Language Evolution: An invisible hand.

A slightly edited "Editor's Summary" from the Oct. 11 Nature:

As a language evolves, grammatical rules emerge and exceptions die out. Lieberman et al. have calculated the rate at which a language grows more regular, based on 1,200 years of English usage. Of 177 irregular verbs, 79 became regular in the last millennium. And the trend follows a simple rule: a verb's half-life scales as the square root of its frequency. Irregular verbs that are 100 times as rare regularize 10 times faster. The emergence of a rule (such as adding –ed for the past tense) spells death for exceptional forms...In a separate study, Pagel et al. looked at changing word meanings. Across the Indo-European languages, words like 'tail' or 'bird' evolve rapidly and are expressed by many unrelated words. Others, like 'two', are expressed by closely related word forms across the whole language family. Data from over 80 modern languages show that the more a word is used, the less it changes.

And, from Fitch's review of the two papers:

The words of language are not inherited biologically, but are passed on culturally through learning. This process of 'cultural evolution' generates a hierarchical tree of relationships among languages, here illustrated by the Indo-European family. Just as descent with modification in biological evolution (phylogeny) leads to phylogenetic trees, so the analogous process in language change (glossogeny) can lead to glossogenetic trees.

Where should we look to gain a deeper understanding of the invisible hand in the cultural evolution of language? A promising future direction is provided by recent attempts to fuse theoretical models of cultural evolution to experimental investigations of social learning in the laboratory. Experimental investigations of 'iterated learning' — similar to the game of Chinese whispers, where one participant's output serves as input for the next — can provide empirical data to inspire, and constrain, our theories. Sophisticated new theoretical models enable language-learning 'agents' to have both innate biases (in the form of so-called bayesian priors) and powerful statistical learning systems capable of discovering and using environmental regularities. Such models demonstrate the possibility of a very indirect and sometimes non-intuitive relationship between the regularities emerging at the level of a whole population and the underlying generating forces. These forces are individual behaviour and learning (social usage) and innate constraints (in Chomsky's terms, a 'language acquisition device', often called universal grammar).

..some of the most persistent 'cultural replicators' — memes — evolve as slowly as some genes. By documenting and quantifying such effects, this work opens the door to a diverse range of theoretical and empirical investigations. If there is ever to be a science of memetics o rival that of genetics, it should proceed along these lines: combining careful quantitative analysis of well-documented linguistic changes with sophisticated theoretical models capable of taking into account the multilayered complexity of cultural evolution.

Emotional enhancement of memory and learning - a molecular mechanism.

Hu et al. have performed an interesting study of how the rush of noradrenaline during an emotional experience might enhance our ability to recall that experience. It causes a long term enhancement of the activity of nerve synapses using the neurotransmitter glutamate by stimulating the phosphorylation of a particular type of glutamate receptor. I'm afraid the molecular details will only make sense to those of you who know something about the chemistry of nerve transmission, but here they are, as outlined in the article's abstract:

Emotion enhances our ability to form vivid memories of even trivial events. Norepinephrine (NE), a neuromodulator released during emotional arousal, plays a central role in the emotional regulation of memory. However, the underlying molecular mechanism remains elusive. Toward this aim, we have examined the role of NE in contextual memory formation and in the synaptic delivery of GluR1-containing α-amino-3-hydroxy-5-methyl-4-isoxazoleproprionic acid (AMPA)-type glutamate receptors during long-term potentiation (LTP), a candidate synaptic mechanism for learning. We found that NE, as well as emotional stress, induces phosphorylation of GluR1 at sites critical for its synaptic delivery. Phosphorylation at these sites is necessary and sufficient to lower the threshold for GluR1 synaptic incorporation during LTP. In behavioral experiments, NE can lower the threshold for memory formation in wild-type mice but not in mice carrying mutations in the GluR1 phosphorylation sites. Our results indicate that NE-driven phosphorylation of GluR1 facilitates the synaptic delivery of GluR1-containing AMPARs, lowering the threshold for LTP, thereby providing a molecular mechanism for how emotion enhances learning and memory.

A Wisconsin fall...

My partner and I went driving and walking in the Wisconsin countryside yesterday afternoon - here is an apple orchard, and the ice age trail.

The well-tempered web

I wanted to pass on this link to an engaging article by New Yorker music critic Alex Ross, on his experience with classical music culture on the web. The Internet may be killing the pop CD, but it's helping classical music. The article contains very useful links to many music sites.

