Professor Nicky Clayton, FRS

Nicky Clayton,

University position


Professor Nicky Clayton, FRS is pleased to consider applications from prospective PhD students.


Department of Psychology


Behavioural and Clinical Neuroscience Institute


Home page (personal home page)

Research Theme

Cognitive and Behavioural Neuroscience


Nicky studies the development and evolution of cognition in members of the crow family (including jackdaws, rooks and jays) and humans. Her work has challenged many of the common-held assumptions that only humans can plan for the future and reminisce about the past, and that only humans can understand other minds as well as other times. This work has led to a radical re-evaluation of animal cognition, and raises important issues about the evolution of cognition.

She is Scientist in Residence at Rambert working with Artistic Direct Mark Baldwin on choreographic works inspired by science e.g. Comedy of Change.

Nicky's most recent collaboration is with artist and write, Clive Wilkins,

who is Artist in Residence in the Psychology department. It arose out of their mutual interest in mental time trave. They also regularly dance tango together.

( ) You can watch there TEDx talk

Conversations without words here (

Research Focus


evolution and development of cognition

episodic memory

future planning

theory of mind


Clinical conditions

Links to amnesia


Behavioural analysis

Video and camera equipment



Anthony Dickinson

Rufous Johnstone

Andrea Manica

Jim Russell

United Kingdom

Rambert Dance Company Web:

Nathan Emery Web:

Murray Shanahan Web:


Cristina Atance Web:

Josep Call Web:

Barney Schlinger Web:

Associated News Items

Key publications

Raby CR, Alexis DM, Dickinson A, Clayton NS (2007), “Planning for the future by Western Scrub-Jays” Nature 445:919-921 Details

Dally JM, Emery NJ, Clayton NS (2006), “Food-caching western scrub-jays keep track of who was watching when” Science 312(5780):1662-1665 Details

Emery NJ, Clayton NS (2004), “The mentality of crows. Convergent evolution of intelligence in corvids and apes” Science 306:1903-1907 Details



Davidson GL, Clayton NS, Thornton A (2014), “Salient eyes deter conspecific nest intruders in wild jackdaws (Corvus monedula).” Biol Lett 10(2):20131077 Details

Jelbert SA, Taylor AH, Cheke LG, Clayton NS, Gray RD (2014), “Using the Aesop's fable paradigm to investigate causal understanding of water displacement by New Caledonian crows.” PLoS One 9(3):e92895 Details

Legg EW, Clayton NS (2014), “Eurasian jays (Garrulus glandarius) conceal caches from onlookers.” Anim Cogn Details

MacLean EL, Hare B, Nunn CL, Addessi E, Amici F, Anderson RC, Aureli F, Baker JM, Bania AE, Barnard AM, Boogert NJ, Brannon EM, Bray EE, Bray J, Brent LJ, Burkart JM, Call J, Cantlon JF, Cheke LG, Clayton NS, Delgado MM, DiVincenti LJ, Fujita K, Herrmann E, Hiramatsu C, Jacobs LF, Jordan KE, Laude JR, Leimgruber KL, Messer EJ, Moura AC, Ostoji? L, Picard A, Platt ML, Plotnik JM, Range F, Reader SM, Reddy RB, Sandel AA, Santos LR, Schumann K, Seed AM, Sewall KB, Shaw RC, Slocombe KE, Su Y, Takimoto A, Tan J, Tao R, van Schaik CP, Virányi Z, Visalberghi E, Wade JC, Watanabe A, Widness J, Young JK, Zentall TR, Zhao Y (2014), “The evolution of self-control.” Proc Natl Acad Sci U S A 111(20):E2140-8 Details

Ostoji? L, Clayton NS (2014), “Behavioural coordination of dogs in a cooperative problem-solving task with a conspecific and a human partner.” Anim Cogn 17(2):445-59 Details

Ostoji? L, Legg EW, Shaw RC, Cheke LG, Mendl M, Clayton NS (2014), “Can male Eurasian jays disengage from their own current desire to feed the female what she wants?” Biol Lett 10(3):20140042 Details

Shaw RC, Clayton NS (2014), “Pilfering Eurasian jays use visual and acoustic information to locate caches.” Anim Cogn Details

