The Shape of the ACC Contributes to Cognitive - IA de Investigación especializada en el MIT

The Shape of the ACC Contributes to Cognitive
Control Efficiency in Preschoolers

Arnaud Cachia1,2*, Grégoire Borst1,2*, Julie Vidal1,2, Clara Fischer3,
Arlette Pineau4, Jean-François Mangin3, and Olivier Houdé1,2,5

Abstracto

■ Cognitive success at school and later in life is supported by
executive functions including cognitive control (CC). The pFC
plays a major role in CC, particularly the dorsal part of ACC or
midcingulate cortex. Genes, ambiente (including school cur-
ricula), and neuroplasticity affect CC. Sin embargo, no study to date
has investigated whether ACC sulcal pattern, a stable brain feature
primarily determined in utero, influences CC efficiency in the
early stages of cognitive and neural development. Using anatom-
ical MRI and three-dimensional reconstruction of cortical folds,
we investigated the effect that ACC sulcal pattern may have on
the Stroop score, a classical behavioral index of CC efficiency, en
5-year-old preschoolers. We found higher CC efficiency, eso es,

lower Stroop interference scores for both RTs and error rates,
in children with asymmetrical ACC sulcal pattern (es decir., diferente
pattern in each hemisphere) compared with children with sym-
metrical pattern (es decir., same pattern in both hemispheres). Criti-
cally, ACC sulcal pattern had no effect on performance in the
forward and backward digit span tasks suggesting that ACC sulcal
pattern contributes to the executive ability to resolve conflicts
but not to the ability to maintain and manipulate information in
working memory. This finding provides the first evidence that
preschoolersʼ CC efficiency is likely associated with ACC sulcal
patrón, thereby suggesting that the brain shape could result in
early constraints on human executive ability. ■

INTRODUCCIÓN
Cognitive control (CC) including inhibitory control—that
es, the ability to overcome conflicts and inhibit a dominant
response—is one of the core executive functions that
enable us to resist habits or automatisms, temptations,
distractions, or interference and allow us to adapt to
complex situations by means of mental flexibility, a saber,
dynamic inhibition/activation of competing cognitive
estrategias (Diamante, 2013). The Stroop Color–Word task
(Stroop, 1935) is a seminal task designed to assess the
ability to process conflicting information, dibujo, en parte,
on CC. In the classical Stroop Color–Word task, par-
ticipants are instructed to name the color of the ink of
printed words that denote colors. In the no-conflict con-
condición, the ink colors match the colors denoted by the
palabras (p.ej., “RED” appears in red ink), whereas in the
conflict condition, the colors denoted by the words inter-
fere with the ink colors to be named (p.ej., “RED” appears
in blue ink). The conflict condition, in contrast to the no-

1CNRS U3521, Laboratory for the Psychology of Child Devel-
opment and Education, Sorbonne, París, Francia, 2Université Paris
Descartes, París, Francia, 3Computer-Assisted Neuroimaging
Laboratory, Neurospin, I2BM, CEA, Gif/ Yvette, Francia, 4Université
Caen Basse Normandie, Caen, Francia, 5Institut Universitaire de
Francia, París, Francia
*These authors contributed equally to this work.

conflict condition, typically results in increased RTs and
error rates (ERs) because of the need to inhibit irrelevant
stimulus feature (es decir., the color denoted by the word) a
focus on an alternative feature of the stimulus (es decir., el
ink color). The Stroop interference score reflect the ability
of CC to overcome perceptual and cognitive conflicts
through the inhibition of a dominant response, a saber
reading when verbal material is presented (MacLeod,
1991). Sin embargo, the Stroop Color–Word task involves
other cognitive processes, such as selective attention, estafa-
flict monitoring, perceptual, semantic interference, re-
sponse interference, and working memory (MacLeod,
Dodd, Sheard, wilson, & bibi, 2003). Functional brain-
imaging studies (Roberts & Sala, 2008; Matthews, Paulus,
Simmons, Nelesen, & Dimsdale, 2004; Arbusto, Luu, &
posner, 2000; Casey et al., 2000; Pardo, Pardo, Janer, &
Raichle, 1990) have demonstrated that the medial pFC
and more precisely the dorsal ACC, also referred to as the
midcingulate cortex (Vogt, 2009), is consistently activated
in Stroop tasks (Petersen & posner, 2012) and other tasks
that involve overriding prepotent responses, selecting re-
sponses in underdetermined contexts, or errors (Petersen
& posner, 2012). According to the conflict-monitoring
hypothesis (Botvinick, 2007; Botvinick, Más valiente, Respeto,
Carretero, & cohen, 2001), one of the core functions of the
dorsal ACC is to signal conflict in information processing
to the CC system supported through dorsolateral prefrontal
cortices. To resolve this conflict, the CC system increases

Revista de neurociencia cognitiva 26:1, páginas. 96–106
doi:10.1162/jocn_a_00459

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the activation of task-relevant information and inhibits
task-irrelevant information (see Egner & Hirsch, 2005).

