Lentic system ecology

Lentic system ecology is the study of the biotic and abiotic interactions within still continental waters.^[1] Together with lotic system ecology, which involves flowing waters such as rivers and streams, these fields form the more general study area of freshwater or aquatic ecology.

Lentic systems are diverse, ranging from a small, temporary rainwater pool a few inches deep to Lake Baikal, which has a maximum depth of 1740 m.^[2] The general distinction between pools/ponds and lakes is vague, but Brown^[1] states that ponds and pools have their entire bottom surfaces exposed to light, while lakes do not. In addition, some lakes become seasonally stratified (discussed in more detail below.) Ponds and pools have two regions: the pelagic open water zone, and the benthic zone, which is comprised of the bottom and shore regions. Since lakes have deep bottom regions not exposed to light, these systems have an additional zone, the profundal.^[3] These three areas can have very different abiotic conditions and, hence, host species that are specifically adapted to live there.^[1]

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Lentic system biota

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.^[3] Bacteria play an important role in system metabolism through nutrient recycling,^[2] which will be discussed in the Trophic Relationships section.

Primary producers

Algae, including both phytoplankton and periphyton are the principle photosynthesizers in ponds and lakes. Phytoplankton are found drifting in the water column of the pelagic zone. Many species have a higher density than water which should making them sink and end up in 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, slowing their descent. A very sophisticated adaptation utilized by a small number of species is a tail-like flagella that can adjust vertical position and allow movement in any direction.^[2] Phytoplankton can also maintain their presence in the water column by being circulated in Langmuir rotations.^[3] Periphytic algae, on the other hand, are attached to a substrate. In lakes and ponds, they can cover all benthic surfaces. Both types of plankton are important as food sources and as oxygen providers.^[2]

Plants, or macrophytes, in lentic systems live in both the benthic and pelagic zones and can be grouped according to their manner of growth: 1) emergent macrophytes = rooted in the substrate but with leaves and flowers extending into the air, 2) floating-leaved macrophytes = rooted in the substrate but with floating leaves, 3) submersed macrophytes = not rooted in the substrate and floating beneath the surface and 4) free-floating macrophytes = not rooted in the substrate and floating on the surface.^[1] These various forms of macrophytes generally occur in different areas of the benthic zone, with emergent vegetation nearest the shoreline, then floating-leaved macrophytes, followed by submersed vegetation. Free-floating macrophytes can occur anywhere on the system’s surface.^[2]

Aquatic plants are more buoyant than their terrestrial counterparts because freshwater has a higher density than air. This makes structural rigidity unimportant in lakes and ponds (except in the aerial stems and leaves). Thus, the leaves and stems of most aquatic plants use less energy to construct and maintain woody tissue, investing that energy into fast growth instead.^[1] In order to contend with stresses induced by wind and waves, plants must be both flexible and tough (Reynolds 2004). Light is the most important factor controlling the distribution of submerged aquatic plants. 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.^[1][4]

Invertebrates

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.^[1] Remaining in the water column may have its advantages in terms of feeding, but this zone’s lack of refugia leaves zooplankton vulnerable to predation. In response, some species, especially Daphnia sp., make daily vertical migrations in the water column by passively sinking to the darker lower depths during the day and actively moving towards the surface during the night. Also, because conditions in a lentic system can be quite variable across seasons, zooplankton have the ability to switch from laying regular eggs to resting eggs when there is a lack of food, temperatures fall below 2 °C, or if predator abundance is high. These resting eggs have a diapause, or dormancy period that should allow the zooplankton to encounter conditions that are more favorable to survival when they finally hatch.^[5] 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.^[2] 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.^[3]

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.^[1] Because the concentration of oxygen within this zone is low, most species construct tunnels or borrows in which they can hide and make the minimum movements necessary to circulate water through, drawing oxygen to them without expending much energy.^[1]

Fishes and other vertebrates

Fishes 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 fishes 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.^[4] 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 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.^[4] The energy created by these primary producers is important for the community because it is transferred to higher trophic levels via consumption.

