drive-controlled reflexes: a theorydrive-controlled reflexes $77 ing reflex is usually...

ACTA NEUROBIOL. EXP. 1973, 35: 475-490

Memorial Paper in Honor of Jerzy Konorski

DRIVE-CONTROLLED REFLEXES: A THEORY

Boguslaw ZERNICKI

Department of Neurophysiology, Nencki Institute of Experimental Biology Warsaw, Poland

Abstract. Drive-controlled (higher) reflexes are divided in two ways: (i) according to the controlling drive into pain, fear and appetite reflexes; (ii) according to the manner of drive elimination into type A and type B reflexes. A type A reflex simply removes the evoking stimulus, whereas a type B reflex provides a drive-inhibiting stimulus. According to different functions performed in an organism, reflexes are grouped into several systems.

I. INTRODUCTION

T h s paper represents a developed version of earlier considerations (28, 29). Drive-controlled forms of neuro-behavioral activity will be analysed and classified. Some of my concepts have been discussed with Professor Jerzy Konorski, and some have been influenced by ideas delivered in his last monograph Integrative activity of the brain. An interdisciplinary approach (14). In particular, Konorski's concept of ani- drive (14, p. 49) has strongly influenced my own speculations.

A number of divisions of reflexes or behavior have been already proposed (1, 2, 10-12, 14, 17, 19, 21) and some were reviewed in the earlier paper (28). However, a problem of a satifsactory reflex classification remains open. A classification proposed in this paper is based on new types of criteria, and thus considers behavior from a different pers- pective.

My considerations have a general character, and I have decided to use some terms (reflex, pain, fear, appetite) in their widest application

476 B. ZERNICKI

(see 6). Although this may be to some degree unconvenient for the read- er, an introduction of new terms would be perhaps even more cumber- some.

11. FUNDAMENTALS

Lower and higher reflexes

The term "reflex" will denote neural processes and effector (behavioral) responses which are evoked by a definite stimulus and form a functional entity. Numerous data indicate that some reflexes are controlled by drives. It is well documented, for example, that alimentary and defensive conditioned reflexes are energized by drives (e.g., see data from Konorski's Laboratory: 14, ch. 9; 16; 24; 27; 31). One can assume that a given reflex is drive-controlled when: (i) it occurs only in the presence of the corresponding drive, its intensity proportional to the strength of drive; (ii) it is followed by drive reduction. Drive-control will be considered a fundamental feature and consequently reflexes will be divided into two large categories: drive-uncontrolled reflexes and drive-controlled reflexes. 'l'hese two groups will be also denoted as lower and higher reflexes, respectively.

Lower reflexes are localized mainly in the segmental part of the central nervous system, and can be observed even in a severely truncat- ed neural system. The light reflex, for example, is fully preserved in the isolated midbrain with hypothalamus of the cat (30). On the contr- ary, for higher reflexes an awake cerebrum is needed. It is reasonable to assume that drives and consequently higher reflexes are absent in decerebrated animals as well as under sleep or narcosis.

A given stimulus can simultaneously evoke both lower and higher reflexes. For example, strong sunlight evokes constriction of pupils (lower reflex) and putting on protective glasses (higher reflex). Simi- larly, a rise of body temperature produces dilatation of skin vessels (lower reflex) and taking a cold shower (higher reflex).

In some instances, a higher reflex is "superimposed" on a lower one. A reflex to a pinprick of the leg is an example; the leg flexion is only partially energized by the drive. In all situations when the pain- ,drive is eliminated (e.g., narcosis), the flexion reflex is reduced t~ the lower reflex. Another example is the locomotor higher reflex, which is superimposed on postural lower reflexes.

A given higher reflex is often preceded by an adjustment of a given analyser (or analysers) to perceive the stimulus. This introductory stage of reflex activity is called orienting reflex. One can assume that orient-

DRIVE-CONTROLLED REFLEXES $77

ing reflex is usually drive-uncontrolled. For example, a predator is re- cognized due to the ocular fixation (orienting reflex), and only then the flight reflex, energized by the fear drive, follows. However, the adjustment of a given analyser can constitute also a part of the higher reflex (see 3). For example, during attack behavior the predator will carefully observe the prey and then ocular fixation is obviously drive-controlled. It should be noted that some authors (see 9) use the term orienting reflex in wider reference.

