Neuroscience and Behavioral Physiology, Vol. 31, No.

1, 2001

The Development of the Scientific Ideas of I. P. Pavlov on the Goal Reflex in Studies of the Mechanisms of Biological Motivation
K. V. Sudakov

Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 85, No. 910, pp. 11631173, SeptemberOctober, 1999. Original article submitted December 25, 1998. Revised version received February 23, 1999. Data obtained in experimental studies developing I. P. Pavlovs concept of the goal reflex are presented. It is shown that purposive behavior in the systems organization of behavioral acts is constructed by the dominant motivation. This article discusses mechanisms determining the energy of biological motivations and their controlling component the results of action acceptor. The role of reinforcement in the mechanisms forming molecular engrams of behavior on structures of the results of action acceptor is assessed.
KEY WORDS: Goal reflex, functional systems, dominant motivation, reinforcement, results of action acceptor engrams.

On January 2, 1916, I. P. Pavlov read a paper on the goal reflex at the Third Congress of Experimental Pedagogy in Petrograd. Analysis of human and animal activity, proposed Pavlov, led me to the conclusion that the set of reflexes must include a special reflex, the goal reflex, which is the tendency to have a defining stimulating object, the meanings of having and object being taken in the widest senses [10, p. 197]. He further wrote: Of all the known reflexes, the goal reflex in human activity is the most typical and therefore especially suitable for analysis, and additionally, the most widespread is the passion to accumulate... The goal reflex has enormous importance in life; it is the main form of vital energy in each of us. Life is only beautiful and strong for those who spend their whole lives striving for the constantly achievable but never achieved goal or jumping from one goal to another with no loss of effort. All life, all its improvements, all its culture, is achieved by the goal reflex, and only by people striving for one or another goal which they place before themselves. Pavlov

P. K. Anokhin Science Research Institute of Normal Physiology, Russian Academy of Medical Sciences, 6 Bolshaya Nikitskaya, 103009 Moscow. 87

regarded this activity as an obscure, primal, irresistible attraction, an instinct, or a reflex. Pavlov constructed his concepts of the goal reflex on the basis of the traditional reflex paradigm, suggesting that the innate basic attractions are regular reflex responses of the body to defined external and internal stimuli. However, the striving of the subject to achieve a defined external object is not included in the traditional concept of the reflex, which is based on the stimulus-response principle. In this case, we do not see the responses of subjects to stimuli but, contrarily, an urgent search and acquisition by subjects of special objects of the external world. This seems to have been an insurmountable obstacle to explanation in Pavlovs time. In subsequent years, he understandably focused on studies of conditioned reflexes as types of response reactions of living beings to external stimuli and never returned to the question of the goal reflex which he had raised. Only at the end of his life, in 1935, at his Wednesday of 13 November did Pavlov say: A monkey building a tower to reach a fruit cannot be called a conditioned reflex. This is a case of forming knowledge, grasping the normal connection of things! This is a different situation. We have to say that the principle of the formation of knowledge, grasping the constant connections between things, is the basis of all scientific activity, the laws of causality, etc. [11].
0097-0549/01/3101-0087$25.00 2001 Plenum Publishing Corporation

88 The scientific ideas of Pavlov on the goal reflex are very closely intertwined with Ukhtomskiis concept of the dominant. As Ukhtomskii wrote: The dominant selects the corresponding stimuli from the environment [30, pp. 235265] and, elsewhere: Each time there is an ongoing symptom-complex dominant, there is also a predefined behavioral vector [29, p. 300]. It is already evident from these quotations that the behavior factor is the dominant excitation formed within the body. The question is to identify the nature of the dominant excitation determining purposive behavioral acts in animals and humans. It is natural to pose the question: which internal mechanisms define the vital energy of attraction and reliably guide subjects to achieving defined objects of the external world? P. K. Anokhins theory of functional systems allowed an answer to this question to be approached [2, 3]. According to functional systems theory, the central architectonics of behavioral acts is made up of the following sequentially replacing important stages: afferent synthesis, decision-taking, predicting the required result (the results of action acceptor), activation of an effector program of action (efferent synthesis), and constant assessment by the subject of measures of the results achieved using reverse afferentation arriving from peripheral receptors to the results of action acceptor. The leading component of the stage of afferent synthesis in the systems architectonics of purposive behavioral acts is the dominant motivation. From the point of view of functional systems theory, dominant motivations, which are a component of the afferent synthesis stage, also take part in forming other sequentially replacing stages of the systems organization of behavioral acts: decision-taking, operation of the results of action acceptor, and the efferent behavioral program efferent synthesis. The dominant motivation thus permeates all stages of the systems organization of behavioral acts, combining the components of their neural elements to achieve a result useful to the organism satisfaction of the initial defining need. Under the influence of the dominant motivation, subjects not only react selectively to external stimuli, but also actively search for special factors (stimuli) in the external environment which would satisfy their initial needs. This search is often conducted extremely persistently, with great energy strength and overcoming various hindrances on the pathway to the goal. Biological motivations, such as hunger, fear, sexual attraction, thirst, etc., are formed predominantly endogenously by means of metabolic processes creating the initial metabolic needs of the body, though their realization in behavior can be affected by external so-called key or defining external factors.

