CHAPTER

Motivation: What have we

learned and what is still
missing?
19
B. Studer1, S. Knecht
Mauritius Hospital, Meerbusch, Germany
Institute of Clinical Neuroscience and Medical Psychology, Medical Faculty, Heinrich-Heine-
€
University Dusseldorf, €
Dusseldorf, Germany
1
Corresponding author: Tel.: +49-2159-679-5114; Fax: +49-2159-679-1535,
e-mail address: bettina.studer@stmtk.de

Abstract
This final chapter deliberates three overarching topics and conclusions of the research pre-
sented in this volume: the endurance of the concept of extrinsic vs intrinsic motivation, the
importance of considering subjective costs of activities when aiming to understand and en-
hance motivation, and current knowledge of the neurobiological underpinnings of motivation.
Furthermore, three topics for future motivation research are outlined, namely the assessment
and determinants of intrinsic benefits, the reconciliation of activity-specific motivation models
with generalized motivation impairments in clinical populations, and the motivational dynam-
ics of groups.

Keywords
Motivation, Conclusions, Open questions, Extrinsic, Intrinsic, Brain, Costs, Groups

The chapters of this volume provided an up-to-date account of motivation theory,

demonstrated novel directions in the assessment of motivation and its determinants
in healthy and clinical populations, presented recent findings on the neurobiological
correlates of motivation and goal-directed behavior, and discussed and tested novel
strategies on enhancing motivation in a variety of application domains. In this final
chapter, we briefly reflect on some overarching topics and insights emerging from
the research presented in the chapters of this volume and outline three topics for fu-
ture motivation research.

Progress in Brain Research, Volume 229, ISSN 0079-6123, http://dx.doi.org/10.1016/bs.pbr.2016.07.001

© 2016 Elsevier B.V. All rights reserved.
441
442 CHAPTER 19 Concluding remarks

1 THREE OVERARCHING INSIGHTS GAINED THROUGH

THIS VOLUME
1.1 ENDURANCE OF THE CONCEPT OF EXTRINSIC VS INTRINSIC
MOTIVATION
The distinction between intrinsic and extrinsic (sources of) motivation (eg, Deci,
1971; Deci and Ryan, 1985; White, 1959) has a long tradition in psychology (for
reviews, see Strombach et al., 2016, this volume; Oudeyer et al., 2016, this
volume) and intuitive appeal. The persistence of this proposition in the field of mo-
tivation research is detectable in several chapters of this volume. For instance, the
benefit–cost framework of motivation for specific activity that we propose in
Studer and Knecht (2016, this volume) distinguishes intrinsic and extrinsic benefits
(and costs), with intrinsic benefits defined as positive feelings experienced during
performance of the activity itself (eg, enjoyment, pleasure, feeling of accomplish-
ment), and extrinsic benefits defined as instrumental gains, rewards, or goals
achieved through performance of the activity. Nafcha et al. (2016, this volume) re-
view previous evidence that (experience of ) personal control over ones environment
is intrinsically rewarding and can drive behaviors with no or only small extrinsic ben-
efits. They postulate that this intrinsic benefit of control is what motivates habitual
behavior, and describe recent empirical findings supporting this hypothesis.
Losecaat Vermeer et al. (2016, this volume) and we (Studer et al., 2016, this
volume) argue that competing against an opponent—or oneself—might carry both
an intrinsic benefit (ie, challenge is enjoyable) and an extrinsic benefit
(ie, winning against someone, or improving once own performance, is rewarding).
The functional magnetic resonance imaging (fMRI) study by Widmer et al.
(2016, this volume) investigated and compared the impact of an extrinsic monetary
reward and performance feedback as an “intrinsic reward” upon motor skill learning
and neural activations. They found that monetary reward and randomly presented
performance feedback, but not performance feedback provided systematically for
good training trials only, were associated with stronger activation of the ventral stri-
atum and better overnight skill consolidation. The interplay between extrinsic, exter-
nally set incentives, intrinsic motivation, and behavioral output, was discussed in
depth by Strang et al. (2016, this volume). This topic was controversially debated
in the extant theoretical and application-focused literature: on one hand, decades
of behavioral research has shown that extrinsic incentives (for instance, food re-
wards) are effective in modulating behavior in animals and humans (eg, Morales
et al., 2016, this volume; Skinner, 1963; Toppen, 1965), and performance-dependent
compensation and other monetary incentives schemes are used widely in our society
(eg, Gerhart and Fang, 2015). On the other hand, it has been argued that extrinsic,
externally set incentives diminish intrinsic motivation for a target activity and could
therefore negatively impact performance of that activity (eg, Deci, 1971; Deci et al.,
1999; Lepper et al., 1973). Through a careful review of previous evidence,
Strang et al. clarify under which conditions extrinsic, externally set incentives
 1 Three overarching insights gained through this volume 443

