Ecology is the scientific study of interactions among organisms and their environment, such as the interactions organisms have with each other and with their abiotic environment. Topics of interest to ecologists include the diversity, distribution, amount (biomass), number (population) of organisms, as well as competition between them within and among ecosystems. Ecosystems are composed of dynamically interacting parts including organisms, the communities they make up, and the non-living components of their environment. Ecosystem processes, such as primary production, pedogenesis, nutrient cycling, and various niche construction activities, regulate the flux of energy and matter through an environment. These processes are sustained by organisms with specific life history traits, and the variety of organisms is called biodiversity. Biodiversity, which refers to the varieties of species, genes, and ecosystems, enhances certain ecosystem services.
Ecology is an interdisciplinary field that includes biology and Earth science. The word “ecology” (“Ökologie”) was coined in 1866 by the German scientist Ernst Haeckel (1834–1919). Ancient Greek philosophers such as Hippocrates and Aristotle laid the foundations of ecology in their studies on natural history. Modern ecology transformed into a more rigorous science in the late 19th century. Evolutionary concepts on adaptation and natural selection became cornerstones of modern ecological theory. Ecology is not synonymous with environment, environmentalism, natural history, or environmental science. It is closely related to evolutionary biology, genetics, and ethology. An understanding of how biodiversity affects ecological function is an important focus area in ecological studies. Ecologists seek to explain:
Life processes, interactions and adaptations
The movement of materials and energy through living communities
The successional development of ecosystems, and
The abundance and distribution of organisms and biodiversity in the context of the environment.
Ecology is a human science as well. There are many practical applications of ecology in conservation biology, wetland management, natural resource management (agroecology, agriculture, forestry, agroforestry, fisheries), city planning (urban ecology), community health, economics, basic and applied science, and human social interaction (human ecology). Organisms and resources compose ecosystems which, in turn, maintain biophysical feedback mechanisms that moderate processes acting on living (biotic) and nonliving (abiotic) components of the planet. Ecosystems sustain life-supporting functions and produce natural capital like biomass production (food, fuel, fiber and medicine), the regulation of climate, global biogeochemical cycles, water filtration, soil formation, erosion control, flood protection and many other natural features of scientific, historical, economic, or intrinsic value.
Integrative levels, scope, and scale of organization
The scope of ecology covers a wide array of interacting levels of organization spanning micro-level to planetary scale phenomena. Ecosystems, for example, contain abiotic resources and interacting life forms (i.e., individual organisms that aggregate into populations 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 critically relevant to organisms living in and on it. Several generations of an aphid population can exist over the lifespan of a single leaf. Each of those aphids, in turn, support diverse bacterial communities. 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 scale of ecological dynamics can operate like a closed system, such as aphids migrating on a single tree, while at the same time remain open with regard to broader scale influences, such as atmosphere or climate. Hence, ecologists classify ecosystems hierarchically by analyzing data collected from finer scale units, such as vegetation associations, climate, and soil types, and integrate this information to identify emergent patterns of uniform organization and processes that operate on local to regional, landscape, 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 genes, to cells, to tissues, to organs, to organisms, to species, to populations, to communities, to ecosystems, to biomes, and up to the level of the biosphere. This framework forms a panarchy and exhibits non-linear 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 (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 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 services which by definition maintain and improve human quality of life. Preventing species extinctions is one way to preserve biodiversity and that goal rests on techniques that preserve genetic diversity, habitat and the ability for species to migrate. Conservation priorities and management techniques require different approaches and considerations to address the full ecological scope of biodiversity. Natural capital that supports populations is critical for maintaining ecosystem services and species migration (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.
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 or alpine 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 lizards (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 and habitat are sometimes used interchangeably, but the former applies to a community’s environment, whereas the latter applies to a species’ environment.
Additionally, some species are ecosystem engineers, altering the environment within a localized region. For instance, beavers manage water levels by building dams which improves their habitat in a landscape.
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 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, 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 states that two species cannot coexist indefinitely by living off the same limiting resource; one will always outcompete 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 coevolution and shared niche occupancy of similar species inhabiting species-rich communities. The habitat plus the niche is called the ecotope, which is defined as the full range of environmental and biological variables affecting an entire species.
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 is an example of holism applied in ecological theory. The Gaia hypothesis states that there is an emergent feedback loop generated by the metabolism of living organisms that maintains the temperature of the Earth and atmospheric conditions within a narrow self-regulating range of tolerance.
Population ecology studies the dynamics of species populations and how these populations interact with the 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.”:18 Simplified population models usually start with four variables: death, birth, immigration, and emigration.
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 processes create the observed data. In these island models, the rate of population change is described by:
where N is the total number of individuals in the population, B is the number of births, D is the number of deaths, b and d are the per capita rates of birth and death respectively, and r is the per capita rate of population change. The formula states that the rate of change in population size (dN/dT) is equal to births minus deaths (B – D).
Using these modelling techniques, Malthus’ population principle of growth was later transformed into a model known as the logistic equation:
where N is the number of individuals measured as biomass density, a is the maximum per-capita rate of change, and K is the carrying capacity of the population. The formula states that the rate of change in population size (dN/dT) is equal to growth (aN) that is limited by carrying capacity (1 – N/K).
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, and survivorship, and these are analysed using mathematical techniques such as matrix algebra. The information is used for managing wildlife stocks and setting harvest quotas. In cases when the use of null hypotheses is not appropriate, 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.”