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Monday, December 20, 2021

Aquaculture

From Wikipedia, the free encyclopedia

Aquaculture
Aquaculture installations in southern Chile
 
Aquaculture production by region

Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lotus). Aquaculture involves cultivating freshwater and saltwater populations under controlled or semi-natural conditions, and can be contrasted with commercial fishing, which is the harvesting of wild fish. Mariculture, commonly known as marine farming, refers specifically to aquaculture practiced in seawater habitats, opposed to in freshwater aquaculture.

Aquaculture can be conducted in completely artificial facilities built on land (onshore aquaculture), as in the case of fish tank, ponds or raceways, where the living conditions rely on human control; on well-sheltered shallow waters nearshore of a body of water (inshore aquaculture), where the cultivated species are subjected to a relatively more naturalistic environments; or on fenced/enclosed sections of open water away from the shore (offshore aquaculture), where the species are exposed to more diverse natural conditions such as ocean currents, diel vertical migration and nutrient cycles.

According to the Food and Agriculture Organization (FAO), aquaculture "is understood to mean the farming of aquatic organisms including fish, molluscs, crustaceans and aquatic plants. Farming implies some form of intervention in the rearing process to enhance production, such as regular stocking, feeding, protection from predators, etc. Farming also implies individual or corporate ownership of the stock being cultivated." The reported

World capture fisheries and aquaculture production
By species group
 
By main producers (2019)

output from global aquaculture operations in 2014 supplied over one half of the fish and shellfish that is directly consumed by humans; however, there are issues about the reliability of the reported figures. Further, in current aquaculture practice, products from several pounds of wild fish are used to produce one pound of a piscivorous fish like salmon.

Particular kinds of aquaculture include fish farming, shrimp farming, oyster farming, mariculture, algaculture (such as seaweed farming), and the cultivation of ornamental fish. Particular methods include aquaponics and integrated multi-trophic aquaculture, both of which integrate fish farming and aquatic plant farming. The Food and Agriculture Organization describes aquaculture as one of the industries most directly affected by climate change and its impacts. Some forms of aquaculture have negative impacts on the environment, such as through nutrient pollution or disease transfer to wild populations.

Overview

Global capture fisheries and aquaculture production reported by FAO, 1990-2030
 
World aquaculture production of food fish and aquatic plants, 1990–2016.

Harvest stagnation in wild fisheries and overexploitation of popular marine species, combined with a growing demand for high-quality protein, encouraged aquaculturists to domesticate other marine species. At the outset of modern aquaculture, many were optimistic that a "Blue Revolution" could take place in aquaculture, just as the Green Revolution of the 20th century had revolutionized agriculture. Although land animals had long been domesticated, most seafood species were still caught from the wild. Concerned about the impact of growing demand for seafood on the world's oceans, prominent ocean explorer Jacques Cousteau wrote in 1973: "With earth's burgeoning human populations to feed, we must turn to the sea with new understanding and new technology."

About 430 (97%) of the species cultured as of 2007 were domesticated during the 20th and 21st centuries, of which an estimated 106 came in the decade to 2007. Given the long-term importance of agriculture, to date, only 0.08% of known land plant species and 0.0002% of known land animal species have been domesticated, compared with 0.17% of known marine plant species and 0.13% of known marine animal species. Domestication typically involves about a decade of scientific research. Domesticating aquatic species involves fewer risks to humans than do land animals, which took a large toll in human lives. Most major human diseases originated in domesticated animals, including diseases such as smallpox and diphtheria, that like most infectious diseases, move to humans from animals. No human pathogens of comparable virulence have yet emerged from marine species.

Biological control methods to manage parasites are already being used, such as cleaner fish (e.g. lumpsuckers and wrasse) to control sea lice populations in salmon farming. Models are being used to help with spatial planning and siting of fish farms in order to minimize impact.

Aquaculture production (2019)

The decline in wild fish stocks has increased the demand for farmed fish. However, finding alternative sources of protein and oil for fish feed is necessary so the aquaculture industry can grow sustainably; otherwise, it represents a great risk for the over-exploitation of forage fish.

Another recent issue following the banning in 2008 of organotins by the International Maritime Organization is the need to find environmentally friendly, but still effective, compounds with antifouling effects.

Many new natural compounds are discovered every year, but producing them on a large enough scale for commercial purposes is almost impossible.

It is highly probable that future developments in this field will rely on microorganisms, but greater funding and further research is needed to overcome the lack of knowledge in this field.

Species groups

Global aquaculture production in million tonnes, 1950–2010, as reported by the FAO
 
Main species groups
Minor species groups
 
World capture fisheries and aquaculture production by main producers (2018), from FAO's Statistical Yearbook 2020

Aquatic plants

Aquatic plants in floating containers
Cultivating emergent aquatic plants in floating containers
 

Microalgae, also referred to as phytoplankton, microphytes, or planktonic algae, constitute the majority of cultivated algae. Macroalgae commonly known as seaweed also have many commercial and industrial uses, but due to their size and specific requirements, they are not easily cultivated on a large scale and are most often taken in the wild.

In 2016, aquaculture was the source of 96.5 percent by volume of the total 31.2 million tonnes of wild-collected and cultivated aquatic plants combined. Global production of farmed aquatic plants, overwhelmingly dominated by seaweeds, grew in output volume from 13.5 million tonnes in 1995 to just over 30 million tonnes in 2016.

Seaweed farming

Underwater Eucheuma farming in the Philippines
 
A person stands in shallow water, gathering seaweed that has grown on a rope.
A seaweed farmer in Nusa Lembongan (Indonesia) gathers edible seaweed that has grown on a rope.

Seaweed farming or kelp farming is the practice of cultivating and harvesting seaweed. In its simplest form, it consists of the management of naturally found batches. In its most advanced form, it consists of fully controlling the life cycle of the algae.

The top seven most cultivated seaweed taxa are Eucheuma spp., Kappaphycus alvarezii, Gracilaria spp., Saccharina japonica, Undaria pinnatifida, Pyropia spp., and Sargassum fusiforme. Eucheuma and K. alvarezii are farmed for carrageenan (a gelling agent); Gracilaria is farmed for agar; while the rest are farmed for food. The largest seaweed-producing countries are China, Indonesia, and the Philippines. Other notable producers include South Korea, North Korea, Japan, Malaysia, and Zanzibar (Tanzania). Seaweed farming has frequently been developed as an alternative to improve economic conditions and to reduce fishing pressure and overexploited fisheries.

Global production of farmed aquatic plants, overwhelmingly dominated by seaweeds, grew in output volume from 13.5 million tonnes in 1995 to just over 30 million tonnes in 2016. As of 2014, seaweed was 27% of all marine aquaculture. Seaweed farming is a carbon negative crop, with a high potential for climate change mitigation . The IPCC Special Report on the Ocean and Cryosphere in a Changing Climate recommends "further research attention" as a mitigation tactic.

Fish

The farming of fish is the most common form of aquaculture. It involves raising fish commercially in tanks, fish ponds, or ocean enclosures, usually for food. A facility that releases juvenile fish into the wild for recreational fishing or to supplement a species' natural numbers is generally referred to as a fish hatchery. Worldwide, the most important fish species used in fish farming are, in order, carp, salmon, tilapia, and catfish.

In the Mediterranean, young bluefin tuna are netted at sea and towed slowly towards the shore. They are then interned in offshore pens (sometimes made from floating HDPE pipe) where they are further grown for the market. In 2009, researchers in Australia managed for the first time to coax southern bluefin tuna to breed in landlocked tanks. Southern bluefin tuna are also caught in the wild and fattened in grow-out sea cages in southern Spencer Gulf, South Australia.

A similar process is used in the salmon-farming section of this industry; juveniles are taken from hatcheries and a variety of methods are used to aid them in their maturation. For example, as stated above, some of the most important fish species in the industry, salmon, can be grown using a cage system. This is done by having netted cages, preferably in open water that has a strong flow, and feeding the salmon a special food mixture that aids their growth. This process allows for year-round growth of the fish, thus a higher harvest during the correct seasons. An additional method, known sometimes as sea ranching, has also been used within the industry. Sea ranching involves raising fish in a hatchery for a brief time and then releasing them into marine waters for further development, whereupon the fish are recaptured when they have matured.

Crustaceans

Commercial shrimp farming began in the 1970s, and production grew steeply thereafter. Global production reached more than 1.6 million tonnes in 2003, worth about US$9 billion. About 75% of farmed shrimp is produced in Asia, in particular in China and Thailand. The other 25% is produced mainly in Latin America, where Brazil is the largest producer. Thailand is the largest exporter.

Shrimp farming has changed from its traditional, small-scale form in Southeast Asia into a global industry. Technological advances have led to ever higher densities per unit area, and broodstock is shipped worldwide. Virtually all farmed shrimp are penaeids (i.e., shrimp of the family Penaeidae), and just two species of shrimp, the Pacific white shrimp and the giant tiger prawn, account for about 80% of all farmed shrimp. These industrial monocultures are very susceptible to disease, which has decimated shrimp populations across entire regions. Increasing ecological problems, repeated disease outbreaks, and pressure and criticism from both nongovernmental organizations and consumer countries led to changes in the industry in the late 1990s and generally stronger regulations. In 1999, governments, industry representatives, and environmental organizations initiated a program aimed at developing and promoting more sustainable farming practices through the Seafood Watch program.

Freshwater prawn farming shares many characteristics with, including many problems with, marine shrimp farming. Unique problems are introduced by the developmental lifecycle of the main species, the giant river prawn.

The global annual production of freshwater prawns (excluding crayfish and crabs) in 2007 was about 460,000 tonnes, exceeding 1.86 billion dollars. Additionally, China produced about 370,000 tonnes of Chinese river crab.

In addition astaciculture is the freshwater farming of crayfish (mostly in the US, Australia, and Europe).

