etymology The term ecosystem was coined in 1930 by Roy Clapham to mean the combined physical and biological components of an environment. British ecologist Arthur Tansley later refined the term, describing it as "The whole system, … including not only the organism-complex, but also the whole complex of physical factors forming what we call the environment".[4] Tansley regarded ecosystems not simply as natural units, but as mental isolates.[4] Tansley later[5] defined the spatial extent of ecosystems using the term ecotope. [edit] Examples of ecosystems

Howard T. Odum

1924– 2002

Co-founder of ecosystem ecology and ecological thermodynamic concepts[25]
[29][37][38][39][40]

Robert MacArthur

1930– 1972

Co-founder on Theory of Island Biogeography and innovator of ecological statistical methods[41]

Environment may refer to:

• •

The natural environment, all living and non-living things that occur naturally on Earth Built environment, constructed surroundings that provide the setting for human activity, ranging from the large-scale civic surroundings to the personal places Environment (biophysical), the physical and biological factors along with their chemical interactions that affect an organism Environment (systems), the surroundings of a physical system that may interact with the system by exchanging mass, energy, or other properties

• •

An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving, physical components of the environment with which the organisms interact, such as air, soil, water, and sunlight.[1] It is all the organisms in a given area, along with the nonliving (abiotic) factors with which they interact; a biological community and its physical environment.[1] [edit] Overview The entire array of organisms inhabiting a particular ecosystem is called a community.[1] In a typical ecosystem, plants and other photosynthetic organisms are the producers that provide the food.[1] Ecosystems can be permanent or temporary. Ecosystems usually form a number of food webs.[2] Ecosystems are functional units consisting of living things in a given area, non-living chemical and physical factors of their environment, linked together through nutrient cycle and energy flow.[citation needed] 1. Natural

• • • • • • • • • • • • • • • • • • • • • • •

agro-ecosystems Agroecosystem Aquatic ecosystem Chaparral Coral reef Desert Forest Greater Yellowstone Ecosystem Human ecosystem Large marine ecosystem Littoral zone Lotic Marine ecosystem Pond Ecosystem Prairie Rainforest Riparian zone Savanna Steppe Subsurface Lithoautotrophic Microbial Ecosystem Taiga Tundra Urban ecosystem

A freshwater ecosystem in Gran Canaria, an island of the Canary Islands. [edit] Biomes

1. 2.

Terrestrial ecosystem Aquatic ecosystem Map of Terrestrial biomes classified by vegetation.

Main article: Biome Biomes are a classification of globally similar areas, including ecosystems, such as ecological communities of plants and animals, soil organisms and climatic conditions.[citation needed] Biomes are in part defined based on factors such as plant structures (such as trees, shrubs and grasses), leaf types (such as broadleaf and needleleaf), plant spacing (forest, woodland, savanna) and climate.[citation needed] Unlike ecozones, biomes are not defined by genetic, taxonomic or historical similarities. Biomes are often identified with particular patterns of ecological succession and climax vegetation. A fundamental classification of biomes is: 1. 2. 3. Terrestrial (land) biomes. Freshwater biomes. Marine biomes.

