Gelatinous zooplankton are fragile animals that live in the water column in the ocean. Their delicate bodies have no hard parts and are easily damaged or destroyed. [2] Gelatinous zooplankton are often transparent. [3] All jellyfish are gelatinous zooplankton, but not all gelatinous zooplankton are jellyfish. The most commonly encountered organisms include ctenophores, medusae, salps, and Chaetognatha in coastal waters. However, almost all marine phyla, including Annelida, Mollusca and Arthropoda, contain gelatinous species, but many of those odd species live in the open ocean and the deep sea and are less available to the casual ocean observer. [4] Many gelatinous plankters utilize mucous structures in order to filter feed. [5] Gelatinous zooplankton have also been called Gelata. [6]
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Jellyfish are slow swimmers, and most species form part of the plankton. Traditionally jellyfish have been viewed as trophic dead ends, minor players in the marine food web, gelatinous organisms with a body plan largely based on water that offers little nutritional value or interest for other organisms apart from a few specialised predators such as the ocean sunfish and the leatherback sea turtle. [7] [1] That view has recently been challenged. Jellyfish, and more gelatinous zooplankton in general, which include salps and ctenophores, are very diverse, fragile with no hard parts, difficult to see and monitor, subject to rapid population swings and often live inconveniently far from shore or deep in the ocean. It is difficult for scientists to detect and analyse jellyfish in the guts of predators, since they turn to mush when eaten and are rapidly digested. [7] But jellyfish bloom in vast numbers, and it has been shown they form major components in the diets of tuna, spearfish and swordfish as well as various birds and invertebrates such as octopus, sea cucumbers, crabs and amphipods. [8] [1] "Despite their low energy density, the contribution of jellyfish to the energy budgets of predators may be much greater than assumed because of rapid digestion, low capture costs, availability, and selective feeding on the more energy-rich components. Feeding on jellyfish may make marine predators susceptible to ingestion of plastics." [1]
According to a 2017 study, narcomedusae consume the greatest diversity of mesopelagic prey, followed by physonect siphonophores, ctenophores and cephalopods. [9] The importance of the so-called "jelly web" is only beginning to be understood, but it seems medusae, ctenophores and siphonophores can be key predators in deep pelagic food webs with ecological impacts similar to predator fish and squid. Traditionally gelatinous predators were thought ineffectual providers of marine trophic pathways, but they appear to have substantial and integral roles in deep pelagic food webs. [9]
Pelagic siphonophores, a diverse group of cnidarians, are found at most depths of the ocean - from the surface, like the Portuguese man of war, to the deep sea. They play important roles in ocean ecosystems, and are among the most abundant gelatinous predators. [10]
Biological oceanic processes, primarily carbon production in the euphotic zone, sinking and remineralization, govern the global biological carbon soft‐tissue pump. [12] Sinking and laterally transported carbon‐laden particles fuel benthic ecosystems at continental margins and in the deep sea. [13] [14] Marine zooplankton play a major role as ecosystem engineers in coastal and open ocean ecosystems because they serve as links between primary production, higher trophic levels, and deep‐sea communities. [15] [14] [16] In particular, gelatinous zooplankton (Cnidaria, Ctenophora, and Chordata, namely, Thaliacea) are universal members of plankton communities that graze on phytoplankton and prey on other zooplankton and ichthyoplankton. They also can rapidly reproduce on a time scale of days and, under favorable environmental conditions, some species form dense blooms that extend for many square kilometers. [17] These blooms have negative ecological and socioeconomic impacts by reducing commercially harvested fish species, [18] limiting carbon transfer to other trophic levels, [19] enhancing microbial remineralization, and thereby driving oxygen concentrations down close to anoxic levels. [20] [11]
The global biomass of gelatinous zooplankton (sometimes referred to as jelly‐C) within the upper 200 m of the ocean amounts to 0.038 Pg C. [21] Calculations for mesozooplankton (200 μm to 2 cm) suggest about 0.20 Pg C. [22] The short life span of most gelatinous zooplankton, from weeks up to 2 to 12 months, [23] [24] suggests biomass‐production rates above 0.038 Pg C year−1, depending on the assumed mortality rates, which in many cases are species‐specific. This is much smaller than global primary production (50 Pg C year−1), [25] which translates into export estimates close to 6 Pg C year−1 below 100 m, [26] [27] depending on the method used. Globally, gelatinous zooplankton abundance and distribution patterns largely follow those of temperature and dissolved oxygen as well as primary production as the carbon source. [21] However, gelatinous zooplankton cope with a wide spectrum of environmental conditions, indicating