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Developmental changes between ages 13 and 21 years in the extent and magnitudeof the BOLD response during decision makingEsther H.H. Keulersa, ? , Peter Stiers b , Jelle Jolles a,caDepartment of Psychiatry and Neuropsychology, Faculty of Health, Medicine and Life Sciences, Maastricht University, The NetherlandsbDepartment of Neuropsychology and Psychopharmacology, Faculty of Psychology and Neuroscience, Maastricht University, The NetherlandscCentre Brain and Learning and AZIRE Research Institute, Faculty of Psychology and Education, VU Amsterdam, The Netherlandsa b s t r a c t a r t i c l e i n f oArticle history:Received 17 March 2010Revised 18 June 2010Accepted 23 August 2010Available online 31 August 2010Developmental neuroimaging results have suggested a progression in focalization in functional activationsfrom childhood to adulthood. The mechanisms underlying this process are thought to be an age-relateddecrease in activation extent as well as an increased magnitude in task-related areas. The present studyaimed to evaluate these notions while controlling for confounders that may bias towards focalization. Weused adolescent subjects in small age ranges. In addition, head motion corrections were incorporated instatistical analyses and regions of interest were identif i ed for each participant separately to overcome inter-individual variability in anatomy and functional organization. Activation patterns of 13-, 17- and 21-year-oldmales were compared during the decision phase of a challenging and complex gambling paradigm. The BOLDamplitude enhanced with increasing age, modulated by task conditions. First, response amplitude duringdiff i cult, endogenous relative to exogenous decisions increased with age. This decision diff i culty effect wasmost pronounced in 21-year-olds, both in areas associated with task execution and default mode areas.Second, deciding to pass as opposed to gamble exerted more effort in inferior frontal and parietal areas onlyby 13- and 17-year-olds. There was neither an age-related decrease in activation extent, nor any qualitativeshifts in activated areas as suggested by the focalization hypothesis. These results suggest that althoughdifferent age groups throughout adolescence engage similar brain areas during decision making, theresponse magnitude in these areas increases with age particularly during diff i cult task conditions, providingthat confounding factors are controlled.© 2010 Elsevier Inc. All rights reserved.IntroductionThe propensity for risk-taking and sensation-seeking behavior hasbeen shown to increase during early and middle adolescence(Steinberg et al., 2008). Poor decision making tendencies in this agegroup might lead to maladaptive outcomes such as increasedsubstance abuse and mortality rates (Dahl, 2004; Paus et al., 2008;Steinberg, 2008). This suggests that decision making and its under-lying neural substrates are subject to development in this period.Neuroimaging research has shown continued maturation andref i nement of brain structures and connectivity throughout adoles-cence and young adulthood. Age-related changes in grey matter showregion-specif i c, inverted U-shaped developmental trajectories as-sumed to ref l ect synaptic pruning and/ or myelination. The peak ofgrey matter volumes is reached earliest in primary sensorimotor areasand latest in higher-order association areas such as prefrontal regions(Giedd et al., 2009; Gogtay et al., 2004). Volumes of white matter andthe integrity of white matter f i bers increase linearly with age,ref l ecting progressive myelination (Giedd, 2004; Giedd et al., 2009;Giorgio et al., 2010). Furthermore, maturational changes have beenshown in functional connectivity within brain networks known tosupport cognitive control, such as prefrontal-parietal and anteriorcingulate-insular connections (Fair et al., 2007; Kelly et al., 2009).These networks continue to specialize and reorganize throughadolescence, with an increased functional connectivity betweenspatially remote cortical areas.Parallel to this structural maturation, socio-emotional and cogni-tive abilities also continue to develop during adolescence (e.g.Choudhury et al., 2006; Huizinga et al., 2006; Keulers et al., 2010).Cognitive functions, which are the focus of the present study, haveshowndifferential rates of maturation. Performance onsimple aspectsof cognition such as response inhibition reaches adult levels in earlyadolescence (Klenberg et al., 2001; Luna et al., 2004; Paus, 2005). Onthe other hand performance on more complex and affective aspects ofcognitionsuch as workingmemory anddecisionmakingreachesadultlevels in middle and late adolescence (Hooper et al., 2004; Luna et al.,2004; Steinberg et al., 2008). The relation between cognitiveNeuroImage 54 (2011) 1442–1454? Corresponding author. Department of Psychiatry and Neuropsychology, Faculty ofHealth, Medicine and Life Sciences, Maastricht University, Dr. Tanslaan 12, 6229 ETMaastricht, The Netherlands. Fax: +31 43 3884092.E-mail address: e.keulers@np.unimaas.nl (E.H.H. Keulers).1053-8119/$ – see front matter © 2010 Elsevier Inc. All rights reserved.doi:10.1016/j.neuroimage.2010.08.059Contents lists available at ScienceDirectNeuroImagejournal homepage: www.elsevier.com/locate/ynimg