Salmon ( /ˈsæmən/) is the common name for several species of fish in the family Salmonidae. Several other fish in the same family are called trout; the difference is often said to be that salmon migrate and trout are resident,[says who?] but this distinction does not strictly hold true. Salmon live along the coasts of both the North Atlantic (one migratory species Salmo salar) and Pacific Oceans (approximately a dozen species of the genus Oncorhynchus), and have also been introduced into the Great Lakes of North America. Salmon are intensively produced in aquaculture in many parts of the world.
Typically, salmon are anadromous: they are born in fresh water, migrate to the ocean, then return to fresh water to reproduce. However, there are populations of several species that are restricted to fresh water through their life. Folklore has it that the fish return to the exact spot where they were born to spawn; tracking studies have shown this to be true, and this homing behavior has been shown to depend on olfactory memory.[1][2]
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Salmon eggs are laid in freshwater streams typically at high latitudes. The eggs hatch into alevin or sac fry. The fry quickly develop into parr with camouflaging vertical stripes. The parr stay for six months to three years in their natal stream before becoming smolts, which are distinguished by their bright silvery colour with scales that are easily rubbed off. It is estimated that only 10% of all salmon eggs survive to this stage.[3] The smolt body chemistry changes, allowing them to live in salt water. Smolts spend a portion of their out-migration time in brackish water, where their body chemistry becomes accustomed to osmoregulation in the ocean.
The salmon spend about one to five years (depending on the species) in the open ocean where they gradually become sexually mature. The adult salmon then return primarily to their natal stream to spawn. In Alaska, the crossing-over to other streams allows salmon to populate new streams, such as those that emerge as a glacier retreats. The precise method salmon use to navigate has not been established, though their keen sense of smell is involved. Atlantic salmon spend between one and four years at sea. (When a fish returns after just one year's sea feeding it is called a grilse in Canada, Britain and Ireland.) Prior to spawning, depending on the species, salmon undergo changes. They may grow a hump, develop canine teeth, develop a kype (a pronounced curvature of the jaws in male salmon). All will change from the silvery blue of a fresh run fish from the sea to a darker colour. Salmon can make amazing journeys, sometimes moving hundreds of miles upstream against strong currents and rapids to reproduce. Chinook and sockeye salmon from central Idaho, for example, travel over 900 miles (1,400 km) and climb nearly 7,000 feet (2,100 m) from the Pacific ocean as they return to spawn. Condition tends to deteriorate the longer the fish remain in fresh water, and they then deteriorate further after they spawn, when they are known as kelts. In all species of Pacific salmon, the mature individuals die within a few days or weeks of spawning, a trait known as semelparity. Between 2% and 4% of Atlantic salmon kelts survive to spawn again, all females. However, even in those species of salmon that may survive to spawn more than once (iteroparity), post-spawning mortality is quite high (perhaps as high as 40 to 50%.)
To lay her roe, the female salmon uses her tail (caudal fin), to create a low-pressure zone, lifting gravel to be swept downstream, excavating a shallow depression, called a redd. The redd may sometimes contain 5,000 eggs covering 30 square feet (2.8 m2).[4] The eggs usually range from orange to red. One or more males will approach the female in her redd, depositing his sperm, or milt, over the roe.[5] The female then covers the eggs by disturbing the gravel at the upstream edge of the depression before moving on to make another redd. The female will make as many as seven redds before her supply of eggs is exhausted.[5]
Each year, the fish experiences a period of rapid growth, often in summer, and one of slower growth, normally in winter. This results in ring formation around an earbone called the otolith, (annuli) analogous to the growth rings visible in a tree trunk. Freshwater growth shows as densely crowded rings, sea growth as widely spaced rings; spawning is marked by significant erosion as body mass is converted into eggs and milt.
Freshwater streams and estuaries provide important habitat for many salmon species. They feed on terrestrial and aquatic insects, amphipods, and other crustaceans while young, and primarily on other fish when older. Eggs are laid in deeper water with larger gravel, and need cool water and good water flow (to supply oxygen) to the developing embryos. Mortality of salmon in the early life stages is usually high due to natural predation and human-induced changes in habitat, such as siltation, high water temperatures, low oxygen concentration, loss of stream cover, and reductions in river flow. Estuaries and their associated wetlands provide vital nursery areas for the salmon prior to their departure to the open ocean. Wetlands not only help buffer the estuary from silt and pollutants, but also provide important feeding and hiding areas.
