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For people with the surname, see Odor (surname).
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"Smell", from Allegory of the Senses by Jan Brueghel the Elder, Museo del Prado

An odor or odour or fragrance is caused by one or more volatilized chemical compounds, generally at a very low concentration, that humans or other animals perceive by the sense of olfaction. Odors are also commonly called scents, which can refer to both pleasant and unpleasant odors. The terms fragrance and aroma are used primarily by the food and cosmetic industry to describe a pleasant odor, and are sometimes used to refer to perfumes, and to describe floral scent. In contrast, malodor, stench, reek, and stink are used specifically to describe unpleasant odor. The term smell (in its noun form) is used for both pleasant and unpleasant odors.

In the United Kingdom, odour refers to scents in general. In the United States and for many non-native English speakers around the world, odor generally has a negative connotation, as a synonym for stink; on the other hand, scent or aroma are used by those people to indicate "pleasant smells".[1]


Odor control covers at a sewage treatment plant: Under these covers, grit and gravel are settled out of the wastewater.

The sense of smell gives rise to the perception of odors, mediated by the olfactory nerve. The olfactory receptor (OR) cells are neurons present in the olfactory epithelium, a small patch of tissue in back of the nasal cavity. There are millions of olfactory receptor neurons that act as sensory signaling cells. Each neuron has cilia in direct contact with air. The olfactory nerve is considered the smell mediator, the axon connects the brain to the external air. Odorous molecules act as a chemical stimulus.[2] Molecules bind to receptor proteins extended from cilia, initiating an electric signal.

The primary sequences of thousands of olfactory receptors are known from the genomes of more than a dozen organisms: they are seven-helix transmembrane proteins, but there are (as of July 2011) no known structures of any OR. There is a highly conserved sequence in roughly three quarters of all ORs that is a tripodal metal ion binding site,[3] and Suslick has proposed that the ORs are in fact metalloproteins (most likely with zinc, copper and possibly manganese ions) that serve as a Lewis Acid site for binding of many odorant molecules. Crabtree, in 1978, had previously suggested that Cu(I) is "the most likely candidate for a metallo-receptor site in olfaction" for strong-smelling volatiles which are also good metal-coordinating ligands, such as thiols.[4] Zhuang, Matsunami and Block, in 2012, confirmed the Crabtree/Suslick proposal for the specific case of a mouse OR, MOR244-3, showing that copper is essential for detection of certain thiols and other sulfur-containing compounds. Thus, by using a chemical that binds to copper in the mouse nose, so that copper wasn’t available to the receptors, the authors showed that the mice couldn't detect the thiols. However, these authors also found that MOR244-3 lacks the specific metal ion binding site suggested by Suslick, instead showing a different motif in the EC2 domain.[5]

When the signal reaches a threshold, the neuron fires, sending a signal traveling along the axon to the olfactory bulb, part of the limbic system of the brain. Interpretation of the smell begins, relating the smell to past experiences and in relation to the substance(s) emitted. The olfactory bulb acts as a relay station connecting the nose to the olfactory cortex in the brain. Olfactory information is further processed and projected through a pathway to the central nervous system (CNS), which controls emotions and behavior as well as basic thought processes.

Odor sensation usually depends on the concentration (number of molecules) available to the olfactory receptors. A single odorant stimulus type is typically recognized by multiple receptors, and different odorants are recognized by combinations of receptors, the patterns of neuron signals helping to identify the smell. The olfactory system does not interpret a single compound, but instead the whole odorous mix, not necessarily corresponding to concentration or intensity of any single constituent.[6][7]

The widest range of odors consists of organic compounds, although some simple compounds not containing carbon, such as hydrogen sulfide and ammonia, are also odorants. The perception of an odor effect is a two-step process. First, there is the physiological part; the detection of stimuli by receptors in the nose. The stimuli are processed by the region of the human brain which is responsible for olfaction. Because of this, an objective and analytical measure of odor is impossible. While odor feelings are very personal perceptions, individual reactions are related to gender, age, state of health, and personal history.

Common odors that people are used to, such as their own body odor, are less noticeable to individuals than external or uncommon odors. This is due to habituation; after continuous odor exposure, the sense of smell fatigues quickly, but recovers rapidly after the stimulus is removed.[8] Odors can change due to environmental conditions, for example odors tend to be more distinguishable in cool dry air.[9]

Habituation affects the ability to distinguish odors after continuous exposure. The sensitivity and ability to discriminate odors diminishes with exposure, and the brain tends to ignore continuous stimulus and focus on differences and changes in a particular sensation. When odorants are mixed, the conditioned odorant is blocked out because of habituation. This depends on the strength of the odorants in the mixture which can change perception and processing of an odor. This process helps classify similar odors as well as adjust sensitivity to differences in complex stimuli.[10]

For most untrained people, the process of smelling gives little information concerning the specific ingredients of an odor. Their smell perception primarily offers information related to the emotional impact.[citation needed] Experienced people, however, such as flavorists and perfumers, can pick out individual chemicals in complex mixes through smell alone.

Odor perception is a primal sense. The sense of smell enables pleasure, can subconsciously warn of danger, help locate mates, find food, or detect predators. Humans have a surprisingly good sense of smell (even though they only have 350 functional olfactory receptor genes compared to the 1,300 found in mice) correlated to an evolutionary decline in sense of smell. Human's remarkable sense of smell is just as good as many animals, and can distinguish a diversity of odors- approximately 10,000 scents. Bushdid et al. reported, however, that humans can distinguish about one trillion odors.[11]

Gordon Shepard proposes that the retro-nasal route of olfaction (odorants introduced to the olfactory mucosa through the oral cavity often as food) was partially responsible for the development of human olfactory acuity. He suggested the evolutionary pressure of diversification of food sources and increased complexity of food preparation presented humans with a broader range of odorants, ultimately leading to a "richer repertoire of smells." However, animals such as dogs show a greater sensitivity to odors than humans especially in studies using short-chained compounds. Higher cognitive brain mechanisms and more olfactory brain regions enable humans to discriminate odors better than other mammals despite fewer olfactory receptor genes.[12]

Different categorizations of primary odors have been proposed, among others this, which relies on seven primary odors (with examples):[13][14][15]

  1. Musky – perfumes/aftershave
  2. Putrid – rotten eggs
  3. Pungent – vinegar
  4. Camphoraceous – mothballs
  5. Ethereal – dry cleaning fluid
  6. Floral – roses (see also floral scent)
  7. Pepperminty – mint gum

Although recently progress has been made, the idea of primary perceptions is disputed, and more so probably the concept of primary odors.[15]


The ability to identify odors varies among people and decreases with age. Studies show there are sex differences in odor discrimination; women usually outperform men.[16] Pregnant women also have increased smell sensitivity, sometimes resulting in abnormal taste and smell perceptions, leading to food cravings or aversions.[17] Deficits in smell also increase with age as well as a prevalence of taste problems (the sense of smell tends to dominate the sense of taste). Chronic smell problems are reported in small numbers for those in their mid-twenties, with numbers increasing steadily with overall sensitivity beginning to decline in the second decade of life, and then deteriorating appreciably as age increases to over 70 years of age.[18]

In Germany, the concentrations of odorants have since the 1870s been defined by "Olfaktometrie", which helps to analyze the human sense of smell using the following parameters: odor substance concentration, intensity of odor, and hedonic assessment.

