Abstract
The goal of this review is to compare two divergent lines of research on signal-centered behavior: the orienting reflex (OR) and autoshaping. A review of conditioning experiments in animals and humans suggests that the novelty hypothesis of the OR is no longer tenable. Only stimuli that represent biological "relevance" elicit ORs. A stimulus may be relevant a priori (i.e., unconditioned) or as a result of conditioning. Exposure to a conditioned stimulus (CS) that predicts a positive reinforcer causes the animal to orient to it throughout conditioning. Within the CS-US interval, the initial CS-directed orienting response is followed by US-directed tendencies. Experimental evidence is shown that the development and maintenance of the conditioned OR occur in a similar fashion both in response-independent (classical) and response-dependent (instrumental) paradigms. It is proposed that the conditioned OR and the signal-directed autoshaped response are identical. Signals predicting aversive events repel the subject from the source of the CS. It is suggested that the function of the CS is not only to signal the probability of US occurrence, but also to serve as a spatial cue to guide the animal in the environment.
Full text
PDF























Selected References
These references are in PubMed. This may not be the complete list of references from this article.
- Anderson M. C., Shettleworth S. J. Behavioral adaptation to fixed-interval and fixed-time food delivery in golden hamsters. J Exp Anal Behav. 1977 Jan;27(1):33–49. doi: 10.1901/jeab.1977.27-33. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Atnip G. W. Stimulus- and response-reinforcer contingencies in autoshaping, operant, classical, and omission training procedures in rats. J Exp Anal Behav. 1977 Jul;28(1):59–69. doi: 10.1901/jeab.1977.28-59. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Badia P., Defran R. H. Orienting responses and GSR conditioning: a dilemma. Psychol Rev. 1970 May;77(3):171–181. doi: 10.1037/h0029086. [DOI] [PubMed] [Google Scholar]
- Barrera F. J. Centrifugal selection of signal-directed pecking. J Exp Anal Behav. 1974 Sep;22(2):341–355. doi: 10.1901/jeab.1974.22-341. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Bernstein A. S., Taylor K. W., Weinstein E. The phasic electrodermal response as a differentiated complex reflecting stimulus significance. Psychophysiology. 1975 Mar;12(2):158–169. doi: 10.1111/j.1469-8986.1975.tb01268.x. [DOI] [PubMed] [Google Scholar]
- Bernstein A. S. The orienting response as novelty and significance detector: reply to O'Gorman. Psychophysiology. 1979 May;16(3):263–273. doi: 10.1111/j.1469-8986.1979.tb02989.x. [DOI] [PubMed] [Google Scholar]
- Bernstein A. S. To what does the orienting response respond? Psychophysiology. 1969 Nov;6(3):338–350. doi: 10.1111/j.1469-8986.1969.tb02911.x. [DOI] [PubMed] [Google Scholar]
- Bilbrey J., Winokur S. Controls for and constraints on auto-shaping. J Exp Anal Behav. 1973 Nov;20(3):323–332. doi: 10.1901/jeab.1973.20-323. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Brown P. L., Jenkins H. M. Auto-shaping of the pigeon's key-peck. J Exp Anal Behav. 1968 Jan;11(1):1–8. doi: 10.1901/jeab.1968.11-1. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Burt J. S., Westbrook R. F. Second-order autoshaped key pecking based on an auditory stimulus. J Exp Anal Behav. 1980 Nov;34(3):305–318. doi: 10.1901/jeab.1980.34-305. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Buzsáki G., Grastyán E., Molnár P., Tveritskaya I. N., Haubenreiser J. Auto-shaping or orienting? Acta Neurobiol Exp (Wars) 1979;39(4):179–200. [PubMed] [Google Scholar]
