Ecology (from Greek: οἶκος, "house", or "environment";
-λογία, "study of")[A] is the branch of biology which studies
the interactions among organisms and their environment. Objects of
study include interactions of organisms with each other and with
abiotic components of their environment. Topics of interest include
the biodiversity, distribution, biomass, and populations of organisms,
as well as cooperation and competition within and between species.
Ecosystems are dynamically interacting systems of organisms, the
communities they make up, and the non-living components of their
Ecosystem processes, such as primary production,
pedogenesis, nutrient cycling, and niche construction, regulate the
flux of energy and matter through an environment. These processes are
sustained by organisms with specific life history traits. Biodiversity
means the varieties of species, genes, and ecosystems, enhances
certain ecosystem services.
Ecology is not synonymous with environmentalism, natural history, or
environmental science. It overlaps with the closely related sciences
of evolutionary biology, genetics, and ethology. An important focus
for ecologists is to improve the understanding of how biodiversity
affects ecological function. Ecologists seek to explain:
Life processes, interactions, and adaptations
The movement of materials and energy through living communities
The successional development of ecosystems
The abundance and distribution of organisms and biodiversity in the
context of the environment.
Ecology has practical applications 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). For example, the Circles of
Sustainability approach treats ecology as more than the environment
'out there'. It is not treated as separate from humans. Organisms
(including humans) and resources compose ecosystems which, in turn,
maintain biophysical feedback mechanisms that moderate processes
acting on living (biotic) and non-living (abiotic) components of the
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.
The word "ecology" ("Ökologie") was coined in
1866 by the German
scientist Ernst Haeckel. Ecological thought is derivative of
established currents in philosophy, particularly from ethics and
politics. Ancient Greek philosophers such as
Aristotle laid the foundations of ecology in their studies on natural
history. Modern ecology became a much more rigorous science in the
late 19th century. Evolutionary concepts relating to adaptation and
natural selection became the cornerstones of modern ecological theory.
1 Levels, scope, and scale of organization
1.5 Niche construction
1.8 Individual ecology
1.10 Metapopulations and migration
1.11 Community ecology
1.13 Food webs
1.14 Trophic levels
1.15 Keystone species
2 Ecological complexity
3 Relation to evolution
3.1 Behavioural ecology
3.2 Cognitive ecology
3.3 Social ecology
3.5.1 r/K selection theory
3.6 Molecular ecology
4 Human ecology
4.1 Restoration and management
5 Relation to the environment
5.1 Disturbance and resilience
Metabolism and the early atmosphere
5.3 Radiation: heat, temperature and light
5.4 Physical environments
Wind and turbulence
Biogeochemistry and climate
6.1 Early beginnings
6.2 Since 1900
7 See also
10 External links
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) to a planetary scale
(e.g., biosphere) phenomena. Ecosystems, for example, contain abiotic
resources and interacting life forms (i.e., individual organisms that
aggregate into populations which aggregate into distinct ecological
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.
Biological organisation and Biological classification
System behaviors must first be arrayed into different levels of
organization. Behaviors corresponding to higher levels occur at slow
rates. Conversely, lower organizational levels exhibit rapid rates.
For example, individual tree leaves respond rapidly to momentary
changes in light intensity, CO2 concentration, and the like. The
growth of the tree responds more slowly and integrates these
O'Neill et al. (1986):76
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
Main article: Biodiversity
Biodiversity refers to the variety of life and its processes. It
includes the variety of living organisms, the genetic differences
among them, the communities and ecosystems in which they occur, and
the ecological and evolutionary processes that keep them functioning,
yet ever changing and adapting.
Noss & Carpenter (1994):5
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
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
Biodiversity plays an important role in ecosystem
services 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 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,
Main article: Habitat
Biodiversity of a coral reef. Corals adapt to and modify their
environment by forming calcium carbonate skeletons. This provides
growing conditions for future generations and forms a habitat for many
Long-tailed Broadbill building its nest
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.":745 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'
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
Main article: Ecological niche
Termite mounds with varied heights of chimneys regulate gas exchange,
temperature and other environmental parameters that are needed to
sustain the internal physiology of the entire colony.
