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Abstract

Fifty years after the term "pheromone" was coined by Peter Karlson and Martin Lüsher the search for these semiochemicals is still an elusive goal of chemical ecology and communication studies. Contrary to what appears in the popular press, the race is still on to capture and define human scents. Over the last several years, it became increasingly clear that pheromone-like chemical signals probably play a role in offspring identification and mother recognition. Recently, we analyzed the volatile compounds in sweat patch samples collected from the para-axillary and nipple-areola regions of women during pregnancy and after childbirth. We hypothesized that, at the time of birth and during the first weeks of life, the distinctive olfactory pattern of the para-axillary area is probably useful to newborns for recognizing and distinguishing their own mother, whereas the characteristic pattern of the nipple-areola region is probably useful as a guide to nourishment.
Chemical basis for mother recognition
www.landesbioscience.com Communicative & Integrative Biology 279
[Communicative & Integrative Biology 2:3, 279-281; May/June 2009]; ©2009 Landes Bioscience
Fifty years after the term “pheromone” was coined by Peter
Karlson and Martin Lüsher the search for these semiochemicals
is still an elusive goal of chemical ecology and communication
studies. Contrary to what appears in the popular press, the race is
still on to capture and define human scents. Over the last several
years, it became increasingly clear that pheromone-like chemical
signals probably play a role in offspring identification and mother
recognition. Recently, we analyzed the volatile compounds in sweat
patch samples collected from the para-axillary and nipple-areola
regions of women during pregnancy and after childbirth. We
hypothesized that, at the time of birth and during the first weeks
of life, the distinctive olfactory pattern of the para-axillary area is
probably useful to newborns for recognizing and distinguishing
their own mother, whereas the characteristic pattern of the nipple-
areola region is probably useful as a guide to nourishment.
Pheromones or Signature Odors?
Throughout the lives of most mammalian species, the sense
of smell plays an important role in response to chemical messen-
gers involved in different behaviors. Within the overall olfactory
communication pheromones are very important actors. The term
‘pheromone’—referred to the chemical compounds used to commu-
nicate between individuals of the same species—was coined by Peter
Karlson and Martin Lüsher in 1959.1 Over the last 50 years, phero-
mones, sending messages between individuals, have been found in
many species across the animal kingdom.
The so-called “individual odors”, learned for recognition, do
not seem to fit Karlson and Lüshers definition. In the past, some
researchers even doubted that humans could have their behavior
altered by something as simple as an instinctive reaction to smell.
Now, after years of debate, it seems clear that these variable odors
are not pheromones and instead are better referred to as ‘signature
odors’.
But species-specific small molecules, that fit the classic phero-
mone definition, have now been identified for mammals. It appears
clear that signature odors and pheromones can be mixed for effect.
Some mammals, including elephants and mice, present their
small-molecule pheromones in the cleft of highly variable lipocalin
proteins. Pheromone signals can also be overlaid and improved with
individual signature odors, the proteins release the small molecules
slowly, making them last longer.