Some higher reflexes are followed by a lower reflex, which creates an important final stage of the reflex sequence producing drive reduction. Such lower reflex will be called consummatory. The examples of consummatory reflexes are swallowing and ejaculation (see below). Thus, in this paper the term consummatory reflex is used in a narrow sense.

In summary, some lower reflexes cooperate closely with higher ones. Lower reflexes can create a functional background, an introductory stage (orienting reflex) and a final stage (consummatory reflex) of the reflex sequence. In the following part of the paper consummatory reflexes will be frequently mentioned, whereas other categories of lower reflexes will be usually neglected'.

Emotional and specific components of higher reflexes

In higher reflexes it is convenient to distinguish between two components: emotional (controlling) component and specific (controlled) component. These terms stress that the specificity of a given reflex is rather not due to its emotional component. In fact, a given emotion can control many different reflexes.

The emotional component is represented almost exclusively by neural processes; direct overt expression of emotions is limited mostly to facial and vocal responses. Conversely, the specific component is composed of neural and behavioral counterparts. The emotional component is localized in the limbic system, whereas localization of the specific component is mainly in the sensory and motor divisions of the central nervous system. Both components will be described in some detail below.

Emotional component. The term "emotion" will denote any unpleasant or pleasant phychic response. Four main types of emotions will be distinguished: pain, pleasure, fear and appetite. The corresponding central processes will be called, respectively, pain, pleasure, fear and appetite processes, and the corresponding stimuli, painful, pleasant, fearful, and appetitive.

There is a clear difference between pain and pleasure emotions on the one hand, and fear and appetite emotions on the other. Pain and

pleasure are evoked by primarily important stimuli, whereas fear and appetite by stimuli signalling the possibility of action of the former.

All of these terms are used in the widest sense of the words (see 6). Thus, pain, which is considered the synonym of "primary" unpleasure, is evoked by a nociceptive cutaneous stimulus, dehydration of the body, a loud noise etc.; pleasure is evoked by tasty food in the mouth, sight of a beautiful painting, good music etc; and fear and appetite by stimuli signalling these pains and pleasures.

I will make two important assumptions: (i) Three of these emotions, namely, pain, fear and appetite are drive emotions (drives), i.e., they facilitate the specific component of the reflex. (ii) Drive emotion is al- ways unpleasant; in other words, a subject acts only when he is "un- happy". These points will be discussed in detail in Section 111.

Although it is beyond the scope of this paper to discuss the pro- blems of learning, three points will be given:

1. Pain and pleasure are usually innate. Surprisingly, however, with individual experience, the number of pleasant stimuli and perhaps also the number of painful stimuli is increased. For example, the pleasure evoked by a sophisticated food, good music or a beautiful painting can be acquired. On the other hand, fear and appetite are usually acquired. Even in man, however, they can also be innate, for example, the sight of a snake may evoke innate fear, and the odor of meat innate appetite (see 25).

2. During conditioning a neutral stimulus "reinforced" by a painful stimulus will become the fearful conditioned stimulus, and a neutral stimulus reinforced by a pleasant stimulus will become the appetitive one. In other words, the emotions evoked by conditioned stimuli do not imitate those produced by the unconditioned painful and pleasant stimuli.

3. With overtraining some higher reflexes (particularly fear reflexes) seem gradually to loose the drive component, and thus become reduced to lower reflexes. Such reflexes are called automatic. A good example is automatic car operation.

Finally, the problem of the rage emotion, which has not been considered basic, should be noted. One can assume that rage is present when we intend to destroy something: the rage central processes facilitate an attack response. Thus, the rage drive can accompany pain, fear and appetite drives.

Specijic component. In accordance with the broad definition of reflex, accepted in this paper in some higher reflexes the specific component is complex. In particular, it can consist of several stages (see Sec-, tior! IV).

