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Encephalon #25 arrives June 20, 2007

Posted by Johan in Developmental Psychology, Links, Social Neuroscience.
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Grab it now at Psyblog. Some favourite posts:

Developing Intelligence reports on some evidence that children have a difficult time telling fantasy from reality. This notion may seem common sense, but this is one of the first empirical demonstrations I’ve heard of.

Omnibrain posted a video of a laughing rat, which almost made me run out and get one for myself.

Finally, Memoirs of a Postgrad explains what embodiment means, as applied in embodied cognition but also in AI research. With mirror neurons being all the rage these days, embodied theories are everywhere. This post is a nice introduction to what the fuss is all about. It’s worth emphasising that embodiment theories preceded  the discovery of mirror neurons, and indeed, it’s not clear that mirror neurons are necessary for embodied representations on a neural level.

Encephalon #23 Arrives May 23, 2007

Posted by Johan in Links.
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Madam Fathom hosts the 23rd issue of Neuroscience blogging carnival Encephalon. It’s quite a good write-up, and I’m feeling the pressure – the next issue will be hosted here, on June 4. Contributions go to encephalon.host(at)gmail.com, as usual.

Some favourites from this issue:

Developing Intelligence offers a scathing attack on reductionism. I’m not sure if I agree with the analysis, but it’s an interesting read.

Memoirs of a Postgrad contributes a review of theories of embodied cognition and how these relate to AI research.

Finally, Neurozone has a post relating mirror neurons to language. I actually found this interesting, which, given my profound disinterest in psycholinguistics, must mean that the post is quite extraordinary.

Object- or Viewer-centered Coding in the Superior Temporal Sulcus April 22, 2007

Posted by Johan in Neuroscience, Sensation and Perception, Social Neuroscience.
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The Superior Temporal Sulcus (STS) is a groove in the temporal lobe (shown in a Macaque monkey above) that appears to play a special role in visual perception: neurons in this area respond selectively to various forms of biological motion. For instance, the figure above shows the location of neurons in the STS that respond to walking and bending of the knees (we’ll get back to this figure shortly).

Undoubtedly, some of you will assume that you’re now reading yet another post about mirror neurons, but this is not true. Mirror neurons are found more anterior (ie to the front) to the STS in the ventral prefrontal cortex, and above the STS in the anterior inferior parietal lobule (all this in macaques, mind). As you’re all painfully aware by now, mirror neurons are so called because they respond both to the perception of (mainly goal-directed) actions in others, and to corresponding actions in the monkey itself. In contrast, neurons in the STS only respond to the actions of others. So it would seem that the monkey (and presumably also human) brain has at least three distinct areas that deal more or less directly with perceiving the actions of other individuals, though it’s far from clear how these areas divide up the work, or how they are inter-connected.

The figure above is from a paper by Jellema and Perrett (2006) that sought to investigate whether STS neurons use object- or viewer-centered coding. These abstract ideas can be boiled down to the following: if the neurons are object-centered, they will respond to the same stimulus no matter which way it’s turned, while viewer-centered neurons would only respond to the stimulus when it is presented in a given orientation. Jellema and Perrett (2006) also argue that a third coding category exists, namely goal-directed coding. Goal-directed coding can be seen in mirror neurons that only respond when the experimenter picks up an object, not to the pantomine of picking up without an object, or to the object viewed in isolation. A STS-relevant example would be the neurons that Jellema and Perrett (2006) found, which respond to bending of the knees only when the legs are standing on a surface. Bending of the knees in the air produced no response.

But how can they know that the neuron responded to this, and not some other aspect of the stimulus? Well, the knee-bending example turns out to provide a nice example of how the work involved in these biological motion single-cell recording studies is carried out:

The simplest explanation for the cell’s responsiveness would of course be that it responded to an object moving downward. [...] Therefore, we presented an agent jumping from a 40 cm high elevation while keeping the knees straight. This constituted a lowering of the body without knee flexing, and produced a very much reduced response (as well as sore knees for the agent!)

As you may have feared, the basic design is to have someone stand in front of the monkey doing stuff, all the while listening to the neuronal firing rate over speakers to spot when the recorded neuron responds (not that I can think of a better way of testing this).

And how was the the goal-directed knee-bending response discovered?

Knee flexion with the principal body axis oriented horizontally was achieved by the experimenter lying on a mobile table, at 2 m distance from the subject. Two different versions were presented. In the first, the feet made contact with the wall of the testing room throughout the flexing and straightening cycles. In this scenario, knee extension was achieved by exerting force onto the wall, thereby pushing the mobile table (with the experimenter on top) away from the wall. To achieve knee bending, a second experimenter (who remained out of sight) pushed the table towards the wall. In the second version, the feet never touched the wall (50 cm minimal distance between feet and wall). The experimenter simply flexed and straightened the knees in the air just above the surface of the table. Interestingly, only knee bending with the feet maintaining contact with the wall produced a response (Fig. 3g), while knee flexing without the feet making contact with the wall produced no response at all.

