The Physiological and Metabolic Adaptations of Salmonids with Respect to Nutrient Requirements and Environmental Influences.


The life of salmonids is greatly dictated by the availability of food and how it is utilised. The nutrition gained from feeding is categorised and then directed down the correct pathways enabling fishes continued growth and reproduction, their sole purpose in life. The environmental influences and changes in lifestyle that create physiological and metabolic adaptations in fish will be discussed in this report along with the basic nutritional requirements of these situations.

A fish needs to be capable of performing chemical processes to survive; these processes are known as their metabolism and measured as a metabolic rate. To calculate this, the carbon dioxide output, the oxygen intake and the excretion of nitrogen together with the calorific value of the excreta needs to be determined (Brown 1957). This happens to be a very complicated process therefore the measurement of oxygen intake alone is commonly used to determine the metabolism. Whilst fish are in a resting state the consumption of oxygen per unit time can be measured, this is known as the basal or minimum metabolic rate. The important factors affecting the metabolism of fish are temperature, oxygen demand and levels of activity. These parameters are closely linked as temperature directly affects the amount of oxygen available in the water and the activity rate of the fish is the largest single factor affecting oxygen consumption.

The metabolic pathway is divided into two distinct routes; the anabolic pathway that uses materials in the formation of new body tissues and the catabolic pathway that assimilates then degrades food into the digestive system. These two pathways are in competition and for growth to occur the anabolic production must be the greater of the two. The supply of materials needed for the basic maintenance of a fish are provided by nutrients, this means that the energy needed for every day activity comes directly from their food. Energy cannot be gained just by consumption of food; firstly, it needs to be absorbed by the digestive system. The energy that can be gained from the food needs releasing by oxidation; this is why the intake and use of oxygen can be a measure of the metabolism.

The main aim of feeding is to provide energy for the metabolic process with the surplus being used for reproduction and growth. The diet of a fish contains various macro-nutrients such as fats, which are available in the form of lipids, proteins (amino acids) and carbohydrates; with micro-nutrients (vitamins and trace elements) being the essential co-factors in the metabolic transfer.

Generally salmonids are predatory so their intake of food consists mainly of animal material therefore carbohydrates are not a substantial part of their diet. When carbohydrates are ingested they have the tendency to become glucose and then stored as glycogen, others are broken down into pyruvate. The completed oxidation process occurs as part of the citric acid cycle, the final progress for most elements of ingested food and in doing so freeing amino acids for growth. Excess carbohydrates will form lipids which can appear as muscle oils, visceral fats or cause fatty degeneration around the liver (Smith 1982). Cheap forms can be added to artificial salmonid diets in quantities of approximately 20% dry weight. Carbohydrates contain around 4.1 Kcal/gram of energy and manipulation of this varies significantly with the ability to digest it efficiently.

The quantities of protein that salmonids require vary with water temperatures and the age of the fish. Smith (1982) suggests that the rapid growth in salmonid fry can only be maximised when a 50% protein content is present in the diet. This compares to a one-year-old fish that only needs a 35% content. His studies on chinook salmon (Oncorhynchus tshawytsha) revealed that 40% of protein at a temperature of 7°c was needed for maximum growth, rising to 50% when the temperature was increased to 15°c. Activity levels increase when oxygen contents rise, which in turn vary with temperature fluctuations. The intake of certain food types must therefore satisfy the metabolism before excess is supplied for growth. The example of the chinook shows that small increases in these variables can lead to huge variations in nutritional demands, which are needed to continue the curve of natural growth. The salmonid can deal efficiently with proteins, for example, there are 4.65 Kcal/gram of energy available with 3.9 Kcal/gram being manipulated, an 85% efficiency rate.

Proteins consist of twenty-one amino acids that are all utilised by the fish. There are ten 'essential' amino acids that determine the quality of the protein produced; these include arginine, histidine, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan and valine (Hoar & Randall 1969). Fish struggle to break down the essential amino acids and in trying to do so they waste more energy that can be gained therefore the non-essential amminos will be the first used for energy production.



