Thin layers (oceanography)

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Visible layers of a red tide, a planktonic algal bloom, off the shores of Southern California La-Jolla-Red-Tide.780.jpg
Visible layers of a red tide, a planktonic algal bloom, off the shores of Southern California

Thin layers are concentrated aggregations of phytoplankton and zooplankton in coastal and offshore waters that are vertically compressed to thicknesses ranging from several centimeters up to a few meters and are horizontally extensive, sometimes for kilometers. Generally, thin layers have three basic criteria: 1) they must be horizontally and temporally persistent; 2) they must not exceed a critical threshold of vertical thickness; and 3) they must exceed a critical threshold of maximum concentration. The precise values for critical thresholds of thin layers has been debated for a long time due to the vast diversity of plankton, instrumentation, and environmental conditions. [1] Thin layers have distinct biological, chemical, optical, and acoustical signatures which are difficult to measure with traditional sampling techniques such as nets and bottles. However, there has been a surge in studies of thin layers within the past two decades due to major advances in technology and instrumentation. Phytoplankton are often measured by optical instruments that can detect fluorescence such as LIDAR, and zooplankton are often measured by acoustic instruments that can detect acoustic backscattering such as ABS. [2] These extraordinary concentrations of plankton have important implications for many aspects of marine ecology (e.g., phytoplankton growth dynamics, zooplankton grazing, behaviour, environmental effects, harmful algal blooms), as well as for ocean optics and acoustics. Zooplankton thin layers are often found slightly under phytoplankton layers because many feed on them. Thin layers occur in a wide variety of ocean environments, including estuaries, coastal shelves, fjords, bays, and the open ocean, and they are often associated with some form of vertical structure in the water column, such as pycnoclines, and in zones of reduced flow. [3]

Contents

Criteria

Persistence

Thin layers persist from hours to weeks while other small-scale patches of plankton exist for minutes. [1] The presence of nutrients as well as coastal fronts, eddies, and upwelling zones greatly increase the persistence of thin layers. One of the main criteria for an aggregation of plankton to be considered a thin layer is that the increased concentration at a certain depth of the water column must appear in subsequently measured profiles. However, thin layers are dynamic and horizontally extensive so their persistence cannot be defined using multiple measurements at only one location. [4] A study on the Karenia brevis algae responsible for more recent and increasingly longer red tide blooms shows that the cellular gene expression patterns are extremely diverse which means that this particular species of plankton are more resilient because they adapt well to changing conditions. Studies also indicate that red tide blooms are often terminated by interactions with other microbes such as viruses and bacteria that may either compete for the same nutrients or adversely impact the algal cells. [5]

Thickness

Some studies have considered the maximum critical threshold for vertical thickness of thin layers as three meters, but more recent data has shown that the criteria can be relaxed to five meters. [1] [2] [4] [6] The horizontal extents of thin layers can reach tens of kilometers, and their horizontal to vertical aspect ratio is usually at least 1000:1. [1]

Intensity

The intensity of a thin layer refers to the maximum concentration of the plankton within the layer relative to the background and the water column. Thin layer concentrations can range between three and 100 times more than the background [1] and up to 75% of the total biomass in the water column. [7]

Formation

Buoyancy

Layers of phytoplankton found in the Arctic Ocean.

Thin layers of non-motile phytoplankton tend to collect at boundaries of strong vertical gradients in salinity (haloclines), temperature (thermoclines), and density (pycnoclines) which often coincide because they are directly proportional. [7] These particular thin layers are formed by sinking non-motile phytoplankton reaching a neutral buoyancy at a pycnocline, and the stifling of vertical turbulent dispersion at these depths. Other studies have shown that gradients in nutrients (nutriclines) also contribute to the formation of thin layers. [8]

Vertical Migration

Many zooplankton normally exhibit a diel vertical migration (DVM) pattern that dictates their depth in the water column based on the time of day. Phytoplankton require sunlight for photosynthesis and protein production, but they are not primarily attracted to light. This is evident by their single move up near the surface prior to sunrise and single move down into deeper waters prior to sunset. Their collective movements may result in the aggregation that form thin layers. These regular movements are thought to be governed by an internal clock in normal nutrient concentrations However, they have also been observed to migrate irregularly when nutrient concentrations are higher or lower than normal. [9]

