Résumé
Ce travail, consacré à l'étude du mécanisme régulateur de la fonction adrénocorticotrophique de l'hypophyse, aborde dans une première partie les preuves expérimentales du contrôle exercé par cette glande sur le cortex surrénalien, les propriétés physiques, chimiques et biologiques de l'ACTH, ainsi que les méthodes qui permettent d'en déceler l'activité. Faisant suite à ces considérations générales, une revue critique de la littérature relative à la libération de l'ACTH sous l'influence du stress, étayée par les observations personnelles de l'auteur, permet l'énoncé des conclusions suivantes:
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1.
Des divers mécanismes proposés par les partisans des théories humorales périphériques et des théories nerveuses de l'activation corticotrophique induite par le stress, aucun n'a su satisfaire aux conditions requises du médiateur unique de cette activation.
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2.
L'inversion expérimentale des altérations métaboliques suspectes, l'administration d'hormones corticales, en vue de prévenir la chute de leur concentration sanguine, l'exclusion du système sympatho-surrénalien par sympathectomie totale, section spinale, dénervation des surrénales, adrénodémédullation ou administration de drogues sympatholytiques, ainsi que la déafférentiation de l'hypophyse par transplantation de la glande peuvent, il est vrai, bloquer, dans certaines circonstances, la réponse au stress du complexe hypophyso-surrénalien; aucune de ces interventions n'a pu la prévenir dans tous les cas.
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3.
Il convient de souligner, cependant, que l'interruption des voies hypothalamo-hypophysaires, réalisée par la transplantation de l'hypophyse dans la chambre antérieure de l'oeil, tout en permettant la libération de l'ACTH suscifée par les stimuli «systémiques», bloque l'effet corticotrophique de la stimulation «neurotrope».
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4.
D'autre part, comme en témoigne la réponse de l'hypophyse transplantée, l'adrenaline et l'histamine exercent sur cette glande un effet corticotrophique direct.
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5.
L'ensemble des faits considérés justifie, selon l'auteur, la conception d'une régulation polyvalente dont la modalité serait conditionnée par la nature du stimulus alarmant.
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6.
Ainsi, les stimuli nerveux ou émotionnels activeraient l'hypophyse par le fruchement des voies vasculo-nerveuses hypothalamo-hypophysaires, alors que l'effet corticotrophique des stimulisystémiques serait médié par diverses altérations humorales (niveau des corticoïdes circulants, libération d'adrénaline par la médullo-surrénale ou d'histamine par les terminaisons nerveuses ou les tissus lésés) produites sous l'influence du stress et agissant indépendamment ou concurrements sur l'hypophyse pour y déclencher la libération de l'ACTH.
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7.
Les deux mécanismes envisagés peuvent vraisemblablement co-exister dans la mesure où le stimulus présente á la fois des propriétéssystémiques et neurotropes.
Zusammenfassung
Diese Arbeit, die dem Regulationsmechanismus der adrenokortikotropen Funktion der Hypophyse gewidmet ist, befaßt sich in ihrem ersten Teil mit den experimentellen Untersuchungen der Kontrolle, welche von dieser Drüse über die Nebennierenrinde ausgeübt wird, mit den physikalischen, chemischen und biologischen Eigenschaften des ACTH, sowie mit den Methoden, die es gestatten, dessen Wirksamkeit zu enthüllen. Diesen allgemeinen Betrachtungen folgend, erlaubt eine kritische Übersicht über die Literatur, welche die Freisetzung des ACTH unter dem Einfluß desStress zum Gegenstand hat und die durch die persönlichen Beobachtungen des Autors unterstützt wird, die Formulierung folgender Schlußfolgerungen:
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1.
Von den verschiedenen Mechanismen, die von den Anhängern der peripheren humoralen und der nervalen Theorien über die durch denStress hervorgerufene kortikotrope Aktivierung vorgeschlagen wurden, konnte keine den vom einzigen Mittler dieser Aktivierung geforderten Bedingungen in befriedigender Weise entsprechen.
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2.
Die experimentelle Inversion der vermuteten metabolischen Veränderungen, die Zuführung kortikaler Hormone, um dem Absinken ihrer Konzentration im Blut vorzubeugen, die Ausschaltung des sympathischen Nebennierensystems durch Totalsympathektomie, Spinalsektion, Denervation der Nebennieren, Adrenodemedullation sowie Deafferenzierung der Hypophyse durch Transplantation dieser Drüse können unter gewissen Umständen tatsächlich dieStress-Reaktion des hypophysären Nebennierenkomplexes blockieren; doch konnte keiner dieser Eingriffe in allen Fällen vorbeugen.
