To be published in Behavioral and Brain Sciences (in press)

Cambridge University Press 2007

Below is the unedited précis of a book that is being accorded BBS multiple book
review. This preprint has been prepared for potential commentators who wish to
nominate themselves for formal commentary invitation. Please do not write a
commentary unless you receive a formal invitation. Invited commentators will receive
full instructions. Commentary must be based on the book.*

*Neuroconstructivism: How the Brain Constructs Cognition, Vols 1 & 2

Denis Mareschal, Mark Johnson, Sylvain Sirois, Michael Spratling, Michael Thomas,
and Gert Westermann

Précis of:

Neuroconstructivism: How the Brain Constructs Cognition

Sylvain Sirois1*, Michael Spratling2,3, Michael S. C. Thomas3, Gert Westermann4,3 , Denis
Mareschal3, and Mark H. Johnson3,
1
School of Psychological Sciences
The University of Manchester
Oxford Road, Manchester
M13 9PL UK
2
Division of Engineering
King's College London
Strand, London
WC2R 2LS UK
3
Centre for Brain and Cognitive Development
School of Psychology
Birkbeck University of London
Malet Street, London
WC1E 7HX UK
4
Department of Psychology
Oxford Brookes University
Oxford
OX3 0BP UK

* author for correspondence

1
Abstract: Neuroconstructivism proposes a unifying framework for the study of development
that brings together (1) constructivism (which views development as the progressive
elaboration of increasingly complex structures), (2) cognitive neuroscience (which aims to
understand the neural mechanisms underlying behaviour), and (3) computational modelling
(which proposes formal and explicit specifications of information processing). The guiding
principle of our approach is context dependence, within and (in contrast to Marr) between
levels of organization. We propose that three mechanisms guide the emergence of
representations: competition, cooperation, and chronotopy, which themselves allow for two
central processes: proactivity and progressive specialization. We suggest that the main
outcome of development is partial representations, distributed across distinct functional
circuits. This framework is derived by examining development at the level of single neurons,
brain systems, and whole organisms. We use the terms encellment, embrainment, and
embodiment to describe the higher-level contextual influences that act at each of these levels
of organization. To illustrate these mechanisms in operation we provide case studies in early
visual perception, infant habituation, phonological development, and object representations in
infancy. Three further case studies are concerned with interactions between levels of
explanation: social development, atypical development and within that, the development of
dyslexia. We conclude that cognitive development arises from a dynamic, contextual change
in neural structures leading to partial representations across multiple brain regions and
timescales.

Keywords: brain, cognition, development, constructivism, embodiment

1. Introduction

Neuroconstructivism draws upon three traditions. The first is the constructivist view
of development attributed to Piaget and his contemporaries, such as the
developmental biologist Waddington. “Neuro” introduces the second tradition: a
commitment to view psychological development as entwined with the mechanistic
and morphological aspects of brain development, from cell to brain to body. Third is
computational modelling, which forces process theories to be explicit about the nature
of information processing, resulting in a level of specification that eludes traditional,
verbal, descriptive theories. Two recent books brought two of these three traditions
together. Rethinking Innateness (Elman, 1996) argued for a connectionist modelling
approach to understanding brain and cognitive development. In contrast, A Dynamic
Systems Approach to the Development of Cognition and Action (Thelen and Smith,
1994) emphasized development as occurring in the context of embodiment. In
Neuroconstructivism: How the Brain Constructs Cognition (Mareschal et al. 2007a)
we add a focus on neural development and the development of representations.

Neuroconstructivism1 emphasizes the interrelation between brain development and

cognitive development. We see constructivist development as a progressive increase

1
We acknowledge that term Neuroconstructivism has been used by others previously,
sometimes with a general meaning very similar to our usage (Karmiloff-Smith 1998, Quartz and
Sejnowksi, 1997, and sometimes with a differing meaning, Sheridan, 1997).

2
in the complexity of representations, with the consequence that new competences can
develop based on earlier, simpler ones. This increase in representational complexity is
realized in the brain by a progressive elaboration of cortical structures. Thus, while
other constructivist theories have emphasized the notion of hierarchical integration of
knowledge (e.g., Piaget, 1970; Karmiloff-Smith, 1992), we explore the relationship
between the elaboration of knowledge and the development of new cognitive abilities.
We assert that increases in representational complexity arise as a natural consequence
of the processes of adaptation typical of complex biological systems like the brain
(see also Quartz and Sejnowski, 1997; Shultz, 2003, for related views).
Neuroconstructivism implies the creation of genuinely new cognitive abilities and not
just the better use of pre-existing abilities.

While our focus is on the development of mental representations, these develop in a

physical and social environment. The body, which changes substantially during
infancy, constrains what the infant can experience, process, and do. This, in turn,
constrains possible learning and development. For example, the limited visual acuity
of babies helps to simplify the visual environment, and could also help with the
integration of modalities as the clearest objects are those within reach. Physical
constraints on action also force a “starting small” situation, whereby the child learns
simpler effective behavior before being able to acquire more complex abilities. The
social environment also constrains what and how the child will learn. The use of
motherese, for instance, simplifies the linguistic input early in language acquisition.
Indeed, most human societies engage in a process of gradually exposing infants and
children to the kinds of problems that they will need to master in order to survive.
The guiding principle behind our approach is context dependence. Representations in
the brain do not emerge or function in isolation, but within the context of co-occurring
molecular, neural, bodily, and social events. This constrains and guides emerging
representation through three mechanisms: cooperation, competition, and chronotopy
(see section 2.4). The mechanisms take different forms at the different levels of
implementation, but together they enable two central developmental processes:
proactivity, which is concerned with the role of internally-generated activity in the
development of function, and progressive specialization, the fact that functions
exhibit a progressive restriction of fate by becoming more specific and less plastic.
We argue that the outcome of these developmental mechanisms and processes are
partial representations. The brain acquires and develops multiple, fragmentary
representations that are just sufficient for on-the-fly processing. The role of
developmental psychology is to understand how and why such partial representations
emerge, how they interact, and the flexibility of their configuration.

One novel aspect of our approach is that our emphasis on context requires consistency
between levels of explanation. We view cognitive functions as inextricably linked to
their neural implementation and to the dynamical environments in which they emerge
and operate, with interactions going both ways across levels. As far as explaining
cognitive development, we see little merit in theories or models that fit data at one
level, however well, yet contradict what is known at other levels. Thus, we argue for a
radical rejection of Marr’s (1982) independent levels of analysis argument. While

3
explanations can be formulated independently at different levels of description, those
levels are not themselves independent. A consequence of our focus on consistency is
parsimony. If a phenomenon can be explained at different levels using a unitary
framework, then this is preferable to an alternative where different and inconsistent
interpretations are used at each level.

In the next section, we lay out the foundations of Neuroconstructivism, culminating

with a proposed set of principles, mechanisms, and processes. Section 3 illustrates
these ideas through the use of case studies in which the Neuroconstructivist
framework is applied to different domains of cognitive development. In a concluding
section, we identify future challenges and briefly discuss Neuroconstructivism:
Volume 2.

2. Foundations

2.1 Encellment

The development of the nervous system is typically described as a two-stage process.

Initially, coarse structure and connectivity is laid out, with little contribution from the
electrical activity of neurons. Then, the firing of neurons becomes crucial in
establishing the finer-grained details of connectivity. Therefore, to examine brain
development at the cellular level, we need to distinguish between context-dependent
and activity-dependent processes (Crowley & Katz, 1999; Herrmann & Shatz, 1995).
In the early stages of neural development activity probably plays the least role in the
outcome. In neurogenesis, precursor cells (neuroblasts) differentiate into neurons,
glial cells, or new precursor cells. The outcome of differentiation is affected both by
the lineage of the cell and, crucially, cell-cell interactions. A new neuron then
migrates to its final position, either by passive displacement (i.e., being pushed by
other emerging neurons) or with the help of radial glial cells that guide neural
migration. Thus, local cellular context plays a key role in the formation of neural
structures.

Neural differentiation begins during, or towards the end of, migration. Axons often
traverse long distances (Purves et al., 1997), facilitated by a mixture of activity-
dependent and activity-independent processes. A neurite (the neuron outgrowth that
will become the axon) develops protrusions that will travel in space in response to
both chemical and physical extracellular events. Particular cues guide axonal growth
while others impair it, and processes can seek specific target locations with a unique
chemical signature (Goodman & Shatz, 1993). Once in a target area, an axon forms
specific connections with dendrites through competition with other dendrites and
other axons. Endogenous electrical activity also plays a role in early, pre-synaptic
neural differentiation such that early in brain development, one observes key roles for
both context and activity-dependence.

Overproduction of cells leads to programmed cell death (Oppenheim, 1991). The

death of a neuron is controlled by a combination of intrinsic and extrinsic factors,

4
whereby an internal “suicide” program will be triggered or suppressed by external
chemical events involved in the guidance and competition for neural differentiation
through trophic factors. Crucially, neural activity appears to have a protective effect
on neurons, as it reduces cell death (Ghosh et al., 1994).

