Biology:Body odor

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Short description: Odor produced by a living animal


Body odor or body odour (BO) is present in all animals and its intensity can be influenced by many factors (behavioral patterns, survival strategies). Body odor has a strong genetic basis, but can also be strongly influenced by various factors, such as sex, diet, health, and medication.[1] The body odor of human males plays an important role in human sexual attraction, as a powerful indicator of MHC/HLA heterozygosity.[2][1] Significant evidence suggests that women are attracted to men whose body odor is different from theirs, indicating that they have immune genes that are different from their own, which may produce healthier offspring.[3]

Causes

In humans, the formation of body odors is caused by factors such as diet, sex, health, and medication, but the major contribution comes from bacterial activity on skin gland secretions.[1] Humans have three types of sweat glands: eccrine sweat glands, apocrine sweat glands and sebaceous glands. Eccrine sweat glands are present from birth, while the latter two become activated during puberty. Among the different types of human skin glands, body odor is primarily the result of the apocrine sweat glands, which secrete the majority of chemical compounds that the skin flora metabolize into odorant substances.[1] This happens mostly in the axillary (armpit) region, although the gland can also be found in the areola, anogenital region, and around the navel.[4] In humans, the armpit regions seem more important than the genital region for body odor, which may be related to human bipedalism. The genital and armpit regions also contain springy hairs which help diffuse body odors.[5]

The main components of human axillary odor are unsaturated or hydroxylated branched fatty acids with E-3-methylhex-2-enoic acid (E-3M2H) and 3-hydroxy-3-methylhexanoic acid (HMHA), sulfanylalkanols and particularly 3-methyl-3-sulfanylhexan-1-ol (3M3SH), and the odoriferous steroids androstenone (5α-androst-16-en-3-one) and androstenol (5α-androst-16-en-3α-ol).[6] E-3M2H is bound and carried by two apocrine secretion odor-binding proteins, ASOB1 and ASOB2, to the skin surface.[7]

Body odor is influenced by the actions of the skin flora, including members of Corynebacterium, which manufacture enzymes called lipases that break down the lipids in sweat to create smaller molecules like butyric acid. Greater bacteria populations of Corynebacterium jeikeium are found more in the armpits of men, whereas greater population numbers of Staphylococcus haemolyticus are found in the armpits of women. This causes male armpits to give off a rancid/cheese-like smell, whereas female armpits give off a more fruity/onion-like smell.[8] Staphylococcus hominis is also known for producing thioalcohol compounds that contribute to odors.[9] These smaller molecules smell, and give body odor its characteristic aroma.[10] Propionic acid (propanoic acid) is present in many sweat samples. This acid is a breakdown product of some amino acids by propionibacteria, which thrive in the ducts of adolescent and adult sebaceous glands. Because propionic acid is chemically similar to acetic acid, with similar characteristics including odor, body odors may be identified as having a pungent, cheesy and vinegar-like smell although certain people might find it pleasant at lower concentrations.[11] Isovaleric acid (3-methyl butanoic acid) is the other source of body odor as a result of actions of the bacteria Staphylococcus epidermidis,[12] which is also present in several types of strong cheese.

Factors such as food, drink, gut microbiome,[13] and genetics can affect body odor.[5]

Function

Animals

In many animals, body odor plays an important survival function. Strong body odor can be a warning signal for predators to stay away (such as porcupine stink), or it can also be a signal that the prey animal is unpalatable.[14] For example, some animals species, who feign death to survive (like opossums), in this state produce a strong body odor to deceive a predator that the prey animal has been dead for a long time and is already in the advanced stage of decomposing. Some animals with strong body odor are rarely attacked by most predators, although they can still be killed and eaten by birds of prey, which are tolerant of carrion odors.[citation needed]

Body odor is an important feature of animal physiology. It plays a different role in different animal species. For example, in some predator species that hunt by stalking (such as big and small cats), the absence of body odor is important, and they spend plenty of time and energy to keep their body free of odor. For other predators, such as those that hunt by visually locating prey and running for long distances after it (such as dogs and wolves), the absence of body odor is not critical. In most animals, body odor intensifies in moments of stress and danger.[15]