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.":519 The ecological niche is a central concept in the
ecology of organisms and is sub-divided into the fundamental and the
realized niche. The fundamental niche is the set of environmental
conditions under which a species is able to persist. The realized
niche is the set of environmental plus ecological conditions under
which a species persists. 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.":71
Biogeographical patterns and range distributions are explained or
predicted through knowledge of a species' traits and niche
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 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, which is defined as
the full range of environmental and biological variables affecting an
Main article: Niche construction
Organisms are subject to environmental pressures, but they also modify
their habitats. The regulatory feedback between organisms and their
environment can affect conditions from local (e.g., a beaver pond) to
global scales, over time and even after death, such as decaying logs
or silica skeleton deposits from marine organisms. The process and
concept of ecosystem engineering is related to niche construction, 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
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, including ants, bees, wasps, and termites. There is an
emergent homeostasis 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 are 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.
Main article: 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, temperate broadleaf and mixed
forest, temperate deciduous forest, taiga, tundra, hot desert, and
polar desert. Other researchers have recently categorized other
biomes, such as the human and oceanic microbiomes. To a microbe, the
human body is a habitat and a landscape. Microbiomes were
discovered largely through advances in molecular genetics, 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.
Main article: Biosphere
See also: Earth's spheres
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
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 core temperature of the
Earth and atmospheric
conditions within a narrow self-regulating range of tolerance.
Life history theory, Ecophysiology, and Metabolic theory of
Understanding traits of individual organisms helps explain patterns
and processes at other levels of organization including populations,
communities, and ecosystems. Several areas of ecology of evolution
that focus on such traits are life history theory, ecophysiology,
metabolic theory of ecology, and Ethology. Examples of such traits
include features of an organisms life cycle such as age to maturity,
life span, or metabolic costs of reproduction. Other traits may be
related to structure, such as the spines of a cactus or dorsal spines
of a bluegill sunfish, or behaviors such as courtship displays or pair
bonding. Other traits include emergent properties that are the result
at least in part of interactions with the surrounding environment such
as growth rate, resource uptake rate, winter, and deciduous vs.
drought deciduous trees and shrubs.
One set of characteristics relate to body size and temperature. The
metabolic theory of ecology provides a predictive qualitative set of
relationships between an organism’s body size and temperature and
metabolic processes. In general, smaller, warmer organisms have higher
metabolic rates and this results in a variety of predictions regarding
individual somatic growth rates, reproduction and population growth
rates, population size, and resource uptake rates.
The traits of organisms are subject to change through acclimation,
development, and evolution. For this reason, individuals form a shared
focus for ecology and for evolutionary ecology.
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See also: Lists of organisms by population
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.":18 Simplified population models
usually start with four variables: death, birth, immigration, and
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
displaystyle frac operatorname d N operatorname d 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 modelling techniques, Malthus' population principle of
growth was later transformed into a model known as the logistic
displaystyle frac dN dT =aNleft(1- frac N K right),
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 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
Main article: Metapopulation
The concept of metapopulations was defined in 1969 as "a
population of populations which go extinct locally and
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 behaviour, stasis, and ranging. Dispersal is
usually distinguished from migration; because, it involves the one way
permanent movement of individuals from their birth population into
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 favourable.
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 favourable. Ecologists use a mixture
of computer models and field studies to explain metapopulation
Interspecific interactions such as predation are a key aspect of
Main article: Community ecology
Community ecology examines how interactions among species and their
environment affect the abundance, distribution and diversity of
species within communities.
Johnson & Stinchcomb (2007):250
Community ecology is the study of the interactions among a collections
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
These ecosystems, as we may call them, are of the most various kinds
and sizes. They form one category of the multitudinous physical
systems of the universe, which range from the universe as a whole down
to the atom.
A riparian forest in the White Mountains,
New Hampshire (USA) is an
example of ecosystem ecology
Ecosystems may be habitats within biomes that form an integrated whole
and a dynamically responsive system having both physical and
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).
attempt to determine the underlying causes of these fluxes. Research
in ecosystem ecology might measure primary production (g C/m^2) in a
wetland 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 and bacteria),
The underlying concept of ecosystem can be traced back to 1864 in the
published work of
George Perkins Marsh
George Perkins Marsh ("Man and Nature").
