BROCA’S AREA , WERNICKE’S AREA, AND OTHER

LANGUAGE-PROCESSING AREAS IN THE BRAIN

Broca’s area is generally defined as comprising Brodmann areas 44 and 45, which lie anterior to the
premotor cortex in the inferior posterior portion of the frontal lobe. Though both area 44 and area
45 contribute to verbal fluency, each seems to have a separate function, so that Broca’s area can be
divided into two functional units.

Area 44 (the posterior part of the inferior frontal gyrus) seems to be involved in phonological
processing and in language production as such; this role would be facilitated by its position close to
the motor centres for the mouth and the tongue. Area 45 (the anterior part of the inferior frontal
gyrus) seems more involved in the semantic aspects of language. Though not directly involved in
accessing meaning, Broca’s area therefore plays a role in verbal memory (selecting and
manipulating semantic elements).

Wernicke’s area lies in the left temporal lobe and, like Broca’s area, is no longer regarded as a
single, uniform anatomical/functional region of the brain. By analyzing data from numerous brain-
imaging experiments, researchers have now distinguished three sub-areas within Wernicke’s area.
The first responds to spoken words (including the individual’s own) and other sounds. The second
responds only to words spoken by someone else but is also activated when the individual recalls a
list of words. The third sub-area seems more closely associated with producing speech than with
perceiving it. All of these findings are still compatible, however, with the general role of Wernicke’s
area, which relates to the representation of phonetic sequences, regardless of whether the individual
hears them, generates them himself or herself, or recalls them from memory.
Wernicke’s area, of which the temporal planum is a key anatomical component, is located on the
superior temporal gyrus, in the superior portion of Brodmann area 22. This is a strategic location,
given the language functions that Wernicke’s area performs. It lies between the primary auditory
cortex (Brodmann areas 41 and 42) and the inferior parietal lobule.

This lobule is composed mainly of two distinct regions: caudally, the angular gyrus (area 39),
which itself is bounded by the visual occipital areas (areas 17, 18, and 19), and dorsally, the
supramarginal gyrus (area 40) which arches over the end of the lateral sulcus, adjacent to the
inferior portion of the somatosensory cortex.
The supramarginal gyrus seems to be involved in phonological and articulatory processing of
words, whereas the angular gyrus (together with the posterior cingulate gyrus) seems more
involved in semantic processing. The right angular gyrus appears to be active as well as the left,
thus revealing that the right hemisphere also contributes to semantic processing of language.

Together, the angular and supramarginal gyri constitute a multimodal associative area that receives
auditory, visual, and somatosensory inputs. The neurons in this area are thus very well positioned to
process the phonological and semantic aspect of language that enables us to identify and categorize
objects.

The language areas of the brain are distinct from the circuits responsible for auditory perception of
the words we hear or visual perception of the words we read. The auditory cortex lets us recognize
sounds, an essential prerequisite for understanding language. The visual cortex, which lets us
consciously see the outside world, is also crucial for language, because it enables us to read words
and to recognize objects as the first step in identifying them by a name.

There are wide variations in the size and position of Broca’s area and Wernicke’s area as
described by various authors.

Brain areas such as these, which perform high-level integration functions, are more
heterogeneous than areas that perform primary functions. This greater heterogeneity might
reflect greater sensitivity to environmental influences and greater plasticity (ability to adapt
to them). The functional organization of language would even appear to vary within the same
individual at various stages of his or her life!
Any attempt to define the precise boundaries of a particular area of the brain, such as Broca’s area
or Wernicke’s area, will involve some serious problems. But we do know that the
cytoarchitectonic areas described by Brodmann provide better anatomical correlates for brain
functions than do the shape of the brain’s convolutions. That said, a cortical area such as Broca’s
cannot be precisely described by reference to Brodmann areas alone. Though many authors regard
Broca’s area as consisting of Brodmann areas 44 and 45, other authors say it consists only of area
44, still others only of area 45, and yet others of areas 44, 45, and 47.

Broca’s area may also include the most ventral portion of Brodmann area 6, as well as other parts
of the cortex lying deep within the lateral sulcus. It is even possible that only certain parts of these
areas are actually dedicated to language.

