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    PA RT I11

    rain Maturation and Cognition

    Brain Development and Cognition: A Reader, Second EditionMark H. Johnson, Yuko Munakata, Rick O. Gilmore

    Copyright 1993 and 2002 by Blackwell Publishers Ltd

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    Editors Introduction to Part

    Perhaps the most obvious way to relate brain development and cognition is to attribute th eonset of a certain cognitive ability to the maturation of underlying neural circuitry. Part I11presents two papers, by Johnson and Nelson, that take this kind of approach. Although thistype of argument has commonly been applied to the onset of an ability, it may also beapplied to the termination of an ability e.g., in sensitive periods, as discussed by Lorenz inPart I, Marler in Part V, and OReilly and Johnson in Part VI). Th e variations on thisgeneral claim usually take one of the following forms: a) sequences of brain maturat ion areused to predict the sequence of development of certain cognitive abilities as evidenced inthe paper by Johnson); b) specific neural developments at a certain age are posited to giverise to a specific computational advance at that same age as evidenced in the paper byNelson).

    T he paper by Johnson uses evidence about patt erns of postnatal brain growth to makepredictions about the sequence of development in visual attention and orienting. Johnsondiscusses both overt orienting head and eye movements that shift gaze) and covert shifts ofatten tion indepe ndent of head and eye movements). I n the case of overt orienting, Johnsonmakes three neuroanatomical observations: a) the primary visual cortex is the maingateway to several pathways that underlie components of visual attention and orienting;b) primary visual cortex, like other areas of cortex, has a layer-specific pattern of connect-

    ivity to these other neural struct ures and pathways; and c) some measures of postnatalcortical growth show a layer-specific pattern of development from deeper layers to moresuperficial ones as we saw in Part 11). Thes e facts support inferences regarding thedevelopment of cortical pathways, such as the earlier development of the visual pathwayinvolving the middle temporal area relative to the pathway involving the frontal eye fields.These pathways underlie particular components of visual orienting and attention, suppor t-ing predictions about sequences of development at the cognitive level, such as smoothvisual tracking preceding anticipatory eye movements. Johnson reviews a large amount ofevidence on the development of visual orienting in human infants to provide support forthe predicted sequences.

    In the case of covert shifts of attention, Johnson focuses on the effects of facilitationfaster responding to a stimulus appearing in a covertly attended location) and inhibition of

    return IOR, slower responding to a stimulus appearing in a covertly attended location).Facilitation generally occurs when a target stimulus appears soon after the offset of the cue

    Brain Development and Cognition: A Reader, Second EditionMark H. Johnson, Yuko Munakata, Rick O. Gilmore

    Copyright 1993 and 2002 by Blackwell Publishers Ltd

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    132 Brain Maturation and Cognition

    that shifts attention, whereas IOR occurs with longer latencies between cue and target.Covert shifts of atten tion appear to involve the parietal lobe, and IOR requires t he superiorcolliculus. Again, Johnson uses sequences of development of such neural structures topredict sequences of development at the behavioral level, in terms of facilitation and IOR,and reviews relevant evidence.

    Nelson takes the first of the two approaches outlined above, reviewing neural develop-ments posited to give rise to advances in distinct types of memory at particular ages. Nelsondiscusses many distinctions made in the memory literature, including explicit versusimplicit, procedural learning and conditioning, and working memory. He suggests thatthe early development of the striatum may support infants success in procedural learningtasks in the first few months of life, and the early development of the cerebellum maysupport conditioning in young infants. Explicit memory may emerge somewhat laterbetween 6 and 12 months) due to the development of temporal lobe structu res, following

    the development of a preexplicit memory dependent on the hippocampus. Workingmemory may also begin to develop within the 6-12 month period, due to developments

    in the prefrontal cortex see the Diamond reading in Part VII for a thorough discussion ofthe role of dopamine in this process). Nelson notes that all of these forms of memory maycontinue to develop for many months and years.

    Nelson emphasizes at the outse t that there are mul tiple caveats to exploring the relation

    between brain development and memory development. For example, because the brainworks as an integrated system, the brain s tructures outlined above do not subserve theirpurported functions alone, but rather as parts of complex circuits. As a result, theinterpretation of behaviors following lesions can be quite difficult. Further, caution mustbe taken in comparing the behaviors of human infants and adults and other species. Wenote that even within the infant literature, there is controversy over the interpretation of

    performance i n various tasks and the implications for memory development cf. Rovee-Colliers 1997) analysis of the development of implicit and explicit memory, Diamonds1991) work on the effects of hippocampal maturation, and Munakatas 1998) discussion of

    working memory development). Resolving these controversies may be a useful step inunderstanding distinct types of memory and their neural substrates.