From a developmental psychology perspective, ex-
ecutive functions including CC are known to support
qualities such as self-control, creativity, and reasoning
that children require to be successful in school and later
in life (Diamante, 2013). Executive function efficiency is a
better predictor of school readiness and future academic
success than intelligence quotient (Blair & Razza, 2007;
Duckworth & Seligman, 2005). Given the critical role
that executive functions play in academic achievement,
numerous studies have focused on how to improve
executive function efficiency. Diverse activities seem
to increase executive function efficiency (Diamante &
Sotavento, 2011), including school curricula (Diamante, barnett,
tomás, & Munro, 2007), attention training (Rueda,
Rothbart, McCandliss, Saccomanno, & posner, 2005),
computerized training (holmes, Gathercole, & Dunning,
2009), noncomputerized games (Mackey, Colina, Piedra, &
Bunge, 2011), aerobics (Hillman, Erickson, & Kramer,
2008), martial arts (Lakes & Hoyt, 2004), yoga, and mindful-
ness (Flook et al., 2010). Por ejemplo, Tools of the Mind—
a school curriculum for preschool kindergarten that empha-
sizes imaginary play—improves executive functions to a
larger extent than high-quality school curricula based on
literacy and thematic units (Diamond et al., 2007).

From a neuroscience perspective, studies have demon-
strated that prolonged learning and specific trainings—
leading to the improvement of cognitive efficiency—can
modify the structure (p.ej., gray matter volume, cortical
thickness) of brain areas functionally related to the pro-
cesses trained (Hyde et al., 2009; Draganski et al., 2004,
2006). Por ejemplo, adults who learned to juggle over
a 3-month period present an increase of the gray matter
volume in the visual motion area. The increase of the gray
matter volume reveals one of the neuroplasticity mecha-
nisms mediating the improvement of cognitive ability
following intense training (Draganski et al., 2004). Esto es
not limited to the motor domain; studies have demon-
strated variation of the structure of the brain in response
to intense learning for medical examinations (Draganski
et al., 2006) and in response to 15 months of musical train-
ing in early childhood—with a correlation between the
structural brain changes induced by training and behav-
ioral improvements (Hyde et al., 2009). Although there
is no study to date that has investigated the structural
change in ACC in response to CC training, estudios previos
have provided evidence that interindividual differences
in adultsʼ CC efficiency are associated with differences
in the structure of ACC, eso es, the local cortical thickness
(Westlye, Grydeland, Walhovd, & Fjell, 2011) o el local
gray matter volume (Takeuchi et al., 2012) of ACC.

Early determined anatomical features of ACC also con-
tribute to interindividual differences in adultsʼ CC efficiency.
The sulcal pattern constitutes one of these early anatomi-
cal factors ( Welker, 1988). This pattern is determined
in utero by genetic and environmental factors (Barkovich,

Guerrini, Kuzniecky, Jackson, & Dobyns, 2012; Rakic,
2004; Molko et al., 2003; Dehay, Giroud, Berland, Killackey,
& Kennedy, 1996). As opposed to quantitative measures
of the cortex morphology, such as the Gyrification Index
(Zilles, Palomero-Gallagher, & Amunts, 2013; Blanco, Su,
Schmidt, Kao, & Sapiro, 2010; Armstrong, Schleicher,
Omran, Curtis, & Zilles, 1995) or cortex structure, semejante
as the thickness, surface, and volume (Giedd et al., 2009;
Gogtay et al., 2004), that vary from childhood through
early adulthood, the sulcal pattern, a qualitative measure
of the cortex morphology, is a stable feature of the brain
anatomy apparently less affected by brain maturation,
training, and learning occurring after birth (Sun et al.,
2012). Two types of ACC sulcal pattern (Ono, Kubik, &
Abarnathey, 1990) are defined between 10 y 15 semanas
of fetal life (Blanco y col., 2010): the “single” type, con
only the cingulate sulcus, and the “double parallel” type,
with an additional paracingulate sulcus (piezas; Paus et al.,
1996). Recent functional data indicate that participants
without a PCS do not lack a particular cortical area, eso
es, “single” and `“double parallel” ACC types are homolo-
gous cortical regions (Amiez et al., 2013). The “double
parallel” is observed in 30–60% of individuals, y esto
ACC type is more frequent in the left hemisphere (Yucel
et al., 2001). In adults, asymmetry in the sulcal pattern of
CAC (es decir., the “single” type in the left hemisphere and
the “double parallel” type in the right hemisphere or vice
versa) is associated with increased CC efficiency (Huster,
Westerhausen, & Herrmann, 2011; Fornito et al., 2004)
and the increased efficiency to manage cognitive conflicts
and inhibit dominant responses as measured by the per-
formance in a Stroop Color–Word task at the behavioral
and electrophysiological level (Huster, Enriquez-Geppert,
Pantev, & Bruchmann, 2012; Huster et al., 2009). Sin embargo,
no study to date has investigated whether the sulcal
pattern of ACC already affects CC efficiency in the early
stages of cognitive and neural development.