Within an ecosystem, organisms are linked to the physical and
biological components of their environment to which they are
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, atmospheric, or marine.
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.
Main article: Food web
See also: Food chain
Generalized food web of waterbirds from Chesapeake Bay
A food web is the archetypal ecological network. Plants capture solar
energy and use it to synthesize simple sugars during photosynthesis.
As plants grow, they accumulate nutrients and are eaten by grazing
herbivores, 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.
The larger interlocking pattern of food chains in an ecological
community creates a complex food web. Food webs are a type of concept
map or a heuristic 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 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) while some are more specialized with fewer
stronger feeding links (e.g., primary predators). Theoretical and
empirical studies identify non-random 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.
Main article: Trophic level
A trophic pyramid (a) and a food-web (b) illustrating ecological
relationships among creatures that are typical of a northern boreal
terrestrial ecosystem. The trophic pyramid roughly represents the
biomass (usually measured as total dry-weight) at each level. Plants
generally have the greatest biomass. Names of trophic categories are
shown to the right of the pyramid. Some ecosystems, such as many
wetlands, do not organize as a strict pyramid, because aquatic plants
are not as productive as long-lived terrestrial plants such as trees.
Ecological trophic pyramids are typically one of three kinds: 1)
pyramid of numbers, 2) pyramid of biomass, or 3) pyramid of
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 adjacent level nearer the abiotic
source.":383 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 consumers), and Detritivores (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 (strict herbivores), secondary consumers
(carnivorous predators that feed exclusively on herbivores), and
tertiary consumers (predators that feed on a mix of herbivores and
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
Trophic levels are part of the holistic or complex systems 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.":815 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
Main article: Keystone species
Sea otters, an example of a keystone species
A keystone species is a species that is connected to a
disproportionately large number of other species in the food-web.
Keystone species have lower levels of biomass in the trophic pyramid
relative to the importance of their role. The many connections that a
keystone species holds means that it maintains the organization and
structure of entire communities. The loss of a keystone species
results in a range of dramatic cascading effects that alters trophic
dynamics, other food web connections, and can cause the extinction of
Sea otters (Enhydra lutris) are commonly cited as an example of a
keystone species; because, they limit the density of sea urchins that
feed on kelp. If sea otters are removed from the system, the urchins
graze until the kelp beds disappear, and this has a dramatic effect on
community structure. Hunting of sea otters, for example, is
thought to have led indirectly to the extinction of the Steller's sea
cow (Hydrodamalis gigas). While the keystone species concept has
been used extensively as a conservation tool, it has been criticized
for being poorly defined from an operational stance. It is 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.
Main article: Complexity
See also: Emergence
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 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 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.":209 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'.[E]
Complexity in ecology is of at least six distinct types: spatial,
temporal, structural, process, behavioral, and geometric.":3 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
Main article: 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, 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
Ecological studies are necessarily holistic as opposed to
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 hierarchy whereby the causal
relations of larger systems are understood without reference to the
smaller parts. Scientific holism differs from mysticism 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 need to reference or understand the biomolecular
properties of the exterior shells.
Relation to evolution
Main article: Evolutionary ecology
Ecology and evolutionary biology are considered sister disciplines of
the life sciences. Natural selection, life history, development,
adaptation, populations, and inheritance are examples of concepts that
thread equally into ecological and evolutionary theory. Morphological,
behavioural, and genetic traits, for example, can be mapped onto
evolutionary trees to study the historical development of a species in
relation to their functions and roles in different ecological
circumstances. In this framework, the analytical tools of ecologists
and evolutionists overlap as they organize, classify, and investigate
life through common systematic principals, such as phylogenetics or
the Linnaean system of taxonomy. The two disciplines often appear
together, such as in the title of the journal Trends in
Evolution. 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
Main article: Behavioural ecology
Social display and colour variation in differently adapted species of
chameleons (Bradypodion spp.).
