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DOI: 10.4324/9781003190912-12
9
THE NEUROLINGUISTICS
OF THE SECOND LANGUAGE
MORPHOLOGICAL SYSTEM
The Role of Grammar- Related and
Speaker- Related Factors
Nicoletta Biondo, Nicola Molinaro, and Simona Mancini
Introduction and Critical Definitions
Morphemes are the smallest units that convey meaning in language. Inflectional morphemes like - i
and - ed in (1) and (2) below are combined with a stem via affixation to generate different forms of the
same lexical item, for example, a verb with different number, person, or tense features.
(1) Re- �
stem- affix1.sg.past/ ‘I laughed’
(2) Laugh- ed
stem- affix.past
A native or proficient non- native speaker of Spanish will be able to extract a composite set of
information from the morpheme - in (1), namely that a single individual laughed at some point in the
past and that this is the speaker of the utterance. Conversely, a reader with little knowledge of Spanish
may be unable to extract the full set of information provided by the verb.
How is morphological knowledge attained in a second language (L2)? Can it become native- like,
and how? We review and critically evaluate recent neuroimaging studies that have addressed these
questions, with the goal of highlighting the role that grammar- and speaker- related factors play in
shaping the L2 morphological system and its neurocognitive underpinnings. Native- like morpho-
logical knowledge refers here to the ability of L2 speakers to process the L2 as efficiently and auto-
matically as their native language. Yet, we clarify that native- likeness does not assume the existence
of a monolingual speaker norm and recognizes that L2 speakers can only become bilingual, by defin-
ition, given that they already possess the knowledge of one language (their mother tongue).
L2 learning occurs either simultaneously (e.g., in early bilinguals) or sequentially (e.g., in late
learners) with respect to the first language (L1). Understanding the (possible) cross- linguistic transfer
of rules, and the conditions under which it occurs is thus pivotal to understanding how the L2 language
system develops (e.g., MacWhinney, 2005). We refer to this factor as the L1– L2 similarity factor (see
Sabourin & Manning, this volume). L1– L2 similarity refers to structurally mappable features, that is,

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Nicoletta Biondo, Nicola Molinaro, and Simona Mancini
116
features that are present both in the L1 and in the L2 (MacWhinney, 2005). Morphological features
can also be similarly or dissimilarly realized/ distributed across constituents in the two languages.
The L1 and the L2 are considered similar here if the morphological feature at issue (e.g., number) is
present and realized on the same constituents (e.g., noun– verb, adjective– noun) in both languages.
However, both aspects of similarity are also considered separately, when possible.
Experience- related factors that pertain to the speakers’ L2 history, such as age of acquisition
(AoA), proficiency, and immersion in an L2 environment are also considered crucial during learning
(e.g., Ullman, 2014, 2020; see Luque & Covey, this volume). AoA refers to the starting point of L2
learning, i.e., when the speaker was first exposed to the language. Proficiency refers to the ability to
use a language, as measured by performance on standardized tests or task- related accuracy scores.
Immersion duration refers to the length of residence in an L2- speaking country. We review evi-
dence about all these factors, mainly from electroencephalography (EEG/ ERPs) and functional
magnetic resonance imaging (fMRI), but also briefly from non- invasive brain stimulation (NIBS),
and comment on their potential to investigate L2 morphological processing and the brain areas that
support it. Because of the few ERP studies available on L2 morphological production, our focus will
be primarily on L2 sentence comprehension studies.
Critical Issues and Theoretical Perspectives
Within neurolinguistics, historical, theoretical, and methodological issues exist in defining how L2
morphological knowledge is attained and whether it can reach native- likeness. Historically, early
experimental research on L2 processing was mainly conducted in English, a language with an
impoverished morphological system. Such findings are therefore difficult to generalize to morpho-
logically richer languages (such as Spanish, Hindi, or Finnish), in which inflection and morpho-
syntactic relations can include a wider set of features (e.g., gender, person, number, and tense) and
constituents (e.g., determiners, adjectives, as well as demonstratives, nouns, and verbs).
Theoretically, models of L2 processing differ in the relevance they attribute to linguistic and experi-
ential factors in L2 processing. Whereas proposals like the competition Model (MacWhinney, 2005,
2018) formalize and discuss how L1– L2 cross- language interaction can affect ultimate L2 attainment,
accounts like the declarative/ procedural model (Ullman, 2014, 2020; see also Ullman & Morgan-
Short, this volume) highlight the importance of experience- related factors such as L2 AoA and
immersion.
In this vein, previous reviews and meta- analyses have failed to reach a consensus on the relative
weight that grammar- and speaker- related factors have on the native- likeness of L2 morphological
processing (e.g., Caffarra et al., 2015, De Diego- Balaguer & Rodriguez- Fornells, 2010; Roncaglia-
Denissen & Kotz, 2016; Tanner et al., 2014). L1– L2 similarity has been indicated as a key factor in
some analyses (e.g., Kotz, 2009) but less crucial in others (e.g., Caffarra et al., 2015; Polczynska &
Bookheimer, 2021). The so- called critical period hypothesis (Birdsong, 2018; Hartshorne et al., 2018;
Johnson & Newport, 1989; Newport et al., 2001), and the assumption that the later the L2 is acquired
the lower its native- likeness is, gave a prominent role to AoA, although more recent work suggests
that this factor may be less crucial than previously considered (e.g., Kotz, 2009; Steinhauer, 2014;
see also Caffarra et al., 2015).