In our study, using anatomical MRI, we investigated
whether an early neurodevelopmental constraint—that
es, the sulcal pattern of ACC—contributes to preschoolersʼ
CC efficiency as measured by their performance on
the “Animal Stroop task” ( Wright, Waterman, Prescott,
& Murdoch-Eaton, 2003)—an adaptation of the Stroop
Color–Word task for young nonreading children. En el
“Animal Stroop task” (Cifra 1), children are required to
name an animalʼs body in a no-conflict condition—that
es, the head and the animalʼs body are matched (p.ej.,
a duckʼs head on a duckʼs body)—and in a conflict con-
dition—that is, the head of the animal is replaced by the
head of a different animal (p.ej., a pigʼs head on a duckʼs
body). As in the classical Stroop Color–Word task, CC
efficiency is reflected by the difference in RTs (or ERs)
between the conflict and the no-conflict conditions.
If the sulcal pattern of ACC contributes to preschoolersʼ
CC efficiency, we expect lower Stroop interference
puntuaciones (es decir., better CC efficiency) for children with asym-
métrico (es decir., the “single” type in the left hemisphere and

Cachia et al.

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Cifra 1. Assessment of CC
efficiency using the “Animal
Stroop task.” (Left) A child
performing the “Animal Stroop
task” in the classroom. (Right)
Example of “conflict” and
“no-conflict” stimuli used in
the “Animal Stroop task” to
assess the CC efficiency of
preschoolers.

the “double parallel” type in the right hemisphere or
viceversa) rather than symmetric (es decir., the “single” type
or “double parallel” type in both hemispheres) sulcal
pattern of ACC (Cifra 2).

Además, to assess the specificity of the effect of
ACC sulcal pattern on the CC efficiency of preschoolers,
the same group of children performed both forward and
backward digit span tasks from the Wechsler Intelligence
Scale for Children ( WISC-IV; see Wechsler, 2003). Nosotros
reasoned that if ACC sulcal pattern contributes specifi-
cally to the CC efficiency and not to the efficiency of
other executive functions, such as verbal working mem-
ory, then ACC sulcal pattern should have no effect on the
performance in the two-digit span tasks, even on the
backward digit span task, which requires more executive

Cifra 2. Morphological patterns of ACC. The two ACC sulcal patterns:
“single” type, with only the cingulate sulcus, and “double parallel”
tipo, with an additional PCS. ACC sulci (azul) are represented on
the cortical surface (gray/white interface).

load (Gathercole, Pickering, Ambridge, & Wearing, 2004;
Baddeley, 2003).

MÉTODOS

Participantes

Nineteen 5-year-old right-handed preschoolers (mean ±
standard deviation age: 5.47 ± 0.18 años; 11 girls) eran
recruited from a public preschool in Caen (Francia).
They had no history of neurological disease and no
cerebral abnormalities. The children were tested in ac-
cordance with national and international norms that
govern the use of human research participants. We ob-
tained written informed consent from the childrenʼs
parents that allowed us to enroll their children in the
estudiar. The ethics committee approved our study (CPP
Nord-Ouest III, Francia).

Behavioral Assessment
The preschoolersʼ CC efficiency was assessed using the
“Animal Stroop task,” an adaptation of the Stroop Color–
Word task for preschoolers ( Wright et al., 2003). Cada
preschooler performed two experimental conditions—
each comprised 24 Animal Stroop stimuli printed on a
sheet of paper. The stimuli were designed and based on
four images of animals: a cow, a duck, a pig, and a sheep.
In both conditions, the children were asked to name the
animalʼs body. In the conflict condition, the head of the
animal was substituted with the head of another animal.
In the no-conflict condition, the head and the animalʼs
body were matched (Cifra 1). All preschoolers named
each of the 24 animal bodies under the no-conflict con-
dition before naming the 24 animal bodies under the con-
flict condition. RTs and ERs were recorded separately for
the conflict and the no-conflict conditions. The Stroop
interference score was defined as the differences in RTs
or ERs between the two experimental conditions. El
higher Stroop interference scores reflected lower CC
efficiency in these children.