Pogue’s Imponderables

Questions that N.Y. Times technical writer David Pogue doesn't have answers to. If you know the answers, fill him in at nytimes.com/pogue.

* Why is Wi-Fi free at cheap hotels, but $14 a night at expensive ones?

* What happens to software programs when their publishers go out of business?

* Would the record companies sell more music online if it weren't copy-protected?

* Do cellphones cause brain cancer?

* What's the real reason you have to turn off your laptop for takeoff?

* Why can't a digital S.L.R. camera record video?

* Wi-Fi on airplanes. What's taking so long?

* Who are the morons who respond to junk-mail offers, thereby keeping spammers in business?

* I'm told that they could make a shirt-pocket digital camera that takes pictures like an S.L.R., but it would cost a lot. So why don't they make one for people who can afford it?

* How come there are still no viruses for Mac OS X? If it has 6 percent of the market, shouldn't it have 6 percent of the viruses?

* Do shareware programmers pay taxes on all those $20 contributions?

* How are we going to preserve all of our digital photos and videos for future generations?

* Why are there no federal rebates or tax credits for solar power?

* Why do you have to take tape camcorders out of your carry-on at airport security, but not the tapeless kind? Couldn't you hide a bomb equally well in either one? (Actually, I have about 500 more logic questions about the rules at airport security, but I have a feeling they'll remain answerless for a very long time.)

* Laptops, cameras and cellphones have improved by a thousand percent in the last ten years. Why not their batteries?

* SmartDisplay, Spot Watch, U.M.P.C., Zune… when will Microsoft realize that it's not a hardware company?

* Why don't public sinks have foot pedals?

* Why don't all hotels have check-in kiosks like airlines do?

* Five billion dollars a year spent on ringtones? What the?

* How come cellphone signal-strength bars are so often wrong?

* Do P.R. people really expect anyone to believe that the standard, stilted, second-paragraph C.E.O. quote was really uttered by a human being?

* Why aren't there recycling bins for bottles and cans where they're most obviously needed, like food courts and cafeterias?

* Why doesn't someone start a cellphone company that bills you only for what you use? That model works O.K. for the electricity, gas and water companies —and people would beat a path to its door.

* Why doesn't everyone have lights that turn off automatically when the room is empty?

* What's the deal with Palm?

* Why are so many people rude on the Internet?

If you know the answers, by all means—fill us in at nytimes.com/pogue.

Genetic dissociation of multiple dopamine roles in learning

Here is the abstract of Frank et al., who show that three genetic variants of the dopamine systems that influence our reactions to positive and negative outcomes (and anticipations) have different effects on human reinforcement learning.

What are the genetic and neural components that support adaptive learning from positive and negative outcomes? Here, we show with genetic analyses that three independent dopaminergic mechanisms contribute to reward and avoidance learning in humans. A polymorphism in the DARPP-32 gene, associated with striatal dopamine function, predicted relatively better probabilistic reward learning. Conversely, the C957T polymorphism of the DRD2 gene, associated with striatal D2 receptor function, predicted the degree to which participants learned to avoid choices that had been probabilistically associated with negative outcomes. The Val/Met polymorphism of the COMT gene, associated with prefrontal cortical dopamine function, predicted participants' ability to rapidly adapt behavior on a trial-to-trial basis. These findings support a neurocomputational dissociation between striatal and prefrontal dopaminergic mechanisms in reinforcement learning. Computational maximum likelihood analyses reveal independent gene effects on three reinforcement learning parameters that can explain the observed dissociations.

A Memory Toolbox

A reader sends this link to 75 tips for going from amnesic to elephantic...

Parallel Distributed Processing and Semantic Cognition

Timothy T. Rogers and James L. McClelland have distilled the essence of the arguments in their book "Semantic Cognition: A Parallel Distributed Processing Approach" for an article (PDF here) to appear in Brain and Behavioral Sciences with peer commentary. Here is their abstract:

In our recent book, we present a parallel distributed processing theory of the acquisition, representation and use of human semantic knowledge. The theory proposes that semantic abilities arise from the flow of activation amongst simple, neuron-like processing units, as governed by the strengths of interconnecting weights; and that acquisition of new semantic information involves the gradual adjustment of weights in the system in response to experience. These simple ideas explain a wide range of empirical phenomena from studies of categorization, lexical acquisition, and disordered semantic cognition. In this précis we focus on phenomena central to the reaction against similarity-based theories that arose in the 1980's and that subsequently motivated the "theory-theory" approach to semantic knowledge. Specifically, we consider i) how concepts differentiate in early development, ii) why some groupings of items seem to form "good" or coherent categories while others do not, iii) why different properties seem central or important to different concepts, iv) why children and adults sometimes attest to beliefs that seem to contradict their direct experience, v) how concepts reorganize between the ages of 4 and 10, and vi) the relationship between causal knowledge and semantic knowledge. The explanations for these phenomena are illustrated with reference to a simple feed-forward connectionist model; and the relationship between this simple model, the broader theory, and more general issues in cognitive science are discussed.

In an Ultimatum Game, Chimps, but not humans, are rational maximizers

Another interesting bit of work from Tomasello's group (PDF here), in which they devised an ingenious apparatus for a mini-ultimatum game, a reduced form of the ultimatum game in which proposers are given a choice between making one of two pre-set offers which the responder can then accept or reject. The proposer had as one option an amount that would typically be rejected by a human responder as unfair, namely 80% for the proposer and 20% for the responder. The most important finding was that responders tended to accept any offer. These results support the hypothesis that other-regarding preferences and aversion to inequitable outcomes, which play key roles in human social organization, distinguish us from our closest living relatives. Here is their abstract, slightly edited:

Traditional models of economic decision-making assume that people are self-interested rational maximizers. Empirical research has demonstrated, however, that people will take into account the interests of others and are sensitive to norms of cooperation and fairness. In one of the most robust tests of this finding, the ultimatum game, individuals will reject a proposed division of a monetary windfall, at a cost to themselves, if they perceive it as unfair. Here we show that in an ultimatum game, humans' closest living relatives, chimpanzees (Pan troglodytes), are rational maximizers and are not sensitive to fairness.

Figure: Illustration of the testing environment. The proposer, who makes the first choice, sits to the responder's left. The apparatus, which has two sliding trays connected by a single rope, is outside of the cages. (A) By first sliding a Plexiglas panel (not shown) to access one rope end and by then pulling it, the proposer draws one of the baited trays halfway toward the two subjects. (B) The responder can then pull the attached rod, now within reach, to bring the proposed food tray to the cage mesh so that (C) both subjects can eat from their respective food dishes (clearly separated by a translucent divider).

Interactions between Declarative and Procedural Memories

Brown and Robertson do a simple experiment showing interaction between two main memory systems usually thought to be independent. Their abstract, slightly edited:

The acquisition of declarative (i.e., facts) and procedural (i.e., skills) memories may be supported by independent systems. This same organization may exist, after memory acquisition, when memories are processed off-line during consolidation. Alternatively, memory consolidation may be supported by interactive systems. This latter interactive organization predicts interference between declarative and procedural memories. Here, we show that procedural consolidation, expressed as an off-line motor skill improvement, can be blocked by declarative learning over wake, but not over a night of sleep. [note: the procedural task was learning a sequence of visually cued button presses at four possible postions.] The extent of the blockade on procedural consolidation was correlated to participants' declarative word recall. Similarly, in another experiment, the reciprocal relationship was found: declarative consolidation was blocked by procedural learning over wake, but not over a night of sleep. [note: The declarative task was learning a list of 16 words each individually presented on a computer screen for two seconds, with the list being presented in the same order five times.] The decrease in declarative recall was correlated to participants' procedural learning. These results challenge the concept of fixed independent memory systems; instead, they suggest a dynamic relationship, modulated by when consolidation takes place, allowing at times for a reciprocal interaction between memory systems.

Facts are no match for gossip...

In yesterday's Science NY Times, John Tierny describes work by Ralf D. Sommerfeld and colleagues on human reciprocity that has a surprising result (PDF here). Here are some clips:

Language, according to the anthropologist Robin Dunbar, evolved because gossip is a more efficient version of the “social grooming” essential for animals to live in groups...Gossip also told people whom to trust, and the prospect of a bad reputation discouraged them from acting selfishly, so large groups could peacefully cooperate. At least, that was the theory: gossip promoted the “indirect reciprocity” that made human society possible.

