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METALEARNING, NEUROMODULATION, AND EMOTION
Information Sciences Division, ATR International
CREST, Japan Science and Technology Corporation
2-2-2 Hikaridai, Seika, Soraku, Kyoto 619-0288, Japan
Recent advances in machine learning and artificial neural networks have made it
possible to build robots and virtual agents that can learn a variety of behaviors.
However, their learning capabilities are strongly dependent on a number of
parameters, such as the learning rate, the degree of exploration, and the time scale of
evaluation. These parameters are often called metaparameters
because they regulate
the way detailed parameters of an adaptive system change with learning. The
permissible ranges of such metaparameters are dependent on particular tasks and
environments, making it necessary for a human expert to tune them usually by trial
and error. This is why most learning robots and agents to date can only work in the
This is in a marked contrast with learning in even the most primitive animals,
which can readily adjust themselves to unpredicted environments without any help
from a supervisor. This commonsense observation suggests that the brain has a
certain mechanism for metalearning
, a capability of dynamically adjusting its own
metaparameters. A candidate of such a regulatory mechanism in the brain is the
neuromodulator systems that project diffusely from the brainstem to the cerebral
cortex, the basal ganglia, and the cerebellum (see, e.g., Role and Kelly, 1991; Robbins,
1997; Katz, 1999). Most notable of such neuromodulators are dopamine (DA),
serotonin (5-HT), noradrenaline (NA), and acetylcholine (ACh) (see Figure 1).
In order to understand the brain mechanism of behavioral learning, the theory of
reinforcement learning (see, e.g., Sutton and Barto, 1998), which has been developed
for artificial agents that learn to optimize their behaviors through interaction with the
environment, can provide a comprehensive computational framework (Doya, 1999).
This paper first reviews basic algorithms of reinforcement learning and
introduces a few metaparameters essential for behavioral learning. Then, based on an
extensive body of neurobiological data, the paper proposes hypotheses on how these
metaparameters are regulated by neuromodulators. The hypotheses allow us to
predict and interpret the interactions between neuromodulators, behaviors, and
environments. They may also allow us to develop a computational model of
Metaparameters of Reinforcement Learning Agents
Central to the theory of reinforcement learning is the value function of a state:
)) = E[ r
) + r
+1) + 2 r
+2) + …]
+2),… denote the reward acquired by following a certain action
starting from the initial state s
). The discount factor 0 ≤ ≤ 1 specifies
how far into the future rewards are taken into account. The optimal policy that
maximizes the above expectation of a cumulative reward is obtained by solving the
) = max [ r
) + V
) is the state reached by taking an action a
at state s
(Sutton and Barto,
1998). What this equation says is that when taking an action a
, both the immediate
) and the future cumulative reward V
)) should be taken into
The relative merit of taking an action a
at state s
) = r
) + V
)) – V
which is called the temporal difference
(TD) signal, can be used both for action selection
and value function learning. A common way of stochastic action selection to facilitate
exploration is the Gibbs sampling method:
) = exp[ (s
)]/Σ exp[ (s
where is a parameter that controls the randomness of the action choice, called the
The estimate of the value function is updated by
)) := V
)) + (s
Control of Metaparameters by Neuromodulators
Based on a large body of neurobiological data and computational modeling
studies, the following hypotheses are proposed:
1) The dopaminergic system encodes the relative merit .
2) The serotonergic system controls the time scale of evaluation .
3) The noradrenergic system controls the inverse temperature .
4) The acetylcholinergic system controls the learning rate .
Given below is the experimental evidence supporting these hypotheses.
The dopaminergic neurons in the substantia nigra pars compacta (SNc) and the
ventral tegmental area (VTA) project extensively to the basal ganglia and the frontal
cortex. It has been shown in monkey conditioning experiments that dopaminergic
neurons respond initially to rewards but later to reward predicting stimuli (Schultz,
1998). Their activity is well characterized by the TD signal in the above formulation
(Houk et al, 1995; Schultz et al., 1997). The TD signal can be used both for action
selection and action reinforcement. This is consistent with the fact that dopamine is
involved in action selection (e.g., in Parkinson’s disease) and action reinforcement
The amplitude of the TD signal also indicates the relevance of the current
action and the state change in terms of the future reward. This is in accordance with
the fact that dopamine facilitates working memory in the frontal cortex (Sawaguchi
In many control tasks, the time scale of evaluation determines the weighting for
future rewards compared to the immediate action cost. Accordingly, setting too
small can make “doing nothing” the best solution.
