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Part II. The Tuning of the Central Nervous System and its Psychological Implication
Summary. The activation of the ergotropic or trophotropic system regardless of the procedures through which it is accomplished results in an alteration of the reactivity of the central nervous system for which the term ergotropic or trophotropic tuning is used. Three states of tuning induced by increasing degrees of activation are distinguished. At the first stage of ergotropic tuning the tone and reactivity of the ergotropic system is increased while that of the trophotropic system is reciprocally inhibited. Corresponding changes in reactivity occur during the tuning of the trophotropic system. At the second stage of ergotropic tuning reversal phenomena occur: a stimulus which under control conditions evokes a trophotropic response elicits an ergotropic response in the tuned state. Thus petting produces a rage reaction in the ergotropically tuned state and neutral stimuli which do not evoke any reactions under control conditions evoke ergotropic and trophotropic responses respectively in states of ergotropic and trophotropic tuning. The alterations in the reactivity of the central nervous system in the tuned state is not restricted to inborn (unconditional) reactions but is also demonstrable in response to conditional stimuli. Whereas the activity of the ergotropic and trophotropic systems shows reciprocal relations in stage I and II of tuning, stage I I I is characterized by simultaneous increased ergotropic and trophotropic discharges. It is believed that at this high degree of ergotropic tuning the ergotropic excitation spills over into the trophotropic system. At the same time pathological symptoms (neurosis, hallucinations) appear. Although changes in the internal environment induced by hypothalamic-hypophyseal discharges modify states of tuning and emotional behavior the chief factor determining mood and emotion is the ergotropic-trophotropic balance. Zusammen/assung. Die Aktivierung des ergotrophischen oder trophotrophischen Systems durch versehicdene Prozeduren ffihrt eine Anderung der Erregbarkeit des Zentralnervensystems herbei, die als ergo- oder trophotrophische Umstimmung bezeichnet wird. Mit stcigender Reizung lassen sich drei verschiedene Stadien solcher Umstimmung unterscheiden. Das erste Stadium der ergotrophischen Umstimmung ist durch erh6hten Tonus und Erregbarkeit des ergotrophischen Systems und gleichzeitiger Hemmung des trophotrophischen Systems charakterisiert. Entsprcchende Andertmgen der Erregbarkeit werden im Zustand der trophotrophischen Umstimmung beobachtet. Mit st/~rkerer Reizung des ergotrophischen Systems-im zweiten Stadium der ergotrophischen Umstimmung - - treten Umkchrungsvorgiinge (reversal) auf. Ein Reiz, der im Kontrollversueh einen trophotrophischen Erfolg hat, ruft eine ergotrophische Antwort unter den Bedingungen der ergotrophischen Umstimmung hervor. Streicheln zum Beispiel ruft im zweiten Stadium der ergotrophischen Umstimmung eine Wutreaktion hervor, und neutrale Reize, die im Kontrollversuch keine Wirkung ausfiben, rufen w/ihrend ergotrophischer Umstimmung eine ergotrophische und w/~hrend trophotrophischer Umstimmung eine trophotrophische Reaktion hervor. Die Ver/inderungen der Erregbarkeit gelten auch fiir bedingte Reize. Im Gegensatz zu dem ersten und zweiten Stadium der Umstimmung, in dem ergotrophische und trophotrophische Prozesse Reziprozit/~t zeigen, treten im dritten Umstimmungsstadiumergotrophische und trophotrophische Entladungen gleichzeitig auf. Es wird angenommen, dab in diesem hohen Grade yon Umstimmung die ergotrophische Erregung in das trophotrophische System iiberflieBt. Gleichzeitig treten pathologische Symptome auf (Neurose, Halluzinationen). Obwohl Jinderungen im imleren Milieu durch hypothalamische und hypo5*

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physeale Entladungen verursacht werden und die Umstimraung und emotionale Erregbarkeit beeinflussen, wird das durch nervSse 1%ozesseregulierte ergotrophischtrophotrophische Gleichgewicht als Hauptfaktor angesehen, der Stimmung und Gefiihlserregung bestimmt.

It was shown in the preceding section that stimuli applied to peripheral receptors or to central structures at various levels of the nervous system from which autonomic responses can be evoked, alter the state of activity of the somatic nervous system at the same time. Autonomic and somatic (motor and sensory) functions are integrated, particularly at hypothalamie and reticular levels and, as the result, ergotropic and trophotropie syndromes are produced. I t will now be shown that when such syndromes are evoked the responsiveness of these systems to various stimuli is altered in a characteristic manner. In order to facilitate the description of these changes the concept of tuning is introduced. This concept clarifies and unifies these changes and makes understandable the fundamental alterations which the organism undergoes when the activity and balance of ergotropic and trophotropic systems is altered. Enhancing the activity of the ergotropic or of the trophotropic systems is said to result in the tuning of the ergotropie and trophotropic system respectively. This state of tuning appears in three different forms which depend on the degree of ergotropie or trophotropic excitation. Under conditions of mild excitation of the ergotropie system, the reactivity of the various parts of the ergotropic system is increased and that of the trophotropic system is reciprocally diminished. Conversely, excitation of the trophotropie system induces increased responsiveness of the trophotropic system while the reactivity of the ergotropic system is reciprocally diminished. When the ergotropic system is activated to a higher degree its reactivity is still further enhanced but that of the trophotropie system is virtually abolished. Stimuli which evoke trophotropic responses under control conditions, elicit at this stage (II) an ergotropic response: a reversal of effects takes place. Finally, at the highest degree of tuning (III) the excitation of the ergotropie system is so great that it spills over into the trophotropic system. The reciprocity relation which characterizes the ergotropic-trophotropie balance under physiological conditions is lost at stage III.

Tuning, Stage I
The changes induced by ergotropic tuning are illustrated for the sympathetic division of the ergotropic system by the increased response of blood pressure, heart rate and nictitating membrane to stimulation

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of the posterior hypothalamus when the pressure in the carotid sinus is lowered. A corresponding rise in excitability of the parasympathetic division of the trophotropic system in response to a trophotropically acting stimulus is demonstrable when a state of trophotropic tuning is induced by a rise in sino-aortic pressure. States of trophotropic tuning m a y also be evoked by stimulation of afferent somatic nerves such as the sciatic with low frequency currents whereas a state of ergotropic tuning results from sciatic excitation with currents of higher frequencies. Even subthreshold currents applied to the hypothalamus alter the state of tuning as indicated by the change in excitability of autonomic
reactions.

