Short profile


FishEthoScore of the species

Abbreviated assessment of the species' likelihood and potential for fish welfare in aquaculture, based on ethological findings for 10 crucial criteria.

Criteria Li Po Ce
1 Home range ?
2 Depth range ?
3 Migration ?
4 Reproduction
5 Aggregation ? ?
6 Aggression
7 Substrate
8 Stress
9 Malformation
10 Slaughter
FishEthoScore 1 2 2
Li = Likelihood that the individuals of the species experience welfare under minimal farming conditions
Po = Potential overall potential of the individuals of the species to experience welfare under improved farming conditions
Ce = Certainty of our findings in Likelihood and Potential
 
                    ?     /  
  High    Medium     Low     Unclear  No findings
 
FishEthoScore = Sum of criteria scoring "High" (max. 10)



General remarks

Despite the fact that Salmo salar is the most farmed fish in Europe, more fine-tuned culture strategies are needed to improve fish welfare and performance of this species. The low FishEthoScore is mainly due to the dependence on fish in the diet, home range needs, high levels of aggression, needs of substrate, stress under farming conditions and high levels of deformations. 
It is recommended to ensure proper space, at least in the vertical sense, according to the biological needs which seems to be satisfied in sea cages, but not in raceways. The development of new rearing strategies to optimise the husbandry practices such as matching the biological rhythms with e.g. feeding activities or unavoidable but often stressful husbandry procedures would be a step forward to solving some specific welfare concerns, to prevent poor welfare and to minimise stress, improving fish welfare, fish performance and reduce stress. Replacing fish meal and fish oil in the feed by plant-based or sustainable sources would ensure an ethical food production. Semi-intensive and extensive farming could be a remediation for some of the current problems and help improving fish welfare and performance. 


1. Are minimal farming conditions likely to provide the home range of the species? What overall welfare potential can be achieved? How certain are these findings?

?
 Likelihood
M
 Potential
M
 Certainty

ALEVINS, FRY: WILD: planktonic as ALEVINS [1]: horizontal movement limited to hydrodynamic displacement. FRY move 1-5 m from the redd [2]. FARM: hatchery: most common, vertical trays: variable size frames: 60 cm wide 44-175 cm high (Aquatec.com).

PARR: WILD: usually 0.1-8 km [3] [4] [5] [1] [6] [7]. FARM: tanks: 0.14 m3 [8].

SMOLTS: WILD ➝ PARR. FARM: tanks: 9 m3 [8], 500-1300 m3, 14.5-20 m diameter [9]; sea cages: 24 x 24 m or 100 m in diameter [10]; 16,000-130,000 m3 [11].

ADULTS: ➝ SMOLTS.

SPAWNERS: WILD: usually 0.1-4.4 km [12]. FARM: 46-90 m3 [8].


2. Are minimal farming conditions likely to provide the depth range of the species? What overall welfare potential can be achieved? How certain are these findings?

?
 Likelihood
H
 Potential
M
 Certainty

ALEVINS and FRY: WILD: usually <1 m [13] [14] [15] [16]. FARM: hatchery trays and tanks: 20-50 cm [17].

PARR and SMOLTS: WILD: <6.5 m [13] [18] [19] [14] [15] [16] [4] [20] [6] [21] [7] [22]. SMOLTS: FARM: tanks: 3.5 to 4.5 m [9];  sea cages: maximum 18 m [10]. 

ADULTS: WILD: further research needed to identify depth range in the wild. FARM: sea cages: 5 m [23], 10 m [24], 15 m [25], 40 m [26], maximum 50 m [11].

KELT: WILD: stay close to the surface [27] [28]. 

SPAWNERS: WILD: usually 0.5-3 m within the spawning season [29]. FARM: no data found yet on spawning culture conditions.


3. Are minimal farming conditions compatible with the migrating or habitat-changing behaviour of the species? What overall welfare potential can be achieved? How certain are these findings?

?
 Likelihood
M
 Potential
M
 Certainty

ANADROMOUS [30] [31] [1] [32] [33] [5]: migrate to the sea in spring-early summer, individuals return as grilse to their rivers of origin to spawn.

ALEVINS, FRY and PARR: WILD: stationary [4] [34] [21]. FARM: rearing in freshwater tanks 0-10 ppm [35] [10]. Stressed by use of artificial light at night [36].

SMOLTS: WILD: migrate along freshwater to the sea [5]. FARM: rearing in halocline and brackish water [37] [25] [38].

ADULTS: WILD: lives in brackish and saltwater [39] [33]. FARM: rearing in brackish and saltwater cages 20-34 ppm [38] [40] [41] [42].

SPAWNERS: WILD: return as GRILSE to natal river to spawn [12]. FARM: rearing in freshwater tanks <10 ppm [43].

All age classes: further research needed on welfare parameters to determine whether presenting species with conditions of different migratory phases indeed satisfies their urge to migrate or whether they need to experience the transition.


4. Is the species likely to reproduce in captivity without manipulation? What overall welfare potential can be achieved? How certain are these findings?

L
 Likelihood
M
 Potential
M
 Certainty

WILD: from June to November [44] [45]. The most dominant male salmon perform the majority of the courting and mating behaviours with the female [46]. Female builds redd [47] [48] [49] [50] [51]. FARM: under farming conditions, eggs and milt are stripped [10] [17]. 


5. Is the aggregation imposed by minimal farming conditions likely to be compatible with the natural behaviour of the species? What overall welfare potential can be achieved? How certain are these findings?

