Bites Joanna of Pye 2001), 92. 79-112 Pron se Goat Brita 9 © 201 The British Poechological Society Surveying the seen: 100 years of British vision Nicholas J. Wade* Department of Prysbolugy, Uneteruity of Danda. UR Vicki Bruce Department o} Piysbology Uanseraty of Sterling UK Perceptual phenomena and their interpretations have fashioned the couse of psychology “This arecle surveys how theories of visual percepaan and methodologies have developed daring the lifetime of che Briush Psychological Society The experimental study ot ‘sion was snstigated by Beteih natural philosophers sn the «aly nineteenth century bue his ampetus way not manncained thereafter Nor uel the 1950 and 19 10s dud research fon perception resume in earnest wichin British universities. The adoption ot concepts ‘uch as schema) porennilly grounded sn neural organtaation, particularly by Barthete and Craik, accelerated experimental. cheoretical aod applied! vision research Brom mid-century the influence uf sntormation provesing models of perception became sncreasingly dominant, and they were often integeated wich che rapudly expanding, understanding of neurophysiological underpinnings The eprtame of these develop ments was Marr’ model of vision which, #9 ou y1e, marked the start ot the modern era of vision research Computers have transformed th response measuremene in percepeual experiments More naturalistic stimule can be presente and manipulated, anx! comple behavioural eeyponses, such as patterns of «ve movements, fractionated Non-invasive recording. of brain acereity to visual semua tion has similarly been transformed with a vanety of methods for imaging brain acuvity Neurormaging has been applied to localiing perceptual and cagniave functions and an studying pasents with known defe es ip visual ro ogmition Howeeer, che eagerness with which the computer has been auopted by perceptual pyc halogises ss Inkely eo be tempered by a growing awareness of the dterences heeween viewing, secs and simulations of them jure of stimulus control sind All that we know of nature, or of exsstence, may be compared tu a tree, which ach 19 oot, trunk, and branches In chis cree of knowledge, percepcion 1 the t00t, commun understanding ts the crunk and the sciences are the branches Thomas Reid (1764, p 120 Perception provides not only the roots to the tree of knowledge, but also to the sapling of psychology. Le was a dominane factor in the development of psychology and it remains one of che domains in which progress can be charted: This review briefly describes 19th century influences on perceptual research during the 20ch century, before highligheing, * Requests tor reprints should be addeesed e Prolessor Nick Wade Deyurtmear of Pychology, Univeriey of Dundee, Dundee DDI 4HIN UK (e-mail NJ WADE@dandee wc uk) Copyright © 2001. All rights reserved.80 Nucholas J, Wade and Vicks Bruce important landmarks in perceptual research during the lifetime of che British Psycho- logical Society. The final sections describe some of the most important contemporary work in visual perception, and speculate about how chese areas may develop in the next few years, Given the enormous volume, and considerable progress, of research in visual percep tion, we have had to be selective in the copies we have been able to elaborate here The motion after-effect and aspects of face perception feature more prominencly than they might if this review had been crafted by others, but we hope our selectivity allows us to achieve more coherence than might be possible with a more even-handed selection of topics. One enduring feature of che scudy of visual perception, well illustrated sn the areas we have chosen to stress, has been ats interdisciphinarity. Ie 1s difficult co discuss specifically psychological research psychologists are among many varieties of vision scienrise. One rather disturbing recene trend has been the tendency for the vision science community set itself apart from the psychological one We demonstrate chat there remain genuinely psychological problems within the field of vision science that require contact beeween vision and other areas of psychology This 1s certainly a lesson that can be drawn from its history Nineteenth century precursors ‘The empiicist philosophers, from Locke onwards, routed the acquisition of knowledge through che senses, and subsequent empirical psychologists have sought to sign che way an greater decail ‘The first stage involved developing experimental procedures that would bring some precision in stimulus control akin co chac adopted successfully in che physical sciences Natural philosophers in Britain devised the prinerples on which che perception of colour, motion and depth could be rendered experimentally tractable T. Young (1802) speculated that colour perception could be based upon the detection of three primaries and Maxwell (1855) provided experimental support for chis trichromatic theory Farad (1831, see Fig 1a) suggested how successive images presented in close temporal sequence could result 1m the perception of movement, setting in crain the long line of research on stroboscopic motion Talbor (1834, see Fig 1b), before he turned co photography, established a lawful relationship beeween apparene brightness and sncermicrent light stimulation Wheatstone (1838, 1852; see Fig 1c) demonstrated char depth could be synthesized from two slightly disparace images presented co separate eyes, dissociating depth perception from sts object base, he also developed the electromagnetic chronoscope thae subsequently was used for reaction time measurements (see Edgell & Symes, 1906) These insights extended the scope of experimental perception, although they were nor actively pursued in Britain to the extent that they were adopted and adapted wichin the German scientific community (see Wade & Heller, 1997) However, chose who sought co control the stimulus had less regard for the response Psychophysics developed almost independently of chese inscruments for stimulus manipulation, and the methods devised by Weber and Fechner were based on very simple tasks like lifting weights (sce Ross & Murray, 1978) William James (1890, pp 226-7) noted thar Wheatstone’s first paper contains the yetm of almost all che methods applied since to che study of optical perception It seems 4 puty that England {sic}, leadeng off so brilliantly che madern epoch of this study, should so quickly Copyright © 2001. All rights reserved.Surveying the won 81 Figure 1. The inscrumental origins of percepeual research, (4) Michael Faraday (1791-1867) alter a portrase trom The World Greas Men (London ‘The London Printing and Publishing Co 185 #) enclosed in a pattern produced by his counter-rotating sectors from Bagaday (1851), (b Walliam Henry Box Talboe 1 800—197) alter a photographic reproduction trom (00 Vans of Phutsgraply 1859-1939 (London The Science Museaam) presented disc chat he employed for incermaccent seamulacion (trom Talbur, 183.1), (¢) Charles Whearsrone (1802-1875) alter a poreiait from Th Hateatad Landon New. 868), 52. 