Reduced Neural Selectivity Increases fMRI Adaptation with Age during Face Discrimination

Key project finally published! This took quite a while, but it was worth it.

[Link to article if you have journal access]
[Link to Pubmed abstract access]

By Joshua O., Goh , Atsunobu, Suzuki , Denise C., Park
Beckman Institute, University of Illinois, Urbana-Champaign, IL, USA; Center for Vital Longevity, University of Texas, Dallas, TX, USA.

Ventral-visual activity in older adults has been characterized by dedifferentiation, or reduced distinctiveness, of responses to different categories of visual stimuli such as faces and houses, that typically elicit highly specialized responses in the fusiform and parahippocampal brain regions respectively in young adults (Park et al., 2004). In the present study, we demonstrate that age-related neural dedifferentiation applies to within-category stimuli (different types of faces) as well, such that older adults process less distinctive representations for individual faces than young adults. We performed a functional magnetic resonance imaging adaptation experiment while young and older participants made same-different judgments to serially presented face-pairs that were Identical, Moderate in similarity through morphing, or Different. As expected, older adults showed adaptation in the fusiform face area (FFA), during the Identical as well as the Moderate conditions relative to the Different condition. Young adults showed adaptation during the Identical condition, but minimal adaptation to the Moderate condition. These results indicate that older adults' FFA treated the morphed faces as Identical faces, reflecting decreased fidelity of neural representation of faces with age.

NeuroImage, In Press, Accepted Manuscript, Available online 6 February 2010

Separation vs. Association

A key function of the brain is to first, process the fact that we are encountering different types of stimuli at every moment, and second, process the simultaneous fact that while there are these different types of stimuli, there are also many consistencies that reflect modifications of the same stimulus at a higher level of abstraction.

One way to evaluate what a cortical region may be doing with respect to this separation/association dichotomy may be to determine the number of neurons at the first level relative to the second level.

If the ratio of neurons at the first relative to second level is large, then the function of the second level is probably to associate. This is a many-to-few limitation. So various permutations and combinations at the first level are funneled into the reduced dimensionality of the second level. Therefore, some combinations are subsumed.

If the ratio of neurons from first to second levels is small, then there is the potential for expansion. The problem becomes a few-to-many scenario. The same combination at the first level may elicit several possible outcomes at the second level. There is information expansion.

...

Agnostic Brain, Biased Mind - what does the FFA do?

Many neuroimaging studies have repeatedly found an area in the human brain that seems to be involved in processing visual faces. This area located in the fusiform gyri in humans, has been affectionately named the fusiform face area or FFA. The FFA is most active when we are looking at pictures of faces, and almost non-responsive to other types of visual items such as objects, houses, scenes, random textures, or a blank screen. Prosopagnosics, who are not able to recognize faces, but are still able to detect the presence of a face and also show no difficulty in processing other types of visual stimuli, have been shown to involve less FFA activity. Even more compelling, patients with lateral occipital lobes lesioned lose some form of object-processing, but show intact face processing. And yet other patients with lesions that have affected the FFA, have problems with face processing (acquired prosopagnosia) but intact processing for other stimuli. The evidence strongly suggests that there is something special about faces, and something about the FFA that deals with this specialization.

The debate regarding the FFA pertain to whether it is the only region or even a critical region that does face processing. Some labs have shown that face processing information can be found in other regions of the brain that are not the FFA. Yet some labs have shown that the FFA is recruited to process fine levels of category distinctions. For example, bird and car experts have been shown to engage some level of FFA activity when processing these stimuli compared to novices. These findings suggest that the FFA is not processing faces per se, but visual representations that have come to require high-levels of fine discrimination through experience, of which faces are the best example of this currently.

I suggest that a more flexible definition is called for when thinking about the FFA and its role in processing visual information. Certainly, it does seem that faces occupy a special place in human experiences. On the other hand, it is difficult to explain why there would be a brain region that codes for faces and faces along based simply based on genetic or biologically determined causes.