A plankton patch in the ocean being dispersed horizontally due to velocity shear Straining due to Shear.png
A plankton patch in the ocean being dispersed horizontally due to velocity shear

Chemotaxis

Motile plankton have been observed to be able to detect and swim towards higher nutrient concentrations and/or light intensities. This mechanism is called chemotaxis and is partly responsible for the formation of thin layers at depths where nutrients are abundant. Another mechanism specific to dinoflagellates is called helical klinotaxis where the algal cell's ability to respond to both positive and negative chemosensory signals is crucial to their motility. If dinoflagellates were not capable of both positive and negative chemotaxis, they would not navigate successfully due to the nature of the transverse and longitudinal flagella causing rotating and translating motions, respectively. [10]

Eddies, Filaments, and Fronts

Gyrotactic trapping of swimming plankton due to sharp changes in flow velocities in the ocean. Gyrotactic Trapping.png
Gyrotactic trapping of swimming plankton due to sharp changes in flow velocities in the ocean.

Another obvious cause of thin layers is the horizontal transport of waters with high plankton concentration into waters with lower concentrations. [1] In this case, upwelled intrusions of nutrient-rich slope water are suggested to be the cause of algal blooms and some thin layers. [11] However, thin layers have been observed to form at the boundaries of more complex fluid mechanisms such as eddies, filaments, and fronts. These thin layers were located at the transition layer, a region of maximum shear and stratification at the base of the mixed layer. [4]

Straining by Shear

A fluid mechanism that contributes to the formation of thin layers is the straining of fluid by the sheared velocity profile which causes the fluid to tilt and disperse horizontally. If a patch of plankton is located at the fluid being sheared, a thin layer could be formed by the straining of the patch by velocity shear. The four phases of plankton distributions caused by straining are: 1) tilting, 2) shear-thinning, 3) decay, and 4) shear-dispersion (dissipation). [12]

Gyrotactic Trapping

A sharp change in flow velocities can also prevent some motile plankton from orienting themselves or swimming vertically. This fluid mechanism is called gyrotactic trapping. [13]

See also

Related Research Articles

<span class="mw-page-title-main">Plankton</span> Organisms that are in the water column and are incapable of swimming against a current

Plankton are the diverse collection of organisms found in water that are unable to propel themselves against a current. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish, and baleen whales.

<span class="mw-page-title-main">Algal bloom</span> Spread of planktonic algae in water

An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term algae encompasses many types of aquatic photosynthetic organisms, both macroscopic multicellular organisms like seaweed and microscopic unicellular organisms like cyanobacteria. Algal bloom commonly refers to the rapid growth of microscopic unicellular algae, not macroscopic algae. An example of a macroscopic algal bloom is a kelp forest.

<span class="mw-page-title-main">Phytoplankton</span> Autotrophic members of the plankton ecosystem

Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words φυτόν, meaning 'plant', and, meaning 'wanderer' or 'drifter'.

<span class="mw-page-title-main">Zooplankton</span> Heterotrophic protistan or metazoan members of the plankton ecosystem

Zooplankton are the animal component of the planktonic community, having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequently, they drift or are carried along by currents in the ocean, or by currents in seas, lakes or rivers.

<span class="mw-page-title-main">Downwelling</span> Process of accumulation and sinking of higher density material beneath lower density material

Downwelling is the downward movement of a fluid parcel and its properties within a larger fluid. It is closely related to upwelling, the upward movement of fluid.

<span class="mw-page-title-main">Spring bloom</span> Strong increase in phytoplankton abundance that typically occurs in the early spring

The spring bloom is a strong increase in phytoplankton abundance that typically occurs in the early spring and lasts until late spring or early summer. This seasonal event is characteristic of temperate North Atlantic, sub-polar, and coastal waters. Phytoplankton blooms occur when growth exceeds losses, however there is no universally accepted definition of the magnitude of change or the threshold of abundance that constitutes a bloom. The magnitude, spatial extent and duration of a bloom depends on a variety of abiotic and biotic factors. Abiotic factors include light availability, nutrients, temperature, and physical processes that influence light availability, and biotic factors include grazing, viral lysis, and phytoplankton physiology. The factors that lead to bloom initiation are still actively debated.

