長時間運動時の脳グリコゲン減少と中枢性疲労 - 上月財団

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Ǟʈǝ Ƕȧ ........................................................................................................................................................................... 2
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Ǟʊǝ ɞůɠɎÒ3ȇUWVGƦę#CʒʂǒǚȲɶʈʃ ............................................... 5
Ǟʋǝ ɎÒů2ȇUWVƦę3œǳůɠ®ČĹʒʂǒǚȲɶʉʃ.................................... 12
Ǟʌǝ ǵäȩȴ ............................................................................................................................................................... 20
Ǟʍǝ ǵŒ ......................................................................................................................................................................... 25
ǒǚłžȤ ......................................................................................................................................................................... 27
ȵɄ............................................................................................................................................................................................. 27
Ûǿţƹ................................................................................................................................................................................... 28
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Table 1 The protocol for habituation to treadmill running exercise.
Day
Running speed and time
1
2
3
4
5
Rest, 10 min + 5 m/min, 10 min + 10 m/min, 10 min
Rest, 5 min + 5 m/min, 10 min + 10 m/min, 10 min + 15 m/min, 10 min
Rest
Rest, 5 min + 10 m/min, 10 min + 15 m/min, 10 min + 20 m/min, 10 min
Rest, 5 min + 15 m/min, 10 min + 20 m/min, 10 min + 25 m/min, 10 min
6
Rest, 5 min + 15 m/min, 10 min + 20 m/min, 10 min + 25 m/min, 10 min
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~ K [ b ʄ Ŭ ƚ Ȟ Ȑ ƃ ħ ¥ ǖ ʄ Japan) G 100µl ǉ 1 ƞ Ė & ʆ ș į 2 ũ ɠ 2 ò
ĳŶɠ2.ĐɼGȘ)&ʆ
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f j G Ɏ Ò É Ǽ ʄɯ Ɏ Ò Ǽ (j f k 1 Ď ǻ )ʄɎ Ò 30 Ä Ǽ ʄɎ Ò 60 Ä Ǽ ʄɎ
Ò 120 Ä Ǽ 2 5 Ǽ 1 Ä ʄ Ä Ɋ 20 mʄ 120 Ä ɠ 2 j f k Ƚ Ɏ Ò G Ș F # & ʆ Ɏ Ò
ɟ Ĉ 0 Ä (Ɏ Ò É )ʄ 30 Ä ʄ 60 Ä ʄ 120 Ä 2 ů Ƴ - ~ K T Ɲ Ƶ Ė (10 kWʄ 1.2 ǘ )1 @ C
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1Ǖ&ʆ
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2ŧƜ1İʄǉȼʄɧʄǷźªʄơɻʄȢģʄȢģɓʄȇʄĘȇʄȇĢ2ʐɓ
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ʉʅʍʆȗƣłÄ2ƨď
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ʉʅʐʆȇ¾pI2ďɛ
Takeda et al. (1990)2 ŧ Ɯ 1 @ B ɾ Ɋ ƣ ª T ~ j U v J (HPLC)- ƨ ď & ʆ
ʉʅʈʇʆǰȨÁƻ
i c 3 ! 7 , Ġ ù ³ ±ƌ Ƭ Ȱ Ğ - Ǖ ʄǰ Ȩ Á ƻ 1 * , 3 · ɔ ǻ Ä Š Ä Ž 2 . ʄ
