Neural Plasticity of Development and Learning Galvan, 2010 Presented by Kristen Morin and Sunil Patel 1 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusions 2 I. Defining Development and Learning What is Development? organismic changes as a result of growth, maturation, and/or experience What is Learning? acquisition of a skill or knowledge through study, instruction, and/or experience The Common Denominator? Experience 3 The Debate: Are they... Inseparable constructs [Casey et al., 2006] • Learning occurs within a developmental framework Distinguishable by time periods [Smith and Thelan, 2003] • Learning: days, hours, months of practice, • Development: weeks, months, years of change On a continuum [Galvan, 2010] • Both are processes that induce neural plasticity via shifts in mechanisms • e.g. Discrimination of phonetic speech sounds in all languages by infants [Kuhl et. al., 1997] Complementary processes [Karmiloff-Smith, 1994] [Greenough et. al. 1987] • Biological predispositions (experience-expectant development) are subsequently modified by experience (experience-dependent learning) • Evolutionary tendency to modify existing systems (avoids redundancy) 4 ... as a Continuum Endpoints receive inputs from both processes Experience-expectant largely influence emergent phenotype (development) utilize environmental information common to all members of a species (common input) throughout evolutionary history (common timeline) neural changes follow phylogenic norm (typical plasticity) Experience-dependent greater influence in learning utilize information from the unique experiences (specific inputs) of each particular individual (fluid timeline) neural changes result from individual experience [unique plasticity) 5 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusions 6 II. Neural Plasticity Functions in Development: - acquisition of new information Thinking and Learning neurogenesis activity dependent synaptic plasticity programmed cell death - adaptations to fluctuating environments or conditions - recuperation from injury Changes in Neural Organization (Functional + Structural) [Johnston, 2009] neuronal circuitry dendritic spines Changes in Behavior 7 Tools in Neuroimaging Cross-Sectional Approach e.g. Cortical representation of fingers in musicians vs. non-musicians [Elbert et al., 1995] [Pros] time and cost vs. [Cons] Lower statistical power, potential confounding variables, cohort effects Longitudinal Method e.g. Increased number of dendrites in visual cortex of trained vs. untrained animals [Greenough et al., 1979] [Pros] high statistical power (within subject comparisons) vs. [Cons] practice effects, increased time and costs, lower retention of subjects Trans-cranial Magnetic Stimulation (TMS) e.g. Repeated TMS delivered to superior temporal cortex causes macroscopic changes in gray matter [May et al., 2007] [Pros] Non-invasive, therapeutic potential vs. [Cons] Subcortical effects difficult to determine, measure, or ensure, potential for inaccuracy when targeting specific ROI 8 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusions 9 III. Progressive and Regressive Changes with Learning functional changes in neural activity following training are contradictory Potential explanations for... A. Regressive changes Dual-processing framework of learning- central resources essential for controlled processing in novices becomes less critical as skill level improves to point of automatic processing [Chein & Schneider, 2005] Developmental process of selective pruning 10 III. Progressive and Regressive Changes with Learning functional changes in neural activity following training are contradictory Potential explanations for... B. Progressive changes Constructivist Manifesto- connectivity progresses from fewer to greater connections [Quartz & Sejnowski, 1997] Recruitment of additional cortical units with practice e.g. increased prefrontal and parietal activation related to working memory capacity in children [Klingberg et. al., 2002] 11 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusion 12 IV. The Plasticity of Developmental Timing Sensitivity to experience-expectant outcomes can be manipulated - Suggested in animal research (e.g. monocular deprivation effects in kittens reared in the dark for 6, 8, 10 months) [Cynader and Mitchell 1980] Processes reliant on experience-expectant mechanisms show experience-related plasticity in developmental onset - Enrichment-experiments conducted with infants (e.g. infants show increased reaching behavior with sticky mittens (and later without mittens) prior to typical motor development allowing for actual grasping of objects) [Needham et. al., 2002] - Help avoid potential confounds of domain general experience with which older children are already equipped. Q: Does accelerating developmental timing result in neural changes? And would these changes have downstream effects? 