Appetite: Psychobiological and behavioral aspects

R.J. Stubbs, G.S. Finlayson, and J.E. Blundell, Appetite Control and Energy Balance Research Group, School of Psychology,
Faculty of Medicine and Health, University of Leeds, Leeds, United Kingdom
© 2023 Elsevier Ltd. All rights reserved.
This is an update of R.J. Stubbs, J.E. Blundell, Appetite: Psychobiological and Behavioral Aspects, Editor(s): Benjamin Caballero, Encyclopedia of
Human Nutrition (Third Edition), Academic Press, 2013, Pages 108–115, ISBN 9780123848857, https://doi.org/10.1016/B978-0-12-375083-9.
00017-9.

Introduction 2
The nature of feeding behavior and appetite control 2
Learned appetites, satieties, and feeding behavior 2
Methodological issues 4
Measuring hunger 4
Measuring food intake 4
Measuring feeding behavior 5
Sensory stimulation and food hedonics 5
Sensory stimuli and body weight 6
Sensory versus nutritional determinants of intake 6
Emotional stimuli and over-consumption 6
Meal patterns, appetite, and energy balance 7
Social and situational influences on feeding behavior 7
Cognitive and social cues 7
Diet composition and appetite 8
How do macronutrients and energy density affect satiety? 8
Diet composition and satiety to prevent weight gain 8
Conclusion 9
References 10

Key points
• Review the nature of human eating behavior and appetite control
• Discuss the role of learning in acquired eating behavior traits.
• Consider methodological issues relevant to the measurement of appetite control and energy balance.
• Discuss the role of environmental factors (meal patterns, social and situational influences) on human appetite control.
• Summarize the role of diet composition in appetite control and weight management.

Glossary
Appetite The subjective expression of willingness or motivation associated with qualitative selection and quantitative ingestion
of specific foods during an ingestive event. Appetites are specific to certain foods. Appetite is not necessarily solely related to
situations of nutritional depletion and can be influenced by a number of physiological and non-physiological factors.
Appetites are often learned and frequently sensory specific
Hunger The subjective sensation and associated processes that can be described as a general motivation to eat. An increase in
subjective hunger usually predicts meal initiation in ad libitum feeding subjects. It does not necessarily predict type or amount
of food eaten
Food hedonics Liking (an experience of pleasure) and wanting (anticipatory motivation), are distinct hedonic processes with
dissociable neural pathways that are thought to serve as a basis for animals (including humans) to acquire through learning,
eating behaviors that lead to the acquisition of energy and nutrients. They may be influenced by the food, the physiological state
of the organism, and the environment in which food and subject interact
Liking The sensory pleasure elicited by contact with food contributing to the hedonic motivation to consume (wanting)
Wanting The motivation to consume a specific food, manifesting explicitly (craving/desire) or implicitly (food cue
responsiveness)

Encyclopedia of Human Nutrition, 4th edition, Volume 2 https://doi.org/10.1016/B978-0-12-821848-8.00111-6 1

2 Appetite: Psychobiological and behavioral aspects

Satiation Processes during a meal that generate the negative feedback leading to its termination (within-meal inhibition)
(strengthened by meal volume and weakened by palatability)
Satiety The motivational process of not wanting to eat and often expressed as degree of satisfaction and/or fullness. This bears
some reciprocal relationship to hunger. Some authors (Le Magnen, Blundell) argue that satiation and satiety are
mechanistically distinct, the former determines meal size; the latter meal frequency

Introduction

Human eating behavior is complex and influenced by physiological, environmental, genetic factors and learned experience. Eating
behavior studies commonly focus on assessment of food intake and its physiological, psychological and environmental determi-
nants (Blundell and Stubbs, 2003). Often, these studies are very short-term and have limited capacity to explain mechanisms
affecting longer-term energy balance. Because overnutrition is increasingly prevalent and undernutrition is still a significant
problem, it is important to consider how appetite control relates to energy balance. It is also necessary to consider some important
methodological issues in appetite research and environmental factors (including the composition of the diet) that influence energy
intake and longer-term energy balance (Stubbs and Finlayson, 2018).

