Uncovering the functional architecture of a historic brain area

In 1840 a patient named Leborgne was admitted to a hospital near Paris: he was only able repeat the word “Tan.” This loss of speech drew the attention of Paul Broca who, after Leborgne’s death, identified lesions in his frontal lobe in the left hemisphere. These results echoed earlier findings from French neurologist Marc Dax. Now known as “Broca’s area,” the roles of this brain region have been extended to mental functions far beyond speech articulation. So much so, that the underlying functional organization of Broca’s area has become a source of discussion and some confusion.

McGovern Investigator Ev Fedorenko is now calling, in a paper at Trends in Cognitive Sciences, for recognition that Broca’s area consists of functionally distinct, specialized regions, with one sub-region very much dedicated to language processing.

“Broca’s area is one of the first regions you learn about in introductory psychology and neuroscience classes, and arguably laid the foundation for human cognitive neuroscience,” explains Ev Fedorenko, who is also an assistant professor in MIT’s Department of Brain and Cognitive Sciences. “This patch of cortex and its connections with other brain areas and networks provides a microcosm for probing some core questions about the human brain.”

Broca’s area, shown in red. Image: Wikimedia

Language is a uniquely human capability, and thus the discovery of Broca’s area immediately captured the attention of researchers.

“Because language is universal across cultures, but unique to the human species, studying Broca’s area and constraining theories of language accordingly promises to provide a window into one of the central abilities that make humans so special,” explains co-author Idan Blank, a former postdoc at the McGovern Institute who is now an assistant professor of psychology at UCLA.

Function over form

Broca’s area is found in the posterior portion of the left inferior frontal gyrus (LIFG). Arguments and theories abound as to its function. Some consider the region as dedicated to language or syntactic processing, others argue that it processes multiple types of inputs, and still others argue it is working at a high level, implementing working memory and cognitive control. Is Broca’s area a highly specialized circuit, dedicated to the human-specific capacity for language and largely independent from the rest high-level cognition, or is it a CPU-like region, overseeing diverse aspects of the mind and orchestrating their operations?

“Patient investigations and neuroimaging studies have now associated Broca’s region with many processes,” explains Blank. “On the one hand, its language-related functions have expanded far beyond articulation, on the other, non-linguistic functions within Broca’s area—fluid intelligence and problem solving, working memory, goal-directed behavior, inhibition, etc.—are fundamental to ‘all of cognition.’”

While brain anatomy is a common path to defining subregions in Broca’s area, Fedorenko and Blank argue that instead this approach can muddy the water. In fact, the anatomy of the brain, in terms of cortical folds and visible landmarks that originally stuck out to anatomists, vary from individual to individual in terms of their alignment with the underlying functions of brain regions. While these variations might seem small, they potentially have a huge impact on conclusions about functional regions based on traditional analysis methods. This means that the same bit of anatomy (like, say, the posterior portion of a gyrus) could be doing different things in different brains.

“In both investigations of patients with brain damage and much of brain imaging work, a lot of confusion has stemmed from the use of macroanatomical areas (like the inferior frontal gyrus (IFG)) as ‘units of analysis’,” explains Fedorenko. “When some researchers found IFG activation for a syntactic manipulation, and others for a working memory manipulation, the field jumped to the conclusion that syntactic processing relies on working memory. But these effects might actually be arising in totally distinct parts of the IFG.”

The only way to circumvent this problem is to turn to functional data and aggregate information from functionally defined areas across individuals. Using this approach, across four lines of evidence from the last decade, Fedorenko and Blank came to a clear conclusion: Broca’s area is not a monolithic region with a single function, but contains distinct areas, one dedicated to language processing, and another that supports domain-general functions like working memory.

“We just have to stop referring to macroanatomical brain regions (like gyri and sulci, or their parts) when talking about the functional architecture of the brain,” explains Fedorenko. “I am delighted to see that more and more labs across the world are recognizing the inter-individual variability that characterizes the human brain– this shift is putting us on the right path to making fundamental discoveries about how our brain works.”

Indeed, accounting for distinct functional regions, within Broca’s area and elsewhere, seems essential going forward if we are to truly understand the complexity of the human brain.

Word Play

Ev Fedorenko uses the widely translated book “Alice in Wonderland” to test brain responses to different languages.

Language is a uniquely human ability that allows us to build vibrant pictures of non-existent places (think Wonderland or Westeros). How does the brain build mental worlds from words? Can machines do the same? Can we recover this ability after brain injury? These questions require an understanding of how the brain processes language, a fascination for Ev Fedorenko.