145, combsned a0 the murtors of his stereoscope (from Wade, 1995) FO Mlusteation by Nicholas Wade Reprinted with permisuon } un have dropped our of the field Almoxe all subsequent progress has been made in Germany, Holland, and, Hongo entervalle, Ameria Helmholtz (1867, 2000) was particularly attracted co the experimental approach and his students developed methods further (see Cahan, 1993). The dominance of German arly, chough indirectly, reflected in what 1s perhaps the most chorough review of vision at the curn of the century in Rivers's (1900) survey more than 759% of references were to German sources arch mn perception 4s cl82 Nicholas J Wade and Vicks Bruce The emergence of British psychology Boring (1942) remarked that Helmholtz carried che corch of philosophical empiricism 1n a hostile Kantian climate, as did his erstwhile assistant Wunde However, their brands of empiricism were quite different Helmholtz borrowed the notion of unconscious inference from Berkeley to account for characteristics of colour and space perception, and the concepe 1s still active in some theories Wunde was more ambitious and applied empiricist and associationist ideas to account for consciousness itself. His ideas were cared co America by the likes of Titchener (see Hilgard, 1987), but his structuralise theory was not widely followed in Britain The demise of steucturalism in the second decade of the 20th century and its gradual replacement by behaviourism on the one hand and Gestalt psychology on the other also seemed to pass relatively unnoticed within British psychology Behaviourism, with ts reliance on associationism, was strongly opposed by James Ward (1886, 1918, see Fig 2a) who maintained an introspeccionist approach In the firse article published in the British Journal of Psychology, Ward wrove ‘phystological and comparative psychology muse fall back on the facts and analysis of our own experience’ (1904, p 25) His widespread influence was one factor in rerarding the spread of experimental psychology, and 1 reflected che neglect of psychology generally 1 British academic inseiturions Relatively lieele research in experimental percep: tion was conducted within Bricain until well into che 20ch century Some of those who were active experimentalists were often discouraged from conducting experimental research For example in 1904, when Wilham McDougall (Fig 2c) took a readership at Oxford University, the cerms of his appointment denied him access co @ laboratory However, he did contribute many arcicles on perception (with experimental studies of afterimages, binocular rivalry, Fechner’s paradox, fluttering hearts, infant colour vision, and visual persistence) co the initial volumes of the Britih Journal of Psychology, before his interests became more hormic Indeed, the initial volumes of ehe journal were weighted heavily 1n favour of perception, particularly stucles of illusions, but this did nor continue ‘Thus probably reflected che influence of W HR Rivers (Fig 2b), who wrote an extensive review of vision at che turn of the cencury (Rivers, 1900) He 1s most noced for instigating cross-cultural investigations of perception, and he also conducted neurological research with Henry Head (see Deregowski, 1998, Whittle, 2000). Rivers delivered lectures on sensory physiology at both Cambridge and University College, London (UCL). McDougall supervised the research ac UCL of Adolf Wohlgemuth (1911) on che motion after-effect (MAE) and acted as a participant in some of the experiments Following observation of a moving surface, like descending water, stationary objects appear to move in che opposite direction This MAE was described graphically by Addams (1834) after observing the Falls of Foyers in Scotland, and it was called che waterfall slluston by Silvanus Thompson (1880) As was che case for stereoscopic depth perception, most of the research on this copic in the 19¢h century was conducted in Germany (see Wade, 1994) Wohlgemuth’s doctoral dissertation was published as the first Monograph Supplement of the British Journal of Piychology he summarized the extensive research on MAES, added novel experimental variations of his own, and advanced a physiological model co account for its occurrence The conservatism and suspicion of British universities to newly established disciplines with Copyright © 2001. All rights reserved.Surveying the sen 83 Figure 2 The edsrorial base of the firse volumes of the Bust / Ward and Rivers were the editors and McDougall was on the editorial board (a) James Ward (18 13—1925) alter a lroneaspuece portrait tn che Britvb Journal of Prycbology (1925), text trom Ward (1918), db) Walliam Halse Rivers Ravers (1864 1922) after a frontispiece porteart 1n Ravers (1923), cext trom Revers (1900), (0X McDougall (1871 1948) after a portsae an sided in signitcant dares for che British Psychological Jona of Pry hub chison (19.26) cmb Society [© Illustration by Nicholas Wade Reyrinced with permission Copyright @ 2001. All rights reserved. 7Nicholas J. Wade and Vuks Brive ‘ina vst sd sntoreual popcorn fo ie ‘tay ety ate sop eed ty ob at ‘Finns i apminant Rafters sed ot "ale" Crate haa ie (a) ‘The Perception of Persons “There 1 one peopl eh gor more probaly set of rete ‘cheat, at die, y Gest fen tne winch we Bae Seay dscmeds 3s fom, wanante ECG eral aera, Spee tnd mova “36 cera wth are aoa ath the et ‘spuca of fgf 07 gl emoonae and ther scone 14, Sp ew ht Irom be / ite yas cen perce her poopie \ fees na tee te Some pce cy he Sanaa wha toe BRN are ay “ea ohn ans ay oan yi ei MO) oy ata a a rend do ome shat ier on, pee Tey ¢ \ cl Copyright © 2001. All rights reserved.Surveying the seen 85 was even evident in the first editorral of che Journal, and regret at the shift from ‘mental philosophy’ to ‘psychology’ was but chinly veiled; the editors (Ward and Rivers, 1904, p.1) wrote In dhis counery half a dozen leccureshups have been Lately founded in difference universities solely £0, promote the study of Psychology as science We have also a society of professed psychologses bach meets frequently for che discussion of princed papers ancl che exhibition af eapersmencal apparatus and resules Psychology was not accorded formal departmental status in Edinburgh and Cambridge until as late as 1931, and che incumbent professors (Drever