Biological Psychology

Volume 186, February 2024, 108741

Review

Active inference as a theory of sentient

behavior
Giovanni Pezzulo a , Thomas Parr b, Karl Friston c d

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Highlights

• We review the history and future of active inference: a unifying

perspective on action and perception.

• We discuss the conceptual roots of active inference, its current

status and promising future directions.

• We highlight the importance of the brain’s generative models to

predict, infer, and direct action.

• We critically discuss how active inference can unify extant

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 psychological theories.

Abstract
This review paper offers an overview of the history and future of active inference—a
unifying perspective on action and perception. Active inference is based upon the
idea that sentient behavior depends upon our brains’ implicit use of internal models
to predict, infer, and direct action. Our focus is upon the conceptual roots and
development of this theory of (basic) sentience and does not follow a rigid
chronological narrative. We trace the evolution from Helmholtzian ideas on
unconscious inference, through to a contemporary understanding of action and
perception. In doing so, we touch upon related perspectives, the neural
underpinnings of active inference, and the opportunities for future development. Key
steps in this development include the formulation of predictive coding models and
related theories of neuronal message passing, the use of sequential models for
planning and policy optimization, and the importance of hierarchical (temporally)
deep internal (i.e., generative or world) models. Active inference has been used to
account for aspects of anatomy and neurophysiology, to offer theories of
psychopathology in terms of aberrant precision control, and to unify extant
psychological theories. We anticipate further development in all these areas and note
the exciting early work applying active inference beyond neuroscience. This suggests
a future not just in biology, but in robotics, machine learning, and artificial
intelligence.

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Keywords
Active inference; Predictive coding; Generative model
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 1. Introduction
Psychologists and neuroscientists are increasingly entertaining the idea of the brain
as a “prediction machine”, which learns an internal (i.e., generative) model of the
lived world – and of the consequences of its actions – to make sense of sensations,
predict how the current situation will unfold (i.e., learning and perception), and to act
in a purposeful manner (i.e., action selection, exploration-exploitation, planning, et
cetera). This idea appears in several guises, including the Bayesian brain, the predictive
brain, predictive processing, predictive coding, active inference and the free energy
principle, to name a few.

Here, we critically review the origins, scope and impact of this idea, in fields like
psychology and neuroscience. For conceptual clarity, we focus specifically on active
inference: a normative theory of sentient behavior that formalizes the “predictive
brain” idea and provides a first-principle account of its computational and neuronal
processes (Parr et al., 2022).

While active inference is still relatively young, it has a growing impact across various
disciplines. It is increasingly used by (for example) neuroscientists interested in the
neural circuits supporting predictions and prediction errors (Bastos et al., 2012, Parr
et al., 2021, Parr and Friston, 2018, Walsh et al., 2020); psychologists interested in
how we deal with uncertainty and cognitive effort during decision-making (Parr et
al., 2023, Rens et al., 2023), modelers interested in the mechanisms of action-
perception, exploration-exploitation and higher cognition (Friston et al., 2017, Friston
et al., 2017, Pezzulo et al., 2015, Pezzulo et al., 2018), clinicians interested in
understanding aberrant behavior in psychopathology (Maisto et al., 2021, Van den
Bergh et al., 2017), roboticists interested in self-supervised learning of world models
and goal-directed behavior (Ahmadi and Tani, 2019, Taniguchi et al., 2023), and
neurophilosophers (Clark, 2015, Hohwy, 2013).

This breadth of application is appealing, but risks creating a fragmented picture and
some uncertainty about its original commitments and conceptual implications. The
aim of this brief manuscript is to help researchers using (or interested in) predictive
coding and active inference to “connect the dots” and orient themselves within a
growing literature. Despite distinct lines of work — that emphasize different aspects
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 of active inference — these applications all rest on the same core principles. To
foreground these core principles, we will look at the historical and conceptual origins
of active inference—to illustrate how its core principles were introduced; then
consider briefly how the scope of active inference has expanded into several
disciplines—and finally look to future developments. Given the brevity of this
treatment, we cannot provide a full introduction to active inference. Rather, we
provide an overview of the narrative in (Parr et al., 2022), which interested readers
can consult.

In the next section, we briefly discuss the conceptual (and historical) roots of active
inference in early views of prediction and action-based cognition. We then review
some key developments of active inference, by focusing on landmark papers that
explain how it stems from a single principle (namely, free energy minimization). We
next consider its scope across perception, action, planning, etc. This brief review
helps us make the point that active inference provides a unifying perspective on
several cognitive topics and theories and across levels of understanding, from
conceptual to neural. Finally, we briefly highlight some promising research directions
that could expand the scope of active inference – and potentially its impact on
psychology and neuroscience.

2. The conceptual and historical roots of active inference

Active inference has roots in various early theories in cognitive science (and beyond,
in fields that would not necessarily use the label “cognitive”). One root is the idea that
the brain carries a small-scale model of the environment and uses it to mentally
simulate what-if actions, instead of (or before) acting on the environment (Craik,
1943). This idea is foundational in cognitive science. For example, (Tolman, 1948)
proposed that humans, rodents and other animals find their way in a maze by first
learning a mental model or “cognitive map”, rather than by considering which of
their navigation actions were previously rewarded the most, as assumed by
behaviorist formulations.