post hoc h ] j (Dunnet)G Ș ) & ʆǍ ɡ Ä Ž 3 Pearson G ƿ & ʆŵ ļ Ɨ Ƭ 3 5%. & ʆ
14
ʊʆǮž
ʊʅʈʆȗǥʄȗɘʄ51ǟȁȇUWVưĥ
Ä Ɋ 20 m ʄ30ʄ60ʄ120 Ä ɠ 2 j f k Ƚ Ɏ Ò ǌ į 2 ȗ ǥ ʄȗ ɘ ʄ 5 1 ǟ
ȁ ȇ U W V ư ĥ G Figure 3 1 Ǖ ! ʆ ȗ ǥ ³ 3 Ɏ Ò 30 Ä Ǽ . 60 Ä Ǽ 1 , Ɏ Ò
É Ǽ . Ɩ Ƀ , Ʀ ę 0 ) & ʄ Ɏ Ò 120 Ä Ǽ - 46 %© & ʆ ǟ ȁ U W V ư
ĥ3ɎÒœǳůɠ®ČǇ1Ʀę&ʆȗǥ.åƋ1ʄǉȼʄơɻʄȢģɓʄĘȇʄȇ
Ģ 2 U W V ư ĥ 3 Ɏ Ò 30 Ä Ǽ . 60 Ä Ǽ 1 , Ɏ Ò É Ǽ . Ɩ Ƀ , Ʀ ę 0 )
Exercise
(20m/min, 120min)
E
30
50
120 min
60
Soleus
40
100
** **
30
**
10
0
0
30
**
60
20
60
40
0
0
15
10
10
**
Glycogen (µmol/g)
0
15 Hypothalamus
10
*
5
0
**
4
2
0
0
30
60
30
**
**
60
120
Cortex
30
60
120
1000
20
8
**
6
4
2
0
0
30
60
Liver
800
**
600
**
400
200
0
**
5
**
0
30
60
120
Hippocampus
**
10
5
0
15 Cerebellum
0
15 Brainstem
10
10
0
120
15
**
5
0
120
D
20
120
5
*
6
Plantaris
80
15 Whole brain
Glycogen (µmol/g)
C
Glycogen (µmol/g)
0
8
Blood lactate
(mM)
B
microwave irradiation
(10 kW, 1.2 s)
Glycogen (µmol/g)
A
Blood glucose
(mM)
& ʄ Ɏ Ò 120 Ä Ǽ - 2 ; Ʀ ę & ʆ
0
30
60
120
**
5
0
0
30
60
120
Fi g ure 3 B l ood pa ram ete rs a nd gl yc oge n l e vel s i n th e s kel eta l m usc le, live r, an d bra in a fter exer ci se
for 0, 30, 6 0, and 120 m i n. A , E xper im en t al pr oced ure. B, B lood gluc ose and lactate levels. C, G lyco gen
l ev els in sk eletal m uscl e s. D , L i ve r gl y cog en levels . E , B rain g lycog en leve ls. D ata repr esent the m ean ±
SE M (n = 5-6 r a t s). *, p < 0. 05 ; **, p < 0. 01 com pare d t o p re-ex ercised rats (D unn ett’s po st hoc test).
15
ʊʅʉʆȇ¾UW]51ɘưĥ
Ä Ɋ 20 mʄ 30ʄ 60ʄ 120 Ä ɠ 2 j f k Ƚ Ɏ Ò ǌ į 2 ȇ ¾ U W ] 5 1 ɘ
ư ĥ G Table 2 1 Ǖ ! ʆ ǉ ȼ ʄ ơ ɻ ʄ Ȣ ģ ɓ ʄ Ę ȇ ʄ ȇ Ģ 2 U W ] ư ĥ 3 ȗ ǥ . å
Ƌ 1 ʄ Ɏ Ò 30 Ä Ǽ . 60 Ä Ǽ 1 , Ɏ Ò É Ǽ . Ɩ Ƀ , Ʀ ę 0 ) & ʄ Ɏ Ò 120
Ä Ǽ - 2 ; Ʀ ę & ʆ ǉ ȼ ʄ ơ ɻ ʄ Ȣ ģ ɓ ʄ Ę ȇ ʄ ȇ Ģ 2 ɘ ư ĥ 3 Ɏ Ò 120 Ä Ǽ ŵļ1ɾ)&ʆ
T a b l e 2 G lu c o s e a n d l a c t a t e l e v e l s i n f i v e b r a in lo c i a f t e r e x e r c is e f o r 0 , 3 0 , 6 0 , a n d 1 2 0 m in ( µ m o l / g
w e t t is s u e ) .
Brain region
Coetex
Hippocampus
Hypothalamus
Cerebellum
Brainstem
0 min
30 min
60 min
120 min
Glucose
2.6 ± 0.1
3.3 ± 0.1*
3.2 ± 0.2
1.3 ± 0.4*
Lactate
Glucose
1.1 ± 0.0
2.8 ± 0.1
1.9 ± 0.3*
3.4 ± 0.1*
1.9 ± 0.3
3.5 ± 0.1*
2.5 ± 0.2**
1.6 ± 0.3*
Lactate
Glucose
1.3 ± 0.0
2.8 ± 0.1
2.2 ± 0.3
3.2 ± 0.1
2.0 ± 0.3
3.0 ± 0.1
2.7 ± 0.4**
1.9 ± 0.3*
Lactate
Glucose
Lactate
Glucose
Lactate
1.1 ± 0.1
3.1 ± 0.1
0.9 ± 0.1
2.7 ± 0.1
1.1 ± 0.1
1.9 ± 0.3*
3.7 ± 0.2
1.5 ± 0.2*
3.2 ± 0.1
1.9 ± 0.3*
1.8 ± 0.3
3.4 ± 0.1
1.5 ± 0.2
3.3 ± 0.2
1.9 ± 0.3
2.4 ± 0.2**
1.9 ± 0.4*
2.0 ± 0.2**
1.6 ± 0.3*
2.5 ± 0.2***
D a ta r e p r e s e n t th e m e a n ± s t a n d a r d e r r o r ( n = 5 - 6 r a t s ) .
te s t) .