Activity-dependence is one part of a feedback loop with morphology, with each

affecting the other. The activity of neurons can also alter subcellular ion channels and
neurotransmitter receptors, so that the response profile of a neuron is adaptive over
time (Turrigiano et al., 1994). Gene expression, too, can be affected by neural activity
(Armstrong & Montminy, 1993). Activity-dependence also affects the connectivity
between cells. A good example is the emergence of ocular dominance columns
(ODC): preventing sensory input to one eye considerably reduces the space occupied
by neurons responding to that eye, illustrating activity-dependent competition.
Two competing theories have been proposed to explain the emergence of cortical
areas. One is the protomap view (Rakic, 1988), which suggests that neurons are
predestined, early in development, to occupy specific functions in specific cortical
areas. The second is the protocortex view (O’Leary, 1989), which argues that
functional areas emerge from interactions with subcortical structures and between
cortical areas. Recent reviews propose a midway view in which patterns of gene
expression create, at a coarse scale, cortical areas more suited to adopting particular
functions as per the protomap view. However, a protocortex account better explains
the emergence of function within those poorly spatially and functionally defined
areas.

2.2 Embrainment

Embrainment refers to the view that functional areas of the brain emerge and exist
within a context of connections to and from other functional areas. This contrasts with
a view of functional brain development in which regions are presumed to mature in
relative isolation of their context, and to the view that particular cognitive operations
can be localized to individual regions in adults. In fact, there is substantial evidence
that the functional properties of specific brain regions are highly constrained by their
past and present interactions with neighboring areas.

For instance, visual ERP components of congenitally deaf people differ markedly
from those of typical hearing participants and participants who became deaf after the
age of four (Neville & Lawson, 1987). These authors proposed that early (but not late)
lack of auditory input allowed a reallocation of resources, such that cortical areas
typically involved in auditory processing were taken over, to a degree, by visual
processing. Similarly, it has been shown that the visual cortex of people blind from an
early age can be activated by tactile stimuli, particularly Braille reading. In both these
examples, the differentiation of the cortex into areas of functional specialization
results from a developmental process and is not functionally encapsulated. If the
context changes during development, then so too can the function associated with a
cortical area.

5
There are three important questions to consider when studying the aetiology of brain
functions. First, does development involve deterministic epigenesis or probabilistic
epigenesis (Gotlieb, 2007)? With the former, the assumption is a unidirectional path
between gene and brain function. With the latter, the relationship between genes,
structure, and function is bidirectional and dynamic. Second, is there a direct mapping
between brain structure and function, and does this change over development? The
third question concerns the nature of brain plasticity, and whether this changes over
development.

A substantial amount of research concerned with mapping brain and behavioral

development has taken a maturational viewpoint, whereby emerging behaviors are
construed to reflect underlying maturing functions of isolated areas of the brain (see
Diamond, 1991). A tacit assumption is that the typical adult brain (and thus behavior)
is prespecified in a protomap (deterministic epigenesis), with a direct mapping
between structure and function. Within this view, plasticity is a special mechanism
activated by brain injury. An alternative to the maturational view is the skill-learning
perspective, which proposes a continuity between infancy and adulthood in the
mechanisms underlying brain learning and plasticity. According to this view,
plasticity is a long-lasting feature of the brain that only appears to be reduced within a
context of the stable constraints that are more likely in adulthood.

The Interactive Specialization viewpoint proposes a middle ground between the

previous two accounts (Johnson, 2005). It proposes that brain regions develop within
the context of other brain regions (embrainment), and that the functional development
of brain regions is shaped in part by interregional interactions. Specifically, cortical
functional brain development is characterized by a process of increased tuning, or
selectivity of functions. Thus, the mapping between structures and function can and
will change during development. Within this viewpoin,t plasticity is retained when a
function is not yet fully specialized.

Functional cortical brain development is best described as progressive localization

and progressive specialization of function, through competition and cooperation
between distinct areas. Representations that emerge within a region are constrained by
existing representations in functionally neighboring areas, consistent with the
interactive specialization view.

2.3 Embodiment

While the distinction between mind and the physical world may have surface appeal,
ethological work has revealed a much closer coupling between behavior and
environment. The brain is best viewed as embedded in its environment, and not
divorced from it. As at other levels of organization, the study of a specific system
must involve consideration of the other systems to which it is coupled. In the case of
the brain, it is unhelpful to ignore the body and the external environment.
While previous work has distinguished embodiment (the constraints of the body on
the brain) from situatedness (the constraints of the environment on the agent), we use

6
embodiment to encompass both types of interaction. Taking embodiment seriously
can reveal simpler solutions to cognitive problems than would be achieved by non-
embodied approaches (e.g., Webb, 1994). Indeed, considering the contribution of both
body and environment can reduce the purported contribution of the nervous system
(Clark, 1997). For some cognitive problems, parts of the solution exist in bodily
constraints and environmental properties. The role of the brain is to coordinate inner
and outer worlds (Ballard et al., 1997). Hence, representations are not independent
from the environment; rather, they contain partial information about the environment,
sufficient to support contextually specific behaviors. Representations serve to cause
behaviors rather than to mirror the environment.

Clark (1997) identifies several important ways in which an embodied perspective

provides benefits to cognitive research. First, it raises awareness to the fact that an
important function for organisms is to harness the environment to their advantage
(Hutchins, 1995). Second, the planning and execution of motor actions must consider
body/environment couplings (Thelen et al., 1996). Third, it stresses the online nature
of information processing (Goldstein & Gigerenzer, 2002), which relies on context-
dependent heuristics for just-in-time adaptations. Fourth, it recognizes how elements
of the environment can act as extensions of the mind, reducing cognitive load. Fifth,
Clark (1997) proposes that language embeds individuals within society allowing
individuals to share representations, and affect one another’s behavior. In terms of
development, language provides a crucial tool to guide the experiences of infants and
children (Rogoff, 1998; Vygotsky, 1986).

Embodied models take one of two forms. In agent modeling, both the organism and
the environment are computer simulations (e.g., Schlesinger, 2004). Alternatively,
researchers may use real robots that function in real environments. An example of this
second approach is ‘Didabot’ (Maris & te Boekhorst, 1996), a simple robot that
avoids objects perceived by sensors to its left or right. The robot has a ‘blind spot’ in
front of it. When several Didabots were placed in an enclosed environment that
contained cubes small enough to fall within the robots’ blind spots the cubes ended up
in heaps at the center and periphery of the arena. This apparently complex “tidying”
behavior emerged from the coupling of a simple mind (avoid obstacles), body (blind
spot), and environment (cubes and other robots).

Developmental embodied cognition focuses on the co-development of the nervous

system and the body within a dynamic environment. This has roots in ecological
psychology (e.g., Gibson, 1979;1982). A key concept of this earlier work is that of
affordances, the fact that particular stimuli invite a specific range of actions in relation
to the agent’s structure and skills. Recent infancy work supports the notion of action
affordance in terms of representing objects (Mareschal & Bremner, 2005; Mareschal
& Johnson, 2003). Thelen and Smith (1994) argued more generally that the interplay
between thought and action is ubiquitous in infancy, consistent with Piaget’s notion of
early sensori-motor development (Piaget, 1952). Importantly, this approach stresses
how the child actively manipulates the environment, with dynamic consequences in
respect to the stimulation encountered. Similarly, the onset of self-locomotion brings

7
about such a major change in the infant’s effective environment that some have
argued it causes a major reorganization of cognitive structures (Campos et al., 2000).

2.4. Principles, mechanisms, and processes

A core principle of the Neuroconstructivist approach is context-dependence. At each

level of description or analysis, the function of interest depends on the context in
which it is realized. Furthermore, context-dependence is particularly important for the
development of those functions and has significant implications for the representations
that emerge.

Context-dependence constrains emerging representations through three domain-

general, level-independent mechanisms: cooperation, competition, and chronotopy (or
timing). The specific implementation of these mechanisms will vary depending on the
level of analysis. The mechanisms themselves make possible two processes that
underlie the development of representations: proactivity and progressive
specialization. The outcome of these processes is the emergence of partial
representations (Figure 1).

8
 Core Principle

Context Dependence

Competition Cooperation Chronotopy

General
mechanisms

Progressive Developmental
Proactivity
specialization
processes

Partial
Outcome
representation

Figure 1. Principles, mechanisms and processes involved in the Neuroconstructivist

framework. Both the mechanisms and processes can be construed as operating at multiple
levels of description. Hence, the arrows do not imply a direction of information flow.

9
Competition implies that from the many initial contributors to an immature function,
only a subset of these will ultimately be involved in the mature function. For example,
over time, the expression of gene A may prevent the expression of gene B. Similarly,
inhibitory neurons or inhibitory brain structures (e.g., frontal lobe inhibition of
subcortical functions) restrict competing processes from participating in a particular
function. At the cognitive level, one representation of a sensory input (e.g. one view
of the ambiguous Necker cube) may compete with another incompatible interpretation
of the same sensory data. Overall, the purpose of competition is to allow for stable,
minimal representations.

Cooperation, however, is a mechanism involved in the integration of multiple

contributors to a function. For example, some genes serve as triggers for other genes,
co-activity of neurons help build circuits, different brain systems may need to be
simultaneously involved in a particular function, and social behavior requires
cooperation between individuals. Unlike competition, but complementary to it,
cooperation strives for overall efficiency through the coordination of interrelated
functions. Together, competition and cooperation help build a system that may be
minimal but involves a degree of redundancy that makes it relatively robust to
damage.

