NEURO-EDUCATION AND NEURO-REHABILITATION EDITED BY : Eduardo Martínez-Montes, Julie Chobert and Mireille Besson PUBLISHED IN : Frontiers in Psychology 1 November 2016 | Neur o-Education and Neuro-Rehabilitation Frontiers in Psychology Frontiers Copyright Statement © Copyright 2007-2016 Frontiers Media SA. All rights reserved. All content included on this site, such as text, graphics, logos, button icons, images, video/audio clips, downloads, data compilations and software, is the property of or is licensed to Frontiers Media SA (“Frontiers”) or its licensees and/or subcontractors. The copyright in the text of individual articles is the property of their respective authors, subject to a license granted to Frontiers. The compilation of articles constituting this e-book, wherever published, as well as the compilation of all other content on this site, is the exclusive property of Frontiers. 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Find out more on how to host your own Frontiers Research Topic or contribute to one as an author by contacting the Frontiers Editorial Office: researchtopics@frontiersin.org 2 November 2016 | Neur o-Education and Neuro-Rehabilitation Frontiers in Psychology NEURO-EDUCATION AND NEURO-REHABILITATION Cover figure designed by Claudia Méndez Romero Topic Editors: Eduardo Martínez-Montes, Cuban Neuroscience Center, Cuba Julie Chobert, Centre National de la Recherche Scientifique, France Mireille Besson, Centre National de la Recherche Scientifique and Aix-Marseille Université, France In the last decade, important discoveries have been made in cognitive neuroscience regarding brain plasticity and learning such as the mirror neurons system and the anatomo-functional organization of perceptual, cognitive and motor abilities.... Time has come to consider the societal impact of these findings. The aim of this Research Topic of Frontiers in Psychology is to concentrate on two domains: neuro-education and neuro-rehabilitation. At the interface between neuroscience, psychology and education, neuro-education is a new inter-disciplinary emerging field that aims at developing new education programs based on results from cogni- tive neuroscience and psychology. For instance, brain-based learning methods are flourishing but few have been rigorously tested using well-controlled procedures. Authors of this Research Topic will present their latest findings in this domain using rigorously controlled experiments. Neuro-rehabilitation aims at developing new rehabilitation methods for children and adults with 3 November 2016 | Neur o-Education and Neuro-Rehabilitation Frontiers in Psychology learning disorders. Neuro-rehabilitation programs can be based upon a relatively low number of patients and controls or on large clinical trials to test for the efficiency of new treatments. These projects may also aim at testing the efficiency of video-games and of new methods such as Transcranial Magnetic Stimulation (TMS) for therapeutic interventions in children or ado- lescents with learning disabilities. This Research Topic will bring together neuroscientists interested in brain plasticity and the effects of training, psychologists working with adults, with normally developing children and children with learning disabilities as well as education researchers directly confronted with the efficiency of education programs. The goal for each author is to describe the state of the art in his/her specific research domain and to illustrate how her/his research findings can impact education in the classroom or rehabilitation of children and adolescents with learning disorders. Citation: Martínez-Montes, E., Chobert, J., Besson, M., eds. (2016). Neuro-Education and Neuro-Rehabilitation. Lausanne: Frontiers Media. doi: 10.3389/978-2-88945-006-0 4 November 2016 | Neur o-Education and Neuro-Rehabilitation Frontiers in Psychology Table of Contents 06 Editorial: Neuro-Education and Neuro-Rehabilitation Eduardo Martínez-Montes, Julie Chobert and Mireille Besson PART I. Neuro-education 09 Neuroscience illuminating the influence of auditory or phonological intervention on language-related deficits Sari Ylinen and Teija Kujala 18 Musical training as an alternative and effective method for neuro-education and neuro-rehabilitation Clément François, Jennifer Grau-Sánchez, Esther Duarte and Antoni Rodriguez-Fornells 33 Music and Dyslexia: A New Musical Training Method to Improve Reading and Related Disorders Michel Habib, Chloé Lardy, Tristan