In addition to L1– L2 similarity and AoA, proficiency has been indicated as a key factor in deter-
mining the temporal dynamics of L2 morphological processing and the degree of neural convergence
between L1 and L2 (e.g., Steinhauer et al., 2009; see also Kotz, 2009; Van Hell & Tokowicz, 2010).
Yet, this factor has also been considered problematic, due to differences in its measurement across
studies and its correlation with other variables such as exposure, AoA, and immersion (Ullman, 2014;
for more on AoA and proficiency, see Fromont, this volume). Finally, the learning environment also

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Neurolinguistics of the Second Language Morphological System
117
plays a significant role in L2 final attainment. Immersive contexts have a positive impact on L2
acquisition (for a recent overview, see Jackson & Schwiter, 2019) because they provide more natur-
alistic and variegated exposure to the L2 compared to formal education settings in the L1- speaking
country. Longer immersion duration can lead to higher grammatical sensitivity (Caffarra et al., 2015).
However, the evidence available is still limited, as few studies have focused on this variable (for more
on learning context, see Bowden & Faretta- Stutenberg, this volume).
From a methodological perspective, the available neuroimaging evidence on L2 morphological
processing mainly comes from EEG/ ERP studies, arguably because this technique is more accessible
across laboratories than (more expensive) techniques such as fMRI, or does not have potential side
effects, as with NIBS. Combining the experimental evidence from these three different techniques
enables the characterization of both the time course and the neural implementation of the mechanisms
that subserve L2 morphological processing, as well as the identification of the brain substrates that
could be causally involved. This is the approach that we undertake in this review.
Thus, the goals of the current review are to provide an updated picture of the role of grammar- and
speaker- related factors based on recent neuroimaging evidence and to identify overlooked aspects
that may deepen our insight into the complexity of L2 morphological processing.
Critical Research Outcomes and Current Empirical Knowledge
Event- Related Potential (ERP) Studies
ERPs have been widely adopted in L2 research (for more on this method, see Dickson & Pelzl, this
volume). ERPs are computed by averaging the portion of the electroencephalogram (EEG) signal
that is time- locked to the presentation of a relevant stimulus (e.g., an isolated word, or a word in a
sentence). The main advantages of ERP data are the ability to record neural activity continuously,
without time delay, and with a high temporal resolution (in the millisecond range).
Sentence comprehension ERP studies offer a comprehensive picture of the role of grammar- and
speaker- related factors during L2 morphosyntactic processing. Sentences are generally presented
visually word- by- word at the center of the screen at a fixed pace, and the participant is required to
provide a grammaticality judgment at the end of each sentence. The electrophysiological activity
generated by a word containing a grammatical error (e.g., *My cat are cute) is then compared with the
activity generated by the same word in a grammatical sentence (e.g., My cats are cute).
Inflection errors tend to elicit early negativities (250– 500 ms after stimulus presentation), followed
by a late positivity peaking about 600 ms post- stimulus onset. An early negativity with anterior and
left- lateralized topography is referred to as left- anterior negativity (LAN), as opposed to the more
broadly distributed negativity that characterizes the N400 effect. These negativities are thought to
reflect automatic morphosyntactic processing (see Molinaro et al., 2011 for a review) and semantic/
lexical integration difficulties (see Lau et al., 2008 for a review). Whereas these early negativities
are not consistently reported across studies and languages, late positive- going deflections represent a
rather stable correlate of morpho- syntactic anomalies. They are typically evident in central- posterior
areas of the scalp and are often referred to as a “P600.” Functionally, P600 effects are generally
interpreted as an index of structure integration or reanalysis (see Molinaro et al., 2011 for a review).
The emergence of early negative and late positive effects has been used to delineate different
stages in the development of L2 morphosyntactic knowledge (e.g., Osterhout et al., 2004; Osterhout
et al., 2006; Steinauer et al., 2009). When the speaker has very little knowledge of the L2 grammar, no
ERP effects arise when comparing grammatically correct and incorrect sentences because the speaker
cannot detect the grammatical error. Subsequently, the speaker may start memorizing salient word/
morphological combinations (e.g., My cats are…) as unanalyzed chunks. At this stage, any violation
of the memorized pattern (e.g., *My cat are…) would result as unfamiliar, thus triggering an N400,

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Nicoletta Biondo, Nicola Molinaro, and Simona Mancini
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which is highly sensitive to word sequence probabilities. Finally, when L2 proficiency is attained, the
speaker is able to abstract morphosyntactic rules successfully, and consequently shows native- like
ERP patterns in response to violations, i.e., left- anterior or bilateral early negativities followed by
a P600. In these accounts, different stages of morphosyntactic acquisition are mainly driven by L2
proficiency.