The notion of chronotopy stresses that time is a dimension of development (cf. Elman
et al., 1996). Some patterns of gene expression are restricted to specific
developmental times, some key aspects of neural development rely on sequences of
events, and adaptive plasticity occurs at different times in different parts of the
developing system. At a cognitive level, this translates to saying that children will
solve restricted problems grounded within a limited domain before solving abstract
general problems that span several domains. Perhaps the most important temporal
aspect concerns restrictions to plasticity. Neural commitment means that some circuits
may be hard to alter once wired. Processing commitment, which is linked to neural
commitment, means that functions become progressively entrenched and selective,
and may lose sensitivity to inputs outside of their current range.

Two developmental processes operate on a larger timescale than the above

mechanisms. Proactivity refers to the idea that, at least in part, representations reflect
internally generated activity. For example, spontaneous neural activity helps to form
certain synaptic connections. Similarly, the child initiates behaviors that have effects
on the environment that, in turn, affect sensory input and ensuing behavior. We view
such a feedback loop, with key involvement from a proactive child, as the engine of
development. There is substantial evidence that, from an early age, infants and
children are selective about the information they process from the environment
(Cohen, 1972; Fantz, 1964; Posner, 1993). There is also evidence revealing a role for
spontaneously generated movements on early development (Goldfield et al., 1993;
Robertson et al., 2001), even in utero (Precht, 2001; Robertson, 1988).

The other developmental process is progressive specialization. In its simplest form,

the state of the system at any given time places constraints on future states of the

10
system. Waddington’s (1957) metaphor of an epigenetic landscape captures the idea
well. According to Waddington, development is like a ball rolling down an uneven
surface, able to take different directions as a function of its direction, inertia, and the
landscape. Typical development would see most balls end up in the same general area
of the landscape, and atypical development would see balls end up in different areas
because of changes to initial direction, inertia, or landscape. A good example of such
restriction-of-fate over time is phonological development. Although newborns can
distinguish speech sounds from all human languages, the ability to discriminate non-
native speech sounds drops substantially after about six months (Stager & Werker,
1997; Trehub, 1976). Importantly, progressive specialization does not simply limit
future adaptations but can also facilitate learning. In a constructivist framework, early
knowledge often provides the building blocks for further knowledge.

We argue that the outcomes of these developmental processes are partial

representations. The brain contains multiple fragmentary and partial representations
that are sufficient to allow successful behavior, for example, in response to a given
object over a range of contexts. Each of these is able to have an independent causal
effect on behavior. This view is consistent with the distributed processing that takes
place in the brain and is a computationally efficient solution to representation. New
representations are thus acquired in the context of existing (also fragmentary)
representations, the current effective learning environment, and the current
developmental state of the body.

3. Case studies

This section presents different areas of research that serve to illustrate how our
approach can be applied in different domains. The first four case studies are concrete
examples of how the mechanisms operate. The last three cases are concerned with the
notion of interactions between the different levels of explanation.

3.1. Early visual perception

Visual information processing in adults involves distinct cortical regions (van Essen
et al., 1992). For example, during object recognition cortical processing begins in area
V1 (the primary visual cortex) and progresses through a series of cortical regions until
object identity per se is processed in the inferotemporal cortex. This portion of the
visual system dealing with object recognition is known as the ventral pathway
(Ungerleider & Mishkin, 1982). Along this pathway early areas are involved in
simpler tasks such as edge, contrast, or orientation detection (e.g., Hubel & Wiesel,
1977), whereas later areas are involved in progressively more abstract representations
(such as, ultimately, identifying an object). However, the pathway is not
unidirectional but involves reciprocal connections between regions (Lamme &
Roelfsema, 2000). Later processes can affect earlier ones, such that the dynamics of
information processing are more important than their specific anatomical locations
(Felleman & van Essen, 1991). Each cortical region is embedded in a network of
other regions and processes information in this dynamical context.

11
Within each region there exists competition between cells, such as that occurring
through lateral inhibition. This competition occurs at each stage of processing and
reflects both bottom-up stimulation and top-down biases (Desimone & Duncan,
1995). Cooperation is also an important process in visual processing. To identify an
object, it is necessary to group features of the visual input that belong to that object
and to segment those from the background. This is achieved through an interactive
process involving feedforward and feedback connections between the different
cortical areas involved in object perception (Driver et al., 2001). Chronotopy is also
important, as the functionality of the visual system emerges from a peripheral to
central ordering over time (Johnson & Vecera, 1996; Shrager & Johnson, 1996). Later
developing neurons in higher-order areas can exploit the earlier, partial
representations developed in lower, peripheral regions. Representations learned at
each level constrain the representational space of subsequent levels, allowing high-
order regularities to be extracted (Clark & Thornton, 1997).
The input selectivity of cells involved in visual processing can be changed through
learning (Desimone, 1996). Moreover, experience leads to a decrease in the
population of cells that respond to a familiar stimulus (Rainer & Miller, 2000).
Changes in performance thus reflect changes in representations, distributed across a
smaller, selective population of neurons (Karni et al., 1995). Some work also reports
that the cortical regions involved in a task may change as a function of expertise
(Walsh et al., 1998), with fewer regions involved after learning than was initially the
case (Petersen et al., 1998). This experience-dependent selectivity can enhance
processing of subsets of visual inputs at the expense of other subsets. Selective
attention implements a form of proactivity, whereby the child preferentially attends to
some stimuli over others.

Overall, cortical regions involved in visual processing carry out contextualized

intraregional competition and interregional cooperation, modulated by a degree of
chronotopy that forces simpler representations to be acquired prior to progressively
more complex representations. The progressive specialization at each level of
processing reflects experience, which is proactive. The outcome is a set of partial
representations across a complex network of cortical areas that together enable object
recognition.

3.2. Infant habituation

Because of the limited perceptual and motor skills of infants, researchers have devised
many indirect methods to assess cognitive abilities in infants. The most popular
method is the use of looking time data (e.g., Thorpe, 1956). Over the repeated
presentation of stimuli, babies show a progressive decrease of interest, reflected by
shorter looking times to the stimuli. They are then deemed to have habituated.
However, they can show renewed interest (assessed from relatively longer looking)
when presented with novel stimuli. This ‘novelty preference’ is an example of
proactive exploration of the environment. By careful manipulation of how the habitual
and novel stimuli differ from each other, researchers can make claims about what

12
infants perceive to be distinct, with implications for the nature of their underlying
representations. A common approach, owing to the pioneering work of Sokolov
(1963), is to suggest that a mental representation of the habitual set of stimuli is
learned, and that the progressive decrease in looking-time reflects an increasing match
between this internal represenataion and the stimuli. Novelty preference is then taken
to reflect a mismatch between the internal representation and the novel stimulus.
In accordance with our theme of consistency across levels of interpretation, Sirois and
Mareschal (2002) argued that models and theories of habituation should reflect the
two key neural mechanisms that support infant habituation. First, the hippocampus is
involved in selective inhibition of high-order features such as color or shape (or
feature relations, such as ‘color+shape’) of the habitual stimuli (Sokolov &
Vinogradova, 1975; Nelson, 1995). Hippocampal inhibition is short-lived. Hence,
given that habituation has lasting effects (Zelazo et al., 1991), the second neural
mechanism of habituation involves long-term storage in hippocampal-related cortical
areas, notably the entorhinal cortex (Nelson, 1995).

In the HAB model of infant habituation (Sirois and Mareschal, 2004), hippocampal
and cortical functions are both implemented by simple autoassociator networks (see
Sirois, 2004); however, they use different learning rules to implement selective
inhibition and long-term storage. Both networks are coupled through reciprocal
connections and both contribute to the overall output of the system. Embedding the
model in a robot and an environment illustrated e how motor learning contributes to
habituation performance (Sirois, 2005).

The HAB model illustrates the main principles of the neuroconstructivist approach in
the following ways. First, learning is driven by context, as each subsystem learns
within the feedback loop of the other, antagonist subsystem. Moreover, as the robot
work illustrated, habituation can be affected by motor learning. Behavior involves a
mixture of cooperation (within subsystems, but also between them when their outputs
are aggregated at the system level) and competition (the hippocampus attempts to shut
down known input signals, whereas the cortex attempts to amplify them). Moreover,
chronotopy was shown by the model’s ability to capture age-related changes in
performance through maturation of outward connections from the cortex (Sirois &
Mareschal, 2004), as observed in infant brains and in absence of prior experience. The
model is proactive, as it seeks maximally stimulating input (Sirois, 2005), and
exhibits progressive specialization as it shifts from an initial familiarity preference to
a novelty preference once known inputs are well learned. The outcome is partial
representations, as the behavior of the model is achieved through the activity of
several interconnected units in two distinct subsystems.

3.3. Phonological development

Infant babbling, the repetition of simple speech sounds, creates a coupling between
the perception and production of language. This view is relatively recent, as early
work on phonological development proposed no such connection between babbling
and speech (Jakobson, 1941; Lenneberg, 1967).