To establish the odor concentration, an olfactometer test is used, which employs a panel of human noses as sensors. In the olfactometry testing procedure, a diluted odorous mixture and an odor-free gas (as a reference) are presented separately from sniffing ports to a group of panelists, who are housed in an odor-neutral room. They are asked to compare the gases emitted from each sniffing port, after which the panelists are asked to report the presence of odor together with a confidence level such as guessing, inkling, or certainty of their assessment. The gas-diluting ratio is then decreased by a factor of two (i.e. chemical concentration is increased by a factor of two). The panelists are then asked to repeat their judgment. This continues for a number of dilution levels. The responses of the panelists over a range of dilution settings are used to calculate the concentration of the odor in terms of European odor units (ouE/m³). The main panel calibration gas used is butan-1-ol, which at a certain diluting gives 1 ouE/m³.

General survey[edit]

The analytic methods could be subdivided into the physical, the gas chromatographical, and the chemosensory method.

When measuring odor, there is a difference between emission and immission measurements. Emission measurement can be conducted by olfactometry using an olfactometer to dilute the odor sample. On the contrary, olfactometry is rarely used for immission measurement because of the low odor concentrations. The same measuring principles are used, but the judgment of the air assay happens without diluting the samples.


Odor measurement is essential for odor regulation and control.[19] An odor emission often consists of a complex mixture of many odorous compounds. Analytical monitoring of individual chemical compounds present in such odor is usually not practical. As a result, odor sensory methods, instead of instrumental methods, are normally used to measure such odor. Odor sensory methods are available to monitor odor both from source emissions and in the ambient air. These two diverse circumstances require different approaches for measuring odor. The collection of odor samples is more easily accomplished for a source emission than for an odor in the ambient air.[20]

Field measurement with portable olfactometers seems more effective, but the use of Field Olfactometers is not regulated in Europe so far, while it is popular in the U.S. and Canada, where several States set limits at the receptor sites or along the perimeter of odor emitting plants, expressed in units of dilution to threshold (D/T).[21]

Different aspects of odor can be measured through a number of quantitative methods, such as assessing concentration or apparent intensity.

Initial entry into a room provides the most accurate sensing of smell, before habituation begins to change perception of odor.

Sensation of odor has 4 properties related to threshold and tolerance: odor concentration, odor intensity, odor quality, and hedonic tone.

Measuring concentration[edit]

Odor concentration is an odor's pervasiveness. To measure odor sensation, an odor is diluted to certain amounts to reach a detection or recognition threshold. The detection threshold is the concentration of an odor in air when 50% of a population can distinguish between the odorous sample and an odor free blank. The recognition threshold is the concentration of an odor in air in which 50% of a population can discern from an odorous sample and odor free blank.The recognition odor threshold is usually a factor of 2 to 5 times higher than the detection threshold.[13]

The measurement of odor concentration is the most widespread method to quantify odors. It is standardized in CEN EN 13725:2003.[22] The method is based on dilution of an odor sample to the odor threshold (the point at which the odor is only just detectable to 50% of the test panel). The numerical value of the odor concentration is equal to the dilution factor that is necessary to reach the odor threshold. Its unit is the European Odour Unit, OUE. Therefore, the odor concentration at the odor threshold is 1 OUE by definition.

To establish the odor concentration, an olfactometer is used which employs a group of panelists. A diluted odorous mixture and an odor-free gas (as a reference) are presented from sniffing ports to a group of panelists. In comparing the odor emitted from each port, the panelists are asked to report if they can detect a difference between the ports. The gas-diluting ratio is then decreased by a factor of 1.4 or two (i.e., the concentration is increased accordingly). The panelists are asked to repeat their judgment. This continues until the panelists respond certain and correct twice in a row. These responses are used to calculate the concentration of the odor in terms of European odor units (OUE/m3).

The test persons must fulfill certain requirements, for example regarding their sensitivity of odor perception. The main panel calibration gas to verify this requirement used is n-Butanol (as 1 OUE/m3≡40 ppb/v n-butanol).[23]

To collect an odor sample, the samples must be collected using specialized sample bags, which are made from an odor free material e.g. Teflon. The most accepted technique for collecting odor samples is the lung technique, where the sample bag is placed in a sealed drum, and a vacuum is placed on the drum, which fills the sample bag as the bag expands, and draws the sample from the source into the bag. Critically, all components which touch the odor sample, must be odor free, which includes sample lines and fittings.

A human's odor detection threshold is variable. Repeated exposure to an odorant leads to enhanced olfactory sensitivity and decreased detection thresholds for a number of different odorants.[24] It was found in a study that humans that were completely unable to detect the odor of androstenone developed the ability to detect it after repeated exposure.[25]

Humans can discriminate between two odorants that differ in concentration by as little as 7%.[26]

There are a number of issues which have to be overcome with sampling, these include: – If the source is under vacuum – if the source is at a high temperature – If the source has high humidity

Issues such as temperature and humidity are best overcome using either pre-dilution or dynamic dilution techniques.


Odor intensity is the perceived strength of odor sensation. This intensity property is used to locate the source of odors and perhaps most directly related to odor nuisance.[7]

Perceived strength of the odor sensation is measured in conjunction with odor concentration. This can be modeled by the Weber-Fechner law: I = a × log(c) + b[27]

I is the perceived psychological intensity at the dilution step on the butanol scale, a is the Weber-Fechner coefficient, C is the chemical concentrations, and b is the intercept constant (0.5 by definition)[27]

Odor intensity can be expressed using an odor intensity scale, which is a verbal description of an odor sensation to which a numerical value is assigned.[27]

Odor intensity can be divided into the following categories according to intensity:

0 – no odor
1 – very weak (odor threshold)
2 – weak
3 – distinct
4 – strong
5 – very strong
6 – intolerable

This method is applied by in the laboratory and is done so by a series of suitably trained panelists/observers who have been trained to appropriately define intensity.

