Edited by: Maria Carmen Usai, University of Genoa, Italy
Reviewed by: Sandra Pellizzoni, University of Trieste, Italy; Chiara Malagoli, University of Florence, Italy
This article was submitted to Developmental Psychology, a section of the journal Frontiers in Psychology
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Musical training is an enrichment activity involving multiple senses, including auditory, visual, somatosensorial, attention, memory, and executive function (EF), all of which are related to cognition. This study examined whether musical training enhances EF in preschool children who had not undergone previous systematic music learning. This study also explored the after-effects 12 weeks after cessation of musical training. Participants were 61 preschool children from a university-affiliated kindergarten in North China. The experimental group underwent 12 weeks of integrated musical training (i.e., music theory, singing, dancing, and role-playing), while the control group performed typical daily classroom activities. The three components (inhibitory control, working memory, cognitive flexibility) of executive functions were evaluated using the Day/Night Stroop, Dimensional Change Card Sort, Dot Matrix Test, and Backward Digit Span Task. In Experiment 1, EFs were tested twice-before (T1) and after (T2) the music training. The results showed that children’s EFs could be promoted by musical training. In addition, EFs were tested again 12 weeks later after the end of the intervention (T3) in Experiment 2. We discovered that integrated musical training demonstrated a sustained promotion effect.
Executive Function (EF) refers to a family of top-down mental processes necessary for concentration, specifically when relying on instinct, intuition, or automatic processing would be ill-advised, insufficient, or impossible (
Executive Function efficiency is an important factor in ensuring physical/mental health, a key predictor of academic/career achievements, and also it plays a vital role in cognitive, social, and psychological development (
To date, researchers have used interventions such as sports, meditation, and gaming to promote children’s EF development (
A synthetic table of EFs.
Computerized training | Working memory | |
Arts training (e.g., martial arts, mindfulness practices, yoga) | Inhibitory control Working memory | |
Sports | Cognitive flexibility, Working memory, Inhibitory control | |
Music training | Cognitive flexibility, Working memory, Inhibitory control | |
Task training (e.g., the delay of gratification task: flanker, go/no-go) | Inhibitory control | |
Add-Ons to Classroom Curricula (e.g., Promoting Alternative Thinking Strategies, the Chicago School Readiness Project) | Inhibitory control Cognitive flexibility |
According to a comprehensive meta-analysis of EF training programs (
At present, there are many training methods to improve EF, but musical training has characteristics of a wide transfer effect, challenging training content, and time-consuming practice. In musical training, children train independently, showing interest, motivation, and pleasure in training. This makes musical training an appropriate method for promoting children’s EF. Compared to previous studies, musical training is more accordant with the characteristics of promoting EF development proposed by
Investigation of differences in brain structure and function between musicians and non-musicians has become an effective way to explore brain plasticity (
It is not clear which mechanisms of musical training affect cognitive ability, but many researchers believe that the transfer effect of musical training involves EF (
In the process of learning music, it is necessary to maintain a high degree of self-control, attention, and memory (
In other words, the transfer effect is determined by the extent to which musical training contributes to EF (
The developmental time point in which a person begins receiving musical training is also a key factor in how music affects individuals. Research suggests that better results are achieved if musical training begins in childhood, rather than adulthood; a finding that has been unanimously recognized (
In 1995, Schlaug et al. studied the neuroanatomical differences between musicians and non-musicians. They found that musicians’ corpora callosa were larger than those of the non-musicians. The corpus callosum is a transverse nerve fiber bundle that connects the two hemispheres of the brain. Maturity of the corpus callosum’s structure and function likely occurs in late childhood to early adolescence; this period corresponds with the development of motor control and coordinated motion (
Although there is a dearth of pre-/post-test experiments, a large body of evidence exists suggesting that children’s EF can be improved after a certain period of musical training. For example,
In the short-term, simply listening to music and structured musical training are representative of interventions studied in the field. Structured musical training generally includes the learning of musical knowledge (e.g., identifying notes, rhythm and beats) and the learning of musical skills (e.g., vocal and keyboard skills;
In the present study, we designed two experiments, based on the conceptualizations of EF proposed by
This study is different from previous studies in several ways. (1) In order to understand the level of musical development and preferences of children, aged three to six, we interviewed professional music teachers who worked in kindergartens. We also referred to previous research and designed musical training curricula (e.g., rhythm, pitch, melody, voice, and basic musical concepts) suitable for 4-year-old children. (2) Although the structure of EF is still debated, this study was based on the view put forth by
The effect of music training is normally influenced by some variables, such as social background of participants. In order to maintain the homogeneity of the variables (kindergarten living environment, daily schedule, daily activities of kindergartens, etc.), we selected two classes of children (average age of 4) in the same university-affiliated kindergarten in northern China: one for the music training group and the other for the control group (
Mean age and gender of participant groups.