The nature of feeding behavior and appetite control

Mammalian feeding occurs regularly and intermittently and despite a general lack of conscious nutritional knowledge on the part
of the animal, usually appears to match energy intake (EI) and nutrient intakes with requirements. How is this achieved? The
common explanation is that appetite, EI or feeding behavior are regulated to ensure that physiological requirements are met.
However, there is a lack of direct evidence for this regulation. Neither feeding behavior, nor appetite are regulated in a strictly
physiological sense because (1) they are not held constant within certain narrow limits and (2) feeding responses are not an inev-
itable response to an altered physiological signal or need. Feeding behavior is responsive to a number of induced states such as
pregnancy, cold exposure, growth and development, emotions, and weight loss. These responses have often been cited as an evidence
of a system that is regulated through homeostatic, symmetrical negative feedback loops. It is probable that some aspects of body
size and composition are regulated and changes in feeding behavior are functionally coupled to those regulatory processes.
Evidence suggests that the regulation of human energy balance is largely asymmetric. Human eating behavior has evolved in
resource-limiting environments to defend against energy deficits (weight loss) rather than excess energy intake (weight gain)
(Stubbs and Turicchi, 2021).
Hunger and satiety often have a large learned, anticipatory component rather than solely being the direct consequences of
unconditioned physiological signals per se, such as reduced gastrointestinal content. Such physiological events can act as important
cues for feeding but they do not necessarily directly determine that behavior.
The mechanism by which feeding behavior is coupled to physiological events (and other events) is the process of learning. To
understand feeding behavior, hunger, and satiety processes, the mechanism by which learning links feeding behavior to physiolog-
ical, sensory, nutritional, situational, and other learning cues, must be appreciated. This mechanism is termed as associative condi-
tioning of preferences, appetites, and satieties (Blundell and Stubbs, 2003; Mela and Rogers, 2013).

Learned appetites, satieties, and feeding behavior

Animals and humans learn (or become conditioned) to associate a given food with the physiological consequences of having
ingested it. They associate certain proximal stimuli such as the smell, color, taste, or texture of a food (the conditioning stimulus),
with a set of sensations that are directly felt (sensory afferent inputs), in relation to the external stimulus and to the endogenous
changes such as physiological and neuroendocrine responses to food. The physiological changes that occur as a result of ingesting
the food are termed as the unconditioned stimulus. The individual forms a learned or conditioned association between the condi-
tioning stimulus and the unconditioned stimulus (the detectable consequence of eating), which informs them of the sensory and
physiological consequences of ingesting that food (Blundell and Stubbs, 2003; Mela and Rogers, 2013). This process is summarized
in Fig. 1. Conditioned or learned associations are most efficiently established if the food is sensorially distinct, if there is a significant
detectable postingestive consequence of ingesting the food, or if a training or learning schedule is encountered (e.g., by repeated
exposure to the food under similar conditions). Learning is facilitated by social interaction.
As regards with the notion of appetite regulation, a problem arises when foods are constructed to look and taste like foods with
a different composition. For sometime, after the initial exposure to the food, subjects will respond to it in a manner that is deter-
mined not by immediate exposure to the food but by what they have learned during the previous period of exposure to the similar
foods upon which the learning was originally based. Only if the food produces a very large unconditioned stimulus will this
 Appetite: Psychobiological and behavioral aspects 3

Fig. 1 The process by which the subject learns to associate the postingestive consequences of eating with the food eaten and the environment in
which it was eaten. Environmental influences can vary in strength from negligible effect to influences so strong that they can constitute the major
factor determining a subject’s subsequent response to that food.

