Pheromones in the Press

Baum MJ. Mammalian animal models of psychosexual differentiation: When is 'translation' to the human situation possible? Horm Behav. 2006 Nov 50(4):579-88. Quote: “…nearly all of the non-primate animals typically studied in the laboratory rely on body odors as important, if not sole, determinants of heterosexual partner selection…”, “If olfactory cues are the critical determinants of ferrets' sex partner preference, it stands to reason that sex differences in the detection and processing of odors from conspecifics may underlie sex differences in partner preference.”

Grammer K, Fink B, Neave N. Human pheromones and sexual attraction.
Eur J Obstet Gynecol Reprod Biol. 2005 Feb 1;118(2):135-42. Review.

Stowers L, Marton TF. What is a pheromone? Mammalian pheromones reconsidered. Neuron. 2005 Jun 2;46(5):699-702. Review.

Wysocki CJ, Preti G. Facts, fallacies, fears, and frustrations with human pheromones. Anat Rec A Discov Mol Cell Evol Biol. 2004 Nov;281(1):1201-11. Review.

Pause BM. Are androgen steroids acting as pheromones in humans? Physiol Behav. 2004 Oct 30;83(1):21-9. Review.

Brennan PA, Keverne EB. Something in the air? New insights into mammalian pheromones. Curr Biol. 2004 Jan 20;14(2):R81-9. Review.

Kohl JV, Atzmueller M, Fink B, Grammer K. Human pheromones: integrating neuroendocrinology and ethology. Neuro Endocrinol Lett. 2001 Oct;22(5):309-21. Review. PDF

Keverne EB. Pheromones, vomeronasal function, and gender-specific behavior.
Cell. 2002 Mar 22; 108(6): 735-8. Review

Savic I. Imaging of brain activation by odorants in humans. Curr Opin Neurobiol. 2002 Aug;12(4):455-61. Review.

Sobel N, Brown WM. The scented brain: pheromonal responses in humans.
Neuron. 2001 Aug 30;31(4):512-4. Review.

Research that helps to detail the role of human pheromones in behaviour

Published in 2006

Rantala, M. J., Enksson, C. J. P., Vainikka, A. and Kortet, R. Male steroid hormones and female preference for male body odor. Evolution and Human Behavior. 2006; 27(4): 259-269. Quote: “…cortisol or its interplay with other causal factors affecting male scent may shape the olfactory cues of mate selection in humans.

Witt M, Wozniak W. Structure and function of the vomeronasal organ.
Adv Otorhinolaryngol. 2006;63:70-83. Review. Quote: “… the most likely binding sites for human pheromone candidates are receptor cells within the olfactory epithelium.”

Liberles SD, Buck LB. A second class of chemosensory receptors in the olfactory epithelium. Nature. 2006 doi:10.1038/nature05066. Key Finding: Genes that code for receptors, called 'trace amine-associated receptors' are present in human, mouse and fish. These receptors, like other odor receptors are expressed in unique subsets of neurons dispersed in the olfactory epithelium. In mice there are at least three of these receptors. One receptor recognizes volatile amines found in urine. Another one detects a stress-related compound, and two others detect compounds that are found in different concentrations in male versus female urine. One of these compounds is reported to be a pheromone. Collectively, these findings indicate that chemical signals that are likely to function as pheromones are processed by the main olfactory system of mammals. Accordingly, a human vomeronasal organ is not required. Downloaded July 31, 2006 from http://www.nature.com/nature/journal/vaop/ncurrent/pdf/nature05066.pdf

Ngai, J. Neuroscience: An extra dimension to olfaction (Comment on Liberles and Buck). Nature. 2006 doi:10.1038/nature05001. Quote: “Evolutionary biologists, as well as physiologists and those studying animal behaviour, will be curious to find out how TAARs evolved for communication in other vertebrate species.”
Downloaded July 31, 2006 from http://www.nature.com/nature/journal/vaop/ncurrent/pdf/nature05001.pdf

Wedekind C,  Seebeck T, Bettens F, Paepke AJ. The Intensity of Human Body Odors and the MHC: Should We Expect A Link?  Evolutionary Psychology 2006 4: 85-94. Quote: “…if we control for MHC-linked perception we find few indications for a possible link between body odor intensity and MHC specificity. It appears that MHC homozygotes produce body odors that are on average perceived as more intense than those of heterozygotes.”  downloaded July 31, 2006 from http://human-nature.com/ep/downloads/ep048594.pdf

Matchock RL, Susman EJ. Family composition and menarcheal age: Anti-inbreeding strategies. Am J Hum Biol. 2006 Jul-Aug;18(4):481-91.Quote: “We believe that, taken together, a more parsimonious explanation of the data is that pheromonal cues modulate sexual maturity so as to enhance mating and prevent inbreeding (Table 2). The father absence-early menarche finding is not just a human anomaly that can be forced into, and explained by, psychological theories. The prevention of inbreeding is so paramount to the successful propagation of healthy genes that anti-inbreeding behaviors and changes in reproductive physiology appear to be highly conserved across species. That is, parents suppress reproduction of their offspring.”

Shepherd GM. Behaviour: smells, brains and hormones. Nature. 2006 Jan 12;439(7073):149-51. Quote: "The traditional distinction that common odours are perceived through the olfactory pathway and pheromones by the vomeronasal pathway is dead." Quote: "We have much more to learn about how intimately neuroendocrine functions, controlled by pheromones, acting through our noses, interact with other operations within the brain to control human behaviour and cognition.”

Pause BM, Krauel K, Schrader C, Sojka B, Westphal E, Muller-Ruchholtz W, Ferstl R. The human brain is a detector of chemosensorily transmitted HLA-class I-similarity in same- and opposite-sex relations. Proc Biol Sci. 2006 Feb 22;273(1585):471-8. Quote: “…HLA-related signals seem to be associated to a negative selection bias in mating behaviour. Moreover, HLA-associated odour signals from same-sex persons are processed differently in males and females, pointing to different behavioural functions in male-to-male (competition) and female-tofemale (communal behaviour) relations.”

Leiblum S, Brezsnyak M. Sexual chemistry: Theoretical elaboration and clinical implications. Sexual and Relationship Therapy. 2006 Feb; 21(1): 55 – 69. Quote: “In a comprehensive review article, Kohl et al. (2001) summarize the many research studies highlighting the influence of odors on human interaction and attraction.”