Music training group | 30 | 20 | 10 | 51.39 | 4.27 |
Control group | 31 | 18 | 13 | 50.35 | 3.38 |
The age difference between the music training group and the control group was not significant
The children engaged in the training programs in one team of 45 min each (10 min for organization and 35 min of training), 5 days a week, for 12 weeks (150 min per week). The music training was based on a combination of motor, perceptual, and cognitive tasks, including training in rhythm, pitch, melody, voice, and basic musical concepts.
For short-term music training participants, the following two methods are more representative. One is to sit and listen to music and experience the Mozart Effect; this could include letting college students listen to Mozart’s double piano sonatas. The other method allows participants to perform structured music training, which generally includes the learning of music knowledge (identifying notes, rhythm, beats, etc.) and the learning of musical skills, such as vocal or keyboard skills, conducted a 4-week structured music training for 32 children aged 4 to 6 years; the training included topics on rhythm, beat, melody, sound, and basic music theory. The research results showed that after the 4 weeks of music training, the experimental group performed better on the control task than the control group (
The selected songs in this experiment are from the
Examples of music activities included in the intervention and associated areas of EFs (Second week).
Day 1 | Clef and Scale: Identify treble and C major, a minor scale | Working memory, cognitive flexibility |
Day 2 | Time signature: Listen to 3/4 beats, and be able to follow them | Working memory, cognitive flexibility |
Day 3 | Termination mark: Identify the termination token, and stop when you see the termination token | Working memory, inhibitory control |
Day 4 | Strong and weak symbol: Identify strong and weak marks(F/P), which can control sound according to strong and weak marks | Working memory, cognitive flexibility |
Day 5 | Repeated mark: Identifying repeated marks, and repeat then according to repeated mark indications | Working memory, inhibitory control, cognitive flexibility |
Examples of music activities included in the intervention and associated areas of EFs (Tenth week).
Day 1 | Solo: A young children sings alone, keeping the pitch and rhythm correct | Working memory, inhibitory control, Cognitive flexibility |
Day 2 | Rotate in turn: Different children rotate and alternately sing the same song | Working memory, inhibitory control, cognitive flexibility |
Day 3 | Sing in silent: Sing without sound | Working memory, inhibitory control |
Day 4 | Role performance: Rhythms was divided into two parts (young frog and old frog) according to pitch, the children who acted as young frog started to sing when the young frog rhythms appeared, the others acting as the old frog should wait quietly, and vice versa. | Working memory, inhibitory control |
Day 5 | Dance: Rhythmic action and action combinations, including clapping, nodding, stamping feet, and so on. | Working memory, Inhibitory control, cognitive flexibility |
The time of effective music training for children was 35 min every day. The order of each music activity was fixed. The melody was active and lively. The difficulty of weekly training is from simple to complex. It gradually increased the difficulty and the training purpose was clear. The purpose of the selected track was clear, the difficulty of music rules gradually increased, the melody was active and lively, the music was mainly about animals and daily life (themes that the children loved), and the rhythm was selected as 2/4 beats and 3/4 beats, emphasizing the enthusiasm, regularity, integration, cheerfulness, and playfulness of the music training. The content was designed to encourage the children to actively participate and experience pleasure in participating. The goal was for children to then follow the rules of music, suppress impulsive behavior, recognize and memorize music symbols, and flexibly use music symbols. All the activities of this music training program were carried out by the same Master of Musicology. This person had a solid theoretical foundation of music teaching, practical experience in early childhood music teaching, relevant knowledge of development and educational psychology, and experimental research experience in kindergarten settings. Furthermore, they could better implement the guiding ideology of music training activities and mobilize the enthusiasm of young children to participate in music training than the experimenter, who only had a psychological background.