DRIVE-CONTROLLED REFLEXES I79

In some reflexes, the specific response consists clearly of two parts, somatic and vegetative. The somatic response obviously plays a basic role in the reflex and it will be mainly discussed in this paper. However, a t a given stage of a reflexes the vegetative response can predominate. A good example is a chain alimentary reflex in the Pavlovian experimental situation. In the first stage of the reflex the dog salivates to the sound of a metronome, and in the second stage it takes the food presented (see 4).

An interesting problem concerns which of the central processes that mediate the specific response are facilitated by the central drive processes. There are two main candidates: (i) Highest motor processes program- ming the specific response; they may be identical with kinesthetic gnostic processes and are localized i~n the premotor cortex (14, p. 190; 16); (ii) Gnostic processes of the evoking stimulus (and of some cue stimuli) localized at the highest level of the sensory systems. We know from introspection that in some reflexes an important role is played by an image, e.g., the image of the signalled painful or pleasant stimulus, or the image of a necessary tool (see 1, 28). There probably exist mutual relationships between image central processes and drive central processes.

111. TYPES OF HIGHER REFLEXES

Two reflex divisions, based on different criteria, will be proposed: 1. In accordance with above considerations, one can assume that

a given higher reflex is driven either by pain, or by fear, or finally by appetite. Consequently, reflexes will be divided into pain, fear and appetite reflexes. The synonyms of these terms are, respectively, escape, avoidance and approach reflexes (see 29). Those reflexes are evoked, respectively, by painful, fearful and appetitive stimuli.

2. One can also assume that one role of a higher reflex is to reduce the corresponding drive. This can be fulfilled in two ways: (i) the evoking stimulus is simply removed, and (ii) a new stimulus inhibiting the drive is provided. For example, in the flight reflex of a child, the con- tact with the mother is such a sti~nulus that inhibits the central fear processes.

Reflexes which work according to the first way will be termed type A, and those which work according to the second way or both ways will be termed type B. The stimulus provided by the type B reflex will be called an antidrive stimulus.

The models of type A and type B reflexes are shown in Fig. 1. One can see that in both reflexes the stimulus evokes two kinds of central

B. ZERNICKI

specific stimulus processes response

drive processes

stimulus specific processes response

processes i Fig. 1. Models of type A and type B reflexes. In the type A reflex the response simply abolishes the evoking stimulus, whereas in the type B reflex an antidrive stimulus is provided. I t is not indicated that in the type B reflex the response can also abolish the evoking stimulus. Key to the symbols for this and subsequent Figures: circle indicates a stimulus, rectangle indicates central processes, octagon indicates an effector response. Arrow denotes evoking central processes or effector response, or providing a stimulus. Line ending with perpendicular dash denotes inhibition of central processes, or abolition (or diminution) of stimulus. Bent arrow

denotes facilitation of central processes.

processes: (i) the processes mediating the specific response and (ii) the drive processes. In the type A reflex the response removes the evoking stimulus. In the type B reflex the response provides the antidrive stimulus inhibiting the central drive processes.

The role of the antidrive stimulus is reasonable. In pain refl-exes the antipain stimulus signals that the painful stimulus will be abolished, in fear reflexes the antifear stimulus signals that the painful stimulus will not appear, and finally in appetite reflexes the role of the antidrive stimulus is played by the pleasant stimulus (see below).

Pain reflexes

In the pain reflex, central specific processes are facilitated by central pain processes. Both, type A and type B reflexes are in operation.

DRIVE-CONTROLLED REFLEXES 481

Due to the type A pain reflex, the painful stimulus is directly abolished or a t least its intensity diminished. For example, a painful stimulus acting on a leg is removed by hand.