The consensus now is that the human use of pheromones, as in
all mammals, appears likely. The armpits are prime source candi-
dates, as their odor emanation develops along with other changes
at puberty. However, both human behavior and chemical emissions
are so complex that at present the research is challenging. Up to
now, no pheromones have been conclusively identified, but a strong
contender for the first identified human pheromone is some elusive
compound in the armpit of women. Apparently this unidentified
pheromone causes menstrual synchrony in females living in close
quarters. It has been speculated that its identification could poten-
tially open the door to sniffable contraceptives.2
Chemical Basis for Olfactory Communication
In animal species, recognition between individuals is an essential
requirement for any kind of further interaction. Recognition between
mother and newborn is a fundamental behavioral interaction that is
worth systematic investigation. The emotional relationship between
a mother and her newborn begins with mutual recognition, which
starts during gestation and continues through birth, augmented by
body contact and lactation. Imprinting takes place through visual,
auditory and olfactory learning, which occurs very early during the
so-called “critical period”. Consequently, from beginning of preg-
nancy olfaction seems to represent an Ariadne’s thread that permits
the infant after birth to find its mother.3
Pheromones regulate reproductive behavior in many animal
species. Once released in the environment, through urine or glan-
dular secretions, these volatile substances reach other individuals
of the same species, signaling, for instance, mating availability and
strengthening ties between mother and offspring, as well as regulating
social relationships.4 In non-human vertebrate species, pheromones
are detected by a specific sensory apparatus, the vomeronasal system,
composed of a peripheral organ located at the base of the nasal
septum, the vomeronasal nerve and a nerve center, the accessory
olfactory bulb. The vomeronasal system is completely separated and
independent from the main olfactory system.5 It is triggered by a
different class of volatile substances and is present in many reptiles
and in almost all mammals, but absent in fishes and birds, even if
they possess a main olfactory system.
Mini-Review
Chemical communication and mother-infant recognition
Stefano Vaglio
Laboratory of Anthropology; Department of Evolutionar y Biology “Leo Pardi”; University of Florence; Florence, Italy
Key words: offspring identification, mother recognition, putative human pheromones, volatile compounds, gas chromatography-mass
spectrometry (GC-MS), solid phase micro-extraction (SPME)
Correspondence to: Stefano Vaglio; Laboratory of Anthropology; Department of
Evolutionary Biology “Leo Pardi”; University of Florence; Via del Proconsolo 12;
Florence 50122 Italy; Email: stefanovaglio@yahoo.it
Submitted: 02/13/09; Accepted: 02/17/09
Previously published online as a Communicative & Integrative Biology E-publication:
http://www.landesbioscience.com/journals/cib/article/8227
Chemical basis for mother recognition
280 Communicative & Integrative Biology 2009; Vol. 2 Issue 3
Primates, long considered functionally non-microsomatic, were
previously thought to show complex olfactory communication only
in prosimians and in some New World monkeys. Now it is well
established that even higher primates use pheromones to recognize
con-specific individuals and for territorial marking.6-8 Furthermore,
protein-pheromone complexes—present in the secretions released by
scent-marking of some non-human primates—were recently shown
to activate vomeronasal receptors particularly for sexually related
behaviors and intra-specific identification of individuals.9-12
Formerly, it was widely held that the human vomeronasal organ
was vestigial and even the existence of pheromonal communication
in humans was contested. However, support for a role of pheromones
in human behavior came from several observational studies (i.e.,
synchronization of the menstrual cycle13), but this role was often
denied because in the absence of a sufficient neuro-anatomic basis
for such a complex behavior.
Recently, a new class of olfactory receptors (trace amino-associ-
ated receptors, TAAR) was discovered in the olfactory epithelium
of mice.14 Genes similar to those responsible for the control of
these receptors in mice have been identified in humans and fishes.
These data suggest a great evolutionary conservation of these
genes and lend support to the hypothesis that the putative human
pheromonal response is mediated by receptors located in the main
olfactory system.
The putative human pheromones are steroids present in the
secretions of exocrine glands.15-20 Estrogen derivatives are present in
females (the so-called “copulins”—mixtures of aliphatic acids such as
acetic, propionic, butyric, isovaleric and isocaproic acid with estratet-
raenol), and androgen derivatives are present in males (androstenol,
androstenone and androstadienone). Recent studies concerning the
most volatile compounds of human sweat21-23 have shown that the
characteristic odor produced by the para-axillary region is due to
the presence of volatile C6-C11 acids, the most abundant being
E-3-methyl-2-hexenoic acid (E-3M2H).