Language acquisition in humans is based on our capacities for abstraction and for applying rules
of syntax—capacities that other animals lack. For example, brain-imaging experiments have
shown that Broca’s area becomes active when subjects are learning actual rules of grammar in
another language, but not when they are exposed to fictitious rules that actually violate the
grammar of that language.
These findings suggest that in Broca’s area, biological constraints interact with experience to make
the acquisition of languages possible. Broca’s area may thus represent the neuronal substrate of
the “universal grammar” shared by all of the world’s languages

MODELS OF SPOKEN AND WRITTEN LANGUAGE

FUNCTIONS IN THE BRAIN

In the 1980s, American neurologist Marsel Mesulam proposed an alternative to the Wernicke-
Geschwind model for understanding the brain’s language circuits. Mesulam’s model posits a
hierarchy of networks in which information is processed by levels of complexity.

For example, when you perform simple language processes such as reciting the months of the year
in order, the motor and premotor areas for language are activated directly. But when you make a
statement that requires a more extensive semantic and phonological analysis, other areas come into
play first.

When you hear words spoken, they are perceived by the primary auditory cortex, then processed by
unimodal associative areas of the cortex: the superior and anterior temporal lobes and the opercular
part of the left inferior frontal gyrus.
According to Mesulam’s
model, these unimodal areas
then send their information on
to two separate sites for
integration. One of these is the
temporal pole of the paralimbic
system, which provides access
to the long-term memory
system and the emotional
system. The other is the
posterior terminal portion of Approximate location of the inferior frontal gyrus. It
the superior temporal sulcus, is divided into three parts: the opercular, triangular,
which provides access to and orbital. The triangular part of the inferior frontal
meaning. The triangular and gyrus forms Broca’s area.
orbital portions of the inferior
frontal gyrus also play a role in
semantic processing.
Mesulam does, however, still believe that there are two “epicentres”for semantic processing, i.e.,
Broca’s area and Wernicke’s area. This new conception of these two areas is consistent with the fact
that they often work synchronously when the brain is performing a word processing task, which
supports the idea that there are very strong connections between them.

Mesulam’s concept of epicentres resembles that of convergence zones as proposed by other

authors: zones where information obtained through various sensory modalities can be combined.
This combining process is achieved through the forming of cell assemblies: groups of
interconnected neurons whose synapses have been strengthened by their simultaneous firing, in
accordance with Hebb’s law. This concept of language areas as convergence zones where neuronal
assemblies are established thus accords a prominent place to epigenetic influences in the process of
learning a language.

Unquestionably, one of these convergence zones is the left inferior parietal lobule, which comprises
the angular gyrus and the supramarginal gyrus. In addition to receiving information from the right
hemisphere, the left inferior parietal lobule also integrates emotional associations from the
amygdala and the cingulate gyrus.

Some scientists believe that over the course of evolution, language remained under limbic control
until the inferior parietal lobule evolved and became a convergence zone that provides a wealth of
inputs to Broca’s area. Some scientists also think that it was the emergence of the inferior parietal
lobule that gave humans the ability to break down the sounds that they heard so as to make sense of
them and, conversely, to express sounds in a sequential manner so as to convey meaning. In this
way, primitive emotional and social vocalizations would have eventually come to be governed by
grammatical rules of organization to create what we know as modern language.

One important idea in Mesulam’s model is that the function of a brain area dedicated to language
is not fixed but rather varies according to the “neural context”. In other words, the function of a
particular area depends on the task to be performed, because these areas do not always activate the
same connections between them. For instance, the left inferior frontal gyrus interacts with different
areas depending on whether it is processing the sound of a word or its meaning.

This networked type of organization takes us beyond the “one area = one function” equation and
explains many of the sometimes highly specific language disorders. For example, some people
cannot state the names of tools or the colours of objects. Other people can explain an object’s
function without being able to say its name, and vice versa.

Brain-imaging studies have shown to what a large extent cognitive tasks such as those involving
language correspond to a complex pattern of activation of various areas distributed throughout the
cortex. That a particular area of the brain becomes activated when the brain is performing certain
tasks therefore does not imply that this area constitutes the only clearly defined location for a
given function. In the more distributed model of cognitive functions that is now increasingly
accepted by cognitive scientists, all it means is that the neurons in this particular area of the brain
are more involved in this particular task than their neighbours. It in no way excludes the
possibility that other neurons located elsewhere, and sometimes even quite far from this area, may
be just as involved.

