MIND, BRAIN, AND EDUCATION
Teachers as Orchestrators of
Neuronal Plasticity: Effects of
Teaching Practices on the Brain
Lorie-Marlène Brault Foisy1,2† , Anna A. Matejko3,4† , Daniel Ansari4 , and Steve Masson1,2
ABSTRACT— Students’ behavioral outcomes are often
used by both researchers and teachers to evaluate the effec-
tiveness of pedagogical interventions. Extensive research
using behavioral metrics has found that some interventions
are more effective than others in certain contexts. However,
there has been less focus on how different interventions
impact the processing of academic skills at a neural level.
To explore this question, we conducted a narrative review
of literature examining two or more interventions related to
the same subject of learning. We discuss five main themes
that encompass different pedagogical practices: (1) orienting
attention toward particular features; (2) teaching a particu-
lar strategy; (3) changing the level of cognitive engagement;
(4) setting an educational context; and (5) interacting with
the learner. We provide examples of how these pedagogical
practices lead to changes in both brain and behavior. This
review provides insights into how teachers orchestrate
neural plasticity through different pedagogical choices.
In recent years, there has been considerable interest in
the idea that understanding the brain may be important
for education (Ansari & Coch, 2006; Goswami, 2006; Sig-
man, Peña, Goldin, & Ribeiro, 2014) and could inform
educational practices (Pasquinelli, 2011; Posner & Roth-
bart, 2005). Although it is not easy to bridge the theoretical
and practical gaps between education and neuroscience
1 Département de didactique, Université du Québec à Montréal, Canada
2 Laboratory for Research in Neuroeducation, Département de didac-
tique, Université du Québec à Montréal, Canada
3 Department of Pediatrics, Georgetown University, USA
4 Department of Psychology, University of Western Ontario, Canada
Address correspondence to Steve Masson, Département de didactique,
Université du Québec à Montréal, C. P. 8888, Succursale Centre-Ville,
Montréal, Québec, Canada; e-mail: masson.steve@uqam.ca
† Both the authors contributed equally to this work.
Volume 14—Number 4 © 2020 International Mind, Brain, and Education Society and Wiley Periodicals LLC
(Bruer, 1997, 2006, 2010; Samuels, 2009; Willingham, 2009),
many researchers from both cognitive neuroscience and
education have argued that a better understanding of the
brain can provide useful insights into education (Ansari,
de Smedt, & Grabner, 2012; Carew & Magsamen, 2010;
Fischer et al., 2007; Masson & Brault Foisy, 2014; The Royal
Society, 2011), and, importantly, that education can also
reveal new information about the developing brain (Bow-
ers, 2016; Pincham et al., 2014). Uncovering the biological
underpinnings of learning might help deepen our under-
standing of how children acquire a variety of academic skills
(Organization for Economic Cooperation and Develop-
ment [OECD], 2007; The Royal Society, 2011). It has also
been argued that a more comprehensive understanding of
how learning takes place in the brain could help inform
teaching (OECD, 2007). To that effect, the intention of the
field of Mind, Brain, and Education (MBE), also referred
to as Neuroeducation (Ansari et al., 2012) or Educational
Neuroscience (McCandliss, 2010), is to establish connec-
tions between education and the brain (Fischer et al., 2007;
Masson, 2012) with three main focuses: (1) examine the
biological processes that support and constrain learning;
(2) explore the neural correlates of academic skills; and (3)
investigate how teaching practices or educational inter-
ventions impact brain function and/or structure. While
much emphasis has been placed on learning and the brain,
there has been comparatively less discussion on how dif-
ferent kinds of teaching practices shape learners’ brains. In
this paper, we provide a narrative review of how different
teaching practices can lead to distinct neural outcomes.
First, we briefly discuss how neuroscience has informed
our understanding of how children learn academic skills.
We then examine how teaching practices can impact how
information is learned.
Learning and the Brain
The concept of neuroplasticity (i.e., the capacity of brain
function and structure to change through learning and
415
 Effects of Teaching Practices on the Brain
experience, see Voss, Thomas, Cisneros-Franco, and de
Villers-Sidani (2017) for a review) links the fields of edu-
cation and neuroscience (Masson & Brault Foisy, 2014;
McCandliss, 2011). Neuroplasticity explains how learning
leads to changes in brain function (Ansari, Garcia, Lucas,
Hamon, & Dhital, 2005; Dehaene et al., 2010) and brain
structure (Draganski et al., 2004; Kwok et al., 2011; Maguire
et al., 2000). Neuroimaging and electrophysiological tech-
niques are essential tools that can help elucidate the neural
changes that arise from education (for a review of strengths
and weaknesses of different neuroimaging techniques to
assess learning, see Matejko & Ansari, 2012 and Vogel,
Matejko, & Ansari, 2016). Many studies have now provided
critical information about the neural correlates of aca-
demic skills such as reading (Cohen et al., 2003; Dehaene
et al., 2010; Dehaene, Clec’H, Poline, Bihan, & Cohen, 2002;
Maurer, Blau, Yoncheva, & McCandliss, 2010; Maurer, Bran-
deis, & McCandliss, 2005; Schlaggar & McCandliss, 2007),
and mathematics (Ansari, 2008; Matejko & Ansari, 2015,
2018). A growing body of literature is also examining how
these networks change as children develop and become
more fluent in reading and math. While neuroscientific
methods cannot replace behavioral research, they can pro-
vide supplemental and converging evidence to elucidate
some of the biological processes supporting the acquisition
of these skills.
A second important concept underlying the relationship
between the brain and learning is that brain architecture
prior to learning (e.g., sulcal geometry, pre-existing neuronal
connections, brain function, etc.) can influence the learn-
ing process (Borst et al., 2016). In other words, biology can
impose constraints on learning. Pre-existing brain architec-
ture’s influence on learning is particularly evident in learning
disabilities such as dyslexia, where impairments to specific
regions of the brain prior to learning result in difficulties
learning to read (Hoeft et al., 2011). There is thus a recip-
rocal relationship between learning and the brain: learning
influences the brain because of its plasticity, and the initial
brain architecture (structure and function prior to learning)
also influences how learning takes place.