Humans

In humans, body odor serves as a means of chemosensory signal communication between members of the species. These signals are called pheromones and they can be transmitted through a variety of mediums. The most common way that human pheromones are transmitted is through bodily fluids. Human pheromones are contained in sweat, semen, vaginal secretions, breast milk, and urine.[1] The signals carried in these fluids serve a range of functions from reproductive signaling to infant socialization.[16] Each person produces a unique spread of pheromones that can be identified by others.[2] This differentiation allows the formation of sexual attraction and kinship ties to occur.[2][17]

Sebaceous and apocrine glands become active at puberty. This, as well as many apocrine glands being close to the sex organs, points to a role related to mating.[5] Sebaceous glands line the human skin while apocrine glands are located around body hairs.[1] Compared to other primates, humans have extensive axillary hair and have many odor producing sources, in particular many apocrine glands.[18] In humans, the apocrine glands have the ability to secrete pheromones. These steroid compounds are produced within the peroxisomes of the apocrine glands by enzymes such as mevalonate kinases.[19]

Sexual selection

Pheromones are a factor seen in the mating selection and reproduction in humans. In women, the sense of olfaction is strongest around the time of ovulation, significantly stronger than during other phases of the menstrual cycle and also stronger than the sense in males.[20][21] Pheromones can be used to deliver information about the major histocompatibility complex (MHC).[2] The MCH in humans is referred to as the Human Leukocyte Antigen (HLA).[22] Each type has a unique scent profile that can be utilized during the mating selection process. When selecting mates, women tend to be attracted to those that have different HLA-types than their own.[2][22] This is thought to increase the strength of the family unit and increase the chances of survival for potential offspring.[2]

Studies have suggested that people might be using odor cues associated with the immune system to select mates. Using a brain-imaging technique, Swedish researchers have shown that homosexual and heterosexual males' brains respond in different ways to two odors that may be involved in sexual arousal, and that homosexual men respond in the same way as heterosexual women, though it could not be determined whether this was cause or effect. When the study was expanded to include lesbian women, the results were consistent with previous findings – meaning that lesbian women were not as responsive to male-identified odors, while responding to female odors in a similar way as heterosexual males.[23] According to the researchers, this research suggests a possible role for human pheromones in the biological basis of sexual orientation.[24]

Kinship communication

Humans can olfactorily detect blood-related kin.[17] Mothers can identify by body odor their biological children, but not their stepchildren. Preadolescent children can olfactorily detect their full siblings, but not half-siblings or step-siblings, and this might explain incest avoidance and the Westermarck effect.[25] Babies can recognize their mothers by smell while mothers, fathers, and other relatives can identify a baby by smell.[5] This connection between genetically similar family members is due to the habituation of familial pheromones. In the case of babies and mothers, this chemosensory information is primarily contained within breastmilk and the mother's sweat.[26] When compared to that of strangers, babies are observed to have stronger neural connections with their mothers.[27] This strengthened neurological connection allows for the biological development and socialization of the infant by their mother. Using these connections, the mother transmits olfactory signals to the infant which are then perceived and integrated.[27]

In terms of biological functioning, olfactory signaling allows for functional breastfeeding to occur. In cases of effective latching, breastfed infants are able to locate their mother's nipples for feeding using the sensory information enclosed in their mother's body odor.[28] While no specific human breast pheromones have been identified, studies compare the communication to that of the rabbit mammary pheromone 2MB2.[29][30] The perception and integration of these signals is an evolutionary response that allows newborns to locate their source of nutrition. Signaling contains a level of precision that allows babies to differentiate their mother's breasts from that of other women.[26] Once the baby recognizes the familiar olfactory signal, the behavioral response of latching follows.[26] Over time the infant becomes habituated to their mother's breast pheromones which increases latch efficiency.[28]

Beyond a biological function, a mother's body odor plays a role in developing a baby's social capabilities. The ability of an infant to evaluate the properties of human faces stems from the olfactory cues given from their mother.[16] Frequent exposure to the pheromones exuded by their mother allows the connection between vision and smell to form in infants.[27] This type of connection is only found between mothers and babies and over time it socializes the ability to recognize the features that distinguish human faces from inanimate objects.[16]

