UNIVERSITY OF BERGEN

Department of
Biological Sciences



How to be a salmon?

This is a depository of material connected to the workshop "What is is like to be an Atlantic salmon?". It is written and maintained by Jarl Giske.



Here is a quite recent version of the ms.


Videos and PDFs of presentations

Unfortunately, the zoom recording of the first day ended abruptly during Knut's talk. The later talks by Magda, Marc and Andy were therefore not recorded. The whole second day is recorded. Sergey has split the whole-day video files into separate files for each talk.


Jarl Giske: To be an Atlantic salmon video PDF
Katja Enberg: The UN perspective video PDF
Knut Wiik Vollset: What senses are used to make life changing decisions video PDF
Magda Dumitru: What is it to be an animal? PDF
Marc Mangel: What Is it like to be (an Atlantic) salmon? PDF
Andy Higginson: A behavioural ecology perspective on what it is to be a salmon. PDF
Steve Railsback: To be a salmon in a river video PDF
Anders Frugård Opdal: How salmon (and its brain) is changed by experiences video PDF
Tore S Kristiansen: The salmon's quality of life .. and the predictive brain video PDF
Ivar Rønnestad: To feed or not to feed - that is my question video PDF
Sergey Budaev: The salmon machine video PDF


Papers mentioned

Nagel 1974: What is it like to be a bat?


Jarl's talk:
Anderson 1972: More is different.
Bubic & al 2010: Prediction, cognition and the brain.
Budaev & al 2018: AHA: A general cognitive architecture for Darwinian agents.
Budaev & al 2019: Decision-making from the animal perspective: Bridging ecology and subjective cognition.
Budaev & al 2020: Computational animal welfare: towards cognitive architecture models of animal sentience, emotion and wellbeing.
Clark 2013: Whatever next? Predictive brains, situated agents, and the future of cognitive science.
FitzGerald & al 2014: Model averaging, optimal inference, and habit formation.
Giske & al 1998: Modelling spatial dynamics of fish.
Grafen 1984: Natural selection, kin selection and group selection.
Hoffmeyer 2008. We do not have this as pdf, but a related text from 2002: Reading Hoffmeyer, rethinking biology.
Jensen & al 2021: Hormonal adjustments to future expectations impact growth and survival in juvenile fish.
Nesse 2013: Tinbergen’s four questions, organized: a response to Bateson and Laland.
Soylu 2016: An embodied approach to understanding: making sense of the world through simulated bodily activity.
Tinbergen 1963: On aims and methods of ethology.
Weidner & al 2020: Hormones as adaptive control systems in juvenile fish.


Magda's talk:
Bshary & Triki 2022: Fish ecology and cognition: insights from studies on wild and wild-caught teleost fishes.
Cleeremans 2011: The radical plasticity thesis: how the brain learns to be conscious.
Cleeremans 2022: Consciousness matters: phenomenal experience has functional value.
Mather & al 2016: Norepinephrine ignites local hotspots of neuronal excitation: How arousal amplifies selectivity in perception and memory.


Marc's talk:
Clark & Mangel 1988: Dynamic Modeling in Behavioral Ecology.
Dobzhansky 1973: Nothing in biology makes sense except in the light of evolution.
Fleming 1998: Pattern and variability in the breeding system of Atlantic salmon (Salmo salar), with comparisons to other salmonids.
Gerhart & Kirschner 1997: Cells, Embryos and Evolution.
Mangel & Clark 2000: Dynamic State Variable Models in Ecology.
Mangel 1994: Climate change and salmonid life history variation.
Nagel 1974: What is it like to be a bat?
Rikardsen & al 2004: Modelling the life-history variation of Arctic charr.
Satterthwaite & al 2009: Steelhead life history on California's Central Coast: Insights from a state-dependent model.
Stearley & Smith 1993: Phylogeny of the Pacific trouts and salmons (Oncorhynchus) and genera of the family Salmonidae.
Thorpe & al 1998: Modelling the proximate basis of salmonid life-history variation, with application to Atlantic salmon, Salmo salar L..