Bacteria

The vast majority of bacteria in lakes and ponds obtain their energy by decomposing vegetation and animal matter. In the pelagic zone, dead fishes and the occasional allochtohnous 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. Nutrients, including those that contain carbon and phosphorus, are reintroduced into the water column at any number of points along this food chain via excretion or organism death, making them available again for bacteria. This regeneration cycle is known as the microbial loop and is a key component of lentic food webs.^[2]

The decomposition of organic materials can continue in the benthic and profundal zones if the matter falls through the water column before being completely digested by the pelagic bacteria. Bacteria are found in the greatest abundance here in sediments, where they are typically 2-1000 times more prevalent than in the water column.^[3]

Invertebrates

Invertebrates can be divided into feeding guilds based on their method of prey capture. Zooplankton rely on a combination of dissolved organic matter and particulates for survival. Zooplankton concentrate unicellular algae, bacteria, and detritus in the pelagic zone by sieving through morphological structures or with the creation of rotary currents that can centrifuge particles into a larger mass for digestion. These processes are energetically efficient, but they do not allow for the selection of usable material. As a result, a portion of the catch is invaluable as a food source and must be discarded. When capturing larger particles, including other zooplankton taxa, each particle is procured individually with a raptorial appendage. This type of feeding is energetically more costly, but the nutritional returns can be greater because selection is involved.^[5]

Benthic invertebrates, due to their high level of species richness, have many methods of prey capture. Filter feeders create currents via siphons or beating cilia, to pull water and its nutritional contents, towards themselves for straining. Grazers use scraping, rasping, and shredding adaptations to feed on periphytic algae and macrophytes. Members of the collector guild browse the sediments, picking out specific particles with raptorial appendages. Deposit feeding invertebrates indiscriminately consume sediment, digesting any organic material it contains. Finally, some invertebrates belong to the predator guild, capturing and consuming living animals.^[2][6] 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.^[1] A small number of invertebrate taxa are predators in the profundal zone. These species are likely from other regions and only come to these depths to feed. The vast majority of invertebrates in this zone are deposit feeders, getting their energy from the surrounding sediments.^[6]

Fish

Fish size, mobility, and sensory capabilities allow them to exploit a broad prey base, covering multiple zonation regions. Like invertebrates, fish feeding habits can be categorized into guilds. In the pelagic zone, herbivores graze on periphyton and macrophytes or pick phytoplankton out of the water column. Carnivores include fishes that feed on zooplankton in the water column (zooplanktivores), insects at the water’s surface, on benthic structures, or in the sediment (insectivores), and those that feed on other fishes (piscivores). Fish that consume detritus and gain energy by processing its organic material are called detritivores. Omnivores ingest a wide variety of prey, encompassing floral, faunal, and detrital material. Finally, members of the parasitic guild acquire nutrition from a host species, usually another fish or large vertebrate.^[2] Fish taxa are flexible in their feeding roles, varying their diets with environmental conditions and prey availability. Many species also undergo a diet shift as they develop. Therefore, it is likely that any single fish occupies multiple feeding guilds within its lifetime.^[7]

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 (eg. 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 dependant 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.^[2]

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.^[8] 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.^[2]

Succession patterns in plankton communities – The PEG model

Phytoplankton and zooplankton communities in lake systems undergo seasonal succession in relation to nutrient availability, predation, and competition. Sommer et al.^[9] described these patterns as part of the Plankton Ecology Group (PEG) model, with 24 statements constructed from the analysis of numerous systems. The following includes a subset of these statements, as explained by Brönmark and Hansson^[2] illustrating succession through a single seasonal cycle:

Winter
1. Increased nutrient and light availability result in rapid phytoplankton growth towards the end of winter. The dominant species, such as diatoms, are small and have quick growth capabilities. 2. These plankton are consumed by zooplankton, which become the dominant plankton taxa.

Spring
3. A clear water phase occurs, as phytoplankton populations become depleted due to increased predation by growing numbers of zooplankton.

Summer
4. Zooplankton abundance declines as a result of decreased phytoplankton prey and increased predation by juvenile fishes. 5. With increased nutrient availability and decreased predation from zooplankton, a diverse phytoplankton community develops. 6. As the summer continues, nutrients become depleted in a predictable order: phosphorus, silica, and then nitrogen. The abundance of various phytoplankton species varies in relation to their biological need for these nutrients. 7. Small-sized zooplankton become the dominant type of zooplankton because they are less vulnerable to fish predation.

Fall
8. Predation by fishes is reduced due to lower temperatures and zooplankton of all sizes increase in number.

Winter
9. Cold temperatures and decreased light availability result in lower rates of primary production and decreased phytoplankton populations. 10. Reproduction in zooplankton decreases due to lower temperatures and less prey.