To test it, researchers at the Max Planck Institute for Evolutionary Biology and the University of Vienna gave 10 Euros apiece to 126 students and had them play a game that put them in a dilemma. On each turn, the players would be paired off, and one of them was offered a chance to give 1.25 Euros to the other. If he agreed, the researchers added a bonus of .75 Euro so that the recipient ended up gaining 2 Euros...If the first player refused to give the money, he’d save 1.25 Euros, but if others found out about his miserliness they might later withhold money from him. As the game progressed, with the players changing partners frequently and alternating between the donor and recipient roles, the players were given information about their partners’ past decisions...Sometimes the donor was shown a record of what the partner had done previously while playing the donor role. The more generous this partner had earlier been toward other players, the more likely the donor was to give him something...Sometimes the donor was shown gossip about the partner from another player. When the partner was paid a compliment like “spendabler spieler!” — generous player! — the donor was more likely to give money. But the donor turned stingy when he saw gossip like “übler geizkragen” — nasty miser.

So far, so good. As predicted, gossip promoted indirect reciprocity. The research, published in the Proceedings of the National Academy of Sciences [Note, I haven't found this article yet in a PNAS search], showed that most people passed on accurate gossip and used it for the common good. They rewarded cooperative behavior even when they themselves weren’t directly affected by the behavior...If a cooperation game like this was played without consequences for the players’ reputations — as has been done in other experiments — most players would be miserly, and cooperation would collapse. In this experiment they were generous most of the time, and on average ended up with twice as much money as they had at the beginning of the game.

But here’s the disconcerting news from the experiment. In a couple of rounds, each donor was given both hard facts and gossip. He was given a record of how his partner had behaved previously as well as some gossip — positive gossip in one round, negative in another...The donor was told that the source of the gossip didn’t have any extra information beyond what the donor could already see for himself. Yet the gossip, whether positive or negative, still had a big influence on the donors’ decisions, and it didn’t even matter if the source of the gossip had a good reputation himself. On average, cooperation increased by about 20 percent if the gossip was good, and fell by 20 percent if the gossip was negative.

Why is this?

It could be,” Sommerfeld suggested, “that we are just more adapted to listen to other information than to observe people, because most of the time we’re not able to observe how other people are behaving. Thus we might believe we have missed something."

Most popular consciousness papers for September 2007

From the ASSC Eprints Archive:

1. Rosenthal, David (2007) Consciousness and its function. In: 11th annual
meeting of the Association for the Scientific Study of Consciousness, 22-25
June 2007, Las Vegas, USA. 1014 downloads from 20 countries.
http://eprints.assc.caltech.edu/293/
2. Koriat, A. (2006) Metacognition and Consciousness. In: Cambridge handbook
of consciousness. Cambridge University Press, New York, USA. 870 downloads
from 18 countries. http://eprints.assc.caltech.edu/175/
3. Windt, Jennifer Michelle and Metzinger, Thomas (2006) The philosophy of
dreaming and self-consciousness: What happens to the experiential subject
during the dream state? In: The new science of dreaming. 834 downloads from
16 countries. http://eprints.assc.caltech.edu/200/
4. Sagiv, Noam and Ward, Jamie (2006) Crossmodal interactions: lessons from
synesthesia. In: Visual Perception, Part 2. Progress in Brain Research,
Volume 155. 814 downloads from 14 countries.
http://eprints.assc.caltech.edu/224/
5. Droege, Paula (2007) ASSC-11 Conference Report. In: 11th annual meeting
of the Association for the Scientific Study of Consciousness, June 22-25,
2007, Las Vegas, Nevada. 798 downloads from 18 countries.
http://eprints.assc.caltech.edu/323/

Motivation alters physical perception

Here is a clip from the Editor's choice section of the Oct. 5 issue of Science:

Students in elementary physics classes are introduced to the concept of frame of reference--the spatial coordinate system used by an observer to describe events--for instance, in the context of the perceived motion of trees by a passenger in a moving automobile. Adding in the dimension of time leads into non-intuitive territory, as in the example of a traveling astronaut who returns to Earth younger than her stay-at-home twin.

Building on previous work that demonstrated that internal physiological states can influence one's perception of physical quantities (such as thirsty people being more likely to characterize objects as transparent; that is, resembling water), Balcetis and Dunning show that internal psychological states are also capable of altering our perception of the external world. They induced states of high or low cognitive dissonance (a mismatch between thought and action) by asking or telling two groups of students to walk across campus wearing various fruit- and vegetable-themed adornments. In order to render a freely chosen yet somewhat embarrassing task less unpleasant to fulfill, the first set of students mentally shortened the distance they had to cover by estimating it to be fully 40% less than the average estimate made by the second group. Intriguingly, the route to ameliorating the state of dissonance appeared to be purely perceptual, as the free-choice students did not shorten their time of exposure by walking faster; in fact, they took about 10% longer.