and Bartlett, see Fig 3) fostered experimental approaches. Both F C. Barcleee and Drever directed psychological laboracortes before these dates, and both were active in che British Psychological Society, Bartlett edited the British Journal of Paycbulngy from 1924 uncil 1949, and he was assisted by Drever for the whole of that period. Nonetheless, in the words of Hearnshaw (1964, p 208) “The total leceuring staff in departments of psychology at the outbreak of Second World War numbered only about thirty: Some perceptual research was able co ride chis wave of indifference because 1¢ was nurtured in a more favorable climate While the ‘New Psychology’ of Wande had been srcually ignored wichin Britain, a “New Physiology’ was actively pursued by Ferrier, Hughlings Jackson, Sherrington and others. Wohlgemuth’s (1911) imrerpretations of che MAE bore closer alleysance co Sherringcon’s physiology than to Wunde's psychology The continuing research on colour vision was driven by the physical control of the samulus, and by increasing understanding of receptor funccion and colour anomalies (see Collins, 1925, Houston, 1932, Parsons, 1915) Indeed, 1¢ was che concepe of schema’, developed within chis new physiology by Head (1920), that was applied by F C Barclert to skilled asks of memory and perception According to Barelett (1932, p> 201),"' Schema’ refers to an active organisation of past reactions, or of past expertences, which must always be supposed to be operacing in the well-adapted organic response The construceave aspects ‘of both memory and perception were emphasized at che expense of their holistic or sequential feacures. Gradually, perception did find a place within Brash psychology, and se was artached to a new type of eheory linking, perception to prediction and accion ‘The emergence of British perception The ewo most prominent British perceptual psychologists throughout the 1930s were F C Bartlett and Maggre Vernon (Fig. 30), although Thouless (193 1a, 1931b) presented his analysis of perceptual constancy ac the beginning of the decade and Craik’s (1939) experiments on visual adaptation appeared at the end ‘The empiricist agenda initially was followed shghtly differently by Bartlett and Vernon, Bartlett (1932) examined tasks in Figure 3. The eablnhmene of perceptual psychology ww Baran (a) Jame Drover (RTIT950 ser a poreeast sn Collins (1951), ext from Drever (1921). (b) Freder: Charles Barcl te (1886-1969) after a portea sn Zanygwill (1970), eext from Barelete (1952), (¢) Magdalen Dorothea Vernon (1901~ 1991) alter a porteat kindly supplied by Halla Beloff, exe from Vernon (19701 LE Mluseeacion by Nicholas Wade Reprinted wich permission }86 Nicholas J. Wade and Vicks Brace which perception goes beyond the information given in the stimulus in a search to extract meaning ftom it, whereas Vernon (1937a, 1937b) investigated the traditional empiricist assue of space perception She also published several books on visual perception (see Vernon, 1937¢, 1952, 1970), the second of which adopted a more Barclectian position on perception. Any percepeual sicuation encouncered could then be firted anto its appropriate schema or chain of schemata, by vistue of the effore afcer meamng’, until w was understood and reacted co an the most appropriate and satisiying manner [1952, pp 258-9] Emphasis on the constructive and individual aspects of perception contradicted approaches chat stressed perceptual constancy, and equations for quancfying this had been proposed by Brunswik (1928) and Thouless (193 1a) Both proposed ratios involved differences between perceived and projected values on the one hand and physical and projected on the other, although Thouless used logarithmic transformations in order to avoid anomalies chat arose with che direce ratios Thouless (193 1a, 1931b) referred to perceptual constancy as ‘phenomenal regression to the real object’, and provided plentiful evidence to support its operation for shape, size, orventacion, brightness and colour perception Applied perception With che onset of war, in 1939, the approach co perception adopted by Bartlece was applied co human operators of complex systems, like flight simulation in the Cambeidge cockpit (see Barelect, 1946, Saito, 2000) The experimental research on perception 19 the 1940s harmonized with developments in cybernecies (Wiener, 1948), and Cratk (1943; see Fig. 4a) conflated the ewo by considering the human operaror as a complex, self-organizing system, His studies of visual adaptacion had indicated thac there was constane feedback from previous and concurrent stimulation, and that it could be modelled by physical processes In the epilogue to his PhD thesis on visual adaptation he wrore some of the flexibility of che percepeual proceys—for instance, the recogmeion of relational sicher than absolure propertics and of changes rather than constane sumulacion, anda primitive type of absteaction—follows from the known properties of the physiological structure and ean be emitated by physical mecharisms [1940, reprinted in Cratk, 1966, p 6] The machine metaphor was co prove particularly attractive to experimental psycho- logists, although Crakk was only able co enlist relatively simple machines He worked with analogue devices as che digital computer was still embryonic Nonetheless, Cratk did formalize che input, processing and output components of servo-systems 1n a manner that could be applied to digital computers ‘The essential feature of the sensory device 1 1s abshiey co translate the change 118 fo measure Ento some form of eneegy which can be ampliied and used co dnve the restoring cause The next parts ‘what may be called the computing device and controller, whch determines the amount and kine of energy co he released feom the effeccor unie co restore equilibrium The final parc 4s che power unt br efector (equivalent (0 the muscles sn men and animals) which restores the state of equulibeium 11966, pp 23-4) Copyright © 2001. All rights reserved. .Surveying the seen 87 After Crask’s untimely death in 1945, computing machines increased 1n speed and complexity so that the tasks that they could simulate became more complex Concepts from engineering, such as informacion and self-organrzation, were also intezraced with a growing knowledge of neurophysiology resulting in the computer becoming a metaphor for the brain The computers mounted for this metaphorical odyssey were digital and sertal, but ar around the same time the ground was being laud for principles of parallel processing McCulloch and Pite’s (1943) model of the neuron provided che foundation for later connectionist models of pattern recognition, and the networks connecting