- Buzsáki G., Grastyán E., Mód L., Winiczai Z. Importance of cue location for intact and fimbria-fornix-lesioned rats. Behav Neural Biol. 1980 Jun;29(2):176–189. doi: 10.1016/s0163-1047(80)90508-7. [DOI] [PubMed] [Google Scholar]
- Buzsáki G., Grastyán E., Winiczai Z., Mód L. Maintenance of signal directed behavior in a response dependent paradigm: a systems approach. Acta Neurobiol Exp (Wars) 1979;39(4):201–217. [PubMed] [Google Scholar]
- Buzsáki G., Haubenreiser J., Grastyán E., Czopf J., Kellényi L. Hippocampal slow wave activity during appetitive and aversive conditioning in the cat. Electroencephalogr Clin Neurophysiol. 1981 Mar;51(3):276–290. doi: 10.1016/0013-4694(81)90141-3. [DOI] [PubMed] [Google Scholar]
- DYKMAN R. A., MACK R. L., ACKERMAN P. T. THE EVALUATION OF AUTONOMIC AND MOTOR COMPONENTS OF THE NONAVOIDANCE CONDITIONED RESPONSE IN THE DOG. Psychophysiology. 1965 Jan;1:209–230. doi: 10.1111/j.1469-8986.1965.tb03238.x. [DOI] [PubMed] [Google Scholar]
- Davey G. C., Oakley D., Cleland G. G. Autoshaping in the rat: Effects of omission on the form of the response. J Exp Anal Behav. 1981 Jul;36(1):75–91. doi: 10.1901/jeab.1981.36-75. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Davol G. H., Steinhauer G. D., Lee A. The role of preliminary magazine training in acquisition of the autoshaped key peck. J Exp Anal Behav. 1977 Sep;28(2):99–106. doi: 10.1901/jeab.1977.28-99. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Dickinson A., Mackintosh N. J. Classical conditioning in animals. Annu Rev Psychol. 1978;29:587–612. doi: 10.1146/annurev.ps.29.020178.003103. [DOI] [PubMed] [Google Scholar]
- FALK J. L. Production of polydipsia in normal rats by an intermittent food schedule. Science. 1961 Jan 20;133(3447):195–196. doi: 10.1126/science.133.3447.195. [DOI] [PubMed] [Google Scholar]
- Gale E. N., Ax A. F. Long term conditioning of orienting responses. Psychophysiology. 1968 Nov;5(3):307–315. doi: 10.1111/j.1469-8986.1968.tb02827.x. [DOI] [PubMed] [Google Scholar]
- Gale E. N., Stern J. A. Classical conditioning of the peripheral vasomotor orienting response. Psychophysiology. 1968 Jan;4(3):342–348. doi: 10.1111/j.1469-8986.1968.tb02777.x. [DOI] [PubMed] [Google Scholar]
- Gale E. N., Stern J. A. Conditioning of the electrodermal orienting response. Psychophysiology. 1967 Jan;3(3):291–301. doi: 10.1111/j.1469-8986.1967.tb02709.x. [DOI] [PubMed] [Google Scholar]
- Gamzu E., Schwam E. Autoshaping and automaintenance of a key-press response in squirrel monkeys. J Exp Anal Behav. 1974 Mar;21(2):361–371. doi: 10.1901/jeab.1974.21-361. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Gamzu E., Williams D. R. Classical conditioning of a complex skeletal response. Science. 1971 Mar 5;171(3974):923–925. doi: 10.1126/science.171.3974.923. [DOI] [PubMed] [Google Scholar]
- Germana J. Response characteristics and the orienting reflex. J Exp Psychol. 1968 Dec;78(4):610–616. doi: 10.1037/h0026626. [DOI] [PubMed] [Google Scholar]
- Glickman S. E., Schiff B. B. A biological theory of reinforcement. Psychol Rev. 1967 Mar;74(2):81–109. doi: 10.1037/h0024290. [DOI] [PubMed] [Google Scholar]
- Grastyán E., Vereczkei L. Effects of spatial separation of the conditioned signal from the reinforcement: a demonstration of the conditioned character of the orienting response or the orientational character of conditioning. Behav Biol. 1974 Feb;10(2):121–146. doi: 10.1016/s0091-6773(74)91725-8. [DOI] [PubMed] [Google Scholar]
- Green L. Are there two classes of classically-conditioned responses? Pavlov J Biol Sci. 1978 Jul-Sep;13(3):154–162. doi: 10.1007/BF03001388. [DOI] [PubMed] [Google Scholar]