The level of serotonin is higher in awake states and lower during sleep. A high
level of serotonin generally stabilize behaviors, while a low level of serotonin can
lead to impulsive behaviors (Buhot, 1997). These facts are consistent with the
hypothesis that serotonin controls the time scale of evaluation. For example,
serotonin-enhancing drugs (e.g., Prozac) can reduce depression and anxiety by
making immediate negative rewards less significant with a longer evaluation time
Noradrenaline for exploration and optimization
Stochastic action selection is helpful for long-term learning by facilitating wide
exploration, while deterministic, greedy action selection is favored in making best
use of what has already been learned. Thus the randomness in action selection
should be actively tuned in reference to the progress of learning and the urgency of
the situation, known as the exploration-exploitation problem.
Noradrenaline has been known to be involved in the control of arousal and
relaxation. The noradrenergic neurons in the locus coeruleus (LC) are activated in
urgent situations (e.g., with aversive stimuli). It was recently shown in monkeys that
the LC neuron activity is correlated closely with the accuracy of action selection
(Usher et al., 1999). Furthermore, noradrenaline sharpens the response tuning of
neurons by increasing the threshold and the gain (Servan-Schreiber et al., 1990).
These facts suggest that noradrenaline regulates the randomness in action
selection, similar to the inverse temperature in the above formulation.
Acetylcholine is known to modulate the synaptic plasticity in the hippocampus
and the cerebral cortex (Hasselmo, 1999). Depletion of acetylcholine leads to memory
disorders like Alzheimer’s disease. These facts point to the possibility that
acetylcholine controls the learning rate , which determines when to learn something
new and when to retain what has been memorized.
The appropriate setting of one of these metaparameters depends on the settings
of the other metaparameters as well as the environmental setting and the progress of
learning. The above hypotheses on the roles of neuromodulators as metaparameters
of learning enable us to predict how the levels of neuromodulators should affect each
other and change with the environment and the behavior.
For example, the equation for the TD signal
) = r
) + V
)) – V
specifies how the activity of dopamine, , should depend on the activity of serotonin,
. It suggests that serotonin can have both facilitatory and inhibitory effects on
dopamine, depending on whether the expected future reward is positive or negative.
A higher level of serotonin favors long-term prediction over short-term outcome
when they are in conflict. This can explain serotonin’s differential effects on dorsal
and ventral striatal dopaminergic pathways (De Deurwaerdére et al., 1998) by
assuming that the two are involved in long- and short-term reward prediction,
Neurobiological studies on emotion have so far focused on the role of emotion as
the “emergency programs” of behaviors, such as escaping and freezing. However,
the role of emotion in modulating cognitive and behavioral learning systems is
highly important; many affective or mental disorders occur as a result of the
“runaway” of learning systems. Consideration of emotion as a metalearning system
enables a novel computational approach in which studies on learning theory,
autonomous agents, and neuromodulatory systems can be bound together.
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vivo dopamine release in the rat nucleus accumbens and striatum that follows
electrical stimulation of dorsal raphe nucleus: role of 5-HT3 receptors. Journal
Doya, K. (1999) What are the computations of the cerebellum, the basal ganglia, and
the cerebral cortex. Neural Networks, 12:961-974.
Hasselmo, M.E. (1999) Neuromodulation: acetylcholine and memory consolidation.
Trends in Cognitive Sciences, 3:351-359.
Houk, J.C., Adams, J.L., and Barto, A.G. (1995) A model of how the basal ganglia
generate and use neural signals that predict reinforcement. In J.C. Houk, J.L.
Davis, and D.G. Beiser (Eds.), Models of Information Processing in the Basal
Ganglia, MIT Press, Cambridge, MA, USA, pp. 249-270.
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for information processing. Oxford University Press, Oxford, UK.
Robbins, T.W. (1997) Arousal systems and attentional processes. Biological
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monoaminergic systems. In E.R. Kandel, J.H. Schwartz, and T.M. Jessel, (Eds.),
Principles of Neural Science, third edition, Appleton & Lange, Norwalk, CT,
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working memory: Local injections of dopamine antagonists into the prefrontal
cortex of rhesus monkeys performing an oculomotor delayed-response task.
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Servan-Schreiber, D., Printz, H., and Cohen, J.D. (1990) A network model of
catecholamine effects: Gain, signal-to-noise ratio, and behavior. Science,
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Usher, M., Cohen, J.D., Servan-Schreiber, D., Rajkowski, J., and Aston-Jones, G. (1999)
The role of locus coeruleus in the regulation of cognitive performance. Science,
Figure 1: The projection of neuromodulators from the brain stem to the cerebral
cortex, the basal ganglia, and the cerebellum. Dopaminergic projection from
substantia nigra, pars compacta (SNc) and ventral tegmental area (VTA).
Serotonergic projection from dorsal raphe nucleus (DR). Noradrenergic projection
from locus coeruleus (LC). Acetylcholinergic projection from septum (S) and Meynert
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