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Anticipatory Goal-Directed Behavior, Limbic System and Endocrine Influences
AN'l'ICIPATORY GOAL-DIREC'I'ED GOAL-DIRECTED BEHAVIOR, LII.illIC LnillIC SYSTEM AND ENDOCRINE INFLUENCES.
E. ENDROCZI, M.D. ,D.Sc. Professor Department of Experimental and Clinical Laboratory Investigations Postgraduate Iliedical Illedical School Budapest, Hungary
A correct performance of the living oreanism may be described as a series of specific responses under control of a series of the situational stimuli. It is known that stimuli produced by responses, regardless of modalities, take a part to correct response performance. The theorist of closed-loop mechanism regards the performer as information processor who compares the response induced sensory feed-back with a stored representation of what feedback from correct response should be. This closed-loop model is known in engineering and was proposed by numerous investigators for animal behavior / Boring /1/, Miller et al.,/2/, Luria /3/, Adams,/4/, Deutsch /5/, Anokhin /6J etc./. Conclusions derived from behavioral studies about the role of a closed-loop mechanism in controlling the animal behavior have sugg~sted sugg~sted that it may be involved in the establlshment of conditioned reflex in which responses are corrected by feed-back stimuli from erroneous responses. Hull /7/ was the first who suggested the a foresight of the performer is essential for performance of a sequence of the responses and the sensory feed-back of a past response is the mediator of an anticipatory goal-directed behavior. The principle that the perceptual image of a past response plays a role in resp?nse resp~nse initiation, ini tiation, was already suggested ln the nineteenth century / Lotze /8/, / 8/, Jarnes James /9/ / 9/ /. The influential behavior theorists rejected rej ected the "ideo-motor" formulation of learning behavior for many decades but it was revived by many neurophysiologists neurophysiolo gists and experimental psychologists in recent decades / Kowrer I.Iowrer /10/, Konorski /11/, Hebb /12/, Beritashvili Beri tashvili /13/, Vandervlolf Vanderv:olf /14/, etc./. Unlike the closed-loop formulation, the "i "ideo-motor" deo-motor" concept regards re gards the necessity of a response image in both response initiation and selection an~ in controlling c ontrolling the performance / Lumsdalne /15/, Deutsch /5/, Bandura /16/, Greenwald and Albert /17/, etc./. From biological point of view, the "ideo-motor" concept
588
may be re regarded garded as a "response-program" with a repertory of anticipated sensory feed-back from past responses; the proepl'Ogram may be initiated by a fragment of the situational stimuli or by changes in the internal environment which produce needs to restore the homeostatic equilibrium through this response. Homeostasis and goal-directed activity. Changes either in the external-orthe internal environment may result in a need of the organism to restore the homeostatic eouilibrium and such drive induc ind~ced ed goal-directed goal -directed response respons e may be innate inna te or acquired. The drives as initiators of goaldirected responses can be divided into to subclasses: 1. evoked by more or less specific changes of chemical environment and manifest themselves by genetically coded responses / e.g. feeding feeding,, sexual, maternal, nest building behavioral reactions/. 2. The drives evoked by noxious stimuli which interfere with the maintenance of homeostasis and lead to response initiation for elimination of such conflicting confli c ting influences. There are at least three main groups of concepts c once pts which have been proposed as possible alternatives alternati ves to drive: these t h ese are arousal, incentive and rei:uorcewcnt. rei:uorcer:;ent. All three thre e seem capable of dealing ~ith ~i t h the effects of conditions c ondit ions of bodily need on behavior, but none assigns the central ioportance i oport ance to vegetative and a nd neuroendocrine processes proces ses they t hey held in drive theory t heory / Bolles /18/, Bower et al.,/19/, Logan /20/, / 20 /, Prenack Premack /21/ Seward /22/, etc./. IItt is knov:n that a re~ponse will be repeated, in the absence of liniting limiting factors, f ac tors, as long as the t he drive persists. A perseveration tendency of the response reSDonse seems to be an intrinsic feature of" of· the central representation and only sensory feed-back inhibition inhi bi tion provides a "time Dattern" pattern" for the "response program". Drive arousal and activation act iva t ion of a response image may be regarded re garded as coherent functions of central representation. representat ion. They are badly defined terms and their meaning ~ere ~ ere frequently revised and criticized by