Anders's talk:
Birse 1980: Neuronal increase in various areas of the nervous system of the nervous system of the guppy, Lebistes.
Braithwaite VA & AGV Salvanes 2005. Environmental variability in the early rearing environment generates behaviourally flexible cod: implications for rehabilitating wild populations. Proc R Soc B 272:1107–1113
Braithwaite et al 1996: Can juvenile Atlantic salmon use multiple cue systems in spatial learning?
Brandão et al 2015: Isolation impairs cognition in a social fish.
Bredy & al 2003: Partial reversal of the effect of maternal care on cognitive function through environmental enrichment.
Bshary & al 2014: Social cognition in fishes.
Burns JG & al 2009. Rearing environment affects the brain size of guppies: Lab-reared guppies have smaller brains than wild-caught guppies. Ethology 115:122–133
Dunlap KD 2016. Fish neurogenesis in context: Assessing environmental influences on brain plasticity within a highly labile physiology and morphology. Brain Behav Evol 87:156–166
Ebbesson and Braithwaite 2012: Environmental effects on fish neural plasticity and cognition.
Gonda et al 2009: Habitat dependent and independent plastic responses to social environment in the nine-spined stickleback (Pungitius pungitius) brain.
Gonda A & al 2010: Predation and competition mediated brain plasticity in Rana temporaria tadpoles.
Gonda A & al 2012: Brain development and predation: plastic responses depend on evolutionary history. Biol Lett 8:249-252
Guo & al 2022: Social isolation affects intra specific interaction behaviour and reduces the size of the cerebellar brain region in juvenile Atlantic salmon Salmo salar.
Herzeg & al 2015: Experimental evidence for sex specific plasticity in adult brain.
Joyce and Brown 2019: Rapid plastic changes in brain morphology in response to acute changes in predation pressure in juvenile Atlantic salmon (Salmo salar) and northern redbelly dace (Phoxinus eos).
Kihslinger RL & Nevitt 2006: Early rearing environment impacts cerebellar growth in juvenile salmon.
Kihslinger RL & al 2006. Environmental rearing conditions produce forebrain differences in wild Chinook salmon Oncorhynchus tshawytscha. Comp Biochem Physiol A 145:145–151
Knudsen 2004: Sensitive periods in the development of the brain and behaviour.
Kotrschal 2012: Inside the heads of David and Goliath: environmental effects on brain morphology among wild and growth-enhanced coho salmon Oncorhynchus kisutch.
Kotrschal and Taborsky 2010: Environmental change enhances cognitive abilities in fish.
Kotrschal et al 1998: Fish brains: evolution and environmental relationships.
Kruska 2005: On the evolutionary significance of encephalization in some eutherian mammals: Effects of adaptive radiation, domestication, and feralization.
Larsen et al 2014: Reduced rearing density increases postrelease migration success of Atlantic salmon (Salmo salar) smolts.
Lema et al 2005: Proliferation zones in the salmon telencephalon and evidence for environmental influence on proliferation rate.
Marchetti and Nevitt 2003: Effects of hatchery rearing on brain structures of rainbow trout, Oncorhynchus mykiss.
Maruska et al 2013: Social descent with territory loss causes rapid behavioral endocrine and transcriptional changes in the brain.
Mayer et el 2011: Domestication causes rapid changes in heart and brain morphology in Atlantic cod (Gadus morhua).
Mes D & al 2018. Neurobiology of wild and hatchery-reared Atlantic salmon: How nurture drives neuroplasticity.
Mes D & al 2019. Effects of environmental enrichment on forebrain neural plasticity and survival success of stocked Atlantic salmon. J Exp Biol 222:jeb212258
Näslund J & al 2012: Early enrichment effects on brain development in hatchery reared salmon Salmo salar: No evidence for a critical period.
Näslund J & al 2017: Environment dependent plasticity and ontogenetic changes in the brain of hatchery-reared Atlantic salmon.
Näslund J & al 2019: Fish density, but not environmental enrichment, affects the size of cerebellum in the brain of juvenile hatchery-reared Atlantic salmon. Environ Biol Fish 102:705–712
Park and Bell 2010: Variation of telencephalon morphology of the threespine stickleback (Gasterosteus aculeatus) in relation to inferred ecology.
Salas et al 2003: Evolution of forebrain and spatial cognition in vertebrates: Conservation across diversity.
Salas et al 2006: Neuropsychology of learning and memory in teleost fish.
Salvanes AGV & al 2007. Effects of early experience on group behaviour in fish.
Salvanes AGV & al 2013. Environmental enrichment promotes neural plasticity and cognitive ability in fish. Proc R Soc B 280:20131331
Schmidt-Hieber et al 2004: Enhanced synaptic plasticity in newly generated granule cells in the adult hippocampus.
Sørensen et al 2007: Social regulation of neurogenesis in teleosts.
von Krogh K & al 2010. Forebrain cell proliferation behaviour and physiology of zebrafish kept in enriched or barren environments. Physiol Behav 101:32–39
Zupanc and Sirbulenscu 2011: Adult neurogenesis and neuronal regeneration in the central nervous system of teleost fish.
Zupanc GHK 2006: Neurogenesis and neuronal regeneration in the adult fish brain. J Comp Physiol A 192:649–670


Tore's talk:
Conant & Ashby 1970: Every good regulator of a system must be a model of that system.
Korte & al 2007: A new animal welfare concept based on allostasis.
Perathoner & al 2016: Potential of zebrafish as a model for exploring the role of the amygdala in emotional memory and motivational behavior.
Sterling 2012: Allostasis: a model of predictive regulation.
Williams & al 2013: The function of fin rays as proprioceptive sensors in fish.