13
In the first year of life, there are substantial changes to the perception of speech
sounds. The best known is how infants, who are initially able to discriminate speech
sounds from any human language (Eimas et al., 1971), progressively lose the ability
to discriminate phonemes from outside their native language in the second half of the
first year (Werker & Tees, 1984). The importance of the linguistic environment is
further stressed by data showing that infants’ speech discrimination ability is
correlated with the clarity of their mother’s speech (Liu et al., 2003).

Changes in speech-sound production can be observed in infant articulation. Before six

months, most speech sounds consist of isolated vowels. However, around six months,
most infants begin to babble. Articulation becomes progressively more complex over
the next few months, and babbling becomes more specific to the infant’s native
language (Boysson-Bardies et al., 1989). This is arguably a key step towards the
development of a phonological inventory, used for words and subsequently more
complex linguistic structures (Vihman, 2002).

The central role of auditory perception for babbling has been emphasized by research
on deaf infants (Oller & Eilers, 1988), who babble later than hearing infants and
produce different sounds. These effects are long lasting and can negatively affect later
speech (Wallace et al., 1998). Auditory feedback is thus necessary for the successful
coordination of phonatory (larynx) and articulatory (vocal tract) speech systems,
essential for babbling and, subsequently, speech (Koopmans-van Beinum et al.,
2001).

Westermann and Miranda (2004) recently proposed a mechanistic model of the

development of the link between speech perception and production. The model
consists of two topographic maps, one each for articulation and perception. Within
these maps, neurons responded to inputs that fell within their respective fields. The
two maps were connected with Hebbian weights, such that units with high covariation
between maps saw their connections strengthened and connections for units with low
covariation were weakened. As a consequence, strongly covarying
articulations/perceptions became prototypical; these prototypes represented vowels
that could be most robustly produced based on articulatory parameters. The model’s
prototypical speech sounds reflected both internally generated activity and
environmental input (see Vihman, 1991, for a similar interpretation called the
Articulatory Filter Hypothesis).

Context is central to the emergence of speech sound prototypes in both maps.

Articulatory representations emerge from a dynamic interaction with perception and
vice versa; moreover, these reflect the context of both internally-generated and
external inputs. Each topographical map implements local competition through lateral
inhibition, but the coupling of articulation and perception requires the cooperation of
both maps. Chronotopy is also important, as both maps require a synchronization of
plasticity. Proactivity is vital, as babbling produces the coupling between perception
and production. Moreover, as this coupling develops, the representations make the

14
model progressively more selective with respect to environmental input. In real
infants, in a real linguistic environment, this would lead to enhanced performance in
the native language, at the expense of discrimination abilities for other languages (see
also McClelland et al., 2002). Ultimately, this progressive specialization within and
between maps leads to partial representations: speech sounds become activations
patterns on both maps, and these cannot be isolated from one another.

3.4. Object representations in infants

Ungerleider and Mishkin (1982) proposed that object processing involves two
separate cortical information processing pathways: the dorsal and ventral streams. It
was proposed that the dorsal stream, terminating in the parietal cortex, processes
object localization (the where function), whereas the ventral system, ending in the
temporal cortex, performs object identification (the what function). The actual degree
of independence of these two streams has recently been the object of some attention
(Fellman & van Essen, 1991; Humphreys & Riddoch, 2003; Merigan & Maunsell,
1993; Puce et al., 1998). Our interest, though, is that both streams process different
types of information and, as such, develop distinct representations.

The fact that object representations are, to a large degree, segregated in functionally
distinct routes, implies that there needs to be a mechanism that integrates these two
sources of information when they are required by some task (such as, for example,
picking up a specific object from several alternatives). Mareschal, Plunkett and Harris
(1999) proposed a model that examines how these two streams of information may be
gradually integrated over development, explaining why successful object retrieval by
infants lags behind successful visual tracking. A key assumption of the model is that
both routes (ventral and dorsal) are exposed to the same input but differ in their
associative learning mechanisms. The object recognition network (ventral stream)
generates a spatially invariant representation of objects, using an unsupervised
competitive learning rule (Foldiak, 1991). The trajectory prediction network (dorsal
stream) uses a partially recurrent feedforward network to track the immediately
anticipated retinal position of moving objects, a proactive process. The response
integration network in Mareschal et al. (1999) represents a measure of infants’
abilities to coordinate and use information about the positions and identities of
objects, analogous to a similar prefrontal cortical function observed in primates (Rao,
Rainer, & Miller, 1997).

Interestingly, young infants show some unusual behaviors when objects are briefly
occluded. For instance, infants can remember spatial properties of occluded objects
but not necessarily identity features (Leslie et al., 1998; Kaldy & Sigala, 2004; Simon
et al., 1995; Wilcox & Schweinle, 2002; Xu & Carey, 1996). Object individuation at
4.5 months relies on shape and size, at 7.5 months on texture, and only at 11.5 months
does it involve color (Wilcox, 1999). Mareschal and Johnson (2003) examined under
which conditions four-month-olds would retain position or identity information
during a five second occlusion. They found that the functional value of objects (their
affordance, in Gibsonian terms) appears to drive a competition between dorsal and

15
ventral streams for object retention. Objects that afford the possibility of actions
maintain dorsally processed information, whereas objects that do not afford action
maintain ventral information.

This and other evidence highlights the contextual nature of object processing,
involving the child, the environment, the affordance of objects and functionally
distinct neural representations. Competition and cooperation occur at different levels
in the distributed system involved in object recognition. The system also exhibits
progressive specialization, in that each stream excludes irrelevant sources of
information (spatial or featural) to carry out its function. Ultimately, the infant brain
must coordinate partial representations in distinct systems to act on specific objects
(Mareschal et al., 1999; Rao et al., 1997).

3.5. Ensocialment

The importance of the social context on cognitive development has a long history
(e.g., Bandura, 1986; Rogoff, 1990; Vygotsky, 1978). Our focus is on the developing
child situated in an environment that includes other humans and in which
development involves a collaboration between the child and those who support and
nurture this development (Rogoff, 2003). While these ideas are not new, it is only
more recently that the importance of social behavior has made forays into the
neurosciences (e.g., Adolphs, 2003).

An early aspect of social brain function is the preference of newborns (as early as
within the first hours after birth) for face-like stimuli (Johnson et al., 1991; Valenza et
al., 1996). Although the specific cues that elicit the preference remain a source of
debate (see Johnson, 2005, for review), it has been suggested that three high-contrast
blobs in the positions of the eyes and mouth may be sufficient (Johnson & Morton,
1991). Hence, the brain does not contain, from birth, a detailed specification of a face
but a skeletal, partial, representation. The preference for face-like stimulation makes
the infant pro-active in seeking stimulation with faces, which places the infant in a
learning context in which other cortical systems will learn about faces. Hence, an
initial bias ensures that later developing areas of the cortex acquire specific
specialization for faces (Johnson 2005). Similarly, evidence shows that infants prefer
to look at faces that show direct gaze towards them (Farroni et al., 2002). Maintaining
eye contact with someone ensures foveation of the face, which may prove essential to
the emergence of a cortical face area (Johnson, 2004).

Children benefit from “ensocialment” in several other ways. For example, Vygotsky
(1978) was probably the first to fully recognize the role of language in shaping
cognitive development. Vygotsky argued that, cognitive change involved moving
from external speech, which instructs the child, to private speech, whereby the child
maintains an internal dialog that takes over from external instruction to guide
behavior. Indeed, Berk and Gavin (1984) observed that most vocalized private speech
in group settings actually takes place when a child is working alone on a difficult task.

16
Vygotsky (1978) further proposed that in relation to the child’s current level of
development, there existed a small window of optimal stimulation within which to
provoke further development: the Zone of Proximal Development (ZPD). Infants
display this spontaneously by exhibiting a preference for stimuli that are moderately
discrepant from their current knowledge or capabilities (McCall, Kennedy, &
Applebaum, 1977). Teaching below this zone provides little enhancement as it is
within the child’s current grasp. Similarly, teaching above this zone would also
provide few gains as the child would fail to see the path between her current level of
competence and the teaching. Thus, Rogoff (1990) proposed the notion of guided
participation, whereby progress is optimal when child and teacher share a focus and
purpose in learning. The main idea in terms of instruction (and, generally, pedagogy)
is that it channels the child’s interaction with the environment (see also Csibra &
Gergely, 2006). The emphasis on joint participation once again highlights the
importance of proactivity from the child.

3.6. Atypical development

In most cases, the outcome of development is relatively predictable. However, how

can we explain variability in developmental outcome? Some variability is observed in
intelligence, much more in cases of developmental disorders. Developmental
disorders can have several causes. Disorders can stem from genetic abnormalities,
such as in Down syndrome, Williams syndrome, and Fragile X. They can be
identified on behavioral grounds, such as autism, Specific Language Impairment
(SLI), Attention Deficit Hyperactivity Disorder (ADHD), and dyslexia. In these latter
cases, some genetic influence is suspected as these conditions can run in families but
the genetic basis is not fully understood. Finally, disorders can be caused by atypical
environments, either biochemical (e.g., mothers taking drugs during pregnancy) or
psychological (e.g., cases of deprivation or abuse). Notably, some developmental
disorders can exhibit very uneven cognitive profiles. For example, there may be
particular problems in language but less so in non-verbal areas (e.g., SLI). Some
abilities appear relatively stronger against a background of low IQ (e.g., face
recognition in Williams syndrome). How should we explain these uneven profiles?

