Multisensory Research (2024) DOI:10.1163/22134808-bja10130 brill.

com/msr

Review

What is the Relation between Chemosensory Perception

and Chemosensory Mental Imagery?

Charles Spence ∗
Department of Experimental Psychology, New Radcliffe House, University of Oxford, Oxford,
OX2 6BW, UK
* E-mail: [email protected]
ORCID iD: 0000-0003-2111-072X

Received 31 March 2024; accepted 8 August 2024; published online 27 August 2024

Abstract
The study of chemosensory mental imagery is undoubtedly made more difficult because of the
profound individual differences that have been reported in the vividness of (e.g.) olfactory mental
imagery. At the same time, the majority of those researchers who have attempted to study peo-
ple’s mental imagery abilities for taste (gustation) have actually mostly been studying flavour mental
imagery. Nevertheless, there exists a body of human psychophysical research showing that chemosen-
sory mental imagery exhibits a number of similarities with chemosensory perception. Furthermore,
the two systems have frequently been shown to interact with one another, the similarities and dif-
ferences between chemosensory perception and chemosensory mental imagery at the introspective,
behavioural, psychophysical, and cognitive neuroscience levels in humans are considered in this nar-
rative historical review. The latest neuroimaging evidence show that many of the same brain areas
are engaged by chemosensory mental imagery as have previously been documented to be involved
in chemosensory perception. That said, the pattern of neural connectively is reversed between the
‘top-down’ control of chemosensory mental imagery and the ‘bottom-up’ control seen in the case of
chemosensory perception. At the same time, however, there remain a number of intriguing questions
as to whether it is even possible to distinguish between orthonasal and retronasal olfactory mental
imagery, and the extent to which mental imagery for flavour, which most people not only describe
as, but also perceive to be, the ‘taste’ of food and drink, is capable of reactivating the entire flavour
network in the human brain.
Keywords
flavour, olfaction, orthonasal, retronasal, taste

Published with license by Koninklijke Brill BV | DOI: 10.1163/22134808-bja10130

© CHARLES SPENCE, 2024 | ISSN: 2213-4794 (print) 2213-4808 (online)
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2 C. Spence / Multisensory Research (2024)

1. Introduction
The vast majority of research on the topic of mental imagery in humans has
been conducted in the visual modality (e.g., see Galton, 1880, 1883; Ganis,
2013; Kosslyn, 1994; Pearson, 2019). That said, mental imagery also operates
in the other senses too, such as audition (e.g., Gissurarson, 1992; Halpern and
Zatorre, 1999; Hubbard, 2013; Intons-Peterson, 1992; Oh et al., 2013; Reis-
berg, 1992; Zatorre et al., 1996), haptic touch (e.g., Gallace, 2013; Jeannerod,
1995; Klatzky et al., 1991), olfaction (Carrasco and Ridout, 1993; Crowder
and Schab, 1995; Herz and Engen, 1996; Stevenson and Case, 2005; Zatorre,
1999), and taste (see Note 1) (e.g., Sauvageot et al., 2000). According to Ben-
safi et al. (2013, p. 77): “Olfactory and gustatory mental images are defined as
short-term memory representations of olfactory or gustatory events that give
rise to the experience of “smelling with the mind’s nose” or “tasting with the
mind’s tongue”. ” (see also Kosslyn, 2003).
Until a few decades ago, however, many researchers still questioned the
very existence of olfactory mental imagery (e.g., see Elmes, 1998; Elmes
and Jones, 1995; Engen, 1982, 1987; Herz, 2000; Schab, 1991). For instance,
Schab (1991, p. 242) claimed that: “There is no unambiguous empirical evi-
dence supporting an ability to image odors”, while Carrasco and Ridout (1993,
p. 287) state that: “Whether a system of imagery for olfaction exists is cur-
rently an unsettled issue” (Note 2). Researchers have though, by now, managed
to provide extensive psychophysical and cognitive neuroscience support for
the existence of an ‘inner nose’ (e.g., Algom and Cain, 1991; Carrasco and
Ridout, 1993; Levy et al., 1999a; Lorig and Roberts, 1990; Lyman, 1988;
Lyman and McDaniel, 1990; though see also Crowder and Schab, 1995; Schif-
ferstein, 1997).
Just as in the visual modality (e.g., see Kosslyn, 1994; Pearson and Kosslyn,
2015; Pylyshyn, 2003), there has also been some debate concerning the ques-
tion of whether olfactory mental imagery is propositional (i.e., semantically-
mediated) or depictive in nature (Rinck et al., 2009; Sugiyama et al., 2006 —
cf. Koenig et al., 2000). Furthermore, some authors (e.g., Crowder and Schab,
1995; Elmes, 1998; Herz, 2000) have argued that it is not altogether clear
whether the alleged odour images in previous research were perceptual or
propositional in nature, that is, whether they were odour-like rather than
verbal-like experiences. At the same time, however, other researchers have
questioned the extent to which tacit knowledge and task demands may explain
the data from a number of the mental imagery studies that have been published
to date (Farah, 1988; Intons-Peterson, 1983; Pylyshyn, 1981; Schifferstein,
1997).
A large body of cognitive neuroscience research now suggests that visual
mental imagery activates many (if perhaps not necessarily all) of the same
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 3

