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Why Do We Eat What We Eat?

Chapter · May 2015

DOI: 10.1093/med/9780199330454.003.0002

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 To appear in: N. Avena (Ed.) Hedonic Eating: How the
Pleasurable Aspects of Food Can Affect Appetite. New York:
Oxford University Press.

2
Why Do We Eat What We Eat?

Kevin Myers

Every August on the rugged high coast of Sweden, the balmy fragrance of seashore and
spruce forest is overtaken by a pungent mixture redolent of rotten eggs, rancid butter,
sweat, and ripe fish. It hisses from bulging cans that are pried open for the annual festival
of Surströmming, a soured herring beloved in the region. “Soured” is a bit euphemistic.
Aggressive anaerobic fermentation swells the cans with hydrogen sulfide and butyric and
propionic acids. Tourists snap photos of uncontrollable grimaces (or worse) triggered by
their cautious first bites. In 1991, a German court hearing a landlord–tenant dispute ruled
the aroma of Surströmming so offensive that it constitutes a nuisance beyond what
neighbors could be reasonably expected to tolerate. Yet many Swedes joyfully gobble up
the fish at summertime parties in an animated affirmation of neighborly fellowship and
national pride.

At the same time of year, market stalls throughout Southeast Asia are piled high
with a heavy, melon-sized fruit called durian. The thick husk studded with spikey thorns
offers visual warning of what is to be found within. Though descriptions of its intense,
complex flavor usually combine terms like sweet, custardy, and caramel with rotten
onion, garbage, turpentine, sewage, and putrid meat, many people snack on it daily and
concoct it into sauces, cakes, pastries, ice cream, and drinks. Particularly rare cultivars
are featured in dishes at exclusive restaurants in Bangkok and Kuala Lumpur. But not
everyone is enamored with durian. It disgusts enough people that passenger airlines
forbid carrying it on board. Signs in the Singapore metro system prohibit smoking,
explosives, and durians.
In my own small town—the very picture of Americana, nestled in the heart of
Pennsylvania farm country—I always offer my out-of-town guests (especially colleagues
interested in the psychology of food preferences) our peculiar regional specialty.
Scrapple is a congealed breakfast loaf made by rendering the bits left over after hog
butchering. It reflects the early Pennsylvania German settlers’ rather utilitarian approach
to nutrition. In years, I do not recall ever watching an adult taste scrapple for the first
time who liked it. But to many of us who grew up in this area, it is the taste of home, and
we miss it when we are away for too long.
What accounts for these striking differences in how individuals evaluate the tastes
and flavors of foods? The tremendous diversity in food preferences reflects a
fundamental feature of the sensory-evaluative processes that govern food choice:
experience-dependent plasticity. Humans (and the rats and mice we use to model the
biopsychological building blocks of our behavior) have migrated to occupy nearly every
ecological zone on the planet. Thus individuals are born into habitats containing such
dissimilar collections of flora and fauna that it would be impossible for any meaningful
food-recognition and -evaluation mechanism to be instinctively wired in by our genetic
instructions. Sensitivity to experience early in development brings preferences roughly in
line with what one can expect to find routinely in one’s macro-environment, or the “food
system,” which Rozin describes as the “massive, behind the scenes” cultural
prearrangement from which one is even capable of choosing.1 Each individual’s unique
history of experiences shapes his or her affective responses to the tastes, flavors, textures,
and appearance of various foods.
Plasticity in food preferences reflects our evolutionary history as generalist
omnivores. There are no nutritionally complete foods, so variety is essential. A few basic
taste sensations from sugars, starches, fats, and proteins only allow rough assessment of
caloric content. Even then, people are fairly inaccurate at judging the energy density of
foods.2,3 There are no universal sensory labels for many essential nutrients, and many
potential toxins exist at low levels, so relying on innate taste recognition would be
infeasible. Experience with a food is necessary to transform sensory properties that were
not innately attractive into “labels” for the food’s quality.
The most obvious way that liking for a food affects its intake is by influencing
which foods are selected, especially when procurement costs are relatively low. Meta-
analysis shows that across a wide range of foods, experimental manipulations of
palatability predictably change relative preference. For any two foods, the difference
between palatability ratings generally predicts the difference in how much participants
will eat of each one.4 But it would be a mistake to view palatability as the sole
determinant of preference. Highly liked foods are often eaten in small quantities or
avoided altogether out of concern for health, weight gain, or for complex ethical or
religious reasons. Thus it is essential to recognize that “liking” is but one influence on
choice and intake. Moreover, causal links between liking and consumption are almost
always bidirectional. In some situations, availability or cost strictly dictates what is to be
eaten, and in turn, sheer familiarity with those foods establishes liking.
A second way hedonics influence intake is through the dynamic changes in
palatability during a meal. Initial exposure to sight, smell, or taste triggers a sensitizing,
“appetizer” effect in proportion to palatability,5,6,7,8 including preparatory digestive
responses and accelerated eating rate. But during a meal, food typically becomes
increasingly unpalatable. This reflects in small part a physiological feedback effect from
gut satiation signals9,10; gut signals, however, are not absolutely necessary. Oral exposure
to a food’s sensory properties causes specific, habituation-like decreases in the hedonic
evaluation of the flavor, and also willingness to work for more of that food.11,12 The
palatability decrement becomes one causal factor in meal cessation, and accumulates
across meals when the same food is eaten on successive occasions.13 Thus hedonic
evaluation comes to influence one’s choice of foods and also influences the dynamic
control of motivational switching between eating and other behaviors.

The Importance of Hedonics in the Psychology of Twenty-first

Century Eating
Choice and intake are intrinsically bound up with hedonic evaluation of food’s sensory
attributes—the pleasures of tasting, smelling, and feeling food as it is consumed.
Understanding hedonic influences becomes especially important now that technologized
food production brings an increasing variety and ease of access to foods that are
specifically designed with sensory appeal in mind. It will always be difficult to elucidate
a causal role of palatability per se in eating patterns, since palatability is neither a
physical property of a food nor a fixed response tendency of the individual, but rather an
interaction between the food, the external context, and internal signals for current
physiological state.14,15,16 Nonetheless, virtually all acts of eating generate some degree of
sensory pleasure.

