Neurobiology of Sensation and Reward (Frontiers in Neuroscience)
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Rewards in operant conditioning are positive reinforcers. Operant behavior gives a good definition for rewards. Anything that makes an individual come back for more is a positive reinforcer and therefore a reward.
Although it provides a good definition, positive reinforcement is only one of several reward functions. Rewards are attractive. They are motivating and make us exert an effort. Rewards induce approach behavior, also called appetitive or preparatory behavior, and consummatory behavior.
Thus any stimulus, object, event, activity, or situation that has the potential to make us approach and consume it is by definition a reward.
Rewarding stimuli, objects, events, situations, and activities consist of several major components. First, rewards have basic sensory components visual, auditory, somatosensory, gustatory, and olfactory A separate form not included in this scheme, incentive salience, primarily addresses dopamine function in addiction and refers only to approach behavior as opposed to learning Third, rewards have a value component that determines the positively motivating effects of rewards and is not contained in, nor explained by, the sensory and attentional components FIGURE 1, right. This component reflects behavioral preferences and thus is subjective and only partially determined by physical parameters.
Only this component constitutes what we understand as a reward. Rewards can also be intrinsic to behavior 31, , They contrast with extrinsic rewards that provide motivation for behavior and constitute the essence of operant behavior in laboratory tests. Intrinsic rewards are activities that are pleasurable on their own and are undertaken for their own sake, without being the means for getting extrinsic rewards.
Intrinsic rewards are genuine rewards in their own right, as they induce learning, approach, and pleasure, like perfectioning, playing, and enjoying the piano. Although they can serve to condition higher order rewards, they are not conditioned, higher order rewards, as attaining their reward properties does not require pairing with an unconditioned reward. These emotions are also called liking for pleasure and wanting for desire in addiction research and strongly support the learning and approach generating functions of reward.
Dialogues Clin. Despite the importance of numerous psychosocial factors, at its core, drug addiction involves a biological process: the ability of repeated exposure to a drug of abuse to induce changes in a vulnerable brain that drive the compulsive seeking and taking of drugs, and loss of control over drug use, that define a state of addiction.
Moreover, there is increasing evidence that, despite a range of genetic risks for addiction across the population, exposure to sufficiently high doses of a drug for long periods of time can transform someone who has relatively lower genetic loading into an addict. Mount Sinai School of Medicine. Department of Neuroscience. Retrieved 9 February Substance-use disorder: A diagnostic term in the fifth edition of the Diagnostic and Statistical Manual of Mental Disorders DSM-5 referring to recurrent use of alcohol or other drugs that causes clinically and functionally significant impairment, such as health problems, disability, and failure to meet major responsibilities at work, school, or home.
Depending on the level of severity, this disorder is classified as mild, moderate, or severe. Addiction: A term used to indicate the most severe, chronic stage of substance-use disorder, in which there is a substantial loss of self-control, as indicated by compulsive drug taking despite the desire to stop taking the drug. In the DSM-5, the term addiction is synonymous with the classification of severe substance-use disorder.
In the prefrontal cortex, recent evidence indicates that the [orbitofrontal cortex] OFC and insula cortex may each contain their own additional hot spots D. Castro et al. In specific subregions of each area, either opioid-stimulating or orexin-stimulating microinjections appear to enhance the number of liking reactions elicited by sweetness, similar to the [nucleus accumbens] NAc and [ventral pallidum] VP hot spots.
Successful confirmation of hedonic hot spots in the OFC or insula would be important and possibly relevant to the orbitofrontal mid-anterior site mentioned earlier that especially tracks the subjective pleasure of foods in humans Georgiadis et al. A brainstem mechanism for pleasure may seem more surprising than forebrain hot spots to anyone who views the brainstem as merely reflexive, but the pontine parabrachial nucleus contributes to taste, pain, and many visceral sensations from the body and has also been suggested to play an important role in motivation Wu et al.
Medical News Today. Retrieved 14 November Social Cognitive and Affective Neuroscience. An Introduction to Brain and Behavior 1st ed. New York: Worth. These glutamatergic inputs make contact on the heads of dendritic spines of the striatal GABAergic medium spiny projection neurons MSNs whereas dopaminergic inputs synapse onto the spine neck, allowing for an important and complex interaction between these two inputs in modulation of MSN activity It should also be noted that there is a small population of neurons in the [nucleus accumbens] NAc that coexpress both D1 and D2 receptors, though this is largely restricted to the NAc shell Bertran- Gonzalez et al.
Neurons in the NAc core and NAc shell subdivisions also differ functionally. The NAc core is involved in the processing of conditioned stimuli whereas the NAc shell is more important in the processing of unconditioned stimuli; Classically, these two striatal MSN populations are thought to have opposing effects on basal ganglia output.
These findings suggest that iMSNs can bidirectionally modulate drug reward. Together these data suggest that iMSNs normally act to restrain drug-taking behavior and recruitment of these neurons may in fact be protective against the development of compulsive drug use. Subst Abuse Rehabil. Regions of the basal ganglia, which include the dorsal and ventral striatum, internal and external segments of the globus pallidus, subthalamic nucleus, and dopaminergic cell bodies in the substantia nigra, are highly implicated not only in fine motor control but also in [prefrontal cortex] PFC function.