Molluscs

Abalone farm
Abalone farm
 
Sturgeon farm
Sturgeon farm

Aquacultured shellfish include various oyster, mussel, and clam species. These bivalves are filter and/or deposit feeders, which rely on ambient primary production rather than inputs of fish or other feed. As such, shellfish aquaculture is generally perceived as benign or even beneficial.

Depending on the species and local conditions, bivalve molluscs are either grown on the beach, on longlines, or suspended from rafts and harvested by hand or by dredging. In May 2017 a Belgian consortium installed the first of two trial mussel farms on a wind farm in the North Sea.

Abalone farming began in the late 1950s and early 1960s in Japan and China. Since the mid-1990s, this industry has become increasingly successful. Overfishing and poaching have reduced wild populations to the extent that farmed abalone now supplies most abalone meat. Sustainably farmed molluscs can be certified by Seafood Watch and other organizations, including the World Wildlife Fund (WWF). WWF initiated the "Aquaculture Dialogues" in 2004 to develop measurable and performance-based standards for responsibly farmed seafood. In 2009, WWF co-founded the Aquaculture Stewardship Council with the Dutch Sustainable Trade Initiative to manage the global standards and certification programs.

After trials in 2012, a commercial "sea ranch" was set up in Flinders Bay, Western Australia, to raise abalone. The ranch is based on an artificial reef made up of 5000 (As of April 2016) separate concrete units called abitats (abalone habitats). The 900 kg abitats can host 400 abalone each. The reef is seeded with young abalone from an onshore hatchery. The abalone feed on seaweed that has grown naturally on the habitats, with the ecosystem enrichment of the bay also resulting in growing numbers of dhufish, pink snapper, wrasse, and Samson fish, among other species.

Brad Adams, from the company, has emphasised the similarity to wild abalone and the difference from shore-based aquaculture. "We're not aquaculture, we're ranching, because once they're in the water they look after themselves."

Other groups

Other groups include aquatic reptiles, amphibians, and miscellaneous invertebrates, such as echinoderms and jellyfish. They are separately graphed at the top right of this section, since they do not contribute enough volume to show clearly on the main graph.

Commercially harvested echinoderms include sea cucumbers and sea urchins. In China, sea cucumbers are farmed in artificial ponds as large as 1,000 acres (400 ha).

Global fish production

Global fish production peaked at about 171 million tonnes in 2016, with aquaculture representing 47 percent of the total and 53 percent if non-food uses (including reduction to fishmeal and fish oil) are excluded. With capture fishery production relatively static since the late 1980s, aquaculture has been responsible for the continuing growth in the supply of fish for human consumption. Global aquaculture production (including aquatic plants) in 2016 was 110.2 million tonnes, with the first-sale value estimated at US$243.5 billion. The contribution of aquaculture to the global production of capture fisheries and aquaculture combined has risen continuously, reaching 46.8 percent in 2016, up from 25.7 percent in 2000. With 5.8 percent annual growth rate during the period 2001–2016, aquaculture continues to grow faster than other major food production sectors, but it no longer has the high annual growth rates experienced in the 1980s and 1990s.

In 2012, the total world production of fisheries was 158 million tonnes, of which aquaculture contributed 66.6 million tonnes, about 42%. The growth rate of worldwide aquaculture has been sustained and rapid, averaging about 8% per year for over 30 years, while the take from wild fisheries has been essentially flat for the last decade. The aquaculture market reached $86 billion in 2009.

Aquaculture is an especially important economic activity in China. Between 1980 and 1997, the Chinese Bureau of Fisheries reports, aquaculture harvests grew at an annual rate of 16.7%, jumping from 1.9 million tonnes to nearly 23 million tonnes. In 2005, China accounted for 70% of world production. Aquaculture is also currently one of the fastest-growing areas of food production in the U.S.

About 90% of all U.S. shrimp consumption is farmed and imported. In recent years, salmon aquaculture has become a major export in southern Chile, especially in Puerto Montt, Chile's fastest-growing city.

A United Nations report titled The State of the World Fisheries and Aquaculture released in May 2014 maintained fisheries and aquaculture support the livelihoods of some 60 million people in Asia and Africa. FAO estimates that in 2016, overall, women accounted for nearly 14 percent of all people directly engaged in the fisheries and aquaculture primary sector.

Category 2011 2012 2013 2014 2015 2016
Production





Capture





Inland 10.7 11.2 11.2 11.3 11.4 11.6
Marine 81.5 78.4 79.4 79.9 81.2 79.3
Total capture 92.2 89.5 90.6 91.2 92.7 90.9
Aquaculture





Inland 38.6 42 44.8 46.9 48.6 51.4
Marine 23.2 24.4 25.4 26.8 27.5 28.7
Total aquaculture 61.8 66.4 70.2 73.7 76.1 80
Total world fisheries and aquaculture 154 156 160.7 164.9 168.7 170.9
Utilization





Human consumption 130 136.4 140.1 144.8 148.4 151.2
Non-food uses 24 19.6 20.6 20 20.3 19.7
Population (billions) 7 7.1 7.2 7.3 7.3 7.4
Per capita apparent consumption (kg) 18.5 19.2 19.5 19.9 20.2 20.3

Over-reporting by China

China overwhelmingly dominates the world in reported aquaculture output, reporting a total output which is double that of the rest of the world put together. However, there are some historical issues with the accuracy of China's returns.

In 2001, scientists Reg Watson and Daniel Pauly expressed concerns that China was over reporting its catch from wild fisheries in the 1990s. They said that made it appear that the global catch since 1988 was increasing annually by 300,000 tonnes, whereas it was really shrinking annually by 350,000 tonnes. Watson and Pauly suggested this may be have been related to Chinese policies where state entities that monitored the economy were also tasked with increasing output. Also, until more recently, the promotion of Chinese officials was based on production increases from their own areas.

China disputed this claim. The official Xinhua News Agency quoted Yang Jian, director general of the Agriculture Ministry's Bureau of Fisheries, as saying that China's figures were "basically correct". However, the FAO accepted there were issues with the reliability of China's statistical returns, and for a period treated data from China, including the aquaculture data, apart from the rest of the world.

Aquacultural methods

Mariculture

Mariculture
Mariculture off High Island, Hong Kong
 
Carp are one of the dominant fishes in aquaculture
 
The adaptable tilapia is another commonly farmed fish
 

Mariculture refers to the cultivation of marine organisms in seawater, usually in sheltered coastal or offshore waters. The farming of marine fish is an example of mariculture, and so also is the farming of marine crustaceans (such as shrimp), mollusks (such as oysters), and seaweed. Channel catfish (Ictalurus punctatus), hard clams (Mercenaria mercenaria) and Atlantic salmon (Salmo salar) are prominent in the U.S. mariculture.

Mariculture may consist of raising the organisms on or in artificial enclosures such as in floating netted enclosures for salmon and on racks for oysters. In the case of enclosed salmon, they are fed by the operators; oysters on racks filter feed on naturally available food. Abalone have been farmed on an artificial reef consuming seaweed which grows naturally on the reef units.

Integrated

Integrated multi-trophic aquaculture (IMTA) is a practice in which the byproducts (wastes) from one species are recycled to become inputs (fertilizers, food) for another. Fed aquaculture (for example, fish, shrimp) is combined with inorganic extractive and organic extractive (for example, shellfish) aquaculture to create balanced systems for environmental sustainability (biomitigation), economic stability (product diversification and risk reduction) and social acceptability (better management practices).

"Multi-trophic" refers to the incorporation of species from different trophic or nutritional levels in the same system. This is one potential distinction from the age-old practice of aquatic polyculture, which could simply be the co-culture of different fish species from the same trophic level. In this case, these organisms may all share the same biological and chemical processes, with few synergistic benefits, which could potentially lead to significant shifts in the ecosystem. Some traditional polyculture systems may, in fact, incorporate a greater diversity of species, occupying several niches, as extensive cultures (low intensity, low management) within the same 2006"/> A working IMTA system can result in greater total production based on mutual benefits to the co-cultured species and improved ecosystem health, even if the production of individual species is lower than in a monoculture over a short-term period.

Sometimes the term "integrated aquaculture" is used to describe the integration of monocultures through water transfer. For all intents and purposes, however, the terms "IMTA" and "integrated aquaculture" differ only in their degree of descriptiveness. Aquaponics, fractionated aquaculture, integrated agriculture-aquaculture systems, integrated peri-urban-aquaculture systems, and integrated fisheries-aquaculture systems are other variations of the IMTA concept.

Netting materials

Various materials, including nylon, polyester, polypropylene, polyethylene, plastic-coated welded wire, rubber, patented rope products (Spectra, Thorn-D, Dyneema), galvanized steel and copper are used for netting in aquaculture fish enclosures around the world. All of these materials are selected for a variety of reasons, including design feasibility, material strength, cost, and corrosion resistance.

Recently, copper alloys have become important netting materials in aquaculture because they are antimicrobial (i.e., they destroy bacteria, viruses, fungi, algae, and other microbes) and they therefore prevent biofouling (i.e., the undesirable accumulation, adhesion, and growth of microorganisms, plants, algae, tubeworms, barnacles, mollusks, and other organisms). By inhibiting microbial growth, copper alloy aquaculture cages avoid costly net changes that are necessary with other materials. The resistance of organism growth on copper alloy nets also provides a cleaner and healthier environment for farmed fish to grow and thrive.

Issues

If performed without consideration for potential local environmental impacts, aquaculture in inland waters can result in more environmental damage than wild fisheries, though with less waste produced per kg on a global scale. Local concerns with aquaculture in inland waters may include waste handling, side-effects of antibiotics, competition between farmed and wild animals, and the potential introduction of invasive plant and animal species, or foreign pathogens, particularly if unprocessed fish are used to feed more marketable carnivorous fish. If non-local live feeds are used, aquaculture may introduce exotic plants or animals with disastrous effects. Improvements in methods resulting from advances in research and the availability of commercial feeds has reduced some of these concerns since their greater prevalence in the 1990s and 2000s .

