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Migration is a deliberate movement taken by animals for survival. It is commonly categorized as spawning, feeding, and refuge seeking migrations. Migration is governed by costs and benefits. Energy production and utilization is one of the greatest challenges of freshwater fish migration. The upstream and long-distance migrants demand more energy. Orientation and navigation mechanisms in fishes have a long history of interest. Different sensory mechanisms for accurate orientation have been suggested, including orientation using sun position, polarized light patterns, and the Earth’s geomagnetic field. Fish morphology plays a significant role in assisting freshwater fish’s migration. Long-distant migrants have streamlined body structure and longer caudal regions, while short-distance migrants are fusiform making them hard to move long distance against water current. Since fish migration may involve two different aquatic environments, all migrant fishes that cross the interface between freshwater and saline water habitats must therefore undergo physiological changes. Fish migration activities are influenced by abiotic factors including variations in water temperature, water level, and light availability. Human activities significantly affect fish migration. A good understanding of the migratory behavior of fishes is important for effective fisheries management. Fishermen and near-shore communities need to become aware about the nature of fish migration.
*Address all correspondence to: gatriaytut@gmail.com
1. Introduction
Migration is the heroic movement of the animal population from one place to another and it may cover some distances [1]. It is not a simple movement, rather it is a special and deliberate journey taken to achieve some survival goals. Animals migrate for many reasons, over long and short distances and in many different ways such as flying, swimming, walking, or drifting [1]. Migration of fishes is observed in migratory species that carryout this life journey for survival and/or to complete their lifecycle. The long distance migration is demonstrated by the movements of salmonid species between different aquatic ecosystems [2]. Whereas juvenile of many fish species experiences a short daily movements within the same water body between the pelagic and littoral zones, for feeding and avoiding predators [3, 4]. Migration occurs principally between feeding and breeding ground and it may have a strong influence on the population dynamics and affect species composition of the area [5, 6].
Migration usually takes various forms depending on its purpose; namely spawning, feeding, and refuge seeking [7]. Fish migration is fundamentally influenced by spawning migration, which is frequently correlated with temperature, a key regulating element, particularly for species that spawn in the spring. When other mature fish are available, adult fish exhibit what is known as homing behavior, returning to their natal streams to reproduce simultaneously. For migratory fish species to have the best chance of reproducing, synchronized migrations are essential [8]. Spring spawning migration is known in some of the common freshwater fishes like Northern Pike (Esox lucius Linnaeus, 1758) [9], Roach (Rutilus rutilus Linnaeus, 1758) [10], Perch (Perca fluviatilis Linnaeus, 1758) [11], Ruffe (Gymnocephalus cernua Linnaeus, 1758), Common bream (Abramis brama Linnaeus, 1758) and Bleak (Alburnus alburnus Linnaeus, 1758) [12]. Temperature is observed to influence the duration and intensity of spawning migration [10, 12]. In the environment with fluctuating temperatures, light plays an important role to induce spawning migration.
Predation is a strong mortality factor for fish [13] and the major driving force of migration especially for smaller species and juveniles. To reduce predation mortality and maximize growth rate, fish migrate to rivers to avoid predators and seek refuges [5, 14]. Such habitats’ change may involve a trade-off between predator avoidance and foraging. If the food accessibility is high, then the fish prey may accept the risk of predation or migrate to an area that gives better protection from predators but low access to food [5, 15].
For migrating fish species, it is important for their survival that they should move freely between feeding and spawning areas. Many fish species cannot complete their life cycles in the lake environment and need access to river habitats [12]. Various management gaps such as natural and artificial barriers can hinder fish to migrate between habitats leading to injuries, decline, or even extinction of fish populations [7]. Manmade barriers particularly have led to fragmentation especially of running water systems. In various freshwater ecosystems, different methods were utilized to overcome these problems. Obstructions like fish ladders, fish elevators, and channel bypasses have been built to facilitate the migration of fish such as cyprinids, esocids, and percids [16]. Misunderstanding of fish migration put migratory fish into a threat for their survival. These fish’s species faced great pressures from overexploitation at their breeding habitat and barriers construction that restrict them from continuing their lifecycle. Therefore, this paper is aimed to present the nature of migration in fish survival for effective fisheries management.
