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Morally significant actions Morally bad actions action of all three might therefore constitute
TOM processing26.
Relatedly, mirror neurons in the PFC of
Betraying
the macaque monkey respond either when
Murdering the monkey makes a specific movement, such
Cheating Stealing as grasping with the index finger and thumb,
Lying
or when it sees another making the same
movement27. This indicates that when the
Tormenting animal sees another make the movement,
Assisting
the premotor cortex generates incipient
Self-sacrifice
motor commands to match the movement.
Defending
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.
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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