A better understanding of pre-existing neural constraints
can help inform how children learn particular academic
skills and may help explain why some teaching practices have
a more positive impact than others. For example, the brain
did not evolve for reading (Wolf, 2008), and neuroscientists
argue that learning to read requires the recruitment of neural
substrates that already perform very similar functions, such
as object recognition, and that are already well connected
to other brain areas involved in phonology and semantic
processing (Dehaene, 2008; Dehaene et al., 2002; Dehaene &
Cohen, 2007). This notion is supported by research show-
ing that the left occipito-temporal cortex (often referred
to as the “visual word form area”), becomes increasingly
416
active during reading acquisition (for a review, see Cohen
& Dehaene, 2004). Our existing brain organization may
therefore constrain the acquisition of skills such as read-
ing (Goswami, 2008). One reason why explicitly teaching
letter and sound correspondences seems to be one of the
most effective practices to teach children to read (Ehri et al.,
2001) might be because it is compatible with our brain
networks’ pre-existing structure. Indeed, some neuroimag-
ing research has shown that teaching letter-sound corre-
spondences mobilizes neural networks related to phonol-
ogy and language in the left hemisphere and facilitates con-
nections between the brain regions responsible for visual
word recognition and comprehension (Brem et al., 2010;
Yoncheva, Blau, Maurer, & McCandliss, 2010; Yoncheva,
Wise, & McCandliss, 2015). This finding illustrates how neu-
roscientific methods can help elucidate the neuroanatomical
constraints on learning, and might help provide neurobi-
ological explanations for why some teaching practices are
more effective than others.
This idea of a bidirectional relationship between learning
and the brain is important for education because it suggests
that some teaching practices may be more compatible with
the pre-existing architecture of the brain and could lead to
more efficient learning (Dehaene, 2008; Masson & Brault
Foisy, 2014).
However, much less is known to date about whether and
how different types of teaching practices impact the acquisi-
tion and processing of academic skills at the neural level. If
one aim of MBE is to contribute to the discussion on edu-
cational practices through research on the brain, it is also
essential to investigate the effects of different educational
practices on the brain.
Teaching and the Brain: Understanding the Neural
Correlates of Different Educational Practices
Neuroscientific research makes several contributions to
understanding the effects of pedagogical practices on
learning. First, MBE may help us better understand the
complexities of neural plasticity in more ecologically valid
settings. Though experimental learning studies can be
informative, they often do not capture the complexities of a
classroom environment. MBE can therefore provide insight
into the neural processes involved in classroom-based
learning, which may unfold differently than training in a
laboratory.
A second important contribution of neuroscience to
education is that it can be used to predict whether learn-
ers (children or adults) will show learning-related gains
based on brain function or structure prior to the onset
of learning (Dresler et al., 2018). Some research has also
found that neural, but not behavioral measures, can pre-
dict learning gains (Hoeft et al., 2011; Supekar, Iuculano,
Volume 14—Number 4
 Lorie-Marlène Brault Foisy et al.
Chen, & Menon, 2015), suggesting that neural measures
can sometimes have predictive power above behavioral
measures.
Third, neuroscience provides a complementary level of
analysis to behavioral evidence and can help differentiate
the effects of pedagogical practices that have similar behav-
ioral, yet different neuronal outcomes. For instance, behav-
ioral studies may show similar or no changes following two
different types of interventions, but the two interventions
may result in substantial differences at the neural level (Lee
et al., 2007; The Royal Society, 2011). This type of observa-
tion is relevant for education because it may reveal that dif-
ferent interventions lead to different types of learning, even
though the interventions are both associated with equivalent
gains at the behavioral level. This could be the case because
(1) some changes might take place in the brain before they
can be noticed behaviorally and (2) behavioral measures
such as accuracy or reaction time are not always truly rep-
resentative of learning. For example, a large body of litera-
ture (e.g., see the seminal paper of Viennot, 1978) has shown
that while a student can perform well at a test in science
by learning how to perform a particular algorithm, that stu-
dent may not have a real conceptual understanding of the
scientific phenomenon. Using neuroscience to understand
learning in such instances can lead to unique insights into
whether behavioral changes are reflective of actual concep-
tual changes (Brault Foisy, Potvin, Riopel, & Masson, 2015;
Vaughn, Brown, & Johnson, 2020).
Finally, neuroscientific research can provide a deeper
understanding of the representational changes (changes in
the way information is processed) that occur as a function
of instruction (e.g., in the case of reading, a representational
change might be a shift from relying on letter-sound corre-
spondences compared to whole-word reading). Neuroscien-
tific methods can be used to examine how different teaching
practices or interventions lead to different neural correlates.
While these methods cannot alone determine which peda-
gogical strategy is more effective in absolute terms, it can
inform whether these teaching practices differently affect
brain function or structure. Examining the neural level of
analysis is therefore another way of querying how teaching
impacts learning. For example, does a particular teaching
strategy strengthen the functional or anatomical connec-
tions between brain regions frequently recruited for that aca-
demic skill? Does one teaching practice modulate networks
involved in reading, arithmetic, working memory, etc. more
than another teaching practice? Do different teaching prac-
tices in the same subject lead to dissimilar neural outcomes
in the learner? Together, the points discussed above highlight
how neuroscience can be a critical tool with which to assess
children’s learning and understand how teaching shapes the
learning brain.
Volume 14—Number 4
The Current Review
On a daily basis, teachers make pedagogical decisions that
impact learning. For example, they can make choices related
to the educational context (e.g., the intensity and timing of
the learning activity), how they interact with the learner (e.g.,
type of feedback they give), what content to teach (e.g., the
type of strategy used to solve a problem), or how to direct
the student’s attention to particular features of a problem.
In this review, we examine whether these types of peda-
gogical choices can have distinct impacts on the brain. Our
aim is to examine whether the way in which information is
taught can impact the way information is learned, and how
this relationship is reflected in brain function or structure.