The PEG model presents an idealized version of this succession pattern, while natural systems are known for their variation.^[2]

Latitudinal patterns

There is a well-documented global pattern that correlates decreasing plant and animal diversity with increasing latitude, that is to say, there are fewer species as one moves towards the poles. The cause of this pattern is one of the greatest puzzles for ecologists today. Theories for its explanation include energy availability, climatic variability, disturbance, competition, etc.^[2] Despite this global diversity gradient, this pattern can be weak for freshwater systems compared to global marine and terrestrial systems.^[10] This may be related to size, as Hillebrand and Azovsky^[11] 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.^[2]

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.^[2]

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^[4] 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.^[4]

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.^[3] These substances dissolve in atmospheric moisture and enter lentic systems as acid rain.^[1] Lakes and ponds that contain bedrock that is rich in carbonates have a natural buffer, resulting in no alteration of pH. Systems without this bedrock, however, are very sensitive to acid inputs because they have a low neutralizing capacity, resulting in pH declines even with only small inputs of acid.^[3] At a pH of 5-6 algal species diversity and biomass decrease considerably, leading to an increase in water transparency – a characteristic feature of acidified lakes. As the pH continues lower, all fauna becomes less diverse. The most significant feature is the disruption of fish reproduction. Thus, the population is eventually composed of few, old individuals that eventually die and leave the systems without fishes.^[3][2] Acid rain has been especially harmful to lakes in the Northeastern United States.

Eutrophication

Eutrophic systems contain a high concentration of phosphorus (~30+µg/L), nitrogen (~1500+µg/L), or both.^[2] Phosphorus enters lentic waters from wastewater treatment effluents, discharge from raw sewage, or from runoff of farmland. Nitrogen mostly comes from agricultural fertilizers from runoff or leaching and subsequent groundwater flow. This increase in nutrients required for primary producers results in a massive increase of phytoplankton growth, termed a plankton bloom. This bloom decreases water transparency, leading to the loss of submerged plants. 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.^[4] Bacteria need large amounts of oxygen to decompose this material, reducing the oxygen concentration of the water. This is especially pronounced in stratified lakes when the thermocline prevents oxygen rich water from the surface to mix with lower levels. Low or anoxic conditions preclude the existence of many taxa that are not physiologically tolerant of these conditions.^[2]

Invasive species

Invasive species have been introduced to lentic systems through both purposeful events (e.g. stocking game and food species) as well as unintentional events (e.g. in ballast water). These organisms can affect natives via competition for prey or habitat, predation, habitat alteration, hybridization, or the introduction of harmful diseases and parasites.^[12] 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.^[2] Examples of prominent invaders of lentic systems include the zebra mussel and sea lamprey in the Great Lakes.

See also

• United States Environmental Protection Agency - Great Lakes Ecosystems
• United States Environmental Protection Agency - Limnology Primer (PDF file)

References

1. ^ ^{a b c d e f g h i j k} Brown, A. L. (1987). Freshwater Ecology. Heinimann Educational Books, London, 163. 
2. ^ ^{a b c d e f g h i j k l m n o p q r s t} Brönmark, C.; L. A. Hansson (2005). The Biology of Lakes and Ponds. Oxford University Press, Oxford, 285. 
3. ^ ^{a b c d e f g h} Kalff, J. (2002). Limnology. Prentice Hall, Upper Saddle, NJ, 592. 
4. ^ ^{a b c d e f} Moss, B. (1998). Ecology of Freshwaters: man and medium, past to future. Blackwell Science, London, 557. 
5. ^ ^{a b} Gliwicz, Z. M. (2004). Zooplankton (in The Lakes Handbook). P. E. O'Sullivan and C.S. Reynolds, 461-516. 
6. ^ ^{a b} Jónasson, P. M. (2003). Benthic Invertebrates (in The Lakes Handbook). P. E. O'Sullivan and C. S. Reynolds, 341-416. 
7. ^ Winfield, I. J. (2003). Fish Population Ecology (in The Lakes Handbook). P.E. Sullivan and C.S. Reynolds, 517-537. 
8. ^ Browne, R. A. (1981). "Lakes as islands: biogeographic distribution, turnover rates, and species composition in the lakes of central New York". Journal of Biogeography 8: 75-83.
9. ^ Sommer, U.; Z. M. Gliwicz, W. Lampert and A. Duncan (1986). "The PEG-model of seasonal succession of planktonic events in freshwaters". Archiv für Hydrobiologie 106: 433-471.
10. ^ Hillebrand, H. (2004). "On the generality of the latitudinal diversity gradient". American Naturalist 163: 192-211.
11. ^ Hillebrand, H.; A. I. Azovsky (2001). "Body size determines the strength of the latitudinal diversity gradient". Ecography 24: 251-256.
12. ^ Giller, S.; B. Malmqvist (1998). The Biology of Streams and Rivers. Oxford University Press, Oxford, 296.