Desiles, Céline Commeiras, Julie Chobert and Mireille Besson 48 Musical plus phonological input for young foreign language readers M. C. Fonseca-Mora, Pilar Jara-Jiménez and María Gómez-Domínguez 57 Engagement in community music classes sparks neuroplasticity and language development in children from disadvantaged backgrounds Nina Kraus, Jane Hornickel, Dana L. Strait, Jessica Slater and Elaine Thompson 66 A groundwork for allostatic neuro-education Lee Gerdes, Charles H. Tegeler and Sung W. Lee PART II. Neuro-rehabilitation 82 Review of neural rehabilitation programs for dyslexia: how can an allophonic system be changed into a phonemic one? Willy Serniclaes, Gregory Collet and Liliane Sprenger-Charolles 90 Hearing and music in unilateral spatial neglect neuro-rehabilitation Alma Guilbert, Sylvain Clément and Christine Moroni 98 New framework for rehabilitation – fusion of cognitive and physical rehabilitation: the hope for dancing Prabhjot Dhami, Sylvain Moreno and Joseph F . X. DeSouza 113 Facing the music: three issues in current research on singing and aphasia Benjamin Stahl and Sonja A. Kotz 117 Intracerebral functional connectivity-guided neurofeedback as a putative rehabilitative intervention for ameliorating auditory-related dysfunctions Stefan Elmer and Lutz Jäncke 5 November 2016 | Neur o-Education and Neuro-Rehabilitation Frontiers in Psychology PART III. Basic research with implications in both fields 124 Basics for sensorimotor information processing: some implications for learning Franck Vidal, Cédric Meckler and Thierry Hasbroucq 138 Perceiving fingers in single-digit arithmetic problems Ilaria Berteletti and James R. Booth 148 Basic and supplementary sensory feedback in handwriting Jérémy Danna and Jean-Luc Velay 159 The differential time course for consonant and vowel processing in Arabic: implications for language learning and rehabilitation Sami Boudelaa 169 Tactile stimulations and wheel rotation responses: toward augmented lane departure warning systems Christophe Tandonnet, Borís Burle, Franck Vidal and Thierry Hasbroucq EDITORIAL published: 22 September 2016 doi: 10.3389/fpsyg.2016.01427 Frontiers in Psychology | www.frontiersin.org September 2016 | Volume 7 | Article 1427 | Edited and reviewed by: Eddy J. Davelaar, Birkbeck, University of London, UK *Correspondence: Eduardo Martínez-Montes eduardo@cneuro.edu.cu Specialty section: This article was submitted to Cognitive Science, a section of the journal Frontiers in Psychology Received: 10 June 2016 Accepted: 06 September 2016 Published: 22 September 2016 Citation: Martínez-Montes E, Chobert J and Besson M (2016) Editorial: Neuro-Education and Neuro-Rehabilitation. Front. Psychol. 7:1427. doi: 10.3389/fpsyg.2016.01427 Editorial: Neuro-Education and Neuro-Rehabilitation Eduardo Martínez-Montes 1 *, Julie Chobert 2 and Mireille Besson 2 1 Neuroinformatics Department, Cuban Neuroscience Center, La Habana, Cuba, 2 Laboratoire de Neurosciences Cognitives, UMR 7291, Centre National de la Recherche Scientifique, Aix-Marseille Université, Marseille, France Keywords: learning disorders, language, musical training, sensorimotor training, neurofeedback The Editorial on the Research Topic Neuro-Education and Neuro-Rehabilitation The latest advances in cognitive psychology, neuropsychology, and cognitive neuroscience has brought from science fiction to reality the possibility of influencing our brain activity (Owen et al., 2010; Glannon, 2014; Gruzelier, 2014a). Better understanding of brain functioning and brain plasticity has allowed neuroscientists to transfer findings from fundamental research to education and to the rehabilitation of learning disabilities (Besson et al., 2011; Goswami, 2016). The emerging fields of neuro-education and neuro-rehabilitation aim at creating effective and safe programs to improve brain functioning related to specific perceptive, cognitive, emotional, and motor abilities. Some attempts to achieve these goals take advantage of the use of natural mechanisms, such as those mediating the interactions between brain and arts (Särkämö et al., 2008; Bringas et al., 2015). Others use experimental designs to make the brain aware of its own activity, creating the so-called neurofeedback loop (Gruzelier, 2014b). Succeeding in these goals would constitute an achievement of high societal impact (Davidson and McEwen, 2012; Vuilleumier et al., 2014). This Frontiers Research Topic brings together 16 articles that cover a broad scope of topics in the relatively