Hedonic tone assessment[edit]

Hedonic assessment is the process of scaling odors on a scale ranging from extremely unpleasant via neutral up to extremely pleasant. It is important to note that intensity and hedonic tone, whilst similar, refer to different things. That is, the strength of the odor (intensity) and the pleasantness of an odor (hedonic tone). Moreover, it is important to note that perception of an odor may change from pleasant to unpleasant with increasing concentration, intensity, time, frequency, and previous experience with a specific odor; all factors determining a response.[28]

The overall set of qualities are sometimes identified as the "FIDOL factors", (short for Frequency, Intensity, Duration, Offensiveness and Location).[29]


The character of an odor is a critical element in assessing an odor. This property is the ability to distinguish different odors and is only descriptive. First a basic description is used such as sweet, pungent, acrid, fragrant, warm, dry, or sour. The odor is then referenced to a source such as sewage or apple which can then be followed by a reference to a specific chemical such as acids or gasoline.[7]

Most commonly, a set of standard descriptors is used, which may range from fragrant to sewer odor.[30] Although the method is fairly simplistic, it is important for the FIDOL factors to be understood by the person recording the character. This method is most commonly used to define the character of an odor which can then be compared to other odors. It is common for olfactometry laboratories to report character as an additional factor post sample analysis.

Interpreting dispersion modeling[edit]

In many countries odor modeling is used to determine the extent of an impact from an odor source. These are a function of modeled concentration, averaging time (over what time period the model steps are run over (typically hourly) and a percentile. Percentiles refer to a statistical representation of how many hours per year, the concentration C may be exceeded based on the averaging period.

Sampling from area sources[edit]

There are two main odor sampling techniques, the direct odor sampling and the indirect odor sampling technique. Indirect refers to collecting samples from the air stream which has already passed over the emitting surface.

Direct sampling[edit]

Direct refers to the placement of an enclosure on or over an emitting surface from which samples are collected, and an odor emission rate is determined.

The most commonly used direct methods include the flux chamber[31] and wind tunnels which include the UNSW wind tunnel.[32] There are many other available techniques, and consideration should be given to a number of factors before selecting a suitable method.

A source which has implications for this method are sources such as bark bed biofilters, which have a vertical velocity component. For such sources, consideration needs to be given as to the most appropriate method. A commonly used technique is to measure the odor concentration at the emitting surface, and combine this with the volumetric flow rate of air entering the biofilter to produce an emission rate.

Indirect sampling[edit]

Indirect sampling is often referred to as back calculation. It involves the use of a mathematical formula to predict an emission rate.

Many methods are used, but all make use of the same inputs which include surface roughness, upwind and down wind concentrations, stability class (or other similar factor), wind speed, and wind direction.

In the indoor environment[edit]

The human sense of smell is a primary factor in the sensation of comfort. Olfaction as a sensory system brings awareness of the presence of airborne chemicals. Some inhaled chemicals are volatile compounds that act as a stimulus, triggering unwanted reactions such as nose, eye, and throat irritation. Perception of odor and of irritation is unique to each person, and varies because of physical conditions or memory of past exposures to similar chemicals. A person's specific threshold before an odor becomes a nuisance depends also on the frequency, concentration, and duration of an odor.

The perception of irritation from odor sensation is hard to investigate because exposure to a volatile chemical elicits a different response based on sensory and physiological signals, and interpretation of these signals influenced by experience, expectations, personality or situational factors. Volatile organic compounds (VOCs) may have higher concentrations in confined indoor environments due to restricted infiltration of fresh air, as compared to the outdoor environment; leading to greater potential for toxic health exposures from a variety of chemical compounds. Health effects of odor are traced to the sensation of an odor or the odorant itself. Health effects and symptoms vary, including eye, nose, or throat irritation, cough, chest tightness, drowsiness, and mood change; all of which decrease as an odor ceases. Odors may also trigger illnesses such as asthma, depression, stress induced illness, or hypersensitivity. Ability to perform tasks may decrease, and other social/behavioral changes may occur.

Occupants should expect remediation from disturbing and unexpected odors that disturb concentration, diminish productivity, evoke symptoms, and generally increase the dislike for a particular environment. It is important to set occupational exposure limits (OELs) to ensure the health and safety or workers as well as comfort, because exposure to chemicals can elicit physiological and biochemical changes in the upper respiratory system. Standards are hard to set when exposures are not reported and can also be hard to measure. Work force populations vary in levels of discomfort from odors because of exposure history or habituation, and they may not realize possible risks of exposure to chemicals that produce specific odors.[33][34]


Some odors such as perfumes and flowers are sought after, with elite varieties commanding high prices. Whole industries have developed around products to remove unpleasant odors (see deodorant). The perception of odors is also very much dependent upon circumstance and culture. The odor of cooking processes may be pleasurable while one is cooking, but not necessarily after the meal.

The odor molecules transmit messages to the limbic system, the area of the brain that governs emotional responses. Some believe that these messages have the power to alter moods, evoke distant memories, raise their spirits, and boost self-confidence. This belief has led to the concept of "aromatherapy" wherein fragrances are claimed to cure a wide range of psychological and physical problems. Aromatherapy claims that fragrances can positively affect sleep, stress, alertness, social interaction, and general feelings of well-being. However, the evidence for the effectiveness of aromatherapy consists mostly of anecdotes and lacks controlled scientific studies to back up its claims.

With some fragrances, such as those found in perfume, scented shampoo, scented deodorant, or similar products, people can be allergic to the ingredients. The reaction, as with other chemical allergies, can be anywhere from a slight headache to anaphylactic shock, which can result in death.[citation needed]

Unpleasant odors play various roles in nature, often to warn of danger, though this may not be known to the subject who smells it.[35] An odor that is viewed as unpleasant by some people or cultures can be viewed as attractive by others where there is more familiarity or a better reputation.[35]

It is commonly viewed that those holding an unpleasant body odor will be unattractive to others. But studies have shown that a person who is exposed to a particular unpleasant odor can be attracted to others who have been exposed to the same unpleasant odor.[35] This includes smells associated with pollution.[35]

What actually causes a substance to smell unpleasant may be different from what one perceives. For example, perspiration is often viewed as having an unpleasant odor, but it is actually odorless. It is the bacteria in the perspiration that cause the odor.[36]

Unpleasant odors can arise from specific industrial processes, adversely affecting workers and even residents downwind of the industry. The most common sources of industrial odor arise from sewage treatment plants, refineries, animal rendering factories, and industries processing chemicals (such as sulfur) which have odorous characteristics. Sometimes industrial odor sources are the subject of community controversy and scientific analysis.

Body odor is present both in animals and humans and its intensity can be influenced by many factors (behavioral patterns, survival strategies). Body odor has a strong genetic basis both in animals and humans, but it can be also strongly influenced by various diseases and psychological conditions.


The study of odors is a growing field but is a complex and difficult one. The human olfactory system can detect many thousands of scents based on only very minute airborne concentrations of a chemical. The sense of smell of many animals is even better. Some fragrant flowers give off odor plumes that move downwind and are detectable by bees more than a kilometer away.

