PERSPECTIVES ON AXILLARY ODOR 201 Androstenone, androstenol, and isovaleric acid are contributing a large part of this odor profile. Thus control measures should be directed at masking these odorants or interfering with their bacterial production. The finding of these two steroids, which incidentally play a key role in the sexual life on the boar (28), as contributors to human axillary odor is also suggestive of a role, possible vestigial, for olfactory communica- tion in humans. ACKNOWLEDGMENTS This research was supported in part by Public Health Service grant 2 RO! AM 26195-03 from the National Institute of Arthritis, Metabolism, and Digestive Diseases. REFERENCES (1) P. Wallace, Individual discrimination of humans by odor, Physiol. and Behav., 19, 577-579 (1977). (2) B. Brady, The sexual significance of the axillae, Psychiatry, 38, 278-289 (1975). (3) A. Dravnieks, Evaluation of human body odors: methods and interpretations, J. Soc. Cosmet. Chem., 26, 551-557 (1975). (4) R. L. Ellin, R. L. Fart, F. W. Oberst, C. L. Crouse, N. B. Billups, W. S. Koon, N. P. Musselman, F. R. Siddell, An apparatus for the detection and quantitation of volatile human effluents, J. Chromatogr., 100, 137-152 (1974). (5) J. N. Labows, G. Preti, E. Hoelze, J. Leyden, A. Kligman, Axillary odors: compounds of exogenous origin, J. Chromatogr.--Biomed. Applic., 163, 294-299 (1979). (6) G. D. Price, N. Smith, D. A. Carlson, The attraction of female mosquitos to stored human emanations in conjunction with adjusted levels of relative humidity, temperature and carbon dioxide, J. Chem. Ecol., 5 (3), 383-395 (1979). (7) S. Sastry, K. Buck, J. Janak, M. Dressier, G. Preti, "Volatiles emitted by humans," Biochemical Applications of Mass Spectrometry, G. Waller and O. C. Dermet, Wiley, p. 1086-1133 (1980). J. N. Labows, K. J. McGinley, J. j. Leyden, G. F. Webster, Characteristic 7-Lactone odor production of the genus Pityrosporum, Appl. and Environ. Micro., 38, 412-415 (1979). J. Hurley, W. Shelley, The human apocrine gland in health and disease, Thomas, Springfield, Illinois (1960). A. Corn'fort, "The likelihood of human pheromones," In: Pheromones, ed., M. C. Birch, (North- Holland Publishing Co., 1974) pp. 386-396. I. Bloch, Odoratus Sexualis (Panurge Press, New York, 1934). M.J. Russell, Human olfactory communication, Nature, 260, 520-521 (1976). R. L. Doty, "A review of recent psychophysical studies examining the possibility of chemical communication of sex and reproductive state in humans," In: Chemical Signals in Vertebrates, Eds. D. Muller-Schwarze and M. M. Mozell (Plenum Press, New York, 1977) pp. 273-286. B. Holdt, M. Schleidt, The importance of human odour in nonverbal communication, Z. Tierpsychol., 43, 225-238 (1977). N. Morris, R. Udry, Pheromonal influences on human sexual behavior: An experimental search, J. Biosoc. Sci., 10, 147-157 (1978). J. Cowley, A. Johnson, B. Brooksbank, The effect of two odorous compounds on performance in an assessment-of-people test, Psychoneuroendocrinology, 2, 159-172 (1977). M. Kirk-Smith, D. Booth, D. Carroll, P. Davies, Human social attitudes affected by androstenol, Res. Comm. in Psychol., Psyvhiatry and Behavior, 3, 379-384 (1978). M.J. Russell, G. M. Switz, K. Thompson, Olfactory influences on the human menstrual cycle, Pharmac. Biochem. Behav., 13, 737-738 (1980). J. E. Amoore, P. Pelosi, J. L. Forrester, Specific anosmias to 5o•-Androst-16-en-3-one and 0)-Pentadecalactone: The urinous and musky primary odors, Chemical Senses and Flavor, 2, 401-425 (1977). H. S. Koelega, Preference for and sensitivity to the odours of androstenone and musk, in Oilaction and Taste VII. H. van der Starre, ed., (Information Retrieval Inc. N.Y., 1980) p. 436. (8) (9) (10) (11) (12) (13) (14) (15) (16) (17) (18) (19) (20a)