The type B pain reflex provides an antipain stimulus and also abolishes the painful stimulus. The antipain stimulus evokes quick, partial or complete relief of pain. Two different situations are possible:

1. Providing an antipain stimulus simultaneously creates a situation abolishing the painful stimulus (Fig. 2). For example, on a hot day

painful specific processes response stimulus

Fig. 2. Type B pain reflex. The response provides an antipain stimulus and simultaneously removes the painful stimulus.

a subject takes a shower, the cold water is the antipain stimulus reduc- ing the feeling of heat and simultaneously it cools the body. In this case, the relief produced by the antipain stimulus can be complete. Another example is when a suffering subject calls for a physician. His sight is the antipain stimulus which can partially inhibit pain. In both examples the antipain stimulus signals that the painful stimulus will be soon abolished.

2. The antipain stimulus evokes a consummatory reflex (lower reflex) which removes the painful stimulus (Fig. 3). An example is when a subject drinks water (thirst can be considered a special category of pain in the wide sense of this word). Water in the mouth is the anti- thirst stimulus (we feel then relief) and evokes swallowing. Water provided into the body removes its dehydration.

It should be noted that consummatory reflex in turn can provide an interoceptive antidrive stimulus (not indicated in Fig. 3). For example, in the alimentary reflex, food provided into the stomach can be such a stimulus. Then both the exteroceptive antihunger stimulus (food in the mouth) and the interoceptive antihunger stimulus collaborate closely,

pain processes

antipain processes stimulus

consummdory consummatory response processes H

Fig. 3. Type B pain reflex followed by consummatory reflex. The antipain stimulus evokes a consummatory reflex which abolishes the painful stimulus. I t is not indicated that the consummatory reflex can provide an interoceptive antipain sti-

mulus.

Fear reflexes

In the fear reflex central specific processes are facilitated by the central fear processes. Type A and type B reflexes are possible. ,

The type A fear reflex abolishes the fearful stimulus. Flight of an animal evoked by the sight of a predator is an example of such a reflex.

Type B fear reflex provides the antifear stimulus, which inhibits the central fear processes; it signals that the painful stimulus will not appear. For example, for a lost traveller the antifear stimulus is the sight of a house.

Appetite reflexes

In the appetite reflex central specific processes are facilitated by central appetite processes. The type B reflex is in operation. The response provides a pleasant stimulus. The central pleasure processes inhibit the appetite processes, i.e., they play the role of a central antidrive processes. Turning on the radio to hear music is the example of such a reflex (this reflex may be evoked by the sight of a radio).

Similarity between appetitive and pleasant stimuli. We know from introspection that an appetitive stimulus can evoke not only appetite but also pleasure, which will be called secondary (Fig. 4). In other words we postulate that the appetitive stimulus may simultaneously evoke ne- gative and positive emotions. This is particularly demonstrative in sexual reflexes: the sight of an attractive member of the opposite sex obviously

DRIVE-CONTROLLED REFLEXES

specific response

stimulus

appetite processes

I pleasure

processes stimulus

Fig. 4. Appetite reflex. Similarities between appetitive and pleasant stimuli are indicated. Interrupted line denotes weak influence.

evokes both appetite and pleasure (the latter explains the popularity of strip-tease). The secondary pleasure, however, is too weak to completely inhibit appetite.

The difference between appetitive and pleasant stimuli may only be in intensity. For example, barely audible music can be the appetitive stimulus, music of an apropriate intensity the pleasant stimulus; the appetite reflex may consist of adjusting the volume dial of the radio. Another example may be taken from the alimentary system, when a por- tion of tasty food is swallowed, the residual food present in the mouth becomes the appetitive stimulus and the subject takes the next morsel of food into the mouth (peanut phenomenon). These examples are very instructive, the appetitive stimulus becomes pleasant a t the moment that the amount of pleasure processes are sufficient to fully inhibit appetite processes.

The above considerations suggest that the pleasant stimulus also has an excitatory connection with the appetite center (Fig. 4). This connection in ineffective because of the simultaneous strong inhibition from the pleasure center.

The appetite reflex followed by a consummatory reflex. Some appetite reflexes are followed by consummatory reflex, which provides an interoceptive antidrive stimulus (Fig. 5). Let us consider a following example of pure appetite alimentary reflex: a tasty food is taken by an unhungry animal. One can assume that food in the mouth has an antiappetite effect and evokes swallowing. The food in the stomach is the interoceptive antiappetite stimulus which preserves overeating. It should be noted that when tasty food is presented to a hungry animal, appetite and hungry reflexes collaborate closely, both being followed by the same consummatory reflex-swallowing (see Section V, alimentary system).