Some of the greatest challenges of freshwater fish migration are energy production and utilization. These physiological challenges are magnified during upstream migration where some fish’s species spend 75 to 82% of their total energy [17]. Moreover, the energy expenditure during fish’s migration increases when the amount of water and its velocity increase at the same time [18]. Binder also added that American shad (Alosa sapidissima Wilson, 1811) does not repeat spawning due to high energy depletion. The reason maybe because the forms in which the energy is stored in a fish’s body lost their physiological capability to spawn for the second time [19].
Based on the behavior of the migrating fish species, the amount of energy to be spent during migration could be inspected. Some species feed during migration to replace the energy spent while others do not do so. The chance for reproduction rate is very rare for those species that do not feed at the time of migration, sometimes even impossible. Both the success of the migrant species and any of their activities extremely rely solely on the amount of energy they stored before migration. Most of the stored energy is used for muscle movement. During fast swimming and long migration, fish’s muscles accumulated lactic acid due to oxygen depletion. The conversion of lactate into glycogen does not last and takes at least 24 hours [17].
The mechanisms of orientation and navigation in fishes have a long history that date back from its discovery in the 1930s. This is exemplified by salmon that return from the ocean to their natal streams to spawn, the phenomenon is termed ‘homing’ [17].
Several orientation and navigation mechanisms have been proposed, and several different cues might likely be involved in orientation during migration. Orientation and navigation activities differ greatly in different water ecosystems. In an open water migration, fishes may probably move in any one of 360° directions. Whereas riverine migrants are located within the channel of the river where they use their added cue of moving either with or against the current, a phenomenon termed positive or negative rheotaxis, respectively.
Quinn [20] suggested various sensory mechanisms for accurate orientation, including orientation using sun’s location, polarized light pattern, and the Earth’s geomagnetic field. Polarized light may provide a good orientation cue for fishes that migrate at sunrise–sunset when the polarized light patterns are strongest [21]. A response to geomagnetic cues has been shown in some species [22] and biogenic magnetite crystals have been found in the olfactory lamellae of rainbow trout (Oncorhynchus mykiss Walbaum, 1792) [22] and in the lateral line of Atlantic salmon [23].
Sockeye salmon (Oncorhynchus nerka, Walbaum, 1792) from the Fraser River is magnetically imprinted when they migrate to sea and use variations in magnetic field intensity to successfully locate the coastal imprinting site during their return migration, according to a recent analysis of a long-term fisheries data set [24]. The map and compass hypothesis is not without controversy, though. Fish cannot utilize a compass since they do not have an accurate biological clock [25]. Instead, they use their highly evolved sense of smell to orient to their natal stream.
It has long been known that salmonids utilize smell to discriminate between different streams, even though there is still significant controversy about the sensory cues fish employ to migrate in open water bodies [26]. The pheromone theory [27] and the olfactory imprinting hypothesis [26] are the two main hypotheses that have been put forth to explain olfactory cues. According to the imprinting hypothesis, each stream has a distinctive composition of chemical elements that come from the soil and flora that surround and form the stream, and even a site-specific odor that is imprinted on the youngster. The salmonids may employ this “stream bouquet” as olfactory cues during their return migration from the sea. According to the pheromone hypothesis, migrating adults use population-specific pheromones that are given off by young relatives in their natal stream. According to research on petromyzontid lampreys, adults seek spawning streams by following the bile acid pheromones of young lampreys [28]. Adult lampreys, however, do not return to their natal stream.