previously learned response be instantly over-ridden. This raises the possibility that the use of food mimetics (e.g., high intensity
sweeteners) may disrupt stable patterns of learned feeding behavior in consumers at large.
The above view of the nature of feeding behavior has implications for the way the appetite system functions in relation to the
development of obesity. Physiological models of body weight (energy balance) regulation suggest that feeding behavior is geared to
the regulation of a stable body weight through homeostatic negative feedback. Obesity has therefore been seen as a consequence of
defects in the regulation of physiology and behaviors related to energy balance. The evidence from behavioral studies suggests the
following: (1) feeding behavior is inherently more responsive to decreases rather than increases in body weight; (2) current secular
trends in body weight suggest that, over time, it is very easy to increase body weight, which infers body weight is not tightly regu-
lated, at least with reference to weight gain (for instance, in the National Health and Nutrition Examination Survey (NHANES) die-
tary surveys of American adults, average weight increases by 0.2–2.0 kg per year), and (3) there is very limited evidence of clear lean/
obese differences in feeding behavior of a type that suggest defects in a regulatory system. For example, evidence suggests that people
living with obesity tend to select a diet that is higher fat, energy dense and containing more highly processed foods, which itself
facilitates over-consumption. However, the tendency to select fat cannot be viewed as a defect in physiological regulation, but rather
a difference in food preferences that subsequently impact body weight (Blundell and Stubbs, 2003; Mela and Rogers, 2013; Stubbs and
Turicchi, 2021). There are interesting individual differences in response to food-based reward stimuli according to body weight
status. Individuals with obesity are more reactive to food cues i.e., their attention is more easily grabbed and held by these cues.
In a hungry state, these “attention grabbing” effects become more pronounced in participants with overweight/obesity compared
to lean. High BMI also correlates with cravings while dieting and subjective cravings in individuals with overweight is associated
with food cue reactivity. Similarly, increased BMI has been associated with more frequent craving, and craving specifically for
high fat foods along with increased intake of them. It may be far more profitable to attempt to understand how feeding responds
to environmental and endogenous stimuli and which of these responses are functional and adaptive, and which are not (Finlayson
et al., 2007). In this context it is important to remember that evolution has selected us to optimize resources in uncertain environ-
ments, bank surplus energy, and compensate for energy deficits. Under conditions of environmental uncertainty there is little need
4 Appetite: Psychobiological and behavioral aspects

to evolve regulatory systems that protect from weight gain. Furthermore, accumulating body fat provides the ecological advantage of
maintaining functional integrity and survival under conditions of food shortage. As a species, humans may have never had the time
nor circumstance to evolve patterns of feeding behavior that are protective against weight gain because such evolved behaviors
would have little functional or evolutionary advantage. Throughout the development of civilization humans have increasingly
manipulated their food production and consumption environment to suit these biological design specifications. This promotes
weight gain. Supermarket society is Optimal Foraging Theory taken to its logical conclusion. We do not tightly regulate energy balance.
Human eating behavior is designed by evolution to feast in times of plenty as protection from famine. Modern industrial marketing
strategies play on the constraints our species has evolved with, as an opportunistic forager. These considerations influence the meth-
odological approaches that attempt to investigate how feeding behavior responds to endogenous and environmental influences
(Stubbs and Tolkamp, 2006; Stubbs and Turicchi, 2021).

Methodological issues
Measuring hunger
Hunger is a subjectively expressed construct that people use to express a motivation to eat. The most appropriate measure of
hunger is its subjective expression at a given time. This is achieved by asking subjects to mark a visual analog scale, which takes
the form of a straight line with two extreme representations of hunger anchored at either end. It is most useful to track changes
in subjective hunger over time and in relation to feeding events, diet composition, or physiological parameters. Hunger itself
exhibits a large learned component (see the Section on Learned appetites, satieties, and feeding behavior, above) as reflected
by the fact that most of the variation in a person’s subjectively expressed hunger is accounted for by time. If hunger is plotted
against time in Western subjects feeding ad libitum then it generally exhibits 3 peaks and troughs, which broadly correspond
to the 3 main meal times of a Western feeding schedule (Fig. 2). Although subjective hunger is a relatively poor proxy for the
amount eaten it is a reasonably good predictor of when eating will occur. It is important to recognize that hunger can be influ-
enced by a large number of factors and so a search for a specific unitary hunger signal is likely to prove fruitless. Thus a large
survey of over 600 men, women, boys, and girls could find no clear constellation of traits, sensations, or characteristics that typi-
fied hunger. A variety of hormones and drugs, the sight and smell of food, its perceived palatability, timing and social situation
can all influence hunger (Blundell et al., 2010).