This is illustrated by the fact that many people will pay an equivalent price for
artificially sweetened versions of liked foods or drinks. Operant studies show a powerful
reinforcement effect of non-nutritive sweeteners,17 which can exceed that of cocaine,18
demonstrating that caloric consequences are not necessary for sweet taste to maintain
food-seeking behavior. Since the spending on artificially sweetened products like diet
soda is not offset by compensatory increased spending on calories in ordinary foods,19 a
behavioral economic perspective suggests that people, at least “health conscious” ones,
are often willing to pay a caloric cost to obtain sweetness per se.
The importance of sensory pleasure is further illustrated by consequences of
removing it. Nutrients infused intravenously or via nasogastric tube (therefore not tasted,
chewed, or swallowed) are generally less satisfying. In studies of healthy volunteers,
tube-fed participants still consume near-normal meals when permitted to do so.20 In long-
term clinical cases, tube-fed individuals become preoccupied by food cravings.21 Healthy
volunteers who consumed nothing for several days except a sweet, nutritionally complete
drink experienced cravings caused by lack of sensory variety. 22 During a chocolate
craving, it is only the tasting, smelling, and eating of chocolate, rather than its post-
ingestive effects, that satisfy the craving.23 Thus, not only will people pay a significant
cost to obtain taste pleasure, but nutritional repletion without oral stimulation is
unsatisfying, and sensory deprivation can powerfully stimulate interest in food and evoke
mental imagery of food’s desirable sensory characteristics.
Understanding hedonic processes is necessary for insights into why individuals
are more or less affected by the modern food environment. People differ in their preferred
amount of sensory stimulation in daily life, known as “sensation-seeking.” But sensory
novelty, complexity, and pleasure can come from both food and non-food sources, and
people differ in the importance of eating as their main source of that sensory stimulation.
Some people describe eating as one of the most pleasurable activities available, while
others would gladly opt to take daily nutrition pills instead of eating at all.24 This is likely
to be a developmentally plastic trait, since it varies across cultures in a manner consistent
with prevailing local attitudes about diet.25
Individual differences in hedonic processes have become a main focus of research
because of the provocative but sometimes contradictory evidence linking food hedonics
to overeating and obesity. Although it is undoubtedly true that a typical person is inclined
to eat more when a given item of food is made to taste better, it is not clear whether or
not obese people perceive food as tasting better in general, nor whether such differences
play a causal role in obesity. Obese individuals may experience more pleasure from
sweetness than do lean individuals, especially when compared to pleasure derived from
non-eating activities.26 Other methods have linked obesity to sensory preferences for fat
and sweet-fat mixtures27,28 or inversely to liking fruits and vegetables.29 But in contrast,
there is also evidence that obese eat more because they experience less reward from
food.30,31 It also needs to be determined whether differences in subjective pleasure from
food act as a preexisting causal factor in overeating, or instead are established by a
history of overeating. Unravelling the complex links between hedonic evaluation, diet
history, and weight status is now a major focus of research on the obesogenic effects of
the modern food environment.

Stimulus Attributes in Food Hedonics

Sweetness and Fat
Scientific and everyday conversations about food enjoyment and preference usually begin
with sweetness and fat, and for good reason. As biologically significant macronutrients,
sugars and fats are laden with intrinsic positive value and contribute directly to
palatability. Sweet–fat combinations have especially potent appeal to many.27,32

The pleasure of sweet taste is generally regarded as “hard-wired” at the start. For
newborns, sugar stimulates mouthing and characteristic hedonic facial expressions,33,34
and has calming, pain-relieving effects.35 This is seen prior to any nursing experience,36
and even in infants delivered prematurely.37 Infant rats respond to sweet taste with
vigorous mouthing and swallowing, with a linear concentration-response function evident
by the time they are six days old.38,39 Even in the womb, embryonic rats respond
positively to saccharin.40 While response to sweet tastes can change with experience,
prior experience eating sweet foods does not appear to be necessary for infants to regard
sugar as palatable.
Liking for sweetness is highest during childhood and declines throughout
adolescence.41 By adulthood, individual differences are evident both in sensitivity to
detecting sweetness and in liking different amounts of sweetness,42 due in part to early
feeding experiences. For example, higher average sweet-liking among African-
Americans has been linked to the cultural practice of providing infants with sweetened
water, as that practice increases sugar-liking in infancy and later in childhood, regardless
of ethnicity.43,44 Parent–child interactions later in childhood also have an effect, ironically
such that stricter parental restriction of sweets is associated with children’s subsequent
choice of higher sweetness levels.45
Despite the presumably innate origins of sweet-liking, large cultural and
geographic differences are evident. In economically advanced countries, sugar
consumption per capita varies nearly twenty-fold, ranging from 3.3 kilograms annually in
South Korea, to 64.4 kg in Israel.46 The United States (33.1 kg) is near the median, and
East Asian countries (Japan, Hong Kong, Taiwan) are consistently the lowest. Cross-
cultural patterns in taste-liking are generally not a consequence of basic differences in
taste detection or intensity perception, but rather in preference,47 and tend to parallel the
frequency of exposure to foods.48 Sweet-liking is largely food-specific, since children
learn through exposure that sweetness is appropriate in some foods but not others.49
 Like sugar, fat also contributes to food palatability. The sensations that make fat
palatable are less well defined, but include gustatory, olfactory, and somatosensory
inputs.50,51,52 Rats avidly consume mineral oil emulsions or diets adulterated with
petroleum jelly,53,54,55 which shows subjectively positive properties independent of
nutritive value, much like the non-caloric sweeteners. Also like sweetness, there are wide
individual differences in fat-liking. This becomes evident in childhood and may be linked
to children’s nutritional or weight status. Children’s fat intake correlates with liking, but
perhaps more importantly, with their parents’ adiposity.56 This suggests a possible link in
the familial transmission of weight status.
It is sometimes assumed that the perceived palatability of fat is innate, like
sweetness, but the evidence for that is rather ambiguous. Animal studies that can
carefully monitor the development of sweet and fat responses show that they differ in
several important respects. First, fat palatability emerges much later in the pre-weaning
period53,57,58 and increases with maturation, unlike sweet-liking, which is initially high.
The central dopaminergic circuits that mediate intake stimulation by sugars and fat in
adulthood are sensitive to sucrose at a much earlier age than to fat.59,60 Also, the shape of
the concentration-response function differs substantially from that for sugars. In sham-
feeding studies of oral reward, intake of sweet solutions increases monotonically as a
function of concentration. In contrast, the function for corn oil has an inverted-U shape.55
Another curious feature is that infant rats are capable of detecting and perceiving
fat well before the time they start treating it as palatable. Ten-day-old pups, which do not
hedonically respond to fat ingestion, do respond to fat taste in other ways, such as
ultrasonic vocalization61 and anti-nociceptive responses.62 Even younger pups learn to
prefer odors paired with intraoral infusion of corn oil.63 Thus, unlike for sugar, which is
palatable as early as sweet can be tasted, for infants (at least in animal models used for
these studies), there may be a period where fat is detectable and possibly rewarding in
some respects, but not especially palatable.
Ultimately, while sugar and fat are calorie sources that form the foundation of
food-liking, the evidence all suggests that responses to sugar and fat have different
origins, and that fat palatability may depend more on early experience. Since infants rely
on nursing as their sole source of nutrition and hydration, it is feasible that ubiquitous
experience may be critical in establishing fat palatability.

