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Bioturbation is defined as the reworking of and by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a primary driver of biodiversity. The formal study of bioturbation began in the 1800s by Charles Darwin experimenting in his garden. The disruption of aquatic sediments and terrestrial soils through bioturbating activities provides significant ecosystem services. These include the alteration of in aquatic sediment and overlying water, shelter to other species in the form of burrows in terrestrial and water ecosystems, and soil production on land.Shaler, N. S., 1891, The origin and nature of soils, in Powell, J. W., ed., USGS 12th Annual report 1890–1891: Washington, D.C., Government Printing Office, p. 213-45.
Bioturbators are deemed ecosystem engineers because they alter resource availability to other species through the physical changes they make to their environments. This type of ecosystem change affects the evolution of cohabitating species and the environment, which is evident in left in marine and terrestrial sediments. Other bioturbation effects include altering the texture of sediments (diagenesis), bioirrigation, and displacement of microorganisms and non-living particles. Bioturbation is sometimes confused with the process of bioirrigation, however these processes differ in what they are mixing; bioirrigation refers to the mixing of water and solutes in sediments and is an effect of bioturbation.
, salmon, and pocket gophers are examples of large bioturbators.Humphreys, G. S., and Mitchell, P. B., 1983, A preliminary assessment of the role of bioturbation and rainwash on sandstone hillslopes in the Sydney Basin, in Australian and New Zealand Geomorphology Group, p. 66-80. Although the activities of these large macrofaunal bioturbators are more conspicuous, the dominant bioturbators are small invertebrates, such as , , Thalassinidea, mud shrimp, and Chironomidae. The activities of these small invertebrates, which include burrowing and ingestion and defecation of sediment grains, contribute to mixing and the alteration of sediment structure.
Microbial communities are greatly influenced by bioturbator activities, as increased transport of more energetically favorable Oxidizing agent, such as oxygen, to typically highly reduced sediments at depth alters the microbial metabolic processes occurring around burrows. As bioturbators burrow, they also increase the surface area of sediments across which oxidized and reduced solutes can be exchanged, thereby increasing the overall sediment metabolism. This increase in sediment metabolism and microbial activity further results in enhanced organic matter decomposition and sediment oxygen uptake. In addition to the effects of burrowing activity on microbial communities, studies suggest that bioturbator fecal matter provides a highly nutritious food source for microbes and other macrofauna, thus enhancing benthic microbial activity. This increased microbial activity by bioturbators can contribute to increased nutrient release to the overlying water column. Nutrients released from enhanced microbial decomposition of organic matter, notably limiting nutrients, such as ammonium, can have bottom-up effects on ecosystems and result in increased growth of phytoplankton and bacterioplankton.
Burrows offer protection from predation and harsh environmental conditions. For example, termites (Macrotermes bellicosus) burrow and create mounds that have a complex system of air ducts and evaporation devices that create a suitable microclimate in an unfavorable physical environment. Many species are attracted to bioturbator burrows because of their protective capabilities. The shared use of burrows has enabled the evolution of Symbiosis between bioturbators and the many species that utilize their burrows. (2025). 9780226467283, University of Chicago Press. ISBN 9780226467283 For example, gobies, scale-worms, and crabs live in the burrows made by innkeeper worms. Social interactions provide evidence of co-evolution between hosts and their burrow symbionts. This is exemplified by shrimp-goby associations. Shrimp burrows provide shelter for gobies and gobies serve as a scout at the mouth of the burrow, signaling the presence of potential danger. In contrast, the blind goby Blind goby lives within the deep portion of Callianassa shrimp burrows where there is not much light. The blind goby is an example of a species that is an obligate Commensalism, meaning their existence depends on the host bioturbator and its burrow. Although newly hatched blind gobies have fully developed eyes, their eyes become withdrawn and covered by skin as they develop. They show evidence of commensal morphological evolution because it is hypothesized that the lack of light in the burrows where the blind gobies reside is responsible for the evolutionary loss of functional eyes.
Bioturbators can also inhibit the presence of other benthic organisms by smothering, exposing other organisms to predators, or resource competition. While thalassinidean shrimps can provide shelter for some organisms and cultivate interspecies relationships within burrows, they have also been shown to have strong negative effects on other species, especially those of bivalves and surface-grazing Gastropoda, because thalassinidean shrimps can smother bivalves when they resuspend sediment. They have also been shown to exclude or inhibit polychaetes, , and Amphipoda. This has become a serious issue in the northwestern United States, as ghost and mud shrimp (thalassinidean shrimp) are considered pests to bivalve aquaculture operations. The presence of bioturbators can have both negative and positive effects on the recruitment of larvae of conspecifics (those of the same species) and those of other species, as the resuspension of sediments and alteration of flow at the sediment-water interface can affect the ability of larvae to burrow and remain in sediments. This effect is largely species-specific, as species differences in resuspension and burrowing modes have variable effects on fluid dynamics at the sediment-water interface. Deposit feeder bioturbators may also hamper recruitment by consuming recently settled larvae.