The musical training consisted of two parts (see
Weeks 1–4: musical theory
Weeks 5–12: singing, dancing, and role-playing
The executive functions of the children from all groups were assessed twice-before (T1) and after (T2) the musical training.
The task was based on study
During the formal experiment, the experimenter presented the opposite condition to the child, this time no feedback was given. The experimenter presented a white card to the child and told the child to say “night” when the child sees the white card. Then the experimenter presented a black card to the child and asked the child to say “day” when the child sees the black card.
The experiment was performed 32 times, and the cards were presented according to a pseudo-random sequence. We recorded the number of times the participant gave the correct “day” or “night” response. In this task, the Cronbach’s Alpha was 0.60.
Before the formal experiment, we conducted a pre-experiment on 3–5-year-old children in other classes of the kindergarten. The pre-experiment results showed that the three-stage experiment was difficult, the cognitive load of the child was too heavy, and the experiment time was long. Therefore, we only carried out the two-stage experiment task, and in the formal experiment, the two-stage experiment task did not reach the ceiling-effect.
The experiment used 16 cards and 2 wooden plates. The cards were 20 cm long and 13 cm wide, each wooden plate is 11.5 cm long, 9.5 cm wide and 2 cm deep. Children had to sort the cards according to a rule involving either color or shape. They were shown cards with boats or rabbits on them, either blue or red in color. The target cards were fixed to the back of each wooden plate, one showing the image of a blue rabbit, and the other a red boat. The experimenter pointed and verbally named the two target cards. In the pre-switch phase, children were asked to sort six cards according to their color, after two demonstrations given by the experimenter. Cards were presented to the child in a pseudo-random order. In the post-switch phase, children were asked to sort the cards by shape. The test was scored according to the number of correct cards. Both the pre-switch phase and post-switch phase tasks were each scored once (
The experimental materials were 16 test cards with red, green, and blue dots. The cards were 20 cm long and 13 cm wide. The dots on the cards were randomly arranged. During the test, children were instructed to count the number of red spots on the card presented. After an initial practice session, children were presented with two cards that were facedown on the table. The experimenter then turned the first card faceup; after the child counted the red spots, this card was turned facedown and the second card was turned faceup. After counting, this card was turned facedown. The experimenter pointed to the first card and then the second, asking the child to recall the number of spots counted on each card. Administration of the test continued until the child made errors on both attempts at a particular span length. This span was recorded as the maximum number of counts recalled in the correct serial order (
Before the formal experiment, we conducted a pre-experiment on 3–5-year-old children in other classes of the kindergarten, and finally selected 1–4 digits as the numerical range of the Backward digit span task.
The experimental materials for this task were the numbers 0–9. The experiment was divided into two phases. In the practice phase, the experimenter said “1, 2” and told the child to say “2, 1,” i.e., reciting the numbers backward. In the formal experiment, the experimenter randomly selected two numbers from 0 to 9, and then let the child say the numbers backward. If the child failed, they scored 1 point. If the child was successful, they scored 2 points, and the experimenter continued on, saying 3 digits. If the child successfully recited the 3 digits backward, they scored 3 points, and then the experimenter moved up to 4 digits. The maximum number of digits used was 4 (
In this experiment, the scores of the 4 tasks were the dependent variables, and the time points (pre-test vs. post-test) and the groups (the experimental group vs. the control group) were independent variables. The experimental design was a 2 (time points: pre-test vs. post-test) × 2 (groups: the experimental group vs. the control group) two-way repeated measures ANOVA, in which the time points were the intra-group variables, and the groups were the inter-group variables. The analysis of variance mainly examined the interaction between time points and groups. In the control of unrelated variables, we took the following measures. First, we conducted a survey of two classes of children during the pre-test to ensure that there were no additional music training activities outside the kindergarten environment. Second, we applied a homogeneity test on the pre-test group to ensure both groups’ developmental level of executive function. While the experimental group underwent music training, the children in the control group engaged in free play. In addition, the daily activities were the same. Uniform requirements were imposed on all teachers, and teachers were not allowed to impose additional activities on the children. Fourth, we informed parents to control additional music training activities.
The scores of children’s executive function tasks before and after music training are shown in
The executive function task data of the experimental group and the control group.