Taylor AH, Cheke LG, Waismeyer A, Meltzoff AN, Miller R, Gopnik A, Clayton NS, Gray RD (2014), “Of babies and birds: complex tool behaviours are not sufficient for the evolution of the ability to create a novel causal intervention.” Proc Biol Sci 281(1787) Details


Cheke LG, Clayton NS (2013), “Do different tests of episodic memory produce consistent results in human adults?” Learn Mem 20(9):491-8 Details

Jozet-Alves C, Bertin M, Clayton NS (2013), “Evidence of episodic-like memory in cuttlefish.” Curr Biol 23(23):R1033-5 Details

Ostoji? L, Clayton NS (2013), “Inequity aversion in human adults: testing behavioural criteria from comparative cognition.” Anim Cogn 16(5):765-72 Details

Ostoji? L, Shaw RC, Cheke LG, Clayton NS (2013), “Evidence suggesting that desire-state attribution may govern food sharing in Eurasian jays.” Proc Natl Acad Sci U S A 110(10):4123-8 Details

Plotnik JM, Pokorny JJ, Keratimanochaya T, Webb C, Beronja HF, Hennessy A, Hill J, Hill VJ, Kiss R, Maguire C, Melville BL, Morrison VM, Seecoomar D, Singer B, Ukehaxhaj J, Vlahakis SK, Ylli D, Clayton NS, Roberts J, Fure EL, Duchatelier AP, Getz D (2013), “Visual cues given by humans are not sufficient for Asian elephants (Elephas maximus) to find hidden food.” PLoS One 8(4):e61174 Details

Shaw RC, Clayton NS (2013), “Careful cachers and prying pilferers: Eurasian jays (Garrulus glandarius) limit auditory information available to competitors.” Proc Biol Sci 280(1752):20122238 Details

Shaw RC, Plotnik JM, Clayton NS (2013), “Exclusion in corvids: the performance of food-caching Eurasian jays (Garrulus glandarius).” J Comp Psychol 127(4):428-35 Details

Thom JM, Clayton NS (2013), “No evidence of temporal preferences in caching by Western scrub-jays (Aphelocoma californica).” Behav Processes Details

Thom JM, Clayton NS (2013), “Re-caching by Western scrub-jays (Aphelocoma californica) cannot be attributed to stress.” PLoS One 8(1):e52936 Details

Watanabe A, Grodzinski U, Clayton NS (2013), “Western scrub-jays allocate longer observation time to more valuable information.” Anim Cogn Details

Ziauddeen H, Chamberlain SR, Nathan PJ, Koch A, Maltby K, Bush M, Tao WX, Napolitano A, Skeggs AL, Brooke AC, Cheke L, Clayton NS, Sadaf Farooqi I, O'Rahilly S, Waterworth D, Song K, Hosking L, Richards DB, Fletcher PC, Bullmore ET (2013), “Effects of the mu-opioid receptor antagonist GSK1521498 on hedonic and consummatory eating behaviour: a proof of mechanism study in binge-eating obese subjects.” Mol Psychiatry 18(12):1287-93 Details


Alpern S, Fokkink R, Lidbetter T, Clayton NS (2012), “A search game model of the scatter hoarder's problem.” J R Soc Interface 9(70):869-79 Details

Cheke LG, Clayton NS (2012), “Eurasian jays (Garrulus glandarius) overcome their current desires to anticipate two distinct future needs and plan for them appropriately.” Biol Lett 8(2):171-5 Details

Cheke LG, Loissel E, Clayton NS (2012), “How do children solve Aesop's Fable?” PLoS One 7(7):e40574 Details

Millan MJ, Agid Y, Brüne M, Bullmore ET, Carter CS, Clayton NS, Connor R, Davis S, Deakin B, DeRubeis RJ, Dubois B, Geyer MA, Goodwin GM, Gorwood P, Jay TM, Joëls M, Mansuy IM, Meyer-Lindenberg A, Murphy D, Rolls E, Saletu B, Spedding M, Sweeney J, Whittington M, Young LJ (2012), “Cognitive dysfunction in psychiatric disorders: characteristics, causes and the quest for improved therapy.” Nat Rev Drug Discov 11(2):141-68 Details