Flavor
Something puzzling about food choice and hedonics is that, despite claims about the
prepotency of sugar and fat palatability’s stemming from our evolutionary history, the
fact is people almost never routinely consume plain sugar or fat, either alone or in
combination. The closest thing—cake frosting—may feature in binge eating episodes, but
is never routinely consumed on its own by healthy people, and is actually made more
attractive by adulteration with vanilla or other flavorings that have no intrinsic biological
significance. As a treat, white chocolate (almost entirely sugar and fat) is considered a
poor substitute for true chocolate, which differs physically only due to cocoa, which adds
complexity of flavor and aroma but has no intrinsic nutritional value to justify its appeal.
In fact, pure cocoa on its own would be initially repellent to naïve individuals due to its
inherent bitterness. Considering sweet beverages like sodas or fruit drinks, without the
proprietary blend of flavorings that turns carbonated sugar-water into cola, or the squirt
of lemon that makes it lemonade, most consumers would just opt for “plain” water or
nothing at all, not unflavored sugar-water. Thus it is genuinely insufficient to view
intrinsic palatability provided by sweetness and fats as sufficient to drive preference.
There needs to be more. People like and seek the complex contributions of flavor in
foods.

Flavor refers to the perceptual experience created by synthesis of the simpler oral
sensations—the basic tastes combined with odors, and also the trigeminal touch senses in
the mouth which convey a food’s textural, temperature, and chemical irritants like hot
pepper. The olfactory system can detect and discriminate an extraordinarily wide range of
distinct odors,64 far outnumbering the few basic tastes, and thus the volatile odor
molecules released from a food contribute much to its unique flavor, and experimental
study of flavor perception usually manipulates odorants. Odors can be detected from a
distance, making them useful for food identification, but they also contribute to flavor
during chewing and swallowing by traveling a retronasal route. When oral sensations and
odors synthetically combine, it creates the perceptions that people mistakenly think of as
“taste,” because the brain refers the experience to the mouth.
This perceptual confusion about the taste vs. olfactory components of food flavor
demonstrates the top-down nature of flavor perception that results from learning. Odors
commonly encountered in the context of food flavorings are typically perceived as having
taste attributes. For instance, the vanilla and strawberry odors seem “sweet,” and lemon
“sour,” although nothing inherent in the odorants triggers coding of those taste
characteristics. Adding one of those “sweet” odors as a flavoring in a sugar solution
causes people to perceive the solution as significantly sweeter. Yet “non-sweet” odors of
similar intensity do not produce this effect, nor do “sweet” odors enhance the perceived
intensity of a non-sweet solution like salt.65,66 Exposure to some odor-based food flavors
is so consistent that it is probably impossible to experience familiar food odors as they
“truly” are, because these inevitably and automatically call up compound memory
representations of other food attributes.
In contrast to the well-established hedonic or aversive responses to basic tastes
like sweetness and bitterness, most food flavors are not strongly attractive or aversive
until experience makes them so. Since odors are a primary constituent of food flavor, the
development of olfactory perception is instructive. Newborns have a fairly well-
developed capacity for olfactory perception (see reviews67,68) and do show autonomic
reactions and behavioral orienting to many odors. But they do not typically respond with
the same behavioral indicators of hedonic valence to food-related odors that adults would
give, even though infants are clearly capable of showing such reactions to basic tastes.69
When infants encounter odors like vanilla in novel situations, responses are correlated
with prior exposure,70 suggesting the critical role of early experience in establishing their
attractiveness. Positive and negative hedonic reactions do begin to emerge in infancy, but
not until later in childhood does the intensity and valence of hedonic responding resemble
that of adults.68,71
Though largely experience-dependent, there are some direct genetic contributions
to individual differences in food evaluation. Most seem to work through basic taste
perception, and broadly speaking, the twin and family studies suggest the biggest
influence is on liking for macronutrient tastes generally (e.g., sweetness), not liking for
specific food items.72 Heritable variation in the taste receptor gene TAS2R38 and in
anatomical differences in taste papillae morphology shapes food preference indirectly,
such as by reducing liking and discouraging vegetable intake among individuals who are
especially sensitive to the bitter tastes in vegetables,73 and by influencing perception of
and liking for fat sensation.74,75 Sensitivity to some food-related odors does differ due to
variation in some olfactory receptor genes, and this may influence acceptability for some
foods.76 Among the very few known examples of genetic differences in evaluation of a
specific food, cilantro is an interesting case, because opinions about cilantro are so
polarized. Variation in a chromosomal region that encodes several olfactory receptors is
strongly correlated with perception of cilantro as either “herbal” or “soapy.”77 But these
genetic differences largely cause different people to perceive things differently, and
therefore cannot explain the more common scenario wherein different people may agree
on how a food tastes and simply have opposite reactions in how much they enjoy it.
In summary, olfactory stimulation is a major input into food flavor perception and
is what enables rather fine discriminations between the flavor characteristics of a very
wide array of actual or potential foods. Consistent with our evolutionary history as
generalist omnivores, hedonic evaluation of flavor is not predetermined, and is mostly
shaped by experience. So what sorts of experience in particular are most critical and
influential?

Determinants of Preference
Prenatal Experience
Experience shapes preference from the time that the olfactory and gustatory systems
become functional in utero. Animal models have established the importance of prenatal
olfactory learning for normal development. In rats, odors present in amniotic fluid guide
orienting and nipple attachment in the critical minutes after birth.78 For humans, olfactory
memories formed from prenatal experience are retained after birth, as neonates can
behaviorally discriminate the odor of their own mother’s amniotic fluid from that of
unfamiliar women.79 Volatile odor and flavor compounds in the food that a pregnant
woman eats appear in the amniotic fluid and alter its odor.80

