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Abstract Article Metrics

Androstenol has been reported to influence judgements of attractiveness and to Citations

affect participants’ mood. In the present study, participants were asked to sniff
Citation Indexes: 3
androstenol or a control odour (pure ethanol) unilaterally with the left or right
nostril. Subsequently, they rated the attractiveness of photographs of the opposite Captures
sex and their own feelings on four mood scales. Participants rated the photographs
Readers: 14
as significantly more attractive after sniffing androstenol compared with the control
odour. This did not depend upon androstenol being perceived as pleasant.
Androstenol made male participants feel more lively, and both male and female View details

participants more sexy, when sniffed through the right compared with the left
nostril. Participants rated themselves as more irritable and aggressive when
exposed to androstenol through the left nostril. The findings are discussed in
relation to the effects of arousal on attraction and in the context of current theories
of hemispheric differences in emotion.

Introduction

Among many non-human animals the sense of smell is very highly developed; for
some species it is the predominant sense (Stevenson, 2010). In The Descent of Man,
Charles Darwin (1871) observed that “The sense of smell is of highest importance to
the greater number of mammals.But [in humans] the sense of smell is of extremely
slight service, if any.” (p.24). Darwin’s view notwithstanding, it is now apparent that
the sense of smell is important to humans (Bushdid et al., 2014, McGann, 2017,
Stevenson, 2010, Williams and Apicella, 2018). In particular, perception of body
odour appears to be an important feature of human interaction (see Carrito et al.,
2017).

Body odour may be especially important within romantic relationships (Bendas et

al., 2018, Croy et al., 2013). When asked to rank in importance the physical
characteristics of a potential (but hypothetical) lover, women have been reported as
rating “how a man smells” as the most important characteristic, whereas men tend
to rank “how a woman looks” as most important (Herz and Cahill, 1997, Herz and
Inzlicht, 2002, Havlíček et al., 2008; but see Foster, 2008).

Substances contributing to body odour include the 16-androstenes, a group of

compounds that includes 5α-androst-16-en-3α-ol (3α-androstenol), 5α-androst-16-
en-3-one (5α-androstenone), and 4,16-androstadien-3-one (androstadienone)
which are found in several mammals, including pigs and humans (for review see
Gower & Ruparelia, 1993). Produced by the boar in urine, saliva and sweat,
presentation of androstenol to a receptive sow is associated with a rigid stance
(lordosis or immobilisation response) that allows mating to occur (Reed, Melrose, &
Patterson, 1974).

The androstenes might also be involved in sexual attraction and reproductive

activity in humans. As well as activating classical olfactory areas of the cortex,
androstenol (and androstenone) activates the hypothalamus, a part of the brain
involved in reproduction, in a sex-differentiated manner in comparison with more
‘ordinary’ and familiar odours, such as butanol, cedar oil and lavender oil (Burke et
al., 2012, Savic and Berglund, 2010, Savic et al., 2001). Shinohara and colleagues
found that androstenol decreased the frequency of the pulsatile secretion of
luteinizing hormone during the follicular phase of their participants’ menstrual
cycles, an effect thought to be mediated by hypothalamic nuclei (Shinohara,
Morofushi, Funabashi, Mitsushima & Kimura, 2000).

The androstenes are believed to originate in the testes and ovaries (Gower &
Ruparelia, 1993) and are secreted by the apocrine glands. They are found as a
relatively small proportion of the axillary sweat and axillary hair (Gower &
Ruparelia, 1993). Axillary secretions (see Wysocki & Preti, 2004 for a review of
axillary chemistry) are odour free and thought to become (mal)odorous as a result
of bacterial activity (Decréau et al., 2003, Gower et al., 1994, Leyden et al., 1981).
Some recent evidence suggests that axillary odour plays a role in human sexual
communication and interaction. The smell of underarm sweat of single men is rated
by women as stronger than that of partnered men (Mahmut & Stevenson, 2019).
Among men, there is close agreement as to the attractiveness of individual women’s
axillary odour (Lobmaier, Fischbacher, Wirthmüller & Knoch, 2018) and they rate the
under-arm odour of sexually aroused women as more attractive than that of
unaroused women (Wisman & Shrira, 2020).

