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Deontologists, exemplified by the Prussian

philosopher Immanuel Kant (17241804), do
not emphasize the consequences of actions, as
utilitarians do. Instead, they focus on the
Moral cognition and its neural maxim of the action the intent-based
principle that plays itself out in an agents
constituents mind. We must do our duty, as derived
from the dictates of pure reason and the
categorical imperative, for dutys sake alone.
William D. Casebeer Deontologists are particularly concerned to
highlight the duties that are owed to each other
Identifying the neural mechanisms of moral Here, I briefly review the neural mechanisms by free and reasonable creatures (paradigmati-
cognition is especially difficult. In part, this is of moral cognition, discuss methodological cally, humans). Maximizing happiness is not
because moral cognition taps multiple pitfalls and consider issues that might inform the goal; instead, ensuring that we do not
cognitive sub-processes, being a highly future experimental work. Ultimately, the cur- violate anothers rights is paramount2.
distributed, whole-brain affair. The rent situation makes the moral psychology Virtue theorists, such as the Greek
assumptions required to make progress in that is required by virtue theory the most philosophers Plato (427347 BC) and Aristotle
identifying the neural constituents of moral neurobiologically plausible, although this is a (384322 BC), make paramount the concept
cognition might simplify morally salient tentative, defeasible conclusion, and more of human flourishing3,4; to be maximally
stimuli to the point that they no longer work is needed to confirm it. moral is to function as well as one can given
activate the requisite neural architectures, ones nature. This involves the cultivation of
but the right experiments can overcome this Moral theories and moral cognition virtues (such as wisdom) and the avoidance
difficulty. The current evidence allows us to To study the neural mechanisms of moral of vices (such as intemperance), and is a
draw a tentative conclusion: the moral cognition, one must delimit the field practical affair.
psychology required by virtue theory is the of inquiry. What does moral cognition Each approach asks different things of us
most neurobiologically plausible. encompass? This depends on how we con- cognitively. What follows is an abbreviated
strue the domain of moral theory. Although discussion of each theorys moral psychology.
Good moral reasoning is extremely important all moral theories claim to speak to what To make the appropriate judgement about
for Homo sapiens. Our lives are more fruitful an agent should do (this is what makes them what one should do, the utilitarian would, at
if we recognize salient ethical norms and distinctively moral), they disagree about the least in morally problematic cases, require that
reason effectively about their application to substance of such recommendations and a moral agent could recognize and compute
our own situations. We are social creatures, the moral psychologies that are required for salient utility functions. We would then be
and if we are to flourish in our social environ- effective reasoning and action. The three moved to act on such judgements by cultiva-
ments, we must learn how to reason well main classic moral theories in the Western tion of appropriate altruistic fellow-feeling or,
about what we should do. Despite its impor- tradition are utilitarianism, deontology and in many cases, merely by self-concern (as
tance for our proper functioning, until virtue theory. utility will often be maximized by having
recently the neural mechanisms of moral The typical utilitarian, such as the British each of us focus on our own happiness as
cognition were not well studied. This is unfor- philosopher John Stuart Mill (18061873), well as the happiness of others). So, in terms
tunate, as co-evolution between the neural thinks that one should take that action (or of raw computations, a utilitarian moral
constituents of moral cognition and the follow that rule) that, if taken (or followed), psychology would require some mechanism
moral psychologies that are required by would produce the greatest amount of happi- for learning what actions or rules would even-
the main ethical theories is necessary if we are ness for the largest number of sentient beings, tually produce happiness. Either implicitly or
to make progress in understanding how effec- where happiness is the presence of pleasure or explicitly, utilitarian computations would
tive ethical reasoning is embodied in the the absence of pain (and where pleasure and constitute the bulk of our moral cognitive
brain. To make such progress requires us to pain are given more sophisticated readings capacity. Whether we act on the outcome
probe the nature of moral judgement and its than mere affective satisfaction). The second of those judgements might require some
relationship to the experimental regimens flavour of utility,rule utilitarianism, is probably derivative character development (such as the
that are used to explore such constituents. the most popular1. cultivation of concern for the happiness of