Flavor exposure in utero produces lasting changes in appetitive and hedonic

reactions to those flavors after birth. Animal studies that allow strict control of the
maternal diet have demonstrated this effect in several species,81,82,,83 and it is observed in
humans as well. Newborn infants born to mothers who routinely consumed anise-
flavored foods and drinks while pregnant exhibit more positive facial expressions,
mouthing, and sucking when tested with anise odor.84 In another study, infants whose
mothers had been randomly assigned to consume carrot during pregnancy reacted more
positively to carrot-flavored cereal at the age of four months.85 Importantly, these studies
find shifts in infants’ facial reactivity to the target flavor, indicating effects on hedonic
evaluation.
As a means for transmitting food acceptance across generations, prenatal flavor-
learning arguably represents the first and most basic way that local culture identifiably
shapes individual habits and preferences. These early effects are especially important
because even subtle shifts in the valence and vigor of flavor responsiveness will affect the
mother–infant dyad during postnatal feedings, amplifying and perpetuating initial
tendencies. Yet the biopsychological mechanisms underlying this learning are unknown.
Exposure of the still-immature olfactory circuitry during fetal development may direct its
connectivity during a unique sensitive period. But there is little evidence that responses to
flavors experienced in utero are qualitatively different in form or function from memories
acquired after birth (e.g., Menella et al. 200185). Another possibility is that simple
familiarity habituates innate neophobic responses. In animal studies, prenatal exposure to
a flavor of the weaning foods does reduce behavioral and physiological indicators of
stress during the weaning period.86
But since fetal rats are also capable of learning aversive flavor associations,87 it is
possible that something more elaborate than simple familiarity is occurring. Dietary odors
appearing in utero come to elicit ingestion-specific facial affective responses, as opposed
to general orienting responses, which implicates neural systems involved in appetite
specifically, rather than a general recognition memory.84 This may represent a form of
operant learning in which the odor/flavor cue comes to elicit motor responses that are
innately triggered by nutrient influx, especially plasma glucose that would follow a
maternal meal.

Neonatal Experience
Immediately after birth, the mother continues to provide experiences preparing the infant
to recognize and accept foods of the local environment, since flavor compounds of the
maternal diet are also transmitted via milk. Infant behavior while nursing shows that they
can perceive and are affected by the changing flavor characteristics of milk.88,89 Specific
flavors consumed by the mother at this stage subsequently elicit more hedonically
positive reactions in solid foods at weaning.85

For non-human mammals, nursing is the obligate source of nutrition for

newborns. But for humans, the availability of manufactured infant formulas creates a
distinct point of developmental divergence for some, since formula feeding produces
lasting effects of its own. One main difference in the experience of formula-fed infants is
reduced sensory variety. Though manipulating the sensory variety of formulas has not
been studied experimentally, effects of reduced variety are likely, since duration of
breastfeeding (which provides variety) is inversely correlated with food neophobia and
picky eating later in childhood.90,91 Experimental studies in nursing rats and in older
children demonstrate that early variety exposure does increase later acceptance of novel
foods.92,93
Another effect of formula feeding comes from the specific sensory characteristics
of the chosen formula. Soy- and hydrolysate-based formulations are more bitter and sour-
tasting and some have odors that are unpleasant to adults. Infants fed these formulas
show decreased aversive reactions to cereals adulterated with sour or bitter tastes, and
increased acceptance of sour or bitter foods and drinks later on. This early experience
also alters childhood preferences for vegetables that share taste or odor characteristics of
the formula.94,95,96
Thus, in some ways, breastfed and formula-fed infants have fundamentally
different experiences, but in each case, sensory experience has lasting effects on
preference. But the effects are complex, since some experiments find more positive
hedonic reactions but reduced intake, at least in short-term test sessions that soon follow
the initial flavor exposure.95,97 Those investigators have proposed this may be an early
form of sensory-specific satiety. Some aspects of learned responding to the flavor may
immediately generalize to other contexts, while other aspects may take more time to
transfer. This is interesting because this is one example of how a developmental analysis
reveals a possible dissociation between aspects of appetite control that would not
necessarily be obvious later in development.

Childhood
Repeated exposure to flavors prenatally and postnatally serves as a bridge to independent
food evaluation; thus, maternal food choice functionally serves as a filter that narrows the
whole range of possible foods to a smaller, “proven safe” category that can guide
acceptance. After children’s transition to solid foods, parents still largely control the food
environment, and parents’ decisions and behaviors surrounding food continue to be the
primary determinant of experience (reviewed in Savage et al. 2007 and Anzman et al.
201098,99).

As the scope of children’s eating experience gradually expands, previous effects

of early exposure continue to be influential, since familiarity explains most of the
variance in self-described food preferences among younger children,100 and adding a
familiar flavor is an effective way to induce acceptance of a novel food.101 But
experience with foods also begins to incorporate more complex social and cognitive
dimensions. For instance, children begin to internalize cultural notions of appropriateness
that make particular foods more or less preferred at different times of day.102
One new influence at this stage is that caregivers begin incorporating foods into
instrumental contingencies, presenting foods either as tasks to complete or as rewards to
earn. Commonly children are instructed to eat a less desirable food to earn a more
desirable food. These strategies usually do not produce the results caregivers intend.103
Making a reward contingent on eating a particular food does increase intake of that food,
but it typically also decreases actual liking for it. Likewise, using a food as a reward for a
task increases liking for the food while potentially reducing intrinsic motivation for the
task. Decreased liking for an already disliked food (e.g., vegetables) that must be eaten to
gain a liked food (e.g., dessert) may be mediated by a perceptual contrast effect between
the two. A mental comparison to the anticipated dessert makes vegetables seem even less
appetizing. A more useful strategy appears to be using non-food rewards or social
rewards for eating less-liked foods, which can better maintain increased preference for
specific vegetables.104
Another important new source of influence is interaction with non-family adults
and peers. As children mature, eating increasingly becomes a social activity involving
explicit information exchange and subtler social modeling. In a variety of species, food
preferences are transmitted socially,105 and in humans, social modeling is seen in
experiments using adult and peer-aged models. Simply observing another person eating a
specific food increases a child’s acceptance of it106 and can induce children to select that
food over another that was initially more preferred.107 However, these social effects may
be rather short-lived on their own.108 The power of social modeling probably arises
because it produces the initial experience of eating a food that is sufficient to begin
establishing familiarity effects and the opportunity for other, more durable learning
processes that can eventually lead to robust preference.

Associative Conditioning: Flavor-Flavor Learning

Much work on food preferences has focused on infancy and childhood because of the
profound importance of the experiences occurring when circumstances relevant to eating
are largely still novel and there is the most to be learned. Nonetheless, specific types of
associative learning experience can and do continue to modify likes and dislikes
throughout life.

