Professor John Mollon

University position


Professor John Mollon is pleased to consider applications from prospective PhD students.


Department of Psychology

Home page (personal home page)

Research Themes

Cognitive and Behavioural Neuroscience

Systems and Computational Neuroscience


My research interests are in the visual perception of colour, motion, form and depth; the genetic basis for individual variations in perception; the nature of the information carried by the cerebral bus; and the history of neuroscience and of colour theory.

A simulation of how a bookcase may look to a person who has the common form of colour blindness called deuteranopia
A simulation of how a bookcase may look to a person who has the common form of colour blindness called deuteranopia
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Research Focus






neural circuit

Clinical conditions


Genetic disorders



Web-based testing


United Kingdom

Gabriele Jordan Web:

Hannah Smithson Web:


Marina Danilova Web:

Associated News Items

Key publications

Smithson HE, Mollon JD (2006), “Do masks terminate the icon?” Quarterly Journal of Experimental Psychology 59:150-160

Bosten JM, Robinson JD, Jordan G, Mollon JD (2005), “Multidimensional scaling reveals a color dimension unique to 'color deficient' observers” Current Biology 15:R950-R952 Details

Danilova MV, Mollon JD (2003), “Comparison at a distance” Perception 32:395-414



Bosten JM, Bargary G, Goodbourn PT, Hogg RE, Lawrance-Owen AJ, Mollon JD (2014), “Individual differences provide psychophysical evidence for separate on- and off-pathways deriving from short-wave cones.” J Opt Soc Am A Opt Image Sci Vis 31(4):A47-54 Details

Bosten JM, Hogg RE, Bargary G, Goodbourn PT, Lawrance-Owen AJ, Mollon JD (2014), “Suggestive association with ocular phoria at chromosome 6p22.” Invest Ophthalmol Vis Sci 55(1):345-52 Details

Danilova MV, Mollon JD (2014), “Is discrimination enhanced at the boundaries of perceptual categories? A negative case.” Proc Biol Sci 281(1785):20140367 Details

Danilova MV, Mollon JD (2014), “Symmetries and asymmetries in chromatic discrimination.” J Opt Soc Am A Opt Image Sci Vis 31(4):A247-53 Details

Goodbourn PT, Bosten JM, Bargary G, Hogg RE, Lawrance-Owen AJ, Mollon JD (2014), “Variants in the 1q21 risk region are associated with a visual endophenotype of autism and schizophrenia.” Genes Brain Behav 13(2):144-51 Details

Lawrance-Owen AJ, Bosten JM, Hogg RE, Bargary G, Goodbourn PT, Mollon JD (2014), “Counterphase modulation flicker photometry: phenotypic and genotypic associations.” J Opt Soc Am A Opt Image Sci Vis 31(4):A226-31 Details

Le Sueur H, Mollon JD, Granzier J, Jordan G (2014), “Counterphase modulation photometry: comparison of two instruments.” J Opt Soc Am A Opt Image Sci Vis 31(4):A34-7 Details


Danilova MV, Chan CH, Mollon JD (2013), “Can spatial resolution reveal individual differences in the L:M cone ratio?” Vision Res 78:26-38 Details

Lawrance-Owen AJ, Bargary G, Bosten JM, Goodbourn PT, Hogg RE, Mollon JD (2013), “Genetic association suggests that SMOC1 mediates between prenatal sex hormones and digit ratio.” Hum Genet 132(4):415-21 Details

McClements M, Davies WI, Michaelides M, Carroll J, Rha J, Mollon JD, Neitz M, MacLaren RE, Moore AT, Hunt DM (2013), “X-linked cone dystrophy and colour vision deficiency arising from a missense mutation in a hybrid L/M cone opsin gene.” Vision Res 80:41-50 Details


Bhardwaj R, Mollon JD, Smithson HE (2012), “Compatible and incompatible representations in visual sensory storage.” J Vis 12(5) Details

Bosten JM, Mollon JD (2012), “Kirschmann's Fourth Law.” Vision Res 53(1):40-6 Details

Danilova MV, Mollon JD (2012), “Foveal color perception: minimal thresholds at a boundary between perceptual categories.” Vision Res 62:162-72 Details