However, in their meta- analysis, Caffarra et al. (2015) found that other factors may play a
role. Immersion duration had an impact on early ERP responses and their underlying cognitive
mechanisms: The probability of reporting a LAN effect was higher when the immersion in an L2-
speaking country was long (> 5 years). Conversely, proficiency had an impact on late mechanisms of
(morpho)syntactic processing: More P600 effects were found when L2 proficiency was high (above
75%). Interestingly, AoA only marginally affected N400 effects, and L1– L2 similarity did not play a
crucial role in any ERP effect.
Table 9.1A summarizes ERP violation studies investigating sentence- level comprehension of L2
inflection published after Caffarra et al.’s meta- analysis. Considering these recent studies, it seems
that the native- likeness of the ERP patterns changes as a function of several factors, as suggested in
previous work and reviewed below.
Grammar- Related Factors
By considering different dimensions of L1– L2 similarity (type of feature, type of relation/ configur-
ation), our review of more recent studies suggests that a finer- grained classification of L1– L2 simi-
larity can capture effects (see Alem�n Ba��n et al., 2017; D�az et al., 2016; Gabriele et al., 2021;
Martinez de la Hidalga et al., 2021) that previous work could not identify, although the evidence
still appears somewhat heterogeneous. Gabriele et al. (2021) found similar ERP responses (P600) in
response to subject– verb and adjective– noun number violations in L1 English L2 Spanish speakers,
thus suggesting that the presence/ absence of a feature in the L1 inventory plays a bigger role
compared to the morphological realization of this feature across languages. In contrast, D�az et al.
(2016) reported data from Basque suggesting that the way a feature is realized is important. They
tested two groups of L1 Spanish L2 Basque speakers with early and later L2 AoA. The violation of a
shared feature (number) that is expressed on different constituents in the L2 (object- verb) triggered
an N400 in the group of late learners and a marginal P600 in early bilinguals. Native speakers showed
P600 effects in response to these violations (D�az et al., 2011).
Speaker- Related Factors
In contexts where the L2 speakers cannot exploit consolidated L1 grammatical knowledge to process
L2 morphology, speaker- related factors seem to become relevant. In a study with L1 Polish/ Russian–
L2 German speakers, Meulman et al. (2015) showed that morphological similarity may determine
the appearance of AoA effects. They found different P600 effects as a function of AoA when pro-
cessing an L1– L2 dissimilar phenomenon (i.e., determiner– noun gender agreement; see also Nichols
& Joanisse, 2019), but not when processing a similar phenomenon (tense/ finiteness marking). These
findings suggest that when the L1 and the L2 share similar morphological properties, both early and
late learners can reach native- like processing. Conversely, when the L1 and the L2 differ morpho-
logically, native- like processing may be attained only if the L2 is acquired earlier in life.
Similarly, L2 proficiency has been shown to interact with L1– L2 similarity. For example, Alem�n
Ba��n et al. (2018) and Gabriele et al. (2021) investigated similarity effects with L1 English speakers
of L2 Spanish. When the morphological patterns differed (e.g., gender, which is not morphologically
realized in adjective– noun relations in English), only highly proficient speakers of Spanish showed

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Neurolinguistics of the Second Language Morphological System
119
P600 effects. When a feature present in both languages was manipulated (number), both low and
high- proficiency speakers showed P600 effects.
Higher proficiency is generally reported to trigger larger P600 effects (Alem�n Ba��n et al., 2018,
2021; Bice & Kroll, 2021; Gabriele et al., 2021; Nichols & Joanisse, 2019; but see Armstrong et al.,
2018). Interestingly, Alem�n Ba��n et al. (2018) compared the impact of two different measures of
proficiency, namely d- prime scores derived from a grammaticality judgment task performed during
EEG recording and overall L2 proficiency derived from standardized grammar tests. Whereas task-
related proficiency correlated with P600 amplitudes in all conditions, overall proficiency scores
did not.
Few ERP studies have investigated immersion duration effects, and just one reported a signifi-
cant effect on the amplitude of late components (Alem�n Ba��n et al., 2021). In line with Caffarra
et al. (2015), other studies that tested immersion (Table 9.1A) showed early negativities (either LAN
or N400, but see Meykadeh, Golfam, Nasrabadi et al., 2021), but only with intermediate or long
immersion duration.
Summary of ERP Findings
The ERP studies presented here show that the emergence of early negativities mirroring automatic
(native- like) processing is not solely driven by L2 morphosyntactic knowledge/ proficiency. LAN
effects have been reported in two studies testing determiner– noun violations (Caffarra et al., 2017;
Nichols & Joanisse, 2019). However, many studies have failed to find LAN effects not only in L2 but
also in native speakers, demonstrating that such findings cannot be related to a low level of morpho-
logical knowledge. N400 effects were found to mirror intermediate stages of L2 knowledge only in
some studies (D�az et al., 2016; Martinez de la Hidalga et al., 2021), whereas in other studies N400
effects (followed by P600s) mirrored native- like processing (Zawiszewski & Laka, 2020; Bice &
Kroll, 2021). The only ERP component that was reliably affected by proficiency was the P600, in line
with Caffarra et al.’s finding. However, the P600 amplitude was also reduced by L1– L2 dissimilar-
ities, later AoA, and shorter immersion duration. In other words, evidence coming from recent ERP
research shows that additional (grammar- and speaker- related) factors should be taken into account
when predicting the time course of L2 morphological processing.