Another root is the idea of (Helmholtz, 1866) that perception is an (unconscious)

inference based on an internal generative model – that uses recurrent (top-down and
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 bottom-up) counter-streams of processing, rather than bottom-up transduction of
external sensations into internal representations (and later actions). This idea was
later developed in psychology (Gregory, 1968, Gregory, 1980) and computational
neuroscience; giving rise to the “Bayesian brain” hypothesis (Doya et al., 2007) and to
formulations of predictive coding as a possible neurobiological implementation of
perception-as-inference in the brain (Friston, 2005, Rao and Ballard, 1999). Beyond
perception, other cognitive functions were later described in terms of inference, i.e.,
planning-as-inference (Botvinick & Toussaint, 2012).

Yet another “root” is the idea of cyberneticists (Miller et al., 1960, Powers, 1973,
Wiener, 1948) that goal-directed action proceeds by firstly setting up a desired state
or observation (e.g., feeling warm), then monitoring the discrepancy – now referred
to as a “prediction error” – between the preferred and sensed state (e.g., feeling
excessively warm), and then selecting a course of action that reduces this discrepancy
– where “action” is a suitcase word and can include any means to exert control over
external stimuli; ranging from simple autonomic reflexes (e.g. thermoregulation) to
sophisticated plans (e.g., visiting one’s favorite ice cream shop). A key result in this
field – which coheres with the Helmholtzian perspective above – is the ‘Good
regulator theorem’ of (Conant & Ashby, 1970), which argues that effective regulatory
systems must [be a] model the environment they regulate. In a similar vein, in
psychology, ideomotor theory proposed that action control is essentially anticipatory
and that action are selected and controlled by their anticipated consequences or
outcomes, not through stimulus-response (Hoffmann, 2003, Hommel, 2003, James,
1890).

Besides cybernetics, there are other influential views that highlight the centrality of
adaptive regulation for behavior and life itself. One example is the idea that living
organisms are autopoietic systems, which create the conditions for their own
existence. More recently, this idea has been framed as ‘self-evidencing’ (Hohwy 2016)
– i.e., creatures seek out sensations that provide evidence for their continued
existence. Intuitively, sensing our body temperature to be around 37 °C offers more
evidence that we are still alive than body temperatures far from this value. The
concept of autopoiesis gave birth to enactive approaches in philosophy (Maturana &
Varela, 1980). From another angle, it has been postulated that a central imperative for
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 living organisms is maintenance of physiological homeostasis (i.e., correction of
deviations from preferred physiological states through reflexive actions) and the
regulation of basic imperatives (Cannon, 1929) – but more modern theories
emphasize that physiological regulation is fundamentally anticipatory (i.e., allostatic)
(Sterling, 2012). Various researchers have proposed that closed-loop adaptive
regulation (and not stimulus-response) is key to understanding not just physiology
but (potentially) all cognitive processing (Cisek, 1999, Pezzulo and Cisek, 2016).

Finally, another root is the idea that cognitive processes, such as learning, perception
and decision-making, require an active engagement of organisms with the
environment. One early example of this action-oriented perspective is the view of
Gibson that perceiving things consists in seeing what to do or not to do with them,
i.e., perceiving affordances (Gibson, 1979). More recently, various researchers
proposed the necessity of a “pragmatic turn” in cognitive science and neuroscience –
and the need to recognize the importance of action as part and parcel of our
cognition (Buzsaki, 2019, Cisek and Kalaska, 2010, Cisek and Pastor-Bernier, 2014,
Engel et al., 2016, Lepora and Pezzulo, 2015, O’Regan and Noe, 2001), rather than just
a way to report “central” decisions, as assumed in conventional (serial) theories.

Interestingly, each of these ideas implies a shift from reactive to predictive, enactive
views of the brain. While a reactive brain waits for incoming stimuli, a predictive and
active brain predicts external events (e.g., predictive coding) and actively gathers
evidence (i.e., active sensing and active learning) to make sense of the world. While a
reactive brain selects actions based on the past and present (e.g., the history of
reinforcement and the current cue), a predictive brain actively imagines its preferred
future and then makes this happen by acting (e.g., acts in a goal-directed manner).
While a reactive brain maintains homeostasis, a predictive brain acts to anticipate
needs and performs anticipatory regulatory (or allostatic) actions.

All these (and other) views contributed to raising the importance of predictive and
enactive views of the brain and of cognition. However, each of these perspectives
were somewhat disconnected from one another and linked to different research
traditions, which are sometimes seen as conflicting with one another (e.g., the
Helmholtzian and the Gibsonian traditions). One benefit of active inference is that it
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 helps unify and thereby advance these traditions, as we explain in the following
Sections.

3. The normative perspective of active inference – and how it has

developed
Active inference provides a normative perspective that unifies and advances the
predictive and enactive views of brain and behavior. It does so by highlighting that
several apparently disconnected accounts – identified by early theories – stem
parsimoniously from the assumption that living organisms obey a single imperative:
namely, they act to minimize their surprise,1 or more formally, their variational free
energy.

The mathematics of variational free energy minimization is beyond the scope of this
article; we suggest to the interested readers to consult (Parr et al., 2022). Here,
instead, we introduce the key concepts of the theory, by briefly reviewing (non-
chronologically) selected landmark papers and linking them to the early theories.