Measuring food intake

Appetite is specific to foods, exhibits wide inter-subject variability, and tends to decline for a specific food as that food is eaten,
leading to selection of other foods. Appetite is therefore, said by Le Magnen to be sensory specific (Magnen, 2012). The sensory
specificity of appetite has been shown to relate inter alia to the postingestive consequences (satiation and satiety) of having ingested
a food. The most objective measure of appetite for a given food in a specific experimental situation is therefore the amount of that
food that a subject chooses to eat. Appetite is not rigidly determined by physiological signals per se although they may greatly influ-
ence it. Both the palatability of a food and the appetite for it tend to co-vary and are often increased subsequent to a period of nega-
tive energy balance. Two examples are dieting and illness, both of which lead to lowered intake and a subsequent rebound in
appetite. As discussed above, the appetite for a food will be learned on the basis of the consequences of having ingested that
food on previous eating occasions. Because of this it is possible to use covertly manipulated foods to deceive subjects into behaving
in a manner largely determined by prior learned experience. If this were not so, such deception would be impossible because the

Fig. 2 Subjective hunger tracked during waking hours in six subjects feeding on isoenergetically dense high-protein, high-fat, and high-carbohydrate
diets. Subjects exhibit the three peaks and troughs of hunger that typify the Western feeding schedule.
 Appetite: Psychobiological and behavioral aspects 5

physiological signals produced by the sensorially similar, yet nutritionally different food, would immediately translate (through
physiological signals) into behavioral compensation. This fact has implications for consumers because food technology can now
dissociate the sensory and nutritional properties of foods, which may undermine the learned basis of food intake control in
some people.

Measuring feeding behavior

There are many different techniques that can be used to measure feeding behavior in a number of different environmental situ-
ations. These techniques are used in naturalistic and/or laboratory studies of feeding. Generally in the laboratory, specific aspects
of the appetite and energy balance system are manipulated at the cognitive, sensory, gastrointestinal, or even the metabolic level,
for example, by deceiving subjects about the energy content of foods, altering the sensory variety of the diet, changing energy
expenditure through systematic alterations in physical activity, administering nasogastric infusions or parenteral infusions, respec-
tively. A number of other manipulations can be achieved. Because the environment can have such a large influence on feeding
behavior, it is important to consider a particular influence on feeding in several environmental contexts. For instance, the effects
of fat on energy intak have been considered in several laboratory and real life contexts. Under most sets of circumstances
increasing the energy density of foods using dietary fat appears to be a risk for excess intake. The relationship between the exper-
imental context and how it influences the investigations made is depicted in Fig. 3 (Blundell et al., 2010; Blundell and Stubbs,
2003; Stubbs et al., 1998).

Sensory stimulation and food hedonics

Food “liking” and “wanting” are emerging constructs in a conceptual approach to appetite where psychological processes of affect
and motivation can be viewed as major influences on food intake. Liking and wanting achieve importance in light of the recognition
of the contrast between homeostatic and hedonic processes that control eating.
The liking and wanting constructs stem from research exploring the neural basis of palatability and addictive behavior. With
principle focus on distinct dopamine and opioid pathways in the brain, the research suggests that processes of liking and wanting
can be separately manipulated to produce patterns of behavior that are either exclusively affective or motivational in conjunction
with a food stimulus. Liking and wanting are thought to reflect processes that can operate without conscious awareness. This
means that they have implicit components (Finlayson et al., 2007). However, their explicit counterparts express themselves
subjectively in the form of hedonic feelings from the ingestion of a specific food (i.e., explicit liking) and the intent or desire
to consume a specific food (i.e., explicit wanting). Under normal circumstances, explicit liking (“I like this”) is closely associated
with explicit wanting (“I want this”). However, there is also evidence to suggest that wanting can be “irrational”; i.e., when
implicit wanting for a food is greater than explicit wanting, and not proportional to experienced or expected liking (Finlayson
et al., 2007).
Liking and wanting appear to have separate and disproportionate roles in promoting overconsumption. In terms of liking, some
individuals at risk of weight gain may experience an exaggerated hedonic response to palatable foods, so that foods are enjoyed