Likewise, the somatic components of the ergotropic and trophotropic systems show the effect of tuning. A heightened excitability of the motor and sensory systems is demonstrable in states of ergotropic tuning (Murphy and Gellhorn, 1945; Gellhorn, Koella and Ballin, 1954, 1955). Stimulation of the hypothalamus at sites and intensities which induce sympathetic effects but no somatic (motor) action greatly increases any movements elicited by stimulation of the motor cortex. Similarly, stimulation of the posterior hypothalamus facilitates the responsiveness of the auditory and visual projection areas to acoustic and optic impulses. These effects occur even outside the confines of the motor cortex and the specific projection areas. They are thought to be due to the interaction between diffuse hypothalamie-cortical discharges and the excitation of cortical motor 1 and sensory neurons activated by electrical stimulation and sensory impulses respectively. Reciprocal relations between the ergotropie and trophotropic systems are characteristic for stage I of tuning. This is illustrated for the autonomic components of these systems by the fact that the sympathetic test stimulus is less effective in a state of parasympathetic tuning than under control conditions; similarly, the action of the parasympathetic test stimulus is reduced in the state of sympathetic tuning (Gellhorn, 1957). The validity of the reciprocity principle for the somatic components of the ergotropie-trophotropic systems in states of ergotropic tuning is apparent from a study of the excitability of the trophotropic system. A measure of this excitability is the "spindle" which may be evoked in the cortex through a single shock applied to the caudate nucleus. The threshold of this synchronizing effect increases with increasing activity of the ergotropie system regardless of whether the state of ergotropic tuning is intensified through stimulation of the reticular formation, the posterior hypothalamus or by other procedures (Moruzzi and Magoun, 1949; Tokizane, Kawakami and Gellhorn, 1957a and b). 1 In addition, a similar facilitation takes place at the spinal level (Rhines and Magoun, 1946; see also Gellhorn, 1957, p. 175).

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Even slight changes in the state of arousal which occur spontaneously may induce significant changes in the state of tuning. An interesting interdependence existing between the state of the hypothalamus, ergotropic and trophotropic reactivity, and awareness is illustrated by experiments on heating the anterior hypothalamus which induces the trophotropic syndrome: synchronization of cortical potentials, relaxation of muscles, increased parasympathetic discharges etc. (yon Euler and SSderberg, 1957). In this state of trophotropic tuning the trophotropic reactivity is heightened ~ and the ergotropie reactivity is lessened: panting and vasodilation are produced whereas the ergotropic response to environmental cold is greatly reduced. A state of trophotropie tuning also accounts for the fact that panting is more easily induced in drowsy than in alert goats (see Gellhorn, 1967 for the literature). On the basis of these and many related observations discussed by Gellhorn (1967-1969) it may be said that ergotropic tuning is characterized by an increased tone and responsiveness of the ergotropie system and by a diminution of tone and excitability of the trophotropie system. Corresponding changes characterize the tuning of the trophotropic system. The ergotropic-trophotropie balance is shifted to the ergotropic and trophotropic sides respectively in states of ergotropic and trophotropie tuning.

Emotion and Tuning

The chief question to be dealt with in this paper concerns the validity of the laws of tuning for the emotions and related behavioral states. To decide this question it is proposed to investigate whether emotions which primarily involve the ergotropic system cause an increase in the responsiveness to ergotropically acting test stimuli and a decrease in the responsiveness to trophotropically acting test stimuli and whether corresponding changes in ergotropic and trophotropic excitability occur when trophotropic emotions are evoked. I t is also expected that a shift in the ergotropic-trophotropic balance regardless of the procedures by which it is induced (via reflexes, direct stimulation of supraspinal structures such as the hypothalamus, or as the result of central lesions) would alter the emotional state in a more or less predictable manner. The study of these problems is beset with great difficulties. Only a rough classification of some emotions in terms of ergotropic or trophotropic involvement is possible. Some intensive emotional states (anxiety, 2 See also Weinberger, Velasco and Lindsley (1967) for the validity of this statement for non-reinforced conditional stimuli which are known to activate the ~rophotropic system (see Gellhorn, 1967, Chapter III).

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Gellhorn, 1965) affect the ergotropic and the trophotropic systems and be dealt with later. Differentiation between pleasant and unpleasant emotions is of no value since both types m a y primarily affect the same system. Postprandial happiness and simple depression show trophotropic dominance whereas the feeling of triumph or extreme happiness as well as the state of rage and aggressiveness are associated with excitation of the ergotropic system. The tone of the striated muscles seems to be a better means of classification since its increase and decrease induce ergotropic and trophotropic excitation respectively (see Gellhorn, 1970). Bearing in mind the reaction of the muscles, the prevalent autonomic discharges, and the behavioral effects resulting from stimulation of the trophotropic and ergotropic division of the hypothalamus, it seems justified to look upon grooming, purring, etc. in the cat as a trophotropic, and rage and aggressiveness as an ergotropic reaction. I n man sudden fear elicits mainly trophotropic effects (often followed by compensatory discharges of the ergotropic system) a whereas states of frustration, resentment and rage evoke chiefly ergotropic effects. I f aggressiveness is a reflection of the tuning of the crgotropie system, one should expect t h a t the ergotropic reactivity is enhanced in this state. Experiments in which the toxicity of amphetamine was determined showed t h a t aggressive mice 4 manifest an increased sensitivity to amphetamine (Consolo et a~., 1965). The toxicity of amphetamine is likewise increased in mice kept in groups in a small space, suggesting t h a t the heightened emotional reactivity of aggregated mice is likewise related to an increased responsiveness of the ergotropic system. Moreover, the release of noradrenaline from the brain b y amphetamine is greater in crowded than in non-crowded animals and drugs which alter the release of noradrenaline cause parallel changes in the toxicity of amphetamine in aggregated animals (Moore, 1963, 1964). These findings seem to indicate a heightened ergotropic reactivity in aggressive animals and in crowded conditions and, consequently a shift in the ergotropictrophotropic balance to the ergotropic side. This alteration in balance is evident from the lessened sleeping time of aggressive as compared to normal mice which results from the administration of barbiturates or barbiturate-chlorpromazine mixtures (Consolo, 1965). Although the study of the action of amphetamine on aggressive and aggregate animals furnishes evidence for the existence of ergotropic tuning under these conditions the global action of this drug does not lend itself to a finer analysis; but such an analysis can be achieved through observations on aggression elicited in the unanesthetized cat b y hypo3 For an analysis of acute and subacute states of fear and their differentiation from anxiety see Gellhorn (1965). 4 Aggressiveness was produced by prolonged isolation.

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thalamic stimulation. Since the effect of rage and aggression on the autonomic nervous system has been investigated extensively by Cannon, Bard, and m a n y others, the changes occurring in somatic functions described b y Flynn (1967) are of particular interest. He placed a cat and a rat in the same cage and used for this work only cats which did not attack rats in the absence of hypothalamie stimulation. This stimulation resulted in a prompt biting attack. However, this attack did not occur when the sensory branches of the trigeminal nerves which supply the area around the mouth were cut. Instead of biting the rat, such eats " r u b b e d their muzzles back and forth over the r a t " . Apparently, there is a central interaction between afferent somatic discharges from the mouth area and hypothalamic stimuli which results in the biting attack. Touching the lips during hypothalamic stimulation induces opening of the mouth followed b y a strong bite. The cutaneous area from which this effect can be ehcited increases with increasing hypothalamic stimulation. A similar interaction takes place when hypothalamus and a part of the motor nucleus of the Vth nerve are stimulated simultaneously: the closure of the jaw is more complete although the hypothalamic stimulus per se did not evoke a motor effect. These experiments confirm Gellhorn's (1954) work on the facilitation of sensory action through hypothalamic stimulation, and Murphy and Gellhorn's study (1945) on a similar facilitation of motor discharges. They demonstrate the increased responsiveness of the sensory and motor components of the ergotropic system in aggression. Ejaculatory speech, expressing anger and joy reported b y Wilson (1940) in eases of aphasia seems to illustrate in m a n the enhanced responsiveness of the motor system in the state of ergotropic tuning. The following experiments illustrate in m a n the change in the reactivity of the autonomic components of the ergotropic and trophotropic systems in emotional states. Immersion of the hand in cold water is accompanied by prolonged vasoconstriction but this phase is interrupted one or more times b y brief periods of dilatation (Lew/s, 1930). This phenomenon is apparently related to the trophotropic rebound which follows an ergotropic excitation as discussed on p. 55. I f this were the case one would expect t h a t a change in the emotional state i.e. in the ergotropic-trophotropic balance would quantitatively alter the trophotropic reaction represented b y the vasodilation. Emotional arousal (anger) should intensify ergotropic and lessen trophotropic reaction. The lessening or disappearance of the periods of dilatation under these circumstances (Teichner, 1966) confirms our interpretation. Wolf's study of the vagal reflex induced by immersion of the face in cold water reveals t h a t this reflex is altered by the emotional state of the experimental subject. I n a state of fear the vagal reflex is greatly