Salmon that are not killed by other means show greatly accelerated deterioration (phenoptosis, or "programmed aging") at the end of their lives. Their bodies rapidly deteriorate right after they spawn as a result of the release of massive amounts of corticosteroids.
The various species of salmon have many names, and varying behaviors.
Eosalmo driftwoodensis, the oldest known salmon fossil in the fossil record, helps scientists figure how the different species of salmon diverged from a common ancestor. The British Columbia salmon fossil provides evidence that the divergence between Pacific and Atlantic salmon had not yet occurred 40 million years ago. Both the fossil record and analysis of mitochondrial DNA suggest that the divergence occurred by 10 to 20 million years ago. This independent evidence from DNA analysis and the fossil record reject the glacial theory of salmon divergence.[6]
The Atlantic Ocean has only one species of salmon, in the genus Salmo:
Pacific species belong to the genus Oncorhynchus, some examples include:
The salmon has long been at the heart of the culture and livelihood of coastal dwellers. Many people of the Northern Pacific shore had a ceremony to honor the first return of the year. For many centuries, people caught salmon as they swam upriver to spawn. A famous spearfishing site on the Columbia River at Celilo Falls was inundated after great dams were built on the river. The Ainu, of northern Japan, trained dogs to catch salmon as they returned to their breeding grounds en masse. Now, salmon are caught in bays and near shore.
Salmon population levels are of concern in the Atlantic and in some parts of the Pacific. Alaska fishery stocks are still abundant, and catches have been on the rise in recent decades, after the state initiated limitations in 1972.[13][14] Some of the most important Alaskan salmon sustainable wild fisheries are located near the Kenai River, Copper River, and in Bristol Bay. Fish farming of Pacific salmon is outlawed in the United States Exclusive Economic Zone, however, there is a substantial network of publicly funded hatcheries,[15] and the State of Alaska's fisheries management system is viewed as a leader in the management of wild fish stocks. In Canada, returning Skeena River wild salmon support commercial, subsistence and recreational fisheries, as well as the area's diverse wildlife on the coast and around communities hundreds of miles inland in the watershed. The status of wild salmon in Washington is mixed. Out of 435 wild stocks of salmon and steelhead, only 187 of them were classified as healthy; 113 had an unknown status, 1 was extinct, 12 were in critical condition and 122 were experiencing depressed populations.[16] The Columbia River salmon population is now less than 3% of what it was when Lewis and Clark arrived at the river.[17] The commercial salmon fisheries in California have been either severely curtailed or closed completely in recent years, due to critically low returns on the Klamath and or Sacramento Rivers, causing millions of dollars in losses to commercial fishermen.[18] Both Atlantic and Pacific salmon are popular sportfish.
Salmon populations now exist in all the Great Lakes. Coho stocks were planted in the late 1960s in response to the growing population of non-native alewife by the state of Michigan. Now Chinook (King), Atlantic, and Coho (silver) salmon are annually stocked in all Great Lakes by most bordering states and provinces. These populations are not self sustaining and do not provide much in the way of a commercial fishery, but have led to the development of a thriving sportfishery.
Salmon aquaculture is the major economic contributor to the world production of farmed fin-fish, representing over US$1 billion annually. Other commonly cultured fish species include: tilapia, catfish, sea bass, carp, bream, and trout. Salmon farming is significant in Chile, Norway, Scotland, Canada and the Faroe Islands, and is the source for most salmon consumed in America and Europe. Atlantic salmon are also, in very small volumes, farmed in Russia and the island of Tasmania, Australia.
Salmon are carnivorous and are currently fed a meal produced from catching other wild fish and other marine organisms. Salmon farming leads to a high demand for wild forage fish. Salmon require large nutritional intakes of protein, and consequently, farmed salmon consume more fish than they generate as a final product. To produce one pound of farmed salmon, products from several pounds of wild fish are fed to them. As the salmon farming industry expands, it requires more wild forage fish for feed, at a time when 75% of the world's monitored fisheries are already near to or have exceeded their maximum sustainable yield.[19] 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.