16
* , p < 0 .0 5 c o m p a r e d t o 0 m i n ( D u n n e t t ’ s p o s t h o c
ʊʅʊʆȗǥʄȇ¾UW]ʄȇ¾ɘ.ȇUWV2Ǎɡ
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1ȇUW].ȇUWV.2ɠ13¼,2ɓ¨-ɯğ1ɾƑ2ǍɡȮ=AD
&ʆȇɘ.ȇUWV.2ɠ13ȢģɓGɦ¼,2ɓ¨-ȷ2ǍɡȮ=AD
&ʆ
A Blood glucose vs. Brain glycogen
Cortex
nr == 210.70
p < 0.001
10
5
0
0 2 4 6 8
!"#$%&'()*+,%"-&).'/)0112'3)
15
Hippocampus
Cerebellum
21
21
nr ==0.54
15nr == 0.67
p < 0.05
p < 0.001
10
10
5
5
0
0
0 2 4 6 8 0 2 4 6 8
Hypothalamus
Medulla oblongata
21
15n = 21
15nr == 0.70
r = 0.64
p < 0.01
p < 0.001
10
10
5
5
0
0
0 2 4 6 8 0 2 4 6 8
Blood glucose (mM)
15
B Brain glucose vs. Brain glycogen
C Brain lactate vs. Brain glycogen
Cortex
8%5/'9
nr == 210.85
p < 0.0001
10
5
0
0 1 2 3 4 5
!"#$%&'()*+,%"-&).'/)0112'3)
15
Hippocampus
"&
"!
&
!
!
n = 21
r = 0.73
p < 0.001
Medulla oblongata
$
%
8'5'<'""2,
'()("1(
*()(+!,0!(
!(.(!,!"
"!
'()("/(
*()(+!,0&(
!(.(!,!"
"
&
#
$
%
:#;%/=6"6,21
15n = 21
nr == 210.59
"&
r = 0.91
p < 0.001
p < 0.0001
10
10
"!
5
& '()(#!(
5
*()(+!,"1(
!
0
0
0 1 2 3 4 5 0 1 2 3 4 5
! "
4567()&"2$%1')*+,%"-&).'/)0112'3
15
#
"&
!
Hypothalamus
"
:7;;%$6,;21
Cerebellum
15
"&
10
10
"!
5
5
&
!
0
0
0 1 2 3 4 5 0 1 2 3 4 5
15
n = 21
r = 0.74
p < 0.0001
'()(#!(
*()(+!,%-(
!(.(!,!&
!
!
"
#
$
%
>'?2""6)%<"%(&6/6
"&
"!
&
#
$
%
'()(#!(
*()(+!,%1(
!(.(!,!&
!
!
"
#
$
%
@6$/6/')*+,%"-&).'/)0112'3)
F i g u r e 5 C o r r e l a t i o n b e t w e e n b r a i n g l y c o g e n le v e ls a n d b lo o d g lu c o s e , b r a in g lu c o s e a n d b r a in la c t a t e .
C o r r e l a t i o n b e t w e e n b r a i n g l y c o g e n l e v e l s a n d A ; b l o o d g l u c o s e , B ; b r a i n g l u c o s e , a n d C ; b r a i n l a c ta te
(Pearson’s product–moment correlation test).
17
ʊʅʋʆǉȼpIưĥʄ51ǉȼpI.UWVưĥ2Ǎɡ
pIkl2 ȵƾƶ-C
A
150 NA
40 MHPG
100
10
0
ng/g wet tissue
(MHPG)3 Ɏ Ò 60 Ä Ǽ - Ɏ Ò É Ǽ . Ɩ
Ƀ , Ā Î ʄ Ɏ Ò 120 Ä Ǽ - 3 Ɏ Ò É
Ǽ ʄ Ɏ Ò 60 Ä Ǽ 2 Ǽ . Ɩ Ƀ , ŵ ļ
- C 5- t k R [ K k ɖ ɘ
(5-HIAA)3 Ɏ Ò 60 Ä Ǽ . 120 Ä Ǽ - Ɏ
0
0
120
60
0
60
400 5-HT
200 5-HIAA
300
200
100
150
100
50
0
0
0
120
60
120
0
60
120
60
120
80 DOPAC
500 DA
400
300
200
100
0
0
0ɾ³GǕ&ʆ_jm2 ȵƾƶ
20
50
jR[tkR[vMmUW
#
30
60
40
20
0
120
60
0
Running time (min)
B
Glycogen (µmol/g wet tissue)
ÒÉǼ.ƖɃ,ŵļ0ɾ³GǕ&ʆ
MHPG vs. glycogen
0Aʄ\tkR[vMmɖ
ɘ (DOPAC) 3 Ā Î ¶ æ 1 C > 2 2 ŵ
ļ0āÔ30)&ʆ$22ƨďɱ
ǋ 1 > ŵ ļ 0 ā Ô 3 ȡ A D 0 ) & (Fig.