positively affect behavior and overall motivation. They concluded that extrinsic in-
centives are most effective in situations where intrinsic motivation (or in other
words, the anticipated intrinsic benefit) is low, the target activity is easy and when
extrinsic benefits are of a social nature (eg, positive verbal feedback). Meanwhile,
extrinsic incentives are less effective, or may even reduce performance of a target
activity, when intrinsic motivation is high and when the target activity is prosocial
behavior. With regard to potential application, it is also noteworthy that removal of
extrinsic incentives often leads to a drop in performance. If used for motivation en-
hancement in long-term interventions (eg, health facilitation), externally set incen-
tives might thus have to be sustained over long periods.

1.2 THE IMPORTANCE OF COSTS

A second overarching conclusion that can be drawn from the work presented in this
volume is that motivation theories, particularly those focusing on motivation for spe-
cific activities, should consider not only the subjective benefits (classically referred
to as motivators) but also the (intrinsic and extrinsic) subjective costs of an activity.
In some psychological theories of motivation (eg, in self-determination theory; Deci,
1980; Deci and Ryan, 2000) as well as in many of the questionnaire-based measures
used to investigate and quantify motivation in healthy and clinical human popula-
tions, the cost dimension has received little attention. Meanwhile, the trade-off be-
tween benefits and costs is a core topic in (neuro-)economic models of human
motivation and behavior (eg, Strang et al., this volume) and motivation research in an-
imals (Hull, 1943; Morales et al., 2016, this volume). Building upon this tradition, re-
cent neuroscience research has often used behavioral arrays of motivation, which test
the willingness of a human or animal to exert physical effort (as an intrinsic cost) for
extrinsic rewards of different magnitudes (see, for instance, Bernacer et al., 2016, this
volume; Morales et al., 2016, this volume; Salamone and Correa, 2012; Salamone
et al., 2007). As reviewed by Chong et al. (2016, this volume), these behavioral mea-
sures have proven very useful and informative in identifying neurobiological under-
pinnings of motivation, are sensitive to individual differences, and hold great
promise as more sensitive tools for the assessment and dissociation of motivation im-
pairments in different clinical conditions. Convergent, a novel study by Bernacer et al.
(2016, this volume) demonstrated that subjective effort costs of treadmill running dif-
ferentiates between individuals with an active lifestyle and individuals with a seden-
tary lifestyle. Relatedly, Kroemer et al. (2016, this volume) discuss how a high body
mass index makes movement metabolically more costly, which is likely to increase
perceived effort and thus decrease motivation for physical activity in obese individ-
uals. As a final example, Morales et al. (2016, this volume) successfully used a behav-
ior paradigm with an effort component to investigate the role of opioid signals in
different aspects of food motivation. Together these chapters demonstrate how consid-
ering subjective expected costs of a target activity, alongside of subjective expected
benefits, is extremely useful for understanding an individual’s motivation for perfor-
mance of that activity and how to enhance it.
444 CHAPTER 19 Concluding remarks