Sergey's talk:
Bossaerts & Murawski 2017: Computational complexity and human decision-making.
Bowers & Davis 2012: Bayesian just-so stories in psychology and neuroscience.
Bubic & al 2010: Prediction, cognition and the brain.
Budaev & al 2020: Computational animal welfare: towards cognitive architecture models of animal sentience, emotion and wellbeing.
Føre & al 2018: Precision fish farming: A new framework to improve production in aquaculture.
Giske & al 2013: Effects of the emotion system on adaptive behavior.
Hesslow 2016: The current status of the simulation theory of cognition.
Higginson & al 2018: Trust your gut: using physiological states as a source of information is almost as effective as optimal Bayesian learning.
Soylu 2016: An embodied approach to understanding: making sense of the world through simulated bodily activity.
Trotsos 1993: The role of computational complexity in perceptual theory.


Other papers:
Adams RA & al 2013. Predictions not commands: Active inference in the motor system. Brain Struct Funct 218:611–643
Anderson DJ & R Adolphs 2014. A framework for studying emotions across species. Cell 157:187-200
Arias Del Angel JA & al 2020. Interplay of mesoscale physics and agent‑like behaviors in the parallel evolution of aggregative multicellularity. EvoDevo 11:21
Baars BJ 1988. A Cognitive Theory of Consciousness. Cambridge UP
Bauer S & al 2011. Cues and decision rules in animal migration. Chapter 6 (pp. 68-87) in Animal Migration – a synthesis, EJ Milner-Gulland & al (eds). Oxford UP
Brown C 2015. Fish intelligence, sentience and ethics. Anim Cogn 18:1–17
Bubic A & al 2010. Prediction, cognition and the brain. Front Human Neurosci 4:1-15
Budaev S & al 2018. AHA: a general cognitive architecture for Darwinian agents. Biol Inspired Cogn Archit 25:51–57
Budaev S & al 2019. Decision-making from the animal perspective: Bridging ecology and subjective cognition. Front Ecol Evol 7:164
Chen C & N Crilly 2014. Modularity, redundancy and degeneracy: Cross-domain perspectives on key design principles. 2014 IEEE International Systems Conference Proceedings 546-553
Clark A 2013. Whatever next? Predictive brains, situated agents, and the future of cognitive science. Behav Brain Sci 36:181-204
Clark A 2017. How to knit your own Markov blanket: Resisting the Second Law with meta- morphic minds. In: Metzinger T, Wiese W (eds), Philosophy and Predictive Processing: 3. MIND Group
Clune J & al 2013. The evolutionary origins of modularity. Proc R Soc B 280:20122863
Conant RC & WR Ashby 1970. Every good regulator of a system must be a model of that system. Int J Systems Sci 1(2):89-97
Crump A & al 2020. Emotion in animal contests. Proc R Soc B 287:20201715
Dehaene S 2014. Consciousness and the Brain: Deciphering How the Brain Codes Our Thoughts. Penguin Random House
Dehaene S & al 2006. Conscious, preconscious, and subliminal processing: a testable taxonomy. Trends Cogn Sci 10:204-211
Del Giudice M 2023. A general motivational architecture for human and animal personality. Neurosci Biobehav Rev 144:104967
Denton DA & al 2009. The role of primordial emotions in the evolutionary origin of consciousness. Consciousness and Cognition 18:500–514
Ebbesson LOE & al 2007. Exposure to continuous light disrupts retinal innervation of the preoptic nucleus during parr-smolt transformation in Atlantic salmon. Aquaculture 273:345–349
Edelman GM & al 2011. Biology of consciousness. Front Psychol 2:4
Fields C & al 2021. Minimal physicalism as a scale-free substrate for cognition and consciousness. Neurosci Cons 7:niab013
Fjelldal PG & al 2011. Continuous light and elevated temperature can trigger maturation both during and immediately after smoltification in male Atlantic salmon (Salmo salar). Aquaculture 321:93-100
Fjelldal PG & al 2018. Sexual maturation and smoltification in domesticated Atlantic salmon (Salmo salar L.) – is there a developmental conflict? Physiol Rep 6:1-18
