Lake ecosystem

Lake ecosystems can be divided into zones. One common system divides lakes into three zones. The first, the littoral zone, is the shallow zone near the shore. This is where rooted wetland plants occur. The offshore is divided into two further zones, an open water zone and a deep water zone. In the open water zone (or photic zone) sunlight supports photosynthetic algae and the species that feed upon them. In the deep water zone, sunlight is not available and the food web is based on detritus entering from the littoral and photic zones. Some systems use other names. The off shore areas may be called the pelagic zone, the photic zone may be called the limnetic zone and the aphotic zone may be called the profundal zone. Inland from the littoral zone, one can also frequently identify a riparian zone which has plants still affected by the presence of the lake—this can include effects from windfalls, spring flooding, and winter ice damage. The production of the lake as a whole is the result of production from plants growing in the littoral zone, combined with production from plankton growing in the open water. Wetlands can be part of the lentic system, as they form naturally along most lake shores, the width of the wetland and littoral zone being dependent upon the slope of the shoreline and the amount of natural change in water levels, within and among years. Often dead trees accumulate in this zone, either from windfalls on the shore or logs transported to the site during floods. This woody debris provides important habitat for fish and nesting birds, as well as protecting shorelines from erosion. ==Abiotic components==

Light Light provides the solar energy required to drive the process of photosynthesis, the major energy source of lentic systems. The amount of light received depends upon a combination of several factors. Small ponds may experience shading by surrounding trees, while cloud cover may affect light availability in all systems, regardless of size. Seasonal and diurnal considerations also play a role in light availability because the shallower the angle at which light strikes water, the more light is lost by reflection. This is known as Beer's law. Once light has penetrated the surface, it may also be scattered by particles suspended in the water column. This scattering decreases the total amount of light as depth increases. Lakes are divided into photic and aphotic regions, the prior receiving sunlight and latter being below the depths of light penetration, making it void of photosynthetic capacity. In relation to lake zonation, the pelagic and benthic zones are considered to lie within the photic region, while the profundal zone is in the aphotic region. Temperature Temperature is an important abiotic factor in lentic ecosystems because most of the biota are poikilothermic, where internal body temperatures are defined by the surrounding system. Water can be heated or cooled through radiation at the surface and conduction to or from the air and surrounding substrate. Shallow ponds often have a continuous temperature gradient from warmer waters at the surface to cooler waters at the bottom. In addition, temperature fluctuations can vary greatly in these systems, both diurnally and seasonally. Temperature regimes are very different in large lakes. In temperate regions, for example, as air temperatures increase, the icy layer formed on the surface of the lake breaks up, leaving the water at approximately 4 °C. This is the temperature at which water has the highest density. As the season progresses, the warmer air temperatures heat the surface waters, making them less dense. The deeper waters remain cool and dense due to reduced light penetration. As the summer begins, two distinct layers become established, with such a large temperature difference between them that they remain stratified. The lowest zone in the lake is the coldest and is called the hypolimnion. The upper warm zone is called the epilimnion. Between these zones is a band of rapid temperature change called the thermocline. During the colder fall season, heat is lost at the surface and the epilimnion cools. When the temperatures of the two zones are close enough, the waters begin to mix again to create a uniform temperature, an event termed lake turnover. In the winter, inverse stratification occurs as water near the surface cools freezes, while warmer, but denser water remains near the bottom. A thermocline is established, and the cycle repeats. Wind In exposed systems, wind can create turbulent, spiral-formed surface currents called Langmuir circulations. Exactly how these currents become established is still not well understood, but it is evident that it involves some interaction between horizontal surface currents and surface gravity waves. The visible result of these rotations, which can be seen in any lake, are the surface foamlines that run parallel to the wind direction. Positively buoyant particles and small organisms concentrate in the foamline at the surface and negatively buoyant objects are found in the upwelling current between the two rotations. Objects with neutral buoyancy tend to be evenly distributed in the water column. This turbulence circulates nutrients in the water column, making it crucial for many pelagic species, however its effect on benthic and profundal organisms is minimal to non-existent, respectively. The degree of nutrient circulation is system specific, as it depends upon such factors as wind strength and duration, as well as lake or pool depth and productivity. Chemistry Oxygen is essential for organismal respiration. The amount of oxygen present in standing waters depends upon: 1) the area of transparent water exposed to the air, 2) the circulation of water within the system and 3) the amount of oxygen generated and used by organisms present. In shallow, plant-rich pools there may be great fluctuations of oxygen, with extremely high concentrations occurring during the day due to photosynthesis and very low values at night when respiration is the dominant process of primary producers. Thermal stratification in larger systems can also affect the amount of oxygen present in different zones. The epilimnion is oxygen rich because it circulates quickly, gaining oxygen via contact with the air. The hypolimnion, however, circulates very slowly and has no atmospheric contact. Additionally, fewer green plants exist in the hypolimnion, so there is less oxygen released from photosynthesis. In spring and fall when the epilimnion and hypolimnion mix, oxygen becomes more evenly distributed in the system. Low oxygen levels are characteristic of the profundal zone due to the accumulation of decaying vegetation and animal matter that “rains” down from the pelagic and benthic zones and the inability to support primary producers. Phosphorus is important for all organisms because it is a component of DNA and RNA and is involved in cell metabolism as a component of ATP and ADP. Also, phosphorus is not found in large quantities in freshwater systems, limiting photosynthesis in primary producers, making it the main determinant of lentic system production. The phosphorus cycle is complex, but the model outlined below describes the basic pathways. Phosphorus mainly enters a pond or lake through runoff from the watershed or by atmospheric deposition. Upon entering the system, a reactive form of phosphorus is usually taken up by algae and macrophytes, which release a non-reactive phosphorus compound as a byproduct of photosynthesis. This phosphorus can drift downwards and become part of the benthic or profundal sediment, or it can be remineralized to the reactive form by microbes in the water column. Similarly, non-reactive phosphorus in the sediment can be remineralized into the reactive form. Sediments are generally richer in phosphorus than lake water, however, indicating that this nutrient may have a long residency time there before it is remineralized and re-introduced to the system. ==Biotic components==

