Developmental Changes in Cognitive Control through Adolescence
Introduction
The main purpose of this review is a better understanding of the developmental transition in behavior from adolescence to adulthood. To distinguish our focus from earlier (infancy) and later (aging) phases of development, we will refer to processes of “maturation,” to emphasize the nature of this phase as a time when improvements are reaching a plateau and stable levels of adult behavior are achieved. During this phase of development many processes are changing that affect decision making including social, emotional, and cognitive aspects of behavior. In this review, I focus on the processes underlying the maturation of voluntary control of behavior (also referred to as cognitive control and executive function) because of its basic role in decision making.
Development refers to the mechanisms of change that ultimately lead to maturity in adulthood. Although much is known about the significant advances that occur in infancy and childhood, relatively less is known of the mechanisms that support the later parts of development in adolescence as mature-level behavior is approached. This chapter focuses on this particular phase of development as we transition from immature mechanism to mature adult-level behavior. Early development involves the acquisition of abilities that significantly change behavior, but as maturity is reached the changes are more subtle and involve the sophistication of abilities. This process begins in adolescence and can be conceptualized as occurring at the bend in the curve of development just before when the curve flattens representing adult stability (Figure 1). An important question during this phase of development is: what are the mechanisms that support this transition into mature levels of cognitive control of behavior? This is a phase of development when much of behavior appears adult-like, yet there is still pervasive evidence for limits in the efficacy of cognitive control. For example, adolescents are vulnerable to psychopathology, such as schizophrenia and mood disorders (Angold, Costello, & Worthman, 1998; Kessler et al., 1994). In addition, sensation and novelty seeking peak in adolescence (Chambers, Taylor, & Petenza, 2003; Spear, 2000) and are associated with extreme sports, drug use, and unprotected sex. These behaviors appear to be characterized by poor judgment and decision making with limited appreciation for long-term consequences and are often impulsive in nature.
A significant landmark of this phase of development is the maturation to adult levels of the flexible and controlled manner of voluntarily guiding goal-directed behavior. In contrast to exogenously driven behavior (i.e., reflexive, automatic, and guided by outside stimuli) that is present early in development, endogenously driven behavior (i.e., voluntary, planned, and driven by internal goals) matures later. Central to endogenous behaviors are executive processes such as voluntary response inhibition and working memory that allow planned responses. When planning a response, alternatives often must be considered and goal inappropriate responses that may be more reflexive need to be suppressed. This process is referred to as response inhibition and is central to cognitive control (Fuster, 1997). Working memory refers to the ‘sketch pad’ that allows us to retain relevant information on-line to make a planned goal-directed response and it is also central to cognitive control (Fuster, 1997). These processes, as described later in this chapter, are present in infancy but continue to improve throughout adolescence into adulthood and may underlie the emergence of adult-like behavior. We discuss what is known to be unique regarding the cognitive control of behavior during adolescence.
The emergence of cognitive control is driven by continuous interaction of environmental and biological factors. Biological mechanisms determine a timeline when environmental factors can have a lasting effect, and the environment establishes the course of biological mechanisms. For example, in the process of synaptic pruning (discussed later), the environment determines which synapses will be kept and which will not be needed but biological mechanisms determine the times in development when different parts of the brain will be most affected. In terms of cognitive control, brain development constrains the processes that can be performed at different ages. Characterizing the state of brain mechanisms during adolescence can help determine the nature of the cognitive tools that are available or not during this period. The interaction of brain and behavior thus becomes crucial in understanding maturation. These interactions have been studied with functional magnetic resonance imaging (fMRI); later we discuss what neuroimaging has revealed regarding onset of adult-like behavior in adolescence.