the ability to adapt and occupy most available ecological niches in a water mass. In terms of Longhurst regions (biogeographical provinces that partition the pelagic environment, [28] [29] the highest densities of gelatinous zooplankton occur in coastal waters of the Humboldt Current, NE U.S. Shelf, Scotian and Newfoundland shelves, Benguela Current, East China and Yellow Seas, followed by polar regions of the East Bering and Okhotsk Seas, the Southern Ocean, enclosed bodies of water such as the Black Sea and the Mediterranean, and the west Pacific waters of the Japan seas and the Kuroshio Current. [30] [31] [21] Large amounts of jelly carbon biomass that are reported from coastal areas of open shelves and semi-enclosed seas of North America, Europe, and East Asia come from coastal stranding data. [32] [11]
Large amounts of jelly carbon are quickly transferred to and remineralized on the seabed in coastal areas, including estuaries, lagoons and subtidal/intertidal zones, [15] shelves and slopes, [33] [34] [35] the deepsea. [36] and even entire continental margins such as in the Mediterranean Sea. [37] Jelly carbon transfer begins when gelatinous zooplankton die at a given "death depth" (exit depth), continues as biomass sinks through the water column, and terminates once biomass is remineralized during sinking or reaches the seabed, and then decays. Jelly carbon per se represents a transfer of "already exported" particles (below the mixed later, euphotic or mesopelagic zone), originated in primary production since gelatinous zooplankton "repackage" and integrate this carbon in their bodies, and after death transfer it to the ocean's interior. While sinking through the water column, jelly carbon is partially or totally remineralized as dissolved organic/inorganic carbon and nutrients (DOC, DIC, DON, DOP, DIN and DIP) [38] [39] [40] and any left overs further experience microbial decomposition or are scavenged by macrofauna and megafauna once on the seabed. [41] [42] Despite the high lability of jelly‐C, [43] [39] a remarkably large amount of biomass arrives at the seabed below 1,000 m. During sinking, jelly‐C biochemical composition changes via shifts in C:N:P ratios as observed in experimental studies. [20] [44] [45] Yet realistic jelly‐C transfer estimates at the global scale remain in their infancy, preventing a quantitative assessment of the contribution to the biological carbon soft‐tissue pump. [11]
Ocean carbon export is typically estimated from the flux of sinking particles that are either caught in sediment traps [47] or quantified from videography, [48] and subsequently modeled using sinking rates. [49] Biogeochemical models [50] [51] [52] are normally parameterized using particulate organic matter data (e.g., 0.5–1,000 μm marine snow and fecal pellets) that were derived from laboratory experiments [53] or from sediment trap data. [50] These models do not include jelly‐C (except larvaceans, [54] [55] not only because this carbon transport mechanism is considered transient/episodic and not usually observed, and mass fluxes are too big to be collected by sediment traps, [27] but also because models aim to simplify the biotic compartments to facilitate calculations. Furthermore, jelly‐C deposits tend not to build up at the seafloor over a long time, such as phytodetritus (Beaulieu, 2002), being consumed rapidly by demersal and benthic organisms [41] or decomposed by microbes. [42] The jelly‐C sinking rate is governed by organism size, diameter, biovolume, geometry, [56] density, [57] and drag coefficients. [58] In 2013, Lebrato et al. determined the average sinking speed of jelly‐C using Cnidaria, Ctenophora, and Thaliacea samples, which ranged from 800 to 1,500 m day−1 (salps: 800–1,200 m day−1; scyphozoans: 1,000–1,100 m d−1; ctenophores: 1,200–1,500 m day−1; pyrosomes: 1,300 m day−1). [59] Jelly‐C model simulations suggest that, regardless of taxa, higher latitudes are more efficient corridors to transfer jelly‐C to the seabed owing to lower remineralization rates. [60] In subtropical and temperate regions, significant decomposition takes place in the water column above 1,500 m depth, except in cases where jelly‐C starts sinking below the thermocline. In shallow‐water coastal regions, time is a limiting factor, which prevents remineralization while sinking and results in the accumulation of decomposing jelly‐C from a variety of taxa on the seabed. This suggests that gelatinous zooplankton transfer most biomass and carbon to the deep ocean, enhancing coastal carbon fluxes via DOC and DIC, fueling microbial and megafaunal/macrofaunal scavenging communities. However, the absence of satellite‐derived jelly‐C measurements (such as primary production) [61] and the limited number of global zooplankton biomass data sets make it challenging to quantify global jelly‐C production and transfer efficiency to the ocean's interior. [11]
Because of its fragile structure, image acquisition of gelatinous zooplankton requires the assistance of computer visioning. Automated recognition of zooplankton in sample deposits is possible by utilising technologies such as Tikhonov regularization, support vector machines and genetic programming. [62]
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.
