Salmon run
A salmon run is an annual fish migration event where many salmonid species, which are typically hatched in fresh water and live most of their adult life downstream in the ocean, swim back against the stream to the upper reaches of rivers to spawn on the gravel beds of small creeks. After spawning, most Atlantic salmon and all species of Pacific salmon die, and the salmon life cycle starts over again with the new generation of hatchlings.
Salmon are anadromous, spending their juvenile life in rivers or lakes, and then migrating out to sea where they spend adult lives and gain most of their body mass. When they reach sexual maturity, the adults return to the upstream rivers to reproduce. Usually they return with uncanny precision to the natal river where they were born, and even to the very spawning ground of their birth. It is thought that, when they are in the ocean, they use magnetoreception to locate the general position of their natal river, and once close to the river, that they use their sense of smell to home in on the river entrance and even their natal spawning ground.
Trout, which are sister species of salmon, also perform similar migrations, although they mostly move potamodromously between creeks and large freshwater lakes, except for some coastal/estuary subspecies such as steelhead and sea trout that migrate seasonally between salty/brackish and fresh water just like salmon do. There are also landlocked populations of some salmon species that have adapted to spend their entire life in freshwater like trout.
In Northwest America, salmons are keystone species, which means the ecological impact they have on other wildlife is greater than would be expected in relation to their biomass. Most salmon species migrate during the autumn, which coincides with the pre-winter activities of many hibernating animals. The annual salmon run can be a major feeding event for predators such as grizzly bears and bald eagles, as well as an important window period for sport fishermen.
The post-spawning death of salmon also has important ecological consequences, because the significant nutrients in their carcasses, rich in nitrogen, sulfur, carbon and phosphorus, are transferred from the ocean and released to inland aquatic ecosystems, terrestrial animals and the wetlands and riparian woodlands adjacent to the rivers. This has knock-on effects not only for the next generation of salmon, but to every wildlife species living in the riparian zones the salmon reach. The nutrients can also be washed downstream into estuaries where they accumulate and provide significant support for invertebrates and estuarine-breeding waterbirds.
Background
Most salmon are anadromous, a term which comes from the Greek anadromos, meaning "running upward". Anadromous fish grow up mostly in the saltwater in oceans. When they have matured they migrate or "run up" freshwater rivers to spawn in what is called the salmon run.Anadromous salmon are Northern Hemisphere fish that spend their ocean phase in either the Atlantic Ocean or the Pacific Ocean. They do not thrive in warm water. There is only one species of salmon found in the Atlantic, commonly called the Atlantic salmon. These salmon run up rivers on both sides of the ocean. Seven different species of salmon inhabit the Pacific, and these are collectively referred to as Pacific salmon. Five of these species run up rivers on both sides of the Pacific, but two species are found only on the Asian side. In the early 19th century, Chinook salmon were successfully established in the Southern Hemisphere, far from their native range, in New Zealand rivers. Attempts to establish anadromous salmon elsewhere have not succeeded.
The life cycle of an anadromous salmon begins and, if it survives the full course of its natural life, usually ends in a gravel bed in the upper reaches of a stream or river. These are the salmon spawning grounds where salmon eggs are deposited, for safety, in the gravel. The salmon spawning grounds are also the salmon nurseries, providing a more protected environment than the ocean usually offers. After 2 to 6 months the eggs hatch into tiny larvae called sac fry or alevin. The alevin have a sac containing the remainder of the yolk, and they stay hidden in the gravel while they feed on the yolk. When the yolk has gone they must find food for themselves, so they leave the protection of the gravel and start feeding on plankton. At this point the baby salmon are called fry. At the end of the summer the fry develop into juvenile fish called parr. Parr feed on small invertebrates and are camouflaged with a pattern of spots and vertical bars. They remain in this stage for up to three years.
As they approach the time when they are ready to migrate out to the sea the parr lose their camouflage bars and undergo a process of physiological changes which allows them to survive the shift from freshwater to saltwater. At this point salmon are called smolt. Smolt spend time in the brackish waters of the river estuary while their body chemistry adjusts their osmoregulation to cope with the higher salt levels they will encounter in the ocean. Smolt also grow the silvery scales which visually confuse ocean predators. When they have matured sufficiently in late spring, and are about 15 to 20 centimetres long, the smolt swim out of the rivers and into the sea. There they spend their first year as a post-smolt. Post-smolt form schools with other post-smolt, and set off to find deep-sea feeding grounds. They then spend up to four more years as adult ocean salmon while their full swimming ability and reproductive capacity develop.