of a bacterial community in a lake Bacteria Bacteria are present in all regions of lentic waters. Free-living forms are associated with decomposing organic material, biofilm on the surfaces of rocks and plants, suspended in the water column, and in the sediments of the benthic and profundal zones. Other forms are also associated with the guts of lentic animals as parasites or in commensal relationships. Phytoplankton are found drifting in the water column of the pelagic zone. Many species have a higher density than water, which should cause them to sink inadvertently down into the benthos. To combat this, phytoplankton have developed density-changing mechanisms, by forming vacuoles and gas vesicles, or by changing their shapes to induce drag, thus slowing their descent. A very sophisticated adaptation utilized by a small number of species is a tail-like flagellum that can adjust vertical position, and allow movement in any direction. Macrophytes are sources of food, oxygen, and habitat structure in the benthic zone, but cannot penetrate the depths of the euphotic zone, and hence are not found there. Invertebrates are predatory insects which rely on surface tension to walk on top of water. They live on the surface of ponds, marshes, and other quiet waters. They can move very quickly, up to 1.5 m/s. Zooplankton are tiny animals suspended in the water column. Like phytoplankton, these species have developed mechanisms that keep them from sinking to deeper waters, including drag-inducing body forms, and the active flicking of appendages (such as antennae or spines). The invertebrates that inhabit the benthic zone are numerically dominated by small species, and are species-rich compared to the zooplankton of the open water. They include: Crustaceans (e.g. crabs, crayfish, and shrimp), molluscs (e.g. clams and snails), and numerous types of insects. These organisms are mostly found in the areas of macrophyte growth, where the richest resources, highly-oxygenated water, and warmest portion of the ecosystem are found. The structurally diverse macrophyte beds are important sites for the accumulation of organic matter, and provide an ideal area for colonization. The sediments and plants also offer a great deal of protection from predatory fishes. Very few invertebrates are able to inhabit the cold, dark, and oxygen-poor profundal zone. Those that can are often red in color, due to the presence of large amounts of hemoglobin, which greatly increases the amount of oxygen carried to cells. Because the concentration of oxygen within this zone is low, most species construct tunnels or burrows in which they can hide, and utilize the minimum amount of movements necessary to circulate water through, drawing oxygen to them without expending too much energy. Fish and other vertebrates Fish have a range of physiological tolerances that are dependent upon which species they belong to. They have different lethal temperatures, dissolved oxygen requirements, and spawning needs that are based on their activity levels and behaviors. Because fish are highly mobile, they are able to deal with unsuitable abiotic factors in one zone by simply moving to another. A detrital feeder in the profundal zone, for example, that finds the oxygen concentration has dropped too low may feed closer to the benthic zone. A fish might also alter its residence during different parts of its life history: hatching in a sediment nest, then moving to the weedy benthic zone to develop in a protected environment with food resources, and finally into the pelagic zone as an adult. Other vertebrate taxa inhabit lentic systems as well. These include amphibians (e.g. salamanders and frogs), reptiles (e.g. snakes, turtles, and alligators), and a large number of waterfowl species. Most of these vertebrates spend part of their time in terrestrial habitats, and thus, are not directly affected by abiotic factors in the lake or pond. Many fish species are important both as consumers and as prey species to the larger vertebrates mentioned above. ==Trophic relationships==