Danilova MV, Mollon JD (2012), “Cardinal axes are not independent in color discrimination.” J Opt Soc Am A Opt Image Sci Vis 29(2):A157-64 Details

Goodbourn PT, Bosten JM, Hogg RE, Bargary G, Lawrance-Owen AJ, Mollon JD (2012), “Do different 'magnocellular tasks' probe the same neural substrate?” Proc Biol Sci 279(1745):4263-71 Details

Mollon JD, Cavonius LR (2012), “The Lagerlunda collision and the introduction of color vision testing.” Surv Ophthalmol 57(2):178-94 Details


Michaelides M, Rha J, Dees EW, Baraas RC, Wagner-Schuman ML, Mollon JD, Dubis AM, Andersen MK, Rosenberg T, Larsen M, Moore AT, Carroll J (2011), “Integrity of the cone photoreceptor mosaic in oligocone trichromacy.” Invest Ophthalmol Vis Sci 52(7):4757-64 Details

Smith WS, Mollon JD, Bhardwaj R, Smithson HE (2011), “Is there brief temporal buffering of successive visual inputs?” Q J Exp Psychol (Colchester) 64(4):767-91 Details

Thomas PB, Formankiewicz MA, Mollon JD (2011), “The effect of photopigment optical density on the color vision of the anomalous trichromat.” Vision Res 51(20):2224-33 Details

To MP, Regan BC, Wood D, Mollon JD (2011), “Vision out of the corner of the eye.” Vision Res 51(1):203-14 Details


Bosten JM, Mollon JD (2010), “Is there a general trait of susceptibility to simultaneous contrast?” Vision Res 50(17):1656-64 Details

Danilova MV, Mollon JD (2010), “Parafoveal color discrimination: a chromaticity locus of enhanced discrimination.” J Vis 10(1)

Jordan G, Deeb SS, Bosten JM, Mollon JD (2010), “The dimensionality of color vision in carriers of anomalous trichromacy.” J Vis 10(8):12 Details


Danilova MV, Mollon JD (2009), “The symmetry of visual fields in chromatic discrimination.” Brain Cogn 69(1):39-46 Details

Formankiewicz MA, Mollon JD (2009), “The psychophysics of detecting binocular discrepancies of luminance.” Vision Res 49(15):1929-38 Details

Lee RJ, Mollon JD, Zaidi Q, Smithson HE (2009), “Latency characteristics of the short-wavelength-sensitive cones and their associated pathways.” J Vis 9(12):5.1-17 Details

Mollon JD (2009), “Unlocked edges or contradictory edges? Is there a master signal for edge locking?” Perception 38(6):832-4; discussion 836 Details

Mollon JD (2009), “A neural basis for unique hues?” Curr Biol 19(11):R441-2; author reply R442-3 Details


Blake Z, Land T, Mollon J (2008), “Relative latencies of cone signals measured by a moving vernier task.” J Vis 8(16):16.1-11 Details


Danilova M, Mollon J (2006), “The gap effect is exaggerated in parafovea.” Vis Neurosci 23(3-4):509-17 Details

Danilova MV, Mollon JD (2006), “The comparison of spatially separated colours.” Vision Res 46(6-7):823-36 Details

Hood SM, Mollon JD, Purves L, Jordan G (2006), “Color discrimination in carriers of color deficiency.” Vision Res 46(18):2894-900 Details

Mollon J (2006), “Monge: The Verriest lecture, Lyon, July 2005.” Vis Neurosci 23(3-4):297-309 Details

Smithson H, Mollon J (2006), “Do masks terminate the icon?” Q J Exp Psychol (Colchester) 59(1):150-60 Details


Michaelides M, Holder GE, Webster AR, Hunt DM, Bird AC, Fitzke FW, Mollon JD, Moore AT (2005), “A detailed phenotypic study of "cone dystrophy with supernormal rod ERG".” Br J Ophthalmol 89(3):332-9 Details

Michaelides M, Johnson S, Bradshaw K, Holder GE, Simunovic MP, Mollon JD, Moore AT, Hunt DM (2005), “X-linked cone dysfunction syndrome with myopia and protanopia.” Ophthalmology 112(8):1448-54 Details