The shape of a fish is a fundamental factor associated with hydrodynamic performance and morphological traits such as a streamlined body can greatly reduce drag thereby assisting swimming during migration [29]. Morphological adaptations for migration in fish have been extensively studied in salmonid migrants. Two key characteristics in the smoltification process are a marked increase in body length relative to mass resulting in streamlining of the body and a decrease in the relative size of pectoral fins [30]. These changes in body form are presumably adaptive for migratory performance. Moreover, variations in body morphology may also occur as a result of adaptation to local conditions and several studies have shown the correlation of body morphology with migration distance and hydrological conditions in the natal habitat [31].
Crossin et al. [32] compared the morphology of short- and long-distance migratory populations of Fraser River sockeye salmon. The study result revealed that populations that undertake difficult migrations had short, fusiform bodies that are favorable for reducing transportation costs. Similarly, Brook trout (Salvelinus fontinalis Mitchill, 1814) populations that undertook longer migrations were more adapted for energy efficient migration with more streamlined bodies and longer caudal regions [33]. Most of the characters and physiology observed in migratory fish species may be an adaptation to increase survival in the alternative habitats and performance during migration.
4.2 Crossing the interface: Osmoregulation
As migration is the movement requiring migrating species to change its original location, fish move between different water ecosystems. Diadromous migrations between hypo-osmotic freshwater and hyper-osmotic seawater place huge demands on osmoregulatory ability of migrating species. To withstand this osmotic difference, all migrants’ fish that cross the interface between freshwater and saline water habitats must therefore undergo physiological changes [34]. The ability to adapt and survive in habitats that differ in salinity is common and has been documented in a wide range of migrating fish species, ranging from Atlantic salmon to pike [35]. However, preparatory physiological adaptations that occur prior to transition to marine habitats differentiate salmonids from other species with euryhaline capability [36]. Though these physiological changes involves the integration of multiple regulations, the increase in euryhalinity is a key characteristic and perhaps the most critical element given the modest osmoregulatory ability of the stream-dwelling parr [36]. Major functional changes in important osmoregulatory organs such as the gill, kidney, gut, urinary bladder, and skin took place prior to the transition between the marine and freshwater ecosystems [37]. Lucas and Baras [38, 39] compiled a list of common long-distance migratory freshwater fish species of Africa, Asia, and South America [Table 1]. Moreover the pictorial representation of migratory freshwater fish species is provided in Figure 1.
The movement and distribution of fishes in freshwater habitats can be influenced by living and nonliving factors, such as hydrological regime, temperature, food resources, predation, and competition. The migration pattern of fish is not likely the same since the conditions in the habitat are not static. Fish migration patterns in large floodplain rivers are highly intricate. Fish move not just laterally (in and out of river tributaries, floodplain streams, and various flooded areas), but also longitudinally (up and down the main channel). Among many possible causal factors, flooding is known to significantly affect patterns of fish migration [40].
5.1 Environmental factors that influence migration
Environmental factors greatly influence the pattern of migration in fish. The relative importance of each environmental factor may be dependent on the local characteristic of the habitat in which migration is occurring [18]. For instance, temperature plays significant role in the correlation between the length of the day (photoperiod) and water temperature. Stephen et al. [41] investigated other three factors that have a direct effect on the migratory activities of fishes.
5.1.1 Light
Photoperiod is a seasonal factor that triggers a complex series of physiological events that prepare migrating species for migration. Binder et al. [17] reported that any change in light intensity is an essential environmental trigger for starting migration. A population’s migratory movement is started and synchronized by the seasonal changes in photoperiod, which give programmed information. Lengthy migrations, as those of Pacific salmon and lampreys for spawning, seem to be principally dependent on photoperiod. Although photoperiod influences the migratory behavior of Pacific salmon smolts, there has been no direct investigation of the role of photoperiod in regulating behavioral changes in Atlantic salmon smolts. In comparison to other vertebrate migrations, photoperiod is observed to increase activity and make fish more responsive to other factors that initiate migration. Hence, photoperiod may function to determine the range of dates when the migration may take place.