Day/Night Stroop | 12.97(4.62) | 23.10(1.16) | 12.00(6.08) | 18.52(4.70) |
DCCS | 12.50(2.93) | 15.53(1.814) | 12.55(2.71) | 13.48(2.49) |
Dot Matrix Test | 0.83(0.87) | 1.83(0.75) | 0.77(0.99) | 0.84(0.86) |
Backward Digit Span Task | 7.30(2.09) | 8.77(1.83) | 6.52(2.31) | 6.90(2.31) |
In order to better control the experimental variables, we performed statistics on the pre-test results. The results showed that there was no significant difference in the scores of the 4 tasks [Day/Night Stroop, Dimensional Change Card Sort (DCCS), Dot Matrix Test, Backward Digit Span Task] between the experimental group and the control group (
The results of the 2 (time points: T1 vs. T2) × 2 (groups: the experimental group vs. the control group) two-way repeated measures ANOVAs are shown in
Analysis of variance analysis before and after music training.
Day/Night Stroop | Time | 1 | 2113.117 | 133.398∗∗∗ | 0.693 |
Group | 1 | 234.851 | 9.257∗∗ | 0.136 | |
Time × Group | 1 | 99.740 | 6.296∗∗ | 0.096 | |
DCCS | Time | 1 | 120.73 | 26.996∗∗∗ | 0.314 |
Group | 1 | 30.525 | 3.691Δ | 0.59 | |
Time × Group | 1 | 33.548 | 7.543∗∗ | 0.113 | |
Dot Matrix Test | Time | 1 | 26.196 | 19.221∗∗∗ | 0.246 |
Group | 1 | 53.424 | 7.370∗∗ | 0.111 | |
Time × Group | 1 | 8.884 | 6.519∗∗ | 0.099 | |
Backward Digit Span Task | Time | 1 | 8.638 | 23.234∗∗∗ | 0.283. |
Group | 1 | 8.465 | 7.343∗∗ | 0.111 | |
Time × Group | 1 | 6.671 | 17.943∗∗∗ | 0.233 |
In the Day/Night Stroop, the interaction between the time points and groups was significant [
In DCCS, the interaction between time points and groups was significant [
In the Dot Matrix Test, the interaction between time points and groups was significant [
In the Backward Digit Span Task, the interaction between the time points and groups was significant [
In order to investigate if there were any difference between the gains of experimental group and control group before and after music training, we took post-assessment scores T2 to minus pre-assessment scores T1, and did independent sample
Day/Night Stroop | Experimental group | 30 | 10.133 | 4.84756 | 2.509∗ | 59 |
Control group | 31 | 6.516 | 6.29217 | |||
DCCS | Experimental group | 30 | 3.033 | 2.98829 | 2.746∗∗ | 59 |
Control group | 31 | 0.935 | 2.97697 | |||
Dot Matrix Test | Experimental group | 30 | 1.466 | 1.77596 | 2.553∗ | 59 |
Control group | 31 | 0.387 | 1.52047 | |||
Backward digit span task | Experimental group | 30 | 1.000 | 0.94686 | 4.236∗∗∗ | 59 |
Control group | 31 | 0.064 | 0.77182 |
The results of Experiment 1 showed that 12 weeks of integrated musical training could promote the development of children’s EF, a finding that was consistent with the results of previous research (
A large number of related studies have shown that short-term or long-term musical training can affect the brain structure, function, and cognitive level of those involved in the training (
The participants were the same as those in Experiment 1.
The stimuli were the same as those in Experiment 1.
In Experiment 2, we used the same performance test materials to conduct after-effects tests on the children in the music training experimental group and the control group at T3 (12 weeks after Experiment 1). Leading up to this period, the researchers asked the parents of the participating children to ensure their children did not have any in-school or extra-curricular music training activities.
In this experiment, the scores of the 4 tasks were the dependent variables, and the time points (T2 vs. T3) and the groups (the experimental group vs. the control group) were independent variables. The experimental design was a 2 (time points: T2 vs. T3) × 2 (groups: the experimental group vs. the control group) two-way repeated measures ANOVAs, in which the time points were the intra-group variables and the groups were the inter-group variables. The analysis of variance mainly examined the interaction between time points and groups.
The results of the after-effect of executive function tasks of each group of children in Experiment 2 at T3 are shown in
After-effect description statistics at T3.