these approaches emphasize different brain

regions: frontal (Kant); pre-frontal, limbic and
sensory (Mill); and the properly coordinated
action of all (Aristotle) (FIG. 1).
Kant would say that moral reasoning is a
robustly rational affair, where rational is
given a strict interpretation. With Aristotle,
however, I think it is more useful to treat
moral judgement in a deflationary manner.
Given that the domain of what constitutes a
moral judgement is itself in contention, we
would be best served by casting our nets
widely, narrowing them appropriately as the
neurobiological, psychological and normative
aspects of morality co-evolve (admittedly,
casting our net so widely might bias us
initially towards a virtue-theoretic moral
psychology; the give and take required by
co-evolution of theories at all three levels of
analysis would, hopefully, correct any such
bias eventually). As a first cut, then, moral
cognition comprises any cognitive act that is
related to helping us ascertain and act on
what we should do. Non-human animals (for
example, primates and other social animals)
might also engage in robust moral reasoning
(see, for example, REF. 6).
Figure 1 | A carnival of philosophers. Kant is shown at lower left, Aristotle at upper right. Reproduced, My push for this deflationary conception
with permission, from REF. 34 (2002) MIT Press.
of moral judgement is driven by the recogni-
tion (but also has as the upshot) that moral
cognition might not be a tightly defined
others), and this would require appropriate shrift to character development and related natural kind in the sense that other cognitive
training of the emotions. issues, although recent work has softened up phenomena might be. For example, the
A Kantian moral psychology would be this position5. What exactly the cognitive domain of the neural mechanisms of visual
different. The ability to reason purely about capacity to reason purely in the Kantian man- cognition, owing to the relatively restricted
the demands of the categorical imperative ner would look like has not been a subject of range of information that is processed by the
(the heuristic that is used by Kant to capture extensive investigation; it would require at visual modality, might be more tightly
our respect for those things that make morality least the ability to check universalized maxims constrained than the domain of the neural
possible autonomy and rationality) would for logical consistency in a manner that is mechanisms of basket-weaving. In that sense,
be the most important part of our cognitive separable from the taint of affect and emotion. the former is a more robust, natural kind
equipment. The best known formulation of Finally, virtue-theoretic moral psychology than the latter, and is therefore an easier target
the categorical imperative requires that we act is often thought to be the richest of the three. for neurobiological study. Moral reasoning
only on maxims that we can will to become a A virtuous person must be able to reason well probably falls somewhere between these
universal law; other maxims are morally about what states of being would be most two extremes and is still worthy of study by
impermissible. For example, you could not conducive to the best life. What type of per- neurobiologists, although this fact might make
universalize the maxim that allows you to lie son must I become if I am to experience it more difficult to progress experimentally7.
to achieve some end; such a maxim requires eudaimonia (variously translated from the Critics might argue that such a co-
that others act with a different intention, of Greek as flourishing, proper functioning or evolutionary strategy commits the naturalistic
delivering and receiving only true utterances happiness)? To act on the outcomes of my fallacy of inferring what should be from what
(or else the lie would not be effective). judgements, I must train my character so that is. The exact nature and status of the natural-
The maxim cannot be made universal; this my appetites and spirit are coordinated istic fallacy is subject to debate (for a summary,
conceptual truth does not require experimen- smoothly with the demands of good reason. see chapter two of REF. 8). Note, however, that
tation to confirm it.We therefore have a perfect Virtuous people are moved to do the appro- the two most famous arguments against
(that is, exceptionless) duty not to form the priate thing at the appropriate time; they naturalism about ethics, Humes law9 and G. E.
intention to lie. Of note, Kant requires that we become angry at unjust events, are sympa- Moores open question argument10, do
be moved to do our duty by the demands of thetic to recipients of wrong-doing, and so not stand up against some contemporary
duty alone; if something else (say, the desire to on. Virtue theorists focus on the appropriate naturalized ethical theories. Both of these
be liked) is moving us, our action is not coordination of properly functioning cogni- arguments rely on an analytic/synthetic
morally praiseworthy as it plays more to our tive sub-entities. Moral reasoning and action distinction that many philosophers agree
animal nature than our rational (and so are therefore whole-psychology, whole-brain collapsed in the twentieth century. In addi-
human) nature. Kant is thought to give short affairs. Jokingly, then, it could be said that tion, Humes argument rules out deductive

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with hunger, thirst, sexual desire and the like