?
 Likelihood
?
 Potential
L
 Certainty

ALEVINS and FRY: WILD: live solitary after emerging from the gravel but remain in the same areas as siblings [2] [52] [1]. FARM: intensive conditions: 50 kg/m3 or higher [10]; 21-86 kg/m3 [53].

PARR: WILD: live solitary or in small groups [54]. Further research needed on extension of groups in the wild. FARM: extensive conditions: 8 kg/m3; intensive conditions: 30 kg/m3 [55], 21-86 kg/m3 [53].

SMOLTS: WILD: live in schools [56] [54] [1] [57]. Further research needed on extension of schools in the wild. FARM: under farming conditions, 15-35 kg/m3 [58] [24]: usually 20 kg/m3 [10]. < 22 kg/m3 best welfare according Salmon Welfare Index model [38].

ADULTS: ➝ SMOLTS.

SPAWNERS: live in schools [56] [54] [1] [57]. Further research needed on extension of schools in the wild. Rearing at low densities with unknown extension [10].


6. Is the species likely to be non-aggressive and non-territorial? What overall welfare potential can be achieved? How certain are these findings?

L
 Likelihood
M
 Potential
M
 Certainty

ALEVINS and FRY: WILD: aggressive and territorial after emerging from the gravel [2]. LAB: aggressive in pairwise interactions [59] [60] and groups of 10 [52]; aggression increases with fish density, from 3 to 9 individuals/m2. FARM: no data found yet on aggression behaviour under farming conditions.

PARR: WILD: territorial and aggressive [61] [62]. LAB: individual differences in metabolism related aggressive behaviour [63]. FARM: more aggressive at high 30 kg/m3 than low 8 kg/m3 density [55], aggression increases with feed restriction [64].

SMOLTS: WILD: schooling behaviour [56] [61] [62] [57]. FARM: lower levels of aggression at feeding time under 25 kg/m3 than under 15 and 30 kg/m3 [58]. Not aggressive in 1,000-1,200 m3 sea-cages of 10 x 10 x 15 m at 0.85 kg/m3 [24]. 

ADULTS: ➝ SMOLTS. More aggressive during matuaration [65], more aggressive at low speed currents [66].

SPAWNERS: WILD: schooling behaviour during migration [56] [61] [62] [57]. Dominance hierarchy during the spawning period in the wild [67] and in the lab [68]. FARM: non-linear dominance hierarchies at 15.2 kg/m3 [69].


7. Are minimal farming conditions likely to match the natural substrate and shelter needs of the species? What overall welfare potential can be achieved? How certain are these findings?

L
 Likelihood
M
 Potential
H
 Certainty

ALEVINS and FRY:LAB: absence of substrate did not affect the ontogeny and behaviour [52], but the use of artificial substrate (polyethylene astro-turf) favoured growth [70]; low survival in sand compared to rocks and stones [71]. FARM: under farming conditions, use artificial hatching substrate [10] [17].

PARR, SMOLTS, ADULTS: WILD: prefer habitats with rocks and stones [6]; use substrate as cover from predators and adverse environmental conditions [21] [72] [73] [74]. FARM: providing cover increased growth rate and improved stress [75]; smoltification process dependent on the number of shelters provided [76]. LAB: vertically-suspended structures provided [77].

SPAWNERS: WILD: use substrate to build redds [49] [21]. FARM: no data found yet on the effect of missing environmental enrichment in farming conditions.


8. Are minimal farming conditions (handling, confinement etc.) likely not to stress the individuals of the species? What overall welfare potential can be achieved? How certain are these findings?

L
 Likelihood
M
 Potential
M
 Certainty

ALEVINS and FRY: stressed by confinement [60] [78]. For stress and light at night ➝ crit. 3.

PARR and SMOLTS: stressed by acute handling [62], temperature shock [79] [80] [81], chasing, netting, noise, sudden darkness with intermittent light, hypoxia and emptying the tank [81]. SMOLTS: stressed more by loading than transport [82].

ADULTS: stressed by chilling, crowding [79] and handling [83].

 


9. Are malformations of this species likely to be rare under farming conditions? What overall welfare potential can be achieved? How certain are these findings?

L
 Likelihood
M
 Potential
M
 Certainty

ALEVINS and FRY: WILD: no data found yet on frequency of malformations. FARM: vertebral column deformities in >10% of individuals [78] [84]. 

PARR and SMOLTS: WILD: no data found yet on frequency of malformations. FARM: vertebral and spinal deformities in >10% of individuals [85] [86] [87] [88] [89] [90].

ADULTS: WILD: vertebral deformities in >10% of individuals, severity of malformations are low compared with farmed salmon [91]. FARM: vertebral and spinal deformities in >10% of individuals [85] [87] [88] [89].


10. Is a humane slaughter protocol likely to be applied under minimal farming conditions? What overall welfare potential can be achieved? How certain are these findings?

H
 Likelihood
H
 Potential
H
 Certainty

Common and high-standard slaughter method: a protocol for electrical and percussive stunning and killing by bleeding is available [92] [93] [94] [95].


Side note: Domestication

DOMESTICATION LEVEL 5 [96] [97], fully domesticated. Cultured since 19th century [10].


Side note: Feeding without components of forage fishery

WILD: carnivorous [10] [29] [98] [99] [100]. FARM: fish meal and fish oil may be mostly* replaced by plant protein [101] [102] [103] [104] [105], but no data found yet for ALEVINS and FRY.