The study of odors can also get complicated because of the complex chemistry taking place at the moment of a smell sensation. For example, iron-containing metallic objects are perceived to have a distinctive odor when touched, although iron's vapor pressure is negligible. According to a 2006 study[37] this smell is the result of aldehydes (for example nonanal) and ketones (example: 1-octen-3-one) released from the human skin on contact with ferrous ions that are formed in the sweat-mediated corrosion of iron. The same chemicals are also associated with the smell of blood, as ferrous iron in blood on skin produces the same reaction.


Pheromones are odors that are used for communication, and are sometimes called "airborne hormones". A female moth may release a pheromone that can entice a male moth that is several kilometers downwind. Honeybee queens constantly release pheromones that regulate the activity of the hive. Workers can release such smells to call other bees into an appropriate cavity when a swarm moves into new quarters, or to "sound" an alarm when the hive is threatened.

Advanced technology[edit]

There are hopes that advanced technology could do everything from testing perfumes to helping detect cancer or explosives by detecting specific scents, but artificial noses are still problematic. The complex nature of the human nose, its ability to detect even the most subtle of scents, is at the present moment difficult to replicate.

Most artificial or electronic nose instruments work by combining output from an array of non-specific chemical sensors to produce a finger print of whatever volatile chemicals it is exposed to. Most electronic noses need to be "trained" to recognize whatever chemicals are of interest for the application in question before it can be used. The training involves exposure to chemicals with the response being recorded and statistically analyzed, often using multivariate analysis and neural network techniques, to "learn" the chemicals. Many current electronic nose instruments suffer from problems with reproducibility subject to varying ambient temperature and humidity. An example of this type of technology is the colorimetric sensor array, which visualizes odor through color change and creates a "picture" of it.[38][39][40][41][42][43]

Behavioral cues[edit]

Odor perception is a complex process involving the central nervous system that can evoke psychological and physiological responses. Because the olfactory signal terminates in or near the amygdala odors are strongly linked to memories and can evoke emotions. The amygdala participates in the hedonic or emotional processing of olfactory stimuli.[44] Odors can disturb our concentration, diminish productivity, evoke symptoms, and, in general, increase a dislike for a particular environment. Odors can impact the liking for a person, place, food, or product as a form of conditioning.[45] Memories recalled by odors are significantly more emotional and evocative than those recalled by the same cue presented visually or auditorily.[46] Odors can become conditioned to experiential states and when later encountered have directional influences on behavior. Doing a frustrating task in a scented room decreases performance of other cognitive tasks with the presence of the same odor.[47] Nonhuman animals communicate their emotional states through changes in body odor and human body odors are indicative of emotional state.[48]

Human body odors influence interpersonal relationships. Human body odors are involved in adaptive behaviors, such as parental attachment in infants or partner choice in adults. "Mothers can discriminate the odor of their own child, and infants recognize and prefer the body odor of their mother over that of another woman. This maternal odor appears to guide infants toward the breast and to have a calming effect." Body odor is involved in the development of infant–mother attachment and is essential to a child’s social and emotional development bringing feelings of security. Reassurance created by familiar parental body odors may contribute significantly to the attachment process.[49] Human body odors can also affect mate choice. Fragrances are commonly used to raise sexual attractiveness and induce sexual arousal. Researchers found that people choose perfume that interacts well with their body odor.[50]

How a man smells is critical for woman to find a lover. Body odor is a sensory cue critical for mate selection because it is a signal of immunological health. Women prefer men with major histocompatibility complex (MHC) genotypes and odor different from themselves especially during ovulation. Different MHC alleles are favorable because different allele combinations would maximize disease protection and minimize recessive mutations in offspring. Biologically females tend to select mates "who are most likely to secure offspring survival and thus increase the likelihood that her genetic contribution will be reproductively viable."[51]

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 gay and straight males' brains respond in different ways to two odors that may be involved in sexual arousal, and that the gay men respond in the same way as straight women, though it could not be determined whether this was cause or effect. 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 their response to female cues was similar to straight males.[52] According to the researchers, this research suggests a possible role for human pheromones in the biological basis of sexual orientation.[53]

An odor can cue recall of a distant memory. Most memories that pertain to odor come from the first decade of life, compared to verbal and visual memories which usually come from the 10th to 30th years of life.[54] Odor-evoked memories are more emotional, associated with stronger feelings of being brought back in time, and have been thought of less often as compared to memories evoked by other cues.[55]

Use in design[edit]

The sense of smell is often overlooked as a way of marketing products. The deliberate and controlled application of scent is used by designers, scientists, artists, perfumers, architects and chefs. Some applications of scents in environments are in casinos, hotels, private clubs and new automobiles. For example, "technicians at New York City’s Sloan-Kettering Cancer Center disperse vanilla-scented oil into the air to help patients cope with the claustrophobic effects of MRI testing. Scents are used at the Chicago Board of Trade to lower the decibel level on the trading floor."[56]

If ingredients are listed on a product, the term "fragrance" can be used in a general sense.

Scent preferences[edit]

Effect of perfume on sexual attractiveness[edit]

Both men and women use perfume to boost their sexual attractiveness to members of the opposite, or same sex. Indeed, when we find that one perfume or aftershave that works for us, we're hard-pressed to change it - perfume can be as much of our personality as our personal style or likes and dislikes. Olfactory communication is completely natural in humans: we don't always realise we've detected people's particular scents when we have. Without perfume or aftershave, we unconsciously detect people's natural scents: in the form of pheromones. Pheromones are usually detected unconsciously, and it is believed that they have an important influence on our social and sexual behaviour [57] Logically then, it follows on that our choice of perfume or aftershave influences how sexually attractive we are to members of the opposite or same sex. Do we choose perfume regardless of our natural scent (as dictated mainly by pheromones) or do we choose to douse ourselves in scents we prefer, regardless of our natural odour? There are a number of hypotheses concerning why we wear perfume or aftershave, and whether it amplifies or reduces our natural scents.

In 2001, a study found that MHC (major histocompatibility complex, a polymorphic set of genes which is important for immune-function in humans, see MHC is correlated with the ingredients found in perfume. This is an important finding because it suggests that humans do, in fact, choose perfumes that complement or enhance their natural scents (their pheromones). This evidence offers much support for the hypothesis that perfume is chosen by individuals to amplify the statement of their physical health . Research suggests that this advertisement of good health will, in fact, enhance females’ attractiveness to the opposite sex as a health markers have been shown to do.[58] While strong evidence has been found to support the hypothesis that wearing perfume enhances females’ attractiveness to males, little research has been done into the effect of aftershave on males’ attractiveness to females. Considerably more research has covered the effect of males’ natural odour and females’ ratings of attractiveness. What is interesting to note is that in many studies (e.g.[59]) is that odour predicted attractiveness when female raters were not on any form of contraceptive pill. For those who were, there was no relation between attractiveness and body odour.