young but very dynamic fields of neuro-education and neuro-rehabilitation. Contributed by world-renowned scientists in cognitive psychology, neuropsychology, and cognitive neuroscience, often experts in different types of learning disorders, this E-book is organized around three main themes: neuro-education, neuro-rehabilitation and basic research with relevance to both fields. Each theme includes Review articles covering the state-of-the-art of knowledge in a specific sub-domain, Original Research articles reporting new discoveries and Opinion and Hypothesis and Theory articles adding exciting new ideas and approaches for neuro-educational and neuro-rehabilitation methods. In the first part, dedicated to neuro-education, Ylinen and Kujala review the impact of auditory or phonological training on the level of performance in various tasks and on the neural basis of behavior in children with dyslexia, children with specific language impairment and children with language-learning impairment. François et al. review the efficacy of musical training for language learning. They highlight several studies showing that learning to play a musical instrument can induce substantial neuro-plastic changes in cortical and subcortical regions of motor, auditory and speech processing networks. They show evidence that musical training can be an alternative, low-cost and effective method for the treatment of language-learning impaired populations, as well as for patients with stroke or Parkinson Disease. Direct support for the use of musical training for the rehabilitation of children with dyslexia is reported by Habib et al. who tested the efficacy of a specially-designed Cognitivo-Musical training (CMT) method. Intensive short-term CMT with dyslexic children yielded significant improvements in categorical and auditory perception 6 Martínez-Montes et al. Editorial: Neuro-Education and Neuro-Rehabilitation of temporal components of speech, while long-term CMT provided additional improvements in auditory attention, phonological awareness, reading abilities and repetition of pseudo words. Along the same lines, Fonseca-Mora et al. also tested the efficacy of a new phonological training program (with and without music), for teaching to read in a foreign language, demonstrating its beneficial effects on early reading skills but without additional improvements linked to music training. Final but not least, Kraus et al. present the results of a longitudinal study examining the impact of a community music program on language development in children from low socio-economic backgrounds. Children more engaged in the music program developed stronger brain encoding of speech and improved reading scores, thereby suggesting that this kind of program provides children with auditory enrichments that may counteract some of the biological consequences of growing up in poverty. To conclude this section, the reader will find a novel approach to neuro-education, as proposed by Gerdes et al. They view learners in terms of their neurodevelopmental trajectories and propose a groundwork for allostatic neuro- education (GANE). Illustrative case studies of the use of GANE in children with Asperger’s syndrome, attention- deficit hyperactivity disorder and reading difficulties are presented. The second part of this E-book is dedicated to neuro- rehabilitation. Serniclaes et al. review the efficacy of remediation methods that tap into core deficits in dyslexia (phonemic, grapho-phonemic, and graphemic) and examine how some of these methods may contribute to the remediation of allophonic perception. Guilbert et al. describe different procedures for sensory training in unilateral spatial neglect (USN) and present recent scientific evidence that makes music a good candidate for USN patients’ rehabilitation. Dhami et al. consider the use of dancing as an intervention tool and as a potential parallel to physical and music therapies, since dancing also engages various perceptive, cognitive, emotional and motor functions. The opinion paper from Stahl and Kotz addresses three relevant issues in current research on singing and aphasia: articulatory tempo, clinical research designs and formulaic language resources. The authors discuss how these issues may reconcile seemingly contradictory findings in the literature and provide guidelines for future research based on holistic and analytic approaches that may help improving the efficacy of music-based aphasia therapy. Finally, in a Hypothesis