the neurophysiologists and experimental r,loreover, there are diffepsychologists. l,loreover, rent forms of arousal I/ electrocortical, autonomic and behavioral I/ and sometimes somet imes it is difficult to use one form of arousal v alid index of another I/ Lacey,/23/, Lacey ,/ 23 /, as a valid Malmo and Belanger Bel anger 1241, / 24/, Berlyne 125/, /25/, I.ialmo etc ./. etc.l. Limbic system and goal-directed goal - direc ted behavior. The appearance of synchronized slow waves in the hippocampus and their spreading to the brainstem and diencephadiencephalic l i c structures during exploration and goal - directed response have bee~ been observed goal-directed by many investigators I/ see reviews rev~ews by /26/, Adey 127/, /27/, Routtenberg 128/, / 28/ , Green 126/, etc.l. etc ./. The hippocampal ablation ablat ion results in an increased response perseveration as a result re sult of the decreased ability of the subject to withdraw responding. Ani-. Ani -. mals Vi with septal-hippocampal exh~ i th septal - hippocampal damages exhlbit both an increased exploratory activity and a decreased habituation to novelty /I Routtenberg 128/, /28/, Vanderwolf 114~ /14~ Kimble 129/, /29/, etc.l. etc ./. Kimble /I 29/ 291 and Endroczi Endroczi 1301 m~~alian hippo/30/ have proposed that mammalian hippo campus constitutes a part of the ne~ral ne~ral machinery necessary for the generatlon generat~on of those brain processes which are functiofunctio nally equivalent to Pavlovian internal inhibition. hibition . Obviously, Obviously , an impairment of the response res ponse selection and the discrimination reversal as result of hippocampal ablation comprises a deficit of the error evaluevaluation and a failure of the decoding of the informations from sensory feed-back I/ see the reviews by Douglas /I 31/ 311,, Kimble 129/, /29/, Endroczi and Fekete 132/, /32/, Lissak and Endroczi 1331, r:ilner 135/, /33/, IIcCleary lvIcCleary 1341, /34/, mlner / 35/, Vanderwolf 114/, /14/, etc.l. etc ./. Goal-directed Goal - directed motor patterns may be regarded as sources of reinforcement which fund&~entally involved in both v/hich is fundaJllentally res ponse increment and decrement of the response tendency and perseveration. ins 1361 perseveration . Perk Perkins /36/ proposed that that "positive reinforcers" both reinforce antecedent re responses sponses and transtrans mit secondary reinforcement properties to antecedent stimuli. stimuli . This formulation of reinforcement does not take into account reinforce~ent that the repetition repetit ion of a goal response is follov!ed by a decrer:lent decrement of the responding, responding , on the one hand, follovled by a hand , and it is followed decrease of the drive or arousal level, level , hand . According to our our stustuon the other hand. dies on the nature of reinforcing value of sensorv feed-back feed - back each reinforcement r einforcement comcomprises"both positive and negative value in relation to the antecedent response. follo~ed A repetition re petition of the response is followed by an increase of the intensity of the. the . negative reinforcing value and results ln ~n a decrenent decrement of the responding. responding . An increase h~p~o of the response perseveration after h~p~o cam ectomy may be attributed to a deflclt def~c~t campectomy in decoding of reinforcing value from
589
/30//. On sensory feed-back /I Endroczi 13011. stimUlation studies the other hand, the stimulation revealed that the appearance of synchroow waves in the hippocampus hippo c aJIlPU S is nized sl slow accompanied by the increase of internal inhibition which whi ch is important i mportant for both response selection and extinction I/ see tenberg 128/,Douglas /28/,Dougl as the reviews by Rout Routtenberg /31/, Endroczi 1301, / 30/, etc./. 