Prehn A, Ohrt A, Sojka B, Ferstl R, Pause BM. Chemosensory anxiety signals augment the startle reflex in humans. Neurosci Lett. 2006 Feb 13;394(2):127-30. Quote: “It has been suggested that the processing of chemosignals—the oldest phylogenetic receptive system shared by all organisms including bacteria—serves survival by generating appropriate behavioral responses to these signals. In line with this consideration, some authors assume that approach-avoidance reactions in animals, elicited by chemical cues, form the phylogenetic basis for the experience of emotions in humans.”

Berglund H, Lindstrom P, Savic I. Brain response to putative pheromones in lesbian women. Proc Natl Acad Sci U S A. 2006 May 23;103(21):8269-74. Quote: "These data support our previous results about differentiated processing of pheromone-like stimuli in humans and further strengthen the notion of a coupling between hypothalamic neuronal circuits and sexual preferences."

Chen D, Katdare A, Lucas N. Chemosignals of fear enhance cognitive performance in humans. Chem Senses. 2006 Jun;31(5):415-23.

Havlıcek J, Dvorakova R, Bartos L, Flegr J. Non-Advertized does not Mean Concealed: Body Odour Changes across the Human Menstrual Cycle. Ethology 112 (2006) 81–90. Quote: “Our results show that both the pleasantness and
attractiveness ratings given to axillary odours were lowest during menstruation and peaked in the follicular phase when the probability of conception is highest.”

Published in 2005

McClintock MK, Bullivant S, Jacob S, Spencer N, Zelano B, Ober C. Human body scents: conscious perceptions and biological effects. Chem Senses. 2005 Jan;30 Suppl 1:i135-i137. Quote: “Some human compounds are experienced as conscious odors, e.g. recognized verbally and explicitly as odors with verbal descriptors. Other compounds have an odor, yet these olfactory properties are not required for them to function as pheromones, modulating hormonal and motivational states.

Savic I, Berglund H, Lindstrom P.  Brain response to putative pheromones in homosexual men. Proc Natl Acad Sci U S A. 2005 May 17;102(20):7356-61.
Quote: "These findings show that our brain reacts differently to the two putative pheromones compared with common odors, and suggest a link between sexual orientation and hypothalamic neuronal processes."

Ebster C, Kirk-Smith, M. The effect of the human pheromone androstenol on product evaluation. Psychology and Marketing. 2005 22(9):739–749. Quote: “… a putative human pheromone can be used to enhance and change product perceptions in a gender-related direction. It remains to be seen if this can be used commercially; however, the results of this experiment suggest that more comprehensive studies would be worthwhile to explore the possible marketing uses of these human chemical signals.

Martins Y, Preti G, Crabtree CR, Runyan T, Vainius AA, Wysocki CJ. Preference for human body odors is influenced by gender and sexual orientation. Psychol Sci. 2005 Sep;16(9):694-701. Quote: “…lesbians and gay males may produce an array of axillary odorants that distinguish them from heterosexuals. Furthermore, gay males may perceive these characteristic odorants differently than their heterosexual counterparts do

Roberts SC, Gosling LM, Spector TD, Miller P, Penn DJ, Petrie M. Body odor similarity in noncohabiting twins. Chem Senses. 2005 Oct;30(8):651-6. Quote: “Our results indicate that odor similarity in pairs of twins can be perceived by the human nose.”

Tokunaga Y, Omoto Y,  Sangu T, Miyazaki M, Kon R, Takada K Sexual differentiation in sensitivity to male body odor. International Journal of Cosmetic Science. 2005 Dec;27(6):333-341. Quote: “Females evaluate androstenone itself as more unpleasant than males do, and furthermore, for only females, androstenone enhances the intensity and unpleasantness of other body-odor constituents such as short-chain fatty acids.”

Lundstrom JN, Olsson MJ.  Subthreshold amounts of social odorant affect mood, but not behavior, in heterosexual women when tested by a male, but not a female, experimenter. Biol Psychol. 2005 Dec;70(3):197-204. KEY FINDING: exposure to a non-detectable amount of a putative human pheromone modulated women’s mood and psychophysiological arousal, but effects were only evident when an experimenter of the opposite sex was present during testing. This suggests that social context can modulate effects of putative human pheromone exposure in women.

Havlicek J, Roberts SC,  Flegr J. (2005). Women’s preference for dominant male odour: effects of menstrual cycle and relationship status. Biol. Lett. doi:10.1098/rsbl.2005.0332 Published online. Quote: “The higher self-confidence of dominant males may also have an impact on the perceived sexiness of their body odour.”

Kohl, JV (2005) Human Pheromones, Neuroscience, and Male Homosexual Orientation. International Behavioural Development Symposium. Minot, ND, Aug 3-6. FULL TEXT available with an introductory article in the Fall 2005/Winter 2006 issue of Entelechy: Mind and Culture.

Published in 2004

Jacob S, Spencer NA, Bullivant SB, Sellergren SA, Mennella JA, McClintock MK. Effects of breastfeeding chemosignals on the human menstrual cycle. Hum Reprod. 2004 Feb;19(2):422-9. KEY FINDING: Because compounds from lactating women and their infants modulated the ovarian cycles of women, as is seen in other mammals, they have the potential to function as pheromones, regulating fertility within groups of women."

Jacquot L, Monnin J, Brand G. Unconscious odor detection could not be due to odor itself. Brain Res. 2004 Mar 26;1002(1-2):51-4. Quote: "These data suggest that unconscious odor detection as defined in the introduction (i.e., different from the common sense experience that perceiving subjects are not always aware of the presence of stimuli in suprathreshold concentration, a phenomenon related to a decrease in attention) could not be due to the odorant, but probably to the trigeminal component of the nasal stimulus."

Pierce JD Jr, Cohen AB, Ulrich PM. Responsivity to two odorants, androstenone and amyl acetate, and the affective impact of odors on interpersonal relationships.
J Comp Psychol. 2004 Mar;118(1):14-9. Quote: "In the present study, we found that responsivity to certain odors is associated with self-reports of the role of odors in influencing humans’ moods, social attractions, and individual preferences."