* partly = <51% – mostly = 51-99% – completely = 100%


Glossary

ADULTS = mature individuals, for details ➝ Findings 10.1 Ontogenetic development
ALEVINS = larvae until the end of yolk sac absorption, for details ➝ Findings 10.1 Ontogenetic development
ANADROMOUS = migrating from the sea into fresh water to spawn
DOMESTICATION LEVEL 5 = selective breeding programmes are used focusing on specific goals [96]
FARM = setting in farm environment
FRY = larvae from external feeding on, for details ➝ Findings 10.1 Ontogenetic development
GRILSE = adults returning from sea to home river to spawn, for details ➝ Findings 10.1 Ontogenetic development
KELT = adults surviving spawning, for details ➝ Findings 10.1 Ontogenetic development
LAB = setting in laboratory environment
PARR = juvenile stage in rivers, for details ➝ Findings 10.1 Ontogenetic development
SMOLTS = juvenile stage migrating to the sea, for details ➝ Findings 10.1 Ontogenetic development
SPAWNERS = adults that are kept as broodstock
WILD = setting in the wild


Bibliography

[1] McCormick, Stephen D, Lars P Hansen, Thomas P Quinn, and Richard L Saunders. 1998. Movement, migration, and smolting of Atlantic salmon (Salmo salar). Canadian Journal of Fisheries and Aquatic Sciences 55: 77–92. https://doi.org/10.1139/d98-011.
[2] Gustafson-Greenwood, Karla I., and John R. Moring. 1990. Territory size and distribution of newly-emerged Atlantic salmon (Salmo salar). Hydrobiologia 206: 125–131. https://doi.org/10.1007/BF00018638.
[3] Hesthagen, T. 1990. Home range of juvenile Atlantic salmon, Salmo salar, and brown trout, Salmo trutta, in a Norwegian stream. Freshwater Biology 24: 63–67. https://doi.org/10.1111/j.1365-2427.1990.tb00307.x.
[4] Erkinaro, J. 1995. The age structure and distribution of Atlantic salmon parr, Salmo salar L., in small tributaries and main stems of the subarctic River Teno, northern Finland. Ecology of Freshwater Fish 4: 53–61. https://doi.org/10.1111/j.1600-0633.1995.tb00117.x.
[5] Elliott, Scott R, Treva A Coe, James M Helfield, and Robert J Naiman. 1998. Spatial variation in environmental characteristics of Atlantic salmon (Salmo salar) rivers. Canadian Journal of Fisheries and Aquatic Sciences 55: 267–280. https://doi.org/10.1139/d98-001.
[6] Girard, Isabelle L, James W.A Grant, and Stefán Ó Steingrímsson. 2004. Foraging, growth, and loss rate of young-of-the-year Atlantic salmon (Salmo salar) in relation to habitat use in Catamaran Brook, New Brunswick. Canadian Journal of Fisheries and Aquatic Sciences 61: 2339–2349. https://doi.org/10.1139/f04-216.
[7] Roussel, Jean-Marc, Richard A. Cunjak, Robert Newbury, Daniel Caissie, and Alexander Haro. 2004. Movements and habitat use by PIT-tagged Atlantic salmon parr in early winter: the influence of anchor ice. Freshwater Biology 49: 1026–1035. https://doi.org/10.1111/j.1365-2427.2004.01246.x.
[8] Wolters, William, Amanda Masters, Brian Vinci, and Steven Summerfelt. 2009. Design, loading, and water quality in recirculating systems for Atlantic Salmon (Salmo salar) at the USDA ARS National Cold Water Marine Aquaculture Center (Franklin, Maine). Aquacultural Engineering 41. Design, Loading, and Water Quality in Recirculating Systems for Salmonids: 60–70. https://doi.org/10.1016/j.aquaeng.2009.06.011.
[9] Summerfelt, Steven T., Frode Mathisen, Astrid Buran Holan, and Bendik Fyhn Terjesen. 2016. Survey of large circular and octagonal tanks operated at Norwegian commercial smolt and post-smolt sites. Aquacultural Engineering 74: 105–110. https://doi.org/10.1016/j.aquaeng.2016.07.004.
[10] Jones, M. 2004. Cultured Aquatic Species Information Programme. Salmo salar. Rome: FAO Fisheries and Aquaculture Department.
[11] Noble, Chris, Kristine Gismervik, Martin Haugmo Iversen, Jelena Kolarevic, Jonatan Nilsson, Lars Helge Stien, and James F Turnbull. 2018. Welfare indicators for farmed Atlantic salmon: Tools for assessing fish welfare.
[12] Baglinière, J. L., G. Maisse, and A. Nihouarn. 1990. Migratory and reproductive behaviour of female adult Atlantic salmon, Salmo salar L., in a spawning stream. Journal of Fish Biology 36: 511–520. https://doi.org/10.1111/j.1095-8649.1990.tb03553.x.
[13] Symons, P. E. K., and M. Heland. 1978. Stream Habitats and Behavioral Interactions of Underyearling and Yearling Atlantic Salmon (Salmo salar). Journal of the Fisheries Research Board of Canada 35: 175–183. https://doi.org/10.1139/f78-029.
[14] Morantz, D. L., R. K. Sweeney, C. S. Shirvell, and D. A. Longard. 1987. Selection of Microhabitat in Summer by Juvenile Atlantic Salmon (Salmo salar). Canadian Journal of Fisheries and Aquatic Sciences 44: 120–129. https://doi.org/10.1139/f87-015.