6 - Acta Neurobiologiae Experimentalis

specific processes response stimulus

antiappetite processes processes

stimulus processes stimulus

consummatory response processes

Fig. 5. Appetite reflex followed by consummatory reflex. The pleasant stimulus evokes consummatory reflex which provides an interoceptive antiappetite stimulus.

IV. CHAIN REFLEXES

Higher reflexes are often in a chain form, i.e., in a given reflex there are two or more clear-cut stages. In a chain reflex the usual role of a given stage is to provide a means stimulus which evokes the next stage of the reflex (Fig. 6). In a two-stage type A pain reflex, evoked by a nociceptive agent, an appropriate tool may be provided in the first stage; the sight of this tool (means stimulus) evokes the second stage

specific processes

I I

processes u I response means stage II

Fig. 6. Two-stage type A reflex. The first stage provides a means stimulus.

removing the evoking stimulus. Another example is the two-stage alimentary appetite reflex evoked by the sound of metronome. The first stage (running to food) provides the means stimulus (sight of food),


which evokes the second stage (eating) providing the pleasant taste stimulus. This stage is followed by the consummatory reflex (swallowing) providing the food into the stomach.

The central specific processes evoked by means stimuli are facilitated by central drive processes. In the pain and fear chain reflexes the drive processes are produced only by the evoking stimulus (Fig. 6 ) . In the pain reflex evoked by a nociceptive stimulus, pain processes produced by this stimulus facilitate specific central processes evoked, for example, by sight of an appropriate tool provided in the course of the reflex. Moreover, in pain and fear reflexes the means stimuli seem to have a weak antidrive effect. In contrast, the means stimuli of appetite reflex produce emotions, which are even stronger than those to the evoking stimulus. For example, in the alimentary reflex mentioned, evoked by the sound of a metronome, appetite and pleasure awakened by this stimulus are obviously weaker than those produced by the sight and smell of food. Thus, in all kinds of reflexes the means stimulus it- self has some "rewarding" effect.

In some chain reflexes a given stage (often the first stage) is not required to provide the means stimulus, but to remove a background inhibitory stimulus (see 29, Fig. 13). For example, micturition may be inhibited in the presence of another person, hence the first stage of the micturition reflex is the locomotor response eliminating this inhibitory stimulus. The specific central processes of this stage are facilitated by the central drive processes awakened by the evoking stimulus. Intro- spective experience suggests that the background inhibitory stimulus has some antidrive effect.

V. SYSTEMS OF HIGHER REFLEXES

A given function of an organism (e.g., alimentary function) is fulfilled by an appropriate system of reflexes, usually including both lower and higher reflexes. Iligher reflexes of different systems are controlled by different drives (hunger, thirst, sexual drive etc.). Some important systems of higher reflexes will be briefly dmcribed. The presence of pain, fear and appetite reflexes in representative systems is shown in Table I.

Ct~taneous unpleasure system (cutaneous nociceptive system). In this system type A pain reflexes and fear reflexes are in operation.

Cutaneous pleasure system. In this system appetite reflexes are in operation. The example of a pleasant stimulus is nonsexual caressing.

Micturition and defecation systems. Both micturition and defecation are type A pain reflexes. Occasionally micturition is a type B fear

486 B. ZERNICKI

Representative systems of reflexes. Some systems described in the text are omitted. Only higher reflexes are indicated.

System

Cutaneous nociceptive Micturition Body temperature Respiratory Alimentary Sexual Wake-sleep transition Curiosity Visual unpleasure Visual pleasure

Pain reflex

Appetite reflex reflex

Symbols: +, important; 0, absent or insignificant.

reflex. An example is micturition before going to sleep: stimuli received during the micturition act (antidrive stimuli) inform the subject that its urinary bladder is empty.