Cornwell RE, Boothroyd L, Burt DM, Feinberg DR, Jones BC, Little AC, Pitman R, Whiten S, Perrett DI. Concordant preferences for opposite-sex signals? Human pheromones and facial characteristics. Proc R Soc Lond B Biol Sci. 2004 Mar 22;271(1539):635-40.  KEY FINDING: "...putative sex pheromones and sexually dimorphic facial characteristics convey common information about the quality of potential mates."

Shepherd GM The human sense of smell: are we better than we think?
PLoS Biol. 2004 May;2(5):E146. Epub 2004 May 11 Quote: “The factors reviewed here suggest that the sense of smell is more important in humans than is generally realized, which in turn suggests that it may have played a bigger role in the evolution of human diet, habitat, and social behavior than has been appreciated.”

Kovacs G, Gulyas B, Savic I, Perrett DI, Cornwell RE, Little AC, Jones BC, Burt DM, Gal V, Vidnyanszky Z. Smelling human sex hormone-like compounds affects face gender judgment of men. Neuroreport. 2004 Jun 7;15(8):1275-7.  KEY FINDING: Odorous sex hormone-like steroids: 5-alpha-androgenst-16-en-3-one (androgen) or oestra-1, 3, 5 (10), 16-tetraen-3-ol (estrogen) bias face gender discrimination. As a result of inhalation of androgen, men perceive faces to be more masculine as compared to when they are exposed to estrogen. Quote: "Our results provide evidence for specific cross-sensory effects of the gender-specific chemosensory cues on the categorization of visual face gender."

Gulyas B, Keri S, O'Sullivan BT, Decety J, Roland PE. The putative pheromone androstadienone activates cortical fields in the human brain related to social cognition. Neurochem Int. 2004 Jun;44(8):595-600. Quote: “…this is the first functional neuroimaging study showing that androstadienone, a putative genderspecific human pheromone, activates inferior PFC and STP in the human brain when compared with pleasant and unpleasant non-pheromone odours. These activated brain areas are, by several other investigations, shown to be heavily involved in other than olfactory functions, including various aspects of attention, visual perception and recognition and social cognition.”

Bensafi M, Brown WM, Khan R, Levenson B, Sobel N. Sniffing human sex-steroid derived compounds modulates mood, memory and autonomic nervous system function in specific behavioral contexts. Behav Brain Res. 2004 Jun 4;152(1):11-22. Quote: "These results suggest that sex-steroidal compounds modulate mood, memory and autonomic nervous system responses and increase their significance within specific behavioral contexts. These findings lend support to a specific role for these compounds in chemical communication between humans."

Kuukasjärvi S, Eriksson CJP, Koskela E, Mappes T, Nissinen K, Rantala MJ. Attractiveness of women's body odors over the menstrual cycle: the role of oral contraceptives and receiver sex. Behavioral Ecology 2004 15: 579-584. Quote: “…our results support the view that the body odors of an ovulating woman increase her attractiveness to men.”

Wilcox AJ, Baird DD, Dunson DB, McConnaughey DR, Kesner JS, Weinberg CR. On the frequency of intercourse around ovulation: evidence for biological influences. Hum Reprod. 2004 Jul;19(7):1539-43. KEY FINDING: "There apparently are biological factors that promote intercourse during a woman's 6 fertile days." COMMENT: The most important biological factor in mammalian female copulation during the fertile period is olfaction/pheromones.

Roberts SC, Havlicek J, Flegr J, Hruskova M, Little AC, Jones BC, Perrett DI, Petrie M. Female facial attractiveness increases during the fertile phase of the menstrual cycle. Proc R Soc Lond B Biol Sci. 2004 Aug 7;271 Suppl 5:S270-2.  KEY FINDING: "This indicates the existence of visible cues to ovulation in the human face, and is consistent with similar cyclical changes observed for preferences of female body odour."

Spencer NA, McClintock MK, Sellergren SA, Bullivant S, Jacob S, Mennella JA.
Social chemosignals from breastfeeding women increase sexual motivation.
Horm Behav. 2004 Sep;46(3):362-70. KEY FINDING: "...natural compounds collected from lactating women and their breastfeeding infants increased the sexual motivation of other women, measured as sexual desire and fantasies." No specific compound was isolated.

Olofsson JK, Nordin S. Gender differences in chemosensory perception and event-related potentials. Chem Senses. 2004 Sep;29(7):629-37. Quote: “The finding that women, compared to men, generated larger amplitudes and shorter latencies for the relatively exogenous P2/P3 component suggests that the observed perceptual gender differences predominantly have their origin at a relatively high level of neural processing.

Bensafi M, Tsutsui T, Khan R, Levenson RW, Sobel N. Sniffing a human sex-steroid derived compound affects mood and autonomic arousal in a dose-dependent manner. Psychoneuroendocrinology. 2004 Nov;29(10):1290-9. Quote: "These findings further implicate AND in chemical communication between humans, but pose questions as to the path by which AND is transduced, whether through chemical sensing or transdermal diffusion."

Pause BM, Ohrt A, Prehn A, Ferstl R. Positive emotional priming of facial affect perception in females is diminished by chemosensory anxiety signals. Chem Senses. 2004 Nov;29(9):797-805. Quote: “… we conclude that the results reported here strongly indicated that anxiety in humans can be chemosensorily communicated.”

Herz RS, Eliassen J, Beland S, Souza T. Neuroimaging evidence for the emotional potency of odor-evoked memory. Neuropsychologia. 2004;42(3):371-8. KEY INDICATION "...the subjective experience of the emotional potency of odor-evoked memory is correlated with specific activation in the amygdala during recall and offers new insights into the affective organization of memory.

Kraft P, Popaj K. Total Synthesis and Olfactory Evaluation of 5β,10-Dimethyl-des-A-18-norandrostan-13β-ol: A Potential Human Pheromone? Eur. J. Org. Chem. 2004, 4995-5002.

Published in 2003

Roney JR, Mahler SV, Maestripieri D.  Behavioral and hormonal responses of men to brief interactions with women. Evol Hum Behav. 2003 24:365–375.
Quote: “…the very existence of a causal nexus between courtship behaviors and neuroendocrine mechanisms suggests the importance of future research on the hormonal correlates of courtship as a possible window onto the design of human mating mechanisms.

Preti G, Wysocki CJ, Barnhart KT, Sondheimer SJ, Leyden JJ. Male axillary extracts contain pheromones that affect pulsatile secretion of luteinizing hormone and mood in women recipients. Biol Reprod. 2003 Jun;68(6):2107-13.