Michaelides M, Johnson S, Simunovic MP, Bradshaw K, Holder G, Mollon JD, Moore AT, Hunt DM (2005), “Blue cone monochromatism: a phenotype and genotype assessment with evidence of progressive loss of cone function in older individuals.” Eye (Lond) 19(1):2-10 Details


Johnson S, Michaelides M, Aligianis IA, Ainsworth JR, Mollon JD, Maher ER, Moore AT, Hunt DM (2004), “Achromatopsia caused by novel mutations in both CNGA3 and CNGB3.” J Med Genet 41(2):e20 Details

Michaelides M, Aligianis IA, Ainsworth JR, Good P, Mollon JD, Maher ER, Moore AT, Hunt DM (2004), “Progressive cone dystrophy associated with mutation in CNGB3.” Invest Ophthalmol Vis Sci 45(6):1975-82 Details

Michaelides M, Holder GE, Bradshaw K, Hunt DM, Mollon JD, Moore AT (2004), “Oligocone trichromacy: a rare and unusual cone dysfunction syndrome.” Br J Ophthalmol 88(4):497-500 Details

Smithson HE, Mollon JD (2004), “Is the S-opponent chromatic sub-system sluggish?” Vision Res 44(25):2919-29 Details

Thomas PB, Mollon JD (2004), “Modelling the Rayleigh match.” Vis Neurosci 21(3):477-82 Details


Danilova MV, Mollon JD (2003), “Comparison at a distance.” Perception 32(4):395-414 Details

Michaelides M, Aligianis IA, Holder GE, Simunovic M, Mollon JD, Maher ER, Hunt DM, Moore AT (2003), “Cone dystrophy phenotype associated with a frameshift mutation (M280fsX291) in the alpha-subunit of cone specific transducin (GNAT2).” Br J Ophthalmol 87(11):1317-20 Details

Sumner P, Mollon JD (2003), “Colors of primate pelage and skin: objective assessment of conspicuousness.” Am J Primatol 59(2):67-91 Details


Kim YG, Mollon JD (2002), “Conditions under which stereopsis and motion perception are blind.” Perception 31(1):65-71 Details

Mollon JD (2002), “The origins of the concept of interference.” Philos Transact A Math Phys Eng Sci 360(1794):807-19 Details

Sumner P, Adamjee T, Mollon JD (2002), “Signals invisible to the collicular and magnocellular pathways can capture visual attention.” Curr Biol 12(15):1312-6 Details


Regan BC, Julliot C, Simmen B, Viénot F, Charles-Dominique P, Mollon JD (2001), “Fruits, foliage and the evolution of primate colour vision.” Philos Trans R Soc Lond B Biol Sci 356(1407):229-83 Details

Simunovic MP, Regan BC, Mollon JD (2001), “Is color vision deficiency an advantage under scotopic conditions?” Invest Ophthalmol Vis Sci 42(13):3357-64 Details


Gregory-Evans K, Kelsell RE, Gregory-Evans CY, Downes SM, Fitzke FW, Holder GE, Simunovic M, Mollon JD, Taylor R, Hunt DM, Bird AC, Moore AT (2000), “Autosomal dominant cone-rod retinal dystrophy (CORD6) from heterozygous mutation of GUCY2D, which encodes retinal guanylate cyclase.” Ophthalmology 107(1):55-61 Details

Sivagnanasundaram S, Morris AG, Gaitonde EJ, McKenna PJ, Mollon JD, Hunt DM (2000), “A cluster of single nucleotide polymorphisms in the 5'-leader of the human dopamine D3 receptor gene (DRD3) and its relationship to schizophrenia.” Neurosci Lett 279(1):13-6 Details

Sumner P, Mollon JD (2000), “Chromaticity as a signal of ripeness in fruits taken by primates.” J Exp Biol 203(Pt 13):1987-2000 Details

Sumner P, Mollon JD (2000), “Catarrhine photopigments are optimized for detecting targets against a foliage background.” J Exp Biol 203(Pt 13):1963-86 Details


Dulai KS, von Dornum M, Mollon JD, Hunt DM (1999), “The evolution of trichromatic color vision by opsin gene duplication in New World and Old World primates.” Genome Res 9(7):629-38 Details

Mollon JD (1999), “Color vision: opsins and options.” Proc Natl Acad Sci U S A 96(9):4743-5 Details