Functional Magnetic Resonance Imaging (fMRI)
The slow temporal resolution that characterizes fMRI makes it unsuitable for capturing the time
course of neurocognitive processes, but its exquisite spatial resolution can give invaluable insights
into the brain areas that are actively involved during a linguistic task (for more on this method, see
Kousaie & Klein, this volume). fMRI is therefore an extraordinary resource to assess the extent of
overlap or segregation in the cortical representations of L1 and L2 and modulation of brain activation
patterns by experience- and grammar- related factors. Yet, fMRI investigations on L2 morphological
processing are still scant, and the evidence for the effect of grammar- and speaker- related variables
appears to be quite fragmented. Moreover, unlike ERPs, there is no meta- analysis or review of fMRI
studies specifically on L2 morphological processing to date. Therefore, the review below is based on
the available fMRI literature that could be identified at the time of writing this chapter.
fMRI studies investigating L2 morphological processing have mostly examined English regular
and irregular tense inflection at the single- word level. Few studies have investigated the processing
of morphosyntactic relations in sentences (e.g., subject– verb agreement, see Table 9.1B for an over-
view). Overall, the available evidence points to substantial neuroanatomical overlap between L1
and L2 morphological systems. Across studies (see Table 9.1B and references therein), consistent

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Nicoletta Biondo, Nicola Molinaro, and Simona Mancini
Table 9.1 Experimental Studies Investigating Morphosyntactic Violations
Table 9.A
Study
L1
L2
Feature
Constituents
L1– L2 AOA
PRF
IMM N400
LAN
P600
Meulman
et al.
(2015)
Polish/
Russian
German
Tense/ Finiteness
Gender
VP
Det- N
S
D
L
L
H
H
L
L
NA
NA
NA
NA
X
X (AOA)
D�az et al.
(2016)
Spanish
Basque
Case
Number
Number
NP
Obj- V
Subj- V
D
D
S
E
L
E
L
E
L
H
L
H
L
H
L
L
L
L
L
L
L
X (delayed)
X (delayed)
X
NA
NA
NA
NA
X (marginal)
X (early)
X
Alem�n
Ba��n
et al.
(2017)
English
Spanish
Number
Gender
NP- Adj
NP- Adj
D
D
L
L
H
H
S
S
X (larger)
X (smaller)
Caffarra et al.
(2017)
Basque
Spanish
Gender
Det- N
D
E
H
L
X
X
Alem�n
Ba��n
et al.
(2018)
English
Spanish
Number
Gender
NP - Adj
NP- Adj
D
D
L
L
L
H
L
H
S
S
S
S
X
X
X
Armstrong
et al.
(2018)
Chinese
English
Number
Subj- V
D
L
L
I
X
Nichols &
Joanisse
(2019)
English
French
Gender
Det- N
D
E
L
I/ L
X (AOA) X* (AOA,
PRF)
Zawiszewski
& Laka
(2020)
Spanish
Basque
Basque
Spanish
Case– ergative
Case– dative
Case– allative
Case– accusative
Case– dative
Case– allative
NP
NP
NP
NP
PP
PP
D
S
S
D
S
S
E
E
E
E
E
E
H
H
H
H
H
H
L
L
L
L
L
L
X
X
X
X
X
X*
X
X*
X*
X*

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121
Neurolinguistics of the Second Language Morphological System
Alem�n
Ba��n
et al.
(2021)
English
Spanish
Person– 1st
Person– 3rd
Subj- V
Subj- V
S
S
L
L
H
H
S
S
X (PRF)
X (IMM)
Bice and
Kroll
(2021)
Spanish
English
Number
Subj- V
S
E
H
L
X
NA
X (PRF)
Cheng et al.
(2021)
Chinese
English
Number
Subj- V
D
L
L
S
X*
Gabriele
et al.
(2021)
English
Spanish
Number
Number
Gender
Subj- V
NP- Adj
NP- Adj
S
D
D
L
L
L
L
L
L
NA
NA
NA
X (PRF)
X (PRF)
(PRF)
Liang et al
(2021)
English/
Dutch/
Italian/
Polish/
Swedish
Chinese
Aspect
VP
D
L
H
S
*
Mart�nez de
la Hidalga
et al.
(2021)
Spanish
Basque
Number
Person
Number
Person
Subj(erg)- V
Subj(erg)- V
Subj(abs)- V
Subj(abs)- V
D
D
S
S
E
E
E
E
H
H
H
H
L
L
L
L
X*
X*
*
*
X* (smaller)
X* (smaller)
X (larger)
X (larger)
Meykadeh
et al.
(2021)
Turkish
Farsi
Number
Subj- V
S
E
H
L
X
Morgan-
Short et al.
