Open ocean habitat
The open ocean, or pelagic ecosystem, comprises marine areas beyond coastal boundaries and above the seabed (Fig. 1). It includes the entire water column beyond the continental shelf, extending from tropical to polar regions and from the sea surface to abyssal depths. The open ocean is not a uniform desert, but a highly structured habitat controlled by light, depth, temperature, stratification, nutrients, oxygen, particle flux, currents and fronts. It is a highly dynamic habitat in which physical and chemical processes strongly influence biological activity, resulting in substantial geographic variability in productivity.
The open ocean plays a major role in regulating Earth’s climate. Ocean currents redistribute heat from equatorial regions to higher latitudes, helping to moderate global temperatures. The ocean is the primary source of atmospheric water vapor and also acts as an important sink for carbon dioxide, absorbing large quantities of CO2 from the atmosphere and thereby reducing the accumulation of greenhouse gases. In addition, open seas influence regional and local weather patterns; winds originating over the ocean often bring rainfall, while tropical cyclones develop through interactions between warm ocean waters and the atmosphere. The open ocean provides habitat for highly specialized pelagic organisms and serves as an important feeding ground for many other species.[1]
This article introduces the main habitat characteristics of the open ocean. It is one of the sub-categories within the section dealing with biodiversity of marine habitats and ecosystems. It gives an overview of the physical structure, zonation, biology and main pressures affecting open-ocean habitats. Some legal aspects are also indicated.
Contents
Zonation
The water column is commonly subdivided into depth zones, which differ in light availability, pressure, temperature and characteristic biological communities. The main zones are (Fig. 2):
- The epipelagic zone ranges from the sea surface to a depth of 100-200 metres. This is also the limit of the photic zone. Light penetrates sufficiently into this surface layer to allow for photosynthesis.
- The mesopelagic zone lies underneath the epipelagic zone and extends to about 1,000 metres.
- The bathypelagic zone is the zone between the 1,000 and 4,000 metres.
- The abyssopelagic zone extends to a water depth of 6,000 metres.
- The hadopelagic zone is deeper than 6,000 metres and is found in deep-sea trenches.
Characteristics
Currents
Currents are driven by density differences, wind forcing and tides. Density depends on the temperature and salinity of seawater, see Seawater density. Cold, saline water is denser and tends to sink, whereas warm, less saline water remains near the surface. Cooling, warming, evaporation and freezing modify density differences between water masses and thereby influence their motion. Large-scale circulation patterns driven by wind and density differences are strongly influenced by topography; see Ocean circulation.
The largest open-ocean ecosystems are the subtropical gyres of the Atlantic, Pacific and Indian Oceans. They are driven by the anticyclonic wind-stress curl generated by the trade winds at low latitudes and the westerlies at mid-latitudes. Convergence associated with Ekman transport generates large downwelling zones within these gyres. The North Pacific Subtropical Gyre is the largest circulation system on Earth and one of the most persistent large-scale oceanic features. Its general position is maintained by the prevailing wind system, although its boundaries and internal structure vary seasonally and interannually. It is characterized by weak surface currents, strong stratification and generally low biological productivity, although substantial mesoscale and seasonal variability occurs[2].
Light transmission and absorption
Life in the ocean depends directly or indirectly on solar energy. Marine plants and protists use chlorophyll and accessory pigments to capture visible light for photosynthesis. Part of the incoming sunlight is reflected at the sea surface, while the remainder penetrates the water column and is progressively absorbed by water molecules and suspended matter.
Infrared and red wavelengths are rapidly absorbed in the upper meters of the ocean, whereas blue and green wavelengths penetrate much deeper, especially in clear oceanic waters (Fig. 3). This selective absorption of wavelengths gives the open ocean its characteristic blue colour. In very clear oceanic water, less than 1% of surface photosynthetically active radiation remains at depths near 100 m. Based on light penetration, the water column can be divided into two zones. The upper photic zone receives sufficient light for photosynthesis, whereas photosynthesis is generally not possible in the deeper aphotic zone.
Nearshore waters are often more turbid because of high concentrations of suspended particles. In these waters, light penetration may be limited to less than 20 m depth. Scattering by suspended particles shifts the underwater light field from blue toward green and yellow wavelengths.
Stratification
The water column is stratified by density differences related to temperature and salinity gradients, as explained in Seawater density. The transition zone between the less dense surface layer and the denser underlying water is called the pycnocline. In many regions, the pycnocline coincides with the thermocline, which separates the warm surface layer from colder deep water (Fig. 4).