O'Donnell S, Logan CJ, Clayton NS (2012), “Specializations of birds that attend army ant raids: an ecological approach to cognitive and behavioral studies.” Behav Processes 91(3):267-74 Details

Taylor AH, Clayton NS (2012), “Evidence from convergent evolution and causal reasoning suggests that conclusions on human uniqueness may be premature.” Behav Brain Sci 35(4):241-2 Details

Thornton A, Clayton NS, Grodzinski U (2012), “Animal minds: from computation to evolution.” Philos Trans R Soc Lond B Biol Sci 367(1603):2670-6 Details


Cheke LG, Bird CD, Clayton NS (2011), “Tool-use and instrumental learning in the Eurasian jay (Garrulus glandarius).” Anim Cogn 14(3):441-55 Details

Taylor AH, Elliffe DM, Hunt GR, Emery NJ, Clayton NS, Gray RD (2011), “New Caledonian crows learn the functional properties of novel tool types.” PLoS One 6(12):e26887 Details


Grodzinski U, Clayton NS (2010), “Problems faced by food-caching corvids and the evolution of cognitive solutions.” Philos Trans R Soc Lond B Biol Sci 365(1542):977-87 Details

Haun DB, Jordan FM, Vallortigara G, Clayton NS (2010), “Origins of spatial, temporal and numerical cognition: Insights from comparative psychology.” Trends Cogn Sci 14(12):552-60 Details

Osvath M, Raby CR, Clayton NS (2010), “What should be compared in comparative mental time travel?” Trends Cogn Sci 14(2):51-2; author reply 52-3 Details

Salwiczek LH, Watanabe A, Clayton NS (2010), “Ten years of research into avian models of episodic-like memory and its implications for developmental and comparative cognition.” Behav Brain Res 215(2):221-34 Details

Teufel C, Alexis DM, Clayton NS, Davis G (2010), “Mental-state attribution drives rapid, reflexive gaze following.” Atten Percept Psychophys 72(3):695-705 Details


Clayton NS, Russell J (2009), “Looking for episodic memory in animals and young children: prospects for a new minimalism.” Neuropsychologia 47(11):2330-40 Details

Emery NJ, Clayton NS (2009), “Tool use and physical cognition in birds and mammals.” Curr Opin Neurobiol 19(1):27-33 Details

Emery NJ, Clayton NS (2009), “Comparative social cognition.” Annu Rev Psychol 60:87-113 Details

Raby CR, Clayton NS (2009), “Prospective cognition in animals.” Behav Processes 80(3):314-24 Details

Salwiczek LH, Emery NJ, Schlinger B, Clayton NS (2009), “The development of caching and object permanence in Western scrub-jays (Aphelocoma californica): which emerges first?” J Comp Psychol 123(3):295-303 Details

Seed AM, Call J, Emery NJ, Clayton NS (2009), “Chimpanzees solve the trap problem when the confound of tool-use is removed.” J Exp Psychol Anim Behav Process 35(1):23-34 Details

Stulp G, Emery NJ, Verhulst S, Clayton NS (2009), “Western scrub-jays conceal auditory information when competitors can hear but cannot see.” Biol Lett 5(5):583-5 Details

Teufel C, Alexis DM, Todd H, Lawrance-Owen AJ, Clayton NS, Davis G (2009), “Social cognition modulates the sensory coding of observed gaze direction.” Curr Biol 19(15):1274-7 Details


Sanford K, Clayton NS (2008), “Motivation and memory in zebra finch (Taeniopygia guttata) foraging behavior.” Anim Cogn 11(2):189-98 Details

Seed AM, Clayton NS, Emery NJ (2008), “Cooperative problem solving in rooks (Corvus frugilegus).” Proc Biol Sci 275(1641):1421-9 Details

Vallortigara G, Snyder A, Kaplan G, Bateson P, Clayton NS, Rogers LJ (2008), “Are animals autistic savants.” PLoS Biol 6(2):e42 Details


Clayton NS (2007), “Nicola S. Clayton.” Curr Biol 17(10):R345-6 Details

Clayton NS, Dally JM, Emery NJ (2007), “Social cognition by food-caching corvids. The western scrub-jay as a natural psychologist.” Philos Trans R Soc Lond B Biol Sci 362(1480):507-22 Details