Thus, only a brief description of the modulatory role of the basal ganglia in addiction-relevant circuits will be mentioned here. The overall output of the basal ganglia is predominantly via the thalamus, which then projects back to the PFC to form cortico-striatal-thalamo-cortical CSTC loops. Three CSTC loops are proposed to modulate executive function, action selection, and behavioral inhibition. In the dorsolateral prefrontal circuit, the basal ganglia primarily modulate the identification and selection of goals, including rewards.
Studies have shown that cravings are underpinned by activation of the reward and motivation circuits McBride et al. According to these authors, the main neural structures involved are: the nucleus accumbens, dorsal striatum, orbitofrontal cortex, anterior cingulate cortex, dorsolateral prefrontal cortex DLPFC , amygdala, hippocampus and insula. Sydor A, Brown RY eds. The neural substrates that underlie the perception of reward and the phenomenon of positive reinforcement are a set of interconnected forebrain structures called brain reward pathways; these include the nucleus accumbens NAc; the major component of the ventral striatum , the basal forebrain components of which have been termed the extended amygdala, as discussed later in this chapter , hippocampus, hypothalamus, and frontal regions of cerebral cortex.
These structures receive rich dopaminergic innervation from the ventral tegmental area VTA of the midbrain. Addictive drugs are rewarding and reinforcing because they act in brain reward pathways to enhance either dopamine release or the effects of dopamine in the NAc or related structures, or because they produce effects similar to dopamine. A macrostructure postulated to integrate many of the functions of this circuit is described by some investigators as the extended amygdala.
The extended amygdala is said to comprise several basal forebrain structures that share similar morphology, immunocytochemical features, and connectivity and that are well suited to mediating aspects of reward function; these include the bed nucleus of the stria terminalis, the central medial amygdala, the shell of the NAc, and the sublenticular substantia innominata. Figure 3: Neural circuits underlying motivated 'wanting' and hedonic 'liking'.
Towards a neuroscience of active sampling and curiosity | Nature Reviews Neuroscience
Pharmacology Biochemistry and Behavior. Nature Reviews. Brain Research. The Journal of Neuroscience. Frontiers in Systems Neuroscience. Annual Review of Psychology. Scientific American. Bibcode : SciAm. Retrieved 17 January So it makes sense that the real pleasure centers in the brain — those directly responsible for generating pleasurable sensations — turn out to lie within some of the structures previously identified as part of the reward circuit.
One of these so-called hedonic hotspots lies in a subregion of the nucleus accumbens called the medial shell. A second is found within the ventral pallidum, a deep-seated structure near the base of the forebrain that receives most of its signals from the nucleus accumbens. On the other hand, intense euphoria is harder to come by than everyday pleasures. The reason may be that strong enhancement of pleasure — like the chemically induced pleasure bump we produced in lab animals — seems to require activation of the entire network at once.
Defection of any single component dampens the high.
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Whether the pleasure circuit — and in particular, the ventral pallidum — works the same way in humans is unclear. Here I discuss how mesocorticolimbic mechanisms generate the motivation component of incentive salience. Incentive salience takes Pavlovian learning and memory as one input and as an equally important input takes neurobiological state factors e. Neurobiological state changes can produce unlearned fluctuations or even reversals in the ability of a previously learned reward cue to trigger motivation. Such fluctuations in cue-triggered motivation can dramatically depart from all previously learned values about the associated reward outcome.
Associative learning and prediction are important contributors to motivation for rewards.
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Learning gives incentive value to arbitrary cues such as a Pavlovian conditioned stimulus CS that is associated with a reward unconditioned stimulus or UCS. Learned cues for reward are often potent triggers of desires. For example, learned cues can trigger normal appetites in everyone, and can sometimes trigger compulsive urges and relapse in addicts.
This is a signature feature of incentive salience. This cue attraction is another signature feature of incentive salience VTA DA neurons play a critical role in motivation, reward-related behavior Chapter 15 , attention, and multiple forms of memory. This organization of the DA system, wide projection from a limited number of cell bodies, permits coordinated responses to potent new rewards.
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Thus, acting in diverse terminal fields, dopamine confers motivational salience "wanting" on the reward itself or associated cues nucleus accumbens shell region , updates the value placed on different goals in light of this new experience orbital prefrontal cortex , helps consolidate multiple forms of memory amygdala and hippocampus , and encodes new motor programs that will facilitate obtaining this reward in the future nucleus accumbens core region and dorsal striatum. In this example, dopamine modulates the processing of sensorimotor information in diverse neural circuits to maximize the ability of the organism to obtain future rewards.
The brain reward circuitry that is targeted by addictive drugs normally mediates the pleasure and strengthening of behaviors associated with natural reinforcers, such as food, water, and sexual contact. Dopamine neurons in the VTA are activated by food and water, and dopamine release in the NAc is stimulated by the presence of natural reinforcers, such as food, water, or a sexual partner. As previously mentioned, the activity of dopaminergic neurons in the VTA appears to be linked to reward prediction. The NAc is involved in learning associated with reinforcement and the modulation of motoric responses to stimuli that satisfy internal homeostatic needs.
The shell of the NAc appears to be particularly important to initial drug actions within reward circuitry; addictive drugs appear to have a greater effect on dopamine release in the shell than in the core of the NAc. Current Opinion in Neurobiology. For instance, mesolimbic dopamine, probably the most popular brain neurotransmitter candidate for pleasure two decades ago, turns out not to cause pleasure or liking at all. Rather dopamine more selectively mediates a motivational process of incentive salience, which is a mechanism for wanting rewards but not for liking them Bibcode : PNAS..
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