The verbal working memory efficiency was assessed
using the forward and backward digit span tasks from the
WISC-IV (Wechsler, 2003). In these two working memory
tareas, the children listened to a series of discrete digits
and subsequently recalled the series of digits in the same

Revista de neurociencia cognitiva

Volumen 26, Número 1

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(es decir., forward digit span task) or reverse (es decir., backward
digit span task) order of presentation. In each task, el
children first performed two series of two digits. The series
of digits were subsequently increased by one digit every
two trials. The task was terminated when a child failed
to recall two consecutive series with the same number of
digits. The working memory span (or score) was defined
as the number of correctly recalled digits in the last series.
The forward and backward digit span tasks were used to
assess the ability to maintain (es decir., forward digit span task)
or maintain and manipulate information (es decir., backward
digit span task), respectivamente, in verbal working memory.

Además, children performed a battery of nonexecu-
tive tasks: three Piagetian logicomathematical tasks (es decir.,
the number conservation task, the substance conservation
tarea, and the class inclusion task), a numerical task (es decir.,
the number estimation line task), and a visual task (es decir.,
Navonʼs local–global task). Por lo tanto, childrenʼs perfor-
mance on these tasks were not analyzed in this study.

MRI Acquisition

We acquired anatomical MRI from the Cyceron biomedical
imaging platform (Caen, Francia, www.cyceron.fr) en
a 3T MRI scanner (Achieva, Philips Medical System, El
Países Bajos), using 3-D T1-weighted spoiled gradi-
ent images (campo de visión = 256 mm, slice thickness =
1.33 mm, 128 slices, matrix size = 192 × 192 vóxeles).
To reduce motion, provide a positive experience, y
decrease wait times, we obtained MRIs as the children
passively watched a cartoon on an MRI-compatible screen
(Lemaire, Moran, & Swan, 2009).

MRI Analysis

An automated preprocessing step skull-stripped T1 MRIs
and segmented the brain tissues. No spatial normalization
was applied to MRIs to overcome potential bias that may
result from the sulcus shape deformations induced by
the warping process. The cortical folds were automatically
segmented throughout the cortex from the skeleton of
the gray matter/cerebrospinal fluid mask, with the cortical
folds corresponding to the crevasse bottoms of the
“landscape,” the altitude of which is defined by its intensity
on the MRIs. This definition provides a stable and robust
sulcal surface definition that is not affected by variations
in cortical thickness or gray matter/white matter contrast
(Mangin et al., 2004). Para cada participante, images at each
processing step were visually checked. No segmentation
error was detected. Image analysis was performed with
the Morphologist toolbox using BrainVISA 4.2 software
(brainvisa.info).

ACC Classification

The sulcal pattern of ACC was visually assessed using 3-D,
mesh-based reconstruction of cortical folds to measure

the occurrence and extent of local sulci (p.ej., leonardo,
Towler, Welcome, & Chiarello, 2009; Huster, Westerhausen,
Kreuder, Schweiger, & Wittling, 2007; Fornito et al., 2004;
Yucel et al., 2001). ACC sulcal pattern was classified as
“single” or “double parallel” type (Ono et al., 1990) based
on the presence or absence of a PCS (Cifra 2). Este
3-D approach was used to overcome methodological
issues inherent to the analysis of the sulcal pattern of
ACC from the two-dimensional sagittal slices. The PCS
was defined as the sulcus located dorsal to the cingulate
sulcus with a course clearly parallel to the cingulate
sulcus ( Yucel et al., 2001; Paus et al., 1996). To reduce
the ambiguity from the confluence of the PCS and the
cingulate sulcus with the superior rostral sulcus (Paus
et al., 1996), we determined the anterior limit of the
PCS as the point at which the sulcus extends posteriorly
from an imaginary vertical line running perpendicular
to the line passing through the anterior and posterior
commissures (AC–PC) and parallel to the anterior com-
missure (Huster et al., 2007; Yucel et al., 2001). The PCS
was considered absent if there were no clearly devel-
oped horizontal sulcus elements parallel to the cingulate
sulcus and extending at least 20 mm (interruptions
or gaps in the PCS course was not taken into account
for the length measure). The finer distinction between
“present” and “prominent” PCS ( Yucel et al., 2001; Paus
et al., 1996), leading to three ACC sulcal pattern types,
was not used here because this distinction is based on
the PCS length of adult brains—that is, greater than
20 mm according to Pausʼs classification (Paus et al.,
1996) or greater than 40 mm according to Yucelʼs classi-
fication (Yucel et al., 2001) for a prominent PCS (leonardo
et al., 2009). Además, the classification of ACC mor-
phology into five categories by grouping the individual
measurements of PCS in 15-mm steps was also proposed
(Huster et al., 2007). Sin embargo, the present/prominent or
five categories of ACC sulcal pattern cannot be applied
to characterize ACC sulcal pattern of developing brains,
as brain size and PCS length increase with age. Notablemente,
the binary classification of ACC sulcal pattern (“single”/
“double parallel” type) used in our study was previously
used in a study on schizophrenia (Proporcionó, Yucel, et al.,
2006).

