Summary
The effects of a number of metabolic inhibitors on aqueous formation in rabbits are described: These are discussed in the light of experimental findings and explanatory hypotheses about the process of aqueous humour formation which are briefly reviewed.
The evidence for active sodium transport is examined and it is shown that support for this hypothesis may arise from experiments using isolated ciliary epithelial tissue in vitro.
Aqueous formation is inhibited by fluoroacetate, dinitrophenol and ouabain whilst the composition of the aqueous undergoes little change. The composition may, however, readily by changed by alterations of plasma osmolarity, and it is suggested that the rate of secretion depends upon:
-
a)
the rate of active Na transport
-
b)
the plasma osmolarity.
Active Na transport is oxygen dependent and subject to inhibition by citrate cycle inhibitors such as malonate and fluoroacetate and by ouabain which inhibits Na+ - K+ - Mg-activated ATP-ase.
Histochemical and biochemical studies on ciliary epithelium suggest that the enzymes succinate dehydrogenase, Na-K-Mg-activated ATP-ase and cytochrome oxidase occur predominantly in the non-pigmented cell layer of the epithelium. It is postulated that the systems: citrate cycle, oxidative phosphorylation, ATP synthesis furnish the substrate for Na-K-Mg-activated ATP-ase which is responsible for extruding sodium from the epithelial cell into the posterior chamber. This system appears co-terminous with the non-pigmented epithelium.
Glycolysis rates in ciliary epithelium are high, but are not efficient as sources of energy for Na transport. The histochemistry suggests that glycolysis may be associated particularly with the ciliary pigment epithelium.
The effects on aqueous secretion of a number of other substances (aldosterone antagonists, agents with selective toxicity to retinal pigment epithelium) are described and discussed, and a conjectural account is given of the metabolism of the ciliary epithelium.
In the final section an hypothesis is advanced which attempts to take account of the role of the ciliary pigment epithelium in aqueous production. It is thought that this layer may function as an electrically charged membrane, the charge being maintained from cell metabolism and acting as a ‘permselective filter’ influencing the final result of the active ‘Na-pump’ in the non-pigmented cell layer.
In an appendix a description is given of a procedure used to determine the surface area of the ciliary epithelium in rabbits.
Résumé
Les effets d'un certain nombre de substances qui inhibent chez le lapin le métabolisme lors de la formation de l'humeur aqueuse sont décrits. On discute ces effets à la lumière de constatations expérimentales et d'hypothèses expliquant le mécanisme de la formation de l'humeur aqueuse.
On examine le transport actif du sodium, et l'on montre que cette hypothèse peut être appuyée par des expériences dans lesquelles on emploie du tissu épithélial ciliaire isolé in vitro.
La formation de l'humeur aqueuse est empêchée par l'acétate de fluor, le dinitrophénol et l'ouabaine, tandis que la composition de l'humeur aqueuse ne subit que peu de changement. Cependant, la composition peut être changée facilement par une modification de la molarité du plasma, et l'on suggère que le taux de sécrétion dépend:
-
a)
du taux de transport actif du sodium;
-
b)
de la molarité du plasma.
Le transport actif du sodium dépend de l'oxygène et il est soumis à l'action des corps inhibiteurs du cycle citrique tels que le malonate et l'acétate de fluor, et l'ouabaine qui empêche l'action de l'ATP-ase activé par Na+ - K+ - Mg.
Des études histochimiques et biochimiques sur l'épithélium ciliaire suggèrent que la succino-déhydrogenase, l'ATP-ase Na-K-Mg-activé et la cytochrome oxydase se trouvent d'une manière prédominante dans la couche des cellules non-pigmentées de l'épithelium. On admet que le système: cycle citrique, phosphorylation oxydative, synthèse de l'ATP fournit le substrat pour l'ATP-ase Na-K-Mg-activé et que celui-ci est chargé de la sortie du sodium hors de la cellule épithéliale vers la chambre postérieure. Il semble que ce système est localisé dans l'épithélium non-pigmenté.
L'importance de la glycolyse dans l'épithélium ciliaire est grande mais elle n'est pas efficace comme source d'énergie pour le transport du Na. L'histochimie fait penser que la glycolyse peut être associée particulièrement à l'épithélium ciliaire du pigmenté.
