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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ȠǨ............................................................................................................................................................................................... 1
Ǟʈǝ Ƕȧ ........................................................................................................................................................................... 2
Ǟʉǝ [email protected] ........................................................................................................................................ 4
Ǟʊǝ ɞůɠɎÒ3ȇU†WV‹GƦę#CʒʂǒǚȲɶʈʃ ............................................... 5
Ǟʋǝ ɎÒů2ȇU†WV‹Ʀę3œdzůɠ®ČĹʒʂǒǚȲɶʉʃ.................................... 12
Ǟʌǝ ǵäȩȴ ............................................................................................................................................................... 20
Ǟʍǝ ǵŒ ......................................................................................................................................................................... 25
ǒǚłžŽȤ ......................................................................................................................................................................... 27
ȵɄ............................................................................................................................................................................................. 27
Ûǿţƹ................................................................................................................................................................................... 28
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ʉʅʈʆțɼÒ[email protected]¤
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Soya et al., 2007a; Nishijima et al., 2011b)ʆ
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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500µl2 6 %ɏ þ ǩ ɘ /1 mM EDTAƭ ƣ G Î Î Ɨ Ä Ȧ Ý Ķ G Ɛ = & ʆ 25,000 gʄ 4 10Ä ɠ ɑ Ĵ Ä ɪ ʄ  Ʈ ; G Ɨ ɘ Ô Q † L € ƭ ƣ (3 MƗ ɘ Ô Q † L € (KOH)ʄ 0.3 M
K  d b  ‡ (imidazole)ʄ 0.4 Mþ Ô Q † L € (KCl))- pH 6ʓ 82 ɠ 1 ȳ Ţ & ʆ
14,000 gʄ4 - 10Ä ɠ ɑ Ĵ Ä ɪ ʄ Ʈ ; G Î Ɨ Ä Ȧ & (U † W V ‹ Ä Ȧ D & )
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Flashʄ Thermo Fisher Scientificʄ USA)G ƿ & ʆ
$ D % D 2 L M ‡ 6 200µl2 Ý Ķ ƣ (50 mMj † ] -þ ɘ ƭ ƣ (Tris-HCL)pH 8.1ʄ0.5 mM
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8
(glucose-6-phosphatede hydrogenase))G Î & ʆ
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. ʄ Ð Ⱦ 355 nmʄ dž º 420 nm- Ǯ Ű & \ t k ‰ m W e ‹ I  k I i m ‹ \ n T
ˆ P e k † ‹ ɘ (NADPH)2 ɛ A U ‡ W  ] ɛ G Ȩ Ǣ & ʆ
U ‡ W  ] 3 molư ĥ - Ț ʄ Ǭ Ǻ 2 Ʃ ɚ ɛ - ȝ Ƒ , µmol/g wet tissue- Ǖ & ʆ
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Y ‹ w ‡ 2 U ‡ W  ] ư ĥ G Ğ Ĩ & ³ G . ʄU ‡ W  ] å Ƌ µmol/g wet tissue - Ǖ
&ʆ
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ĥ G Figure 2 1 Ǖ ! ʆȗ ǥ ³ 3 Exercise Ǽ 1 , Sedentary Ǽ . Ɩ 7 , 45 %© ) & (p
< 0.01)ʆǟ . ȁ Ȋ 2 U † W V ‹ 3 Exercise Ǽ 1 , sedentary Ǽ . Ɩ 7 , Ǩ 90 %Ʀ ę & ʆlj ȼ ʄơ ɻ ʄȢ ģ ‘ ɓ ʄĘ ȇ ʄȇ Ģ 2 U † W V ‹ ư ĥ exercise Ǽ 1 , sedentary
Ǽ . Ɩ 7 , Ǩ 50 %Ʀ ę & (p < 0.05)ʆ Ž ŧ ʄ – ɧ ʄ Ƿ ź ª ʄ Ȣ ģ ʄ – ȇ 2 U † W V ‹ ư
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50
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800
600
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400
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Exercise
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5
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Reduction rate of brain glycogen
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100
80
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e x e r c is e .
represent
( u n p a ir e d
B l o o d g l u c o s e , a n d g l y c o g e n l e v e l s i n t h e l i v e r , s k e l e ta l m u s c l e s , a n d t h e b r a i n a f t e r 2 h o f
A , B lo o d g l u c o s e ; B , L i v e r g l y c o g e n ; C , S k e l e t a l m u s c l e s g l y c o g e n ; D , B r a i n g l y c o g e n . D a ta
th e m e a n ± s t a n d a r d e r r o r ( n = 5 - 1 1 r a t s ) . * , p < 0 .0 5 ; * * , p < 0 .0 1 c o m p a r e d t o s e d e n t a r y r a ts
t - t e s t) .