Mother-Child Identification
Human chemical signals may also play a role in offspring iden-
tification. Odor cues from newborns are absolutely salient to their
mothers.24 Mothers are able to distinguish the odor of their own
newborn baby from that of other newborns.25 Experiments also have
demonstrated that adults can even recognize gender and individuality
of non-related children.26 Thus, body odors can provide humans
with important information about the individual identity of their
offspring.27-32
On the other hand, children usually prefer parts of clothes that
were in contact with the axilla and worn by their own mothers to
clothes worn by other mothers.25 Therefore, chemical signals seem
to have a fundamental role in the mechanism of mother-child iden-
tification.30,33 Breast-feed versus bottle-feed infants show different
reactions to maternal odors. Breast-feed infants are exposed to salient
maternal odors and rapidly become familiarized with their mothers’
unique olfactory signature.34 Apparently, orientation to lactating-
breast odors is an inborn adaptive response of newborns.35
It seems an inescapable conclusion that naturally occurring odors
play an important role in mediating infant behavior. Even fetal olfac-
tory learning seems to occur and breast odors from the mother exert
a pheromone-like effect at the newborn’s first attempt to locate the
nipple. Newborns are generally responsive to breast odors produced
by lactating women.33 Olfactory recognition may be implicated in
the early stages of the mother-infant attachment process, when the
newborns learn to recognize their own mother’s unique odor signa-
ture: this process is possibly made easier by the high norepinephrine
release and the arousal of the locus coeruleus at birth.36 Human infants
are responsive to maternal odors beginning shortly after birth. They
show an attraction to amniotic fluid odor that may reflect fetal expo-
sure to that substance (i.e., prenatal olfactory learning37). Moreover,
human amniotic fluid seems to carry individualized odor properties,
which are hypothesized to initiate parent-infant interactions.38
Not only Chemical Signals: Some Socio-Biological Remarks
It is clear that, as any other organism, humans are subject to
invisible but potentially irresistible influences of metabolic mate-
rials on our muscle, motive and motor actions, both directly and
indirectly.39,40
Obviously, mutual recognition among organisms is related to many
aspects of the personal profile, both of metabolic-material biological
similarity and morphologic-motor ecological familiarity.41
Mothers recognize the sounds of their own babies’ chortles and
cries, and may differentiate among their causes, from tiredness to
hunger, from pain to scare. Yet mothers may pause for a moment to
listen harder to cries from an unseen child that are similar enough to
their own babies’ cries to require more attention before discounting
and dismissing them as indeed being from someone else baby.42
Yet it is appreciated that the cry from any newborn can cause
agitation and distress in any mother. Perhaps this reflects the
process of acquiring maternal memories: that is, at the time a
mother has a newborn, she does not yet know her babies cry
well enough to discriminate with certainty. Moreover, mothers of
newborns are in states of hormonally heightened metabolic arousal,
and are thus readily responsive to both biologically and ecologically
conditioned cues.43
Immediacy of recognition of a simple similar-enough sound in
a noisy place, such as a familiar name, is well enough known (the
“cocktail party syndrome”), even if that name was not in fact said,
and so what was heard was only ‘similar enough’ to seem ‘familiar
enough’.
The connection among all aspects of recognition is surely not
sets of discrete ‘signals’ but, instead, a composition of several sorts
of similarities and familiarities, including biochemical features.
Probably these features serve to substantiate and sustain mutual
recognition.44
Volatile Compounds Behind Mother Recognition
In our recent study3 we hypothesized that women probably
develop a volatile profile, through pregnancy and childbirth that
enables identification of the mother by the newborn. The aim was to
understand, through an analytical approach, how the volatile pattern
of pregnant women changes during pregnancy and, consequently,
to verify the effective role played by volatile chemical signals in the
mechanism of mother-infant recognition.
We identified the volatiles compounds in sweat patch samples
collected from the para-axillary and nipple-areola regions of women
during pregnancy and after childbirth. Results showed that during
pregnancy women developed a distinctive pattern of five volatile
Chemical basis for mother recognition
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compounds common to the para-axillary and nipple-areola regions
(1-dodecanol, 1-1'-oxybis octane, isocurcumenol, α-hexyl-cinnamic
aldehyde and isopropyl myristate).
