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Ecology () is the study of the relationships between living
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s, including
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s, and their
physical environment A biophysical environment is a biotic and abiotic surrounding of an organism or population, and consequently includes the factors that have an influence in their survival, development, and evolution. A biophysical environment can vary in scale ...
. Ecology considers organisms at the individual,
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction usi ...
,
community A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, t ...
,
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
, and
biosphere The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also ...
level. Ecology overlaps with the closely related sciences of
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
,
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
,
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
,
ethology Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objecti ...
, and natural history. Ecology is a branch of
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
, and it is not synonymous with
environmentalism Environmentalism or environmental rights is a broad Philosophy of life, philosophy, ideology, and social movement regarding concerns for environmental protection and improvement of the health of the environment (biophysical), environment, par ...
. Among other things, ecology is the study of: * The abundance,
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...
, and distribution of organisms in the context of the environment * Life processes,
antifragility Antifragility is a property of systems in which they increase in capability to thrive as a result of stressors, shocks, volatility, noise, mistakes, faults, attacks, or failures. The concept was developed by Nassim Nicholas Taleb in his book, '' ...
, interactions, and
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
s * The movement of materials and
energy In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of ...
through living communities * The successional development of ecosystems * Cooperation, competition, and predation within and between
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
* Patterns of
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
and its effect on ecosystem processes Ecology has practical applications in
conservation biology Conservation biology is the study of the conservation of nature and of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction and the erosion of biotic interactions. It is an in ...
,
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
management,
natural resource management Natural resource management (NRM) is the management of natural resources such as land, water, soil, plants and animals, with a particular focus on how management affects the quality of life for both present and future generations (stewardship) ...
(
agroecology Agroecology (US: a-grō-ē-ˈkä-lə-jē) is an academic discipline that studies ecological processes applied to agricultural production systems. Bringing ecological principles to bear can suggest new management approaches in agroecosystems. Th ...
,
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
,
forestry Forestry is the science and craft of creating, managing, planting, using, conserving and repairing forests, woodlands, and associated resources for human and environmental benefits. Forestry is practiced in plantations and natural stands. ...
,
agroforestry Agroforestry is a land use management system in which trees or shrubs are grown around or among crops or pastureland. Trees produce a wide range of useful and marketable products from fruits/nuts, medicines, wood products, etc. This intentional ...
,
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, ...
,
mining Mining is the extraction of valuable minerals or other geological materials from the Earth, usually from an ore body, lode, vein, seam, reef, or placer deposit. The exploitation of these deposits for raw material is based on the econom ...
,
tourism Tourism is travel for pleasure or business; also the theory and practice of touring (disambiguation), touring, the business of attracting, accommodating, and entertaining tourists, and the business of operating tour (disambiguation), tours. Th ...
),
urban planning Urban planning, also known as town planning, city planning, regional planning, or rural planning, is a technical and political process that is focused on the development and design of land use and the built environment, including air, water, ...
(
urban ecology Urban ecology is the scientific study of the relation of living organisms with each other and their surroundings in an urban environment. An urban environment refers to environments dominated by high-density residential and commercial building ...
),
community health Community health refers to simple health services that are delivered by laymen outside hospitals and clinics. Community health is also the subset of public health that is taught to and practiced by clinicians. Community health volunteers and commu ...
,
economics Economics () is the social science that studies the production, distribution, and consumption of goods and services. Economics focuses on the behaviour and interactions of economic agents and how economies work. Microeconomics anal ...
,
basic BASIC (Beginners' All-purpose Symbolic Instruction Code) is a family of general-purpose, high-level programming languages designed for ease of use. The original version was created by John G. Kemeny and Thomas E. Kurtz at Dartmouth College ...
and
applied science Applied science is the use of the scientific method and knowledge obtained via conclusions from the method to attain practical goals. It includes a broad range of disciplines such as engineering and medicine. Applied science is often contrasted ...
, and human social interaction (
human ecology Human ecology is an interdisciplinary and transdisciplinary study of the relationship between humans and their natural, social, and built environments. The philosophy and study of human ecology has a diffuse history with advancements in ecolog ...
). The word ''ecology'' (german: Ökologie) was coined in 1866 by the German scientist
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
. The science of ecology as we know it today began with a group of American botanists in the 1890s.
Evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary concepts relating to adaptation and
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
are cornerstones of modern ecological theory. Ecosystems are dynamically interacting systems of organisms, the communities they make up, and the non-living ( abiotic) components of their environment. Ecosystem processes, such as
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...
,
nutrient cycling A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cycli ...
, and
niche construction Niche construction is the process by which an organism alters its own (or another species') local environment. These alterations can be a physical change to the organism’s environment or encompass when an organism actively moves from one habita ...
, regulate the flux of energy and matter through an environment. Ecosystems have biophysical feedback mechanisms that moderate processes acting on living ( biotic) and abiotic components of the planet. Ecosystems sustain life-supporting functions and provide
ecosystem service Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...
s like
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...
production (food, fuel, fiber, and medicine), the regulation of
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
, global
biogeochemical cycle A biogeochemical cycle (or more generally a cycle of matter) is the pathway by which a chemical substance cycles (is turned over or moves through) the biotic and the abiotic compartments of Earth. The biotic compartment is the biosphere and th ...
s,
water filtration A water filter removes impurities by lowering contamination of water using a fine physical barrier, a chemical process, or a biological process. Filters cleanse water to different extents, for purposes such as: providing agricultural irrigation ...
, soil formation,
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is d ...
control, flood protection, and many other natural features of scientific, historical, economic, or intrinsic value.


Levels, scope, and scale of organization

The scope of ecology contains a wide array of interacting levels of organization spanning micro-level (e.g.,
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
) to a planetary scale (e.g.,
biosphere The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also ...
)
phenomena A phenomenon ( : phenomena) is an observable event. The term came into its modern philosophical usage through Immanuel Kant, who contrasted it with the noumenon, which ''cannot'' be directly observed. Kant was heavily influenced by Gottfried ...
. Ecosystems, for example, contain abiotic
resources Resource refers to all the materials available in our environment which are technologically accessible, economically feasible and culturally sustainable and help us to satisfy our needs and wants. Resources can broadly be classified upon their av ...
and interacting life forms (i.e., individual organisms that aggregate into
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction usi ...
s which aggregate into distinct ecological communities). Ecosystems are dynamic, they do not always follow a linear successional path, but they are always changing, sometimes rapidly and sometimes so slowly that it can take thousands of years for ecological processes to bring about certain successional stages of a forest. An ecosystem's area can vary greatly, from tiny to vast. A single tree is of little consequence to the classification of a forest ecosystem, but is critically relevant to organisms living in and on it. Several generations of an
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
population can exist over the lifespan of a single leaf. Each of those aphids, in turn, supports diverse
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
l communities. The nature of connections in ecological communities cannot be explained by knowing the details of each species in isolation, because the emergent pattern is neither revealed nor predicted until the ecosystem is studied as an integrated whole. Some ecological principles, however, do exhibit collective properties where the sum of the components explain the properties of the whole, such as birth rates of a population being equal to the sum of individual births over a designated time frame. The main subdisciplines of ecology,
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction usi ...
(or
community A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, t ...
) ecology and
ecosystem ecology Ecosystem ecology is the integrated study of living ( biotic) and non-living (abiotic) components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components ...
, exhibit a difference not only in scale but also in two contrasting paradigms in the field. The former focuses on organisms' distribution and abundance, while the latter focuses on materials and energy fluxes.


Hierarchy

The scale of ecological dynamics can operate like a closed system, such as aphids migrating on a single tree, while at the same time remaining open with regard to broader scale influences, such as atmosphere or climate. Hence, ecologists classify
ecosystems An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
hierarchically by analyzing data collected from finer scale units, such as vegetation associations, climate, and
soil type A soil type is a taxonomic unit in soil science. All soils that share a certain set of well-defined properties form a distinctive soil type. Soil type is a technical term of soil classification, the science that deals with the systematic categori ...
s, and integrate this information to identify emergent patterns of uniform organization and processes that operate on local to regional,
landscape A landscape is the visible features of an area of land, its landforms, and how they integrate with natural or man-made features, often considered in terms of their aesthetic appeal.''New Oxford American Dictionary''. A landscape includes the ...
, and chronological scales. To structure the study of ecology into a conceptually manageable framework, the biological world is organized into a nested hierarchy, ranging in scale from
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s, to
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
, to tissues, to organs, to
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s, to
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, to
populations Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
, to
communities A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, to ...
, to
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s, to
biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
s, and up to the level of the
biosphere The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also ...
. This framework forms a
panarchy Panarchy may refer to: * Panarchy (Dartmouth), student society at Dartmouth College * Panarchy (ecology) A social-ecological system consists of 'a bio-geo-physical' unit and its associated social actors and institutions. Social-ecological system ...
and exhibits
non-linear In mathematics and science, a nonlinear system is a system in which the change of the output is not proportional to the change of the input. Nonlinear problems are of interest to engineers, biologists, physicists, mathematicians, and many other ...
behaviors; this means that "effect and cause are disproportionate, so that small changes to critical variables, such as the number of nitrogen fixers, can lead to disproportionate, perhaps irreversible, changes in the system properties."


Biodiversity

Biodiversity (an abbreviation of "biological diversity") describes the diversity of life from genes to ecosystems and spans every level of biological organization. The term has several interpretations, and there are many ways to index, measure, characterize, and represent its complex organization. Biodiversity includes species diversity, ecosystem diversity, and
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
and scientists are interested in the way that this diversity affects the complex ecological processes operating at and among these respective levels. Biodiversity plays an important role in
ecosystem service Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...
s which by definition maintain and improve human quality of life. Conservation priorities and management techniques require different approaches and considerations to address the full ecological scope of biodiversity.
Natural capital Natural capital is the world's stock of natural resources, which includes geology, soils, air, water and all living organisms. Some natural capital assets provide people with free goods and services, often called ecosystem services. All of t ...
that supports populations is critical for maintaining
ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...
and species
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
(e.g., riverine fish runs and avian insect control) has been implicated as one mechanism by which those service losses are experienced. An understanding of biodiversity has practical applications for species and ecosystem-level conservation planners as they make management recommendations to consulting firms, governments, and industry.