are powerful motivators; as our brains cortical
capacity expanded during evolution,newer
frontal brain regions remained connected to
and were innervated by this regulatory core.
No wonder then that these basic, survival-
laden emotions serve as the platform on which
the moral emotions (and effective moral
reasoning) are built.
Exploring the role of the prefrontal regions
in connecting limbic areas to frontal areas
is difficult; studies of humans with focal
brain damage and experimental lesions in
monkeys have indicated a relationship between
prefrontal cortex (PFC) and planning,
decision-making, emotion, attention, memory
for spatio-temporal patterns, and recognition
of a mismatch between intention and execu-
tion. The precise nature of the relationship
between these functions and the contribution
of the PFC remains unclear, in part because the
temporal and spatial resolution of functional
PFC PC BFB ATC magnetic resonance imaging (fMRI) is limited,
MFC Insula MOFC? STS and because we are still ascertaining the
ACC TH/MB Amygdala
network-level properties of this area.
Figure 2 | Five horizontal slices and one sagittal cross-section of a brain showing many morally Despite these difficulties, the link between
relevant neural areas. ACC, anterior cingulate cortex; ATC, anterior temporal cortex; BFB, basal moral decision-making, social cognition and
forebrain; FPC, frontopolar cortex; MFC, medial frontal cortex; MOFC, medial orbitofrontal cortex; PC,
the emotions is becoming clearer. Converging
precuneous; STS, superior temporal sulcus; TH/MB, thalamus/midbrain. Images courtesy of J. Moll.
results from lesion and imaging studies
indicate that damage to the ventral and
medial PFC is consistently associated with
relationships between facts and norms, this can be done, then the naturalistic fallacy impairments in practical and moral decision-
but not necessarily abductive (inference is not actually a fallacy (it merely amounts to making1215. Patients with focal ventromedial
to the best explanation style) relationships. saying that you dont have a good naturalized lesions show abnormally flat (emotionless)
And Moore himself admitted, in the second ethical theory yet). responses when shown emotional pictures,
version of his Principa Ethica, that his and perform poorly on tasks where feelings are
argument best applies to the two forms Emotion and affect needed to guide complex self-directed choices.
of naturalized ethics that he attacks in the The rich and diverse literature on the neural Electroencephalogram (EEG) studies of
book: Spencers evolutionary ethic and mechanisms of moral cognition can be use- children with self-control disorders16 also
hedonism. I agree that both of these natural- fully divided into three branches: the moral support a link between ventromedial PFC
ized ethics are poor moral theories, but emotions, theory of mind and abstract and moral emotions, as does fMRI of normal
I disagree that Moore has offered an moral reasoning. I will briefly discuss the subjects. For example, viewing scenes that
argument that is general to all attempts to connections between cortical areas and evoke moral emotions produces activation
naturalize ethics. the limbic system that are necessary for in the ventromedial PFC and the superior
Keep in mind that no good naturalized good moral judgement, the neural correlates temporal sulcus17,18.
ethical theory will say that all facts are norma- of theory of mind (TOM) and how they Within the ventral PFC, orbitofrontal
tive facts, nor that all existing states of affairs manifest themselves in moral judgement, cortex (OFC) is crucial for cueing morally
merely because they are natural in the and useful conceptual tools for thinking appropriate behaviour in adulthood and
sense that they were produced by natural about abstract moral reasoning. acquiring moral knowledge during childhood;
processes are good. Unlike Greene in his The moral emotions are crucial for effec- although patients with adult-onset and
companion piece in this issue11, I think that tive moral cognition. They motivate action, childhood-onset OFC damage showed similar
the neurobiological facts support a version of serve as markers of value, are vital for coordi- abnormal socio-moral behaviour, their scores
relational moral realism, but this discussion is nating group activity, and help to filter out and on standardized tests of moral reasoning
beyond the scope of my paper. Ultimately, highlight certain aspects of the moral calculus. differed. Those with early damage performed
even the most ardent anti-naturalist would In mammals and reptiles, the brains regula- poorly on the tests, exhibiting the egoistic
admit that, at the very least, our moral theo- tory core is situated in the brainstem/limbic reasoning that is typical of a ten year old,
ries must require us to carry out cognitive acts axis, and it subserves important activities such whereas adult-onset subjects performed
that are also possible for us to implement. The as breathing, arousal and the coordination of normally despite their abnormal behaviour15.
goal of naturalized ethics is to show that drives (for food, sex, oxygen and so on) with The PFC receives important inputs from
norms are natural, and that they arise from perceptions (turkey sandwich there, attractive both sensory and limbic areas. The limbic
and are justified by purely natural processes. If mate here). The basic emotions associated system is a highly interconnected set of