- Grings W. W. Orientation, conditioning, and learning. Psychophysiology. 1977 Jul;14(4):343–350. doi: 10.1111/j.1469-8986.1977.tb02962.x. [DOI] [PubMed] [Google Scholar]
- Gruzelier J. H., Venables P. H. Skin conductance responses to tones with and without attentional significance in schizophrenic and nonschizophrenic psychiatric patients. Neuropsychologia. 1973 May;11(2):221–230. doi: 10.1016/0028-3932(73)90011-0. [DOI] [PubMed] [Google Scholar]
- HUDSON B. B. One-trial learning in the domestic rat. Genet Psychol Monogr. 1950 Feb;41(FIRST):99–145. [PubMed] [Google Scholar]
- Herrnstein R. J., Loveland D. H. Food-avoidance in hungry pigeons, and other perplexities. J Exp Anal Behav. 1972 Nov;18(3):369–383. doi: 10.1901/jeab.1972.18-369. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hillyard S. A., Galambos R. Effects of stimulus and response contingencies on a surface negative slow potential shift in man. Electroencephalogr Clin Neurophysiol. 1967 Apr;22(4):297–304. doi: 10.1016/0013-4694(67)90199-x. [DOI] [PubMed] [Google Scholar]
- Holland P. C. Conditioned stimulus as a determinant of the form of the Pavlovian conditioned response. J Exp Psychol Anim Behav Process. 1977 Jan;3(1):77–104. doi: 10.1037//0097-7403.3.1.77. [DOI] [PubMed] [Google Scholar]
- Jenkins H. M., Boakes R. A. Observing stimulus sources that signal food or no food. J Exp Anal Behav. 1973 Sep;20(2):197–207. doi: 10.1901/jeab.1973.20-197. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jenkins H. M., Moore B. R. The form of the auto-shaped response with food or water reinforcers. J Exp Anal Behav. 1973 Sep;20(2):163–181. doi: 10.1901/jeab.1973.20-163. [DOI] [PMC free article] [PubMed] [Google Scholar]
- KAMIN L. J. Apparent adaptation effects in the acquisition of a conditioned emotional response. Can J Psychol. 1961 Sep;15:176–188. doi: 10.1037/h0083217. [DOI] [PubMed] [Google Scholar]
- KARMOS G., GRASTYAN E., LOSONCZY H., VERECZKEY L., GROSZ J. THE POSSIBLE ROLE OF THE HIPPOCAMPUS IN THE ORGANIZATION OF THE ORIENTATION REACTION. Acta Physiol Acad Sci Hung. 1965;26:131–141. [PubMed] [Google Scholar]
- Kovach J. K. Interaction of innate and acquired: color preferences and early exposure learning in chicks. J Comp Physiol Psychol. 1971 Jun;75(3):386–398. doi: 10.1037/h0030940. [DOI] [PubMed] [Google Scholar]
- Lawicka W. Auditory targeting reflexes: their determining role in directional instrumental responding. Acta Neurobiol Exp (Wars) 1979;39(6):537–552. [PubMed] [Google Scholar]
- Lockhart R. A. Comments regarding multiple response phenomena in long interstimulus interval conditioning. Psychophysiology. 1966 Oct;3(2):108–114. doi: 10.1111/j.1469-8986.1966.tb02687.x. [DOI] [PubMed] [Google Scholar]
- Locurto C., Terrace H. S., Gibbon J. Autoshaping, random control, and omission training in the rat. J Exp Anal Behav. 1976 Nov;26(3):451–462. doi: 10.1901/jeab.1976.26-451. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Maltzman I. Orienting reflexes and significance: a reply to O'Gorman. Psychophysiology. 1979 May;16(3):274–282. doi: 10.1111/j.1469-8986.1979.tb02990.x. [DOI] [PubMed] [Google Scholar]
- O'Gorman J. G. The orienting reflex: Novelty or significance detector? Psychophysiology. 1979 May;16(3):253–262. doi: 10.1111/j.1469-8986.1979.tb02988.x. [DOI] [PubMed] [Google Scholar]
- Ohman A. Differentiation of conditioned and orienting response components in electrodermal conditioning. Psychophysiology. 1971 Jan;8(1):7–22. doi: 10.1111/j.1469-8986.1971.tb00432.x. [DOI] [PubMed] [Google Scholar]