131/, The role r ole of hippocampus hippocaJIlPUS in neuronal activities activiti es underlying the consolidation of memory traces is striking st riking in huhu Milner and Penfield /3 6/, Penfield mans /I l.1ilner 1361, Milner 13711.Following /37// .Foll owing bilateral leand 1lilner hippocampus , the papasions invading the hippocampus, tients show a deficit to transit memory traces from short-term short -t erm to long-term stosto rage. A sensory feed-back induced inhibition of the response initiation is necess8XY to shift the attention to one or more 8Xy conseouential environmental events and this process is important important for memory consolidation I/ see the reviews by Walke Walker r 138/, /38/, Glickman 139/, / 39/ , McGaugh 1401,etc.l. /4 0/, etc ./. An integrity i ntegri t y of the neuronal circuit between hippocampus and brainstem is i s required for integration of internal inhibitory processes and memory consolidation. consolidation . Limbic system and anticipated behavior. behavior . In studying the appearance of hippocampal theta rhythm during passive passi ve avoidance avo idance learning Vie Vi e have found that it could be observed before the motor perperformance I/ Endroczi,/301. Endroczi ,/ 30/. In further experiments by the t he use of intermittent and unsignalled water reward reVlard for one minute in every hour we have found that the waves appear 5 to 10 min. hippocampal slow ,,,aves min . prior to the wat water er administration after 20 to 30 drinking sessions. sessions . It is worth to mentioning that the appearance of the hippocampal hippocampal theta rhythm r hy thm was accompanied by searching activity which may be regarded as a sign of the expectation or . of the "time conditioning". conditioning" . These behav~ behavl oral observations led to assume that the appearance of hippocampal theta rhythm is an electrical correlate of the response initiation. initiation . This formulation is in accordance with vd th the view of Vanderwolf 114/ /14/ that the appearance of synchronized slov, waves is correlated to voluntary slow activities. activities . Limbic Limb ic system and hormonal conditioning . Rostral and basal limbic strucstruc ditioning. tures play an important modifying role in controlling the neuroendocrine processes I/ see the reviews by Harris and Donovan 141/, /41/, Sawyer and Gorski 142/, /42/, etc.l. etc ./. This part of the brain is a specific target organ for hormones and other chemical messages of the body body.. Thus, Thus , the hippohippo campal cells have a property to accumuaccumulate l a te steroid hormones selectively and in a greater quantity quant ity than other brain regions. r:oreover, regions . Mo reover, the electrophysioloelectrophysiological gic al studies revealed that pituitary
hormones exert an influence on the hippocampus and the amygdaloid complex of nuclei which is not mediated via peripheral target organs I see the reviews by Kawakami et al. 143/, 1431, Sawyer and Gorski/4211. Moreover, the recent observations indicated that the steroid hormones are accumulated in the cells of the dentate gyrus which constitute an essential part to the sensory input of pyramidal cell layer. Studying the effect of pituitaryadrenal hormones on exploratory activity, response initiation and selection in both active and passive avoidance situations, we have found that the hippocampus-septal -diencephalic complex plays a basic role in the corticosteroid-adjusted behavior of rats,cats and dogs I see the reviews by Lissak and Endroczi 1331 and Endroczi 13011. These findings may be summarized as follows: 1. The corticosterone induced suppression of exploratory activity is absent in hippocampectomized rats. 2. Corticosterone administration produced an increase of the extinction of approaching response which could not be observed after lesioning the septal-hippocampal system in cats and dogs. 3. In a conflict situation I e.g. hunger vs. fearl the corticosteroid treatment evoked neurotic behavior and paradox conditioned responses in dogs. This unbalanced state in controling the response initiation and selection may be attributed to a failure of the comparator function of the limbic system for decoding the informations of sensory feedback. With regard that the pituitaryadrenal function is activated by nonspecific stimuli of either internal or external environment, the adrenal cortical hormones cannot be designated as specific humoral factors to conditioning of either innate or acquired acqUired behavioral patterns. This is in contrast to the effect of sex steroids or prolactin which are primarily involved in the integration of such behavioral responses as sexual or maternal, respectively. The adrenal cortical steroids produce changes in the excitatory state of the central nervous system which manifest by a facilitation of the internal inhibitory processes necessary for decoding of sensory feedback and for integration of an adequate anticipated response initiation. The central nervous system is considered to arrange a transformation from the "shape-time-space" to a neural network space. This transformation includes signal recoenition, continuous comparison of arriving signals with innate or acquired ones, development of short-term memory loops and transition of short-term memory to a permanent storage. Recently, experiments were performed to study the time mer.lOry transition in rats. pattern of memory