<span class="mw-page-title-main">Pycnocline</span> Layer where the density gradient is greatest within a body of water

A pycnocline is the cline or layer where the density gradient is greatest within a body of water. An ocean current is generated by the forces such as breaking waves, temperature and salinity differences, wind, Coriolis effect, and tides caused by the gravitational pull of celestial bodies. In addition, the physical properties in a pycnocline driven by density gradients also affect the flows and vertical profiles in the ocean. These changes can be connected to the transport of heat, salt, and nutrients through the ocean, and the pycnocline diffusion controls upwelling.

<span class="mw-page-title-main">Critical depth</span> Hypothesized depth at which phytoplankton growth is matched by losses

In biological oceanography, critical depth is defined as a hypothetical surface mixing depth where phytoplankton growth is precisely matched by losses of phytoplankton biomass within the depth interval. This concept is useful for understanding the initiation of phytoplankton blooms.

<i>Karenia brevis</i> Species of dinoflagellate

Karenia brevis is a microscopic, single-celled, photosynthetic organism in the genus Karenia. It is a marine dinoflagellate commonly found in the waters of the Gulf of Mexico. It is the organism responsible for the "Florida red tides" that affect the Gulf coasts of Florida and Texas in the U.S., and nearby coasts of Mexico. K. brevis has been known to travel great lengths around the Florida peninsula and as far north as the Carolinas.

<span class="mw-page-title-main">Langmuir circulation</span> Series of shallow, slow, counter-rotating vortices at the oceans surface aligned with the wind

In physical oceanography, Langmuir circulation consists of a series of shallow, slow, counter-rotating vortices at the ocean's surface aligned with the wind. These circulations are developed when wind blows steadily over the sea surface. Irving Langmuir discovered this phenomenon after observing windrows of seaweed in the Sargasso Sea in 1927. Langmuir circulations circulate within the mixed layer; however, it is not yet so clear how strongly they can cause mixing at the base of the mixed layer.

The deep chlorophyll maximum (DCM), also called the subsurface chlorophyll maximum, is the region below the surface of water with the maximum concentration of chlorophyll. The DCM generally exists at the same depth as the nutricline, the region of the ocean where the greatest change in the nutrient concentration occurs with depth.

<i>Karenia</i> (dinoflagellate) Genus of single-celled organisms

Karenia is a genus that consists of unicellular, photosynthetic, planktonic organisms found in marine environments. The genus currently consists of 12 described species. They are best known for their dense toxic algal blooms and red tides that cause considerable ecological and economical damage; some Karenia species cause severe animal mortality. One species, Karenia brevis, is known to cause respiratory distress and neurotoxic shellfish poisoning (NSP) in humans.

<span class="mw-page-title-main">Ecosystem of the North Pacific Subtropical Gyre</span> Major circulating ecosystem of ocean currents

The North Pacific Subtropical Gyre (NPSG) is the largest contiguous ecosystem on earth. In oceanography, a subtropical gyre is a ring-like system of ocean currents rotating clockwise in the Northern Hemisphere and counterclockwise in the Southern Hemisphere caused by the Coriolis Effect. They generally form in large open ocean areas that lie between land masses.

<span class="mw-page-title-main">Sea foam</span> Foam created by the agitation of seawater

Sea foam, ocean foam, beach foam, or spume is a type of foam created by the agitation of seawater, particularly when it contains higher concentrations of dissolved organic matter derived from sources such as the offshore breakdown of algal blooms. These compounds can act as surfactants or foaming agents. As the seawater is churned by breaking waves in the surf zone adjacent to the shore, the surfactants under these turbulent conditions trap air, forming persistent bubbles that stick to each other through surface tension.