Fish waste is organic and composed of nutrients necessary in all components of aquatic food webs. In-ocean aquaculture often produces much higher than normal fish waste concentrations. The waste collects on the ocean bottom, damaging or eliminating bottom-dwelling life. Waste can also decrease dissolved oxygen levels in the water column, putting further pressure on wild animals. An alternative model to food being added to the ecosystem, is the installation of artificial reef structures to increase the habitat niches available, without the need to add any more than ambient feed and nutrient. This has been used in the "ranching" of abalone in Western Australia.

Impacts on wild fish

Some carnivorous and omnivorous farmed fish species are fed wild forage fish. Although carnivorous farmed fish represented only 13 percent of aquaculture production by weight in 2000, they represented 34 percent of aquaculture production by value.

Farming of carnivorous species like salmon and shrimp leads to a high demand for forage fish to match the nutrition they get in the wild. Fish do not actually produce omega-3 fatty acids, but instead accumulate them from either consuming microalgae that produce these fatty acids, as is the case with forage fish like herring and sardines, or, as is the case with fatty predatory fish, like salmon, by eating prey fish that have accumulated omega-3 fatty acids from microalgae. To satisfy this requirement, more than 50 percent of the world fish oil production is fed to farmed salmon.

Farmed salmon consume more wild fish than they generate as a final product, although the efficiency of production is improving. To produce one pound of farmed salmon, products from several pounds of wild fish are fed to them - this can be described as the "fish-in-fish-out" (FIFO) ratio. In 1995, salmon had a FIFO ratio of 7.5 (meaning 7.5 pounds of wild fish feed were required to produce 1 pound of salmon); by 2006 the ratio had fallen to 4.9. Additionally, a growing share of fish oil and fishmeal come from residues (byproducts of fish processing), rather than dedicated whole fish. In 2012, 34 percent of fish oil and 28 percent of fishmeal came from residues. However, fishmeal and oil from residues instead of whole fish have a different composition with more ash and less protein, which may limit its potential use for aquaculture.

As the salmon farming industry expands, it requires more wild forage fish for feed, at a time when seventy-five percent of the world's monitored fisheries are already near to or have exceeded their maximum sustainable yield. The industrial-scale extraction of wild forage fish for salmon farming then impacts the survivability of the wild predator fish who rely on them for food. An important step in reducing the impact of aquaculture on wild fish is shifting carnivorous species to plant-based feeds. Salmon feeds, for example, have gone from containing only fishmeal and oil to containing 40 percent plant protein. The USDA has also experimented with using grain-based feeds for farmed trout. When properly formulated (and often mixed with fishmeal or oil), plant-based feeds can provide proper nutrition and similar growth rates in carnivorous farmed fish.

Another impact aquaculture production can have on wild fish is the risk of fish escaping from coastal pens, where they can interbreed with their wild counterparts, diluting wild genetic stocks. Escaped fish can become invasive, out-competing native species.

Animal welfare

As with the farming of terrestrial animals, social attitudes influence the need for humane practices and regulations in farmed marine animals. Under the guidelines advised by the Farm Animal Welfare Council good animal welfare means both fitness and a sense of well-being in the animal's physical and mental state. This can be defined by the Five Freedoms:

  • Freedom from hunger & thirst
  • Freedom from discomfort
  • Freedom from pain, disease, or injury
  • Freedom to express normal behaviour
  • Freedom from fear and distress

However, the controversial issue in aquaculture is whether fish and farmed marine invertebrates are actually sentient, or have the perception and awareness to experience suffering. Although no evidence of this has been found in marine invertebrates, recent studies conclude that fish do have the necessary receptors (nociceptors) to sense noxious stimuli and so are likely to experience states of pain, fear and stress. Consequently, welfare in aquaculture is directed at vertebrates; finfish in particular.

Common welfare concerns

Welfare in aquaculture can be impacted by a number of issues such as stocking densities, behavioural interactions, disease and parasitism. A major problem in determining the cause of impaired welfare is that these issues are often all interrelated and influence each other at different times.

Optimal stocking density is often defined by the carrying capacity of the stocked environment and the amount of individual space needed by the fish, which is very species specific. Although behavioural interactions such as shoaling may mean that high stocking densities are beneficial to some species, in many cultured species high stocking densities may be of concern. Crowding can constrain normal swimming behaviour, as well as increase aggressive and competitive behaviours such as cannibalism, feed competition, territoriality and dominance/subordination hierarchies. This potentially increases the risk of tissue damage due to abrasion from fish-to-fish contact or fish-to-cage contact. Fish can suffer reductions in food intake and food conversion efficiency. In addition, high stocking densities can result in water flow being insufficient, creating inadequate oxygen supply and waste product removal. Dissolved oxygen is essential for fish respiration and concentrations below critical levels can induce stress and even lead to asphyxiation. Ammonia, a nitrogen excretion product, is highly toxic to fish at accumulated levels, particularly when oxygen concentrations are low.

Many of these interactions and effects cause stress in the fish, which can be a major factor in facilitating fish disease. For many parasites, infestation depends on the host's degree of mobility, the density of the host population and vulnerability of the host's defence system. Sea lice are the primary parasitic problem for finfish in aquaculture, high numbers causing widespread skin erosion and haemorrhaging, gill congestion, and increased mucus production. There are also a number of prominent viral and bacterial pathogens that can have severe effects on internal organs and nervous systems.

Improving welfare

The key to improving welfare of marine cultured organisms is to reduce stress to a minimum, as prolonged or repeated stress can cause a range of adverse effects. Attempts to minimise stress can occur throughout the culture process. Understanding and providing required environmental enrichment can be vital for reducing stress and benefit aquaculture objects such as improved growth body condition and reduced damage from aggression. During grow-out it is important to keep stocking densities at appropriate levels specific to each species, as well as separating size classes and grading to reduce aggressive behavioural interactions. Keeping nets and cages clean can assist positive water flow to reduce the risk of water degradation.

Not surprisingly disease and parasitism can have a major effect on fish welfare and it is important for farmers not only to manage infected stock but also to apply disease prevention measures. However, prevention methods, such as vaccination, can also induce stress because of the extra handling and injection. Other methods include adding antibiotics to feed, adding chemicals into water for treatment baths and biological control, such as using cleaner wrasse to remove lice from farmed salmon.

Many steps are involved in transport, including capture, food deprivation to reduce faecal contamination of transport water, transfer to transport vehicle via nets or pumps, plus transport and transfer to the delivery location. During transport water needs to be maintained to a high quality, with regulated temperature, sufficient oxygen and minimal waste products. In some cases anaesthetics may be used in small doses to calm fish before transport.

Aquaculture is sometimes part of an environmental rehabilitation program or as an aid in conserving endangered species.

Coastal ecosystems

Aquaculture is becoming a significant threat to coastal ecosystems. About 20 percent of mangrove forests have been destroyed since 1980, partly due to shrimp farming. An extended cost–benefit analysis of the total economic value of shrimp aquaculture built on mangrove ecosystems found that the external costs were much higher than the external benefits. Over four decades, 269,000 hectares (660,000 acres) of Indonesian mangroves have been converted to shrimp farms. Most of these farms are abandoned within a decade because of the toxin build-up and nutrient loss.

Pollution from sea cage aquaculture

Salmon aquaculture, Norway

Salmon farms are typically sited in pristine coastal ecosystems which they then pollute. A farm with 200,000 salmon discharges more fecal waste than a city of 60,000 people. This waste is discharged directly into the surrounding aquatic environment, untreated, often containing antibiotics and pesticides." There is also an accumulation of heavy metals on the benthos (seafloor) near the salmon farms, particularly copper and zinc.

In 2016, mass fish kill events impacted salmon farmers along Chile's coast and the wider ecology. Increases in aquaculture production and its associated effluent were considered to be possible contributing factors to fish and molluscan mortality.

Sea cage aquaculture is responsible for nutrient enrichment of the waters in which they are established. This results from fish wastes and uneaten feed inputs. Elements of most concern are nitrogen and phosphorus which can promote algal growth, including harmful algal blooms which can be toxic to fish. Flushing times, current speeds, distance from the shore and water depth are important considerations when locating sea cages in order to minimize the impacts of nutrient enrichment on coastal ecosystems.

The extent of the effects of pollution from sea-cage aquaculture varies depending on where the cages are located, which species are kept, how densely cages are stocked and what the fish are fed. Important species-specific variables include the species' food conversion ratio (FCR) and nitrogen retention.

Freshwater ecosystems

Whole-lake experiments carried out at the Experimental Lakes Area in Ontario, Canada have displayed the potential for cage aquaculture to source numerous changes in freshwater ecosystems. Following the initiation of an experimental rainbow trout cage farm in a small boreal lake, dramatic reductions in mysis concentrations associated with a decrease in dissolved oxygen were observed. Significant increases in ammonium and total phosphorus, a driver for eutrophication in freshwater systems, were measured in the hypolimnion of the lake. Annual phosphorus inputs from aquaculture waste exceeded that of natural inputs from atmospheric deposition and inflows, and phytoplankton biomass has had a fourfold annual increase following the initiation of the experimental farm.

Genetic modification

A type of salmon called the AquAdvantage salmon has been genetically modified for faster growth, although it has not been approved for commercial use, due to controversy. The altered salmon incorporates a growth hormone from a Chinook salmon that allows it to reach full size in 16–28 months, instead of the normal 36 months for Atlantic salmon, and while consuming 25 percent less feed. The U.S. Food and Drug Administration reviewed the AquAdvantage salmon in a draft environmental assessment and determined that it "would not have a significant impact (FONSI) on the U.S. environment."

Fish diseases, parasites and vaccines

A major difficulty for aquaculture is the tendency towards monoculture and the associated risk of widespread disease. Aquaculture is also associated with environmental risks; for instance, shrimp farming has caused the destruction of important mangrove forests throughout southeast Asia.