[15] Heggenes, Jan. 1990. Habitat utilization and preferences in juvenile atlantic salmon (salmo salar) in streams. Regulated Rivers: Research Management 5: 341–354. https://doi.org/10.1002/rrr.3450050406.
[16] Crisp, D. Trevor. 1993. The environmental requirements of salmon and trout in fresh water. Freshwater Forum 3: 176–202.
[17] Stead, Selina M., and Lindsay Laird. 2002. The Handbook of Salmon Farming. Food Sciences. London: Springer-Verlag.
[18] Rimmer, D. M., U. Paim, and R. L. Saunders. 1984. Changes in the Selection of Microhabitat by Juvenile Atlantic Salmon (Salmo salar) at the Summer–Autumn Transition in a Small River. Canadian Journal of Fisheries and Aquatic Sciences 41: 469–475. https://doi.org/10.1139/f84-056.
[19] Degraaf, D. A., and L. H. Bain. 1986. Habitat Use by and Preferences of Juvenile Atlantic Salmon in Two Newfoundland Rivers. Transactions of the American Fisheries Society 115: 671–681. https://doi.org/10.1577/1548-8659(1986)115671:HUBAPO2.0.CO;2.
[20] Bremset, Gunnbjørn, and Ole Kristian Berg. 1999. Three-dimensional microhabitat use by young pool-dwelling Atlantic salmon and brown trout. Animal Behaviour 58: 1047–1059. https://doi.org/10.1006/anbe.1999.1218.
[21] Guay, J C, D Boisclair, M Leclerc, and M Lapointe. 2003. Assessment of the transferability of biological habitat models for Atlantic salmon parr (Salmo salar). Canadian Journal of Fisheries and Aquatic Sciences 60: 1398–1408. https://doi.org/10.1139/f03-120.
[22] Davidsen, J. G., N. Plantalech Manel-la, F. Økland, O. H. Diserud, E. B. Thorstad, B. Finstad, R. Sivertsgård, R. S. McKinley, and A. H. Rikardsen. 2008. Changes in swimming depths of Atlantic salmon Salmo salar post-smolts relative to light intensity. Journal of Fish Biology 73: 1065–1074. https://doi.org/10.1111/j.1095-8649.2008.02004.x.
[23] NOT FOUND
[24] Dempster, Tim, Jon-Erik Juell, Jan Erik Fosseidengen, Arne Fredheim, and Pål Lader. 2008. Behaviour and growth of Atlantic salmon (Salmo salar L.) subjected to short-term submergence in commercial scale sea-cages. Aquaculture 276: 103–111. https://doi.org/10.1016/j.aquaculture.2008.01.018.
[25] Johansson, David, Kari Ruohonen, Jon-Erik Juell, and Frode Oppedal. 2009. Swimming depth and thermal history of individual Atlantic salmon (Salmo salar L.) in production cages under different ambient temperature conditions. Aquaculture 290: 296–303. https://doi.org/10.1016/j.aquaculture.2009.02.022.
[26] Cardia, Francesco, and Alessandro Lovatelli. 2015. Aquaculture operations in floating HDPE cages: a field handbook. FAO Fisheries and Aquaculture Technical Paper 593. Rome: Food and Agriculture Organization of the United Nations.
[27] Hubley, P. Bradford, Peter G. Amiro, A. Jamie F. Gibson, Gilles L. Lacroix, and Anna M. Redden. 2008. Survival and behaviour of migrating Atlantic salmon (Salmo salar L.) kelts in river, estuarine, and coastal habitat. ICES Journal of Marine Science: Journal du Conseil 65: 1626–1634. https://doi.org/10.1093/icesjms/fsn129.
[28] Halttunen, Elina, Audun H. Rikardsen, Jan G. Davidsen, Eva B. Thorstad, and J. Brian Dempson. 2009. Survival, Migration Speed and Swimming Depth of Atlantic Salmon Kelts During Sea Entry and Fjord Migration. In Tagging and Tracking of Marine Animals with Electronic Devices, ed. Jennifer L. Nielsen, Haritz Arrizabalaga, Nuno Fragoso, Alistair Hobday, Molly Lutcavage, and John Sibert, 35–49. Reviews: Methods and Technologies in Fish Biology and Fisheries 9. Springer Netherlands.
[29] Froese, R., and D. Pauly. 2014. FishBase. World Wide Web electronic publication. www.fishbase.org.
[30] Moore, A., E. C. E. Potter, N. J. Milner, and S. Bamber. 1995. The migratory behaviour of wild Atlantic salmon (Salmo salar) smolts in the estuary of the River Conwy, North Wales. Canadian Journal of Fisheries and Aquatic Sciences 52: 1923–1935. https://doi.org/10.1139/f95-784.
[31] Armstrong, John D, James WA Grant, Harvey L Forsgren, Kurt D Fausch, Richard M DeGraaf, Ian A Fleming, Terry D Prowse, and Isaac J Schlosser. 1998. The application of science to the management of Atlantic salmon (Salmo salar): integration across scales. Canadian Journal of Fisheries and Aquatic Sciences 55: 303–311. https://doi.org/10.1139/d98-014.
[32] Hedger, Richard D., Francois Martin, Daniel Hatin, Francois Caron, F. G. Whoriskey, and Julian J. Dodson. 2008. Active migration of wild Atlantic salmon Salmo salar smolt through a coastal embayment. ResearchGate 355: 235–246. https://doi.org/10.3354/meps07239.
[33] Cunjak, R. A., and J. Therrien. 1998. Inter-stage survival of wild juvenile Atlantic salmon, Salmo salar L. Fisheries Management and Ecology 5: 209–223. https://doi.org/10.1046/j.1365-2400.1998.00094.x.