6A)ʆ MHPG @ 5 5-HIAA . U W V
.2ɠ13ɾȷ2ǍɡȮ=AD
15 n = 15
r = -0.66
p < 0.01
10
5
0
0
10
20
30
40
MHPG (ng/g wet tissue)
5-HIAA vs. glycogen
DOPAC vs. glycogen
15 n = 15
r = -0.68
p < 0.01
10
15
13
nr == -0.49
10
5
5
0
0
50
100
150
200
00
5-HIAA (ng/g wet tissue)
20
40
60
80
DOPAC (ng/g wet tissue)
Fi g ure 6 M on oam in es an d th eir m etab olites in
the co rtex aft er e xerc ise for 0, 6 0, and 1 20 m i n.
A, M on oam i ne s an d t he i r m et abo l i t es. D ata
rep resen t the m ean ± st an dard error (n = 4 -6 ra t s).
*, p < 0 . 05; ** , p < 0 . 01 co mp ared t o 0 m i n; #, p <
0. 0 5 com par ed t o 60 m i n of ex erci se (T uk ey’s po st
ho c test ). B , C orrelatio n be twee n m ono am ine
m etabo lites a nd glyco gen leve ls (Pe arson ’s
pro duc t–m om ent c orrel at i o n t est ).
& ʄ DOPAC . U W V 2 ɠ 1 3 Ǎ
ɡ 3 ȡ A D 0 ) & (Fig. 6B)ʆ
ʋʆǿĔ
Ź Đ ɼ - 3 ʄǒ ǚ Ȳ ɶ 1 - ǔ ǜ & ȇ U W V ď ɛ Ɯ G ƿ , ʄ30ʄ60ʄ120 Ä ɠ 2
Ƚ Ɏ Ò f j 2 ȇ U W V ư ĥ 1 Ü 9 ! ī ɰ G Ɔ ȩ & ʆ 30 @ 5 60 Ä ɠ 2 Ƚ Ɏ
Ò - 3 ȗ ǥ 3 Ʀ ę # " 1 ȗ ɘ > Ā Î 0 ʄ Ɏ Ò 120 Ä - 3 ȗ ǥ 46 %© ȗ
ɘ 3 Ǩ ʌ ² 1 Ā Î & ʆ 2 . ǟ . ȁ Ȋ 2 U W V 3 œ ǳ ů ɠ ® Č Ǉ 1 Ǩ 90 %
Ʀ ę & ʆ Ɏ Ò 30ʄ 60 Ä 2 ů Ƴ - 3 ǟ ȁ U W V 3 © , C ʄ ȗ ǥ © , A " ʄǄ Ñ # " 1 Ɏ Ò G ǲ ǳ , C v M ^ - C . ǿ A D C ʆ ŧ ʄɎ Ò 120
18
Ä2ůƳ-3©ȗǥȾBʄǟ?ȁȊ2UWV>ƀƥ,CʆD3ǒǚȲɶ
ʉʅʈ2Ƹƛ.89åƋ-ɎÒ1@BǄÑ,CvM^-C.ĸFDCʆ
2 . ʄǉ ȼ ʄơ ɻ ʄȢ ģ ɓ ʄĘ ȇ ʄȇ Ģ 2 U W V ư ĥ 3 Ļ 1 Ý Ɏ Ò 30ʄ
60 Ä - 3 Ʀ ę # " ʄ Ɏ Ò 120 Ä - 2 ; Ǩ 50ʁ Ʀ ę & ʆ D 1 @ B ʄ ȇ U W V 3 ©
ȗǥG§ɞůɠɎÒů12;Ʀę!C.Æ=,ŬA.0)&ʆ2.Ʀę
&ȗǥ51ȇUW].ȇUWV2ɠ1ɯğ1ɾƑ2ǍɡǔȮD&ʆ
2.Aʄȗǥ?ȇUW]3ɎÒů2ȇUWVƦę2ƙďóċ-CáȄĹ
CʆA1ʄ2.ȇ¾-ĀÎ&ɘ.ȇUWV2ɠ1Ȣģɓ¢ă2ʋ
ɓ¨-ȷ2ǍɡȡAD&ʆ2Ǯž3ʄɎÒů1ȇUWVɘ1ÄȦDʄm
1ǯD&áȄĹGǕ!ʆ
Î,ʄŹĐɼ-3ǉȼ¾pI.$2 ȵƾƶ>ďɛ&.EʄpIk
l 2 ȵ ƾ ƶ - C MHPG . _ j m 2 ȵ ƾ ƶ - C 5-HIAA Ɏ Ò 120 Ä Ǽ
-ĀÎ&ʆpIkl?_jm3ȇUWV2ÄȦ°ɍóċ-B
(Benington & Heller, 1995a; Brown, 2004; Benarroch, 2010)ʄ Ɏ Ò ů 1 Ā Î ! C . > ü ê
D , C (Newsholme et al., 1992; Pagliari & Peyrin, 1995)ʆ Ź Đ ɼ - 3 ǉ ȼ 2 MHPG .