In terms of a neural network, if indeed the brain consists of many different sub-types of neural networks that conglomerate to form one large complex network, the FFA is a sub-network specialized to perform a specific operation that is maximized and specialized (trained) for a specific information domain - faces. This or these specific operation(s) could involve identification, discrimination, recognition, or all of these, or even a yet unknown operation. Certainly neural network non-linearities can surprise us! Moreover, these operations have been tuned for a specialized class of stimuli that consists of eyes, nose, mouths, and other visual characteristics of faces when occurring together as a whole (whether from external input, or through internal imagination or retrieval).

What this means is that if you were able to "remove" the FFA, and plug it into a computer so that you can feed this FFA network with inputs and measure its outputs, you could theoretically feed it anything, but the information would be most meaningful or organized when the inputs correspond to information about a face. Of course, this would require us to know what is the language of the input to perform such an experiment.

Other types of inputs may elicit some level of meaningful output of the FFA. Neural network do that. Yet other types may elicit nothing at all. This does not necessarily mean that the FFA outputs from such inputs is useless, nor does necessarily mean that it is used! It is just output. What higher-level brain mechanisms do with the output depends on the task, and how the brain is wired to treat outputs from its sub-networks. It may be ignored, or it may actually incorporate relevant information. That is, the FFA is agnostic to the incoming information. It does not care. It will process it anyway. But other regions decide whether what is it saying needs to be incorporated or not, or if it should be further modified even.

Such a view would reconcile why the FFA is special for faces, yet seems to be carrying some information about other stimuli. It would also be consistent with the idea that information about faces is certainly also available to a certain extent in non-FFA regions, the same principles being applied to these other sub-networks. It would also be consistent with how self-organizing behavior in neural network (see von der Malsburg article [link]) can lead to a consistent topology across every person that processes a particular stimulus in a particular way in a particular spatial location.

This is probably not a new idea, but needs to be clarified in the literature I think.

VSS Conference Day 4: My Poster

This poster was presented at VSS Conference 2009 [link to VSS website], morning session [download poster pdf]. This study investigated the effect of task instructions on repetition suppression in the brain. Repetition suppression refers to the phenomenon that the brain response to repeated stimuli is usually reduced or attenuated. It is thought that such reduction in brain response reflects less neuronal recruitment, and hence, a more "efficient" way of processing the same information.
In this study, however, I postulated that under certain circumstances, the brain requires more neuronal recruitment in order to effectively process information for task demands. That is, repetition suppression becomes inefficient because it reduces the degrees of freedom that the brain can use to manipulate existing representations.

The study evaluated brain response in the fusiform region to face-pairs morphed at different levels of similarity. The idea is that the more similar face-pairs are, the more repetition suppression should be observed in the fusiform face area. Participants viewed the face-pairs under two different task instructions. The first task made face-pair similarity irrelevant. In this task, repetition suppression was observed to repeated faces. In the second task, face-pairs were made critical as participants had to make same-different judgments about the pairs. In this task, repetition suppression was eliminated.

The idea here is that in the same-different judgment task, the brain has to represent faces as distinctinctively as possible so that subtle morph differences can be detected. Thus, repetition suppression is prevented, possibly from executive function areas that process task instruction and exert a top-down modulatory control in the fusiform area.

The study also shows that there are individual differences in participants ability to exert this top-down modulation to regulate repetition suppression in the fusiform regions. This study was also performed in older adults, which will be reported in a subsequent research article. Briefly though, it is thought that older adults show declines in behavioral performance because of less distinctiveness in cognitive representations. This design is thus useful as a means to measure and related distinctinveness of representations in the brain and how that affects behavior.

A structural model of aging, brain and behavior

Possible working structural model that can be tested with measures of stimulus, behavior, neuro-functional, neuro-anatomical variables. The dynamic influences of age and "culture" can also be tested. Culture here refers to long-term experiences of any kind. More complex models can be postulated from this current framework by adding more factors, or measures, and by also constraining the specific weights and covariances. In the broadest sense, the weights and covariances are modeled linearly. However, certainly, non-linear functions can be imposed. The result of such impositions would be a neural network with non-linear activation functions.