    F U R T H E R R E A D I NG

    Atkinson, J. 1984). Human visual development over the first six months of life: A review and ahypothesis. Human Neurobiology 3: 61-74. A comprehensive review of the development ofvision and its neural basis.)

    Bachevalier, J. and Mishkin, M. 1984). An early and a late developing system for learning andretention in infant monkeys. Behavioral Neuroscience98: 770-8.

    Bates, E., Thal , D. , et al. 1992). Early language development and its neural correlates.Handbook o Neuropsychology. I. Rapin and S. Segalowitz eds.). Amsterdam: Elsevier. Vol6: Child Neurology. A review of neurodevelopmental correlates of language acquisition.)

    Bronson, G. W. 1974). The postnatal growth of visual capacity. Child Development45: 873-90.The now classic original statement proposing that the development of visually guided

    behavior in the human infant can be viewed in terms of a transition from subcortical tocortical processing.)

    Carey, S. 1980). Maturational factors in human development. Biological Studies o Menta l

    Processes. D. Caplan. Cambridge, MA: MI T Press: 1-7. An introduction to some of theissues about the relation between brain development and cognitive development.)

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    Brain Maturation and Cognition 133

    Carey, S. and D iamon d, R. 1980). Ma turational determination of the developmental course offace enc oding. Biological Stud ies of Menta l Processes.Cambridge, MA : M I T Press. Evidencefor a maturationally determined dip in face recognition abilities around the time ofpuberty.)

    Dehaene, S. and C hangeux, J. P. 1989). A simple mode l of prefrontal cortex function indelayed-response tasks.Jonrnal of Cognitive Neurosciencel 3): 244-61.

    Dia mo nd, A. 1991). Neuropsychological insights into the meaning of object concept develop-ment. The Epigenesis of Mind : Essays on Biology and Cognition.s. Carey and R. Gelman.Hillsdale, NJ: Law rence E rlbau m Associates: 67-1 10.

    Goldman-Rakic, P. S 1987). Development of cortical circuitry and cognitive function.ChildDevelopment58: 601-22. A review of the possible effects of prefrontal cortex matu ration oncognition.)

    Goldman-Rakic, P. S. and Isseroff, A ,, et al. 1983). T h e neurobiology of cognitive develop-ment. Handbook of Child Psychology: Biology and Infancy Development.P. Mussen ed.) . NewYork: Wiley: 281-334. A comprehensive overview of the relation between th e anatomicaldevelopment of the cortex, especially the prefrontal cortex, and some aspects of behavioral

    development.)He ld, R. 1985). Binocular vision: Behavioral and neuronal d evelopment.Neonate Cognition:

    Beyond the Blooming Buzzing Confusion.. M ehler and R. Fox eds.). Hillsdale, NJ: LawrenceErlbaum.

    Johnson, M . H . 1990). Cortical maturation and the development of visual attention in earlyinfancy. Jonrnal of Cognitive Neuroscience2 2): 81-95.

    Mu ir, D . W. and Clifton, R.K., et al. 1989). T h e development of a hum an auditory localizationresponse: A U-shaped function. Canadian 30nrnal of Psychology43: 199-216. A review ofexperiments indicating that auditory orienting in the human infant goes through a similarsubcortical to cortical shift to that described in the visual system.)

    Mu nakata, Y . 1998). Infan t perseveration and implications for object permanence theories: AP D P Mod el of the A-not-B task.Developmental Sciencel 2): 161-84.

    Nelson, C. A. 1995). T h e ontogeny of hum an mem ory: A cognitive neuroscience perspective.Developmental Psychology31 5): 723-38.

    New port, E . L. 1990). Ma turational constraints o n language learning.Cognitive Science14:11-28. Arg ues for matu rational constrain ts of language acqu isition.)

    Rovee-Collier, C . 1997). Dissociations in Infan t Mem ory: Rethinking th e Developm ent ofImplicit and Explicit Mem ory.Psychological Review104 : 467-98.

    Witelson, S . 1987). Neurobiological aspects of language in child ren.Child Development58:653-8. Explores some parallels between aspects of brain and language development.)