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! ɞ ů ɠ (120 Ä ɠ )2 Ư Ɏ Ò … f j 2 ȇ U † W V ‹ ư ĥ 1 Ü 9 ! ī ɰ G Ɔ ȩ & ʆɎ Ò
1 @ B ȗ ǥ ³ 3 45 %© ‘ ʄǟ . ȁ Ȋ 2 U † W V ‹ 3 90 %Ʀ ę & ʆɎ Ò ů 2 © ȗ ǥ ?
ǟ Œ ȁ U † W V ‹ 2 ƀ ƥ 3 DŽ Ñ 2 Ŕ ƌ . D C . A (Nybo & Secher, 2004)ʄ Ź Đ ɼ ƿ&ɎÒ3ɯğ1ÚɎÒź¤-)&.ÄCʆ2.ʄljȼʄơɻʄȢ
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10
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Ò – 1 $ 2 Ɏ Ò ĩ ĥ ? œ dz ů ɠ 1 ® Č , Ʀ ę ! C (Gollnick et al., 1974)ʆ Ž ŧ ʄ ǒ ǚ Ȳ
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Ɏ Ò ů ʄ ȇ 2 ȗ – ˜ ɘ ß B Ʌ ; 3 ȗ – ˜ ɘ ư ĥ Ā Î ! C ˜ ɘ Ĺ « Ƈ ɢ ³ (lactate
threshold, LT)G ɀ C ĩ ĥ 1 , Ⱦ C ʄ LT @ B © ĩ ĥ - 3 Ⱦ A 0 . Ä
) , C (Ide et al., 2000)ʆ Ž ŧ ʄ Ń 2 ǒ ǚ đ - 3 ʄ ~ K T ‰ d K I † [ ] Ɯ G ƿ & ǒ ǚ 1 @ B ʄȽ Ɏ Ò ů 2 … f j ơ ɻ ¾ ˜ ɘ ȗ – ˜ ɘ 2 Ā Î 0 LT @ B © ĩ ĥ 2
Ɏ Ò 1 , > Ā Î ! C . G ȡ  & (ŷ ½ Ț i  c )ʆ 2 . A ʄ LT @ B © ĩ
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3 đ ¾ Ƨ ĥ 22±2 ʄ Ʃ ĥ 60±10%ʄ 7:00ʓ 19:00 G Ŭ Ŷ . & Ŭ ű Y K T ‡ G Ǵ œ & ʆ
ɹ Ť 1 3 Ò ƶ ƿ ÷ Ī ɹ Ť (MFʄ P † N ‹ c ‡ ɗ ƕ ʄ Japan)G ʄ ɸ Ť Ɨ 1 3 Ȓ ǂ Ɨ G $ D %
12
D ƿ ʄ . > 1 24 ů ɠ ȋ ǀ ś ß . & ʆ
ʉʅʉʆȽȘčǾ
… f j 1 3 1 Ɍ ɠ 2 š µ ɹ Ȃ 2 . ʄj ˆ f k  ‡ Ƚ Ɏ Ò 1 Ŀ D # C & = ʄ6 ũ ɠ 2
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1 ũ 30 Ä ɠ G Ȩ 5 ũ ɠ Ș ) & (Ț 1)ʆ 2 w ‰ j W  ‡ - Ƚ Ș č Ǿ G Ș ) & … f j 2 LT
3 ʄ @ $ 15ʓ 20 m/min - C . ǔ Ȯ D , C (Nishijima & Soya, 2006; Soya et al.,
2007a; Nishijima et al., 2011b)ʆ
ʉʅʊʆăɵɮȅQhh‡ǂǻņș
1 Ɍ ɠ 2 š µ ɹ Ȃ 2 . ʄ] j ˆ ] 2 0 ɿ ɕ Ȕ 2 ɮ ȅ ʼn “ G á Ȅ . ! C & = 1 ʄz ‹ j
r‡uc‡Ǽ2ăɵɮȅ1[†W‹ȞQhh‡Gǂǻ&ʆ…fj1z‹jruc
 ‡ ɿ ɕ (50 mg/kg B.W ., i.p.)G Ũ ʄ â ɝ ɽ  ɓ 2 lj Ȉ G Å ɟ & ʆ â ă ɵ ɮ ȅ G ɭ Â
į ʄ $ 2 Ž ɓ G ~ K T ‰ # H à - Å ɟ ʄ $ 2 Å ɟ ɓ A â Ĵ ń 1 < 10%x s † ‹ ƽ
ƻ ɷ þ Ɨ - ƪ & & Q h  h ‡ G 32 mm Ŗ » & ʆ dz , Q h  h ‡ G Ǧ - ă ɵ ɮ ȅ 1
÷ ď į ʄ į ɴ ɽ nj ‘ Ǩ 1 cm G Å ɟ ʄ $ A Q h  h ‡ G ɭ  # & ʆ ȗ ƣ ŋ C