Sudakov Our studies, conducted over many years, have allowed us to discover several mechanisms determining the energy basis of biological motivations. Studies of overall electrical activity (EEG) in various brain structures in animals have shown that dominant biological motivations are constructed on the basis of specific ascending activatory influences from the hypothalamic motivation centers on other brain structures, including the cerebral cortex. These influences are manifest as EEG desynchronization or activation responses [16, 18]. The leading pacemaker role of the hypothalamic centers in organizing cortical-subcortical architectonics of dominant motivations has been demonstrated. Suppression of activity in these centers eliminates motivational excitation in all brain structures connected to them. The motivation centers of the hypothalamus thus maintains, in a defined morphofunctional system, all other brains structures connected to them, up to and including the cerebral cortex [19]. Our studies have shown that the motivational state is constructed on the basis of the structural and neurochemical integration of brain formations. The structural integration of dominant motivations, along with ascending activatory influences of the motivation centers of the hypothalamus, activate specific descending excitatory and inhibitory effects of the cortex on the intiatory hypothalamus motivation centers. Thus, the rostral parts of the cerebral cortex had predominantly inhibitory influences on the formation of the food-related and defensive motivations in rabbits evoked by electrical stimulation of the lateral and ventromedial parts of the hypothalamus respectively. The occipital regions of the cortex have facilitatory influences on the formation of the food-related motivation and inhibitory effects on the development of the defensive motivation. Coagulation of the dorsal areas of the hippocampus related inhibitory corticofugal influences on the food-related and defensive motivations, while lesioning of the mesencephalic reticular formation has facilitatory effects on the formation of the food-related motivation in animals [8]. Dynamic reverberant interactions have been demonstrated between the cortex and subcortical structures, including the motivation centers of the hypothalamus [23]; these evidently also determine the animals tonic tension and the duration of the period during which the dominant motivation is manifest. These experiments provide evidence that motivational states are based on a mosaic of dynamically integrated excitatory and inhibitory brain structures. Along with the structural integration of dominant biological motivations, specific neurochemical integration of motivations has also been demonstrated. Unlike researches outside Russia, who have attempted to link biological motivations with some common neu-

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Fig. 1. Responses of neurons in the hunger center of the lateral hypothalamus to electrical stimulation of the motor area of the cerebral cortex in hungry (A) and fed (B) rabbits. A) A rabbit after one day of food deprivation: electrical stimulation of the motor cortex enhances trains of spike activity from a neuron in the lateral hypothalamus; B) in a fed rabbit: in conditions of similar motor cortex stimulation, the neuron fails to respond to stimulation of the lateral hypothalamus. The moment of stimulation is indicated by the black bar half way along each trace. a) Neuron activity trace; b) 200-msec time marker; c) interval histogram of neuron activity before (1) and after (2) stimulation of the motor cortex. The abscissa shows the duration of interspike intervals, msec; the ordinate shows the number of intervals of a given duration as a percentage of the total number of intervals analyzed.

rochemical mechanism [34], we have shown that dominant motivations are constructed by means of the integration of numerous chemical processes simultaneously interplaying in a variety of brain structures. The neurochemical integration of dominant motivations has been shown to involve the same neuromediators and neuropep-

tides, but in different ratios [9]. However, one or several chemical components can be involved in the same motivation in different subjects, which determines their selective relationships to specific blockers. Our studies showed that the formation of dominant motivations involves a variety of oligopeptides. Intrac-