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increased indicating the enhanced trophotropie reactivity in the trophotropically tuned state. Contrariwise, harassing a subject, a procedure known to induce a state of ergotropie tuning, may delay or abolish the immersion reflex, thus exemplifying reciprocal inhibition of the trophotropic system in this state. I t was shown in the preceding pages that various emotions such as fear, resentment, and rage are associated with characteristic changes in ergotropie and trophotropie reactivity which imply a shift in the ergotropic-trophotropic balance since the reciprocity between ergotropie and trophotropie processes is preserved at this stage of tuning. The following data show indeed that sufficiently marked changes in ergotropic-trophotropic balance, induced by intracerebral stimulations or lesions, evoke emotions which are fundamentally different in states of ergotropic and trophotropie dominance. Shifting the ergotropic-trophotropie balance to the trophotropic side occurs following amygdalectomy and leads to an "increase in pleasure reactions to stroking and petting" and to a rise in the threshold of the aggressive reaction in response to painful and emotionally arousing stimuli such as barking dogs (Kling and Hurt, 1958). Similar effects are produced by stimulation of the caudate nucleus, or of the septum and preoptie area, with currents of low frequency. In aggressive monkeys stimulation of the latter areas often elicits penile erection, afterdischarges in the hippocampus, and a change in mood toward placidity which may last for hours (MacLean, 1962). That these effects are based on a trophotropie shift is supported by the observation that shivering or excitement (indicating a state of ergotropic tuning) lessens penile erection in response to septal stimulation (MacLean and Ploog, 1962). On the other hand, rage-like reactions and emotional hyperactivity result from excitation of the posterior hypothalamus (Hess, 1949). A similar effect is obtained when the tonic activity of the trophotropie system is diminished. Thus, a release of the ergotroloic system follows bilateral lesions in the septal area (Brady and Nauta, 1953), or in the ventromedial nuclei of the hypothalamus (Wheatley, 1944). This shift in the ergotropic-trophotropie balance to the ergotropie side can be also accomplished in animals in which trophotropic dominance and placidity has been produced by amygdalectomy. Thus amygdalectomized oats displaying marked trophotropic pleasure reactions, show a reversal in emotional behavior and react, following ventro-medial hypothalamie lesions, with rage in response to minimal stimuli (Kling and Hurt, 1958). Release of the ergotropie system via reflexes induces similar emotional effects. The lowering of the pressure in the sine-aortic area reduces the tonic discharges of the baroreceptors and evokes attacks of sham rage

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in the decorticate cat ~ (Guazzi, Malliani and Zanchetti, 1964). Well directed attacks of rage in response to a nociceptive stimulus are abolished in the nnanesthemonkey when the pressure in the carotid sinus is raised (Koch, 1932). I t may, therefore, be said t h a t sham rage and rage are intensified by lowering, and inhibited b y increasing the sino-aortic pressure and that the former effect is associated with signs of the release and the latter with signs of inhibition of the ergotropie system and corresponding changes in the ergotropic-trophotropic balance. I n spite of large gaps in our knowledge it seems to follow from the evaluation of the experiments presented in this and earlier papers (Gellhorn, 1967-1969) t h a t a shift in the ergotropic-trophotropic balance due to stimulation of peripheral nerves or central structures, such as the hypothalamic and the limbic system, alters the emotional state. Similar changes in balance and in the emotional state occur as the result of a release of the ergotropic system from the tonic inhibitory action of the trophotropic system and vice versa. The heightened excitability of the stimulated or released system, and the lessened excitability of the antagonistic system, account for the physiological and psychological changes induced by the tuning of the ergotropic or the trophotropic system.

Reversal: The Second Stage o/ Tuning

Reflexes, stimulation of central structures such as the limbic areas and the hypothalamus, or lesions m a y induce states of tuning in which the responsiveness of the ergotropic and trophotropic systems is altered not only quantitatively but also qualitatively: Whereas in the control test an ergotropic hypothalamic stimulus evokes a rise in blood pressure and heart rate it m a y produce a fall in pressure and rate after an adequate state of trophotropic tuning has been produced through increased discharges from the baroreceptors. Tuning, stage I and I I seem to be based on summation processes. In stage I the increased central ergotropic discharges in ergotropic tuning summate with the ergotropic excitation evoked b y the ergotropic test stimulus. The result is a greater excitation and reactivity than under control conditions. The opposite holds for the effect of a trophotropic test stimulus in ergotropic tuning since due to reciprocal innervation the central trophotropic discharges are diminished. With the further increase in tuning in stage l I the reciprocally inhibited trophotropic receptors become virtually unresponsive. Now it has to be borne in mind t h a t our test stimuli activate both ergotropic and trophotropic systems as seen by the fact t h a t low and high frequency stimulations of a peripheral or central structure tend to 5 A similar effect is produced by stimulation of the chemoreceptors of the sino-aortic area.

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produce trophotropic and ergotropic effects respectively (see Gellhorn, 1967, Chapter I for the literature and Roitbak and Eristavi, 1966). However, due to the reciprocity principle, only one effect prevails which is determined by the system which is in a state of increased activity (tuned). From the application of these principles it follows that a trophotropie stimulus induces an ergotropic effect when the trophotropic excitability is virtually nil and the ergotropic reactivity is great, as in the second stage of ergotropic tuning. Corresponding changes account for the reversal, in the second stage of trophotropic tuning, of the action of stimuli which induce ergotropic effects under control conditions. The following discussion will show that tuning-induced states of reversal can be demonstrated not only in autonomic reflexes but in more complex reactions of the ergotropic and trophotropic systems. This work contributes to our understanding of the physiology of the emotions and of behavior in general. Fundamental changes in the response of the E E G to light in different states of awareness seem to illustrate tuning and reversal. The blocking of alpha potentials through light is thought to be due to desynchronization of cortical potentials and is part of the ergotropically mediated arousal. The reverse effect, production of alpha potentials (due to synchronization) in response to the same stimulus, suggests that a shift to the trophotropie side has taken place. The fact that this phenomenon occurs in the drowsy state induced by hypnosis, as the result of deprivation of sleep, and also in narcolepsy supports this interpretation (Armington and Mitniek, 1959; Goldie and Green, 1960; Malmo, 1959). Numerous examples of reversal were demonstrated by Pschonik (1956) in man and experimental animals on the basis of conditioned vasomotor reactions induced by two conditioned stimulus (CS) unconditioned stimulus (US) combinations such as Bell -~ 63 ~ C (A) evoking pain and vasoconstriction and Light-~43~ C (B) evoking vasodilatation and warmth. A striking reversal occurs when gradually the temperature of the thermode serving as US is lowered in A to 43 ~ C and raised hi B to 53 ~ or 63 ~ C. In these tests the US (43 ~ C) causes vasoconstriction and pain when applied after the Bell as in A and the US (63 ~ C) causes vasodilatation and warmth when applied after the Light as in B. In other words, the US reinforces the vasomotor reflex and the sensation which had been evoked by the preceding CS as if the US had been unchanged. As this work shows, it should be amphasized that the concept of tuning is applicable to a wider area than was first anticipated. Its validity for conditional processes implies that experiential factors in general may produce tuning. Further, autonomic reversal is associated with a reversal of sensory perceptions. Obviously, in these experiments