Figure 1: A graph showing the relationship between the increase of weight against variations of fat supplements in a salmonids diet. Source: Halver et al. (1973), p.11
  The final group of macro nutrients needed by salmonids is the fats; these come in the form of lipids. The majority of work done in sustained swimming will be created by the fishes red muscle; this muscle requires energy from the oxidation of these lipids. The need for lipids in the diet of the salmonid is essential as their daily living programme receives routine energy from lipid metabolism (Smith, 1982). Fats contain the highest energy content of the macro nutrients with up to 9.45 Kcal/gram, 8.0 Kcal/gram of which is utilised by salmonids (Smith 1982). The growth of the rainbow trout (Oncorhynchus mykiss) tends to be very slow when on a fat free diet and Smith (1982) suggests that 3-15% of lipid content is required in their natural diet. This diet should contain at least 1% of linolenic acid which is essential for growth, and linoleic acid which is not of the same importance should not exceed 2% in total. Figure 1 represents the relationship of the average weekly growths with and without these acids. The importance of linolenic acid is clear if compared to the average weight when fed on a fat free diet.
Fish are the same as all animals in their requirements for certain vitamins. The physiological status of the fish dictates the demand for each vitamin, influences such as stress and illness being the common factors requiring an increase in supply. Table one outlines the specific vitamins needed by salmon and trout, giving a brief description of the symptoms if they are lacking from their diet. It is clear that all are essential co-factors in the metabolic transfer with biotin, folic acid, chlorine and inositol having a very important affect in the process of growth.
  Table 1: The Essential Vitamins and there requirements by Trout and Salmon.
Source: Adapted from Hoar & Randall, (1969), p.409. With previous references to Phillips and Brockway (1957), Poston (1964, 1965, 1967), Phillips (1963), Kitamura et al. (1965), Halver (1957), DeLong et al. (1958) and Coates and Halver (1958).

 
 
Vitamin Required Symptoms if missing from diet
Tocopherol Trout - yes Trout - Increased mortality
  Salmon - yes Salmon - Anemia, Poor growth
p-Aminobenzoic acid Trout - yes Trout - Increased severity of anemia when deficient in folic acid
  Salmon - no Salmon - none
Biotin Trout - yes Trout - No growth, high mortality
  Salmon - yes Salmon - Muscle atrophy
Vitamin B12 Trout - yes Trout - Reduced growth
  Salmon - yes Salmon - Erratic hemoglobin and erythrocyte counts
Folic Acid Trout - yes Trout - Anemia, reduced growth
  Salmon - yes Salmon - Anemia, reduced growth
Ascorbic Acid Trout - yes Trout - Internal hemorrhaging
  Salmon - ? Salmon - ?
Nicotinic acid Trout - yes Trout - Reduced growth, susceptible to suburn
  Salmon - yes Salmon - Muscle spasms, susceptible to suburn
Riboflavin Trout - yes Trout - little growth, opaque eyes
  Salmon - yes Salmon - Cloudy lens
Chlorine Trout - yes Trout - Reduced growth
  Salmon - yes Salmon - Poor growth
Thiamine Trout - yes Trout - High mortality
  Salmon - yes Salmon - Loss of equilibrium, muscle atrophy
Vitamin K Trout - yes Trout - Retarded blood coagulation
  Salmon - ? Salmon - ?
Inositol Trout - yes Trout - Reduced growth
  Salmon - yes Salmon - Poor growth
Pyridoxine Trout - yes Trout - Complete mortality in 6-12 weeks
  Salmon - yes Salmon - Anemia, nervous disorders
Pantothenic acid Trout - yes Trout - Poor growth, high losses
  Salmon - yes Salmon - Clubbed gills
 


Figure 2: The effect of water temperature on the metabolic rate of trout and salmon. Source: Halver et al. (1973), p.2
 

The important areas of nutrition have been discussed, but the changes in the surrounding environment will have an impact on quantities of these nutrients that are needed. The fish must be sure that when under resting conditions, minimal energy required is available to support the body.

One of the main influencing factors on metabolism is temperature, Schaeperclaus (1933) found that a 10°c increase almost doubled the activity of the metabolic rate in fish. When the temperature of the water was increased by one degree celsius, Phillips et al. (1960b) discovered that brook trout (Salvelinus fontinalis), whilst being starved, increased their weight loss by 10%. Figure 2 shows the relationship of water temperature to metabolic rate. The percentage of metabolic rate was measured at the standard environmental temperatures of 10°c for salmon and 15°c for trout (Halver et al. 1973). This graph clearly shows the influence that temperature has on metabolism, with small temperature increases almost doubling the salmon's metabolic rate.