Work continues on substituting vegetable proteins for animal proteins in the salmon diet. Unfortunately, though, this substitution results in lower levels of the highly valued omega-3 fatty acid content in the farmed product.
Intensive salmon farming now uses open-net cages, which have low production costs, but have the drawback of allowing disease and sea lice to spread to local wild salmon stocks.[20]
On a dry weight basis, 2–4 kg of wild-caught fish are needed to produce one kg of salmon.[21]
Another form of salmon production, which is safer, but less controllable, is to raise salmon in hatcheries until they are old enough to become independent. They are then released into rivers, often in an attempt to increase the salmon population. This system is referred to as ranching, and was very common in countries such as Sweden before the Norwegians developed salmon farming, but is seldom done by private companies, as anyone may catch the salmon when they return to spawn, limiting a company's chances of benefiting financially from their investment. Because of this, the method has mainly been used by various public authorities and nonprofit groups such as the Cook Inlet Aquaculture Association as a way of artificially increasing salmon populations in situations where they have declined due to overharvesting, construction of dams, and habitat destruction or fragmentation. Unfortunately, there can be negative consequences to this sort of population manipulation, including genetic "dilution" of the wild stocks, and many jurisdictions are now beginning to discourage supplemental fish planting in favour of harvest controls and habitat improvement and protection. A variant method of fish stocking, called ocean ranching, is under development in Alaska. There, the young salmon are released into the ocean far from any wild salmon streams. When it is time for them to spawn, they return to where they were released where fishermen can then catch them.
An alternative method to hatcheries is to use spawning channels. These are artificial streams, usually parallel to an existing stream with concrete or rip-rap sides and gravel bottoms. Water from the adjacent stream is piped into the top of the channel, sometimes via a header pond, to settle out sediment. Spawning success is often much better in channels than in adjacent streams due to the control of floods, which in some years can wash out the natural redds. Because of the lack of floods, spawning channels must sometimes be cleaned out to remove accumulated sediment. The same floods which destroy natural redds also clean them out. Spawning channels preserve the natural selection of natural streams, as there is no benefit, as in hatcheries, to use prophylactic chemicals to control diseases.
Farm-raised salmon are fed the carotenoids astaxanthin and canthaxanthin to match their flesh color to wild salmon.[22]
According to Canadian biologist Dorothy Kieser, the myxozoan parasite Henneguya salminicola is commonly found in the flesh of salmonids. It has been recorded in the field samples of salmon returning to the Queen Charlotte Islands. The fish responds by walling off the parasitic infection into a number of cysts that contain milky fluid. This fluid is an accumulation of a large number of parasites.
Henneguya and other parasites in the myxosporean group have a complex life cycle, where the salmon is one of two hosts. The fish releases the spores after spawning. In the Henneguya case, the spores enter a second host, most likely an invertebrate, in the spawning stream. When juvenile salmon migrate to the Pacific Ocean, the second host releases a stage infective to salmon. The parasite is then carried in the salmon until the next spawning cycle. The myxosporean parasite that causes whirling disease in trout, has a similar life cycle.[23] However, as opposed to whirling disease, the Henneguya infestation does not appear to cause disease in the host salmon — even heavily infected fish tend to return to spawn successfully.
According to Dr. Kieser, a lot of work on Henneguya salminicola was done by scientists at the Pacific Biological Station in Nanaimo in the mid-1980s, in particular, an overview report[24] which states that "the fish that have the longest fresh water residence time as juveniles have the most noticeable infections. Hence in order of prevalence coho are most infected followed by sockeye, chinook, chum and pink." As well, the report says that, at the time the studies were conducted, stocks from the middle and upper reaches of large river systems in British Columbia such as Fraser, Skeena, Nass and from mainland coastal streams in the southern half of B.C. "are more likely to have a low prevalence of infection." The report also states "It should be stressed that Henneguya, economically deleterious though it is, is harmless from the view of public health. It is strictly a fish parasite that cannot live in or affect warm blooded animals, including man".