Clayton NS, Emery NJ (2007), “The social life of corvids.” Curr Biol 17(16):R652-6 Details

Clayton NS, Salwiczek LH, Dickinson A (2007), “Episodic memory.” Curr Biol 17(6):R189-91 Details

Correia SP, Dickinson A, Clayton NS (2007), “Western scrub-jays anticipate future needs independently of their current motivational state.” Curr Biol 17(10):856-61 Details

de Kort SR, Correia SP, Alexis DM, Dickinson A, Clayton NS (2007), “The control of food-caching behavior by Western scrub-jays (Aphelocoma californica).” J Exp Psychol Anim Behav Process 33(4):361-70 Details

Emery NJ, Seed AM, von Bayern AM, Clayton NS (2007), “Cognitive adaptations of social bonding in birds.” Philos Trans R Soc Lond B Biol Sci 362(1480):489-505 Details

Seed AM, Clayton NS, Emery NJ (2007), “Postconflict third-party affiliation in rooks, Corvus frugilegus.” Curr Biol 17(2):152-8 Details

Tebbich S, Seed AM, Emery NJ, Clayton NS (2007), “Non-tool-using rooks, Corvus frugilegus, solve the trap-tube problem.” Anim Cogn 10(2):225-31 Details


de Kort SR, Clayton NS (2006), “An evolutionary perspective on caching by corvids.” Proc Biol Sci 273(1585):417-23 Details

Helme AE, Call J, Clayton NS, Emery NJ (2006), “What do bonobos (Pan paniscus) understand about physical contact?” J Comp Psychol 120(3):294-302 Details

Helme AE, Clayton NS, Emery NJ (2006), “What do rooks (Corvus frugilegus) understand about physical contact?” J Comp Psychol 120(3):288-93 Details

Seed AM, Tebbich S, Emery NJ, Clayton NS (2006), “Investigating physical cognition in rooks, Corvus frugilegus.” Curr Biol 16(7):697-701 Details


Clayton NS, Dally J, Gilbert J, Dickinson A (2005), “Food caching by western scrub-jays (Aphelocoma californica) is sensitive to the conditions at recovery.” J Exp Psychol Anim Behav Process 31(2):115-24 Details

Clayton NS, Hen R (2005), “Neural circuits and behaviour: developmental and evolutionary perspectives.” Curr Opin Neurobiol 15(6):683-5 Details

Emery NJ, Clayton NS (2005), “Evolution of the avian brain and intelligence.” Curr Biol 15(23):R946-50 Details

Healy SD, de Kort SR, Clayton NS (2005), “The hippocampus, spatial memory and food hoarding: a puzzle revisited.” Trends Ecol Evol 20(1):17-22 Details


Dally JM, Emery NJ, Clayton NS (2004), “Cache protection strategies by western scrub-jays (Aphelocoma californica): hiding food in the shade.” Proc Biol Sci 271 Suppl 6:S387-90 Details

Emery NJ, Dally JM, Clayton NS (2004), “Western scrub-jays ( Aphelocoma californica) use cognitive strategies to protect their caches from thieving conspecifics.” Anim Cogn 7(1):37-43 Details

Lucas JR, Brodin A, de Kort SR, Clayton NS (2004), “Does hippocampal size correlate with the degree of caching specialization?” Proc Biol Sci 271(1556):2423-9 Details

Oberlander JG, Schlinger BA, Clayton NS, Saldanha CJ (2004), “Neural aromatization accelerates the acquisition of spatial memory via an influence on the songbird hippocampus.” Horm Behav 45(4):250-8 Details

Pravosudov VV, Kitaysky AS, Wingfield JC, Clayton NS (2004), “No latitudinal differences in adrenocortical stress response in wintering black-capped chickadees (Poecile atricapilla).” Comp Biochem Physiol A Mol Integr Physiol 137(1):95-103 Details


Clayton NS, Bussey TJ, Dickinson A (2003), “Can animals recall the past and plan for the future?” Nat Rev Neurosci 4(8):685-91 Details