Statistical Analysis

We conducted separately 2 (ACC Sulcal Pattern, es decir.,
“symmetric” vs. “asymmetric”) × 2 (Stroop Condition,
es decir., “conflict” vs. “no-conflict”) mixed-design ANOVAs
on the RTs and ERs of the Stroop task. Además, nosotros
ran a 2 (ACC Sulcal Pattern, es decir., “symmetric” vs. “asym-
metric”) × 2 (Working Memory Task, es decir., “forward” vs.
“backward”) mixed-design ANOVA on the scores in
the forward and backward digit span tasks. When we
compared two means, we computed two-tailed t tests
or Welchʼs t tests in cases of unequal variances. For each
análisis, we reported the effect size either in the ANOVA

Cachia et al.

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Cifra 3. Interindividual
variability of ACC sulcal pattern.
Superimposition of the 3-D
mesh-based reconstructions of
the cingulate sulcus (turquoise)
and PCS (azul) for all children
included in the study. Sulci
were represented on the
cortical surface (gray/white
interface). The reconstructions
of the sulci of each child were
linearly aligned in a common
referential (MNI space) para
visualization purpose.

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(partial η2) or in terms of the difference of the means
(Cohenʼs d).

RESULTADOS

Participants were divided into two groups based on the
asymmetry of the sulcal pattern of ACC: 11 children with
symmetrical ACC sulcal pattern—“single” (norte = 9) o
“double parallel” (norte = 2) type in both hemispheres—
y 8 children with asymmetrical ACC sulcal pattern—
“single” type in the left hemisphere and “double parallel”
type in the right hemisphere (norte = 4) or vice versa (norte =
4; ver figura 3 for the interindividual variability of ACC
sulcal pattern). These groups were matched for age, sexo,
household income as a proxy indicator for socioeconomic
estado, scores on the Edinburgh Handedness Inventory
(Oldfield, 1971), and raw scores on the colored progres-
sive matrices of Raven as a proxy indicator for general
intelligence (Raven, Raven, & Court, 1976; ver tabla 1).
The two-way mixed-design ANOVA in the RTs demon-
strated that, irrespective of ACC sulcal pattern, niños
needed more time to name the animalsʼ bodies in the
conflicto (65.1 ± 28.3 segundo) than in the no-conflict (37.5 ±
10.5 segundo) condiciones, F(1, 17) = 32.57, pag < .0001, ηp 2 = .66, revealing a classical Stroop-like interference effect on the RTs. Furthermore, we found no main effect of ACC Sulcal Pattern—that is, RTs averaged over the Stroop con- ditions did not differ between children with symmetric ACC and children with asymmetric ACC, F(1, 17) = 1.67, p = .26. Finally, the difference in RTs between the conflict and the no-conflict conditions was greater for children with symmetric ACC sulcal pattern (74.2 ± 31.8 sec vs. 37.3 ± 10.8 sec) than for children with asymmetric ACC sulcal pattern (52.5 ± 17.6 sec vs. 37.7 ± 10.8 sec) as witnessed by a significant two-way interaction, F(1, 17) = 5.87, p < .025, ηp 2 = .26. Critically, the Stroop interference scores (i.e., RT difference between the conflict and no-conflict conditions) were lower; thus, CC efficiency was higher in children with asymmetrical ACC (14.9 ± 13.2 sec) than in children with symmetrical ACC (36.8 ± 22.9 sec), Welch-t (16.32) = 2.64, p < .025, d = 1.17 (Figure 4). ACC asym- metry explained 21% of Stroop interference score variabil- ity based on RTs. Finally, we note that children in the two groups required the same amount of time to name the animal body in the no-conflict condition (37.3 sec vs. 37.7 sec), t(17) = .06, p = .95 (see Table 2). A two-way mixed-design ANOVA on the ERs demon- strated a similar pattern of results. Children had more errors in the conflict (8.1 ± 4.7%) than in the no-conflict conditions (3.5 ± 3.7%), F(1, 17) = 22.37, p < .0001, ηp 2 = .57; the sulcal pattern of ACC had no main effect on the ERs, F < 1. Finally, similar to the findings for the RTs, the difference in ERs between the conflict and the no-conflict conditions was greater for children with symmetric ACC sulcal pattern (8.7 ± 4.4% vs. 2.3 ± 3.9%) than for children with asymmetric ACC sulcal pattern (7.3 ± 5.3% vs. 5.2 ± 2.9%), as demonstrated by a significant two-way interaction, F(1, 17) = 5.84, p < .025, ηp 2 = .26. Table 1. Demographic Characteristics of the Sample of Preschoolers (n = 19) Age (years) Sex (female/male) Household (low/high income) Handedness (Oldfield score) Raven (raw score) Sym. n = 11 5.48 (0.2) 5/6 5/6 91.9 (26.8) 27.9 (3.2) Asym. n = 8 5.47 (0.15) 5/3 6/2 94.4 (15.7) 29.3 (1.1) Welch-t/χ2 t = 0.10 χ2 = 0.12 χ2 = 1.66 t = .025 t = .087 p .91 .73 .19 .80 .40 100 Journal of Cognitive