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Published in 2007

Books and book chapters

van den Hurk, R. (2007). Intraspecific chemical communication in vertebrates with special attention to its role in reproduction. Pheromone Information Centre, Zeist, The Netherlands. (available from author for 15 euro or $20 US -- Paypal accepted: contact r.vandenhurk@planet.nl) Note: This is the most recent comprehensive review that is currently available.

Herz, R. (2007).  The Scent of Desire: Discovering Our Enigmatic Sense of Smell. William Morrow, New York. Note: This is a well-written easy to understand primer on the role that odor plays in life and, perhaps, in love.

Kohl, JV (2007). The Mind's Eyes: Human pheromones, neuroscience, and male sexual preferences. In M. R. Kauth (Ed.), Handbook of the Evolution of Human Sexuality (pp. 313-369). Binghamton: Haworth Press. 

Hoffmann, H. (2007). The role of classical conditioning in sexual arousal. In E. Janssen (Ed.), The psychophysiology of sex. Bloomington: Indiana University Press.

Articles and presentations

Gottfried JA. What can an orbitofrontal cortex-endowed animal do with smells? Ann N Y Acad Sci. 2007 Dec;1121:102-20. Quote: “Specifically, as an epitome of the neuro-behavioral interface between sensation, learning, and experience, the following section describes two recent functional magnetic resonance imaging (fMRI) experiments from our laboratory demonstrating the role of human OFC in olfactory perceptual learning and associative (Pavlovian) conditioning.”

Berglund, H., Lindstrom, P., Dhejne-Helmy, C., & Savic, I. (2007). Male-to-Female Transsexuals Show Sex-Atypical Hypothalamus Activation When Smelling Odorous Steroids. Cereb Cortex, 3, 3. Quote: "...suggests that in transsexuals the organization of certain sexually dimorphic circuits of the anterior hypothalamus could be sex atypical. It adds a new dimension to our previous reports by showing that the observed effects are not necessarily learned and that a sex-atypical activation by the 2 putative pheromones may reflect neuronal reorganization." 

Chen, D., and Zhou, W. (2007). Relationship between Olfactory and Emotional Competencies, Association for Chemoreception Sciences 29th Annual Meeting, Sarasota, Florida.

Dematte, M. L., Osterbauer, R., and Spence, C. (2007). Olfactory Cues Modulate Facial Attractiveness. Chem. Senses, 32(6):603-10. Quote:"... the results of the present study add to a growing list of studies demonstrating that the presence of olfactory cues can exert a small but significant crossmodal influence on people’s judgments of a variety of nonolfactory stimulus attributes/qualities..."

Doucet, S., Soussignan, R., Sagot, P., and Schaal, B. (2007). The "smellscape" of mother's breast: effects of odor masking and selective unmasking on neonatal arousal, oral, and visual responses. Dev Psychobiol. 49(2), 129-38. Quote: "… the findings of the present study confirm that the effluvium of the human mother’s breast… delays the onset of crying and activates appetitive behavior, and (at least in males) tends to promote visual input in 3–4 day-old newborns. Further, the odor of the whole breast and the isolated odors of the areola, of the nipple or of milk appear comparable in releasing these behavioral effects." 

Goubet, N., Strasbaugh, K., and Chesney, J. (2007). Familiarity breeds content? Soothing effect of a familiar odor on full-term newborns. J Dev Behav Pediatr. 28(3), 189-94. Abstract excerpt: “A familiar odor is effective in significantly reducing crying and grimacing during a minor painful procedure.” 

Jacob, T., Wang, L., Boulkroune, N., March, A., and Walker, N. (2007). Changes in olfactory threshold, hedonics and brain activity in response to repetitive exposure to androstadienone, Association for Chemoreception Sciences 29th Annual Meeting, Sarasota, Florida. 

Kelahan, LC, Hoffmann, H, & Kohl, JV (2007). Putative human pheromones may condition a human hormonal effect/behavioral affect. Poster presented at the Society for the Scientific Study of Sexuality Annual Meeting, Indianapolis, IN, Nov 7-11. 

Kelahan, LC, Hoffmann, H, Kohl, JV, & Shea, A. (2007). Putative human pheromones may condition a human hormonal effect/behavioral affect. Poster presented at the Association for Chemoreception Sciences 29th Annual Meeting, Sarasota, Florida, Apr 25-29. 

Keller, A., Zhuang, H., Chi, Q., Vosshall, L. B., & Matsunami, H. (2007). Genetic variation in a human odorant receptor alters odour perception. Nature,  449(7161):468-72. Quotes: “We identify OR7D4 as a significant heritable factor influencing androstenone and androstadienone perception, thus providing the first reported link between genetic polymorphisms in an odorant receptor gene and altered perception of the ligands that activate this receptor.” “In this study we investigated only the olfactory percept reported when odorous steroids were sniffed, but olfactory exposure to androstenone and androstadienone has also been shown to induce several physiological responses in both men and women.” 