. G ǔ Ȯ & . ʄ Å ɟ ɓ G Ǹ ä & ʆ $ 2 į ʄ Ľ Ɓ Dž G ɣ & = 1 Ŋ ƽ ƶ ȼ (Ò ƶ ƿ
~ 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
ʉʅʋʆɎÒĐɼ
… f j G Ɏ Ò É Ǽ ʄɯ Ɏ Ò Ǽ (j ˆ 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 ˆ 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
ĜƔ2.ťɴʄªĢȗ.ȇʄǟʄȁȊGŗß&ʆĐɼ2 2 ůɠÉA…fjGDZ
ɷ Ƹ ľ 1 ʄ Đ ɼ 3 ! 7 , × É – 1 Ș F D & ʆ Ƚ Ɏ Ò Đ ɼ 2 w ‰ j W  ‡ G Figure 12A
1Ǖ&ʆ
13
ʉʅʌʆǬǺ2ŗß
ǒ ǚ Ȳ ɶ ʈ . å Ƌ 1 ~ K T ‰ Ɲ Ƶ Ė Ȝ ǻ (NJE-2606ʄ Ŧ ũ Ź ƴ Ƿ ƃ ħ ¥ ǖ ʄ Japan) G ƿ
ʄ 10 kW 2 ~ K T ‰ Ɲ G 1.2 ǘ ɠ Ƶ Ė & . ȇ G ŗ ß & ʆ ȇ 3 Hirano et al. (2006)
2ŧƜ1İʄljȼʄ–ɧʄǷźªʄơɻʄȢģʄȢģ‘ɓʄ–ȇʄĘȇʄȇĢ2ʐɓ
¨1Äǁ&ʆåů1t…ǟʄɁĤǟʄȁȊ>ŗß&ʆŗß&ǬǺ3ƣªǛǩÀ Ǯ # ʄ U † W V ‹ 2 ď ɛ 1 ƿ C : - ʅ 80 - ± Č & ʆ Đ ɼ 1 3 lj ȼ ʄ ơ ɻ ʄ Ȣ
ģ‘ɓʄĘȇʄȇĢ2ʌɓ¨Gƿ&ʆ
ʉʅʍʆȗƣłÄ2ƨď
ǂ ǻ & Q h  h ‡ A 2 ɮ ȅ ȗ (x s † ‹ Á ƻ )G ƿ , ʄ 1 U ‡ W  ] /… T h  j
I l … K Z  (2300 STAT PLUSʄ YSI, USA)- ȗ ǥ ³ . ȗ – ˜ ɘ ³ G ƨ ď & ʆ
[email protected]]ưĥ2ďɛ
ȇ 2 U † W V ‹ @ 5 U ‡ W  ] 2 Ō  3 Kong et al. (2002)1 ʄ U ‡ W  ] ư ĥ 2 ƨ ď
3 Passonneau & Lauderdale (1974)2 ŧ Ɯ 1 İ ) & ʆ ņ ɲ 3 ǒ ǚ Ȳ ɶ ʈ . å Ƌ 1 Ș ) & ʆ
ʉʅʏʆȇ˜ɘưĥ2ďɛ
Passonneau & Lauderdale (1974)2 ŧ Ɯ 1 @ C R f j (DiaSysʄGermany)1 @ B ď ɛ & ʆ
ʉʅʐʆȇ¾‚pI‹2ďɛ
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 Ž · 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
ʊʅʈʆȗǥʄȗ–˜ɘʄ•51ǟŒȁŒȇU†WV‹ưĥ
Ä Ɋ 20 m ʄ30ʄ60ʄ120 Ä ɠ 2 j ˆ f k  ‡ Ƚ Ɏ Ò nj į 2 ȗ ǥ ʄȗ – ˜ ɘ ʄ• 5 1 ǟ Œ
ȁ Œ 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 Œ ȁ U † W V ‹ ư
ĥ3ɎÒœdzůɠ®ČLJ1Ʀę&ʆȗǥ.åƋ1ʄljȼʄơɻʄȢģ‘ɓʄĘȇʄȇ
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
ʊʅʉʆȇ¾U‡W]•51˜ɘưĥ
Ä Ɋ 20 mʄ 30ʄ 60ʄ 120 Ä ɠ 2 j ˆ 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
ʊʅʊʆȗǥʄȇ¾U‡W]ʄȇ¾˜ɘ.ȇU†WV‹2Ǎɡ
ȗ ǥ ʄ ȇ ¾ U ‡ W  ] ʄ ȇ ¾ ˜ ɘ . ȇ U † W V ‹ 2 Ǎ ɡ G Figure 5 1 Ǖ ! ʆ ȗ ǥ • 5
1ȇU‡W].ȇU†WV‹.2ɠ13¼,2ɓ¨-ɯğ1ɾƑ2ǍɡȮ=AD
&ʆȇ˜ɘ.ȇU†WV‹.2ɠ13Ȣģ‘ɓGɦ¼,2ɓ¨-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