Chameleons change their skin colour to
match their background as a behavioural defence mechanism and also use
colour to communicate with other members of their species, such as
dominant (left) versus submissive (right) patterns shown in the three
species (A-C) above.
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 of plants, mobile phytoplankton,
zooplankton swimming toward the female egg, the cultivation of fungi
by weevils, the mating dance of a salamander, or social gatherings of
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, "[t]he 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 "[t]he optimal
flight initiation distance occurs where expected postencounter fitness
is maximized, which depends on the prey's initial fitness, benefits
obtainable by not fleeing, energetic escape costs, and expected
fitness loss due to predation risk."
Symbiosis: Leafhoppers (Eurymela fenestrata) are protected by ants
(Iridomyrmex purpureus) in a symbiotic relationship. The ants protect
the leafhoppers from predators and in return the leafhoppers feeding
on plants exude honeydew from their anus that provides energy and
nutrients to tending ants.
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. These displays
serve a dual purpose of signalling healthy or well-adapted individuals
and desirable genes. The displays are driven by sexual selection as an
advertisement of quality of traits among suitors.
Cognitive ecology integrates theory and observations from evolutionary
ecology and neurobiology, 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
Main article: Social ecology
Social ecological behaviours are notable in the social insects, slime
moulds, social spiders, human society, and naked mole-rats where
eusocialism has evolved. Social behaviours include reciprocally
beneficial behaviours among kin and nest mates and
evolve from kin and group selection.
Kin selection explains altruism
through genetic relationships, whereby an altruistic behaviour leading
to death is rewarded by the survival of genetic copies distributed
among surviving relatives. The social insects, including ants, bees,
and wasps are most famously studied for this type of relationship
because the male drones are clones that share the same genetic make-up
as every other male in the colony. In contrast, group
selectionists find examples of altruism among non-genetic relatives
and explain this through selection acting on the group; whereby, it
becomes selectively advantageous for groups if their members express
altruistic behaviours to one another. Groups with predominantly
altruistic members beat groups with predominantly selfish
Main article: Coevolution
Bumblebees and the flowers they pollinate have coevolved so that both
have become dependent on each other for survival.
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 within a species that
are mutually or reciprocally beneficial are called mutualisms.
Examples of mutualism include fungus-growing ants employing
agricultural symbiosis, bacteria living in the guts of insects and
other organisms, the fig wasp and yucca moth pollination complex,
lichens with fungi and photosynthetic algae, and corals with
photosynthetic algae. If there is a physical connection
between host and associate, the relationship is called symbiosis.
Approximately 60% of all plants, for example, have a symbiotic
relationship with arbuscular mycorrhizal fungi living in their roots
forming an exchange network of carbohydrates for mineral
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;
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.
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
Parasitism: A harvestman arachnid being parasitized by mites. The
harvestman is being consumed, while the mites benefit from traveling
on and feeding off of their host.
Main article: 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
Biogeography was established in 1974.
ecology share many of their disciplinary roots. For example, the
theory of island biogeography, published by the
Robert MacArthur and
Edward O. Wilson
Edward O. Wilson in 1967 is considered one of the fundamentals of
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 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 explains much of the modern distribution
of the Earth's biota. The splitting of lineages in a species is called
vicariance biogeography and it is a 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.
r/K selection theory
Main article: r/K selection theory
A population ecology concept is r/K selection theory,[D] one of the
first predictive models in ecology used to explain life-history
evolution. The premise behind the r/K selection model is that natural
selection pressures change according to population density. For
example, when an island is first colonized, density of individuals is
low. The initial increase in population size is not limited by
competition, leaving an abundance of available resources for rapid
population growth. These early phases of population growth experience
density-independent forces of natural selection, which is called
r-selection. As the population becomes more crowded, it approaches the
island's carrying capacity, thus forcing individuals to compete more
heavily for fewer available resources. Under crowded conditions, the
population experiences density-dependent forces of natural selection,
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 in r-selected species. Many kinds of insects and invasive
species exhibit r-selected characteristics. In contrast, a K-selected
species has low rates of fecundity, high levels of parental investment
in the young, and low rates of mortality as individuals mature. Humans
and elephants are examples of species exhibiting K-selected
characteristics, including longevity and efficiency in the conversion
of more resources into fewer offspring.