590
After a 5-day habituation period to the experimental environment in which the animals were provided with water after 23-hour water deprivation, each drinking attempt was associated with electrical shocks on the 6th day. The animals were removed from this conflict situation in the 10, 60, 120, 180 and 300th seconds after the third drinking attempt and the latency of the first drinking was tested on the next day. With Vlith regard that the intervals between subsequent drinking attempts showed an exponential increase, the fourth could be e::pected about 320 to 400 seconds. It was found that the shortest latency of first responding was shown by the rats which were removed at 10 and 300 sec, and the longest latency by those which were removed near to the middle of the subsequent intertrial interval. 'rhis This inverse U-shape increase and decrement of the response inhibition was markedly impaired in rats with septal-hippocampal damage. These observations led to assume that the development of short-term memory loops requires the integrity of limbic system, on the one hand, and that the transition of short-term memory to a longterm form is impaired in hippocampectornihippocampectomized rats. This function of the limbic system is under a partial control of hwnoral influences; both hormones and drugs given in the transition time may produce changes in the memory function. REFEREliJCES REFEREIiJCES 1. Boring,E.C.: Sensation and perceEtion in the histor of ex erimental s c 010 tew or: App e on- en ury- ro , 2. Lliller,G.A. ,Galanter,E. ,Pribram,K.H.: Plans and the structure of behavior. New York: Holt,Rinehart and Winston,1960. 3. Luria,A.R.: Higher corticGl functions in man. New York:Basic Eooks,196l. 4. Adams,J.A.: Psychol. Bull., 70:486, 1968. 5. Deutsch,J.A.: Brit.J.Psychol., 47:115, 1956. J.F.Delufresnaye IEdl 6. Anokhin,P.K.: In J.F.Delafresnaye SprincErain rr:echa!'1isms and learning. Sprinc:,Charles C '~\!lomas,1961. '~\1l0mas,1961. field, Ill. Ill.,Charles Hull ,C .L.: 'The The Drinciple Drinciple of behavior. behe,vior. 7. Hull,C.L.: York:;"ppleton-Century, 1943. New York:Appleton-Century, ~edizinische Psychologie. 8. Lotze,R.H.: ~edizinische Leip zig :Yieidemann. Leipzig:Weidemann. J2nes, VI.: 9. Jenes, Vi.: Principles of gs"chOl2£:L' vol.2. New York:Holt,189 .t Lear!'1ing theory ond be10. r.:oVirer,O.E.: Lear:1ing hayior. Eew Kew York:Wiley, 1960. 11. Koncrski,J.: Integrative activity of the brain. Chicago Univ. Press,1967. 12. Hebb,D.O.: Concernin~ Concernin~ icagery icagery.. 75:4b6 ,1968. Psychol. Bull., 75:4b6,1968. 13. Beritashvili,I.S.: In: ~.Cole,I.~altz ~.Cole,I.~altz IEds/, A handbook of cont errJDorary eUJDorary man IEdsl,A psychOlogy. I:eVl York :Basic Books. Book s . soviet psychology. 1969.
14. Vanderwolf,C.H.: Vanderwo1f,C.H.: Psychol.Rev.,j§:83, Psycho1.Rev.,j§:83, 43. Kawakami,M. et al.: In C.H.Sawyer, 1971. R.Gorski /Eds/ IEds/ Steroid hormones and brain 15. Lumsdaine,A.A.: In Symposium Symp osium on expefunction. UCLA Forum Medical Sciences, rimental studies of .ue and response facN.15,1972. tors in group and individual learning. Washington:Nat.Acad.Sci.Res.Council,196l. Washington:N at .Acad.Sci .Res .Counci1 ,1961. 16. Bandura,A.: Principles of behavioral modification. New York: Holt,Rlnehart Ho1t,Rlneliart and Wlnston, 1965. 17. Greenwald,A.G.,Albert,S.M.: Greenwa1d,A.G. ,A1bert ,S.~.1.: J. exp. expo Psychol., Psycho1., 76:267,1968. 18. Bolles:R.C.: Bol1es:R.C.: Theor~ Theor~ of motivation. New York:Harper,Row, 1 67. 19. Bower,G. ,r.rcLean,J. ,r.rcLean, J. ,!,!eachem,J.: ,I,leachem,J.: J.comp. Physiol.Psychol.,62:184,1966. Physiol.Psycho1.,62:184,1966. 20. Logan,F.A.: Logan ,F.A.: Incentive. Yale Univ.PresB Univ.Press 1960. 21. Premack,D.: Nebraska Prelllack,D.: In D.Levine lEd! I Ed/ Ne braska moti vation. Univ. Nebraska Symposium on motivation. Press, 1965,p.123. 