Active inference starts from a simple consideration: that to maintain their existence
and integrity, all living organisms need to remain in a bounded set of characteristic
states that basically define their place within an ecological niche; for example, a fish
cannot live out of water. Using the lexicon of Bayesian inference, being out of water
for a fish is a “surprising” state. Clearly a fish should avoid this surprise, and the idea
generalizes to suggest that living organisms must avoid surprising states (Friston et
al., 2010). If they did not, they would not be living organisms for long. Another way of
looking at this is that everything (including me) is defined by being in some
characteristic (attracting) set of states. Conversely, I am defined by the kinds of states
I cannot be in. These are surprising states.

A computationally tractable solution to surprise minimization is the minimization of

an information-theoretic quantity – variational free energy – which is a function of
two things: a generative model (i.e., a statistical model that describes how sensations
are generated) and observed sensory data. This implies that a living organism must be
equipped with a generative model – or in the lexicon of (Craik, 1943), a small-scale
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 model – to predict the sensations generated by the world (and by the organism’s
place in it). In Bayesian terms, a generative model comprises two things: a prior over
the hidden (i.e., unobserved) variables of interest and a likelihood function that maps
the hidden variables to observables (Bishop, 2006). See Fig. 1 for a schematic
illustration of the organism’s generative model of the world and its relation with the
generative process: the true environmental contingencies that generate its
observations, which is inaccessible to the organism.

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Fig. 1. Generative model and generative process in active inference. This Figure—
reproduced from (Parr et al., 2022)—illustrates the structure of active inferential
theories of brain function. Our worlds evolve according to some dynamical process
that generates observations (y) from hidden states (x * ). Our internal models account
for observations in terms of hypothetical hidden states (x). Our inferences about these
states based upon our observations then drive actions (u) that intervene on the
processes generating our sensations.

Put simply, an organism can minimize variational free energy by aligning the
predictions of its generative model and the data it observes. In different settings, this
minimization has been described in various ways, such as the minimization of
surprise, of prediction errors, or of the discrepancy between the model and the world.
All of these are equivalent to the minimization of variational free energy under
specific sets of assumptions.

Interestingly, aligning the predictions derived from a generative model and data can
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 be achieved in two ways: by changing the model predictions and by changing the
observed data. The former corresponds to revising the agent’s beliefs (used in the
technical sense of probability distributions over hidden variables) if they do not
explain the data well. This is exactly the inferential view of perception of (Helmholtz,
1866). The latter corresponds to acting in the world to change the data that will be
sampled next – to render them more like the organism’s prior predictions. This latter
perspective on action – and on its dependence on expected outcomes – is highly
congruent with cybernetics (Miller et al., 1960, Powers, 1973, Wiener, 1948) and
ideomotor theory (Hoffmann, 2003, Hommel, 2003, James, 1890).

In sum, changing beliefs about the causes of data (i.e., perception) and changing the
data (i.e., action) are two aspects of free energy minimization. In formal terms, they
map to its two components: the minimization of divergence and the maximization of
evidence, see Fig. 2. Recognizing that action and perception can be unified within a
single formal imperative – the minimization of free energy – is one of the key
innovations of active inference, which helps integrate and extend the early theories
reviewed above.

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Fig. 2. Perception and action play complementary roles in the minimization of

variational free energy. This Figure— reproduced from (Parr et al., 2022)—highlights
the relationship between action and perception via free energy (F). Perception
involves minimizing free energy by changing our beliefs (Q) about states (x). This
effectively minimizes the divergence (DKL) between our beliefs and the probability of
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 these states given sensory data (y). Action minimizes free energy through changing
those parts of the free energy that depend upon sensory data—notably, the evidence
or probability of data under our internal model.

Regarding neural implementation, one of the most widely entertained hypotheses –

about how the brain might implement perceptual inference – is predictive coding
(Rao & Ballard, 1999). Fig. 3 shows the architecture of a predictive coding scheme as it
might manifest in the cerebral cortex. In this predictive coding network, inference is
realized by propagating predictions and prediction errors through top-down and
bottom-up pathways, respectively, and by minimizing prediction errors across all
levels. Interesting, predictive coding can be derived as a special case of variational
free energy minimization (Friston, 2005).

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Fig. 3. The architecture of predictive coding. This Figure—reproduced from (Parr et

al., 2022)—shows the message passing between populations of neurons under a
predictive coding scheme as it might manifest in the layers of the cerebral cortex
(separated into superficial layers I-III, layer IV, and deep layers V-VI). This shows
predictions based upon expectations (µ) being subtracted from ascending signals to
compute errors (ε), which are used to update expectations. The subscripts indicate
whether we are dealing with fast changing dynamical variables (x) or more slowly
changing contextual variables (v) which act to link together different hierarchical
levels, with hierarchy indicated by the bracketed superscripts. As we ascend the
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 hierarchy, the variables we deal with become slower, such that the contextual
variables at one hierarchical level evolve over the same timescale as the dynamical
variables at the level above.

While predictive coding is a model of perception, it can be readily extended to

encompass the role of action in the minimization of free energy (described above).
The move from predictive coding to active inference can be realized by equipping
predictive coding networks with simple motor reflexes. In this perspective, the motor
system works by generating proprioceptive predictions (in the same way standard
predictive coding generates exteroceptive predictions) — and not motor commands,
as conventionally proposed – and these proprioceptive predictions are realized
through the motor reflexes (Adams et al., 2013).