This perspective of niche allows for the existence of ecological equivalents and also empty niches. For example, the anolis lizards of the Greater Antilles are a rare example of convergent evolution, adaptive radiation, and the existence of ecological equivalents--the anolis lizards evolved in similar microhabitats independently of each other and resulted in the same ecomorphs across all four islands. [edit] Eltonian Niche In 1927 Charles Sutherland Elton, a British ecologist, gave the first working definition of the niche concept. He is credited with saying: "[W]hen an ecologist says 'there goes a badger,' he should include in his thoughts some definite idea of the animal's place in the community to which it belongs, just as if he had said, 'there goes the vicar.'"[4] The Eltonian niche encompasses the idea that the niche is the role a species plays in a community, rather than a habitat. [edit] Hutchinsonian Niche The Hutchinsonian Niche views niche as an n-dimensional hypervolume, where the dimensions are environmental conditions that define the range in which a species can persist. The niche concept was popularized by the zoologist G. Evelyn Hutchinson in 1957[5]. Hutchinson wanted to know why there are so many different types of organisms in any one habitat. The full range of environmental conditions (biological and physical) under which an organism can exist describes its fundamental niche. As a result of pressure from, and interactions with, other organisms (e.g. superior competitors), species are usually forced to occupy a niche that is narrower than this, and to which they are mostly highly adapted. This is termed the realized niche. The ecological niche has also been termed by G.E. Hutchinson a "hypervolume." This term defines the multi-dimensional space of resources (e.g., light, nutrients, structure, etc.) available to (and specifically used by) organisms. The term adaptive zone was coined by the paleontologist, George Gaylord Simpson, and refers to a set of ecological niches that may be occupied by a group of species that exploit the same resources in a similar manner. (Simpson, 1944; After Root, 1967.)[citation needed] Hutchinson's "niche" (a description of the ecological space occupied by a species) is subtly different from the "niche" as defined by Grinnell (an ecological role, that may or may not be actually filled by a species—see vacant niches). Different species cannot occupy the same niche (or guild). A niche is a very specific segment of ecospace occupied by a single species. Species can however share a 'mode of life' or 'autecological strategy' which are broader definitions of ecospace[6]. For example, Australian grasslands species, though different from those of the Great Plains grasslands, occupy the similar modes of life.[7] Once a niche is left vacant, other organisms can fill that position. For example, the niche that was left vacant by the extinction of the tarpan has been filled by other animals (in particular a small horse breed, the konik). Also, when plants and animals are introduced into a new environment, they have the potential to occupy or invade the niche or niches of native organisms, often outcompeting the indigenous species. Introduction of non-indigenous species to non-native habitats by humans often results in biological pollution by the exotic or invasive species.

Branches of Environmental Science Environmental science deals with many areas as is comprehensible as environmental factors work in determining the way life would work in synchronization with the natural factors. For example pollution is related to population, green house effects is related to global warming, pollution is again related to ozone depletion which also contributes to Global warming. There is no definite demarcation between the branches of study of Environmental Science, each overlaps on the scope of the other, yet to make things easier to grasp and to facilitate work upon them, the study of Environment Science has been divided into the following divisions:

• • • • • •

Environmental Chemistry Ecology Geosciences Atmospheric Science Environmental Assessment Environmental Microbiology

n ecology, a niche (pronounced /ˈniːʃ/ or /ˈnɪtʃ/)[1] is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food resources and foraging methods.[1] A shorthand definition of niche is how an organism makes a living. The ecological niche describes how an organism or population responds to the distribution of resources and competitors (e.g., by growing when resources are abundant, and when predators, parasites and pathogens are scarce) and how it in turn alters those same factors (e.g., limiting access to resources by other organisms, acting as a food source for predators and a consumer of prey).[2] [edit] Grinnelian Niche The word "niche" is derived from the Middle French word nicher, meaning to nest. The term was coined by the naturalist Joseph Grinnell in 1917, in his paper "The niche relationships of the California Thrasher."[3] The Grinnelian niche concept embodies the idea that the niche of a species is determined by the habitat in which is lives in. In other words, the niche is the sum of the habitat requirements that allow a species to persist and produce offspring. For example, the behavior of the California Thrasher is consistent with the chaparral habitat it lives in--it breeds and feeds in the underbrush; and escapes from predators by shuffling from underbrush to underbrush.

The mathematical representation of a species' fundamental niche in ecological space, and its subsequent projection back into geographic space, is the domain of niche modelling. Environmental chemistry is the scientific study of the chemical and biochemical phenomena that occur in natural places. It should not be confused with green chemistry, which seeks to reduce potential pollution at its source. It can be defined as the study of the sources, reactions, transport, effects, and fates of chemical species in the air, soil, and water environments; and the effect of human activity on these. Environmental chemistry is an interdisciplinary science that includes atmospheric, aquatic and soil chemistry, as well as heavily relying on analytical chemistry and being related to environmental and other areas of science. Environmental chemistry involves first understanding how the uncontaminated environment works, which chemicals in what concentrations are present naturally, and with what effects. Without this it would be impossible to accurately study the effects humans have on the environment through the release of chemicals. Environmental chemists draw on a range of concepts from chemistry and various environmental sciences to assist in their study of what is happening to a chemical species in the environment. Important general concepts from chemistry include understanding chemical reactions and equations, solutions, units, sampling, and analytical techniques.[1] Habitat From Wikipedia, the free encyclopedia Jump to: navigation, search For other uses of "Habitat", see Habitat (disambiguation).