Hypothetically, the differentiation of the olfactory pattern among
pregnant women helps newborns to recognize their own mother
and distinguish her from other individuals. At the time of birth
and during the first weeks of life, the distinctive olfactory pattern of
the para-axillary area might be useful to newborns to recognize and
distinguish their mother, whereas the characteristic pattern of the
nipple-areola region is probably useful as a guide to nourishment.
Future Challenges
These recent results show the effectiveness of the methodology
used. Through the collection and analysis of the secretion released at
the level of the para-axillary area and in the nipple areola region, it is
possible to investigate the volatile compounds in a proper way. Up to
now these phenomena have often been investigated with inadequate
methodologies and, as a consequence, the role of volatile compounds
as regulators of mother-infant recognition has been underestimated.
Moreover, recently developed technical instruments and procedures—
as Solid Phase Micro-Extraction (SPME), Dynamic Head-Space
Extraction (DHE), and Maldi TOF/TOF—in addition to the classic
ones—as Gas Chromatography-Mass Spectrometry (GC-MS) and
Liquid Chromatography-Mass Spectrometry (LC-MS)—now allow
investigators to characterize volatile and non-volatile compounds
with high reliability.45
The study of the mechanism of mother-infant recognition is
important not only for the acquisition of new knowledge concerning
the emission of signal molecules essential for mother-child identifica-
tion, but also for its clear practical consequences. This information
can be helpful for setting up the proper conditions to establish solid
mother-child bonding. It can indicate the behavior and conduct to
maintain during gestation and the initiation period of life of the
newborn. Therefore an understanding of the mechanisms of newborns
recognition of their mother could have practical health implications.
Acknowledgements
I am deeply in debt to Professor Roscoe Stanyon for his editorial
work on the article. Moreover, I would like to thank Leaf Lovejoy
for useful suggestions. I also thank Professor Brunetto Chiarelli and
Professor Giorgio Mello for encouragement.
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www.landesbioscience.com Communicative & Integrative Biology 281
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... The neonate at milestone 4 cannot put into words the proposition that "this voice is the one most familiar to me and it is coming from the face of that bodily person", but conceptually the baby realizes that the familiar voice -a very special one from prior experience -is being produced by that person whom the baby knew before it was born, but now meets up with in the present and now visible external world (Sai, 2005). If mom nurses the baby, the research is clear that mom's smell and taste, like the sound of her voice along with the familiar rhythms of her movements, are also known to the infant from before birth (Vaglio, 2009;Marin et al., 2015;Crenshaw, 2019;Schaal et al., 2020). Moreover, given the uniqueness of the inborn, innate, inherited human capacity for language, perhaps it should not surprise us to discover that the unborn human child has a natural interest in figuring out the peculiar rhythms and sounds of its native language from well before it is born (Spitzer, 2001;Partanen et al., 2013). ...