Environmental threats

The connection between olfactory and visual cues has also been observed outside of familial relationships. Evolutionarily, body odor has been used to communicate messages about potentially dangerous stimuli in the environment.[1] Body odor produced during particularly stressful situations can produce a cascade of reactions in the brain. Once the olfactory system is activated by a threatening stimuli, heightened activity in the amygdala and occipital cortex is triggered.[31][1] This chain reaction serves to help assess the nature of the threat and increase chance of survival.

Humans have few olfactory receptor cells compared to dogs and few functional olfactory receptor genes compared to rats. This is in part due to a reduction of the size of the snout in order to achieve depth perception as well as other changes related to bipedalism. However, it has been argued that humans may have larger brain areas associated with olfactory perception compared to other species.[18]

Genes affecting body odor

World map of the distribution of the A allele of the single nucleotide polymorphism rs17822931 in the ABCC11 gene. The proportion of A alleles in each population is represented by the white area in each circle.

MHC

Body odor is influenced by major histocompatibility complex (MHC) molecules. These are genetically determined and play an important role in immunity of the organism. The vomeronasal organ contains cells sensitive to MHC molecules in a genotype-specific way.[citation needed]

Experiments on animals and volunteers have shown that potential sexual partners tend to be perceived more attractive if their MHC composition is substantially different. Married couples are more different regarding MHC genes than would be expected by chance. This behavior pattern promotes variability of the immune system of individuals in the population, thus making the population more robust against new diseases. Another reason may be to prevent inbreeding.[5]

ABCC11

The ABCC11 gene determines axillary body odor and the type of earwax.[6][32][33][34] The loss of a functional ABCC11 gene is caused by a 538G>A single-nucleotide polymorphism, resulting in a loss of body odor in people who are specifically homozygous for it.[34][35] Firstly, it affects apocrine sweat glands by reducing secretion of odorous molecules and its precursors.[6] The lack of ABCC11 function results in a decrease of the odorant compounds 3M2H, HMHA, and 3M3SH via a strongly reduced secretion of the precursor amino-acid conjugates 3M2H–Gln, HMHA–Gln, and Cys–Gly–(S) 3M3SH; and a decrease of the odoriferous steroids androstenone and androstenol, possibly due to the reduced secretion of dehydroepiandrosterone sulfate (DHEAS) and dehydroepiandrosterone (DHEA), possibly bacterial substrates for odoriferous steroids; research has found no difference, however, in testosterone secretion in apocrine sweat between ABCC11 mutants and non-mutants.[6] Secondly, it is also associated with a strongly reduced/atrophic size of apocrine sweat glands and a decreased protein (such as ASOB2) concentration in axillary sweat.[6]

The non-functional ABCC11 allele is predominant among East Asians (80–95%), but very low among European and African populations (0–3%).[6] Most of the world's population has the gene that codes for the wet-type earwax and average body odor; however, East Asians are more likely to inherit the allele associated with the dry-type earwax and a reduction in body odor.[6][32][34] The reduction in body odor may be due to adaptation to colder climates by their ancient Northeast Asian ancestors.[32]

However, research has observed that this allele is not solely responsible for ethnic differences in scent. A 2016 study analyzed differences across ethnicities in volatile organic compounds (VOCs), across racial groups and found that while they largely did not differ significantly qualitatively, they did differ quantitatively. Of the observed differences, they were found to vary with ethnic origin, but not entirely with ABCC11 genotype.[36]

One large study failed to find any significant differences across ethnicity in residual compounds on the skin, including those located in sweat.[37] If there were observed ethnic variants in skin odor, one would find sources to be much more likely in diet, hygiene, microbiome, and other environmental factors.[38][36][39]

Research has indicated a strong association between people with axillary osmidrosis and the ABCC11-genotypes GG or GA at the SNP site (rs17822931) in comparison to the genotype AA.[34]