ʊʅʋʆljȼ‚pI‹ưĥʄ•51ljȼ‚pI‹.U†WV‹ưĥ2Ǎɡ
p‡Ikˆl†‹2 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_‰jm‹2 ȵƾƶ
20
50
jR[tk‰R[vMm‡U†W
#
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\tk‰R[vMm‡ɖ
ɘ (DOPAC) 3 Ā Î ¶ æ 1 C > 2 2 ŵ
ļ0āÔ30)&ʆ$2Ÿ2ƨďɱ
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 Œ 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 ʆŽ ŧ ʄɎ Ò 120
18
Ä2ůƳ-3©ȗǥȾBʄǟ?ȁȊ2U†WV‹>ƀƥ,CʆD3ǒǚȲɶ
ʉʅʈ2Ƹƛ.89åƋ-ɎÒ[email protected]Ñ,CvM^-C.ĸFDCʆ
2 . ʄlj ȼ ʄơ ɻ ʄȢ ģ ‘ ɓ ʄĘ ȇ ʄȇ Ģ 2 U † W V ‹ ư ĥ 3 š 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
&ȗǥ•51ȇU‡W].ȇU†WV‹2ɠ1ɯğ1ɾƑ2ǍɡǔȮD&ʆ
2.Aʄȗǥ?ȇU‡W]3ɎÒů2ȇU†WV‹Ʀę2ƙďóċ-CáȄĹ
CʆA1ʄ2.ȇ¾-ĀÎ&˜ɘ.ȇU†WV‹2ɠ1Ȣģ‘ɓ¢ă2ʋ
ɓ¨-ȷ2ǍɡȡAD&ʆ2Ǯž3ʄɎÒů1ȇU†WV‹˜ɘ1ÄȦDʄm
„‰‹1­ǯD&áȄĹGǕ!ʆ
Î,ʄŹĐɼ-3ljȼ¾‚pI‹.$2 ȵƾƶ>ďɛ&.Eʄp‡Ikˆ
l † ‹ 2 ȵ ƾ ƶ - C MHPG . _ ‰ j m ‹ 2 ȵ ƾ ƶ - C 5-HIAA Ɏ Ò 120 Ä Ǽ
-ĀÎ&ʆp‡Ikˆl†‹?_‰jm‹3ȇU†WV‹2ÄȦ°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!ɞůɠɎÒ[email protected]_‰j
m‹2 ȵɍɡ“,CáȄĹCʆ
ɎҖ2©ȗǥ?ȇ¾_‰jm‹ưĥ2ĀÎ3ɎÒů2–ſDŽÑ2Ùó.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?_‰jm‹2ĀÎ1§ȇU
†WV‹2ƦęɎÒů2–ſDŽÑ2ǰäȠó-AáȄĹGǕï!Cʆžįʄ›É1
ȇU†WV‹ưĥGɾ=&ýä1œ—ĹsvO~‹]æ!C/Ɔȩ!CĵȠ
Cʆ
19
Ǟʌǝ ǵäȩȴ
ɎÒůʄȇ3ǟ.åƋ1ƟĹÔʄȇ1CNo‡S?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 ˆ y ‡ (Ɏ Ò 2 ĩ ĥ ? œ dz
ů ɠ )1 ® Č , Ʀ ę ! C (Gollnick et al., 1974)ʆŽ 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-3ʄȇU†WV‹ďɛ2X‡i‹]c‹dk-C~KT‰ƝƵĖƜG
ė»ʄɞůɠɎÒů1ȇU†WV‹Ʀę!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!.AʄȇU†W
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§ɞůɠɎÒů2ȇU†WV‹ưĥG~KT‰ƝƵĖƜGƿ,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?ǟŒȁU†WV‹2ƀƥ3DŽÑ2Ŕƌ.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
ȇU†WV‹GƦę#C.Æ=,ŬA10)&ʆ©ȗǥ3ȇU†WV‹GƦ
ę#CȠó.,@ǑAD,C.Aʄ2Ǯž3ʄɞůɠɎÒů1ĨȾ
D&©ȗǥȗƣAȇ62No‡S­ǯ’ɁGőʄ$2’ɁGȝ&=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 .