brain areas as visual perception (e.g., Kosslyn et al., 1995; though see also
Spagna et al., 2021). Intriguingly, however, the patterns of connectivity appear
to operate in opposing directions — ‘bottom-up’ in the case of visual per-
ception and ‘top-down’ in the case of visual mental imagery (see Dijkstra et
al., 2017). For those interested in the chemical senses, it is natural to wonder
whether the same story — namely, that imagery and perception activate the
same network of brain areas — also describes the case of chemosensory per-
ception and chemosensory mental imagery (in all its various guises, stretching
from olfactory mental imagery through gustatory and flavour mental imagery;
see also Royet et al., 2013).
In this narrative review (Ferrari, 2015; Furley and Goldschmied, 2021),
I examine the similarities and differences between chemosensory perception
and chemosensory mental imagery. In the following sections, this question is
addressed at a number of different levels (Finke, 1980), from phenomenol-
ogy (i.e., based on introspection) through to perceptual/psychophysical inter-
actions and behavioural similarities. The degree of overlap in terms of the
underlying neural substrates is also assessed. Ultimately, it is argued that
while there are many similarities between chemosensory mental imagery and
chemosensory perception at these various levels, there are also a number of
salient differences. Furthermore, it remains unclear as to whether a meaning-
ful distinction can be made between orthonasal and retronasal olfactory mental
imagery, and whether the entire flavour network in the human brain (Suen et
al., 2021) is reactivated in the case of people mentally imagining the flavour
of food and drink (see also Seubert, Ohla, Yokomukai, Kellermann and Lund-
ström, 2015; Stevenson and Prescott, 1997).

2. Comparing Chemosensory Mental Imagery to Chemosensory

Perception
2.1. Introspection: Individual Differences in the Vividness of Chemosensory
Mental Imagery
The results of those studies that have used introspective reports to assess the
vividness of people’s mental imagery tend to suggest that visual images are
experienced as most vivid, whereas olfactory and gustatory images tend to
be least vivid (e.g., Ashton and White, 1980; Betts, 1909; Marsella and Qui-
jano, 1974; Schifferstein, 2009; Sheehan, 1967; White et al., 1977 — see
also Gilbert et al., 1998; Lawless, 1997; Lindauer, 1969; Royet et al., 2013;
Young, 2020). Lawless reported that 25% of the participants whom he quizzed
reported never having experienced olfactory mental imagery. When Brower
and Brower (1947) asked participants to imagine the sight, sound, and smell of
frying onions, 57% reported being unable to imagine the smell, whereas only
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4 C. Spence / Multisensory Research (2024)

3% were unable to visualize it. When Lindauer (1969) repeated the test, all his
participants could visualize frying onions, but 30% reported being unable to
imagine the smell.
At the same time, however, the published research also suggests the exis-
tence of large interindividual differences in the ability to create (and/or the
vividness of) olfactory mental images (Schifferstein, 2009; Sulfaro et al.,
2024; Switras, 1978, 1979 — though see also Bensafi et al., 2005; and Taka-
hashi et al., 2023; Zeman, 2024, on the emerging study of aphantasia) (Note 3).
According to research from Fantin et al. (2022), age and gender may affect the
vividness of people’s olfactory mental imagery, at least when it is assessed by
self-report. Meanwhile, another study reported that those individuals who are
classified as good olfactory imagers (by questionnaire) sniffed odorants for
longer and judged them to be more edible and familiar, thus suggesting a bet-
ter semantic knowledge associated with odours (Rouby et al., 2009; see also
Bensafi and Rouby, 2007; Stevenson et al., 2007). People sometimes appear to
recruit visual cortex when they find themselves in the ‘tip-of-the-nose’ state —
that is, visual imagery is seemingly recruited to help try and identify the source
of (or name for) an odour (Qureshy et al., 2000).
In contrast to the well-documented individual differences in the vividness
of olfactory mental imagery, far fewer studies have provided evidence relevant
to addressing the question of whether such individual differences also exist
in terms of people’s ability to imagine basic tastes or flavours. The ability of
people to imagine specifically oral trigeminal stimuli, such as, for example,
the pungency of pepper (Spence, 2024) or ginger (Spence, 2023a), the burn
of chilli (Spence, 2018), and/or the cooling of menthol have also received
virtually no attention from researchers either (Simons and Carstens, 2008;
Stevenson and Prescott, 1997). Meanwhile, studies of craving (a largely sep-
arate area of empirical research; see Kemps and Tiggeman, 2013) (Note 4)
appear to suggest that these two modalities of mental imagery come close
behind vision in terms of driving people’s cravings. For instance, according
to the results of one survey in which people were asked to assign percent-
ages to each of five senses involved in an imagined food craving experience,
vision (40%) scored top, followed by gustatory (31%) (Note 5) and olfactory
(16%), with the senses of touch (10%) and audition (4%) apparently little used
(Tiggemann and Kemps, 2005). The vividness of olfactory mental imagery
responsiveness to food cues has also recently been established as a risk factor
for weight gain due to its role in supporting craving (Perszyk et al., 2023).
Dou et al. (2018) reported that even when participants are instructed to try
and suppress their mental imagery, food items (be they presented as ortho-
graphic stimuli or as drawings) nevertheless still involuntarily elicit olfactory
(an effect in ∼40% of trials), taste (∼54%), touch (∼60%), and visual/auditory
(∼79%) mental imagery. At the same time, some researchers have suggested
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 5