It stands to reason that odour can increase or decrease ratings of attractiveness because the olfactory receptors in the brain are directly linked with the limbic system, the part of the brain that is thought to be most involved with emotion. This link is an important one, because if an individual associates positive affect (elicited by pheromones[60]), with a potential mate, their liking for, and attraction to, that potential mate will be increased.[61] Although not a typically evolutionary hypothesis, this hypothesis is one that acknowledges how humans have adapted their mating strategies to modern-day societal norms.

Major histocompatibility complex (MHC) and body odor preferences[edit]

Major histocompatibility complex (MHC) is a genotype found in vertebrates including humans. MHC is thought to contribute to mate choice in animals and humans. In sexual selection, females opt for mates with MHC which differs from their own, optimising genes for their offspring.[62] The ‘heterozygote advantage’ and ‘Red Queen’ explanations for these findings fall under the ‘pathogen hypothesis’. Due to differences in MHC alleles’ resistance to pathogens, preference for mates with a dissimilar MHC composition has been argued to act as a mechanism to avoid infectious diseases. According to the ‘heterozygote advantage’ hypothesis, diversity within the MHC genotype is beneficial for the immune system due to a greater range of antigens available to the host. Therefore, the hypothesis proposes that MHC heterozygotes will be superior to MHC homozygotes in fighting off pathogens. Experimental research has shown mixed findings for this idea.[63] The ‘Red Queen’ or ‘rare-allele’ hypothesis suggests that diversity in the MHC gene provides a moving target for pathogens, making it more difficult for them to adapt to MHC genotypes in the host.[64] Another hypothesis suggests that preferences for MHC-dissimilar mates could serve to avoid inbreeding.[65]

Body odor can provide MHC information. Although less is known about how odor is influenced by MHC genes, possible explanations have been that microbial flora[66] or volatile acids[67] are affected by the gene, which can be detected in body odor. Female mice and humans have both shown odor preferences for males with MHC-dissimilarity.[68] Research has shown that women prefer the scent of men with dissimilar MHC genes. In a study, women rated the scent of T-shirts, worn over two nights by men, as more pleasant when smelling those of MHC-dissimilar men.[69] It has also been found that women were reminded more of current or prior partners when smelling odors from men whose MHC was dissimilar to that of the smeller. A study of married couples found that MHC haplotypes differed between spouses more than chance expectations.[70] Taking oral contraceptives has been found to reverse the MHC-dissimilarity odor preference.[71]

How women's preferences for scent change across the cycle[edit]

Women’s preferences for body odour change across their menstrual cycle.[72] The ovulatory shift hypothesis argues that women experience elevated immediate sexual attraction on high relative to low fertile days of the cycle to men with characteristics that reflect good genetic quality.[73] Body odour may provide significant cues about a potential sexual partner’s genetic quality, reproductive status and health, with preferences for particular body odours becoming heightened during a woman’s most fertile days.[74] ). As certain body odours can reflect good genetic quality, woman are more likely to prefer these scents when they are fertile as this is when they are most likely to produce offspring with any potential mates, with conception risk being related to a preference for the scent of male symmetry.[72] Men also prefer the scent of woman at their fertile cycle points.[75]

There are several scents that reflect good genetic quality that females prefer during the most fertile phase of their cycles. Women prefer the scent of symmetrical men more during the fertile phases of their menstrual cycle than during their infertile phases,[76] with estrogen positively predicting women’s preferences for the scent of symmetry.[77] Women’s preferences for masculine faces is greatest when their fertility is at its highest,[78] and so is the preference for attractive faces.[79] Other scents found to be preferred by women in the most fertile phase of their cycle are, the scent for developmental stability,[80] and the scent for dominance.[81]

If women are taking the contraceptive pill the changes in mate scent preferences over the menstrual cycle are not expressed.[82] If odour plays a role in human mate choice then the contraceptive pill could disrupt disassortative mate preferences.[83] Those taking the contraceptive pill show no significant preference for the scent of either symmetrical or asymmetrical men whereas normally cycling women prefer the scent of shirts worn by symmetrical men.[84] Males’ preferences for women’s scent may also change if the woman is taking oral contraceptives. When women take the contraceptive pill this has been found to demolish the cycle attractiveness of odours than men find attractive in normally ovulating women.[85] Therefore, the contraceptive pill affects both women’s preferences for scent and also affects their own scents, making their scents less attractive to males than the scent of normally cycling women.

See also[edit]