Neuroscience Volume 26, Number 1 Figure 4. Asymmetry of ACC and cognitive control efficiency in preschoolers (n = 19). Average Stroop interference scores (RTs and ERs) in preschoolers with symmetrical ACC (“single” type or “double parallel” type in both hemispheres; light gray; n = 11) or asymmetrical ACC (“single” type in the left hemisphere and “double parallel” type in the right hemisphere or vice versa; hashed; n = 8). Error bars denote SEM. Consistent with the results reported on the RTs, the Stroop interference scores computed on the ERs were lower in children with asymmetrical ACC sulcal pattern (2.1 ± 3.9%) than in children with symmetrical ACC sulcal pattern (6.4 ± 3.9%), Welch-t (15.32) = 2.42, p < .05, d = 1.1. As for the RTs, 21% of the variance of the Stroop interference score computed on the ERs is explained by the sulcal pat- Table 2. Mean (M ) and Standard Deviation (SD) of the RTs and ERs in the Stroop Color Word Task for Children with Symmetrical (Sym.) and Asymmetrical (Asym.) Sulcal Pattern of ACC Sym. Asym. M SD M SD 37.3 74.2 36.8 2.3 8.7 6.4 10.8 31.8 22.9 3.9 4.4 3.9 37.6 52.5 14.9 5.2 7.3 2.1 10.7 17.6 13.4 2.9 5.3 3.9 RTs (sec) No conflict Conflict Stroop score ERs ( %) No conflict Conflict Stroop score tern of ACC. As for the RTs, children with asymmetrical (5.2 ± 2.9%) and symmetrical (2.3 ± 3.9%) sulcal pattern of ACC committed approximately the same number of errors in the no-conflict condition, t(17) = 1.79, p = .09. A 2 (ACC Sulcal Pattern, i.e., symmetric vs. asymmetric) × 2 (Working Memory Task, i.e., forward vs. backward) mixed- design ANOVA revealed that irrespective of ACC sulcal pattern, the scores in the backward digit span tasks were smaller (M = 2.32 ± .58) than those obtained in the forward (M = 3.79 ± .92) digit span tasks, F(1, 17) = 30.85, p < .0001, ηp 2 = .65. There was no main effect of ACC Sulcal Pattern on the scores averaged over the two conditions, F < 1. Critically, the differences between the forward and backward scores were similar for children with symmetric (4.09 ± 1.04 vs. 2.18 ± .41) and asymmetric (3.38 ± .52 vs. 2.5 ± .76) ACC patterns, as the lack of sig- nificant two-way interactions suggests, F(1, 17) = 4.26, p = .06. Moreover, planned comparisons revealed that the scores did not differ between the children with sym- metric and asymmetric ACC sulcal patterns in the for- ward, t(17) = 1.77, p = .09, and backward, t(17) = 1.19, p = .25, digit span tasks. DISCUSSION As expected, children with an asymmetrical sulcal pat- tern of ACC are less sensitive to the interference in a Cachia et al. 101 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / e j d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 6 / 1 2 6 9 / 6 1 1 / 9 9 4 6 6 / 9 1 2 7 3 8 o 0 c 4 n 4 _ 6 a / _ j 0 o 0 c 4 n 5 _ 9 a p _ d 0 0 b 4 y 5 g 9 u . e p s d t o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j / t . f u s e r o n 1 7 M a y 2 0 2 1 Stroop-like task than children with a symmetrical sulcal pattern of ACC. Thus, preschoolersʼ CC efficiency includ- ing inhibitory control—that is, the ability to overcome a cognitive conflict and inhibit a dominant response—is directly related to their ACC sulcal pattern. Critically, the effect of the sulcal pattern of ACC on the Stroop interference score unlikely results from a difference in the ability to name the animals, given that we found no difference in the no-conflict condition between the two groups. Furthermore, we note that analyses of the RTs and ERs revealed a similar pattern of results, with strong effect sizes providing evidence (a) of the robustness of the behavioral effects of ACC sulcal pattern, despite the sample size, and (b) the lack of speed accuracy trade-off. Moreover, ACC sulcal pattern had no effect on perfor- mances in the forward and backward digit span tasks. Taken together, the results suggest that ACC sulcal pat- tern contributes to the ability to resolve conflicts (i.e., as in the Animal Stroop task) but did not contribute to the ability to maintain and manipulate information in verbal working memory (i.e., as in the forward and the backward digit span tasks) or manage the increasing dif- ficulty of a task (i.e., the difference between the scores in the backward and the forward digit span tasks). A potential limitation of this study is the small sample size, reflecting the difficulty to conduct brain imaging studies in young children. This study focuses on the general construct of execu- tive functions and the inhibition of prepotent responses; therefore, our findings do not provide information on the specific cognitive processes affected by ACC sulcal pattern. Indeed, although