According to Klaus Schallie, Molluscan Shellfish Program Specialist with the Canadian Food Inspection Agency, "Henneguya salminicola is found in southern B.C. also and in all species of salmon. I have previously examined smoked chum salmon sides that were riddled with cysts and some sockeye runs in Barkley Sound (southern B.C., west coast of Vancouver Island) are noted for their high incidence of infestation."
Sea lice, particularly Lepeophtheirus salmonis and various Caligus species, including C. clemensi and C. rogercresseyi, can cause deadly infestations of both farm-grown and wild salmon.[25][26] Sea lice are ectoparasites which feed on mucus, blood, and skin, and migrate and latch onto the skin of wild salmon during free-swimming, planktonic nauplii and copepodid larval stages, which can persist for several days.[27][28][29] Large numbers of highly populated, open-net salmon farms can create exceptionally large concentrations of sea lice; when exposed in river estuaries containing large numbers of open-net farms, many young wild salmon are infected, and do not survive as a result.[30][31] Adult salmon may survive otherwise critical numbers of sea lice, but small, thin-skinned juvenile salmon migrating to sea are highly vulnerable. On the Pacific coast of Canada, the louse-induced mortality of pink salmon in some regions is commonly over 80%.[32]
The population of wild salmon declined markedly in recent decades, especially North Atlantic populations which spawn in the waters of western Europe and eastern Canada, and wild salmon in the Snake and Columbia River system in northwestern United States. The decline is attributed to the following factors:
Several governments and nongovernmental organizations (NGOs) are sharing in research and habitat restoration efforts to relieve this situation.
Results overall show estuary problems exist for some rivers, but issues involving feeding grounds at sea are impacting populations as well. In 2008, returns were markedly improved for Atlantic salmon on both sides of the Atlantic Ocean, but no one knows if this is a temporary improvement or sign of a trend.
One proposed alternative to the use of wild-caught fish as feed for the salmon, is the use of soy-based products. This should be better for the local environment of the fish farm, however, producing soy beans has a high environmental cost for the producing region.
Another promising alternative is a yeast-based, proteinaceous fermentation biomass (PFB). PFB is a co-product of bio-ethanol production. Studies have shown that the substitution of PFB products for engineered feed have resulted in equal (sometimes enhanced) growth in fish.[41] With its increasing availability, this would address the problems of rising costs for buying hatchery fish feed.
Yet another attractive alternative is the increased use of seaweed. Seaweed provides essential minerals and vitamins for growing organisms. It offers the advantage of providing natural amounts of dietary fiber and having a lower glycemic load than grain-based fish meal.[41] In the best-case scenario, widespread use of seaweed could yield a future in aquaculture that eliminates the need for land, freshwater, or fertilizer to raise fish.[42]
A great threat to Pacific salmon conservation is commercial fishing. There are many methods of harvest for the commercial salmon fishing industry, such as trolling, seining, and gillnetting. Gillnets are an extremely size-selective method of harvest, where a long net is placed in the path of the salmon’s migration to their natal stream in hopes of entangling the salmon for commercial harvest. Fish too small to be caught pass through the net, fish too large cannot be entangled, only catching fish that fall somewhere in between. By selectively harvesting certain sizes of fish, governed by the mesh-size of the gillnet, some age-class and length-class fish are selectively removed from the population, progressively leaving phenotypically smaller spawners. Fecundity generally decreases with length. So, smaller fish produce fewer eggs than larger fish.[43] There is also concern regarding the genetic information passed down from the fish. If the majority of spawning fish in a particular salmon run has gotten smaller due to the size-selective fishing methods, the run could eventually evolve to become smaller.