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 πλαγκτός, meaning 'wanderer' or 'drifter'.
Zooplankton are the animal component of the planktonic community. 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.
The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. In other words, it is a biologically mediated process which results in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).
The mesopelagiczone, also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins at the depth where only 1% of incident light reaches and ends where there is no light; the depths of this zone are between approximately 200 to 1,000 meters below the ocean surface.
The bathypelagic zone or bathyal zone is the part of the open ocean that extends from a depth of 1,000 to 4,000 m below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypelagic is also known as the midnight zone because of the lack of sunlight; this feature does not allow for photosynthesis-driven primary production, preventing growth of phytoplankton or aquatic plants. Although larger by volume than the photic zone, human knowledge of the bathypelagic zone remains limited by ability to explore the deep ocean.
Picoplankton is the fraction of plankton composed by cells between 0.2 and 2 μm that can be either prokaryotic and eukaryotic phototrophs and heterotrophs:
Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word "diel" comes from Latin: diēs, lit. 'day', and means a 24-hour period. The migration occurs when organisms move up to the uppermost layer of the sea at night and return to the bottom of the daylight zone of the oceans or to the dense, bottom layer of lakes during the day. It is important to the functioning of deep-sea food webs and the biologically driven sequestration of carbon.
The microbial loop describes a trophic pathway where, in aquatic systems, dissolved organic carbon (DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by phytoplankton-zooplankton-nekton. In soil systems, the microbial loop refers to soil carbon. The term microbial loop was coined by Farooq Azam, Tom Fenchel et al. in 1983 to include the role played by bacteria in the carbon and nutrient cycles of the marine environment.
Calanus finmarchicus is a species of copepod and a component of the zooplankton, which is found in enormous amounts in the northern Atlantic Ocean.
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.
In the deep ocean, marine snow is a continuous shower of mostly organic detritus falling from the upper layers of the water column. It is a significant means of exporting energy from the light-rich photic zone to the aphotic zone below, which is referred to as the biological pump. Export production is the amount of organic matter produced in the ocean by primary production that is not recycled (remineralised) before it sinks into the aphotic zone. Because of the role of export production in the ocean's biological pump, it is typically measured in units of carbon. The term was coined by explorer William Beebe as observed from his bathysphere. As the origin of marine snow lies in activities within the productive photic zone, the prevalence of marine snow changes with seasonal fluctuations in photosynthetic activity and ocean currents. Marine snow can be an important food source for organisms living in the aphotic zone, particularly for organisms that live very deep in the water column.
Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλανκτος, meaning "wanderer" or "drifter", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and freshwater.
Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 millimeters (53 μm) to 2 millimeters.
Jelly-falls are marine carbon cycling events whereby gelatinous zooplankton, primarily cnidarians, sink to the seafloor and enhance carbon and nitrogen fluxes via rapidly sinking particulate organic matter. These events provide nutrition to benthic megafauna and bacteria. Jelly-falls have been implicated as a major “gelatinous pathway” for the sequestration of labile biogenic carbon through the biological pump. These events are common in protected areas with high levels of primary production and water quality suitable to support cnidarian species. These areas include estuaries and several studies have been conducted in fjords of Norway.
The viral shunt is a mechanism that prevents marine microbial particulate organic matter (POM) from migrating up trophic levels by recycling them into dissolved organic matter (DOM), which can be readily taken up by microorganisms. The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.
Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers is larger than the biomass of primary producers. This happens because the ocean's primary producers are tiny phytoplankton which grow and reproduce rapidly, so a small mass can have a fast rate of primary production. In contrast, many significant terrestrial primary producers, such as mature forests, grow and reproduce slowly, so a much larger mass is needed to achieve the same rate of primary production.
Benthic-pelagic coupling are processes that connect the benthic zone and the pelagic zone through the exchange of energy, mass, or nutrients. These processes play a prominent role in both freshwater and marine ecosystems and are influenced by a number of chemical, biological, and physical forces that are crucial to functions from nutrient cycling to energy transfer in food webs.
Jellyfish blooms are substantial growths in population of species under the phyla Cnidaria and Ctenophora.
Helle Ploug is marine scientist known for her work on particles in seawater. She is a professor at the University of Gothenburg, and was named a fellow of the Association for the Sciences of Limnology and Oceanography in 2017.