To understand whether different educational interventions
impact the brain differently, we examined papers that inves-
tigated interventions related to the same subject of learn-
ing (e.g., two different types of reading interventions, or two
types of math instruction). This allowed us to explore if the
way information is taught can impact learning and whether
two different teaching practices produce different outcomes
in brain and behavior. Due to differences in experimental
design, it is difficult to compare teaching practices across
studies that examine only one teaching practice compared to
an unrelated control intervention or a passive control condi-
tion. A direct comparison of teaching practices within the
same study is necessary to understand how they lead to dif-
ferent learning outcomes. To our knowledge, no study has
yet integrated findings across fields to better understand how
different pedagogical practices affect the brain. By reviewing
literature that directly compares the effects of two educa-
tional interventions on the brain, we aim not only to advance
our theoretical understanding of learning in the brain, but
also to provide insights into how neuroscience can be used
to guide evidence-based instruction.
We selected papers by searching PsychInfo, Web of Sci-
ence, & Google Scholar for several keywords: interven-
tion(s), learning, teaching, reading, mathematics, science,
Magnetic Resonance Imaging (MRI), functional Magnetic
Resonance Imaging (fMRI), functional Near-Infrared Spec-
troscopy (fNIRS), EEG, ERP, and brain. We also searched the
reference sections of relevant papers. After forming an initial
list of papers, we read the abstracts and method sections to
determine whether they met several criteria: (1) compared
two or more interventions on the same subject; (2) used
a neuroscientific methodology (MRI, electroencephalogra-
phy [EEG], fNIRS, EEG, etc.) to assess the response to the
interventions; (3) the intervention trained an educationally
relevant skill (e.g., reading, math, science, language); (4) the
study did not include pharmacological intervention or neu-
rofeedback; and (5) participants were typically developing
children or adults. It is important to note that this paper
is not a systematic and exhaustive review of the literature.
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 Effects of Teaching Practices on the Brain
Rather, this narrative review uses examples from the litera-
ture to explore how different types of pedagogical practices
affect neural systems.
After examining the papers, five main themes encompass-
ing different pedagogical practices emerged: (1) orienting
attention toward particular features of a problem or situa-
tion; (2) teaching a particular strategy; (3) changing the level
of cognitive engagement; (4) setting an educational context;
and (5) interacting with the learner. In this review, we discuss
each of these themes and provide examples from the liter-
ature to illustrate how pedagogical practices are associated
with changes in the brain. The purpose of this review is not
to identify “best” pedagogical practices or to provide direct
pedagogical implications in the light of the neural results
presented. Rather, this review aims to evaluate and discuss
whether the way in which a given subject is taught can influ-
ence neural outcomes.
ORIENTING ATTENTION TOWARD PARTICULAR
FEATURES OF A PROBLEM OR SITUATION
Several studies have investigated whether orienting the
learner’s attention to different features of a problem changes
the way information is learned. Changing the attentional
focus of the learner could elicit different learning strategies,
which could subsequently lead to different behavioral and
neural outcomes. The first example concerns the develop-
ment of logic and reasoning skills, which can be influenced
by the use of misleading strategies or biases leading to sys-
tematic errors. A classic example of such type of systematic
error is illustrated in Piaget’s number conservation task
where children are asked to identify whether two rows of
tokens have the same number. Children under 7 often rely
on a misleading “length equals number” strategy that leads
them to think that the longer row of tokens contains more
tokens than a shorter one, without considering the spacing
between those tokens (Houdé & Guichart, 2001).
Several other systematic errors in logical reasoning have
been identified in the scientific literature, such as the “per-
ceptual matching bias”, which is one of the most reliable and
robust biases in the psychology of reasoning (Evans, 2003).
According to Evans (2003), a matching bias is a “tendency to
see as relevant information which matches the lexical con-
tent described in the statement about which one is reason-
ing, and conversely to neglect logically relevant information
which fails to match” (p. 456). It can thus represent an obsta-
cle in tasks requiring logical reasoning. This specific bias
was explored in an intervention study by Houdé et al. (2001)
in which they examined whether the pedagogical strategy
of shifting attentional focus might influence the capacity to
overcome a perceptual matching bias. The first type of train-
ing was a logic-only training, in which adult participants’
418
attention was oriented toward the logical rule to respect (i.e.,
the logical explanation). The second type of training was a
logicoemotional training in which participants were given
instructions about the logical task, but were also explicitly
alerted to the potential trap (the misleading strategy) in the
task. Their attention was thus directed toward the difficulty
they had to overcome in order to solve the task. Results of
this study show that the logicoemotional training was the
only type of instruction to induce a significant shift in perfor-
mance. Indeed, participants who received this kind of train-
ing were able to respond logically on more than 90% of the
items presented in the task after the training, whereas the
logic-only group still exhibited the initial bias for more than
90% of the items. It thus appears that in this situation, direct-
ing the attention of the learner toward the logical explanation
only would not be sufficient to trigger a biased-to-logical
shift. The two types of training compared in the Houdé
et al. (2001) study also resulted in the recruitment of dif-
ferent brain networks. Specifically, logicoemotional training
was associated with greater activation in a right ventrome-
dial prefrontal area that extended to the junction with the
anterior cingulate cortex. Prior literature has suggested that
the right ventromedial prefrontal area can be associated with
emotional and cognitive regulation (see, for example, Bush,
Luu, & Posner, 2000 and the meta-analysis of de la Vega,
Chang, Banich, Wager, & Yarkoni, 2016). One possible inter-
pretation of these results could be that directing the learner’s
attention toward the logical explanation is not sufficient to
trigger a biased-to-logical shift, which is reflected in the lack
of activation in the right ventromedial prefrontal cortex. In
this study, neuroimaging provided an explanatory hypothe-
sis of the performance differences observed in participants
after both types of interventions.
Other studies have investigated the impact of educational
interventions that direct the attention of the learner to
different features during reading. For example, a teacher
could direct attention to different unit sizes, ranging from
grapheme-phoneme (letter-sound) to whole word units.
Since attentional focus during learning might influence
brain mechanisms that are recruited during reading, some
studies have investigated how different teaching interven-
tions lead to differences in brain activity during reading
(Bitan, Manor, Morocz, & Karni, 2005; Xue, Chen, Jin, &
Dong, 2006; Yoncheva et al., 2010, 2015). In many of these
studies, participants learn new alphabets, characters, or
words (created by the researchers), placing them in a learn-
ing situation similar to that of novice readers. Participants
are then instructed to focus on different levels of the words
(for example grapheme-to-phoneme or whole-word read-
ing) when learning them. The design of these studies could
therefore isolate the influence of attentional focus during
teaching. For instance, Yoncheva et al. (2010) compared the
effect of grapheme-phoneme instructions to whole-word
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 Lorie-Marlène Brault Foisy et al.
instructions in adult participants learning an artificial script.