Day/Night Stroop | 21.73 | 1.143 | 19.19 | 2.315 |
DCCS | 15.37 | 1.474 | 14.42 | 1.432 |
Dot Matrix Test | 8.87 | 2.063 | 7.84 | 1.934 |
Backward Digit Span Task | 1.63 | 0.718 | 1.10 | 0.597 |
We used the 4 experimental scores as the dependent variables. The time points (T2 vs. T3) and the groups (the experimental group vs. the control group) were independent variables. The results of the 2 time points (T2 vs. T3) × 2 groups (the experimental group vs. the control group) two-way repeated-measure ANOVAs are shown in
Music training after-effect repeated measurement analysis of variance.
Day/Night Stroop | Time | 1 | 3.621 | 1.150 | 0.019 |
Group | 1 | 386.837 | 31.908∗∗∗ | 0.351 | |
Time × Group | 1 | 31.851 | 10.110∗∗ | 0.146 | |
DCCS | Time | 1 | 4.506 | 2.556 | 0.042 |
Group | 1 | 68.459 | 13.388∗∗ | 0.185 | |
Time × Group | 1 | 9.260 | 5.252∗ | 0.082 | |
Dot Matrix Test | Time | 1 | 8.174 | 6.671∗ | 0.102 |
Group | 1 | 63.729 | 9.755∗∗ | 0.142 | |
Time × Group | 1 | 5.321 | 4.343∗ | 0.069 | |
Backward Digit Span Task | Time | 1 | 0.026 | 0.062 | 0.001 |
Group | 1 | 17.872 | 26.582∗∗∗ | 0.311 | |
Time × Group | 1 | 1.599 | 3.873Δ | 0.062 |
In the after-effects test of the Day/Night Stroop, the interaction between the time points and the groups was significant [
In the after-effects test of the DCCS, the interaction between the time points and the groups was significant [
In the after-effects test of the Dot Matrix Test, the interaction between the time points and the groups was significant [
In the after-effects test of the Backward Digit Span Task, the interaction between the time points and the groups was significant [
Twelve weeks following the cessation of musical training, we again tested the EF of the two groups of children and explored the duration of the transfer effect of musical training. Leading up to this period, the parents of the children who participated in the experiment were asked to ensure that their child avoided any additional musical training. Our results showed that in the absence of additional musical training in both groups, the scores of the EF tasks in the experimental group remained significantly higher than those in the control group. While scores on the Day/Night Stroop decreased significantly, scores on the other three tasks did not; thus, the effect of musical training had a sustained effect.
Scores on the DCCS and the Dot Matrix Test in the control group increased significantly with age, while scores on the Day/Night Stroop and Digit Span Task did not. While this shows that EF improved due merely to the development of age and other factors, the effect of this natural improvement was still not as pronounced as that seen in the musical training group. The results of Experiment 2 showed that the transfer effect continues to play a role after cessation of musical training.
As the number of EF intervention studies increase, it is progressively more important to explore the most appropriate method of increasing EF. According to
In musical training, whether children can understand music rules, memorize, recognize music symbols, sing, and/or tap a beat according to music rules and music symbols is the key to exhibiting melody. A musical melody is composed of notes according to the rules of music writing and using music symbols. To accomplish this, children need to understand the rules of music, including the symbols, and must be able to sing or play musical melodies according to these rules. Moreover, carrying out musical training requires cooperation from many people, and involves, for example, alternate singing, a chorus, and part singing. Therefore, children need to conform to singing order, detect the singing order of other children, and supervise/regulate their performance appropriately. This process requires a high degree of restraint, control, working memory, and cognitive flexibility. For example, in the musical training called “Finding Notes,” children need to quickly and accurately recognize different types of notes according to rules and test instructions. With understanding of the rules of notes, children not only can quickly and accurately find the corresponding notes, but also point out the mistakes of others.
In musical training, it is also important to be able to suppress and adjust behavior according to changes in musical symbols. Silent singing means singing without sound. We used this form of “singing” to train the inhibitory control of young children. In the early stage of musical training, it was difficult for children to sing without sound, while with the understanding of “silent singing” rules, young children could gradually suppress their impulse to sing out loud. For example, when learning the basics of music, the child would learn to terminate singing at a particular mark. When this mark appeared in the music, the child could suppress their urge to continue singing, and stop singing. In the case of beat practice, the test would add a “decrescendo” or “crescendo” symbol to the exercise according to the change rule of the beat. The child must then suppress their dominant reaction and produce either fortissimo or pianissimo beats.
