Nara Women's University
Negative BOLD responce during motor execution
and imagery: an fMRI study
メタデータ

言語: English
出版者:
奈良女子大学生活環境学部スポーツ健康科学コース
公開日: 2016-06-14
キーワード (Ja): デフォルトモードネットワーク,
機能的時期共鳴画像法, 上前頭回, 内側前頭回, 楔全部
キーワード (En): deactivation, default mode network,
fMRI, MFG, precuneus, SFG
作成者: 中田,大貴, 松田,宗大, 水口,暢章, 坂本,貴和子,
彼末,一之メールアドレス:所属:

URL

http://hdl.handle.net/10935/4243

Negative BOLD responses during motor execution and imagery: an fMRI study
Hiroki Nakata 1)2)

Norihiro Matsuda 3)

Kiwako Sakamoto 4)

Nobuaki Mizuguchi 2)

Kazuyuki Kanosue 2)

Abstract
The aim of this functional magnetic resonance imaging (fMRI) study was to evaluate
negative deflections in blood oxygen level-dependent (BOLD) signals during motor executionof left or right hand movements, and motor imagery of left or right hand movements.

We

observed the Negative BOLD responses during left and right hand movements in the
superior frontal gyrus (SFG), medial frontal gurus (MFG), precuneus, ipsilateral primary
motor cortex (MI), premotor area (PM), dorsolateral prefrontal cortex (DLPFC), and middle
occipital gyrus (MOG), while those of Negative BOLD during motor imagery of left and righthand movements were found in the SFG/MFG and precuneus.

These findings indicate that

the SFG/MFG and precuneus, which involved regions similar to those of the default mode
network, were involved in the neural networks of Negative BOLD during both motor
execution and motor imagery.

Also, deactivations in the ipsilateral MI, PM, DLPFC, and

MOG during motor execution were added to the default neural networks.

The present

findings extend the applicability of BOLD signals by comparing Negative and Positive
signals in order to understand the brain functions involved in motor execution and motor
imagery.
（Research Journal of Sport Science in Nara Women's University, 18: 65-77, 2016）
Keywords: fMRI; deactivation; SFG; MFG; precuneus; default mode network
キーワード：機能的磁気共鳴画像法；deactivation；上前頭回；内側前頭回；楔前部；デフォル
トモードネットワーク
1)

Corresponding author: Hiroki Nakata
奈良女子大学 研究院生活環境科学系 〒630-8506

奈良県奈良市北魚屋西町

Department of Health Sciences, Faculty of Human Life and Environment, Nara Women’s University
Kitauoya-Nishi Machi, Nara, 630-8506, Japan.
2)

早稲田大学 スポーツ科学学術院

3)

早稲田大学 スポーツ科学部

4)

自然科学研究機構

生理学研究所E-mail: hiroki-nakata@cc.nara-wu.ac.jp

感覚運動調節研究部門－ 65 －

Introduction

and Positive BOLD responses share blood
supplied by the same artery.

However, if the

Recent neuroimaging studies using functional

distance between Negative and Positive BOLD

magnetic resonance imaging (fMRI) have shownfoci is large (e.g. left and right hemispheres, andnot only an increase but also a decrease in blood

frontal and occipital cortices), it is difficult tooxygen level-dependent (BOLD) signals during

explaintasks.

This decrease is often referred to as

responses with the “blood steal” phenomenon 20).

‘Negative BOLD’ or deactivation, and several

A fourth mechanism could involve the “default

mechanisms

mode

phenomenon.

have

been

implicated

in

this

The first involves transcallosal

inhibition from one hemisphere to the other.

In

the

Negative

network”.

and

This

Positive

network

BOLD

includestask-independent deactivation regions as a baseline
or resting state of brain function involving ageneral, whereas BOLD signals are increased in

specific set of mental operations 3), 10), 15).

the primary motor area (MI) of the contralateralphysiological basis of Negative BOLD has

hemisphere during voluntary movement of a limb

recently been researched in detail, and non-human26)

studies have described a coupling between, BOLD signals are decreased in the MI

ipsilateral to the movement 1), 16), 28), 37).

The

The

Negative BOLD signals and decreases in neuralsecond mechanism involves the task-related

activity 2), 34) as well as enhanced inhibition within

deactivation of associated areas belonging to an

sensory systems 7).

irrelevant sensory modality.