perception to its underlying physiology were further woven by Hebb (Fig 6a) in his speculative synchesis of perception and learning Hebb proposed that perceptual learning rakes place when assemblies of cells fire cogether, their revetberaeing, activity resulted in synaptic changes which further increased the probability of the nerves firing together The functions of cell-assemblies and phase sequences were based on his neurophysiological postulace When an axon of cell Avs near enough to excite Cell B and repeatedly or persistently cakes pare an Firing 16, some growth process ur mcrabobe change cakes place an one or bork cells such that A'S efficiency, as one of the cells firing Bay ancreased [Hebb, 1919, p» 621 Hebb's postulate 1s taken as providing the foundation for currence connectionist models of recognition and learning despite the fact chat the principle had been enunciated over 70. years earlier by Buin (1873, see Wilkes & Wade, 1997) Hebb later applied che concepts to account for a wide range of phenomena, from stabilized retinal images to sensory deprivation Bartlece’s emphasis on the constructive nature of perception found an echo in America in the New Look experiments, like those reported by Bruner and Postman (19 £7), where motivation was considered ro interact with perception Similar experimental investiga~ tions had been undertaken earlier by Brunswick (1934, 1935) who examined the perceived sizes of postage stamps of different value Ames’s many demonscracions of the ambiguities of stimulation and their perceptual resolutions (see Ieelson, 1952) were also accorded renewed attention in this cognitive climate In these heady post-war years personality flirted with perception, but their larson was not lasting Cestain subthreshold recognition phenomena were brought co their perceptual defence (see McGinnies, 19 19), but the sober verdict was not in their favour Te would seem wise co reyard with zrear caution che existence of limstacions on speud and accaracy ot percepe on imposed by personaly factors, ac lease n normal observers fVernon, 1970p 2 ‘The affair did focus actention on che lability of the men, which was scrurimzed 1 che theory of signal detection (Tanner & Swets, 195-1) Fechner's (1860) psychophysics had survived for almost a century bue st did nor provide a secure platform for predicting target derection in life-threatening situations Radar operators could report signals below their classically determined thresholds, buc they also made more mistakes Accordingly, che cask of detecting signals within background noise was analysed as a detasion process. The principal factors affeceang the dec ns reached were the payoffs for huts and false alarms and the signal probability (see Green & Swers, 1966) The acher assaule was on Fechner’s logarithmic law linking Copyright © 2001. All rights reserved.88 Nudulas J. Wade and Vuks Brive sensors and stimulus intensities {t hac been supported by indirect category scaling of sensory magnitude, but novel direce methods, such as magnitude escamation and cross- modality marching, cast doubt on its validity An alternative formulation in terms of a power function was proposed by Stevens (1951), although a variety of models can be plied to data from magnitude estimation (see Poulton, 1968) Dissatisfaction with classical psychophysics was also voiced by those who tried to apply it to skilled tasks ike flying (Gibson, 1950) Gibson also poured scorn on the artificiality of the laboratory for studying vision in comparison to the natural world, where objects and observers do not retain the same spatiotemporal relations to one another Gibson's ‘psychophysical theory of perception’ had little in common with Fechner, and he discarded the distinction beeween sensation and perception Information and pattern processing Information theory was developed in the context of celecommunications, and the mathematical measurement of information was formalized by Shannon and Weaver (1949), its powerful impact on perception was felt in the 1950s Miller (1957) linked the concept of limited information capacity co absolute perceptual judgments Actneave (1954) devised procedures to determine the locations of highest informacion in sumple patterns They corresponded to boundartes of brightness (contours) and particularly co abrupt changes 1n contour direction (corners) Support for the significance of concours in perception derived from two other sources single unit recordings from various levels in the visual pathway, and scanning eye movements Indeed, carly attempts to stabilize the retinal image by compensating for any involuntary eye movements resulted in disap- pearance of che target (Ditchburn & Ginsborg, 1952, Riggs & Racliff, 1952) However, it was the qualitative concept of information processing rather than quantitative information measures that was to have lasting appeal The perceiver was conceived of as a limited capacity informanion processor and the information could be filtered, filed or reformulated on the basis of stored events. Broadbent's (1958) model was among the first to formalize and represent pictorsally the putative processing stages He stated thar che “advantage of informacion theory rerms 15 that they emphasize che tionship between the stimulus now present and the others that might have been present bur are noc (1958, pp 306-7) Thus, Broadbent (Fig. 4b) combined Bartlere’s approach of examining skilled asks with Crauk’s modelling mecaphor. Theoretical attention was shifted towards pattern recognition by both humans and computers because they were both thought of as informacion processors or manipulators of symbolic information, The paccerns were typically outline figures or alphanumeric symbols, and rival theories, based on template marching and feature analysis (see Uhr, 1966), vied for simulaced supremacy at recognition and one result was pandemonium, (Selfridge, 1959)! Sutherland (Fig Gb) sounded a cautionary note on this endeavour chat was not generally heeded chen nor has been subsequently rel Patterns are of imporrance co animals and man only in so far as they sugmify objects Ie 1s the recognition of abjeces chat 15 vital for survival and asa gaude co accion, and the parrerned stim df our receptors 18 oF use only because 14s posseble co conseruct from 1 the nature of the objece from which 1 emanated {1975, p 157 Copyright © 2001. All rights reserved.Starveying the seen 89 Monten of eral come ene tat te “The arto whch geo bal ee ch coer 0 weal fe than ag TE ne ft eer nk of tern any gape wha tts rn HE Sere pmo rg aE Aten itfao eller smultancoudy, but then ere one emai Ge ony wh the het Aa etre Ge — *, Sagas oe hd Heng Sinan hee ae ‘Ming dau arb oo at se ates : th Fear “ely ERE In te she cae “he oft tts scp Set ew, Wingmen fd that conn Ue aed baes PAT hgh e sper rans a mano pedo Tene carb Scar nd Wee Yo three loudest wore Pete than one ib) Figure 4.