13 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusion 14 V. Neural Mechanism- Same or Different? Same Learning Process: Prediction Error Signal (PES) Neural activation in adults, accompanies learning by trial and error [Hollerman & Schultz, 1998] Thought to be mediated primarily through dopamine as a “learning signal” event unexpected: increase firing rate/increase activation of dopamine rich-regions event expected: decrease in dopamine signal [Fiorillo et al., 2003; 2008] Different emergent plasticity across development Infants: provides basic architecture for the neural system. Adults: modifies existing system 15 Utility of “Noise”: Performance confounds between children and adults Alternative neural strategies engaged by children to compensate for system maturity of adults: confound or useful? Dynamic Systems Theory utilizes variability in performance [Smith & Thelan, 2003] - developmental trajectories of change over short time periods (inter-subject longitudinal) and/or over longer developmental span (cross-sectional) Developmental vs. performance-related neural activity post-hoc - performance-matched subgroups, age subgroups, or neither [Schlagger et al., 2002] [Brown et al.,2005] - Identified progressive and regressive changes associated with age - potential to investigate training-related, age-related or independent neural changes in future experiments 16 Longitudinal training studies across development “The only way to identify the root of neural plasticity as either experiential or developmental is to conduct a longitudinal study” Recent studies have proven feaseability despite inherent challenges - [Durston et al., 2006] combined cross-sectional and longitudinal study of response inhibition task [Hyde et al., 2009] Implemented a training component building upon existing studies in adult musicians and matched non musicians that showed structural and functional differences in brain areas associated with music production. 17 18 Longitudinal training studies across development Recent studies have proven feasibility despite inherent challenges - [Durston et al., 2006] Longitudinal results: Attenuation of dorsolateral prefrontal cortex with age Cross-sectional results: increased activation in ventral prefrontal cortex with improved task performance [Hyde et al., 2009] Trained children showed greater behavioral improvements on music discrimination and related tasks. - - - Structural changes in right perecentral gyrus, corpus collosum, and the primary auditory region. Findings consistent with same findings in adults, provide new evidence for traininginduced structural brain pasticity in early childhood. 19 Questions to be Addressed - Which neural systems show greater or less training-related plasticity earlier in development? - Greater plasticity in learning and receiving language input during infancy. - How do the timescales of neural plasticity change across development? - Which behaviors cannot be “sped up” by exposure earlier in development because of time locked experience-expectant mechanisms? 20 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusion 21 VI. Methodological Considerations and Potential Confounds Resources: Steep scanner costs and the need for a healthy sample size of both children and adults, along with the need for specialized training staff call for a very expensive experiment. Subject attrition: Subject burden for training is high. Participants are required to maintain cognitive and training engagement as they visit the lab for multiple sessions. Paying closer attention and being sensitive to families’ needs can help achieve successful subject retention. 22 VI. Methodological Considerations and Potential Confounds • Scanner Related Anxiety: Subjects, particularly children, may have increased head movements as well as increased vigilance which may skew results. • Performance Differences: Without addressing performance differences the differences in brain activity between children and adults can inappropriately be misinterpreted as maturational differences. 23 Neural Plasticity of Development and Learning I. Defining Development and Learning II. Neural Plasticity III. Progressive and Regressive Changes with Learning IV. Plasticity of Developmental Timing V. Neural Mechanism- Same or Different? VI. Methodological Considerations/Potential Confounds VII. Conclusion 24 VII. Conclusions - Consider if and how development differs from learning. - How does this distinction translates to differences or similarities in neural mechanisms? - The local neural environment in which plasticity occurs is a critical component of change. - Inherent challenges in developmental study have prevented significant advances in addressing plasticity-related questions. - It’s IMPORTANT to embrace longitudinal studies in order to uncover mechanisms underlying plasticity across development. 25