Flavor-flavor conditioning (hereafter, “F-F conditioning”) can occur when a

flavor (the conditioned stimulus [CS]) is consumed along with another taste or flavor that
is already hedonically significant (the unconditioned stimulus [US]). For example, a
novel flavor may itself may be neither liked nor disliked, but when it occurs in a palatable
dessert, it may become associated with the dessert’s sweet taste, and the flavor CS will be
more positively evaluated when encountered again. Evaluation can also shift negatively
when the CS flavor is paired with an unconditionally aversive US-like bitterness.
There are numerous studies demonstrating this effect in rats.109,110,111,112,113
Human studies have shown, for example, increased liking for a non-sweet herbal tea
flavor after it was previously consumed with sugar,114 and decreased liking for a specific
fruit flavor after it was consumed in an unpleasant, soapy-tasting mixture.115 F-F
conditioning appears to be a specific instance of the learning process more generally
called “evaluative conditioning,” which makes it resemble the unconscious shifts in
emotional responses and feelings of attraction or disdain that can be produced by
advertising, political propaganda, and even some interpersonal processes.116,117
Human studies of F-F conditioning as a whole suggest that it is easier to produce
negative shifts for flavors paired with aversive US’s than to produce positive shifts with
sweet pairing.118,119 An adaptive account based on negativity bias in evaluative
conditioning more broadly120 does make some sense for food learning, since poison
avoidance would be a pressing concern for survival, but that is unlikely to completely
explain why positive F-F conditioning is more difficult in humans, since F-F preferences
with sweetness are robust and easy to produce in rat experiments.
It may be important for F-F conditioning studies to distinguish effects on
perception from effects on liking. Pairing a flavor with sweetness can cause the flavor
itself to be perceived as sweet, but that would not necessarily cause a person to like it
better. Preexisting differences in sweetness-liking do predict the direction and strength of
hedonic shifts for a sweet-paired odor.121 Learning may also be moderated by cognitive
or personality factors, like dietary restraint. One study found that learned preference shift
for a flavor paired with eating a sweet candy was attenuated in restrained eaters.122 It
could be that restrained eaters did not perceive the candy as pleasurable, or that eating it
triggers concerns about dieting that interfere with conditioning. Thus, as a general
principle, perception and evaluation of a flavor is altered by sensory associations with
other tastes or flavors that have accompanied it, but there are clear individual differences
in the case of positive conditioning (especially with sweet tastes). Further work on this
topic could benefit from extending studying positive F-F conditioning with a range of
positive US’s other than sweetness.

Associative Conditioning: Flavor-Nutrient Learning

Another associative learning process that shapes preference reflects the value of
Pavlovian conditioning for using sensory cues to predict impending, biologically
significant events. Since a food’s flavor is always experienced before its post-ingestive
effects, associations between specific flavors and nutrients sensed in the gut or post-
absorptively make flavors a useful guide to food choice and the amount consumed—or at
least that would be the function for animals and our human ancestors searching for
nutrition in an environment where foods were of variable quality. Flavor-nutrient
conditioning (hereafter “F-N conditioning”) is a Pavlovian learning process in which
evaluation of a food’s flavor (the CS) is altered because of its history of pairing with
post-ingestive nutritional effects (US’s).