Phycotoxins are complex allelopathic chemicals produced by eukaryotic and prokaryotic algal secondary metabolic pathways. More simply, these are toxic chemicals synthesized by photosynthetic organisms. These metabolites are not harmful to the producer but may be toxic to either one or many members of the marine food web. This page focuses on phycotoxins produced by marine microalgae; however, freshwater algae and macroalgae are known phycotoxin producers and may exhibit analogous ecological dynamics. In the pelagic marine food web, phytoplankton are subjected to grazing by macro- and micro-zooplankton as well as competition for nutrients with other phytoplankton species. Marine bacteria try to obtain a share of organic carbon by maintaining symbiotic, parasitic, commensal, or predatory interactions with phytoplankton. Other bacteria will degrade dead phytoplankton or consume organic carbon released by viral lysis. The production of toxins is one strategy that phytoplankton use to deal with this broad range of predators, competitors, and parasites. Smetacek suggested that "planktonic evolution is ruled by protection and not competition. The many shapes of plankton reflect defense responses to specific attack systems". Indeed, phytoplankton retain an abundance of mechanical and chemical defense mechanisms including cell walls, spines, chain/colony formation, and toxic chemical production. These morphological and physiological features have been cited as evidence for strong predatory pressure in the marine environment. However, the importance of competition is also demonstrated by the production of phycotoxins that negatively impact other phytoplankton species. Flagellates are the principle producers of phycotoxins; however, there are known toxigenic diatoms, cyanobacteria, prymnesiophytes, and raphidophytes. Because many of these allelochemicals are large and energetically expensive to produce, they are synthesized in small quantities. However, phycotoxins are known to accumulate in other organisms and can reach high concentrations during algal blooms. Additionally, as biologically active metabolites, phycotoxins may produce ecological effects at low concentrations. These effects may be subtle, but have the potential to impact the biogeographic distributions of phytoplankton and bloom dynamics.

<span class="mw-page-title-main">Planktivore</span> Aquatic organism that feeds on planktonic food

A planktivore is an aquatic organism that feeds on planktonic food, including zooplankton and phytoplankton. Planktivorous organisms encompass a range of some of the planet's smallest to largest multicellular animals in both the present day and in the past billion years; basking sharks and copepods are just two examples of giant and microscopic organisms that feed upon plankton. Planktivory can be an important mechanism of top-down control that contributes to trophic cascades in aquatic and marine systems. There is a tremendous diversity of feeding strategies and behaviors that planktivores utilize to capture prey. Some planktivores utilize tides and currents to migrate between estuaries and coastal waters; other aquatic planktivores reside in lakes or reservoirs where diverse assemblages of plankton are present, or migrate vertically in the water column searching for prey. Planktivore populations can impact the abundance and community composition of planktonic species through their predation pressure, and planktivore migrations facilitate nutrient transport between benthic and pelagic habitats.

<i>Cochlodinium polykrikoides</i> Species of single-celled organism

Cochlodinium polykrikoides is a species of red tide producing marine dinoflagellates known for causing fish kills around the world, and well known for fish kills in marine waters of Southeast Asia. C. polykrikoides has a wide geographic range, including North America, Central America, Western India, Southwestern Europe and Eastern Asia. Single cells of this species are ovoidal in shape, 30-50μm in length and 25-30μm in width.

<span class="mw-page-title-main">Mixotrophic dinoflagellate</span> Plankton

Dinoflagellates are eukaryotic plankton, existing in marine and freshwater environments. Previously, dinoflagellates had been grouped into two categories, phagotrophs and phototrophs. Mixotrophs, however include a combination of phagotrophy and phototrophy. Mixotrophic dinoflagellates are a sub-type of planktonic dinoflagellates and are part of the phylum Dinoflagellata. They are flagellated eukaryotes that combine photoautotrophy when light is available, and heterotrophy via phagocytosis. Dinoflagellates are one of the most diverse and numerous species of phytoplankton, second to diatoms.

<span class="mw-page-title-main">Marine protists</span> Protists that live in saltwater or brackish water

Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.

Eddy pumping is a component of mesoscale eddy-induced vertical motion in the ocean. It is a physical mechanism through which vertical motion is created from variations in an eddy's rotational strength. Cyclonic (Anticyclonic) eddies lead primarily to upwelling (downwelling) in the Northern Hemisphere and vice versa in the Southern hemisphere. It is a key mechanism driving biological and biogeochemical processes in the ocean such as algal blooms and the carbon cycle.