Fig. 3 The constraints and limitations that the experimental environment places on studies of human feeding. In general the environment ranges
from totally free living, which is realistic but very difficult to make measurements into the laboratory where measurements are easy but may be
contaminated by artifacts due to the artificiality of the laboratory surroundings.
6 Appetite: Psychobiological and behavioral aspects

more and therefore eaten in greater amounts for longer periods of time. Conversely, susceptible individuals may have a diminished
ability to experience pleasure from food and therefore consumption of palatable food is driven up to satisfy an optimum level of
stimulation. Processes of wanting may also bring about vulnerability to weight gain through increased reactivity toward cues
signaling the availability of food. Moreover, a reduced ability to resist the motivation to eat when satiated may promote non-
homeostatic overconsumption. A widely held notion is that wanting rather than liking may be the crucial process in maintaining
an obese state. Moreover, food liking is often a rather stable characteristic within an individual and appears relatively uninfluenced
by increasing weight status. The implication is that liking may be important in establishing the motivational properties of food, but
once these are retained it is the upregulation of wanting in an obesogenic environment “insensitivity to homeostatic signals but
over-reactivity to external cues” that promotes overconsumption by influencing what and possibly how much is eaten from
moment to moment (Finlayson et al., 2007).

Sensory stimuli and body weight

It was originally proposed that obese subjects are more susceptible to external stimuli such as sensory stimuli than lean subjects who
were more reliant upon internal physiological cues. This predicted that in a Western context the numerous external food stimuli
would promote excess EI in susceptible individuals. As can be appreciated from much of this text, the interplay between “external”
and “internal” signal is much more complex and interactive than was initially supposed. Nevertheless, given the multiplicity of
afferent inputs that can influence feeding it is possible that some subjects have learned to base feeding predominantly on some
cues rather than others. However, dividing these cues into simply internal and external sources is perhaps oversimplified. Current
questionnaires which attempt to characterize subject’s responsiveness to food attempt to dissociate “externality,” “restraint,” and
“emotionality.” It is becoming increasingly clear that emotional and reward-based responses to food are critically important in facil-
itating over-consumption. A theoretical framework that integrates these eating behavior traits into coherent domains that have
methodological and predictive validity is still needed (Mela and Rogers, 2013).
Systematic differences have been found in sensory preference profiles, but not perceptions of intensity of various tastes
between lean and obese subjects. Work has suggested that subjects with a history of weight fluctuation have an enhanced
sensory preference for high-fat stimuli that are sweet. It has also been shown that sensory preference for fats is associated
with degree of overweight in adults and children. This may be important. Although there is little evidence that sensory
stimuli alone promote positive energy balances, there is evidence that people select foods they prefer. Preferential selection
of energy-dense foods can lead to excess EI without any apparent change in the amount of food eaten. Thus sensory factors
are likely to play a role in the selection of foods, which are conducive to weight gain (Blundell and Stubbs, 2003; Mela
and Rogers, 2013).

Sensory versus nutritional determinants of intake

The major problems with the concept of sensory preference or liking as determinants of hyperphagia and obesity in humans are that
(1) there is little direct evidence for this effect per se (because the appropriate experiments are very difficult to do) and (2) both
animals and humans appear to acquire sensory preferences for foods dense in readily available energy. Dissociation of the sensory
characteristics and postingestive consequences of ingesting a food becomes difficult and perhaps artificial. It seems that at the
present time the data from animal studies tends to suggest that maximal sensory preference for a food or diet is achieved when
the sensory stimulus is reinforced by the metabolic consequences that form part of the satiety sequence. Indeed, there is controver-
sial evidence that ingestion of one sensory stimulus (sweetness) without the associated nutrient (carbohydrate) promotes ingestion
of energy shortly afterward. The contribution of sensory and nutritional determinants of feeding is still poorly understood in
humans (Blundell and Stubbs, 2003; Mela and Rogers, 2013).