In the grapheme-phoneme group, the instructions explicitly
directed participants’ attention toward the letter-like figures
embedded in each visual character (similar to a line draw-
ing) and their association with phonemes in English spoken
words. The embedded letter-like figures were stacked in
a vertical manner and were evident only when instruc-
tions drew attention to them. In the whole-word group,
participants’ attention was directed toward the whole
visual character and its association with an entire English
word. After training, participants were asked to perform a
reading verification task during which EEG recording was
acquired. Both groups were successful in associating the
trained visual characters with the corresponding spoken
words during the task. At a neural level, results showed
distinct patterns of brain activity: the grapheme-phoneme
group exhibited a predominantly left-lateralized topog-
raphy in the occipito-temporal region, relative to a more
right-lateralized topography of the whole-word group. The
left occipito-temporal region, often referred to as the “vi-
sual form area” (Dehaene & Cohen, 2007), is known to be
involved in the visual recognition of written words (Cohen
et al., 2000; Dehaene et al., 2002), and its activation has
been shown to be positively correlated with reading skills
(Maurer et al., 2005, 2010; Schlaggar & McCandliss, 2007;
Shaywitz et al., 2002). Yoncheva and colleagues also repli-
cated their previous findings in a more recent study using
a within-subjects design (Yoncheva et al., 2015). Together,
these results indicate that different instructional approaches
can impact brain function differently.
Similar investigations (Bitan et al., 2005; Xue et al., 2006)
have examined the effects of different pedagogical prac-
tices on learning to read using fMRI. For example, Xue
et al. (2006) investigated how visual form, phonological, and
semantic training shaped brain activity in the fusiform cor-
tex for adults learning to read an artificial language. They
found that when the attention was oriented toward the visual
form of the characters (visual form training), brain activity
in the bilateral fusiform cortex and the left inferior occip-
ital cortex decreased following training. These regions are
both known to be associated with diverse neural pathways
of visual recognition (including objects and words). In con-
trast, phonological training led to increased activation of the
same regions. Moreover, phonological and semantic training
also led to greater activation in other regions involved in lan-
guage processing. Consequently, all types of training (visual,
phonological, and semantic) appear to make significant but
different contributions to shaping fusiform activation during
reading acquisition of a novel language.
Altogether, the results described above are relevant for
education because they highlight the central role of atten-
tional focus in learning and show that attention can be a
powerful tool for teachers (McCandliss, 2016). Indeed, the
Volume 14—Number 4
studies presented above show that teaching practices that
modulate the learner’s attention to different features can lead
to different ways of learning and subsequently elicit differ-
ent patterns of neural activity. When a teacher directs a stu-
dent’s attention during learning, it will most likely influence
how the brain works to accomplish a task and learn new
abilities. Although these results do not allow us to formu-
late direct pedagogical recommendations, they highlight the
idea that different teaching practices that differ in how they
direct the learner’s attention result in different neuronal out-
comes.
TEACHING A PARTICULAR STRATEGY
One pedagogical approach that teachers often employ to
modify the content being learned is to teach a particular
strategy to achieve a solution. This approach is particu-
larly common in mathematics where teachers can instruct
different ways to solve arithmetic or algebraic problems.
For example, the same algebra problem might be solved
by drawing pictorial diagrams or by using alphanumeric
equations (Lee et al., 2007). In one context, a particular strat-
egy may seem more efficient and effective than another
(Siegler, 2005), whereas in other contexts, different strate-
gies may lead to behaviorally equivalent outcomes (Sohn
et al., 2004). Even in cases where there are no behavioral dif-
ferences in performance, it is possible that two strategies
may elicit fundamentally different patterns of brain activ-
ity. To address this question, several studies have explored
how instructing one strategy over another leads to different
behavioral and neural effects, particularly in the context of
arithmetic and algebra.
A seminal paper by Sohn et al. (2004) demonstrated how
different methods of solving equivalent algebraic problems
resulted in similar behavioral performance but distinct pat-
terns of brain activity. The authors found that college stu-
dents were equally accurate when they solved algebraic prob-
lems in either a verbal story-problem format or a symbolic
algebraic equation format. Although it is possible that stu-
dents could have been abstracting the information from each
format to form a common representation, an investigation
of the patterns of brain activity suggested that each format
recruited different neural circuits. In particular, story prob-
lems recruited the left anterior dorsolateral prefrontal cor-
tex (Brodmann Area 9) more than the equation condition,
whereas the symbolic equation format recruited the bilat-
eral posterior parietal cortex (Brodmann Area 7) more than
the story condition. Similar findings have been observed
when instructing individuals to use either pictorial strate-
gies or alphanumeric equations to solve algebraic problems;
even though behavioral performance on these strategies was
equal, utilizing these strategies resulted in both distinct and
419
 Effects of Teaching Practices on the Brain
overlapping neural circuits (Lee et al., 2007). These findings
indicate that similar behavioral outcomes can be supported
by different neural pathways, and that using neuroscience to
examine such differences can inform our understanding of
the underlying cognitive differences that result from differ-
ent pedagogical practices.