Kohl, JV (2007). The Mind's Eyes: Modeling the Development of Diverse Sexual Preferences. Paper presented at the Society for the Scientific Study of Sexuality Annual Meeting, Indianapolis, IN, Nov 7-11. (powerpoint presentation available for $10.00 US via paypal to jvkohl@bellsouth.net) 

Li, W., Moallem, I., Paller, K. A., & Gottfried, J. A. (2007). Subliminal smells can guide social preferences. Psychol Sci., 18(12), 1044-1049. Quote:”… the time-honored belief that scents play an important role in human social settings appears to withstand scientific scrutiny. Furthermore, our data suggest that it is in the absence of conscious awareness that odors best exert their effects.”

Oberzaucher, E., Grammer, K., Zimmer, K., Fischer, G., Soini, M. V., Dixon, S. J., Xu, Y., Zomer, S., Brereton, R. G., and Penn, D. J. (2007). The Identification of Compounds in Human Sweat - Signals of Individuality, Gender and Genes, Association for Chemoreception Sciences 29th Annual Meeting, Sarasota, Florida.

Roney, J. R., Lukaszewski, A. W., & Simmons, Z. L. (2007). Rapid endocrine responses of young men to social interactions with young women. Horm Behav, 52, 326-333. Quote: “…the similarities between the hormonal responses reported here and those seen in many nonhuman species support the possibility that phylogenetically conserved brain mechanisms may play important roles in the regulation of men's reactions to social encounters with potential mates.”

Sergeant, M. J., Dickins, T. E., Davies, M. N., & Griffiths, M. D. (2007). Women's Hedonic Ratings of Body Odor of Heterosexual and Homosexual Men. Arch Sex Behav., 36(3), 395-401. Quote: “…it appears that an individual’s sexual orientation has a significant impact on their olfactory function, both in terms of their body odor production and perceptions of certain odorants.”

Timon Andrada, R., Maynar Marino, M., Munoz Marin, D., Olcina Camacho, G. J., Caballero, M. J., and Maynar Marino, J. I. (2007). Variations in urine excretion of steroid hormones after an acute session and after a 4-week programme of strength training. Eur J Appl Physiol. 99(1), 65-71.KEY FINDING: Strength training increases the production of putative human pheromones associated with masculinity.

Thevis, M., Geyer, H., Mareck, U., Sigmund, G., Henke, J., Henke, L., et al. (2007). Detection of manipulation in doping control urine sample collection: a multidisciplinary approach to determine identical urine samples. Anal Bioanal Chem., 388(7), 1539-1543. KEY FINDING: Putative human pheromones and other measures of steroid hormone metabolism were used to identify one urine sample amoung 14,700 other samples. These authors found 3 indentical samples that were supposedly submitted by three different individuals, and "busted" the offenders for "doping." 

Wang, Z., Nudelman, A., & Storm, D. R. (2007). Are pheromones detected through the main olfactory epithelium? Mol Neurobiol., 35(3), 317-323. Conclusion: In light of recent studies in mice which indicate that main olfactory inputs can modulate pheromone responses in gonadotropin releasing hormone neurons in the hypothalamus, it is plausible that pheromone responses to androstadienone and estratetraenol also originate in the main olfactory epithelium. 

Wedekind C, et al.,  (2007). The major histocompatibility complex and perfumers’ descriptions of human body odors. Evolutionary Psychology 5 (2), 330-343. PDF.  KEY FINDING: It seems obvious that both the MHC and volatile steroids conrtibute to human body odor. 

Whittle, C. L., Fakharzadeh, S., Eades, J., and Preti, G. (2007). Human Breath Odors and Their Use in Diagnosis. Ann NY Acad Sci 1098(1), 252-266. KEY ISSUE: The authors developed a protocol to help differentiate individuals with chronic halitosis from those with the genetic, odor-producing metabolic disorder trimethylaminuria (TMAU). Quote: "…the assumption that the individual with TMAU will always smell “like fish” is incorrect…”

Penn DJ, Oberzaucher E, Grammer K, Fischer G, Soini HA, Wiesler D, Novotny MV, Dixon SJ, Xu Y, Brereton RG. Individual and gender fingerprints in human body odour. J R Soc Interface. 2007 Apr 22;4(13):331-340  Quote: “We found that although the axillary sweat of men and women had remarkably similar GC–MS profiles, we could statistically discriminate the sexes, and we identified the chemical structures of 12 of these marker compounds characteristic of gender.”

Kline JP, Schwartz GE, Dikman ZV. Interpersonal defensiveness and diminished perceptual acuity for the odor of a putative pheromone: Androstenone. Biol Psychol. 2007 Mar;74(3):405-13. PDF Key Issue: The pattern of associations with androstenone sensitivity is sexually dimorphic. Dislike for androstenone has been associated with sensitivity to it in women, but not in men . This suggests that the quality and degree of perception of androstenone may manifest quite differently for women and men. The nature of these sex differences is poorly understood.