U W V ʄ @ 5 5-HIAA . U W V 2 ɠ 1 ȷ 2 Ǎ ɡ ȡ A D & ʆ D 3 ʄȇ U W
V ȵ 1 ɡ 0 k s 2 ȵ ƾ ƶ - C DOPAC - 3 ȡ A D 0 . A ʄ©
ȗǥGĨȾ!ɞůɠɎÒ1@CȇUWV2Ʀę13pIkl._j
m2 ȵɍɡ,CáȄĹCʆ
ɎÒ2©ȗǥ?ȇ¾_jmưĥ2ĀÎ3ɎÒů2ſǄÑ2Ùó.DC
(Nybo & Secher, 2004)ʆ Ź Đ ɼ 1 , ʄ ȗ ǥ ? _ j m Ɏ Ò ů 2 ȇ U W V Ʀ ę
2ƙďóċ.,ȡÂD&ʆDA2Ǯž3ʄ©ȗǥ?_jm2ĀÎ1§ȇU
WV2ƦęɎÒů2ſǄÑ2ǰäȠó-AáȄĹGǕï!CʆįʄÉ1
ȇUWVưĥGɾ=&ýä1œĹsvO~]æ!C/Ɔȩ!CĵȠ
Cʆ
19
Ǟʌǝ ǵäȩȴ
ɎÒůʄȇ3ǟ.åƋ1ƟĹÔʄȇ1CNoS?ɘǩ2ɬȠ3ĀÎ!C
(Nybo & Secher, 2004)ʆ ȇ 3 ǥ ȼ 2 ; G N o S û ȼ . , C . D C ʄ Ɏ Ò 1 @
Bȇ¾2ǥ ȵ/ƍȄʄɒĶ!C38.H/ŬA1D,0ʆ
Ɏ Ò ů 2 ɽ Ƅ ǟ - 3 ʄ Ⱥ ȓ ǥ ȼ - C U W V ư ĥ Ɵ Ò y (Ɏ Ò 2 ĩ ĥ ? œ ǳ
ů ɠ )1 ® Č , Ʀ ę ! C (Gollnick et al., 1974)ʆ ŧ ʄȇ 1 > U W V 3 Č ø (Wender
et al., 2000)ʄ $ D 3 m 2 Ɵ Ĺ Ô ? ȗ ƣ ǀ Ż 2 U W ] ǯ Ɂ ů 1 Ç ƿ D
Ʀ ę ! C (Brown, 2004)ʆ Ɏ Ò 3 m G Ɵ Ĺ Ô (Vissing et al., 1996; Saito & Soya,
2004; Nishijima & Soya, 2006; Ohiwa et al., 2006; Soya et al., 2007a; Soya et al., 2007b;
Nishijima et al., 2011b)ʄ ɞ ů ɠ Ɏ Ò 3 © ȗ ǥ G Ĩ Ⱦ ! . A (Tabata et al., 1984;
Winder et al., 1987)ʄ ȇ U W V G ǟ U W V å Ƌ 1 Ʀ ę # C á Ȅ Ĺ C ʆ $ -Źǒǚ-3ʄȇUWVďɛ2Xi]cdk-C~KTƝƵĖƜG
ė»ʄɞůɠɎÒů1ȇUWVƦę!C/GŬA1!C.GǋǇ.
&ʆ
I ] j Y K j 1 Č ø ! C ȇ 2 U W V 3 ʄȗ ƣ ǀ Ż 2 U W ] ǯ Ɂ (© ȗ ǥ )
ů1ÇƿDƦę!C.DCʆɞůɠɎÒ3©ȗǥGĨȾ!.AʄȇUW
V Ç ƿ D Ʀ ę ! C . ǿ A D C ʄ D 3 ¼ Ŭ ' ) & ʆ$ - ǒ ǚ Ȳ ɶ ʉ ʈ
-3ʄ©ȗǥG§ɞůɠɎÒů2ȇUWVưĥG~KTƝƵĖƜGƿ,Ɔȩ
& ʆ 120 Ä ɠ 2 ɞ ů ɠ Ɏ Ò 1 @ B ȗ ǥ ³ 3 45 %© ʄ ǟ . ȁ Ȋ 2 U W V 3 90 %
Ʀę&ʆɎÒů2©ȗǥ?ǟȁUWV2ƀƥ3ǄÑ2Ŕƌ.DC.A
(Nybo & Secher, 2004)ʄŹ Đ ɼ - ƿ & Ɏ Ò 3 ɯ ğ 1 Ú Ɏ Ò ź ¤ - ) & . Ä C ʆ 2 . ʄȇ U W V 3 ȇ ¼ ª - Ʀ ę ¶ æ G Ǖ ʄǉ ȼ ʄơ ɻ ʄȢ ģ ɓ ʄĘ ȇ ʄ
ȇ Ģ 2 U W V ư ĥ ŵ ļ 1 Ǩ 50ʁ Ʀ ę & ʆ D 1 @ B ʄ © ȗ ǥ G § ɞ ů ɠ Ɏ Ò
ȇUWVGƦę#C.Æ=,ŬA10)&ʆ©ȗǥ3ȇUWVGƦ
ę#CȠó.,@ǑAD,C.Aʄ2Ǯž3ʄɞůɠɎÒů1ĨȾ
D&©ȗǥȗƣAȇ62NoSǯɁGőʄ$2ɁGȝ&=1I]
20
?