Lundstrom JN, Hummel T, Olsson MJ. Individual differences in sensitivity to the odor of 4,16-androstadien-3-one. Chem Senses. 2003 Sep;28(7):643-50. KEY FINDING Women tend to be more sensitive to the odor than men; "olfactory sensitivity to androstadienone is bimodally distributed in the population with a subgroup consisting of highly sensitive people."

Lundstrom JN, Goncalves M, Esteves F, Olsson MJ. Psychological effects of subthreshold exposure to the putative human pheromone 4,16-androstadien-3-one. Horm Behav. 2003 Dec; 44(5): 395-401.

Bensafi M, Brown WM, Tsutsui T, Mainland JD, Johnson BN, Bremner EA, Young N, Mauss I, Ray B, Gross J, Richards J, Stappen I, Levenson RW, Sobel N. Sex-steroid derived compounds induce sex-specific effects on autonomic nervous system function in humans. Behav Neurosci. 2003 Dec;117(6):1125-34. KEY INDICATION "...AND's opposite effects on physiology in men and women further implicate this compound in chemical communication between humans."

Knecht M, Lundstrom JN, Witt M, Huttenbrink KB, Heilmann S, Hummel T.
Assessment of olfactory function and androstenone odor thresholds in humans with or without functional occlusion of the vomeronasal duct. Behav Neurosci. 2003 Dec; 117(6): 1135-41. KEY INDICATION: The human vomeronasal duct, and therefore, the human VNO does not play a major role in sensitivity toward odorants or the perception of a putative human pheromone.

Lundstrom JN, Goncalves M, Esteves F, Olsson MJ. Psychological effects of subthreshold exposure to the putative human pheromone 4,16-androstadien-3-one. Horm Behav. 2003 Dec;44(5):395-401. KEY FINDING "...this study corroborates earlier findings suggesting that androstadienone exposure yields effects on women's mood; the feeling of being focused. The mood effects were not dependent on menstrual cycle phase. Further, these effects are replicable and occur also when androstadienone detection is rigorously controlled for across variation in menstrual cycle."

Published before 2003

Thorne F, Neave N, Scholey A, Moss M, Fink B. Effects of putative male pheromones on female ratings of male attractiveness: influence of oral contraceptives and the menstrual cycle. Neuro Endocrinol Lett. 2002 Aug;23(4):291-7. Quote: “…the results of this study once again demonstrate that male axillary secretions cause changes in females’ emotional evaluations.”

Gangestad SW, Thornhill R, Garver CE. Changes in women's sexual interests and their partners' mate-retention tactics across the menstrual cycle: evidence for shifting conflicts of interest. Proc Biol Sci. 2002 May 7;269(1494):975-82. KEY FINDING: Men are able to perceive their partner's hormonal state, and adjust their behavior accordingly.

Ackerl K, Atzmueller M, Grammer K. The scent of fear. Neuro Endocrinol Lett. 2002 Apr;23(2):79-84. KEY FINDING: Women are able to detect the scent of fear. PDF.

Jacob S, McClintock MK, Zelano B, Ober C. Paternally inherited HLA alleles are associated with women's choice of male odor. Nat Genet. 2002 Feb;30(2):175-9. Quote: "Our data indicate that paternally inherited  HLA-associated odors influence odor preference and may serve as social cues."

Woodson JC. Including 'learned sexuality' in the organization of sexual behavior.
Neurosci Biobehav Rev. 2002 Jan;26(1):69-80. Review. KEY CONCEPT: "...learning from experience plays a critical role in the organization of sexual motivation and psychosexual differentiation...[and] ... provide[s] an essential link between biological predispositions and mature sexual preferences..."

Jacob S, Hayreh DJ, McClintock MK. Context-dependent effects of steroid chemosignals on human physiology and mood. Physiol Behav. 2001 Sep 1-15;74(1-2):15-27. KEY FINDING: Autonomic nervous system (ANS) responsivity to the human chemosignals D4,16-androstadien-3-one and 1,3,5,(10),16-estratetraen-3-ol is context dependent (e.g.) modulated by the presence of a man or woman.

Savic I, Berglund H, Gulyas B, Roland P. Smelling of odorous sex hormone-like compounds causes sex-differentiated hypothalamic activations in humans.
Neuron. 2001 Aug 30;31(4):661-8. KEY FINDING: Pheromones either from men or from women are processed in sexually dimorphic regions male and female brain. (see also Savic et al., 2005).

Jacob S, Kinnunen LH, Metz J, Cooper M, McClintock MK. Sustained human chemosignal unconsciously alters brain function. Neuroreport. 2001 Aug 8;12(11):2391-4.

Singh D, Bronstad PM. Female body odour is a potential cue to ovulation.
Proc Biol Sci. 2001 Apr 22;268(1469):797-801. KEY FINDING: Fertile (i.e., ovulating) women produce the most pleasant odor: evidence against the "concealed ovulation," which is a basic premise in evolutionary psychology.

Shinohara K, Morofushi M, Funabashi T, Kimura F. Axillary pheromones modulate pulsatile LH secretion in humans. Neuroreport. 2001 Apr 17;12(5):893-5.

Schaefer ML, Young DA, Restrepo D. Olfactory fingerprints for major histocompatibility complex-determined body odors. J Neurosci. 2001 Apr 1;21(7):2481-7. Quote: “Recognition of individual body odors is analogous to human face recognition in that it provides information about identity.”

Chen D, Haviland-Jones J. Human olfactory communication of emotion. Percept Mot Skills. 2000 Dec;91(3 Pt 1):771-81. KEY FINDING Nonhuman animals communicate their emotional states through changes in body odor. Information in human body odors also is indicative of emotional state. This finding introduces new complexity in how humans perceive and interact.

Wedekind C, Penn D. MHC genes, body odours, and odour preferences.
Nephrol Dial Transplant. 2000 Sep;15(9):1269-71. Review. “Besides work on the MHC, there are other findings that indicate human odours play a role in sexual behaviour: …, (ii) pheromones influence women's reproductive synchrony, and (iii) women prefer the odour of physically symmetrical men.”