Thus, just because the content of a word is encoded in a particular neuronal assembly does not
necessarily mean that all of the neurons in this assembly are located at the same place in the brain.
On the contrary, understanding or producing a spoken or written word can require the
simultaneous contribution of several modalities (auditory, visual, somatosensory, and motor).
Hence the interconnected neurons in the assembly responsible for this task may be distributed
across the various cortexes dedicated to these modalities.
In contrast, the neuronal assemblies involved in encoding grammatical functions appear to be less
widely distributed.

It may therefore be that the brain processes language functions in two ways simultaneously: in
parallel mode by means of distributed networks, and in serial mode by means of localized
convergence zones.

Lastly, a number of researchers now

reject classic locationist models of
language such as Geschwind’s and
Mesulam’s. Instead, they conceptualize
language, and cognitive functions in
general, as being distributed across
anatomically separate areas that process
information in parallel (rather than
serially, from one “language area” to
another).

Even those researchers who embrace

this view that linguistic information is
processed in parallel still accept that the
primary language functions, both
auditory and articulatory, are localized
to some extent.

This concept of a parallel, distributed

processing network for linguistic
information constitutes a distinctive
epistemological paradigm that is
leading to the reassessment of certain
functional brain imaging studies.

The proponents of this paradigm

believe that the extensive activation of
various areas in the left hemisphere and
the large number of psychological
processes involved make it impossible
to associate specific language functions
with specific anatomical areas of the
brain. For example, the single act of
recalling words involves a highly
distributed network that is located
primarily in the left brain and that
includes the inferolateral temporal lobe,
the inferior posterior parietal lobule, the
premotor areas of the frontal lobe, the
anterior cingulate gyrus, and the
supplementary motor area. According
to this paradigm, with such a widely
distributed, parallel processing network,
there is no way to ascribe specific
functions to each of these structures that
contribute to the performance of this
task.
 The brain does seem to access meanings by way of categories that it stores in different
physical locations. For example, if the temporal pole (the anterior end of the temporal lobe) is
damaged, the category “famous people” is lost; if a lesion occurs in the intermediate and
inferior parts of the temporal lobe, the category “animals” disappears. It also seems that the
networks involved in encoding words activate areas in the motor and visual systems. The task
of naming tools activates the frontal premotor areas, while that of naming animals activates
the visual areas. But in both cases, Broca’s area and Wernicke’s area are not even activated.

Among those scientists who argue that the brain’s language processing system is distributed
across various structures, some, such as Philip Lieberman, believe that the basal ganglia play
a very important role in language. These researchers further believe that other subcortical
structures traditionally regarded as involved in motor control, such as the cerebellum and the
thalamus, also contribute to language processing. These views stand in opposition to
Chomsky’s on the exceptional nature of human language and fall squarely within an
adaptationist, evolutionary perspective.

HANDEDNESS, LANGUAGE, AND BRAIN

LATERALIZATION

The brain’s anatomical asymmetry, its lateralization for language, and the phenomenon of
handedness are all clearly interrelated, but their influences on one another are complex. Though
about 90% of people are right-handed, and about 95% of right-handers have their language areas on
the left side of their brains, that still leaves 5% of right-handers who are either right-lateralized for
language or have their language areas distributed between their two hemispheres. And then there are
the left-handers, among whom all of these patterns can be found, including left-lateralization.

Some scientists suggest that the left hemisphere’s dominance for language evolved from this
hemisphere’s better control over the right hand. The circuits controlling this “skilful hand” may
have evolved so as to take control over the motor circuits involved in language. Broca’s area, in
particular, is basically a premotor module of the neocortex and co-ordinates muscle contraction
patterns that are related to other things besides language.

Even in many species that are quite distant from humans in evolutionary terms (frogs, for
example), the brain is left-lateralized for the vocalization function.
In chimpanzees, lateralization for the anatomical areas corresponding to Broca’s and
Wernicke’s areas already exists, even though it does not yet correspond to the language
function. And like the majority of humans, the majority of chimpanzees use their right hand
in preference to their left.