Brain
GLUT3
?
Glucose
GLUT1
?
Pyruvate
Lactate
Mitochondria
?
MCT2 Serotonin
Noradrenaline
?
Lactate
MCT4 ?
Pyruvate
MCT1 Glucose
Glycolysis
G-6-P
5-HIAA
MHPG
Neuron
MCT1 Glycogen
Lactate
?
GLUT4
Glucose
?
?
GLUT1
GLUT1
Astrocyte
Vessel
Prolonged exhaustive exercise
F i g u r e 7 B r a in g l y c o g e n m e t a b o l i s m d u r i n g p r o lo n g e d e x h a u s tiv e e x e r c is e . G - 6 - P ; g l u c o s e - 6 - p h o s p h a te ,
G L U T ; G l u c o s e t r a n s p o r t e r , M C T ; m o n o c a r b o x y lic a c id t r a n s p o r t e r . E n e r g y s o u r c e s f o r n e u r o n s i n c l u d e n o t
o n ly b l o o d g lu c o s e b u t a l s o l a c t a t e . A s t r o c y t i c g l y c o g e n i s s y n t h e s i z e d f r o m b l o o d g l u c o s e a n d d e g r a d e d in to
la c t a t e b y e x c ita t o r y n e u r o t r a n s m i t t e r s s u c h a s n o r a d r e n a lin e a n d s e r o to n in . L a c ta te is u p ta k e n n e u r o n s a n d
c h a n g e d t o p y r u v a t e , w h i c h i s u s e d f o r A T P s y n th e s is in th e m ito c h o n d r ia . T h e e f f e c t o f e x e r c is e o n G L U T s
a n d M C T s in th e b r a i n i s n o t e l u c i d a t e d y e t .
jYKj-UWVÄȦɍ&áȄĹGǕ!ʆ0AʄŹĐɼ1,
ŵ ļ 0 U W V Ʀ ę ǔ Ȯ D & 2 3 ȇ ¼ ª - 3 0 ʄǉ ȼ ʄơ ɻ ʄȢ ģ ɓ ʄĘ ȇ ʄ
ȇĢ2ʌɓ¨-)&ʆǉȼ3ǟ62Þǹí¡?ɎÒ2wUUʄơɻ3ɎÒů
2ȮǑʄȢģɓ3ªƧ?NoS ȵ2ȳǤʄĘȇ3ǟ2Øȳ?ĊÓ2ǴœʄȇĢ
3ìé?Ĵŏ2ȳǤ0/ʄɎÒů1ƟĹÔ!C.ǿADCɓ¨-Cʆ&),ʄ
ɞůɠɎÒů2ȇUWVƦę13©ȗǥ'-0ɎÒů2m2ƟĹÔ>
ɡ,C2>D0ʆ0Aʄ2Ƴ1ɡ,3ŬA-0)&ʆ
2 ñ ɶ 1 Ɉ C & = ʄ ǒ ǚ Ȳ ɶ ʉ ʉ - 3 ʄ ǃ 0 C œ ǳ ů ɠ (30ʄ 60ʄ 120 Ä ɠ )2 Ɏ Ò
fj2ȇUWVưĥ1Ü9!īɰGƆȩ&ʆȇUWVɎÒů2m
21
ƟÒ1Ķ
, Ç ƿ D Ʀ ę ! C 0 A ʄ ǟ U W V Ɏ Ò œ ǳ ů ɠ (ǟ Ɵ Ò y )
1®Č,Ʀę!C@1ʄȇUWV>ɎÒœǳůɠ®ČǇ1Ʀę!C.ǿAD
C ʆ © ȗ ǥ 3 Ɏ Ò ɟ Ĉ A 30ʄ 60 Ä 2 ů Ƴ - 3 Ⱦ A " ʄ 120 Ä 2 ů Ƴ - 2 ; ƽ
&ʆ
ǟ . ȁ Ȋ 2 U W V 3 ¹ Ș ǒ ǚ 2 ɉ B ʄ Ɏ Ò œ ǳ ů ɠ 1 ® Č , Ʀ ę & ʆ ȇ (ǉ ȼ ʄ
ơ ɻ ʄ Ȣ ģ ɓ ʄ Ę ȇ ʄ ȇ Ģ )2 U W V 3 ¼ , 2 ɓ ¨ 1 , ʄ © ȗ ǥ 2 ƽ
0Ɏ
Ò ɟ Ĉ A 30ʄ60 Ä 2 ů Ƴ - 3 Ʀ ę # " ʄ© ȗ ǥ G ő & 120 Ä 2 ů Ƴ - 2 ; Ʀ ę & ʆ
2.ʄƦę&ȗǥ.ȇUWV2ɠ1ɯğ1ɾƑ2ǍɡǔȮD&ʆD
A2Ǯž3ʄȗǥɎÒů2ȇUWVƦę2ƙďóċ-CáȄĹGǕï!Cʆ
A1ʄ2.Ȣģɓ¢ă2ʋɓ¨1,ʄȇ¾-ĀÎ&ɘ.ȇUWV.
