Ecosystem ecology
Encyclopedia
Ecosystem ecology is the integrated study of biotic
Life
Life is a characteristic that distinguishes objects that have signaling and self-sustaining processes from those that do not, either because such functions have ceased , or else because they lack such functions and are classified as inanimate...

 and abiotic components of ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....

s and their interactions within an ecosystem framework. This science
Science
Science is a systematic enterprise that builds and organizes knowledge in the form of testable explanations and predictions about the universe...

 examines how ecosystems work and relates this to their components such as chemicals, bedrock
Bedrock
In stratigraphy, bedrock is the native consolidated rock underlying the surface of a terrestrial planet, usually the Earth. Above the bedrock is usually an area of broken and weathered unconsolidated rock in the basal subsoil...

, soil
Soil
Soil is a natural body consisting of layers of mineral constituents of variable thicknesses, which differ from the parent materials in their morphological, physical, chemical, and mineralogical characteristics...

, plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s, and animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s.

Ecosystem ecology examines physical and biological structures and examines how these ecosystem characteristics interact with each other. Ultimately, this helps us understand how to maintain high quality water and economically viable commodity production. A major focus of ecosystem ecology is on functional processes, ecological mechanisms that maintain the structure and services produced by ecosystems. These include primary productivity (production of biomass
Biomass (ecology)
Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...

), decomposition
Decomposition
Decomposition is the process by which organic material is broken down into simpler forms of matter. The process is essential for recycling the finite matter that occupies physical space in the biome. Bodies of living organisms begin to decompose shortly after death...

, and trophic
Trophic level
The trophic level of an organism is the position it occupies in a food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism...

 interactions.

Studies of ecosystem function have greatly improved human understanding of sustainable production of forage
Fodder
Fodder or animal feed is any agricultural foodstuff used specifically to feed domesticated livestock such as cattle, goats, sheep, horses, chickens and pigs. Most animal feed is from plants but some is of animal origin...

, fiber
Fiber
Fiber is a class of materials that are continuous filaments or are in discrete elongated pieces, similar to lengths of thread.They are very important in the biology of both plants and animals, for holding tissues together....

, fuel
Fuel
Fuel is any material that stores energy that can later be extracted to perform mechanical work in a controlled manner. Most fuels used by humans undergo combustion, a redox reaction in which a combustible substance releases energy after it ignites and reacts with the oxygen in the air...

, and provision of water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

. Functional processes are mediated by regional-to-local level climate
Climate
Climate encompasses the statistics of temperature, humidity, atmospheric pressure, wind, rainfall, atmospheric particle count and other meteorological elemental measurements in a given region over long periods...

, disturbance
Disturbance
In ecology, a disturbance is a temporary change in average environmental conditions that causes a pronounced change in an ecosystem. Outside disturbance forces often act quickly and with great effect, sometimes resulting in the removal of large amounts of biomass...

, and management. Thus ecosystem ecology provides a powerful framework for identifying ecological mechanisms that interact with global environmental problems, especially global warming
Global warming
Global warming refers to the rising average temperature of Earth's atmosphere and oceans and its projected continuation. In the last 100 years, Earth's average surface temperature increased by about with about two thirds of the increase occurring over just the last three decades...

 and degradation of surface water.

This example demonstrates several important aspects of ecosystems:
  1. Ecosystem boundaries are often nebulous and may fluctuate in time
  2. Organisms within ecosystems are dependent on ecosystem level biological and physical processes
  3. Adjacent ecosystems closely interact and often are interdependent for maintenance of community structure and functional processes that maintain productivity and biodiversity
    Biodiversity
    Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...



These characteristics also introduce practical problems into natural resource management. Who will manage which ecosystem? Will timber cutting in the forest degrade recreational fishing in the stream? These questions are difficult for land managers to address while the boundary between ecosystems remains unclear; even though decisions in one ecosystem will affect the other. We need better understanding of the interactions and interdependencies of these ecosystems and the processes that maintain them before we can begin to address these questions.