Subsequently, this theory was extended to also model autonomic control (Barrett and
Simmons, 2015, Pezzulo, 2014, Seth et al., 2012). The general idea is that autonomic
control might work by generating interoceptive predictions (i.e., homeostatic
setpoints) and then fulfilling them through autonomic reflexes, in much the same
way motor control might work by generating proprioceptive predictions and then
fulfilling them through motor reflexes. This development of active inference helps
connect it with theories of allostatic control (Sterling, 2012) and paves the way to a
better understanding of our ability to model and control the internal milieu, not just
the external environment. This stream of research underwrote novel approaches to
psychopathology – as deficits of interoceptive processing (Paulus et al., 2019).

So far, we have discussed active inference using generative models that characterize
processes that unfold in continuous time (e.g., predictive coding networks) and use
continuous variables (i.e., the formal framework of dynamical systems and state-
space models). However, many cognitive problems can be characterized at a distinct
level: as (sequences of) discrete decisions. These include problems that require the
selection of discrete responses during psychology experiments, the targets for
saccades, or navigational trajectories in discretized environments (Friston et al., 2017,
Friston et al., 2017). These problems can be modeled in active inference, using
generative models that use discrete variables (and the formal framework of Partially
Observable Markov Decision Processes).
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 In addition to the two aforementioned components (priors and likelihood function),
the generative models for active inference in discrete time often include a third
component: the transition function, which describes the way in which hidden states
change depending upon the agent’s actions (or sequences of actions, called policies).
Crucially, these generative models have temporal depth and afford a novel capability
that was not available in simpler models: namely, planning. In simple terms, planning
involves using the generative model to predict the consequences of different policies,
scoring the policies according to how much they are expected to minimize free energy
in the future and then (with some simplifications) select the best policy.

This planning process induces a novel quantity – expected free energy – that is the
functional that active inference uses to evaluate (and assign a prior to) policies and it
is distinct from the notion of variational free energy discussed so far (Friston et al.,
2017). The notion of expected free energy has been very useful in the development of
active inference models of things like (bounded) decision-making, planning,
exploration-exploitation and curiosity (Friston et al., 2017, Parr and Pezzulo, 2021,
Schwartenbeck et al., 2019). This is because this notion is richer than the common
optimization objectives used in other formal frameworks (e.g., economic theory and
reinforcement learning). This is because expected free energy considers jointly a
pragmatic imperative (utility maximization) and an epistemic imperative (information
gain, or the resolution of the uncertainty). Indeed, as Fig. 4 illustrates, it is possible to
map expected free energy to various other formal notions (e.g., Bayesian surprise,
Risk-sensitive control, Expected utility theory), by removing one or more of its terms.
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Fig. 4. Expected free energy and the way it can be mapped to different formal
notions (e.g., Bayesian surprise, Risk-sensitive control, Expected utility theory) by
removing one or more terms, denoted with numbers. This Figure—reproduced
from (Parr et al., 2022)—expresses expected free energy in terms of beliefs about
trajectories (indicated by the tilde ∼). The additional symbols here, not in previous
figures, are the π for policies and the C for preferences. Note that some terms
(including term 1) are expressed in terms of expectations—i.e., averages under the
subscripted distribution.

Active inference is a general scheme that can be applied to address various cognitive
processes. Crucially, the functioning of active inference is the same across all
problems: what differs is the generative model, which is task specific. This implies
that by designing the appropriate generative models, it is possible to address a variety
of cognitive tasks with the same approach – and to pass from the normative
perspective of active inference to specific implementations that have biological
plausibility (Friston et al., 2017, Parr and Friston, 2018).

Here, a worked example may be helpful. To illustrate some of the principles we have
outlined so far, we will consider how we might go about developing a model for a
ubiquitous task in cognitive neuroscience—a delay period oculomotor task. This is a
relatively simple task that can be performed by humans—and some animals—and that
is designed to probe working memory function (Funahashi et al., 1989). The task
sequence is as follows. First, a cross is presented on screen and our subject maintains
fixation on this cross. A target then appears at one of several possible locations
towards the periphery of the screen, but our subject still maintains fixation. The
target then disappears and, after a ‘delay period’, a stimulus appears to signify that
the subject should make a saccadic eye movement to the location of the target.
Successful performance of this task relies upon retaining a memory of the target
location during the delay and response phases.

To model this task, we must consider the data available to the subject. In this case,
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 these are the visual stimuli and proprioceptive inputs, and whether the correct action
was chosen. To do so, we need to take account of the causes of these data. The causes
of proprioceptive data are simply the direction in which our subject’s eyes are
pointing. Visual outcomes, depend upon a combination of (1) gaze direction, (2) the
intended target location, and (3) the current stage of the task (i.e., the fixation, target
presentation, delay, or response stage). For each of these three variables, we must
then specify how we expect them to evolve throughout the task. The gaze direction
will transition from one step to the next based upon the decisions our subject makes.
The intended target location will be fixed (although initially unknown) throughout
the task. The task stage evolves predictably through a sequence of steps. Together,
these beliefs about the way in which data are generated and the dynamics of the
causes allow our subject to predict what will be observed next, and to update these
beliefs when these predictions are violated.