Habitat

The habitat of a species describes the environment over which a species is known to occur and the type of community that is formed as a result. 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." For example, a habitat might be an aquatic or terrestrial environment that can be further categorized as a
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial ...
or
alpine Alpine may refer to any mountainous region. It may also refer to: Places Europe * Alps, a European mountain range ** Alpine states, which overlap with the European range Australia * Alpine, New South Wales, a Northern Village * Alpine National P ...
ecosystem. Habitat shifts provide important evidence of competition in nature where one population changes relative to the habitats that most other individuals of the species occupy. For example, one population of a species of tropical lizard (''Tropidurus hispidus'') has a flattened body relative to the main populations that live in open savanna. The population that lives in an isolated rock outcrop hides in crevasses where its flattened body offers a selective advantage. Habitat shifts also occur in the developmental life history of amphibians, and in insects that transition from aquatic to terrestrial habitats.
Biotope A biotope is an area of uniform environmental conditions providing a living place for a specific assemblage of plants and animals. ''Biotope'' is almost synonymous with the term "habitat", which is more commonly used in English-speaking countr ...
and habitat are sometimes used interchangeably, but the former applies to a community's environment, whereas the latter applies to a species' environment.


Niche

Definitions of the niche date back to 1917, but G. Evelyn Hutchinson made conceptual advances in 1957 by introducing a widely adopted definition: "the set of biotic and abiotic conditions in which a species is able to persist and maintain stable population sizes." 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. The Hutchinsonian niche is defined more technically as a " Euclidean hyperspace whose ''dimensions'' are defined as environmental variables and whose ''size'' is a function of the number of values that the environmental values may assume for which an organism has ''positive fitness''."
Biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
patterns and range distributions are explained or predicted through knowledge of a species' traits and niche requirements. Species have functional traits that are uniquely adapted to the ecological niche. A trait is a measurable property,
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (biology), morphology or physical form and structure, its Developmental biology, developmental proc ...
, or characteristic of an organism that may influence its survival. Genes play an important role in the interplay of development and environmental expression of traits. Resident species evolve traits that are fitted to the selection pressures of their local environment. This tends to afford them a competitive advantage and discourages similarly adapted species from having an overlapping geographic range. The
competitive exclusion principle In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the sligh ...
states that two species cannot coexist indefinitely by living off the same limiting
resource Resource refers to all the materials available in our environment which are technologically accessible, economically feasible and culturally sustainable and help us to satisfy our needs and wants. Resources can broadly be classified upon thei ...
; one will always out-compete the other. When similarly adapted species overlap geographically, closer inspection reveals subtle ecological differences in their habitat or dietary requirements. Some models and empirical studies, however, suggest that disturbances can stabilize the co-evolution and shared niche occupancy of similar species inhabiting species-rich communities. The habitat plus the niche is called the
ecotope Ecotopes are the smallest ecologically distinct landscape features in a landscape mapping and classification system. As such, they represent relatively homogeneous, spatially explicit landscape functional units that are useful for stratifying land ...
, which is defined as the full range of environmental and biological variables affecting an entire species.


Niche construction

Organisms are subject to environmental pressures, but they also modify their habitats. The
regulatory feedback There are many types of artificial neural networks (ANN). Artificial neural networks are computational models inspired by biological neural networks, and are used to approximate functions that are generally unknown. Particularly, they are inspired ...
between organisms and their environment can affect conditions from local (e.g., a
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers a ...
pond A pond is an area filled with water, either natural or artificial, that is smaller than a lake. Defining them to be less than in area, less than deep, and with less than 30% emergent vegetation helps in distinguishing their ecology from tha ...
) to global scales, over time and even after death, such as decaying logs or
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
skeleton deposits from marine organisms. The process and concept of ecosystem engineering are related to
niche construction Niche construction is the process by which an organism alters its own (or another species') local environment. These alterations can be a physical change to the organism’s environment or encompass when an organism actively moves from one habita ...
, but the former relates only to the physical modifications of the habitat whereas the latter also considers the evolutionary implications of physical changes to the environment and the feedback this causes on the process of natural selection. 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." The ecosystem engineering concept has stimulated a new appreciation for the influence that organisms have on the ecosystem and evolutionary process. The term "niche construction" is more often used in reference to the under-appreciated feedback mechanisms of natural selection imparting forces on the abiotic niche. An example of natural selection through ecosystem engineering occurs in the nests of
social insects Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
, including ants, bees, wasps, and termites. There is an emergent
homeostasis In biology, homeostasis ( British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...
or homeorhesis 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 is subject to the forces of natural selection. Moreover, a nest can survive over successive generations, so that progeny inherit both genetic material and a legacy niche that was constructed before their time.


Biome

Biomes are larger units of organization that categorize regions of the Earth's ecosystems, mainly according to the structure and composition of vegetation. There are different methods to define the 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. Biomes include
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...
,
temperate broadleaf and mixed forest Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These ...
, temperate deciduous forest,
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, sp ...
,
tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mou ...
,
hot desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one ...
, and
polar desert Polar deserts are the regions of Earth that fall under an ice cap climate (''EF'' under the Köppen classification). Despite rainfall totals low enough to normally classify as a desert, polar deserts are distinguished from true deserts (' or ' un ...
. Other researchers have recently categorized other biomes, such as the human and oceanic
microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably wel ...
s. To a
microbe A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
, the human body is a habitat and a landscape. Microbiomes were discovered largely through advances in
molecular genetics Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
, which 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.


Biosphere

The largest scale of ecological organization is the biosphere: the total sum of ecosystems on the planet. Ecological relationships regulate the flux of energy, nutrients, and climate all the way up to the planetary scale. For example, the dynamic history of the planetary atmosphere's CO2 and O2 composition has been affected by the biogenic flux of gases coming from respiration and photosynthesis, with levels fluctuating over time in relation to the ecology and evolution of plants and animals. Ecological theory has also been used to explain self-emergent regulatory phenomena at the planetary scale: for example, the
Gaia hypothesis The Gaia hypothesis (), also known as the Gaia theory, Gaia paradigm, or the Gaia principle, proposes that living organisms interact with their inorganic surroundings on Earth to form a synergistic and self-regulating, complex system that help ...
is an example of
holism Holism () is the idea that various systems (e.g. physical, biological, social) should be viewed as wholes, not merely as a collection of parts. The term "holism" was coined by Jan Smuts in his 1926 book '' Holism and Evolution''."holism, n." OED O ...
applied in ecological theory. The Gaia hypothesis states that there is an emergent
feedback loop Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause-and-effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handled c ...
generated by the
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...
of living organisms that maintains the core temperature of the Earth and atmospheric conditions within a narrow self-regulating range of tolerance.


Population ecology

Population ecology studies the dynamics of species populations and how these populations interact with the wider environment. A population consists of individuals of the same species that live, interact, and migrate through the same niche and habitat. A primary law of population ecology is the Malthusian growth model which states, "a population will grow (or decline) exponentially as long as the environment experienced by all individuals in the population remains constant." Simplified population
models A model is an informative representation of an object, person or system. The term originally denoted the plans of a building in late 16th-century English, and derived via French and Italian ultimately from Latin ''modulus'', a measure. Models c ...
usually starts with four variables: death, birth,
immigration Immigration is the international movement of people to a destination country of which they are not natives or where they do not possess citizenship in order to settle as permanent residents or naturalized citizens. Commuters, tourists, ...
, and
emigration Emigration is the act of leaving a resident country or place of residence with the intent to settle elsewhere (to permanently leave a country). Conversely, immigration describes the movement of people into one country from another (to permanent ...
. An example of an introductory population model describes a closed population, such as on an island, where immigration and emigration does not take place. Hypotheses are evaluated with reference to a null hypothesis which states that
random In common usage, randomness is the apparent or actual lack of pattern or predictability in events. A random sequence of events, symbols or steps often has no order and does not follow an intelligible pattern or combination. Individual ran ...
processes create the observed data. In these island models, the rate of population change is described by: : \frac = bN(t) - dN(t) = (b - d)N(t) = rN(t), where ''N'' is the total number of individuals in the population, ''b'' and ''d'' are the per capita rates of birth and death respectively, and ''r'' is the per capita rate of population change. Using these modeling techniques, Malthus' population principle of growth was later transformed into a model known as the
logistic equation A logistic function or logistic curve is a common S-shaped curve (sigmoid curve) with equation f(x) = \frac, where For values of x in the domain of real numbers from -\infty to +\infty, the S-curve shown on the right is obtained, with the ...
by
Pierre Verhulst Pierre is a masculine given name. It is a French form of the name Peter. Pierre originally meant "rock" or "stone" in French (derived from the Greek word πέτρος (''petros'') meaning "stone, rock", via Latin "petra"). It is a translatio ...
: : \frac = rN(t) - \alpha N(t)^2 = rN(t)\left(\frac\right), where ''N(t)'' is the number of individuals measured as
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...
density as a function of time, ''t'', ''r'' is the maximum per-capita rate of change commonly known as the intrinsic rate of growth, and \alpha is the crowding coefficient, which represents the reduction in population growth rate per individual added. The formula states that the rate of change in population size (\mathrmN(t)/\mathrmt) will grow to approach equilibrium, where (\mathrmN(t)/\mathrmt = 0), when the rates of increase and crowding are balanced, r/\alpha. A common, analogous model fixes the equilibrium, r/\alpha as ''K'', which is known as the "carrying capacity." Population ecology builds upon these introductory models to further understand demographic processes in real study populations. Commonly used types of data include life history,
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
, and survivorship, and these are analyzed using mathematical techniques such as
matrix algebra In abstract algebra, a matrix ring is a set of matrices with entries in a ring ''R'' that form a ring under matrix addition and matrix multiplication . The set of all matrices with entries in ''R'' is a matrix ring denoted M''n''(''R'')Lang, '' ...
. The information is used for managing wildlife stocks and setting harvest quotas. In cases where basic models are insufficient, ecologists may adopt different kinds of statistical methods, such as the Akaike information criterion, or use models that can become mathematically complex as "several competing hypotheses are simultaneously confronted with the data."