1.3 MOTIVATION IN THE BRAIN: DOPAMINE AND BEYOND

A third overarching topic of this volume is the neurobiological underpinnings of mo-
tivation and goal-directed behavior. Through careful reviews of extant neuroscience
findings and novel original investigations, the chapters included in Section 3 (and
elsewhere) of this volume provided an up-to-date account of the brain structures
and neurotransmitter systems implicated in motivation and goal-directed behavior.
Let us briefly recount some core insights. First, with regard to brain structures, extant
neuroscience research revealed that a distributed network of brain areas, including
the midbrain, ventral and dorsal striatum, pallidum, prefrontal cortex, anterior cin-
gulate cortex, and basolateral amygdala, is implicated in the computation of the sub-
jective expected benefits and costs of an actions and benefits–costs comparison. As
reviewed in Chong et al. (2016, this volume) and Kroemer et al. (2016, this volume),
lesions of structures of this network, in particular of the nucleus accumbens, medial
prefrontal cortex, and amygdala, reduce willingness to work for a reward and lead to
a preference of actions with low effort costs. Lesions to areas of this network also
affect sensitivity to the (extrinsic) benefits or rewards outside of an effort context
(eg, Clarke et al., 2008; Coutureau et al., 2009; Manohar and Husain, 2016;
Studer et al., 2015). These lesion results are corroborated by a large body of func-
tional neuroimaging studies on decision making. For instance, as Bernacer et al.
(2016, this volume) point out, fMRI studies using effort-based decision-making par-
adigms find that activity patterns in the striatum, anterior cingulate, and ventrome-
dial prefrontal cortex, as well as the motor cortex and supplementary motor areas,
reflect the level of required physical effort associated with choice options
(eg, Croxson et al., 2009; Prevost et al., 2010). In their own novel study, Bernacer
and colleagues also find sensitivity for required effort in the ventrolateral prefrontal
cortex. Furthermore, as we review in Studer and Knecht (2016, this volume), activity
patterns in this network were found to covary with factors determining the expec-
tancy and value of extrinsic rewards and their integrated overall benefit. Finally,
Kroemer et al. (2016, this volume) presented an intriguing neurocomputational
model in which fluctuations in motivation and behavior over time are caused by sig-
nal fluctuations in key nodes of this network.
Second, several chapters examined the role of the neurotransmitter dopamine in
motivation. As pointed out by Chong and Husain (2016, this volume), dopaminergic
neocortical and nigrostriatal pathways are at the core of the network described ear-
lier. And, a large body of research in nonhuman animals shows that (systemic or
region-targeted) disruption of dopaminergic transmission leads to a decreased mo-
tivation to work for a given reward, or in other words, a shift toward a reduced benefit
and increased costs evaluation (see Chong and Husain, 2016, this volume; Kroemer
et al., 2016, this volume for reviews). Chong and Husain (2016, this volume) rea-
soned that this implied central role of dopamine in signaling (extrinsic) incentives
to act and in facilitating the overcoming of effort costs is consistent with the finding
that apathy is common in neurological disorders affecting the dopaminergic system
(such as Parkinson disease). Kroemer and colleagues (2016, this volume) highlighted
 2 Outlook: Three topics for future research 445

the convergence between the aforementioned results from animal studies and recent
human data on dopaminergic transmission during an effort-based decision-making
paradigm by Treadway et al. (2012). But, Kroemer et al. also point out that while
the importance of dopaminergic signaling for effort-related aspects of motivation
is now well established, much less is known about the neurobiological substrates
of other motivational dimensions. Relevant to this point is the novel study by
Morales et al. (2016, this volume) on the functions of opioid signaling in food mo-
tivation, which provides a highly interesting addition to previous knowledge. Mo-
rales and colleagues build on an argument originally made by Salamone and
colleagues (Salamone and Correa, 2012) that dopamine is primarily implicated
in activational and directional aspects of food motivation (or in other words, instru-
mental behavior), rather than in hedonic aspects of food consumption. In two well-
designed experiments, Morales et al. tested the effects of the opioid receptor an-
tagonist naloxone upon instrumental and consummatory behavior in rats. They find
that disruption of opioid signaling reduces rats’ liking of a preferred food and de-
crease their willingness to lever press (ie, exert effort) for that preferred food. This
suggests that the opioid system might be involved in computing anticipated plea-
sure of food consumption, or more generally speaking, expected hedonic value
(ie, intrinsic benefit) of an activity.
Finally, this volume also demonstrated how knowledge of the neurobiological
underpinnings may be used for clinical and nonclinical application. For instance,
in their fMRI study, Widmer et al. (2016, this volume) found that providing concur-
rent performance feedback and monetary reward during a motor learning task raised
ventral striatum activation, and that stronger responsiveness in the striatum to these
incentives was associated with better overnight skill consolidation. These results
suggest that increasing ventral striatal activity during motor training through verbal
or monetary reward could help improve consolidation of the motor skill. Further,
Chong and Husain (2016, this volume) argue that the aforementioned findings from
animal research on dopaminergic functions in effort-related motivation imply dopa-
mine agonists as a primary candidate for pharmacological treatment of apathy.
Reviewing extant research using this intervention, they conclude that the studies con-
ducted to date offer some evidence for (selective) dopamine agonist therapy being
effective in ameliorating apathy in human patients, but also highlight that more
well-controlled clinical studies are required.