As outlined above, active inference equips models with prior beliefs about the relative
plausibility of different choices based upon their relative expected free energies. In
this model, the key part of the expected free energy is a preference for receiving the
‘correct’ feedback outcome which is only available during the response phase of the
task (see (Mirza et al., 2016) for a similar setup in the context of scene categorisation,
in which the main role of the expected free energy is to promote information
seeking). It is this that prompts a saccade to the remembered target location. Finally,
the predicted action must be executed. This depends upon resolving the error
between the anticipated proprioceptive information given the inferred saccade and
current proprioceptive input. The result is the sequence of steps shown in Fig. 5.
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Fig. 5. A simulated oculomotor delay period task. This figure, taken from (Parr &
Friston, 2019b) (published under a CC BY 4.0 license), shows simulated performance
of a simple working memory task under active inference. Although simple, this task
calls for planning (of our next saccade), recall (of the target location), and movement
execution. The upper left images show a series of frames taken from the simulation,
as if we were observing our participant’s eyes. The black arrows link these
behavioural responses to the view of the stimulus screen from the time of the target
(red) presentation to the response phase. A series of black dots show the (cumulative)
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 trajectory of gaze direction. Because this model is formulated to have both
continuous (prediction-error minimising) and discrete (sequential planning) parts,
we can plot the trajectory both in terms of position and velocity (lower left) and in
terms of the sequence of actions taken.

The oculomotor control example illustrates how active inference can be concretely
applied to study cognitive tasks, by designing (or learning) the appropriate generative
models. Generative models represent formal hypotheses about how cognitive tasks
are accomplished – hypotheses that can be validated with empirical data. A useful
illustration of the design principles to realize (or train) generative models for
different cognitive problems is provided in (Parr et al., 2022). This treatment makes a
distinction between generative models in continuous time (that are useful to address
motor control tasks) and discrete time (that are useful to address decision and
planning tasks) and explains how these two types can be combined to form so-called
hybrid or mixed generative models, in which discrete-time models are placed on top
of continuous-time models. Furthermore, the generative models of active inference
can be extended hierarchically, to model processes that unfold at different timescales.
One example is the model of active listening processes, in which (for example) lower
hierarchical levels deal with words and higher levels deal with sentences (Friston et
al., 2021). Another example is a model of hierarchical action recognition that
recognizes actions at different levels, from low level kinematics to higher level goals
and intentions (Proietti et al., 2023). It is also possible to use hierarchical models to
model hierarchies of control, in which lower-to-higher levels deal with autonomic
imperatives (e.g., ensure a correct basic temperature) in increasingly complex ways
(e.g., from thermoregulation to the goal-directed plan to buy water before a long run)
(Pezzulo et al., 2015, Tschantz et al., 2021). These developments – from simple to
more sophisticated (e.g., hierarchically and temporally deep) generative models has
extended the range of cognitive models that have been addressed using active
inference over the years.

Another interesting realization is the fact that it is possible to derive a biologically

motivated “process theory” for active inference in discrete time, by interpreting the
specific operations (variational updates) required to minimize free energy as signals
that are computed or exchanged across neurons (Friston et al., 2017). This is
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 important because it permits crossing levels of explanation – from normative to
mechanistic and neuronal – and to use active inference to simulate neuronal activity
that would ensue from the performance of cognitive tasks (Friston et al., 2017, Parr
and Friston, 2018).

Another important development of active inference regards precision control and its
role in psychopathologies. In predictive coding, variables are encoded as Gaussian
distributions and precision simply refers to the inverse of the variance of a
distribution (Friston, 2005). Precision control refers to a mechanism that optimizes
the precision of (the distribution of) each variable of an organism’s generative model.
It is important since it regulates the relative importance of top-down predictions and
bottom-up prediction errors across the hierarchy. This is because prediction errors
that are assigned greater (lower) precision have greater (lower) impact on the belief
updating and the ensuing inference. Veridical inference requires the precision of (the
distribution of) each variable to be optimized, to reflect the signal-to-noise ratio of
sensory signals – therefore highlighting a link between precision control and
attention as gain control (Parr & Friston, 2019a) – or the importance of an organism’s
prior preferences – reflecting the fact that an organism’s innate drives or goal states
can be encoded as highly precise priors (Pezzulo et al., 2015). Interestingly, when
precision control fails, it can produce excessively rigid forms of inference (when
priors fail to be updated in the light of novel evidence) or excessive sensitivity to
stimuli (when belief revision follows the sensory input and its random fluctuations
too closely—i.e., it overfits). These forms of aberrant inference, which depend
sensitively on predicted precision, have been adopted to explain several
psychopathological conditions, such as delusions, depression, psychosis, and many
others (Barrett et al., 2016, Corlett and Fletcher, 2015, Edwards et al., 2012). In turn,
these theories also speak to aberrant neuromodulation, since the precision of (the
distribution of) different variables might be encoded by different neuromodulators,
e.g., acetylcholine for the precision of the likelihood, noradrenaline for the precision
of transitions, dopamine for the precision of policies, etc. (Parr & Friston, 2018).