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a stage has been reached which borders on pathology. These matters will be pursued in the discussion of the third stage of tuning. As was mentioned earlier, ventromedial hypothalamic lesions release the lateral hypothalamus. Even the mildest stimulus (touch) elicits a rage reaction. Pleasurable reactions are absent (Wheatley, 1944). Obviously a state of heightened ergotropie tuning exists in which any stimulus evokes only an ergotropie response. Similar is the effect of frontal lesions which likewise tend to release the ergotropic system (Brutkowski, 1963). A reversal in an ergotropieally tuned animal is also seen on near threshold stimulation of the amygdala (Delgado, 1964). Although this stimulus does not evoke any overt changes in behavior the cat reacts to petting, not with rubbing the head against the hand of the observer, but with aggressiveness. This illustrates t h a t amygdaloid stimulation induces ergotropic tuning and reverses the petting response: a trophotropic reaction is replaced by an ergotropic one. Ergotropic reversal reactions have been observed in phobic states indicated by the fact t h a t positive and negative conditional stimuli produce a strong psyehogalvanic reflex (Alexander, 1961), whereas this effect is restricted to positive conditional stimuli under control conditions. (It has been shown t h a t positive and negative conditional stimuli activate the ergotropic and trophotropic systems respectively, Gellhorn, 1967). Reversal of behavior in a trophotropically tuned state is illustrated by the following experimental and clinical findings. Most afferent stimuli contribute to arousal. The removal of the major sense organs tends therefore to shift the ergotropic-trophotropic balance to the trophotropic side. Thus, animals deprived of the organs of smell, vision, and hearing show an increased reactivity of the trophotropie system indicated by a lesser general activity, an increase in the duration of sleep (Hagamen, 1959), and a marked acceleration of extinction (by lack of reinforcement) of the conditional reaction (CI~) (Koranyi, EndrSczi and Lisss 1963). Such animals show behavioral reversal. Nociceptive stimuli, which in the normal cat induce ergotropic effects, exert trophotropic actions in cats which are blind, deaf, and anosmic. " W h e n a cat was placed in the pen, sleep occurred more quickly after a strong noxious stimulus than when the cat was not so stimulated ''s (Hagamen, 1959). 6 Lindsley et al. (1964) showed that monkeys deprived of light for years show reversal of cortical (and hippocampal) reactions to light. Light induces the production and darkness the disappearance of alpha potentials. I suggest that the former is a reversal of the normal alpha blocking effect of light due to the trophotropic tuning of the nervous system resulting from light deprivation.

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Here are some clinical examples of trophotropic reversal phenomena: 1. Conditioned reflex (C1~) studies in which a sound serves as the CS and a shock to the finger as the US show in depressions t h a t the CS produce synchronization in the E E G instead of desynchronization as found in normal persons. 2. Emotional excitement (laughter, orgasm) m a y precipitate a cataplectic attack (loss of muscle tone) in cases of narcolepsy (see Gellhorn, 1967, for further details). The following three experiments illustrating reversals were chosen to show t h a t : 1. the same stimulus evokes totally different but completely reversible behavioral reversal reactions in the ergotropic and trophotropic states of tuning; 2. the phenomenon is demonstrable at the hypothalamic level for widely different types of behavior. Kopa, Szabo and Grastyan studied cats which had learned to escape a shock from the floor grid by jumping onto a ledge. When placed on the grid, the cat was restless and apprehensive and obviously in a state of heightened ergotropic activity. On the contrary, the cat was quiet on the ledge where it had never been shocked and sometimes even showed grooming and purring (indicating a state of trophotropic tuning). If in these two situations reflecting different experiences the cat's centrum medianum 7 is stimulated, the effect is, in the grid situation, one of increased arousal leading to a reaction as seen in avoidance conditioning: the eat jumps onto the ledge; but in the ledge situation, the same stimulus evokes relaxation, lowering of the head,, and sleeping posture. The second example of reversal demonstrates fundamental alterations in neurophysiological and behavioral responsiveness in hunger and satiety which represent ergotroic and trophotropic states respectively regulated b y the lateral and ventromedial hypothalamus s. When hungry the cat in which an alimentary CR had been established responds to the CS with desynchronization in the E E G , a negative shift in the cortical d.c. (direct current) response and feeding. If, however, through frequent application of the CS reinforced b y feeding, a state of satiation is reached the E E G , the d.c. response and the behavior are reversed. Synchronization in the E E G and a positive shift in the d.c. response occur while the animal refuses to eat (Rowland and Goldstone, 1963). 7 Later Grastyan (1968) stated that the pretectal area was stimulated; in earlier work the hypothalamus was involved. 8 See Grossman, 1967 for the literature.

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I t is suggested t h a t a shift from ergotropic to trophotropic dominance and vice versa is accompanied by electrophysiological and behavioral reversal. The third experiment (Brown, Hunsperger and Rosvold, 1969) is a physiological model of the tuning process. These authors stimulated hypothalamic points which produced chiefly ergotropic (aggressive or arousing) and trophotropic (sniffing) reactions. If the cat was presented with a d u m m y cat the d u m m y was ignored in the absence of hypothalamic stimulation. Stimulation of the ergotropic points resulted in intensive attacks on the dummy, but t h a t of the trophotropic points produced sniffing which was directed not only to the nose as seen on hypothalamic stimulation in the absence of a d u m m y but also to other parts of the dummy. As Brown et al expressed it: "When bhe hypoJhalsmus is not stimulated the d u m m y is usually regarded as a neutral object; when, however, stimulation is applied, the d u m m y is viewed as a menacing object to be fended off or attacked, or a friendly object to be greeted. Thus, the significance of a social stimulus m a y be determined b y selective activation of a particular site in the hypothalamus." I t was shown earlier t h a t hypothalamic stimulation enhances cortically induced motor action (Murphy and Gellhorn, 1945) and the effect of sensory impulses on the cortex within and without the sensory projection areas (Gellhorn, 1954). During rage and attack a similar facilitation of motor and sensory processes takes place which is thought to be due to the increased ergotropic discharges under these conditions (Flynn, 1967). This interaction between hypothalamic and somatic excitation is the basis of the tuning process. I n addition, a shift of hypothalamie excitation from trophotropic to ergotropic sites and vice versa determines type of tuning and reversal as in Hunsperger's experiment 9. The work of Brown, ttunsperger and Rosvold and t h a t of Kopa et al. needs repetition and expansion. At present it strongly suggests t h a t the behavioral response even to a neutral stimulus 1~ is determined b y the state of excitation of the ergotropic and trophotropic systems and, particularly, by the ergotropic-trophotropic balance. 9 It is probable that even subthreshold hypothalamic stimuli have similar effects since such stimuli produce states of ergotropic and trophotropic tuning (Gellhorn, 1957, 1959). The experimental verification of this assumption for the emotions would be valuable since weak, near threshold stimuli applied repeatedly over long periods are apt to cause states of ergotropic-trophotropic imbalanet in the human patient. 10 i.e. a stimulus which is ineffective (as the sight o5 the dummy withouc hypothalamic stimulation) or as in Kopa's experiment evokes only an insignificant orienting reaction.