22. Seward,J.P.: Psychol. Psycho1. Rev., 22:195, 1956. N.Y.Acad.Sci.,£1: 23. Lacey,J.I.: Ann. N.Y.Acad.Sci.,21: 123, 1956. Malmo ,R. B. , Belanger, D.: In Sleep and 24. Malmo,R.B.,Belanger,D.: cons ciousness. Assoc. altered states of consciousness. 228,1 96 7. Res.Nerv.Ment.Dis., .,42: 32:228,1967. 25. Berlyne,D.E.: In D.Levine IEdl Nebrasi.lot ivation. Univ. Nebraska Symposium on iJotivation. ka Press, 11967,1-110. 967 ,1-110. 26. Green,J.D.: Physiol.Rev.,~:561,1964. Physiol.Rev.,~:561,1964. 27. Adey,W.R.: W.R.Adey and T.Tokizane IIEdsl Eds / Progress in brain research: structure and function of the limbic system. Amsterdam:Elsevier,1967. 28. Routtenberg,A.: Psychol.Rev., Psycho1.Rev., ]2: 51,1968. 29. Kimble,D.O.: Psychol.Bull.,lQ: Psychol.Bull.,1Q: 28 5,1968. 30. Endroczi,E.: Limbic system,Iearning and ~ituitary-adrenal ~ituitary-adrenal functl0n. Akaae~a Akaae~a Kiado,Budapest,1972. Dougl as ,R.J.: Psychol.Bull.,67:416, Psyc ho1.Bull.,67:416, 31. Douglas,R.J.: 19b7. K.Lissak IEdl 32. Endroczi,E.,Fekete,T.: In K.Lissik Recent Prog. Brain Res. Res . in Hungary, vol.3, 1972. Lissak ,K. ,Endroczi,E.: The neuroendo33. Lissak,K.,Endroczi,E.: crine control of adaptation. Pergamon Press OX! ord,I965. ord,1965. Physiol.Psycho1., 34. !
t
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591
E. ENDROCZI, M.D. ,D.Sc. Professor Department of Experimental and Clinical Laboratory Investigations Postgraduate Iliedical Illedical School Budapest, Hungary
A correct performance of the living oreanism may be described as a series of specific responses under control of a series of the situational stimuli. It is known that stimuli produced by responses, regardless of modalities, take a part to correct response performance. The theorist of closed-loop mechanism regards the performer as information processor who compares the response induced sensory feed-back with a stored representation of what feedback from correct response should be. This closed-loop model is known in engineering and was proposed by numerous investigators for animal behavior / Boring /1/, Miller et al.,/2/, Luria /3/, Adams,/4/, Deutsch /5/, Anokhin /6J etc./. Conclusions derived from behavioral studies about the role of a closed-loop mechanism in controlling the animal behavior have sugg~sted sugg~sted that it may be involved in the establlshment of conditioned reflex in which responses are corrected by feed-back stimuli from erroneous responses. Hull /7/ was the first who suggested the a foresight of the performer is essential for performance of a sequence of the responses and the sensory feed-back of a past response is the mediator of an anticipatory goal-directed behavior. The principle that the perceptual image of a past response plays a role in resp?nse resp~nse initiation, ini tiation, was already suggested ln the nineteenth century / Lotze /8/, / 8/, Jarnes James /9/ / 9/ /. The influential behavior theorists rejected rej ected the "ideo-motor" formulation of learning behavior for many decades but it was revived by many neurophysiologists neurophysiolo gists and experimental psychologists in recent decades / Kowrer I.Iowrer /10/, Konorski /11/, Hebb /12/, Beritashvili Beri tashvili /13/, Vandervlolf Vanderv:olf /14/, etc./. Unlike the closed-loop formulation, the "i "ideo-motor" deo-motor" concept regards re gards the necessity of a response image in both response initiation and selection an~ in controlling c ontrolling the performance / Lumsdalne /15/, Deutsch /5/, Bandura /16/, Greenwald and Albert /17/, etc./. From biological point of view, the "ideo-motor" concept
588
may be re regarded garded as a "response-program" with a repertory of anticipated sensory feed-back from past responses; the proepl'Ogram may be initiated by a fragment of the situational stimuli or by changes in the internal environment which produce needs to restore the homeostatic equilibrium through this response. Homeostasis and goal-directed activity. Changes either in the external-orthe internal environment may result in a need of the organism to restore the homeostatic eouilibrium and such drive induc ind~ced ed goal-directed goal -directed response respons e may be innate inna te or acquired. The drives as initiators of goaldirected responses can be divided into to subclasses: 1. evoked by more or less specific changes of chemical environment and manifest themselves by genetically coded responses / e.g. feeding feeding,, sexual, maternal, nest building behavioral reactions/. 