[34] Carr, J. W., J. M. Anderson, F. G. Whoriskey, and T. Dilworth. 1997. The occurrence and spawning of cultured Atlantic salmon (Salmo salar) in a Canadian river. ICES Journal of Marine Science: Journal du Conseil 54: 1064–1073. https://doi.org/10.1016/S1054-3139(97)80010-0.
[35] European Food Safety Authority (EFSA). 2008. Animal welfare aspects of husbandry systems for farmed Atlantic salmon - Scientific Opinion of the Panel on Animal Health and Welfare. European Food Safety Authority. July 22.
[36] Newman, Rhian C., Tim Ellis, Phil I. Davison, Mark J. Ives, Rob J. Thomas, Sian W. Griffiths, and William D. Riley. 2015. Using novel methodologies to examine the impact of artificial light at night on the cortisol stress response in dispersing Atlantic salmon (Salmo salar L.) fry. Conservation Physiology 3: cov051. https://doi.org/10.1093/conphys/cov051.
[37] Johansson, David, Kari Ruohonen, Anders Kiessling, Frode Oppedal, Jan-Erik Stiansen, Mark Kelly, and Jon-Erik Juell. 2006. Effect of environmental factors on swimming depth preferences of Atlantic salmon (Salmo salar L.) and temporal and spatial variations in oxygen levels in sea cages at a fjord site. Aquaculture 254: 594–605. https://doi.org/10.1016/j.aquaculture.2005.10.029.
[38] NOT FOUND
[39] Heggenes, J., J. L. Baglinière, and R. A. Cunjak. 1999. Spatial niche variability for young Atlantic salmon (Salmo salar) and brown trout (S. trutta) in heterogeneous streams. Ecology of Freshwater Fish 8: 1–21. https://doi.org/10.1111/j.1600-0633.1999.tb00048.x.
[40] Bœuf, Gilles, and Patrick Payan. 2001. How should salinity influence fish growth? Comparative Biochemistry and Physiology Part C: Toxicology Pharmacology 130: 411–423. https://doi.org/10.1016/S1532-0456(01)00268-X.
[41] Oppedal, Frode, Tone Vågseth, Tim Dempster, Jon-Erik Juell, and David Johansson. 2011. Fluctuating sea-cage environments modify the effects of stocking densities on production and welfare parameters of Atlantic salmon (Salmo salar L.). Aquaculture 315: 361–368. https://doi.org/10.1016/j.aquaculture.2011.02.037.
[42] Oppedal, Frode, Tim Dempster, and Lars H. Stien. 2020. Environmental drivers of Atlantic salmon behaviour in sea-cages: A review. Aquaculture 311: 1–18. https://www.academia.edu/15253709/Environmental_drivers_of_Atlantic_salmon_behaviour_in_sea_cages_A_review. Accessed August 10.
[43] Taranger, G. L., and T. Hansen. 1993. Ovulation and egg survival following exposure of Atlantic salmon, Salmo salar L., broodstock to different water temperatures. Aquaculture Research 24: 151–156. https://doi.org/10.1111/j.1365-2109.1993.tb00535.x.
[44] Froese, R., and D. Pauly. 2020. Salmo salar, Atlantic salmon summary page. www.fishbase.org. https://www.fishbase.se/summary/salmo-salar.html. Accessed August 10.
[45] FAO. 2014. FAO Fisheries Aquaculture - Species Fact Sheets - Salmo salar (Linnaeus, 1758). World Wide Web electronic publication. www.fao.org.
[46] Fleming, Ian A., and Sigurd Einum. 2010. Reproductive Ecology: A Tale of Two Sexes. In Atlantic Salmon Ecology, 33–65. John Wiley Sons, Ltd. https://doi.org/10.1002/9781444327755.ch2.
[47] Crisp, D. T., and P. A. Carling. 1989. Observations on siting, dimensions and structure of salmonid redds. Journal of Fish Biology 34: 119–134. https://doi.org/10.1111/j.1095-8649.1989.tb02962.x.
[48] Cowx, I. G., K. T. O’Grady, W. C. K. O’Connor, and T. E. Andrew. 1998. The effects of siltation on Atlantic salmon, Salmo salar L., embryos in the River Bush. Fisheries Management and Ecology 5: 393–401. https://doi.org/10.1046/j.1365-2400.1998.550393.x.
[49] Moir, H. J., C. Soulsby, and A. Youngson. 1998. Hydraulic and sedimentary characteristics of habitat utilized by Atlantic salmon for spawning in the Girnock Burn, Scotland. Fisheries Management and Ecology 5: 241–254. https://doi.org/10.1046/j.1365-2400.1998.00105.x.
[50] Soulsby, C., A. F. Youngson, H. J. Moir, and I. A. Malcolm. 2001. Fine sediment influence on salmonid spawning habitat in a lowland agricultural stream: a preliminary assessment. Science of The Total Environment 265: 295–307. https://doi.org/10.1016/S0048-9697(00)00672-0.
[51] Julien, H. P., and N. E. Bergeron. 2006. Effect of Fine Sediment Infiltration During the Incubation Period on Atlantic Salmon (Salmo salar) Embryo Survival. Hydrobiologia 563: 61–71. https://doi.org/10.1007/s10750-005-1035-2.
[52] Dill, Peter A. 1977. Development of behaviour in alevins of atlantic salmon, Salmo salar, and rainbow trout, S. gairdneri. Animal Behaviour 25, Part 1: 116–121. https://doi.org/10.1016/0003-3472(77)90073-2.