Metapopulations and migration

The concept of metapopulations was defined in 1969 as "a population of populations which go extinct locally and recolonize". Metapopulation ecology is another statistical approach that is often used in conservation research. Metapopulation models simplify the landscape into patches of varying levels of quality, and metapopulations are linked by the migratory behaviours of organisms. Animal migration is set apart from other kinds of movement because it involves the seasonal departure and return of individuals from a habitat. Migration is also a population-level phenomenon, as with the migration routes followed by plants as they occupied northern post-glacial environments. Plant ecologists use pollen records that accumulate and stratify in wetlands to reconstruct the timing of plant migration and dispersal relative to historic and contemporary climates. These migration routes involved an expansion of the range as plant populations expanded from one area to another. There is a larger taxonomy of movement, such as commuting, foraging, territorial behavior, stasis, and ranging. Dispersal is usually distinguished from migration because it involves the one-way permanent movement of individuals from their birth population into another population. In metapopulation terminology, migrating individuals are classed as emigrants (when they leave a region) or immigrants (when they enter a region), 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; the population at the site will disappear unless rescued by an adjacent source patch or environmental conditions become more favorable. Metapopulation models examine patch dynamics over time to answer potential 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 are sources when conditions are more favorable. Ecologists use a mixture of computer models and field studies to explain metapopulation structure.


Community ecology

Community ecology is the study of the interactions among a collection of species that inhabit the same geographic area. Community ecologists study the determinants of patterns and processes for two or more interacting species. Research in community ecology might measure species diversity in grasslands in relation to soil fertility. It might also include the analysis of predator-prey dynamics, competition among similar plant species, or mutualistic interactions between crabs and corals.


Ecosystem ecology

Ecosystems may be habitats within biomes that form an integrated whole and a dynamically responsive system having both physical and biological complexes. Ecosystem ecology is the science of determining the fluxes of materials (e.g. carbon, phosphorus) between different pools (e.g., tree biomass, soil organic material). Ecosystem ecologists attempt to determine the underlying causes of these fluxes. Research in ecosystem ecology might measure
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...
(g C/m^2) in a
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
in relation to decomposition and consumption rates (g C/m^2/y). This requires an understanding of the community connections between plants (i.e., primary producers) and the decomposers (e.g.,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
and bacteria), The underlying concept of an ecosystem can be traced back to 1864 in the published work of
George Perkins Marsh George Perkins Marsh (March 15, 1801July 23, 1882), an American diplomat and philologist, is considered by some to be America's first environmentalist and by recognizing the irreversible impact of man's actions on the earth, a precursor to the ...
("Man and Nature"). Within an ecosystem, organisms are linked to the physical and biological components of their environment to which they are adapted. Ecosystems are complex adaptive systems where the interaction of life processes form self-organizing patterns across different scales of time and space. Ecosystems are broadly categorized as terrestrial,
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
, atmospheric, or
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military ...
. Differences stem from the nature of the unique physical environments that shapes the biodiversity within each. A more recent addition to ecosystem ecology are technoecosystems, which are affected by or primarily the result of human activity.


Food webs

A food web is the archetypal ecological network. Plants capture
solar energy Solar energy is radiant light and heat from the Sun that is harnessed using a range of technologies such as solar power to generate electricity, solar thermal energy (including solar water heating), and solar architecture. It is an essen ...
and use it to synthesize simple sugars during
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
. As plants grow, they accumulate nutrients and are eaten by grazing
herbivores A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, and the energy is transferred through a chain of organisms by consumption. The simplified linear feeding pathways that move from a basal trophic species to a top consumer is called the
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
. The larger interlocking pattern of food chains in an ecological community creates a complex food web. Food webs are a type of
concept map A concept map or conceptual diagram is a diagram that depicts suggested relationships between concepts. Concept maps may be used by instructional designers, engineers, technical writers, and others to organize and structure knowledge. A conc ...
or a
heuristic A heuristic (; ), or heuristic technique, is any approach to problem solving or self-discovery that employs a practical method that is not guaranteed to be optimal, perfect, or rational, but is nevertheless sufficient for reaching an immediate ...
device that is used to illustrate and study pathways of energy and material flows. Food webs are often limited relative to the real world. Complete empirical measurements are generally restricted to a specific habitat, such as a cave or a pond, and principles gleaned from food web
microcosm Microcosm or macrocosm, also spelled mikrokosmos or makrokosmos, may refer to: Philosophy * Microcosm–macrocosm analogy, the view according to which there is a structural similarity between the human being and the cosmos Music * Macrocosm (al ...
studies are extrapolated to larger systems. Feeding relations require extensive investigations into the gut contents of organisms, which can be difficult to decipher, or stable isotopes can be used to trace the flow of nutrient diets and energy through a food web. Despite these limitations, food webs remain a valuable tool in understanding community ecosystems. Food webs exhibit principles of ecological emergence through the nature of trophic relationships: some species have many weak feeding links (e.g.,
omnivores An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
) while some are more specialized with fewer stronger feeding links (e.g., primary predators). Theoretical and empirical studies identify
non-random In common usage, randomness is the apparent or actual lack of pattern or predictability in events. A random sequence of events, symbols or steps often has no order and does not follow an intelligible pattern or combination. Individual rando ...
emergent patterns of few strong and many weak linkages that explain how ecological communities remain stable over time. Food webs are composed of subgroups where members in a community are linked by strong interactions, and the weak interactions occur between these subgroups. This increases food web stability. Step by step lines or relations are drawn until a web of life is illustrated.


Trophic levels

A trophic level (from Greek ''troph'', τροφή, trophē, meaning "food" or "feeding") is "a group of organisms acquiring a considerable majority of its energy from the lower adjacent level (according to
ecological pyramid An ecological pyramid (also trophic pyramid, Eltonian pyramid, energy pyramid, or sometimes food pyramid) is a graphical representation designed to show the biomass or bioproductivity at each trophic level in a given ecosystem. A ''pyramid of ...
s) nearer the abiotic source." Links in food webs primarily connect feeding relations or trophism among species. Biodiversity within ecosystems can be organized into trophic pyramids, in which the vertical dimension represents feeding relations that become further removed from the base of the food chain up toward top predators, and the horizontal dimension represents the abundance or biomass at each level. When the relative abundance or biomass of each species is sorted into its respective trophic level, they naturally sort into a 'pyramid of numbers'. Species are broadly categorized as autotrophs (or primary producers), heterotrophs (or
consumer A consumer is a person or a group who intends to order, or uses purchased goods, products, or services primarily for personal, social, family, household and similar needs, who is not directly related to entrepreneurial or business activities. ...
s), and
Detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...
s (or decomposers). Autotrophs are organisms that produce their own food (production is greater than respiration) by photosynthesis or chemosynthesis. Heterotrophs are organisms that must feed on others for nourishment and energy (respiration exceeds production). Heterotrophs can be further sub-divided into different functional groups, including
primary consumers A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
(strict herbivores), secondary consumers (
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...
predators that feed exclusively on herbivores), and tertiary consumers (predators that feed on a mix of herbivores and predators). 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 because compared to herbivores, they are relatively inefficient at grazing. Trophic levels are part of the
holistic Holism () is the idea that various systems (e.g. physical, biological, social) should be viewed as wholes, not merely as a collection of parts. The term "holism" was coined by Jan Smuts in his 1926 book '' Holism and Evolution''."holism, n." OED On ...
or
complex systems A complex system is a system composed of many components which may interact with each other. Examples of complex systems are Earth's global climate, organisms, the human brain, infrastructure such as power grid, transportation or communication sy ...
view of ecosystems. Each trophic level contains unrelated species that are grouped together because they share common ecological functions, giving a macroscopic view of the system. While the notion of trophic levels provides insight into energy flow and top-down control within food webs, it is troubled by the prevalence of omnivory in real ecosystems. This has led some ecologists to "reiterate that the notion that species clearly aggregate into discrete, homogeneous trophic levels is fiction." Nonetheless, recent studies have shown that real trophic levels do exist, but "above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores."


Keystone species

A keystone species is a species that is connected to a disproportionately large number of other species in the
food-web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one o ...
. 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 (termed ''trophic cascades'') that alters trophic dynamics, other food web connections, and can cause the extinction of other species. The term keystone species was coined by Robert Paine in 1969 and is a reference to the keystone architectural feature as the removal of a keystone species can result in a community collapse just as the removal of the keystone in an arch can result in the arch's loss of stability.
Sea otter The sea otter (''Enhydra lutris'') is a marine mammal native to the coasts of the northern and eastern North Pacific Ocean. Adult sea otters typically weigh between , making them the heaviest members of the weasel family, but among the smal ...
s (''Enhydra lutris'') are commonly cited as an example of a keystone species because they limit the density of
sea urchins Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) of ...
that feed on
kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "underwa ...
. 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. Hunting of sea otters, for example, is thought to have led indirectly to the extinction of the
Steller's sea cow Steller's sea cow (''Hydrodamalis gigas'') is an extinct sirenian described by Georg Wilhelm Steller in 1741. At that time, it was found only around the Commander Islands in the Bering Sea between Alaska and Russia; its range extended across ...
(''Hydrodamalis gigas''). While the keystone species concept has been used extensively as a
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and manageme ...
tool, it has been criticized for being poorly defined from an operational stance. It is difficult to experimentally determine what species may hold a keystone role in each ecosystem. Furthermore, food web theory suggests that keystone species may not be common, so it is unclear how generally the keystone species model can be applied.


Complexity

Complexity is understood as a large computational effort needed to piece together numerous interacting parts exceeding the iterative memory capacity of the human mind. Global patterns of biological diversity are complex. This
biocomplexity Biocomplexity is the study of complex structures and behaviors that arise from nonlinear interactions of active biological agents, which may range in scale from molecules to cells to organisms. Almost every biological system exhibits complexity ...
stems from the interplay among ecological processes that operate and influence patterns at different scales that grade into each other, such as transitional areas or
ecotones An ecotone is a transition area between two biological communities, where two communities meet and integrate. It may be narrow or wide, and it may be local (the zone between a field and forest) or regional (the transition between forest and gras ...
spanning landscapes. Complexity stems from the interplay among levels of biological organization as energy, and matter is integrated into larger units that superimpose onto the smaller parts. "What were wholes on one level become parts on a higher one." Small scale patterns do not necessarily explain large scale phenomena, otherwise captured in the expression (coined by Aristotle) 'the sum is greater than the parts'. "Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral, and geometric." From these principles, 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 explanations at each integrative level. Ecological complexity relates to the dynamic resilience of ecosystems that transition to multiple shifting steady-states directed by random fluctuations of history. Long-term ecological studies provide important track records to better understand the complexity and resilience of ecosystems over longer temporal and broader spatial scales. These studies are managed by the International Long Term Ecological Network (LTER). The longest experiment in existence is the Park Grass Experiment, which was initiated in 1856. Another example is the Hubbard Brook study, which has been in operation since 1960.