For example, bioturbating animals are hypothesized to have affected the cycling of sulfur in the early oceans. According to this hypothesis, bioturbating activities had a large effect on the sulfate concentration in the ocean. Around the Cambrian-Precambrian boundary (539 million years ago), animals begin to mix reduced sulfur from to overlying water causing sulfide to oxidize, which increased the sulfate composition in the ocean. During large extinction events, the sulfate concentration in the ocean was reduced. Although this is difficult to measure directly, seawater sulfur isotope compositions during these times indicates bioturbators influenced the sulfur cycling in the early Earth.
Bioturbators have also altered phosphorus cycling on geologic scales. Bioturbators mix readily available particulate organic phosphorus (P) deeper into ocean sediment layers which prevents the precipitation of phosphorus (mineralization) by increasing the sequestration of phosphorus above normal chemical rates. The sequestration of phosphorus limits oxygen concentrations by decreasing production on a geologic time scale. This decrease in production results in an overall decrease in oxygen levels, and it has been proposed that the rise of bioturbation corresponds to a decrease in oxygen levels of that time. The negative feedback of animals sequestering phosphorus in the sediments and subsequently reducing oxygen concentrations in the environment limits the intensity of bioturbation in this early environment.
, such as earth worms and small mammals, form passageways for air and water transport which changes the soil properties, such as the vertical particle-size distribution, soil porosity, and nutrient content. Invertebrates that burrow and consume plant detritus help produce an organic-rich topsoil known as the soil biomantle, and thus contribute to the formation of Soil horizon Small mammals such as pocket gophers also play an important role in the production of soil, possibly with an equal magnitude to abiotic processes. Pocket gophers form above-ground mounds, which moves soil from the lower soil horizons to the surface, exposing minimally weathered rock to surface erosion processes, speeding soil formation. Pocket gophers are thought to play an important role in the downslope transport of soil, as the soil that forms their mounds is more susceptible to erosion and subsequent transport. Similar to tree root effects, the construction of burrows-even when backfilled- decreases soil density. The formation of surface mounds also buries surface vegetation, creating nutrient hotspots when the vegetation decomposes, increasing soil organic matter. Due to the high metabolic demands of their burrow-excavating subterranean lifestyle, pocket gophers must consume large amounts of plant material. Though this has a detrimental effect on individual plants, the net effect of pocket gophers is increased plant growth from their positive effects on soil nutrient content and physical soil properties.
Lake and pond sediments often transition from the aerobic (oxygen containing) character of the overlaying water to the anaerobic (without oxygen) conditions of the lower sediment over sediment depths of only a few millimeters, therefore, even bioturbators of modest size can affect this transition of the chemical characteristics of sediments. By mixing anaerobic sediments into the water column, bioturbators allow aerobic processes to interact with the re-suspended sediments and the newly exposed bottom sediment surfaces.
Macroinvertebrates including Chironomidae (non-biting midges) larvae and Tubifex tubifex (detritus worms) are important agents of bioturbation in these ecosystems and have different effects based on their respective feeding habits. Tubificid worms do not form burrows, they are upward conveyors. Chironomids, on the other hand, form burrows in the sediment, acting as bioirrigators and aerating the sediments and are downward conveyors. This activity, combined with chironomid's respiration within their burrows, decrease available oxygen in the sediment and increase the loss of nitrates through enhanced rates of denitrification.
The increased oxygen input to sediments by macroinvertebrate bioirrigation coupled with bioturbation at the sediment-water interface complicates the total flux of phosphorus . While bioturbation results in a net flux of phosphorus into the water column, the bio-irrigation of the sediments with oxygenated water enhances the adsorption of phosphorus onto iron-oxide compounds, thereby reducing the total flux of phosphorus into the water column.
The presence of macroinvertebrates in sediment can initiate bioturbation due to their status as an important food source for benthivorous fish such as Common carp. Of the bioturbating, benthivorous fish species, carp in particular are important ecosystem engineers and their foraging and burrowing activities can alter the water quality characteristics of ponds and lakes. Carp increase water turbidity by the re-suspension of benthic sediments. This increased turbidity limits light penetration and coupled with increased nutrient flux from the sediment into the water column, inhibits the growth of macrophytes (aquatic plants) favoring the growth of phytoplankton in the surface waters. Surface phytoplankton colonies benefit from both increased suspended nutrients and from recruitment of buried phytoplankton cells released from the sediments by the fish bioturbation. Macrophyte growth has also been shown to be inhibited by displacement from the bottom sediments due to fish burrowing.