Frequencies of ABCC11 allele c.538 (One nonsynonymous SNP 538G > A)[40]
Ethnic groups Tribes or inhabitants AA GA GG
Korean Daegu city inhabitants 100% 0% 0%
Chinese Northern and southern Han Chinese 80.8% 19.2% 0%
Mongolian Khalkha tribe 75.9% 21.7% 2.4%
Japanese Nagasaki people 69% 27.8% 3.2%
Thai Central Thai in Bangkok 63.3% 20.4% 16.3%
Vietnamese People from multiple regions 53.6% 39.2% 7.2%
Native American 30% 40% 30%
Filipino Palawan 22.9% 47.9% 29.2%
Kazakh 20% 36.7 43.3%
Russian 4.5% 40.2% 55.3%
White Americans From CEPH families without the French and Venezuelans 1.2% 19.5% 79.3%
African From various sub-Saharan nations 0% 8.3% 91.7%
African Americans 0% 0% 100%
Amino-acid conjugates of key human body odorants in sweat samples of panelists with different genotypes, determined by liquid chromatography-mass spectrometry[41]
Genotype
ABCC11 		Sex Ethnic population Age Net weight
sweat (g)/2 pads 		HMHA–Gln
(µmol/2 pads) 		3M2H–Gln
(µmol/2 pads) 		Cys–Gly conjugate

of 3M3SH (µmol/2 pads)

AA F Chinese 27 2.05 ND' ND ND
AA F Filipino 33 2.02 ND ND ND
AA F Korean 35 1.11 ND ND ND
GA F Filipino 31 1.47 1.23 0.17 Detectable, < 0.03 µmol
GA F Thai 25 0.90 0.89 0.14 Detectable, < 0.03 µmol
GA F German 25 1.64 0.54 0.10 Detectable, < 0.03 µmol
GG F Filipino 45 1.74 0.77 0.13 Detectable, < 0.03 µmol
GG F German 28 0.71 1.30 0.19 0.041
GG F German 33 1.23 1.12 0.16 0.038

* ND indicates that no detectable peak is found on the [M+H]+ ion trace of the selected analyte at the correct retention time.
* HMHA: 3-hydroxy-3-methyl-hexanoic acid; 3M2H: (E)-3-methyl-2-hexenoic acid; 3M3SH: 3-methyl-3-sulfanylhexan-1-ol.

Alterations

Body odor may be reduced or prevented or even aggravated by using deodorants, antiperspirants, disinfectants, underarm liners, triclosan, special soaps or foams with antiseptic plant extracts such as ribwort and liquorice, chlorophyllin ointments and sprays topically, and chlorophyllin supplements internally. Although body odor is commonly associated with hygiene practices, its presentation can be affected by changes in diet as well as the other factors.[42] Skin spectrophotometry analysis found that males who consumed more fruits and vegetables were significantly associated with more pleasant smelling sweat, which was described as "floral, fruity, sweet and medicinal qualities".[43]

Industry

As many as 90% of Americans and 92% of teenagers use antiperspirants or deodorants.[44][45] In 2014, the global market for deodorants was estimated at US$13.00 billion with a compound annual growth rate of 5.62% between 2015 and 2020.[46]

Medical conditions

Osmidrosis or bromhidrosis is defined by a foul odor due to a water-rich environment that supports bacteria, which is caused by an abnormal increase in perspiration (hyperhidrosis).[33] This can be particularly strong when it happens in the axillary region (underarms). In this case, the condition may be referred to as axillary osmidrosis.[33] The condition can also be known medically as apocrine bromhidrosis, ozochrotia, fetid sweat, body smell, or malodorous sweating.[47][48]

Trimethylaminuria (TMAU), also known as fish odor syndrome or fish malodor syndrome, is a rare metabolic disorder where trimethylamine is released in the person's sweat, urine, and breath, giving off a strong fishy odor or strong body odor.[49]