Wind forcing and convective overturning caused by surface cooling and evaporation mix the upper ocean above the thermocline, creating a surface mixed layer whose temperature and biological communities vary seasonally. Stratification can also result from salinity differences; the corresponding transition zone is called the halocline. High surface salinity results from strong evaporation.
In many subtropical gyres, the upper ocean consists of a warm, well-mixed surface layer, typically 40–100 m deep, overlying colder and denser interior waters. Strong density gradients suppress vertical mixing across the pycnocline and restrict the upward transport of nutrients from deeper waters. Downwelling and stratification both contribute to reduce the supply of nutrients, which become the limiting factor for primary production, leading to oligotrophic conditions and low surface chlorophyll concentrations[4] (Fig. 5).
High chlorophyll concentrations along the eastern margins of ocean basins, particularly off western Africa and South America, are associated with coastal upwelling driven by Ekman transport. In highly productive regions, decomposition of sinking organic matter may lead to oxygen depletion within the thermocline; under low-oxygen conditions, denitrification can also occur. Major oxygen minimum zones occur in the eastern tropical North Pacific and the Arabian Sea. Both areas are characterized by relatively long residence times due to weak lateral transport and mixing.
In high-latitude oceans, cold surface waters reduce vertical density gradients, allowing deep vertical mixing. This mixing supplies abundant nutrients to surface waters, but phytoplankton growth is often limited by low light availability and, in some regions, by iron limitation. Nevertheless, surface chlorophyll concentrations in high-latitude oceans are generally higher than in the subtropical gyres.
Biology
The epipelagic or photic zone
In large parts of the open ocean, primary production is limited by the availability of inorganic nitrogen. Nitrogen fixation by cyanobacteria is therefore an important source of bioavailable nitrogen. Nitrogen-fixing cyanobacteria have a particularly high iron demand because the nitrogenase enzyme complex requires iron. Iron limitation is especially important in high-nutrient, low-chlorophyll regions such as the Southern Ocean, the equatorial Pacific and the subarctic North Pacific[7]. See also Biogeochemical cycles in the coastal marine environment.
In oligotrophic open-ocean regions, primary production in the sunlit upper layer is constrained by nutrient availability. Near the base of the photic zone, nutrients are supplied by turbulent diffusion from the mesopelagic layer below, where mineralization of sinking organic matter (marine snow) releases inorganic nutrients. Enhanced phytoplankton biomass and productivity therefore often occur near the base of the photic zone, forming a subsurface chlorophyll maximum (Fig. 6). Phytoplankton optimize growth by adjusting resource allocation[8]. Under low-light and nutrient-replete conditions, they preferentially allocate resources to light harvesting at the expense of nutrient uptake[6].
Despite the enormous size of the open ocean, biomass concentrations are generally low, although species diversity is high. Pelagic primary producers consist mainly of phytoplankton, including cyanobacteria, diatoms, dinoflagellates and coccolithophores. Diatoms are important food sources for herbivorous zooplankton, especially copepods and krill, which in turn are consumed by fishes and other predators.
Many neustonic organisms, which live at or near the sea surface and drift with currents, are blue or transparent, providing camouflage and protection against intense solar and ultraviolet radiation. [9].
The epipelagic biome hosts a specialized fish fauna, including flying fishes and their relatives (needlefishes and halfbeaks), flotsam-associated fishes (such as triggerfishes, filefishes, jacks and dolphinfish), schooling fishes (herrings and anchovies), larval and juvenile fishes of many species, and highly migratory predators such as tunas, billfishes and mako sharks[10].
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Fig. 7a. Neuston Physalia physalis[11] |
Fig. 7c. Copepod [12] |
The mesopelagic or dysphotic zone
The mesopelagic or dysphotic zone extends approximately from 200 m to 1,000 m depth and receives only faint light. Oxygen concentrations may become very low in this zone because of bacterial decomposition of sinking organic material from surface water. Seasonal temperature variability is much weaker than at the surface, resulting in relatively stable environmental conditions.
The mesopelagic fauna is dominated by crustaceans such as copepods, shrimps, amphipods, ostracods and euphausiids, together with squids and fishes. Many species are red or black because red wavelengths are rapidly absorbed with depth, rendering these colors effectively invisible. Many organisms possess large, highly sensitive eyes adapted to dim light conditions. Others possess photophores, organs that produce bioluminescence (Fig. 8). In some fishes, the light is produced by symbiotic bacteria living within specialized photophores.