In the 1990s, disease wiped out China's farmed Farrer's scallop and white shrimp and required their replacement by other species.

Needs of the aquaculture sector in vaccines

Aquaculture has an average annual growth rate of 9.2%, however the success and continued expansion of the fish farming sector is highly dependent on the control of fish pathogens including a wide range of viruses, bacteria, fungi, and parasites. In 2014, it was estimated that these parasites cost the global salmon farming industry up to 400 million Euros. This represents 6-10% of the production value of the affected countries, but it can go up to 20% (Fisheries and Oceans Canada, 2014). Since pathogens quickly spread within a population of cultured fish, their control is vital for the sector. Historically, the use of antibiotics was against bacterial epizootics but production of animal proteins has to be sustainable, which means that preventive measures that are acceptable from a biological and environmental point of view should be used to keep disease problems in aquaculture at an acceptable level. So, this added to the efficiency of vaccines resulted in an immediate and permanent reduction in the use of antibiotics in the 90s. At the beginning there were fish immersion vaccines efficient against the vibriosis but proved ineffective against the furunculosis, hence the arrival of injectable vaccines: first water-based and after oil-based, much more efficient (Sommerset, 2005).

Development of new vaccines

It is the important mortality in cages among farmed fish, the debates around DNA injection vaccines, although effective, their safety and their side effects but also societal expectations for cleaner fish and security, lead research on new vaccine vectors. Several initiatives are financed by the European Union to develop a rapid and cost-effective approach to using bacteria in feed to make vaccines, in particular thanks to lactic bacteria whose DNA is modified (Boudinot, 2006). In fact, vaccinating farmed fish by injection is time-consuming and costly, so vaccines can be administered orally or by immersion by being added to feed or directly into water. This allows to vaccinate many individuals at the same time, while limiting the associated handling and stress. Indeed, many tests are necessary because the antigens of the vaccines must be adapted to each species or not present a certain level of variability or they will not have any effect. For example, tests have been done with 2 species: Lepeophtheirus salmonis (from which the antigens were collected) and Caligus rogercresseyi (which was vaccinated with the antigens), although the homology between the two species is important, the level of variability made the protection ineffective (Fisheries and Oceans Canada, 2014).

Recent vaccines development in aquaculture

There are 24 vaccines available and one for lobsters. The first vaccine was used in the USA against enteric red mouth in 1976. However, there are 19 companies and some small stakeholders are producing vaccines for aquaculture nowadays. The novel approaches are a way forward to prevent the loss of 10% aquaculture through disease. Genetically modified vaccines are not being used in the EU due to societal concerns and regulations. Meanwhile, DNA vaccines are now authorised in the EU (Adams, 2019). There are challenges in fish vaccine development, immune response due to lack of potent adjScientists are considering microdose application in future. But there are also exciting opportunities in aquaculture vaccinology due to low cost of technology, regulations change and novel antigen expression and delivery systems. In Norway subunit vaccine (VP2 peptide) against infectious pancreatic necrosis is being used. In Canada, a licensed DNA vaccine against Infectious hematopoietic necrosis has been launched for industry use. Fish have large mucosal surfaces, so the preferred route is immersion, intraperitoneal and oral respectively. Nanoparticles are under progress for delivery purposes. The common antibodies produced are IgM and IgT. Normally booster is not required ifn Fish because more memory cells are produced in response to booster rather than increased level of antibodies. mRNA vaccines are alternative to DNA vaccines because they are more safe, stable, easily producible at large scale and mass immunization potential. Recently these are used in cancer prevention and therapeutics. Studies in rabies has shown that efficacy depends on dose and route of administration. These are still in infancy (Adams, 2019).

Economic gains

In 2014, the aquaculture produced fish overtook wild caught fish, in supply for human food. This means there is a huge demand for vaccines, in prevention of diseases. The reported annual loss fish, calculates to >10 billion usd. This is from approximately 10% of all fishes dying from infectious diseases. (Adams, 2019). The high annual losses increases the demand for vaccines. Even though there are about 24 traditionally used vaccines, there is still demand for more vaccines. The breakthrough of DNA-vaccines has sunk the cost of vaccines (Adams, 2019).

The alternative to vaccines would be antibiotics and chemotherapy, which are more expensive and with bigger drawbacks. DNA-vaccines have become the most cost-efficient method of preventing infectious diseases. This bouts well for DNA-vaccines becoming the new standard both in fish vaccines, and in general vaccines (Ragnar Thorarinsson, 2021).

Salinization/acidification of soils

Sediment from abandoned aquaculture farms can remain hypersaline, acidic and eroded. This material can remain unusable for aquaculture purposes for long periods thereafter. Various chemical treatments, such as adding lime, can aggravate the problem by modify the physicochemical characteristics of the sediment.

Ecological benefits

While some forms of aquaculture can be devastating to ecosystems, such as shrimp farming in mangroves, other forms can be very beneficial. Shellfish aquaculture adds substantial filter feeding capacity to an environment which can significantly improve water quality. A single oyster can filter 15 gallons of water a day, removing microscopic algal cells. By removing these cells, shellfish are removing nitrogen and other nutrients from the system and either retaining it or releasing it as waste which sinks to the bottom. By harvesting these shellfish the nitrogen they retained is completely removed from the system. Raising and harvesting kelp and other macroalgae directly remove nutrients such as nitrogen and phosphorus. Repackaging these nutrients can relieve eutrophic, or nutrient-rich, conditions known for their low dissolved oxygen which can decimate species diversity and abundance of marine life. Removing algal cells from the water also increases light penetration, allowing plants such as eelgrass to reestablish themselves and further increase oxygen levels.

Aquaculture in an area can provide for crucial ecological functions for the inhabitants. Shellfish beds or cages can provide habitat structure. This structure can be used as shelter by invertebrates, small fish or crustaceans to potentially increase their abundance and maintain biodiversity. Increased shelter raises stocks of prey fish and small crustaceans by increasing recruitment opportunities in turn providing more prey for higher trophic levels. One study estimated that 10 square meters of oyster reef could enhance an ecosystem's biomass by 2.57 kg The shellfish acting as herbivores will also be preyed on. This moves energy directly from primary producers to higher trophic levels potentially skipping out on multiple energetically-costly trophic jumps which would increase biomass in the ecosystem.

Seaweed farming is a carbon negative crop, with a high potential for climate change mitigation. The IPCC Special Report on the Ocean and Cryosphere in a Changing Climate recommends "further research attention" as a mitigation tactic. Regenerative ocean farming is a polyculture farming system that grows a mix of seaweeds and shellfish while sequestering carbon, decreasing nitrogen in the water and increasing oxygen, helping to regenerate and restore local habitat like reef ecosystems.

Prospects

Global wild fisheries are in decline, with valuable habitat such as estuaries in critical condition. The aquaculture or farming of piscivorous fish, like salmon, does not help the problem because they need to eat products from other fish, such as fish meal and fish oil. Studies have shown that salmon farming has major negative impacts on wild salmon, as well as the forage fish that need to be caught to feed them. Fish that are higher on the food chain are less efficient sources of food energy.

Apart from fish and shrimp, some aquaculture undertakings, such as seaweed and filter-feeding bivalve mollusks like oysters, clams, mussels and scallops, are relatively benign and even environmentally restorative. Filter-feeders filter pollutants as well as nutrients from the water, improving water quality. Seaweeds extract nutrients such as inorganic nitrogen and phosphorus directly from the water, and filter-feeding mollusks can extract nutrients as they feed on particulates, such as phytoplankton and detritus.

Some profitable aquaculture cooperatives promote sustainable practices. New methods lessen the risk of biological and chemical pollution through minimizing fish stress, fallowing netpens, and applying Integrated Pest Management. Vaccines are being used more and more to reduce antibiotic use for disease control.

Onshore recirculating aquaculture systems, facilities using polyculture techniques, and properly sited facilities (for example, offshore areas with strong currents) are examples of ways to manage negative environmental effects.

Recirculating aquaculture systems (RAS) recycle water by circulating it through filters to remove fish waste and food and then recirculating it back into the tanks. This saves water and the waste gathered can be used in compost or, in some cases, could even be treated and used on land. While RAS was developed with freshwater fish in mind, scientists associated with the Agricultural Research Service have found a way to rear saltwater fish using RAS in low-salinity waters. Although saltwater fish are raised in off-shore cages or caught with nets in water that typically has a salinity of 35 parts per thousand (ppt), scientists were able to produce healthy pompano, a saltwater fish, in tanks with a salinity of only 5 ppt. Commercializing low-salinity RAS are predicted to have positive environmental and economical effects. Unwanted nutrients from the fish food would not be added to the ocean and the risk of transmitting diseases between wild and farm-raised fish would greatly be reduced. The price of expensive saltwater fish, such as the pompano and combia used in the experiments, would be reduced. However, before any of this can be done researchers must study every aspect of the fish's lifecycle, including the amount of ammonia and nitrate the fish will tolerate in the water, what to feed the fish during each stage of its lifecycle, the stocking rate that will produce the healthiest fish, etc.

Some 16 countries now use geothermal energy for aquaculture, including China, Israel, and the United States. In California, for example, 15 fish farms produce tilapia, bass, and catfish with warm water from underground. This warmer water enables fish to grow all year round and mature more quickly. Collectively these California farms produce 4.5 million kilograms of fish each year.

National laws, regulations, and management

Laws governing aquaculture practices vary greatly by country and are often not closely regulated or easily traceable.

In the United States, land-based and nearshore aquaculture is regulated at the federal and state levels; however, no national laws govern offshore aquaculture in U.S. exclusive economic zone waters. In June 2011, the Department of Commerce and National Oceanic and Atmospheric Administration released national aquaculture policies to address this issue and "to meet the growing demand for healthy seafood, to create jobs in coastal communities, and restore vital ecosystems." Large aquaculture facilities (i.e. those producing 20,000 pounds (9,100 kg) per year) which discharge wastewater are required to obtain permits pursuant to the Clean Water Act. Facilities that produce at least 100,000 pounds (45,000 kg) of fish, molluscs or crustaceans a year are subject to specific national discharge standards. Other permitted facilities are subject to effluent limitations that are developed on a case-by-case basis.