Hunt DM, Dulai KS, Cowing JA, Julliot C, Mollon JD, Bowmaker JK, Li WH, Hewett-Emmett D (1998), “Molecular evolution of trichromacy in primates.” Vision Res 38(21):3299-306 Details

Jordan G, Mollon JD (1998), “Shift in Rayleigh matches after adaptation to monochromatic light of various intensities.” Vision Res 38(21):3253-7 Details

Mollon JD, Regan BC, Bowmaker JK (1998), “What is the function of the cone-rich rim of the retina?” Eye (Lond) 12 ( Pt 3b):548-52 Details

Regan BC, Freudenthaler N, Kolle R, Mollon JD, Paulus W (1998), “Colour discrimination thresholds in Parkinson's disease: results obtained with a rapid computer-controlled colour vision test.” Vision Res 38(21):3427-31 Details

Regan BC, Julliot C, Simmen B, Viénot F, Charles-Dominique P, Mollon JD (1998), “Frugivory and colour vision in Alouatta seniculus, a trichromatic platyrrhine monkey.” Vision Res 38(21):3321-7 Details

Simunovic MP, Votruba M, Regan BC, Mollon JD (1998), “Colour discrimination ellipses in patients with dominant optic atrophy.” Vision Res 38(21):3413-9 Details


Brettel H, Viénot F, Mollon JD (1997), “Computerized simulation of color appearance for dichromats.” J Opt Soc Am A Opt Image Sci Vis 14(10):2647-55 Details

Gaitonde EJ, Sivagnanasundaram S, Morris AG, McKenna PJ, Mollon JD, Hunt DM (1997), “The number of triplet repeats in five brain-expressed loci with CAG repeats is not associated with schizophrenia.” Schizophr Res 25(2):111-6 Details

Jordan G, Mollon JD (1997), “Adaptation of colour vision to sunlight.” Nature 386(6621):135-6 Details

Mollon JD (1997), “"... On the basis of velocity clues alone": some perceptual themes 1946-1996.” Q J Exp Psychol A 50(4):859-78 Details

Webster MA, Mollon JD (1997), “Adaptation and the color statistics of natural images.” Vision Res 37(23):3283-98 Details

Webster MA, Mollon JD (1997), “Motion minima for different directions in color space.” Vision Res 37(11):1479-98 Details


Bonnardel V, Bellemare H, Mollon JD (1996), “Measurements of human sensitivity to comb-filtered spectra.” Vision Res 36(17):2713-20 Details

Gaitonde EJ, Morris A, Sivagnanasundaram S, McKenna PJ, Hunt DM, Mollon JD (1996), “Assessment of association of D3 dopamine receptor MscI polymorphism with schizophrenia: analysis of symptom ratings, family history, age at onset, and movement disorders.” Am J Med Genet 67(5):455-8 Details

Mollon JD, Danilova MV (1996), “Three remarks on perceptual learning.” Spat Vis 10(1):51-8 Details

Mollon JD, Perkins AJ (1996), “Errors of judgement at Greenwich in 1796.” Nature 380(6570):101-2 Details


Hunt DM, Cowing JA, Patel R, Appukuttan B, Bowmaker JK, Mollon JD (1995), “Sequence and evolution of the blue cone pigment gene in Old and New World primates.” Genomics 27(3):535-8 Details

Hunt DM, Dulai KS, Bowmaker JK, Mollon JD (1995), “The chemistry of John Dalton's color blindness.” Science 267(5200):984-8 Details

Jordan G, Mollon JD (1995), “Rayleigh matches and unique green.” Vision Res 35(5):613-20 Details

Morris AG, Gaitonde E, McKenna PJ, Mollon JD, Hunt DM (1995), “CAG repeat expansions and schizophrenia: association with disease in females and with early age-at-onset.” Hum Mol Genet 4(10):1957-61 Details

Shyue SK, Hewett-Emmett D, Sperling HG, Hunt DM, Bowmaker JK, Mollon JD, Li WH (1995), “Adaptive evolution of color vision genes in higher primates.” Science 269(5228):1265-7 Details

Viénot F, Brettel H, Ott L, Ben M'Barek A, Mollon JD (1995), “What do colour-blind people see?” Nature 376(6536):127-8 Details