that imagery training may facilitate people’s ability to mentally imagine the
flavour of food (Richardson and Patterson, 1986; see also Bensafi et al., 2013).
As Stevenson and Case (2005, p. 244) note: “self-reports of olfactory imagery
can resemble those obtained for actual perception”.
2.2. Perceptual Interactions between Chemosensory Mental Imagery and
Perception
Researchers have long been aware that visual imagery and visual perception
interact as, for example, in the well-known Perky effect (Perky, 1910; see also
Ishai and Sagi, 1995; Pearson et al., 2008) (Note 6). For instance, in her clas-
sic early study, Cheves Perky reported that those participants who had been
asked to imagine coloured objects (such as a banana) on the wall of a pitch
black room incorporated elements of a faint colour patch that had been sur-
reptitiously back-projected onto the wall of the experimental chamber. This
phenomenon subsequently came to be known as ‘incorporation’ — though
see Craver-Lemley and Reeves (1992), for the suggestion that this may sim-
ply reflect nothing more than an attentional distraction effect. More recently,
Berger and Ehrsson (2013; see also Berger, 2020) reported that perception and
mental imagery interacted, such that, for example, imagining a visual stimulus
in one location could lead to the ventriloquism of a sound that had simul-
taneously been physically presented from a different location. At the same
time, however, in a number of these areas there is a question of discriminating
between the mental imagery of a stimulus in a particular location, say, from
merely attending to that location (or modality, Segal and Fusella, 1971 — see
also Royet et al., 2013).
Over the last 30 years or so, various researchers have investigated whether
chemosensory mental imagery interacts with chemosensory perception. For
example, Carrasco and Ridout (1993) conducted two studies, one with a set of
16 largely familiar olfactory stimuli (such as chocolate and leather) presented
to participants in pairs. Another group of participants merely imagined pairs
of odorants. In both cases, the participants had to rate the degree of similarity
between various pairs of olfactory stimuli. Multidimensional scaling (MDS)
suggested that three-dimensional solutions with similar stimulus dimensions,
such as fruitiness, strength, and familiarity, defined the psychological space of
both imagined and perceived odours, hinting at the similarity between imagery
and perception at this level (see also Sugiyama et al., 2006).
Algom and Cain (1991) reported similar interaction patterns for perceived
and imaginary mixtures of odorants. A couple of years later, Algom et al.
(1993) studied perceptual and mental mixtures of odour and taste. They used
taste–smell interactions to study olfactory mental imagery. The participants
in their study were either presented with tastant–odorant mixtures (sucrose
and orange), or asked to construct these mixtures mentally (note that both
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6 C. Spence / Multisensory Research (2024)

groups had previously been familiarized with the mixture constituents). The
participants’ task was to rate the intensity of presented or imagined mixtures.
Both perceived and mentally constructed mixtures (of different concentrations
of sucrose and orange) revealed the same, approximately additive, pattern of
(multisensory) integration. These results were therefore taken to show that
imagined smells interact with both imagined and perceived tastants in the same
way as real perceived smells do.
By contrast, when Schifferstein (1997) studied sucrose–citric acid mixtures,
mixture suppression was observed when tastants were actually combined,
while a very different pattern of results was observed when participants merely
imagined what a mixture of tastants would taste like. The latter results would
appear to suggest that the phenomenon of mixture suppression does not oper-
ate when people imagine mixtures of basic tastes. Indeed, such results led
Schifferstein to question whether Algom et al.’s (1993) earlier results might
have reflected people’s tacit knowledge of olfaction and gustation rather than
the consequences of olfactory plus gustatory mental imagery being combined.
However, to the extent that mixture suppression is a phenomenon that reflects
interactions taking place at the taste receptor level, rather than centrally (cf.
Bell et al., 1987, for a receptor-level explanation of odour mixture suppres-
sion), Schifferstein’s results should perhaps not come as such a surprise.
Stevenson and Prescott (1997) reported a study in which the participants
had to make judgements concerning the sweetness, sourness, and intensity
in memory (i.e., using gustatory mental imagery) or with combinations of
sucrose and citric acid that were actually presented. Participants’ performance
in the imagery and perception conditions was equivalent. Debriefing revealed
some explicit knowledge about the way such mixtures interact (cf. Schiffer-
stein, 1997). In a second experiment, Stevenson and Prescott investigated the
relationship between such explicit knowledge and performance on the mem-
ory task. The participants once again made the same ratings, but sampled, on
separate days; (1) capsaicin alone, (2) a flavoured, sucrose–citric acid solution
and imagined adding capsaicin, (3) the solution alone, and finally (4) the solu-
tion with capsaicin. Performance was equivalent across the semi-mental and
real mixture conditions. In particular, only sweetness was suppressed. Intrigu-
ingly, however, debriefing of the participants revealed that they thought that all
of the tastes and flavours had been affected by capsaicin, thus arguing against
Schifferstein’s tacit knowledge account in this case (see also Cain and Algom,
1997).
The evidence that has been published to date demonstrates that imagined
chemosensory stimuli (such as imagined odours and imagined basic tastes)
can interact with the perception of actual olfactory and gustatory stimuli (see
Spence, 2023b, for a review). Odour-induced taste enhancement (OITE) has
been observed in response to a remembered/imagined olfactory stimulus (see
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 7

Djordjevic et al., 2004a,b). For instance, the presentation of a strawberry odour

has been reported to enhance the perceived sweetness of a sucrose solution
whereas the presence of a soy sauce odour enhanced the perceived saltiness
of a solution of sodium chloride instead. That is, the presence of an odour
enhanced participants’ perception of the congruent taste (where congruency
is determined by the cooccurrence of the taste and smell in foods; Spence,
2022). A similar, but rather limited effect was also documented with odours
that were merely imagined: that is, imagined strawberry enhanced perceived
sweetness of water solutions, and imagined soy sauce enhanced the perceived
saltiness of weak sodium chloride solutions. In other words, imagined odours
can induce changes in perceived taste intensity that are comparable in kind to
those elicited by perceived odours.
Djordjevic et al. (2004a) reported that simply imagining a ham odour was
sufficient to reduce people’s ability to detect sucrose in solution when com-
pared to their performance when instructed to imagine strawberry odour, or
when no odour imagery instructions were provided. Note that the latter two
conditions gave rise to an equivalent level of performance, thus arguing that
it was imagining the incorrect odorant that selectively impaired performance
(i.e., rather than imagining the correct odorant somehow facilitating it). In this
study, significant effects were reported for both the perception and imagery
conditions. Meanwhile, Djordjevic et al. (2004c) demonstrated that olfactory
imagery enhanced poeple’s ability to detect a congruent odourant, whereas
visual imagery did not. In this case, the participants were presented with two
bottles, one with no odour, and the other with a threshold level concentration
of phenyl ethyl alcohol (PEA, which smells like roses) and citral, a substance
that smells like lemons, and had to try and identify the bottle that contained
the aroma.
Taken together, therefore, the evidence that has been published to date
highlights a number of similarities between chemosensory mental imagery
and chemosensory perception (e.g., Algom and Cain, 1991; Algom et al.,
1993; Baird and Harder, 2000; Carrasco and Ridout, 1993; Djordjevic et al.,
2004a,b,c, 2005; Stevenson and Prescott, 1997), as well as a few putative dif-
ferences (e.g., Barker and Weaver, 1983; Osaka, 1987; Schifferstein, 1997).