“I’ve always been interested in language. Early on, I wanted to found a company that teaches kids languages that share structure — Spanish, French, Italian — in one go,” says Fedorenko, an associate investigator at the McGovern Institute and an assistant professor in brain and cognitive sciences at MIT.

Her road to understanding how thoughts, ideas, emotions, and meaning can be delivered through sound and words became clear when she realized that language was accessible through cognitive neuroscience.

Early on, Fedorenko made a seminal finding that undermined dominant theories of the time. Scientists believed a single network was extracting meaning from all we experience: language, music, math, etc. Evolving separate networks for these functions seemed unlikely, as these capabilities arose recently in human evolution.

Language Regions
Ev Fedorenko has found that language regions of the brain (shown in teal) are sensitive to both word meaning and sentence structure. Image: Ev Fedorenko

But when Fedorenko examined brain activity in subjects while they read or heard sentences in the MRI, she found a network of brain regions that is indeed specialized for language.

“A lot of brain areas, like motor and social systems, were already in place when language emerged during human evolution,” explains Fedorenko. “In some sense, the brain seemed fully occupied. But rather than co-opt these existing systems, the evolution of language in humans involved language carving out specific brain regions.”

Different aspects of language recruit brain regions across the left hemisphere, including Broca’s area and portions of the temporal lobe. Many believe that certain regions are involved in processing word meaning while others unpack the rules of language. Fedorenko and colleagues have however shown that the entire language network is selectively engaged in linguistic tasks, processing both the rules (syntax) and meaning (semantics) of language in the same brain areas.

Semantic Argument

Fedorenko’s lab even challenges the prevailing view that syntax is core to language processing. By gradually degrading sentence structure through local word swaps (see figure), they found that language regions still respond strongly to these degraded sentences, deciphering meaning from them, even as syntax, or combinatorial rules, disappear.

The Fedorenko lab has shown that the brain finds meaning in a sentence, even when “local” words are swapped (2, 3). But when clusters of neighboring words are scrambled (4), the brain struggles to find its meaning.

“A lot of focus in language research has been on structure-building, or building a type of hierarchical graph of the words in a sentence. But actually the language system seems optimized and driven to find rich, representational meaning in a string of words processed together,” explains Fedorenko.

Computing Language

When asked about emerging areas of research, Fedorenko points to the data structures and algorithms underlying linguistic processing. Modern computational models can perform sophisticated tasks, including translation, ever more effectively. Consider Google translate. A decade ago, the system translated one word at a time with laughable results. Now, instead of treating words as providing context for each other, the latest artificial translation systems are performing more accurately. Understanding how they resolve meaning could be very revealing.

“Maybe we can link these models to human neural data to both get insights about linguistic computations in the human brain, and maybe help improve artificial systems by making them more human-like,” says Fedorenko.

She is also trying to understand how the system breaks down, how it over-performs, and even more philosophical questions. Can a person who loses language abilities (with aphasia, for example) recover — a very relevant question given the language-processing network occupies such specific brain regions. How are some unique people able to understand 10, 15 or even more languages? Do we need words to have thoughts?

Using a battery of approaches, Fedorenko seems poised to answer some of these questions.

Speaking many languages

Ev Fedorenko studies the cognitive processes and brain regions underlying language, a signature cognitive skill that is uniquely and universally human. She investigates both people with linguistic impairments, and those that have exceptional language skills: hyperpolyglots, or people that are fluent in over a dozen languages. Indeed, she was recently interviewed for a BBC documentary about superlinguists as well as the New Yorker, for an article covering people with exceptional language skills.

When Fedorenko, an associate investigator at the McGovern Institute and assistant professor in the Department of Brain and Cognitive Sciences at MIT, came to the field, neuroscientists were still debating whether high-level cognitive skills such as language, are processed by multi-functional or dedicated brain regions. Using fMRI, Fedorenko and colleagues compared engagement of brain regions when individuals were engaged in linguistic vs. other high level cognitive tasks, such as arithmetic or music. Their data revealed a clear distinction between language and other cognitive processes, showing that our brains have dedicated language regions.

Here is my basic question. How do I get a thought from my mind into yours?

In the time since this key study, Fedorenko has continued to unpack language in the brain. How does the brain process the overarching rules and structure of language (syntax), as opposed to meanings of words? How do we construct complex meanings? What might underlie communicative difficulties in individuals diagnosed with autism? How does the aphasic brain recover language? Intriguingly, in contrast to individuals with linguistic difficulties, there are also individuals that stand out as being able to master many languages, so-called hyperpolyglots.