Aging increases fMR-Adaptation to repeated faces and limits discrimination ability

Age-related ventral-visual activity is characterized by reduced selectivity between categories of visual stimuli such as faces and houses that typically elicit highly specialized responses in the fusiform and parahippocampal brain regions respectively of young adults (Park et al., 2004). This study demonstrates that older adults’ less selective neural response (dedifferentiation) to faces is due to a coarser neural representation than that of young adults for individual faces. In this event-related fMRI adaptation study, 20 young and 20 older participants made same-different judgments to serially presented face-pairs that were either identical (repetition of the same face), moderately different (second face was morphed with 40% of prior face), or completely different (faces from two individuals). In the fusiform regions, older adults showed greatest adaptation during the exact-repetition condition and intermediate adaptation during the moderately-different condition relative to the completely-different condition. Young adults showed a similar pattern of adaptation, but with reduced adaptation magnitudes. This suggests that older adults’ fusiform area was not able to represent facial differences at the same level of sensitivity as young adults. Individual subjects’ adaptation magnitudes positively correlated with behavioral face discrimination thresholds for morphed faces (r = .38). Greater age-related adaptation during the exact-repetition condition suggests older adults’ fusiform face region is particularly sensitive to similarity, whereas young adults have a neural response that is more sensitive to facial differences. The data provide strong evidence for an age-related decrease in representational contrast in the fusiform area that is linked to behavioral differences during perceptual discrimination. [pdf]

I Think That I Shall Never See, A Brain as Pretty as a Tree

What if, sunlight to a tree, is as information to the brain? A tree needs sunlight to survive, to produce food. In response to this basic need, the tree spends a lot of its effort to maximize its ability to obtain sunlight. It does this by forming more leaves, and by spreading those leaves out at widely as possible to cover as much area as possible. Pushing this idea even further, to the extent that the tree covers a portion of area, that is the amount of sunlight it can absorb. Sunlight falling on other areas will be lost to the tree (albeit there might be secondary or tertiary transfer of energy via light reflection, diffusion, and other means). One important parameter that would determine the success of sunlight absorption for a tree would then be leaf surface area. Specifically, greater surface area would increase sunlight absorption rate.

Now, we project this idea onto the brain, of course being well aware that the brain is much different from a tree, although, probably not very very different. The brain is in the business of representing information. Its very function is the processing and retaining of all the information fed into it from the moment of its development. It would be interesting to pursue when this onsets, but that is a digression for later. Nevertheless, the brain develops in tandem with its experience with information. Some of that information is hardwired, or genetic. Some of that information is nurtured, or environmentally experienced. The role of each neuron then, in cooperation with all the other neurons in the brain, is to keep EVERY SINGLE EXPERIENCE, whether internal or external.

Why does the brain want to do that? Well, that's the same as answering why does a tree want so much sunlight for? We can only provide partial understanding here, because this borders on the domain of philosophical and religious pursuits. Biologically, a tree seeks sunlight as part of its nutritional source for the purpose of ultimately creating more trees. That is about as far as we can describe based on observation. This is, in a way, a tautology. Because what we impose as the purpose of the tree, is in fact, what we see the tree already doing. Therefore, such an answer may not satisfy some, but it is a partial answer at the least. Turning back to the brain, again, only a partial answer is given. The brain seeks to contain as much information as possible, because that's what is already observed that it is doing, and perhaps, this information helps the organism to survive, and to produce more organisms of this kind.

More importantly here, we shall consider how does the brain perform this function of representing as much information as possible. The tree does it by increasing surface area exposed to the sunlight. The brain's equivalent would be to increase the number of neurons it has, the connections between these neurons, the variability in the way these neurons can activate. Some smart person might be able to come up with an equation that tells us how much information a given brain with a given number of neurons and connections, and variability in activity, can hold. This could somehow be mathematically related to the concept of surface area...