Main article: Molecular ecology
The important relationship between ecology and genetic inheritance
predates modern techniques for molecular analysis. Molecular
ecological research became more feasible with the development of rapid
and accessible genetic technologies, such as the polymerase chain
reaction (PCR). The rise of molecular technologies and influx of
research questions into this new ecological field resulted in the
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
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. In a biogeographical context, the marriage
between genetics, ecology, and evolution resulted in a new
sub-discipline called phylogeography.
Main article: Human ecology
The history of life on
Earth has been a history of interaction between
living things and their surroundings. To a large extent, the physical
form and the habits of the earth's vegetation and its animal life have
been molded by the environment. Considering the whole span of earthly
time, the opposite effect, in which life actually modifies its
surroundings, has been relatively slight. Only within the moment of
time represented by the present century has one species man acquired
significant power to alter the nature of his world.
Rachel Carson, "Silent Spring"
Ecology is as much a biological science as it is a human science.
Human ecology is an interdisciplinary 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.":3 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 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-natural capital, 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.
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.
Restoration and management
Main article: Restoration ecology
Natural resource management
Ecosystem management is not just about science nor is it simply an
extension of traditional resource management; it offers a fundamental
reframing of how humans may work with nature.
Ecology is an employed science of restoration, repairing disturbed
sites through human intervention, in natural resource management, and
in environmental impact assessments.
Edward O. Wilson
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, for
example, employ ecologists to develop, adapt, and implement ecosystem
based methods into the planning, operation, and restoration phases of
land-use. 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
Relation to the environment
Main article: Natural environment
The environment of ecosystems includes both physical parameters and
biotic attributes. It is dynamically interlinked, and contains
resources 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.":62 The physical
environment is external to the level of biological organization under
investigation, including abiotic factors such as temperature,
radiation, light, chemistry, climate and geology. The biotic
environment includes genes, cells, organisms, members of the same
species (conspecifics) and other species that share a habitat.
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, 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 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 approach to ecology. The dialectical approach examines
the parts, but integrates the organism and the environment into a
dynamic whole (or umwelt). Change in one ecological or environmental
factor can concurrently affect the dynamic state of an entire
Disturbance and resilience
Main article: Resilience (ecology)
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
Metabolism and the early atmosphere
Metabolism – the rate at which energy and material resources are
taken up from the environment, transformed within an organism, and
allocated to maintenance, growth and reproduction – is a fundamental
Ernest et al.:991
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 to one composed mostly of methane and
ammonia. Over the next billion years, the metabolic activity of life
transformed the atmosphere into a mixture of carbon dioxide, nitrogen,
and water vapor. These gases changed the way that light from the sun
hit the Earth's surface and greenhouse effects trapped heat. There
were untapped sources of free energy within the mixture of reducing
and oxidizing gasses that set the stage for primitive ecosystems to
evolve and, in turn, the atmosphere also evolved.
The leaf is the primary site of photosynthesis in most plants.
Throughout history, the Earth's atmosphere and biogeochemical cycles
have been in a dynamic equilibrium with planetary ecosystems. The
history is characterized by periods of significant transformation
followed by millions of years of stability. The evolution of the
earliest organisms, likely anaerobic methanogen microbes, started the
process by converting atmospheric hydrogen into methane (4H2 + CO2 →
CH4 + 2H2O).
Anoxygenic photosynthesis reduced hydrogen concentrations
and increased atmospheric methane, by converting hydrogen sulfide into
water or other sulfur compounds (for example, 2H2S + CO2 + hv → CH2O
+ H2O + 2S). Early forms of fermentation also increased levels of
atmospheric methane. The transition to an oxygen-dominant atmosphere
(the Great Oxidation) did not begin until approximately
2.4–2.3 billion years ago, but photosynthetic processes started
0.3 to 1 billion years prior.
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. Temperature
is largely dependent on the incidence of solar radiation. The
latitudinal and longitudinal spatial variation of temperature greatly
affects climates and consequently the distribution of biodiversity and
levels of primary production in different ecosystems or biomes across
the planet. Heat and temperature relate importantly to metabolic
activity. Poikilotherms, for example, have a body temperature that is
largely regulated and dependent on the temperature of the external
environment. In contrast, homeotherms regulate their internal body
temperature by expending metabolic energy.