(2022)
English
Spanish
Number
Subj- V
S
L
L
NA
X
(Continued)

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Nicoletta Biondo, Nicola Molinaro, and Simona Mancini
Table 9.1B
Study
L1
L2
Feature
Constituents
L1– L2 AOA
PRF
IMM Main findings
Wartenburger
et al.
(2003)
Italian
German
Person/ Number/ Case Subject- verb
S
E/ L
H/ L
L/ I/ S L1– L2 overlap in left IFG at early
AoA. Increased IFG activation in
the late- AoA group.
Sakai et al.
(2004)
Japanese English
Tense
Single word (verb) D
L
(During
experiment
training)
L
NA
L1– L2 overlap in left IFG
Tatsuno
and Sakai
(2005)
Japanese English
Tense
Single word (verb) D
L
(During
experiment
training)
L/ H (two
groups)
NA
Less left IFG activation
corresponding to higher proficiency
Hernandez
et al.
(2007)
Spanish
English
Gender
Single word (noun) D
E/ L
H
NA
Greater activation for L2 late learners.
Lehtonen
et al.
(2009)
Finnish/
Swedish
Finnish/
Swedish
Inflected vs. non-
inflected nouns
Single word (noun) D
E
H
NA
Greater left IFG activation for Finnish
vs. Swedish inflected words
Pliatsikas
et al.
(2014)
Greek
English
Tense
Single word (verb) S
L
H
I
Overlapping activation patterns for
natives and non- natives.
L2 immersion modulates activation
difference between regular and
irregular in left IFG. L2 proficiency
modulates activation difference
between regular and irregular in left
caudate modulated.
Yan et al.
(2016)
Chinese
English
Tense
Single word (verb) D
L
H
NA
Increased activation in left IFG,
anterior/ posterior STG, MTG, IPL,
and BG for regular compared to
irregular tense.
Table 9.1 (Continued)

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Neurolinguistics of the Second Language Morphological System
123
Meykhadeh
et al.
(2021)
Turkish
Persian
Person, Number
Subject- verb
S
L
H
I
L1– L2 overlap
L1: Left pars opercularis more
sensitive to ungrammatical (relative
to grammatical conditions).
L2: Greater activation in left STG
for ungrammatical compared to
grammatical conditions.
Notes: We report the speakers’ L1, L2, morphosyntactic Feature tested, Constituents involved. L1– L2 (S: similar; D: different): “similar” when the feature at issue
was present in both languages and the violation involved the same constituents in both languages, “different” in all the other cases. AOA (E: early; L: late): “early” for
a mean AoA value below 10 years; “late” for a mean AoA above 10. PRF (L: low; H: high) was “low” for mean proficiency values below 75%; “high” for mean values
above 75%; IMM (S: short; I: intermediate; L: long) was “short” for mean length of residence in the L2- speaking country less than 2 years, “intermediate” for mean
values between 2 and 5 years, “long” for mean values above 5 years. Table 9.1 A summarizes the ERP literature. The grey boxes identify the factors that were formally
analyzed, either in traditional group- analyses or in correlation/ regression analyses in relation with ERP effect amplitudes (in this last case, we report their effect in
parentheses). In the N400/ LAN/ P600 columns X stands for the presence of an ERP effect (violation– control condition). The asterisk (*) stands for smaller amplitude
compared to native speakers. Table 9.1 B summarizes the fMRI literature. Legend: IFG: Inferior Frontal Gyrus; STG: Superior Temporal Gyrus; IPL: Inferior Parietal
Lobule; BG: Basal Ganglia; NA= not available.

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124
activation patterns have been identified both cortically, especially in left frontal regions, and
subcortically, in the left caudate nucleus, the basal ganglia, and the cerebellum.
Research on native and non- native processing offers two different functional interpretations of
the involvement of the left inferior frontal gyrus (IFG) during morphological processing. Some
proposals emphasize the domain- specific nature of the morphosyntactic computations supported by
this region (e.g., Friederici, 2017), especially with regard to the pars opercularis. In contrast, other
accounts associate this region with domain- general cognitive control mechanisms (e.g., Bornkessel-
Schlesewsky & Schlesewsky, 2013) that support linguistic processing, in light of the white- matter
tracts that connect this area with subcortical regions such as the caudate nuclei. Interestingly, studies
with bilingual speakers have functionally associated these subcortical regions with the monitoring
and control functions that are triggered when the linguistic system is required to switch between two
languages (Branzi et al., 2021; Crinion et al., 2006; Lehtonen et al., 2005; see Abutalebi, 2008 for
a review).
Grammar– and speaker- related variables can influence the amount of language control necessary to
monitor L1 interference, and thus the degree of automaticity with which L2 processing mechanisms
are performed (see Abutalebi, 2008; Polczynska & Bookheimer, 2021; Roncaglia- Denissen & Kotz,
2016 for reviews). Neurophysiologically, such factors can modulate the degree of efficiency of a
given area in supporting a specific cognitive function. In fact, the neuronal organization of a brain
region could be optimized for native language processing and its automatized mechanisms, but not
for the L2 and its more controlled computations (Abulalebi, 2008; Indefrey, 2006). The involvement
of larger neuronal populations, and so the emergence of stronger activation patterns, could there-
fore compensate for the lower neuronal and computational efficiency of the L2. As L2 processing
becomes more automatic, the neuronal organization of the involved region(s) could become more
efficient, leading to a change in the linkage between performance and strength of activation (Indefrey,
2006), i.e., a decrease in activation.
