Limnology
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Limnology ( ; from Greek λίμνη, ''limne'', "lake" and λόγος, ''logos'', "knowledge") is the study of inland aquatic ecosystems. The study of limnology includes aspects of the biological,
chemical A chemical substance is a form of matter having constant chemical composition and characteristic properties. Some references add that chemical substance cannot be separated into its constituent elements by physical separation methods, i.e., w ...
, physical, and
geological Geology () is a branch of natural science concerned with Earth and other astronomical objects, the features or rocks of which it is composed, and the processes by which they change over time. Modern geology significantly overlaps all other E ...
characteristics of fresh and saline, natural and man-made bodies of water. This includes the study of
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s, reservoirs, ponds, rivers, springs, streams,
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The p ...
s, and
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidated ...
.Wetzel, R.G. 2001. Limnology: Lake and River Ecosystems, 3rd ed. Academic Press () Water systems are often categorized as either running (lotic) or standing (lentic). Limnology includes the study of the drainage basin, movement of water through the basin and biogeochemical changes that occur en route. A more recent sub-discipline of limnology, termed landscape limnology, studies, manages, and seeks to conserve these
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s using a landscape perspective, by explicitly examining connections between an aquatic ecosystem and its
drainage basin A drainage basin is an area of land where all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
. Recently, the need to understand global inland waters as part of the Earth System created a sub-discipline called global limnology. This approach considers processes in inland waters on a global scale, like the role of inland aquatic ecosystems in global biogeochemical cycles. Limnology is closely related to
aquatic ecology An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The tw ...
and hydrobiology, which study aquatic organisms and their interactions with the abiotic (non-living) environment. While limnology has substantial overlap with freshwater-focused disciplines (e.g., freshwater biology), it also includes the study of inland salt lakes.


History

The term limnology was coined by
François-Alphonse Forel François-Alphonse Forel (February 2, 1841 – August 7, 1912) was a Swiss physician and scientist who pioneered the study of lakes, and is thus considered the founder, and the Father of limnology.Prof. F. A. Forel. Nature 89, 638–639 (1912). ...
(1841–1912) who established the field with his studies of Lake Geneva. Interest in the discipline rapidly expanded, and in 1922
August Thienemann August Friedrich Thienemann (7 September 1882 in Gotha – 22 April 1960 in Plön) was a German limnologist, zoologist and ecologist. He studied zoology at the University of Greifswald. He was an associate Professor of Hydrobiology at the Univer ...
(a German zoologist) and
Einar Naumann Einar Christian Leonard Naumann (13 August 1891 – 22 September 1934) was a Swedish botanist and limnologist who was professor of limnology at the University of Lund. Naumann worked during the summers at the Fishery Station in Aneboda (Smålan ...
(a Swedish botanist) co-founded the International Society of Limnology (SIL, from
Societas Internationalis Limnologiae The International Society of Limnology (SIL) is an international scientific society that disseminates information among limnologists, those who study all aspects of inland waters, including their physics, chemistry, biology, geology, and management ...
). Forel's original definition of limnology, "the oceanography of lakes", was expanded to encompass the study of all inland waters, and influenced
Benedykt Dybowski Benedykt Tadeusz Dybowski (12 May 183331 January 1930) was a Polish naturalist and physician. Life Benedykt Dybowski was born in Adamaryni, within the Minsk Governorate of the Russian Empire to Polish nobility. He was the brother of naturalis ...
's work on Lake Baikal. Prominent early American limnologists included G. Evelyn Hutchinson and Ed Deevey. At the University of Wisconsin-Madison, Edward A. Birge, Chancey Juday, Charles R. Goldman, and Arthur D. Hasler contributed to the development of the Center for Limnology.