History

Photo of dripping, cup-shaped net, approximately 6 feet (1.8 m) in diameter and equally tall, half full of fish, suspended from crane boom, with four workers on and around larger, ring-shaped structure in water
Workers harvest catfish from the Delta Pride Catfish farms in Mississippi

The Gunditjmara, the local Aboriginal Australian people in south-western Victoria, Australia, may have raised short-finned eels as early as about 4,580 BCE. Evidence indicates they developed about 100 km2 (39 sq mi) of volcanic floodplains in the vicinity of Lake Condah into a complex of channels and dams, and used woven traps to capture eels, and preserve them to eat all year round. The Budj Bim Cultural Landscape, a World Heritage Site, is thought to be one of the oldest aquaculture sites in the world.

Oral tradition in China tells of the culture of the common carp, Cyprinus carpio, as long ago as 2000–2100 BCE (around 4,000 years BP), but the earliest significant evidence lies in the literature, in the earliest monograph on fish culture called The Classic of Fish Culture, by Fan Li, written around 475 BCE (c.2475 BP). Another ancient Chinese guide to aquaculture was by Yang Yu Jing, written around 460 BCE, showing that carp farming was becoming more sophisticated. The Jiahu site in China has circumstantial archeological evidence as possibly the oldest aquaculture locations, dating from 6200BCE (about 8,200 years BP), but this is speculative. When the waters subsided after river floods, some fish, mainly carp, were trapped in lakes. Early aquaculturists fed their brood using nymphs and silkworm faeces, and ate them.

Ancient Egyptians might have farmed fish (especially Gilt-head bream) from Lake Bardawil about 1,500 BCE (3,520 years BP), and they traded them with Canaan.

Gim cultivation is the oldest aquaculture in Korea. Early cultivation methods used bamboo or oak sticks, which were replaced by newer methods that utilized nets in the 19th century. Floating rafts have been used for mass production since the 1920s.

Japanese cultivated seaweed by providing bamboo poles and, later, nets and oyster shells to serve as anchoring surfaces for spores.

Romans bred fish in ponds and farmed oysters in coastal lagoons before 100 CE.

In central Europe, early Christian monasteries adopted Roman aquacultural practices. Aquaculture spread in Europe during the Middle Ages since away from the seacoasts and the big rivers, fish had to be salted so they did not rot. Improvements in transportation during the 19th century made fresh fish easily available and inexpensive, even in inland areas, making aquaculture less popular. The 15th-century fishponds of the Trebon Basin in the Czech Republic are maintained as a UNESCO World Heritage Site.

Hawaiians constructed oceanic fish ponds. A remarkable example is the "Menehune" fishpond dating from at least 1,000 years ago, at Alekoko. Legend says that it was constructed by the mythical Menehune dwarf people.

In the first half of the 18th century, German Stephan Ludwig Jacobi experimented with external fertilization of brown trouts and salmon. He wrote an article "Von der künstlichen Erzeugung der Forellen und Lachse" (On the Artificial Production of Trout and Salmon) summarizing his findings, and is regarded as the founder of artificial fish rearing in Europe. By the latter decades of the 18th century, oyster farming had begun in estuaries along the Atlantic Coast of North America.

The word aquaculture appeared in an 1855 newspaper article in reference to the harvesting of ice. It also appeared in descriptions of the terrestrial agricultural practise of sub-irrigation in the late 19th century before becoming associated primarily with the cultivation of aquatic plant and animal species.

In 1859, Stephen Ainsworth of West Bloomfield, New York, began experiments with brook trout. By 1864, Seth Green had established a commercial fish-hatching operation at Caledonia Springs, near Rochester, New York. By 1866, with the involvement of Dr. W. W. Fletcher of Concord, Massachusetts, artificial fish hatcheries were underway in both Canada and the United States. When the Dildo Island fish hatchery opened in Newfoundland in 1889, it was the largest and most advanced in the world. The word aquaculture was used in descriptions of the hatcheries experiments with cod and lobster in 1890.

By the 1920s, the American Fish Culture Company of Carolina, Rhode Island, founded in the 1870s was one of the leading producers of trout. During the 1940s, they had perfected the method of manipulating the day and night cycle of fish so that they could be artificially spawned year around.

Californians harvested wild kelp and attempted to manage supply around 1900, later labeling it a wartime resource.

Deep-sea fish

From Wikipedia, the free encyclopedia
 

Deep-sea fish are animals that live in the darkness below the sunlit surface waters, that is below the epipelagic or photic zone of the sea. The lanternfish is, by far, the most common deep-sea fish. Other deep sea fishes include the flashlight fish, cookiecutter shark, bristlemouths, anglerfish, viperfish, and some species of eelpout.

Only about 2% of known marine species inhabit the pelagic environment. This means that they live in the water column as opposed to the benthic organisms that live in or on the sea floor. Deep-sea organisms generally inhabit bathypelagic (1000–4000m deep) and abyssopelagic (4000–6000m deep) zones. However, characteristics of deep-sea organisms, such as bioluminescence can be seen in the mesopelagic (200–1000m deep) zone as well. The mesopelagic zone is the disphotic zone, meaning light there is minimal but still measurable. The oxygen minimum layer exists somewhere between a depth of 700m and 1000m deep depending on the place in the ocean. This area is also where nutrients are most abundant. The bathypelagic and abyssopelagic zones are aphotic, meaning that no light penetrates this area of the ocean. These zones make up about 75% of the inhabitable ocean space.

The epipelagic zone (0–200m) is the area where light penetrates the water and photosynthesis occurs. This is also known as the photic zone. Because this typically extends only a few hundred meters below the water, the deep sea, about 90% of the ocean volume, is in darkness. The deep sea is also an extremely hostile environment, with temperatures that rarely exceed 3 °C (37.4 °F) and fall as low as −1.8 °C (28.76 °F) (with the exception of hydrothermal vent ecosystems that can exceed 350 °C, or 662 °F), low oxygen levels, and pressures between 20 and 1,000 atmospheres (between 2 and 100 megapascals).

Environment

Scale diagram of the layers of the pelagic zone

In the deep ocean, the waters extend far below the epipelagic zone, and support very different types of pelagic fishes adapted to living in these deeper zones. In deep water, marine snow is a continuous shower of mostly organic detritus falling from the upper layers of the water column. Its origin lies in activities within the productive photic zone. Marine snow includes dead or dying plankton, protists (diatoms), fecal matter, sand, soot and other inorganic dust. The "snowflakes" grow over time and may reach several centimetres in diameter, travelling for weeks before reaching the ocean floor. However, most organic components of marine snow are consumed by microbes, zooplankton and other filter-feeding animals within the first 1,000 metres of their journey, that is, within the epipelagic zone. In this way marine snow may be considered the foundation of deep-sea mesopelagic and benthic ecosystems: As sunlight cannot reach them, deep-sea organisms rely heavily on marine snow as an energy source. Since there is no light in the deep sea (aphotic), there is a lack of primary producers. Therefore, most organisms in the bathypelagic rely on the marine snow from regions higher in the vertical column.

Some deep-sea pelagic groups, such as the lanternfish, ridgehead, marine hatchetfish, and lightfish families are sometimes termed pseudoceanic because, rather than having an even distribution in open water, they occur in significantly higher abundances around structural oases, notably seamounts and over continental slopes. The phenomenon is explained by the likewise abundance of prey species which are also attracted to the structures.

Hydrostatic pressure increases by 1 atmosphere for every 10m in depth. Deep-sea organisms have the same pressure within their bodies as is exerted on them from the outside, so they are not crushed by the extreme pressure. Their high internal pressure, however, results in the reduced fluidity of their membranes because molecules are squeezed together. Fluidity in cell membranes increases efficiency of biological functions, most importantly the production of proteins, so organisms have adapted to this circumstance by increasing the proportion of unsaturated fatty acids in the lipids of the cell membranes. In addition to differences in internal pressure, these organisms have developed a different balance between their metabolic reactions from those organisms that live in the epipelagic zone. David Wharton, author of Life at the Limits: Organisms in Extreme Environments, notes "Biochemical reactions are accompanied by changes in volume. If a reaction results in an increase in volume, it will be inhibited by pressure, whereas, if it is associated with a decrease in volume, it will be enhanced". This means that their metabolic processes must ultimately decrease the volume of the organism to some degree.

Humans seldom encounter frilled sharks alive, so they pose little danger (though scientists have accidentally cut themselves examining their teeth).

Most fish that have evolved in this harsh environment are not capable of surviving in laboratory conditions, and attempts to keep them in captivity have led to their deaths. Deep-sea organisms contain gas-filled spaces (vacuoles). Gas is compressed under high pressure and expands under low pressure. Because of this, these organisms have been known to blow up if they come to the surface.

Characteristics

An annotated diagram of the basic external features of an abyssal grenadier and standard length measurements.
 
Rhinochimera atlantica
 
Gigantactis is a deep-sea fish with a dorsal fin whose first filament has become very long and is tipped with a bioluminescent photophore lure.
 
Bigeye tuna cruise the epipelagic zone at night and the mesopelagic zone during the day

The fish of the deep-sea have evolved various adaptations to survive in this region. Since many of these fish live in regions where there is no natural illumination, they cannot rely solely on their eyesight for locating prey and mates and avoiding predators; deep-sea fish have evolved appropriately to the extreme sub-photic region in which they live. Many of these organisms are blind and rely on their other senses, such as sensitivities to changes in local pressure and smell, to catch their food and avoid being caught. Those that aren't blind have large and sensitive eyes that can use bioluminescent light. These eyes can be as much as 100 times more sensitive to light than human eyes. Rhodopsin (Rh1) is a protein found in the eye’s rod cells that helps animals see in dim light. While most vertebrates usually have one Rh1 opsin gene, some deep sea fish have several Rh1 genes, and one species, the silver spinyfin (Diretmus argenteus), has 38. This proliferation of Rh1 genes may help deep sea fish to see in the depths of the ocean. Also, to avoid predation, many species are dark to blend in with their environment.