2. The drives evoked by noxious stimuli which interfere with the maintenance of homeostasis and lead to response initiation for elimination of such conflicting confli c ting influences. There are at least three main groups of concepts c once pts which have been proposed as possible alternatives alternati ves to drive: these t h ese are arousal, incentive and rei:uorcewcnt. rei:uorcer:;ent. All three thre e seem capable of dealing ~ith ~i t h the effects of conditions c ondit ions of bodily need on behavior, but none assigns the central ioportance i oport ance to vegetative and a nd neuroendocrine processes proces ses they t hey held in drive theory t heory / Bolles /18/, Bower et al.,/19/, Logan /20/, / 20 /, Prenack Premack /21/ Seward /22/, etc./. IItt is knov:n that a re~ponse will be repeated, in the absence of liniting limiting factors, f ac tors, as long as the t he drive persists. A perseveration tendency of the response reSDonse seems to be an intrinsic feature of" of· the central representation and only sensory feed-back inhibition inhi bi tion provides a "time Dattern" pattern" for the "response program". Drive arousal and activation act iva t ion of a response image may be regarded re garded as coherent functions of central representation. representat ion. They are badly defined terms and their meaning ~ere ~ ere frequently revised and criticized by
the neurophysiologists and experimental r,loreover, there are diffepsychologists. l,loreover, rent forms of arousal I/ electrocortical, autonomic and behavioral I/ and sometimes somet imes it is difficult to use one form of arousal v alid index of another I/ Lacey,/23/, Lacey ,/ 23 /, as a valid Malmo and Belanger Bel anger 1241, / 24/, Berlyne 125/, /25/, I.ialmo etc ./. etc.l. Limbic system and goal-directed goal - direc ted behavior. The appearance of synchronized slow waves in the hippocampus and their spreading to the brainstem and diencephadiencephalic l i c structures during exploration and goal - directed response have bee~ been observed goal-directed by many investigators I/ see reviews rev~ews by /26/, Adey 127/, /27/, Routtenberg 128/, / 28/ , Green 126/, etc.l. etc ./. The hippocampal ablation ablat ion results in an increased response perseveration as a result re sult of the decreased ability of the subject to withdraw responding. Ani-. Ani -. mals Vi with septal-hippocampal exh~ i th septal - hippocampal damages exhlbit both an increased exploratory activity and a decreased habituation to novelty /I Routtenberg 128/, /28/, Vanderwolf 114~ /14~ Kimble 129/, /29/, etc.l. etc ./. Kimble /I 29/ 291 and Endroczi Endroczi 1301 m~~alian hippo/30/ have proposed that mammalian hippo campus constitutes a part of the ne~ral ne~ral machinery necessary for the generatlon generat~on of those brain processes which are functiofunctio nally equivalent to Pavlovian internal inhibition. hibition . Obviously, Obviously , an impairment of the response res ponse selection and the discrimination reversal as result of hippocampal ablation comprises a deficit of the error evaluevaluation and a failure of the decoding of the informations from sensory feed-back I/ see the reviews by Douglas /I 31/ 311,, Kimble 129/, /29/, Endroczi and Fekete 132/, /32/, Lissak and Endroczi 1331, r:ilner 135/, /33/, IIcCleary lvIcCleary 1341, /34/, mlner / 35/, Vanderwolf 114/, /14/, etc.l. etc ./. Goal-directed Goal - directed motor patterns may be regarded as sources of reinforcement which fund&~entally involved in both v/hich is fundaJllentally res ponse increment and decrement of the response tendency and perseveration. ins 1361 perseveration . Perk Perkins /36/ proposed that that "positive reinforcers" both reinforce antecedent re responses sponses and transtrans mit secondary reinforcement properties to antecedent stimuli. stimuli . This formulation of reinforcement does not take into account reinforce~ent that the repetition repetit ion of a goal response is follov!ed by a decrer:lent decrement of the responding, responding , on the one hand, follovled by a hand , and it is followed decrease of the drive or arousal level, level , hand . According to our our stustuon the other hand. dies on the nature of reinforcing value of sensorv feed-back feed - back each reinforcement r einforcement comcomprises"both positive and negative value in relation to the antecedent response. follo~ed A repetition re petition of the response is followed by an increase of the intensity of the. the . negative reinforcing value and results ln ~n a decrenent decrement of the responding. responding . An increase h~p~o of the response perseveration after h~p~o cam ectomy may be attributed to a deflclt def~c~t campectomy in decoding of reinforcing value from
