Current Biology Vol 22 No 23
R994
4. Nudds, R.L., and Dyke, G.J. (2010). Narrow
primary feather rachises in Confuciusornis and
Archaeopteryx suggest poor flight ability.
Science 328, 887–889.
5. Longrich, N.R. (2006). Structure and function
of hindlimb feathers in Archaeopteryx
lithographica. Paleobiology 32, 417–431.
6. Xu, X., Zhou, Z.-H., Wang, X.-L., Kuang, X.-W.,
Zhang, F.-C., and Du, X.-K. (2003).
Four-winged dinosaurs from China. Nature
421, 335–340.
7. Hu, D.Y., Hou, L.-H., Zhang, L.J., and Xu, X.
(2009). A pre-Archaeopteryx troodontid from
China with long feathers on the metatarsus.
Nature 461, 640–643.
8. Longrich, N.R., Vinther, J., Meng, Q.J., Li, Q.,
and Russell, A.P. (2012). Primitive wing feather
arrangement in Archaeopteryx lithographica
and Anchiornis huxleyi. Curr. Biol. 22,
2262–2267.
9. Lucas, A.M., and Stettenheim, P.R. (1972).
Avian anatomy: integument (Washington. D.C.:
United States Department of Agriculture).
10. Rietschel, S. (1985). Feathers and wings of
Archaeopteryx, and the question of her flight
ability. In The beginnings of birds, M.K. Hecht,
J.H. Ostrom, G. Viohl, and P. Wellnhofer, eds.
Neuroscience: A More Dynamic View
of the Social Brain
Information relevant for social interactions is thought to be processed in
specific neural circuits. Recent studies shed new light on how that social
information is encoded and processed by different brain areas.
J. Sallet1, R.B. Mars1,2,
and M.F.S. Rushworth1,2
Human and non-human primates are
social animals and living in complex
social environments has an impact on
brain structure and function [1–3].
Social information is thought to be
processed by a specific set of neural
circuits often referred to as the ‘social
brain’ [4–6]. How social information is
encoded and the nature of the
computations performed by different
brain areas is nevertheless still
debated. Papers in this issue of Current
Biology by Watson and Platt [7] and
Santiesteban et al. [8], focusing on
the orbitofrontal cortex (OFC) and
the temporo-parietal junction (TPJ),
respectively (Figure 1), begin to unpick
some of these issues. These two
studies nicely complement recent
observations on the roles of the OFC
and TPJ from other laboratories [9,10].
Altogether, the results suggest that
the context in which a social decision
is taken strongly affects how the
information is processed, suggesting
a quite dynamic view of how social
information is encoded and used by the
social brain.
(Eichstatt: Freunde des Jura-Museums),
pp. 251–260.
11. Hoyle, F., Wickramasinghe, N.C., and
Rabilizirov, R. (1985). Archaeopteryx: Problems
arise – and a motive. British Journal of
Photography 132, 693–695.
12. Feduccia, A., and Tordoff, H.B. (1979). Feathers
of Archaeopteryx: asymmetric vanes indicate
aerodynamic function. Science 203,
1021–1022.
13. Senter, P. (2006). Scapular orientation in
theropods and basal birds and the origin of
flapping flight. Acta Palaeontol. Polonica 51,
305–313.
14. Xu, X., You, H., Du, K., and Han, F. (2011). An
Archaeopteryx-like theropod from China and
the origin of Avialae. Nature 475, 465–470.
15. Tucker, V.A. (1993). Gliding birds: reduction of
induced drag by wing tip slots between the
primary feathers. Journal of Experimental
Biology 180, 285–310.
16. Xu, X., and Guo, Y. (2009). The origin and early
evolution of feathers: insights from recent
paleontological and neontological data.
Vertebrata Pal Asiatica 47, 311–329.
17. Prum, R.O., and Brush, A.H. (2002). The
evolutionary origin and diversification of
In their study, Watson and Platt [7]
presented macaque monkeys with
a simple decision-making task in which
they could sacrifice juice to watch
social images — pictures of other
animals of various social statuses or
pictures of female macaque perinea
that male macaques appear to find
intrinsically interesting. They found that
neural activity in the OFC is modulated
by both social and reward information,
but most of the OFC neurons that show
such modulated activity are either
sensitive to reward or to social
information [7]. This is one of the first
tests of OFC single neuron activity in
the social domain, but the result is
broadly in line with previous reports
that only a small proportion of OFC
neurons multiplex values across
reward dimensions [11].