Grammar- Related Factors
Contrary to electrophysiological studies, inherently linguistic factors such as L1– L2 similarity have
received scarce attention in the fMRI literature on morphological processing. Previous reviews have
suggested that, across linguistic domains, typologically similar languages are more likely to show a
convergent neuroanatomical representation (Polczynska & Bookheimer, 2021; Roncaglia- Denissen
& Kotz, 2016). However, the studies reviewed in Table 9.1B suggest that neuroanatomical overlap
can also occur across typologically dissimilar languages. Converging patterns of activation in left
IFG regions have been found in comparisons between languages that differ in their degree of mor-
phological richness (e.g., agglutinating Finnish vs. fusional Swedish morphology in simultaneous
bilingual speakers, Lehtonen et al., 2005), or in the way a morphological feature is expressed on a
verb (e.g., tense in English L2 vs. Japanese and Chinese as L1, Sakai et al., 2004; Tatsuno & Sakai,
2005; Yan et al., 2016). These results are similar to contrasts between languages that share morpho-
syntactic features (tense in Greek and English, Pliatsikas et al., 2014; person/ number in German and
Italian, as in Wartenburger et al., 2003, and in Persian and Turkish, Meykadeh, Golfam, Batouli, &
Sommer, 2021).
To complicate matters further, in both morphologically similar and dissimilar L1– L2 pairs, the
neural overlap can be accompanied by either quantitative or qualitative differences. For example,
Lehtonen et al. (2009) showed that the processing of the complex agglutinative morphology of
Finnish yielded stronger activation of the left IFG compared to the simpler fusional morphology
of Swedish. Meykadeh et al. (2021) tested Turkish and Persian, which similarly encode person and
number agreement on the verb (and showed that within their common left fronto- temporal network,

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the left IFG was more engaged by grammatical sentences in the L1 (Turkish), whereas the left pos-
terior superior temporal gyrus was more sensitive to ungrammaticality in the L2 (Persian).
Part of the difficulty in identifying the contribution of cross- linguistic similarity to the neural
representation of L2 morphology is likely to be methodological in nature. Although L1– L2 similarity
is usually acknowledged in the rationale of the functional neuroimaging studies we reviewed, it is
neither entered in the experimental design so as to test the effect of shared vs. non- shared morpho-
logical features, nor is a precise definition of cross- linguistic (dis- )similarity provided, contrary to the
ERP studies previously described. It is therefore difficult to disentangle the effect of L1– L2 similarity
from that elicited by speaker- related variables. For example, Wartenburger et al. (2003) reported
overlapping patterns of activation between Italian L1 and German L2 (both languages similarly mark
number and gender information in determiners, nouns, adjectives, and verbs), but only at high levels
of proficiency and early AoA (i.e., when more automatic and efficient processing is expected). With a
later AoA and lower proficiency, quantitative differences between L1 and L2 emerged. Similarly, the
partial L2 specialization in left temporal areas reported by Meykadeh et al. (2021) could be modulated
by the different AoA of Persian and the different immersion that characterized the languages (natural-
istic for L1 and a mixture of naturalistic and formal for L2). These factors could affect the efficiency
and native- likeness of the response, even at very high levels of L2 proficiency. In this respect, fMRI
findings align with ERP evidence on the interaction between L1– L2 similarity and speaker- related
factors.
Speaker- Related Factors
Clearer but not fully consistent scenarios emerge for the role of proficiency and AoA. A negative
correlation is usually identified between proficiency and neural activation: As proficiency progresses
towards a native- like level, L2 activation in a given area decreases (Tatsuno & Sakai, 2005) and
becomes indistinguishable from the L1. Pliatsikas et al. (2014) reported a positive correlation between
the size of the regularity effect (regular > irregular forms) in the left caudate nucleus and the L2 pro-
ficiency of a group of late Greek- English bilinguals. As English proficiency increased, so did the
efficiency with which English morphologically complex words were processed, such that left- caudate
activation became statistically indistinguishable from that of native speakers.
When proficiency is kept constant across groups, AoA has been found to influence morphological
processing in the L2 and lead to quantitative modulations in activation patterns, as evidenced by the
greater left IFG involvement associated with categorizing gender- irregular nouns for late learners
compared to native speakers of Spanish (Hernandez et al., 2007). Likewise, highly proficient Italian–
German speakers who had acquired their L2 late (> 6 years) showed stronger left IFG activation for
the L2 compared to the L1 in response to morphosyntactic violations (Wartenburger et al., 2003). In
both studies, following Indefrey (2006), the stronger involvement of left IFG in the L2 could be due
to this region’s lower efficiency in processing a later- learned language. However, in contrast to these
studies, Pliatsikas et al. (2014) observed no quantitative differences between native speakers and
late- learners of English in the patterns of activation elicited by regular and irregular past- tense pro-
cessing in a lexical decision task. This suggests that efficient, native- like neuronal organization can
be achieved even when the L2 is acquired late.