For example,

In the present study, we investigated Negative

deactivation of the visual cortex occurs during

BOLD responses during motor execution andsomatosensory (tactile) discrimination tasks 19), 26),

imagery, and set two objectives.

30),

objective was to clarify how neural networks

31).

The third mechanism entails local

The first

When BOLD

showing Negative BOLD were recruited in thesignals in some parts of the primary visual cortex

whole brain during voluntary movement of theincrease after particular types of visual stimulation,

limbs.

signals in other parts of the visual cortex decrease

have17), 33)

A possible explanation for the above

contralateral to voluntary movement of the limb

results could involve “blood steal” phenomenon.

and Negative BOLD in the MI ipsilateral to the

Adjacent regions between the areas of Negative

limb.

reallocation of perfusion resources.

.

－ 66 －As mentioned above, previous studies
reported

Positive

BOLD

in

the

MI

However, the responses of the neural

Methods

networks of Negative BOLD in the whole brain
during voluntary movement remain unclear.

The

second objective was to examine the Negative

Subjects

BOLD responses during ‘motor imagery’ of rightFifteen normal right-handed subjects (threeand left hand movement.

Motor imagery isfemales and twelve males; mean age 23.0 years,

defined as the mental execution of a movement

range 21-28 years) participated in this study.

without any overt movement.

of them met the criteria of to the Edinburgh

fMRI studies have

31)

All

shown that brain activity during motor imagery is

Inventory

similar to the activity that occurs during motor

neurological or psychiatric disorders.

execution 14), 18).

Many studies have report that

protocol was approved by the Human Research

the supplemental motor area (SMA), premotor

Ethics Committee of Waseda University, Japan.

area (PM), primary motor cortex (MI), posterior

Written informed consent was obtained from allparietalsubjects.

cortex(PPC),

basal

ganglia,and.

Subjects had no history ofThecerebellum are activated during motor imagery 5), 8),14), 18), 25).

We suspected that if the neuronal

Tasks
Recordings

networks of Positive BOLD responses in motor

wereconducted

under

four

imagery were similar to those involved with motor

conditions; (1) right hand motor execution

execution, the same relationship could hold for the(RH-exe), (2) left hand motor execution (LH-exe),

Negative BOLD responses.

(3) right hand motor imagery (RH-image), and (4)In order to extend knowledge in this area, theleft hand motor imagery (LH-image).

Subjects

current study included an investigation of thewere asked to repeat extension and flexion of thecharacteristics of Negative BOLD responses in theright hand in the RH-exe condition, and of the left

whole brain during right or left hand movement

hand in the LH-exe condition.

and also during motor imagery of right or left hand

LH-image conditions, subjects were instructed to

movement utilizing fMRI.

imagine repetitive extension and flexion of theright and left hand, respectively.

In the RH- andSubjects werealso asked to match the pace of real and imaginary
movements in all conditions.
Each condition consisted of five alternate－ 67 －

repetitions of task and rest periods (total 5 min).

All images were acquired using a 1.5 T MR

The duration of the task and rest period was 30 sec

scanner (Signa, General Electric, Wisc., USA).

for each.

The four conditions were performed

BOLD contrast functional images were acquired

separately.

Subjects were informed only at theusing T2*-weighted echo planar imaging (EPI)

start and end of the task period via a tactile cue.

free induction decay (FID) sequences with the

The cue was presented by an experimenter to thefollowing parameters: TR 3000 ms, TE 50 ms,

dorsum of the right hand during RH-exe and

FOV 22 cm × 22 cm, flip angle 90°, slice thickness

RH-image conditions, and to the dorsum of the left

5 mm, gap 1 mm, and 25 slices.

hand during LH-exe and LH-image conditions.

volumes of each fMRI session were discarded due

The order of the four conditions was randomized

to unstable magnetization.

for each subject and counterbalanced across all(TR 30 ms, TE 6 ms, FOV 24 cm × 24 cm, flip

subjects.

We asked the subjects to open theirangle 90°, slice thickness 1 mm, no gap, and 124eyes and relax their hands during both rest andslices) were also obtained for each subject fortask periods.

anatomical reference.

The first fourT1-weighted images

All subjects practiced the right and left hand
movements several times at their own pace before

Data analysis

recording was commenced.