There is a relationship between light, primary production, and
ecological energy budgets.
Sunlight is the primary input of energy
into the planet's ecosystems. Light is composed of electromagnetic
energy of different wavelengths.
Radiant energy from the sun generates
heat, provides photons of light measured as active energy in the
chemical reactions of life, and also acts as a catalyst for genetic
mutation. Plants, algae, and some bacteria absorb light
and assimilate the energy through photosynthesis.
Organisms capable of
assimilating energy by photosynthesis or through inorganic fixation of
H2S are autotrophs.
Autotrophs — responsible for primary production
— assimilate light energy which becomes metabolically stored as
potential energy in the form of biochemical enthalpic
Main article: Aquatic ecosystem
Wetland conditions such as shallow water, high plant productivity, and
anaerobic substrates provide a suitable environment for important
physical, biological, and chemical processes. Because of these
processes, wetlands play a vital role in global nutrient and element
Cronk & Fennessy (2001):29
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 where anaerobic
bacteria 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) 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,
brackish, or oceanic environments. Anaerobic soil microorganisms in
aquatic environments use nitrate, manganese ions, ferric ions,
sulfate, carbon dioxide, 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 gradients that mediate salt excretion in salt water
and uptake in fresh water.
The shape and energy of the land is 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 as well as influencing geomorphic
processes such as orogeny and erosion. 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
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 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).
Climatic and osmotic pressure places physiological 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 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 whales, dolphins, and seals are specially adapted to
deal with changes in sound due to water pressure differences.
Differences between hagfish species provide another example of
adaptation to deep-sea pressure through specialized protein
Wind and turbulence
The architecture of the inflorescence in grasses is subject to the
physical pressures of wind and shaped by the forces of natural
selection facilitating wind-pollination (anemophily).
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.
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
and influencing the environmental profile to create thermally layered
zones, affecting how fish, algae, and other parts of the aquatic
ecosystem are structured.
Wind speed and turbulence also
influence evapotranspiration rates and energy budgets in plants and
Wind speed, temperature and moisture content can
vary as winds travel across different land features and elevations.
For example, the westerlies come into contact with the coastal and
interior mountains of western
North America to produce a rain shadow
on the leeward side of the mountain. The air expands and moisture
condenses as the winds increase in elevation; this is called
orographic lift and can cause precipitation.[clarification needed]
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
ecosystems (e.g., of the Columbia Basin in western North America) to
intermix with sister lineages that are segregated to the interior
Main article: Fire ecology
Forest fires modify the land by leaving behind an environmental mosaic
that diversifies the landscape into different seral stages and
habitats of varied quality (left). Some species are adapted to forest
fires, such as pine trees that open their cones only after 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), 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
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 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 of ecosystems.
Main article: Soil ecology
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 of dead organic matter (for
example, leaves on the forest floor), results in soils containing
minerals 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 detritivores 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. This
aerates soils and stimulates heterotrophic growth and production. Soil
microorganisms are influenced by and feed 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 in the
Devonian period played a significant role in the early development of
ecological trophism in soils.
Biogeochemistry and climate
Main article: Biogeochemistry
Nutrient cycle and Climate
Ecologists study and measure nutrient budgets to understand how these
materials are regulated, flow, and recycled 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, carbon, nitrogen,
oxygen, sulfur, and phosphorus; 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 cycles 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, 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, from twenty-five to thirty-two million years ago,
there was another significant restructuring of the global carbon cycle
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, 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 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 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
Main article: History of ecology
By ecology, we mean the whole science of the relations of the organism
to the environment including, in the broad sense, all the "conditions
of existence. "Thus the theory of evolution explains the housekeeping
relations of organisms mechanistically as the necessary consequences
of effectual causes and so forms the monistic groundwork of ecology.