Gillnets are designed to harvest a specific sized fish. For example, Washington Department of Fish and Wildlife's 2010 Commercial Regulations had a 7-inch (180 mm) minimum mesh size restriction for Chinook, and a 5-inch (130 mm) minimum – 5.5-inch (140 mm) maximum for sockeye.[44] Possible problems arising from this selective harvest are smaller reproducing adult fish, as well as the unexpected mortality of the fish which sustain injuries from the gillnet but are not retained in the fishery. Most salmon populations include several age classes, allowing for fish of different ages, and sizes, to reproduce with each other. A recent 2009 study looked at 59 years of catch and escapement data of Bristol Bay sockeye to determine age and size at maturity trends attributable to the selectivity of commercial gillnet harvests. The study found that the larger females (>550 mm) of all age classes were most susceptible to harvest.[43] The study suggests that smaller, younger fish were more likely to successfully traverse the gillnet fishery and reproduce than the larger fish. The study also found that the average length of sockeye harvested from 1946–2005 was 8 mm larger than the sockeye who escaped the gillnet fishery to spawn, reducing the fecundity of the average female by 5%, or 104 eggs.[43]
If a salmon enters a gillnet, but manages to escape, it often sustains injuries. These injuries can lead to a lower degree of reproductive success. A study aimed at quantifying mortality of Bristol Bay sockeye salmon due to gillnet-related injuries found that 11 - 29% of sockeye sustained fishery-related injuries attributable to gillnets, 51% of those fish were expected to not reproduce.[45]
Beavers are archetypal ecosystem engineers; in the process of clear cutting and damming, beavers alter their ecosystem extensively. Beaver ponds can provide critical habitat for juvenile salmon. An example of this was seen in the years following 1818 in the Columbia River Basin. In 1818, the British government made an agreement with the U.S. government to allow U.S. citizens access to the Columbia catchment (see Treaty of 1818). At the time, the Hudson's Bay Company sent word to trappers to extirpate all furbearers from the area in an effort to make the area less attractive to U.S. fur traders. In response to the elimination of beavers from large parts of the river system, salmon runs plummeted, even in the absence of many of the factors usually associated with the demise of salmon runs. Salmon recruitment can be affected by beavers' dams because dams can:[46][47][48]
Beavers' dams are able to nurture salmon juveniles in Estuarine tidal marshes where the salinity is less than 10 ppm. Beavers build small dams of generally less than 2 feet (60 cm) high in channels in the Myrtle zone. These dams can be overtopped at high tide and hold water at low tide. This provides refuges for juvenile salmon so they do not have to swim into large channels where they are subject to predation.[49]
Salmon is a popular food. Classified as an "oily fish",[50] salmon is considered to be healthy due to the fish's high protein, high omega-3 fatty acids, and high vitamin D[51] content. Salmon is also a source of cholesterol, with a range of 23–214 mg/100 g depending on the species.[52] According to reports in the journal Science, however, farmed salmon may contain high levels of dioxins. PCB (polychlorinated biphenyl) levels may be up to eight times higher in farmed salmon than in wild salmon.[53] Omega-3 content may also be lower than in wild-caught specimens, and in a different proportion to what is found naturally. Omega-3 comes in three types, alpha-linolenic acid (ALA), docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA); wild salmon has traditionally been an important source of DHA and EPA, which are important for brain function and structure, among other things. The body can itself convert ALA omega-3 into DHA and EPA, but at a very inefficient rate (2–15%). Nonetheless, according to a 2006 study published in the Journal of the American Medical Association, the benefits of eating even farmed salmon still outweigh any risks imposed by contaminants.[54] The type of omega-3 present may not be a factor for other important health functions.
A simple rule of thumb is that the vast majority of Atlantic salmon available on the world market are farmed (almost 99%[55]), whereas the majority of Pacific salmon are wild caught (greater than 80%).
Salmon flesh is generally orange to red, although there are some examples of white-fleshed wild salmon. The natural colour of salmon results from carotenoid pigments, largely astaxanthin but also canthaxanthin, in the flesh.[56] Wild salmon get these carotenoids from eating krill and other tiny shellfish. Because consumers have shown a reluctance to purchase white-fleshed salmon, astaxanthin (E161j), and very minutely canthaxanthin (E161g), are added as artificial colorants to the feed of farmed salmon, because prepared diets do not naturally contain these pigments. In most cases, the astaxanthin is made chemically; alternatively it is extracted from shrimp flour. Another possibility is the use of dried red yeast, which provides the same pigment. However, synthetic mixtures are the least expensive option. Astaxanthin is a potent antioxidant that stimulates the development of healthy fish nervous systems and enhances the fish's fertility and growth rate. Research has revealed canthaxanthin may have negative effects on the human eye, accumulating in the retina at high levels of consumption.[56] Today, the concentration of carotenoids (mainly canthaxanthin and astaxanthin) exceeds 8 mg/kg of flesh, and all fish producers try to reach a level that represents a value of 16 on the "Roche Color Card", a colour card used to show how pink the fish will appear at specific doses. This scale is specific for measuring the pink colour due to astaxanthin and is not for the orange hue obtained with canthaxanthin. The development of processing and storage operations, which can be detrimental on canthaxanthin flesh concentration, has led to an increased quantity of pigments added to the diet to compensate for the degrading effects of the processing. In wild fish, carotenoid levels of up to 25 mg are present, but levels of canthaxanthin are, in contrast, minor.[56]
Canned salmon in the U.S. is usually wild Pacific catch, though some farmed salmon is available in canned form. Smoked salmon is another popular preparation method, and can either be hot or cold smoked. Lox can refer either to cold smoked salmon or to salmon cured in a brine solution (also called gravlax). Traditional canned salmon includes some skin (which is harmless) and bone (which adds calcium). Skinless and boneless canned salmon is also available.