3. Behavioural Similarities: Sniffing and Salivating in Response to

Imagined Stimuli
Overt eye movements seen while people imagine particular visual stimuli have
been shown to match those expected were people to be actually looking at
a perceptual image (e.g., Laeng and Teodorescu, 2002). Such findings have
been taken to highlight the behavioural similarities that exist between visual
mental imagery and visual perception (see also Laeng and Sulutvedt, 2014).
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8 C. Spence / Multisensory Research (2024)

Researchers have also reported an overlap between perception and imagery in

the case of people’s olfactory behaviours. In particular, people tend to inhale
when imagining odours (Bensafi et al., 2003) (Note 7). What is more, they
sniff (inhale) more deeply when imaging a pleasant odour (e.g., chocolate)
than when imagining an unpleasant one (e.g., urine; Bensafi et al., 2003).
Meanwhile, Bensafi et al. (2007) reported that hedonic-specific activity in pir-
iform cortex during odour mental imagery appears to mimic that seen during
olfactory perception
At the same time, blocking people’s nostrils has been shown to reduce the
vividness of olfactory mental imagery in a group of individuals with a high
olfactory mental imagery ability (Arshamian et al., 2008). Blocking partic-
ipants’ noses so as to prevent sniffing reduced the emotional impact of the
olfactory mental image and made the imagined quality of pleasant odours
less distinct from unpleasant odours. Kleemann et al. (2009, p. 1) summarized
their findings by suggesting that: “We conclude from our results that olfactory
perception and olfactory imagery both have effects on the human respiratory
profile and that these effects are based on a common underlying mechanism”
(Note 8).
In terms of gustatory responses (or behaviours), people tend to salivate
when eating sour-tasting foods, such as a lemon, or even seeing lemons being
sliced and/or other people biting into sour foods, such as (a slice of) lemon (see
Spence, 2011, for a review). Furthermore, simply imagining tasting something
sour has also been shown to lead to salivation. For instance, Winer et al. (1965,
p. 876) verbally suggested to their participants that a solution that participants
had in their mouth (which was a pure tastant, either citric acid or sucrose dis-
solved in water) to: “Disregard the actual taste of the solution you are using
and vividly imagine, trying as best you can, all during the collection interval,
that the solution is a strong, sour/sweet solution (like the one you tasted ear-
lier)”. The results provided some (albeit weak) evidence to suggest that verbal
suggestion had a measurable impact on parotid secretion rate in response to
the actual gustatory stimuli. In particular, introducing the suggestion that water
was sour and that citric acid was tasteless, effectively elevated and depressed
the rate of parotid secretion (see also Jenkins and Dawes, 1966; White, 1978;
Wooley and Wooley, 1973).
However, a somewhat different pattern of results was reported by Drum-
mond (1995). In the latter study, 24 participants both imagined, and actually
tasted, something sour on one side of their tongue. Increased parotid salivation
was observed in both cheeks during the actual unilateral gustatory stimulation,
with a greater response documented on the stimulated side than contralater-
ally. However, although the participants were able to imagine the sour taste
more clearly after having actually experienced it, salivation did not increase
significantly during the imagery trials (either before or after exposure to the
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 9

sour taste). In fact, salivation during the imagery condition actually decreased
below baseline after exposure. More recent research has though demonstrated
that vivid visual mental imagery of what a desirable food smells like can give
rise to an increase in salivation (Keesman et al., 2016; Krishna et al., 2014 —
though see also Wang et al., 2017).
Taken together, therefore, the evidence in terms of observable behavioural
responses to chemosensory mental imagery once again shows both a number
of similarities (Bensafi et al., 2003, 2005, 2007; Jenkins and Dawes, 1966;
Kleemann et al., 2009; White, 1978; Winer et al., 1965; Wooley and Wooley,
1973) as well as a few differences (Drummond, 1995). While the majority
of the research has tended to focus on participants imagining ambient odours
(such as ham, strawberry, or soy sauce), or basic tastes (such as sour or sweet),
one might expect to find that people would exhale through their noses, and/or
engage in some sort of masticatory behaviours (i.e., over-and-above simply
salivating), when mentally imagining the taste/flavour of food. However, we
are not aware of anyone who has reported on this directly/explicitly. In their
excellent and very thorough review of olfactory imagery from a couple of
decades ago, Stevenson and Case (2005) highlight the fact that at least two
separate sensory systems, taste and smell, are involved in eating and drinking.
They write that “The distinction between taste and smell is an important one
for two reasons: First, many participants regard eating and drinking as involv-
ing only taste stimuli and do not understand the role of smell. This perception
has been exploited in some imagery experiments. Second, several of the stud-
ies described below involve both taste and smell, so it is important to bear
in mind that these are separate sensory systems”. (Stevenson and Case, 2005,
p. 245).