Where uneven cognitive profiles are observed in typical adults who have experienced
brain damage, the usual recourse is to infer that certain parts of the adult cognitive
structure have sustained damage. Some researchers have attempted to apply this
explanatory framework to developmental disorders, inferring that isolated cognitive
components have failed to develop, while the rest of the cognitive system has
developed typically. Examples include a Theory of Mind module in autism (Baron-
Cohen, 1999; Baron-Cohen et al., 1993) and a syntax module in SLI (van der Lely,
1997). Where the disorder has a genetic basis, there has been a further temptation to
view uneven cognitive profiles as evidence for direct links between genes and
particular cognitive mechanisms.

Explaining developmental deficits with reference to the typical adult cognitive system
is, however, problematic. The adult structure is not pre-specified but is itself a product

17
of development. Yet strong analogies with adult brain damage produce accounts of
developmental deficits with no role for development at all. In contrast empirical
evidence supports the role of development in producing atypical cognitive profiles.
When Paterson et al. (1999) explored the language and number abilities of toddlers
with Down syndrome and Williams syndrome, they found a different relative pattern
to that observed in adults with these disorders. The profile in early childhood was not
a miniature version of the adult profile.

The influence of genetic variation and genetic mutation on brain development is not
yet fully understood. However, current data suggests that these effects are typically
graded and diffuse, and have not been found to co-occur with the regions of the cortex
associated with specialised higher cognitive functions in typical adults. For example,
the British KE family were initially identified as having a specific speech and
language deficit caused by mutation to a single gene (FOXP2). However, subsequent
research revealed that there were widespread structural and functional brain
differences in family members possessing the mutation, while cognitive deficits
extended outside the domain of language to negatively affect, for example,
performance on non-verbal, rapid associative learning tasks (Watkins et al., 2002;
Watkins, Dronkers & Vargha-Khadem, 2002). In a comparison of a number of genetic
syndromes, Kaufmann and Moser (2000) confirmed that diffuse effects on brain
development are the norm.

The neuroconstructivist approach places the developmental process at the heart of

explanations of developmental disorders (Karmiloff-Smith, 1998). Empirically, the
framework encourages researchers to focus on trajectories of development rather than
static snapshots. Theoretically, disorders are viewed as cases of atypically constrained
trajectories. A disordered system is still adaptive, yet it may not possess the
neurocomputational constraints that are appropriate to acquire a domain. In some
circumstances, apparently typical behaviors may be generated by atypical underlying
processes (see, e.g., Deruelle et al., 1999; Karmiloff-Smith et al., 2004, for work on
face recognition in Williams syndrome). In other cases, the atypical constraints may
produce better than typical performance in a domain, such as in some aspects of
perception in autism. In Waddington’s (1957) metaphor, the epigenetic landscape has
changed.

Several of the core ideas of neuroconstructivism are emphasized by the study of

atypical development. For example, in some cases interactive specialization of
cortical areas appears atypical. Adults with Williams syndrome exhibit face
recognition skills in the typical range, but examination of ERPs reveal reduced
evidence of specialization and localization of neural activity (e.g., Grice et al., 2001,
2003). Neuroimaging data have suggested differences in the constraints of
chronotypy, in terms of the changes in connectivity (and associated plasticity) over
time in disorders such as autism and Down syndrome (e.g., Becker et al., 1986;
Chugani et al., 1999).

18
Differences in input encoding have been proposed to have cascading effects on the
context in which other cognitive abilities are acquired (e.g., in autism, SLI, and
dyslexia). Alterations in the level of abstraction achieved in forming internal
representations, or in the dimensions of similarity that those representations encode,
can play a material role in the ability of other brain systems to employ this
information to drive other processes. It is proposed that in autism, SLI, and dyslexia,
for example, the consequence of atypical similarity structure in the input
representations results in a processing deficit much higher up in a hierarchy of
representational systems. Differences in embodiment may also impact on the
trajectory of development. For example, Sieratzki and Woll (1994) proposed that in
children with spinal muscular atrophy, a disorder that reduces early mobility,
language development might be accelerated as a compensatory way for the young
child to control their environment. Lastly, an atypical child co-specifies an atypical
social environment, for example, in the expectations and reactions of parents and
peers, which has also been observed to influence these children’s development.

Of course, when we place an emphasis on development as a trajectory, and atypical

development as an atypically constrained trajectory, it becomes increasingly
important to specify what is different about the constraints and mechanisms of
change. Here computational modeling has offered exciting avenues for progress in the
study of disorders (e.g., disorders of infant gaze perception: Triesch et al., 2006;
disorders of language: Thomas & Karmiloff-Smith, 2003).

3.7. Dyslexia

Reading is a relatively recent human invention, going back only a few thousand years.
It is highly unlikely that evolution has produced domain-specific constraints on the
cognitive systems involved (McCandliss, Cohen, & Dehaene, 2003). The existence of
a specialised reading system in a brain must therefore represent an adaptation of more
general functions to reflect the specific environmental demands of cultures that
encourage expertise in this area.

Neuroimaging suggests that reading recruits up to a dozen distinct brain areas

(Dehaene, 2003). One particular area, the visual word form area, shows comparably
high levels of activity for words and pseudowords (seemingly plausible nonwords
made up of standard letters), but not for illegal letter symbols, suggesting an area that
processes visual stimuli that look like words, prior to establishing whether they have a
meaning (Dehaene et al., 2002; Posner et al., 1988). The location of this area is
relatively consistent across individuals and languages (and thus scripts). Because it
couldn’t have been selected by evolution, this area must be one that happens to have
appropriate computational properties for processing words; i.e., fine foveal
discrimination and invariant recognition of letter- and word-sized stimuli (McCandliss
et al., 2003).

The emergence of reading requires a mapping to be established between the linguistic

phonological discriminations of the pre-literate child and the letters or letter clusters

19
he or she must learn: i.e., the child must learn the relation between graphemes (new)
and phonemes (old). Some languages, such as English and French, have inconsistent
mappings between graphemes and phonemes (e.g., the letter ‘i’ sounds different in the
English words bit and bite). Compared to languages with consistent mappings (e.g.,
Italian and Spanish), inconsistent mappings can delay proficient grapheme-phoneme
decoding by up to two years (Goswami, 2002).

Reading disabilities in English affect between 5 and 17 percent of school aged

children (Shaywitz & Shaywitz, 1994). Two main subtypes of developmental dyslexia
have been proposed (Castles & Coltheart, 1993; Manis et al., 1996). Phonological
developmental dyslexia involves difficulties reading novel or pseudowords, whereas
surface developmental dyslexia, describes the difficulty in reading irregular words for
which the pronunciation cannot be predicted from the sounds of the individual letters
(words such as aisle or yacht).

Twin studies have suggested a significant genetic contribution to developmental

dyslexia (Pennington, 1999; Plomin & Dale, 2000; Plomin & Rutter, 1998). However,
a direct mapping of gene to cognitive function is highly unlikely. Therefore, one
might expect to observe more widespread effects than dyslexia alone if there is a
genetic aetiology. Although there is no consensus and substantial variability in the
literature, especially for surface dyslexia, various concurrent cognitive deficits have
been observed in individuals with dyslexia, suggesting a more general sensorimotor
syndrome (Stein & Walsh, 1997).

Most computational models of typical and atypical reading development assume that
the problem is learning to map representational codes for written words, spoken
words, and word meaning (e.g., Harm & Seidenberg, 2004; Plaut et al., 1996;
Seidenberg & McClelland, 1989). Surface dyslexia has been simulated by alteration
of initial constraints reducing the ability to learn the mapping between orthography
and phonology, in some cases through the semantic route. Phonological dyslexia has
been simulated in two major ways. The first method is to degrade the properties of the
phonological representations (e.g., Harm & Seidenberg, 2004). The second method
degrades the ability to learn the mapping between orthography and phonology. It is
interesting that similar manipulations can produce surface or phonological dyslexia,
as many people with either dyslexia show symptoms common to both types (Manis et
al., 1996).

Developmental dyslexia serves to illustrate the principles of neuroconstructivism in

the following ways. Reading is a specialisation of a more general system, reflecting
the context of a particular environment. In typical development, reading involves
dynamic interactions between multiple functions with suitable computational
properties. Partial representations interact during development, as highlighted by the
restructuring of phonology when the mapping with graphemes is learned. In
developmental dyslexia, the somewhat heterogeneous clusters of difficulties reflect
the different ways crucial mappings between phonology, orthography, and semantics
can be disrupted, as illustrated by computational work. This is consistent with the

20
suggestion of a genetic aetiology, which would be unlikely to selectively disrupt a
function that evolution cannot have selected in the first place. Rather, genetic
anomalies are expected to show more general, diffuse effects with differential rather
than specific effects in particular domains.

4. Conclusions

Our framework has many implications for developmental theory. One major
implication is that our proposal of multi-level isomorphism is a radical rejection of
Marr (1982). Cognition cannot be studied independently of the brain and body.
Another important implication is the central role of developmental trajectories in the
interpretation of adult cognition. There is no teleology involved in development;
mature, normative cognition is an outcome of development, not a pre-specified target
(Thomas & Karmiloff-Smith, 2003).