(a) Kenneth Job William Ceask (1914-1945) alece a portrase in Bartlet (19146), (bi Danalel Bre Broadbent (1926~1993) after a porteaxt in Lundzey (1980), text from Broadbent (195K), (c) Horace Bas Barlow (b 1921) combined with a diagram of espace features sn che visual pathway uf the cat, as Mlusteated sn Barlow, Narasimban, & Rosenfeld (1972) [© Illustration by Nicholas Wade Repeinted wach permssson ] Copyright @ 2001. All rights reserved.90 Nuholas J. Wade and Vuks Brive Pattern recognition was the heme thae unified physiologists, psychologists and computer scientists, many of whom were assembled for a seminal symposium, held in 1959, on sensory communication (Rosenblich, 1961) While there were dangers in the oversimplification of che stimulus, che approach also allowed smportant tools to be developed co probe discrete visual achievements. One example was the random dot stereogram (RDS) developed by Julesz (1960). Wheatstone (1838) had employed outline figures for his stereoscope 1n order to reduce any monocular cues to depch, but he was acutely aware chat some remained. Julesz employed the dawning power of the computer to produce pairs of matrices of black and white dots, the central areas of which were displaced with respece to the common backgrounds, and hence disparate The displays looked amorphous when viewed by each eye alone, but when viewed binocularly patterns gradually arose or descended from the background This not only spawned a new area called ‘cyclopean perception’ (Julesz, 1971), but the technique was adopred in che clinic as a test for stereoscopic depeh perception. Analogous developments in the temporal domain produced random dot kunematograms which were used by Braddick (1974) and others to make important discoveries about different types of motion processing. Developments im rnsual nerorctence Research on patterned scamulacion at che receptor level had proceeded throughout the first half of the century, but its pace quickened thereafter ‘The glimmerings of pattern processing beyond the receptors emerged in the 1950s, and were amplified in che 1960s When recordings of nerve impulses could be made from individual cells in the visual pathway their adequate stimuli could be determined Adrian (1928) coined the term receptive field’ and Hartline (1938) applied ic co describe the region of the receptor surface over which the action of light modified the activity of a neuron It came as something of a surprise that retinal ganglion cells of frog responded to quite complex features of simulation (ike moving dark regions of a specific visual angle, resembling a bug), and stimulus properties that excited or inhibited neurons were generally called trigger features’ (Barlow, 1953, sce Fig fe) Retinal ganglion cells of cat, on the other hand, were excited by rather simpler stimulus arrangements Kuffler (1953) found thac they were concentrically and antagonistically organized, 1f the centre was excited by light the surround was inhibited, and vice versa Such an arrangement served the detection of differences in luminance well, but steady states would have cele effect, since excitation nullified inhibition. This pattern of neural activity was retamed in the lateral geniculate, bur ic underwent a radical change at the level of che visual cortex. Hubel and Wiesel (1962, 1968) found that single cells in primary visual cortex (V1), first of cat then of monkey, responded to specifically oriented edges, they had different recepeive field properties which were called simple, complex and hypercomplex. Physiologists refined the stimulus characteristics of trigger features throughout che 1960s, while psychologists sought their phenomenal counterparts. Almost any experi~ ment involving concours paid lip service to Hubel and Wiesel, despite che renuousness of the links between particular phenomena and their underlying physiology At least an appeal co trigger features was considered preferable co reliance on the speculacive neurophysiology advanced by Kohler (1940) Spatial sllustons, for example, attempted fo rise above their enigmatic seats by adopting this reductionsse parh, despite its Copyright © 2001. All rights reserved.Surveying the seen 91 attractions the greatest success was found for contour repulsion (Blakemore, Carpenter, & Georgeson, 1970) The alternative lure of illusions was to relate them to the traditional empiricist concept of constancy (Gregory, 1963) ‘The links between perception and physiology were made explicit for the motion after-effece (Barlow & Hill, 1963, Sutherland, 1961) resulting in an explosion of empirical studies examining their consequences. Barlow (1963) also investigated the link beeween visibility and rerinal image motion using afterimages and optically stabilized retinal umages The concept of channels or spacial filters emerged during the decade, and 1t was applied with particular rigour by Fergus Campbell (Fig. 6b) and his colleagues to the detection of and adaptation to sine-wave yratings (see Campbell & Robson, 1968) The attraction of gratings was thac they provided at one and the same came a definition of the stimulus and theory of the response to 1c. Cratk (1966; pp 4-5) characteristically foresaw the principle behind these developments the action of various physical devices which recognize or respond identically to certain simple objects can be treated an terms of such [mathematical] teansformacions ‘Thus the esentutl part of physical ‘recognizing’ snscruments 1s usually « filter-—wherher 4 be a mechanrcal sieve, an opeteal filter, of a tuned elecerical extcuit—which ‘passes’ only quantities of the kind 4 required to identify and eeyects all others Establishing the ceichromacy of vision also proved amenable to this approach, boch psychophysically (Rushton, 1964, Stiles, 1959) and physiologically (MacNichol, 1964) Visual development ‘The cortical mapping of visual receptive fields had an unexpecced influence on che age old nativisempiricise debate, providing fuel for borh sides Hubel and Wiesel (1963) demonstrated char receptive fields were present prior co visual experience but that chey could be modified by 1¢ This applied co both binocularity and orventation selectivity For example, the responsiveness of cortical cells to stumulacion by either eye, present at birth, could be modified by monocular deprivation from birch The timing of such modsfication was critical; Blakemore and van Sluyters (1974) demonstrated that