In addition, musical training activities, such as turn singing, choral singing, and voice-singing have numerous rules. We created a situation for young children to perform a role, which allowed the children to become interested in our training, and ultimately, the constraints of music rules. Improving EF can also be promoted. For example, in the musical training called “Frog Chorus,” children can freely choose to role-play as a small frog or an old frog. Different characters sing a different melody. When the little frog sings, the old frog wants to listen quietly. When the old frog sings, the little frog wants to listen quietly. After the little frog and the old frog sing, all the frogs sing together. In the course of this training, the children are very active. After practicing a few times, most of the children can follow the order of the characters singing. When they are not singing the melody, they wait quietly and pay attention. Over time, musical training with clear rules and full of gameplay gradually improves performance.
Intensity is an important factor in musical training. For the auditory cortex, the intensity of musical training is positively correlated with left transverse temporal gyrus volume (
Although the role of musical training in promoting EF has been supported by empirical research, most of the previous studies on the subject have not tracked the effects of musical training; thus it is not clear how long the effects last. To address this gap in the research, we tracked the children who participated in Experiment 1. 12 weeks after the end of musical training (T3), the EF level of the two groups of children was tested to explore the duration of the transfer effect of musical training. The results showed that scores in three major subcomponents of EF in the control group showed significant improvement, while the scores of the children in the musical training group slightly decreased; however, the level of EF in children in the musical training group was still significantly higher than that of children in the control group. This result shows that after musical training (T3), the transfer effect still plays a role. In a return visit to the teachers and parents of the children tested, we found that the children in the musical training group always thoroughly enjoyed music. These teachers and parents stated that many of the children practiced the songs learned in musical training independently in the kindergartens or at home. When the children saw some music symbols, they would also actively explain the meaning of the symbols to their parents. Music exercises spontaneously carried out by children were very frequent, and parents paid special attention to the music that appeared in their daily life.
In the Digit Span Task, the interaction between measurement time points and groups was significant. The reason for this result may be related to the characteristics of the task itself. Although both the Dot Matrix Test and the Digit Span Task are working memory tasks, the Dot Matrix Test is more of a spatial memory task. Some researchers believe that there is a correlation between music processing and spatial processing. For example,
Additionally, long-term training in five-line reading may improve the local processing ability of individuals who have been musically trained.
Moreover, to ensure ecological validity, this research had chosen natural classes as training and control groups. Thus, training effects observed occurred in natural environments. EF is more sensitive to environmental stimulation. The experimental group participated in 12 weeks of group musical learning in a regulated and structured environment set by researchers. In this group setting, observational learning likely took place, with the subjects possibly being the trainers (i.e., music teachers) or their peers in the same class. This interchangeable nature of observational learning may amplify the effects of musical training, resulting in the sustained effects observed.
Previous studies have preliminarily explored the core subcomponents of musical training that affect EF (
There are three main limitations to this study. First, this study used only behavioral experiments to measure the effect of musical training. The influence of musical training on EF is not reflected by activation intensity of a single brain region, but also may be accompanied by changes in the spatial pattern of EF-related brain region activation, as well as the functional linking pattern of related brain regions. Accordingly, future research should not only use ERP, fMRI, and other technologies, but also use multi-modal brain imaging technology to explore the role of musical training on the neural basis of EF. Additionally, EF-related brain structures and functional changes in musical training transfer effects can also be explored. Second, this study tracked the after-effects of musical training. However, the scope of the tracking was limited and only provided a preliminary explanation of the continuous effect of musical training. Finally, we only chose control and experimental groups in this study. In future studies, we will increase the number of different training groups to examine differences among various interventions.
Results showed that musical training can promote children’s inhibitory control, working memory, and cognitive flexibility. Furthermore, 12 weeks after the experiment, integrated musical training demonstrated a sustained promotion effect. Ultimately, musical training is an appropriate means to promote the development of children’s EF.
The datasets for this manuscript are not publicly available because the topic is not finished. Requests to access the datasets should be directed to corresponding author.
This study was approved by the local ethics committees of Liaoning Normal University. Written informed consent had been obtained from the parents/legal guardians of all participants.
LF and YS designed the experiment. YS, LF, and GL prepared the materials and performed the experiment. YS, LF, YL, and SL analyzed the data and wrote the manuscript.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.