Most of our understanding of F-N conditioning is based on work with laboratory

animal models. Conditioning typically involves a novel CS flavor of modest initial
acceptability, or even one that would be initially disliked. Consumption of that CS is
accompanied by intragastric or intraduodenal infusion through a catheter of a nutrient
such as glucose, the US. The purpose of this flavor+infusion method is to separately
manipulate what is detected in the gut from what is tasted in the mouth, so that the flavor
is becoming associated with the post-ingestive actions of the nutrient rather than its own
inherently attractive taste (i.e., F-F conditioning). In alternating sessions, a different
flavor is accompanied by plain water infusion, to control for simpler effects like mere
exposure. With experience, rats and mice generally show dramatically increased
attraction to that flavor paired with nutrient infusion, and strongly prefer it in a choice
relative to the unpaired flavor, even when it is no longer accompanied by the nutrient.
F-N conditioning occurs rapidly. In fact, several experiments show that positive
preference shift for a flavor paired with nutrition can be learned in a single trial.123,124
Prior to this finding, it was already clear that one-trial learning could occur for flavors
paired with long-delayed illness, but unlike aversion learning, F-N conditioning involves
post-ingestive US events that onset rapidly, within the first few minutes of a meal. Soon
after the first bite, food begins emptying from the stomach into the duodenum, and
various sensors there respond to its macronutrient content. This detection is
psychologically significant. In studies where rats or mice are accustomed to consuming a
solution accompanied by intragastric water infusion, the first time a new flavor is
accompanied by nutrient infusion, they accelerate intake within minutes and end up
consuming a significantly larger meal.125,126,127 That positive feedback response to
nutrients arriving in the gut is not merely a general stimulation of appetite. Rather, it
reflects specific enhancement in evaluation of the accompanying flavor.125 If the flavor is
removed and replaced with a different one, rats no longer are inclined to continue
consuming it. In other words, within minutes of a flavor accompanied by nutrition, rats
already act as if that flavor seems “better” because of it.
There are several other attributes of F-N conditioning that underscore its
important role. F-N conditioning is probably involved early in development, such as
increased preference for flavors experienced in mothers’ milk, as F-N conditioning has
been demonstrated in pre-weanling rats.128 Preferences acquired through F-N
conditioning are enduring and long-lasting.123,129 Preference shifts produced by F-N
conditioning can be strong enough that rats will begin avidly consuming an initially
aversive taste or flavor (e.g., bitterness, vinegar) that they would have otherwise
avoided.123,130
The rewarding US signal (or signals) supporting F-N conditioning stems from
physiological effect of nutrients, but it is important to recognize that reward is not a
detection of calories per se, because different nutrients are not equally effective.131
Glucose and maltodextrin in particular support the strongest and most rapid preference
learning, with fructose being relatively ineffective. Fat can produce learned flavor
preferences, but when equated for calories with glucose, post-ingestive reward signals
from fat are weaker, onset more slowly, and require more experience to shift flavor
preferences.124,132,133
Extensive work by Sclafani and colleagues aimed at identifying the precise site of
nutrient detection that supports F-N conditioning has made use of surgical
deafferentations, genetic knockouts of nutrient-sensitive receptors, and non-
metabolizable nutrient analogues. The transduction mechanisms and afferent pathways
have proven difficult to isolate, though pre-absorptive detection in the proximal intestines
through a still-unknown signaling pathway appears to be important (see Sclafani &
Ackroff 2012134 for a complete account of this line of work). There may also be
secondary, longer-delayed mechanisms of post-ingestive reward, since portal glucose
infusion also activates brain areas relevant to food reward and preference evaluation.135
This is consistent with behavioral evidence that rats appear to be affected by at least two
phases of post-ingestive events in F-N conditioning: a rapid-acting signal affecting flavor
perception and evaluation within the early minutes of a meal (thus presumably pre-
absorptive), and then a longer-delayed signal or signals arising late in the meal or after its
cessation.136
The speed of acquisition and powerful, lasting effects on choice and intake all
suggest that F-N conditioning may play an important role in overeating by steering
choice towards more energy-dense foods and by stimulating increased intake once they
are obtained. But does learning a F-N association produce a true hedonic shift, causing
that flavor to be perceived as more palatable? Though this has often been a presumed
mechanism when preference behavior is so robust, the answer to this question should not
be taken for granted. An alternative, for instance, could be that a flavor has become more
capable of capturing attention and eliciting approach from a distance, thus leading to
meal initiation without necessarily eliciting more pleasure.
The most direct evidence for a hedonic shift in F-N conditioning comes from
taste-reactivity measurements.137,138 One study found that pairing a minimally sweet
flavor with a post-ingestive nutrient reward produced a positive shift in hedonic taste
reactivity that was comparable in magnitude to increasing the sweetness of a solution
from 3% to 16% sugar. Other studies have shown the same pattern of changes in lick
microstructure seen when making a sugar solution sweeter.139 Some human studies also
show an apparent palatability effect, especially for flavors associated with the satisfaction
of hunger.140,141 In several experiments with children, the young participants begin to
place a flavor higher in a rank-ordering of different flavored foods after they have
experienced that flavor paired with added calories.142,143
 But the evidence for positive hedonic shifts in F-N conditioning is not entirely
straightforward. In one study,144 participants reported increased liking for a distinctive
flavor in sorbet after previously experiencing that flavor in a drink containing sucrose
(sweet and caloric). But participants trained with the flavor in a drink with maltodextrin
(caloric but not sweet) did not show the same pleasantness-increase, even though F-N
conditioning was still evident as increased consumption of the flavored sorbet. This
suggests F-N conditioning can produce a hedonic shift under some circumstances, but a
hedonic shift is not necessary to stimulate increased intake. That interpretation concurs
with the animal literature. Some rat studies have found positive taste-reactivity shifts, but
other studies have found large increases in intake and preference without any apparent
taste-reactivity change.145 A common factor across the sorbet study with human
participants and in the rat taste-reactivity studies may be that positive hedonic shifts were
only evident when a sweetened CS flavor was followed by caloric consequences. In any
case, we can conclude that F-N conditioning can shape food intake and meal size through
both hedonic and non-hedonic processes, though more work is necessary to understand
why a hedonic shift only sometimes occurs.
The work on F-N conditioning in animal models produces such robust effects on
flavor evaluation and on meal patterns, and there are enough examples of F-N
conditioning in human experiments, that it is difficult to doubt that this is a fundamental
influence in human food preference. But reliably producing and describing F-N
conditioning effects with human participants in controlled studies has been challenging,
and some studies fail to detect effects. There are a number of reasons this may be. First,
people already have extensive exposure to different foods and established likes and
dislikes for flavors. Conditioning is likely to be much less effective with familiar flavors.
It is also theoretically debatable whether “liking” should be the main dependent variable,
since animal experiments sometimes find effects on choice and intake without liking. To
the extent that liking is involved, the self-report response scales commonly used have
dubious psychometric properties.146
Another possibility involves developmental changes. There are several clear
demonstrations of F-N conditioning in children (reviewed in Birch 1999147). It is easier to
test children with novel foods and flavors for which they have few prior associations.
But, as mentioned previously, from an ecological and adaptive perspective, early
childhood is a critical time for learning. There may therefore be gradual reductions in
plasticity in the neural circuitry involved in F-N conditioning after childhood. It is widely
accepted that such maturational constraints occur in other domains of human learning,
like language acquisition.148 But there are no obvious developmental reductions in the F-
N learning abilities of laboratory animals.
Additionally, there may be individual differences in F-N conditioning that are
unaccounted for. One study trained participants with two differently flavored desserts,
with the flavors corresponding to differences in energy density. Only participants who
were low in dietary restraint subsequently rated the high energy-paired flavor as better
tasting. Restrained eaters showed no evidence of differential conditioning.149 It is unclear
if some aspect of dietary restraint made those participants less sensitive to the caloric
consequence, or made them respond differently to it. But either way, it shows that
cognitive factors may impair or interfere with effects of F-N conditioning.
 Some conceptual proposals are more speculative, but warrant serious
consideration, especially with regard to the limits of animal models for human nutritional
psychology. Some have questioned whether F-N conditioning should be as influential in
governing human appetite as it is in animals, or, at least whether it should operate in the
same way, given major shifts in the organization of eating behavior that coincide with the
advent of human culture.150 For instance, the invention of cooking offered proto-humans
considerable advantages, which depended to some degree on mental capacity for
advanced planning. Cultural changes that allow more intentional control over the food
environment and eating occasions may mark a quantum leap that required a fundamental
shift in controls of choice and meal size to incorporate explicit decisions about future
states, such as expected satiety, rather than lower-level, more automatic associative shifts
in immediate evaluation of foods’ sensory attributes.151 Thus perhaps an aim of research
on flavor-nutrient conditioning in humans should be to study its effects on more complex
beliefs and attitudes about foods, like expectations that a food will be “satisfying,” or its
economic utility, rather than sensory enjoyment as such.
Another possibility is that, for most adult humans living in the modern food
environment, the processes underlying acquisition of new F-N associations are simply
little used and poorly rehearsed. This would be consistent with the evidence that F-N
conditioning is easier to produce in children, but that adults still can learn F-N
preferences under some circumstances. It may be the case that in the modern food
environment, beyond the foundational preferences learned in childhood, there remains
little opportunity or even need to learn new preferences in adulthood. There are at least
two observations about modern eating that support this idea.
First, for most people most of the time, we really only encounter foods of fairly
high quality. Unlike our ancestors (and in spite of the dubious claims of many “natural”
health gurus), we do not need to be concerned that a food from the supermarket will
poison us, nor that we will fail to get enough calories. The current public concern about
industrial sugar production stems from the fact that foods are usually likely to have more
calories than we expect them to. If one is selecting from a wide range of (mostly
processed) foods, there is little need to learn to prefer one food over another based on its
nutrient content because it would be nearly impossible to self-select a diet that failed to
provide adequate calories. The real risk is the potential adverse effect of
overconsumption, but it is not clear the basic mechanisms of F-N conditioning are
sensitive to that input. Second, the ubiquity of eating opportunities means there is very
little cost to getting too few calories on any one occasion, since another meal is usually
available for relatively little cost in money or effort. Thus there is less need to remember
which flavors signal the biggest nutritive payoff.
If true, this by no means diminishes the importance of F-N conditioning in food
preference. The totality of the evidence shows that F-N preferences are readily learned in
childhood, which is when most foods are first encountered and when many long-lasting
habits are established, and those preferences can be modified in adulthood. Ultimately,
these proposals attempt to address why F-N conditioning experiments in adults obtain
mixed or weak results relative to the animal models. This suggests research on animal
models of F-N conditioning should incorporate experimental manipulations that model
specific features of the modern food environment, and effects of that environment on
basic learning processes.
Looking Forward: Applications to the Modern Obesogenic
Environment
The ways that food preferences—of cultural groups, of families, of individuals—are
fundamentally shaped by learning reflect the challenges and constraints on food
availability during our evolutionary history. Now, the propensity to overeat and maintain
extra weight is commonly viewed as resulting from the mismatch between our inherited
tendencies for coping with scarcity, and the new, biologically bizarre circumstances in
which high-calorie food is easy to come by. Thus it becomes important to identify the
ways in which the learning mechanisms that promoted adaptive control of food
recognition, choice, and meal size in ancestral environments are affected by dietary
excess.