Holism

Holism remains a critical part of the theoretical foundation in contemporary ecological studies. Holism addresses the biological organization of life that self-organizes into layers of emergent whole systems that function according to non-reducible properties. This means that higher-order patterns of a whole functional system, such as an
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
, cannot be predicted or understood by a simple summation of the parts. "New properties emerge because the components interact, not because the basic nature of the components is changed." Ecological studies are necessarily holistic as opposed to
reductionistic Reductionism is any of several related philosophical ideas regarding the associations between phenomena which can be described in terms of other simpler or more fundamental phenomena. It is also described as an intellectual and philosophical pos ...
. Holism has three scientific meanings or uses that identify with ecology: 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 Metaphysics is the branch of philosophy that studies the fundamental nature of reality, the first principles of being, identity and change, space and time, causality, necessity, and possibility. It includes questions about the nature of conscio ...
hierarchy whereby the causal relations of larger systems are understood without reference to the smaller parts. Scientific holism differs from
mysticism Mysticism is popularly known as becoming one with God or the Absolute, but may refer to any kind of ecstasy or altered state of consciousness which is given a religious or spiritual meaning. It may also refer to the attainment of insight in ...
that has appropriated the same term. An example of metaphysical holism is identified in the trend of increased exterior thickness in shells of different species. The reason for a thickness increase can be understood through reference to principles of natural selection via predation without the need to reference or understand the biomolecular properties of the exterior shells.


Relation to evolution

Ecology and evolutionary biology are considered sister disciplines of the life sciences.
Natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
, life history, development,
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
,
populations Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
, and
inheritance Inheritance is the practice of receiving private property, titles, debts, entitlements, privileges, rights, and obligations upon the death of an individual. The rules of inheritance differ among societies and have changed over time. Of ...
are examples of concepts that thread equally into ecological and evolutionary theory. Morphological, behavioural, and 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 principles, such as
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
or the Linnaean system of taxonomy. The two disciplines often appear together, such as in the title of the journal ''
Trends in Ecology and Evolution A fad or trend is any form of collective behavior that develops within a culture, a generation or social group in which a group of people enthusiastically follow an impulse for a short period. Fads are objects or behaviors that achieve short- ...
''. 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 processes operating across different spatial or temporal scales of organization. While the boundary between ecology and evolution is not always clear, ecologists study the abiotic and biotic factors that influence evolutionary processes, and evolution can be rapid, occurring on ecological timescales as short as one generation.


Behavioural ecology

All organisms can exhibit behaviours. Even plants express complex behaviour, including memory and communication. Behavioural ecology is the study of an organism's behaviour in its environment and its ecological and evolutionary implications. Ethology is the study of observable movement or behaviour in animals. This could include investigations of motile
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
of plants, mobile
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. ...
,
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
swimming toward the female egg, the cultivation of fungi by weevils, the mating dance of a
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
, or social gatherings of
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
. Adaptation is the central unifying concept in behavioural ecology. Behaviours can be recorded as traits and inherited in much the same way that eye and hair colour can. Behaviours can evolve by means of natural selection as adaptive traits conferring functional utilities that increases reproductive fitness. Predator-prey interactions are an introductory concept into food-web studies as well as behavioural ecology. 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 principles of conservation, optimization, or efficiency. For example, " e threat-sensitive predator avoidance hypothesis predicts that prey should assess the degree of threat posed by different predators and match their behaviour according to current levels of risk" or " e optimal
flight initiation distance The flight zone of an animal is the area surrounding an animal that if encroached upon by a potential predator or threat, including humans, will cause alarm and escape behavior. The flight zone is determined by the animal's flight distance, someti ...
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." Elaborate sexual displays and posturing are encountered in the behavioural ecology of animals. The birds-of-paradise, for example, sing and display elaborate ornaments during
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
. These displays serve a dual purpose of signalling healthy or well-adapted individuals and desirable genes. The displays are driven by
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
as an advertisement of quality of traits among suitors.


Cognitive ecology

Cognitive ecology integrates theory and observations from
evolutionary ecology Evolutionary ecology lies at the intersection of ecology and evolutionary biology. It approaches the study of ecology in a way that explicitly considers the evolutionary histories of species and the interactions between them. Conversely, it can ...
and
neurobiology Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, developme ...
, primarily cognitive science, in order to understand the effect that animal interaction with their habitat has on their cognitive systems and how those systems restrict behavior within an ecological and evolutionary framework. "Until recently, however, cognitive scientists have not paid sufficient attention to the fundamental fact that cognitive traits evolved under particular natural settings. With consideration of the selection pressure on cognition, cognitive ecology can contribute intellectual coherence to the multidisciplinary study of cognition." As a study involving the 'coupling' or interactions between organism and environment, cognitive ecology is closely related to enactivism, a field based upon the view that "...we must see the organism and environment as bound together in reciprocal specification and selection...".


Social ecology

Social-ecological behaviours are notable in the
social insects Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
, slime moulds,
social spider A social spider is a spider species whose individuals form relatively long-lasting aggregations. Whereas most spiders are solitary and even aggressive toward other members of their own species, some hundreds of species in several families show a ...
s, human society, and
naked mole-rat The naked mole-rat (''Heterocephalus glaber''), also known as the sand puppy, is a burrowing rodent native to the Horn of Africa and parts of Kenya, notably in Somali regions. It is closely related to the blesmols and is the only species in th ...
s where eusocialism has evolved. Social behaviours include reciprocally beneficial behaviours among kin and nest mates and evolve from kin and group selection.
Kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution ...
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,
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s, and
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s 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. In contrast,
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behavi ...
ists 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 with predominantly altruistic members survive better than groups with predominantly selfish members.


Coevolution

Ecological interactions can be classified broadly into a host and an associate relationship. A host is any entity that harbours another that is called the associate. Relationships between species that are mutually or reciprocally beneficial are called
mutualisms Mutualism describes the ecological interaction between two or more species where each species has a net benefit. Mutualism is a common type of ecological interaction. Prominent examples include most vascular plants engaged in mutualistic intera ...
. Examples of mutualism include
fungus-growing ants Fungus-growing ants (tribe Attini) comprise all the known fungus-growing ant species participating in ant–fungus mutualism. They are known for cutting grasses and leaves, carrying them to their colonies' nests, and using them to grow fungus o ...
employing agricultural symbiosis, bacteria living in the guts of insects and other organisms, the
fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while the ...
and yucca moth pollination complex,
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
, and
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
s with photosynthetic algae. If there is a physical connection between host and associate, the relationship is called
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
. Approximately 60% of all plants, for example, have a symbiotic relationship with
arbuscular mycorrhizal fungi An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscule ...
living in their roots forming an exchange network of carbohydrates for mineral nutrients. Indirect mutualisms occur where the organisms live apart. For example, 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 Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fr ...
because many others receive the benefits of clean air at no cost or harm to trees supplying the oxygen. If the associate benefits while the host suffers, the relationship is called
parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
. Although parasites impose a cost to their host (e.g., via damage to their reproductive organs or propagules, denying the services of a beneficial partner), their net effect on host fitness is not necessarily negative and, thus, becomes difficult to forecast. Co-evolution is also driven by competition among species or among members of the same species under the banner of reciprocal antagonism, such as grasses competing for growth space. The Red Queen Hypothesis, for example, posits that parasites track down and specialize on the locally common genetic defense systems of its host that drives the evolution of sexual reproduction to diversify the genetic constituency of populations responding to the antagonistic pressure.


Biogeography

Biogeography (an amalgamation of ''biology'' and ''geography'') is the comparative study of the geographic distribution of organisms and the corresponding evolution of their traits in space and time. The ''
Journal of Biogeography The ''Journal of Biogeography'' is a peer-reviewed scientific journal in biogeography that was established in 1974. It covers aspects of spatial, ecological, and historical biogeography. The founding editor-in-chief was David Watts, followed by ...
'' was established in 1974. Biogeography and ecology share many of their disciplinary roots. For example,
the theory of island biogeography ''The Theory of Island Biogeography'' is a 1967 book by the ecologist Robert MacArthur and the biologist Edward O. Wilson. It is widely regarded as a seminal piece in island biogeography and ecology. The Princeton University Press reprinted the ...
, published by the Robert MacArthur and Edward O. Wilson in 1967 is considered one of the fundamentals of ecological theory. Biogeography has a long history in the natural sciences concerning the spatial distribution of plants and animals. Ecology and evolution provide the explanatory context for biogeographical studies. Biogeographical patterns result from ecological processes that influence range distributions, such as
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
and dispersal. and from historical processes that split populations or species into different areas. The biogeographic processes that result in the natural splitting of species explain much of the modern distribution of the Earth's biota. The splitting of lineages in a species is called
vicariance biogeography Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
and it is a sub-discipline of biogeography. 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 the context of
global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
.


r/K selection theory

A population ecology concept 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 Population density (in agriculture: standing stock or plant density) is a measurement of population per unit land area. It is mostly applied to humans, but sometimes to other living organisms too. It is a key geographical term.Matt RosenberPopu ...
. 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 Resource refers to all the materials available in our environment which are technologically accessible, economically feasible and culturally sustainable and help us to satisfy our needs and wants. Resources can broadly be classified upon their av ...
for rapid population growth. These early phases of
population growth Population growth is the increase in the number of people in a population or dispersed group. Actual global human population growth amounts to around 83 million annually, or 1.1% per year. The global population has grown from 1 billion in 1800 to ...
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. 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. Different species evolve different life-history strategies spanning a continuum between these two selective forces. An ''r''-selected species is one that has high birth rates, low levels of parental investment, and high rates of mortality before individuals reach maturity. Evolution favours high rates of
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
in ''r''-selected species. Many kinds of insects and
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...
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.