[53] Hosfeld, Camilla Diesen, Jannicke Hammer, Sigurd O. Handeland, Sveinung Fivelstad, and Sigurd O. Stefansson. 2009. Effects of fish density on growth and smoltification in intensive production of Atlantic salmon (Salmo salar L.). Aquaculture 294: 236–241. https://doi.org/10.1016/j.aquaculture.2009.06.003.
[54] Gibson, R. J. 1983. Water velocity as a factor in the change from aggressive to schooling behaviour and subsequent migration of Atlantic salmon smolt (Salmo salar). Le Naturaliste canadien.
[55] Cañon Jones, Alberto Hernán. 2011. Social network analysis of behavioural interactions influencing the development of fin damage in Atlantic salmon (Salmo salar). Thesis, University of Cambridge.
[56] Fried, Stephen M., James D. McCleave, and George W. LaBar. 1978. Seaward Migration of Hatchery-Reared Atlantic Salmon, Salmo salar, Smolts in the Penobscot River Estuary, Maine: Riverine Movements. Journal of the Fisheries Research Board of Canada 35: 76–87. https://doi.org/10.1139/f78-011.
[57] Riley, W. D., A. T. Ibbotson, D. L. Maxwell, P. I. Davison, W. R. C. Beaumont, and M. J. Ives. 2014. Development of schooling behaviour during the downstream migration of Atlantic salmon Salmo salar smolts in a chalk stream. Journal of Fish Biology 85: 1042–1059. https://doi.org/10.1111/jfb.12457.
[58] Adams, C E, J F Turnbull, A Bell, J E Bron, and F A Huntingford. 2007. Multiple determinants of welfare in farmed fish: stocking density, disturbance, and aggression in Atlantic salmon (Salmo salar). Canadian Journal of Fisheries and Aquatic Sciences 64: 336–344. https://doi.org/10.1139/f07-018.
[59] Metcalfe, N. B., and J. E. Thorpe. 1992. Early predictors of life-history events: the link between first feeding date, dominance and seaward migration in Atlantic salmon, Salmo salar L. Journal of Fish Biology 41: 93–99. https://doi.org/10.1111/j.1095-8649.1992.tb03871.x.
[60] Vaz-Serrano, J., M. L. Ruiz-Gomez, H. M. Gjoen, P. V. Skov, F. A. Huntingford, Øyvind Øverli, and E. Höglund. 2011. Consistent boldness behaviour in early emerging fry of domesticated Atlantic salmon (Salmo salar): Decoupling of behavioural and physiological traits of the proactive stress coping style. Physiology Behavior 103: 359–364. https://doi.org/10.1016/j.physbeh.2011.02.025.
[61] Gibson, R. John. 1995. Regulation of the fitness of altantic salmon ( Salmo salar ) by intra-specific competition amongst the juveniles. Freshwater Forum A Journal of the Freshwater Biological Association 5: 54–72.
[62] McCormick, S. D, J. M Shrimpton, J. B Carey, M. F O’Dea, K. E Sloan, S Moriyama, and B. Th Björnsson. 1998. Repeated acute stress reduces growth rate of Atlantic salmon parr and alters plasma levels of growth hormone, insulin-like growth factor I and cortisol. Aquaculture 168: 221–235. https://doi.org/10.1016/S0044-8486(98)00351-2.
[63] Cutts, C. J., N. B. Betcalfe, and A. C. Caylor. 1998. Aggression and growth depression in juvenile Atlantic salmon: the consequences of individual variation in standard metabolic rate. Journal of Fish Biology 52: 1026–1037. https://doi.org/10.1111/j.1095-8649.1998.tb00601.x.
[64] Jones, Hernán Alberto Cañon, Chris Noble, Børge Damsgård, and Gareth P. Pearce. 2017. Evaluating the effects of a short-term feed restriction period on the behavior and welfare of Atlantic salmon, Salmo salar, parr using social network analysis and fin damage. Journal of the World Aquaculture Society 48: 35–45. https://doi.org/10.1111/jwas.12322.
[65] Taranger, Geir Lasse, Manuel Carrillo, Rüdiger W. Schulz, Pascal Fontaine, Silvia Zanuy, Alicia Felip, Finn-Arne Weltzien, et al. 2010. Control of puberty in farmed fish. General and Comparative Endocrinology 165: 483–515. https://doi.org/10.1016/j.ygcen.2009.05.004.
[66] Solstorm, Frida, David Solstorm, Frode Oppedal, Rolf Erik Olsen, Lars Helge Stien, and Anders Fernö. 2016. Not too slow, not too fast: water currents affect group structure, aggression and welfare in post-smolt Atlantic salmon Salmo salar. 1869-215X. https://doi.org/10.3354/aei00178.
[67] Fleming, I. A. 2017. Reproductive strategies of Atlantic salmon: ecology and evolution. Rev Fish Biol Fish 6: 379-416. Reviews in Fish Biology and Fisheries 6: 379–416. https://doi.org/http://dx.doi.org/10.1007/BF00164323.
[68] Järvi, Torbjörn. 1990. The Effects of Male Dominance, Secondary Sexual Characteristics and Female Mate Choice on the Mating Success of Male Atlantic Salmon Salmo salar. Ethology 84: 123–132. https://doi.org/10.1111/j.1439-0310.1990.tb00789.x.
[69] Cubitt, K Fiona, Svante Winberg, Felicity A Huntingford, Sunil Kadri, Vivian O Crampton, and Øyvind Øverli. 2008. Social hierarchies, growth and brain serotonin metabolism in Atlantic salmon (Salmo salar) kept under commercial rearing conditions. Physiology behavior 94: 529–35. https://doi.org/10.1016/j.physbeh.2008.03.009.