These asymmetries in the other primates represent persuasive evidence of the ancient
phylogenetic origin of lateralization in the human brain. The expansion of the prefrontal
cortex in humans might in part reflect its role in the production of language.
 Women have the reputation of being able to talk and listen while doing all sorts of things at
the same time, whereas men supposedly prefer to talk or hear about various things in
succession rather than simultaneously. Brain-imaging studies may now have revealed an
anatomical substrate for this behavioural difference, by demonstrating that language
functions tend to place more demands on both hemispheres in women while being more
lateralized (and mainly left-lateralized) in men. Women also have more nerve fibres
connecting the two hemispheres of their brains, which also suggests that more information is
exchanged between them.

Brain-imaging studies have

shown that several structures
involved in language
processing are larger in the
left hemisphere than in the
right. For instance, Broca’s
area in the left frontal lobe is
larger than the homologous
area in the right hemisphere.
But the greatest asymmetries
are found mainly in the
posterior language areas,
such as the temporal planum
and the angular gyrus.
Two other notable asymmetries are the larger protrusions of the frontal lobe on the right side and the
occipital lobe on the left. These protrusions might, however, be due to a slight rotation of the
hemispheres (counterclockwise, as seen from above) rather than to a difference in the volume of
these areas. These protrusions are known as the right-frontal and left-occipital petalias (“petalias”
originally referred to the indentations that these protrusions make on the inside of of the skull).

The structures involved in producing and understanding language seem to be laid down in
accordance with genetic instructions that come into play as neuronal migration proceeds in the
human embryo. Nevertheless, the two hemispheres can remain just about equipotent until language
acquisition occurs. Normally, the language specialization develops in the left hemisphere, which
matures slightly earlier. The earlier, more intense activity of the neurons in the left hemisphere
would then lead both to right-handedness and to the control of language functions by this
hemisphere.

But if the left hemisphere is damaged or defective, language can be acquired by the right
hemisphere. An excess of testosterone in newborns due to stress at the time of birth might well be
one of the most common causes of slower development in the left hemisphere resulting in greater
participation by the right.
This hypothesis of a central role for testosterone is supported by experiments which showed that in
rats, cortical asymmetry is altered if the rodents are injected with testosterone at birth. This
hormonal hypothesis would also explain why two-thirds of all left-handed persons are males.

Interindividual variations, which are essential for natural selection, are expressed in various
ways in the human brain. Volume and weight can vary by a factor of two or even more. The
brain’s vascular structures are extremely variable; the deficit caused by an obstruction at a
given point in the vascular system can vary greatly from one individual to another. At the
macroscopic anatomical level, the folds and grooves in the brain also vary tremendously from
individual to individual, especially in the areas associated with language. Variability in the
language areas can also be observed at the microscopic level, for example, in the synaptic
 structure of the neurons in Wernicke’s area.
Interindividual variability is also expressed in the brain’s functional organization, and
particularly in the phenomenon of hemispheric asymmetry. For instance, some data indicate
that language functions may be more bilateral in women than in men. The percentage of
atypical lateralization for language also varies with handedness: it is considerably higher among
left-handers than among right-handers.

Lastly, as if all this were not enough, there is also such a thing as intraindividual variability. In
the same individual, a given mental task can sometimes activate different neuronal assemblies
in different circumstances—for instance, when the individual is performing this task for the
first time, as opposed to when he or she has already performed it many times before.

THE RIGHT HEMISPHERE’S CONTRIBUTION TO

LANGUAGE

To follow a conversation, a written document, or an exchange of witticisms, you must be able not
only to understand the syntax of sentences and the meanings of words, but also to interrelate
multiple elements and interpret them with respect to a given context. While various types of damage
to the left hemisphere produce the many documented forms of aphasia, right hemisphere damage
(RHD) causes a variety of communication deficits involving the interpretation of context. These
deficits can be divided into two main categories.

The first category of RHD-induced deficits affect communication indirectly, by disrupting people’s
ability to interact effectively with their environment.
One example of a
deficit that can be
caused by RHD is
hemineglect, in which
an individual pays no
attention to stimuli
presented to the
various sensory
modalities on the left
side of the body.
Drawings 2, 4, 5, and 6 were made by a patient with hemineglect.