and Theory article, Elmer and Jäncke consider the use of a neurofeedback approach for auditory rehabilitation. They first stress the advantages of using intracerebral functional connectivity (IFC) instead of quantitative EEG for interventional applications and then propose concrete interventional IFC applications that may improve auditory-related dysfunctions such as developmental dyslexia. In the third part, we compiled some interesting studies which contribute both to a better comprehension of basic psychophysiological mechanisms and to the development of potential applications for neuro-education and neuro- rehabilitation. Vidal et al. review the role of sensorimotor information for motor learning. They discuss the effects of several factors known to influence information processing in sensorimotor activities based on the distinction between extrinsic (e.g., quantity and quality of information, level of instruction and motor program learning) and intrinsic factors (e.g., prior information, individual strategies and capabilities for fast error detection). In a more specific context, Berteletti and Booth investigated the extent to which somatosensory information from the fingers contributes to numerical sense in children. Their work provided first neurological evidence for a functional role of the somatosensory finger area in proficient arithmetic problem solving, thereby encouraging educational practices to integrate finger-based strategies as a tool for instilling stronger numerical sense. Still in another context, writing, Danna and Velay review studies that use natural sensory and supplementary feedback to help the writer learn how to write and to control writing. They discuss the role of each sensory modality, how information is used in handwriting control and how this control changes with practice and learning. Turning from writing to reading, Boudelaa reports, in an original research article, that the processing time course in auditory modality is different for consonants and vowels in Arabic. The implications of this work for neuro-education and neuro-rehabilitation in Arabic are discussed. Finally, Tandonnet et al. illustrate how basic approaches in cognitive science may benefit human factors engineering and potentially improve man- machine interfaces. We hope this compilation of articles describing the latest research in the field of neuro-education and neuro-rehabilitation will be of interest to the readers and will impulse even more research in these fascinating new fields with strong societal impact. AUTHOR CONTRIBUTIONS All authors listed, have made substantial, direct and intellectual contribution to the work, and approved it for publication. REFERENCES Besson, M., Chobert, J., and Marie, C. (2011). Transfer of training between music and speech: common processing, attention, and memory. Front. Psychology 2:94. doi: 10.3389/fpsyg.2011.00094 Bringas, M. L., Zaldivar, M., Rojas, P. A., Martinez-Montes, K., Chongo, D. M., Ortega, M. A. et al. (2015). Effectiveness of music therapy as an aid to neurorestoration of children with severe neurological disorders. Front. Neurosci. 9:427. doi: 10.3389/fnins.2015.00427 Davidson, R. J., and McEwen, B. S. (2012). Social influences on neuroplasticity: stress and interventions to promote well-being. Nat. Neurosci. 15, 689–695. doi: 10.1038/nn.3093 Glannon, W. (2014). Neuromodulation, agency and autonomy. Brain Topogr. 27 , 46–54. doi: 10.1007/s10548-012-0269-3 Frontiers in Psychology | www.frontiersin.org September 2016 | Volume 7 | Article 1427 | 7 Martínez-Montes et al. Editorial: Neuro-Education and Neuro-Rehabilitation Goswami, U. (2016). Educational neuroscience: neural structure-mapping and the promise of oscillations. Curr. Opin. Behav. Sci. 10, 89–96. doi: 10.1016/j.cobeha.2016.05.011 Gruzelier, J. H. (2014a). EEG-neurofeedback for optimising performance. I: a review of cognitive and affective outcome in healthy participants. Neurosci. Biobehav. Rev. 44, 124–141. doi: 10.1016/j.neubiorev.2013.09.015 Gruzelier, J. H. (2014b). EEG-neurofeedback for optimising performance. III: A review of methodological and theoretical considerations. Neurosci. Biobehav. Rev. 44, 159–182. doi: 10.1016/j.neubiorev.2014.03.015 Owen, A. M., Hampshire, A., Grahn, J. A., Stenton, R., Dajani, S., Burns, A. S., et al. (2010). Putting brain training to the test. Nature 465, 775–778. doi: 10.1038/nature09042 Särkämö, T., Tervaniemi, M., Laitinen, S., Forsblom, A., Soinila, S., Mikkonen, M., et al. (2008). Music listening enhances cognitive recovery and mood after middle cerebral artery stroke. Brain 131, 866–876. doi: 10.1093/brain/awn013 Vuilleumier, P., Sander, D., and Baertschi, B. (2014). Changing the brain, changing the society: clinical and ethical implications of neuromodulation techniques in neurology and psychiatry. Brain Topogr. 