4. Cognitive Neuroscience of Chemosensory Perception/Mental Imagery

The cognitive neuroscience research that has been published over the last quar-
ter of a century or so has identified a number of neural substrates involved in
various aspects of chemosensory perception. The question to be addressed in
this section of the review concerns the extent to which the same areas are also
involved (i.e., activated) in the case of chemosensory mental imagery. The
anterior insula and frontal operculum (AI/FO), known as the primary gusta-
tory cortex (though see below for other discrete regions within the insula that
might also deserve this label), receive direct projections from the ventropos-
terior medial nucleus of the thalamus, and project to the orbitofrontal cortex
(e.g., Börnstein, 1940; Frey and Petrides, 1999; Shenkin and Lewey, 1944;
Small, 2010; Small et al., 1999). Intriguingly, taste-specific representations
have been discerned in human insular cortex (IC) and the overlying opercu-
lum (Chikazoe et al., 2019).
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10 C. Spence / Multisensory Research (2024)

The IC receives multimodal sensory inputs including gustatory, visceral,

thermal, and pain via the sensory thalamus. However, despite its designation,
the gustatory area in the IC involves a relatively small percentage (10%) of
gustatory neurons. In fact, the majority of the population responds to jaw and
mouth movements. The IC does, however, have dense reciprocal connections
with the amygdala, in which 7% of the neurons respond to gustatory stimuli.
Additionally, some gustatory information is sent directly to the orbitofrontal
cortex (OFC), designated as the secondary gustatory cortex. The OFC is
believed to process the identification of food, and to represent information
concerning satiety and food preference. Recalling the taste of food has been
reported to give rise to an asymmetric prefrontal cortex activation (Okamoto
et al., 2011 — see also Okamoto et al., 2006).
Neuroimaging research from Levy et al. (1999b) indicated that the brain
regions activated for mentally imagined tastes are consistent with those for
actual taste perception. However, using the technology available at the time, it
was simply not possible to distinguish the regional brain localization for salt
or sweet taste memories. However, the story has now changed, given the dra-
matically higher field-strength scanners (7T) that have become available for
research in recent years and which offer much better spatial resolution (see
below). In a functional magnetic resonance imaging (fMRI) study, Japanese
volunteers were instructed to think about pickled plums (umeboshi), a tradi-
tional Japanese food with a strong and sour taste (Kikuchi et al., 2005). A
significant increase in neural activation was documented in the right insula,
the bilateral opercula, the bilateral orbitofrontal cortices, and the left Broca’s
area. Although activation in the primary gustatory area (insula) was very weak
and detected only unilaterally, the pattern of neural activation was nevertheless
similar to that observed during taste perception. Here, it is interesting to note
that like in a number of other studies, the objective existence of chemosensory
mental imagery (specifically mentally imagining tastes) is taken to come from
activation of the relevant brain areas rather than necessarily from the intro-
spective reports of the participants involved (see also Djordjevic et al., 2005).
As the authors themselves put it: “If the cerebral activation pattern during con-
centration on strong-tasting foods is similar to that of taste perception, we may
assume that the human brain has the potential for taste imagery. In particular,
the aim of their functional magnetic resonance imaging (fMRI) study was to
investigate whether the cerebral activation while participants concentrated on
strong-tasting food was similar to that of taste perception” (Kikuchi et al.,
2005, p. 281) (Note 9).
There are two other points to make about Kikuchi et al.’s (2005) intrigu-
ing study. First, the participants were not explicitly asked to mentally imagine
the taste (or flavour) of sour plums. Rather they merely looked at three pick-
led plums while repeating the phrase: ‘Pickled plums are sour’ in Japanese
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 11

to themselves. So while the participants stated that they could produce taste
imagery, it is unclear whether they actually did so while they were lying
in the brain scanner. Second, the researchers were unable to rule out the
impact of salivation, either on brain activity, or on the possibility of imaging
the taste of this strong-tasting food (Note 10). Note here that other research
has demonstrated that simply reading the word ‘cinnamon’ activates olfactory
brain regions (González et al., 2006), while reading the word ‘salt’ can acti-
vate gustatory brain regions (Barrós-Loscertales et al., 2012 (Note 11) — see
also Pomp et al., 2018).
Turning now to the case of olfaction, Wiesmann et al. (2001, p. 237) note
that: “Olfactory information is projected from the olfactory bulb to the pri-
mary olfactory cortex, which is composed of the anterior olfactory nucleus,
the olfactory tubercle, the piriform cortex, the amygdala, the periamygdaloid
region, and the entorhinal cortex. From there, the primary olfactory cortex
projects to secondary olfactory regions including the hippocampus, ventral
striatum and pallidum, hypothalamus, thalamus, orbitofrontal cortex, agranu-
lar insular cortex, and cingulate gyrus” (see also Boot et al., 2024).
Primary piriform cortex (PC) is typically activated when non-experts try
to imagine an odour (e.g., Djordjevic et al., 2005; Kettenmann et al., 2000;
Levy et al., 1999a). That said, Royet et al. (2013) have argued that the neu-
ral activity that has been reported in the PC where non-experts try to imagine
olfactory stimuli might have other causes, such as the neural activity that is
associated with sniffing, attention/expectation, and/or other top-down influ-
ences, and crossmodal associations (Karunanayaka et al., 2015). Sniffing also
results in PC activation, perhaps linked to somatosensory stimulation that is
caused by airflow through the nostrils (Koritnik et al., 2009; Sobel et al., 1998).
A meta-analysis of human neuroimaging studies conducted by McNor-
gan (2012) identified a number of brain areas that were modality-specific, as
well as several others that were modality-general (see also Daselaar et al.,
2010; Olivetti Belardinelli et al., 2004 — see Note 12). In particular, over-
lapping brain regions that were activated during olfactory mental imagery and
olfactory perception included the primary olfactory cortex, anterior cingulate
cortex, hippocampus, insula and superior parietal lobule. Olivetti Belardinelli
et al. (2009), meanwhile, reported greater activity of sensory–specific cortices
in high- vs low-vivid participants for visual, gustatory, kinesthetic, tactile, and
the somatic modalities, but not for either auditory or olfactory imagery. Men-
tal imagery generation in different modalities activates sensory-motor areas
(Palmiero et al., 2009). Research from Han et al. (2022) suggests that different
neural strategies may sometimes be used by those individuals who are high vs
low in olfactory mental imagery abilities (by self-report questionnaire). The
latter group had stronger brain activation of the supplementary motor area
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12 C. Spence / Multisensory Research (2024)