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3.
Man muß indessen betonen, daß die Unterbrechung der hypothalamo-hypophysären Bahnen, die durch die Transplantation der Hypophyse in die vordere Augenkammer bewirkt wird, welche dabei die von den “systemischen” Stimuli hervorgerufene Freisetzung des ACTH ermöglicht, die kortikotrope Wirkung der “neurotropen” Stimulation blockiert.
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4.
Anderseits üben das Adrenalin und das Histamin — wie durch die Reaktion der transplantierten Hypophyse bezeugt wird —, auf diese Drüse eine direkte kortikotrope Wirkung aus.
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5.
Die Gesamtheit der in Betracht gezogenen Tatsachen rechtfertigt — nach der Ansicht des Autors —, die Annahme einer polivalenten Regulation, deren Modalität durch die Natur des alarmierenden Stimulus bedingt ist.
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6.
Somit würden die nervalen oder emotionalen Stimuli die Hypophyse vermittels der vaskulo-nervalen hypothalamo-hypophysären Bahnen aktivieren, während die kortikotrope Wirkung dersystemischen Stimuli durch verschiedene humorale Veränderungen vermittelt würde (Niveau der zirkulierenden Kortikoïde, Freisetzung von Adrenalin durch das Nebennierenmark oder von Histamin durch die Nervenendigungen oder die lädierten Gewebe), die unter dem Einfluß desStress zustandekommen und unabhängig voneinander oder miteinander auf die Hypophyse einwirken und dort die Freisetzung des ACTH auslösen.
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7.
Die beiden in Betracht gezogenen Mechanismen können wahrscheinlich in dem Maß zusammen existieren, als der Stimulus gleichzeitigsystemische undneurotrope Eigenschaften aufweist.
Summary
Following a review of the experimental proofs of the control exerted by the hypophysis over the adrenal cortex, the physical, chemical and biological properties of ACTH, as well as the indices of its secretion, the author attempts a critical evaluation of the factors responsible for its release under the influence of stress. From a survey of the literature, including the author's own observations, the following conclusions are reached:
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1.
Of the various mechanisms, independently shown to play a role in the activation of the pituitary adrenocorticotrophic function, none proved essential to the release of ACTH under all conditions of stress.
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2.
Experimental inversion of suspected metabolic alterations; administration of cortical hormones to prevent the lowering of their venous titer; exclusion of the sympathoadrenal system through complete sympathectomy, spinal section, denervation of the adrenal gland, adrenodemedullation or the use of adrenolytic agents; or the deafferentiation of the hypophysis through pituitary-stalk section or hypophyseal transplantation may admittedly interfere, under certain conditions, with the stress-induced release of ACTH from the anterior pituitary. However, none of these interventions, taken singly, can prevent it in all cases.
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3.
This is compatible with the author's view that the pituitary may be activated through more than one pathway; the nature of the required pathway or mediator being determined by the very nature of the stressing situation.
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4.
It was observed indeed that interruption of the hypothalamo-hypophyseal pathways, while allowing a normal response to “systemic” stimuli, suppressed the release of ACTH consequent to“neurotropic” stimulation.
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5.
Adrenaline and histamine were shown moreover, to exert a direct ACTH-releasing effect on pituitary transplants.
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6.
It is suggested, on the basis of present evidence, that the hypothalamohypophyseal pathways are required for pituitary activation in response toneurotropic (nervous or emotional) stimuli, while the corticotrophic effect ofsystemic stressing agents is mediated through various humoral changes (such as variations of the level of circulating adrenal cortical hormones, or the discharge of adrenaline or histamine), induced by stress, and acting either independently or concurrently on the pituitary to elicit the release of ACTH.
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7.
The above two mechanisms should not be necessarily considered as mutually exclusive, and it is logical to assume that they are simultaneously called into play under the many stress conditions which partake of both nervous and systemic components.
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Fortier, C. Facteurs humoraux et nerveux de la réponse hypophyso-surrénalienne au stress. Acta Neurovegetativa 5, 55–131 (1952). https://doi.org/10.1007/BF01232006
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DOI: https://doi.org/10.1007/BF01232006