27, 1–3. doi: 10.1007/s10548-013- 0325-7 Conflict of Interest Statement: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Copyright © 2016 Martínez-Montes, Chobert and Besson. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms. Frontiers in Psychology | www.frontiersin.org September 2016 | Volume 7 | Article 1427 | 8 MINI REVIEW ARTICLE published: 17 February 2015 doi: 10.3389/fpsyg.2015.00137 Neuroscience illuminating the influence of auditory or phonological intervention on language-related deficits Sari Ylinen 1 * and Teija Kujala 1,2 1 Cognitive Brain Research Unit, Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland 2 Cicero Learning, University of Helsinki, Helsinki, Finland Edited by: Julie Chobert, Centre National de la Recherche Scientifique, France Reviewed by: April A. Benasich, Rutgers University, USA Anna J. Simmonds, Imperial College London, UK L. Robert Slevc, University of Maryland, College Park, USA Fabio Richlan, University of Salzburg, Austria *Correspondence: Sari Ylinen, Cognitive Brain Research Unit, Institute of Behavioural Sciences, University of Helsinki, P .O. Box 9, Helsinki, FIN-00014, Finland e-mail: sari.ylinen@helsinki.fi Remediation programs for language-related learning deficits are urgently needed to enable equal opportunities in education. To meet this need, different training and intervention programs have been developed. Here we review, from an educational perspective, studies that have explored the neural basis of behavioral changes induced by auditory or phonological training in dyslexia, specific language impairment (SLI), and language-learning impairment (LLI). Training has been shown to induce plastic changes in deficient neural networks. In dyslexia, these include, most consistently, increased or normalized activation of previously hypoactive inferior frontal and occipito-temporal areas. In SLI and LLI, studies have shown the strengthening of previously weak auditory brain responses as a result of training. The combination of behavioral and brain measures of remedial gains has potential to increase the understanding of the causes of language-related deficits, which may help to target remedial interventions more accurately to the core problem. Keywords: language deficit, dyslexia, neuroscience, training, remediation, intervention INTRODUCTION Finding the most effective techniques to remediate language- related impairments, such as dyslexia, specific language impair- ment (SLI), or language-learning impairment (LLI, cf. Tallal, 2001), would be of crucial importance to educators, who try to help children struggling with these learning difficulties. This raises a question, whether understanding the neurobiological underpin- nings of language impairments facilitates their efficient treatment. In this review, we discuss how neuroscience illuminates the effects of auditory or phonological intervention on dyslexia, SLI, and LLI. We focus on auditory or phonological interventions, because in many cases dyslexia, SLI, and LLI are all characterized by phono- logical (or auditory) deficits (Tallal, 2001; Shaywitz and Shaywitz, 2005; Pennington and Bishop, 2009; Ramus et al., 2013), despite their complex etiology. Whereas detailed brain areas influenced by reading interventions can be found in a recent meta-analysis by Barquero et al. (2014), here we address whether neuroscientific research on the remediation of language-related deficits is useful for educators and whether it has something to add over behavioral research from an educational perspective. In the current review, the selection of publications was based on the following criteria: the research should concern dyslexia, SLI, or LLI, include testing before and after an auditory or phonological intervention or training, involve brain research measures [(func- tional) magnetic resonance imaging (MRI/fMRI), magnetic source imaging (MSI) or magnetoencephalography (MEG), event-related potentials (ERP), or electroencephalography (EEG)], and com- pare two or more groups of participants to control for the effects of repeated testing and maturation (McArthur, 2009). Searches from Web of Science