Book
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The human language capacity stands at the very top of the intellectual abilities of us human beings, and it ranks incommensurably higher than the intellectual powers of any other organism or any robot. It vastly exceeds the touted capacities of "artificial intelligence" with respect to creativity, freedom of will (control of thoughts and words), and moral responsibility. These are traits that robots cannot possess and that can only be understood by human beings. They are no part of the worlds of robots and artificial intelligences, but those entities, and all imaginable fictions, etc., are part of our real world... True narrative representations (TNRs) can express and can faithfully interpret every kind of meaning or form in fictions, errors, lies, or nonsensical strings seeming in any way to be representations. None of the latter, however, can represent even the simplest TNR ever created by an intelligent person. It has been proved logically, in the strictest forms of mathematical logic, that all TNRs that seem to have been produced by mechanisms, robots, or artificial intelligence, must be contained within a larger and much more far-reaching TNR that cannot be explained mechanistically by any stretch of imagination. These unique constructions of real intelligence, that is, genuine TNRs, (1) have the power to determine actual facts; (2) are connected to each other in non-contradictory ways, and (3) are generalizable to all contexts of experience to the extent of the similarities of those contexts up to a limit of complete identity. What the logicomathematical theory of TNRs has proved to a fare-thee-well is that only TNRs have the three logical properties just iterated. No fictions, errors, lies, or any string of nonsense has any of those unique formal perfections. The book is about how the human language capacity is developed over time by human beings beginning with TNRs known to us implicitly and actually even before we are born. All scientific endeavors, all the creations of the sciences, arts, and humanities, all the religions of the world, and all the discoveries of experience utterly depend on the prior existence of the human language capacity and our power to comprehend and produce TNRs. Without it we could not enjoy any of the fruits of human experience. Nor could we appreciate how things go wrong when less perfect representations are mistaken, whether accidentally or on purpose, for TNRs. In biology, when DNA, RNA, and protein languages are corrupted, the proximate outcome is disorder, followed by disease if not corrected, and, in the catastrophic systems failures known as death in the long run. The book is about life and death. Both are dependent on TNRs in what comes out to be an absolute dependency from the logicomathematical perspective. Corrupt the TNRs on which life depends, and death will follow. Retain and respect TNRs and life can be preserved. However, ultimate truth does not reside in material entities or the facts represented by TNRs. It resides exclusively in the TNRs themselves and they do not originate from material entities. They are from God Almighty and do not depend at all on any material thing or body. TNRs outrank the material facts they incorporate and represent. It may seem strange, but the result is more certain, I believe, than the most recent findings of quantum physics. Representations are connected instantaneously. Symbol speed is infinitely faster than the speed of light. In the larger perspective of history, when TNRs are deliberately corrupted, the chaos of wars, pestilence, and destruction follows as surely as night follows day. The human language capacity makes us responsible in a unique manner for our thoughts, words, and actions. While it is true that no one ever asked us if we wanted to have free will or not, the fact that we have it can be disputed only by individuals who engage in a form of self-deception that borders on pathological lying, the kind that results when the deceiver can no longer distinguish between the actions he or she actually performed in his or her past experience and the sequences of events that he or she invented to avoid taking responsibility for those events, or to take credit for actions he or she never performed. On the global scale such misrepresentations lead to the sort of destruction witnessed at Sodom in the day of Abraham. That historical destruction has recently been scientifically revealed at the site of Tall el-Hammam in Jordan. More about that and all of the foregoing in the book. If you encounter errors, please point them out to the author at joller@bellsouth.net. Thank you.
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Chemosignals are mediators of social interactions in mammals, providing con‐ and hetero‐specifics with information on fixed (e.g., species, sex, group, and individual identity) and variable (e.g., social, reproductive, and health status) features of the signaler. Yet, methodological difficulties of recording and quantifying odor signals, especially in field conditions, have hampered studies of natural systems. We present the first use of the Torion® portable gas chromatography‐mass spectrometry (GC‐MS) instrument for in situ chemical analysis of primate scents. We collected and analyzed swab samples from the scent glands and skin from 13 groups (57 individuals) of two sympatric species of wild emperor tamarins, Saguinus imperator, and Weddell's saddleback tamarins, Leontocebus weddelli (Callitrichidae). In total, 11 compounds of interest (i.e., probably derived from the animals) could be detected in the samples, with 31 of 215 samples containing at least one compound of interest. The composition of these 31 samples varied systematically with species, group, sex, and breeding status. Moreover, we tentatively identified seven of the compounds of interest as methyl hexanoate, benzaldehyde, ethyl hexanoate, acetophenone, a branched C15 alkane, 4‐methoxybenzaldehyde, and hexadecan‐1‐ol. As the field of primate semiochemistry continues to grow, we believe that portable GC‐MS instruments have the potential to help make progress in the study of primate chemosignaling in field conditions, despite limitations that we encountered. We further provide recommendations for future use of the Torion® portable GC‐MS for in situ analyses. Research Highlights • We report the first use of the Torion® portable GC‐MS for in situ chemical analysis of animal scents. • The chemical composition of scent‐gland swabs from wild callitrichids differs among species, group, sex, breeding status, and between scent glands.