It has been suggested that the androstene compounds in general might act as
human pheromones (for reviews of the effects of substances argued to be human
pheromones see Grammer, Fink, & Neave, 2005; Mostafa, El Khouly, & Hassan,
2012). However, the question of whether pheromones exist in humans is much
debated. Research in this area has been subject to considerable criticism and
controversy (see Bensafi, Brown, Khan, Levenson & Sobel, 2004; Ferdenzi, Ortegón,
Delplanque, Baldovini & Bensafi, 2020; Havlíček, Murray, Saxton & Roberts, 2010;
Wyatt, 2015, Wyatt, 2020; Wysocki & Preti, 2004).

Pheromones may be defined as chemical signals (molecules) that when released by

an organism elicit a specific reaction, such as a stereotyped behaviour or
developmental process, in members of the same species. According to Wyatt (2015),
claims that there are human pheromones are flawed because they do not arise from
a methodical, bioassay-led approach to their identification. Other authors take a
rather broader view, preferring the term chemical signal defined as “any chemical
emitted by one individual that alters either the behavior or physiology of another
organism” (Williams & Apicella, 2018., p. 122–123).

Numerous studies have been conducted in which the effects of androstenes on

human sexual attraction and mood have been investigated (for review see Havlíček
et al., 2010; Wysocki & Preti, 2004). These fall into two distinct waves or phases.
During the 1980s and 1990s interest centred on androstenol and androstenone;
more recently, for no conspicuous scientific reason, attention has turned to
androstadienone (Havlíček et al., 2010, Wyatt, 2020). However, the focus of the
present paper is on androstenol.

The results of studies investigating the effect of androstenol on ratings of

attractiveness are mixed. Some researchers have reported that androstenol
enhances judgements of physical attractiveness. Kirk-Smith, Booth, Carroll, and
Davies (1978) presented photographs to 12 male and 12 female undergraduates.
When wearing a mask impregnated with androstenol both sexes rated photographs
of other young men and women as more attractive than in the control condition,
when the mask was free of odour (see also Maiworm & Langthaler, 1992). The effect
was strongest for ratings of photographs of women. Filsinger, Braun, & Monte (1985)
asked participants to sniff a bag containing either an androstenol solution or one of
three other solutions. Following this, they read descriptions of a fictional male
character and a fictional female character presented along with photographs
supposedly of the individuals described. Male ratings of the sexual attractiveness of
the male target, but not the female target, were higher in the androstenol condition
than in the other conditions. In a study in which a male and a female confederate of
the experimenters “emanated the odour of androstenol”, a control substance or no
odour, Black & Biron (1982) found no effect of androstenol on ratings of
attractiveness of the opposite sex confederate with whom each participant had
been in close proximity for 15 minutes.

It is possible that the results of the study by Kirk-Smith et al. (1978) were due to a
Hawthorne-like effect (see Jacob, Garcia, Hayreh & McClintock, 2002), in that the
androstenol condition was compared with a no-odour condition. On the other hand,
the negative findings of Black & Biron (1982) might be attributable to participants
spending sufficiently long time in the company of the experimenters’ confederates
for the latters’ personalities to have over-ridden any effect of the odours.

Conceivably, the positive effects of androstenol on ratings of attractiveness that have

been reported thus far would have been produced by any pleasant-smelling odour.
In a ‘classic’ experiment Baron (1981) investigated whether male participants’
ratings of attractiveness of two female confederates with whom they interacted
briefly would be influenced by whether the confederates wore perfume. He found
that wearing perfume led to an increase in attractiveness of the female confederate
to male participants (but only if she was dressed informally rather than more
formally). Given this finding (see also Kirk-Smith & Booth, 1987; Marinova & Moss,
2014), a positive effect of androstenol on attractiveness ratings may be attributable
simply to the odour being perceived as pleasant.