Fleming IA 1988. Pattern and variability in the breeding system of Atlantic salmon (Salmo salar), with comparisons to other salmonids. Can J Fish Aquat Sci 55(S1):59-76
Friston K 2019. A free-energy principle for a particular physics. Preprint arxiv:1906.10184
Friston KJ 2005. A theory of cortical responses. Philos Trans R Soc B 360:815–836
Friston KJ 2010. The free-energy principle: a unified brain theory? Nat Rev Neurosci 11:127–138
Friston KJ 2013. Life as we know it. J R Soc Interface 10:20130475
Friston KJ & al 2006. A free energy principle for the brain. J Physiol Paris 100:70–87
Friston KJ & al 2017. Deep temporal models and active inference. Neurosci Biobehav Rev 77:388–402
Friston KJ & KE Stephan 2007. Free-energy and the brain. Synthese 159:417–458
Gallup GG & JR Anderson 2020. Self-recognition in animals: where do we stand 50 years later: Lessons from cleaner wrasse and other species. Psychol Conscious 7:46–58
Giske J & al 2013. Effects of the emotion system on adaptive behavior. Am Nat 182:689-703
Giske J & al 2014. The emotion system promotes diversity and evolvability. Proc R Soc B 281:20141096
Glimcher PW 2003. Decisions, uncertainty, and the brain. The science of neuroeconomics. MIT Press
Godfrey-Smith P 2016. Animal evolution and the origins of experience. In How Biology Shapes Philosophy: New Foundations for Naturalism. DL Smith (ed), Cambridge UP
Good C & J Davidson 2016. A review of factors influencing maturation of Atlantic Salmon, Salmo salar, with focus on water recirculation aquaculture system environments. J World Aquacult Soc 47:605-632
Gregory RL 1980. Perceptions as hypotheses. Phil Trans Royal Soc Lond B 290:181–197
Hesslow G 2012. The current status of the simulation theory of cognition. Brain Res 1428:71-79
Hutchinson JMC & G Gigerenzer 2005. Simple heuristics and rules of thumb: Where psychologists and behavioural biologists might meet. Behav Proc 69:97-124
Iversen M & al 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
Johnson D & al 2013. The evolution of error: Error management, cognitive constraints, and adaptive decision-making biases. TREE 28:474-481
Kauffman S 2007. Question 1: Origin of life and the living state. Orig Life Evol Biosph 37:315–322
Kim SS & al 2017. Ring attractor dynamics in the Drosophila central brain. Science 356:849-853
Knudsen EI 2007. Fundamental components of attention. Annu Rev Neurosci 30:57–78
Kohda M & al 2019. If a fish can pass the mark test, what are the implications for consciousness and self-awareness testing in animals? PLoS Biol 17:e3000021
Kohda M & al 2022. Further evidence for the capacity of mirror self-recognition in cleaner fish and the significance of ecologically relevant marks. PLoS Biol 20:e3001529
Kuchling F & al 2020. Morphogenesis as Bayesian inference: A variational approach to pattern formation and control in complex biological systems. Phys Life Rev 33:88–108
Kull K 2003. Ladder, tree, web: The ages of biological understanding. Sign Systems Studies 31.2
Kull K 2022. Choices by organisms: on the role of freedom in behaviour and evolution. Biol J Linn Soc XX:1–8
LeDoux J 1996. The Emotional Brain. Simon and Schuster
LeDoux J 2012. Rethinking the emotional brain. Neuron 73:653
LeDoux JE 2014. Coming to terms with fear. PNAS 111:2871–2878
Lennox RJ & al 2021. The quest for successful Atlantic salmon restoration: perspectives, priorities, and maxims. ICES J Mar Sci 78:3479–3497
Levin M 2020. Life, death, and self: fundamental questions of primitive cognition viewed through the lens of body plasticity and synthetic organisms. Biochem Biophys Res Comm 564:114–133
Lotka AJ 1925. Elements of Physical Biology. Williams & Wilkins Company
Mangel M & CW Clark 1986. Towards a unified foraging theory. Ecology 67:1127-1138
Mathis A & al 1996. Cultural transmission of predator recognition in fishes: intraspecific and interspecific learning. Anim Behav 51:185–201
Maynard Smith J & GR Price 1973. The logic of animal conflict. Nature 246:15-18