Then, in one of the animal kingdom's more extreme migrations, the salmon return from the saltwater ocean back to a freshwater river to spawn afresh.
Return from the ocean
After several years wandering huge distances in the ocean, most salmon return to the natal rivers where they were spawned. Then most of them swim up the rivers until they reach the very spawning ground that was their original birthplace.There are various theories about how this happens. One theory is that there are geomagnetic and chemical cues which the salmon use to guide them back to their birthplace. The fish may be sensitive to the Earth's magnetic field, which could allow the fish to orient itself in the ocean, so it can navigate back to the estuary of its natal stream.
Salmon have a strong sense of smell. Speculation about whether odours provide homing cues go back to the 19th century. In 1951, Hasler hypothesised that, once in vicinity of the estuary or entrance to its birth river, salmon may use chemical cues which they can smell, and which are unique to their natal stream, as a mechanism to home onto the entrance of the stream. In 1978, Hasler and his students found that the way salmon locate their home rivers with such precision was because they could recognise its characteristic smell. They further demonstrated that the smell of their river becomes imprinted in salmon when they transform into smolts, just before they migrate out to sea. Homecoming salmon can also recognise characteristic smells in tributary streams as they move up the main river. They may also be sensitive to characteristic pheromones given off by juvenile conspecifics. There is evidence that they can "discriminate between two populations of their own species".
The recognition that each river and tributary has its own characteristic smell, and the role this plays as a navigation aid, led to a widespread search for a mechanism or mechanisms that might allow salmon to navigate over long distances in the open ocean. In 1977, Leggett identified, as mechanisms worth investigating, the use of the sun for navigation, and orientation to various possible gradients, such as temperature, salinity or chemicals gradients, or geomagnetic or geoelectric fields.
There is little evidence salmon use clues from the sun for navigation. Migrating salmon have been observed maintaining direction at nighttime and when it is cloudy. Likewise, electronically tagged salmon were observed to maintain direction even when swimming in water much too deep for sunlight to be of use.
In 1973, it was shown that Atlantic salmon have conditioned cardiac responses to electric fields with strengths similar to those found in oceans. "This sensitivity might allow a migrating fish to align itself upstream or downstream in an ocean current in the absence of fixed references." In 1988, researchers found iron, in the form of single domain magnetite, resides in the skulls of sockeye salmon. The quantities present are sufficient for magnetoception.
Tagging studies have shown a small number of fish do not find their natal rivers, but travel instead up other, usually nearby streams or rivers. It is important some salmon stray from their home areas; otherwise new habitats could not be colonized. In 1984, Quinn hypothesized there is a dynamic equilibrium, controlled by genes, between homing and straying. If the spawning grounds have a uniform high quality, then natural selection should favour the descendants that home accurately. If the spawning grounds have a variable quality, then natural selection should favour a mixture of the descendants that stray and the descendants that home accurately.
Prior to the run up the river, the salmon undergo profound physiological changes. Fish swim by contracting longitudinal red muscle and obliquely oriented white muscles. Red muscles are used for sustained activity, such as ocean migrations. White muscles are used for bursts of activity, such as bursts of speed or jumping. As the salmon comes to end of its ocean migration and enters the estuary of its natal river, its energy metabolism is faced with two major challenges: it must supply energy suitable for swimming the river rapids, and it must supply the sperm and eggs required for the reproductive events ahead. The water in the estuary receives the freshwater discharge from the natal river. Relative to ocean water, this has a high chemical load from surface runoff. Researchers in 2009 found evidence that, as the salmon encounter the resulting drop in salinity and increase in olfactory stimulation, two key metabolic changes are triggered: there is a switch from using red muscles for swimming to using white muscles, and there is an increase in the sperm and egg load. "Pheromones at the spawning grounds a second shift to further enhance reproductive loading."
The salmon also undergo radical morphological changes as they prepare for the spawning event ahead. All salmon lose the silvery blue they had as ocean fish, and their colour darkens, sometimes with a radical change in hue. Salmon are sexually dimorphic, and the male salmon develop canine-like teeth and their jaws develop a pronounced curve or hook. Some species of male salmon grow large humps.