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Tuning, Stage I I I
In contradistinction to the tuning in stage I and II in which the ergotropic and trophotropic systems are reciprocally related the stage III occurring at still higher degrees of tuning is characterized by simultaneous ergotropie and trophotropie discharges. This is illustrated by the behavior of animals as they pass from the phase of conditioning to that of experimental neurosis. Whereas during conditioning ergotropic activity is increased but behavior normal, a fundamental change occurs when differentiation of conditioned reflexes is carried too far. Then a variety of somatic symptoms indicates neurotic behavior, the ergotropic discharges persist and, in addition, strong trophotropic symptoms such as licking, retching, vomiting and diarrhea become prominent. It is thought that the "spilling over" of crgotropic excitation into the trophotropic system is responsible for these symptoms and the pathological behavior. This interpretation is supported by the following facts: 1. Spontaneous cures of neuroses are accompanied by the disappearance of the abnormal trophotropic symptoms. Moreover, benaetyzine which is a tranquilizer with strong anticholinergic effects (Shcrrod, 1963) relieves experimental neurosis while at the same time, symptoms of strong trophotropic discharge (nausea, vomiting) disappear. Since only those derivatives of benactyzine which exert central anticholinergic effects are able to restore neurotic animals (Holten and Sonne, 1955) the data suggest that these drugs counteract the spilling over of ergotropic excitation into the trophotropic system and contribute thereby to behavioral resitution. 2. Inescapable shock leads to experimental neurosis with massive trophotropie discharges (Overmicr and Se]igman, 1967). 3. Rats forced to s ~ m in conditions in which neither a rest (through floating) nor an escape was possible, died after relatively brief periods. The heart rate first accelerated but declined markedly later. At death there was a standstill of the heart in diastole. Cholinergic drugs aggravated and atropine retarded these effects (Richter, 1957). These data and the finding of gross hypothalamic lesions at autopsy suggest that in this condition intensive trophotropic discharges are associated with states of strong ergotropic excitation and that the hypothalamic system is primarily involved. Clinical and experimental studies have demonstrated in man a pathological state of ergotropic tuning with enhanced ergotropic and trophotropic discharges: 1. Neurotic patients show a combination of ergotropic and trophotropic symptoms as seen in animals with experimental neurosis. 2. If after establishment of conditional vasodilator and vasoconstrictor reflexes the CS for warmth and pain were applied in quick succession,

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the vasodilator and constrictor responses were abolished while nearneurotic symptoms appeared. The disappearance of these vasomotor reactions seems to be due to the simultaneous activation of the ergotropic and trophotropic conditional reflexes. When, after a period of rest, the neurotic symptoms improved tests showed that the vasomotor CI~ had been restored [see Gellhorn's (1967) interpretation of Pschonik's (1956) work]. 3. E E G studies on spontaneous and experimentally induced delirium offer striking evidence for the simultaneity of ergotropic and trophotropic discharges. Ditran produces an increase in the slow (theta and delta) as well as in the fast (beta) potentials. The former is primarily associated with clouding of consciousness, the latter with increased psychomotor activity (Itil and Fink, 1966). The participation of both systems in the production of delirium is supported by the findings that improvement of the delirium is associated with a decline of slow and fast potentials in the E E G and "a progressive increase in the amount of alpha activity". 4. Certain amphetamine derivatives produce in man hallucinogenic effects while amphetamine causes only alerting. Quantitative E E G studies of these drugs in unanesthetized cats disclose that the hallucinogenic drugs cause, in addition to strong ergotropic effects, a synchronizing action on the cortex whereas amphetamh~e induces desynchronization (Fairchild, Alles, Jenden and Mickey, 1967). I t seems not unlikely that the differential effects on E E G and behavior are due to the fact that amphetamine activates the ergotropie system only, whereas the hallucinogenic derivatives activate the ergotropic and trophotropie systems with the latter dominant on the EEG. Since pathology often involves physiological reactions under quantitatively altered conditions the question arises as to whether a lack of reciprocity between the ergotropic and trophotropic systems occurs under certain physiological conditions. I suggest that the paradoxical state of sleep 11 represents such a state. Judged by the fall in the blood pressure and the complete loss of tone of the neck muscles this state shows very intensive trophotropic discharges. On the other hand the desynchronization in the EEG, the appearance in the hippocampus of the theta rhythm characteristic for arousal, and the response of action potentials in the visual cortex to optic nerve stimulation disclose, particularly during the phase of rapid eye movements, strong ergotropic activity, equal to, or surpassing that, seen during awakening (Okuma and Aklmoto, 1966). Apparently, simultaneous ergotropic and trophotropic discharges which occur in neurosis and in experimentally 11 See numerous symposia on sleep and Gellhorn (1967b) for the literature.

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induced abnormal mental states such as delirium are also seen in the paradoxical phase of sleep. It does not seem accidental that under these circumstances dreaming takes place which although a physiological process definitely involves gross deviations from normal perceptions. A borderline case between physiology and pathology seems to be the state of a person subjected to a prolonged vigil. As pointed out elsewhere (Gellhorn, 1967b) various tests show that ergotropic and trophotropie discharges are greatly increased and here again this condition is associated with mental symptoms such as hallucinations, illusions, disorientation and temporal and cognitive disturbances.