and the superior frontal cortex. Variation in the activation levels and connec-
tion strength between olfactory areas has also been reported by Bensafi et al.
(2013).
The neural mechanisms underpinning olfactory mental imagery can appar-
ently also be activated by verbal descriptors [‘floral bouquet’; e.g., in the
context of scent marketing (Meng et al., 2020); as well as by non-figurative
visual cues (Hossu et al., 2024)]. Importantly, however, odour coding in human
PC is category-specific (Howard et al., 2009), meaning that olfactory informa-
tion processing does not break down odour objects into specific features, like
colour, motion, and shape (as has been documented in the visual modality,
say).
Viewing pictures of appetizing foods appear to activate the gustatory cor-
tices for taste and reward, namely the right insular/opercular and left OFC
(Simmons et al., 2005). Research from Avery et al. (2021) demonstrated that
the IC activity when viewing images of foods can be decoded by the inferred
taste quality associated with those foods. In fact, subsequent research from
Avery et al. (2023) suggests that there may be a common neural code for rep-
resenting imagined and inferred tastes. The participants in the latter study were
presented with the words ‘SUGAR’, ‘SALT’, ‘LEMON JUICE’ and ‘WATER’
and were instructed to “imagine to the best of your ability that someone has
placed a spoonful of that substance on your tongue” (Avery et al., 2023, p. 6).
In another task, the same participants saw images of dominantly sweet, salty,
and sour foods. High-field fMRI revealed that imagined tastes elicited activ-
ity in the bilateral dorsal mid-insula which was reliably similar to that elicited
when viewing images of food with the corresponding dominant taste quality,
thus suggesting a common neural code for representing taste quality regardless
of whether explicitly imagined or automatically inferred when viewing food.
However, while the involvement of chemosensory cortical areas in mental
imagery does indeed suggest that those mental images are depictive or percep-
tual in nature, one possibility that should be considered is that imagined and
experienced tastes or smells may be represented in those same regions using
a different format, or at the least within different neural populations. Avery
et al. (2021, 2023) attempted to address this issue of format by using cross-
task decoding analyses. Avery et al. (2023) used just such an analysis in order
to demonstrate that the mid-insula activity evoked by the mental imagery of
taste and the activity evoked by viewing food pictures shared a common for-
mat. By contrast, when the same type of analysis was performed in Avery et
al.’s (2021) study, the results suggested that while they both activate that same
mid-insula region, respondes to food pictures responses and those of experi-
enced tastes (which were delivered during scanning) did not appear to share a
common format.
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 13

Japanese researchers prompted sweet, sour, salty, and bitter-tasting foods

by presented words (e.g., ‘lemon’) or else pictures of foods dominantly asso-
ciated with those basic tastes. Participants were instructed “to recall the taste of
the presented food or beverage” (Kobayashi et al., 2004, p. 1273). During the
perception task, orange, apple, grapefruit and grape juice were delivered to the
participants’ mouths. The left insula was predominantly activated when partic-
ipants engaged in gustatory mental imagery. The middle and superior frontal
gyri were also involved in the generation of gustatory ‘hallucinations’, but
were not activated by gustatory perception. Kobayashi et al. went on to sug-
gest that these regions in the frontal cortex may mediate the ‘top-down’ control
involved in retrieving gustatory information from long-term memory storage.
The suggestion was that the source of the top-down signals may be in middle
and superior frontal gyri, and that these signals may affect the neural activities
in the insula, orbitofrontal, and precentral gyri. A few years later, magnetoen-
cephalography (MEG) research from Kobayashi et al. (2011) demonstrated
that the PFC was indeed activated prior to the IC when the participants in their
study engaged in gustatory mental imagery (Note 13) — what the authors
describe as “the “top-down” modulation of taste perception” (Kobayashi et
al., 2011, p. 1).

4.1. Unanswered Questions Concerning the Mental Imagery of Flavours

What happens neurally when people image the flavour of food and drink
(assuming, that is, that that is something they are able to do)? In terms of mul-
tisensory flavour perception, superadditive opercular activation in response to
congruent food flavour is mediated by enhanced temporal and limbic cou-
pling (Seubert et al., 2015). One would therefore imagine that flavour mental
imagery should also give rise to enhanced coupling between these brain areas
(see also Suen et al., 2021). However, we are not aware of anyone who has
conducted a similar analysis to that reported by Seubert et al. but for imagined
odours, tastes, and imagined flavours. One might also want to know about
the cortical processing of pungent trigeminal stimuli, such as, for example,
delivered by chilli, black pepper, or ginger (see Spence, 2018, 2023a, 2024).
Note that trigeminal inputs are included as part of flavour perception by many,
if not necessarily all, researchers (ISO, 1992/2008). Relevant here, Japanese
researchers have reported that tasting and ingesting capsaicin (the pungent
active that gives chilli its burn) activates the ventral part of the middle and
posterior short gyri (M/PSG) of the insula which is known as the primary gus-
tatory area (though see below on the several regions within the insula that have
been described as the primary gustatory cortex), suggesting that capsaicin is
recognized as a taste (Kawakami et al., 2015). One might also wonder whether
chemesthetic oral experiences which are in some sense metameric (cf. Ravia
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14 C. Spence / Multisensory Research (2024)