Yet another development regards the analysis of generative models during sleep or
other ‘offline’ periods. It has long been hypothesized that learning generative models
benefits from alternating on-line and off-line periods (Hinton et al., 1995). While on-
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 line generative modelling maximises accuracy (under complexity constraints), during
off-line activity – in the absence of sensory data to “explain away” – model
optimisation can focus on minimising complexity; for example, by removing
redundant parameters (Friston et al., 2017, Pezzulo et al., 2021). From a neuronal
perspective, generative modelling during offline periods could be associated with
(generative) replay activity in the hippocampus, the prefrontal cortex and other brain
areas; but these links remain to be fully established (Foster, 2017, Schwartenbeck et
al., 2023, Stoianov et al., 2022).

Finally, an interesting development regards the realization of active inference in

which the free energy minimization extends “beyond the skull”, to model the ways
multiple active inference agents engage in cooperative or competitive tasks (Friston
and Frith, 2015, Maisto et al., 2023) or construct their own niches (Constant et al.,
2022). These and other works illustrate that the concept of free energy minimization
can readily extend to multi-agent settings – including settings that go beyond the
standard scope of cognitive science, such as morphogenesis (Friston & Levin,
Sengupta, et al., 2015) and autopoiesis (Friston, 2013) – and hence potentially shed
light on the relations between multiple nested levels of (self-)organization, from
individual to social and cultural levels.

In sum, we have highlighted various developments of active inference, which

encompass the complementary roles of perception and action in minimizing an
organism’s variational free energy (and ensuring that it successfully avoids
“surprising” and characteristic states), the proposal of biologically plausible
architectures for continuous time predictive coding and action control, the realization
of generative models for discrete decisions that afford planning and the minimization
of expected free energy, the hierarchical extension of these models, the importance of
precision control, and beyond. For each of these topics, we have cited some selected
papers that the interested readers might want to consult for more detailed
information. Clearly, this is not an exhaustive list, but each of these developments has
been useful to develop models of increasingly complex cognitive and social functions;
see (Parr et al., 2022) for a more exhaustive treatment of active inference.
:
 4. The benefits of unification
In the previous Section, we saw that the scope of active inference touches several
domains of psychology and neuroscience. Here, we foreground a benefit of this rapid
expansion: namely, unification.

Arguably, a main goal of cognitive psychology and neuroscience is explaining

behavior and its neural foundations, in a comprehensive (if not a unified) way. Yet, to
ensure methodological rigor, these disciplines usually adopt restricted laboratory
settings that tend to isolate cognitive functions and obfuscate their relations (Maselli
et al., 2023). Consider for example a mundane task that we solve almost every day:
crossing a busy road. Even this relatively simple task engages several cognitive
processes in a coordinated manner, such as perception (of the situation), memory (of
past street crossing episodes), planning and action selection (of the best route),
motivation (and the “why” of crossing), attention (to select the most relevant stimuli),
etc. These processes are often studied in isolation using different paradigms leading
to a proliferation of hypothesis and theories that assign each of them a distinct
computational objective (and perhaps brain area) – therefore determining a very
fragmented theoretical landscape.

Active inference proceeds the other way around: it starts from a single principle and
asks how far one can go with it. And to what extent it is possible to derive from that
principle empirically testable hypotheses about behavior and its cognitive and neural
mechanisms? This approach brings the benefits of unification, in at least six ways.

First, active inference assumes that everything, from perception to action selection
and learning ultimately serves to minimize variational free energy. A consequence of
this is that one can align the (sometimes vague) conceptual terms used in psychology
with crisp formal terms of free energy minimization. For example, one can assign
things like attention to precision control. At the neuronal level, the fast updates –
mediated by synaptic activity – might correspond to inferential processes that
minimize free energy at a fast time scale, whereas the slower updates – at the level of
synaptic efficacy – might correspond to learning processes that minimize free energy
at a slower timescale. Precision dynamics might correspond to the activity of
neuromodulators, which finesse the inference at multiple levels, for example, by
:
 increasing the post-synaptic gain of sensory or prediction error-units (Feldman &
Friston, 2010). Oscillatory dynamics that are ubiquitous (and that often occur in
synchrony) both within and across brain area might be signatures of temporal
prediction and of the exchange of top-down and bottom-up information across
hierarchical levels of the brain’s generative model (Arnal & Giraud, 2012).

Second, active inference suggests that cognitive functions – usually addressed in

isolation – might be instead better understood by appealing to a unique process
theory. For example, in prominent computational neuroscience theories, perception
and action are two separate functions with different objectives and neural substrates.
According to Bayesian decision theory (Robert, 2007), the goal of perception is to
provide an accurate estimate of the agent’s state, whereas the goal of action selection
is to maximize its expected utility. The former process is a precondition for the latter,
implying an outdated, serial view of cognitive processing. Active inference holds that
perception and action cooperate to minimize free energy, by minimizing divergence
and maximizing evidence, respectively (Parr et al., 2022). Another example is the fact
that in 20th-century cognitive science, working memory was considered as a separate
storage that can be assessed by other components when needed; therefore, imposing
a separation between information storage and information processing. In contrast,
active inference models of hierarchical perception and action (Friston et al., 2021,
Pezzulo et al., 2018) treat memory of the previous state as intrinsic to the belief
updating under generative or world models, across multiple timescales, which is in
keeping with 21st-century accounts of working memory (Hasson et al., 2015).