See also


References

  1. ↑ 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7 Lundström, Johan N.; Olsson, Mats J. (2010). "Functional Neuronal Processing of Human Body Odors". Vitamins & Hormones 83: 1–23. doi:10.1016/S0083-6729(10)83001-8. ISBN 9780123815163. PMID 20831940. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 "Human pheromones and sexual attraction". European Journal of Obstetrics, Gynecology, and Reproductive Biology 118 (2): 135–142. February 2005. doi:10.1016/j.ejogrb.2004.08.010. PMID 15653193. 
  3. ↑ Everts, Sarah (21 July 2021). "What Your Body Odor Says About You" (in en). Time. https://time.com/6082321/your-body-odor-says-about-you/. ""In one study about smell and romance, straight women preferred the body odor of straight men whose immune systems were different enough that any offspring would have healthy immune systems. For most of human history, infectious disease has been our greatest threat. In modern times we may seek life-partners that satisfy a multitude of needs, but more fundamentally, if you could produce babies with immune systems able to fight a potpourri of pathogens, then your progeny—and your genes—stand a better chance at survival."". 
  4. ↑ The encyclopedia of skin and skin disorders (3rd ed.). New York: Facts on File. 2007. pp. 363. ISBN 978-0-8160-6403-8. https://archive.org/details/encyclopediaskin00turk. 
  5. ↑ 5.0 5.1 5.2 5.3 5.4 "Body Odours and Body Odour Preferences in Humans". Oxford Handbook of Evolutionary Psychology. 2007. pp. 315–320. doi:10.1093/oxfordhb/9780198568308.013.0022. ISBN 978-0-19-174365-8. 
  6. ↑ 6.0 6.1 6.2 6.3 6.4 6.5 6.6 "A functional ABCC11 allele is essential in the biochemical formation of human axillary odor". The Journal of Investigative Dermatology 130 (2): 529–540. February 2010. doi:10.1038/jid.2009.254. PMID 19710689. 
  7. ↑ "A human axillary odorant is carried by apolipoprotein D". Proceedings of the National Academy of Sciences of the United States of America 93 (13): 6626–6630. June 1996. doi:10.1073/pnas.93.13.6626. PMID 8692868. Bibcode: 1996PNAS...93.6626Z. 
  8. ↑ (in Dutch) De microbemens: Het belang van het onzichtbare leven.. Amsterdam: Athenaeum, Polak & Van Gennep. September 2017. ISBN 978-90-253-0692-2. 
  9. ↑ "Bacterial genetic pathway involved in body odor production discovered" (Press release). Society for General Microbiology. 30 March 2015.
  10. ↑ Human Wildlife: The Life That Lives On Us.. Baltimore, Md.: The Johns Hopkins University Press. 2003. pp. 93–94. ISBN 978-0-8018-7407-9. 
  11. ↑ Charles, M.; Martin, B.; Ginies, C.; Etievant, P.; Coste, G.; Guichard, E. (2000). "Potent aroma compounds of two red wine vinegars". Journal of Agricultural and Food Chemistry 48 (1): 70–77. doi:10.1021/jf9905424. PMID 10637054. https://pubmed.ncbi.nlm.nih.gov/10637054/. 
  12. ↑ "Foot odor due to microbial metabolism and its control". Canadian Journal of Microbiology 52 (4): 357–364. April 2006. doi:10.1139/w05-130. PMID 16699586. 
  13. ↑ Gabashvili, Irene S. (2020). "Cutaneous Bacteria in the Gut Microbiome as Biomarkers of Systemic Malodor and People Are Allergic to Me (PATM) Conditions: Insights from a Virtually Conducted Clinical Trial". JMIR Dermatology 3: e10508. doi:10.2196/10508. https://derma.jmir.org/2020/1/e10508. 
  14. ↑ Avoiding Attack: The Evolutionary Ecology of Crypsis, Aposematism, and Mimicry. Oxford University Press. 2018. ISBN 978-0-19-186849-8. [page needed]