Tuning, Hormones and the Hypothalamus

In 1940 Bard reported that vaginal stimulation elicits a sexual reaction consisting of crouching and treading followed by after-effects involving squirming and purring in the estrous cat. On the contrary, when not in heat the same stimulus evokes "angry vocalization and various forms of struggling". I suggest that under the influence of gonadotropic hormones the secretion of which is regulated by the hypothalamus, the cat in heat is in a state of trophotropic tuning and responds to sexually arousing stimuli with acceptance of the male and typical trophotropic reactions such as purring whereas during anestrus ergotropic effects prevail. As the anestrous state is changed into the estrous state the ergotropie-trophotropic balance is shifted to the trophotropic side and a reversal phenomenon occurs similar to those described earlier in which this shift was effeeted by alterations in neuronal discharges. This interpretation is supported by the fact that: 1. vaginal stimuli under optimal hormonal conditions (injection of estrogen and progesterone) activate anterior hypothalamus and preoptic area (Porter et al., 1957); 2. mating is abolished by lesions in this area; 3. injection of stilbesterol in this but in no other parts of the hypothalamus elicits mating in ovariectomized cats (Michael, 1965) ; 4. gonadotropic hormones which induce mating cause changes in ergotropic and trophotropic thresholds indicating an early increase and a late decrease in excitability (Kawakami and Sawyer, 1959; Sawyer, 1960). Since the changes in the trophotropic reaction resembling the paradoxical phase of sleep are greater than those involving ergotropic arousal it is suggested that these hormones cause a shift in the trophotropie-ergotropic balance to the trophotropie side which accounts for the reversal phenomenon described above. This interpretation is supported by the fact that this trophotropic-reaction is blocked by lesions in the ventromedial nucleus of the hypothalamus (Kawakami, 1966) and elicited by low
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frequency stimulation of the septum (Sawyer, Kawakami and Kanematsu, 1966). Moreover, injection of progesterone into the preoptic area induces sleep (Heuser, Ling and Kluver, 1967). The relation of the hypothalamus to the endocrine system is complex and cannot be reviewed here. Suffice it to say that the secretion of pituitary hormones is regulated by specific parts of the hypothalamus through neurohumoral releasing factors. In addition, other areas of the hypotha]amus sense the titre of the hormones in the blood and maintain homeostasis by negative feed-back. Nevertheless, diurnal variations of hypothalamica]ly regulated hormones exist and their concentration in the blood shows marked changes under the influence of different forms of stress. Since these hormones alter brain functions in general and hypotha]amic activity in particular (Beyer et al., 1967; Woodbury, 1954) it is probable that the reactivity of the ergotropic and trophotropic systems is altered when endocrine activity and balance are changed. This is illustrated by the following data. When a frog is placed into the cage of a lactating rat and her litter, the rat kills the frog. Since this reaction does not occur in non-lactating females or in lactating females in the absence of the litter (EndrSczi, Lissak and Telegdy, 1958) i~ seems that the excitability of the crgotropic system is sufficiently raised under the influence of hormonal and neural (optic ?) stimuli to produce aggression. The importance of the hormonal balance for " m a t e r n a l " aggressiveness is evident from the fact that oestrone treatment reduces this form of aggressiveness and may even convert it into a fear reaction, suggesting a shift from crgotropic to trophotropic dominance since trophotropic tuning is characteristic for fear (Gellhorn, 1965). The gonads have also a profound influence on aggressiveness in males. Thus, aggressiveness in male rats kept in isolation for a long period, is abolished by castration whereas administration of male hormones enhances aggressiveness in various species and raises the position in the hormone-treated animals in the social hierarchy (see Moyer, 1968, for the literature). From the extensive literature showing that adreno-eortical hormones are liberated following emotional excitement and stimulation of hypothalamus and limbie system (see Mason, 1968, for the literature) only one group of data particularly important for the problems discussed in this paper may be mentioned. It concerns the relation between the secretion of the 17-hydroxycorticosteroids (17-OH-CS) and the activity of the ergotropic system and shows that: 1. Aggressive animals excrete more 17-OH-CS than non-aggressive cats;

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2. Conditioning enhances ergotropic activity and corticoid secretion whereas non-reinforced CS inhibit these functions; 3. Stimulation of the anterior hypothalamus decreases and that of the posterior hypothalamus increases the corticosteroid concentration in the blood (EndSczi, Hartmann and Lissak, 1967). 4. The severity of the somatic symptoms in experimental neurosis is reflected in the quantity of the 17-OH-CS secretion but also in a change of the hydrocortisone-corticosterone quotient (Lissak, EndrSczi and Medgyesi, 1957). Obviously, the adrenocortical hormones parallel tone and responsiveness of the ergotropic system. They are intimately related to the emotional state: Persons with low levels of corticoid excretion are "overtly depressed, apathetic and withdrawn" whereas those with high levels are "conflict-ridden", restless and irritable (Holmes and Ripley, 1955; Mason, 1968). Moreover, emotional stimuli such as struggle induced by restraint lead to increased corticoid excretion in normal rats but not in animals with pituitary grafts in the eye (Fortier, 1951). Apparently, the liberation of the corticotropin releasing factor is an essential neurovascular link between the hypothalamus and the secretion of ACTH. I t is concluded that the tuning of the central nervous system depends not only on neuronal stimuli which act on the ergotropic and trophotropic systems reflexly or directly at reticular, hypothalamic and limbie levels but also on chemical stimuli due to changes in the hormonal balance. The two groups of stimuli interact: certain hormones when reaching a critical level in the blood sensitize the ergotropic or trophotropic systems to environmental stimulation and induce specific behavioral reactions. Heightened responsiveness (tuning) but also reversal may result. Discussion. Through stimulation of afferent nerves or central structures, lesions in hypothalamus and limbic brain and through experiential factors (conditioning) the ergotropie/trophotropic balance and, consequently, the reactivity of the organism is changed in a predictable manner expressed in the laws of tuning. They account for the quantitative changes in sensori-motor, visceral and behavioral reactions seen on activation of the ergotropic or the trophotropie system. At higher degrees of tuning qualitative changes (reversal phenomena) occur which are demonstrable in somatic and autonomic responses as well as in behavior. Mood and emotions change in a dramatic manner as the state of tuning is altered from one of ergotropic dominance to one of trophotropic dominance and vice versa. Although a distinction between mood and
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emotion is often difficult it has frequently been observed t h a t stimuli which arouse emotions change the mood in a similar manner. Thus chlorpromazine induces a state of contentment (inferred from purring, rubbing, etc. in cats and playing and grooming in monkeys) whereas amphetamine produces a hostile mood. Similar observations have been made in man. The experimental analysis of the action of these and other drugs, of related therapeutic procedures such as electroshock (Ploog, 1950) and hypothalamic tumors on mood (see Gellhorn and Loofbourrow, 1963, for the literature) seems to show t h a t moods are based on tonic and emotions on phasic reactions of the ergotropictrophotropic systems. I n a given mood certain emotions are facilitated, others are inhibited (Bollnow, 1956). This phenomenon depends on the state of tuning of the nervous system. I n an ergotropic mood emotions acting primarily on the ergotropic system are facilitated and those chiefly involving the trophotropic system are inhibited. Corresponding rules apply to trophotropic moods 12. These phenomena are closely related to observations of Maranon (1924) who on injection of adrenaline, particularly following administration of thyroid extracts, evoked emotions such as great happiness or anxiety. Schachter (1964) resuming this work showed t h a t adrenalineinjected subjects are readily manipulated into states of euphoria, anger, and amusement. Apparently, the injection of adrenaline heightens the excitability of centers of the ergotropic system (Bonvallet, Dell and tIiebcl, 1954) and increases, therefore, the sensitivity of the subject to certain forms of emotional excitement. This sensitization is not specific for a single emotion but rather for a group of emotions which are linked to the ergotropic state. Although the peripheral effects of the injection of adrenaline are similar the evoked emotions m a y differ widely. This indicates to Schachter and Singer (1960) that "cognitions arising from the immediate situation as interpreted by past experience provide the framework within which one understands and labels his feelings". Apparently such diametrically oppposed emotions as joy and anger are easily evoked foliowing the injection of adrenaline in doses which cause palpitation, tremor, increased rate of breathing, and other sympathetic effects. Physiologically, these observations suggest t h a t a variety of hypothalamic-cortical patterns is compatible with a typical increased sympathetic discharge. I t is, therefore, assumed t h a t at a given 12 As Bollnow (1956) expresses it ,,Wie ich reich einem Ding zuwende und wie es mir erscheint, ist yon vornherein durch die Stimmungslage bedingt, in der ieh reich befinde. Nut in ~ngstlieher Stimmung begegnet mir Bedrohliches, und nur in einer heiteren Gemfitsveffassung kommen mir die beglfickenden Erlebnisse wie yon selbst entgegen. Datum ist es so sehwer, den zu tr5sten, der ernstlich betriibt ist".