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 the influx of research questions into this new ecological field resulted in the publication '' Molecular Ecology'' in 1992. Molecular ecology uses various analytical techniques to study genes in an evolutionary and ecological context. In 1994, John Avise also played a leading role in this area of science with the publication of his book, ''Molecular Markers, Natural History and Evolution''. 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
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s. Molecular ecology engendered a new research paradigm for investigating ecological questions considered otherwise intractable. Molecular investigations revealed previously obscured details in the tiny intricacies of nature and improved resolution into probing questions about behavioural and biogeographical ecology. For example, molecular ecology revealed promiscuous sexual behaviour and multiple male partners in tree swallows previously thought to be socially
monogamous Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time ( serial monogamy) — as compared to the various forms of non-monogamy (e.g., pol ...
. In a biogeographical context, the marriage between genetics, ecology, and evolution resulted in a new sub-discipline called phylogeography.


Human ecology

Ecology is as much a biological science as it is a human science. Human ecology is an
interdisciplinary Interdisciplinarity or interdisciplinary studies involves the combination of multiple academic disciplines into one activity (e.g., a research project). It draws knowledge from several other fields like sociology, anthropology, psychology, ec ...
investigation into the ecology of our species. "Human ecology may be defined: (1) from a bioecological standpoint as the study of man as the ecological dominant in plant and animal communities and systems; (2) from a bioecological standpoint as simply another animal affecting and being affected by his physical environment; and (3) as a human being, somehow different from animal life in general, interacting with physical and modified environments in a distinctive and creative way. A truly interdisciplinary human ecology will most likely address itself to all three." The term was formally introduced in 1921, but many sociologists, geographers, psychologists, and other disciplines were interested in human relations to natural systems centuries prior, especially in the late 19th century. The ecological complexities human beings are facing through the technological transformation of the planetary biome has brought on the
Anthropocene The Anthropocene ( ) is a proposed geological epoch dating from the commencement of significant human impact on Earth's geology and ecosystems, including, but not limited to, anthropogenic climate change. , neither the International Commissio ...
. The unique set of circumstances has generated the need for a new unifying science called coupled human and natural systems that builds upon, but moves beyond the field of human ecology. Ecosystems tie into human societies through the critical and all-encompassing life-supporting functions they sustain. In recognition of these functions and the incapability of traditional economic valuation methods to see the value in ecosystems, there has been a surge of interest in
social Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not. Etymology The word "social" derives from ...
-
natural capital Natural capital is the world's stock of natural resources, which includes geology, soils, air, water and all living organisms. Some natural capital assets provide people with free goods and services, often called ecosystem services. All of t ...
, which provides the means to put a value on the stock and use of information and materials stemming from ecosystem goods and services. Ecosystems produce, regulate, maintain, and supply services of critical necessity and beneficial to human health (cognitive and physiological), economies, and they even provide an information or reference function as a living library giving opportunities for science and cognitive development in children engaged in the complexity of the natural world. Ecosystems relate importantly to human ecology as they are the ultimate base foundation of global economics as every commodity, and the capacity for exchange ultimately stems from the ecosystems on Earth. Ecology is an employed science of restoration, repairing disturbed sites through human intervention, in natural resource management, and in
environmental impact assessment Environmental Impact assessment (EIA) is the assessment of the environmental consequences of a plan, policy, program, or actual projects prior to the decision to move forward with the proposed action. In this context, the term "environmental imp ...
s. Edward O. Wilson predicted in 1992 that the 21st century "will be the era of restoration in ecology". Ecological science has boomed in the industrial investment of restoring ecosystems and their processes in abandoned sites after disturbance. Natural resource managers, in
forestry Forestry is the science and craft of creating, managing, planting, using, conserving and repairing forests, woodlands, and associated resources for human and environmental benefits. Forestry is practiced in plantations and natural stands. ...
, for example, employ ecologists to develop, adapt, and implement ecosystem based methods into the planning, operation, and restoration phases of land-use. Another example of conservation is seen on the east coast of the United States in Boston, MA. The city of Boston implemented the Wetland Ordinance, improving the stability of their wetland environments by implementing soil amendments that will improve groundwater storage and flow, and trimming or removal of vegetation that could cause harm to water quality. Ecological science is used in the methods of sustainable harvesting, disease, and fire outbreak management, in fisheries stock management, for integrating land-use with protected areas and communities, and conservation in complex geo-political landscapes.


Relation to the environment

The environment of ecosystems includes both physical parameters and biotic attributes. It is dynamically interlinked and contains
resources Resource refers to all the materials available in our environment which are technologically accessible, economically feasible and culturally sustainable and help us to satisfy our needs and wants. Resources can broadly be classified upon their av ...
for organisms at any time throughout their life cycle. Like ecology, the term environment has different conceptual meanings and overlaps with the concept of nature. Environment "includes the physical world, the social world of human relations and the built world of human creation." The physical environment is external to the level of biological organization under investigation, including abiotic factors such as temperature, radiation, light, chemistry,
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
and geology. The biotic environment includes genes, cells, organisms, members of the same species (
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
s) and other species that share a habitat. The distinction between external and internal environments, however, is an abstraction parsing life and environment into units or facts that are inseparable in reality. There is an interpenetration of cause and effect between the environment and life. The laws of
thermodynamics Thermodynamics is a branch of physics that deals with heat, work, and temperature, and their relation to energy, entropy, and the physical properties of matter and radiation. The behavior of these quantities is governed by the four laws ...
, for example, apply to ecology by means of its physical state. With an understanding of metabolic and thermodynamic principles, a complete accounting of energy and material flow can be traced through an ecosystem. In this way, the environmental and ecological relations are studied through reference to conceptually manageable and isolated
material Material is a substance or mixture of substances that constitutes an object. Materials can be pure or impure, living or non-living matter. Materials can be classified on the basis of their physical and chemical properties, or on their geolo ...
parts. After the effective environmental components are understood through reference to their causes; however, they conceptually link back together as an integrated whole, or ''holocoenotic'' system as it was once called. This is known as the
dialectical Dialectic ( grc-gre, διαλεκτική, ''dialektikḗ''; related to dialogue; german: Dialektik), also known as the dialectical method, is a discourse between two or more people holding different points of view about a subject but wishing t ...
approach to ecology. The dialectical approach examines the parts but integrates the organism and the environment into a dynamic whole (or
umwelt In the semiotic theories of Jakob von Uexküll and Thomas A. Sebeok, ''umwelt'' (plural: umwelten; from the German '' Umwelt'' meaning "environment" or "surroundings") is the "biological foundations that lie at the very epicenter of the stu ...
). Change in one ecological or environmental factor can concurrently affect the dynamic state of an entire ecosystem.


Disturbance and resilience

Ecosystems are regularly confronted with natural environmental variations and disturbances over time and geographic space. A disturbance is any process that removes biomass from a community, such as a fire, flood, drought, or predation. Disturbances occur over vastly different ranges in terms of magnitudes as well as distances and time periods, and are both the cause and product of natural fluctuations in death rates, species assemblages, and biomass densities within an ecological community. These disturbances create places of renewal where new directions emerge from the patchwork of natural experimentation and opportunity. Ecological resilience is a cornerstone theory in ecosystem management. Biodiversity fuels the resilience of ecosystems acting as a kind of regenerative insurance.


Metabolism and the early atmosphere

The Earth was formed approximately 4.5 billion years ago. As it cooled and a crust and oceans formed, its atmosphere transformed from being dominated by
hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-to ...
to one composed mostly of
methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane ...
and
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous ...
. Over the next billion years, the metabolic activity of life transformed the atmosphere into a mixture of
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
,
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
, 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. Throughout history, the Earth's atmosphere and
biogeochemical cycles A biogeochemical cycle (or more generally a cycle of matter) is the pathway by which a chemical substance cycles (is turned over or moves through) the biotic and the abiotic compartments of Earth. The biotic compartment is the biosphere and the ...
have been in a
dynamic equilibrium In chemistry, a dynamic equilibrium exists once a reversible reaction occurs. Substances transition between the reactants and products at equal rates, meaning there is no net change. Reactants and products are formed at such a rate that the co ...
with planetary ecosystems. The history is characterized by periods of significant transformation followed by millions of years of stability. The evolution of the earliest organisms, likely anaerobic
methanogen Methanogens are microorganisms that produce methane as a metabolic byproduct in hypoxic conditions. They are prokaryotic and belong to the domain Archaea. All known methanogens are members of the archaeal phylum Euryarchaeota. Methanogens are c ...
microbes, started the process by converting atmospheric hydrogen into methane (4H2 + CO2 → CH4 + 2H2O). Anoxygenic photosynthesis reduced hydrogen concentrations and increased atmospheric methane, by converting
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The under ...
into water or other sulfur compounds (for example, 2H2S + CO2 + h''v'' → CH2O + H2O + 2S). Early forms of
fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food p ...
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.


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, behaviour, and
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...
. Temperature is largely dependent on the incidence of
solar radiation Solar irradiance is the power per unit area ( surface power density) received from the Sun in the form of electromagnetic radiation in the wavelength range of the measuring instrument. Solar irradiance is measured in watts per square metre ...
. The latitudinal and longitudinal spatial variation of
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied o ...
greatly affects climates and consequently the distribution of
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
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. 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 In physics, electromagnetism is an interaction that occurs between particles with electric charge. It is the second-strongest of the four fundamental interactions, after the strong force, and it is the dominant force in the interactions o ...
of different
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tr ...
s.
Radiant energy Radiant may refer to: Computers, software, and video games * Radiant (software), a content management system * GtkRadiant, a level editor created by id Software for their games * Radiant AI, a technology developed by Bethesda Softworks for ''Th ...
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 In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
. Plants, algae, and some bacteria absorb light and assimilate the energy through
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
. Organisms capable of assimilating energy by photosynthesis or through inorganic fixation of H2S are autotrophs. Autotrophs—responsible for primary production—assimilate light energy which becomes metabolically stored as
potential energy In physics, potential energy is the energy held by an object because of its position relative to other objects, stresses within itself, its electric charge, or other factors. Common types of potential energy include the gravitational potenti ...
in the form of biochemical
enthalpic Enthalpy , a property of a thermodynamic system, is the sum of the system's internal energy and the product of its pressure and volume. It is a state function used in many measurements in chemical, biological, and physical systems at a constant p ...
bonds.