Rodriguez I, Greer CA, Mok MY, Mombaerts P. A putative pheromone receptor gene expressed in human olfactory mucosa. Nat Genet. 2000 Sep;26(1):18-9.

Morofushi M, Shinohara K, Funabashi T, Kimura F. Positive relationship between menstrual synchrony and ability to smell 5alpha-androst-16-en-3alpha-ol.
Chem Senses. 2000 Aug;25(4):407-11. KEY FINDING: Women whose menstrual cycles synchronized with room-mates had higher olfactory acuity for androstenol.

Shinohara K, Morofushi M, Funabashi T, Mitsushima D, Kimura F. Effects of 5alpha-androst-16-en-3alpha-ol on the pulsatile secretion of luteinizing hormone in human females. Chem Senses. 2000 Aug;25(4):465-7. KEY FINDING: "androstenol retards the growth and maturation of ovarian follicles and consequently delays the timing of ovulation."

Grosser BI, Monti-Bloch L, Jennings-White C, Berliner DL. Behavioral and electrophysiological effects of androstadienone, a human pheromone.
Psychoneuroendocrinology. 2000 Apr;25(3):289-99.  KEY FINDING: Administration of this steroid [androstadienone] under these conditions results in a significant reduction of nervousness, tension and other negative feeling states. Concordant changes were observed in autonomic physiology.

Jacob S, McClintock MK. Psychological state and mood effects of steroidal chemosignals in women and men. Horm Behav. 2000 Feb;37(1):57-78. KEY FINDING: Putative human pheromones are " psychologically potent, mandating future work delineating their function - i.e., whether these steroids are communicative chemosignals, context specific, or related to unconscious associations."

Barni T, Maggi M, Fantoni G, Granchi S, Mancina R, Gulisano M, Marra F, Macorsini E, Luconi M, Rotella C, Serio M, Balboni GC, Vannelli GB. Sex steroids and odorants modulate gonadotropin-releasing hormone secretion in primary cultures of human olfactory cells. J Clin Endocrinol Metab. 1999 Nov;84(11):4266-73. Quote: “This is the first report on GnRH production in human olfactory cells exposed to a selective odorant. The regulation of GnRH expression by sex steroids and odorants can help elucidate the complex neuroendocrine network that controls human reproductive behavior.”

Sobel N, Prabhakaran V, Hartley CA, Desmond JE, Glover GH, Sullivan EV, Gabrieli JD.  Blind smell: brain activation induced by an undetected air-borne chemical. Brain. 1999 Feb;122 ( Pt 2):209-17. Quote: "These findings localize human brain activation that was induced by an undetectable air-borne chemical (the low concentration compound).”

Thornhill, R & Gangestad, SW (1999) The Scent of Symmetry: A Human Sex Pheromone that Signals Fitness? Evolution and Human Behavior 20, 175-201.

Porter RH, Winberg J. Unique salience of maternal breast odors for newborn infants. Neurosci Biobehav Rev. 1999;23(3):439-49. Review. Quote: "Early odor-based recognition may be an important factor in the development of the infant-mother bond.”

Gangestad SW, Thornhill R. Menstrual cycle variation in women's preferences for the scent of symmetrical men. Proc Biol Sci. 1998 May 22;265(1399):927-33. KEY FINDING: Women use olfactory cues as honest signals about certain qualities of men, especially when conception is possible.

Stern K, McClintock MK. Regulation of ovulation by human pheromones.
Nature. 1998 Mar 12;392(6672):177-9. Quote: "By showing in a fully controlled experiment that the timing of ovulation can be manipulated, this study provides definitive evidence of human pheromones." KEY FINDING: The pheromones of women change hormone levels (LH and FSH) in other women.

Winberg J, Porter RH. Olfaction and human neonatal behaviour: clinical implications. Acta Paediatr. 1998 Jan;87(1):6-10. Review. Quote: "New knowledge about human odour physiology may have diagnostic and therapeutic implications…"

Grammer K, Jutte A. [Battle of odors: significance of pheromones for human reproduction] Gynakol Geburtshilfliche Rundsch. 1997;37(3):150-3. Review. German. KEY CONCEPT: Female pheromones (copulins), which are present in vaginal secretions, influence male perception of females and may induce hormonal changes in males.

Ober C, Weitkamp LR, Cox N, Dytch H, Kostyu D, Elias S. HLA and mate choice in humans. Am J Hum Genet. 1997 Sep;61(3):497-504.   Quote: "These results are consistent with the conclusion that Hutterite mate choice is influenced by HLA haplotypes, with an avoidance of spouses with haplotypes that are the same as one's own."

Diamond M, Binstock T, Kohl JV.  From fertilization to adult sexual behavior:
Nonhormonal influences on sexual behavior. Horm Behav. 1996 Dec;30(4):333-53.

Kohl, JV.  Human pheromones: Mammalian olfactory, genetic, neuronal, hormonal and behavioral reciprocity, and human sexuality. Advances in Human Behavior and Evolution. 1996.

Wedekind C, Seebeck T, Bettens F, Paepke AJ. MHC-dependent mate preferences in humans. Proc Biol Sci. 1995 Jun 22;260(1359):245-9.

Fan W, Liu YC, Parimoo S, Weissman SM. Olfactory receptor-like genes are located in the human major histocompatibility complex. Genomics. 1995 May 1;27(1):119-23.

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In other mammals, pheromones influence the onset of puberty, and are also important in establishing nurturing behaviour. These articles, despite no significant mention of pheromones, strongly suggest that human pheromones influence the onset of puberty:

Ellis BJ, Garber J. Psychosocial antecedents of variation in girls' pubertal timing: maternal depression, stepfather presence, and marital and family stress.
Child Dev. 2000 Mar-Apr;71(2):485-501. Quote: "Stepfather presence, rather than biological father absence, best accounted for earlier pubertal maturation in girls living apart from their biological fathers."

Ellis BJ, McFadyen-Ketchum S, Dodge KA, Pettit GS, Bates JE. Quality of early family relationships and individual differences in the timing of pubertal maturation in girls: a longitudinal test of an evolutionary model. J Pers Soc Psychol. 1999 Aug;77(2):387-401. Quote: "In total, the quality of fathers' investment in the family emerged as the most important feature of the proximal family environment relative to daughters' pubertal timing."

In other mammals, pheromones from the female alter levels of testosterone in the male. Here are three studies that strongly suggest pheromones from women alter levels of testosterone in men.