Many deep-sea fish are bioluminescent, with extremely large eyes adapted to the dark. Bioluminescent organisms are capable of producing light biologically through the agitation of molecules of luciferin, which then produce light. This process must be done in the presence of oxygen. These organisms are common in the mesopelagic region and below (200m and below). More than 50% of deep-sea fish, as well as some species of shrimp and squid, are capable of bioluminescence. About 80% of these organisms have photophores – light producing glandular cells that contain luminous bacteria bordered by dark colourings. Some of these photophores contain lenses, much like those in the eyes of humans, which can intensify or lessen the emanation of light. The ability to produce light only requires 1% of the organism's energy and has many purposes: It is used to search for food and attract prey, like the anglerfish; claim territory through patrol; communicate and find a mate, and distract or temporarily blind predators to escape. Also, in the mesopelagic where some light still penetrates, some organisms camouflage themselves from predators below them by illuminating their bellies to match the colour and intensity of light from above so that no shadow is cast. This tactic is known as counter-illumination.

The lifecycle of deep-sea fish can be exclusively deep water although some species are born in shallower water and sink upon maturation. Regardless of the depth where eggs and larvae reside, they are typically pelagic. This planktonic — drifting — lifestyle requires neutral buoyancy. In order to maintain this, the eggs and larvae often contain oil droplets in their plasma. When these organisms are in their fully matured state they need other adaptations to maintain their positions in the water column. In general, water's density causes upthrust — the aspect of buoyancy that makes organisms float. To counteract this, the density of an organism must be greater than that of the surrounding water. Most animal tissues are denser than water, so they must find an equilibrium to make them float. Many organisms develop swim bladders (gas cavities) to stay afloat, but because of the high pressure of their environment, deep-sea fishes usually do not have this organ. Instead they exhibit structures similar to hydrofoils in order to provide hydrodynamic lift. It has also been found that the deeper a fish lives, the more jelly-like its flesh and the more minimal its bone structure. They reduce their tissue density through high fat content, reduction of skeletal weight — accomplished through reductions of size, thickness and mineral content — and water accumulation makes them slower and less agile than surface fish.

Due to the poor level of photosynthetic light reaching deep-sea environments, most fish need to rely on organic matter sinking from higher levels, or, in rare cases, hydrothermal vents for nutrients. This makes the deep-sea much poorer in productivity than shallower regions. Also, animals in the pelagic environment are sparse and food doesn't come along frequently. Because of this, organisms need adaptations that allow them to survive. Some have long feelers to help them locate prey or attract mates in the pitch black of the deep ocean. The deep-sea angler fish in particular has a long fishing-rod-like adaptation protruding from its face, on the end of which is a bioluminescent piece of skin that wriggles like a worm to lure its prey. Some must consume other fish that are the same size or larger than them and they need adaptations to help digest them efficiently. Great sharp teeth, hinged jaws, disproportionately large mouths, and expandable bodies are a few of the characteristics that deep-sea fishes have for this purpose. The gulper eel is one example of an organism that displays these characteristics.

Fish in the different pelagic and deep water benthic zones are physically structured, and behave in ways, that differ markedly from each other. Groups of coexisting species within each zone all seem to operate in similar ways, such as the small mesopelagic vertically migrating plankton-feeders, the bathypelagic anglerfishes, and the deep water benthic rattails.

Ray finned species, with spiny fins, are rare among deep sea fishes, which suggests that deep sea fish are ancient and so well adapted to their environment that invasions by more modern fishes have been unsuccessful. The few ray fins that do exist are mainly in the Beryciformes and Lampriformes, which are also ancient forms. Most deep sea pelagic fishes belong to their own orders, suggesting a long evolution in deep sea environments. In contrast, deep water benthic species, are in orders that include many related shallow water fishes.

Mesopelagic fish

Mesopelagic fish
Most mesopelagic fishes are small filter feeders which ascend at night to feed in the nutrient rich waters of the epipelagic zone. During the day, they return to the dark, cold, oxygen deficient waters of the mesopelagic where they are relatively safe from predators. Lanternfish account for as much as 65% of all deep sea fish biomass and are largely responsible for the deep scattering layer of the world's oceans.
 
Most of the rest of the mesopelagic fishes are ambush predators, like this sabertooth fish which uses its telescopic, upward-pointing eyes to pick out prey silhouetted against the gloom above. Their recurved teeth prevent a captured fish from backing out.
 
The Antarctic toothfish have large, upward looking eyes, adapted to detecting the silhouettes of prey fish.
 
The barreleye has barrel-shaped, tubular eyes which are generally directed upwards but can be swivelled forward
 
The telescopefish has large, forward-pointing telescoping eyes with large lenses

Below the epipelagic zone, conditions change rapidly. Between 200 metres and about 1000 metres, light continues to fade until there is almost none. Temperatures fall through a thermocline to temperatures between 3.9 °C (39 °F) and 7.8 °C (46 °F). This is the twilight or mesopelagic zone. Pressure continues to increase, at the rate of one atmosphere every 10 metres, while nutrient concentrations fall, along with dissolved oxygen and the rate at which the water circulates.

Sonar operators, using the newly developed sonar technology during World War II, were puzzled by what appeared to be a false sea floor 300–500 metres deep at day, and less deep at night. This turned out to be due to millions of marine organisms, most particularly small mesopelagic fish, with swim bladders that reflected the sonar. These organisms migrate up into shallower water at dusk to feed on plankton. The layer is deeper when the moon is out, and can become shallower when clouds pass over the moon. This phenomenon has come to be known as the deep scattering layer.

Most mesopelagic fish make daily vertical migrations, moving at night into the epipelagic zone, often following similar migrations of zooplankton, and returning to the depths for safety during the day. These vertical migrations often occur over large vertical distances, and are undertaken with the assistance of a swim bladder. The swim bladder is inflated when the fish wants to move up, and, given the high pressures in the messoplegic zone, this requires significant energy. As the fish ascends, the pressure in the swim bladder must adjust to prevent it from bursting. When the fish wants to return to the depths, the swim bladder is deflated. Some mesopelagic fishes make daily migrations through the thermocline, where the temperature changes between 50 °F (10 °C) and 69 °F (20 °C), thus displaying considerable tolerances for temperature change.

These fish have muscular bodies, ossified bones, scales, well developed gills and central nervous systems, and large hearts and kidneys. Mesopelagic plankton feeders have small mouths with fine gill rakers, while the piscivores have larger mouths and coarser gill rakers.

Mesopelagic fish are adapted for an active life under low light conditions. Most of them are visual predators with large eyes. Some of the deeper water fish have tubular eyes with big lenses and only rod cells that look upwards. These give binocular vision and great sensitivity to small light signals. This adaptation gives improved terminal vision at the expense of lateral vision, and allows the predator to pick out squid, cuttlefish, and smaller fish that are silhouetted against the gloom above them.

Mesopelagic fish usually lack defensive spines, and use colour to camouflage themselves from other fish. Ambush predators are dark, black or red. Since the longer, red, wavelengths of light do not reach the deep sea, red effectively functions the same as black. Migratory forms use countershaded silvery colours. On their bellies, they often display photophores producing low grade light. For a predator from below, looking upwards, this bioluminescence camouflages the silhouette of the fish. However, some of these predators have yellow lenses that filter the (red deficient) ambient light, leaving the bioluminescence visible.

The brownsnout spookfish, a species of barreleye, is the only vertebrate known to employ a mirror, as opposed to a lens, to focus an image in its eyes.

Sampling by deep trawling indicates that lanternfish account for as much as 65% of all deep-sea fish biomass. Indeed, lanternfish are among the most widely distributed, populous, and diverse of all vertebrates, playing an important ecological role as prey for larger organisms. The estimated global biomass of lanternfish is 550–660 million tonnes, several times the entire world fisheries catch. Lanternfish also account for much of the biomass responsible for the deep scattering layer of the world's oceans.

Bigeye tuna are an epipelagic/mesopelagic species that eats other fish. Satellite tagging has shown that bigeye tuna often spend prolonged periods cruising deep below the surface during the daytime, sometimes making dives as deep as 500 metres. These movements are thought to be in response to the vertical migrations of prey organisms in the deep scattering layer.

Bathypelagic fish

Bathypelagic fish
The humpback anglerfish is a bathypelagic ambush predator, which attracts prey with a bioluminescent lure. It can ingest prey larger than itself, which it swallows with an inrush of water when it opens its mouth.
 
Many bristlemouth species, such as the "spark anglemouth" above, are also bathypelagic ambush predators which can swallow prey larger than themselves. They are among the most abundant of all vertebrate families.
 
Young, red flabby whalefish make nightly vertical migrations into the lower mesopelagic zone to feed on copepods. When males make the transition to adults, they develop a massive liver, and then their jaws fuse shut. They no longer eat, but continue to metabolise the energy stored in their liver.
 
The Sloane's viperfish can make nightly migrations from bathypelagic depths to near surface waters.
 
The widespread fangtooth has the largest teeth of any fish, proportionate to body size. Despite their ferocious appearance, bathypelagic fish are usually weakly muscled and too small to represent any threat to humans.

Below the mesopelagic zone it is pitch dark. This is the midnight (or bathypelagic zone), extending from 1000 metres to the bottom deep water benthic zone. If the water is exceptionally deep, the pelagic zone below 4000 metres is sometimes called the lower midnight (or abyssopelagic zone). Temperatures in this zone range from 1 to 4 degrees celsius and is completely aphotic.

Conditions are somewhat uniform throughout these zones; the darkness is complete, the pressure is crushing, and temperatures, nutrients and dissolved oxygen levels are all low.