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Sudakov

Fig. 2. Responses of a visual cortex neuron to electrical stimulation of the proximal end of the sectioned pyramidal tract using combined stimulation of the pyramidal tract and subsequent electrocutaneous stimulation. 1) No neuron response to stimulation of the pyramidal tract (indicated by the long line); 2) first combination of pyramidal tract stimulation with subsequent electrocutaneous stimulation (shown by the short line); 3) tenth combination; 4) 16th combination; 5) 63rd combination; 69) extinction of the response to stimulation of the pyramidal tract in the absence of electrocutaneous reinforcement (from [12]).

erebroventricular doses of some oligopeptide inhibit, while other facilitate biological motivations in conditions of electrical stimulation of the motivation centers of the hypothalamus [17, 36]. Thus, angiotensin II, arginine-vasopressin, delta sleep-inducing peptide, substance P, leu- and met-enkephalins, and -endorphin, given into the lateral ventricles, block the food-related and defensive motivations in rabbits in conditions of electrical stimulation of the lateral and ventromedial hypothalamus respectively. Administration of -lipotropin, ACTH1014, melanocyte-stimulating hormone (MSH47), and pentagastrin into the lateral ventricles activated food-related motivation in fed animals. In the same experimental conditions, bradykinin and renin facilitated defensive responses during electrical stimulation of the ventromedial hypothalamus [17]. Special studies conducted in our laboratory showed that the dominant motivation significantly alters the convergent and discriminant pros of neurons in various parts of the brain, changing their sensitivity to neuromediators and oligopeptides. The most clear changes affect the sensitivity of neurons to the effects of reinforcing stimuli [21]. Experiments conducted in our institute by Zhuravlev et al. [6] showed that dominant biological motivations are specifically manifest in the patterns of interspike intervals in the trains of neuron activity in various parts of the brain. Thus, the dominant motivation of hunger in rabbits is manifest in the train activity of neurons in various parts

of the brain with dominant interspike intervals of 10 and 150 msec. In the thirst motivation in rabbits, the discharge activity of brain neurons was dominated by interspike intervals of 25 and 150 msec. The defensive motivation is dominated by neurons with interspike intervals of 40 and 150 msec. Different brain structures in different motivations show a marked gradient in the distribution of neurons with dominant interspike intervals. The proportion of such neurons is higher in the stem areas of the brain and in the hypothalamus, and decreases towards the thalamic formations and cerebral cortex [22]. Cortical neurons in the rabbit brain, as shown by Shvyrkova [31] are involved primarily in the fine mechanisms of the animals interactions with surrounding factors in the environment. The dominant role of motivation in organizing brain functions and behavior has clearly been demonstrated by experiments performed in our institute by Valadov [4]. We would suggest that this author has used an experimental model reminiscent of the classical experiment of Ukhtomskii. Rabbits were immobilized in a cage and the surface of the motor cortex was exposed under local anesthesia and stimulated with an electrical current. The stimulation criterion was a movement response in the contralateral paw. Microelectrodes were placed close to individual neurons in the hunger center of the lateral hypothalamus and baseline activity was recorded in hungry and fed animals, along with the responses of these neurons to electrical stimulation of the motor cortex. These experiments showed that electrical stimulation of motor cortex generally strengthened the pattern of inter-

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Fig. 3. Decreases in the expression of the c-fos gene during the acquisition of a productive defensive habit. Left: decreases in the mean time taken to perform the escape response in mice during daily training. The vertical axis shows the mean time taken to perform the response, sec. 19) Training days. Right: levels of c-fos mRNA in the mouse forebrain. A) Passive controls; B) day one of escape response training; C) ninth day of escape response training.

spike interval distribution of lateral hypothalamus neurons in hungry animals, this pattern being characteristic of the natural hunger motivational state (Fig. 1, A). These animals showed no contraction of the contralateral paw. In fed rabbits, the most typical response to electrical stimulation of the motor cortex consisted of activation or inhibition of the initial spike activity of neurons in the lateral hypothalamus (Fig. 1, B). These animals showed contractions of the contralateral paw. These experiments provided evidence that stimulation of other brain structures in conditions of a dominant motivation increased the strength of this dominant motivation, which suppressed the specific response to stimulation of the motor cortex. This example is also evidence that the dominant motivation subordinates other brain areas and functions, mobilizing them to organize behavior directed to satisfy the initial need. All of these processes which we have observed determine the energy basis of the dominant motivation. In addition, they are only a blind toning force for motivation. Hunger, wrote Sechenov, can raise an animal to its feet, can confer a singular character on searches, but contains no elements directing movements in one or another direction or modifying the need-defined locality or affecting random encounters [13]. It is appropriate to raise the question: what are the mechanisms which ultimately convert motivation from a blind force into an emphatic, seeing, rigid, and error-free motivation controlling the need of experienced subject for special objects in the external world? Our own studies have shown that in the systems organization of behavioral acts, motivation, as a compo-