common, affecting the identity of the participants and their degree of cohesiveness. In sociology, the concept of community has led to significant debate, and sociologists are yet to reach agreement on a definition of the term. There were ninety-four discrete definitions of the term by the mid-1950s.[1] Traditionally a "community" has been defined as a group of interacting people living in a common location. The word is often used to refer to a group that is organized around common values and is attributed with social cohesion within a shared geographical location, generally in social units larger than a household. The word can also refer to the national community or global community. The word "community" is derived from the Old French communité which is derived from the Latin communitas (cum, "with/together" + munus, "gift"), a broad term for fellowship or organized society.[2] Biomes are climatically and geographically defined as similar climatic conditions on the Earth, such as communities of plants, animals, and soil organisms,[1] and are often referred to as ecosystems.Some parts of the earth have more or less the same kind of abiotic and biotic factors spread over a large area creating a typical ecosystem over that area. Such major ecosystems are termed as biomes. Biomes are defined by factors such as plant structures (such as trees, shrubs, and grasses), leaf types (such as broadleaf and needleleaf), plant spacing (forest, woodland, savanna), and climate. Unlike ecozones, biomes are not defined by genetic, taxonomic, or historical similarities. Biomes are often identified with particular patterns of ecological succession and climax vegetation (quasi-equilibrium state of the local ecosystem). An ecosystem has many biotopes and a biome is a major habitat type. A major habitat type, however, is a compromise, as it has an intrinsic inhomogeneity. The biodiversity characteristic of each biome, especially the diversity of fauna and subdominant plant forms, is a function of abiotic factors and the biomass productivity of the dominant vegetation. In terrestrial biomes, species diversity tends to correlate positively with net primary productivity, moisture availability, and temperature.[2] Ecoregions are grouped into both biomes and ecozones. A fundamental classification of biomes is: 1. 2. Terrestrial (land) biomes Aquatic biomes (including Freshwater biomes and Marine biomes)

A habitat (which is Latin for "it inhabits") is an ecological or environmental area that is inhabited by a particular species of animal, plant or other type of organism.[1][2] It is the natural environment in which an organism lives, or the physical environment that surrounds (influences and is utilized by) a species population.[citation needed] [edit] Definition The term "population" is preferred to "organism" because, while it is possible to describe the habitat of a single black bear, it is also possible that we may not find any particular or individual bear but the grouping of bears that constitute a breeding population and occupy a certain biogeographical area. Further, this habitat could be somewhat different from the habitat of another group or population of black bears living elsewhere. Thus it is neither the species nor the individual for which the term habitat is typically used. [edit] Microhabitat The term microhabitat is often used to describe the smallscale physical requirements of a particular organism or population. A microhabitat is often a smaller habitat within a larger one. For example, a fallen log inside a forest can provide microhabitat for insects that are not found in the wider forest habitat outside such logs. Microenvironment is the immediate surroundings and other physical factors of an individual plant or animal within its pool. A microhabitait can be big or small depending on how much it varies. biological terms, a community is a group of interacting organisms sharing a populated environment. In human communities, intent, belief, resources, preferences, needs, risks, and a number of other conditions may be present and

Biomes are often known in English by local names. For example, a temperate grassland or shrubland biome is known commonly as steppe in central Asia, prairie in North America, and pampas in South America. Tropical grasslands are known as savanna in Australia, whereas in Southern Africa it is known as veldt (from Afrikaans). Sometimes an entire biome may be targeted for protection, especially under an individual nation's Biodiversity Action Plan. Climate is a major factor determining the distribution of terrestrial biomes. Among the important climatic factors are:

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latitude: Arctic, boreal, temperate, subtropical, tropical. humidity: humid, semi-humid, semi-arid, and arid.