Task- related differences may explain the seemingly inconsistent conclusions reached by the three
studies. Compared to lexical decision tasks (as in Pliatsikas et al., 2014), gender categorization or
grammaticality judgments (as in Hernandez et al., 2007, and Wartenburger et al., 2003, respectively)
involve additional rule- based mechanisms aimed at verifying inflectional consistency across lexical
items (e.g., noun– verb, determiner– noun). Such mechanisms are activated even when not explicitly

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required, as in gender- decision tasks (see Cubelli et al., 2005). Several single- word production studies
have shown that patterns of cortical activations during lexical access are modulated by the speakers’
L2 proficiency (low > high proficiency), but not by AoA, which appears to have a more sizeable
impact on rule- based morphosyntactic processing (Abutalebi, 2008; Indefrey, 2006; Wartenburger
et al., 2003). Thus, the distinct effects of AoA across the three studies could be due to the different
sensitivity that behavioral tasks have to this speaker- related factor.
To our knowledge, no hemodynamic study has so far systematically tested whether L2 immersion
duration has any effect on the neuroanatomical correlates of L2 morphological processing. Of the
eight functional neuroimaging studies included in this review, only three report their participants
to have been naturalistically immersed in the L2 environment, ranging from short to long periods
(Meykhadeh, Golfam, Nasrabadi et al., 2021; Pliatsikas et al., 2014; Wartenburger et al., 2003).
Both Pliatsikas et al. (2014) and Wartenburger et al. (2003) found L1– L2 overlap with increasing
immersion, thus suggesting that naturalistic exposure can guide the L2 towards reaching a native-
like neural representation. This is in line with ERP studies showing that learning in immersion- like
contexts leads to electrophysiological signatures more fully typical of native speakers (Morgan- Short
et al., 2012). However, more research is necessary to determine whether immersion duration is a
strong and independent predictor of the cortical representation of L2 morphology (for the effect of
immersion on volumetric brain changes, see Korenar & Pliatsikas, this volume).
Summary of fMRI Findings
The findings reviewed above suggest that L1– L2 similarity alone is not a robust and independent
predictor of the neural representation of L2 morphology. However, research that systematically
manipulates this factor or uses techniques with the finest spatial resolution, such as invasive brain
stimulation in neurosurgical studies, would confirm this conclusion. Speaker- related factors have the
most tangible effects on the neural representation of L2 morphology. However, as will be discussed
below, when examining the role of such variables and their predictive power, it is important to con-
sider the degree of collinearity that exists between AoA, proficiency, and immersion duration (i.e.,
early exposure to the L2 is usually associated with naturalistic immersion, thus higher proficiency).
Current Trends and Future Directions
Native- likeness refers to the ability of speakers to process stimuli in an L2 (almost) as efficiently and
automatically as in their native language. The degree of overlap between L1 and L2 electrophysio-
logical and neuroanatomical correlates is taken to be a marker of the level of native- likeness achieved
by a group of speakers. The greater their similarity, the more native- like, and so the more efficient
and automatic, L2 processing is assumed to be. Our review shows that quantitative and qualitative
modulations relative to native patterns of reference can depend on grammar- and speaker- related
factors.
Inherent to the definition of native- likeness provided above is a controversial assumption that
characterizes the profile of native speakers, namely their homogeneity in terms of processing routines,
as well as the homogeneity of the electrophysiological and hemodynamic responses to stimuli in
their native language. A legitimate question is whether it makes sense to consider native speakers as
a homogeneous group, given the increasing bi/ multilingual status and globalization of the world’s
population. Many studies have suggested reconsidering the “gold standard” for nativelikeness in
ERPs and investigating L1 and L2 processing along a continuum (e.g., Tanner et al., 2013, 2014;
see also Freunberger et al., 2022). Future challenges in the field could therefore be to identify the
variables that can account for inter- individual variability both in native and non- native processing, as

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well as the appropriate methodology to investigate and account for such variability (see for example,
Fromont, and Luque & Covey, this volume).
Factors Affecting Morphological Processing
Our review suggests that to date, none of the different grammar- and speaker- related factors considered
here are independent predictors of L2 native- likeness. This is partly due to differences in the depth
with which the different techniques have been used to examine the effects of specific predictors, as in
the case of L1– L2 similarity and proficiency, as well as the number of studies adopting these different
methodologies to investigate L2 morphosyntactic processing.
ERP research has proceeded as far as to investigate (dis- )similarities in single features and the
position in which they are expressed within morphosyntactic relations across languages. In contrast,
fMRI studies have predominantly focused on the regular vs. irregular past tense opposition and how it
is handled across broadly defined typologically similar or dissimilar languages. Future hemodynamic
studies should therefore go beyond single- word processing and assess L1– L2 (dis- )similarities at
the sentence level. In this respect, a fruitful testing ground is the contrast between morphological
features and their presence/ absence in L1 and L2 languages, thus extending and complementing
existing single- word (e.g., Hernandez et al., 2007) and sentence- level L1 studies (Carreiras et al.,
2015; Mancini et al., 2017; Qui�ones et al., 2014, 2018).