Salmon function as bioturbators on both gravel to sand-sized sediment and a nutrient scale, by moving and re-working sediments in the construction of redds (gravel depressions or "nests" containing eggs buried under a thin layer of sediment) in rivers and streams and by mobilization of nutrients. The construction of salmon redds functions to increase the ease of fluid movement (hydraulic conductivity) and porosity of the stream bed. In select rivers, if salmon congregate in large enough concentrations in a given area of the river, the total sediment transport from redd construction can equal or exceed the sediment transport from flood events. The net effect on sediment movement is the downstream transfer of gravel, sand and finer materials and enhancement of water mixing within the river substrate.
The construction of salmon redds increases sediment and nutrient fluxes through the hyporheic zone (area between surface water and groundwater) of rivers and effects the dispersion and retention of marine derived nutrients (MDN) within the river ecosystem. MDN are delivered to river and stream ecosystems by the fecal matter of spawning salmon and the decaying carcasses of salmon that have completed spawning and died. Numerical modeling suggests that residence time of MDN within a salmon spawning reach is inversely proportional to the amount of redd construction within the river. Measurements of respiration within a salmon-bearing river in Alaska further suggest that salmon bioturbation of the river bed plays a significant role in mobilizing MDN and limiting primary productivity while salmon spawning is active. The river ecosystem was found to switch from a net to system in response to decreased primary production and increased respiration. The decreased primary production in this study was attributed to the loss of benthic primary producers who were dislodged due to bioturbation, while increased respiration was thought to be due to increased respiration of organic carbon, also attributed to sediment mobilization from salmon redd construction. While marine derived nutrients are generally thought to increase productivity in riparian and freshwater ecosystems, several studies have suggested that temporal effects of bioturbation should be considered when characterizing salmon influences on nutrient cycles.
The effects of bioturbation on the nitrogen cycle are well-documented. Coupled denitrification and nitrification are enhanced due to increased oxygen and nitrate delivery to deep sediments and increased surface area across which oxygen and nitrate can be exchanged. The enhanced nitrification-denitrification coupling contributes to greater removal of biologically available nitrogen in shallow and coastal environments, which can be further enhanced by the excretion of ammonium by bioturbators and other organisms residing in bioturbator burrows. While both nitrification and denitrification are enhanced by bioturbation, the effects of bioturbators on denitrification rates have been found to be greater than that on rates of nitrification, further promoting the removal of biologically available nitrogen. This increased removal of biologically available nitrogen has been suggested to be linked to increased rates of nitrogen fixation in microenvironments within burrows, as indicated by evidence of nitrogen fixation by sulfate-reducing bacteria via the presence of nifH (nitrogenase) genes.
Bioturbation by walrus feeding is a significant source of sediment and biological community structure and nutrient flux in the Bering Sea. Walruses feed by digging their muzzles into the sediment and extracting clams through powerful suction. By digging through the sediment, walruses rapidly release large amounts of organic material and nutrients, especially ammonium, from the sediment to the water column. Additionally, walrus feeding behavior mixes and oxygenates the sediment and creates pits in the sediment which serve as new habitat structures for invertebrate larvae.
Parameterization of bioturbation, however, can vary, as newer and more complex models can be used to fit tracer profiles. Unlike the standard biodiffusion model, these more complex models, such as expanded versions of the biodiffusion model, random walk, and particle-tracking models, can provide more accuracy, incorporate different modes of sediment transport, and account for more spatial heterogeneity.
An alternate, less widely accepted hypothesis for the origin of bioturbation exists. The trace fossil Nenoxites is thought to be the earliest record of bioturbation, predating the Cambrian Period. The fossil is dated to 555 million years, which places it in the Ediacaran The fossil indicates a 5 centimeter depth of bioturbation in muddy sediments by a burrowing worm. This is consistent with food-seeking behavior, as there tended to be more food resources in the mud than the water column. However, this hypothesis requires more precise Geochronology to rule out an early Cambrian origin for this specimen.
The evolution of trees during the Devonian enhanced soil weathering and increased the spread of soil due to bioturbation by tree roots. Root penetration and uprooting also enhanced soil carbon storage by enabling mineral weathering and the burial of organic matter.
Important from bioturbation have been found in marine sediments from tidal, coastal and deep sea sediments. In addition sand dune, or Eolian dust, sediments are important for preserving a wide variety of fossils. Evidence of bioturbation has been found in deep-sea sediment cores including into long records, although the act extracting the core can disturb the signs of bioturbation, especially at shallower depths. Arthropods, in particular are important to the geologic record of bioturbation of Eolian sediments. Dune records show traces of burrowing animals as far back as the lower Mesozoic (250 Million years ago), although bioturbation in other sediments has been seen as far back as 550 Ma.
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