[70] Hansen, Lars P., and Bror Jonsson. 1985. Downstream migration of hatchery-reared smolts of Atlantic salmon (Salmo salar L.) in the River Imsa, Norway. Aquaculture 45. Salmonid Smoltification II: 237–248. https://doi.org/10.1016/0044-8486(85)90273-X.
[71] Lapointe, Michel F., Normand E. Bergeron, F. Bérubé, M.-A. Pouliot, and P. Johnston. 2004. Interactive Effects of Substrate Sand and Silt Contents, Redd-Scale Hydraulic Gradients, and Interstitial Velocities on Egg-to-Emergence Survival of Atlantic Salmon (Salmo Salar). Can. J. Fish. Aquat. Sci. 61: 2271–2277. https://doi.org/10.1139/f04-236.
[72] Beland, K. F., J. G. Trial, and J. F. Kocik. 2004. Use of Riffle and Run Habitats with Aquatic Vegetation by Juvenile Atlantic Salmon. North American Journal of Fisheries Management 24: 525–533. https://doi.org/10.1577/M02-196.1.
[73] Amiro, Peter G. 2006. A synthesis of fresh water habitat requirements and status for Atlantic salmon (Salmo salar) in Canada. Research Document 2006/017. Dartmouth, Canada: Canadian Science Advisory Secretariat.
[74] Mitchell, J., R. S. McKinley, G. Power, and D. A. Scruton. 1998. Evaluation of Atlantic salmon parr responses to habitat improvement structures in an experimental channel in Newfoundland, Canada. Regulated Rivers: Research Management 14: 25–39. https://doi.org/10.1002/(SICI)1099-1646(199801/02)14:125::AID-RRR4743.0.CO;2-1.
[75] Pickering, A. D., R. Griffiths, and T. G. Pottinger. 1987. A comparison of the effects of overhead cover on the growth, survival and haematology of juvenile Atlantic salmon, Salmo salar L., brown trout, Salmo trutta L., and rainbow trout, Salmo gairdneri Richardson. Aquaculture 66: 109–124. https://doi.org/10.1016/0044-8486(87)90226-2.
[76] Pavlov, D. S., I. V. Nechaev, V. V. Kostin, and N. I. Shindavina. 2008. Influence of shelters and food resources on smoltification of juveniles of the Atlantic salmon Salmo salar. Journal of Ichthyology 48: 605–609. https://doi.org/10.1134/S0032945208080067.
[77] NOT FOUND
[78] Eriksen, M. S., M. Bakken, Å. Espmark, B. O. Braastad, and R. Salte. 2006. Prespawning stress in farmed Atlantic salmon Salmo salar: maternal cortisol exposure and hyperthermia during embryonic development affect offspring survival, growth and incidence of malformations. Journal of Fish Biology 69: 114–129. https://doi.org/10.1111/j.1095-8649.2006.01071.x.
[79] Skjervold, Per Olav, Svein Olav Fjæra, Per Braarød Østby, and Olai Einen. 2001. Live-chilling and crowding stress before slaughter of Atlantic salmon (Salmo salar). Aquaculture 192: 265–280. https://doi.org/10.1016/S0044-8486(00)00447-6.
[80] Foss, A., E. Grimsbø, E. Vikingstad, R. Nortvedt, E. Slinde, and B. Roth. 2012. Live chilling of Atlantic salmon: physiological response to handling and temperature decrease on welfare. Fish Physiology and Biochemistry 38: 565–571. https://doi.org/10.1007/s10695-011-9536-6.
[81] Madaro, Angelico, Rolf E. Olsen, Tore S. Kristiansen, Lars O. E. Ebbesson, Tom O. Nilsen, Gert Flik, and Marnix Gorissen. 2015. Stress in Atlantic salmon: response to unpredictable chronic stress. Journal of Experimental Biology 218: 2538–2550. https://doi.org/10.1242/jeb.120535.
[82] Iversen, Martin, Bengt Finstad, Robert S. McKinley, Robert A. Eliassen, Kristian Tuff Carlsen, and Tore Evjen. 2005. Stress responses in Atlantic salmon (Salmo salar L.) smolts during commercial well boat transports, and effects on survival after transfer to sea. Aquaculture 243: 373–382. https://doi.org/10.1016/j.aquaculture.2004.10.019.
[83] Gatica, M. C., G. E. Monti, T. G. Knowles, P. D. Warriss, and C. B. Gallo. 2010. Efecto del transporte comercial y manejo ante mortem sobre constituyentes sanguíneos de salmones del Atlántico. Archivos de medicina veterinaria 42: 73–78. https://doi.org/10.4067/S0301-732X2010000100010.
[84] Sánchez, R. C., E. B. Obregón, and Mariana Rojas Rauco. 2011. Vertebral Column Deformity and Hypoxia in Salmo salar. ResearchGate 29: 1291–1295.
[85] Gjerde, Bjarne, Ma. Josefa R. Pante, and Grete Baeverfjord. 2005. Genetic variation for a vertebral deformity in Atlantic salmon (Salmo salar). Aquaculture 244: 77–87. https://doi.org/10.1016/j.aquaculture.2004.12.002.
[86] Berg, Arne, Odd Magne Rødseth, Arild Tangerås, and Tom Hansen. 2006. Time of vaccination influences development of adhesions, growth and spinal deformities in Atlantic salmon Salmo salar. Diseases of Aquatic Organisms 69: 239–248. https://doi.org/10.3354/dao069239.