che cortical mono- cularity could be reversed if the eye occluded was reversed in the fourth week of che kurten’s Infe Similar sensitive periods have been shown to operate for exposure to orientation (Blakemore & Mitchell, 1973) The relevance of chis research to develop- mental abnormalities in humans (like serabismus and astigmatism) has been stressed by many (see Blakemore, 1978, for a review of the early research) Contour extraction was also considered co be one of the first tasks tackled by che visual systems of newborns, and many novel methods were devised to study them For example, Fantz (1961) inferred infant perceptual discrimination from the patterns of differential fixation, and Bower (1966) used operant conditioning techniques for investigaeing the emergence of percept ncies Fantz even suggested that there was an innate preference for viewing human faces alchough che ourline figures he used as stimuli had very little ecological validicy, hus suggestions have been confirmed in recent work (e x see Johnson, Dzrurawrec, Ellis, & Morton, 1991) Salapatek and Kessen (1966) recorded eye ‘movements of infants only a few days old and found that they were concentrated on contours or corners Habituation co repeated presentations of a stamulus provided another Copyright © 2001. All rights reserved.92 Nuholas J. Wade and Vick: Brace source of inference regarding discrimination (Bridger, 1961) These methods were refined and the course of perceptual development began co be charted (see Cohen & Salapacek, 1975). From the late 1960s the principal stimulus for vision research became the sine wave grating, and these were presented to infants, too (eg Atkinson, Braddick, & Braddick. 1974; Atkinson, Braddick, & Moar, 1977) (Other, more recent British contributions to the study of visual developmen mentioned by Goswami in this issue.) Perception, cognition and action The informacion-processing agenda was at the heart of che ‘cognitive revolution’ which engulfed experimental psychology in the 1960s (see Gardner, 1987, Nersser, 1967), although chis had more marked repercussions for perceptual research in America than 1 Britain The distinctions beeween perception and thought became even more blurred, problems of pattern recognition were considered paramount, representational processes proliferated, and support for particular theories derived increasingly from computer simulations The erend was epitomized by Gregory's (1963, see Fig. 5a) analysis of visual illustons, and re was expressed eloquently in the many editions of hus Eye and brain (1966; pp 222-3) Perception becomes a macter of suggeseing. and testing hypotheses The continual searching for che bese incerpretacion 1s good evidence for the zeneral amportance of augmenting the limaations of che senses by imporcing erher knowledee This hypothesis testing theory was applied to a range of visual phenomena like ambiguous figures, size illusions and subjective contours Gregory was critical of many accempts co link perception with its underlying physiology, as chis reflected a passive rather chan an active theory All perceptions were considered co be cognitive fictions Though generally predictive, and so essentially correct, cognitive fictions may be wrong —to drive us into error On chis Acave view, both versal (cortecr-predictive) and allusory (alse-predicaved perceptions arc equally hetiams. To porceave ts to read the prose Concrol che future {197 4, xin erm of the past to predict snd He likened petcepeion co the operation of programs in computers with the same basic units performing many different tasks. This, in eurn, was used as an accack on accempts to localize bran functions by means of lesions, and in so doing he drew a discinccion between serial and parallel systems In a serial system che various tcentihable clemencs of che output are nor separately represented by discrete purts of the system che bratn cunsisted ofa series of independent parallel elements with separare ourpur cerminals for each, like piano, st might he possible eo ideneify behavioural elements with parcicalar parts of the system T1961, p 3217 Ic as precisely this distinction that has fuelled che modular approach outlined below Sutherland (1968, 1973, see Fiz 5b) developed the cognitive agenda in his advocacy of an alliance beeween perception and language. With regard to objece recognition he re ked that e must exist a set of rules chat makes 1¢ possible co map fram one domain ante another For ple i che case of solid objects compased of plane surfaces, each surface must be joined ac its Copyright © 2001. All rights reserved.Surveying the veen 93 o : encompassed ever more processes, like decision and scress (Broadbent 1971) and spawned offspring, hike logogens (Morton, 1969) Driver (this issue) provides further discussion of Broadbent's work, and Altmann (this issue) elaborates Morton's model. a ee ee94 Nubolas J. Wade and Vickr Brace However, almost all cognitive copics were systematically eschewed by Howard and Templeton (1966) in their integrative survey of human spatial orientation. They placed the spatial senses in an external framework and examined their interactions; particular importance was given to the many links between the visual and vestibular systems Howard (Fig 5c) continued his analytic approach to perception with detailed commen- anes on visual ortentation (Howard, 1982) and binocular vision (Howard & Rogers, 1995) Gabson (1966) sought to stem the cognitive current and developed a novel cheory that owed more to Thomas Reid (1764) chan to his own contemporaries When the senses are considered as perceptual systems, all theories of perception become at one stroke unnecessary Ir 1s no longer a question of how the mind operates on the deliverances of sense, or how past experience can organize the data, or cven how the brain can process the inputs of the nerves, but simply how informacion 1s puckedl up (Gibson, 1966, p 319} Despite Gibson’s pejorative purview of conventional perceptual experiments, the strongest support for his position derived therefrom: simplified dynamic dot patterns could be recognized far more easily chan static ones (Johannson, 1964) Gibson's ideas established a new field of ‘ecological’ optics which has been tilled by many Eor example, Lee (1976, Lee & Reddish, 1981) has examined perception in complex natural conditions, like driving and diving (See Haggard, this issue, for more on this topic ) Thus, the 1960s saw the beginnings of a split between a ‘cognitive’ approach, where the goal of vision could be seen as an abstract categorization of the objects of vision, and an ‘action’ approach, where vision was part of an integrated system