This must have been one confused monkey. Do note the subtle distinction in who’s carrying out the task: the comfy knee-bending is done by an “experimenter,” while the 40 cm knees-straight drop is performed by an “agent,” which I presume is a euphemism for “undergraduate research assistant.”

Jellema and Perrett (2006) report on a range of STS neurons that respond to various behaviours, but their principal findings are well summarised by the figure at the top of this post. The series of coronal STS slices goes from anterior (left) to posterior (right). Note that Jellema and Perrett (2006) only recorded from the anterior part of the STS. As you can see, it’s a bit of a mixed bag – some cells are object-centered, others viewer-centered. The principal finding is the large group of object-centered neurons that can be seen as grey rings and triangles in the slices marked +18 and +16. Previous investigations had not found as large groups of object-centered neurons, which Jellema and Perrett (2006) attribute to the relatively anterior recording site that they used. This matches the overall trend in their data for viewer-centered coding to appear in the posterior sections, while object-centered coding appears in the anterior. This is also supported by previous findings that response latencies are longer at anterior sites, and that the main outputs from the STS are in the anterior region. The authors suggest that since object-centered representations likely build on viewer-centered representations, processing in the STS may move back to front.

Another thing to note from the figure above is that all the responding neurons were in the upper bank of the STS. Jellema and Perrett (2006) did record from both, so it’s an open question what those neurons might respond to – apparently it isn’t knee-bending and shattered knees, at least.

Finally, Jellema and Perrett (2006) argue that the goal-centered neurons they found may represent basic understanding of causal relations (“he picked up the ball”), if the STS projects on to mirror neurons in premotor cortex. They also note, however, that behavioural evidence shows that monkeys have little capacity to understand or imitate the actions of others – this is apparently the territory of primates. With this in mind, the neurons they’ve found may, in concert with previously reported mirror neurons, form the basis of what rudimentary social learning and imitation that monkeys are capable of. But the fact that primates are capable of visually-guided imitation suggests that we have something that the macaques lack. Thus, it would seem that macaque monkeys (on whom the bulk of the mirror neuron research was carried out, incidentally) are somewhat lacking in this area as models of human neuroanatomy.

References
Jellema, T., & Perrett, D.I. (2006). Neural representations of perceived bodily actions using a categorical frame of reference. Neuropsychologia, 44, 1535-1546.

Why mirror neurons isn’t the whole story April 16, 2007

Posted by Johan in Developmental Psychology, Neuroscience, Social Neuroscience.
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How do we understand the thoughts and actions of others? Traditionally, this is explained by positing that humans use a kind of naive theory of psychology to make sense of the often limited and contradictory input that we receive in social situations. This naive theory is far from perfect. For example, we tend to assume that the behaviour of others is far more likely to have been caused by dispositional than situational factors (the fundamental attribution error). So as naive theorists, we are inclined to jump to conclusions when it comes to explaining the behaviour of others.

More recently, this “theory” view of predicting and explaining actions has met some challenge from a group of theories that I will refer to collectively as simulation theory. Fueled by the recent discovery of mirror neurons in monkeys (e.g., Rizzolatti & Craighero, 2004; see a previous post), the basic idea of simulation theory is that others are understood by simulating their minds in one’s own mind. When I see a woman on the subway with her head in her hands and a stricken expression, I need not refer to poorly defined and unparsimonious concepts or naive theories about how this behaviour is to be interpreted. I simply simulate (with the aid of a mirroring system) how I would feel if I were in the same position, with the same facial expression. Simulation theory draws a lot of its appeal from the apparent parsimony of using our intuitive understanding of our own mental states to understand the mental states of others.

The basic idea of simulation theory is simple and parsimonious, yet it seeks to explain something as complex and seemingly-irrational as human social behaviour. Unsurprisingly, it runs into some problems. A paper by Saxe (2005) gets at many of these problems by showing how systematic errors in perception and reasoning (for example, the previously-mentioned fundamental attribution error) can’t be explained by simulation theory. Saxe (2005) concedes that a mirror system may well underpin “simple actions and some basic emotions” (p. 174), but maintains that a naive theory must be used for more complex attributions and predictions. To make this point, she draws on empirical examples of cognitive errors and biases:

  1. Children frequently confuse “not knowing” and “getting it wrong.” When 4-year-olds perform a task where they but not an adult observer know the colour of a bead (of a known set of two colours) that has been moved into a container, the children generally reason that the adult will believe that the colour is the wrong one. This is incorrect, as the adult is equally likely to guess that the colour is the right or the wrong one, since he or she does not know which colour it really was.
  2. 3-year-olds fail to realize that all properties of an object are not known automatically by a person who interacts with it. Thus, they do not understand that an object’s colour can be known if the object is seen, but not if it is merely touched, or that the firmness of an object can be known through touching, but not seeing.
  3. These egocentric errors do not all disappear at the same age. For instance, children are able to understand that other people have different desires and perceptions at two years, but will not understand that beliefs can also differ until a year later. This implies that different concepts underlie the different tasks, which simulation theory does not allow.
  4. Adults are generally convinced that their reasoning is entirely rational and free from such errors as the self-serving bias, but when this is probed experimentally, actual behaviour is a far cry from normative logic. If we use simulation to predict how we will reason, why is the actual behaviour so different from the simulation?
  5. In situations where the self-serving bias is known, adults engage in”naive cynicism,” that is, an overly pessimistic estimation of another person’s self-serving bias. For instance, when married couples were asked how often they were responsible for desirable and undesirable marital events, and how often they thought that the other spouse would assume responsibility for desirable and undesirable events, the spouses estimated that the other would be self-serving, even though the actual data showed that both took credit equally for desirable and undesirable events. It’s difficult to explain why this occurs without again invoking different concepts.

All the above examples probe the same inconsistency: our failure to consider others in the same way that we consider ourselves. In its simplest form, simulation theory predicts that these social biases simply should not occur.

Perhaps imperfect simulation can explain these discrepancies? It’s reasonable to suggest that any simulation is going to be inferior to first-hand experiences, and we do not lose much of the ever-alluring parsimony of simulation theory by positing that the simulated experience is weaker or somehow degraded. Saxe (2005) doesn’t buy this explanation. For example, the case of the beads outlined in point 1 above cannot be explained in this way. To re-iterate, the child sees the colour of a bead being put into a container, while an adult does not. The child is then asked what colour the adult will think that the bead is. Either the child simulates the mind of the other appropriately, in which case performance should be good, or it fails to simulate the mind of the other, and assumes that the adult knows what the child knows. But this isn’t what happens – as described above, the child insists that the adult will always pick the wrong bead, instead of guessing (proper simulation) or picking the right bead (failed simulation). Similar problems arise with the self-serving bias examples in points 4 and 5. Why is performance so different from the norm in the couple situation?

Fundamentally, the problem may go back to the tired old controlled/automatic distinction. Simulation theory posits a bottom-up, automatic process, which is unaffected by beliefs, expectations, and other aspects of a naive theory. As such, simulation theory works very well when explaining behaviour in paradigms such as facial expression identification, where people seem to have an uncanny ability to identify expressions extremely quickly, and with little effort. The “theory” approach has a better shot at explaining top-down influences in reasoning. It’s worth noting that Saxe’s paradigm, where children are essentially told stories and are then asked what different actors in the story might know or think, is one well-suited to this approach. The examples Saxe (2005) cites as evidence against simulation theory (in points 1-5 above) rely on making inferences in this controlled manner.

Perhaps it is too much to ask that one theory should explain both the fast, automatic processing required in typical simulation paradigms, and the thoughtful, controlled processing in typical “theory” paradigms.

References
Rizzolatti, G., & Craighero, L. (2004). The Mirror-Neuron System. Annual Review of Neuroscience, 27, 169-192.

Saxe, R. (2005). Against Simulation: The Argument from Error. Trends in Cognitive Sciences, 9, 174-179.

Mirror Neurons and Communicative Actions January 9, 2007

Posted by Johan in Neuroscience, Raves.
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Ah, what one is not willing to go through in the name of science. This figure comes from an article by Ferrari et al (2003), which investigated mirror neuron responses to mouth actions. Mirror neurons are neurons which have been found in the premotor cortex of monkeys and apes, and are believed to exist in humans as well. These neurons fire when the animal performs a specific action (e.g., making a face with portruded lips, as above), but also when the animal sees someone else perform a similar action (Rizzolatti & Craighero, 2004).

This is terribly interesting because it appears to offer an explanation for how we make sense of the actions of other people: in this view, we understand actions by “mirroring” them in our own motor system.

The study by Ferrari et al (2003) is interesting because it contradicts the previously canonical knowledge that mirror neurons only respond to object-oriented actions, i.e., miming a grasping action yields no response, nor does the object itself. Only when an object is grasped does the neuron respond (Rizzolatti & Craighero, 2004). Ferrari et al identified a group of mirror neurons in the macaque monkey’s area F5 which responded to mouth movements. The bulk of these “mouth” neurons responded to eating-related actions, but a small group responded to communicative mouth movements, such as the lip portrusion pictured above. While you could argue that eating by necessity is an object-oriented action, the communicative movements are not so easily put into this framework.

The implication for what I’m working with right now is that mirror neurons may play a role in detecting facial expressions in humans. There is considerable evidence that faces and in particular facial expressions are perceived quite differently from other stimuli. Mirror neurons may play some part in explaining these differences.

References

Ferrari, P.F., Gallese, V., Rizzolatti, G., & Fogassi, L. (2003). Mirror Neurons Responding to the Observation of Ingestive and Communicative Mouth Actions in the Monkey Ventral Premotor Cortex. European Journal of Neuroscience, 17, 1703-1714.

Rizzolatti, G., & Craighero, L. (2004). The Mirror-Neuron System. Annual Review of Neuroscience, 27, 169-192.