Morally significant actions Morally bad actions action of all three might therefore constitute
TOM processing26.
Relatedly, mirror neurons in the PFC of
the macaque monkey respond either when
Murdering the monkey makes a specific movement, such
Cheating Stealing as grasping with the index finger and thumb,
or when it sees another making the same
movement27. This indicates that when the
Tormenting animal sees another make the movement,
the premotor cortex generates incipient
motor commands to match the movement.
These signals might be detected as off-line
0 intentions that are used to interpret what is
Morally seen (for example, the dominant male
insignificant actions Morally praiseworthy actions intends to attack me). Mirror neurons might
Figure 3 | A moral state-space. The axes of this reduced state-space would correspond to functionally therefore bootstrap full-blown TOM into
salient groups of neurons, and regions of the state space might correspond to moral theories or they existence through an inner simulation of the
might help us to identify undiscovered moral concepts. Modified, with permission, from REF. 35 (1998) behaviour of others. TOM is probably what
Kluwer Academic Publishers.
allows a chimpanzee to know whether a high-
ranking male can see the food she is grabbing
or whether the food is occluded from view
subcortical regions (including the hippo- The cingulate cortex has a number of and can be taken without fear28. Robust TOM
campus, amygdala, hypothalamus and basal subregions with different functions: regulation is necessary for healthy moral judgement; it is
forebrain) and the cingulate cortex. The of selective attention, regulation of motivation, also associated with our ability to lie, but is
activity of this system is modulated by the and detection of malcoordinated intention nonetheless probably necessary if a whole
neurotransmitters dopamine, serotonin, and execution are associated with anterior host of morally important cognitive abilities
noradrenaline and acetylcholine, and changes regions (anterior cingulate cortex, ACC). are to be realized. For an excellent review of
in the levels of these substances can greatly Rostral ACC activation (along with the TOM mechanisms, see REF. 29.
affect sex drive, moods, emotions and aggres- nucleus accumbens, the caudate nucleus
siveness. The proper operation of the system and ventromedial (VM)/OFC) is needed for Abstract moral reasoning
as a whole is crucial for effective moral judge- cooperative behaviour among subjects playing Probably the most difficult aspect of the
ment. FIGURE 2 shows some of the brain areas a version of the prisoners dilemma; the neural mechanisms of moral cognition is
that are important in moral cognition. hyperscanning methodology used to obtain the constituents of abstract moral reasoning.
The amygdala, for example, is part of the these results is especially promising21 (see later The most important forms of moral reasoning
complex reward circuitry involving the positive in text). Other work supports the conclusion that we rely on daily, involve background
emotions19. It is likely that the amygdaloid that the ACC is crucial for identifying times social skills, tacit use of TOM, ready-at-hand
complex modulates the storage of emotionally when the organism needs to be more strongly action patterns and interpretive schema, and
important and arousing memories; events that engaged in controlling its behaviour2225. the like. Much of our day-to-day moral
are important to survival provoke specific reasoning does not involve highly convoluted
emotions and, with amygdala activity, are Theory of mind and moral judgement moral modelling; mostly, we can rely on skills
more likely to be permanently stored than This brief exploration of limbic areas and and habits of character as informed by condi-
neutral events. The amygdala is also crucial for their connections undervalues the role of PFC tioned emotion and affect (indeed, Haidt
aiding retrieval of socially relevant knowledge in the second important research area in claims that abstract moral reasoning is a
about facial appearance20; three subjects with moral cognition: social judgement and TOM. completely post-hoc affair and is almost never
total bilateral amygdala damage were asked to Our ability to know what others are thinking the direct cause of moral judgements)30.
judge the trustworthiness of unfamiliar people, so that we can interact fruitfully with them is Nonetheless, abstract moral reasoning is
and all three judged unfamiliar people to be vital it underlies our ability to empathize sometimes necessary. It probably depends on
more approachable and trustworthy than did with others, to judge how they might react in brain structures that subserve morally neutral
control subjects20. response to our actions, and to predict the abstract thought (such as a capacity to model
Hippocampal structures are essential for subjective consequences of our actions for the consequences of an action) and practical
learning and remembering specific events or conspecifics. Studies of children with autism reasoning about how to accomplish things.
episodes, although permanent memory indicate that TOM might be subserved by the For example, in a classic moral dilemma, such
storage lies elsewhere in the cortex. The aggregate neural activity of the OFC, the as the trolley problem (in which one has to
hippocampus, parahippocampal cortex, medial structures of the amygdala and the decide whether to allow an out-of-control
entorhinal cortex and perirhinal cortex all superior temporal sulcus (STS). The circuit trolley to continue down a track where it will
seem to be important for the processing and that is formed by the last two structures might strike five people or whether to throw a switch
retrieval of salient me-relevant memories. In mediate direction-of-gaze detection (a crucial diverting it onto a track where it will strike only
moral judgement, the hippocampus might component of our ability to infer what others one person, explored in detail in fMRI work
facilitate conscious recollection of schemas might be thinking about), all three locations by Greene et al.31), higher-order cognitive abili-
and memories that allow past events to affect are probably involved in mediating shared ties such as planning, executive flexibility and
current decisions. attention, and the specially coordinated strategy application (see REF. 32) are needed.