General limnology


Physical properties

Physical properties of aquatic ecosystems are determined by a combination of heat, currents, waves and other seasonal distributions of environmental conditions. The
morphometry Morphometrics (from Greek μορϕή ''morphe'', "shape, form", and -μετρία ''metria'', "measurement") or morphometry refers to the quantitative analysis of ''form'', a concept that encompasses size and shape. Morphometric analyses are c ...
of a body of water depends on the type of feature (such as a lake, river, stream, wetland, estuary etc.) and the structure of the earth surrounding the body of water. Lakes, for instance, are classified by their formation, and zones of lakes are defined by water depth. River and stream system morphometry is driven by underlying geology of the area as well as the general velocity of the water. Stream morphometry is also influenced by topography (especially slope) as well as precipitation patterns and other factors such as vegetation and land development. Connectivity between streams and lakes relates to the landscape drainage density, lake surface area and lake shape. Other types of aquatic systems which fall within the study of limnology are estuaries. Estuaries are bodies of water classified by the interaction of a river and the ocean or sea.
Wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The p ...
s vary in size, shape, and pattern however the most common types, marshes, bogs and swamps, often fluctuate between containing shallow, freshwater and being dry depending on the time of year.


Light interactions

Light zonation is the concept of how the amount of sunlight penetration into water influences the structure of a body of water. These zones define various levels of productivity within an aquatic ecosystems such as a lake. For instance, the depth of the water column which sunlight is able to penetrate and where most plant life is able to grow is known as the photic or euphotic zone. The rest of the water column which is deeper and does not receive sufficient amounts of sunlight for plant growth is known as the
aphotic zone The aphotic zone (aphotic from Greek prefix + "without light") is the portion of a lake or ocean where there is little or no sunlight. It is formally defined as the depths beyond which less than 1 percent of sunlight penetrates. Above the aphot ...
.


Thermal stratification

Similar to light zonation, thermal
stratification Stratification may refer to: Mathematics * Stratification (mathematics), any consistent assignment of numbers to predicate symbols * Data stratification in statistics Earth sciences * Stable and unstable stratification * Stratification, or st ...
or thermal zonation is a way of grouping parts of the water body within an aquatic system based on the temperature of different lake layers. The less turbid the water, the more light is able to penetrate, and thus heat is conveyed deeper in the water. Heating declines exponentially with depth in the water column, so the water will be warmest near the surface but progressively cooler as moving downwards. There are three main sections that define thermal stratification in a lake. The
epilimnion The epilimnion or surface layer is the top-most layer in a thermally stratified lake. It sits above the deeper metalimnion and hypolimnion. It is typically warmer and has a higher pH and higher dissolved oxygen concentration than the hypolimnion ...
is closest to the water surface and absorbs long- and shortwave radiation to warm the water surface. During cooler months, wind shear can contribute to cooling of the water surface. The
thermocline A thermocline (also known as the thermal layer or the metalimnion in lakes) is a thin but distinct layer in a large body of fluid (e.g. water, as in an ocean or lake; or air, e.g. an atmosphere) in which temperature changes more drastically with ...
is an area within the water column where water temperatures rapidly decrease. The bottom layer is the
hypolimnion The hypolimnion or under lake is the dense, bottom layer of water in a thermally- stratified lake. The word hypolimnion is derived from the Greek "limnos" meaning "lake". It is the layer that lies below the thermocline. Typically the hypolimni ...
, which tends to have the coldest water because its depth restricts sunlight from reaching it. In temperate lakes, fall-season cooling of surface water results in turnover of the water column, where the thermocline is disrupted, and the lake temperature profile becomes more uniform. In cold climates, when water cools below 4oC (the temperature of maximum density) many lakes can experience an inverse thermal stratification in winter. These lakes are often dimictic, with a brief spring overturn in addition to longer fall overturn. The relative thermal resistance is the energy needed to mix these strata of different temperatures.Wetzel, R. G. (2001). Limnology: Lake and river ecosystems. San Diego: Academic Press. p74, 86


Lake Heat Budget

An annual heat budget, also shown as θa, is the total amount of heat needed to raise the water from its minimum winter temperature to its maximum summer temperature. This can be calculated by integrating the area of the lake at each depth interval (Az) multiplied by the difference between the summer (θsz) and winter (θwz) temperatures or \displaystyle \intAzszwz)