and PubMed (keywords dyslexia/SLI/LLI, intervention/remediation/training, fMRI/MEG/ERP) were used in finding literature. Additional publications were found in the reference lists of relevant studies. IS NEUROSCIENTIFIC RESEARCH USEFUL FOR EDUCATORS? Research on remedial interventions for learning deficits may have important applicability to education (Tallal, 2012). In this area, collaboration between education and neuroscience could result in mutual benefits (Sigman et al., 2014). However, the value of the neuroscientific approach in such research has been questioned by Bishop (2013) because of methodological and interpretive reasons. She argued that neuroscientific studies often use small subject groups, which may decrease their reliability and result in small sta- tistical power (cf. Button et al., 2013). Furthermore, Bishop (2013) noted that some studies lack an adequate control group, which is important to control for the effects of repeated testing and matura- tion (see also McArthur, 2009). Indeed, future intervention studies should not only aim at having larger subject groups (Bishop, 2013) and adequate control groups (McArthur, 2009; Bishop, 2013), but also control for placebo effects (Boot et al., 2013). Bishop (2013) also argued that the critical test of the effective- ness of interventions is the change of behavior rather than that of brain function; changes in the brain should not be considered more important than changes in behavior. However, rather than emphasizing the brain over behavior, neuroscientific intervention studies typically aim to determine the links between brain function and behavior. Importantly, understanding the link or correlation between brain activation and skills as a result of training may help to explain how and why remedial gains take place. Since the combination of neuroscientific and behavioral measures has been www.frontiersin.org February 2015 | Volume 6 | Article 137 | 9 Ylinen and Kujala Neuroscience in language-related interventions shown to be a better predictor of reading skills than behavioral measures alone (Hoeft et al., 2007; Maurer et al., 2009), this com- bination has potential to outperform mere behavioral measures in the study of remedial gains. Cognitive neuroscience has, in our opinion, also some advantages over behavioral research that were not mentioned by Bishop (2013). Especially when working with children whose motivation and skills can affect their performance considerably, a possibility to study the effects of intervention with- out subject’s active effort or attention is a clear advantage. This is possible, for example, by recording mismatch negativity (MMN) brain response (Näätänen et al., 2007; Kujala and Näätänen, 2010). From educators’ perspective, neuroscientific research is seldom directly applicable in the assessment of remedial interventions. Importantly, however, educators may benefit from neuroscientific research by obtaining a more detailed picture of relevant processes underlying behavior. For example, brain measures may help to dis- entangle whether behaviorally observed improvement is due to the normalization of the core deficit or some compensatory strategy (e.g., Eden et al., 2004; Shaywitz et al., 2004), which is not evident in behavioral data. If, hypothetically, some intervention resulted in the formation of a compensatory function to solve some task, it may improve behavior to a certain degree but might not compete in effectiveness with the optimal function for solving that task. Still, in a large subject group, this compensatory improvement in behavior may be taken to reflect a successful intervention, if statis- tically significant improvement is achieved. Thus, neuroscientific research can potentially give some valuable information to educa- tors about the deficits, which may help to target the contents of interventions more accurately. OVERVIEW OF STUDIES ON NEUROBIOLOGICAL CHANGES FOLLOWING PHONOLOGICAL OR AUDITORY INTERVENTIONS As shown by Tables 1 and 2, the majority of studies on phono- logical or auditory interventions focused on dyslexia or related problems in reading, writing, or spelling. Furthermore, the major- ity of studies have focused on children. Older age groups should not be neglected in remediation and its research, however: as noted by Eden et al. (2004), most dyslexics are adults, who may suffer from the socio-economic consequences of their reading deficit. There seem to be no constraints with respect to brain plasticity