In 2013, she came across a young adult that had mastered over 30 languages, a prodigy in languages. To facilitate her analysis of processing of different languages Fedorenko has collected dozens of translations of Alice in Wonderland, for her ‘Alice in the language localizer Wonderland‘ project. She has already found that hyperpolyglots tend to show less activity in linguistic processing regions when reading in, or listening to, their native language, compared to carefully matched controls, perhaps indexing more efficient processing mechanisms. Fedorenko continues to study hyperpolyglots, along with other exciting new avenues of research. Stay tuned for upcoming advances in our understanding of the brain and language.

Evelina Fedorenko

Exploring Language

Evelina (Ev) Fedorenko aims to understand how the language system works in the brain. Her lab is unpacking the internal architecture of the brain’s language system and exploring the relationship between language and various cognitive, perceptual, and motor systems. To do this, her lab employs a range of approaches – from brain imaging to computational modeling – and works with a diverse populations, including polyglots and individuals with atypical brains. Language is a quintessential human ability, but the function that language serves has been debated for centuries. Fedorenko argues that language serves is primarily as a tool for communication, contrary to a prominent view that language is essential for thinking.

Ultimately, this cutting-edge work is uncovering the computations and representations that fuel language processing in the brain.

Can we think without language?

As part of our Ask the Brain series, Anna Ivanova, a graduate student who studies how the brain processes language in the labs of Nancy Kanwisher and Evelina Fedorenko, answers the question, “Can we think without language?”

Anna Ivanova headshot
Graduate student Anna Ivanova studies language processing in the brain.

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Imagine a woman – let’s call her Sue. One day Sue gets a stroke that destroys large areas of brain tissue within her left hemisphere. As a result, she develops a condition known as global aphasia, meaning she can no longer produce or understand phrases and sentences. The question is: to what extent are Sue’s thinking abilities preserved?

Many writers and philosophers have drawn a strong connection between language and thought. Oscar Wilde called language “the parent, and not the child, of thought.” Ludwig Wittgenstein claimed that “the limits of my language mean the limits of my world.” And Bertrand Russell stated that the role of language is “to make possible thoughts which could not exist without it.” Given this view, Sue should have irreparable damage to her cognitive abilities when she loses access to language. Do neuroscientists agree? Not quite.

Neuroimaging evidence has revealed a specialized set of regions within the human brain that respond strongly and selectively to language.

This language system seems to be distinct from regions that are linked to our ability to plan, remember, reminisce on past and future, reason in social situations, experience empathy, make moral decisions, and construct one’s self-image. Thus, vast portions of our everyday cognitive experiences appear to be unrelated to language per se.

But what about Sue? Can she really think the way we do?

While we cannot directly measure what it’s like to think like a neurotypical adult, we can probe Sue’s cognitive abilities by asking her to perform a variety of different tasks. Turns out, patients with global aphasia can solve arithmetic problems, reason about intentions of others, and engage in complex causal reasoning tasks. They can tell whether a drawing depicts a real-life event and laugh when in doesn’t. Some of them play chess in their spare time. Some even engage in creative tasks – a composer Vissarion Shebalin continued to write music even after a stroke that left him severely aphasic.

Some readers might find these results surprising, given that their own thoughts seem to be tied to language so closely. If you find yourself in that category, I have a surprise for you – research has established that not everybody has inner speech experiences. A bilingual friend of mine sometimes gets asked if she thinks in English or Polish, but she doesn’t quite get the question (“how can you think in a language?”). Another friend of mine claims that he “thinks in landscapes,” a sentiment that conveys the pictorial nature of some people’s thoughts. Therefore, even inner speech does not appear to be necessary for thought.

Have we solved the mystery then? Can we claim that language and thought are completely independent and Bertrand Russell was wrong? Only to some extent. We have shown that damage to the language system within an adult human brain leaves most other cognitive functions intact. However, when it comes to the language-thought link across the entire lifespan, the picture is far less clear. While available evidence is scarce, it does indicate that some of the cognitive functions discussed above are, at least to some extent, acquired through language.

Perhaps the clearest case is numbers. There are certain tribes around the world whose languages do not have number words – some might only have words for one through five (Munduruku), and some won’t even have those (Pirahã). Speakers of Pirahã have been shown to make mistakes on one-to-one matching tasks (“get as many sticks as there are balls”), suggesting that language plays an important role in bootstrapping exact number manipulations.

Another way to examine the influence of language on cognition over time is by studying cases when language access is delayed. Deaf children born into hearing families often do not get exposure to sign languages for the first few months or even years of life; such language deprivation has been shown to impair their ability to engage in social interactions and reason about the intentions of others. Thus, while the language system may not be directly involved in the process of thinking, it is crucial for acquiring enough information to properly set up various cognitive domains.