j‰YKj-U†WV‹ÄȦɍ&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¡?ɎÒ2w‰U…‹Uʄơɻ3ɎÒů
2ȮǑʄȢģ‘ɓ3ªƧ?No‡S ȵ2ȳǤʄĘȇ3ǟ2Øȳ?ĊÓ2ǴœʄȇĢ
3ìé?Ĵŏ2ȳǤ0/ʄɎÒů1ƟĹÔ!C.ǿADCɓ¨-Cʆ&),ʄ
ɞůɠɎÒů2ȇU†WV‹Ʀę13©ȗǥ'-0ɎÒů2m„‰‹2ƟĹÔ>
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
…fj2ȇU†WV‹ưĥ1Ü9!īɰGƆȩ&ʆȇU†WV‹ɎÒů2m„
21
‰‹ƟÒ1Ķ
, Ç ƿ D Ʀ ę ! C 0 A ʄ ǟ U † W V ‹ Ɏ Ò œ dz ů ɠ (ǟ Ɵ Ò ˆ y ‡ )
1®Č,[email protected]>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.ȇU†WV‹2ɠ1ɯğ1ɾƑ2ǍɡǔȮD&ʆD
A2Ǯž3ʄȗǥɎÒů2ȇU†WV‹Ʀę2ƙďóċ-CáȄĹGǕï!Cʆ
A1ʄ2.Ȣģ‘ɓ¢ă2ʋɓ¨1,ʄȇ¾-ĀÎ&˜ɘ.ȇU†WV‹.
2ɠ1ȷ2ǍɡȡAD&ʆ2Ǯž3ʄɞůɠɎÒů1ȇU†WV‹˜ɘ1ÄȦ
D & . G Ǖ ʄ ɞ ů ɠ Ɏ Ò ů 2 I ] j ‰ Y K j m „  ‰ ‹ ˜ ɘ [ ƒ j ‡ (Pellerin &
Magistretti, 1994)1 C ˜ ɘ 2 ­ ǯ ƫ . , U † W V ‹ ȹ ƹ , C á Ȅ Ĺ G Ǖ ï
!CʆÎ,ʄI]j‰YKj2U†WV‹ÄȦ°ɍóċ.,ǑADCp‡Ikˆl
†‹._‰jm‹2 ȵGljȼ1,ďɛ&.Eʄp‡Ikˆl†‹2 ȵƾƶ
- C MHPG . _ ‰ j m ‹ 2 ȵ ƾ ƶ - C 5-HIAA Ɏ Ò 120 Ä Ǽ - Ā Î ʄ Ʀ ę &U†WV‹ưĥ.ȷ2ǍɡGǕ&ʆp‡Ikˆl†‹?_‰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!CʆA1ʄȇU†WV‹ ȵ1ɡ“0ks‹2 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
ȇU†WV‹Ʀę13ȌćĹǗǭ¦ɐƶȼ-Cp‡Ikˆl†‹._‰jm‹2 ȵ
 ɍ ɡ “ , C á Ȅ Ĺ C (Fig. 7)ʆ
ŹĐɼ1,ʄȗǥ?_‰jm‹ɎÒů2ȇU†WV‹Ʀę2ƙďóċ-Cá
ȄĹȡÂD&ʄDA3ɞůɠɎÒů2–ſDŽÑ2Ƞó.,>ǑADCʆ&
),ʄȇU†WV‹3ɞůɠɎÒů2ȇ¾_‰jm‹ ȵ2ɍ?ȗǥ³2©‘1@
BÇƿDƦęʄ$2ƦęɞůɠɎÒů2–ſDŽÑ2ǰäóċ.0C2>D0
(Fig. 8)ʆ D 1 * , 3 ʄ ž 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 .
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ǒǚłžŽȤ
Ùȑȴţ
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 œ ĝ ʄĮ ǐ ȏ Ů (2010)ʆ
797-804.
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1
2
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3
2010 ACSM International Student Award, American College of Sports Medicine ACSM (June 2010)
ȵɄ
Ź ǒ ǚ 3 ʄǞ ʎ ò  Ŵ ] }  g ʔş Ȃ ȸ ô ] }  g ǒ ǚ Ï ł › Ƈ 2 Ś Ï 1 @ B Ș F D :
&ʆ1ƤĽȵ2ļGȚ:!ʆ
27
Ûǿţƹ
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