2 OUTLOOK: THREE TOPICS FOR FUTURE RESEARCH

2.1 UNDERSTANDING AND ENHANCING THE INTRINSIC BENEFIT
OF AN ACTIVITY
Intrinsic motivation—or, in the terminology of the benefit–cost framework proposed
in Studer and Knecht (2016, this volume), the intrinsic benefit of an activity—is a
popular topic in the psychological, educational, rehabilitation, and healthcare
446 CHAPTER 19 Concluding remarks

literature. And, what determines intrinsic motivation and how intrinsic motivation
can be enhanced remain two hot topics in current motivation research (see for in-
stance Nafcha et al., 2016, this volume; Oudeyer et al., 2016, this volume). Yet, the
vast majority of extant studies investigating the neural correlates of motivation
have used extrinsic incentives. Therefore, intrinsic benefits and their neurobiolog-
ical underpinnings remain less well understood. Furthermore, although several be-
havioral (eg, for how long an activity is performed during a free-choice period) and
questionnaire assays of intrinsic motivation have been established through labora-
tory and field studies (eg, Deci et al., 1999; Pelletier et al., 1995; Vallerand et al.,
1992), intrinsic benefits or motives such as enjoyment, curiosity, control over the
environment, novelty (perceived), competence, and interest are—in our opinion—
more difficult to identify, quantify, and understand in real-life scenarios than ex-
trinsic benefits, due to their more abstract nature. We hope that future research will
continue to shed light on the determinants of intrinsic benefits and how intrinsic
motives can be integrated into current neurobiological and (neuro-)economic
models of human motivation.

2.2 ACTIVITY-SPECIFIC MODELS VS GENERALIZED MOTIVATION

IMPAIRMENTS IN CLINICAL POPULATIONS
Most chapters of this volume discussed motivation in specific contexts and for
specific activities or tasks and assumed that the degree of motivation in such sit-
uation is determined by (subjective evaluation of) activity-specific aspects
(ie, possible rewards, effort requirements, perceived autonomy and control,
etc.). One open question is how such activity-specific models of motivation,
and the findings obtained through research utilizing these models, can be recon-
ciled with the more generalized (ie, situation-unspecific) and enduring motivation
deficits observed in clinical contexts (including apathy and the motivation-
affecting syndromes depression and fatigue). One current proposition is that such
disorders are characterized by a systematic shift in the subjective evaluation of
benefits and/or of (effort) costs, and/or a change in the benefit–cost comparison,
and that such alterations could be captured and dissociated through use of effort-
based decision-making paradigms (see Chong et al., 2016, this volume; Chong and
Hussain, 2016, this volume). As reviewed by Chong and colleagues, results from
first studies using this methodology in patient samples are promising; however,
how suitable and insightful such effort-based decision-making paradigms are as
diagnostic tools in real-life clinical application has yet to be established. Future
research should also further investigate how different neurobiological correlates
of these clinical conditions, for instance hyperarousal in depression, hypoarousal
in fatigue (Hegerl and Ulke, 2016, this volume), and dopaminergic dysfunctions in
Parkinson disease and other neurological conditions with high prevalence of ap-
athy (Chong and Husain, 2016, this volume), relate to dissociable or shared alter-
ations in motivation.
 References 447

2.3 SOCIAL INFLUENCES ON MOTIVATION AND GROUP BEHAVIOR

Psychological theories and contemporary economic models recognize the impor-
tance of social factors in determining motivation and guiding behavior. Social ap-
proval, fairness, reciprocity, social status, and reputation have all been postulated
to be strong motivators of human behavior (see Losecaat Vermeer et al., 2016,
this volume; Strombach et al., 2016, this volume, for reviews of the relevant litera-
ture). Furthermore, social comparison is known to affect the subjective valuation of
extrinsic benefits and costs (eg, Báez-Mendoza and Schultz, 2013; Bault et al., 2011;
Grygolec et al., 2012) and perception of one’s own skill and performance (Corcoran
et al., 2011; Vostroknutov et al., 2012). Previous research has also shown that
humans do not only take their own benefits and costs into account, but also what
benefits and costs their actions bring for others (eg, Crockett et al., 2014). Note, how-
ever, that these lines of research focus on how social factors affect motivation and
behavior of an individual. A topic that received less attention in this volume, and in
neuroscience research on motivation in general, is what drives motivation and behav-
ior of groups. Of course, a group can simply be seen as multiple individuals, each
with their own motivation. But in every-day life, we also observe situations where
the motivation and behavior of group members appear to arise, or at least be en-
hanced, though an interactive dynamic. Think for instance of eruptions of violence
in crowds of soccer supporters. Such group dynamics are deliberated in the social
psychology literature. For instance, it has been shown that group interaction en-
hances individuals’ propensity to take risks (“risky-shift phenomenon”; Kogan and
Wallach, 1967) and polarize individuals’ attitudes (eg, Keating et al., 2016; Myers
and Lamm, 1976). Future research might examine how such group dynamics fit into
current psychological and economic models of motivation, and how they might be uti-
lized for motivation enhancement. Further, it would be interesting for future studies to
assess the neurobiological correlates of such interactional social influences upon mo-
tivation and behavior.

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