Pergamon
Psychoneuroendocrinology, Vol. 19, No. 2, pp. 209-232, 1994
Copyright © 1994 Elsevier Science Ltd
Printed in the USA. All rights reserved
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0306-4530(93)E0010-S
REVIEW
ACTIVATION OF THE
HYPOTHALAMUS-PITUITARY-ADRENAL AXIS BY
BACTERIAL ENDOTOXINS: ROUTES AND
INTERMEDIATE SIGNALS
FRED J.H. TILDERS, l ROEL H. DERIJK, 1.
ANNE-MARIE VAN DAM, I VALERIE A.M. VINCENT, I
KAREL SCHOTANUS, 1 and JEK H.A. PERSOONS 2
Departments of IPharmacology and 2Cell Biology, Research Institute Neurosciences Vrije
Universiteit, Amsterdam, The Netherlands
SUMMARY
Peripheral administration of endotoxin induces brain-mediated responses, including activation of
the hypothalamus-pituitary-adrenal (HPA) axis and changes in thermoregulation. This paper re-
views the mechanisms by which endotoxin affects these responses. The effects on theroregulation
are complex and include macrophage-dependent hyperthermic and hypothermic responses. Low
doses of endotoxin, given IP, activate peripheral macrophages to produce interleukin (IL)-lfl,
which enters the circulation and acts as a hormonal signal. IL-I may pass fenestrated endothelium
in the median eminence to stimulate corticotropin-releasing hormone (CRH) secretion from the
CRH nerve-terminals. In addition, IL-1 may activate brain endothelial cells to produce IL-I, IL-
6, prostaglandins, etc., and secrete these substances into the brain. By paracrine actions, these
substances may affect neurons (e.g., CRH neurons) or act on microglial cells, which show IL-1-
induced IL- 1 production and therefore amplify and prolong the intracerebral IL- 1 signal. In contrast,
high doses of endotoxin given IV may directly stimulate endothelial cells to produce IL-1, IL-6,
and prostaglandin-E 2 (PGE2) and thereby activate the HPA axis in a macrophage-independent
manner.
INTRODUCTION
THE STRESS CONCEPT of Hans Selye was primarily based on stimuli that disturbed the
physical integrity of the individual. In his studies, administering toxins to animals, induc-
ing tissue damage, or exposing animals to X-rays played an important role. Selye noted
that this type of stimuli resulted in a general and stereotyped somatic response, known
as the GeneralAdaptation Syndrome, which involves enlargement of the adrenals, involu-
tion of the thymus, lymphopenia, and gastric ulcerations. These changes are associated
with increased activity of the hypothalamus-pituitary-adrenal (HPA) axis and played a
Address correspondence and reprint requests to: F.J.H. Tilders, Research Institute Neurosciences
Vrije Universiteit, Faculty of Medicine, Department of Pharmacology, Van der Boechorststraat
7, 1081 BT Amsterdam, The Netherlands.
* Present address: NIMH, Dept. Intramural Res. Prog., Clin. Neuroendocrinology Branch, Bldg
10 Room 3S231, 9000 Rockville Pike, Bethesda, Maryland 20892, U.S.A.
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