[87] Witten, P. Eckhard, Alex Obach, Ann Huysseune, and Grete Baeverfjord. 2006. Vertebrae fusion in Atlantic salmon (Salmo salar): Development, aggravation and pathways of containment. Aquaculture 258: 164–172. https://doi.org/10.1016/j.aquaculture.2006.05.005.
[88] Fjelldal, Per Gunnar, Tom Johnny Hansen, and Arne Erik Berg. 2007. A radiological study on the development of vertebral deformities in cultured Atlantic Salmon (Salmo salar L.). ResearchGate 273: 721–728. https://doi.org/10.1016/j.aquaculture.2007.07.009.
[89] Aunsmo, A, A Guttvik, P J Midtlyng, R B Larssen, Ø Evensen, and E Skjerve. 2008. Association of spinal deformity and vaccine-induced abdominal lesions in harvest-sized Atlantic salmon, Salmo salar L. Journal of Fish Diseases 31: 515–524. https://doi.org/10.1111/j.1365-2761.2007.00899.x.
[90] Berg, A., A. Yurtseva, T. Hansen, D. Lajus, and P. G. Fjelldal. 2012. Vaccinated farmed Atlantic salmon are susceptible to spinal and skull deformities. Journal of Applied Ichthyology 28: 446–452. https://doi.org/10.1111/j.1439-0426.2012.01988.x.
[91] Sambraus, F., K. A. Glover, T. Hansen, T. W. K. Fraser, M. F. Solberg, and P. G. Fjelldal. 2014. Vertebra deformities in wild Atlantic salmon caught in the Figgjo River, southwest Norway. Journal of Applied Ichthyology 30: 777–782. https://doi.org/10.1111/jai.12517.
[92] Robb, D. H. F., S. B. Wotton, J. L. McKinstry, N. K. Sørensen, S. C. Kestin, and N. K. Sørensen. 2000. Commercial slaughter methods used on Atlantic salmon: determination of the onset of brain failure by electroencephalography. Veterinary Record 147: 298–303. https://doi.org/10.1136/vr.147.11.298.
[93] Roth, Bjorn, Erik Slinde, and David H. F. Robb. 2007. Percussive stunning of Atlantic salmon (Salmo salar) and the relation between force and stunning. Aquacultural Engineering 36: 192–197. https://doi.org/10.1016/j.aquaeng.2006.11.001.
[94] European Food Safety Authority (EFSA). 2009. Species-specific welfare aspects of the main systems of stunning and killing of farmed Atlantic Salmon. EFSA Journal 7: 1011. https://doi.org/10.2903/j.efsa.2009.1011.
[95] Lambooij, E., E. Grimsbø, J. W. van de Vis, H. G. M. Reimert, R. Nortvedt, and B. Roth. 2010. Percussion and electrical stunning of Atlantic salmon (Salmo salar) after dewatering and subsequent effect on brain and heart activities. Aquaculture 300: 107–112. https://doi.org/10.1016/j.aquaculture.2009.12.022.
[96] Teletchea, Fabrice, and Pascal Fontaine. 2012. Levels of domestication in fish: implications for the sustainable future of aquaculture. Fish and Fisheries 15: 181–195. https://doi.org/10.1111/faf.12006.
[97] Teletchea, Fabrice. 2015. Domestication of Marine Fish Species: Update and Perspectives. Journal of Marine Science and Engineering 3: 1227–1243. https://doi.org/10.3390/jmse3041227.
[98] Hendry, K, and D Cragg-Hine. 2003. Ecology of the Atlantic Salmon - IN106. 7. Conserving Natura 2000 Rivers Ecology.
[99] Amundsen, Per-Arne, Heidi-Marie Gabler, and Lars Sigvald Riise. 2001. Intraspecific food resource partitioning in Atlantic salmon ( Salmo salar) parr in a subarctic river. Aquatic Living Resources 14: 257–265. https://doi.org/10.1016/S0990-7440(01)01127-5.
[100] Orlov, Alexander V., Yuri V. Gerasimov, and Oleg M. Lapshin. 2006. The feeding behaviour of cultured and wild Atlantic salmon, Salmo salar L., in the Louvenga River, Kola Peninsula, Russia. ICES J. Mar. Sci. 63: 1297–1303. https://doi.org/10.1016/j.icesjms.2006.05.004.
[101] Carter, Chris. 1993. Fish Meal Replacement In Aquaculture Feeds For Atlantic Salmon. 93/120–05. University of Tasmania.
[102] Bell, J. Gordon, John McEvoy, Douglas R. Tocher, Fiona McGhee, Patrick J. Campbell, and John R. Sargent. 2001. Replacement of Fish Oil with Rapeseed Oil in Diets of Atlantic Salmon (Salmo salar) Affects Tissue Lipid Compositions and Hepatocyte Fatty Acid Metabolism. The Journal of Nutrition 131: 1535–1543.
[103] Torstensen, B. E., M. Espe, M. Sanden, I. Stubhaug, R. Waagbø, G. -I. Hemre, R. Fontanillas, et al. 2008. Novel production of Atlantic salmon (Salmo salar) protein based on combined replacement of fish meal and fish oil with plant meal and vegetable oil blends. Aquaculture 285: 193–200. https://doi.org/10.1016/j.aquaculture.2008.08.025.
[104] Espe, Marit, Adel El-Mowafi, and Kari Ruohonen. 2012. Replacement of Fishmeal with Plant Protein Ingredients in Diets to Atlantic Salmon (Salmo salar) – Effects on Weight Gain and Accretion. https://doi.org/10.5772/29975.
[105] Couture, Jessica L., Roland Geyer, Jon Øvrum Hansen, Brandon Kuczenski, Margareth Øverland, Joseph Palazzo, Christian Sahlmann, and Hunter Lenihan. 2019. Environmental Benefits of Novel Nonhuman Food Inputs to Salmon Feeds. Environmental Science Technology. World. https://doi.org/10.1021/acs.est.8b03832.