Third, active inference has the potential to unify different “levels of understanding” of
cognitive processes. Marr famously introduced a distinction between computational,
algorithmic and neural implementation levels and argued that progress can be made
within each level and by connecting different levels (Marr, 1982). Establishing links
between theories that operate at different levels is often challenging. Active inference
helps establish firm relations across levels of description. Rather than Marr’s
tripartite distinction, in active inference it is more common to appeal to a distinction
between normative theory and process theory (Friston et al., 2017). Free energy
minimization is the normative objective of living organisms, whereas predictive
coding and variational message passing are process-level theories that describe how
:
 the brain might support free energy minimization. Importantly, as shown by (Friston,
2005), under certain assumptions predictive coding can be directly derived by the
minimization of variational free energy, connecting the two levels of explanation. A
similar case can be made for the variational message passing schemes proposed to
support discrete active inference in neural circuits (Friston et al., 2017).

Fourth, unification endows existing constructs with validity, via the application of
active inference across domains. One example is the development of theories of
interoceptive inference and autonomic control (Barrett and Simmons, 2015, Pezzulo,
2014, Seth et al., 2012) by analogy with the functioning of action control (Adams et
al., 2013). In this perspective, autonomic control works exactly like action control –
namely, it aims to minimize a discrepancy between a predicted and a sensed signal –
except that the “signal” refers to interoceptive streams rather than proprioceptive
streams. Another example can be found in computational psychiatry, where
numerous accounts of psychopathology appeal to a single mechanism: namely,
aberrant precision control.

Fifth, active inference has the potential to reconcile (or at least to contextualize)
theoretical perspectives that have long been considered at odds in psychology,
neuroscience and philosophy. One example is the Helmholtzian view that perception
constitutes an inference about the entities of the external world that cause our
sensations (Helmholtz, 1866) and the Gibsonian view that perceiving consists in
seeing action opportunities and affordances, not reconstructing a model of the
external reality within the brain (Gibson, 1979). This apparent dialectic could be
dissolved by considering that there are multiple ways to design generative models;
specifically, a relevant distinction is between generative models that explicitly model
the ways external states produce sensations (a.k.a., environmental models) or the
ways actions produce sensations (a.k.a., sensorimotor models) (Sims and Pezzulo,
2021, Pezzulo et al., 2023). Some active inference studies use generative models that
include explicit beliefs about entities in the external world that cause sensations,
such as one’s location in space (Friston et al., 2017). Other active inference studies use
generative models that only consider the sensory consequences of one’s action, such
as touch sensations that follow whisking at a given amplitude, but not explicit beliefs
about objects ‘out there’ (Mannella et al., 2021). The latter generative models adhere
:
 more closely to the notions of affordance (Gibson, 1979) and of sensorimotor
contingency (O’Regan & Noe, 2001), despite the fact they still entail inferential
dynamics. Besides this specific topic, there is a vivid debate in philosophy that
concerns the most appropriate way to consider active inference, in relation to
internalist (Hohwy, 2013), externalist (Clark, 2013) or enactivist theories (Bruineberg
et al., 2018).

Finally, and importantly, the integrative perspective of active inference could be

valuable in characterising of sentient behaviour – considered here to be the capacity
to infer states of the world and to act upon it with a sense of purpose (Friston, Da
Costa, et al., 2023). This operational definition is satisfied by active inference when,
and only when the generative model includes the consequences of action
(mathematically, when the generative model includes priors over policies based upon
expected free energy). This notion of sentience is does not have any
phenomenological commitments and is probably best read as ‘basic sentience’ in the
sense of (Clark, 2023).

Recently, there has been a proliferation of advanced Generative AI systems that

process language, images and videos with very high accuracy. However, in most cases,
these systems learn passively from large predefined datasets and disregard agency –
and the possibility to act upon the world with a purpose – to develop genuine
understanding (Pezzulo et al., 2023). Active inference suggests a different path to
understand and simulate sentient behaviour, which focuses on the development of
grounded world (i.e., generative) models, by actively engaging with the environment
and by predicting the consequences of the requisite interactions. An open question
for future research is whether the enactive and embodied approach of active
inference has the potential to complement and advance the development and
deployment of Generative AI.

5. Opportunities for the future

It's Difficult To Make Predictions, Especially About the Future. Niels Bohr.

The compass of active inference is expanding rapidly, but the landscape of future
:
 opportunities may be even ampler. Here, we focus on some of the developments that
we consider most promising and most likely in the near future.

The first and perhaps most obvious direction for the future regards a deeper empirical
scrutiny of active inference. A question that is sometimes asked of active inference is
whether any empirical findings could offer evidence for or against the framework.
This can be a vexed question to answer as it constitutes a category error. A framework
is not in itself a hypothesis. It is a way of formulating hypotheses. The relationship
between active inference and empirical psychology is that we can formalize
psychological theories in terms of the generative models that underwrite
neurophysiological and behavioural responses. Equipped with a proposed model, the
framework can be used to express a hypothesis, to predict the behaviour expected
under that hypothesis, and to fit to measured data to formally compare alternative
hypotheses. In other words, while active inference is an application of the free energy
principle – which is a principle (i.e., method) rather than a theory (Friston, 2010) –
theories tested under the active inference framework (e.g., those considered in this
article) make specific empirical predictions that can (and need to) be empirically
validated. One example of this is the oculomotor delay period model shown in Fig. 5,
which generate empirically testable predictions about oculomotor performance as a
function of varying delay periods (Parr & Friston, 2019b). Various empirical studies
are already addressing the empirical predictions of predictive coding, such as how
top-down and bottom-up dynamics support predictions and prediction errors,
respectively (Walsh et al., 2020). However, active inference makes a number of
specific predictions about (for example) the way the motor system works (Shipp et
al., 2013) and the way higher cognitive functions are implemented (Pezzulo et al.,
2018) that differ from mainstream theories and could be increasingly scrutinized by
future studies.