Bathypelagic fish have special adaptations to cope with these conditions – they have slow metabolisms and unspecialized diets, being willing to eat anything that comes along. They prefer to sit and wait for food rather than waste energy searching for it. The behaviour of bathypelagic fish can be contrasted with the behaviour of mesopelagic fish. Mesopelagic fish are often highly mobile, whereas bathypelagic fish are almost all lie-in-wait predators, normally expending little energy in movement.

The dominant bathypelagic fishes are small bristlemouth and anglerfish; fangtooth, viperfish, daggertooth and barracudina are also common. These fishes are small, many about 10 centimetres long, and not many longer than 25 cm. They spend most of their time waiting patiently in the water column for prey to appear or to be lured by their phosphors. What little energy is available in the bathypelagic zone filters from above in the form of detritus, faecal material, and the occasional invertebrate or mesopelagic fish. About 20 percent of the food that has its origins in the epipelagic zone falls down to the mesopelagic zone, but only about 5 percent filters down to the bathypelagic zone.

Bathypelagic fish are sedentary, adapted to outputting minimum energy in a habitat with very little food or available energy, not even sunlight, only bioluminescence. Their bodies are elongated with weak, watery muscles and skeletal structures. Since so much of the fish is water, they are not compressed by the great pressures at these depths. They often have extensible, hinged jaws with recurved teeth. They are slimy, without scales. The central nervous system is confined to the lateral line and olfactory systems, the eyes are small and may not function, and gills, kidneys and hearts, and swim bladders are small or missing.

These are the same features found in fish larvae, which suggests that during their evolution, bathypelagic fish have acquired these features through neoteny. As with larvae, these features allow the fish to remain suspended in the water with little expenditure of energy.

Despite their ferocious appearance, these beasts of the deep are mostly miniature fish with weak muscles, and are too small to represent any threat to humans.

The swim bladders of deep sea fish are either absent or scarcely operational, and bathypelagic fish do not normally undertake vertical migrations. Filling bladders at such great pressures incurs huge energy costs. Some deep sea fishes have swim bladders which function while they are young and inhabit the upper epipelagic zone, but they wither or fill with fat when the fish move down to their adult habitat.

The most important sensory systems are usually the inner ear, which responds to sound, and the lateral line, which responds to changes in water pressure. The olfactory system can also be important for males who find females by smell. Bathypelagic fish are black, or sometimes red, with few photophores. When photophores are used, it is usually to entice prey or attract a mate. Because food is so scarce, bathypelagic predators are not selective in their feeding habits, but grab whatever comes close enough. They accomplish this by having a large mouth with sharp teeth for grabbing large prey and overlapping gill rakers which prevent small prey that have been swallowed from escaping.

It is not easy finding a mate in this zone. Some species depend on bioluminescence, where bioluminescent patterns are unique to specific species. Others are hermaphrodites, which doubles their chances of producing both eggs and sperm when an encounter occurs. The female anglerfish releases pheromones to attract tiny males. When a male finds her, he bites on to her and never lets go. When a male of the anglerfish species Haplophryne mollis bites into the skin of a female, he releases an enzyme that digests the skin of his mouth and her body, fusing the pair to the point where the two circulatory systems join up. The male then atrophies into nothing more than a pair of gonads. This extreme sexual dimorphism ensures that, when the female is ready to spawn, she has a mate immediately available.

Many forms other than fish live in the bathypelagic zone, such as squid, large whales, octopuses, sponges, brachiopods, sea stars, and echinoids, but this zone is difficult for fish to live in.

Adaptation to high pressure

As a fish moves deeper into the sea, the weight of the water overhead exerts increasing hydrostatic pressure on the fish. This increased pressure amounts to about one standard atmosphere for every 10 meters in depth (one atmosphere is the pressure exerted at the surface of the sea by the atmosphere above). For a fish at the bottom of the bathypelagic zone, this pressure amounts to about 400 atmospheres (nearly 6000 pounds per square inch).

Deep sea organisms possess adaptations at cellular and physiological levels that allow them to survive in environments of great pressure. Not having these adaptations limits the depths at which shallow-water species can operate. High levels of external pressure affects how metabolic processes and biochemical reactions proceed. The equilibrium of many chemical reactions is disturbed by pressure, and pressure can inhibit processes which result in an increase in volume. Water, a key component in many biological processes, is very susceptible to volume changes, mainly because constituents of cellular fluid have an effect on water structure. Thus, enzymatic reactions that induce changes in water organization effectively change the system's volume. Proteins responsible for catalyzing reactions are typically held together by weak bonds and the reactions usually involve volume increases.

To adapt to this change, the protein structure and reaction criteria of deep sea fish have been adapted to withstand pressure in order to perform reactions in these conditions. In high pressure environments, bilayer cellular membranes experience a loss of fluidity. Deep-sea cellular membranes favor phospholipid bilayers with a higher proportion of unsaturated fatty acids, which induce a higher fluidity than their sea-level counterparts.

The rattail Coryphaenoides armatus (abyssal grenadier) on the Davidson Seamount at 2253 meters depth.

Deep sea species exhibit lower changes of entropy and enthalpy compared to surface level organisms, since a high pressure and low temperature environment favors negative enthalpy changes and reduced dependence on entropy-driven reactions. From a structural standpoint, globular proteins of deep sea fish due to the tertiary structure of G-actin are relatively rigid compared to those of surface level fish. The fact that proteins in deep sea fish are structurally different from surface fish is apparent from the observation that actin from the muscle fibers of deep sea fish are extremely heat resistant; an observation similar to what is found in lizards. These proteins are structurally strengthened by modification of the bonds in the tertiary structure of the protein which also happens to induce high levels of thermal stability. Proteins are structurally strengthened to resist pressure by modification of bonds in the tertiary structure. Therefore, high levels of hydrostatic pressure, similar to high body temperatures of thermophilic desert reptiles, favor rigid protein structures.

Na+/K+ -ATPase is a lipoprotein enzyme that plays a prominent role in osmoregulation and is heavily influenced by hydrostatic pressure. The inhibition of Na+/K+ -ATPase is due to increased compression due to pressure. The rate-limiting step of the Na+/K+ -ATPase reaction induces an expansion in the bilayer surrounding the protein, and therefore an increase in volume. An increase in volume makes Na+/K+ -ATPase reactivity susceptible to higher pressures. Even though the Na+/K+ -ATPase activity per gram of gill tissue is lower for deep sea fishes, the Na+/K+ -ATPases of deep sea fishes exhibit a much higher tolerance of hydrostatic pressure compared to their shallow-water counterparts. This is exemplified between the species C. acrolepis (around 2000m deep) and its hadalpelagic counterpart C. armatus (around 4000m deep), where the Na+/K+ -ATPases of C. armatus are much less sensitive to pressure. This resistance to pressure can be explained by adaptations in the protein and lipid moieties of Na+/K+ -ATPase.

Lanternfish

Lantern fish

Sampling via deep trawling indicates that lanternfish account for as much as 65% of all deep-sea fish biomass. Indeed, lanternfish are among the most widely distributed, populous, and diverse of all vertebrates, playing an important ecological role as prey for larger organisms. With an estimated global biomass of 550–660 million metric tons, several times the entire world fisheries catch, lanternfish also account for much of the biomass responsible for the deep scattering layer of the world's oceans. In the Southern Ocean, Myctophids provide an alternative food resource to krill for predators such as squid and the king penguin. Although these fish are plentiful and prolific, currently only a few commercial lanternfish fisheries exist: these include limited operations off South Africa, in the sub-Antarctic, and in the Gulf of Oman.

Endangered species

A 2006 study by Canadian scientists has found five species of deep-sea fish – blue hake, spiny eel – to be on the verge of extinction due to the shift of commercial fishing from continental shelves to the slopes of the continental shelves, down to depths of 1600 meters. The slow reproduction of these fish – they reach sexual maturity at about the same age as human beings – is one of the main reasons that they cannot recover from the excessive fishing.

 

Intertidal ecology

From Wikipedia, the free encyclopedia
 
Anjajavy Forest on Tsingy rocks jutting into the Indian Ocean.

Intertidal ecology is the study of intertidal ecosystems, where organisms live between the low and high tide lines. At low tide, the intertidal is exposed whereas at high tide, the intertidal is underwater. Intertidal ecologists therefore study the interactions between intertidal organisms and their environment, as well as between different species of intertidal organisms within a particular intertidal community. The most important environmental and species interactions may vary based on the type of intertidal community being studied, the broadest of classifications being based on substrates—rocky shore and soft bottom communities.

Organisms living in this zone have a highly variable and often hostile environment, and have evolved various adaptations to cope with and even exploit these conditions. One easily visible feature of intertidal communities is vertical zonation, where the community is divided into distinct vertical bands of specific species going up the shore. Species ability to cope with abiotic factors associated with emersion stress, such as desiccation determines their upper limits, while biotic interactions e.g.competition with other species sets their lower limits.

Intertidal regions are utilized by humans for food and recreation, but anthropogenic actions also have major impacts, with overexploitation, invasive species and climate change being among the problems faced by intertidal communities. In some places Marine Protected Areas have been established to protect these areas and aid in scientific research.

Types of intertidal communities

Intertidal habitats can be characterized as having either hard or soft bottoms substrates. Rocky intertidal communities occur on rocky shores, such as headlands, cobble beaches, or human-made jetties. Their degree of exposure may be calculated using the Ballantine Scale. Soft-sediment habitats include sandy beaches, and intertidal wetlands (e.g., mudflats and salt marshes). These habitats differ in levels of abiotic, or non-living, environmental factors. Rocky shores tend to have higher wave action, requiring adaptations allowing the inhabitants to cling tightly to the rocks. Soft-bottom habitats are generally protected from large waves but tend to have more variable salinity levels. They also offer a third habitable dimension: depth. Thus, many soft-sediment inhabitants are adapted for burrowing.