We have also argued that brain regions do not utilize complete representations. In
support of this view, we discussed, firstly, how a particular input signal is typically
processed separately along distinct dimensions (e.g., shape, color, motion in object
perception). Secondly, functional brain systems are interrelated with other functional
systems, and are located and within a body and an environment. All these contexts
provide varying levels and sources of information such that only fragments of
information need to be represented. An implication is that cognition will be
compositional and systematic (e.g., Fodor, 1975; Fodor & Pylyshyn, 1988) only if the
context permits it.

Progressive specialization, and notions such as emergent modularity (e.g., Karmiloff-

Smith, 1998) imply that, as a result of development, the cognitive architecture will
exhibit a progressive lack of flexibility to the novel. This can be observed when
emergent specialized systems have well-delineated functions (Johnson & Munakata,
2005). These specialized systems should not be construed as mere imprinting from
environmental pressures and regularities. The child, from birth (e.g., Robertson et al.,
2004), is an active contributor to his or her development. Our emphasis on context-
dependence may be taken as implying that it is impossible to make general claims
about cognition. This is not the case. Instead, we argue that the key to understanding
contextualised function is to identify those contexts that are central to the function of
interest, while ignoring those contexts that may have a peripheral rather than central
role in determining the function of interest.

A recurrent theme in our book is that we need to have causal theories of what makes
complex behaviors emerge. These theories need to explain behaviors at multiple time
scales. They must explain how and why behaviors unfold as we observe them in real
time, as well as how they unfold in developmental time. To do this, we need more
than just a very detailed description of the behaviors that can be observed at any point
in time. Certainly, such descriptions are essential for the advancement of causal
theories of development, but they are unsatisfactory to the extent that they are unable
to explain or predict new behaviors. The clearest example of this is with regards to

21
explaining the behavior of children with developmental disorders. Unless one has a
mechanistic theory of what is causing behaviors to unfold, and a causal theory of what
is atypical in such children’s processing, it is impossible to explain or predict why one
set of atypically developing children may show a delay at one behavior and excel at
another, while a second set of atypically developing children will excel at the former
behavior but have a developmental delay in the latter behavior.

In the companion volume, Neuroconstructivism Volume2: Perspectives and Prospects

(Mareschal et al. 2007b), we invited nine different research labs with objectives
broadly consistent what the Neuroconstructivist approach to present their
computational modeling work. The questions that the models are built to investigate
differ both in the level of description and in time scale over which the relevant
behaviors operate. Some models focus on relatively rapid adaptation occurring
(perhaps) at the cellular level of description, whereas other models focus on relatively
slow adaptation occurring at the cognitive level.

We asked all contributors to the companion volume to emphasize the following

aspects of their contributions when describing their work:

1. What functional brain constraints operate on the process of representation

development?
2. What embodiment or situatedness constraints operate on the process of
representation development?

Ultimately, computational models are tools to help us reflect on questions of process

and mechanisms. Therefore, we also asked the contributing authors to answer the
following questions:

3. How does the model embody these constraints?

4. What concrete predictions does the model make?

In Neurocontructivism: How the Brain Constructs Cognition we set out to investigate

how the representations that underlie cognition emerge in the brain during
development. We argued that the emergence of such representations is the outcome of
a constructivist process involving constraints that operate at all levels from the
cellular environment to the social environment. To truly understand how these
representations emerge, it is necessary to locate our theories at the point where the
constraints of brain, body and environment come together.

Acknowledgements

The writing of this paper was supported by EC grants 516542(NEST) and 029088
(NEST), ESRC grant RES-061-23-0129, MRC grants G9715587 and G0300188 and
British Academy grant SG-40400

22
References

Ballard, DH, Hayhoe, MM, Pook, PK & Rao, RPN (1997) Deictic codes for the
embodiment of cognition. Behavioral and Brain Sciences, 20, 723-767

Bandura A (1986) The social foundations of thought and action. Engelwood Cliffs,
NJ: Prentice-Hall.

Baron-Cohen, S, Tager-Flusberg, H and Cohen, HJ Eds. (1993). Understanding

Other Minds: Perspectives from Autism. Oxford: Oxford University Press.

Baron-Cohen, S (1999) Does the study of autism justify minimalist innate

modularity? Learning and Individual Differences, 10, 179-191.

Bogartz, RS, Shinskey, JL & Speaker, CJ (1997). Interpreting infant looking: The
event set * event set design. Developmental Psychology, 33, 408-422.

Campos, JJ, Anderson, DI, Barbu-Roth, MA, et al. (2000). Travel broadens the mind.
Infancy, 1, 149-219.

Cashon, CH & Cohen, LB (2000). Eight-month-old infants' perceptions of possible

and impossible events. Infancy, 1, 429-446.

Castles, A & Coltheart, M (1993). Varieties of developmental dyslexia. Cognition, 47,

149-180.

Clahsen, H & Almazan, M (1998). Syntax and morphology in Williams Syndrome.

Cognition, 68, 167-198.

Clark, A (1997). Being There, MIT Press: Cambridge.

Clark, A and Thornton, C (1997). Trading spaces: computation, representation and the
limits of uninformed learning. Behavioural and Brain Sciences, 20, 57–66.

Clifton, RK & Nelson, MN (1976). Developmental study of habituation in infants: the

importance of paradigm, response system, and state. In TJ Tighe & RN Leaton (Eds.),
Habituation: Perperspectives from child development, animal behavior and
neurophysiology. Hillsdale: Erlbaum. p. 159-205.

Cohen, LB (1972). Attention-getting and attention-holding process of infant visual

preferences. Child Development, 43, 869-879.

Cohen, LB & Cashon, CH (2003). Infant perception and cognition. In R. Lerner, M.

Easterbrooks, M. Ann et al. (eds.) Handbook of psychology: Developmental
psychology, Vol. 6. New York, NY, US: John Wiley & Sons, Inc. pp. 65-89.

23
Cohen, LB & Marks, KS (2002). How infants process addition and subtraction events.
Developmental Science, 5, 186-201.

Crowley, JC and Katz, LC (1999). Development of ocular dominance columns in the

absence of retinal input. Nature Neuroscience, 2, 1125-1130.

Csibra, G & Gergely, G (2006). Social learning and social cogniton: The case for
pedagogy. In M. H. Johnson & Y. Munakata (Eds.), Processes of Change in Brain
and Cognitive Development. Attention and Performance XXI. Oxford: Oxford
University Press.

Dehaene, S (2003). Natural born readers. New Scientist, 5th July 2003, no. 2402, 30-
33.

Dehaene, S, Le Clec’H, Poline, J-P, Le Bihan, D & Cohen, L (2002). The visual word
form area: A prelexical representation of visual words in the fusiform gyrus.
NeuroReport, 13, 321-325.

Desimone, R (1996). Neural mechanisms for visual memory and their role in
attention. Proceedings of the National Academy of Sciences USA, 93, 13494–13499.

Desimone, R and Duncan, J (1995). Neural mechanisms of selective visual attention.

Annual Review of Neuroscience, 18, 193–222.

Diamond, A (1991) Frontal lobe involvement in cognitive changes during the first
year of life, In KR Gibson & AC Petersen (Eds.), Brain Maturation and Cognitive
Development: A Comparative and Cross-cultural Perspective pp. 127-180. New
York: Aldine de Gruyter.

Driver, J, Davis, G, Russell, C, Turatto, M, and Freeman, E (2001). Segmentation,

attention and phenomenal visual objects. Cognition, 80, 61–95.

Eimas, PD, Siqueland, ER, Jusczyk, P & Vigorito, J (1971). Speech perception in
infants. Science, 171, 303-306.

Elman, J, Bates, E, Johnson, MH, Karmiloff-Smith, A, Parisi, D, & Plunkett, K

(1996) Rethinking Innateness: A connectionist perspective on development.
Cambridge, MA: MIT Press.

Fantz, RL (1964). Visual experience in infants: Decreased attention familar patterns

relative to novel ones. Science, 146, 668-670.

Farroni, T, Csibra, G, Simion, F & Johnson, MH (2002). Eye contact detection in

humans from birth. Proceedings of the National Academy of Sciences, 99, 9602-9605

24
Felleman, DJ and Van Essen, DC (1991). Distributed hierachical processing in the
primate cerebral cortex. Cerebral Cortex, 1, 1-47.

Fodor, JA (1975). The Language of Thought, Cambridge, Massachusetts: Harvard

University Press.

Fodor, JA, and Pylyshyn, Z (1988) Connectionism and cognitive architecture: A

critique. Cognition, 28, 3-71. .

Foldiak, P (199l). Learning invariance from transformation sequences. Neural

Computation, 3, 194-200.

Ghosh, A, Carnahan, J and Greenberg, ME (1994). Requirement for BDNF in

activity-dependent survival of cortical neurons. Science, 263, 1618-1623.

Gibson, EJ (1979). The ecological approach to visual perception, Houghton Mifflin:

Boston.

Gibson, EJ (1982) The Concept of Affordances in Development: The Renascence of

Functionalism. In Collins, W. A., (Ed.), The Concept of Development. The Minnesota
Symposia On Child Psychology, pp. 55-82. Hillsdale, N.J.: Lawrence Erlbaum.