nent of the afferent synthesis stage, is simultaneously closely associated with another stage in the systems organization of behavior the results of action acceptor. While the dominant motivation at the afferent synthesis stage has an energetic, toning role, at the stage of the results of action acceptor it forms the directing component of motivation the goal, in the widest sense of the word. Not only is the behavioral program constructed and controlled, but it is also actively adjusted by peripheral receptors in terms of the special parameters of the required results [5, 15, 35]. From 1979, we started to develop concepts of the imprinting mechanism for forming the results of action acceptor. According to these concepts, the reinforcing excitation, propagating to the results of action acceptor, influenced by various parameters of the results of the behavior as perceived by the bodys external receptors, is addressed to the brain structures initially excited by the dominant motivation, leaving traces or impressions on these structures these are molecular engrams [20]. These engrams are extracted by the dominant motivation in conditions of sequentially occurring analogous needs. Subsequent experiments supported our theoretical position. Experiments performed by our colleagues Kotov and Zhuravlev [14] showed that motivational and reinforcing excitations are found in the very same brain neurons. This was displayed by up to 50% of neurons in the orbital surface of the cortex in rabbits, while up to 30% of neurons in the sensorimotor cortex showed this property. The clearest interaction between motivation and reinforcement was observed by Zhuravlev [6], which

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Sudakov Our studies over recent years have shown that in conditions of reinforcement, the engram of the results of action acceptor constructs its molecular mechanisms using special protein molecules. In naive animals in which stimulation of the motivation centers of the hypothalamus was not accompanied by reinforcement, the protein synthesis inhibitor cycloheximide had no effect on the searching orientational-investigative response or on the reactions of neurons in various parts of the brain to stimulation of these hypothalamic centers. However, stimulation of motivation-producing hypothalamic centers in combination with adequate reinforcement, such as provision of food to the animal just after stimulation of the lateral hypothalamus, has the effect that subsequent stimulation of the motivation-producing centers and the responses of neurons in various parts of the brain to this stimulation are subject to significant inhibition by cycloheximide. These responses were restored when animals given the protein synthesis inhibitor were also given certain oligopeptides into the lateral ventricles. Thus, food-related motivation in conditions of lateral hypothalamus stimulation in rabbits after administration of cycloheximide is restored by additional administration of pentagastrin. The defensive motivation could not be suppressed by cycloheximide and is restored by additional administration of bradykinin [27]. All of this evidence shows that reinforcement alters the genetic apparatus of neurons involved in the results of action acceptor and that sequential appearance of the corresponding need causes these cells, under the influence of the dominant motivation, to start expressing special effector oligopeptides which organize purposive behavior. Our experiments [1, 33] showed that in the absence of reinforcement, orientational-investigative behavior evoked by the dominant motivation is associated with increased expression to c-fos gene in animals brains. After reinforcement, c-fos expression in the brain decreases (Fig. 3). These profound molecular processes may provide the basis for the mechanism by which the dominant motivation extracts the engram from memory after its formation by effector oligopeptides. In addition, our experiments have demonstrated that the interaction of motivation and reinforcement in structures mediating the results of action acceptor also involves immunological mechanisms. Sudakov showed [26] that food-related motivation in rabbits can be blocked by administration of antigastrin immunoglobulins into the lateral ventricles. Zhuravlev et al. [7] have demonstrated that immunomodulators such as neurotropin, interleukin-1, and interferon fragment 2 (RITLY) improve learning of a defensive habit in rats and improve the prediction of the properties of useful, adaptive results.

