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Einige endokrinologische Besonderheiten der Vögel

Some endocrine specializations of the birds

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Zusammenfassung

Anhand neuer embryologischer, morphologischer, physiologischer und biochemischer Befunde werden die vogelspezifischen Eigentümlichkeiten von drei endokrinen Systemen diskutiert: Pinealkomplex, hypothalamo-hypophysäres System und gastrointestinales Endokrinium.

Trotz der allgemein besonders starken Beeinflußbarkeit der Vögel durch photoperiodische Veränderungen ist bei ihnen der Pinealkomplex zu einer intracranialen endokrinen Drüse reduziert. Diese Epiphyse (Pinealorgan) zeigt unter experimentellen Bedingungen eine außerordentliche Unabhängigkeit von photoperiodischer Steuerung. Das völlige Fehlen des Pinealkomplexes bei den relativ nahe verwandten Krokodilen deutet auf die Möglichkeit hin, daß dies wohl eher phylogenetisch als funktionell zu erklären ist.

Das hypothalamo-hypophysäre System ist im wesentlichen reptilienartig, zeigt aber drei auffallende Spezialisierungen: 1. Eine hochentwickelte Eminentia mediana, welche aus zwei getrennten Regionen besteht. 2. Die Pars distalis ist mit der Eminentia so verbunden, daß ihre beiden Unterabteilungen (cephaler und caudaler Lappen) von beiden Regionen der Eminentia über getrennte Portalgefäße versorgt werden. 3. Den Vögeln als Gruppe fehlt die Pars intermedia.

Die auffallendsten Besonderheiten des gastrointestinalen Endokriniums stehen mit dem Inselorgan in Verbindung: insulinproduzierende B-Zellen und glukagonbildende A-Zellen sind weitgehend in besondere Inseln getrennt. Im Gegensatz zu dem der anderen Tetrapoden ist das Fettgewebe der Vögel insulin-unempfindlich. Die endokrine Kontrolle des Fettgewebes erfolgt über eine indirekte anabole Route (durch prolaktinstimulierte Lipogenese in der Leber) sowie eine direkte katabole Route durch Pankreasglukagon.

Bei einem Vergleich mit anderen Gnathostomengruppen zeigen Epiphyse, Adenohypophyse und Inselorgan einen besonders starken Trend zur Unabhängigkeit von direkter Innervation. In bezug auf Adenohypophyse und Inselorgan repräsentieren dabei die Vögel das entgegengesetzte Extrem zu den Knochenfischen (Teleostier), bei denen diese Drüsen besonders stark innerviert sind.

Die Vögel und Teleostier werden kurz mit den Myxiniden verglichen, deren endokrines System besonders schwach entwickelt ist. Die Unterschiede zwischen den drei Gruppen werden durch drei verschiedene Wege der Anpassung an Umweltbedingungen gedeutet: 1. Aktive Anpassung durch ein hochentwickeltes neuro-endokrines Kontrollsystem (Teleostier). 2. Passive, osmokonforme Anpassung, welche ein stark entwickeltes Endokrinium überflüssig macht (Myxiniden). 3. Vermeidung ungünstiger Umweltbedingungen durch Flugvermögen; eine verfeinerte, sekretomotorische Kontrolle der endokrinen Drüsen des normalen Stoffwechsels ist unnötig (Vögel).

Summary

Against the background of recent embryological, morphological, physiological and biochemical studies, peculiarities of three different avian endocrine systems are discussed: pineal complex, hypothalamo-hypophysial system, and gastrointestinal endocrine system.

Despite the marked photosensitivity of the birds in general, their pineal complex is reduced to an intracranial pineal gland, which shows a very high degree of functional autonomy when released from photoperiodic control. The complete absence of the pineal complex in the alligators suggests that this may be due to phylogenetic rather than adaptive reasons.

The hypothalamo-hypophysial system is basically of reptilian type, but shows three marked specializations: (1) the highly developed median eminence consists of two distinct regions; (2) the Pars distalis is located so, that both of its subdivisions (cephalic and caudal lobe) appear to receive distinctly separate vascularisation from the median eminence; (3) the birds as a group lack a Pars intermedia.

Within the gastrointestinal endocrine system of birds, the most obvious peculiarities identified so far are connected with the islet organ: insulin-producing B-cells and glucagonproducing A-cells are largely separated in different islets, and contrary to the situation in other tetrapodes, the avian adipose tissue is insulin-insensitive. The endocrine control of the adipose tissue involves an indirect anabolic route via prolactin-induced hepatic lipogenesis, and a direct catabolic route, via pancreatic glucagon.

When compared with other groups of gnathostomes, the pineal gland, adenohypophysis and islet organ of the birds show a particularly strong trend towards independence from secretomotor innervation. With respect to the adenohypophysial and islet innervation, the birds represent the opposite extreme to the teleosts, whose adenohypophysis and islet organ are particularly well innervated.

Birds and teleosts are compared with the Myxinidae, whose endocrine system is particularly poorly developed. It is hypothesized that the differences between the endocrine systems of these three groups reflect three different ways of adjustment to environmental conditions: (1) teleosts actively adjust to a continuous challenge from external osmolarity, whose control has to be integrated with the “milieu interne” and nutritional factors. They developed a highly sophisticated control system of their endocrines, which is capable of overriding humoral stimuli by nervous interference. (2) The Myxinidae passively conform with the osmotic environment, which makes a highly developed endocrine control system unnecessary. (3) The birds are shielded from environmental factors by a specialized body surface, and they can easily avoid unfavorable conditions by flight. This makes a sophisticated and fast acting (i.e. nervous) control of such endocrines unnecessary, which are involved in the maintenance of the normal “milieu interne”.

Wir danken MissKathleen Matt für die Hilfe bei der Anfertigung der Zeichnungen.

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  1. Daniel Baugh Inst. of Anatomy, Thomas Jefferson University, 19 107, Philadelphia, Penna., USA

    August Epple & Milton H. Stetson

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  1. August Epple
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Epple, A., Stetson, M.H. Einige endokrinologische Besonderheiten der Vögel. J Ornithol 117, 257–278 (1976). https://doi.org/10.1007/BF01640487

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