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The Chemistry of Light-Production in Luminous Organisms

oxygen, dubois, cypridina, substance and light

THE CHEMISTRY OF LIGHT-PRODUCTION IN LUMINOUS ORGANISMS The literature on the subject of animal phosphorescence or biolum inescence or biophotogenesis is so voluminous that no attempt can be made in the present article to record a complete list. A fairly com plete bibliography will be found in Mangold's article, "Die Production von Licht," in Winterstein's Vergleichende Physiologie. The names of Beijerinck, Dubois, Ehrenberg, Krukenberg, Mangold, McDermott, Molish, Panceri, Phipson, and Quatrefages are intimately connected with the chemical and physiological side of light-production. The work which has been accomplished up to the present time indicates that light-production by animals is to be classified among the chemilum inescent reactions which have been studied from the purely chemical side, especially by Radziszewski and Trautz. All of the evidence now indicates that oxygen is necessary for this reaction—at least the state ment is true for the following investigated forms: the fireflies; lumi nous bacteria and fungi; Noctiluca (2); a pennatulid (Cavernularia) ; and an ostracod crustacean (Cypridina). It has been objected by some that if we place the organisms, say luminous bacteria, in an atmosphere devoid of oxygen and find that no light is produced, that merely means that certain functions of the cell are interfered with, including light production, but does not indicate that oxygen is actually used up in the photogenic process. The objection will not hold, however, for extracts of luminous cells or the luminous secretion of Cypridina, which contains the light-producing substance in solution (so that it can not be removed by filtration through a porcelain or siliceous filter) will give no light if the oxygen be removed, and light returns when oxygen is readmitted.

We may therefore say that a definite substance is burned (oxidized), and in some forms it burns as granules and in others as a homogeneous and probably colloidal solution. This substance is stable and non luminous when dried and is not a fat or any substance soluble in fat solvents. It has been called noctilucin by Phipson (s and 7), photogen

by Molish (8), luciferin by Dubois (6), and photogenin by myself. Phipson and Molish did not recognize that in many forms two sub stances are concerned in light-production, but Dubois (9) pointed out that in the elaterid beetles and in the mollusc Photos dactyl= such was the case and he showed that these substances could be separated by heat. I can confirm this statement for certain other forms (lampyrid beetles and Cypridina), but find that the substances in Cypridina have many characteristics different from those described by Dubois for Photos and that the mechanism of light-production is quite different from that which Dubois (10) supposes.' A record of the experiments and a discussion of the chemistry of light-production in all its aspects will be found in the following pages. Cypridina, as the most favor able species, will be considered first and then the other forms which I have studied—the luminous beetles, the pennatulid Cavernularia, lum inous bacteria, the squid Watasenia scintillans, and a fish (Monocentris japonica)—together with a consideration of certain chemiluminescent reactions which involve the taking up of oxygen.

Most of the studies reported herein were made in Japan, a region unusually rich in luminous forms, through the kindness of Dr. Alfred G. Mayer and under the auspices of the Department of Marine Biology of the Carnegie Institution of Washington.

The success of my investigations was largely due to the interest and kindness of Professor C. Ishikawa, of the Zoology Department, Agri cultural College, Imperial University of Tokyo, and to Professor I. Ijima, of the Imperial University, who extended to me the hospitality of the marine biological laboratory at Misaki. I am also indebted to Dean Kozai and Professor Aso, of the Agricultural College, for the use of many chemicals and apparatus. I extend my cordial thanks to all of these men.