The abstract of the original article;

Two studies demonstrated that the motivation to resolve cognitive dissonance affects the visual perception of physical environments. In Study 1, subjects crossed a campus quadrangle wearing a costume reminiscent of Carmen Miranda. In Study 2, subjects pushed themselves up a hill while kneeling on a skateboard. Subjects performed either task under a high-choice, low-choice, or control condition. Subjects in the high-choice conditions, presumably to resolve dissonance, perceived the environment to be less aversive than did subjects in the low-choice and control conditions, seeing a shorter distance to travel (Study 1) and a shallower slope to climb (Study 2). These studies suggest that the impact of motivational states extends from social judgment down into perceptual processes.

Baboon Metaphysics

Nicholas Wade comments (PDF here) on the work of Cheney and Seyfarth, whose most recent book has the title of this post. They have carried out an ingenious series of experiments in the wild to probe how the baboon's mind keeps track of transient relationships:

Royal is a cantankerous old male baboon whose troop of some 80 members lives in the Moremi Game Reserve in Botswana. A perplexing event is about to disturb his day....From the bushes to his right, he hears a staccato whoop, the distinctive call that female baboons always make after mating. He recognizes the voice as that of Jackalberry, the current consort of Cassius, a male who outranks Royal in the strict hierarchy of male baboons. No hope of sex today....But then, surprisingly, he hears Cassius’s signature greeting grunt to his left. His puzzlement is plain on the video made of his reaction. You can almost see the wheels turn slowly in his head:...“Jackalberry here, but Cassius over there. Hmm, Jackalberry must be hooking up with some one else. But that means Cassius has left her unguarded. Say what — this is my big chance!”...The video shows him loping off in the direction of Jackalberry’s whoop. But all that he will find is the loudspeaker from which researchers have played Jackalberry’s recorded call.

Although Baboons excel at the skills required for maintaining social networks regulated by matrilineal lines and dominance hierarchies, there is no evidence that they attribute beliefs or ideas to other animals, or that 'they know that they know.' They provide an example of what sort of social and cognitive complexity is possible in the absence of language and a theory of mind.

The prospective brain

I've been meaning to point you to a nice review article by Daniel Schacter and colleagues (PDF here). Here is the abstract and a summary figure.

A rapidly growing number of recent studies show that imagining the future depends on much of the same neural machinery that is needed for remembering the past. These findings have led to the concept of the prospective brain; an idea that a crucial function of the brain is to use stored information to imagine, simulate and predict possible future events. We suggest that processes such as memory can be productively re-conceptualized in light of this idea.

The core brain system that is consistently activated while remembering the past, envisioning the future and during related forms of mental simulation is illustrated schematically. Prominent components of this network include medial prefrontal regions, posterior regions in the medial and lateral parietal cortex (extending into the precuneus and the retrosplenial cortex), the lateral temporal cortex and the medial temporal lobe. Moreover, regions within this core brain system are functionally correlated with each other and, prominently, with the hippocampal formation. We suggest that this core brain system functions adaptively to integrate information about relationships and associations from past experiences, in order to construct mental simulations about possible future events.

New phases of our lives...

NY Times columnist David Brooks muses on how the phases of our life have increased in number (PDF here). Some clips:

There used to be four common life phases: childhood, adolescence, adulthood and old age. Now, there are at least six: childhood, adolescence, odyssey, adulthood, active retirement and old age. Of the new ones, the least understood is odyssey, the decade of wandering that frequently occurs between adolescence and adulthood...People who were born before 1964 tend to define adulthood by certain accomplishments — moving away from home, becoming financially independent, getting married and starting a family....In 1960, roughly 70 percent of 30-year-olds had achieved these things. By 2000, fewer than 40 percent of 30-year-olds had done the same.

...you can see the spirit of fluidity that now characterizes this stage....everything seems to give way to a less permanent version of itself...Dating gives way to Facebook and hooking up. Marriage gives way to cohabitation....Graduating seniors don’t find corporations offering them jobs that will guide them all the way to retirement. Instead they find a vast menu of information economy options...what we’re seeing is the creation of a new life phase, just as adolescence came into being a century ago...European nations are traveling this route ahead of us... Europeans delay marriage even longer than we do and spend even more years shifting between the job market and higher education...Someday people will look back and wonder at the vast social changes wrought by the emerging social group that saw their situations first captured by “Friends” and later by “Knocked Up.”