Physical environments


Water

Diffusion of carbon dioxide and oxygen is approximately 10,000 times slower in water than in air. When soils are flooded, they quickly lose oxygen, becoming hypoxic (an environment with O2 concentration below 2 mg/liter) and eventually completely
anoxic The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of diss ...
where
anaerobic bacteria An anaerobic organism or anaerobe is any organism that does not require molecular oxygen for growth. It may react negatively or even die if free oxygen is present. In contrast, an aerobic organism (aerobe) is an organism that requires an oxygenat ...
thrive among the roots. Water also influences the intensity and spectral composition of light as it reflects off the water surface and submerged particles. Aquatic plants exhibit a wide variety of morphological and physiological adaptations that allow them to survive, compete, and diversify in these environments. For example, their roots and stems contain large air spaces ( aerenchyma) that regulate the efficient transportation of gases (for example, CO2 and O2) used in respiration and photosynthesis. Salt water plants (
halophytes A halophyte is a salt-tolerant plant that grows in soil or waters of high salinity, coming into contact with saline water through its roots or by salt spray, such as in saline semi-deserts, mangrove swamps, marshes and sloughs and seashores. T ...
) have additional specialized adaptations, such as the development of special organs for shedding salt and osmoregulating their internal salt (NaCl) concentrations, to live in
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
,
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
, or
ocean The ocean (also the sea or the world ocean) is the body of salt water that covers approximately 70.8% of the surface of Earth and contains 97% of Earth's water. An ocean can also refer to any of the large bodies of water into which the wor ...
ic environments. Anaerobic soil
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s in aquatic environments use
nitrate Nitrate is a polyatomic ion with the chemical formula . Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are soluble in water. An example of an insolu ...
, manganese ions, ferric ions,
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ...
,
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
, and some organic compounds; other microorganisms are facultative anaerobes and use oxygen during respiration when the soil becomes drier. The activity of soil microorganisms and the chemistry of the water reduces the oxidation-reduction potentials of the water. Carbon dioxide, for example, is reduced to methane (CH4) by methanogenic bacteria. The physiology of fish is also specially adapted to compensate for environmental salt levels through osmoregulation. Their gills form
electrochemical gradient An electrochemical gradient is a gradient of electrochemical potential, usually for an ion that can move across a membrane. The gradient consists of two parts, the chemical gradient, or difference in solute concentration across a membrane, and ...
s that mediate salt excretion in salt water and uptake in fresh water.


Gravity

The shape and energy of the land are significantly affected by gravitational forces. On a large scale, the distribution of gravitational forces on the earth is uneven and influences the shape and movement of
tectonic plates Plate tectonics (from the la, label=Late Latin, tectonicus, from the grc, τεκτονικός, lit=pertaining to building) is the generally accepted scientific theory that considers the Earth's lithosphere to comprise a number of large ...
as well as influencing geomorphic processes such as
orogeny Orogeny is a mountain building process. An orogeny is an event that takes place at a convergent plate margin when plate motion compresses the margin. An '' orogenic belt'' or ''orogen'' develops as the compressed plate crumples and is uplifted ...
and
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is d ...
. These forces govern many of the geophysical properties and distributions of ecological biomes across the Earth. On the organismal scale, gravitational forces provide directional cues for plant and fungal growth ( gravitropism), orientation cues for animal migrations, and influence the
biomechanics Biomechanics is the study of the structure, function and motion of the mechanical aspects of biological systems, at any level from whole organisms to organs, cells and cell organelles, using the methods of mechanics. Biomechanics is a branch of ...
and size of animals. 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. The cardiovascular systems of animals are functionally adapted to overcome the pressure and gravitational forces that change according to the features of organisms (e.g., height, size, shape), their behaviour (e.g., diving, running, flying), and the habitat occupied (e.g., water, hot deserts, cold tundra).


Pressure

Climatic and
osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in a pure ...
places
physiological Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemica ...
constraints on organisms, especially those that fly and respire at high altitudes, or dive 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. For example, oxygen levels decrease with decreasing pressure and are a limiting factor for life at higher altitudes.
Water transportation Water transportation is the international movement of water over large distances. Methods of transportation fall into three categories: * Aqueducts, which include pipelines, canals, tunnels and bridges * Container shipment, which includes trans ...
by plants is another important ecophysiological process affected by osmotic pressure gradients. Water pressure in the depths of oceans requires that organisms adapt to these conditions. For example, diving animals such as
whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s,
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the b ...
s, and
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
are specially adapted to deal with changes in sound due to water pressure differences. Differences between
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, ...
species provide another example of adaptation to deep-sea pressure through specialized protein adaptations.


Wind and turbulence

Turbulent forces in air and water affect the environment and ecosystem distribution, form, and dynamics. On a planetary scale, ecosystems are affected by circulation patterns in the global
trade winds The trade winds or easterlies are the permanent east-to-west prevailing winds that flow in the Earth's equatorial region. The trade winds blow mainly from the northeast in the Northern Hemisphere and from the southeast in the Southern Hemisp ...
. Wind power and the turbulent forces it creates can influence heat, nutrient, and biochemical profiles of ecosystems. For example, wind running over the surface of a lake creates turbulence, mixing the
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
and influencing the environmental profile to create thermally layered zones, affecting how fish, algae, and other parts of the
aquatic ecosystem An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The t ...
are structured. Wind speed and turbulence also influence evapotranspiration rates and energy budgets in plants and animals. Wind speed, temperature and moisture content can vary as winds travel across different land features and elevations. For example, the
westerlies The westerlies, anti-trades, or prevailing westerlies, are prevailing winds from the west toward the east in the middle latitudes between 30 and 60 degrees latitude. They originate from the high-pressure areas in the horse latitudes and tren ...
come into contact with the
coastal The coast, also known as the coastline or seashore, is defined as the area where land meets the ocean, or as a line that forms the boundary between the land and the coastline. The Earth has around of coastline. Coasts are important zones in ...
and interior mountains of western North America to produce a
rain shadow A rain shadow is an area of significantly reduced rainfall behind a mountainous region, on the side facing away from prevailing winds, known as its leeward side. Evaporated moisture from water bodies (such as oceans and large lakes) is ca ...
on the leeward side of the mountain. The air expands and moisture condenses as the winds increase in elevation; this is called
orographic lift Orographic lift occurs when an air mass is forced from a low elevation to a higher elevation as it moves over rising terrain. As the air mass gains altitude it quickly cools down adiabatically, which can raise the relative humidity to 100% and cr ...
and can cause precipitation. This environmental process produces spatial divisions in biodiversity, as species adapted to wetter conditions are range-restricted to the coastal mountain valleys and unable to migrate across the
xeric Deserts and xeric shrublands are a biome defined by the World Wide Fund for Nature. Deserts and xeric (ancient Greek xērós, “dry") shrublands form the largest terrestrial biome, covering 19% of Earth's land surface area. Ecoregions in this h ...
ecosystems (e.g., of the Columbia Basin in western North America) to intermix with sister lineages that are segregated to the interior mountain systems.


Fire

Plants convert carbon dioxide into biomass and emit oxygen into the atmosphere. By approximately 350 million years ago (the end of the
Devonian period The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, w ...
), photosynthesis had brought the concentration of atmospheric oxygen above 17%, which allowed combustion to occur. Fire releases CO2 and converts fuel into ash and tar. Fire is a significant ecological parameter that raises many issues pertaining to its control and suppression. While the issue of fire in relation to ecology and plants has been recognized for a long time, Charles Cooper brought attention to the issue of forest fires in relation to the ecology of forest fire suppression and management in the 1960s. Native North Americans were among the first to influence fire regimes by controlling their spread near their homes or by lighting fires to stimulate the production of herbaceous foods and basketry materials. Fire creates a heterogeneous ecosystem age and canopy structure, and the altered soil nutrient supply and cleared canopy structure opens new ecological niches for seedling establishment. Most ecosystems are adapted to natural fire cycles. Plants, for example, are equipped with a variety of adaptations to deal with forest fires. Some species (e.g., '' Pinus halepensis'') cannot
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
until after their seeds have lived through a fire or been exposed to certain compounds from smoke. Environmentally triggered germination of seeds is called serotiny. Fire plays a major role in the persistence and
resilience Resilience, resilient, resiliency, or ''variation'', may refer to: Science Ecology * Ecological resilience, the capacity of an ecosystem to recover from perturbations ** Climate resilience, the ability of systems to recover from climate change * ...
of ecosystems.


Soils

Soil is the living top layer of mineral and organic dirt that covers the surface of the planet. It is the chief organizing centre of most ecosystem functions, and it is of critical importance in agricultural science and ecology. The
decomposition Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and ...
of dead organic matter (for example, leaves on the forest floor), results in soils containing
minerals In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed ...
and nutrients that feed into plant production. The whole of the planet's soil ecosystems is called the pedosphere where a large biomass of the Earth's biodiversity organizes into trophic levels. Invertebrates that feed and shred larger leaves, for example, create smaller bits for smaller organisms in the feeding chain. Collectively, these organisms are the
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...
s that regulate soil formation. 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. Soils form composite phenotypes where inorganic matter is enveloped into the physiology of a whole community. As organisms feed and migrate through soils they physically displace materials, an ecological process called
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a ...
. This aerates soils and stimulates heterotrophic growth and production. Soil
microorganisms A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
are influenced by and are fed back into the trophic dynamics of the ecosystem. No single axis of causality can be discerned to segregate the biological from geomorphological systems in soils. 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 the
colonization of land The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''gigaannum'') and evide ...
in the Devonian period played a significant role in the early development of ecological trophism in soils.