allowing manipula tion of and navigation through the world. This distinction has matured in contemporary approaches to vision, both through the influence of David Marr and through further developments in neuroscience and neuropsychology The remainder of this article describes the past alongside present-day achievements and, where appropriate, dares to speculate about the furure Computers and computational theory Craik, as well as ‘Turing (see Millican & Clark, 1996), anticipated that the computer would be a powerful cool to simulate theores of perception, as well as providing a metaphor for the processes of perception and cognition themselves Since the lace 1960s the study of visual perception has been influenced profoundly by computers As well as allowing scientists to collect or to analyse data more quickly, the digital computer provided a cool for the laboratory scientist co develop new ways of cesting the visual system with novel kinds of visual displays ‘The move away from reliance on oscilloscopes o present sine wave and other simple pacterns facilitated the increasing use of more naturalistic patterns, as well as those which can be constructed and manipulated in controlled ways Computer developments also enabled the better recording of eye movements and the linkage of eye movements to changes 1n display features, allowing a number of groups to conduct ingenious experiments into che control of eye movements, in reading (e g Rayner, 1978) Computer developments also created new problems Unknown non-linearities in screen display properties or cheir cemporal characteristics present problems that were not initially foreseen (Kennedy, Brysbaere, & Murray, 1998), Copyright © 2001. All rights reserved. .Surveying the ven 95 te) Figure 6. (a) Donald Olding Hebb (1901-1985) alter a porctart in Lindsey (1980) with his eyes st che hidden Layer of a neural neework, (b) Fergus Wallium Campbell (19219936 after 4 photograph kindly supplied by Johar Rabson, presented en low conerase crossed! aracings, (C1 Daval Coureaey Marr 1345-1980 alter a photograph kindly supplied by Kent Stevens, embedded na pattern of ancerwccing Joes [© Iluseeason by Nichols Wade Reprinted with permssion J Copyright @ 2001. All rights reserved.96 Nudolas | Wade and Vicks Bruce 1n addition co a variety of problems chac have been termed ‘visual seress’ (Wilkins, 1995) During the 1970s David Marr (Fig 6c) worked with a number of outstanding collaborators, including Ellen Hildreth, Tommy Poggio and Shimon Ullman, on research which has significantly shaped our understanding of —and approach co—vision Marr set out to develop a complete framework for vision, spanning the very lowest level processes within che retina (Marr, 1976; Marr & Hildreth, 1980) up to the process of visual objece recognition (Marr & Nashthara, 1978) Sadly, Marr died very young, and his overall theory was published posthumously (Marr, 1982) The key feature of Marr's theory was that vision can be understood at different levels The first ‘computational’ level 1s a theory of the cask that the visual system 1s to solve, and an understanding of the constraints that can enable solution of that task ‘The second level, of ‘representation and algorithm’, 1s a means of achieving che cask, and the final hardware implementation’ level describes how the brain, or a computer, actually implements these algorithms in neural assue or silicon Marr (1982, p_5) argued chat For the subject of vision, there 2+ no single equation or view that explains everything Bach prob hhas co be addressed from several points of view—as a problem an represencany informacion, as computation capable of denving that representation, and 1s a problen computer capable of carryang out bork things quickly and seluably nthe archicecture of a The distinctions between Marr's three levels are not always quite as clear as Marr proposed, but there are important examples which make the distinctions clear For instance, Marr and Pogyao's orginal (1976) approach to stereopsis tackled che question of how the visual system can solve the correspondence problem in perceiving forms in depth from RDSs Somehow che brain 1s able co pair up the dots in each eye’s view of an RDS so that che three-dimensional solution 1s revealed, but s¢ 1s nor at all clear how this process can work given the enormous number of different potential solucrons Marr and Poggio’s computational theory specified three simple constraints on che matching process which cut down che number of potential matches, and which are sensible assumprions co make in che world of natural objects In the natural world, the same surface patch looks similar to each eye (so in an RDS, black dots muse match black dots), a surface patch can only be sn one place at a time (so 1n an RDS, each dot can match only one other), and discontinuities in depth at the edges of obyects are rare (so in an RDS disparity shifts should be rare—disparity should vary smoothly almost everywhere except at che rare boundartes between objects) Starting from this computational level Marr and Poggio (1976, 1979) produced ewo different algorithms for solving RDSs based upon rather different representations of images. Marr and Poggio’s first algorithm (1976) was cast 1n terms of @ neural network model in which the steady state for an RDS emerged from the balance of excitation and mbubition between simple processing units. Their second algorithm (1979) accommodated observations on che way in which stereopsis 1s guided by informacion in distance spatial frequency bands, making links with both psychophysics and physiological evidence for the separate analysis in early vision of information from different spatial scales In addition to presenting a unified approach to different topics within vision, Marr and hus colleagues also presented a complete theory of the different stages of representation {primal skecch, 2 5D sketch, 3D models) snvolved in the ncerprerarion of retinal images cremel Copyright © 2001. All rights reserved.Surveying the ven 97 In so doing, Marr distinguished a stage which made explicit the three-dimensional layout of the world with respect to the viewer (the 2 5D sketch), potentially useful for action 1n the world from the more abstrace 3D models which allowed object recognition Despre this distinction, however, his framework fails co anticipate the more recent idea of a separation of visual pathways for action and for object categorization, which we discuss facer Marr's framework for visual object recognition was developed and extended by Biederman and colleagues (Biederman, 1987, Hummel & Biederman, 1992), and sn the UK Humphreys and Riddoch made important experimental