Webster MA, Mollon JD (1995), “Colour constancy influenced by contrast adaptation.” Nature 373(6516):694-8 Details


Dulai KS, Bowmaker JK, Mollon JD, Hunt DM (1994), “Sequence divergence, polymorphism and evolution of the middle-wave and long-wave visual pigment genes of great apes and Old World monkeys.” Vision Res 34(19):2483-91 Details

Regan BC, Reffin JP, Mollon JD (1994), “Luminance noise and the rapid determination of discrimination ellipses in colour deficiency.” Vision Res 34(10):1279-99 Details

Webster MA, Mollon JD (1994), “The influence of contrast adaptation on color appearance.” Vision Res 34(15):1993-2020 Details


Hunt DM, Williams AJ, Bowmaker JK, Mollon JD (1993), “Structure and evolution of the polymorphic photopigment gene of the marmoset.” Vision Res 33(2):147-54 Details

Jordan G, Mollon JD (1993), “A study of women heterozygous for colour deficiencies.” Vision Res 33(11):1495-508 Details

Jordan G, Mollon JD (1993), “The Nagel anomaloscope and seasonal variation of colour vision.” Nature 363(6429):546-9 Details

Mollon JD (1993), “Colour vision: mixing genes and mixing colours.” Curr Biol 3(2):82-5 Details

Webster MA, Mollon JD (1993), “Contrast adaptation dissociates different measures of luminous efficiency.” J Opt Soc Am A 10(6):1332-40 Details


Ibbotson RE, Hunt DM, Bowmaker JK, Mollon JD (1992), “Sequence divergence and copy number of the middle- and long-wave photopigment genes in Old World monkeys.” Proc Biol Sci 247(1319):145-54 Details

Mollon J (1992), “Colour vision. Worlds of difference.” Nature 356(6368):378-9 Details

Mollon JD, Astell S, Cavonius CR (1992), “A reduction in stimulus duration can improve wavelength discriminations mediated by short-wave cones.” Vision Res 32(4):745-55 Details

Mollon JD, Bowmaker JK (1992), “The spatial arrangement of cones in the primate fovea.” Nature 360(6405):677-9 Details

Morgan MJ, Adam A, Mollon JD (1992), “Dichromats detect colour-camouflaged objects that are not detected by trichromats.” Proc Biol Sci 248(1323):291-5 Details

Tovée MJ, Bowmaker JK, Mollon JD (1992), “The relationship between cone pigments and behavioural sensitivity in a New World monkey (Callithrix jacchus jacchus).” Vision Res 32(5):867-78 Details

Williams AJ, Hunt DM, Bowmaker JK, Mollon JD (1992), “The polymorphic photopigments of the marmoset: spectral tuning and genetic basis.” EMBO J 11(6):2039-45 Details


Bowmaker JK, Astell S, Hunt DM, Mollon JD (1991), “Photosensitive and photostable pigments in the retinae of Old World monkeys.” J Exp Biol 156:1-19 Details

Mollon J (1991), “G protein-coupled receptors. Hue and the heptahelicals.” Nature 351(6329):696-7 Details

Webster MA, Mollon JD (1991), “Changes in colour appearance following post-receptoral adaptation.” Nature 349(6306):235-8 Details


Mollon JD (1990), “Neurobiology. The club-sandwich mystery.” Nature 343(6253):16-7 Details


Mollon JD (1989), “"Tho' she kneel'd in that place where they grew..." The uses and origins of primate colour vision.” J Exp Biol 146:21-38 Details


Mollon JD, Estévez O (1988), “Tyndall's paradox of hue discrimination.” J Opt Soc Am A 5(1):151-9 Details

Travis DS, Bowmaker JK, Mollon JD (1988), “Polymorphism of visual pigments in a callitrichid monkey.” Vision Res 28(4):481-90 Details


Bowmaker JK, Jacobs GH, Mollon JD (1987), “Polymorphism of photopigments in the squirrel monkey: a sixth phenotype.” Proc R Soc Lond B Biol Sci 231(1264):383-90 Details

Mollon JD (1987), “History of optics: John Elliot MD (1747-1787).” Nature 329(6134):19-20 Details

Mollon JD, Stockman A, Polden PG (1987), “Transient tritanopia of a second kind.” Vision Res 27(4):637-50 Details