ACC is consistently activated in the Stroop task (Roberts & Hall, 2008; Nee, Wager, & Jonides, 2007), the precise role of this region remains elusive (Botvinick, Cohen, & Carter, 2004; MacLeod & MacDonald, 2000). ACC is critical for monitoring conflicts (Kerns et al., 2004; Botvinick et al., 2001; Carter et al., 2000), selecting responses in underdetermined contexts (Palmer et al., 2001), detecting errors (Braver, Barch, Gray, Molfese, & Snyder, 2001; Falkenstein, Hoormann, Christ, & Hohnsbein, 2000; Carter et al., 1998), making reward- based decisions (Nieuwenhuis, Yeung, Holroyd, Schurger, & Cohen, 2004; Bush et al., 2002), and encoding cogni- tive efforts (Rushworth, Walton, Kennerley, & Bannerman, 2004; Botvinick et al., 2001). Additional studies are needed to determine the precise cognitive process affected by the morphology of ACC, which in turn might shed light on the cognitive processes critical to perform the Animal Stroop task and the role of ACC in supporting these pro- cesses. For instance, future researches should study the effect of ACC sulcal pattern on the performance in differ- ent executive tasks tackling different types of executive processes. Thus, our results support the hypothesis that CC efficiency in preschoolers is rooted in early neurodevel- opmental processes. The cortical folding patterns are primarily determined in utero ( Welker, 1988) and are robust to changes induced by maturation after birth and experience-dependent factors (Sun et al., 2012). The sulcal pattern results from early neurodevelopmental processes, starting as early as 10 weeks of fetal life, that shape the cortex anatomy from a smooth lissencephalic structure to a highly convoluted surface (Welker, 1988). In particular, the development of ACC sulcal pattern occurs between 10 and 15 weeks of fetal life (Feess- Higgins & Larroche, 1987; Chi, Dooling, & Gilles, 1977). Such long-term effect of early neurodevelopmental constraints on the subsequent development of cognitive capacities is in line with previous studies in adults showing that variations in ACC sulcal patterns are related to indi- vidual differences in CC efficiency (Huster et al., 2009; Fornito et al., 2004) as well as in four core temperament dimensions, that is, effortful control, negative affectivity, surgency, and affiliation (Whittle et al., 2009). An impor- tant contribution of our findings is that such long-term effect was observed for the first time in preschoolers, namely in the early stages of cognitive and neural devel- opment. However, because of the sample size, we could not statistically determine whether children with a left- ward asymmetry of the sulcal pattern of ACC (i.e., PCS in the left but not in the right hemispheres) have greater CC efficiency than children with a rightward asymmetry (i.e., PCS in the left but not in the right hemispheres) as in adults (Huster et al., 2009; Whittle et al., 2009; Fornito et al., 2004).1 Several factors contribute to the neurodevelopmental processes that influence the shape of the folded cerebral cortex (Mangin, Jouvent, & Cachia, 2010), including struc- tural connectivity through axonal tension forces (Hilgetag & Barbas, 2006; Van Essen, 1997). These mechanical constraints lead to a compact layout that optimizes the transmission of neuronal signals between brain regions (Klyachko & Stevens, 2003) and thus brain network func- tioning. In this context, we speculate that the differences in CC efficiency observed in preschoolers with symmetrical or asymmetrical ACC might reflect differences in brain network efficiency because of differences in long-range (i.e., interhemispheric) and short-range (i.e., intrahemi- spheric) brain connectivity. Increased cognitive efficiency in asymmetric brains might be associated with hemispheric specialization, as it is more efficient to transfer informa- tion between close areas within the same hemisphere rather than between distant areas distributed in the two hemispheres (Deary, Penke, & Johnson, 2010; Toga & Thompson, 2003). The association between hemispheric specialization and the asymmetry of the brain morphology is supported by studies of the corpus callosum, a large bundle of interhemispheric fibers, showing that asym- metrical brains have fewer and/or thinner fibers connecting the two hemispheres relative to more symmetrical brain, as evidenced by a reduced midsagittal area ( Witelson, 1985) and microstructural integrity measured using dif- fusion MRI (Putnam, Wig, Grafton, Kelley, & Gazzaniga, 2008). Individuals with no corpus callosum (i.e., complete 102 Journal of Cognitive Neuroscience Volume 26, Number 1 