A second interesting direction for the future is assessing to what extent active
inference – and more broadly, the free energy principle – can help us understand the
evolution of complex neural circuits and life forms from simpler ones. Active
inference suggests a possible path from the simple mechanisms that supported
prediction and control in our earlier evolutionary ancestors to the more sophisticated
abilities of our species (Pezzulo et al., 2022), but a comprehensive account of the
:
 evolution and “phylogenetic refinement” (Cisek, 2019) of living organisms remains to
be fully developed (Friston et al., 2023, Friston et al., 2023).

A third interesting direction for the future regards the realization of advanced
artefacts, such as AIs and robots, based on active inference. There have already been
several successful robotic implementations of active inference, but the full potential
of the theory has not yet been reached (Ahmadi and Tani, 2019, Lanillos et al., 2021,
Priorelli, Pezzulo, & Stoianov, 2023, Taniguchi et al., 2023). Interestingly, some of the
central concepts of active inference, such as the importance of generative models and
self-supervised, predictive learning, are becoming central in mainstream research in
AI, as testified by the recent successes in generative AIs such as large language
models. This creates an important opportunity, since (apart for their obvious
technological impact), state-of-the-art AI systems can be precious in advancing our
understanding of living organisms, providing that they incorporate appropriate
(design) principles (Pezzulo et al., 2023).

Box 1
Glossary of technical terms.

Active Inference: A normative framework that elucidates the neural and

cognitive processes underlying sentient behavior, beginning with first
principles. This framework posits that perception and action work in concert
to minimize a shared functional known as variational free energy.

Expected Free Energy: This is the quantity that is used in active inference to
score action sequences or policies (and then to select between them). It takes
into consideration both the pragmatic value of policies – or how close a
policy’s expected outcomes are to the preferred outcomes – and their
epistemic value (or information gain) – or how much the policy is expected
to reduce uncertainty.

Generative Model: A statistical model designed to explain the generation of

observable content from unobservable, hidden (latent) causes. For instance,
it clarifies the process by which a visual object gives rise to an image on the
:
 retina. Generative models serve a dual purpose: they allow the generation of
novel, synthetic content and support the inference of hidden causes from
observable data. From a technical standpoint, generative models encode the
joint probability distribution governing both observables and hidden causes.

Latent (or Hidden) Variable: An internal variable within a generative model,

referred to as "latent" or "hidden" due to the fact that it cannot be directly
observed, but must be inferred.

Precision and precision-weighting: Precision denotes the inverse of

variance or standard error, serving as a measure of the reliability or certainty
associated with sensory information. Precision-weighting refers to the fact
that in predictive coding and active inference, prediction errors are weighted
by their respective precisions, therefore determining the extent to which
sensory observations influence the process of updating beliefs.

Predictive Coding: A computational framework in neuroscience that

provides a possible neural implementation for the idea that perception
consists in a process of inference. In hierarchical predictive coding networks,
inference is realized by minimizing (precision-weighted) prediction errors
across all hierarchical levels. In turn, this requires bidirectional loops
between top-down processes (conveying predictions) and bottom-up
processes (conveying prediction errors).

Variational Free Energy: This is the functional (function of a function) that

is minimized within the framework of active inference. It is also widely
utilized in utilized in probabilistic modeling, statistical inference and
machine learning. In its simplest instantiation, it corresponds to a
summation of prediction errors, which quantifies the deviation of observed
data from the predictions of the generative model. More formally, variational
free energy serves as an upper bound on the negative logarithm of the
evidence, which is the probability of observed data given a model.
:
 Funding and acknowledgements
This research received funding from the European Union's Horizon 2020 Framework
Programme for Research and Innovation under the Specific Grant Agreements No.
945539 (Human Brain Project SGA3) to GP and KF and No. 952215 (TAILOR) to GP; the
European Research Council under the Grant Agreement No. 820213 (ThinkAhead) to
GP; the PNRR MUR projects PE0000013-FAIR and IR0000011–EBRAINS-Italy to GP; for
the Wellcome Centre for Human Neuroimaging (Ref: 205103/Z/16/Z) to KF, a Canada-
UK Artificial Intelligence Initiative (Ref: ES/T01279X/1) to KF. The funders had no role
in study design, data collection and analysis, decision to publish, or preparation of the
manuscript.

The authors did not use generative AI technologies for preparation of this work.

Declaration of Competing Interest

The authors declare no conflict of interest.

Special issue articles Recommended articles

Data availability
No data was used for the research described in the article.

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1 Technically, surprise here refers to self-information (a.k.a., surprisal); namely, the implausibility of
some (sensory) outcome under a (generative) model of how that outcome was generated.

© 2023 The Author(s). Published by Elsevier B.V.

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