- Ohman A. Factor analytically derived components of orienting, defensive, and conditioned behavior in electrodermal conditioning. Psychophysiology. 1972 Mar;9(2):199–209. doi: 10.1111/j.1469-8986.1972.tb00754.x. [DOI] [PubMed] [Google Scholar]
- POLIDORA V. J., FLETCHER H. J. AN ANALYSIS OF THE IMPORTANCE OF S-R SPATIAL CONTIGUITY FOR PROFICIENT PRIMATE DISCRIMINATION PERFORMANCE. J Comp Physiol Psychol. 1964 Apr;57:224–230. doi: 10.1037/h0043928. [DOI] [PubMed] [Google Scholar]
- POLIDORA V. J., THOMPSON W. J. ORIENTING BEHAVIOR AND THE S-R SPATIAL DISCONTIGUITY EFFECT IN MONKEYS. J Comp Physiol Psychol. 1965 Apr;59:240–245. doi: 10.1037/h0021842. [DOI] [PubMed] [Google Scholar]
- Peterson G. B., Ackilt J. E., Frommer G. P., Hearst E. S. Conditioned Approach and Contact Behavior toward Signals for Food or Brain-Stimulation Reinforcement. Science. 1972 Sep 15;177(4053):1009–1011. doi: 10.1126/science.177.4053.1009. [DOI] [PubMed] [Google Scholar]
- Powell R. W., Kelly W. Responding under positive and negative response contingencies in pigeons and crows. J Exp Anal Behav. 1976 Mar;25(2):219–225. doi: 10.1901/jeab.1976.25-219. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Pribram K. H., McGuinness D. Arousal, activation, and effort in the control of attention. Psychol Rev. 1975 Mar;82(2):116–149. doi: 10.1037/h0076780. [DOI] [PubMed] [Google Scholar]
- RAZRAN G. The observable unconscious and the inferable conscious in current Soviet psychophysiology: interoceptive conditioning, semantic conditioning, and the orienting reflex. Psychol Rev. 1961 Mar;68:1–147. [PubMed] [Google Scholar]
- Ray R. D., Brown D. A. The behavioral specificity of stimulation: a systems approach to procedural distinctions of classical and instrumental conditioning. Pavlov J Biol Sci. 1976 Jan-Mar;11(1):3–23. doi: 10.1007/BF03000534. [DOI] [PubMed] [Google Scholar]
- Reberg D., Mann B., Innis N. K. Superstitious behavior for food and water in the rat. Physiol Behav. 1977 Dec;19(6):803–806. doi: 10.1016/0031-9384(77)90318-3. [DOI] [PubMed] [Google Scholar]
- Rescorla R. A. Pavlovian conditioning and its proper control procedures. Psychol Rev. 1967 Jan;74(1):71–80. doi: 10.1037/h0024109. [DOI] [PubMed] [Google Scholar]
- Rescorla R. A., Solomon R. L. Two-process learning theory: Relationships between Pavlovian conditioning and instrumental learning. Psychol Rev. 1967 May;74(3):151–182. doi: 10.1037/h0024475. [DOI] [PubMed] [Google Scholar]
- SIMPSON G. G. The status of the study of organisms. Am Sci. 1962 Mar;50:36–45. [PubMed] [Google Scholar]
- Schneider G. E. Two visual systems. Science. 1969 Feb 28;163(3870):895–902. doi: 10.1126/science.163.3870.895. [DOI] [PubMed] [Google Scholar]
- Schwartz B. Maintenance of key pecking by response-independent food presentation: the role of the modality of the signal for food. J Exp Anal Behav. 1973 Jul;20(1):17–22. doi: 10.1901/jeab.1973.20-17. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Schwartz B., Williams D. R. The role of the response-reinforcer contingency in negative automaintenance. J Exp Anal Behav. 1972 May;17(3):351–357. doi: 10.1901/jeab.1972.17-351. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Sheafor P. J. "Pseudoconditioned" jaw movements of the rabbit reflect associations conditioned to contextual background cues. J Exp Psychol Anim Behav Process. 1975 Jul;1(3):245–260. doi: 10.1037//0097-7403.1.3.245. [DOI] [PubMed] [Google Scholar]
- Shettleworth S. J. Reinforcement and the organization of behavior in golden hamsters: Pavlovian conditioning with food and shock unconditioned stimuli. J Exp Psychol Anim Behav Process. 1978 Apr;4(2):152–169. doi: 10.1037//0097-7403.4.2.152. [DOI] [PubMed] [Google Scholar]