McCallum E 2019. Are fish self-aware? J Exp Biol 222:jeb192781
McNamara JM & AI Houston 1986. The common currency for behavioral decisions. Am Nat 127:358-378
Metcalfe NB & JE Thorpe 1992. Anorexia and defended energy levels in over-wintering juvenile salmon. J Anim Ecol 61:175–181
Mobley KB & al 2021. Maturation in Atlantic salmon (Salmo salar, Salmonidae): a synthesis of ecological, genetic, and molecular processes. Rev Fish Biol Fisher 31:523–571
Nesse RM 2013. Tinbergen’s four questions, organized: a response to Bateson and Laland. TREE 28: 681-682
Newman SA 2022. Inherency and agency in the origin and evolution of biological functions. Biol J Linnean Soc (in press)
Nilsen TO & al 2007. Differential expression of gill Na+, K+- ATPase alpha- and beta-subunits, Na+, K+, 2Cl(-) cotransporter and CFTR anion channel in juvenile anadromous and landlocked Atlantic salmon Salmo salar. J Exp Biol 210:2885–2896
Nisembaum LG & al 2020. Effects of a temperature rise on melatonin and thyroid hormones during smoltification of Atlantic salmon, Salmo salar. J Comp Physiol B 190:731–748
Pino Martinez E & al 2023. Interaction of temperature and feed ration on male postsmolt maturation of Atlantic salmon (Salmo salar L.). Aquaculture 562:738877
Pylyshyn Z 1984. Computation and Cognition: Toward a Foundation for Cognitive Science. MIT Press
Ramstead MJD & al 2018. Answering Schrödinger's question: A free-energy formulation. Phys Life Rev 24:1–16
Ramstead MJ & al 2019. Variational ecology and the physics of sentient systems. Phys Life Rev 31:188–205
Scholl BJ & PD Tremoulet 2000. Perceptual causality and animacy. Trends Cogn Sci 4:299–309
Schulkin J 2003. Rethinking homeostasis: allostatic regulation in physiology and pathophysiology. MIT Press
Schulz RW & al 2010. Spermatogenesis in fish. Gen Comp Endocrinol 165:390-411
Seth AK & M Tsakiris 2018. Being a Beast Machine. The Somatic Basis of Selfhood. Trends Cogn Sci 22:969-981
Soylu F 2016. An embodied approach to understanding: making sense of the world through simulated bodily activity. Front Psychol 7:1914
Stanley KO & J Lehman 2015. Why greatness cannot be planned: the myth of the objective. Springer
Stearley RF & GR Smith 1993. Phylogeny of the Pacific trouts and salmons (Oncorhynchus) and genera of the family Salmonidae. Trans Am Fish Soc 122:1-33
Stefansson SO & al 2007. Molecular mechanisms of continuous light inhibition of Atlantic salmon parr-smolt transformation. Aquaculture 273:235–245
Sterling P & S Laughlin 2015. Principles of Neural Design. MIT Press
Sterling P 2012. Allostasis: A model of predictive regulation. Physiology & Behavior 106:5–15
Svendsen E & al 2021. Heart rate and swimming activity as stress indicators for Atlantic salmon (Salmo salar). Aquaculture 531:735804
Tang H & al 2018. Recurrent computations for visual pattern completion. PNAS 115:8835-8840
Taranger GL & al 2010. Control of puberty in farmed fish. Gen Comp Endocrinol 165:483-515
Thorpe JE & al 1982. Bimodality of growth and smolting in Atlantic salmon, Salmo salar L. Aquaculture 28:123–132
Thorpe JE & al 1998. Modelling the proximate basis of salmonid life-history variation, with application to Atlantic salmon, Salmo salar L. Evol Ecol 12:581-599
Thorpe JE 1994. Reproductive strategies in Atlantic salmon, Salmo salar L. Aquac Fish Manag 25:77-87
Tinbergen N 1963. On aims and methods in ethology. Z Tierpsychol 20:410-433
Wagner GP 1996. Homologues, natural kinds and evolution of modularity. Am Zool 36:36–43
Wang XJ 2008. Decision making in recurrent neuronal circuits. Neuron 60:215-34
Whitacre JM 2010. Degeneracy: a link between evolvability, robustness and complexity in biological systems. Theor Biol Med Model 7:6
Whyte CJ & R Smith 2021. The predictive global neuronal workspace: A formal active inference model of visual consciousness. Prog Neurobiol 199:101918
Woodruff ML 2017. Consciousness in teleosts: There is something it feels like to be a fish. Animal Sentience 13:1

Webmaster:  Christian Jørgensen.