Gray PB, Yang CF, Pope HG Jr. Fathers have lower salivary testosterone levels than unmarried men and married non-fathers in Beijing, China.
Proc Biol Sci. 2006 Feb 7;273(1584):333-9.

Gray PB, Campbell BC, Marlowe FW, Lipson SF, Ellison PT.
Social variables predict between-subject but not day-to-day variation in the testosterone of US men. Psychoneuroendocrinology. 2004 Oct;29(9):1153-62.
KEY FINDING: Married men had lower evening T levels than unmarried men.

Junk science

Cutler WB, Friedmann E, McCoy NL. Pheromonal influences on sociosexual behavior in men. Arch Sex Behav. 1998 Feb;27(1):1-13. Quote: "These initial data need replication but suggest that human male pheromones affected the sexual attractiveness of men to women." Results refuted in Wysocki CJ, Preti G. Pheromonal influences. Arch Sex Behav. 1998 Dec;27(6):627-34.

McCoy NL, Pitino L. Pheromonal influences on sociosexual behavior in young women. Physiol Behav. 2002 Mar;75(3):367-75. Quote: "Three or more sociosexual behaviors increased over baseline for 74% of pheromone users compared with 23% of placebo users." Results refuted in Winman A. Do perfume additives termed human pheromones warrant being termed pheromones? Physiol Behav. 2004 Sep 30;82(4):697-701.

Friebely J, Rako S. Pheromonal influences on sociosexual behavior in postmenopausal women. J Sex Res. 2004 Nov;41(4):372-80. Quote: "These results suggest that the pheromone formulation worn with perfume for a period of 6 weeks has sex-attractant effects for postmenopausal women." Note: WB Cutler's proprietary "pheromone formulation" was used. Non-disclosure of the pheromone formulation means that the study cannot be replicated by independent research.

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Recent mammalian studies supporting the likelihood that olfactory input conditions our visual response.

Published in 2006

Coureaud G, Moncomble AS, Montigny D, Dewas M, Perrier G, Schaal B. A Pheromone That Rapidly Promotes Learning in the Newborn. Curr Biol. 2006 Oct 10;16(19):1956-1961. Key Issue: The unconscious affect of a rabbit pheromone can act either as a reinforcing agent, or a one-trial conditioning agent. Reinforcement of the most likely hormone response (e.g., a downstream effect of gonadotropin-releasing hormone) could occur when the pheromone converts an associated secondary odor into a conditioned stimulus. Extension to humans of this mammalian model indicates that either maternal natural body odor, or the body odor of other people, may become a conditioned stimulus that evokes a behavioral response, even in the absence of the pheromone during subsequent encounters. This suggests that a single exposure to a pheromone may elicit powerful long-term behavioural affects.

Gelez H, Fabre-Nys C. Role of the olfactory systems and importance of learning in the ewes' response to rams or their odors. Reprod Nutr Dev. 2006 Jul-Aug;46(4):401-15. Quote: “ Our findings support the idea that processes of olfactory cues detected by the main olfactory system can involve cognitive or learning mechanisms. The response to the ram odor does not correspond to preprogrammed or reflex responses, but rather responses to a stimulus that had acquired a meaning.

Alekseyenko OV, Baum MJ, Cherry JA. Sex and gonadal steroid modulation of pheromone receptor gene expression in the mouse vomeronasal organ.
Neuroscience. 2006 Jul 18;140(4):1349-57.

Baxi KN, Dorries KM, Eisthen HL. Is the vomeronasal system really specialized for detecting pheromones? Trends Neurosci. 2006 Jan;29(1):1-7.

Spehr M, Kelliher KR, Li XH, Boehm T, Leinders-Zufall T, Zufall F. Essential role of the main olfactory system in social recognition of major histocompatibility complex peptide ligands. J Neurosci. 2006 Feb 15;26(7):1961-70. Quote: “Our finding that MHC peptide ligands can gain access to and be recognized by sensory neurons of the MOE will prompt further investigations aimed at understanding whether evolutionary conserved MHC peptide ligands also function as odor signals in humans.”

Laska M, Wieser A, Salazar LT. Sex-specific differences in olfactory sensitivity for putative human pheromones in nonhuman primates. J Comp Psychol. 2006 May;120(2):106-12. Quote: "...—at least within the order of primates—allometric comparisons of olfactory brain structures or differences in the number of
functional olfactory receptor genes do not allow us to draw generalizing
conclusions as to olfactory sensitivity of any two species."

Kaminski RM, Marini H, Ortinski PI, Vicini S, Rogawski MA. The pheromone androstenol (5 alpha-androst-16-en-3 alpha-ol) is a neurosteroid positive modulator of GABAA receptors. J Pharmacol Exp Ther. 2006 May;317(2):694-703. Quote: "Androstenol and androstenone are among the best-accepted mammalian pheromones (Grammer et al., 2005). The present results raise the possibility that effects on GABAA receptors could contribute to
their pheromonal activity."

Spehr M, Spehr J, Ukhanov K, Kelliher KR, Leinders-Zufall T, Zufall F.

Parallel processing of social signals by the mammalian main and accessory olfactory systems. Cell Mol Life Sci. 2006 May 29; [Epub ahead of print]
Quote: “… the main and accessory olfactory systems should be viewed as complementary rather than separate pathways for chemical communication. Taken together, these developments add up to what has been called ‘a revolution in our understanding of the role of smell in controlling the neuroendocrine brain’.

Keller M, Douhard Q, Baum MJ, Bakker J. Destruction of the main olfactory epithelium reduces female sexual behavior and olfactory investigation in female mice. Chem Senses. 2006 May;31(4):315-23. KEY FINDING: The main olfactory epithelium is primarily involved in the detection and processing of odors that are used to localize and identify the sex and hormone status of conspecifics.

Gelez H, Fabre-Nys C. Neural pathways involved in the endocrine response of anestrous ewes to the male or its odor. Neuroscience. 2006;140(3):791-800. Quote: The demonstration of the role of the main olfactory system, the role of sexual experience, and the importance of other sensory cues other than olfaction in the ‘‘male effect’’ support the idea that the endocrine response elicited by the ram or its odor does not involve basic mechanisms corresponding to reflex or automatic responses, but rather elaborate treatment integration of olfactory or other sensory cues and resulting behavioral changes and learning processes.