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state of arousal, i.e. at a given degree of reticular discharge, various patterns of hypothalamic "upward" discharges occur. This inference is supported by the finding that pleasant and unpleasant emotions may be associated with a similar degree of heightened ergotropic excitation as indicated by the excretion of cateeholamines (Levi, 1965). On the other hand, a transition from ergotropic emotions to trophotropic emotions is difficult. This holds true for hypnosis. If the posture accompanying " t r i u m p h " is suggested and " l o c k e d " under hypnosis, a depressive mood cannot be brought about unless the postural setting is changed (Pasquarelli and Bull, 1951). A state of heightened ergotropic reactivity prevents the occurrence of emotions which involve strong trophotropie discharges and vice versa (Gellhorn, 1967). From the experiments discussed in this paper it is obvious that the tone of the skeletal muscles has a decisive influence on whether a state of trophotropic or ergotropic dominance will prevail. A passive attitude tends to promote trophotropic, an active attitude ergotropic dominance. These different attitudes account for the fact different persons may show ergotropic or trophotropic reactions in response to a particular stress (psychiatric interview) or the same person may react at one time with an ergotropic and another time with a trophotropie reaction. This "action-oriented" and "non-action oriented" behavior (Dudtey, Martin and Holmes, 1964) seems to underlie the ergotropic effects associated with mental work under stress and the trophotropic effects accompanying the exposure to a flash of light (with the instruction to attend to it) which Lacey et al. (1963) describe. I t has been mentioned that pleasant as well as unpleasant emotions of the ergotropic type may be accompanied by similar degrees of arousal. Similarly, sadness and postprandial happiness involve two psychologically opposed states associated with trophotropic dominance and muscular relaxation. These observations suggest that a variety of hypothalamic upward discharges may occur at a given degree of ergotropic or trophotropic excitation. Factors which are thought to play a role in producing this variety seem to be 1. multiform expressive movements particularly those involving the face which through cutaneous and proprioceptive discharges impose various patterns of excitation upon the hypothalamus (Gellhorn, 1964; Miller, Caul and Mirsky, 1967); 2. alterations in the hormonal milieu resulting from hypothalamically induced secretions of the anterior and posterior hypophysis which modify behavior and emotions; 3. partial as well as diffuse hypothalamic-cortical discharges occurring in states of ergotropic and trophotropic tuning. Different forms of partial

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trophotropic excitation indicated by the appearance of synchrony in restricted cortical areas have been reported by l~oth, Sterman and Clemente (1967) and Torii et al. (1965) whereas diffuse trophotropie cortical excitation resulted from stimulation of the basal forebrain area (Sterman and Clemente, 1962). On the other hand, partial activity of the sympathetic system has long been known (Gcllhorn, Cortell and Morphy, 1946; Gellhorn, 1967, p. 204, and 1968) is. Recently Di Cara (1969) has shown that partial autonomic discharges may be induced in conditioning experiments through positive reinforcement. In view of the "close association between increased arousal and the rewarding effects" (Berlyne, 1967), and the role of the hypothalamus and the thalamie reticular system in conditioning it is suggested that a variety of hypothalamic-cortical patterns based on partial activation of ergotropie and trophotropic division of the hypothalamus is involved in Di Cards experiments and that this mechanism is related to the great variety of moods and emotions. Moods and emotions can be divided into two major groups in which either the ergotropie system or the trophotropic system is dominant while the nondominant system is reciprocally inhibited. Omitting for the moment the paradoxical phase of sleep the quotient ergotrophie discharges falls continually from a state of a high degree trophotropie discharges of awareness and emotional arousal to a state of sleep. The turning point lies between the state of moderate arousal as seen in a person who is resting but awake and a person practicing meditation in a condition of nearly complete muscular relaxation. In the former state there is a slight ergotropic, in the latter a slight trophotropic dominance. As one passes from a resting state to that of contemplation the heart rate and palmar conduction decrease, the muscles become relaxed and the E E G shows signs of trophotropic dominance: the alpha potentials decrease in frequency and increase in amplitude (Wallace, 1970). The various states of mood, arousal and emotion which accompany a gradual rise of the ergotropic/trophotropie quotient from deep sleep to rage are characterized by reciprocal relations between the ergotropie and trophotropic systems but beyond this range simultaneous discharges of both systems occur. They are associated with pathological or nearly pathological phenomena such as dreams and hypnagogic hallucinations in narcolepsy in a state of trophotropie dominance, and hallucinations and delusions in delirium in a state of ergotropic dominance. In each instance the perception of reality is interfered with. 13 See also Lacey and Lacey's (1958) observations on autonomic response specificity.

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One of the important tasks for future research is the detailed characterization of the neurological substate on which the various emotions depend. The activation theory of Lindsley (1951) is in spite of its historic significance unsatisfactory since arousal and emotion are not identical. The finding that reticular lesions interfere with cortical desynchronization but not with behavioral arousal whereas lesions in the posterior hypothalamus produce the reverse effect (Feldman and Waller, 1962) and numerous related data discussed elsewhere point to the hypothalamus as the crucial structure involved in emotion (see Gellhorn, 1967, 1968; Gellhorn and Loofbourrow, 1963). I t is hoped that the study of the effects of tuning through activation of the ergotropic and trophotropic systems and their influence on hypothalamic functions and behavior has established some general concepts of the mechanisms underlying normal and some pathological forms of emotional behavior although the specific contributions of various levels of the cerebrospinal axis to these systems remain to be worked out (see Bronson, 1965). At present we know that emotions of the ergotropic type or stimulation of the ergotropic division of the hypothalamus, for instance, are associated with cortical desynchronization, increase in sympathetic discharges, and enhanced reactivity of motor and sensory functions. The specific contributions to the emotions of these various physiological changes are ill understood. In spite of the importance of proprioceptive discharges for hypothalamic excitability and emotional behavior high degrees of emotional excitation are compatible with muscular relaxation as seen in the Yoga trance. Cortical desynchronization and synchrony may be produced by several drugs without inducing behavioral changes (see Wikler, 1957). The human brain distingttishes between two intestinal stretch stimuli if one is habituated: the blocking of the alpha potentials in the E E G occurs only in response to the non-habituated stimulus; but this fine discrimination takes place without awareness indicating that desynchronization induced by the reticular formation is not the mechanism on which consciousness is based (Adam, 1967; see also Feldman and Waller, 1962). Perhaps the clarification of the interaction between reticular formation and hypothalamus which Bonvallet and D'Anna (1966) are studying will contribute to the solution of these important problems. References _~d~m, G.: Interoeeption and behavior. Budapest: Akademiai Kiado 1967. Alexander, L.: Effects of psychotropic drugs on conditional responses in man. In: Neuro-psychopharmacology, E. Rothlin, ed., vol. 2, p. 93-123. Proc. 2rid Internat. Meeting, Basle: Elsevier 1961. Andersson, ]3, Gale, C. C., I-Iofkelt, B., Ohga, A. : l~elation of preoptic temperature to the function of the sympathico-adrenomedullary system and the adrenal cortex. Acta physiol, scan& 61, 182-191 (1963).