- Siddle D. A., O'Gorman J. G., Wood L. Effects of electrodermal lability and stimulus significance on electrodermal response amplitude to stimulus change. Psychophysiology. 1979 Nov;16(6):520–527. doi: 10.1111/j.1469-8986.1979.tb01514.x. [DOI] [PubMed] [Google Scholar]
- Stepień I. The magnet reaction, a symptom of prefrontal ablation. Acta Neurobiol Exp (Wars) 1974;34(1):145–160. [PubMed] [Google Scholar]
- Stiers M., Silberberg A. Lever-contact responses in rats: automaintenance with and without a negative response-reinforcer dependency. J Exp Anal Behav. 1974 Nov;22(3):497–506. doi: 10.1901/jeab.1974.22-497. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Testa T. J. Effects of similarity of location and temporal intensity pattern of conditioned and unconditioned stimuli on the acquisition of conditioned suppresion in rats. J Exp Psychol Anim Behav Process. 1975 Apr;1(2):114–121. doi: 10.1037//0097-7403.1.2.114. [DOI] [PubMed] [Google Scholar]
- Thompson R. F. The search for the engram. Am Psychol. 1976 Mar;31(3):209–227. doi: 10.1037//0003-066x.31.3.209. [DOI] [PubMed] [Google Scholar]
- Tinbergen N. On war and peace in animals and man. An ethologist's approach to the biology of aggression. Science. 1968 Jun 28;160(3835):1411–1418. doi: 10.1126/science.160.3835.1411. [DOI] [PubMed] [Google Scholar]
- Tomie A. Retardation of autoshaping: control by contextual stimuli. Science. 1976 Jun 18;192(4245):1244–1246. doi: 10.1126/science.192.4245.1244. [DOI] [PubMed] [Google Scholar]
- Wasserman E. A. Pavlovian conditioning with heat reinforcement produces stimulus-directed pecking in chicks. Science. 1973 Aug 31;181(4102):875–877. doi: 10.1126/science.181.4102.875. [DOI] [PubMed] [Google Scholar]
- Wesp R. K., Lattal K. A., Poling A. D. Punishment of autoshaped key-peck responses of pigeons. J Exp Anal Behav. 1977 May;27(3):407–418. doi: 10.1901/jeab.1977.27-407. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Wessells M. G. The effects of reinforcement upon the prepecking behaviors of pigeons in the autoshaping experiment. J Exp Anal Behav. 1974 Jan;21(1):125–144. doi: 10.1901/jeab.1974.21-125. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Whittington D. A., Hepp-Reymond M. C., Flood W. Eye and head movements to auditory targets. Exp Brain Res. 1981;41(3-4):358–363. doi: 10.1007/BF00238893. [DOI] [PubMed] [Google Scholar]
- Wilcoxon H. C., Dragoin W. B., Kral P. A. Illness-induced aversions in rat and quail: relative salience of visual and gustatory cues. Science. 1971 Feb 26;171(3973):826–828. doi: 10.1126/science.171.3973.826. [DOI] [PubMed] [Google Scholar]
- Williams D. R., Williams H. Auto-maintenance in the pigeon: sustained pecking despite contingent non-reinforcement. J Exp Anal Behav. 1969 Jul;12(4):511–520. doi: 10.1901/jeab.1969.12-511. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Woodard W. T., Ballinger J. C., Bitterman M. E. Autoshaping: further study of "negative automaintenance". J Exp Anal Behav. 1974 Jul;22(1):47–51. doi: 10.1901/jeab.1974.22-47. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Woodruff G., Williams D. R. The associative relation underlying autoshaping in the pigeon. J Exp Anal Behav. 1976 Jul;26(1):1–13. doi: 10.1901/jeab.1976.26-1. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Zentall T. R., Hogan D. E. Key pecking in pigeons produced by pairing keylight with inaccessible grain. J Exp Anal Behav. 1975 Mar;23(2):199–206. doi: 10.1901/jeab.1975.23-199. [DOI] [PMC free article] [PubMed] [Google Scholar]