Different pedagogical approaches can also be used to
promote one strategy over another. For instance, Delazer
et al. (2005) compared two different learning methods in
adults: learning by strategies (e.g., using a sequence of arith-
metic operations) versus learning by drill (e.g., memoriz-
ing the solution of a problem). Though both methods led
to significant improvements in speed and accuracy follow-
ing training, the authors found that accuracy was higher on
problems that were learned through using strategies com-
pared to drill, and reaction times were the same across
the learning conditions. Neuroimaging data also revealed
that these different learning strategies led to distinct pat-
terns of brain activation. Learning by strategy compared
to drill resulted in greater activation in frontal and pari-
etal regions including the bilateral cingulate gyrus, right
inferior frontal gyri, and the bilateral precuneus extend-
ing to the cuneus. In contrast, learning by drill compared
to strategies resulted in greater activation in bilateral pari-
etal cortex extending into the angular gyrus, as well as the
bilateral superior and inferior frontal gyri. These findings
indicate that different patterns of brain activity are asso-
ciated with different teaching practices, even though both
instructional practices improve performance. This discrep-
ancy is likely a result of different cognitive systems being
engaged. For instance, learning by drill may promote the use
of more memory-based systems, whereas learning by strat-
egy may result in the use of brain systems that are more
involved in working memory and numerical magnitude pro-
cessing. Wirebring et al. (2015) also found a similar pattern
of findings when teaching young adults to learn mathemati-
cal problems through repeated practice or by letting the stu-
dents create a solution method themselves. Students were
more accurate on mathematical test questions when they
created a solution method themselves and had relatively less
activation in the angular gyrus in this condition, which may
suggest less reliance on retrieving the solutions from mem-
ory. These findings indicate that how different arithmetic
problems are taught can also influence the strategies stu-
dents use, which is reflected in distinct neural activation fol-
lowing learning.
Even though most of the literature discussed here has
focused on learning calculation skills, such strategies could
be applied to other domains such as reading or science learn-
ing. These findings highlight how teaching different strate-
gies to solve the same problem can result in different cogni-
tive processes, which can be observed at the neural level of
analysis even when behavioral performance is comparable.
420
CHANGING THE LEVEL OF COGNITIVE
ENGAGEMENT
Several studies have explored different pedagogical
approaches that change the level of cognitive engage-
ment and how this is reflected at the neural level. Changing
the level of cognitive engagement could be achieved by
adjusting the complexity of the task (e.g., tracing letters
versus writing them) and may lead students to process
information in a way that leads to greater retention of infor-
mation or greater generalization to other tasks. Cognitive
neuroscience can provide insights into how changing the
level of cognitive engagement results in different neural
outcomes. For instance, James and Engelhardt (2012) exam-
ined how preliterate 5-year-old children learned letters by
either having them type letters, handwrite letters in free
form, or trace letters on a template. The brain regions that
were activated for letters learned through handwriting were
similar to those typically engaged during reading (e.g., infe-
rior frontal gyrus, posterior parietal cortex, and fusiform
gyrus), but typing and tracing letters did not have this
effect. These findings may indicate that learning letters by
handwriting them has important implications for reading
because this type of learning led to the greatest recruitment
of reading networks. These data highlight how fMRI can
provide insights into why a particular pedagogical approach
may have longer-lasting effects and why it may result in
greater generalization to other skills (e.g., in this example,
handwriting engages reading-related brain networks which
may subsequently help promote reading-acquisition).
Using pedagogical strategies to change the level of cogni-
tive engagement can be applied to multiple domains. Other
research has examined how adjusting cognitive engagement
can impact students’ understanding of scientific concepts.
Participants were taught scientific concepts and were either
presented the scientific hypothesis in a didactic fashion and
needed to understand the reasoning behind them (hypoth-
esis understanding), or were required to generate their own
hypothesis about the same scientific topic (hypothesis gen-
eration) (Lee & Kwon, 2011, 2012). These two methods differ
in the level of cognitive engagement because an immediate
solution is provided to the scientific question in hypoth-
esis understanding, whereas the student has to actively
engage creating possible solutions in hypothesis genera-
tion. Hypothesis generation and understanding had differ-
ent behavioral outcomes, where hypothesis generation led
to a stronger ability to explain hypotheses. The two types of
training also led to differences in brain activity. Hypothesis
generation led to greater activation in a set of left-lateralized
regions including the middle frontal gyrus, parahippocam-
pal gyrus, putamen, and superior and middle temporal gyri
(Lee & Kwon, 2011). Regions such as the middle frontal
Volume 14—Number 4
 Lorie-Marlène Brault Foisy et al.
gyrus may be involved in the integration of abstract infor-
mation, whereas the parahippocampal and the superior tem-
poral gyrus may be related to form contextual associations
and semantic integration, respectively. In contrast, hypoth-
esis understanding resulted in greater activation within the
right middle frontal gyrus and right precuneus, which the
authors posit may be related to causal effect recognition and
visuospatial working memory. These findings indicate that
adjusting the depth under which information is processed
can lead to distinct differences in brain function.
MODIFYING THE EDUCATIONAL CONTEXT
There are several pedagogical practices that alter the con-
text in which information is presented, which subsequently
impacts how information is learned (e.g., explicit vs. implicit
instruction, timing or frequency of instruction). Such dif-
ferences in the educational context can have a significant
impact on both behavioral and neural indices of learning.
There exist several examples where neuroscience has sig-
nificantly contributed to our understanding of how certain
modifications to the educational context may promote learn-
ing and help individuals retain information.
Second language acquisition provides an ideal example of
how the pedagogical environment can change how infor-
mation is learned. One might learn a second language in
a didactic classroom setting (i.e., explicit instruction) but
could also learn a language through more implicit instruc-
tion within an immersion setting. Though it has often been
assumed that a second language is best learned through
implicit instruction (immersion-like settings), several behav-
ioral studies have demonstrated an advantage to explicit
instruction (grammar-focused didactic instruction) or equal
gains in both kinds of settings (Norris & Ortega, 2000). Neu-
roscience has also provided additional evidence that the con-
text in which a second language is learned changes process-
ing at the neural level, even when the language is learned
to equal proficiency. In a set of studies, Morgan-Short
et al. (2010) and Morgan-Short, Steinhauer, Sanz, & Ull-
man, 2012) revealed that when adults learned an artifi-
cial language using implicit or explicit instruction, they
became equally proficient in the language. However, when
the authors measured event-related potentials (ERPs) dur-
ing syntactic violations, the two teaching strategies led to
fundamental differences at the neural level; adults who were
trained using implicit instruction had patterns of brain
activity that were more similar to native-language speak-
ers (Morgan-Short et al., 2012). Differences in brain activ-
ity do not necessarily determine which method of language
learning is better, but it does show that these two teaching
methods result in different neurobiological changes. Even
though these studies were conducted on adults, this research
Volume 14—Number 4
highlights that investigating underlying neural signatures
can provide information about the neurocognitive differ-
ences between different pedagogical techniques, even when
behavioral metrics are the same.