Like L1– L2 similarity, proficiency encompasses distinct components— such as global and task-
related proficiency— that can differentially affect the response of interest. Empirical evidence
concerning the impact that distinct proficiency assessments have on morphological processing comes
from ERP research, while to the best of our knowledge, fMRI studies have not addressed this issue.
Task- related proficiency reflects a specific and less naturalistic measure of proficiency, which, unlike
global proficiency, cannot provide a complete measure of the real linguistic abilities of L2 speakers
in everyday life. Consequently, the role that this type of proficiency has in the modulation of late ERP
components should be interpreted with more caution. One possibility, albeit more radical, would be
to abandon the general concept of proficiency and rely on more specific indices of L2 experience such
as the type and amount of L2 exposure and immersion (e.g., Ullman, 2014) and/ or the frequency of
L2 use (e.g., Martinez de la Hidalga et al., 2021; Osterhout et al., 2006).
A Note on Non- Invasive Brain Stimulation
Research on non- invasive brain stimulation (NIBS) investigates the effects that magnetic and electric
stimulation, namely TMS (transcranial magnetic stimulation) and tDCS (transcranial direct current
stimulation), have on participants’ performance (see Pandža, this volume). In language research,
participants are generally asked to produce a word/ sentence or to perform a metalinguistic task (e.g.,
grammaticality judgments) while specific brain regions are stimulated. The stimulation can either
have disruptive effects, mirrored by longer reaction times and/ or lower task accuracy, or it can posi-
tively enhance the participants’ performance, leading to shorter reaction times and/ or higher accuracy.
NIBS research on L2 acquisition is still in its early stages (see Pandža, this volume). For instance,
studies of picture naming (Tussis et al., 2017) and language switching (Holtzheimer et al., 2005) have
been reported. Although these studies did not manipulate morphology or morphosyntax specifically,
their results suggest that research on L2 processing could benefit from non- invasive brain stimulation
techniques.
Available NIBS evidence on native language processing with healthy and brain- damaged speakers
points to a causal role of the left IFG in morphosyntactic processing (for a review see Maran et al.,
2022) and of the temporal lobe in the retrieval of irregular past tense verbs in English (Holland &

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Lambon- Ralph, 2010). These findings lend support to the correlational evidence provided by the
hemodynamic studies discussed above, suggesting that NIBS could provide useful insights into the
neural representation of the L1 and L2.
Final Remarks
In the introduction to this chapter, we highlighted theoretical, historical, and methodological issues
behind the lack of a comprehensive neurolinguistic account of L2 morphological processing. Our
review of recent ERP, fMRI, and NIBS literature shows that while historical limitations have been
overcome, there are still some methodological aspects that should be considered in future research.
A case in point is represented by the use of rigid categories to assess the effects of experience- related
variables (i.e., division into distinct low- and high- proficiency groups). L2 data are subject to a
high degree of variability, as it is notoriously difficult to find bilingual speakers with homogeneous
linguistic profiles, and this variability is not accounted for in the statistical analyses. New methodo-
logical approaches are thus needed (see Fromont, this volume).
Finally, ERP studies have provided a fine- grained picture of the temporal dynamics related to the
processing of different morphosyntactic features and relations, while a more fragmented scenario
emerges from fMRI and NIBS. Brain areas and networks involved during L2 morphological pro-
cessing have been identified but, to date, it is still hard to define whether these networks change as a
function of different morphosyntactic features/ relations.
Further Readings
This article attempts to provide a set of global principles that determine the neuroanatomical overlap of languages
in the brain.
Połczyńska, M.M., & Bookheimer, S.Y. (2021). General principles governing the amount of neuroanatomical
overlap between languages in bilinguals. Neuroscience & Biobehavioral Reviews, 130, 1– 14. https:// doi.org/
10.1016/ j.neubio rev.2021.08.005
This article discusses issues related to a rigid concept of native- likeness in ERP research.
Freunberger, D., Bylund, E., & Abrahamsson, N. (2022). Is it time to reconsider the “gold standard” for
nativelikeness in ERP studies on grammatical processing in a second language? A critical assessment based
on qualitative individual differences. Applied Linguistics, 43(3), 433– 452. https:// doi.org/ 10.1093/ app lin/
amab 058
Acknowledgments
The authors acknowledge funding from the Basque Government (BERC 2022– 2025 program), the Spanish
State Research Agency (BCBL Severo Ochoa excellence accreditation CEX2020- 001010- S, grants PSI2015-
65694- P, RTI2018- 096311- B- I00, and PCI2022- 135031- 2 to NM, grants RYC- 2017- 22015 and PID2020-
113945RB- I00 to SM), Marie Sklodowska- Curie grant agreement No 101028370 to NB.
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