2ɠ1ȷ2ǍɡȡAD&ʆ2Ǯž3ʄɞůɠɎÒů1ȇUWVɘ1ÄȦ
D & . G Ǖ ʄ ɞ ů ɠ Ɏ Ò ů 2 I ] j Y K j m ɘ [ j (Pellerin &
Magistretti, 1994)1 C ɘ 2 ǯ ƫ . , U W V ȹ ƹ , C á Ȅ Ĺ G Ǖ ï
!CʆÎ,ʄI]jYKj2UWVÄȦ°ɍóċ.,ǑADCpIkl
._jm2 ȵGǉȼ1,ďɛ&.EʄpIkl2 ȵƾƶ
- C MHPG . _ j m 2 ȵ ƾ ƶ - C 5-HIAA Ɏ Ò 120 Ä Ǽ - Ā Î ʄ Ʀ ę &UWVưĥ.ȷ2ǍɡGǕ&ʆpIkl?_jm.$DA2 ȵ
3 ʄ Ɏ Ò ů 1 Ā Î ! C . ¹ Ș ǒ ǚ 1 , ü ê D , B (Pagliari & Peyrin, 1995)ʄ
Źǒǚ2ǮžGŞœ!CʆA1ʄȇUWV ȵ1ɡ0ks2 ȵƾ
ƶ - C DOPAC . ȇ U W V ư ĥ 1 Ǎ ɡ 3 0 . A ʄ© ȗ ǥ G § ɞ ů ɠ Ɏ ů 2
ȇUWVƦę13ȌćĹǗǭ¦ɐƶȼ-CpIkl._jm2 ȵ
ɍ ɡ , C á Ȅ Ĺ C (Fig. 7)ʆ
ŹĐɼ1,ʄȗǥ?_jmɎÒů2ȇUWVƦę2ƙďóċ-Cá
ȄĹȡÂD&ʄDA3ɞůɠɎÒů2ſǄÑ2Ƞó.,>ǑADCʆ&
),ʄȇUWV3ɞůɠɎÒů2ȇ¾_jm ȵ2ɍ?ȗǥ³2©1@
BÇƿDƦęʄ$2ƦęɞůɠɎÒů2ſǄÑ2ǰäóċ.0C2>D0
(Fig. 8)ʆ D 1 * , 3 ʄ į ʄ Ɏ Ò É 1 ȇ U W V ư ĥ G ɾ = & ý ä 1 œ Ĺ s
vO~]æ!C/GƆȩ!CĵȠCʆ
22
Central fatigue
<An integrative factor?>
Brain glycogen decrease
<Central factors>
Hypoglycaemia
Increase in
brain monoamines
(serotonin hypothesis)
Increase in
brain temperature
(hot brain) Increase in
tryptophan/BCAA ratio
in blood
Increase in
body temperature
with dehydration
<Peripheral factors>
Glycogen depletion
in the muscle and liver
Prolonged exhaustive exercise
F ig u r e 8 H y p o t h e t i c a l d i a g r a m s h o w i n g t h e b r a i n g l y c o g e n d e c r e a s e a s a n i n t e g r a t i v e f a c t o r o f
c e n t r a l f a t ig u e d u r i n g p r o l o n g e d e x e r c i s e . P r o l o n g e d e x e r c i s e i n d u c e s g l y c o g e n d e p l e t i o n i n t h e m u s c le s
a n d liv e r , a n d h y p o g l y c a e m i a , w h i c h c a u s e s p e r ip h e r a l f a tig u e . H y p o g ly c a e m ia e lic its e n e r g y s h o r ta g e s in
th e b r a i n , a n d li k e l y i n d u c e s c e n t r a l f a t i g u e . I n c r e a s e i n b r a i n s e r o t o n i n d u e t o r i s e i n t r y p t o p h a n / B C A A
r a t i o i n b lo o d a l s o i n d u c e s c e n t r a l f a t i g u e b y e l i c i t i n g l a s s i t u d e ( s e r o t o n i n h y p o t h e s i s ) . F u r t h e r m o r e ,
in c r e a s e s i n b o d y a n d b r a i n t e m p e r a t u r e a t t r i b u t e d d e h y d r a t i o n i n d u c e c e n t r a l f a t i g u e d i r e c t l y a n d /o r
in d ir e c t l y th r o u g h i n c r e a s e s i n b r a i n n o r a d r e n a l i n e a n d s e r o t o n i n . H y p o g l y c a e m i a a n d s e r o t o n i n a r e n o t o n ly