5.1.2 Water flow
Changes in water levels to higher volume may function as an initiating factor for migration. When the snow is melting in the spring, water levels become higher and water flow increase as well. Increase in water flow may help the migrating fish species to find the entrance to the stream. However, if flows are very high they can instead be a hinder for fish migration [42]. Very high water flows can also lead to displacements of fish eggs and larvae. Roger and Katherine [43] reported that in spawning migration, the greatest egg deposition occurred when the flood tide is slow. High water flow in rivers may stimulate downstream movement in a large number of fish species and the downstream migration of smolts has been linked to increased water flow. This may be because they migrated at a faster speed and were closer to the water surface, using high turbidity in fast flowing water to make them less visible to predators.
5.1.3 Water temperature
Temperature is another environmental factor that can trigger and synchronize migratory activity in fishes. When temperature function as a stimulus, migration can be viewed as a form of behavioral thermoregulation [17]. This occur under two conditions. Firstly, in thermally heterogeneous environments, the temperature may deviate from the range of thermal tolerance for the migrating species. Thus, fish are forced to migrate for search of new thermally optimal habitat. Secondly, the thermal requirements of migrating fish species may change in demand to undertake various survival activities. For instance, the ideal temperature for growth differs from that of reproduction. In this case, the migration to spawning sites is a result of the temperature. But during spawning migrations, temperature is also a well-known synchronization trigger. Most species with relatively short migrations can attest to this. The springtime upstream migration of sea lampreys (Petromyzon marinus Linnaeus, 1758) in the Laurentian Great Lakes is a good illustration of this; the migration does not start until stream temperatures are above −10°C. As a result, the migration starts later in springs that are cooler than usual and sooner in springs that are warmer than average. Thermal thresholds for this kind of spawning migrating fishes is common and believed to have evolved in response to the strict thermal requirements of the developing embryos.
Marine water temperature may have a significant effect on the spawning activities of migratory fishes [43]. Piecuch et al. [44] reported that the trout migrating from the reservoir to the main tributary started when the water temperature in the reservoir decreased below 8°C and the water temperature in the tributary is in the range between 5 and 7°C.
5.2 Anthropogenic impact on fish’s migration
For their sustenance, migratory species depend on an interconnected chain of intact habitats, a requirement which exposed them to human disturbances in both their habitats and also their migratory routes [45]. Migratory fish species need movement between different habitats, clear migratory pathways between these habitats and appropriate migratory cues (which initiate migration or guide direction) to complete their life cycle or locating valuable resources [46]. Anthropogenic activities have a long history of interfering with fish migration. The complex nature of their life history makes migratory species extremely susceptible to human related threats and climate change, with different impacts at different life stages. Generally, anthropogenic threats to freshwater ecosystems include developmental structures (e.g. dams, weirs, road intersections, hydropower plants), habitat degradation (e.g., land use changes, pollution, channelization), and flow modification (e.g., water extraction, flow regulation, flood control), overfishing, and invasive species (predation, competition, disease, gene transfer, etc.) [39].
The most obvious way in which human activities disrupt migration is through the construction of developmental structures such as dams and weirs that acts as barriers by blocking access to the desired habitats leading to increased migration mortality [38]. Dam construction leaves migratory fishes without any choice since they never pass over it [18]. Therefore, the power of nature to select migratory fishes for their survival may be altered [29]. Human activities in upstream areas blocked with barriers can change migratory cues and downstream habitat through flow control [47]. For instance, tidal regulation can limit sea water incursion into estuaries, capable of reducing suitable breeding grounds for Australian bass (Percalates novemaculeata Steindachner, 1866) [48]. Upstream fish migration in watersheds containing such dams has been facilitated by the introduction of fish ladders. Fish ladders, however, only permit migration of fish species with particular phenotypic traits that can overcome the very strong currents [49]. Barriers disrupt stream continuity thereby reducing the abundance and quality of suitable stream habitat. Species hindered by barriers from migration may also be exposed to additional predators when they are enforced to assemble together with other predatory organisms within the same habitat. Another problem caused by the dam is genetic isolation.