Emotional stimuli and over-consumption

As an intensely gregarious species we are bound together by numerous relationships ranging from mother–offspring bonds devel-
oped at birth, through kin relations to culturally defined positions in any given social structure. Positioning in social structures is
also common among the primates and is inextricably intertwined with access to resources, safety (e.g., predatory avoidance), and
food. This is critically important to the survival of groups of animals under ecological conditions. Humans have not lost the neuro-
anatomical bases of our emotions and behavior with which we evolved. Hence, food is as, if not more emotionally important today
as it has always been during our evolutionary heritage. However, the social and emotional meanings of food have changed. Food is
still important for sharing social emotions and creating social bonds through ceremonies, rituals, and traditions. Food is a major
source of reward for achievements or simply as treats. Interestingly, in children, there is evidence that more energy-dense foods are
more preferred. More energy-dense foods are also generally cheaper than less energy-dense but nutritionally balanced foods, making
the fast food outlet phenomenon self perpetuating. Food is a major source of soothing for emotional distress in others, for example,
when parents comfort children. In the context of this discussion it is notable that food is also a major source of reward, soothing,
and comfort in binge eaters, and is a common theme that emerges, in people struggling to control their weight. Thus, the emotional
drives and social facilitators of excess energy intake are powerful, if not more powerful than ever in modern Western society (De
Castro, 1997; Finlayson et al., 2007; Mela and Rogers, 2013).
 Appetite: Psychobiological and behavioral aspects 7

Meal patterns, appetite, and energy balance

The effect of meal patterns on appetite and energy balance is also an unresolved issue. It has been noted that snacking and commer-
cially available snack food are often believed to elevate EI. However, there is considerably less evidence that meal or snack patterns
contribute to the development of obesity. It is important to note at this point that the relationship between a meal and a snack
relates to timing and size of ingestive events in meal feeding animals. In nonhuman species (and indeed humans) that engage
in numerous small feeding bouts throughout their diurnal cycle there is little if any distinction between a meal and a snack.
Meal-feeding animals are conditioned to ingest the majority of their EI in a few large ingestive events in their diurnal cycle, at
approximately the same time points. Under these conditions, a snack can be defined as an energetically small, intermeal ingestive
event. To avoid confusion with a common use of the word to describe a certain type of “commercially available food,” we use the
phrase “commercially available snack foods” to describe those specific foods. Commercially available snack foods tend to differ
from the rest of the diet as they are more energy dense, high in fat and carbohydrate and low in protein and usually contain a large
fraction of their edible mass as dry matter. They are by no means the only food eaten as a small inter-meal ingestive event by many
people at large.
The evidence in relation to meal patterns, appetite, EI, and body weight is indirect and fragmentary. On aggregate, cross-
sectional studies tend to support no, or a negative, relationship between meal frequency and BMI. However it has been convinc-
ingly argued that examinations of the relationship between snacking and energy balance in free-living subjects are extensively
flawed by misreporting, misclassification of meals and snacks, and potentially by reverse causality. Under these conditions it
is difficult to draw clear conclusions about the effects of snacking in cross-sectional studies. It is therefore important to conduct
controlled laboratory interventions over a number of days in humans. These studies suggest that in the short-to-medium term
adding mandatory snacks to the diet leads to over-consumption. This effect is most pronounced in those who do not habitually
snack and least pronounced in those who do. It is also of note that rats tend to be “snackers” and Western humans tend to be meal
feeders. The rat tends to adjust EI by varying meal frequency; the human by varying meal size. However, if rats are meal fed, they
learn to adjust EI by varying meal size. Humans placed in time isolation begin to adjust intake by varying meal frequency. These
comparisons illustrate the fact that adjustment of intake to energy or nutrient requirements occurs within a conditioned time
framework, which itself is variable depending on the conditioning environment. Despite large changes in the pattern of feeding,
EI can still be adjusted to satisfy requirements.