that would hinder remediation in adults or older children (Simos et al., 2002; Eden et al., 2004). Nevertheless, the earlier the inter- ventions are conducted, the more benefit to individuals is gained, because learning is cumulative. The early gains may help to prevent difficulties not only in academic but socio-emotional domain. The optimal timing of intervention is, however, determined by matu- rity and acquired skills. For example, if a new skill is scaffolded by previous skills, it cannot be adapted before they are mastered (cf. Jolles and Crone, 2012). The studies listed in Tables 1 and 2 suggest that in addition to behavior, the remedial gains of phonological or auditory inter- ventions are consistently reflected in different aspects of brain functioning. These include increased or normalized brain acti- vation as a result of training in previously hypoactive areas as measured with fMRI (Aylward et al., 2003; Temple et al., 2003; Eden et al., 2004; Shaywitz et al., 2004; Gaab et al., 2007; Meyler et al., 2008; Heim et al., 2014) and MSI or MEG (Simos et al., 2002; Pihko et al., 2007) during different cognitive tasks. MRI- based proton MR spectroscopy has shown normalized metabolism in certain brain areas after interventions (Richards et al., 2000, 2002). Training-induced changes in strength and timing of neu- ral responses to stimulation have been demonstrated with ERPs (Kujala et al., 2001; Hayes et al., 2003; Stevens et al., 2008, 2013; Jucla et al., 2010; Lovio et al., 2012; Hasko et al., 2014). Also the time-frequency analysis of EEG has revealed amplitude increases in the oscillatory brain activity after training (Heim et al., 2013). In addition to brain function, interventions have been found to change brain anatomy, such as white matter integrity (Keller and Just, 2009). Tables 1 and 2 also show that remedial gains, if any, consistently manifest in both behavioral and brain measures: in 16 out of 17 studies of Table 1 and in all four studies of Table 2 , remedial gains were found in both brain activation and skills tar- geted by intervention (note that Jucla et al., 2010, failed to find different behavioral improvement and similar brain response pat- terns between their treatment group and controls). The strong coupling of training gains in behavior and brain activation sug- gests that most likely the observed changes in the brain drive the changes in the behavior. As neuroscientific research may reveal the neural dynamics of processes related to behavioral perfor- mance and allows localize the deficient brain functions, it may enable to specify the neural mechanisms underlying language- related impairments and to determine brain functions and areas altered by interventions, which surface in behavior as improved skills. However, it is noteworthy that Tables 1 and 2 lists published studies, whereas studies failing to find changes in behavior or brain activation may remain unpublished. This may cause bias toward systematically finding the coupling between neural and behavioral gains. A recent meta-analysis of neuroscientific research exploring reading networks in the brain has suggested that dyslexia is characterized by the dysfunction of left occipito-temporal cor- tex, left inferior frontal gyrus, and the inferior parietal lobule (Richlan, 2012; see also Richlan et al., 2011). These brain areas are involved in phonological encoding, phonological representa- tions, and attention, respectively (Richlan, 2012). Barquero et al.’s (2014) meta-analysis of the neuroimaging of reading interven- tions, in turn, suggests intervention-induced functional changes in the left thalamus, left middle occipital gyri, bilateral inferior frontal gyri, right insula, and right posterior cingulate gyrus. Thus, both Richlan’s (2012) and Barquero et al.’s (2014) findings point toward the central role of inferior frontal and occipito- temporal/occipital dysfunction in dyslexia. Correspondingly, the neuroscientific dyslexia studies included in Table 1 , involving auditory or phonological intervention, have shown normalized brain activation, metabolism, or anatomy as a result of interven- tions in the occipito-temporal (Aylward et al., 2003; Heim et al., 2014) and inferior frontal (Richards et al., 2000, 2002; Aylward et al., 2003; Shaywitz et al., 2004; Heim et al., 2014) areas. In addition, normalized activation following interventions has been repeatedly observed in inferior parietal (Temple et al., 2003; Eden et al., 2004; Meyler et al., 