Chemical properties

The chemical composition of water in aquatic ecosystems is influenced by natural characteristics and processes including precipitation, underlying
soil Soil, also commonly referred to as earth or dirt Dirt is an unclean matter, especially when in contact with a person's clothes, skin, or possessions. In such cases, they are said to become dirty. Common types of dirt include: * Debri ...
and bedrock in the
drainage basin A drainage basin is an area of land where all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
,
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is dis ...
, evaporation, and
sedimentation Sedimentation is the deposition of sediments. It takes place when particles in suspension settle out of the fluid in which they are entrained and come to rest against a barrier. This is due to their motion through the fluid in response to the ...
. All bodies of water have a certain composition of both organic and
inorganic In chemistry, an inorganic compound is typically a chemical compound that lacks carbon–hydrogen bonds, that is, a compound that is not an organic compound. The study of inorganic compounds is a subfield of chemistry known as ''inorganic chemist ...
elements and compounds. Biological reactions also affect the chemical properties of water. In addition to natural processes, human activities strongly influence the chemical composition of aquatic systems and their water quality. ''Allochthonous'' sources of carbon or nutrients come from outside the aquatic system (such as plant and soil material). Carbon sources from within the system, such as algae and the microbial breakdown of aquatic particulate organic carbon, are ''autochthonous''. In aquatic food webs, the portion of biomass derived from allochthonous material is then named "allochthony". In streams and small lakes, allochthonous sources of carbon are dominant while in large lakes and the ocean, autochthonous sources dominate.


Oxygen and carbon dioxide

Dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
and dissolved
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
are often discussed together due their coupled role in
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
and photosynthesis. Dissolved oxygen concentrations can be altered by physical, chemical, and biological processes and reaction. Physical processes including wind mixing can increase dissolved oxygen concentrations, particularly in surface waters of aquatic ecosystems. Because dissolved oxygen solubility is linked to water temperatures, changes in temperature affect dissolved oxygen concentrations as warmer water has a lower capacity to "hold" oxygen as colder water. Biologically, both photosynthesis and aerobic respiration affect dissolved oxygen concentrations. Photosynthesis by autotrophic organisms, such as phytoplankton and aquatic algae, increases dissolved oxygen concentrations while simultaneously reducing carbon dioxide concentrations, since carbon dioxide is taken up during photosynthesis. All
aerobic organism Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics Aerobics is a form of physical exercise that combines rhythmic aerobic exe ...
s in the aquatic environment take up dissolved oxygen during aerobic respiration, while carbon dioxide is released as a byproduct of this reaction. Because photosynthesis is light-limited, both photosynthesis and respiration occur during the daylight hours, while only respiration occurs during
dark Darkness, the direct opposite of lightness, is defined as a lack of illumination, an absence of visible light, or a surface that absorbs light, such as black or brown. Human vision is unable to distinguish colors in conditions of very low lu ...
hours or in dark portions of an ecosystem. The balance between dissolved oxygen production and consumption is calculated as the aquatic metabolism rate. Vertical changes in the concentrations of dissolved oxygen are affected by both wind mixing of surface waters and the balance between photosynthesis and respiration of organic matter. These vertical changes, known as profiles, are based on similar principles as thermal stratification and light penetration. As light availability decreases deeper in the water column, photosynthesis rates also decrease, and less dissolved oxygen is produced. This means that dissolved oxygen concentrations generally decrease as you move deeper into the body of water because of photosynthesis is not replenishing dissolved oxygen that is being taken up through respiration. During periods of thermal stratification, water density gradients prevent oxygen-rich surface waters from mixing with deeper waters. Prolonged periods of stratification can result in the depletion of bottom-water dissolved oxygen; when dissolved oxygen concentrations are below 2 milligrams per liter, waters are considered hypoxic. When dissolved oxygen concentrations are approximately 0 milligrams per liter, conditions are anoxic. Both hypoxic and anoxic waters reduce available habitat for organisms that respire oxygen, and contribute to changes in other chemical reactions in the water.