Escalating water flow is associated with increases in oxygen levels, but requirements for energy will also rise. Although the initial thought would be directed towards energy requirements needed for the swimming against currents, it is commonly forgotten that energy is also required when fish are just trying to hold position in their chosen habitat.

An interesting part of some salmonid lifecycles includes migration, which entails periods of starvation. An experiment conducted on starved brook trout found that energy requirements reduced daily for the first three days, then stabilised for the following ten days. Beamish (1964c) concluded that he had discovered a minimum energy level occurring at the point of stabilisation.

The process of reproduction causes fluctuations in the metabolism for various reasons. Energy is required in the formation of ova and milt, which can be up to 33% of the body weight in females, and 3.7% in males (Hoar & Randall 1969). With these organs being so large, energy gained from food that would have been utilised in other areas of the body system will be lost in the form of calories stored in the ova and milt. This will increase the energy requirements of the fish with extra energy needed during spawning and recovery. Salmonids can be categorised into having indeterminate growth patterns. Although at ever decreasing rates, growth will continue through out the life of the fish, in contrast, fish that have a fixed final size are said to have determinate patterns (Purdom 1993). The difference being salmonids will use surplus energy in creating body mass, which will be turned into sexual organs when necessary. This is compared to the direct metabolism of food immediately into gametes, which is the practise of fish with determinate growth patterns. All of these points will contribute to significant changes in the nutrient requirements and oxygen demands, both affecting the metabolism of the fish.

The role of cortisol production has been found highly important to a salmon when running to its spawning grounds. Due to the high stress levels encountered when travelling such distances, the fish will respond by releasing cortisol, the main interrenal steroid of teleosts. The secondary and tertiary responses involved with energy metabolism can be a direct or indirect result of this hormone (Iwama & Nakanishi 1996). The cortical tissue secretes gluco-corticoids, which have an effect on the level of blood sugars produced via carbohydrate metabolism. There is also some influence on the osmoregulation; this is down to the secretion of mineral-corticoids (Lagler et al. 1977). While Smith (1982) was investigating the cortisol levels in salmon he discovered that fish caught offshore and held in captivity increased their levels 4-5 times over a period of 24 hours. These results were compared to running salmon that increased their levels of cortisol naturally as they migrated upstream. The levels peaked at the spawning grounds and the quantities were found to be the same as his fish being held in captivity. He concluded that the starting level of cortisol in salmon was not important as the final quantity was fixed to reach a standard maximum. Cortisol functions in intermediary metabolism are fundamental, as teleost fish require them as an essential factor of life (Hoar & Randall 1988). It is suggested by Hoar & Randall (1988) and Iwama & Nakanishi (1996) that the immune system and related stress actions may need cortisol as a major catalyst.

At sometime in a salmon's life cycle, they feel the need to turn downstream and head towards the sea. Unlike the mortal spawning migrations of many species of salmon, smolt migration must be successful so the fish are available to return to spawn. Major physiological changes occur, which involve the loss of their parr marks that make way for silvery body coloration's. The parr tend to become restless just before the migration; Smith (1982) suggested that an increased thyroxin level that increases the basal metabolism was accountable for this behaviour. Table 2 shows many of the physiological characteristics of smolts in comparison to their original state as parr.


  Table 2: The general physiological changes of parr to smolt transformation of salmon. Source: From Smith (1982) with previous reference to Wedemeyer and Yasutake (1977).

 
 
Physiological Characteristic Level in smolt, compared with that in parr
Thyroid section Increases
Gill microsome, NA+ -K+ ATPase enzyme activity Increases
Buoyancy (swim bladder, Atlantic salmon) Increases
Liver glycogen Decreases
Body silvering Increases
Oxygen consumption Increases
Migratory behaviour Increases
Weight per unit length (condition factor) Decreases
Hyposmotic regulatory capability Increases
Blood glucose Increases
Body total lipid content Decreases
Ammonia production Increases
Salinity tolerance and preference Increases
Ability to grow well in full strength sea water (salinity 35%) Increases
 

These changes in physiology are essential in gaining the smolt the correct balance in their body system when having to face sudden increases in salinity. Although smolts and adults are both subjected to these changes in salinity, lower stress levels in smolts could suggest that they adapt more efficiently to the new environment. The age of the fish does not seem to dictate the time of smoltification, but their size seems to be very influential. Smith (1982) realised that coho salmon (Oncorhynchus kisutch) could start running safely as soon as they reach a fork length of 68-70mm. This length can be gained in 1.5 years, but some have been found to run in the same year that they have hatched.