Published in 2005

Roth TL, Sullivan RM. Memory of early maltreatment: neonatal behavioral and neural correlates of maternal maltreatment within the context of classical conditioning. Biol Psychiatry. 2005 Apr 15;57(8):823-31. Quote: "RESULTS: Odor-maternal maltreatment pairings within a seminatural setting and odor-shock pairings both resulted in paradoxical odor preferences. Learning-induced gene expression was altered in the olfactory bulb and anterior piriform cortex (part of olfactory cortex) but not the amygdala. CONCLUSIONS: Infants appear to use a unique brain circuit that optimizes learned odor preferences necessary for attachment."

Coria-Avila GA, Ouimet AJ, Pacheco P, Manzo J, Pfaus JG. Olfactory conditioned partner preference in the female rat. Behav Neurosci. 2005 Jun;119(3):716-25. Quote:" An odour paired with the ability of females to
pace copulation becomes a sexual incentive, such that males bearing the odor are preferred when females are given a choice between scented and unscented males. Thus, just like male rats, female rats can learn to modify their sexual behavior and partner preferences on the basis of experience with sexual reward."

Yoon H, Enquist LW, Dulac C. Olfactory inputs to hypothalamic neurons controlling reproduction and fertility. Cell. 2005 Nov 18;123(4):669-82. KEY FINDING: There is a lack of anatomical and functional connection between VNO and the central regulators of mammalian reproduction: the GnRH neurons. There is a direct link these GnRH neurons and the olfactory and somatosensory pathways that play an essential role in reproductive behavior and in the modulation of GnRH neuronal activity.

Boehm U, Zou Z, Buck LB. Feedback loops link odor and pheromone signaling with reproduction. Cell. 2005 Nov 18;123(4):683-95. Abstract excerpt::  “…approximately 800 GnRH neurons communicate with approximately 50,000 neurons in 53 functionally diverse brain areas, with some connections exhibiting sexual dimorphism. These studies reveal a complex interplay between reproduction and other functions in which GnRH neurons appear to integrate information from multiple sources and modulate a variety of brain functions.”

Published in 2004

Roselli CE, Larkin K, Resko JA, Stellflug JN, Stormshak F. The volume of a sexually dimorphic nucleus in the ovine medial preoptic area/anterior hypothalamus varies with sexual partner preference. Endocrinology. 2004 Feb; 145(2): 478-83. KEY INDICATION: A mammalian model linking pheromones and olfaction to neuroanatomy and neuroendocrinology may extend to encompass aspects of human sexual orientation.

Reyes R, Mendoza J, Ballesteros J, Moffatt C. Male chemosignals inhibit the neural responses of male mice to female chemosignals. Brain Res Bull. 2004 May 30;63(4):301-8. KEY FINDING: "...male urine inhibited the responses of cells within the MOS and AOS to female urine."

Meredith M, Westberry JM. Distinctive responses in the medial amygdala to same-species and different-species pheromones. J Neurosci. 2004 Jun 23;24(25):5719-25. "This is the first evidence for an important role of the amygdala, a limbic structure known to be involved in social and emotional behavior, in discrimination of species specificity in chemosignals."

Richardson HN, Nelson AL, Ahmed EI, Parfitt DB, Romeo RD, Sisk CL.
Female pheromones stimulate release of luteinizing hormone and testosterone without altering GnRH mRNA in adult male Syrian hamsters (Mesocricetus auratus). Gen Comp Endocrinol. 2004 Sep 15;138(3):211-7. Quote: "The absence of a demonstrable change in GnRH mRNA in response to the presumed increased release of GnRH suggests either that the magnitude of the GnRH response to female pheromones does not require significant increases in GnRH synthesis for replenishment of released peptide, or that GnRH mRNA and release are impacted by pheromones in a relatively small number of GnRH neurons."

Gelez H, Archer E, Chesneau D, Campan R, Fabre-Nys C. Importance of learning in the response of ewes to male odor. Chem Senses. 2004 Sep;29(7):555-63. KEY ISSUES: Ram exposure activates the LH response in ewes; sexual experience is a factor. After pairing lavendar scent with ram exposure; the lavendar scent activated the LH response in ewes. Simply put, ewes can learn to associate an arbitrary odor with the affect of pheromones on their level of LH.

Gelez H, Fabre-Nys C. The "male effect" in sheep and goats: a review of the respective roles of the two olfactory systems. Horm Behav. 2004 Sep;46(3):257-71. Review. KEY FINDING: The VNO is not required for rams to elicit an LH response from ewes.

Published before 2004

Westberry J, Meredith M. The influence of chemosensory input and gonadotropin releasing hormone on mating behavior circuits in male hamsters. Brain Res. 2003 Jun 6;974(1-2):1-16. KEY FINDING "...the combination of pheromone exposure and intracerebrally-injected GnRH increases Fos expression in the MPOA above the increase seen in pheromone-exposed males, or in males given only the exogenous GnRH. In males with vomeronasal organs removed (VNX), there was an also an increment in Fos expression in the MPOA when these pheromone exposed males were injected with GnRH, provided they had previous sexual experience. Males with vomeronasal organs removed and without sexual experience showed increased Fos expression in the medial amygdala when pheromone exposure and GnRH injection were combined, but not in the medial preoptic area."

Westberry JM, Meredith M. Pre-exposure to female chemosignals or intracerebral GnRH restores mating behavior in naive male hamsters with vomeronasal organ lesions. Chem Senses. 2003 Mar;28(3):191-6.  RATIONALE: Hamster vaginal fluid and  intracerebroventricular injections of GnRH eliminate mating deficits normally seen in naive male hamsters with vomeronasal organs removed. KEY FINDING: The action of  female pheromones on GnRH  is the most likely link to restoration of  naive males who have had their vomeronasal organs removed.

Roselli CE, Resko JA, Stormshak F.  Estrogen synthesis in fetal sheep brain: effect of maternal treatment with an aromatase inhibitor. Biol Reprod. 2003 Feb; 68(2): 370-4. KEY INDICATION: Aromatase activity might be linked to estradiol receptor content in the amygdala (an olfactory processing center) which varies with sexual orientation in rams. This fits well with what is known about sexual differentiation of the mammalian olfactory system(s).