  15. ↑ "Olfactory systems and neural circuits that modulate predator odor fear". Frontiers in Behavioral Neuroscience 8: 72. 11 March 2014. doi:10.3389/fnbeh.2014.00072. PMID 24653685. 
  16. ↑ 16.0 16.1 16.2 "Olfaction in the Multisensory Processing of Faces: A Narrative Review of the Influence of Human Body Odors". Frontiers in Psychology 12: 750944. 2021-10-05. doi:10.3389/fpsyg.2021.750944. PMID 34675855. 
  17. ↑ 17.0 17.1 "Odor signatures and kin recognition". Physiology & Behavior 34 (3): 445–448. March 1985. doi:10.1016/0031-9384(85)90210-0. PMID 4011726. 
  18. ↑ 18.0 18.1 "Evolutionary psychology and perfume design". Applied Evolutionary Psychology. 2011. pp. 330–348. doi:10.1093/acprof:oso/9780199586073.003.0020. ISBN 978-0-19-958607-3. 
  19. ↑ "Peroxisomes in the apocrine sweat glands of the human axilla and their putative role in pheromone production". Cellular and Molecular Life Sciences 58 (9): 1344–1349. August 2001. doi:10.1007/PL00000946. PMID 11577991. 
  20. ↑ Lundström & Olsson 2010:"In addition, the impact that biological factors have on our percept of body odors has recently been indirectly demonstrated by several experiments. Our percept of body odors is dependent on the sexual orientations of both the donor and the perceiver (Martins et al., 2005), and heterosexual women's percept of men's body odor varies over their menstrual cycle (Roberts et al., 2004)."
  21. ↑ "Lower olfactory threshold during the ovulatory phase of the menstrual cycle". Biological Psychology 63 (3): 269–279. July 2003. doi:10.1016/s0301-0511(03)00076-0. PMID 12853171. 
  22. ↑ 22.0 22.1 "Influence of HLA on human partnership and sexual satisfaction". Scientific Reports 6: 32550. August 2016. doi:10.1038/srep32550. PMID 27578547. Bibcode: 2016NatSR...632550K. 
  23. ↑ "Brain response to putative pheromones in lesbian women". Proceedings of the National Academy of Sciences of the United States of America 103 (21): 8269–8274. May 2006. doi:10.1073/pnas.0600331103. PMID 16705035. Bibcode: 2006PNAS..103.8269B. 
  24. ↑ "Gay Men Are Found to Have Different Scent of Attraction". The New York Times. 9 May 2005. https://www.nytimes.com/2005/05/09/science/09cnd-smell.html. 
  25. ↑ "Possible olfaction-based mechanisms in human kin recognition and inbreeding avoidance". Journal of Experimental Child Psychology 85 (3): 279–295. July 2003. doi:10.1016/s0022-0965(03)00061-4. PMID 12810039. 
  26. ↑ 26.0 26.1 26.2 "Transcriptional profiling of lung cell populations in idiopathic pulmonary arterial hypertension". Pulmonary Circulation 10 (1): 803. August 1989. doi:10.2307/1131020. PMID 32166015. 
  27. ↑ 27.0 27.1 27.2 "Maternal chemosignals enhance infant-adult brain-to-brain synchrony". Science Advances 7 (50): eabg6867. December 2021. doi:10.1126/sciadv.abg6867. PMID 34890230. Bibcode: 2021SciA....7.6867E. 
  28. ↑ 28.0 28.1 "Does the newborn baby find the nipple by smell?" (in English). Lancet 344 (8928): 989–990. October 1994. doi:10.1016/S0140-6736(94)91645-4. PMID 7934434. 
  29. ↑ "Pheromones for Newborns", Neurobiology of Chemical Communication, Frontiers in Neuroscience (Boca Raton (FL): CRC Press/Taylor & Francis), 2014, ISBN 978-1-4665-5341-5, PMID 24830031, http://www.ncbi.nlm.nih.gov/books/NBK200997/, retrieved 2022-11-27 
  30. ↑ "Pheromone From Mother's Milk May Rapidly Promote Learning In Newborn Mammals" (in en). https://www.sciencedaily.com/releases/2006/10/061010022813.htm. 
  31. ↑ "Chemosensory cues to conspecific emotional stress activate amygdala in humans". PLOS ONE 4 (7): e6415. July 2009. doi:10.1371/journal.pone.0006415. PMID 19641623. Bibcode: 2009PLoSO...4.6415M. 