in d u c i n g f a c to r s o f c e n t r a l f a t i g u e b u t a l s o e n h a n c i n g f a c t o r s o f a s t r o c y t i c g l y c o g e n d e g r a d a t i o n . I n d e e d , w e
o b s e r v e d t h a t b r a i n g l y c o g e n l e v e l s a f t e r r u n n i n g w e r e c o r r e la te d w ith th e r e s p e c tiv e b lo o d g lu c o s e a n d
in c r e a s e d s e r o to n i n m e t a b o l i s m ( M a t s u i e t a l., 2 0 1 1 ) . E x e r c is e - i n d u c e d b r a i n g l y c o g e n d e c r e a s e c o u l d b e a n
in te g r a tiv e f a c to r o f c e n t r a l f a t i g u e .
Źǒǚ1@BʄɞůɠɎÒ1@Bȇ3ȗƣǀŻ2UW].ɘ'-0ʄȇ¾
2Ⱥȓǥȼ-CUWVGÇƿʄA1ʄɎÒį2ȇUWVȿòĳǟU
WV.åƋ1ȾBʄjmU1§ȇ2 ȵɒĶ1ɚȠ-CáȄĹŬA
10)&ʆD3ɞůɠɎÒů2ſǄÑ?ɎÒjmU1CȇƍȄæ2Qm
^2ȦŬ1*0C>D0ʆ0AʄŹǒǚ1,ʄɎÒ1@CȇU
WVƦę51ȿòĳ2ÄċƍƊ?ƽƻǇļǽGǌŘǇ1Ɔȩ!C.3-0
) & ʆ į 3 ȇ U W V ȵ 1 ɡ ! C ó ċ (p I k l ʄ _ j m ʄ K ]
0 / )2 ɤ Ē Ê G ƿ & Ɏ Ò 1 @ C ȇ U W V Ʀ ę 5 1 ȿ ò ĳ 2 Ä ċ ƍ Ɗ G Ȧ
23
Ŭ!Cǒǚʄ$,ȇUWVÄȦɗǩɤĒȔGƿ&ɎÒů2ȇUWV2ƽƻ
ǇļǽGŬA1!CǒǚĵȠ.0CʆǒǚA1ɍ=4ʄȇUWVGŔƌ.
&ȇƍȄ2ǴœĀɍ2&=2ɎÒÁŧ?Ywj2ɟǆ1*0C>D0
ʆ
24
Ǟʍǝ ǵŒ
Ɏ Ò ů 2 ɽ Ƅ ǟ - 3 ʄ Ⱥ ȓ ǥ ȼ - C U W V ư ĥ Ɵ Ò y (Ɏ Ò 2 ĩ ĥ ? œ ǳ
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ǒǚłžȤ
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1.
Takashi M atsui, Taro Ishikawa, Hitoshi Ito, Masahiro Okamoto, Koshiro Inoue,
Min-chul Lee, Takahiko Fujikawa, Yukio Ichitani, Kentaro Kawanaka, and Hideaki
Soya (2012). Brain glycogen supercompensation following exhaustive exercise. J Physiol
590, 607–616.
2.
Takashi Matsui, Shingo Soya, Masahiro Okamoto, Yukio Ichitani, Kentaro Kawanaka,
and Hideaki Soya (2011). Brain glycogen decreases during prolonged exercise. J Physiol
589, 3383-3393.
ǵȱȴţ
1.
Takashi Matsui and Hideaki Soya (2012). Brain glycogen metabolism and central fatigue
during prolonged exercise. Physiol News In press.
2.
ʆȇ ¾ U W V 2 Ʀ ę . ſ Ĺ Ǆ Ñ ʆ 60ʄ
ż ĝ ʄĮ ǐ ȏ Ů (2010)ʆ
797-804.
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1
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3
2010 ACSM International Student Award, American College of Sports Medicine ACSM (June 2010)
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27
Ûǿţƹ
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31

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