Mollon J (1986), “Walter Stanley Stiles 1901-1985.” Perception 15(6):657-66 Details

Mollon JD (1986), “Perception. Questions of sex and colour.” Nature 323(6089):578-9 Details

Mollon JD (1986), “Molecular genetics: understanding colour vision.” Nature 321(6065):12-3 Details

Stockman A, Mollon J (1986), “The spectral sensitivities of the middle- and long-wavelength cones: an extension of the two-colour threshold technique of W S Stiles.” Perception 15(6):729-54 Details


Bowmaker JK, Jacobs GH, Spiegelhalter DJ, Mollon JD (1985), “Two types of trichromatic squirrel monkey share a pigment in the red-green spectral region.” Vision Res 25(12):1937-46 Details


Mollon JD, Bowmaker JK, Jacobs GH (1984), “Variations of colour vision in a New World primate can be explained by polymorphism of retinal photopigments.” Proc R Soc Lond B Biol Sci 222(1228):373-99 Details


Dartnall HJ, Bowmaker JK, Mollon JD (1983), “Human visual pigments: microspectrophotometric results from the eyes of seven persons.” Proc R Soc Lond B Biol Sci 220(1218):115-30 Details


Mollon JD (1982), “Color vision.” Annu Rev Psychol 33:41-85 Details


Jacobs GH, Bowmaker JK, Mollon JD (1981), “Behavioural and microspectro-photometric measurements of colour vision in monkeys.” Nature 292(5823):541-3 Details


Bowmaker JK, Dartnall HJ, Mollon JD (1980), “Microspectrophotometric demonstration of four classes of photoreceptor in an old world primate, Macaca fascicularis.” J Physiol 298:131-43 Details

Mollon JD (1980), “Post-receptoral processes in colour vision.” Nature 283(5748):623-4 Details

Mollon JD, Polden PG (1980), “A curiosity of light adaptation.” Nature 286(5768):59-62 Details

Polden PG, Mollon JD (1980), “Reversed effect of adapting stimuli on visual sensitivity.” Proc R Soc Lond B Biol Sci 210(1179):235-72 Details


Bowmaker JK, Dartnall HJ, Mollon JD (1979), “The violet-sensitive receptors of primate retinae [proceedings].” J Physiol 292:31P Details

Mollon JD, Polden PG (1979), “Post-receptoral adaptation.” Vision Res 19(4):435-40 Details

Pugh EN Jr, Mollon JD (1979), “A theory of the pi1 and pi3 color mechanisms of Stiles.” Vision Res 19(3):293-312 Details


Bowmaker JK, Dartnall HJ, Lythgoe JN, Mollon JD (1978), “The visual pigments of rods and cones in the rhesus monkey, Macaca mulatta.” J Physiol 274:329-48 Details


Mollon JD, Polden PG (1977), “An anomaly in the response of the eye to light of short wavelengths.” Philos Trans R Soc Lond B Biol Sci 278(960):207-40 Details

Mollon JD, Polden PG (1977), “Saturation of a retinal cone mechanism.” Nature 265(5591):243-6 Details


Mollon JD, Polden PG (1976), “Absence of transient tritanopia after adaptation to very intense yellow light.” Nature 259(5544):570-2 Details

Mollon JD, Polden PG (1976), “Proceedings: Transient tritanopia: evidence for interaction between colour mechanisms.” J Physiol 254(1):66P-67P Details

Mollon JD, Polden PG (1976), “Proceedings: Some properties of the blue cone mechanism of the eye.” J Physiol 254(1):1P-2P Details


Mollon JD, Polden PG (1975), “Colour illusion and evidence for interaction between cone mechanisms.” Nature 258(5534):421-2 Details

Mollon JD, Ross H (1975), “Gregory on the sun illusion.” Perception 4(1):115-8 Details


Mollon JD, Krauskopf J (1973), “Reaction time as a measure of the temporal response properties of individua colour mechanisms.” Vision Res 13(1):27-40 Details


Krauskopf J, Mollon JD (1971), “The independence of the temporal integration properties of individual chromatic mechanisms in the human eye.” J Physiol 219(3):611-23 Details


Mollon JD (1968), “Fixation and perception.” Opt Acta (Lond) 15(3):295-7 Details