Many mesopelagic organisms undertake diurnal vertical migration. They ascend into the epipelagic zone at night to feed and descend again into deeper waters during daytime[9].
The bathypelagic or aphotic zone
The bathypelagic or aphotic zone extends from approximately 1,000 m to 4,000 m depth. Sunlight no longer penetrates into this region, which is permanently dark and cold. Many species are black or red in color and possess bioluminescent organs. Common organisms include copepods, ostracods, jellyfishes, amphipods, mysids, worms and deep-sea fishes.
Many bathypelagic fishes possess adaptations that enhance prey detection and capture under extremely low-light conditions. They are often small, with bioluminescent organs, large mouths and expandable jaws that allow them to consume relatively large prey. Some species possess luminous lures, as in female deep-sea anglerfishes (Fig. 9). Metabolic rates and body musculature are generally reduced, reflecting the low availability of food. Many deep-sea species are slow growing, long lived and late to mature.
The deeper ocean and seabed ecosystems are sustained mainly by organic matter exported from surface waters, including marine snow, fecal pellets and occasional large carcasses (see Ocean carbon sink, Coastal and marine sediments). Turbidity currents may also transport organic detritus downslope into the deep ocean. Bacteria consume dissolved organic matter and are subsequently grazed by heterotrophic nanoflagellates, ciliates and other microzooplankton. This recycling pathway is known as the microbial loop.
Increasing depth exposes organisms to strong physiological stress, especially hydrostatic pressure. Pressure increases by approximately 1 atmosphere for every 10 m increase in depth[13].
Pressures
- Overfishing. Industrial fishing has caused major declines in many populations of large pelagic fishes and sharks. While several shark populations have declined by more than 70% globally, many tuna stocks have stabilized or partially recovered under improved international fisheries management.[14][15] More about this subject can be found in the articles on Effects of fisheries on marine biodiversity and Overexploitation.
- Pollution. This causes the loss of many species or a degradation of the environment, see the article Coastal pollution and impacts.
- Alien species. These are species that are introduced from another area and can compete with the indigenous species. This introduction can be done through ballast water from cargo ships or on hulls of vessels. More specific information is given in the article Non-native species invasions.
Global warming
Global warming affects most strongly the animals at the top of the food web and to a lesser degree the lower trophic levels. However, the intermediate trophic levels (zooplankton) can be seriously affected by the indirect effect of cascading trophic interactions from the top of the food web.[16] Superimposed on gradual warming, marine heatwaves can expose open-ocean communities to unusually high temperatures for periods of days to months. Such events can alter plankton composition, bloom timing and food availability, and may cause displacement or increased mortality of temperature-sensitive species. Their ecological consequences depend on season, duration, depth, spatial extent and the sensitivity or mobility of the organisms present.[17][18]
Legal aspect
In the United Nations Convention on the Law of the Sea (UNCLOS 1982), the definition of the open ocean or High Sea is: ‘The high seas are open to all States, whether coastal or land-locked. Freedom of the high seas is exercised under the conditions laid down by this Convention and the rules of international law [19] (see Legislation for the sea). It comprises, inter alia, both for coastal and land-locked States:
- freedom of navigation
- freedom of overflight
- freedom to lay submarine cables and pipelines
- freedom to construct artificial islands and other installations permitted under international law
- freedom of fishing
- freedom of scientific research
Governance of areas beyond national jurisdiction
In areas beyond national jurisdiction, the conservation and sustainable use of marine biodiversity are now also addressed by the Agreement on Biodiversity Beyond National Jurisdiction, commonly called the High Seas Treaty or BBNJ Agreement[20]. This agreement, adopted under the framework of UNCLOS and in force since 17 January 2026, provides for international cooperation on marine genetic resources, area-based management tools including marine protected areas, environmental impact assessments, and capacity-building and transfer of marine technology. For open-ocean habitats, its importance is that it provides a legal framework for conservation and sustainable use of biodiversity in ocean areas that fall outside national jurisdiction.
Related articles
- Shelf sea exchange with the ocean
- Biogeochemical cycles in the coastal marine environment
- Ocean carbon sink
- Deep sea habitat
- Deep sea bottom
- Plankton bloom
- Ekman transport
- Ocean circulation
- Coastal and marine sediments
References
- ↑ Göltenboth, F., Schoppe, S. and Widmann, P. 2006. “Open Ocean.” In: Ecology of Insular SE Asia – The Indonesian Archipelago. Elsevier, pp. 74–81.