It is interesting to compare the
results of the Watson and Platt study [7]
with those from the one other recent
investigation of single neuron activity
in OFC during social cognition. Azzi
et al. [9] taught monkeys a simple
oculomotor task in which their choices
could lead to rewards either for just
themselves or for a second animal too.
The authors argue that the receipt of
feathers. Quarterly Review of Biology 77,
261–295.
18. Xu, X., Zheng, X.T., and You, H.L. (2010).
Exceptional dinosaur fossils show ontogenetic
development of early feathers. Nature 464,
1338–1341.
19. Lee, M.S.Y., and Worthy, T.H. (2011). Likelihood
reinstates Archaeopteryx as a primitive bird.
Biology letters.
20. Agnolı́n, F.L., and Novas, F.E. (2011).
Unenlagiid theropods: are they members of the
Dromaeosauridae (Theropoda, Maniraptora)?
Anais Da Academia Brasileira de Ciencias 83,
117–162.
Institute of Vertebrate Paleontology and
Paleonanthropology, Key Laboratory of
Evolutionary Systematics of Vertebrates,
Chinese Academy of Sciences,
142 Xiwai Street, Beijing, 100044, China.
E-mail: xuxing@ivpp.ac.cn
http://dx.doi.org/10.1016/j.cub.2012.10.015
a reward by the second macaque
apparently diminishes the value of
rewards received by the first macaque.
They found that lateral OFC neurons
also reflect this apparent diminution of
the reward value for the experimental
animal when rewards are
simultaneously given to another
macaque. In other words, Azzi et al. [9]
report the consequences of integrating
both social and reward information in
the firing rates of individual OFC
neurons.
These two studies, by Watson and
Platt [7] and by Azzi et al. [9], can be
seen to be complementary if one
remembers that, in order to be able to
learn the links that exist between
choices and their outcomes, it is crucial
to compute not only the scalar values of
choice outcomes but also the identities
of choice outcomes [12,13]. In other
words, it is not just important whether
an outcome is rewarding, but it is
important to know what type of reward
it is. One of the other key results
reported by Watson and Platt [7] is the
observation of a strong modulation of
OFC activity by social categories
(images with a dominant animal,
a subordinate animal or with sexual
content) — just what would be needed
if the monkey is to learn about the
precise category of outcome that is to
be expected from a choice.
But while it is essential to understand
the identity of the outcome that is
expected from a choice, it is also
important to be able to compare the
values of different outcomes on a single
Dispatch
R995
Figure 1. Localization of the temporo-parietal junction on a human brain (left) and the orbito-
frontal cortex on a macaque brain (right).
The two brains are not represented at the same scale. Note that the localization in the macaque
of an analogous area of the human TPJ is still debated. STS: superior temporal sulcus.
scale so that the preferred outcome
can be ascertained. Both macaque and
human studies are beginning to
suggest that this second process might
be more closely linked to medial OFC
and adjacent ventromedial prefrontal
cortex rather than lateral OFC [14–16].
Some evidence that such a process
operates even in the social domain can
be gleaned from the recent OFC
recording studies, from the observation
of a weak modulation of OFC activity by
the value of social stimuli by Watson
and Platt [7] and from the effect of the
social status of the second macaque in
the study by Azzi et al. [9].
One of the most dramatic ways in
which the social context can affect
cognition is when a person actually
takes on the perspective of another
person at the expense of their own
perspective. The second study in
Current Biology, by Santiesteban et al.
[8], suggests that the TPJ is critical for
such a change in social perspective.
The authors applied transcranial direct
current stimulation (tDCS), with the
anode placed over the TPJ, while
people performed various behavioral
tasks in which either their own ‘self
perspective’ or another’s perspective
had to be considered. Such an
electrode arrangement is thought likely
to facilitate the operation of the TPJ. In
one task, the subjects were instructed
to imitate a finger movement they saw
on a screen or to make a different finger
movement. In a second task, they
were asked to suppress their own
perspective on a scene and to adopt
that of another person. In both cases,
tDCS over TPJ was seen to induce
improvements in performance.
The findings reported by
Santiesteban et al. [8] complement
another recent study [10] in which
subjects were asked to choose
between two gambles to reward either
themselves or someone else. The
results of this experiment show that
TPJ signals (as well as dorsomedial
frontal signals) can flip between
encoding the value to one’s own self of
the choices that are being taken, or the
value of a choice for another person
[10]. The on-line control over which
representation to use might depend on
a learning mechanism that reports
deviation from expectations. Such a
mechanism has been associated with
the TPJ [17].