Fig. 4. Responses of a sensorimotor cortex neuron to electrical stimulation of the hunger center of the lateral hypothalamus (lHT) before (A) and after (B) microiontophoretic application of anti-fibroblast acidic growth factor antibodies (current 4 nA), and 3 min (C) after microiontophoretic application of anti-fibroblast acidic growth factor antibodies. The abscissa shows time, msec; the ordinate shows the mean number (three lHT stimulations) of spikes per 50 msec.

showed that reinforcement propagated trains of neuron activity to the various parts of the brain involved in the dominant motivation, with a consistent dominance of some particular interspike interval. Our experiments have shown that reinforcement significantly alters the responses of brain neurons to motivational influences. This involves changes in the sensitivity of these neurons to neuromediators and oligopeptides [28]. The studies of Pravdivtsev [12] showed that motivational and reinforcing excitations interact in brain interneurons, which according to P. K. Anokhin make up the apparatus of the results of action acceptor. Antidromal stimulation of the central end of the sectioned pyramidal tract is known to generate responses in the interneurons in animals brains. In Pravdivtsevs experiments on cats under urethane anesthesia, combined antidromal stimulation of the pyramidal tract and subsequent electrocutaneous stimulation of the animals paw produced clear responses in interneurons to the antidromal stimulation of the pyramidal tract: they started to anticipate the responses typical of the nociceptive stimulus. This response was extinguished when reinforcement was withdrawn (Fig. 2).

The Development of the Scientific Ideas of I. P. Pavlov on the Goal Reflex Changes in the responses of brain neurons during stimulation of the motivational hunger center of the lateral hypothalamus in rabbits in conditions of microiontophoretic administration of antibodies to nerve growth factor fibroblast acidic factor were demonstrated in experiments performed by our colleague Kravtsov (Fig. 4). These experiments provide evidence for an immunological mechanism of imprinting of reinforced excitations on neurons involved in the results of action acceptor. Nonetheless, these mechanisms require further study. In recent years we have developed the concept of the informational properties of the bodys functional systems [23]. We have formulated the hypothesis that the results of action acceptor is a holographic screen for assessing needs and their satisfaction in the brain. Informational protein molecules undoubtedly play a leading role in these processes. From these points of view, motivations seeing endogenously, along with the accompanying emotionogenic percepts, are part of the ideal (informational) aspect of behavior [25]. The emotional aspect of motivations has received very little study.

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CONCLUSIONS The factual material presented here, obtained by ourselves over many years, provides evidence that the physiological mechanisms of purposive behavioral acts become more understandable from the point of view of the functional systems theory proposed by P. K. Anokhin. The purposive acts of animals and humans are organized by the dominant motivation, its energetic and controlling component, which in the systems organization corresponds to the results of action acceptor. The most typical feature of dominant biological motivations is the fact that their organizing pacemaker motivation centers are morphofunctionally dependent on all other brain structures involved in motivation. This property underlies the formation of pathological motivations, including alcoholism and drug addiction [24]. The use of ethanol and narcotics functionally transforms the motivation-producing centers in the hypothalamus such that they start to exert pathogenic influences neuron the brain, leading to the pathologically altered behavior of the alcoholic and drug addict. Particularly tight interactions in the systems organization of behavioral acts occur between motivation and the apparatus responsible for predicting the properties of the required results the results of action acceptor. The structures of the results of action acceptor, excited by the dominant motivation, dynamically leave an impression of the activity these are the properties of the required

results and the means of achieving them. As a result, the subjects behavior acquires a character which is directed at achieving the needed objects in the external world. These processes determine the movement of the hungry subject to food and direct the collector to the next item for his collection. The goal reflex proposed by Pavlov has acquired new features and new concrete physiological content from functional systems theory. Motivations are dynamically fulfilled during an individuals life. In the systems organization of behavioral acts, subjects are enriched by the materials and methods of satisfaction of the underlying needs. In 1927, Ukhtomskii wrote: The most life-enriched character is the person whose attention is open to ongoing reality, ready to take action, whatever it might be, the enthusiastic seeker of truth, who does not seize the initially selected coordinate axis, because he understands their relativity and, to the end, the final moment, does not stick to positions which seem to be the most economically favorable, but goes on and on, but persists in considering inadequately evaluated details and might joyfully use these details to accept radical changes in the initial pathway. This again is the dominant we might say the youth dominant in which nothing is yet the victim of sclerosis or necrosis, but is a wide life open to what is ahead ([29], p. 312). The goal reflex about which Pavlov wrote and spoke reorganizes itself in such a way as to construct, within the overall systems organization, the dominant motivation.

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