Another way of adjusting the educational context is by
changing the frequency with which information is pre-
sented. For example, an extensive body of behavioral liter-
ature has shown that learning information through repeated
but periodic presentation (spaced learning) is more effec-
tive than learning information all at once (massed learn-
ing) (Smolen, Zhang, & Byrne, 2016). Moreover, children
who learn information through spaced learning are more
likely to retain this information, generalize to new concepts,
and develop greater conceptual knowledge of the mate-
rial (Gluckman, Vlach, & Sandhofer, 2014; Kelley & What-
son, 2013; Vlach, 2014). Spaced learning is thought to be
more effective than massed learning because it provides
sufficient time to consolidate memory traces and provides
more opportunities for cognitive rehearsals of the infor-
mation being learned (see Smolen et al., 2016 for a more
detailed description of the cognitive theories of spaced learn-
ing). More recently, neuroscience has provided evidence to
explain this effect at the cellular level. Memory formation is
thought to occur through changes in the synaptic strength
of neurons, and spaced learning may optimize memory for-
mation by allowing the memory traces to recover after a
refractory period (see Smolen et al., 2016 for a more detailed
review). Neuroimaging and electrophysiological research
have also provided evidence for the efficacy of spaced learn-
ing at a more macrolevel (Callan & Schweighofer, 2010;
Zhao et al., 2015). These findings suggest that spaced learn-
ing, relative to massed learning, results in greater activa-
tion in the inferior frontal cortex, which could result in bet-
ter encoding through verbal maintenance rehearsal (Callan
& Schweighofer, 2010). Behavioral research converges with
neuroscientific evidence to suggest that learning informa-
tion in spaced intervals optimizes long-term memory for-
mation and leads to greater information retention in a class-
room environment (Gluckman et al., 2014; Kelley & What-
son, 2013; Sobel, Cepeda, & Kapler, 2011). This provides an
example of where neuroscience evidence converges with
existing behavioral research. Though neuroscience cannot
determine whether one teaching practice is more effective
than another, it does provide an explanation of the mecha-
nisms of a particular teaching strategy and the correspond-
ing differences in behavioral performance.
Neuroscience has also provided important insights into
how changing the educational and learning context may have
effects on the learning process, but little effect on learning
outcomes. For example, Lee, Fincham, and Anderson (2015)
investigated how different types of instruction impact brain
activity during the instructional and solution periods in
mathematical problem solving (i.e., when participants are
421
 Effects of Teaching Practices on the Brain
learning versus solving simple arithmetic computations).
Learning these mathematical problems through verbal direc-
tions (a set of instructions on how the problem should be
completed) compared to learning through examples (pre-
senting an example of how a procedure is applied to a sim-
ilar problem) had only minor behavioral differences in the
learning and solution phases. Examining brain activity dur-
ing the instruction period demonstrated that the two types
of instructions had marked differences; learning through
examples showed greater activation in prefrontal and pari-
etal regions (left lateral inferior prefrontal cortex, and left
intraparietal sulcus), whereas learning through verbal direc-
tions showed greater activation in motor and visual regions
(e.g., bilateral precentral/postcentral gyrus as well as the
bilateral cuneus and lingual gyri; for a full list of regions see
Lee et al., 2015). However, once the information was learned,
neural responses were nearly identical within the solution
phase. These findings indicate that different types of learn-
ing contexts may result in different ways of encoding infor-
mation, but once the information is encoded, the execution
of the problem appears to be similar regardless of how the
information was learned.
Together, this literature demonstrates not only how
long-standing behavioral findings can inform our under-
standing of basic neurobiological mechanisms (e.g., spaced
learning), but also how neuroscientific findings inform our
understanding of different learning environments (e.g.,
implicit vs. explicit instruction). Importantly, these studies
illustrate again how behavioral equivalence does not always
mean neural equivalence.
INTERACTING WITH THE LEARNER
Every day, teachers have countless interactions with their
students. How teachers interact with students (for example,
by providing different types of feedback) can influence how
successful learning will be (Hattie & Timperley, 2007). A cru-
cial aspect of successful learning is indeed based on the abil-
ity to process performance feedback and to adjust behav-
ior on subsequent occasions (Holroyd & Coles, 2002). It has
been demonstrated that adults adjust their behavior more
successfully than children when receiving negative feedback
on their performance (Crone, Richard Ridderinkhof, Worm,
Somsen, & Van Der Molen, 2004; Crone, Zanolie, Van Lei-
jenhorst, Westenberg, & Rombouts, 2008; Huizinga, Dolan,
& van der Molen, 2006), whereas children respond better
to positive feedback (Qu & Zelazo, 2007). An fMRI study
by Van Duijvenvoorde, Zanolie, Rombouts, Raijmakers, and
Crone (2008) specifically investigated the age-related dif-
ferences in feedback learning. Children, adolescents, and
adults were asked to perform a rule shift task. This type of Teachers can be thought of as “orchestrators” of neuronal
task assesses the ability to respond correctly to a rule and plasticity, and this review highlights the idea that the peda-
to shift from one rule to another. After first completing a gogical choices that they make can impact how we learn and
422
guess trial, participants were given positive or negative feed-
back on their performance. Following feedback, they had
to answer to a repetition trial based on the feedback they
had received on the guess trial. Behavioral results from the
three age groups showed that compared with adults, children
made more mistakes after receiving negative feedback than
positive feedback. Neuroimaging results revealed that brain
regions associated with cognitive control (dorsolateral pre-
frontal cortex, superior parietal cortex and anterior cingulate
cortex) were differentially engaged during feedback-based
learning across development, which mainly provides infor-
mation on how children and adults respectively use perfor-
mance feedback. Adults mobilized these regions after nega-
tive feedback (i.e., signals of response adjustment), whereas
young children engaged these regions after positive feed-
back (i.e., signals of response continuation). Adolescents pre-
sented no differential feedback sensitivity in these regions,
which according to the authors could indicate that a transi-
tion may occur around this age toward an increased influ-
ence of negative feedback on performance adjustment.