Wyart C, Webster WW, Chen JH, Wilson SR, McClary A, Khan RM, Sobel N. Smelling a Single Component of Male Sweat Alters Levels of Cortisol in Women. J. Neurosci. 2007 Feb; 27(6):1261-1265. Key Finding: Androstadienone exposure influences cortisol levels, mood, and sexual arousal in women. 

Porter J, Craven B, Khan RM, Chang SJ, Kang I, Judkewicz B, Volpe J, Settles G, Sobel N. Mechanisms of scent-tracking in humans. Nat Neurosci. 2007 Jan;10(1):27-9. Quote: “Here we found that not only are humans capable of the demanding macrosmatic behavior of scent-tracking, but they spontaneously mimic the tracking patterns of macrosmatic mammals.” 

Haselton MG, Mortezaie M, Pillsworth EG, Bleske-Rechek A, Frederick DA. Ovulatory shifts in human female ornamentation: Near ovulation, women dress to impress. Horm Behav. 2007 Jan;51(1):40-5. Quote: “…ovulatory cycle shifts in female sexual interests and male mate guarding are themselves dependent upon contextual cues, including characteristics of the partner. It is not at all evident that this adaptive information processing could be achieved by general cognitive abilities not structured or specialized to solve particular adaptive problems specific to mating…”

Published in 2006

Kohl, JV. The Mind's Eyes: Human Pheromones, Neuroscience, and Male Sexual Preferences. Journal of Psychology & Human Sexuality, 2006 Dec 18(4):313-369.

Brennan PA, Kendrick KM Mammalian social odours: attraction and individual recognition. Philosophical Transactions of the Royal Society B: Biological Sciences. 2006 Dec; 361(1476) 2061–2078. Quote: "Finally, although the sense of smell is often relegated to a minor role in human social communication, evidence is starting to accumulate that odours have not lost the ability to influence human behaviour. The extent to which social odours play a role in modern human society is an open but intriguing question for future research." 

Brennan PA, Zufall F. Pheromonal communication in vertebrates. Nature. 2006 Nov 16;444(7117):308-15. Quote: "More problematic for advocates of the existence of a human VNO are the consistent failure to find vomeronasal nerves projecting to the brain and the failure to find an AOB in adult humans."

Garver-Apgar CE, Gangestad SW, Thornhill R, Miller RD, Olp JJ. Major histocompatibility complex alleles, sexual responsivity, and unfaithfulness in romantic couples. Psychol Sci. 2006 Oct;17(10):830-5. Quote: “…as MHC sharing increases, women report more extrapair partners (but only in the current relationship) and experience greater attraction to extrapair men relative to their partners, particularly on fertile days of their cycles. These effects may be mediated by scent,…”

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. PDF 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 Aug 10;442(7103):645-50. 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.

Ngai J. Neuroscience: an extra dimension to olfaction. Nature. 2006 Aug 10;442(7103):637-8. (Comment on Liberles and Buck). 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.” 

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.” 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. PDF 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.” 

Jacob TJ, Wang L, Jaffer S, McPhee S. Changes in the odor quality of androstadienone during exposure-induced sensitization. Chem Senses. 2006 Jan;31(1):3-8. Key Finding: Detection of androstadienone is experience dependent. detection thresholds during repetitive exposure During repetitive exposure detection thresholds decrease. This sensitization is accompanied by perceptual odor quality changes.

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. 

Hummel T, Krone F, Lundstrom JN, Bartsch O. Androstadienone odor thresholds in adolescents. Horm Behav. 2005 Mar;47(3):306-10. Key finding: Androstadienone sensitivity decreases with puberty in males but not females.

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 PDF 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 Behavioral 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."

Pillsworth EG, Haselton MG, Buss DM  Ovulatory shifts in female sexual desire.

J Sex Res. 2004 Feb;41(1):55-65. Key Finding: Women are more attracted to their primary partner at peak fertility within the cycle.

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. 

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.

Pause BM. Is the human skin a pheromone-producing organ? J Cosmet Dermatol. 2004 Dec;3(4):223-8. Quote: “It has been suggested that humans might have a special organ within their nose that can transmit such chemosensory information. However, the evidence for this organ is highly questionable. In any case, the main olfactory system is a highly diverse system, capable of transmitting pheromonal information."

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.

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 The Scent of Symmetry: A Human Sex Pheromone that Signals Fitness? Evolution and Human Behavior. 1999 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 behavior. 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.

Storey AE, Walsh CJ, Quinton RL, Wynne-Edwards KE. Hormonal correlates of paternal responsiveness in new and expectant fathers. Evol Hum Behav. 2000 Mar 1;21(2):79-95. KEY FINDING: Testosterone levels in men decreased as pregnancy in their mate progressed. "This pattern of hormonal change in men suggests that hormones may play a role in priming males to provide care for young."

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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 behavioral 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 Psychiatr