Schaefer ML, Yamazaki K, Osada K, Restrepo D, Beauchamp GK. Olfactory fingerprints for major histocompatibility complex-determined body odors II: relationship among odor maps, genetics, odor composition, and behavior.
J Neurosci. 2002 Nov 1;22(21):9513-21. Quote: “…modification of a single gene… of the major histocompatibility locus… results in a subtle change in the odiferous quality of urine, causes a small but significant change in the composition of urine volatiles and consequently the evoked glomerular activation pattern in the MOB. [This change]… is predictive of the extent of (1) the genetic difference among the urine donors, (2) the difference in the chemical composition of urine, and (3) the odor detector's ability to discriminate…individual recognition.”

Del Punta K, Leinders-Zufall T, Rodriguez I, Jukam D, Wysocki CJ, Ogawa S, Zufall F, Mombaerts P. Deficient pheromone responses in mice lacking a cluster of vomeronasal receptor genes. Nature. 2002 Sep 5;419(6902):70-4.

Moncho-Bogani J, Lanuza E, Hernandez A, Novejarque A, Martinez-Garcia F. Attractive properties of sexual pheromones in mice: innate or learned?
Physiol Behav. 2002 Sep;77(1):167-76. KEY INDICATION  Pavlovian-like associative learning (i.e., classical conditioning) in which previously neutral volatiles (very likely odorants) acquire attractive properties by association with the nonvolatile, innately attractive pheromone(s) is the likely basis for not only the sexual but also the 'chemical' experience (previous experience with sexual pheromones), which must be taken into account to interpret the role of chemicals as releaser or primer pheromones.

Takami S. Recent progress in the neurobiology of the vomeronasal organ.
Microsc Res Tech. 2002 Aug 1;58(3):228-50. Review.

Beckman M. Pheromone reception. When in doubt, mice mate rather than hate.
Science. 2002 Feb 1;295(5556):782.

Kendrick KM, Haupt MA, Hinton MR, Broad KD, Skinner JD. Sex differences in the influence of mothers on the sociosexual preferences of their offspring.
Horm Behav. 2001 Sep;40(2):322-38.

Kelliher KR, Baum MJ. Nares occlusion eliminates heterosexual partner selection without disrupting coitus in ferrets of both sexes. J Neurosci. 2001 Aug 1;21(15):5832-40.

Johnson BN, Mainland JD, Sobel N. Rapid olfactory processing implicates subcortical control of an olfactomotor system. J Neurophysiol. 2003 Aug;90(2):1084-94. KEY INDICATION The time course of sniffing appears to be more rapid than a response that requires cortical control. "Considering that odorant transduction takes around 150 ms and odorant-induced cortical evoked potentials have latencies of around 300 ms, the rapid motor adjustments measured here suggest that olfactomotor sniff feedback control is subcortical and may rely on neural mechanisms similar to those that modulate eye movements to accommodate vision and ear movements to accommodate audition."

Kippin TE, Pfaus JG. The nature of the conditioned response mediating olfactory conditioned ejaculatory preference in the male rat. Behav Brain Res. 2001 Jul;122(1):11-24.

Brennan PA, Schellinck HM, de la Riva C, Kendrick KM, Keverne EB.
Changes in neurotransmitter release in the main olfactory bulb following an olfactory conditioning procedure in mice. Neuroscience. 1998 Dec; 87(3): 583-90. KEY INDICATION: Changes in synaptic connectivity occur in response to odors during classically conditioned behavior; which may be a general feature of olfactory learning. COMMENT: If the above is true, it becomes more important for others to realize that pheromones cause changes in hormone levels during development, and these changes cause changes in synaptic connectivity that varies in males and females..

Katz LS, Price EO, Wallach SJ, Zenchak JJ.
Sexual performance of rams reared with or without females after weaning.
J Anim Sci. 1988 May; 66(5): 1166-73. KEY INDICATION: Pheromone exposure during development might play an important role in adult  sexual orientation and in sexual behavior.

McClintock MK. On the nature of mammalian and human pheromones.
Ann N Y Acad Sci. 1998 Nov 30;855:390-2.

Johnston RE.  Pheromones, the vomeronasal system, and communication. From hormonal responses to individual recognition. Ann N Y Acad Sci. 1998 Nov 30;855:333-48. Review. Quote: "In sum, it is important to maintain a broad, balanced view and to avoid oversimplifications if we are to advance our understanding of the sensory mechanisms underlying responses to chemical signals and other odors."

Meredith M. Vomeronasal, olfactory, hormonal convergence in the brain. Cooperation or coincidence? Ann N Y Acad Sci. 1998 Nov 30;855:349-61. Review. Quote:  "Despite evidence [in hamsters] that the release of LHRH in response to female chemosignals is dependent on vomeronasal input, no activation of LHRH neurons has been demonstrated in males exposed to such chemosignals."

Wood RI. Integration of chemosensory and hormonal input in the male Syrian hamster brain. Ann N Y Acad Sci. 1998 Nov 30;855:362-72. Review. Quote: "According to our current model, hormones may act as a gating signal to strengthen synaptic contacts along the chemosensory pathway, thereby permitting or enhancing transmission of chemosensory cues."

Monti-Bloch L, Jennings-White C, Berliner DL. The human vomeronasal system. A review.Ann N Y Acad Sci. 1998 Nov 30;855:373-89. Review. Quote: "These findings present new information supportive of a functional vomeronasal system in adult humans."

Monti-Bloch L, Diaz-Sanchez V, Jennings-White C, Berliner DL. Modulation of serum testosterone and autonomic function through stimulation of the male human vomeronasal organ (VNO) with pregna-4,20-diene-3,6-dione. J Steroid Biochem Mol Biol. 1998 Apr;65(1-6):237-42. KEY FINDING: Humans have a functional VNO and respond to pheromones with hormonal change.

Berliner DL, Monti-Bloch L, Jennings-White C, Diaz-Sanchez V. The functionality of the human vomeronasal organ (VNO): evidence for steroid receptors. J Steroid Biochem Mol Biol. 1996 Jun;58(3):259-65. KEY FINDING: existence of a human vomeronasal-hypothalamic-pituitary-gonadal system.

Monti-Bloch L, Jennings-White C, Dolberg DS, Berliner DL. The human vomeronasal system. Psychoneuroendocrinology. 1994;19(5-7):673-86.