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seasonal variation: Rainfall may be distributed evenly throughout the year or be marked by seasonal variations. dry summer, wet winter: Most regions of the earth receive most of their rainfall during the summer months; Mediterranean climate regions receive their rainfall during the winter months.

In 1804 , for example, he reported an impressive number of species, particularly plants, for which he sought to explain their geographic distribution with respect to Geological data. One of Humboldt's famous works was "Idea for a Plant Geography" ( 1805 ). Other important botanists of the time included Aimé Bonpland and Eugenius Warming . The notion of biocoenosis: Charles Darwin and Alfred Wallace Towards model. Alfred Russel Wallace , contemporary and competitor to Darwin, was first to propose a "geography" of animal species. Several authors recognized at the time that species were not independent of each other, and grouped them into plant species, animal species, and later into communities of living beings or Biocoenosis . This term was coined in 1877 by Karl Möbius . EARLY 20TH CENTURY ~ EXPANSION OF ECOLOGICAL THOUGHT The biosphere - Eduard Suess, Henry Chandler Cowles, and Vladimir Vernadsky By the 19th Century , ecology blossomed due to new discoveries in Chemistry by Lavoisier and De Saussure , notably the Nitrogen Cycle . After observing the fact that life developed only within strict limits of each compartment that makes up the Atmosphere , Hydrosphere , and Lithosphere , the Austrian geologist Eduard Suess proposed the term Biosphere in 1875 . Suess proposed the name biosphere for the conditions promoting life, such as those found on Earth , which includes Flora , Fauna , Minerals , Matter Cycles , et cetera. In the 1920s Vladimir I. Vernadsky , a Russian geologist who had defected to France, detailed the idea of the biosphere in his work "The biosphere" ( 1926 ), and described the fundamental principles of the Biogeochemical Cycle s. He thus redefined the biosphere as the sum of all Ecosystem s. First ecological damages were reported in the 18th Century , as the multiplication of colonies caused Deforestation . Since the 19th Century , with the Industrial Revolution , more and more pressing concerns have grown about the impact of human activity on The Environment . The term Ecologist has been in use since the end of the 19th Century . The ecosystem: Arthur Tansley Over the 19th Century , botanical geography and zoogeography combined to form the basis of Biogeography . This science, which deals with habitats of species, seeks to explain the reasons for the presence of certain species in a given location. It was in 1935 that Arthur Tansley , the British Ecologist , coined the term Ecosystem , the interactive system established between the

elevation: Increasing elevation causes a distribution of habitat types similar to that of increasing latitude.

The most widely used systems of classifying biomes correspond to latitude (or temperature zoning) and humidity. Biodiversity generally increases away from the poles towards the equator and increases with humidity. Earth science (also known as geoscience, the geosciences or the Earth sciences), is an all-embracing term for the sciences related to the planet Earth.[1] It is arguably a special case in planetary science, the Earth being the only known life-bearing planet. There are both reductionist and holistic approaches to Earth sciences. The formal discipline of Earth sciences may include the study of the atmosphere, oceans and biosphere, as well as the solid earth. Typically Earth scientists will use tools from physics, chemistry, biology, chronology and mathematics to build a quantitative understanding of how the Earth system works, and how it evolved to its current state. Ecology is generally spoken of as a new science, having only become prominent in the second half of the 20th Century. Nonetheless, ecological thinking at some level has been around for a long time, and the principles of ecology have developed gradually, closely intertwined with the development of other biological disciplines. Thus, one of the first ecologists may have been Aristotle or perhaps his student, Theophrastus , both of whom had interest in many species of animals. Theophrastus described interrelationships between animals and between animals and their environment as early as the 4th century BC (Ramalay, 1940). 18TH AND 19TH CENTURY ~ ECOLOGICAL MURMURS The botanical geography and Alexander von Humboldt Throughout the 18th and the beginning of the 19th Century , the great maritime powers such as Britain, Spain, and Portugal launched many world exploratory expeditions to develop Maritime Commerce with other countries, and to discover new natural resources, as well as to catalog them. At the beginning of the 18th Century , about twenty thousand plant species were known, versus forty thousand at the beginning of the 19th Century , and almost 400,000 today. These expeditions were joined by many scientists, including Botanists , such as the German explorer Alexander Von Humboldt . Humboldt is often considered a father of ecology. He was the first to take on the study of the relationship between organisms and their Environment . He exposed the existing relationships between observed plant species and Climate , and described vegetation zones using Latitude and Altitude , a discipline now known as Geobotany .

