1 繊毛虫テトラヒメナ(Tetrahymena thermophila)のセリンラセマーゼの酵素特性 宇田幸司・山城由也・西願麻以 〒780-8520 高知市曙町 2−5−1 高知大学理工学部比較生化学研究室 Enzymatic properties of the serine racemase from Tetrahymena thermophila Kouji Uda, Yuya Yamashiro and Mai Saigan Laboratory of Comparative Biochemistry, Faculty of Science and Technology, Kochi University, Kochi 780-8520 Japan Key Words; amino acid racemase, D-amino acid, D-serine, serine racemase. 要旨 セリンラセマーゼ(SerR)は光学異性体である L-セリンと D-セリンの相互変換反応と,L-セリンまたは D-セ リンをピルビン酸に分解する反応を触媒する酵素である。 SerR は動物及び植物に広く分布しており,生体内 D- セリンの合成酵素であると考えられている。本研究では,繊毛虫テトラヒメナ(Tetrahymena thermophila)から 発見された SerR 遺伝子について,そのリコンビナント酵素を用いた詳細な酵素特性の解明を行った。また, テトラヒメナの増殖が D-セリンによって阻害されること等から,テトラヒメナにおいては,SerR が外部から 取り込まれた D-セリンの分解酵素として働く可能性を指摘した。 Abstract Serine racemase (SerR) is an enzyme that catalyzes the conversion of L-serine and D-serine and the degradation of L-serine or D-serine to pyruvate. SerR is widely distributed in animals and plants and is thought to be involved in the synthesis of D-serine in vivo. In this study, we constructed the recombinant protein of the ciliate Tetrahymena thermophila SerR gene and measured its detailed enzymatic properties. Since the growth of Tetrahymena thermophila is inhibited by D-serine, we suggested that SerR might act as a degrading enzyme for D-serine taken up from the outside in vivo. 序論 アミノ酸には L 体と D 体の 2 種類の鏡像異性体が存在するが,タンパク質の合成材料となるのは L 体のアミ ノ酸のみである。そのため,生体内では遊離アミノ酸として L-アミノ酸のみが存在し,D-アミノ酸とその代謝 酵素はごく限られた生物にしか存在しないと長い間考えられてきた。しかしながら,近年の研究によって,D- アミノ酸が多くの生物に存在し,多様な生理機能を担うことが明らかとなっている(宇田, 2016)。アミノ酸ラ Scientific and Educational Reports of the Faculty of Science and Technology, Kochi University Vol. 3 (2020), No. 2 Received: March 27, 2020 Reviewed by anonymous referee(s), and accepted: March 30, 2020 Published: April 9, 2020
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1
繊毛虫テトラヒメナ(Tetrahymena thermophila)のセリンラセマーゼの酵素特性
宇田幸司山城由也西願麻以
780-8520 高知市曙町 2minus5minus1
高知大学理工学部比較生化学研究室
Enzymatic properties of the serine racemase from Tetrahymena thermophila
Kouji Uda Yuya Yamashiro and Mai Saigan
Laboratory of Comparative Biochemistry Faculty of Science and Technology Kochi University
Kochi 780-8520 Japan
Key Words amino acid racemase D-amino acid D-serine serine racemase
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
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25 mM L-Ser
50 mM L-Ser
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11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
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Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
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15 mM D-Ser
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2 mM L-Ala
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2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
3
曲線を作製した
結果と考察
テトラヒメナ SerR 遺伝子の合成とリコンビナント酵素の作製
テトラヒメナでは一般的な生物において終止コドンとして用いられる TAA 及び TAG がグルタミンをコー
ドしているテトラヒメナの SerR 遺伝子のコード領域にも 15 個の TAA または TAG でコードされるグルタミ
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
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Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
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025
030
0 5 10 15 20
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Den
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[5
40 n
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Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
6
Table 2 真核生物SerRの酵素活性
K m k cat k catK m
[mM] [10-3s-1
] [10-3s-1mM-1]
オニクマムシSerR L-Ser rarr D-Ser Uda et al 2016 284 plusmn 42 175 plusmn 17 0621 plusmn 0046
(Milnesium tardigradum ) D-Ser rarr L-Ser Uda et al 2016 190 plusmn 19 970 plusmn 088 0513 plusmn 0037
プラナリアSerR L-Ser rarr D-Ser Uda et al 2016 112 plusmn 18 947 plusmn 102 851 plusmn 062
(Dugesia japonica ) D-Ser rarr L-Ser Uda et al 2016 683 plusmn 040 655 plusmn 35 960 plusmn 006