The olfactory system in humans is predominantly ipsilateral, fibres from each nostril
projecting to the olfactory bulb and cortex on the same side (Gordon and Sperry,
1969, Smith and Bhatnagar, 2019). Presentation of an odour to only one nostril
therefore initially stimulates the ipsilateral cerebral hemisphere. In what appears to
have been one of the earliest systematic studies of olfactory asymmetry with
neurologically intact participants, Toulouse and Vaschide (1900) presented solutions
of camphor to blindfolded subjects who had each nostril blocked in turn. They
reported that in 56 of 64 male and female participants (including four young
children) the asymmetry favoured the left nostril for both detection and recognition
of camphor. Koelega (1979) using n-amyl acetate in a discrimination task could not
replicate this nostril effect, finding no difference between left and right nostrils.

Toulouse and Vaschide (1899) had found detection thresholds (for camphor) to be
lower on the left side for right-handers and lower on the right side for left-handers.
In re-examining the question of lateral sensitivity in olfaction, similar findings were
obtained by Frye, Doty, & Shaman (1992). However, Youngentob, Kurtz, Leopold,
Mozell, and Hornung (1982) reported the opposite pattern; the left nostril of left-
handers and the right nostril of right-handers showed greater sensitivity in
detecting n-butanol.

In a number of other studies the effects of handedness have been found to interact
with side of nostril in various aspects of odour perception (Bensafi et al., 2003,
Betchen and Doty, 1998, Cain and Gent, 1991, Dijksterhuis et al., 2002, Doty and Kerr,
2005, Gilbert et al., 1989, Hummel et al., 1998, Koelega, 1979, Royet et al., 2003,
Zatorre and Jones-Gotman, 1990) or with side of hemispheric activation (Royet, et
al., 2003), not always consistently. Lübke, Gottschlich, Gerber, Pause, and Hummel
(2012) refer to the confusing literature on handedness-related differences in
olfaction, pointing out that the reported effects seem to vary with task demands
(see also Royet et al., 2001). Conceivably, they also vary with the nature of the
olfactory stimulus. However, the fact that such effects occur at all reinforces the
view that there exists some hemispheric differentiation of olfactory functions.

Evidence from patients with lateralised brain damage or dysfunction, supported by

functional imaging and experimental studies with neurologically intact participants,
reveals that there is little or no difference between the two sides of the brain in
simple detection of odours (Bensafi et al., 2003, Carroll et al., 1993, Jones-Gotman
and Zatorre, 1988, Royet and Plailly, 2004). However, the right cerebral hemisphere
may be dominant for certain other aspects of odour processing, such as
discrimination and recognition of odours (Abraham and Mathai, 1983, Jones-
Gotman and Zatorre, 1993, Melero et al., 2019, Zald and Pardo, 2000) and evaluation
of intensity, familiarity and edibility of associated substances (Royet et al., 2001).

In addition, some evidence indicates that the right hemisphere is preferentially

involved in processing pleasant odours (Bensafi et al., 2003, Dijksterhuis et al., 2002,
Herz et al., 1999, Katata et al., 2009, Kobal et al., 1992) while other evidence
implicates the left hemisphere (Henkin & Levy, 2001). Royet et al. (2003) suggested
that judgement of hedonic valence involves the left hemisphere more than the right
regardless of whether an odour is perceived as pleasant or unpleasant. (For review
and critique of PET and fMRI studies of olfactory hedonicity see Mantel, Ferdenzi,
Roy & Bensafi, 2019).