Raw data were analyzed

The difference

between the first person perspective and third
person perspective of the motor imagery

38)

was

Parametric

Mappingusing Statistical(SPM5,

Wellcome

Department of Cognitive Neurology, London, UK)
11)

Matlab(Mathworks,

subsequently instructed to imagine repetitive

Sherborn, Massachusetts, USA).

The effect ofextension and flexion of the hand with a firsthead motion was corrected by realigning all scans

person perspective at their own pace during theto the initial one.

task periods for the RH-image and LH-image

normalized to a standard stereotaxic spaceconditions, and told not to change the pace during(Montreal Neurological Institute (MNI) template),the experiment.

This procedure followed that ofand images were then spatially smoothed (full25)

width, half maximum = 8 mm).

then explained to the subjects.

our previous study.

They were

implemented

with

The functional scans were

Statistical analysis was performed at two levels.
fMRI data acquisition

First,－ 68 －

individualtask-related

activation

wasevaluated.

We constituted a statistical parametric

LH-imagemap of the t-statistic for the four contrasts ofConjunction analysis for common regions

Negative BOLD, (1) RH-exe, (2) LH-exe, (3)deactivated during RH-exe and LH-exe revealed

RH-image, and (4) LH-image.

Subject-specificsignificant deactivation in the left superior frontal

contrast images of parameter estimates were thengyrus (SFG) (Brodmann’s areas: BA 8 and 9),

used

a

dorsolateral prefrontal cortex (DLPFC) (BA 9),

to make inferences at theventrolateral prefrontal cortex (VLPFC) (BA 45),

A groupwas

premotor area (PM) (BA 6 and 8), precuneus (BAperformed by entering contrast images into a7), superior parietal lobule (BA 7), superior

one-sample t-test, and mainly focused on thetemporal gyrus (STG) (BA 39), middle occipitalconjunction

regionsgyrus (MOG) (BA 18), cuneus (BA 19), lingual

deactivated during both RH-exe and LH-exe

gyrus (BA 18), posterior cingulate (BA 30), andconditions as well as during both RH-image andcingulate gyrus (BA 30), and in the right SFG (BALH-image conditions. The threshold was set at p

8 and 9), DLPFC (BA 9 and 10), PM (BA 6 and 8),< 0.05 uncorrected (spatial extent > 10 voxels).

precuneus (BA 7 and 31), STG (BA 39), MOG

The locations of brain activity were transformed(BA 19), cuneus (BA 19), lingual gyrus (BA 19),from MNI coordinates into Talairach standard

parahippocampal gyrus (BA 19), and posteriorforsecond-level
12)

random-effect model
populationlevel.

data

brain coordinates

to

39)

.

analysisdetectusing

analysisbrain

In addition, to investigate

cingulate (BA 29 and 30) (Figure 1A, Table 1).

the deactivations under four conditions separately

Conjunction analysis for common regions

as trends results, we set the statistical threshold ofdeactivated during RH-image and LH-image

one sample T-tests at P < 0.05, uncorrected, based

demonstrated significant deactivation in the left

on previous data showing Negative BOLD signals

MFG (BA 9), cuneus (BA 17), and posterior16), 27)

cingulate (BA 31), and in the right precuneus (BA.

7) (Figure 1B, Table 2).

Results

Negative BOLD in right hand motor execution(RH-exe) and left hand motor execution (LH-exe)

Conjunction analysis for Negative BOLD during

Regions deactivated in the left hemisphere by

RH-exe and LH-exe, and during RH-image andRH-exe were located in the SFG (BA 6, 8, and 9),－ 69 －

Figure 1: (A) Brain regions commonly deactivated in RH-exe and LH-exe.

(B) Brain regions commonly deactivatedin the RH-image and LH-image. VLPFC = ventrolateral prefrontal cortex; DLPFC = dorsolateral prefrontal cortex;
MOG = middle occipital gyrus; PM= premotor area; SFG = superior frontal gyrus; MFG = medial frontal gyrus; PCC= posterior cingulate cortex.
Figure 2: Group activation map showing deactivated brain
regions during (A) right and (B) left hand movements, and(C) motor imagery of the right hand and (D) the left hand.
PM = premotor area; MI = primary motor area; MOG =middle occipital gyrus; SFG = superior frontal gyrus;
VLPFC = ventrolateral prefrontal cortex; DLPFC =dorsolateral prefrontal cortex; SG = supramarginal gyrus;
MFG = medial frontal gyrus

PM (BA 6), DLPFC (BA 9), VLPFC (BA 45),
medial frontal gyrus (MFG) (BA 9), precuneus(BA 7), MOG (BA 18), and cingulate cortex (BA32).