Social and situational influences on feeding behavior

There are a number of social and situational influences on food intake in humans. In general, the shorter the time period of
measurement, the greater the effect of situational and social influences. Thus there are a large number of factors that can influence
single meal size in humans. These factors are summarized in Fig. 4.
Time of day appears to influence meal size in that the amount eaten and the EI increases on going from breakfast, to lunch, to
the evening meal. Meal size also increases across the feeding period in rats. It has been suggested that this occurs in learned antic-
ipation of the energy requirements in the fasting period (night for humans, day for rats). Meal size and EI tend to be greater at
weekends than on weekdays, in Western adults. Meal size also varies as a power function of the number of people present at
a meal. DeCastro has termed this effect “social facilitation” of feeding. Social facilitation and daily routine account for much
of this effect.
Seasonality can influence feeding. A number of studies also suggest that EI, meal size, and eating rate are all elevated in the
autumn. In one particular study hunger was associated with meal size in winter and spring, but not so clearly in the autumn
(Mela and Rogers, 2013; Stubbs, 2014).

Cognitive and social cues

Throughout the 1960s and 1970s a large number of behavioral studies examined the effect of cognitive and social cues (perceived
energy content of foods, salience of cues, eating behavior of others present) on feeding in relation to the externality hypothesis.
Although a large number of studies found that so called external cues do relate to short-term feeding behavior, a large number
of others did not. However, the presence of external cues alone does not reliably predict how much food people will eat. Neither
does the presence of external cues always relate to lean/obese differences in feeding patterns. Some of these differences in relation to
cognitive and social cues are better explained in relation to dietary restraint. Restraint is a term used to describe people who are
attempting to limit or reduce their body weight by means of cognitive energy restriction (dieting). In doing so it is proposed
that they are placing their motivation in relation to feeding at odds with physiological feeding stimuli. Placing cognition at
odds with physiological drives can result in pathologies of eating since the normal “regulatory” processes are cognitively under-
mined. Furthermore it is argued that restraint will increase the probability that a person will break a diet. It has been shown
that an intervention (usually a preload) that breaks the rules of restraint, almost paradoxically induces a greater intake. This
phenomenon has been termed counter-regulation. This effect is cognitive, because it can be induced by deceiving a restrained eater
into believing that a preload was high in calories (Mela and Rogers, 2013). Because the concept of restraint has predictable behavioral
outcomes it is a useful tool in characterizing different people with respect to their feeding behavior. However, it is now generally
8 Appetite: Psychobiological and behavioral aspects

Fig. 4 Major factors known to affect single meal size in humans. In general, the shorter the time interval of measurement, the greater the influence
these factors have on the observed feeding behavior.

accepted that restraint is not a unitary construct and people who attempt to lose weight and who show flexible restraint are more
successful than those who are rigid in their patterns of cognitive restraint.
It is useful to consider the role of dual process models of behavior in the context of energy balance (weight change). Models of
behavior and behavior change have focused historically on social cognition (e.g., beliefs, intentions, attitudes and decisions),
emphasizing pathways of reasoned action in which pre-decisional motivation leads to the formation of intentions and the imple-
mentation of those intentions as volitional action. Automatic processes (emotions, desires, habits resulting from associative
learning and physiological states) may also impact eating behavior and behavior change. These processes tend to be relatively rapid,
impulsive (less conscious) and habitual in comparison to the slow, deliberative processes of motivation and cognitive self-
regulation. Furthermore, in the context of appetite and energy balance behaviors, the development of self-regulatory behavior
change is effortful, particularly in the face of physiological responses to weight loss, while unconscious or automatic components
of EB behaviors are rapid and effortless. Physiological mediators of homeostatic and hedonic appetitive drives, and changes in phys-
ical activity that are triggered by weight loss may feed into such automatic process of behavior change to undermine self-regulation
of eating behavior.

Diet composition and appetite

Food and nutrient ingestion influence human appetite through multiple feedbacks at several levels, which can be traced through the
processes of food location, ingestion, digestion, absorption, and metabolism. Satiety is therefore maintained by a functional
sequence or cascade of sequential physiological events that reinforce each other. Removing parts of a food or nutrient’s effects
on this sequence will therefore diminish its impact on satiety.

How do macronutrients and energy density affect satiety?