Body temperature system. Two subsystems can be distinguished that involve one for decreasing and the other for increasing body temperature. The type B pain reflex and fear reflexes are in operation. In the above mentioned example of a type B pain reflex, a cold shower taken by a subject immediately abolishes the feeling of heat (antipain stimulus) and slowly cools the body.

A subject can also regulate body temperature by drinking liquid of different temperature than the body. Then the type B pain reflex is followed by a consummatory reflex. Liquid in the mouth is the antipain stimulus and swallowing provides liquid into the body.

Respiratory system. The appropriate humoral stimulus, if strong enough to produce pain, evokes the type B pain reflex. Fresh air is the antipain stimulus and it also restores normal carbon dioxide content in the circulation. The specific component may involve respiratory muscles (strong action) as well as other effectors (e.g., the subject opens a win- dow).

Fear reflexes occur in particular situations. For example, the subject makes a deep inspiration before diving (type B fear reflex).

Alimentary system. Alimentary pain is called hunger. Type B pain and appetite reflexes are in operation. Both are followed by consummatory reflex (swallowing). Mixed', hunger-appetite reflexes are often present.

DRIVE-CONTROLLED REFLEXES 48 7

In the hunger reflex the antihunger stimulus is tasteless food in the mouth, which inhibits the central hunger processes (short-term satiety) (see 8; 14, ch. 1, 6; 24). When a sufficient amount of food is consumed, the hunger stimulus is eliminated (long-term satiety).

Solid food in the mouth evokes masticatory movements and sali- vation. These responses seem to be controlled by hunger only to some degree (see 23). In other words, they belong partially to the consummatory component of the reflex.

The appetite reflex provides tasty food in the absence of the hunger stimulus. This is the normal mechanism of eating a dessert. The extra amount of swallowed food provides the interoceptive antiappetite stimulus (presumably dilatation of stomach) inhibiting the reflex.

The hunger-appetite reflex is usually evoked by an appetitive stimulus appearing in the background of the hunger stimulus: such as a tasty food that is presented to the hungry animal. The specific response is then facilitated by both hunger and appetite, and both of these drives are inhibited by the pleasant stimulus.

Neural mechanisms of alimentary system are better known than other systems. A few main data will be presented to show that at least they are not contradictory to the concepts developed in this paper. Al- though in the decerebrate cat, mastication and swallowing can be evoked, the consummatory reflex is not integrated and the animal must be fed by a stomach tube. A chronic thalamic cat can be fed by hand, and a chronic striatal cat can find food placed on the floor but with some difficulty (5, 26). These acts seem to be abortive hunger reflexes. As shown by experiments with local lesions or electrical stimulation, the long-term satiety is controlled by the center in the ventromedial hypothalamus. On the other hand, both the short-term satiety (hunger antidrive) and alimentary pleasure are presumably controlled by units scat- tered among the hunger units in the lateral hypothalamus (see 14, p. 50), This is suggested at least by two facts: (i) after the ventromedial hypothalamic lesion the instrumental conditioned reflexes are saved, thus the food preserves its reinforcing value; (ii) stimulation of the lateral hypothalamus seems to evoke both hunger and pleasure (e.g., see 13, 18, 20). Hypothalamic alimentary functions seem to be duplicated in the amygdala (7). On the other hand, an antidrive center located in the me- dial part of the prefrontal cortex (see 15) plays a different role: its re- moval produces disinhibition of conditioned reflexes. Finally, alimentary pleasure, and consequently appetite, seem to be impaired after ablation of the gustatory cortex (32).

Water system. Everything said about alimentary reflexes seems to be true for water reflexes. The painful stimulus evoking the water re-

488 B. ZERNICKI

flex is called a thirst stimulus. Tasteless water of body temperature is provided by the thirst reflex, whereas tasty water (e.g., soda water) by the appetite reflex in the unthirsty subject and by mixed thirst-appetite reflex in the thirsty one.

It should be noted that food and water intake also play an important role in the temperature system (see above). Food and water, only if of body temperature, have purely alimentary or "water" meaning.