The areas deactivated in the righthemisphere were in the MI (BA 4), PM (BA 6),
SFG (BA 8 and 9), DLPFC (BA 9 and 10),－ 70 －

precuneus (BA 7), and MOG (BA 18) (Figure 2A).
Negative activities in LH-exe were found in the

Table 1: Conjunction analysis for regions deactivated during
RH-exe and LH-exe conditions

left MI (BA 4), PM (BA 6), SFG (BA 6), DLPFC(BA 9), supramarginal gyrus (SG) (BA 40), IPL(BA 40), MTG (BA 39), precuneus (BA 7), cuneus(BA 19), MOG (BA 19), insula (BA 13),parahippocampal gyrus, and cerebellar uvula.

Inthe right hemisphere, deactivated regions weredetected in the right PM (BA 6), DLPFC (BA 9and 10), SFG (BA 6 and 8), SG (BA 40), STG (BA22), precuneus (BA 19), MOG (BA 19), andcerebellar declive (Figure 2B).

Negative BOLD in right hand motor imagery(RH-image)

and

left

hand

motor

imagery(LH-image)
Regions deactivated by RH-image were foundin the left SFG (BA 9), precuneus, and cuneus, andin the right SFG (BA 9), precuneus, and STG (BA39) (Figure 2C).
Negative BOLD effects in LH-image wereBA, Brodmann’s area; L, left hemisphere; R, righthemisphere.

found in the left medial frontal gyrus (BA 8),
cuneus (BA 18), and superior occipital gyrus (BA

Table 2: Conjunction analysis for regions deactivated during19), and in the right precuneus (BA 19) and cuneus

RH-image and LH-image conditions(BA 18) (Figure 2D).

－ 71 －

Discussion

involved in voluntary hand movements.

In aprevious study, we recorded Negative BOLD
32)

Negative BOLD in motor execution of right andresponses during tongue movement

left hands

general, tongue movement showed a bilateral

As mentioned in the Introduction section,.

Inpattern of activation in the MI as Positive BOLD 9),

several studies have demonstrated deactivation in

22)

the

hand

observed on movement-related cortical areas such

To the best of our knowledge, only

as the MI, SMA, PM, and cerebellum, but was

ipsilateral

movements.

MIduringvoluntary.

A Negative BOLD response was notobserved

deactivation as well as the ipsilateral MI during

Sakamoto and colleagues suggest that, in addition

these movements.

Hamzei and colleagues

to transcallosal inhibition, there are alternative

showed deactivation in the precuneus whileneural mechanisms that mediate the Negative

16).

BOLD response during voluntary movements.

However, since the number of subjects in their

The results of the present study also indicated thestudy was small (i.e. six right-handed subjects),

deactivation

deactivation regions need to be confirmed using a

unilateral hand movements.

larger number of subjects.

The authors also didthat the Negative BOLD responses that occur

not discuss why the precuneus was deactivatedduring voluntary movements consisted of specificwhile performing the pinch grip.

neural networks.

performingaright-handed

pinch

grip

In the RH-exe

on

the

in

precuneus

some

brain

and

PPC

32)

one study reported the other brain regions ofregions.

duringThis is an indication

condition of the present study, the Negative BOLD
response was found in the ipsilateral MI, SFG, PM,

Negative BOLD in motor imagery of the right and

DLPFC, VLPFC, precuneus, MOG, and cingulate

left hands

In the LH-exe condition,

Conjunction analysis demonstrated that thedeactivation was identified in the ipsilateral MI,

common Negative BOLD response occurred in thePM, SFG, DLPFC, SG, IPL, precuneus, cuneus,

MFG, precuneus, cuneus, and posterior cingulate

MOG, insula, STG, and cerebellum (Figure 2B).

(Figure 1B, Table 2).

cortex (Figure 2A).

These findings suggest that neural networks in

Why did deactivation occur in the SFG/MFG,

addition to transcallosal inhibition from theprecuneus, and posterior cingulate during motor

contralateralimagery?

toipsilateralhemisphere

were－ 72 －

Our hypothesis holds that it occurredin ‘the context of a default mode network’ that is

The Negative BOLD response in the precuneus

associated with motor imagery.