2008, see also Richlan, 2012), superior parietal (Aylward et al., 2003; Eden et al., 2004; Meyler et al., 2008), and temporal (Simos et al., 2002; Aylward et al., 2003; Temple et al., 2003; Shaywitz et al., 2004) areas. Although inferior frontal and Frontiers in Psychology | Cognitive Science February 2015 | Volume 6 | Article 137 | 10 Ylinen and Kujala Neuroscience in language-related interventions Table 1 | Publications including neuroscientific research on phonological or auditory remediation of dyslexia (or its risk). Reference Age of participants (years; mean or range) Participant N (treatment; control) Impairment or problem Content of training Duration of training Brain research method Task in testing Behavioral improvement (pre-test vs. post-test) Normalization of brain activation Aylward et al. (2003) 11 10; 11 Dyslexia Linguistic awareness, alphabetic principle, fluency, reading comprehension 2 weeks (28 h) fMRI Phoneme mapping, morpheme mapping Yes Yes Eden et al. (2004) 41–44 19; 19 Dyslexia Sound awareness, establishment of the rules for letter-sound organization, sensory stimulation, articulatory feedback 8 weeks (112 h) fMRI Repeating words, sound deletion Yes Yes Gaab et al. (2007) 10 22; 23 Dyslexia FastForWord* 8 weeks (about 67 h) fMRI Pitch discrimination Yes Yes Hasko et al. (2014) 8 28 (11 improvers, 17 non-improvers); 25 Dyslexia Phoneme discrimination and orthographic knowledge; phonics training 6 months (30 h) ERP Phonological lexical decision Yes (improvers); no (non-improvers) Yes (improvers); no (non- improvers) Heim et al. (2014) 8–11 35 (12 training phonology, 7 training attention; 14 training reading); 10 Dyslexia Phonological (Würzburger Trainingsprogramm, Kieler Leseaufbau), attentional (CogniPlus, Celeco), reading (Blitzschnelle Worterkennung) 4 weeks (10 h) fMRI Reading Yes Yes Jucla et al. (2010) 9–11 24; 10 Dyslexia Phonological training; visual and orthographic training 2 months (about 16 h) ERP Visual lexical decision Yes (but also in controls) Mixed (treatment group showed a different pattern than controls) (Continued) www.frontiersin.org February 2015 | Volume 6 | Article 137 | 11 Ylinen and Kujala Neuroscience in language-related interventions Table 1 | Continued Reference Age of par- ticipants (years; mean or range) Participant N (treatment; control) Impairment or problem Content of training Duration of training Brain research method Task in testing Behavioral improvement (pre-test vs. post-test) Normalization of brain activation Keller and Just (2009) 8–10 35 treated poor readers; 12 non-treated poor readers; 25 non-treated good readers Poor reading Corrective Reading, Wilson Reading, Spell Read Phonological Auditory Training, Failure Free Reading 6 months (100 h) DTI – Yes Yes Kujala et al. (2001) 7 24; 24 Dyslexia Non-linguistic audiovisual matching 7 weeks (about 3 h) ERP Passive listening, attention directed elsewhere Yes Yes Lovio et al. (2012) 6–7 10; 10 Difficulties in reading- related skills GraphoGame: letter–sound correspondences (vs. number-knowledge game for controls) 3 weeks (3 h) ERP Passive listening, attention directed elsewhere Yes Yes Meyler et al. (2008) 10 23; 12 Poor reading Corrective Reading, Wilson Reading, Spell Read Phonological Auditory Training, Failure Free Reading 6 months (100 h) fMRI Sentence comprehen- sion Yes Yes Richards et al. (2000) 10–13 8; 7 Dyslexia Phonological and morphological reading instruction 3 weeks (30 h) Proton MR spec- troscopy Phonological and lexical access and a non-linguistic tone task Yes Yes (Continued) Frontiers in Psychology | Cognitive Science February 2015 | Volume 6 | Article 137 | 12 Ylinen and Kujala Neuroscience in language-related interventions Table 1 | Continued Reference Age of par- ticipants (years; mean or range) Participant N (treatment; control) Impairment or problem Content of training Duration of training Brain research method Task in testing Behavioral improvement (pre-test vs. post-test) Normalization of brain activation Richards et al. (2002) 9–12 10; 8 Dyslexia Phonological vs. morphological reading instruction 3 weeks (30 h) Proton MR spec- troscopy Phonological and lexical tasks, passive listening Yes Yes Shaywitz et al. (2004) 6–9 37 (experimental intervention); 12 (community intervention); 28 (control) Reading disability Phonological intervention: sound–symbol associations, phoneme analysis, timed reading, oral story reading, dictation (vs. community intervention in school) 8 months (50 min/day) fMRI Cross-m