Biogeochemistry and climate

Ecologists study and measure nutrient budgets to understand how these materials are regulated, flow, and
recycled Recycling is the process of converting waste materials into new materials and objects. The recovery of energy from waste materials is often included in this concept. The recyclability of a material depends on its ability to reacquire the p ...
through the environment. This research has led to an understanding that there is global feedback between ecosystems and the physical parameters of this planet, including minerals, soil, pH, ions, water, and atmospheric gases. Six major elements (
hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-to ...
,
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...
,
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
,
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
,
sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formul ...
, and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ea ...
; H, C, N, O, S, and P) form the constitution of all biological macromolecules and feed into the Earth's geochemical processes. From the smallest scale of biology, the combined effect of billions upon billions of ecological processes amplify and ultimately regulate the
biogeochemical cycle A biogeochemical cycle (or more generally a cycle of matter) is the pathway by which a chemical substance cycles (is turned over or moves through) the biotic and the abiotic compartments of Earth. The biotic compartment is the biosphere and th ...
s of the Earth. Understanding the relations and cycles mediated between these elements and their ecological pathways has significant bearing toward understanding global biogeochemistry. The ecology of global carbon budgets gives one example of the linkage between biodiversity and biogeochemistry. It is estimated that the Earth's oceans hold 40,000 gigatonnes (Gt) of carbon, that vegetation and soil hold 2070 Gt, and that fossil fuel emissions are 6.3 Gt carbon per year. There have been major restructurings in these global carbon budgets during the Earth's history, regulated to a large extent by the ecology of the land. For example, through the early-mid Eocene volcanic
outgassing Outgassing (sometimes called offgassing, particularly when in reference to indoor air quality) is the release of a gas that was dissolved, trapped, frozen, or absorbed in some material. Outgassing can include sublimation and evaporation (which ...
, the oxidation of methane stored in wetlands, and seafloor gases increased atmospheric CO2 (carbon dioxide) concentrations to levels as high as 3500  ppm. In the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but t ...
, from twenty-five to thirty-two million years ago, there was another significant restructuring of the global
carbon cycle The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth. Carbon is the main component of biological compounds as well as a major compon ...
as grasses evolved a new mechanism of photosynthesis, C4 photosynthesis, and expanded their ranges. This new pathway evolved in response to the drop in atmospheric CO2 concentrations below 550 ppm. The relative abundance and distribution of biodiversity alters the dynamics between organisms and their environment such that ecosystems can be both cause and effect in relation to climate change. Human-driven modifications to the planet's ecosystems (e.g., disturbance,
biodiversity loss Biodiversity loss includes the worldwide extinction of different species, as well as the local reduction or loss of species in a certain habitat, resulting in a loss of biological diversity. The latter phenomenon can be temporary or permanent, de ...
, agriculture) contributes to rising atmospheric greenhouse gas levels. Transformation of the global carbon cycle in the next century is projected to raise planetary temperatures, lead to more extreme fluctuations in weather, alter species distributions, and increase extinction rates. The effect of global warming is already being registered in melting glaciers, melting mountain ice caps, and rising sea levels. Consequently, species distributions are changing along waterfronts and in continental areas where migration patterns and breeding grounds are tracking the prevailing shifts in climate. Large sections of
permafrost Permafrost is ground that continuously remains below 0 °C (32 °F) for two or more years, located on land or under the ocean. Most common in the Northern Hemisphere, around 15% of the Northern Hemisphere or 11% of the global surface ...
are also melting to create a new mosaic of flooded areas having increased rates of soil decomposition activity that raises methane (CH4) emissions. There is concern over increases in atmospheric methane in the context of the global carbon cycle, because methane is a
greenhouse gas A greenhouse gas (GHG or GhG) is a gas that absorbs and emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse gases in Earth's atmosphere are water vapor (), carbon dioxide (), methane ...
that is 23 times more effective at absorbing long-wave radiation than CO2 on a 100-year time scale. Hence, there is a relationship between global warming, decomposition and respiration in soils and wetlands producing significant climate feedbacks and globally altered biogeochemical cycles.


History


Early beginnings

Ecology has a complex origin, due in large part to its interdisciplinary nature. Ancient Greek philosophers such as
Hippocrates Hippocrates of Kos (; grc-gre, Ἱπποκράτης ὁ Κῷος, Hippokrátēs ho Kôios; ), also known as Hippocrates II, was a Greek physician of the classical period who is considered one of the most outstanding figures in the history o ...
and
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
were among the first to record observations on natural history. However, they viewed life in terms of
essentialism Essentialism is the view that objects have a set of attributes that are necessary to their identity. In early Western thought, Plato's idealism held that all things have such an "essence"—an "idea" or "form". In ''Categories'', Aristotle sim ...
, where species were conceptualized as static unchanging things while varieties were seen as aberrations of an idealized type. This contrasts against the modern understanding of ecological theory where varieties are viewed as the real phenomena of interest and having a role in the origins of adaptations by means of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
. Early conceptions of ecology, such as a balance and regulation in nature can be traced to
Herodotus Herodotus ( ; grc, , }; BC) was an ancient Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire (now Bodrum, Turkey) and a later citizen of Thurii in modern Calabria (Italy). He is known fo ...
(died ''c''. 425 BC), who described one of the earliest accounts of mutualism in his observation of "natural dentistry". Basking
Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed throughout sub-Saharan Africa, occurring mostly in the central, eastern, ...
s, he noted, would open their mouths to give
sandpiper Sandpipers are a large family, Scolopacidae, of waders. They include many species called sandpipers, as well as those called by names such as curlew and snipe. The majority of these species eat small invertebrates picked out of the mud or soil ...
s safe access to pluck
leech Leeches are segmented parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
es out, giving nutrition to the sandpiper and oral hygiene for the crocodile. Aristotle was an early influence on the philosophical development of ecology. He and his student
Theophrastus Theophrastus (; grc-gre, Θεόφραστος ; c. 371c. 287 BC), a Greek philosopher and the successor to Aristotle in the Peripatetic school. He was a native of Eresos in Lesbos.Gavin Hardy and Laurence Totelin, ''Ancient Botany'', Routle ...
made extensive observations on plant and animal migrations, biogeography, physiology, and their behavior, giving an early analogue to the modern concept of an ecological niche.
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
(left) and Eugenius Warming (right), two founders of ecology
Ecological concepts such as food chains, population regulation, and productivity were first developed in the 1700s, through the published works of microscopist
Antoni van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
(1632–1723) and botanist Richard Bradley (1688?–1732). Biogeographer
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a German polymath, geographer, naturalist, explorer, and proponent of Romantic philosophy and science. He was the younger brother of the Prussian minister ...
(1769–1859) was an early pioneer in ecological thinking and was among the first to recognize ecological gradients, where species are replaced or altered in form along environmental gradients, such as a cline forming along a rise in elevation. Humboldt drew inspiration from
Isaac Newton Sir Isaac Newton (25 December 1642 – 20 March 1726/27) was an English mathematician, physicist, astronomer, alchemist, Theology, theologian, and author (described in his time as a "natural philosophy, natural philosopher"), widely ...
, as he developed a form of "terrestrial physics". In Newtonian fashion, he brought a scientific exactitude for measurement into natural history and even alluded to concepts that are the foundation of a modern ecological law on species-to-area relationships. Natural historians, such as Humboldt,
James Hutton James Hutton (; 3 June O.S.172614 June 1726 New Style. – 26 March 1797) was a Scottish geologist, agriculturalist, chemical manufacturer, naturalist and physician. Often referred to as the father of modern geology, he played a key role ...
, and
Jean-Baptiste Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
(among others) laid the foundations of the modern ecological sciences. The term "ecology" (german: Oekologie, Ökologie) was coined by
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
in his book ''Generelle Morphologie der Organismen'' (1866). From p. 286: ''"Unter Oecologie verstehen wir die gesammte Wissenschaft von den Beziehungen des Organismus zur umgebenden Aussenwelt, wohin wir im weiteren Sinne alle "Existenz-Bedingungen" rechnen können."'' (By "ecology" we understand the comprehensive science of the relationships of the organism to its surrounding environment, where we can include, in the broader sense, all "conditions of existence".) Haeckel was a zoologist, artist, writer, and later in life a professor of comparative anatomy. Opinions differ on who was the founder of modern ecological theory. Some mark Haeckel's definition as the beginning; others say it was Eugenius Warming with the writing of Oecology of Plants: An Introduction to the Study of Plant Communities (1895), or
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
' principles on the economy of nature that matured in the early 18th century. Linnaeus founded an early branch of ecology that he called the economy of nature. His works influenced Charles Darwin, who adopted Linnaeus' phrase on the ''economy or polity of nature'' in '' The Origin of Species''. Linnaeus was the first to frame the
balance of nature The balance of nature, also known as ecological balance, is a theory that proposes that ecological systems are usually in a stable equilibrium or homeostasis, which is to say that a small change (the size of a particular population, for example) w ...
as a testable hypothesis. Haeckel, who admired Darwin's work, defined ecology in reference to the economy of nature, which has led some to question whether ecology and the economy of nature are synonymous. From Aristotle until Darwin, the natural world was predominantly considered static and unchanging. Prior to ''The Origin of Species'', there was little appreciation or understanding of the dynamic and reciprocal relations between organisms, their adaptations, and the environment. An exception is the 1789 publication ''Natural History of Selborne'' by
Gilbert White Gilbert White FRS (18 July 1720 – 26 June 1793) was a " parson-naturalist", a pioneering English naturalist, ecologist, and ornithologist. He is best known for his ''Natural History and Antiquities of Selborne''. Life White was born on ...
(1720–1793), considered by some to be one of the earliest texts on ecology. While
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
is mainly noted for his treatise on evolution, he was one of the founders of
soil ecology Soil ecology is the study of the interactions among soil organisms, and between biotic and abiotic aspects of the soil environment. It is particularly concerned with the cycling of nutrients, formation and stabilization of the pore structure, the ...
, and he made note of the first ecological experiment in ''The Origin of Species''. Evolutionary theory changed the way that researchers approached the ecological sciences.