In line with the work of van Duijvenvoorde and col-
leagues, another study by Peters, Braams, Raijmakers,
Koolschijn, and Crone (2014) investigated age-related
changes in the frontoparietal network during feedback
learning. Using fMRI, they asked 268 participants (aged 8 to
25 years old) to perform a feedback learning task in which
they had to identify the correct location of different stimuli
on the computer screen. Behavioral results indicated that
the learning rate (number of trials to complete the task)
was positively correlated with age. Neuroimaging results
revealed that the prefrontal cortex showed more activation
following negative compared with positive feedback with
increasing age. In contrast, activity in the parietal cortex
demonstrated an age-related shift in sensitivity; there was
a greater response to positive feedback in young children,
but a greater response to negative feedback in adolescents
and adults. These findings suggest that the type of feed-
back would mobilize different regions of the frontoparietal
network depending on age.
The results of both studies are of great interest for teaching
because they offer an explanation as to how different types of
feedback may impact learning differently over development.
This idea not only expands our understanding of the mech-
anisms of feedback learning, but also has implications for
teaching in the classroom in terms of differentiated instruc-
tion across grade levels.
DISCUSSION
Volume 14—Number 4
 Lorie-Marlène Brault Foisy et al.
retain information. Importantly, understanding how peda-
gogical choices influence the brain could ultimately provide
useful insights for teaching by unveiling the neurocogni-
tive mechanisms of learning. In this narrative review, we
have provided instances where neuroscience has informed
our understanding about teachers’ pedagogical choices by
adding a complementary level of analysis. We identified five
interrelated themes of pedagogical practices where there
have been significant contributions from neuroscientific evi-
dence: (1) attention orienting; (2) teaching particular strate-
gies; (3) changing the level of cognitive engagement; (4)
modifying the educational context; and (5) interacting with
the learner. Using neuroscientific studies, we illustrated how
these pedagogical practices lead to different neural out-
comes. For all of these themes, we identified studies that
compared two different pedagogical approaches within the
same subject of learning. Such a method, as opposed to com-
paring across studies, allows for a better comparison of the
similarities and differences of these pedagogical approaches
at the neural and behavioral levels of analysis.
Several common themes cut across the literature reviewed
here. First, many of these studies highlighted that pedagogi-
cal decisions influence learning, which can often be seen in
behavioral changes, but also in changes to brain function.
For example, Delazer et al. (2005) found that teaching com-
plex arithmetic by drill or through strategies led to different
behavioral and neural outcomes. Second, there were many
instances where different teaching practices produced simi-
lar behavioral performance, but different neural signatures.
This was illustrated in Sohn et al. (2004), where individu-
als were equally accurate when presented with mathematical
problems in either a story or equation format, but very differ-
ent neural pathways were used in each format to achieve the
same answer. Similarly, Morgan-Short et al. (2010) demon-
strated that implicit instruction leads to neural signatures
more typical of one’s primary language, even when the
teaching practices have similar behavioral outcomes. Such
examples underscore the added value of using neuroscience
to understand teaching and learning, because neural changes
may not always be evident at the behavioral level but could
ultimately provide more information about the nature of the
learning taking place. Third, neuroscience can be a way to
test, validate, and understand the mechanisms of pedagog-
ical practices often employed by teachers. This was shown
in the literature of spaced learning, wherein cellular mecha-
nisms have been proposed to help explain why information
is better retained when it is presented in a spaced rather than
a massed format (Smolen et al., 2016). Even though it has
long been known that spaced learning is more efficient than
massed practice, insights into the neural underpinnings of
spaced learning still have some implications for education.
Out of a desire to implement evidence-based pedagogical
practices, discussions often revolve around “what works” or
Volume 14—Number 4
“what works best.” However, understanding “why” a practice
is more effective can also help teachers by providing them
with more information with which to make evidence-based
decisions. Indeed, teachers may be more likely to rely on ped-
agogical practices that are known to be effective if they have
a better understanding of why they are effective, which could
be achieved through neuroscientific evidence. As suggested
by Dehaene (2008), it may be beneficial for teachers to “un-
derstand the initial state, the developmental trajectory, and
the end state of the brain changes that they are trying to
teach” (p. 245).
Finally, the studies discussed in this review demonstrate
how comparing two pedagogical approaches against each
other, rather than against a dissimilar control intervention
or a “business-as-usual” group, can provide a richer under-
standing of the similarities and differences of these pedagog-
ical approaches. This method can, for instance, determine
which pedagogical approach shows the greatest changes
in brain and behavior, and whether different pedagogical
approaches lead to greater long-term gains. It can also pro-
vide insight into the cognitive changes that result from edu-
cational interventions, and into which systems are involved
in these cognitive changes. Together, these themes highlight
the ways in which neuroscience can help deepen our under-
standing of how teaching practices lead to similar or different
learning outcomes.
LIMITATIONS AND CONSIDERATIONS FOR FUTURE
RESEARCH
Although we discuss many aspects of how different peda-
gogical practices change the brain, there are some factors
that need to be taken into consideration when interpreting
this literature. First, although this literature review is narra-
tive and nonsystematic in nature, we wish to acknowledge
that the selection of articles is necessarily limited by publi-
cation bias. Considering that studies with statistically signifi-
cant results are much more likely to be published than papers
with null results (Rothstein, Sutton, & Borenstein, 2005) it
is possible that some unpublished research found no neural
differences between different pedagogical approaches. The
importance of each of these studies should therefore not
be overestimated. The purpose of this review was to select
examples from the literature to explore the idea that educa-
tional choices can have an influence on the brain. Of course,
this does not mean that this is always the case: there are most
likely studies in which, conversely, neuroscience fails to bring
valuable information to education.
Second, some interpretations of neuroimaging findings
are made through reverse inferences. A reverse inference
is where a particular cognitive process is inferred from
activation of a particular brain region during a task, without
423
 Effects of Teaching Practices on the Brain
explicitly measuring this cognitive process (Aguirre, 2003;
Hutzler, 2014; Poldrack, 2006). Reverse inferences have
been widely criticized because they are not considered
to be deductively valid (Pinal & Nathan, 2017). Although
it is undisputed that the careless use of reverse infer-
ences is a problematic practice in neuroimaging, some
researchers have argued that they can still be informative
(Poldrack, 2006) and can even have high predictive power of
the presence of a specific cognitive process (Hutzler, 2014).