202 JOURNAL OF THE SOCIETY OF COSMETIC CHEMISTS (2oh) (21) (22a) (22b) (23) M. Kirk-Smith, D. Booth, Effect of androstenone on choice of location in others' presence, ibid, pp. 397-400. N. Shehadeh, A. Kligman, The bacteria responsible for apocrine odor, part II, J. oflnv. Derm., 41, 1-5 (1963). J. Leyden, K. McGinley, E. Holzle, J. Labows, A. Kligman, The microbiology of the human axilla and its relationship to axillary odor, J. Invest. Derm., 77,413-416 (1981). P.J.H. Jackman, (personal communication) in Microbiology of Human Skin, 2nd ed., W. C. Noble (Lloyd-Luke LTD., London, 1981) p. 94. M. G.J. Beets, E. T. Theimer, "Odor similarity between structurally related odorants," in Ciba Foundation Symposium on Taste and Smell Vertebrates, G. E. W. Wolstenholme andJ. Knight, Eds. (J & A Churchill, London, 1970), pp. 313-323. (24) S. Bird, D. B. Gower, The validation and use of a radioimmunoassay for 5c•-androst-16-en-3-one in human axillary collections, J. Ster. Biochem., 14, 213-219 (1981). (25) J. N. Labows, G. Preti, E. Hoelzle, J. Leyden, A. Kligman, Steroid analysis of human apocrine secretion, Steroids, 3, 249-258 (1979). (26) D. B. Gower, M. R. Hancock and L. H. Bannister, Biochemical Studies on the boar pheromones, in Biochemistry of Taste and Olfaction, R. Cagan and M. Kate, Eds. (Academic Press, N.Y., 1981) pp. 7-31. (27) J. C. Do, E. Kitatsuji, E. Yoshii, Study on the components of musk. I. Ether soluble components, Chem. Pharm. Bull, 23 (3), 629-635 (1975). (28) H. Reed, O. Melrose, R. L. S. Patterson, Androgen steroids as an aid to the detection of oestrus in pig artificial insemination, Brit. Vet. J., 130, 61-67 (1974). (29) B. S. Goodrich, E. R. Hesterman, K. E. Murray, R. Mykytowycz, G. Stanley, G. Sugowdz, Identification of behaviorally significant volatile compounds in the anal gland of the rabbit, Oryctolagus cuniculus, J. Chem. Ecol., 4 ( 5 ), 581-594 (1978). (30) Z. Valenta, A. Khaleque, The structure of castoramine, Tetrahedron Letters, 12, 1-5 (1959). (31) J. Adams, A. Garcia, C. S. Foote, Some chemical constituents of the secretion from the Temporal Gland of the African Elephant, J. Chem. Ecol., 4, 17-25 (1978). (32) G. Anderson, A. Brundin, K. Anderson, Volatile compounds from the interdigital gland of reindeer (Rangifer t. tarandus L.), J. ofChem. Ecol., 5 (3), 321-333 (1978). (33) A. Brundin, G. Anderson, K. Anderson, T. Mossing, L. Kallquist, Short chain aliphatic acid in the interdigital gland secretion in reindeer (Rangifer tarandus L.), and their discrimination by reindeer, J. Chem. Ecol., 4, 613-622 (1978). (34) D. Miiller-Schwarze, C. Miiller-Schwarze, A. G. Singer, R. M. Silverstein, Mammalian pheromone: identification of active component in the subauricular scent of the male pronghorn, Science, 183, 860-861 (1976). (35) G. Anderson, K. Anderson, A. Brundin, C. Rappe, Volatile compounds from the tarsal scent gland of reindeer (Rangifer tarandus),J. Chem. Ecol., 1 (2), 275-281 (1975). (36) G. Epple, N. F. Golob, A. B. Smith, Odor communication in the Tamarin Saguinus fuscicollis (Callitridridae): behavioral and chemical studies, in Chemical Ecology.' Odor Communication in Animals, F.J. Ritter, ed. (North-Holland Biomedical Press, 1979)pp. 117-130. (37) D. H. McBumey, J. M. Levine, P. H. Cabanaugh, Psychophysical and social ratings of human body odor, Personality--Soc. Psych. Bull., 3, 135-138 (1977). (38) J. Amoore, Specific anosmia and the concept of primary odors, Chemical Senses and Flavor, 2, 267-281 (1977). (39) S. Puhvel, R. Reisner, M. Sakamoto, Analysis of lipid composition of isolated human sebaceous gland homogenates after incubation with cutaneous bacteria: thin-layer chromatography, J. Inv. Derre., 64, 406-410 (1975). (40) M. Julesz, New advances in the field of androgenic steroidogenesis of the human skin, Acta Med. Acad. Sci. Hungat, 25,273-285 (1968). (41) B. W. L. Brooksbank, Labelling of steroids in axillary sweat after administration of 3H- AS-Pregnenolone and •4C-Progesterone to a healthy man, Experentia, 26, 1012-1014 (1970). (42) B. W. L. Brooksbank, R. Brown, J. A. Gustafusson, The detection of 5c•-Androst-16-en-3c•-ol in human male axillary sweat, Experentia, 30, 864-865 (1974). (43) R. Claus, W. Alsing, Occurrence of 5c•-Androst-16-en-3-one, a boar pheromone, in man and its relationship to testosterone,J. Endocr., 68, 483 (1976). (44) J. N. Labows, Human odors--what can they tell us?, Perfumer and Flavorist, 4, 12-17 (1979). (45) F. Ayorinde, J. W. Wheeler, C. Wemmer, J. Murtaugh, Volatile components of the occipital gland secretion of the Bactrian camel,J. Chem. Ecol., 8, 177-183 (1982).