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / e j d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 6 / 1 2 6 9 / 6 1 1 / 9 9 4 6 6 / 9 1 2 7 3 8 o 0 c 4 n 4 _ 6 a / _ j 0 o 0 c 4 n 5 _ 9 a p _ d 0 0 b 4 y 5 g 9 u . e p s d t o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j . f / t u s e r o n 1 7 M a y 2 0 2 1 agenesis) exhibit an intact Stroop interference effect (Brown, Thrasher, & Paul, 2001), suggesting that the pro- cesses involved in performing the Stroop task are highly lateralized in the brain. If these processes are lateralized, then it is reasonable to expect that morphological asym- metry in the regions involved in performing the Stroop task, such as ACC, would produce better efficiency by reinforcing hemispheric specialization. This hypothesis is supported by multimodal brain imaging of the cortex morphology and the white matter connectivity in the same individuals, revealing that participants with different sulcal patterns have distinct short-range white matter con- nectivity (Leonard, Eckert, & Kuldau, 2006). Therefore, the increased CC efficiency observed in children with asymmetric ACC in our study might reflect the asymmetry of the underlying white matter connectivity of ACC. Faulty executive functions including CC can account for learning difficulties in children, such as errors, reasoning biases, and maladjustment, both in cognitive (Borst, Poirel, Pineau, Cassotti, & Houdé, 2013; Poirel et al., 2012; Houdé, 2000) and social (Steinberg, 2005) domains. Childrenʼs executive function efficiency actually predicts, for in- stance, health and professional success later in life (Moffitt et al., 2011). The predictive nature of childrenʼs executive functions including CC efficiency on cognitive and social development may be partly related to the early neuro- developmental constraints induced by the sulcal pattern of ACC (Dubois et al., 2008). However, brain–behavior associations are not fixed in all cases. For instance, previous studies reported that CC efficiency, assessed either in a visual discrimination task (Casey et al., 1997) or in a Flanker task (Fjell et al., 2012), is related to the cortical surface area of ACC (Fjell et al., 2012; Casey et al., 1997). Critically, the relationship is stronger for younger children (under 12 years old) and decreases linearly with age (Fjell et al., 2012). Because the cortical surface area of ACC is determined in part by ACC sulcal pattern (Fornito et al., 2008; Fornito, Whittle, et al., 2006), the relationship between ACC sulcal pattern and the CC efficiency reported in our study might vary with age. Hence, early neuroanatomical constraints might be overcome during cognitive development by training CC. Indeed, CC continues to mature from childhood to adolescence (Luna, 2009; Luna, Garver, Urban, Lazar, & Sweeney, 2004) and can be modified by training and practice (Diamond, 2013). Longitudinal studies in large-scale samples should investigate the possible interactions between the sulcal pattern of ACC and intense preschool interventions that have shown to improve executive functions including CC efficiency (Diamond et al., 2007). An early objective assessment of CC in children is critical, given that (a) executive function training is more beneficial for children with faulty executive function efficiency (Diamond et al., 2007) and (b) executive functions can demonstrate im- proved results in children as early as 4 or 5 years (Moffitt et al., 2011). Hence, our brain-imaging findings may ultimately contribute to shape specific educational inter- ventions designed to help children overcome their CC deficits, particularly those with symmetrical sulcal pattern of ACC who are at increased risk of developing faulty CC efficiency. In conclusion, this study provides the first evidence that preschoolersʼ CC efficiency is likely associated with ACC sulcal pattern, thereby suggesting that the brain shape could result in early constraints on human executive ability. Acknowledgments We thank the CNRS for financial support and the French Board of Education for their collaboration. 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D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / e j d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 6 / 1 2 6 9 / 6 1 1 / 9 9 4 6 6 / 9 1 2 7 3 8 o 0 c 4 n 4 _ 6 a / _ j 0 o 0 c 4 n 5 _ 9 a p _ d 0 0 b 4 y 5 g 9 u . e p s d t o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j / f t . u s e r o n 1 7 M a y 2 0 2 1 106 Journal of Cognitive Neuroscience Volume 26, Number 1 The Shape of the ACC Contributes to Cognitive image

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