The individual may also suffer from

anosognosia: unawareness of such deficits.
For instance, some people who have
damage just posterior to the central sulcus
in their right hemispheres cannot even
recognize certain parts of their own bodies
as being their own. Thus this type of RHD
produces a kind of indifference that is the
opposite of the minimum emotional
investment required to establish
harmonious communication.
The other major family of RHD-induced deficits affect communication and cognition directly.
These deficits can be grouped under the heading of pragmatic communication disorders, pragmatics
being the discipline that studies the relationships between language and the way that people use it in
context. Pragmatic disorders can be subdivided into disorders in prosody, discourse organization,
and understanding of non-literal language.
Prosody refers to the intonation and stress with which the
phonemes of a language are pronounced. People with
aprosodia—RHD that impairs their use of prosody—
cannot use intonation and stress to effectively express the
emotions they actually feel. As a result, they speak and
behave in a way that seems flat and emotionless.

The second category of pragmatic communication

Image of the brain of a woman
who is deciding whether or not
certain words rhyme. As can be
seen, the right hemisphere is
very active.
Source: Shaywitz and Shaywitz,
Yale Medical School
disorders that can be caused by RHD affect the
organization of discourse according to the rules that
govern its construction. In some individuals, these
disorders take the form of a reduced ability to interpret the
signs that establish the context for a communication, or the
nuances conveyed by certain words, or the speaker’s
intentions or body language, or the applicable social
conventions. With regard to social conventions, for
example, people generally do not address their boss the
same way they would their brother, but people with certain
kinds of RHD have difficulty in making this distinction.

Last but not least among the types of pragmatic communication disorders caused by RHD are
disorders in the understanding of non-literal language. It is estimated that fewer than half of the
sentences that we speak express our meaning literally, or at least they do not do so entirely. For
instance, whenever we use irony, or metaphors, or other forms of indirect language, people’s ability
to understand our actual meaning depends on their ability to interpret our intentions.

To understand irony, for example, people must apply two levels of awareness, just as they must do
to understand jokes. First, they must understand the speaker’s state of mind, and second, they must
understand the speaker’s intentions as to how his or her words should be construed. Someone who
is telling a joke wants these words not to be taken seriously, while someone who is speaking
ironically wants the listener to perceive their actual meaning as the opposite of their literal one.

Metaphors too express an intention that belies a literal interpretation of the words concerned. If a
student turns to a classmate and says “This prof is a real sleeping pill”, the classmate will
understand the implicit analogy between the pill and the prof and realize that the other student finds
this prof boring. But someone with RHD that affects their understanding of non-literal language
might not get this message.
Lastly, the various indirect ways that we commonly use language in everyday life can cause
problems for people with RHD. In such cases, the speaker’s actual intention underlies their oral
statement as such. For example, someone who says “I wonder what the time is now ” is indirectly
asking for someone to tell them the time, but a person with RHD may not understand that.
Though the left hemisphere is still regarded as the dominant hemisphere for language, the role of
the right hemisphere in understanding the context in which language is used is now well
established. We know that in the absence of the left hemisphere (for example, when Wada’s test is
performed), the right hemisphere can produce some rudimentary language. But lesion studies have
shown that the right hemisphere’s role in language appears to be far wider—so much so that it is
now more accurate to think of the two hemispheres’ language specializations not as separate
functions, but rather as a variety of abilities that operate in parallel and whose interaction makes
human language in all its complexity possible.

Many theories have been offered to explain people’s ability to adapt their use of language to the
interpersonal context. One of these is the theory of mind. According to Premack and Woodruff
(1978), the theory of mind is the ability that lets people ascribe mental processes to other people,
to reason on the basis of these ascribed processes, and to understand the behaviours that arise from
them. Premack and Woodruff were the first authors to use the term “theory of mind”. They did so
in a study on the ability of chimpanzees to ascribe beliefs and intentions to human beings. Since
the time of this study, the theory of mind has been applied mainly in studies comparing the
cognitive development of normal children and autistic children, because the latter represent a
population that is known to display deficits in social reasoning from the very earliest age.

When experimental subjects are asked to identify the emotional content of recorded sentences that
are played back into only one of their ears, they perform better if these sentences are played into
their left ear (which sends them to the right hemisphere) then into their right (which sends them to
the left hemisphere).