Features of the Modern Food Environment

There are several important features of the modern food environment that pose a
challenge. Many modern foods are not grown or produced so much as they are
“designed,” and this process is aimed at maximizing inherent palatability. The global
food manufacturing enterprise uses techniques of chemistry, engineering, psychology,
and mathematics to produce formulations that optimize consumer enjoyment. The vast
scale of food production and retailing makes food less expensive than ever before. Even
though people are buying and eating more food, it represents only 15% of Americans’
spending, compared to over 40% in the 1960s.152 Doubly problematic is that food
processing, which adds sugar and fat, also makes foods cost less per calorie.153 Food is
also easier to obtain. Shelf-stable processed foods can be purchased ahead, allowing
people to keep a pantry full of snacks and meal items in their homes at any given time.
Fast food restaurants and supermarkets have proliferated, and many are now open 24
hours a day. With this combination of low cost and high accessibility, the time and effort
needed to stop whatever you are doing to have a snack is often minimal. Food now has
more sensory variety than ever before. The Kellogg brothers developed corn flakes in
1894 as part of the bland, abstemious diet Dr. John Harvey Kellogg prescribed for moral
well-being. The company bearing their name now offers about 55 varieties of breakfast
cereal. Finally, reminders of food are ubiquitous, as advertising continually makes us
aware of food’s sensory attributes and availability.

Given that we are endowed with a learning system that steers choice and
promotes intake of foods with flavors that predict positive caloric consequences, and is
tuned to pick out environmental cues that predict food availability, this combination of
factors creates problems. One claim is that, at least for some people, this pattern of
experience makes foods “drug-like,” resulting in a syndrome of learned physiological and
behavioral adaptations that resemble drug addiction, especially in regard to the excess
attention paid to food cues, compulsive food-seeking, and subjectively experiencing loss
of control over intake. To evaluate the usefulness of this framework, it may be instructive
to consider what it means to be a “drug.” Drugs of abuse that are consumed recreationally
for their desirable sensory, perceptual, and emotional effects are in that category
specifically because they directly, powerfully, and supernormally stimulate brain
circuitry that would ordinarily process “natural” rewards: the taste of a berry, a flirtatious
display from a potential mate, a friendly embrace, a sip of water on a hot day. We should
expect that neural circuitry for processing such stimuli should be tuned in sensitivity to
the range of stimulus intensity it would ordinarily handle. But drugs, being
“supernormal” stimuli, stimulate an exaggerated version of the ordinary response
patterns. That is, drugs are drugs because they are like food, only more so. It is fitting to
view modern food similarly. Modern foods are like foods, only more so, especially in the
sensory cues that drive hedonic evaluation and preference. First, there are beyond-natural
levels of added sugars and fat, the fundamental stimuli for palatability. Second, added
flavors and flavor enhancers mean that the sensory cues that become learned signals for
post-ingestive consequences are themselves more salient.
As will be discussed in detail in the subsequent chapters of this book, the sensory
and nutritional characteristics of modern foods, especially within the context of ubiquity
and convenience, may for some individuals contribute to a pattern of neurobehavioral
responses in motivation, attention, affect, and memory that results in escalating intake,
craving, compulsive seeking, and loss of control that is commonly recognized as
addiction. Research on the fundamental motivational drivers of both appetite and
substance abuse have much of value to share with each other, recognizing that (like with
drugs) these consequences ultimately stem from an extensive history of learning
experiences.

Interactions with the Modern Food Environment

There are currently two main perspectives on how the modern food environment interacts
with the learning mechanisms governing food choice, especially flavor-nutrient learning,
that can contribute to overeating. At first they appear to offer potentially contradictory
explanations for overeating, though it is possible to reconcile them.

Supporting the Preference Learning Systems

The first main view is that overeating is the predictable consequence of the food
preference learning systems’ doing what they are supposed to do. This view is consistent
with the idea that these learning mechanisms function to steer choice and promote intake
of calorically dense options, but also, importantly, to anticipate post-ingestive
consequences so as to simultaneously trigger adaptive physiological preparations in
digestion and metabolism that enable the organism to cope with such large meals.154 In
this view, it could be said that learning functions too well in the modern food
environment that offers more opportunities to encounter nutrient-dense foods. The
appearance, smell, taste, and flavor of preferred foods, established by prior learning
experiences described previously, come to direct both appetitive and evaluative
responses. These stimuli attract, hold attention, and elicit approaches leading to increased
choice of calorically dense foods specifically. The increased palatability from sugar and
fat is influential on its own, but also establishes conditioned responses to the initially
arbitrary flavors in the food. F-F conditioning and F-N conditioning act to enhance
palatability, which serves both to stimulate vigorous eating and promote cephalic
responses that prepare the digestive system for nutrient influx, delaying satiation and
enabling rapid deposition of ingested fuels.
Obscuring the Preference Learning Systems
A different view of food–environment interactions is that certain aspects of the modern
food environment prevent learning from working as it should. Most of the research on
preference learning has focused on the ways learning promotes choice and increased
intake of calorically dense foods. But this second view proposes that sensory cues for
nutrient density also enable individuals to appropriately adjust and limit their intake in
response to anticipated nutrient loads, to avoid overconsumption and, ultimately, the
physiological costs of chronic positive energy balance. Modern overconsumption,
according to this view, occurs in part because aspects of the modern food environment
make sensory cues routinely uninformative or unreliable.155