Adolescence commonly is considered to encompass 12–17 years of age, with the precise timing varying with gender and puberty (Spear, 2000). Puberty occurs when secretion of hormones in the pituitary gland stimulate ovarian development in females and spermatogenesis in males (Ojeda, Ma, & Rage, 1995). The timing is determined by age as well as metabolic and neuronal factors. Gonad hormone levels have direct effects on molecular mechanisms throughout the brain (McEwen, 2001) including influencing cortical development. Mood and social processes are affected by hormonal changes (Alsaker, 1996) which influence sex steroid receptors in the hippocampus and are associated with the regulation of the neurotransmitter dopamine (DA) especially in the part of the midbrain, namely the nucleus accumbens (NAcc), known to be critical in reward processing (Chambers et al., 2003). There is no clear association between puberty and cognitive processes. Some studies have found a link between spatial abilities and pubertal timing (Petersen, 1976; Waber, Mann, Merola, & Moylan, 1985), but others have not (Orr, Brack, & Ingersoll, 1988; Strauss & Kinsbourne, 1981). Puberty may influence the degree to which one can exert cognitive control in situations of high arousal but in itself may not be directly linked to the development of cognitive control. Still, it is a pivotal aspect of this stage of development that must not be over-looked.
In this chapter, I analyze available evidence regarding the behavioral and brain processes present during adolescence. I discuss immaturities in the brain and how these may limit voluntary control. I also discuss the evidence indicating continuing development of cognitive control through adolescence, assessing the nature of what is still immature during this period. A critical review of the results of neuroimaging studies provides a forum to integrate the mechanisms of how brain and behavior interact during this time. A developmental theory emphasizing the role of the integration of widely distributed circuitry underlying the transition from adolescence to adulthood is contrasted against the traditional view that cognitive development is primarily driven by the unique protracted maturation of prefrontal cortex. Finally, I delineate a theory of maturation that is specific to this time of development and can be used to make predictions of behavioral performance.
Section snippets
What is Maturing in the Brain during Adolescence?
Identifying the progression of structural brain maturation in adolescence can further understanding of the brain functional systems that are available to support complex behavior such as cognitive control. The gross morphology of the brain is apparent early in life. The delineation of cortical folding in the brain is in place by birth (Armstrong et al., 1995). The brain rapidly changes from a smooth surface to a convoluted one postnatally with well-defined sulci and gyri that are organized in
What Executive Processes Improve during Adolescence?
In parallel with maturation of brain mechanisms, behavior is becoming more controlled and voluntary during adolescence. Executive function is used to define the processes that allow for cognitive or voluntary control of behavior including response planning and preparation, response inhibition, and working memory that support cognitive flexibility, abstract thought, and rule-guided behavior. Executive function is apparent early in development but continues to improve through adolescence.
What have Developmental Neuroimaging Studies Revealed about Maturation of the Cognitive System?
The co-occurrence in adolescence of changes in brain processes such as synaptic pruning and myelination that support cognition and speed of processing as well as changes in cognition and behavior suggest an important link between brain and behavior through development. However, investigating the association between changes in brain structure and behavior does not provide information regarding the maturity of dynamic brain function. That is, although pruning and myelination throughout cortical
Limitations in Developmental Studies of Brain Function
A number of factors underlying differences across studies need to be taken into consideration when interpreting the results reported here. The most controversial in the literature has been the issue of how to interpret age-related differences in brain function when performance also differs by age. On the one hand, differences in performance could simply reflect that unique strategies are adopted and may not capture developmental change in the brain processes themselves. On the other hand,
The Transition to Widely Distributed Circuitry
Adolescence differs qualitatively from childhood in that adolescents can appear to have mature cognitive control. Our developmental studies on both response inhibition and working memory show that adolescents typically behave at adult-like levels (Luna et al., 2004; Scherf et al., 2006). Adolescents recruit many of the same circuits that adults use, yet there are important differences that are telling of the adolescent period. Adolescents appear to use dorsolateral prefrontal cortex to a higher
How does Immaturity in Executive Processing Inform us about Real Life Decision Making?
Adolescence is the peak of sensation seeking and risk-taking behavior across species (Hodes & Shors, 2005; Stansfield & Kirstein, 2006). Sensation seeking is believed to be important for obtaining experience in independent decision making needed for survival as an adult. Risk-taking behavior occurs when sensation-seeking involves decision making that results in a risk to survival. Drug use, unprotected sex, and extreme sports are examples of risk-taking behavior. During adolescence, sensation
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