Goldfield, EC, Kay, B & Warren, W (1993). Infant bouncing: The assembly and
tuning of an action system. Child Development, 64, 1128-1142.

Goldstein, DG & Gigerenzer, G (2002). Models of Ecological Rationality: The

Recognition Heuristic. Psychological Review, 109, 75-90

Goswami, U (2002). Phonology, reading development and dyslexia: A cross-

linguistic perspective. Annals of Dyslexia, 52, 1-23.

Goswami, U (2003). Phonology, learning to read and dyslexia: A cross-linguistic

analysis. In V Csepe (Ed.), Dyslexia: Different Brain, Different Behavior, NL: Kluwer
Academic. pp. 1-40.

Gotltlieb, G. (2007) Probabilistic Epigenesis. Developmental Science, 10, 1-11.

Haith, MM (1998). Who put the cog in infant cognition? Is rich interpretation too
costly? Infant Behavior & Development, 21, 167-179.

Harm, MW & Seidenberg, MS (2004). Computing the meaning of words in reading:

Cooperative division of labor between visual and phonological processes.
Psychological Review, 111, 662-720.

Hubel, DH and Wiesel, TN (1963). Shape and arrangement of columns in cat’s striate
cortex. Journal of Physiology, 165, 559–568.

25
Humphreys, GW & Riddoch, M (2003). From what to where. Psychological Science,
14, 487-492.

Hutchins, E (1995) Cognition in the Wild. Cambridge, MA: MIT press.

Jakobson, R (1941). Child Language, Aphasia and Phonological Universals. The

Hague: Mouton. English translation 1968.

Johnson, MH (2005) Developmental cognitive neuroscience (2nd Edition). Oxford,

UK: Blackwells

Johnson, MH (2004) Plasticity and functional brain development: The case of face
processing. In Kanwisher, N. and Duncan, J., (Eds.), Attention & Performance XX:
Functional neuroimaging of visual cognition, pp. 257- 263: Oxford University Press.

Johnson, MH, Dziurawiec, S, Ellis, HD and Morton, J (1991). Newborns' preferential

tracking of face-like stimuli and its subsequent decline. Cognition, 40, 1-19.

Johnson, M.H. and Munakata, Y. (2005). Processes of change in brain and cognitive
development. Trends in Cognitive Sciences, 9, 152-158.

Johnson, MH and Vecera, SP (1996). Cortical differentiation and neurocognitive

development: the parcellation conjecture. Behavioural Processes, 36, 195–212.

Kaldy, Z & Sigala, N (2004) The neural mechanisms of object working memory:
what is where in the infant brain? Neuroscience and Biobehavioural Reviews, 28,
113-121.

Karmiloff-Smith, A (1998) Development itself is the key to understanding

developmental disorders. Trends in Cognitive Sciences, 2, 389 - 398.

Karmiloff-Smith, A (1992) Beyond modularity: A developmental perspective on

cognitive science. Cambridge, MA: MIT Press.

Karni, A, Meyer, G, Jezzard, P, Adams, MM, Turner, R & Ungerleider, LG (1995)

Functional MRI Evidence for Adult Motor Cortex Plasticity During Motor Skill
Learning. Nature, 377, 155-158.

Koopmans-van Beinum, FJ, Clement, CJ & van den Dikkenberg-Pot, I (2001)

Babbling and the lack of auditory speech perception: a matter of coordination?
Developmental Science, 4, 61-70.

Lamme, VAF & Roelfsema, PR (2000). The distinct modes of vision offered by
feedforward and recurrent processing. Trends in Neurosciences, 23, 571–579.

26
Lenneberg, E (1967). Biological Foundations of Language. New York: Wiley.

Leslie, A, Xu, F, Tremoulet, P & Scholl, B (1998). Indexing and the object concept:
"what" and "where" systems in infancy. Trends in Cognitive Sciences, 2, 10-18.

Liu, H-M, Kuhl, P & Tsao, F-M (2003). An association between mothers' speech
clarity and infants' speechdiscrimination skills. Developmental Science, 6, F1-F10.

Manis, FR, Seidenberg, MS, Doi, LM, McBride-Chang, C &, Petersen, A (1996). On
the bases of two subtypes of developmental dyslexia. Cognition, 58, 157-195.

Mareschal, D & Bremner A J (2005). When do 4-month-olds remember the “what”

and “where” of hidden objects? In MH Johnson & Y Munakata (eds.) Attention &
Performance XXI: Processes of Change in Brain and Cognitive Development. Oxford,
UK: Oxford University Press.

Mareschal, D., Johnson, M.H., Sirois, S., Spratling, M., Thomas, M. & Westermann,
G. (2007a). Neuroconstructivism, Vol. I: How the brain constructs cognition. Oxford,
UK: Oxford University Press

Mareschal, D and Johnson, MH (2003). The “What” and “Where” of infant object
representations. Cognition, 88, 259-276.

Mareschal, D and Shultz, TR (1996). Generative connectionist architectures and

constructivist cognitive development. Cognitive Development, 11, 571-605.

Mareschal, D., Sirois, S., & Westermann, G. (2007b). Neuroconstructivism, Vol. II:
Perspectives and Prospects. Oxford, UK: Oxford University Press.

Maris, M & te Boekhorst, R (1996). Exploiting physical constraints: heap formation

through behavioural error in a group of robots. In Proc. IROS’96, IEEE/RSJ
International Conference on Intelligent Robots and Systems.

Marr, D (1982). Vision. San Francisco: W. Freeman.

McCall, RB, Kennedy, CB and Applebaum, MI (1977). Magnitude of discrepancy

and the distribution of attention in infants. Child Development, 48, 772-786.

McCandliss, BD, Cohen, L & Dehaene, S (2003). The Visual Word Form Area:
Expertise for reading in the fusiform gyrus. Trends in Cognitive Sciences, 7, 293-299.

McClelland, JL, Fieza, JA, & McCandliss, BD (2002). Teaching the /r/-/l/
discrimination to Japanese adults: behavioral and neural aspects. Physiology &
Behavior, 77, 657-662.

27
Merigan, WH and Maunsell, JHR (1993). How parallel are the primate visual
pathways. Annual Review of Neuroscience, 16, 369-402.

Morton, J and Johnson, MH (1991). CONSPEC and CONLERN: A two-process

theory of infant face recognition. Psychological Review, 98, 164-181.

Nelson, CA (1995). The ontogeny of human memory: A cognitive neuroscience

perspective. Developmental Psychology, 31, 723-738.

Oliver, A, Johnson, MH, Karmiloff-Smith, A, & Pennington, B (2000). Deviations

in the emergence of representations: A neuroconstructivist framework for analysing
developmental disorders. Developmental Science, 3, 1-23.

Oller, DK & Eilers, RE (1988). The role of audition in infant babbling. Child
Development, 59, 441-449.

Pennington, BF (1999). Dyslexia as a neurodevelopmental disorder. In H Tager-

Flusberg (Ed.), Neurodevelopmental disorders, pp. 307-330. Cambridge, Mass.: MIT
Press.

Petersen, SE, Van Mier, H, Fiez, JA, and Raichle, ME (1998). The effects of practice
on the functional anatomy of task performance. Proceedings of the National Academy
of Sciences USA, 95, 853–860.

Piaget, J (1970) Genetic epistemology (translated by Eleanor Duckworth), New York:

Columbia University Press.

Piaget, J (1952). The Origins of Intelligence in the Child. New York: International
Universities Press.

Plaut, DC, McClelland, JL, Seidenberg, MS & Patterson, K (1996). Understanding

normal and impaired word reading: Computational principles in quasi-regular
domains. Psychological Review, 103, 56-115.

Plomin, R, & Dale, PS (2000). Genetics and early language development: A UK study
of twins. In D. V. M. Bishop & L. B. Leonard (Eds.), Speech and Language
Impairments in Children: Causes, Characteristics, Intervention and Outcome, pp. 35-
51. Hove, UK: Psychology Press.

Plomin, R & Rutter M (1998). Child development, molecular genetics, and what to do
with genes once they are found. Child Development, 69, 1221–1240.

Posner, MI (1993) Attention before and during the decade of the brain. In D Meyers
& S Kornblum (eds.) Synergies in experimental psychology, Artificial Intelligence,
and cognitive neuroscience: Vol XIV, Attention and Performance, pp. 343- 350.
Cambridge, MA: MIT Press.

28
Posner, MI. Petersen, SE, Fox. PT & Raichle, ME (1988). Localization of cognitive
functions in the human brain. Science, 240, 1627-1631.

Precht, HFR (2001) Prenatal and early postnaal development of human motor
behaviour. In AF Kalverboer, & A Gramsbergen (Eds.). Handbook of Brain and
Behaviour in Human Development, pp.415-427. Kluwer Academic Press.

Puce, A, Allison, T, Bentin, S, Gore, JC & McCarthy, G (1998) Temporal cortex

activation in humans viewing eye and mouth movements. Journal of Neuroscience,
18, 2188-2199.

Quinlan, PT (1988) Structural change and development in real and artificial neural
networks. Neural Network, 11, 577-599.

Rainer, G and Miller, EK (2000). Effects of visual experience on the representation of

objects in the prefrontal cortex. Neuron, 27, 179–89.