Ecosystem ecology is an inherently interdisciplinary field of study. An individual ecosystem is composed of populations of organisms, interacting within communities, and contributing to the cycling of nutrients and the flow of energy
Energy
In physics, energy is an indirectly observed quantity. It is often understood as the ability a physical system has to do work on other physical systems...

. The ecosystem is the principal unit of study in ecosystem ecology.

Population, community, and physiological ecology provide many of the underlying biological mechanisms influencing ecosystems and the processes they maintain. Cycling of energy and matter at the ecosystem level are often examined in ecosystem ecology, but, as a whole, this science is defined more by subject matter than by scale. Ecosystem ecology approaches organisms and abiotic pools of energy and nutrients as an integrated system which distinguishes it from associated sciences such as biogeochemistry
Biogeochemistry
Biogeochemistry is the scientific discipline that involves the study of the chemical, physical, geological, and biological processes and reactions that govern the composition of the natural environment...

.

Biogeochemistry and hydrology
Hydrology
Hydrology is the study of the movement, distribution, and quality of water on Earth and other planets, including the hydrologic cycle, water resources and environmental watershed sustainability...

 focus on several fundamental ecosystem processes such as biologically mediated chemical cycling of nutrients and physical-biological cycling of water. Ecosystem ecology forms the mechanistic basis for regional or global processes encompassed by landscape-to-regional hydrology, global biogeochemistry, and earth system science.

History

Ecosystem ecology is philosophically and historically rooted in terrestrial ecology. The ecosystem concept has evolved rapidly during the last 100 years with important ideas developed by Frederic Clements
Frederic Clements
Frederic Edward Clements was an American plant ecologist and pioneer in the study of vegetation succession.-Biography:...

, a botanist who argued for specific definitions of ecosystems and that physiological processes were responsible for their development and persistence. Although most of Clements ecosystem definitions have been greatly revised, initially by Henry Gleason
Henry Gleason
Henry Allan Gleason was a noted American ecologist, botanist, and taxonomist, most recognized for his endorsement of the individualistic/open community concept of ecological succession.- Life and work :...

 and Arthur Tansley
Arthur Tansley
Sir Arthur George Tansley FRS was an English botanist who was a pioneer in the science of ecology. He obtained his degree in Biological Science in 1896, with specialization in botany and zoology. From the start, he was much influenced by the Danish plant ecologist Eugenius Warming. He championed...

, and later by contemporary ecologists, the idea that physiological processes are fundamental to ecosystem structure and function remains central to ecology.

Later work by Eugene Odum
Eugene Odum
Eugene Pleasants Odum was an American scientist known for his pioneering work on ecosystem ecology. He wrote the first ecology textbook: Fundamentals of Ecology....

 and Howard T. Odum
Howard T. Odum
Howard Thomas Odum was an American ecologist...

 quantified flows of energy and matter at the ecosystem level, thus documenting the general ideas proposed by Clements and his contemporary Charles Elton
Charles Sutherland Elton
Charles Sutherland Elton FRS was an English zoologist and animal ecologist. His name is associated with the establishment of modern population and community ecology, including studies of invasive organisms.-Personal life:...

. See Figure 3.

In this model, energy flows through the whole system were dependent on biotic and abiotic interactions of each individual component (species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

, inorganic pools of nutrients, etc.). Later work demonstrated that these interactions and flows applied to nutrient cycles, changed over the course of succession
Ecological succession
Ecological succession, is the phenomenon or process by which a community progressively transforms itself until a stable community is formed. It is a fundamental concept in ecology, and refers to more or less predictable and orderly changes in the composition or structure of an ecological community...

, and held powerful controls over ecosystem productivity. Transfers of energy and nutrients are innate to ecological systems regardless of whether they are aquatic or terrestrial. Thus, ecosystem ecology has emerged from important biological studies of plants, animals, terrestrial, aquatic
Aquatic ecosystem
An aquatic ecosystem is an ecosystem in a body of water. Communities of organisms that are dependent on each other and on their environment live in aquatic ecosystems. The two main types of aquatic ecosystems are marine ecosystems and freshwater ecosystems....