ウシエビSerR L-Ser rarr D-Ser Uda et al 2016 146 plusmn 5 817 plusmn 056 00559 plusmn 00038
(Penaeus monodon ) D-Ser rarr L-Ser Uda et al 2016 119 plusmn 16 772 plusmn 113 00651 plusmn 00012L-Ser rarr Pyruvate Uda et al 2017 474 plusmn 34 489 plusmn 23 104 plusmn 009
マガキSerR L-Ser rarr D-Ser Uda et al 2017 215 plusmn 06 178 plusmn 001 00827 plusmn 00022
(Crassostrea gigas ) D-Ser rarr L-Ser Uda et al 2017 293 plusmn 46 211 plusmn 025 00728 plusmn 00047L-Ser rarr Pyruvate Uda et al 2017 226 plusmn 069 291 plusmn 21 140 plusmn 39
ハイマツミドリイシSerR L-Ser rarr D-Ser Uda et al 2016 771 plusmn 065 414 plusmn 25 538 plusmn 012
(Acropora millepora ) D-Ser rarr L-Ser Uda et al 2016 848 plusmn 019 551 plusmn 16 650 plusmn 009
マウスSerR L-Ser rarr D-Ser Uda et al 2016 149 plusmn 07 668 plusmn 36 447 plusmn 09
(Mus musculus ) D-Ser rarr L-Ser Uda et al 2016 209 plusmn 07 840 plusmn 34 402 plusmn 05L-Ser rarr Pyruvate Uda et al 2017 479 plusmn 130 285 plusmn 28 621 plusmn 97D-Ser rarr Pyruvate Uda et al 2017 261 plusmn 030 395 plusmn 34 152 plusmn 09
ラットSerR L-Ser rarr D-Ser Wolosker et al 1999 98 50 51
(Rattus norvegicus ) D-Ser rarr L-Ser Wolosker et al 1999 60 220 37
ヒトSerR L-Ser rarr D-Ser Hoffman et al 2009 41 plusmn 02 691 plusmn 25 166 plusmn 133
(Homo sapiens ) D-Ser rarr L-Ser Hoffman et al 2009 108 plusmn 08 1415 plusmn 70 131 plusmn 15L-Ser rarr Pyruvate Hoffman et al 2009 47 plusmn 04 1660 plusmn 90 353 plusmn 45D-Ser rarr Pyruvate Hoffman et al 2009 95 plusmn 06 128 plusmn 5 135 plusmn 15
キイロタマホコリカビSerR L-Ser rarr D-Ser Ito et al 2012 304 plusmn 27 1460 plusmn 60 476
(Dictyostelium discoideum ) D-Ser rarr L-Ser Ito et al 2012 340 plusmn 52 1160 plusmn 80 34L-Ser rarr Pyruvate Ito et al 2013 67 plusmn 11 655 plusmn 25 977D-Ser rarr Pyruvate Ito et al 2013 224 plusmn 38 718 plusmn 42 317
イネSerR L-Ser rarr D-Ser Gogami et al 2010 100 310 31
(Oryza sativa ) D-Ser rarr L-Ser Gogami et al 2010 89 200 22L-Ser rarr Pyruvate Gogami et al 2010 180 360 20D-Ser rarr Pyruvate Gogami et al 2010 63 120 18
シロイヌナズナSerR L-Ser rarr D-Ser Fujitani et al 2006 25 29 117
(Arabidopsis thaliana ) D-Ser rarr L-Ser Fujitani et al 2006 077 064 083L-Ser rarr Pyruvate Fujitani et al 2006 20 140 70D-Ser rarr Pyruvate Fujitani et al 2006 50 15 30
オオムギSerR L-Ser rarr D-Ser Fujitani et al 2007 26 12 46
(Hordeum vulgare ) D-Ser rarr L-Ser Fujitani et al 2007 90 265 29L-Ser rarr Pyruvate Fujitani et al 2007 27 17 063D-Ser rarr Pyruvate Fujitani et al 2007 83 96 12
キャッサバSerR L-Ser rarr D-Ser Uda et al 2020 906 plusmn 090 60 plusmn 77 676 plusmn 141
(Manihot esculenta ) D-Ser rarr L-Ser Uda et al 2020 823 plusmn 020 517 plusmn 51 628 plusmn 061L-Ser rarr Pyruvate Uda et al 2020 347plusmn 18 203 plusmn 2 587 plusmn 029D-Ser rarr Pyruvate Uda et al 2020 435 plusmn 77 110 plusmn 7 263 plusmn 066
トマトSerR L-Ser rarr D-Ser Uda et al 2020 117 plusmn 31 287 plusmn 49 251 plusmn 023
(Solanum lycopersicum) D-Ser rarr L-Ser Uda et al 2020 233 plusmn 22 428 plusmn 38 184 plusmn 005L-Ser rarr Pyruvate Uda et al 2020 330 plusmn 18 343 plusmn 31 104 plusmn 01D-Ser rarr Pyruvate Uda et al 2020 421 plusmn 120 159 plusmn 25 389 plusmn 043
分裂酵母SerR L-Ser rarr D-Ser Yamauchi et al 2009 32 18 056(Schizosaccharomyces pombe ) D-Ser rarr L-Ser Yamauchi et al 2009 96 41 043
L-Ser rarr Pyruvate Yamauchi et al 2009 36 510 14D-Ser rarr Pyruvate Yamauchi et al 2009 10 30 30
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
9
25 mM まで変化させても増殖曲線に影響は見られず50 mM まで増加したときのみ増殖の阻害が確認され
た(Fig 7A)一方でD-セリンを添加した場合には終濃度 8 mM を超えると増殖が阻害されることがわか
った(Fig 7B)またL-アラニンD-アラニン及びグリシンを添加した場合はL-セリン同様に終濃度 50 mM
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
宇田幸司 (2016) セリンアスパラギン酸ラセマーゼの構造と機能の進化 比較生理生化学 33(2) 68-76 Fujitani Y Nakajima N Ishihara K Oikawa T Ito K amp Sugimoto M (2006) Molecular and biochemical characterization of a
serine racemase from Arabidopsis thaliana Phytochemistry 67(7) 668-674