and neuropsychological tests of many of the claims about representation and sequence (eg Humphreys & Riddoch, 1984, 1987) Much recent theory about objece recognition, however, departs from Marr's economical object-centred representations, to develop representational ideas based around che representation of muluple viewpoints (¢ ¢ Bulehoff & Edelman, 1992; Tarr, 1995; see Biederman, Subramaniam, Bar, Kalocsar, & Fiser, 1999, tor a recente discussion) Thus 1¢ 1s not the details of Marr's cheory which have so far stood! the test of tume, but the approach itself In chis spirie, a great deal of important British research buile upon clus approach co develop new analyses of early visual processing (c g Watt & Morgan, 1985), stereopsis (eg. Mayhew & Frisby, 1981) and particularly motion processing, where importane concribucions have been made by a large aumber of UK scientists, Modular visual processing Another key feature of Marr's theory (1982, p_ 102) was the principle of modularity Computer scientists call che separate piecey of a process its moduiles, and the ides thar a large compuration can be sphe up anil implemented asa collection of parts that are as nearly andepenclene of ‘one another ay the overall ask allows, 18 0 amaportant that L was mowed to eleva principle of modular design ere to 4 principle the If a complex task like ‘vision’ comprises many discrete modules, more-or-less mutually independent, then failure sn one aspect of vision (¢ g the stereo ‘module’) 1s not fatal for other aspects of the task This means that modular designs make sense, both for the construction and debugging of computer programs and also for brains in che course of evolution. Indeed, there 15 good evidence from single cases of brain-inyured patients: increasingly studied by cognitive neuropsychologists since the 1970s thac visual perception may indeed be modular in the manner proposed by Marr Evidence for modularity of visual processing systems has been «round since the 19th century, but the neurological theory that there was a single area of primary visual cortex for the reception of retinal inputs, leading ro a subsequent stage of assoctacion with stored knowledge in the association cortex, held sway uncl the 1970s Zeki (1993) provides a fascinating description of how and why evidence for specialized subsystems of cortical visual processing was ignored for so long The 1970s and 1980s saw a revival in che use by perceptual theorists of patterns of perceptual and cognitive impairment observed in individual patients who had suffered brain injury The single case method of cognitive neuropsychology was pursued particularly in Europe Marr himself was influenced greatly by findings described by Copyright © 2001. All rights reserved.98 Nucbalas J. Wade and Vicks Bruce Elizabeth Warrington on dissociations between the perception and understanding of visual shapes (Warrington & Taylor, 1978, see Marr, 1982). Interest was revived in much earlier reported cases of apparent dissociations between the perception of form and colour (c.g Verrey, 1888, cited in Zeki, 1993) and beeween form and motion (Riddoch, 1917), Zeks (1993) provided a detailed history of how early cases of acquired colour blindness (cerebral achromatopsta) were described and how, eventually, che sdea became accepted chat there was an area of cortex where colour was analysed (now conventionally labelled V4) beyond the primary visual cortex (V1) One reason why 16 cook so long for the idea of a ‘colour centre’ co be accepted was that it can sometimes be argued that selective deficits arise because some casks are just coo difficule for a mildly injured brain to perform Cerebral achromatopsta, for example, often accompanying selective inability to recognize faces (prosopagnosia), might just show that injury co the visual cortex makes complex processing of colours and fine detail of faces impossible However, where a double disvocuation 1s Seen, where two different patients show opposite pacterns of deficient and spared ability, 1¢ 1s difficule co argue chat one or other task 1s incrinsically more difficule (see Shallice, 1988) During che 1970s, cognitive neuropsychologists began to understand the power of the double dissociation in highlighting potentially distinct modules. A particularly dramatic example 1s provided in che area of motion perception In 1917, George Riddoch (grandfather of Jane Riddoch, herself a neuropsychologise of distinction, eg see Humphreys & Riddoch, 1984, 1987) noticed char brain-injured soldiers from the Great War could sometimes see movements in otherwise blind visual fields His observations led him co suggest chat ‘the elementary visual perceptions, of light, of movement, and of an object, are dissociated’ (p 15) This idea was well ahead of its time, yet even in Riddoch’s day such an argument could have been supported by examples of the opposite condition, in which a person could see reasonably normally when chings were stl, but was blind co movement (see Zihl, von Cramon, & Mat, 1983) In passing, Riddoch also noted among his cases one patient who was unable to appreciate depth in or solidity of objects which he could ocherwise see quite well in his untact fields, ehus demonstraung a further selective deficit the physiological origin of which remains poorly understood The theoretical climate had changed significantly by che ime chat che aksnecopsic patient LM was studied by Zh! and colleagues in Germany (Zbl ef af , 1983) Following brain injury as a result of thrombosis, LM reported chat she could no longer see movement, making st difficult for her to cope with a number of everyday tasks including pouring drinks and crossing the road safely Testing showed that she was blind ro fast movements but retained some ability ro see slower movements of 10 deg/s or less She did not report MAES after prolonged viewing of a moving spiral, and reporced such effects only rarely, and briefly, after viewing a slowly moving pattern of stripes However, she did report seeing the spiral and che stripes move in the adaptation phase, suggesting that different processes are snvolved in adaptation and cest A similar conclusion has been applied to the MAE in normal observers (Wade, Spillmann, & Swanston, 1996): adapration 1s ¢o local motion whereas test involves global seructure Consistent with residual ability to see only certain kinds of motion, McLeod, Diterick, Driver, Perrect, and Zihl (1996) reported that LM 1s able co weneify che moving human figures in Johannson stimuli, though unable co describe how the figures are moving, and unable to see them at all af they are presented in textured nosse Copyright © 2001. All rights reserved.