Primary producers Lentic systems gain most of their energy from photosynthesis performed by aquatic plants and algae. This autochthonous process involves the combination of carbon dioxide, water, and solar energy to produce carbohydrates and dissolved oxygen. Within a lake or pond, the potential rate of photosynthesis generally decreases with depth due to light attenuation. Photosynthesis, however, is often low at the top few millimeters of the surface, likely due to inhibition by ultraviolet light. The exact depth and photosynthetic rate measurements of this curve are system-specific and depend upon: 1) the total biomass of photosynthesizing cells, 2) the amount of light attenuating materials, and 3) the abundance and frequency range of light absorbing pigments (i.e. chlorophylls) inside of photosynthesizing cells. Bacteria The vast majority of bacteria in lakes and ponds obtain their energy by decomposing vegetation and animal matter. In the pelagic zone, dead fish and the occasional allochthonous input of litterfall are examples of coarse particulate organic matter (CPOM>1 mm). Bacteria degrade these into fine particulate organic matter (FPOM<1 mm) and then further into usable nutrients. Small organisms such as plankton are also characterized as FPOM. Very low concentrations of nutrients are released during decomposition because the bacteria are utilizing them to build their own biomass. Bacteria, however, are consumed by protozoa, which are in turn consumed by zooplankton, and then further up the trophic levels. Elements other than carbon, particularly phosphorus and nitrogen, are regenerated when protozoa feed on bacterial prey and this way, nutrients become once more available for use in the water column. This regeneration cycle is known as the microbial loop and is a key component of lentic food webs. The profundal zone is home to a unique group of filter feeders that use small body movements to draw a current through burrows that they have created in the sediment. This mode of feeding requires the least amount of motion, allowing these species to conserve energy. Lentic food webs As noted in the previous sections, the lentic biota are linked in complex web of trophic relationships. These organisms can be considered to loosely be associated with specific trophic groups (e.g. primary producers, herbivores, primary carnivores, secondary carnivores, etc.). Scientists have developed several theories in order to understand the mechanisms that control the abundance and diversity within these groups. Very generally, top-down processes dictate that the abundance of prey taxa is dependent upon the actions of consumers from higher trophic levels. Typically, these processes operate only between two trophic levels, with no effect on the others. In some cases, however, aquatic systems experience a trophic cascade; for example, this might occur if primary producers experience less grazing by herbivores because these herbivores are suppressed by carnivores. Bottom-up processes are functioning when the abundance or diversity of members of higher trophic levels is dependent upon the availability or quality of resources from lower levels. Finally, a combined regulating theory, bottom-up:top-down, combines the predicted influences of consumers and resource availability. It predicts that trophic levels close to the lowest trophic levels will be most influenced by bottom-up forces, while top-down effects should be strongest at top levels. ==Community patterns and diversity==

Local species richness The biodiversity of a lentic system increases with the surface area of the lake or pond. This is attributable to the higher likelihood of partly terrestrial species of finding a larger system. Also, because larger systems typically have larger populations, the chance of extinction is decreased. Additional factors, including temperature regime, pH, nutrient availability, habitat complexity, speciation rates, competition, and predation, have been linked to the number of species present within systems. This may be related to size, as Hillebrand and Azovsky found that smaller organisms (protozoa and plankton) did not follow the expected trend strongly, while larger species (vertebrates) did. They attributed this to better dispersal ability by smaller organisms, which may result in high distributions globally. ==Natural lake lifecycles==

Lake creation Lakes can be formed in a variety of ways, but the most common are discussed briefly below. The oldest and largest systems are the result of tectonic activities. The rift lakes in Africa, for example are the result of seismic activity along the site of separation of two tectonic plates. Ice-formed lakes are created when glaciers recede, leaving behind abnormalities in the landscape shape that are then filled with water. Finally, oxbow lakes are fluvial in origin, resulting when a meandering river bend is pinched off from the main channel. Natural extinction All lakes and ponds receive sediment inputs. Since these systems are not really expanding, it is logical to assume that they will become increasingly shallower in depth, eventually becoming wetlands or terrestrial vegetation. The length of this process should depend upon a combination of depth and sedimentation rate. Moss gives the example of Lake Tanganyika, which reaches a depth of 1500 m and has a sedimentation rate of 0.5 mm/yr. Assuming that sedimentation is not influenced by anthropogenic factors, this system should go extinct in approximately 3 million years. Shallow lentic systems might also fill in as swamps encroach inward from the edges. These processes operate on a much shorter timescale, taking hundreds to thousands of years to complete the extinction process. ==Human impacts==

Acidification Sulfur dioxide and nitrogen oxides are naturally released from volcanoes, organic compounds in the soil, wetlands, and marine systems, but the majority of these compounds come from the combustion of coal, oil, gasoline, and the smelting of ores containing sulfur. The resultant reduction in habitat structure has negative impacts on the species that utilize it for spawning, maturation, and general survival. Additionally, the large number of short-lived phytoplankton result in a massive amount of dead biomass settling into the sediment. With regard to native species, invaders may cause changes in size and age structure, distribution, density, population growth, and may even drive populations to extinction. Examples of prominent invaders of lentic systems include the zebra mussel and sea lamprey in the Great Lakes. ==See also==