... These advances have particularly been supported in humans for evolutionary and medical purposes (e.g. Havlicek and Roberts, 2009;Penn and Potts, 1998;Roberts et al., 2011;Vaglio, 2009;Winternitz and Abbate, 2015 Jolly, 1966;and callitrichids, Heymann, 1998; Lazaro-Perea, Snowdon and Arruda, 1999), only after the turn of the century did it become a quite strong focus of research (Heymann, 2006b). ...
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Olfactory communication is an important mediator of social interactions in mammals, providing information about an individual’s identity and current social, reproductive, and health status. Callitrichids (i.e. marmosets and tamarins) constitute a good model for the study of olfactory communication, as they make use of a range of odour signals. Callitrichids conspicuously deposit odorous secretions, produced by specialized scent-glands, on branches in their environment, a behaviour called scent-marking. Several functions have been attributed to callitrichid scent-marking behaviour, including advertisement of reproductive and dominance status, and of identity, territorial defence, and spatial orientation and signalling of food resource location. The present doctoral project combined behavioural and chemical information to investigate callitrichid olfactory communication. The study explored how environmental, social, and reproductive aspects might influence patterns of callitrichid scent-marking behaviour, as well as the chemical composition of scent-gland secretions and urine used to convey chemosignals. Behavioural observations, along with swabs of scent-glands, and of naturally deposited scent-marks and urine, were collected from captive groups of bearded emperor tamarins, Saguinus imperator subgrisescens, cotton-top tamarins, Saguinus oedipus, and silvery marmosets, Mico argentatus, in three British zoos. Chemical samples were analysed using headspace gas chromatography-mass spectrometry (GC-MS). In addition, scent-gland secretion samples were collected from a wild population of sympatric emperor tamarins and Weddell’s saddleback tamarins, Leontocebus weddelli, during an annual capture-and-release programme in the south-eastern Peruvian Amazon. These samples were analysed using both in situ and laboratory-based GC-MS techniques. I established the existence of unique chemical signatures of species, groups, sex, reproductive status, and the individual, in callitrichid scent samples, which were matched with differences in scent-marking behaviour. My results support the assumption that chemosignalling plays an important role in the advertisement of identity, reproductive state/status and dominance in this taxon. Moreover, I showed that the social context, as well as spatiotemporal aspects of scent-marking deposition, influenced scent-marking activity. Further differences in the characteristics of scent-marking deposition revealed in this study, both at behavioural and chemical levels, may reflect variable strategies of communication to ensure that signals are transmitted to the intended receivers, which is especially relevant for sympatric species. I identified a number of putative semiochemicals (i.e. chemicals involved in communication) from the scent samples of captive and wild callitrichids. Notably, I presented results from the first use of the Torion® portable GC-MS for in situ analysis of wild mammal scent samples. In addition, I revealed differences in the chemical composition of tamarin scent-gland secretion samples between wild and captive conditions, which may indicate an effect of captivity on the chemicals produced. This study provides knowledge of mammalian olfactory communication systems, applicable to captive husbandry practices, including conservation breeding programmes of rare species.