However, there is a problem in terms of space. While the brain is fantastic and has way superior computational capacity, it is still finite. That is, there may come a time when a given person's brain can no longer process anymore new information. Maybe it has come already, just that we don't know it or that its not as big of a problem as we might think, given we have external aids for our memory now, through things such as computers, books, paper, language, and symbols. This finite capacity is indeed a problem, but our brains have a rather interesting way of solving it, at least to a great extent. Lets turn back to the tree for a moment, because its a greener thought. Lets say that to get more sunlight, the tree has two ways of doing it given a fixed amount of material. It can send more branches out with many leaves, or it can make fewer but bigger leaves. If it sends out more branches, it would have to content with using some of that material to make tree-parts that don't absorb sunlight (branches). If it makes bigger leaves, it may have to content with those leaves blocking each other out, since they will be close together as there are no branches to help spread them out. In the same way, the brain might have two ways of holding information within a limited amount of material. It can create more and more connections with more neurons, or it can use the existing neurons and connections in different ways. Here is where the tree analogy might break down. Unlike information, sunlight to a tree is a one-dimensional problem in the sense that it only needs to worry about expose area. Information, however, is obviously multi-dimensional, with auditory, visual, tactile, odor, taste modalities in the sensory domains, and countless of types of dimensions when you think about concepts and their associations, temporal information etc. Another dissimilarity between a tree and the brain is that most trees only make one kind of leaf, or grow with a certain fixed physical structure. The brain is able to flexibly use neuronal connections to group neurons in very dynamic ways. So, while a tree either has small or big leaves, the brain may use both small groups of neurons encoding some type of information, as well as bigger groups encoding other types of information.

The maximum surface area of the brain (meaning the physical ability of the brain to differentiate between its billions of states of activity using its neuronal connections), limits the total amount of different information the that brain can keep. This will be developed in later blogs. Here are some teasers. One brilliant way of reducing the space needed would be to encode information in terms of similarities and differences. And also, unlike a tree, the brain in the organism makes decisions about what the organism should do, affecting the environment and modifying subsequent experiences, as opposed to being completely at the mercy of the experiences.

Next time...."Similarities and Differences", and "The Brain, The Tree, Intentions, and Decisions".

HBM: Ideas: Visual Discrimination Project: General Questions

1. Is there poorer behavioral perceptual discrimination with age?
2. If so, what are the neural correlates? Is it a perceptual representation problem (ventral visual dedifferentiation)? Or is it a selection/decision-making/control process problem (noise, non-selectivity in frontal cortex)?
3. Is this the same across all types of stimuli (eg faces, patterns, random shapes)?
4. Is there a constant in terms of brain activation pattern across all individuals which is necessary for discrimination?
5. Which parts of the brain are predictive of whether the individual is able to discriminate visual stimuli (classifier algorithms on whole brain)?
6. What leads to individual differences in performance? And if all individuals are equated at some level of performance, do the individual differences disappear?

Quality of representation: PhD Proposal

Here is an idea that I am toying with for my PhD research.

In aging, the idea is that neural representations change in several ways. Right now, we know there are changes, but we don't know what they are exactly, or why they happen the way they do. One obvious objective change we are pretty certain of is that, with age, processing speed slows down.

Now, is this a result of nerve conduction changes? Or is this more to do with changes in the processes themselves. That is, neuronal connectivity is changing, so the process computation changes as well, and changes in a way that results in slowing of the process. In addition, connectivity may also be changing in terms of a reduction in neural plasticity, the ability of neurons to form (or prune) connections based on experience.

One test I propose is related to ascertaining if the observations about neural changes with age relate to changes in processes vis a vis changes in connectivity. If neural representations are poorer with age, then it also means that they are less able to dissociate between similar representations. That is, aging reduces distinctiveness between neural representations. We should be able to measure this using the adaptation paradigm. In theory, if two representations are similar, the adaptation should be greater. To the extent that two representations are distinct, there will be less adaptation. Thus, with younger adults, there should be less adaptation to similar but different stimuli, or, there should be adaptation only when stimuli are very similar. In older adults, adaptation should happen at a lower threshold of similarity, for stimuli that are in fact quite distinct compared to that for the young.

In addition, since the neural representational quality in the lower perceptual areas feeds the cognitive processes that operate on them downstream (e.g. perceptual matching, target identification, memory, attention, decision-making etc) then it stands that if the neural representation is poorer, the cortex involved in working on these representations will either work harder to produce the same result, or be incapable of producing the same result if the representations were clearer. So, this could be measured as a correlation of frontal cortex activity with the degree of adaptation in the posterior, more perceptual areas.