- ↑ Grassle, J.F. 2001. Marine ecosystems. Encyclopedia of Biodiversity, Volume 4. Academic Press
- ↑ http://disc.sci.gsfc.nasa.gov/oceancolor/scifocus/oceanColor/oceanblue.shtml
- ↑ Sarmiento, J.L. and Gruber, N. 2006. Ocean Biogeochemical Dynamics. Princeton University Press
- ↑ Sigman, D. M. and Hain, M. P. 2012. The Biological Productivity of the Ocean. Nature Education Knowledge 3(10), 21
- ↑ 6.0 6.1 Masuda, Y., Yamanaka, Y., Smith, S.L., Hirata, T, Nakano, H., Oka, A. and Sumata, H. 2021. Photoacclimation by phytoplankton determines the distribution of global subsurface chlorophyll maxima in the ocean. Communications Earth & Environment 2, 128
- ↑ Falkowski, P. G., Barber, R. T. and Smetacek, V. 1998. Biogeochemical controls and feedbacks on ocean primary production. Science 281: 200–206
- ↑ Pahlow, M. 2005. Linking chlorophyll-nutrient dynamics to the Redfield N:C ratio with a model of optimal phytoplankton growth. Mar. Ecol. Prog. Ser. 287: 33–43
- ↑ 9.0 9.1 Pinet P.R. 1992. Oceanography: An introduction to the Planet Oceanus. West Publishing Company. p. 571
- ↑ Sutton, T.T. and Milligan, R.J. 2019. Deep-Sea Ecology. Encyclopedia of Ecology, 2nd edition, Vol. 1, pp. 35-45
- ↑ http://commons.wikimedia.org/wiki/Image:Portuguese_Man-O-War_(Physalia_physalis).jpg
- ↑ http://en.wikipedia.org/wiki/Copepod
- ↑ Kaiser M. et al. 2005. Marine ecology: Processes, systems and impacts. Oxford University Press. pp.584
- ↑ International Seafood Sustainability Foundation (ISSF) 2023. Status of the World Fisheries for Tuna: March 2023. ISSF Technical Report 2023-01. Pittsburgh, PA, USA, 120 pp.
- ↑ Pacoureau, N., Rigby, C.L., Kyne, P.M., Sherley, R.B., Winker, H., Carlson, J.K., Fordham, S.V., Barreto, R., Fernando, D., Francis, M.P., Jabado, R.W., Herman, K.B., Liu, K.-M., Marshall, A.D., Pollom, R.A., Romanov, E.V., Simpfendorfer, C.A., Yin, J.S., Kindsvater, H.K. and Dulvy, N.K. 2021. Half a century of global decline in oceanic sharks and rays. Nature 589: 567–571
- ↑ Murphy, G.E.P., Romanuk, T.N. and Worm, B. 2020. Cascading effects of climate change on plankton community structure. Ecology and Evolution 10: 2170–2181.
- ↑ Oliver, E.C.J., Donat, M.G., Burrows, M.T., Moore, P.J., Smale, D.A., Alexander, L.V., Benthuysen, J.A., Feng, M., Gupta, A.S., Hobday, A.J., Holbrook, N.J., Perkins-Kirkpatrick, S.E., Scannell, H.A., Straub, S.C. and Wernberg, T. 2018. Longer and more frequent marine heatwaves over the past century. Nature Communications 9: 1324.
- ↑ Holbrook, N.J., Scannell, H.A., Sen Gupta, A., Benthuysen, J.A., Feng, M., Oliver, E.C.J., Alexander, L.V., Burrows, M.T., Donat, M.G., Hobday, A.J., Moore, P.J., Perkins-Kirkpatrick, S.E., Smale, D.A., Straub, S.C. and Wernberg, T. 2019. A global assessment of marine heatwaves and their drivers. Nature Communications 10: 2624.
- ↑ http://www.un.org/Depts/los/convention_agreements/texts/unclos/closindx.htm
- ↑ United Nations, 2023. Agreement under the United Nations Convention on the Law of the Sea on the conservation and sustainable use of marine biological diversity of areas beyond national jurisdiction. New York, 19 June 2023. United Nations Treaty Collection, Chapter XXI, No. 10.
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