Sexual system. In men, the appetite reflex (followed by consummatory reflex) seems to be usually in operation. The stimulus is often the sight of a member of the opposite sex. Tactile stimuli acting during caressing and coitus have a double meaning. In the beginning they play a role as the means stimuli (they evoke appetite and secondary pleasure), and they rapidly become the pleasant stimulus during the orgasm (the pleasure processes becomc very strong and fully inhibit the appetite processes). Then the consummatory reflex (in male - ejaculation) provides the interoceptive antidrive stimulus (its nature is not well known) inhibiting the reflex.

After deprivation of sexual activity for a considerable periods of time, the painful (hunger) sexual stimulus appears, and the pain-appetite sexual reflex is in operation. This reflex occurs in female animals in the estrus state.

Wake-sleep transition system. In favorable conditions (comfortable body position, quiet place etc.) no behavioral response is necessary for the transition from wakefulness to sleep. In fact, sleep-waking cycle is present even in the isolated cerebrum (22). When interferring stimuli (e.g., noise) are present, the painful stimulus evoking somnolence-drive appears and an appropriate place for sleep is provided by type A pain locomotor reflex (see 14, p. 301).

Curiosity system. In this system, the type A pain reflex is presumed to be mainly in operation. It is called the investigatory reflex and should be clearly distinguisned from the orienting reflex. An example of the investigatory reflex is the locomotor behavior of a dog in a new experimental room.

Visual and auditory unpleasure systems. Type A pain reflexes and fear reflexes are in operation. The obvious examples of painful stimuli are a strong light and a noise. Possibly some patterned visual and auditory stimuli have also primary unpleasant meaning, i.e., pain meaning in the wide sense of this term. We can speculate that in phylogeny some important fearful stimuli (heights, sight of a snake etc.) have become painful stimuli (see Section 11).

Visual and auditory pleasure systems. In these systems appetite reflexes are in operation. The examples of pleasant stimuli are, respecti-


vely, the sight of beautiful mountains, and good music. In the visual and auditory systems the response often consists exclusively of the long-last- ing adjustment of a given analyser to perceive the stimulus well. Thus, the receptor adjustment is first the orienting reflex (lower reflex) and then higher reflex energized by visual or auditory drive (see Section 11).

These two systems probably exist only in men. They represent numerous human systems of higher reflexes, which are controlled by sophisticated drives, including social ones.

REFERENCES

1. BERITOV, I. S. 1965. Neural mechanisms of higher vertebrate behavior. Little Brown Co., Boston. 384 p.

2. CRAIG, W. 1918. Appetites and aversions as constituents of instincts. Biol. Bull. 34: 91-107.

3. DREHER, B. and ZERNICKI, B. 1969. Visual fixation reflex: behavioral pro- perties and neural mechanism. Acta Biol. Exp. 29: 359-383.

4. ELLISON, G. D. and KONORSKI, J. 1965. An investigation on the relations between salivary and motor responses during instrumental performance. Acta Biol. Exp. 25: 297-315.

5. EMMERS, R., CHUN, R. W. M. and WANG, G. H. 1965. Behavior and reflexes of chronic thalamic cats. Arch. Ital. Biol. 103: 178-193.

6. ENGLISH, H. B. and ENGLISH, A. C. 1958. A comprehensive dictionary of psychological and psychoanalytical terms. Longmans Green Co., New York. 594 p.

7. FONBERG, E. 1974. Amygdala functions within the alimentary system. Acta Neurobiol. Exp. 34: 435-466.

8. GALPERIN, S. I. and RUBINOV, I. S. 1968. The effect of unconditioned and conditioned stimuli on periodic motor activity of the stomach and duode- num in dogs. Acta Biol. Exp. 28: 93-106.