... The occurrence of such an impulse in human beings has been discussed very controversially (e.g., Maslow, 1943Maslow, , 1954Birney and Teevan, 1961), but observations that human infants (and in part older individuals) respond fearfully to (pictures of) snakes (Headland and Greene, 2011;Hoehl et al., 2017;Denzer, 2018) represent at present the best indication of such operating instincts in humans. However, the situation is quite different in the social bonding of an infant to its mother (or, more generallyspeaking to its care-giver), because an infant can differentiate other individuals by means of olfactory communication (e.g., Vaglio, 2009), and such an ability requires learning processes. Thus, the mother-infant tie is classified here as a drive, 31 albeit not a Freudian one because this attachment drive operates 29 Wright and Panksepp noted: "The ability to process and 'decide' between the drives might be lost if each drive is not also an independent generator. ...
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... Or des recherches comme celles de (Li, 2014) ; nous utilisons les odeurs dans nos choix alimentaires-l' odeur nous donne une indication de l'état de préservation d'une denrée, mais aussi si on l' aime ou non ; l'odorat semble avoir un rôle crucial dans le comportement parental (Okamoto et al., 2016) dans la reconnaissance par les nouveau-nés des odeurs maternelles et de leur environnement familier (Vaglio, 2009), dans les interactions des jeunes enfants avec leur environnement (Cameron, 2018). Le rôle de l'odorat a également été démontré comme participant au choix de partenaires sexuels (Herz and Inzlicht, 2002 (Engen, 1991, p. 7). ...
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Modern humans are currently experiencing a transformation of their environment and of their way of life that are impacting the way they can experience nature in their daily life. These experiences of nature are of great importance for the well-being and health of individuals. They are complex phenomena, anchored in an environmental context, but also socio-cultural, which are based on closely intertwined components that are emotions, memories, knowledge, but also all the sensory stimuli perceived by the human body. Among the senses mobilized, the sense of smell, by its memory and emotional importance, seems to play a singular part in the relationship that weaves the human to nature. The work conducted in this thesis, articulating around three axes of research and reflection, concerns this role of the olfaction in the experience of nature. L’humain moderne vit une transformation de son environnement et de son mode de vie qui impactent la façon dont il peut vivre des expériences de nature au quotidien. Or, ces expériences de nature sont d'une grande importance pour le bien-être et la santé des individus. Ce sont des phénomènes complexes, ancrés dans un contexte environnemental, mais aussi socio-culturel, qui reposent sur des composantes étroitement mêlées que sont les émotions, les souvenirs, les connaissances, mais aussi tous les stimuli sensoriels perçus. Parmi les sens mobilisés, l’odorat, de par son importance mémorielle et émotionnelle, semble jouer un rôle à part dans la relation que tisse l’humain à la nature. C’est à ce rôle de l’olfaction dans l’expérience de nature, à cette part olfactive de l’expérience, que s’intéresse cette thèse s’articulant autour de trois axes de recherches et de réflexion. Dans un premier chapitre, ce manuscrit aborde comment caractériser la part olfactive de l’expérience de nature d’un point de vue individuel. La réflexion de ce chapitre se base sur la mise en œuvre et l’analyse des réponses à un questionnaire articulant entre eux l’olfaction, l’identité et les usages sensoriels d’espaces de nature déclarés par les individus. Dans le deuxième chapitre, c’est la façon dont l’expérience olfactive de nature s’ancre dans un environnement et un contexte qui est abordée, et comment cet ancrage influence la façon dont l’individu vit et décrit son expérience olfactive. La réflexion de ce chapitre s’appuie sur des témoignages recueillis lors de parcours olfactifs commentés et des questionnaires in situ. Enfin, en se basant sur les résultats des études précédentes, le troisième et dernier chapitre s’intéresse à la façon dont l’expérience olfactive de nature peut avoir un rôle transformateur sur l’individu dans le cadre particulier des environnements restaurateurs. En guise de conclusion, ce manuscrit aborde des réflexions, des ouvertures théoriques et pratiques, et des applications que peuvent apporter les résultats du travail de thèse, notamment la place que le sensoriel, l’incarné et l’olfactif pourraient prendre à l’avenir dans la façon de penser et d’enrichir l’expérience de nature.
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