Clayton NS, Bussey TJ, Emery NJ, Dickinson A (2003), “Prometheus to Proust: the case for behavioural criteria for 'mental time travel'.” Trends Cogn Sci 7(10):436-7; author reply 437-8 Details

Clayton NS, Yu KS, Dickinson A (2003), “Interacting Cache memories: evidence for flexible memory use by Western Scrub-Jays (Aphelocoma californica).” J Exp Psychol Anim Behav Process 29(1):14-22 Details

de Kort SR, Emery NJ, Clayton NS (2003), “Food offering in jackdaws ( Corvus monedula).” Naturwissenschaften 90(5):238-40 Details

Pravosudov VV, Mendoza SP, Clayton NS (2003), “The relationship between dominance, corticosterone, memory, and food caching in mountain chickadees (Poecile gambeli).” Horm Behav 44(2):93-102 Details


Dwyer DM, Clayton NS (2002), “A reply to the defenders of the faith.” Trends Cogn Sci 6(3):109-111 Details

Pravosudov VV, Clayton NS (2002), “A test of the adaptive specialization hypothesis: population differences in caching, memory, and the hippocampus in black-capped chickadees (Poecile atricapilla).” Behav Neurosci 116(4):515-22 Details

Pravosudov VV, Kitaysky AS, Saldanha CJ, Wingfield JC, Clayton NS (2002), “The effect of photoperiod on adrenocortical stress response in mountain chickadees (Poecile gambeli).” Gen Comp Endocrinol 126(2):242-8 Details

Pravosudov VV, Lavenex P, Clayton NS (2002), “Changes in spatial memory mediated by experimental variation in food supply do not affect hippocampal anatomy in mountain chickadees (Poecile gambeli).” J Neurobiol 51(2):142-8 Details


Clayton NS (2001), “Hippocampal growth and maintenance depend on food-caching experience in juvenile mountain chickadees (Poecile gambeli).” Behav Neurosci 115(3):614-25 Details

Clayton NS, Griffiths DP, Emery NJ, Dickinson A (2001), “Elements of episodic-like memory in animals.” Philos Trans R Soc Lond B Biol Sci 356(1413):1483-91 Details

Clayton NS, Yu KS, Dickinson A (2001), “Scrub jays (Aphelocoma coerulescens) form integrated memories of the multiple features of caching episodes.” J Exp Psychol Anim Behav Process 27(1):17-29 Details

Emery NJ, Clayton NS (2001), “Effects of experience and social context on prospective caching strategies by scrub jays.” Nature 414(6862):443-6 Details

Griffiths DP, Clayton NS (2001), “Testing episodic memory in animals: a new approach.” Physiol Behav 73(5):755-62 Details

Lee DW, Smith GT, Tramontin AD, Soma KK, Brenowitz EA, Clayton NS (2001), “Hippocampal volume does not change seasonally in a non food-storing songbird.” Neuroreport 12(9):1925-8 Details

Pravosudov VV, Clayton NS (2001), “Effects of demanding foraging conditions on cache retrival accuracy in food-caching mountain chickadees (Poecile gambeli).” Proc Biol Sci 268(1465):363-8 Details

Pravosudov VV, Kitaysky AS, Wingfield JC, Clayton NS (2001), “Long-term unpredictable foraging conditions and physiological stress response in mountain chickadees (Poecile gambeli).” Gen Comp Endocrinol 123(3):324-31 Details


Saldanha CJ, Schlinger BA, Clayton NS (2000), “Rapid effects of corticosterone on cache recovery in mountain chickadees (Parus gambeli).” Horm Behav 37(2):109-15 Details

Suzuki WA, Clayton NS (2000), “The hippocampus and memory: a comparative and ethological perspective.” Curr Opin Neurobiol 10(6):768-73 Details


Clayton NS, Dickinson A (1999), “Scrub jays (Aphelocoma coerulescens) remember the relative time of caching as well as the location and content of their caches.” J Comp Psychol 113(4):403-16 Details

Hauber ME, Clayton NS, Kacelnik A, Reboreda JC, DeVoogd TJ (1999), “Sexual dimorphism and species differences in HVC volumes of cowbirds.” Behav Neurosci 113(5):1095-9 Details