Animal Luminescence 593 



Beccari and collaborators (1724) , using Pholas, perhaps confirmed 

 by Spallanzani in 1788 with medusae, although it is doubtful if 

 Spallanzani's jellyfish were thoroughly dried. Spallanzani (1796), 

 Carradori (1798), and Macaire (1821) demonstrated that after the 

 glowworm lantern is dried, light will again appear on moistening. 



Although many had supposed that animal light might be due to 

 the element phosphorus, or a compound of phosphorus, the first 

 indication that some organic compound might emit the light came 

 from Macaire (1821), who suggested an albumin-like material re- 

 sponsible for the glowworm light. He did not give it a name, an 

 oversight remedied by later workers. Phipson (1872) thought in 

 terms of a fat, called noctilucine, as responsible for the light of dead 

 fish (luminous bacteria), and Dubois (1885, 1887) of a protein, 

 luciferin, present in the beetle Pyrophorus and in the mollusc, 

 Pholas. Luciferin was next discovered both in the firefly and in the 

 ostracod crustacean, Cypridina, by E. N. Harvey (1916) , who also 

 demonstrated the reduction of the oxidized luciferin by any pro- 

 cedure which adds hydrogen to the compound (1918) . 



Although seventeenth-century scientists attributed the cause of 

 many bioluminescences to " fermentation," this word had no spe- 

 cific meaning at that time. The first demonstration that an enzyme 

 in the modern sense is involved in luminescence is due to R. Dubois 

 (1885, 1887) , who named the enzyme luciferase. The part played by 

 adenosinetriphosphate and magnesium ions in firefly luminescence 

 was discovered by W. D. McElroy (1947) . That light-emitting ma- 

 terials are associated with granules while in the cell, appears to have 

 been first recognized by Malpighi (1688) for the firefly and by 

 Quatrefages (1850) for Noctiluca. 



Physical study of a bioluminescence involving observation of its 

 spectrum was made by Achard (1783) , using luminous wood and a 

 glass prism. Achard was rather surprised to find no colors and re- 

 garded the light as irrefrangible. Murray (1826) also examined the 

 glowworm light and reported " that it seems monochromatic and 

 incapable of further decomposition." These results were due to 

 experimental difficulties, as both Lehmann (1862) and Schnauss 

 (1862) observed red, yellow, and green components in glowworm 

 light. Louis Pasteur (1864) appears to be the next observer, on 

 the beetle, Pyrophorus, and E. Ray Lancaster next (1870) , using 

 the worm, Chaetopterus. All these men found a short continuous 

 spectrum not crossed by bright or dark lines, in the visible region. 

 The spectral energy curve was first measured by Ives and Coblentz 

 (1909). 



Both Dubois (1886) and Langley and Very (1890) attempted to 