Flavor-nutrient learning can only occur to the extent that specific sensory cues
reliably correspond to post-ingestive consequences. But modern food processing can
explicitly uncouple flavor–nutrient links, using added flavors, flavor enhancers, non-
caloric sweeteners, thickeners, and textural manipulations. When a sensory cue
sometimes does and sometimes does not predict post-ingestive consequences, it is no
longer useful for anticipatory adjustments in physiology or behavior. Impairment would
be especially pernicious if one’s early life experience was characterized by sensory-
nutrient consistency, with a maturing appetite control system that comes to rely on such
high-fidelity information, only then to have that ability degraded as one’s food world
expanded. This could be increasingly commonplace for modern children raised with
mostly wholesome, minimally processed foods early on, who then later discover sodas,
processed snacks, and other sources of sensory entertainment.
Much of the work stemming from this view has focused on the effects of
incorporating non-nutritive artificial sweeteners into one’s habitual diet, since sweetness
is an especially salient sensory cue that is intentionally manipulated in modern foods.156
Artificial sweeteners allow consumers to obtain sweetness without added calories or
other physiological effects of sugar. Presumably, considerable effort by food product
designers is devoted to making the sugar-sweetened and artificially sweetened versions of
snack foods similar in their flavor and texture, though the artificially sweetened versions
may represent anything from a modest to a dramatic reduction in calories, thus potentially
obscuring the sensory–nutrient link that would be otherwise informative for frequent
consumers. The same may become true for a range of other sensory attributes as
advances in food technology permit creation of various treats whose rich flavors and
textures disguise the absence of expected calories.
There is a variety of circumstantial evidence consistent with the notion that a
history of consuming non-nutritive sweeteners is associated with disturbances in appetite.
In functional magnetic resonance imaging (fMRI) studies, habitual diet-soda consumers
showed a more complete pattern resembling sugar ingestion when tasting saccharin,
unlike non-consumers, for whom saccharin activated fewer taste-reward areas than real
sucrose.157 Use of non-caloric sweeteners of all forms (not just sodas) is associated with
attenuated amygdala response to sugar taste.158 Amygdala activation could be interpreted
as indicating perceived biological relevance of a stimulus, so a reduction is consistent
with the proposal that artificial-sweetener history changes the consequence-signaling
value of sweetness. Compared to non-consumers, people who routinely consume diet soft
drinks experience less stimulation of appetite by a sweet taste, which has been interpreted
as a weakened learned expectation of energy consequences.159
These studies of artificial sweetener consumption, however, must rely on self-
selected samples, whose habits and histories with weight and diet must certainly differ in
various other ways, making it impossible to infer causality. This makes animal studies of
diet history especially useful. These studies have shown ways that sensory-nutrient
inconsistency is problematic for weight maintenance. One study exposed rats to chow
diets with varied flavors added. For different groups, the flavors sometimes did and
sometimes did not correspond to different energy densities. Rats gained significantly
more weight when flavor–energy relationships were unpredictable.160 This study did not
involve sweeteners, but still demonstrated the principle that some salient sensory property
that is informative about nutritive consequences can be protective against overweight.
Another line of research comparing rats reared with repeated exposure to non-caloric
sweeteners to those with only exposure to nutritive sugars paradoxically found greater
weight gain with non-caloric sweeteners. The effect on weight gain appears to be due to
reduced satiating effectiveness of sweetness, as rats failed to reduce their intake of a test
meal after a sweet preload with caloric or non-caloric sweeteners.161,162 The effects
persist after non-caloric sweeteners are removed from the diet, suggesting a lasting effect
on learned responses.

Combining Perspectives
It may at first seem that these two views offer different, contradictory explanations for
the obesogenic influence of learning in the modern food environment. Is it that
successfully learning about flavor–nutrient relationships (which would have been
essential for adaptation to a lean environment) now drives excess energy intake when the
environment provides an unnatural surplus? Or is it that the inconsistent nature of the
modern, processed food supply confuses and prevents appropriate learned adjustments in
behavior and physiology and in coping with excess energy? These theories are both
feasible and in fact complementary, to the extent that these experiences affect different
aspects of eating behavior. The learned responses to sensory cues that predict post-
ingestive consequences reflect the joint contribution of two opposing response
tendencies: preferential choice and enhanced evaluation stimulates intake, but learned
anticipatory satiation responses put the brakes on intake.163,164 The modern food
environment could inadvertently be acting to promote the former of these while
preventing the latter.

An environment that provides occasional links between specific sensory cues and
post-ingestive nutritional consequences may be sufficient to establish increased seeking,
preferential choice, and positive evaluation of those cues, even if those links are irregular.
Enhanced preference for a flavor paired with nutrients can be learned in a single trial, and
it remains fairly resistant to extinction even after only minimal training experience.123 But
the learned, intake-limiting effects of satiation expected to follow a learned flavor cue
appear to extinguish fairly rapidly when post-ingestive consequences do not consistently
occur.165 Thus the modern food environment could be doubly problematic by providing
situations where the correspondence between sensory cues like flavors, sweetness,
texture, and caloric consequences is reliable enough to maintain learned intake-promoting
responses, but not reliable enough to maintain learned intake-limiting responses.

Summary
At the core of appetite, beneath layers of cognitive, emotional, and sociocultural
elaboration, the biopsychological controls of food choice are tied to hedonic evaluation of
food’s sensory attributes, including the basic tastes of nutrients and complex flavors. But
how individuals evaluate and respond to different foods is diverse enough that in just
about every cuisine there’s something savored by locals that seems repulsive to visitors.
The range of food acceptability stretches so far that staple foods in some diets are simply
rejected as non-foods by outsiders (e.g., insects, blood, beef). But it is not even necessary
to traverse wide cultural or geographic distances to document these differences. On a
local scale, looking casually at one’s own family, friends, and neighbors makes the
idiosyncratic and individualized nature of food preference evident.

All of this diversity reflects the predominant role of experience in shaping

preference. The environment is influential from the time that gustatory and olfactory
systems are functional in utero. Through experience, which can be as simple as mere
exposure, and broad enough to encompass associative, cognitive, and social influences,
the sensory properties of food become labels for its identity, quality, and suitability to
meet nutritional needs. Bottom-up coding of a food’s physical properties elicits stored
memories of prior experiences, and this information is integrated with signals about
current bodily states. Although there are innate influences (e.g., the early response to
sweetness), the power of learning is demonstrated by the fact that, in contests between
innate responses and learning, learning typically wins. Culture and individual experiences
modify the inherent liking for sweet taste, and individuals acquire liking through
exposure and conditioning for tastes that would be otherwise aversive.
Learning systems form the nexus between the environment and food hedonics,
and therefore become especially important in our efforts to understand how the modern
food environment leads to overeating. The major shifts in how food is made, distributed,
and marketed, and the concomitant changes in food variety, composition, and availability
can be expected to have several effects on appetite through learning mechanisms that
evolved to promote adaptive behavior in a very different environment. On one hand,
learning to like flavors that signaled food quality enabled individuals to cope with
environments where quality was variable, but this now acts to promote selection of foods
that are of unnaturally high quality (in terms of caloric density, at least). At the same
time, some aspects of the modern environment may fool these learning mechanisms by
providing inconsistent, unreliable information about foods’ sensory properties, and those
failures may particularly impair appropriate cessation of eating. In sum, why we eat what
we eat is generally explained by what we learn about what we eat. Why many people are
now eating so much can be attributable both to what we learn and to what we fail to
learn.

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