A common theme of dynamic
change — both of how outcome values
are represented when alone and in
a social context [7] and of how
perspectives can be shifted between
the personal and the social [8] — runs
through the two new Current Biology
papers. Such dynamism is not likely to
be restricted to the ways in which
a single brain area works but it is likely
to be a feature of how areas interact in
different configurations in order to
mediate social cognition.
References
1. Sallet, J., Mars, R.B., Noonan, M.P.,
Andersson, J.L., O’Reilly, J.X., Jbabdi, S.,
Croxson, P.L., Jenkinson, M., Miller, K.L., and
Rushworth, M.F. (2011). Social network size
affects neural circuits in macaques. Science
334, 697–700.
2. Bickart, K.C., Wright, C.I., Dautoff, R.J.,
Dickerson, B.C., and Barrett, L.F. (2011).
Amygdala volume and social network size in
humans. Nat. Neurosci. 14, 163–164.
3. Dunbar, R.I., and Shultz, S. (2007). Evolution in
the social brain. Science 317, 1344–1347.
4. Adolphs, R. (2009). The social brain: neural
basis of social knowledge. Annu. Rev. Psychol.
60, 693–716.
5. Frith, C.D. (2007). The social brain? Philos.
Trans. R. Soc. Lond. B. Biol. Sci. 362, 671–678.
6. Behrens, T.E., Hunt, L.T., and Rushworth, M.F.
(2009). The computation of social behavior.
Science 324, 1160–1164.
7. Watson, K.K., and Platt, M.L. (2012). Social
signals in primate orbitofrontal cortex. Curr.
Biol. 22, 2268–2273.
8. Santiesteban, I., Banissy, M.J., Catmur, C., and
Bird, G. (2012). Enhancing social ability by
stimulating the right temporoparietal junction.
Curr. Biol. 22, 2274–2277.
9. Azzi, J.C., Sirigu, A., and Duhamel, J.R. (2012).
Modulation of value representation by social
context in the primate orbitofrontal cortex.
Proc. Natl. Acad. Sci. USA 109, 4020.
10. Nicolle, A., Klein-Flügge, M.C., Hunt, L.T.,
Vlaev, I., Dolan, R.J., and Behrens, T.E.J. (2012).
An independent axis for executed and modeled
choice in medial prefrontal cortex. Neuron 75,
1114–1121.
11. Kennerley, S.W., Behrens, T.E., and Wallis, J.D.
(2011). Double dissociation of value
computations in orbitofrontal and anterior
cingulate neurons. Nat. Neurosci. 14,
1581–1589.
12. Niv, Y., and Schoenbaum, G. (2008). Dialogues
on prediction errors. Trends Cogn. Sci. 12,
265–272.
13. Noonan, M.P., Mars, R.B., and Rushworth, M.F.
(2011). Distinct roles of three frontal cortical
areas in reward-guided behavior. J. Neurosci.
31, 14399–14412.
14. Noonan, M.P., Walton, M.E., Behrens, T.E.,
Sallet, J., Buckley, M.J., and Rushworth, M.F.
(2010). Separate value comparison and learning
mechanisms in macaque medial and lateral
orbitofrontal cortex. Proc. Natl. Acad. Sci. USA
107, 20547–20552.
15. Rudebeck, P.H., and Murray, E.A. (2011).
Balkanizing the primate orbitofrontal cortex:
distinct subregions for comparing and
contrasting values. Ann. N. Y. Acad. Sci. 1239,
1–13.
16. Bouret, S., and Richmond, B.J. (2010).
Ventromedial and orbital prefrontalneurons
differentially encode internally and externally
driven motivational values in monkeys.
J. Neurosci. 30, 8591–8601.
17. Suzuki, S., Harasawa, N., Ueno, K., Gardner, J.L.,
Ichinohe, N., Haruno, M., Cheng, K., and
Nakahara, H. (2012). Learning to simulate other’s
decisions. Neuron 74, 1125–1137.
1Department of Experimental Psychology,
University of Oxford, Oxford, OX1 3UD, UK.
2Oxford Centre for Functional Magnetic
Resonance Imaging of the Brain (FMRIB),
University of Oxford, Oxford, OX1 3UD, UK.
E-mail: jerome.sallet@psy.ox.ac.uk
http://dx.doi.org/10.1016/j.cub.2012.10.051