589
/30//. On sensory feed-back /I Endroczi 13011. stimUlation studies the other hand, the stimulation revealed that the appearance of synchroow waves in the hippocampus hippo c aJIlPU S is nized sl slow accompanied by the increase of internal inhibition which whi ch is important i mportant for both response selection and extinction I/ see tenberg 128/,Douglas /28/,Dougl as the reviews by Rout Routtenberg /31/, Endroczi 1301, / 30/, etc./. 131/, The role r ole of hippocampus hippocaJIlPUS in neuronal activities activiti es underlying the consolidation of memory traces is striking st riking in huhu Milner and Penfield /3 6/, Penfield mans /I l.1ilner 1361, Milner 13711.Following /37// .Foll owing bilateral leand 1lilner hippocampus , the papasions invading the hippocampus, tients show a deficit to transit memory traces from short-term short -t erm to long-term stosto rage. A sensory feed-back induced inhibition of the response initiation is necess8XY to shift the attention to one or more 8Xy conseouential environmental events and this process is important important for memory consolidation I/ see the reviews by Walke Walker r 138/, /38/, Glickman 139/, / 39/ , McGaugh 1401,etc.l. /4 0/, etc ./. An integrity i ntegri t y of the neuronal circuit between hippocampus and brainstem is i s required for integration of internal inhibitory processes and memory consolidation. consolidation . Limbic system and anticipated behavior. behavior . In studying the appearance of hippocampal theta rhythm during passive passi ve avoidance avo idance learning Vie Vi e have found that it could be observed before the motor perperformance I/ Endroczi,/301. Endroczi ,/ 30/. In further experiments by the t he use of intermittent and unsignalled water reward reVlard for one minute in every hour we have found that the waves appear 5 to 10 min. hippocampal slow ,,,aves min . prior to the wat water er administration after 20 to 30 drinking sessions. sessions . It is worth to mentioning that the appearance of the hippocampal hippocampal theta rhythm r hy thm was accompanied by searching activity which may be regarded as a sign of the expectation or . of the "time conditioning". conditioning" . These behav~ behavl oral observations led to assume that the appearance of hippocampal theta rhythm is an electrical correlate of the response initiation. initiation . This formulation is in accordance with vd th the view of Vanderwolf 114/ /14/ that the appearance of synchronized slov, waves is correlated to voluntary slow activities. activities . Limbic Limb ic system and hormonal conditioning . Rostral and basal limbic strucstruc ditioning. tures play an important modifying role in controlling the neuroendocrine processes I/ see the reviews by Harris and Donovan 141/, /41/, Sawyer and Gorski 142/, /42/, etc.l. etc ./. This part of the brain is a specific target organ for hormones and other chemical messages of the body body.. Thus, Thus , the hippohippo campal cells have a property to accumuaccumulate l a te steroid hormones selectively and in a greater quantity quant ity than other brain regions. r:oreover, regions . Mo reover, the electrophysioloelectrophysiological gic al studies revealed that pituitary
hormones exert an influence on the hippocampus and the amygdaloid complex of nuclei which is not mediated via peripheral target organs I see the reviews by Kawakami et al. 143/, 1431, Sawyer and Gorski/4211. Moreover, the recent observations indicated that the steroid hormones are accumulated in the cells of the dentate gyrus which constitute an essential part to the sensory input of pyramidal cell layer. Studying the effect of pituitaryadrenal hormones on exploratory activity, response initiation and selection in both active and passive avoidance situations, we have found that the hippocampus-septal -diencephalic complex plays a basic role in the corticosteroid-adjusted behavior of rats,cats and dogs I see the reviews by Lissak and Endroczi 1331 and Endroczi 13011. These findings may be summarized as follows: 1. The corticosterone induced suppression of exploratory activity is absent in hippocampectomized rats. 2. Corticosterone administration produced an increase