The final point on smoltification involves the possibility that migration is not completed. Any of the changes a parr may have experienced will revert to the conditions suited to freshwater. The metabolic rate that had increased will reduce and the displaying of unsettled behaviour patterns will cease, completing the full return to parr characteristics.

It is clear that fish have many situations to overcome when trying to complete their journey through life. As with most animals the careful selection of correct food types will dictate their existence, reproductive qualities and growth rates. This report has opened my eyes to the complex world of fish nutrition, metabolic rates, and the effects they have on their life styles and a good understanding of previously unknown areas has been the result.


References

Beamish, F.W.H. (1964c). Influence of starvation on standard and routine oxygen consumption. Trans. Am. Fisheries Society, 93, 103-107
Brown, M.E. (1957). The Physiology of Fishes Volume 1:Metabolism. Academic Press, New York. Coates, J.A., and Halver, J.E. (1958). Water soluble vitamin requirements of silver salmon. U.S. Fish Wildlife Serv., Spec. Sci. Rept., Fisheries, 281, 1-9.
DeLong, D.C., Halver, J.E., and Mertz, E.T. (1958). Nutrition of salmonid fishes. VI. Protein requirements of chinook salmon at two water temperatures. Journal of Nutrition, 65, 589- 599. Halver, J.E. (1957). Nutrition of salmonid fishes. III. Water soluble vitamin requirements of chinook salmon. Journal of Nutrition, 62, 225-243.
Halver, J.E., Coates, J.A., Deyoe, C.W., Dupree, H.K., Post, G., and Sinnhuber, R.O. (1973). Nutrient requirements of trout, salmon and catfish. National Academy of Sciences, Washington, D.C. Hoar, W.S., and Randall, D.J. (1969). Fish Physiology Volume 1: Excretion, Ionic regulation, and Metabolism. Academic Press, New York.
Hoar, W.S., and Randall, D.J. (1988). Fish Physiology Volume XI, Part B: The Physiology of developing fish. Academic Press, New York
Iwama, G., and Nakanishi, T. (1996). Fish Physiology Volume XV: The fish immune system. Academic Press, New York.
Kitamura, S., Ohara, S., Suwa, T., and Nakagawa, K. (1965). Studies on vitamin requirements of rainbow trout. I. On the ascorbic acid. Bull. Japan. Soc. Sci. Fisheries, 31, 818-826.
Lagler, K.F., Bardach, J.E., Miller, R.R., and Passino, D.R. (1977). Ichthyology. John Wiley & Sons, New York.
Phillips, A.M., and Brockway, D.R. (1957). The nutrition of trout. IV. Vitamin requirements. Progressive Fish Culturist, 19, 119-123.
Phillips, A.M. (1963). Folic acid as an anti-anemia factor for brook trout. Progressive Fish Culturist, 25, 132-134.
Phillips, A.M., Livingston, D.L., and Dumas, R.F. (1960b). Effect of starvation and feeding on the chemical composition of brook trout. Progressive Fish Culturist, 22, 147-154.
Poston, H.A. (1964). Effect of dietary vitamin K and sulfaguanidine on blood coagulation time, microhematocrit and growth of immature brook trout. Progressive Fish Culturist, 26, 59-64.
Poston, H.A. (1965). Effect of dietary vitamin E on microhematocrit, mortality and growth of immature brown trout. Cortland Hatchery Report 33 for the year 1964. Fisheries Research. Bulletin. 28, 6-10. Poston, H.A. (1967). Effect of L-ascorbic acid on immature brook trout. Cortland Hatchery Report 35 for the year 1964. . Fisheries Research. Bulletin. 30, 46-51.
Purdom, C.E. (1993). Genetics and fish breeding. Chapman & Hall, London.
Schaeperclaus, W. (1933). Textbook of pond culture. U.S Fish Wildlife Services, Fishery Leaflet, 311, 1-240.
Smith, L.S. (1982). Introduction to Fish Physiology. TFH Publications Ltd, Seattle.
Wedemeyer, G.A. and Yasutake, W.T. (1977). Clinical methods for the assessment of the effects of environmental stress on fish health. Technical Papers of the U.S. Fish and Wildlife Service, 89, p18

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