Biocoenosis (the group of living creatures), and their Biotope , the environment in which they live. Ecology thus became the science of ecosystems. Tansley's concept of the ecosystem was adopted by the energetic and influential biology educator Eugene Odum . Along with his brother, Howard Odum , Eugene P. Odum wrote a textbook which (starting in 1953) educated more than one generation of biologists and ecologists in North America. Ecological Succession - Henry Chandler Cowles At the turn of the 20th Century , Henry Chandler Cowles was one of the founders of the emerging study of "dynamic ecology", through his study of Ecological Succession at the Indiana Dunes , sand dunes at the southern end of Lake Michigan . Here Cowles found evidence of Ecological Succession in the Vegetation and the Soil with relation to age. Ecological Succession is the process by which a natural community moves from a simpler level of organisation to a more complex community (e.g., from bare sand, to grass growing on the sand, to grass growing on dirt produced from dead grass, to trees growing in the dirt produced by the grass). MODERN ECOLOGICAL THEORY AND RESEARCH

James Lovelock and the Gaia hypothesis The Gaia Theory , proposed by James Lovelock , in his work ''Gaia: A New Look at Life on Earth'', advanced the view that the Earth should be regarded as a single living macro-organism. In particular, it argued that the ensemble of living organisms has jointly evolved an ability to control the global environment — by influencing major physical parameters as the composition of the atmosphere, the evaporation rate, the chemistry of soils and oceans — so as to maintain conditions favorable to life. This vision was largely a sign of the times, in particular the growing perception after the , was "becoming sick from humans and their activities". Conservation and environmental movements Since the 19th century, Environmentalists and other Conservationists have used ecology and other sciences (e.g., Climatology ) to support their Advocacy Positions . Environmentalist views are often controversial for political or economic reasons. As a result, some scientific work in ecology directly influences policy and political debate; these in turn often direct ecological research. Ecology and global policy Ecology became a central part of the World's politics as early as 1971 , UNESCO launched a research program called '' Man And Biosphere '', with the objective of increasing knowledge about the mutual relationship between humans and nature. A few years later it defined the concept of Biosphere Reserve . In 1972 , the United Nations held the first international conference on the human environment in Stockholm , prepared by Rene Dubos and other experts. This conference was the origin of the phrase " Think Globally, Act Locally ". The next major events in ecology were the development of the concept of biosphere and the appearance of terms "biological diversity" -- or now more commonly Biodiversity -- in the 1980s . These terms were developed during the Earth Summit in Rio De Janeiro in 1992 , where the concept of the biosphere was recognized by the major international organizations, and risks associated with reductions in biodiversity were publicly acknowledged. Then, in 1997 , the dangers the biosphere was facing were recognized from an international point of view at the conference leading to the Kyoto Protocol . In particular, this conference highlighted the increasing dangers of the Greenhouse Effect -- related to the increasing concentration of Greenhouse Gas es in the atmosphere, leading to Global Changes In Climate . In Kyoto , most of the world's nations recognized the importance of looking at ecology from a global point of view, on a worldwide scale, and to take into account the impact of humans on the Earth's environment. Subcategories