Environment

A rock, seen at low tide, exhibiting typical intertidal zonation.
 
A specimen of the shell Pinna nobilis exposed by low tide

Because intertidal organisms endure regular periods of immersion and emersion, they essentially live both underwater and on land and must be adapted to a large range of climatic conditions. The intensity of climate stressors varies with relative tide height because organisms living in areas with higher tide heights are emersed for longer periods than those living in areas with lower tide heights. This gradient of climate with tide height leads to patterns of intertidal zonation, with high intertidal species being more adapted to emersion stresses than low intertidal species. These adaptations may be behavioral (i.e. movements or actions), morphological (i.e. characteristics of external body structure), or physiological (i.e. internal functions of cells and organs). In addition, such adaptations generally cost the organism in terms of energy (e.g. to move or to grow certain structures), leading to trade-offs (i.e. spending more energy on deterring predators leaves less energy for other functions like reproduction).

Intertidal organisms, especially those in the high intertidal, must cope with a large range of temperatures. While they are underwater, temperatures may only vary by a few degrees over the year. However, at low tide, temperatures may dip to below freezing or may become scaldingly hot, leading to a temperature range that may approach 30 °C (86 °F) during a period of a few hours. Many mobile organisms, such as snails and crabs, avoid temperature fluctuations by crawling around and searching for food at high tide and hiding in cool, moist refuges (crevices or burrows) at low tide. Besides simply living at lower tide heights, non-motile organisms may be more dependent on coping mechanisms. For example, high intertidal organisms have a stronger stress response, a physiological response of making proteins that help recovery from temperature stress just as the immune response aids in the recovery from infection.

Intertidal organisms are also especially prone to desiccation during periods of emersion. Again, mobile organisms avoid desiccation in the same way as they avoid extreme temperatures: by hunkering down in mild and moist refuges. Many intertidal organisms, including Littorina snails, prevent water loss by having waterproof outer surfaces, pulling completely into their shells, and sealing shut their shell opening. Limpets (Patella) do not use such a sealing plate but occupy a home-scar to which they seal the lower edge of their flattened conical shell using a grinding action. They return to this home-scar after each grazing excursion, typically just before emersion. On soft rocks, these scars are quite obvious. Still other organisms, such as the algae Ulva and Porphyra, are able to rehydrate and recover after periods of severe desiccation.

The level of salinity can also be quite variable. Low salinities can be caused by rainwater or river inputs of freshwater. Estuarine species must be especially euryhaline, or able to tolerate a wide range of salinities. High salinities occur in locations with high evaporation rates, such as in salt marshes and high intertidal pools. Shading by plants, especially in the salt marsh, can slow evaporation and thus ameliorate salinity stress. In addition, salt marsh plants tolerate high salinities by several physiological mechanisms, including excreting salt through salt glands and preventing salt uptake into the roots.

In addition to these exposure stresses (temperature, desiccation, and salinity), intertidal organisms experience strong mechanical stresses, especially in locations of high wave action. There are myriad ways in which the organisms prevent dislodgement due to waves. Morphologically, many mollusks (such as limpets and chitons) have low-profile, hydrodynamic shells. Types of substrate attachments include mussels' tethering byssal threads and glues, sea stars' thousands of suctioning tube feet, and isopods' hook-like appendages that help them hold on to intertidal kelps. Higher profile organisms, such as kelps, must also avoid breaking in high flow locations, and they do so with their strength and flexibility. Finally, organisms can also avoid high flow environments, such as by seeking out low flow microhabitats. Additional forms of mechanical stresses include ice and sand scour, as well as dislodgment by water-borne rocks, logs, etc.

For each of these climate stresses, species exist that are adapted to and thrive in the most stressful of locations. For example, the tiny crustacean copepod Tigriopus thrives in very salty, high intertidal tidepools, and many filter feeders find more to eat in wavier and higher flow locations. Adapting to such challenging environments gives these species competitive edges in such locations.

Food web structure

During tidal immersion, the food supply to intertidal organisms is subsidized by materials carried in seawater, including photosynthesizing phytoplankton and consumer zooplankton. These plankton are eaten by numerous forms of filter feedersmussels, clams, barnacles, sea squirts, and polychaete worms—which filter seawater in their search for planktonic food sources. The adjacent ocean is also a primary source of nutrients for autotrophs, photosynthesizing producers ranging in size from microscopic algae (e.g. benthic diatoms) to huge kelps and other seaweeds. These intertidal producers are eaten by herbivorous grazers, such as limpets that scrape rocks clean of their diatom layer and kelp crabs that creep along blades of the feather boa kelp Egregia eating the tiny leaf-shaped bladelets. Crabs are eaten by goliath grouper, which are then eaten by sharks. Higher up the food web, predatory consumers—especially voracious starfish—eat other grazers (e.g. snails) and filter feeders (e.g. mussels). Finally, scavengers, including crabs and sand fleas, eat dead organic material, including dead producers and consumers.

Species interactions

Tide pools with sea stars and sea anemone in Santa Cruz, California

In addition to being shaped by aspects of climate, intertidal habitats—especially intertidal zonation patterns—are strongly influenced by species interactions, such as predation, competition, facilitation, and indirect interactions. Ultimately, these interactions feed into the food web structure, described above. Intertidal habitats have been a model system for many classic ecological studies, including those introduced below, because the resident communities are particularly amenable to experimentation.

One dogma of intertidal ecology—supported by such classic studies—is that species' lower tide height limits are set by species interactions whereas their upper limits are set by climate variables. Classic studies by Robert Paine established that when sea star predators are removed, mussel beds extend to lower tide heights, smothering resident seaweeds. Thus, mussels' lower limits are set by sea star predation. Conversely, in the presence of sea stars, mussels' lower limits occur at a tide height at which sea stars are unable to tolerate climate conditions.

Competition, especially for space, is another dominant interaction structuring intertidal communities. Space competition is especially fierce in rocky intertidal habitats, where habitable space is limited compared to soft-sediment habitats in which three-dimensional space is available. As seen with the previous sea star example, mussels are competitively dominant when they are not kept in check by sea star predation. Joseph Connell's research on two types of high intertidal barnacles, Balanus balanoides, now Semibalanus balanoides, and a Chthamalus stellatus, showed that zonation patterns could also be set by competition between closely related organisms. In this example, Balanus outcompetes Chthamalus at lower tide heights but is unable to survive at higher tide heights. Thus, Balanus conforms to the intertidal ecology dogma introduced above: its lower tide height limit is set by a predatory snail and its higher tide height limit is set by climate. Similarly, Chthamalus, which occurs in a refuge from competition (similar to the temperature refuges discussed above), has a lower tide height limit set by competition with Balanus and a higher tide height limit is set by climate.

Hermit crabs and live Tegula snails on a dead gumboot chiton, Cryptochiton stelleri, in a tide pool at low tide in central California

Although intertidal ecology has traditionally focused on these negative interactions (predation and competition), there is emerging evidence that positive interactions are also important. Facilitation refers to one organism helping another without harming itself. For example, salt marsh plant species of Juncus and Iva are unable to tolerate the high soil salinities when evaporation rates are high, thus they depend on neighboring plants to shade the sediment, slow evaporation, and help maintain tolerable salinity levels. In similar examples, many intertidal organisms provide physical structures that are used as refuges by other organisms. Mussels, although they are tough competitors with certain species, are also good facilitators as mussel beds provide a three-dimensional habitat to species of snails, worms, and crustaceans.

All of the examples given so far are of direct interactions: Species A eat Species B or Species B eats Species C. Also important are indirect interactions where, using the previous example, Species A eats so much of Species B that predation on Species C decreases and Species C increases in number. Thus, Species A indirectly benefits Species C. Pathways of indirect interactions can include all other forms of species interactions. To follow the sea star-mussel relationship, sea stars have an indirect negative effect on the diverse community that lives in the mussel bed because, by preying on mussels and decreasing mussel bed structure, those species that are facilitated by mussels are left homeless. Additional important species interactions include mutualism, which is seen in symbioses between sea anemones and their internal symbiotic algae, and parasitism, which is prevalent but is only beginning to be appreciated for its effects on community structure.

Current topics

Humans are highly dependent on intertidal habitats for food and raw materials, and over 50% of humans live within 100 km of the coast. Therefore, intertidal habitats are greatly influenced by human impacts to both ocean and land habitats. Some of the conservation issues associated with intertidal habitats and at the head of the agendas of managers and intertidal ecologists are:

1. Climate change: Intertidal species are challenged by several of the effects of global climate change, including increased temperatures, sea level rise, and increased storminess. Ultimately, it has been predicted that the distributions and numbers of species will shift depending on their abilities to adapt (quickly!) to these new environmental conditions. Due to the global scale of this issue, scientists are mainly working to understand and predict possible changes to intertidal habitats.

2. Invasive species: Invasive species are especially prevalent in intertidal areas with high volumes of shipping traffic, such as large estuaries, because of the transport of non-native species in ballast water. San Francisco Bay, in which an invasive Spartina cordgrass from the east coast is currently transforming mudflat communities into Spartina meadows, is among the most invaded estuaries in the world. Conservation efforts are focused on trying to eradicate some species (like Spartina) in their non-native habitats as well as preventing further species introductions (e.g. by controlling methods of ballast water uptake and release).

3. Marine protected areas: Many intertidal areas are lightly to heavily exploited by humans for food gathering (e.g. clam digging in soft-sediment habitats and snail, mussel, and algal collecting in rocky intertidal habitats). In some locations, marine protected areas have been established where no collecting is permitted. The benefits of protected areas may spill over to positively impact adjacent unprotected areas. For example, a greater number of larger egg capsules of the edible snail Concholepus in protected vs. non-protected areas in Chile indicates that these protected areas may help replenish snail stocks in areas open to harvesting. The degree to which collecting is regulated by law differs with the species and habitat.

Introduction to entropy

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