, and marine
Marine (ocean)
Marine is an umbrella term. As an adjective it is usually applicable to things relating to the sea or ocean, such as marine biology, marine ecology and marine geology...

 ecosystems.

Ecosystem services

Ecosystem services are ecologically mediated functional processes essential to sustaining healthy human
Human
Humans are the only living species in the Homo genus...

 societies. Water provision and filtration, production of biomass
Biomass (ecology)
Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...

 in forestry
Forestry
Forestry is the interdisciplinary profession embracing the science, art, and craft of creating, managing, using, and conserving forests and associated resources in a sustainable manner to meet desired goals, needs, and values for human benefit. Forestry is practiced in plantations and natural stands...

, agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

, and fisheries, and removal of greenhouse gas
Greenhouse gas
A greenhouse gas is a gas in an atmosphere that absorbs and emits radiation within the thermal infrared range. This process is the fundamental cause of the greenhouse effect. The primary greenhouse gases in the Earth's atmosphere are water vapor, carbon dioxide, methane, nitrous oxide, and ozone...

es such as carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 (CO2) from the atmosphere
Atmosphere
An atmosphere is a layer of gases that may surround a material body of sufficient mass, and that is held in place by the gravity of the body. An atmosphere may be retained for a longer duration, if the gravity is high and the atmosphere's temperature is low...

 are examples of ecosystem services essential to public health
Public health
Public health is "the science and art of preventing disease, prolonging life and promoting health through the organized efforts and informed choices of society, organizations, public and private, communities and individuals" . It is concerned with threats to health based on population health...

 and economic opportunity. Nutrient cycling is a process fundamental to agricultural and forest production.

However, like most ecosystem processes, nutrient cycling is not an ecosystem characteristic which can be “dialed” to the most desirable level. Maximizing production in degraded systems is an overly simplistic solution to the complex problems of hunger and economic security. For instance, intensive fertilizer
Fertilizer
Fertilizer is any organic or inorganic material of natural or synthetic origin that is added to a soil to supply one or more plant nutrients essential to the growth of plants. A recent assessment found that about 40 to 60% of crop yields are attributable to commercial fertilizer use...

 use in the midwestern United States has resulted in degraded fisheries in the Gulf of Mexico
Gulf of Mexico
The Gulf of Mexico is a partially landlocked ocean basin largely surrounded by the North American continent and the island of Cuba. It is bounded on the northeast, north and northwest by the Gulf Coast of the United States, on the southwest and south by Mexico, and on the southeast by Cuba. In...

. Regrettably, a “Green Revolution
Green Revolution
Green Revolution refers to a series of research, development, and technology transfer initiatives, occurring between the 1940s and the late 1970s, that increased agriculture production around the world, beginning most markedly in the late 1960s....

” of intensive chemical fertilization has been recommended for agriculture in developed and developing countries. These strategies risk alteration of ecosystem processes that may be difficult to restore, especially when applied at broad scales without adequate assessment of impacts. Ecosystem processes may take many years to recover from significant disturbance.

For instance, large-scale forest clearance in the northeastern United States during the 18th and 19th centuries has altered soil texture, dominant vegetation, and nutrient cycling in ways that impact forest productivity in the present day. An appreciation of the importance of ecosystem function in maintenance of productivity, whether in agriculture or forestry, is needed in conjunction with plans for restoration of essential processes. Improved knowledge of ecosystem function will help to achieve long-term sustainability and stability in the poorest parts of the world.

How do ecosystems work?

Biomass productivity is one of the most apparent and economically important ecosystem functions. Biomass accumulation begins at the cellular level via photosynthesis. Photosynthesis requires water and consequently global patters of annual biomass production are correlated with annual precipitation. Amounts of productivity are also dependent on the overall capacity of plants to capture sunlight which is directly correlated with plant leaf area and N content.