ECOLOGY'S INFLUENCE IN THE SOCIAL SCIENCES AND HUMANITIES

Human ecology Human Ecology began in the 1920s , through the study of changes in Vegetation Succession in the city of Chicago . It became a distinct field of study in the 1970s . This marked the first recognition that humans, who had colonized all of the Earth's Continent s, were a major Ecological Factor . Humans greatly modify the environment through the development of the habitat (in particular Urban Planning ), by intensive exploitation activities such as Logging and Fishing , and as side effects of Agriculture , Mining , and Industry . Besides ecology and biology, this discipline involved many other natural and social sciences, such as Anthropology and Ethnology , Economics , Demography , Architecture and Urban Planning , Medicine and Psychology , and many more. The development of human ecology led to the increasing role of ecological science in the design and management of Cities . In recent years human ecology has been a topic that has interested organizational researchers. Hannan and Freeman (''Population Ecology of Organizations (1977)'', American Journal of Sociology) argue that organizations do not only adapt to an environment. Instead it is also the environment that selects or rejects populations of Organizations . In any given environment (in Equilibrium ) there will only be one form of organization ( Isomorphism ). Organizational Ecology has been a prominent theory in accounting for diversities of organizations and their changing composition over time.

As an example study of soils erosion, calculations would be made of surface runoff by soil scientists. Hydrologists would assist in examining sediment transport in overland flow. Physicists would contribute by assessing the changes in light transmission in the receiving waters. Biologists would analyze subsequent impacts to aquatic flora and fauna from increases in water turbidity. Environmental assessment is the process of appraisal through which environmental protection and sustainable development may be considered. Environmental assessments typically involve collection of field data, this can be from stakeholders and the ambient environment, and serves to harmonize the linkages between the different branches of the environment and development. Environmental microbiology is the study of the composition and physiology of microbial communities in the environment. The environment in this case means the soil, water, air and sediments covering the planet and can also include the animals and plants that inhabit these areas. Environmental microbiology also includes the study of microorganisms that exist in artificial environments such as bioreactors. Environmental Biology is a sub-category of environmental science that focuses specifically on the effects of environmental conditions on biological systems. Although it incorporates aspects of environmental science such as geochemistry and ecology, studies are focused on individual organisms, their biological processes, and their genetics. In addition, environmental biology incorporates the ideas of global change and conservation biology to encourage the conservation of biodiversity. Tulane University of New Orleans offers a B.S. in Environmental Biology distinct from the B.S. in Environmental Science.

Environmental Scientist sampling water. Atmospheric sciences examine the new phenomenology of the Earth's gaseous outer layer with emphasis upon interrelation to other systems. Atmospheric sciences comprises meteorological studies, greenhouse gas phenomena, atmospheric dispersion modeling of airborne contaminants, sound propagation phenomena related to noise pollution, and even light pollution. Taking the example of the global warming phenomena, physicists create computer models of atmospheric circulation and infra-red radiation transmission, chemists examine the inventory of atmospheric chemicals and their reactions, biologists analyze the plant and animal contributions to carbon dioxide fluxes, and specialists such as meteorologists and oceanographers add additional breadth in understanding the atmospheric dynamics. Ecology studies typically analyze the dynamics among an interrelated set of populations, or a population and some aspect of its environment. These studies could address endangered species, predator/prey interactions, habitat integrity, effects upon populations by environmental contaminants, or impact analysis of proposed land development upon species viability. An interdisciplinary analysis of an ecological system which is being impacted by one or more stressors might include several related environmental science fields. For example one might examine an estuarine setting where a proposed industrial development could impact certain species by water pollution and air pollution. For this study biologists would describe the flora and fauna, chemists would analyze the transport of water pollutants to the marsh, physicists would calculate air pollution emissions and geologists would assist in understanding the marsh soils and bay muds. Environmental chemistry is the study of chemical alterations in the environment. Principal areas of study include soil contamination and water pollution. The topics of analysis involve chemical degradation in the environment, multi-phase transport of chemicals (for example, evaporation of a solvent containing lake to yield solvent as an air pollutant), and chemical effects upon biota. As an example study, consider the case of a leaking solvent tank which has entered the soil upgradient of a habitat of an endangered species of amphibian. Physicists would develop a computer model to understand the extent of soil contamination and subsurface transport of solvent, chemists would analyze the molecular bonding of the solvent to the specific soil type and biologists would study the impacts upon soil arthropods, plants and ultimately pond dwelling copepods who are the food of the endangered amphibian. Geosciences include environmental geology, environmental soil science, hydrology, physical geography, climatology, and geomorphology. It may also embrace oceanography and other related fields.