9. GRASTYAN, E. and VERECZKEI, L. 1974. 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. 10: 121-146.

10. HEBB, D. 0. 1958. A textbook of psychology. Saunders Co., Philadelphia. 353 p. 11. HILGARD, E. R. and MARQUIS, D. G. 1940. Conditioning and learning. Ap-

pleton-Century-Crofts, New York. 550 p. 12. HINDE, R. A. 1966. Animal behaviour. A synthesis of ethology and compara-

tive psychology. McGraw-Hill. London. 574 p. 13. HUSTON, J. P. 1971. Relationship between motivating and rewarding stimula-

tion of the lateral hypothzlamus. Physiol. Behav. 6: 7111-716. 14. KONORSKI, J. 1967. Integrative activity of the brain. An interdisciplinary

approach. Univ. Chicago Press, Chicago. 531 p. 15. KONORSKI, J. 1572. Some hypotheses concerning the functional organization

of prefrontal cortex. Acta Neurobiol. Exp. 32: 595-613. 16. KONORSKI, J. 1974. Classical and instrumental conditioning: The general laws

of connections between "centers". Acta Neurobiol. Exp. 34: 5-13. 17. KOZAK, W. and WESTERMAN, R. 1965. Basic patterns of plastic change in

the mammalian nervous system. Symp. Soc. Exp. Biol. 20: 509444.

490 B. ZERNICKI

18. MARGULES D. L. and OLDS, J. 1962. Identical "feeding" and "rewarding" systems in the lateral hypothalamus of rats. Science 135: 374-375.

19. PAVLOV, I. P. 1928. Lectures on conditioned reflexes. Int. Pub., New York. 20. SADOWSKI, B. 1973. Negativism to food during self-stimulation in the ante-

rior part of the basal forebrain in dogs. Physiol. Behav. 13: 645-651. 21. SECHENOV, I. M. 1935. Selected works. State Publ. Hous. Biol. Med. Lit.,

Moscow. 489 p. 22. SLOSARSKA, M. and ZERNICKI, B. 1973. Chronic pretrigeminal and cerveau

is016 cats. Acta Neurobiol. Exp. 33: 811-827. 23. SOETYSIK, S. 1971. The effect of satiation expon conditioned and uncondition-

ed salivary responses. Acta Neurobiol. Exp. 31: 59-63. 24. SOETYSIK, S. 1975. Post-consummatory arousal of drive as a mechanism of

incentive motivation. Acta Neurobiol. Exp. 35: 475-502. 25. TSITOVICH, J. 1911. Origin and elaboration of the natural conditioned reflexes

(in Russian). Ph. D. Thesis. Quoted, 19. 26. WANG, G. H. and AKERT, K. 1961. Behavior and reflexes of chronic striatal

cats. Arch. Ital. Biol. 100: 48-85. 27. WYRWICKA, W., DOBRZECKA, C. and TARNECKI, R. 1960. The effects of

electrical stimulation of the hypothalamic feeding centre in satiated goats on alimentary conditioned reflexes type 11. Acta. Biol. Exp. 20: 121-136.

28. SERNICKI, B. 1968. Reflex as the unit of neuro-behavioral activity: A theore- tical multidisciplinary approach to the reflex activity in higher animals. Acta Biol. Exp. 28: 291-315.

29. ZERNICKI, B. 1972. Higher reflexes: An attempt a t comprehensive classification of neuro-behavioral activity. Acta Neurobiol. Exp. 32: 87-112.

30. ZERNICKI, B., DOTY, R. W. and SANTIBANEZ-H., G. 1970. Isolated midbrain in cats. Electroenceph. Clin. Neurophysiol. 28: 221-235.

31. ZERNICKI, B. and EKEL, J. 1959. Elaboration and mutual relations between alimentary and water instrumental conditioned reflexes in dogs. Acta Biol. Exp. 19: 313-325.

32. ZERNICKI, B. and SANTIBANEZ-H., G. 1971. Alimentary and acid avoidancs instrumental conditioned reflexes in dogs after ablation of the anterior composite gyrus (gustatory cortex). Acta Neurobiol. Exp. 31: 365-371.

Received 28 November 1974

Boguslaw ZERNICKI, Department of Neurophysiology, Nencki Institute of Experimental Bio- logy, Pasteura 3, 02-093 Warsaw, Poland.

drive-controlled reflexes: a theorydrive-controlled reflexes $77 ing reflex is usually...

Documents