Because the control of food intake in humans is imprecise, alterations in the energy density of the diet can lead to changes in energy
intake, even over quite prolonged periods. Macronutrients are not all equal when it comes to how satisfied one feels after eating
them. Protein has the greatest effect on satiety, followed by carbohydrate, followed by fat. Studies using smaller preloads (typically
of 250 kcal or less) do not always find this effect. Although fat is the most energy-dense macronutrient it is also the least satisfying.
The effect macronutrients have on satiety is illustrated in Fig. 5. Foods that are high in protein include lean meat, fish, beans, peas,
lentils, mycoprotein, tofu, and eggs. Foods rich in carbohydrates and fiber include wild rice, potatoes, cereals and oats, bread, beans,
peas, lentils, and couscous. Fat-rich foods include savory snacks, pies, biscuits, cakes, cream, and most cheese (Stubbs et al., 2010).

Diet composition and satiety to prevent weight gain

Any weight reduction strategy that focuses on one simple aspect of diet alone will have limited effects on weight control. Dietary mono-
therapies are very limited in what they will achieve. Several aspects of diet composition can be used simultaneously to influence
 Appetite: Psychobiological and behavioral aspects 9

Fig. 5 Effect of increasing energy content of macronutrient loads on satiety Index subjectively expressed over 3.25 h.

satiety and help people navigate to a healthy body weight. In other words, we should be taking a package definition of what consti-
tutes a reasonable dietary approach to weight management and that should be integrated with evidence-based behavior change
approaches. Dietary factors that help with self-management of eating behavior include a low fat content, tolerably low energy
density of 1.1–1.3 kcal g 1, a high water content; avoiding where possible, caloric beverages, fiber content should be tolerably
high (which parenthetically is not much more than 20–30 g per day). Fiber holds water in foods, which lowers energy density,
and decreases the rates at which those foods are digested. To enhance satiety, protein content of the diet can be relatively, tolerably,
and reasonably high (probably not exceeding 25–30% of an energy-reduced diet, for example, a diet that is followed to induce
a negative energy balance. This translates into less than 20–25% of energy requirements from protein). Protein intake from red
and processed meats should be limited. The carbohydrate content depends very much on type. There is a good deal in the literature
that suggests sweet, short-chain carbohydrates can elevate energy intake, especially when combined with fat. This combination is
most common in commercial processed and snack foods. The orosensory properties of foods need to be maintained to encourage
people to select those foods, so that they become a practical option for the development of healthy eating habits (Stubbs et al.,
2010).

Conclusion

Human appetite is influenced by both physiological and psychological cues which themselves are shaped by the environment.
Appetite control and energy balance regulation are geared to defend against weight loss and less tightly geared to defend against
weight gain. Many of the environmental eating-related cues that are part of modern society also act as stimuli that promote higher
levels of food and energy intake, reinforced by an interaction of the sensory properties of foods with liking and wanting circuits.
Because many aspects of eating behavior are learned through associative conditioning they are often habitual and not under imme-
diate, direct conscious control. These factors facilitate weight gain in modern environments, but may also provide some potential
opportunities for obesity prevention and long-term management of body weight. Gaining weight and excess adipose tissue across
the adult lifespan requires little conscious effort. Losing weight and preventing weight regain requires considerable effort and weight
loss attempts are often subject to subsequent weight regain. Because most of energy intake and less than half of energy expenditure is
behavior, appetite control and self-management of eating behaviors are major targets for many weight management interventions.
The effects of behavior change interventions on weight management are still relatively modest and our understanding of the factors
that disrupt and undermine self-management of eating behaviors is limited. These factors include physiological resistance to weight
loss, gradual compensatory changes in eating and physical activity and reactive processes related to stress, emotions, rewards and
desires that influence eating to meet psychological needs. In the future it is likely that more interdisciplinary research between the
areas of appetite control, energy balance and behavior change research, together with methodological developments in these areas
10 Appetite: Psychobiological and behavioral aspects

will lead to more effective interventions to improve weight and health outcomes in the general population. Better matching
evidence-based intervention content to the needs of individuals and better objective tracking of appetite and energy balance behav-
iors may improve outcomes in the future.

See Also: Dietary intake measurement: Methodology; Food choice: Behavioral aspects

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