of the extinction of approaching response which could not be observed after lesioning the septal-hippocampal system in cats and dogs. 3. In a conflict situation I e.g. hunger vs. fearl the corticosteroid treatment evoked neurotic behavior and paradox conditioned responses in dogs. This unbalanced state in controling the response initiation and selection may be attributed to a failure of the comparator function of the limbic system for decoding the informations of sensory feedback. With regard that the pituitaryadrenal function is activated by nonspecific stimuli of either internal or external environment, the adrenal cortical hormones cannot be designated as specific humoral factors to conditioning of either innate or acquired acqUired behavioral patterns. This is in contrast to the effect of sex steroids or prolactin which are primarily involved in the integration of such behavioral responses as sexual or maternal, respectively. The adrenal cortical steroids produce changes in the excitatory state of the central nervous system which manifest by a facilitation of the internal inhibitory processes necessary for decoding of sensory feedback and for integration of an adequate anticipated response initiation. The central nervous system is considered to arrange a transformation from the "shape-time-space" to a neural network space. This transformation includes signal recoenition, continuous comparison of arriving signals with innate or acquired ones, development of short-term memory loops and transition of short-term memory to a permanent storage. Recently, experiments were performed to study the time mer.lOry transition in rats. pattern of memory
590
After a 5-day habituation period to the experimental environment in which the animals were provided with water after 23-hour water deprivation, each drinking attempt was associated with electrical shocks on the 6th day. The animals were removed from this conflict situation in the 10, 60, 120, 180 and 300th seconds after the third drinking attempt and the latency of the first drinking was tested on the next day. With Vlith regard that the intervals between subsequent drinking attempts showed an exponential increase, the fourth could be e::pected about 320 to 400 seconds. It was found that the shortest latency of first responding was shown by the rats which were removed at 10 and 300 sec, and the longest latency by those which were removed near to the middle of the subsequent intertrial interval. 'rhis This inverse U-shape increase and decrement of the response inhibition was markedly impaired in rats with septal-hippocampal damage. These observations led to assume that the development of short-term memory loops requires the integrity of limbic system, on the one hand, and that the transition of short-term memory to a longterm form is impaired in hippocampectornihippocampectomized rats. This function of the limbic system is under a partial control of hwnoral influences; both hormones and drugs given in the transition time may produce changes in the memory function. REFEREliJCES REFEREIiJCES 1. Boring,E.C.: Sensation and perceEtion in the histor of ex erimental s c 010 tew or: App e on- en ury- ro , 2. Lliller,G.A. ,Galanter,E. ,Pribram,K.H.: Plans and the structure of behavior. New York: Holt,Rinehart and Winston,1960. 3. Luria,A.R.: Higher corticGl functions in man. New York:Basic Eooks,196l. 4. Adams,J.A.: Psychol. Bull., 70:486, 1968. 5. Deutsch,J.A.: Brit.J.Psychol., 47:115, 1956. J.F.Delufresnaye IEdl 6. Anokhin,P.K.: In J.F.Delafresnaye SprincErain rr:echa!'1isms and learning. Sprinc:,Charles C '~\!lomas,1961. '~\1l0mas,1961. field, Ill. Ill.,Charles Hull ,C .L.: 'The The Drinciple Drinciple of behavior. behe,vior. 7. Hull,C.L.: York:;"ppleton-Century, 1943. New York:Appleton-Century, ~edizinische Psychologie. 8. Lotze,R.H.: ~edizinische Leip zig :Yieidemann. Leipzig:Weidemann. J2nes, VI.: 9. Jenes, Vi.: Principles of gs"chOl2£:L' vol.2. New York:Holt,189 .t Lear!'1ing theory ond be10. r.:oVirer,O.E.: Lear:1ing hayior. Eew Kew York:Wiley, 1960. 11. Koncrski,J.: Integrative activity of the brain. Chicago Univ. Press,1967. 12. Hebb,D.O.: Concernin~ Concernin~ icagery icagery.. 75:4b6 ,1968. Psychol. Bull., 75:4b6,1968. 13. Beritashvili,I.S.: In: ~.Cole,I.~altz ~.Cole,I.~altz IEds/, A handbook of cont errJDorary eUJDorary man IEdsl,A psychOlogy. I:eVl York :Basic Books. Book s . soviet psychology. 1969.
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