  1. ^ Spectrum Language Arts, Grade 8. Carson-Dellosa Publishing. 2014. p. 159. ISBN 1483814246. Retrieved July 21, 2015. 
  2. ^ de march, Claire A.; Ryu, sangEun; Sicard, Gilles; Moon, Cheil; Golebiowski, Jérôme (September 2015). "Structure–odour relationships reviewed in the postgenomic era". Flavour and Fragrance Journal. 30 (5): 342–361. doi:10.1002/ffj.3249. 
  3. ^ Wang, J.; Luthey-Schulten, Z.; Suslick, K. S. (2003). "Is the Olfactory Receptor A Metalloprotein?". Proc. Natl. Acad. Sci. U.S.A. 2003 (100): 3035–3039. Bibcode:2003PNAS..100.3035W. doi:10.1073/pnas.262792899. PMC 152240free to read. PMID 12610211. 
  4. ^ Crabtree, R.H. (1978). "Copper(I) – Possible Olfactory Binding-Site". J. Inorg. Nucl. Chem. 1978 (40): 1453. doi:10.1016/0022-1902(78)80071-2. 
  5. ^ Duan, Xufang; Block, Eric; Li, Zhen; Connelly, Timothy; Zhang, Jian; Huang, Zhimin; Su, Xubo; Pan, Yi; Wu, Lifang; Chi, Qiuyi; Thomas, Siji; Zhang, Shaozhong; Ma, Minghong; Matsunami, Hiroaki; Chen, Guo-Qiang; Zhuang, Hanyi (2012). "Crucial role of copper in detection of metal-coordinating odorants". Proc. Natl. Acad. Sci. U.S.A. 2012 (109): 3492–3497. Bibcode:2012PNAS..109.3492D. doi:10.1073/pnas.1111297109. PMC 3295281free to read. PMID 22328155. 
  6. ^ Axel, Richard (1995). "The molecular logic of smell". Scientific American. 273 (4): 154–159. Bibcode:1995SciAm.273d.154A. doi:10.1038/scientificamerican1095-154. 
  7. ^ a b c Spengler, p. 492
  8. ^ Chaudhury, D; Manella, L; Arellanos, A; Escanilla, O; Cleland, T. A.; Linster, C (2010). "Olfactory bulb habituation to odor stimuli". Behavioral Neuroscience. 124 (4): 490–9. doi:10.1037/a0020293. PMC 2919830free to read. PMID 20695648. 
  9. ^ Salthammer, Tunga; Bahadir, Müfit (2009). "Occurrence, Dynamics and Reactions of Organic Pollutants in the Indoor Environment". CLEAN - Soil, Air, Water. 37 (6): 417–435. doi:10.1002/clen.200900015. 
  10. ^ Devriese, S; Winters, W; Stegen, K; Diest, I Van; Veulemans, H; Nemery, B; Eelen, P (2000). "Generalization of acquired somatic symptoms in response to odors: a pavlovian perspective on multiple chemical sensitivity". Psychosom Med. 62 (6): 751–759. doi:10.1097/00006842-200011000-00003. PMID 11138993. 
  11. ^ C. Bushdid, M. O. Magnasco, L. B. Vosshall, A. Keller, C.; Magnasco, M. O.; Vosshall, L. B.; Keller, A. (21 March 2014). "Humans Can Discriminate More than 1 Trillion Olfactory Stimuli". Science. 343 (6177): 1370–1372. Bibcode:2014Sci...343.1370B. doi:10.1126/science.1249168. PMID 24653035. 
  12. ^ Shepherd, Gordon M. (2004). "The Human Sense of Smell: Are We Better Than We Think?". PLoS Biology. 2 (5): e146. doi:10.1371/journal.pbio.0020146. PMC 406401free to read. PMID 15138509. 
  13. ^ a b Spengler, p. 483
  14. ^ Oracle Education Foundation (25 Aug 2010). "Your Sense of Smell – The Senses". ThinkQuest Library. 
  15. ^ a b Auffarth, B. (2013). "Understanding smell – the olfactory stimulus problem. Neuroscience & Biobehavioral Reviews". Neuroscience & Biobehavioral Reviews. 37 (8): 1667–1679. doi:10.1016/j.neubiorev.2013.06.009. 
  16. ^ Doty, Richard L.; Applebaum, Steven; Zusho, Hiroyuki; Settle, R.Gregg (1985). "Sex differences in odor identification ability: A cross-cultural analysis". Neuropsychologia. 23 (5): 667–72. doi:10.1016/0028-3932(85)90067-3. PMID 4058710. 
  17. ^ Nordin, Steven; Broman, Daniel A.; Olofsson, Jonas K.; Wulff, Marianne (2004). "A Longitudinal Descriptive Study of Self-reported Abnormal Smell and Taste Perception". Pregnant Women Chem. Senses. 29 (5): 391–402. doi:10.1093/chemse/bjh040. PMID 15201206. 
  18. ^ Hoffman, H. J.; Cruickshanks, K. J.; Davis, B (2009). "Perspectives on population-based epidemiological studies of olfactory and taste impairment". Annals of the New York Academy of Sciences. 1170 (1): 514–30. Bibcode:2009NYASA1170..514H. doi:10.1111/j.1749-6632.2009.04597.x. PMC 2760342free to read. PMID 19686188. 
  19. ^ Ueno, H; Amano, S; Merecka, B; Kośmider, J (2009). "Difference in the odor concentrations measured by the triangle odor bag method and dynamic olfactometry" (PDF). Water Science & Technology. 59 (7): 1339–42. doi:10.2166/wst.2009.112. PMID 19380999. 
  20. ^ "GUIDELINES ON ODOUR POLLUTION & ITS CONTROL" (PDF). Ministry of Environment & Forests, Govt. of India. May 2008. 
  21. ^ Benzo, Maurizio; Mantovani, Alice; Pittarello, Alberto (2012). "Measurement of Odour Concentration of Immissions using a New Field Olfactometer and Markers' Chemical Analysis" (PDF). Chemical Engineering Transactions. 30: 103. 
  22. ^ CEN EN 13725:2003, Air quality – Determination of odour concentration by dynamic olfactometry. sipe-rtd.info
  23. ^ Van Harreveld, A. P.; Heeres, P.; Harssema, H. (1999). "A review of 20 years of standardization of odor concentration measurement by dynamic olfactometry in Europe". Journal of the Air & Waste Management Association. 49 (6): 705–715. 
  24. ^ Cain, W. S.; Gent, J. F. (1991). "Olfactory sensitivity: Reliability, generality, and association with aging". Journal of experimental psychology. Human perception and performance. 17 (2): 382–91. doi:10.1037/0096-1523.17.2.382. PMID 1830082. 
  25. ^ Wysocki, CJ; Dorries, KM; Beauchamp, GK. (1989). "Ability to perceive androstenone can be acquired by ostensibly anosmic people". Proc. Natl. Acad. Sci. USA. 86 (20): 7976–7978. Bibcode:1989PNAS...86.7976W. doi:10.1073/pnas.86.20.7976. PMC 298195free to read. PMID 2813372. 
  26. ^ Cain, WS. (1977). "Differential sensitivity for smell: "noise" at the nose". Science. 195 (4280): 796–798. Bibcode:1977Sci...195..796C. doi:10.1126/science.836592. PMID 836592. 
  27. ^ a b c Jiang, J; Coffey, P; Toohey, B (2006). "Improvement of odor intensity measurement using dynamic olfactometry". Journal of the Air & Waste Management Association (1995). 56 (5): 675–83. doi:10.1080/10473289.2006.10464474. PMID 16739805. 
  28. ^ Spengler, p. 486
  29. ^ "F.i.d.o.l.". Retrieved 2011-11-30. 
  30. ^ "Odour Assessment". MFE.govt.nz. Retrieved 2012-12-30. 
  31. ^ "Flux Chamber Measurements: Defensible Analytical Data for Evaluating Human Health Risk". Ceschmidt.com. Retrieved 2012-12-30. 