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Hyperscan experiment tioned memories and insight into the minds of

others, so as to think about and actually behave
in a maximally functional manner. There is
clear consilience between contemporary
neuroethics and Aristotelian moral psychology.
A co-evolutionary strategy, then, would suggest
that some version of pragmatic Aristotelian
virtue theory is most compatible with the
neurobiological sciences.
The localization work that is mentioned in
Client Client this paper uses various techniques and experi-
mental regimens. The stimuli normally range
from sentences to small photographs. The
Hyperscan server
and database constraints of rigorous experimental design
mean that tests are sometimes conducted in
highly artificial situations; ecological validity
for moral reasoning is difficult. Moral cogni-
Client Client tion exhibits several characteristics that make
it difficult to capture in the fMRI chamber37.
A list of these characterisitics follows.

Figure 4 | A schematic of four subjects engaged in a hyperscanning experiment, interacting Moral cognition is hot. Owing to evolutionary
through an internet connection. Such a setup allows simultaneous functional neuroimaging of subjects
history, affective and conative states are
as they interact with each other. The data collected by each functional magnetic image is passed, through
a client, to a hyperscan server and database, which is regulated by an experiment controller. Image
part and parcel of effective moral judgement.
courtesy of Read Montague. This is endemic to moral reasoning (consider
the Damasios patient E.V.R., whose PFC
damage, like that of Phineas Gage, disrupted
These capacities might be realized in cerebral neurons or relevant populations of neurons, the connections between limbic somatic
cortex by transient cortical networks which and tag each of them as being a dimension markers and frontal cortex, resulting in poor
Fuster calls cognits33. of the space, using the right statistical tools moral judgement14). Unfortunately, hot cogni-
The difficulty we have in understanding the (principal or independent components tion is difficult to capture in artificial settings.
neural basis of moral reasoning is indicative of analysis primarily), we can reduce the dimen-
two things: first, that we still need both better sionality of the space to something that is Moral cognition is social. Several crucial
theoretical frameworks to understand higher- more manageable. components of the neural constituents of
order cognitive capacities and better network- To behave morally would be to have this moral cognition aim to achieve appropriate
level tools for probing activity; and second, state-space allocated appropriately (presum- behaviour in social and group settings. This
that such capacities might be overvalued ably by the conjunction of experience and the is no accident: animal and human groups
relative to the work that they perform in our ontogeny of native neurobiological equip- are social groups. Social environments are
cognitive economy (in some respects, then, ment) so that one is maximally moral (which, difficult to simulate in the scanner. A notable
eliminativism might be called for; see REF. 34). in the case of virtue theory, means being max- methodological improvement in this area
One device that might be useful for helping imally functional). The axes of this reduced is Montagues use of multi-scanner hyper-
us to organize abstract moral reasoning is a state-space would correspond to functionally scanning methodology21, in which several
moral state-space (a concept first articulated by salient groups of neurons, and regions of the subjects can interact simultaneously while
P. M. Churchland)35. We can think of much of state-space might correspond to the big three being scanned (FIG. 4). Although the technology
the activity of frontal cortex and the limbic/ traditional moral theories that were discussed is currently used to link-up subjects from
brainstem axis as consisting of a moving point earlier, or they might help us to identify multiple states concomitantly, it could be used
in an n-dimensional space, where n could (in undiscovered moral concepts. The idea of a to monitor the interactions of subjects who
complex cases) be determined by making the moral state-space is one way to conceptually are within sight of each other. Minimally, this
activity of every neuron that is involved in unify disparate brain activity that is related to technology offers added efficiency in studying
the system an axis of that space (in some cases, moral cognition. social interactions; maximally, however,
an axis might be constituted by a single neuron, it adds another dimension to the study of
which might explain results about how single Consilience with virtue theory the neural mechanisms of social reasoning.
neurons in PFC can seem to encode rules36). The evidence, albeit tentative, that we have As Montague et al. note, studying social
Reducing the dimensions of this space enables discussed lends more credence to the moral interactions by scanning the brain of just one
us to capture its principal components, which psychology that is required by virtue theory. person is analogous to studying synapses
might themselves correspond to traditional Empirically successful moral cognition on the while observing either the presynaptic neuron
moral concepts that have been explored by part of an organism requires the appropriate or the postsynaptic neuron, but never both
ethicists for the last 2,500 years. The idea of coordination of multi-modal signals conjoined simultaneously synapses, like socially
a moral state-space (FIG. 3) allows us to aggre- with appropriately cued executive systems interacting people, are best understood
gate various cortical regions involved in the that share rich connections with affective and by simultaneously studying the interacting
processing of moral concepts: if we identify conative brain structures that draw on condi- components21 (FIG. 4).



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