Robertson, SS (1988) Mechanism and function of cyclicity in spontaneous

movement. In WP Smotherman & SR Robinson (Eds.) Behavior of the Fetus, pp. 77-
94. Caldwell, NJ: Telford.

Robertson SS, Guckenheimer J, Masnick AM & Bacher LF (2004) The dynamics of

infant visual foraging, Developmental science, 7, 194-200.

Robertson, SS, Bacher, L. F & Huntington, NJ (2001) The integration of body

movement and attention in young infants. Psychological Science, 12, 523-526.

Rogoff, B (2003) The Cultural Nature of Human Development. New York: Oxford
University Press.

Rogoff, B (1998) Cognition as a Collaborative Process. In Damon, W., (Ed.),

Handbook of Child Psychology: Cognition, Perception and Language, pp. 679-744.
New York: John Wiley.

Rogoff, B (1990). Apprenticeship in Thinking, Cognitive Development in Social

Contexts, Oxford University Press: Oxford.

Salin, PA and Bullier, J (1995). Corticocortical connections in the visual-system:

structure and function. Physiological Reviews, 75, 107-154.

Schilling, TH (2000). Infants' looking at possible and impossible screen rotations: The
role of familiarization. Infancy, 1, 389-402.

Schlesinger, M (2004). Evolving agents as a metaphor for the developing child.

Developmental Science, 7, 154-168.

29
Schoner, G & Thelen, E. (In press) Using Dynamic Field Theory to Rethink Infant
Habituation. Psychological Review

Seidenberg, MS & McClelland, JL (1989). A distributed, developmental model of

word recognition and naming. Psychological Review, 96, 523-568.

Shaywitz, BA & Shaywitz, SE (1994). Learning disabilities and attention disorders. In

K. Swaiman (Ed.), Principles of pediatric neurology. St. Louis: Mosby. p. 1119-1151.

Sheridan, S. R. (1997). 1997 Drawing/Writing and the new literacy. Amherst, MA:
Drawing/Writing Publications.

Shrager, J and Johnson, MH (1996). Dynamic plasticity influences the emergence of

function in a simple cortical array. Neural Networks, 9, 1119–1129.

Shultz, TR (2003) Computational Developmental Psychology. Cambridge,

Massachusetts: MIT Press.

Simon, TJ, Hespos, SJ and Rochat, P (1995). Do infants understand simple

arithmetic? A replication of Wynn (1992). Cognitive Development, 10, 253-269.

Sirois, S (2004). Autoassociator networks and insights into infancy. Developmental

Science, 7, 133-140.

Sirois, S. (2005). Hebbian motor control in a robot-embedded model of habituation,

Proceedings of the International Joint Conference on Neural Networks (IJCNN 2005)
2772-2777: IEEE.

Sirois, S & Mareschal, D (2004) An Interacting Systems model of infant habituation.

Journal of Cognitive Neuroscience, 16, 1352-1362.

Sirois, S & Mareschal, D (2002) Models of infant habituation Trends in Cognitive

Sciences, 6, 293-298.

Slater, A, Morison, V, Somers, M, Mattock, A, Brown, E, & Taylor, D (1990).

Newborn and older infants’ perception of partly occluded objects. Infant Behavior
and Development, 13, 33-49.

Sloutsky, VM & Lo, Y-F (1999). How much does a shared name make things similar?
Part 1: Linguistic labels and the development of similarity judgement. Developmental
Psychology, 6, 1478-1492.

Sloutsky, VM, Lo, Y-F & Fisher, A (2001). How much does a shared name make
things similar? Linguistic Labels, Similarity and the Development of Inductive
Inference. Child Development, 72, 1695-1709.

30
Sokolov, EN & Vinogradova, OS (1975). Neuronal mechanisms of the orienting
reflex. Hillsdale, NJ: Erlbaum.

Spelke, ES, Breinlinger, K, Macomber, J & Jacobson, K (1992). Origins of

knowledge. Psychological Review, 99, 605-632.

Sporns, O, Tononi, G and Edelman, GM (2000). Theoretical neuroanatomy: relating

anatomical and functional connectivity in graphs and cortical connection matrices.
Cerebral Cortex, 10, 127-141.

Stein, J & Walsh, V (1997). To see but not to read: the magnocellular theory of
dyslexia. Trends in Neurosciences, 20, 147-152.

Thelen, E, Corbetta, D and Spencer, JP (1996). Development of reaching during the

first year: role of movement speed. Journal of Experimental Psychology: Human
Perception and Performance, 22, 1059–1076.

Thelen, E & Smith, LB (1994). A dynamic systems approach to the development of

cognition and action. Cambridge, MA: MIT Press.

Thomas, MSC & Karmiloff-Smith, A (2005). Can developmental disorders reveal

the component parts of the human language faculty? Language Learning and
Development, 1, 65-92.

Thomas, MSC & Karmiloff-Smith, A (2003a). Modelling language acquisition in

atypical phenotypes. Psychological Review, 110, 647-682.

Thompson, RF & Spencer, WA (1966). Habituation: A model phenomenon for the

study of neuronal substrates of behavior. Psychological Review, 73, 16-43.

Thorpe, WH (1956). Learning and instinct in animals. London: Methuen.

Tucker, M and Ellis, R (2001).The potentiation of grasp types during visual object
categorization. Visual Cognition, 8, 769-800.

Ungerleider, LG and Mishkin, M (1982). Two cortical visual systems. In Ingle, D J,

Goodale, MA, and Mansfield, RJW, (eds), Analysis of Visual Behavior, pages 549–
86. MIT Press, Cambridge, MA.

Valenza, E, Simion, F, Cassia, VM and Umilta, C (1996). Face preference at birth.

Journal of Experimental Psychology: Human Perception and Performance, 22, 892-
903.

Van der Lely, HKJ (1997). Language and cognitive development in a grammatical
SLI boy: Modularity and innateness. Journal of Neurolinguistics, 10, 75-107.

31
Van der Meer, ALH, Van der Weel, FR & Lee, DN (1995). The Functional
Significance of Arm Movements in Neonates. Science, 267, 693-695.

Van Essen, DC, Anderson, CH, & Felleman, D J (1992). Information processing in
the primate visual system: An integrated systems perspective. Science, 255, 419-423.

Vihman, MM (2002). The role of mirror neurons in the ontogeny of speech. In

[Link] & [Link] (Eds.), Mirror Neurons and the Evolution of Brain and
Language, Amsterdam: John Benjamins. pp. 305-314.

Vihman, MM (1991). Ontogeny of phonetic gestures. In I Mattingly & M Studdert-

Kennedy (Eds.), Modularity and the Motor Theory of Speech Perception, Hillsdale
NJ: Lawrence Erlbaum. pp. 69-84.

Vygotsky, L (1986). Thought and Language. Boston: MIT Press.

Vygotsky, LS (1978) Mind in Society. The developmental of higher psychological

processes. Harvard, MA: Harvard University Press.

Waddington, CH (1957). The Strategy of the Genes. London. UK: Allen and Unwin.

Waddington, CH (1953). Genetic assimilation of an acquired character. Evolution, 7,

118-126.

Walsh, V, Ashbridge, E, and Cowey, A (1998). Cortical plasticity in perceptual

learning demonstrated by transcranial magnetic stimulation. Neuropsychologia, 36,
363–367.

Watkins, KE, Dronkers, NF & Vargha-Khadem, F (2002). Behavioural analysis

of an inherited speech and language disorder: comparison with acquired aphasia.
Brain, 125, 452-464.

Watkins, KE, Vargha-Khadem, F, Ashburner, J et al. (2002). MRI analysis of an

inherited speech and language disorder: structural brain abnormalities. Brain, 125,
465-478.

Webb, B (1994) Robotic Experiments in Cricket Phonotaxis. In Cliff, D., Husbands,

P., Meyer, J.-A. and Wilson, S. W., (Eds.), From Animals to Animats 3: Proceedings
of the Third International Conference on the Simulation of Adaptive Behaviour
Brighton, pp. 45-54. Cambridge, MA: MIT Press.

Werker, J & Tees, R (1984).Cross-language speech-perception - evidence for

perceptual reorganization during the 1st year of life. Infant Behavior and
Development, 7, 49-63.

32
Westermann, G, & Mareschal, D (2003). Models of atypical development must also
be models of normal development. Behavioral and Brain Sciences, 25, 771-772

Westermann, G & Miranda, ER (2004). A new model of sensorimotor coupling in the

development of speech. Brain and Language. 89, 393-400.

Wilcox, T (1999). Object Individuation: Infants’ use of shape, size, pattern, and color.
Cognition, 72, 125-166.

Wilcox, T & Schweinle, A (2002). Object individuation and event mapping:

developmental changes in infants' use of featural information. Developmental Science,
5, 132-150.

Xu, F & Carey, S (1996). Infants' metaphysics: the case of numerical identity.
Cognitive Psychology, 30, 111-153.

Zelazo, PR, Weiss, MJS & Tarquinio, N (1991). Habituation and recovery of neonatal
orienting to auditory stimuli. In MJS Weiss & PR Zelazo (Eds.) Newborn attention:
Biological constraints and the influence of experience. Norwood, NJ: Ablex. pp. 120-
141.

33