Fishing activities and habitat degradation also threaten migratory species particularly diadromous species in estuarine, coastal and marine ecosystems [46]. Large-scale commercial fishing activities commonly occur in these habitats causing overfishing that led to a remarkable decline in many migratory fish populations [45]. Although several studies have reported the importance of estuary and coastal habitats to fishery productions [50, 51], they are vulnerable to anthropogenic activities that may affect water and sediment quality [52, 53].
Land-use changes such as urbanization or agricultural activities can also modify or pollute important habitats along streams [54]. Some pollutants can cause the sensory cells that regulate orientation to sustain physical harm. For instance, heavy metals are extremely toxic to the lateral line and olfactory organs. The metabolism can also be affected by contaminants. Numerous toxicants have been demonstrated to impair fishes’ ability to swim by reducing oxygen intake and diverting energy from the swimming muscles. Additionally, this elevated metabolic burden may result in early death [17]. Toxicants and other chemical contaminants [17] may mask the odors that some fishes use to identify the home stream. If a disease or any poison occurs in the water bodies that are dammed, the population of many fishes and other organisms will certainly decline [18].
Climate change as a result of human activities is thought to affect the distribution and productivity of migratory fishes species by interrupting growth, survivorship, habitat availability, migration and [47, 55]. For example, reducing freshwater flow due to climate change might decrease the recruitment and growth of barramundi (Lates calcarifer Bloch, 1790) [56] and disrupt the link between sea water and freshwater ecosystems for diadromous fish [47, 57]. Moreover, climate change can reduce survival of diadromous fish in the ocean stage [58] thereby influencing recruitment [59] and decreasing population density. In addition, sea level rise cuased by climatic change may alter the nature of estuarine and reduce freshwater habitats suitability for salmonids [60].
The consequence of climate change on migratory fish particularly diadromous fish species is thought to be complicated for the reason that it influence populations in various ways in different life stages. For instance, Piou and Prevost [61] reported that increasing river discharge and decreasing oceanic growth stage of Atlantic salmon (Salmo salar Linneaus, 1758) due to climate change surge their risk of local extinction. Moreover, Hilborn [58] revealed that both the change in ocean climate and anthropogenic activities (dams, fisheries and habitat degradation) contributed to the population decline of Chinook salmon (Oncorhynchus tshawytscha Walbaum, 1792) and European eel (Anguilla Anguilla Linneaus, 1758).
Fishes are widely distributed in both freshwater and marine environment. Fishes like any other animals move from one habitat to another to sustain their life. They undertake this journey for various reasons including spawning, feeding, avoiding adverse environmental condition and predators avoidance. Like other living organisms, fish migrate to cope with their survival needs and requirements. As resources and habitats are not always stable, migration in fishes is one mechanism to compromise with unstable situation in particular habitat. Different age groups in fish have different habitat requirements suitable for their survival. As a problem experienced by many migratory fishes, migratory routes are being blocked by various construction facility such as dams and weirs. To this end, provision of fish’s ladder and migratory ways becomes mandatory and should be considered when constructing these developmental facilities. Fisheries experts and managers have focused their attention on these migratory route obstructions to gain good results in fisheries management. A good understanding of the migratory behavior of fishes is an important means of controlling overexploitation of fish stock in particular and enhancing effective fisheries management in general. Catches in the feeding area may increase following the reduction of harvest at the spawning area. Moreover, human activities such as dams and weirs construction obstructing migratory behavior in fishes should be compromised by providing ladders and fish ways, which allow migratory fish to change habitat thereby completing their life cycle.
Awareness regarding the nature of fish migration should be expanded especially to the fishermen and the communities inhabiting the areas near the water bodies.
Developmental construction like dams and weirs should be provided with fish ladders and way to allow fish effective fish migrations with ease.
Spawning habitats need not to be exploited to avoid premature harvest of young fishes and brood adult females.
The author declares that there are no conflicts of interest regarding the publication of this paper.
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Written By
Gatriay Tut Deng and Birtukan Tsegaye Demisse
Submitted: 24 November 2022Reviewed: 16 January 2023Published: 12 February 2023
Open access peer-reviewed chapter