The default

was considered to be part the “irrelevant cortex” asmode network has been identified to occur during

well as functioning as a default mode network.

task-induced deactivations, or during brain activity

Moreover, since the precuneus has reciprocal

associated with a passive fixation baseline

corticocortical connections with SFG/MFG andcondition related to specific attention-demanding

posterior cingulate 4), it is little wonder that thesevisual tasks 15), 36).

regions were simultaneously deactivated during

This network involves theMFG, medial and lateral parietal areas, andmotor imagery.

posterior cingulate 10), and is related to active
cognitive processing that includes mind-wandering

Neural network of the regions showing Negative

23), and autobiographical remembering 35).

BOLD between motor execution and motor

The SFG/MFG is known as one of the key

imagery

regions involved with the default mode network 6).

In the present study, Negative BOLD responses

It is associated with self-reflective thought andwere clearly different between motor execution

judgments that depend on social and emotionaland motor imagery.

content 3) as well as stimulus-related thought suchcommon deactivated regions, independent ofas enhanced watchfulness toward the external

motor execution, imagery, and the right and left

environment

13).

The precuneus has beenhands.

However, there were

Deactivation of the SFG/MFG and

implicated in a wide variety of highly integrated

precuneus was observed under all conditions.

tasks, including visuo-spatial imagery, episodic

mentioned above, these regions are closely related

memory retrieval and self-processing operations 4).

to the default mode network.

This implicates the precuneus in the function ofneural networks of Negative BOLD seen during

human higher-order cognitive and integrative

voluntary movements involve three mechanisms;

function, rather than with lower-order earlytranscallosal inhibition from the contralateral tocognitive processing.

In the present study, theipsilateral hemisphere, the default mode network,

precuneus might not have played an important roleand deactivation of task-related networks thatin the performance of voluntary hand movements,

belong to an irrelevant sensory modalities.

since our hand movement tasks were simple andcontrast, neural networks that exhibited a Negative

did not require higher-order cognitive function.

BOLD during motor imagery may well be－ 73 －

As

We suggest that theInassociated only with the default mode network.

and deactivation during finger movement. Neurology, 54:

While previous studies have reported that Positive

135-142.

BOLD responses during motor imagery are similar

2) Boorman, L., Kennerley, A.J., Johnston, D., Jones, M.,

to those that occur during motor execution 14), 18),

Zheng, Y., Redgrave, P. and Berwick, J. (2010) Negative

our findings indicated that the Negative BOLD

blood oxygen level dependence in the rat: a model for

responses that occurred during motor imagery

investigating the role of suppression in neurovascularwere clearly different from those that occurredcoupling. Journal of Neuroscience, 30: 4285-4294.

during motor execution.

In this regard, an3) Buckner, R.L., Andrews-Hanna, J.R. and Schacter, D.L.

investigation on the difference in Negative BOLD

(2008) The brain's default network. Annals of New York

responses between motor execution and imagery

Academy of Sciences, 1124: 1-38.

would contribute to an understanding of how these4) Cavanna, A.E. and Trimble, M.R. (2006) The precuneus:

important processes differ.

a review of its functional anatomy and behaviouralcorrelates. Brain, 129: 564-583.

In conclusion, this is the first study to5) Decety, J., Perani, D., Jeannerod, M., Bettinardl, V.,investigate Negative BOLD responses in the whole

Tadary, B., Woods, R., Mazziotta, J.C. and Fazlo, F. (1994)

brain during voluntary motor execution involving

Mapping motor representations with positron emission

the right or left hand as well as motor imagery of

tomography. Nature, 371: 600-602.

the right or left hand.

The present findings

6) Deshpande, G., Santhanam, P. and Hu, X. (2011)

extend the applicability of BOLD signals by

Instantaneous and causal connectivity in resting state brain

comparing Negative and Positive signals andnetworks derived from functional MRI data. NeuroImage,utilizing

54: 1043-1052.

thisinformationtofurtheranunderstanding of the brain functions involved in7) Devor, A., Tian, P., Nishimura, N., Teng, I.C., Hillman,

motor execution and motor imagery.

E.M., Narayanan, S.N., Ulbert, I., Boas, D.A., Kleinfeld, D.
and Dale, A.M. (2007) Suppressed neuronal activity andconcurrent arteriolar vasoconstriction may explain negative

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