How to get health facts completely wrong....

John Tierney reviews Gary Taubes account (in "“Good Calories, Bad Calories”; Knopf, 2007) of how the health science establishment got it completely wrong, from the basic science on through official government pronouncements on dietary fat and hearth disease (PDF here). Some clips:

The notion that fatty foods shorten your life began as a hypothesis based on dubious assumptions and data; when scientists tried to confirm it they failed repeatedly.

Even so, in 1988, the surgeon general, C. Everett Koop, proclaimed ice cream to a be public-health menace right up there with cigarettes.This happened because of what social scientists call a cascade:

We like to think that people improve their judgment by putting their minds together, and sometimes they do. The studio audience at “Who Wants to Be a Millionaire” usually votes for the right answer. But suppose, instead of the audience members voting silently in unison, they voted out loud one after another. And suppose the first person gets it wrong...If the second person isn’t sure of the answer, he’s liable to go along with the first person’s guess. By then, even if the third person suspects another answer is right, she’s more liable to go along just because she assumes the first two together know more than she does. Thus begins an “informational cascade” as one person after another assumes that the rest can’t all be wrong...Because of this effect, groups are surprisingly prone to reach mistaken conclusions even when most of the people started out knowing better... If, say, 60 percent of a group’s members have been given information pointing them to the right answer (while the rest have information pointing to the wrong answer), there is still about a one-in-three chance that the group will cascade to a mistaken consensus.

Your Money and Your Brain

I thought I would pass on this brief commentary by Joe Nocera in the NY Times (PDF here), which emphasizes comments by Jason Zweig on how our emotional brains are not very well suited for making rational financial decisions.

A neural marker of consciousness

Here is the author's summary of an interesting article by Del Cul et al. in PlosBiology.

Understanding the neural mechanisms that distinguish between conscious and nonconscious processes is a crucial issue in cognitive neuroscience. In this study, we focused on the transition that causes a visual stimulus to cross the threshold to consciousness, i.e., visibility. We used a backward masking paradigm in which the visibility of a briefly presented stimulus (the “target”) is reduced by a second stimulus (the “mask”) presented shortly after this first stimulus. (Human participants report the visibility of the target.) When the delay between target and mask stimuli exceeds a threshold value, the masked stimulus becomes visible. Below this threshold, it remains nonvisible. During the task, we recorded electric brain activity from the scalp and reconstructed the cortical sources corresponding to this activity. Conscious perception of masked stimuli corresponded to activity in a broadly distributed fronto-parieto-temporal network, occurring from about 300 ms after stimulus presentation. We conclude that this late stage, which could be clearly separated from earlier neural events associated with subliminal processing and mask-target interactions, can be regarded as a marker of consciousness.


Figure: Top, depth of cortical processing: subliminal stimuli (left panel) should evoke a strong activation in extrastriate visual cortex, but their intensity should quickly decrease in higher visual areas; only conscious stimuli (right panel) should trigger a late surge of activation in a global prefronto-parietal network. Bottom, schematic time course of activation as a function of masking strength. Masking is expected to have little effect on early visual activation but to modulate the strength of activation in higher visual areas. Furthermore, there should be a nonlinear effect of masking strength in prefrontal cortex, with a similar late top-down activation peak occurring simultaneously in visual areas

Where the brain decides how much we are willing to pay.

Plassmann et al. show some brain correlates of our willingness to pay:

An essential component of every economic transaction is a willingness-to-pay (WTP) computation in which buyers calculate the maximum amount of financial resources that they are willing to give up in exchange for the object being sold. Despite its pervasiveness, little is known about how the brain makes this computation. We investigated the neural basis of the WTP computation by scanning hungry subjects' brains using functional magnetic resonance imaging while they placed real bids for the right to eat different foods. We found that activity in the medial orbitofrontal cortex and in the dorsolateral prefrontal cortex encodes subjects' WTP for the items. Our results support the hypothesis that the medial orbitofrontal cortex encodes the value of goals in decision making.
Neural correlates of WTP. A, B, Activity in the medial OFC and the DLPFC was positively correlated with WTP at the time of evaluation in the free trials more than in the forced trials.