  32. ^ UNSW wind tunnel dimensions. Odour.unsw.edu.au
  33. ^ Young, Christopher A. (2010). "What Smells?". Pollution Engineering. 42 (5). 
  34. ^ Dalton, P (2002). "Odor, irritation and perception of health risk". International Archives of Occupational and Environmental Health. 75 (5): 283–90. doi:10.1007/s00420-002-0312-x. PMID 11981666. 
  35. ^ a b c d Engen, Trygg (1991). Odor sensation and memory. New York: Praeger. ISBN 0-275-94111-6. 
  36. ^ What's Happening to My Body? Book for Boys: Revised Edition – Lynda Madaras, Area Madaras, Simon Sullivan – Google Boeken. Books.google.com. 2007-06-08. ISBN 9781557047694. Retrieved 2012-12-30. 
  37. ^ Communication The Two Odors of Iron when Touched or Pickled: (Skin) Carbonyl Compounds and Organophosphines Dietmar Glindemann, Andrea Dietrich, Hans-Joachim Staerk, Peter Kuschk Angewandte Chemie International Edition web release 2006 doi:10.1002/anie.200602100 PMID 17009284
  38. ^ Rakow, N. A.; Suslick, K. S. (2000). "A Colorimetric Sensor Array for Odour Visualization". Nature. 406 (6797): 710–714. doi:10.1038/35021028. 
  39. ^ Suslick, Kenneth S. (2011). "An Optoelectronic Nose:"Seeing" Smells by Means of Colorimetric Sensor Arrays" (PDF). MRS Bulletin. 29 (10): 720–5. doi:10.1557/mrs2004.209. PMID 15991401. 
  40. ^ Lim, S. H.; Feng, L; Kemling, J. W.; Musto, C. J.; Suslick, K. S. (2009). "An optoelectronic nose for the detection of toxic gases". Nature Chemistry. 1 (7): 562–7. Bibcode:2009NatCh...1..562L. doi:10.1038/nchem.360. PMC 2761044free to read. PMID 20160982. 
  41. ^ Suslick, B. A.; Feng, L.; Suslick, K. S. (2010). "Discrimination of Complex Mixtures by a Colorimetric Sensor Array: Coffee Aromas". Anal. Chem. 2010 (82): 2067–2073. doi:10.1021/ac902823w. 
  42. ^ Feng, Liang; Musto, Christopher J.; Kemling, Jonathan W.; Lim, Sung H.; Suslick, Kenneth S. (2010). "A colorimetric sensor array for identification of toxic gases below permissible exposure limits" (PDF). Chemical Communications. 46 (12): 2037–9. doi:10.1039/B926848K. PMID 20221484. 
  43. ^ Feng, L.; Musto, C.J.; Suslick, K. S. (2010). "A Simple and Highly Sensitive Colorimetric Detection Method for Gaseous Formaldehyde". J. Am. Chem. Soc. 2010 (132): 4046–4047. doi:10.1021/ja910366p. PMC 2854577free to read. PMID 20218682. 
  44. ^ Zald, David H.; Pardo, J. V. (1997). "Emotion, olfaction, and the human amygdala: Amygdala activation during aversive olfactory stimulation". PNAS. 94 (8): 4119–4124. Bibcode:1997PNAS...94.4119Z. doi:10.1073/pnas.94.8.4119. PMC 20578free to read. PMID 9108115. 
  45. ^ Wrzesniewski, Amy; McCauley, Clark; Rozin, Paul (1999). "Odor and Affect: Individual Differences in the Impact of Odor on Liking for Places, Things and People". Chem. Senses. 24 (6): 713–721. doi:10.1093/chemse/24.6.713. PMID 10587506. 
  46. ^ Herz, Rachel S. (2004). "A Naturalistic Analysis of Autobiographical Memories Triggered by Olfactory Visual and Auditory Stimuli". Chem. Senses. 29 (3): 217–224. doi:10.1093/chemse/bjh025. PMID 15047596. 
  47. ^ Epple, Gisela; Herz, Rachel S. (1999). "Ambient odors associated to failure influence cognitive performance in children". Developmental Psychobiology. 35 (2): 103–107. doi:10.1002/(sici)1098-2302(199909)35:2<103::aid-dev3>3.0.co;2-4. PMID 10461124. 
  48. ^ Chen, D; Haviland-Jones, J. (2000). "Human olfactory communication of emotion" (PDF). Percept Mot Skills. 91 (3 Pt 1): 771–81. doi:10.2466/pms.2000.91.3.771. PMID 11153847. 
  49. ^ Ferdenzi, Camille; Schaal, Benoist; Roberts, S. Craig (2010). "Family Scents: Developmental Changes in the Perception of Kin Body Odor?". Journal of Chemical Ecology. 36 (8): 847–854. doi:10.1007/s10886-010-9827-x. PMID 20640943. 
  50. ^ Lenochová, Pavlína; Vohnoutová, Pavla; Roberts, S. Craig; Oberzaucher, Elisabeth; Grammer, Karl; Havlíček, Jan (2012-03-28). "Psychology of Fragrance Use: Perception of Individual Odor and Perfume Blends Reveals a Mechanism for Idiosyncratic Effects on Fragrance Choice". PLoS ONE. 7 (3): e33810. doi:10.1371/journal.pone.0033810. PMC 3314678free to read. PMID 22470479. 
  51. ^ Herz, Rachel S.; Inzlicht, Michael (2002). "Sex differences in response to physical and social factors involved in human mate selection: The importance of smell for women". Evolution and Human Behavior. 23 (5): 359–364. doi:10.1016/s1090-5138(02)00095-8. 
  52. ^ Berglund, H.; Lindstrom, P.; Savic, I. (2006). "Brain response to putative pheromones in lesbian women". Proceedings of the National Academy of Sciences. 103 (21): 8269–74. Bibcode:2006PNAS..103.8269B. doi:10.1073/pnas.0600331103. PMC 1570103free to read. PMID 16705035. 
  53. ^ Wade, Nicholas (May 9, 2005) "Gay Men are found to have Different Scent of Attraction". NY Times
  54. ^ Larsson, M.; Willander, J. (2009). "Autobiographical odor memory". Ann. N. Y. Acad. Sci. International Symposium on Olfaction and Taste. 1170: 318–323. 
  55. ^ Larsson, M. & J. Willander. 2009. Autobiographical odor memory. Ann. N. Y. Acad. Sci. International Symposium on Olfaction and Taste.
  56. ^ "Miller, Tabitha M.A. Smell". Tabithamiller.com. Retrieved 2012-12-30. 
  57. ^ Grammer, Karl (2005). "Human pheromones and sexual attraction" (PDF). European Journal of Obstetrics and Gynecology and Reproductive Biology. 
  58. ^ Weeden, Jason (2005). "Physical Attractiveness and Health in Western Societies: A Review". Psychological Bulletin. 
  59. ^ Foster, Joshua (2008). "Beauty Is Mostly in the Eye of the Beholder: Olfactory Versus Visual Cues of Attractiveness". The Journal of Social Psychology. 
  60. ^ Jacob, Suma; McClintock, Martha K. (2000-02-01). "Psychological State and Mood Effects of Steroidal Chemosignals in Women and Men". Hormones and Behavior. 37 (1): 57–78. doi:10.1006/hbeh.1999.1559. 
  61. ^ Kohl, James (2001). "Human Pheromones: Integrating Neuroendocrinology and Ethology". Neuroendocrinology Letters. 
  62. ^ Grammer, Karl; Fink, Bernhard; Neave, Nick (February 2005). "Human pheromones and sexual attraction". European Journal of Obstetrics & Gynecology and Reproductive Biology. 118 (2): 135–142. doi:10.1016/j.ejogrb.2004.08.010. 