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Armington, J.C., Mimick, L.L.: Electroencephalogram and sleep deprivation. J. appl. Physiol. 14, 247-250 (1959). Bandura, A. : Principles of behavior modification. New York: Holt 1969. Bard, P. : The hypothalamus and sexual behavior. Res. Publ. Ass. nerv. ment. Dis. 20, 551-579 (1940). Barterelli, C., Bizzi, E., Libretti, A., Zanchetti, A.: Inhibitory control of sinocarotid pressoreceptive afferents on hypothalamic autonomic activity and sham rage behavior. Arch. ital. Biol. 98, 308-326 (1960). Baust, W., Heinemann, H.: The role of the baroreceptors and of blood pressure in the regulation of sleep and wakefulness. Exp. Brain Res. 3, 12-24 (1967). Berlueci, G., Moruzzi, G., Salvi, G., Strata, P.: Pupil behavior and ocular movements during synchronized and desynchronized sleep. Arch. ital. Biol. 10g, 230-244 (1964). Berlyne, D. E. : Arousal and reinforcement. In: Nebraska Symposium on Motivation, D. Levine, ed., p. 1-110. Lincoln, Nebraska: U. of Nebraska Press 1967. Bernhaut, 1~I., Gellhorn, E., Rasmussen, A. T. : Experimental contributions to the problem of consciousness. J. Neurophysiol. 16, 21-35 (1953). Beyer, C., Ramirez, V. D., Whitmoyer, D. I., Sawyer, C. H. : Effects of hormones on the electrical activity of the brain in the rat and rabbit. Neurol. 18, 313-326 (1967). Bloch, V.: Le eontrole central de l'activit~ 616ctrodermale. ]~tude neurophysiologique et psychophysiologique d'un indice sympathique de l'activation r~ticulaire. J. Physiol. (Paris) 57, Suppl. 13, 1-132 (1965). - - V a l a t , IV[., Roy, J.-C.: Influences des afferences musculaires sur le tonus r~ticulaire. J. Physiol. (Paris) 57, 561-562 (1965). Bollnow, 0. F. : Das Wesen der Stimmungen, 3rd edition. Frankfurt: Klostermann 1956. Bonvallet, M., Allen, 1~. B., Jr.: Prolonged spontaneous and evoked reticular activation following discret bulbar lesions. Electronceph. clin. Neurophysiol. 15, 969-988 (1963). -D'Anna, L. : Responses r6ticulaires immediates, responses r6ticulaires tardives et oscillations spontanees du tonus r6ticulaire. Arch. ital. Biol. 164, 280-306 (1956). - - D e l l , P., Hiebel, G.: Tonus sympathique et activit6 ~l~ctrique corticale. Electroenceph. clin. Neurophysiol. 6, 119-144 (1954). -Zbrozyna, A. : Les commandes r6tieulaires du syst~me autonome et en particulier de l'innervation sympathiqne et parasymapthique de la pupille. Arch. ital. Biol. 161, 174--207 (1963). Born, H.: Die Umkehrbarkeit psyehosomatischer Abl~ufe als Grundlage fiir ein Prinzip allgemeiner Therapie. Z. ~rztl. Fortbild. 7, 23-31 (1953). - - Der Umkehreffekt in der Ausdrucksmotorik als bedingter Reflex. Z. ~rztl. Fortbild. 7, 699-700 (1953). Brady, J. V, Nauta, W. J . H . : Subcortical mechanisms in emotional behavior: affective changes following septal forebrain lesions in the albino rat. J. comp. physiol. Psyehol. 46, 339-346 (1953). Bronson, G.: The hierarchical organization of the central nervous system: implications for learning processes and critical periods in early development. Behav. Sei. 10, 7-25 (1965). Brown, J.L., Hunsperger, R.W., Rosvold, H . E . : Defence, attack and flight elicited by electrical stimulation of the hypothalamus of the cat. Exp. Brain Res. 8, 113-129 (1969). Brutkowski, S., Dabrowska, J. : Disinhibition after prefrontal lesions as a function of duration of intertrial intervals. Science 139, 505-506 (1963).

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]3uchwald, N . A . , Hull, C. D., Trachtenberg, M. C. : Concomitant behavioral and neural inhibition and disinhibition in response to subcorticM stimulation. Exp. Brain Res. 4, 58-72 (1967). Clemente, C. D., Sterman, M. B. : ]3asal forebrain mechanisms for internal inhibition and sleep. Res. Publ. Ass. herr. ment. Dis. 45, 127-147 (1967). Consolo, S. : Sensitivity of aggressive mice to centrally acting drugs. J. Pharm. Pharmacol. 17, 594 (1965). - - Garattini, S., Valzelli, L. : Amphetamine toxicity in aggressive mice. J. Pharm. Pharmacol. 17, 53-55 (1965). Delgado, J. M. R. : Free behavior and brain stimulation. Int. Rev. Neurobiol. 6, 349449 (1964). DiCara, L. V., Miller, N. E. : Heart-rate learning in the noncurarized state, transfer to the curarized state, and subsequent retraining in the noncurarized state. Physiol. ]3ehav. 4, 621-624 (1969). Ditfurth, H. von: Zur Problemlage der Pharmakopsychiatrie. In: ]3efinden und Verhalten, G. S. Achelis, ed. Stuttgart: Thieme 1961. Duchenne, G. ]3. : Physiology of motion. Philadelphia: Lippincott 1949. Dudley, D. L., Martin, C. J., Holmes, T. H. : Psychophysiologic studies of pulmonary ventilation. Psychosom. Med. 26, 645-660 (1964). Duffy, E. : Activation and behavior. New York: Wiley 1962. Eiff, A. W. yon: Der EinfluB seelischer ]3elastungen auf Stoffwechsel und Muskeltonus. Verb. dtsch. Ges. inn. Med. ~8, 468471 (1952). Endroczi, E., Hartmann, G., Lissak, A. : Meso-diencephalic activatory and inhibitory mechanisms. Acta physiol. Acad. Sci. hung. 31, 115-126 (1967). - - Lissak, K. : Interrelations between palaeocortical activity and pituitary-adrenocortical function. Aeta physiol. Acad. Sci. hung. 21, 257-263 (1962). - - - - Telegdy, G. : hlfluence of sexual and adrenocortical hormones on the maternal aggressivity. Acta physiol. Acad. Sci. hung. 14, 353-357 (1958). Euler, C. yon: The influence of hypothalamic thermoceptive structures on the electroencephalogram and gamma motor activity. Electroenceph. olin. Neurophysiol. 9, 391-408 (1957). Fairchild, M. D., Alles, G. A., Jenden, D. J., Mickey, M. R. : The effects of mescaline, amphetamine and four-ring substituted amphetamine derivatives on spontaneous brain electrical activity in the cat. Int. J. Neuropharmacol. 6, 151-167 (1967). Feldman, S.M., Waller, H . J . : Dissociation of electrocortical activation and behavioural arousal. Nature (Lond.) 196, 1320-1322 (1962). Flynn, J . P . : The neural basis of aggression in eats. In: Neurophysiology and emotion, New York: Rockefeller U. Press, D. C. Glass, ed., p. 40-60, 1967. Fortier, C. : Dual control of adrenocortieotrophin release. Endocrinology 49, 782788 (1951). Freeman, G.L., Pathman, J . H . : The relation of overt muscular discharge to physiological recovery from experimentally induced displacement. J. exp. Psyehol. 36, 161-174 (1942). Gel]horn, E.: Physiological processes related to consciousness and perception. Brain 77, 401-415 (1954). --Autonomic imbalance and the hypothalamus. Minneapolis: U. Minnesota Press 1957. - - The influence of curare on hypothalamic excitability and the electroencephalogram. Electroenceph. clin. l~europhysiol. 10, 697-703 (1958). -Sympathetic and parasympathetic summations. Acta. neuroveg. (Wien) 20, 181-194 (1959).

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-

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