et al., 2020), such as of heat or cooling, which may be delivered by ther-

mal or chemesthetic stimuli, recruit the same network of brain areas when
participants directly perceive vs merely imagine such complex chemosensory
stimuli.
It is, though, possible to imagine that attending to the flavour of food might
not necessarily activate exactly the same network of brain areas activated as
when actually tasting food, given that people consider the flavour of food and
drink to originate from the oral cavity, and explicitly and ubiquitously refer
to it as ‘taste’ (see Auvray and Spence, 2008; Prescott, 1999; Spence et al.,
2015), due to the phenomenon of oral referral (Small, 2010; Spence, 2016).
It is further worth noting that our attention is typically directed to the oral
cavity while eating (i.e., when savouring the ‘taste’ of food; Spence, 2019;
Stevenson et al., 2011), not to the nose where the majority of the information
is actually transduced (see Spence, 2015). Relevant here, modulation of early
gustatory cortex (IC and frontal operculum), and perceptual enhancement has
been reported a result of attention being directed to taste (e.g., Marks and
Wheeler, 1998a,b; Spence, 2019; Veldhuizen et al., 2007; Zelano et al., 2005,
2011).
Finally, here, and coming from a rather different research direction, Bar-
ron et al. (2013) assessed the neural changes in the hippocampus and medial
prefrontal cortex that were associated with exposure to a novel combination
of flavours/ingredients, i.e., such as avocado and raspberry smoothie or tea
jelly. In this case, the researchers concerned were interested in the question
of whether the co-activation of multiple relevant memories might provide a
training signal to the valuation system that allows the consequences of new
experiences to be imagined and acted on. However, it would seem at least
worth considering whether the performance of such a task might require the
manipulation of mental imagery for flavour.
4.2. Interim Summary
Taken together, the cognitive neuroscience research on the neural substrates of
chemosensory perception and chemosensory mental imagery appear to show
a great deal of overlap in terms of the networks of primary and secondary
areas that are involved, though with the patterns of connectivity seemingly
reversed — bottom-up in the case of chemosensory perception and top-down
in the case of chemosensory mental imagery. The research that has been pub-
lished to date has primarily focused on taste (or perhaps flavour — Note 14)
and olfaction (presumably orthonasal olfaction, but as has been mentioned the
distinction between orthonasal and retronasal olfaction has not been devel-
oped for the case of mental imagery in quite the way that it has been in the
case of chemosensory perception; e.g., Rozin, 1982; Small et al., 2008; Wil-
son, 2021). As such, it will be intriguing to find out whether a similar pattern
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 15

emerges when multisensory flavour mental imagery is compared to multisen-

sory flavour perception (cf. Nanay, 2018). Finally, here, another aspect of our
multisensory experience of food and drink that is often very important con-
cerns mouthfeel. While mental imagery questionnaires have recently started
to incorporate questions about this attribute of food experience (Note 15), it
would be interesting to know more about the degree of overlap in terms of the
relevant brain areas in the case of both perception and imagery.
One important point to consider here concerns the involvement of chemo-
sensory cortical areas in chemosensory mental imagery. Multiple studies that
have been discussed thus far claim to have documented activation of the pri-
mary gustatory cortex. However, closer examination reveals that those studies
appear to point to different areas of the insula, such as the AI/FO (dorsal ante-
rior insula; e.g., Small, 2010; Small et al., 1999), the dorsal mid-insula (Avery
et al., 2021, 2023), and the ventral anterior insula in Kawakami et al. (2015).
As pointed out by a helpful reviewer, a large body of research on the functional
organization of the insula shows that these regions have different functional
roles across a variety of cognitive tasks (for instance, see Kurth et al., 2010),
and differential connectivity to the wider brain (e.g., Cauda et al., 2011; Kelly
et al., 2012). As such, it does not seem likely that these different areas have
equal claim to the designation of ‘primary gustatory cortex’. This obviously
bears on the question of the involvement of these areas in chemosensory men-
tal imagery. This point is likely to be especially relevant when it comes to
flavour mental imagery, given that two of these insula regions (ventral anterior
and dorsal mid-insula) have elsewhere been implicated in the integration of
taste and smell to produce flavour (see Seubert et al., 2015; Small, 2012).

5. Experience/Expertise and its Effect on Chemosensory Mental

Imagery/Perception
The chemical senses also an area where expertise is known to exert a profound
influence. At the perceptual level, Sezille et al. (2014) assessed the hedo-
nic appreciation and verbal description of a range of pleasant and unpleasant
odours in untrained, trainee cooks, flavourists, and perfumers. They reported
no group level differences in hedonic ratings for pleasant and unpleasant
odours. On a verbal level, smell descriptions were richer and did not refer
to pleasantness in experts compared to untrained participants who used terms
that referred to the odour sources (e.g., candy) accompanied by terms referring
to odour hedonics (see also Gilbert et al., 1998). Perfumers, as a representative
category of olfactory experts (Gilbert et al., 1998), are significantly better at
olfactory imagery (in terms of generating vivid imagery) than novices.
Olfactory experience has been found to induce functional reorganization
in brain regions involved in odour imagery in perfumers (Plailly et al., 2012)
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16 C. Spence / Multisensory Research (2024)

(Note 16). Perfumers exhibit both cortical and functional changes in the pri-
mary and secondary olfactory areas (Delon-Martin et al., 2013). Specifically,
expert perfumers exhibited less activation in the hippocampus and PC during
olfactory imagery as their level of expertise increased (presumably because
less effort is required to generate the olfactory images). As a group, perfumers
showed an increase in grey-matter volume in the bilateral gyrus rectus/medial
orbital gyrus (GR/MOG), an orbitofrontal area that surrounds the olfactory
sulcus. Additionally, grey-matter volume in the anterior PC and left GR/MOG
was found to be positively correlated with experience in professional per-
fumers. These findings led Delon-Martin and colleagues to conclude that the
extensive olfactory knowledge acquired as a result of prolonged olfactory
training leads to the structural reorganization of olfactory brain areas. Accord-
ing to Royet et al. (2013), the strength of connection between areas may also
vary as a function of the level of expertise of odour experts. Here, it is inter-
esting to note that when perfumers create a olfactory mental image, the right
middle frontal gyrus, a key region in the neural signature of retrieval strongly
co-activated with olfactory and memory regions but not in non-experts (Lep-
age et al., 2000). Intriguing changes have also been reported in the case of
expert wine-tasters (e.g., Carreiras et al., 2024).