Net primary productivity
Primary production
400px|thumb|Global oceanic and terrestrial photoautotroph abundance, from September [[1997]] to August 2000. As an estimate of autotroph biomass, it is only a rough indicator of primary production potential, and not an actual estimate of it...

 (NPP) is the primary measure of biomass accumulation within an ecosystem. Net primary productivity can be calculated by a simple formula where the total amount of productivity is adjusted for total productivity losses through maintenance of biological processes:
NPP = GPP – Rproducer


Where GPP is gross primary productivity and Rproducer is photosynthate (Carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

) lost via cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

.

NPP is difficult to measure but a new technique known as eddy co-variance has shed light on how natural ecosystems influence the atmosphere. Figure 4 shows seasonal and annual changes in CO2 concentration measured at Mauna Loa
Mauna Loa
Mauna Loa is one of five volcanoes that form the Island of Hawaii in the U.S. state of Hawaii in the Pacific Ocean, and the largest on Earth in terms of volume and area covered. It is an active shield volcano, with a volume estimated at approximately , although its peak is about lower than that...

, Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...

 from 1987 to 1990. CO2 concentration steadily increased, but within-year variation has been greater than the annual increase since measurements began in 1957.

These variations were thought to be due to seasonal uptake of CO2 during summer months. A newly developed technique for assessing ecosystem NPP has confirmed seasonal variation are driven by seasonal changes in CO2 uptake by vegetation. This has led many scientists and policy makers to speculate that ecosystems can be managed to ameliorate problems with global warming
Global warming
Global warming refers to the rising average temperature of Earth's atmosphere and oceans and its projected continuation. In the last 100 years, Earth's average surface temperature increased by about with about two thirds of the increase occurring over just the last three decades...

. This type of management may include reforesting or altering forest harvest schedules many parts of the world.

Decomposition and nutrient cycling

Decomposition
Decomposition
Decomposition is the process by which organic material is broken down into simpler forms of matter. The process is essential for recycling the finite matter that occupies physical space in the biome. Bodies of living organisms begin to decompose shortly after death...

 and nutrient cycling are fundamental to ecosystem biomass production. Most natural ecosystems are nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

 (N) limited and biomass production is closely correlated with N turnover.
Typically external input of nutrients is very low and efficient recycling of nutrients maintains productivity. Decomposition of plant litter accounts for the majority of nutrients recycled through ecosystems (Figure 3). Rates of plant litter decomposition are highly dependent on litter quality; high concentration of phenolic compounds, especially lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...

, in plant litter has a retarding effect on litter decomposition. More complex C compounds are decomposed more slowly and may take many years to completely breakdown. Decomposition is typically described with exponential decay.

Globally, rates of decomposition are mediated by litter quality and climate. Ecosystems dominated by plants with low-lignin concentration often have rapid rates of decomposition and nutrient cycling (Chapin et al. 1982). Simple carbon (C) containing compounds are preferentially metabolized by decomposer
Decomposer
Decomposers are organisms that break down dead or decaying organisms, and in doing so carry out the natural process of decomposition. Like herbivores and predators, decomposers are heterotrophic, meaning that they use organic substrates to get their energy, carbon and nutrients for growth and...

 microorganisms which results in rapid initial rates of decomposition, see Figure 5A, models that depend on constant rates of decay; so called “k” values, see Figure 5B.

However, these models do not reflect simultaneous linear and non-linear decay processes which likely occur during decomposition. For instance, proteins, sugars and lipids decompose exponentially, but lignin decays at a more linear rate Thus, litter decay is inaccurately predicted by simplistic models.

A simple alternative model presented in Figure 5C shows significantly more rapid decomposition that the standard model of figure 4B. Better understanding of decomposition models is an important research area of ecosystem ecology because this process is closely tied to nutrient supply and the overall capacity of ecosystems to sequester CO2 from the atmosphere.

Trophic dynamics

Trophic dynamics refers to process of energy and nutrient
Nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...

 transfer between organisms. Trophic dynamics is an important part of the structure and function of ecosystems. Figure 3 shows energy transferred for an ecosystem at Silver Springs, Florida. Energy gained by primary producers (plants, P) is consumed by herbivores (H), which are consumed by carnivores (C), which are themselves consumed by “top- carnivores”(TC).