Ernst Haeckel (1866):140 [B]
Ecology has a complex origin, due in large part to its
interdisciplinary nature. Ancient Greek philosophers such as
Aristotle were among the first to record observations
on natural history. However, they viewed life in terms of
essentialism, 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. Early conceptions of ecology, such as a
balance and regulation in nature can be traced to
Herodotus (died c.
425 BC), who described one of the earliest accounts of mutualism in
his observation of "natural dentistry". Basking Nile crocodiles, he
noted, would open their mouths to give sandpipers safe access to pluck
leeches out, giving nutrition to the sandpiper and oral hygiene for
Aristotle was an early influence on the
philosophical development of ecology. He and his student Theophrastus
made extensive observations on plant and animal migrations,
biogeography, physiology, and on their behaviour, giving an early
analogue to the modern concept of an ecological niche.
Nowhere can one see more clearly illustrated what may be called the
sensibility of such an organic complex,--expressed by the fact that
whatever affects any species belonging to it, must speedily have its
influence of some sort upon the whole assemblage. He will thus be made
to see the impossibility of studying any form completely, out of
relation to the other forms,--the necessity for taking a comprehensive
survey of the whole as a condition to a satisfactory understanding of
Stephen Forbes (1887)
Ernst Haeckel (left) and
Eugenius Warming (right), two founders of
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
Antoni van Leeuwenhoek (1632–1723) and
botanist Richard Bradley (1688?–1732). Biogeographer Alexander
von Humboldt (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
Isaac Newton 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, and
Jean-Baptiste Lamarck (among others) laid the
foundations of the modern ecological sciences. The term "ecology"
(German: Oekologie, Ökologie) is of a more recent origin and was
first coined by the German biologist
Ernst Haeckel in his book
Generelle Morphologie der Organismen (1866). Haeckel was a zoologist,
artist, writer, and later in life a professor of comparative
Opinions differ on who was the founder of modern ecological theory.
Some mark Haeckel's definition as the beginning; others say it
Eugenius Warming with the writing of Oecology of Plants: An
Introduction to the Study of
Plant Communities (1895), or Carl
Linnaeus' 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 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.
The layout of the first ecological experiment, carried out in a grass
Woburn Abbey in 1816, was noted by
Charles Darwin in The
Origin of Species. The experiment studied the performance of different
mixtures of species planted in different kinds of soils.
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 (1720–1793), considered by some to be
one of the earliest texts on ecology. While
Charles Darwin is
mainly noted for his treatise on evolution, he was one of the
founders of soil ecology, and he made note of the first
ecological experiment in The Origin of Species. Evolutionary
theory changed the way that researchers approached the ecological
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).
Ellen Swallow Richards
Ellen Swallow Richards may have first introduced the
term "oekology" (which eventually morphed into home economics) in the
U.S. as early 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 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, a South African general and
polarizing historical figure who was inspired by Clements'
superorganism concept.[C] Around the same time, Charles Elton
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 J. Lotka brought in many
theoretical concepts applying thermodynamic principles to ecology.
Raymond Lindeman wrote a landmark paper on the trophic
dynamics 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
Ecology also has developed through
contributions from other nations, including Russia's Vladimir
Vernadsky and his founding of the biosphere concept in the 1920s
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.
This whole chain of poisoning, then, seems to rest on a base of minute
plants which must have been the original concentrators. But what of
the opposite end of the food chain—the human being who, in probable
ignorance of all this sequence of events, has rigged his fishing
tackle, caught a string of fish from the waters of Clear Lake, and
taken them home to fry for his supper?
Rachel Carson (1962):48
Ecology surged in popular and scientific interest during the
1960–1970s environmental movement. 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, such as Aldo
Leopold and Arthur Tansley, 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) and brought about changes in environmental
legislation. Women such as
Ellen Swallow Richards
Ellen Swallow Richards and Julia Lathrop,
among others, were precursors to the more popularized environmental
movements after the 1950s.
In 1962, marine biologist and ecologist Rachel Carson's book Silent
Spring helped to mobilize the environmental movement by alerting the
public to toxic pesticides, 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.
Main article: Outline of ecology
Circles of Sustainability
Glossary of ecology
Index of biology articles
List of ecologists
Outline of biology
Terminology of ecology
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