On décrit et discute les effets sur la sécrétion aqueuse d'un certain nombre d'autres substances (antagonistes de l'aldostérone, agents sélectivement toxiques pour l'épithélium pigmenté ciliaire), et on en déduit quel doit être le métabolisme de l'épithélium ciliaire.
Dans la partie finale on émet une hypothèse qui tâche de tenir compte du rôle de l'épithélium ciliaire pigmenté dans la production de l'humeur aqueuse. On pense que cette couche peut fonctionner comme membrane chargée électriquement, la charge étant maintenue par le métabolisme de la cellule. Elle agirait comme un ‘filtre permsélectif’, et influerait sur le résultat final de la ‘pompe-Na’ dans la couche non-pigmentée.
Dans un appendice on décrit une méthode destinée à mesurer la superficie de l'épithélium ciliaire chez le lapin.
Zusammenfassung
Es wird eine Anzahl von Stoffwechselhemmern der Kammerwasserproduktion beim Kaninchen beschrieben: Diese werden im Licht der experimentellen Befunde und der erläuternden Hypothesen über den Prozess der Kammerwasserproduktion, die kurz durchgesehen werden, erörtert.
Der Beweis für einen aktiven Natriumtransport wird geprüft, und es wird gezeigt, dass diese Hypothese möglicherweise durch Experimente mit isoliertem Epithelgewebe vom Ciliarkörper in vitro gestützt werden könnte.
Kammerwasserproduktion wird von Fluoracetat, Dinitrophenol und Ouabain gehemmt, während die Zusammensetzung des Kammerwassers sich wenig verändert. Die Zusammensetzung kann jedoch durch Änderungen der Plasmaosmolarität leicht verändert werden, und es wird angenommen, dass die Sekretionsgrösse a) von dem Ausmass des aktiven Natriumtransportes und b) von der Plasmaosmolarität abhängt.
Aktiver Natriumtransport ist sauerstoffbedingt und unterliegt einer Hemmung durch Inhibitionen des Zitronensäurezyklus wie Malonat und Fluoracetat und Ouabain, welches die Na+-K+-Mg-aktivierte ATP-ase hemmt.
Histochemische und biochemische Untersuchungen am Ciliarepithel lassen vermuten, dass die Enzyme Succinatdehydrogenase, Na-K-Mg-aktivierte ATP-ase und Cytochromoxydase vorwiegend in der pigmentfreien Zellage des Epithels vorkommen. Es wird postuliert, dass die Systeme Zitratzyklus, oxydative Phosphorylation, ATP-Synthese das Substrat für die Na-K-Mg-aktivierte ATP-ase bilden, welches für die Verdrängung vom Natrium aus den Epithelzellen in die hintere Augenkammer verantwortlich ist. Dieses System scheint mit dem pigmentfreien Epithel zusammenzufallen.
Die Glykolyserate im Ciliarepithel ist zwar gross, ist aber als Energiequelle für den Natriumtransport nicht genug wirksam. Es lässt sich auf Grund der Histochemie vermuten, dass Glykolyse besonders mit dem Pigmentepithel des Ciliarkörpers verbunden ist.
Es werden die Wirkungen einer Anzahl anderer Substanzen (Aldosteron Antagonisten, Agenzien mit selektiver auf das Pigmentepithel gerichteter Toxizität) auf die Kammerwassersekretion beschrieben und erörtert, und ein Bericht über den mutmasslichen Stoffwechsel des Ciliarepithels erteilt.
Im letzten Abschnitt wird eine Hypothese vorgeschlagen, welche die Rolle des Ciliarepithels bei der Kammerwasserproduktion in Rechnung zu stellen versucht. Es wird angenommen, dass diese Zellschicht als eine elektrisch geladene Membran funktioniere, deren Ladung vom Zellstoffwechsel aufrechterhalten werde, und die als ein ‘permselektives Filter’ wirken und so das Endresultat der aktiven ‘Natriumpumpe’ im pigmentfreien Zellager beeinflussen könnte.
In einem Anhang wird ein Verfahren beschrieben, die Oberfläche des Ciliarepithels beim Kaninchen zu bestimmen.
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Department of Physiology - Institute of Ophthalmology - Judd Street - W. C.1
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Cole, D.F. Aqueous humour formation. Doc Ophthalmol 21, 116–238 (1966). https://doi.org/10.1007/BF00184135
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DOI: https://doi.org/10.1007/BF00184135