Raw salmon flesh may contain Anisakis nematodes, marine parasites that cause Anisakiasis. Before the availability of refrigeration, Japanese did not consume raw salmon. Salmon and salmon roe have only recently come into use in making sashimi (raw fish) and sushi.
159g (0.5 fillet) of raw salmon contains:
Ordinary types of cooked salmon contain 500–1500 mg DHA and 300–1000 mg EPA per 100 grams.[57]
The salmon is an important creature in several strands of Celtic mythology and poetry, which often associated them with wisdom and venerability. In Irish mythology, a creature called the Salmon of Wisdom (or the Salmon of Knowledge)[58] plays key role in the tale known as The Boyhood Deeds of Fionn. The Salmon will grant powers of knowledge to whoever eats it, and has been sought by the poet Finn Eces for seven years. Finally Finn Eces catches the fish and gives it to his young pupil, Fionn mac Cumhaill, to prepare it for him. However, Fionn burns his thumb on the salmon's juices, and he instinctively puts it in his mouth. As such, he inadvertently gains the Salmon's wisdom. Elsewhere in Irish mythology, the salmon is also one of the incarnations of both Tuan mac Cairill[59] and Fintan mac Bóchra.[60]
Salmon also feature in Welsh mythology. In the prose tale Culhwch and Olwen, the Salmon of Llyn Llyw is the oldest animal in Britain, and the only creature who knows the location of Mabon ap Modron. After speaking to a string of other ancient animals who do not know his whereabouts, King Arthur's men Cai and Bedwyr are led to the Salmon of Llyn Llyw, who lets them ride its back to the walls of Mabon's prison in Gloucester.
In Norse mythology, after Loki tricked the blind god Höðr into killing his brother Baldr, Loki jumped into a river and transformed himself into a salmon in order to escape punishment from the other gods. When they held out a net to trap him he attempted to leap over it but was caught by Thor who grabbed him by the tail with his hand, and this is why the salmon's tail is tapered.[61]
Salmon are central to Native American mythology on the Pacific coast, from the Haida to the Nootka.
In the Pacific Northwest, one of the most notable relief efforts is the Puget Sound Partnership.[62] The Puget Sound Partnership is currently working to implement policy change at the local level to alter the fate of salmon. Salmon recovery is guided by implementation of the Puget Sound Salmon Recovery Plan, adopted by the National Oceanic and Atmospheric Administration (NOAA) in January 2007. This recovery plan was developed by Shared Strategy, a grassroots collaborative effort to protect and restore salmon runs across Puget Sound. The Puget Sound Partnership has now rehabilitated over 800 acres of salmon habitat and it plans to continue to fight nearshore development and human impact affecting important salmon ecosystems. The Partnership's Action Agenda plans to instigate the Elwha Dam removal, begin restoration of the Nisqually Estuary and removal of derelict Fishing gear, and continue with the current salmon Recovery Plan. In order to ensure the future of Pacific Northwest salmon, the Partnership continues to encourage Stormwater & Low Impact Development, and advocates the "Puget Sound Starts Here" public education program.
Another notable local relief effort is the People for Puget Sound. People for Puget Sound is a citizen group founded by Kathy Fletcher in 1991 working to restore the health of our local land and waters with help from volunteers in the Puget Sound basin.[63]
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