One of the most obvious patterns in Figure 3 is that as one moves up to higher trophic level
Trophic level
The trophic level of an organism is the position it occupies in a food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism...

s (i.e. from plants to top-carnivores) the total amount of energy decreases. Plants exert a “bottom-up” control on the energy structure of ecosystems by determining the total amount of energy that enters the system.

However, predators can also influence the structure of lower trophic levels from the top-down. These influences can dramatically shift dominant species in terrestrial and marine systems The interplay and relative strength of top-down vs. bottom-up controls on ecosystem structure and function is an important area of research in the greater field of ecology.

Trophic dynamics can strongly influence rates of decomposition and nutrient cycling in time and in space. For example, herbivory can increase litter decomposition and nutrient cycling via direct changes in litter quality and altered dominant vegetation. Insect herbivory has been shown to increase rates of decomposition and nutrient turnover due to changes in litter quality and increased frass
Frass
Frass is the fine powdery material phytophagous insects pass as waste after digesting plant parts. It causes plants to excrete chitinase due to high chitin levels, it is a natural bloom stimulant, and has high nutrient levels. Frass is known to have abundant amoeba, beneficial bacteria, and fungi...

 inputs

However, insect outbreak does not always increase nutrient cycling. Stadler showed that C rich honeydew produced during aphid outbreak can result in increased N immobilization by soil microbes thus slowing down nutrient cycling and potentially limiting biomass production. North atlantic marine ecosystems have been greatly altered by overfishing of cod. Cod stocks crashed in the 1990s which resulted in increases in their prey such as shrimp and snow crab Human intervention in ecosystems has resulted in dramatic changes to ecosystem structure and function. These changes are occurring rapidly and have unknown consequences for economic security and human well-being.

Lessons from two Central American cities

The biosphere has been greatly altered by the demands of human societies. Ecosystem ecology plays an important role in understanding and adapting to the most pressing current environmental problems. Restoration ecology and ecosystem management are closely associated with ecosystem ecology. Restoring highly degraded resources depends on integration of functional mechanisms of ecosystems.

Without these functions intact, economic value of ecosystems is greatly reduced and potentially dangerous conditions may develop in the field. For example, areas within the mountainous western highlands of Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...

 are more susceptible to catastrophic landslides and crippling seasonal water shortages due to loss of forest resources. In contrast, cities such as Totonicapán
Totonicapán
Totonicapán is a city in Guatemala. It serves as the capital of the department of Totonicapán and as the administrative seat for the surrounding municipality of Totonicapán.- External links :# #...

 that have preserved forests through strong social institutions have greater local economic stability and overall greater human well-being.

This situation is striking considering that these areas are close to each other, the majority of inhabitants are of Mayan
Maya civilization
The Maya is a Mesoamerican civilization, noted for the only known fully developed written language of the pre-Columbian Americas, as well as for its art, architecture, and mathematical and astronomical systems. Initially established during the Pre-Classic period The Maya is a Mesoamerican...

 descent, and the topography and overall resources are similar. This is a case of two groups of people managing resources in fundamentally different ways. Ecosystem ecology provides the basic science needed to avoid degradation and to restore ecosystem processes that provide for basic human needs.

See also

  • Biogeochemistry
    Biogeochemistry
    Biogeochemistry is the scientific discipline that involves the study of the chemical, physical, geological, and biological processes and reactions that govern the composition of the natural environment...

  • Community ecology
  • Landscape ecology
    Landscape ecology
    Landscape ecology is the science of studying and improving relationships between urban development and ecological processes in the environment and particular ecosystems...

  • Systems ecology
    Systems ecology
    Systems ecology is an interdisciplinary field of ecology, taking a holistic approach to the study of ecological systems, especially ecosystems. Systems ecology can be seen as an application of general systems theory to ecology. Central to the systems ecology approach is the idea that an ecosystem...

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