References

  1. 1 2 3 4 5 6 Durham, William M.; Stocker, Roman (2012-01-15). "Thin Phytoplankton Layers: Characteristics, Mechanisms, and Consequences". Annual Review of Marine Science. 4 (1): 177–207. Bibcode:2012ARMS....4..177D. doi:10.1146/annurev-marine-120710-100957. ISSN   1941-1405. PMID   22457973.
  2. 1 2 Benoit-Bird, Kelly J.; Shroyer, Emily L.; McManus, Margaret A. (2013-08-02). "A critical scale in plankton aggregations across coastal ecosystems". Geophysical Research Letters. 40 (15): 3968–3974. Bibcode:2013GeoRL..40.3968B. doi: 10.1002/grl.50747 . ISSN   0094-8276. S2CID   16586461.
  3. McManus, M. A., Cheriton, O. M., Drake, P. J., Holliday, D. V., Storlazzi, C. D., Donaghay, P. L., et al. (2005). Effects of physical processes on structure and transport of thin zooplankton layers in the coastal ocean. Marine Ecology Progress Series, 301, 199-215.
  4. 1 2 3 Johnston, T.M. Shaun; Cheriton, Olivia M.; Pennington, J. Timothy; Chavez, Francisco P. (February 2009). "Thin phytoplankton layer formation at eddies, filaments, and fronts in a coastal upwelling zone". Deep Sea Research Part II: Topical Studies in Oceanography. 56 (3–5): 246–259. Bibcode:2009DSRII..56..246J. doi:10.1016/j.dsr2.2008.08.006. ISSN   0967-0645.
  5. Van Dolah, Frances M.; Lidie, Kristy B.; Monroe, Emily A.; Bhattacharya, Debashish; Campbell, Lisa; Doucette, Gregory J.; Kamykowski, Daniel (March 2009). "The Florida red tide dinoflagellate Karenia brevis: New insights into cellular and molecular processes underlying bloom dynamics". Harmful Algae. 8 (4): 562–572. doi:10.1016/j.hal.2008.11.004. ISSN   1568-9883.
  6. Greer, Adam T.; Cowen, Robert K.; Guigand, Cedric M.; McManus, Margaret A.; Sevadjian, Jeff C.; Timmerman, Amanda H.V. (2013-06-04). "Relationships between phytoplankton thin layers and the fine-scale vertical distributions of two trophic levels of zooplankton". Journal of Plankton Research. 35 (5): 939–956. doi: 10.1093/plankt/fbt056 . ISSN   1464-3774.
  7. 1 2 McManus, M. A.; Woodson, C. B. (2012-02-22). "Plankton distribution and ocean dispersal". Journal of Experimental Biology. 215 (6): 1008–1016. doi: 10.1242/jeb.059014 . ISSN   0022-0949. PMID   22357594.
  8. Churnside, James H.; Marchbanks, Richard D. (2015-06-22). "Subsurface plankton layers in the Arctic Ocean". Geophysical Research Letters. 42 (12): 4896–4902. Bibcode:2015GeoRL..42.4896C. doi: 10.1002/2015gl064503 . ISSN   0094-8276.
  9. Yamazaki, Atsuko K.; Kamykowski, Daniel (September 2000). "A dinoflagellate adaptive behavior model: response to internal biochemical cues". Ecological Modelling. 134 (1): 59–72. doi:10.1016/s0304-3800(00)00336-7. ISSN   0304-3800.
  10. FENCHEL, T (December 2001). "How Dinoflagellates Swim". Protist. 152 (4): 329–338. doi:10.1078/1434-4610-00071. ISSN   1434-4610. PMID   11822661.
  11. Walsh, John J. (2003). "Phytoplankton response to intrusions of slope water on the West Florida Shelf: Models and observations". Journal of Geophysical Research. 108 (C6): 3190. Bibcode:2003JGRC..108.3190W. doi:10.1029/2002jc001406. ISSN   0148-0227.
  12. Birch, Daniel A.; Young, William R.; Franks, Peter J.S. (March 2008). "Thin layers of plankton: Formation by shear and death by diffusion". Deep Sea Research Part I: Oceanographic Research Papers. 55 (3): 277–295. Bibcode:2008DSRI...55..277B. doi:10.1016/j.dsr.2007.11.009. ISSN   0967-0637.
  13. Guasto, Jeffrey S.; Rusconi, Roberto; Stocker, Roman (2012-01-21). "Fluid Mechanics of Planktonic Microorganisms". Annual Review of Fluid Mechanics. 44 (1): 373–400. Bibcode:2012AnRFM..44..373G. doi:10.1146/annurev-fluid-120710-101156. ISSN   0066-4189.