A D
B E
C
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ser
Control (15 mM L-Ser)
8 mM L-Ser
15 mM L-Ser
25 mM L-Ser
50 mM L-Ser
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ser
2 mM D-Ser
8 mM D-Ser
15 mM D-Ser
25 mM D-Ser
50 mM D-Ser
Control
000
005
010
015
020
025
030
035
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM L-Ala
2 mM L-Ala
8 mM L-Ala
15 mM L-Ala
25 mM L-Ala
50 mM L-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM D-Ala
2 mM D-Ala
8 mM D-Ala
15 mM D-Ala
25 mM D-Ala
50 mM D-Ala
Control
000
005
010
015
020
025
030
0 5 10 15 20
Op
tical
Den
sity
[5
40 n
m]
Time[h]
05 mM Gly
2 mM Gly
8 mM Gly
15 mM Gly
25 mM Gly
50 mM Gly
Control
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424
11
Fujitani Y Horiuchi T Ito K amp Sugimoto M (2007) Serine racemases from barley Hordeum vulgare L and other plant species represent a distinct eukaryotic group gene cloning and recombinant protein characterization Phytochemistry 68(11) 1530-1536
Gogami Y Ito K Kamitani Y Matsushima Y amp Oikawa T (2009) Occurrence of D-serine in rice and characterization of rice serine racemase Phytochemistry 70(3) 380-387
Gogami Y Kobayashi A Ikeuchi T amp Oikawa T (2010) Site‐Directed Mutagenesis of Rice Serine Racemase Evidence That Glu219 and Asp225 Mediate the Effects of Mg2+ on the Activity Chemistry amp biodiversity 7(6) 1579-1590
Goto M Yamauchi T Kamiya N Miyahara I Yoshimura T Mihara H amp Esaki N (2009) Crystal structure of a homolog of mammalian serine racemase from Schizosaccharomyces pombe Journal of Biological Chemistry 284(38) 25944-25952
Hoffman H E Jiraacuteskovaacute J Ingr M Zvelebil M amp Konvalinka J (2009) Recombinant human serine racemase enzymologic characterization and comparison with its mouse ortholog Protein expression and purification 63(1) 62-67
Ito T Murase H Maekawa M Goto M Hayashi S Saito H amp Yoshimura T (2012) Metal ion dependency of serine racemase from Dictyostelium discoideum Amino acids 43(4) 1567-1576
Ito T Maekawa M Hayashi S Goto M Hemmi H amp Yoshimura T (2013) Catalytic mechanism of serine racemase from Dictyostelium discoideum Amino Acids 44(3) 1073-1084
Kumar S Stecher G amp Tamura K (2016) MEGA7 molecular evolutionary genetics analysis version 70 for bigger datasets Molecular biology and evolution 33(7) 1870-1874
Radkov A D amp Moe L A (2014) Bacterial synthesis of D-amino acids Applied microbiology and biotechnology 98(12) 5363-5374 Střıšovskyacute K Jiraacuteskovaacute J Bařinka C Majer P Rojas C Slusher B S amp Konvalinka J (2003) Mouse brain serine racemase
catalyzes specific elimination of L-serine to pyruvate FEBS letters 535(1-3) 44-48 Szablewski L Andreasen P H Tiedtke A Florin‐Christensen J Florin‐Christensen M amp Rasmussen L (1991) Tetrahymena
thermophila Growth in synthetic nutrient medium in the presence and absence of glucose The Journal of protozoology 38(1) 62-65 Uda K Edashige Y Nishimura R Shikano Y Matsui T Radkov A D amp Moe L A (2020) Distribution and evolution of the
serineaspartate racemase family in plants Phytochemistry 169 112164 Uda K Abe K Dehara Y Mizobata K Sogawa N Akagi Y amp Moe L A (2016) Distribution and evolution of the
serineaspartate racemase family in invertebrates Amino Acids 48(2) 387-402 Uda K Abe K Dehara Y Mizobata K Edashige Y Nishimura R amp Moe L A (2017) Triple serine loop region regulates the
aspartate racemase activity of the serineaspartate racemase family Amino acids 49(10) 1743-1754 Wolosker H Sheth K N Takahashi M Mothet J P Brady R O Ferris C D amp Snyder S H (1999) Purification of serine racemase
biosynthesis of the neuromodulator D-serine Proceedings of the National Academy of Sciences 96(2) 721-725 Yamauchi T Goto M Wu H Y Uo T Yoshimura T Mihara H amp Esaki N (2009) Serine racemase with catalytically active
lysinoalanyl residue Journal of biochemistry 145(4) 421-424