  32. ↑ 32.0 32.1 32.2 "A SNP in the ABCC11 gene is the determinant of human earwax type". Nature Genetics 38 (3): 324–330. March 2006. doi:10.1038/ng1733. PMID 16444273. 
  33. ↑ 33.0 33.1 33.2 "Body malodours and their topical treatment agents". International Journal of Cosmetic Science 33 (4): 298–311. August 2011. doi:10.1111/j.1468-2494.2011.00649.x. PMID 21401651. 
  34. ↑ 34.0 34.1 34.2 34.3 "A strong association of axillary osmidrosis with the wet earwax type determined by genotyping of the ABCC11 gene". BMC Genetics 10 (1): 42. August 2009. doi:10.1186/1471-2156-10-42. PMID 19650936. 
  35. ↑ "Genetic influences on human body odor: from genes to the axillae". The Journal of Investigative Dermatology 130 (2): 344–346. February 2010. doi:10.1038/jid.2009.396. PMID 20081888. 
  36. ↑ 36.0 36.1 "The Effect of Ethnicity on Human Axillary Odorant Production". Journal of Chemical Ecology 42 (1): 33–39. January 2016. doi:10.1007/s10886-015-0657-8. PMID 26634572. 
  37. ↑ "Effect of ethnicity, gender and age on the amount and composition of residual skin surface components derived from sebum, sweat and epidermal lipids". Skin Research and Technology 20 (1): 97–107. February 2014. doi:10.1111/srt.12091. PMID 23865719. 
  38. ↑ "Grease and Sweat: Race and Smell in Eighteenth-Century English Culture". Cultural and Social History 13 (3): 307–322. 2 July 2016. doi:10.1080/14780038.2016.1202008. 
  39. ↑ "The influence of age, gender and race/ethnicity on the composition of the human axillary microbiome". International Journal of Cosmetic Science 41 (4): 371–377. August 2019. doi:10.1111/ics.12549. PMID 31190339. 
  40. ↑ "Pharmacogenetics of human ABC transporter ABCC11: new insights into apocrine gland growth and metabolite secretion". Frontiers in Genetics 3: 306. 2012. doi:10.3389/fgene.2012.00306. PMID 23316210. 
  41. ↑ "A functional ABCC11 allele is essential in the biochemical formation of human axillary odor". The Journal of Investigative Dermatology 130 (2): 529–540. February 2010. doi:10.1038/jid.2009.254. PMID 19710689. 
  42. ↑ "Learn How to Fight Body Odor". MD Health Network. http://www.mdhealthnetwork.org/BodyOder.html. 
  43. ↑ "Diet quality and the attractiveness of male body odor" (in en). Evolution and Human Behavior 38 (1): 136–143. January 2017. doi:10.1016/j.evolhumbehav.2016.08.002. ISSN 1090-5138. 
  44. ↑ "Antiperspirants Alter Your Armpit Bacteria and Could Actually Make You Smell Worse". RealClearScience. 10 August 2014. http://www.realclearscience.com/blog/2014/08/antiperspirants_alter_your_armpit_bacteria_and_could_actually_make_you_smell_worse.html. 
  45. ↑ "Genetically, Some of Us Never Have Body Odor, But We Still Think We're Smelly". Vice. 17 January 2013. https://www.vice.com/en_us/article/wnnyz9/even-if-you-dont-smell-you-probably-use-deodorant. 
  46. ↑ "Global Deodorants Market is Expected to Reach USD 17.55 Billion by 2020". gosreports.com. http://www.gosreports.com/global-deodorants-market-is-expected-to-reach-usd-17-55-billion-by-2020/. 
  47. ↑ Andrews' Diseases of the Skin: Clinical Dermatology (10th ed.). Saunders. 2005. p. 779. ISBN 978-0-7216-2921-6. 
  48. ↑ Fitzpatrick's Dermatology in General Medicine (6th ed.). McGraw-Hill. 2003. p. 707. ISBN 978-0-07-138076-8. 
  49. ↑ "Body Odor: Causes, Prevention, Treatments" (in en). Medical News Today. http://www.medicalnewstoday.com/articles/173478.php. 

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