Nitrogen and phosphorus

Nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
and phosphorus are ecologically significant nutrients in aquatic systems. Nitrogen is generally present as a
gas Gas is one of the four fundamental states of matter (the others being solid, liquid, and plasma). A pure gas may be made up of individual atoms (e.g. a noble gas like neon), elemental molecules made from one type of atom (e.g. oxygen), or ...
in aquatic ecosystems however most water quality studies tend to focus on nitrate, nitrite and
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
levels. Most of these dissolved nitrogen compounds follow a seasonal pattern with greater concentrations in the
fall Autumn, also known as fall in American English and Canadian English, is one of the four temperate seasons on Earth. Outside the tropics, autumn marks the transition from summer to winter, in September (Northern Hemisphere) or March ( Southe ...
and
winter Winter is the coldest season of the year in polar and temperate climates. It occurs after autumn and before spring. The tilt of Earth's axis causes seasons; winter occurs when a hemisphere is oriented away from the Sun. Different cultur ...
months compared to the spring and summer. Phosphorus has a different role in aquatic ecosystems as it is a limiting factor in the growth of phytoplankton because of generally low concentrations in the water. Dissolved phosphorus is also crucial to all living things, is often very limiting to primary productivity in freshwater, and has its own distinctive ecosystem cycling.


Biological properties


Role in ecology

Lakes "are relatively easy to sample, because they have clear-cut boundaries (compared to terrestrial ecosystems) and because field experiments are relatively easy to perform.", which make then especially useful for ecologists who try to understand ecological dynamics.Lampert, W., & Sommer, U. 2007. Limnoecology.


Lake trophic classification

One way to classify lakes (or other bodies of water) is with the
trophic state index The Trophic State Index (TSI) is a classification system designed to rate water bodies based on the amount of biological productivity they sustain. Although the term "trophic index" is commonly applied to lakes, any surface water body may be inde ...
. An oligotrophic lake is characterized by relatively low levels of
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...
and low levels of nutrients. A eutrophic lake has high levels of primary productivity due to very high nutrient levels. Eutrophication of a lake can lead to algal blooms. Dystrophic lakes have high levels of
humic matter In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Latin ...
and typically have yellow-brown, tea-coloured waters. These categories do not have rigid specifications; the classification system can be seen as more of a spectrum encompassing the various levels of aquatic productivity.


Professional organizations

People who study limnology are called limnologists. These scientists largely study the characteristics of inland fresh-water systems such as lakes, rivers, streams, ponds and wetlands. They may also study non-oceanic bodies of salt water, such as the Great Salt Lake.There are many professional organizations related to limnology and other aspects of the aquatic science, including the Association for the Sciences of Limnology and Oceanography, the Asociación Ibérica de Limnología, the International Society of Limnology, the Polish Limnological Society, the Society of Canadian Limnologists, and the
Freshwater Biological Association The Freshwater Biological Association (FBA) is an independent scientific organisation founded in 1929 in Cumbria by Felix Eugen Fritsch, William Harold Pearsall, Francis Balfour-Browne, and Robert Gurney among others. Whilst originally created to ...
.


See also

* Hydrology *
Lentic ecosystems A lake ecosystem or lacustrine ecosystem includes biotic (living) plants, animals and micro-organisms, as well as abiotic (non-living) physical and chemical interactions. Lake ecosystems are a prime example of lentic ecosystems (''lentic'' ref ...
* Limnological tower * Lotic ecosystems * Paleolimnology


References


Further reading

* Gerald A. Cole, ''Textbook of Limnology'', 4th ed. (Waveland Press, 1994) * Stanley Dodson, ''Introduction to Limnology'' (2005), * A.J.Horne and C.R. Goldman: ''Limnology'' (1994), * G. E. Hutchinson, ''A Treatise on Limnology'', 3 vols. (1957–1975) - classic but dated * H.B.N. Hynes, ''The Ecology of Running Waters'' (1970) * Jacob Kalff, ''Limnology'' ( Prentice Hall, 2001) * B. Moss, ''Ecology of Fresh Waters'' ( Blackwell, 1998) * Robert G. Wetzel and Gene E. Likens, ''Limnological Analyses'', 3rd ed. ( Springer-Verlag, 2000) * Patrick E. O'Sullivan and Colin S. Reynolds ''The Lakes Handbook: Limnology and limnetic ecology'' {{Authority control .01 Hydrography Lakes Rivers Systems ecology Aquatic ecology Water