6. Conclusions
The study of chemosensory mental imagery has proved particularly challeng-
ing because of the profound individual differences that have been reported
in the vividness of, for instance, olfactory mental imagery (e.g., Arshamian
and Larsson, 2014; Djordjevic et al., 2004c; Gilbert et al., 1998; Zhou et al.,
2022). At the same time, the majority of those researchers who have attempted
to study people’s mental imagery abilities for taste (gustation) have actually
mostly been studying flavour mental imagery. Nevertheless, there exists a
body of human psychophysical research showing that chemosensory mental
imagery exhibits a number of similarities with chemosensory perception. Fur-
thermore, the two systems frequently interact with one another (see Spence,
2023b, for a review). In this narrative historical review (Ferrari, 2015; Furley
and Goldschmied, 2021), the many similarities between chemosensory per-
ception and chemosensory mental imagery at the introspective, behavioural,
psychophysical, and cognitive neuroscience levels in humans are highlighted,
as well as the differences, such as there are.
The latest neuroimaging evidence show that many of the same brain areas
are engaged by chemosensory mental imagery as have previously been docu-
mented to be involved in chemosensory perception. That said, the pattern of
neural connectively appears to be reversed between the ‘top-down’ control of
chemosensory mental imagery and the ‘bottom-up’ control seen in the case of
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 Multisensory Research (2024) DOI:10.1163/22134808-bja10130 17

chemosensory perception (Kobayashi et al., 2011). However, as shown by the

research published by Avery et al. (2021, 2023), just because the same neural
areas are activated that does not mean that the format for imagined and per-
ceived chemosensory stimuli is necessarily always the same. At the same time,
however, there remain a number of intriguing questions. For instance, it is cur-
rently unclear to what extent it makes sense to try and distinguish between
orthonasal and retronasal olfactory mental imagery. It also remains unclear
whether mental imagery for flavour, which most people not only describe but
also perceive to be the ‘taste’ of food and drink, is capable of reactivating the
entire flavour network in the human brain (e.g. including retronasal olfaction).
These, then, both represent intriguing areas for future research in the study of
chemosensory mental imagery.

Notes

1. Though, as we will see later, taste typically refers to people imagining

flavours rather than basic tastes.
2. One also needs to be sensitive to the suggestion that an investigator’s intu-
itions and introspections about imagery, in particular the vividness of their
own mental imagery experience, may shape their theoretical views (see
Reisberg, Pearson and Kosslyn, 2003). In other words, rather than simply
accepting the fact that there may be some profound individual differences
in the vividness of olfactory mental imagery, researchers with presum-
ably weak olfactory mental imagery typically appear to assume that those
whom they study must experience the world similarly.
3. Köster (2002) uses the term ‘olfactorise’ to mean ‘the capacity to imagine
smells’.
4. It is sometimes hard to know the extent to which flavour mental imagery is
actually involved when the participants in cravings studies are instructed
to: “Imagine you are eating your favourite food” (Harvey et al., 2005).
5. Though, what people describe as gustatory most likely actually refers to
flavour mental imagery.
6. Though note that in the case of the latter two studies this is more retaining
an image in memory than the spontaneous generation of something from
within, as in the original Perky (1910) effect.
7. But not when mentally imagining a visual stimulus.
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18 C. Spence / Multisensory Research (2024)

8. Looking to the future, it would be interesting to know whether olfactory

mental imagery could be harnessed to deliver some of the well-being bene-
fits that exposure to actual pleasant ambient odours has been shown to have
(Boot et al., 2024; Bromberg and Schilder, 1934; Spence, 2020; Weber and
Heuberger, 2008; see also Schlintl et al., 2022).
9. One can see such a position as responding to the scepticism expressed by
some researchers about the meaning, or usefulness, of questionnaire data
(see Ahsen, 1995). Or, as Crowder and Schab (1995, p. 94) once succinctly
put it: “experience is not evidence”.
10. As the authors note: “Most Japanese people salivate just on looking at
pickled plums because of their long familiarity with this sour food”.
(Kikuchi et al., 2005, p. 281).
11. Areas activated in response to reading taste words included the anterior
insula, frontal operculum, lateral orbitofrontal gyrus, and thalamus among
others. Given that these areas comprise the primary and secondary gusta-
tory cortices, the researchers concluded that the meaning of taste words
is grounded in distributed cortical circuits reaching into areas that process
taste sensations.
12. This, note, is one of the very few studies that describes odours and flavours
(i.e., rather than tastes).
13. The participants in this study were asked to recall the taste (sweet, sour,
bitter, or salty) of flavourful stimuli (five stimuli were associated with each
taste, as per the authors’ earlier study).
14. Note here that what is described as mental imagery for taste (or gustation)
is really more likely to be mental imagery for flavour, given that something
like 75–95% of what people think of as taste really comes from the sense
of smell (see Spence, 2015).
15. Hazebroek and Croijmans (2023), for instance, recently conducted a study
assessing the strength of people’s mental imagery for the aroma, mouth-
feel, and taste of coffee.
16. Note that at the other extreme, those suffering from olfactory loss show
a decline in activity in olfactory areas that correlates with the duration of
the loss (Flohr et al., 2014).

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