Inferring a cognitive process from the observed pattern of
brain activity is not a fallacy per se, but the validity of this
practice depends on the cognitive process of interest, the
specificity of the brain region activated, and the task-setting
used (Hutzler, 2014; Poldrack, 2011). In this review, it was
not possible to discuss in great detail the a priori hypotheses
of the researchers, the design of the cognitive tasks that
were used, or even the specificity of the brain regions that
were identified in the results. It is important to keep this
limitation in mind and to approach some of these cognitive
interpretations with caution; it would not be wise to infer
that a particular educational intervention has led to a change
in a particular cognitive process from this single literature
review.
A third consideration when interpreting the literature dis-
cussed above is that some pedagogical practices may be
more effective in one specific domain (e.g., rote learning
in mathematics), whereas other pedagogical practices (e.g.,
spaced learning) may be beneficial for learning in multiple
domains. Just because one study demonstrates that a partic-
ular teaching practice changes behavior and alters brain net-
works in one domain does not mean it would be as effective
in another. Comparing these pedagogical practices across
domains would be an interesting avenue for future research.
Another caveat to the literature discussed above is that
many studies examined the effect of different teaching
practices in adults rather than in children. Children and
adults may learn material in different ways, especially
because adults can scaffold new knowledge onto previously
learned material and largely have fully developed systems
for learning and cognitive control. These types of differences
were demonstrated by Van Duijvenvoorde et al. (2008) who
showed evidence for developmental changes in the neural
response to positive and negative performance feedback.
It is also important to acknowledge that every learner has
different characteristics, capabilities, and experiences. An
intervention that works well on average may not work well
with all children. In the future, neuroscience examining
effects at both the group and individual levels might help
guide our understanding of why some children benefit from
certain interventions while others do not. An identification
and understanding of individual neural differences could
ultimately make it possible to ensure a better differentiation
424
of pedagogical practices and to respond to the specific needs
of each student in a more targeted way.
Finally, all the discussed studies have examined how
different pedagogical practices affect brain function, but
none have investigated how instruction affects brain struc-
ture, or whether brain structure and function prior to the
start of an intervention impact how information is learned.
This might be due to a traditionally held view that the
anatomical structure of the human brain does not sub-
stantially change. From this perspective, and considering
that educational interventions are generally of short dura-
tion (often a few weeks), researchers likely do not antici-
pate significant effects on brain structure and tend to focus
on interventions’ functional effects. However, some find-
ings indicate that learning-induced neuroplasticity can also
be reflected at a structural level (such as in the density
of gray matter), as a result of training over periods of
several weeks (Draganski et al., 2004) or longer (Carreiras
et al., 2009; Maguire et al., 2000; Mechelli et al., 2004), and
even after a few hours of training (Kwok et al., 2011). These
findings suggest that distinct pedagogical practices could
indeed be associated with different structural changes. Stud-
ies have also shown that the sulcal pattern of certain brain
regions, which is primarily affected by early neurodevelop-
mental factors, can predict subsequent cognitive skills such
as inhibitory control (Borst et al., 2014; Cachia et al., 2014)
and reading (Cachia et al., 2018). These differences in sul-
cation may partly explain why some students seem less
receptive than others to certain educational interventions. It
might be of interest to examine whether brain structure (in
this case the sulcation) affects the response to different edu-
cational interventions. The link between teaching and brain
structure remains a fruitful avenue for future exploration.
Future research will need to explore whether structure and
function show similar or different changes following inter-
vention, and how structure and function of the brain shape
subsequent learning.
More generally, it is important to keep in mind that while
neuroeducational work brings us closer to understanding
the neural correlates of learning in more ecologically valid
contexts, neuroscientific methods still place significant con-
straints and challenges. The first constraint is that neuro-
science studies are most often conducted in experimental
settings that bear little resemblance to the classroom (Ansari
et al., 2012; De Smedt & Verschaffel, 2010). Most of the avail-
able neuroimaging methods available (for example fMRI)
require participants to remain very still and to respond to
rigorously controlled stimuli via response pads. This con-
straint is not unique to neuroeducational research; it is also
often the case for research projects in psychology (and some-
times education) that are conducted in more controlled envi-
ronments. This limitation does not in itself invalidate the
relevance of the results obtained but must be acknowledged.
Volume 14—Number 4
 Lorie-Marlène Brault Foisy et al.
Another methodological challenge concerns the sam-
ple sizes used in neuroscientific research, which are much
smaller than those generally used in educational research.
For instance, a sample of 20–30 participants per group in
an fMRI study is often considered sufficient (Desmond &
Glover, 2002; Murphy & Garavan, 2004). However, such a
small sample size raises a number of problems for the rep-
resentativeness of results, primarily with regards to their
generalization. Moreover, small sample sizes can limit a
researcher’s ability to randomly assign participants to con-
ditions, which is considered necessary to identify and com-
pare the effects of different educational interventions. These
methodological constraints thus highlight the importance of
considering neuroscientific data alongside behavioral data.
CONCLUSIONS
Despite these methodological limitations, neuroscience
can provide both unique and complementary insights into
children’s learning and instruction. Indeed, investigating the
brain can sometimes provide information about the cogni-
tive changes in learning that may not have been predicted
by behavior alone (Hoeft et al., 2011; Supekar et al., 2015),
indicating the importance of investigating learning at multi-
ple levels of analysis. The literature reviewed here highlights
how neuroscience can be used to examine the effects of
different pedagogical choices, and how these instructional
practices can lead to drastically different outcomes in the
brain. The different neural outcomes are likely the result
of different cognitive operations being performed by the
learner, leading to different brain systems being engaged
and interacting with one another. Investigating these neu-
ral underpinnings sheds light on how different teaching
interventions lead pupils to process and retain informa-
tion differently by either strengthening existing networks
or developing connections with new brain areas. Beyond
informing our understanding of teaching, these studies also
help us understand the complexities of neuronal plasticity
and how culturally acquired skills come to be processed
within the brain. Furthermore, different pedagogical prac-
tices may fundamentally change how the brain learns
information and how receptive it is to learning. Together,
this narrative review illustrates how neuroscience can be
used to understand how different pedagogical practices
affect children’s learning, and how teachers orchestrate
neural plasticity through different pedagogical choices. We
therefore consider this review a step toward bridging the
gap between neuroscience and education.
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