Since 1900

Modern ecology is a young science that first attracted substantial scientific attention toward the end of the 19th century (around the same time that evolutionary studies were gaining scientific interest). The scientist
Ellen Swallow Richards Ellen Henrietta Swallow Richards (December 3, 1842 – March 30, 1911) was an American industrial and safety engineer, environmental chemist, and university faculty member in the United States during the 19th century. Her pioneering work i ...
adopted the term " oekology" (which eventually morphed into
home economics Home economics, also called domestic science or family and consumer sciences, is a subject concerning human development, personal and family finances, consumer issues, housing and interior design, nutrition and food preparation, as well as texti ...
) in the U.S. as early as 1892. In the early 20th century, ecology transitioned from a more descriptive form of natural history to a more analytical form of ''scientific natural history''.
Frederic Clements Frederic Edward Clements (September 16, 1874 – July 26, 1945) was an American plant ecologist and pioneer in the study of plant ecology and vegetation succession. Biography Born in Lincoln, Nebraska, he studied botany at the University of Neb ...
published the first American ecology book in 1905, presenting the idea of plant communities as a superorganism. This publication launched a debate between ecological holism and individualism that lasted until the 1970s. Clements' superorganism concept proposed that ecosystems progress through regular and determined stages of seral development that are analogous to the developmental stages of an organism. The Clementsian paradigm was challenged by Henry Gleason, who stated that ecological communities develop from the unique and coincidental association of individual organisms. This perceptual shift placed the focus back onto the life histories of individual organisms and how this relates to the development of community associations. The Clementsian superorganism theory was an overextended application of an idealistic form of holism. The term "holism" was coined in 1926 by
Jan Christiaan Smuts Field Marshal Jan Christian Smuts, (24 May 1870 11 September 1950) was a South African statesman, military leader and philosopher. In addition to holding various military and cabinet posts, he served as Prime Minister of South Africa, prime m ...
, a South African general and polarizing historical figure who was inspired by Clements' superorganism concept. Around the same time,
Charles Elton Charles Elton may refer to: *Charles Elton (Born, 1993) Professional Rugby Player for Otago Rugby * Charles Isaac Elton (1839–1900), English lawyer, politician, writer and antiquarian * Charles Sutherland Elton (1900–1991), English biologist ...
pioneered the concept of food chains in his classical book ''Animal Ecology''. Elton defined ecological relations using concepts of food chains, food cycles, and food size, and described numerical relations among different functional groups and their relative abundance. Elton's 'food cycle' was replaced by 'food web' in a subsequent ecological text.
Alfred J. Lotka Alfred James Lotka (March 2, 1880 – December 5, 1949) was a US mathematician, physical chemist, and statistician, famous for his work in population dynamics and energetics. An American biophysicist, Lotka is best known for his proposa ...
brought in many theoretical concepts applying thermodynamic principles to ecology. In 1942,
Raymond Lindeman Raymond Laurel Lindeman (1915 – June 29, 1942) was an ecologist whose graduate research is credited with being a seminal study in the field of ecosystem ecology, specifically on the topic of trophic dynamics. Graduate research work Lindeman ...
wrote a landmark paper on the
trophic dynamics A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...
of ecology, which was published posthumously after initially being rejected for its theoretical emphasis. Trophic dynamics became the foundation for much of the work to follow on energy and material flow through ecosystems. Robert MacArthur advanced mathematical theory, predictions, and tests in ecology in the 1950s, which inspired a resurgent school of theoretical mathematical ecologists. Ecology also has developed through contributions from other nations, including Russia's Vladimir Vernadsky and his founding of the biosphere concept in the 1920s and Japan's Kinji Imanishi and his concepts of harmony in nature and habitat segregation in the 1950s. Scientific recognition of contributions to ecology from non-English-speaking cultures is hampered by language and translation barriers. Ecology surged in popular and scientific interest during the 1960–1970s
environmental movement The environmental movement (sometimes referred to as the ecology movement), also including conservation and green politics, is a diverse philosophical, social, and political movement for addressing environmental issues. Environmentalists a ...
. There are strong historical and scientific ties between ecology, environmental management, and protection. The historical emphasis and poetic naturalistic writings advocating the protection of wild places by notable ecologists in the history of
conservation biology Conservation biology is the study of the conservation of nature and of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction and the erosion of biotic interactions. It is an in ...
, such as
Aldo Leopold Aldo Leopold (January 11, 1887 – April 21, 1948) was an American writer, philosopher, naturalist, scientist, ecologist, forester, conservationist, and environmentalist. He was a professor at the University of Wisconsin and is best known for his ...
and
Arthur Tansley Sir Arthur George Tansley FLS, FRS (15 August 1871 – 25 November 1955) was an English botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phy ...
, have been seen as far removed from urban centres where, it is claimed, the concentration of pollution and environmental degradation is located. Palamar (2008) notes an overshadowing by mainstream environmentalism of pioneering women in the early 1900s who fought for urban health ecology (then called
euthenics Euthenics () is the study of improvement of human functioning and well-being by improvement of living conditions. "Improvement" is conducted by altering external factors such as education and the controllable environments, including environmen ...
) and brought about changes in environmental legislation. Women such as
Ellen Swallow Richards Ellen Henrietta Swallow Richards (December 3, 1842 – March 30, 1911) was an American industrial and safety engineer, environmental chemist, and university faculty member in the United States during the 19th century. Her pioneering work i ...
and
Julia Lathrop Julia Clifford Lathrop (June 29, 1858 – April 15, 1932) was an American social reformer in the area of education, social policy, and children's welfare. As director of the United States Children's Bureau from 1912 to 1922, she was the first wo ...
, among others, were precursors to the more popularized environmental movements after the 1950s. In 1962, marine biologist and ecologist
Rachel Carson Rachel Louise Carson (May 27, 1907 – April 14, 1964) was an American marine biologist, writer, and conservationist whose influential book '' Silent Spring'' (1962) and other writings are credited with advancing the global environmental ...
's book ''
Silent Spring ''Silent Spring'' is an environmental science book by Rachel Carson. Published on September 27, 1962, the book documented the environmental harm caused by the indiscriminate use of pesticides. Carson accused the chemical industry of spreading d ...
'' helped to mobilize the environmental movement by alerting the public to toxic
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and ...
s, such as DDT, bioaccumulating in the environment. Carson used ecological science to link the release of environmental toxins to human and ecosystem health. Since then, ecologists have worked to bridge their understanding of the degradation of the planet's ecosystems with environmental politics, law, restoration, and natural resources management.


See also

*
Carrying capacity The carrying capacity of an environment is the maximum population size of a biological species that can be sustained by that specific environment, given the food, habitat, water, and other resources available. The carrying capacity is defined as ...
* Chemical ecology *
Climate justice Climate justice is a concept that addresses the just division, fair sharing, and equitable distribution of the burdens of climate change and its mitigation and responsibilities to deal with climate change. "Justice", "fairness", and "equity" ar ...
*
Circles of Sustainability Circles of Sustainability is a method for understanding and assessing sustainability, and for project management directed towards socially sustainable outcomes. It is intended to handle 'seemingly intractable problems' such as outlined i ...
*
Cultural ecology Cultural ecology is the study of human adaptations to social and physical environments. Human adaptation refers to both biological and cultural processes that enable a population to survive and reproduce within a given or changing environment. Thi ...
*
Dialectical naturalism Murray Bookchin (January 14, 1921 – July 30, 2006) was an American social theorist, author, orator, historian, and political philosopher. A pioneer in the environmental movement, Bookchin formulated and developed the theory of social ...
* Ecological death * Ecological psychology *
Ecology movement The environmental movement (sometimes referred to as the ecology movement), also including conservation and green politics, is a diverse philosophical, social, and political movement for addressing environmental issues. Environmentalists advoc ...
* Ecosophy * Ecopsychology *
Human ecology Human ecology is an interdisciplinary and transdisciplinary study of the relationship between humans and their natural, social, and built environments. The philosophy and study of human ecology has a diffuse history with advancements in ecolog ...
* Industrial ecology *
Information ecology Information ecology is the application of ecological concepts for modeling the information society. It considers the dynamics and properties of the increasingly dense, complex and important digital informational environment. "Information ecology" of ...
*
Landscape ecology Landscape ecology is the science of studying and improving relationships between ecological processes in the environment and particular ecosystems. This is done within a variety of landscape scales, development spatial patterns, and organizatio ...
*
Natural resource Natural resources are resources that are drawn from nature and used with few modifications. This includes the sources of valued characteristics such as commercial and industrial use, aesthetic value, scientific interest and cultural value. ...
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Normative science In the applied sciences, normative science is a type of information that is developed, presented, or interpreted based on an assumed, usually unstated, preference for a particular outcome, policy or class of policies or outcomes. Regular or tradit ...
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Philosophy of ecology Philosophy of ecology is a concept under the philosophy of science, which is a subfield of philosophy. Its main concerns centre on the practice and application of ecology, its moral issues, and the intersectionality between the position of humans a ...
* Political ecology *
Theoretical ecology Theoretical ecology is the scientific discipline devoted to the study of ecological systems using theoretical methods such as simple conceptual models, mathematical models, computational simulations, and advanced data analysis. Effective models im ...
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Sensory ecology Sensory ecology is a relatively new field focusing on the information organisms obtain about their environment. It includes questions of what information is obtained, how it is obtained (the mechanism), and why the information is useful to the org ...
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Sexecology Sexecology, also known as ecosexuality, is a radical form of environmental activism based around nature fetishism, the idea of the earth as a lover. It invites people to treat the earth with love rather than see it as an infinite resource to explo ...
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Spiritual ecology Spiritual ecology is an emerging field in religion, conservation, and academia recognizing that there is a spiritual facet to all issues related to conservation, environmentalism, and earth stewardship. Proponents of Spiritual Ecology assert a nee ...
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Sustainable development Sustainable development is an organizing principle for meeting human development goals while also sustaining the ability of natural systems to provide the natural resources and ecosystem services on which the economy and society depend. The ...
; Lists * Glossary of ecology * Index of biology articles *
List of ecologists This is a list of notable ecologists. A-D * John Aber (USA) * Aziz Ab'Saber ( Brazil) * Charles Christopher Adams (USA) * Warder Clyde Allee (USA) * Herbert G. Andrewartha ( Australia) * Sarah Martha Baker ( UK) * Fakhri A. Bazzaz (USA) * ...
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Outline of biology Biology – The natural science that studies life. Areas of focus include structure, function, growth, origin, evolution, distribution, and taxonomy. History of biology *History of anatomy *History of biochemistry * History of biotechn ...
* Terminology of ecology


Notes

, meaning "dwelling place, distributional area" —quoted from Stauffer (1957).


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