Even after her stroke, our patient Sue will have access to a wide range of cognitive abilities. She will be able to think by drawing on neural systems underlying many non-linguistic skills, such as numerical cognition, planning, and social reasoning. It is worth bearing in mind, however, that at least some of those systems might have relied on language back when Sue was a child. While the static view of the human mind suggests that language and thought are largely disconnected, the dynamic view hints at a rich nature of language-thought interactions across development.

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Brain’s language center has multiple roles

A century and a half ago, French physician Pierre Paul Broca found that patients with damage to part of the brain’s frontal lobe were unable to speak more than a few words. Later dubbed Broca’s area, this region is believed to be critical for speech production and some aspects of language comprehension.

However, in recent years neuroscientists have observed activity in Broca’s area when people perform cognitive tasks that have nothing to do with language, such as solving math problems or holding information in working memory. Those findings have stimulated debate over whether Broca’s area is specific to language or plays a more general role in cognition.

A new study from MIT may help resolve this longstanding question. The researchers, led by Nancy Kanwisher, the Walter A. Rosenblith Professor of Cognitive Neuroscience, found that Broca’s area actually consists of two distinct subunits. One of these focuses selectively on language processing, while the other is part of a brainwide network that appears to act as a central processing unit for general cognitive functions.

“I think we’ve shown pretty convincingly that there are two distinct bits that we should not be treating as a single region, and perhaps we shouldn’t even be talking about “Broca’s area” because it’s not a functional unit,” says Evelina Fedorenko, a research scientist in Kanwisher’s lab and lead author of the new study, which recently appeared in the journal Current Biology.

Kanwisher and Fedorenko are members of MIT’s Department of Brain and Cognitive Sciences and the McGovern Institute for Brain Research. John Duncan, a professor of neuroscience at the Cognition and Brain Sciences Unit of the Medical Research Council in the United Kingdom, is also an author of the paper.

A general role

Broca’s area is located in the left inferior frontal cortex, above and behind the left eye. For this study, the researchers set out to pinpoint the functions of distinct sections of Broca’s area by scanning subjects with functional magnetic resonance imaging (fMRI) as they performed a variety of cognitive tasks.

To locate language-selective areas, the researchers asked subjects to read either meaningful sentences or sequences of nonwords. A subset of Broca’s area lit up much more when the subjects processed meaningful sentences than when they had to interpret nonwords.

The researchers then measured brain activity as the subjects performed easy and difficult versions of general cognitive tasks, such as doing a math problem or holding a set of locations in memory. Parts of Broca’s area lit up during the more demanding versions of those tasks. Critically, however, these regions were spatially distinct from the regions involved in the language task.

These data allowed the researchers to map, for each subject, two distinct regions of Broca’s area — one selectively involved in language, the other involved in responding to many demanding cognitive tasks. The general region surrounds the language region, but the exact shapes and locations of the borders between the two vary from person to person.

The general-function region of Broca’s area appears to be part of a larger network sometimes called the multiple demand network, which is active when the brain is tackling a challenging task that requires a great deal of focus. This network is distributed across frontal and parietal lobes in both hemispheres of the brain, and all of its components appear to communicate with one another. The language-selective section of Broca’s area also appears to be part of a larger network devoted to language processing, spread throughout the brain’s left hemisphere.

Mapping functions

The findings provide evidence that Broca’s area should not be considered to have uniform functionality, says Peter Hagoort, a professor of cognitive neuroscience at Radboud University Nijmegen in the Netherlands. Hagoort, who was not involved in this study, adds that more work is needed to determine whether the language-selective areas might also be involved in any other aspects of cognitive function. “For instance, the language-selective region might play a role in the perception of music, which was not tested in the current study,” he says.

The researchers are now trying to determine how the components of the language network and the multiple demand network communicate internally, and how the two networks communicate with each other. They also hope to further investigate the functions of the two components of Broca’s area.

“In future studies, we should examine those subregions separately and try to characterize them in terms of their contribution to various language processes and other cognitive processes,” Fedorenko says.

The team is also working with scientists at Massachusetts General Hospital to study patients with a form of neurodegeneration that gradually causes loss of the ability to speak and understand language. This disorder, known as primary progressive aphasia, appears to selectively target the language-selective network, including the language component of Broca’s area.

The research was funded by the Eunice Kennedy Shriver National Institute of Child Health and Human Development, the Ellison Medical Foundation and the U.K. Medical Research Council.