Despite increasing recognition that components of body odour are important in

human interaction and mate choice (Groyecka et al., 2017, Hofer et al., 2020,
Mahmut and Croy, 2019, Thornhill and Gangestad, 1999), and despite a long history
of research into laterality effects in olfaction (for reviews see Brancucci, Lucci,
Mazzatenta & Tommasi, 2009; Royet & Plailly, 2004), to date no study appears to
have looked at whether there is a laterality effect in perception of androstenes or in
their influence on judgements of physical attraction or modulation of mood. This is
somewhat surprising (see Brancucci et al., 2009) given that neuroimaging studies
have demonstrated lateralised activation patterns in response to stimulation by
certain androstene compounds (Savic et al., 2001, Savic and Lindström, 2008, Savic
and Berglund, 2010) and that laterality effects in socio-affective domains are well
documented in humans for the visual and auditory modalities (for reviews see
Gainotti, 2019; Palomero-Gallagher & Amunts, 2021). Furthermore, olfactory
laterality differences have been reported in relation to approach-avoidance
(emotional?) behaviour in some animal species (for reviews see Gainotti, 2021;
Leliveld, Langbein, & Puppe, 2013; Rutherford & Lindell, 2011; Siniscalchi, d’Ingeo, &
Quaranta, 2021).

Although in recent years research interest has been directed towards the possible
influence of androstadienone rather than androstenol on judgements of
attractiveness (see e.g. Ferdenzi, Delplanque, Atanassova & Sander, 2016; Hare,
Schlatter, Rhodes & Simmons, 2017; Parma, Tirindelli, Bisazza, Massaccesi &
Castiello, 2012; Saxton, Lyndon, Little & Roberts, 2008), we believe it is not yet time
to draw a veil over the potential effects of androstenol. In particular, the issue of
whether any effects may be due to pleasantness of odour has not been addressed.

We therefore decided to re-examine the influence of androstenol on ratings of

attractiveness and mood. We also sought to determine whether perceived
attractiveness is influenced by how pleasant the compound is deemed to be. Finally,
we investigated whether there are any laterality differences in the effect of
androstenol. On the basis of previous research, our main hypothesis was that
androstenol would enhance ratings of attractiveness of photographs of the opposite
sex. We further predicted that sniffing androstenol through the right nostril would
lead to stronger effects than sniffing through the left nostril.

Section snippets

Method

The experiment was run in accordance with the principles of the Helsinki
Declaration (2013) and approval for the study was given by the appropriate
University Departmental Ethics Committee. All participants provided written
informed consent.…

Descriptive statistics

Mean (and SD) ratings of attractiveness, strength and pleasantness of odour, and for
each of the mood scales are shown in Table 1.…

Ratings of attractiveness

Each participant rated a different pair of photographs in each of the four conditions.
As each photograph was given a very similar mean rating by the panel of judges in
the pilot study, ratings for the two photographs in each condition were combined
and the mean rating taken as the response measure for that participant and
condition. Group mean ratings of…

Discussion

The present study compared the effects of sniffing a solution containing androstenol
with a control solution (ethanol) through the left or right nostril. We predicted that
androstenol would increase ratings of attractiveness of photographs of the opposite
sex and that any effects of androstenol on mood would be greater for the right
nostril than the left. The findings can be summarised as follows.

Androstenol was rated as having a stronger and more pleasant smell than the
control solution.…

Conclusion

The results of our study show an effect of androstenol on ratings of attractiveness of

opposite-sex photographs. This could not be attributed to the smell of androstenol
being perceived as pleasant. We also found that androstenol influenced self-ratings
of liveliness, sexiness, aggression, and irritability. A novel aspect of our results is
that there were significant differences between right and left nostrils, implying
hemispheric differences in psychological effects after exposure to…

Funding

This research did not receive any specific grant from funding agencies in the public,
commercial, or not-for-profit sectors.…

Declaration of Competing Interests

The authors declare no competing interests.…

Acknowledgements
We acknowledge with thanks the helpful comments of two anonymous referees.…

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