  63. ^ Penn, D. J.; Potts, W. K. (1999). "The evolution of mating preferences and major histocompatibility complex genes." (PDF). The American Naturalist. 153 (2): 145–164. doi:10.1086/303166. 
  64. ^ Wedekind, C.; Penn, D. (2000). "MHC genes, body odours, and odour preferences" (PDF). Nephrology Dialysis Transplantation. 15 (9): 1269–1271. doi:10.1093/ndt/15.9.1269. 
  65. ^ Potts, W. K; Manning, C. J.; Wakeland, E. K.; Hughes, A. L. (1994). "The role of infectious disease, inbreeding and mating preferences in maintaining MHC genetic diversity: an experimental test" (PDF). Philosophical Transactions of the Royal Society of London B: Biological Sciences. 346: 369–378. doi:10.1098/rstb.1994.0154. 
  66. ^ Singh, P. B.; Herbert, J.; Roser, B.; Arnott, L.; Tucker, D. K.; Brown, R. E. (1990). "Rearing rats in a germ-free environment eliminates their odors of individuality". Journal of Chemical Ecology. 16 (5): 1667–1682. doi:10.1007/bf01014099. 
  67. ^ Singer, A. G.; Beauchamp, G. K.; Yamazaki, K. (1997). "Volatile signals of the major histocompatibility complex in male mouse urine" (PDF). Proceedings of the National Academy of Sciences. 94 (6): 2210–2214. doi:10.1073/pnas.94.6.2210. 
  68. ^ Dunbar, Robin Ian MacDonald; Barrett, Louise (2007). Oxford handbook of evolutionary psychology (1 ed.). Oxford: Oxford University Press. p. 317. ISBN 9780198568308. 
  69. ^ Wedekind, C.; Seebeck, T.; Bettens, F.; Paepke, A. J. (22 June 1995). "MHC-Dependent Mate Preferences in Humans" (PDF). Proceedings of the Royal Society B: Biological Sciences. 260 (1359): 245–249. doi:10.1098/rspb.1995.0087. 
  70. ^ Ober, Carole; Weitkamp, Lowell R.; Cox, Nancy; Dytch, Harvey; Kostyu, Donna; Elias, Sherman (September 1997). "HLA and Mate Choice in Humans". The American Journal of Human Genetics. 61 (3): 497–504. doi:10.1086/515511. 
  71. ^ Thorne, Frances, Fink, Bernhard (2002). "Effects of putative male pheromones on female ratings of male attractiveness: influence of oral contraceptives and the menstrual cycle.". Neuroendocrinology Letters. 23 (4): 291–297. 
  72. ^ a b Thornhill, R., Chapman, J. F., & Gangestad, S. W. (2013). Women’s preferences for men’s scents associated with testosterone and cortisol levels: Pattens across the ovulatory cycle. Evolution and Human Behaviour, 34.3, 216-221
  73. ^ Glidersleeve, K., Haselton, M. G., & Fales, M. R. (2014). Do women’s mate preferences change across the ovulatory cycle? A meta-analytic review. Psychological Bulletin, 140.5, 1205-1259
  74. ^ Havlicek, J., Roberts, C. S, & Flegr, J. (2005). Women’s preference for dominant male odour: effects of mistral cycle and relationship status. Biology Letters, 3.1
  75. ^ Thornhill, R., Gangastad, S. W., Miller, R., Scheyd, G., McCollongh, J. K., & Franklin, M. (2003). Major histocompatibility complex geens, symmetry, and body scent attractiveness in men and women. Behavioural Ecology, 14.5, 668-678
  76. ^ Gangestad, S. W., Simpson, J. A., Cousins, A. J., Garver- Apgar, C. E., & Christensen, P. N. (2004). Women’s preferences for male behavioural displays change across the menstrual cycle. Psychological Science, 15.3, 203-207
  77. ^ Garver- Aprgar, C. E., Gangestad, S. W. & Thornhill, R. (2008). Hormonal correlates of women’s mid-cycle preference for the scent of symmetry. Evolution and Human Behaviour, 29.4, 223-232
  78. ^ Gangestad, S. W., Simpson, J. A., Cousins, A. J., Garver- Apgar, C. E., & Christensen, P. N. (2004). Women’s preferences for male behavioural displays change across the menstrual cycle. Psychological Science, 15.3, 203-207.
  79. ^ Thornhill, R., & Gangestad, S. W. (1999). The scent of symmetry: A human sex pheromone that signals fitness? Evolution and Human Behaviour, 20.3, 175-201
  80. ^ Rikowski, K. Grammer. (1999). Human body odour, symmetry and attractiveness. Proceedings of the Royal Society of London B, 266, 869–874
  81. ^ Havlicek, J., Roberts, C. S, & Flegr, J. (2005). Women’s preference for dominant male odour: effects of mistral cycle and relationship status. Biology Letters, 3.1, DOI: 10.1098/rsbl.2005.0332
  82. ^ Alvergne, A. & Lummaa, V. (2010). Does the contraceptive pill alter mate choice in humans? Trends in Ecology and Evolution, 25.3, 171-179
  83. ^ Roberts, C. S., Gosling, L. M., Carter, V., & Petrie, M. (2008). MHC-correlated ocour preferences in humans and the use of oral contraceptives. Biological Sciences, 275.1652, 2715-2722
  84. ^ Gangestad, S. W. & Thornhill, R. (1998). Menstrual cycle variation in women’s preferences for the scent of symmetrical men. Biological Sciences, 265,1399, 927- 933
  85. ^ Kuukasjarvi, S., Eriksson, P. C. J., Koskela, E., Mappes, T., Nissinen, K., & Rantala, M. J. (2004). Attractiveness of women’s body odours over the menstrual cycle: the role of oral contraceptives and receiver sex. Behavioural Ecology, 15.4, 579-584


  • Spengler, John D.; McCarthy, John F; Samet, Jonathan M. (2000). Indoor Air Quality Handbook. New York, NY, USA: McGraw-Hill Professional Publishing. ISBN 978-0-07-445549-4. 

Further reading[edit]

  • Gilbert, Avery (2008). What the nose knows : the science of scent in everyday life (1st ed.). New York: Crown Publishers. ISBN 978-1-4000-8234-6. 
  • Kaye, Joseph Nathaniel (May 2001). "Symbolic Olfactory Display (Master's Thesis)" (PDF). Symbolic Olfactory Display. Massachusetts Institute of Technology. Retrieved 2011-06-25.  — A survey of current olfactory knowledge, experimental investigation of computer-based olfactory interfaces. Includes extensive reference list, partially annotated.
  • Samet, edited by Jonathan M.; Spengler, John D. (1991). Indoor air pollution : a health perspective. Baltimore: Johns Hopkins University Press. ISBN 978-0-8018-4125-5. 
  • Watson, Lyall (2000). Jacobson's organ and the remarkable nature of smell (1st American ed.). New York: W.W. Norton. ISBN 978-0-393-04908-4. 
  • Majid, Asifa (February 2015). "Olfaction: Scent off". The Economist. 

External links[edit]