このページはRCSBの David S. Goodsell博士による「Molecule of the Month」2002年12月の記事を日本語に訳したものです。転載・引用については利用規約をご覧下さい。
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:翻訳 工藤高裕 (PDBj)
シトクロムc(PDB:3cyt)

電気は現代における最も一般的な現象の一つで、部屋の明かりからあなたの目の前にあるコンピュータまであらゆるものに動力を供給している。その電気とは、金属線など導電性物質の中を電子が通ることである。電子は、線を伝って原子と原子の間の曲がりくねった空洞を流れる。細胞も様々な反応過程に動力を供給するのに電気を使うが、電子の移動方法はかなり違っている。電子は細胞サイズの線を滑らかには流れない。その代わり電子は一度に1つずつ、タンパク質からタンパク質へと飛びながら輸送される。このようにして電子はある特定の場所から拾い上げ、必要な場所へと正確に届けることができるのである。

電子伝達

ここに示したシトクロムc(cytochrome c)は PDBエントリー 3cyt から得られた電子の運び屋である。電子を運ぶその他多くのタンパク質と同様、つかまえにくい電子を扱うのに特別な補欠分子族(prosthetic group)を持っている。シトクロムcは、図中では赤色で示した鉄イオンを含むヘム基(heme group)を内側に強くつかんで持っている。この鉄イオンは簡単に電子の取り込みと解放を行う。そして周りのタンパク質は、どれだけしっかり電子をつかんでいるかを調整し電子にとって完全な環境を作り出す。また次に示すように、シトクロムcが細胞の電子回路全体におけるどの位置に当てはまるかについても決めている。

古くからある群

シトクロムcは古くからあるタンパク質で、生命進化の早い段階に発達した。このなくてはならないタンパク質は、細胞の動力生産において重要な段階を担っており、ここ数百万年の間ほとんど変化してこなかった。だから、酵母、植物、そして我々の細胞どれを探しても非常に似た型のシトクロムcが見つかるのである。ところがPDBを見渡すと、他にも様々な電子運搬分子があることが分かる。シトクロムcには様々な変異があり、いずれも電子運搬にはヘムと鉄イオンを使うが、担当する仕事の違いに応じて電子を囲むタンパク質を変化させている。他の電子運搬体はまた別の補欠分子族を使って電子を運んでいる。そのような補欠分子族としては、フェレドキシン(ferredoxin)で使われている鉄硫黄クラスター(iron-sulfur cluster)や、アズリン(azurin)やプラストシアニン(plastocyanin)で使われる銅イオンのほか、より珍しい金属イオンを使ったものも存在する。シトクロムcのように、これらのタンパク質もそれぞれ単独で細胞電子回路に接続していて、ある地点から別の地点へ電子を運搬している。

細胞の回路

シトクロムbc 1 複合体(左:PDB:1kyo)とシトクロムc酸化酵素(右:PDB:1oco) ピンクの分子はシトクロムc

シトクロムcは細胞内に配線された電気回路に1つの接点を形成する。そして、細胞のエネルギー生産における最終段階で電子を運搬する。この電子は元々糖の分解を通じて得られたもので、最終的には酸素に付加されて水になる(これが、我々が呼吸で取り入れた酸素の最終的な運命である)。シトクロムcは、2つの大きなタンパク質複合体の間で個々の電子を輸送する。つまり、上図左(PDBエントリー 1kyo )の シトクロムbc 1 (cytochrome bc 1 )複合体から電子を集め、上図右(PDBエントリー 1oco )に示す シトクロムc酸化酵素 に電子を渡している(2000年5月、今月の分子5番も参照のこと)。この2つの複合体がエネルギー生産という重労働を行っている。電子が電子輸送グループ(赤色)を通って流れるのに従い、黄色い帯で示した膜の反対側へプロトン(水素イオン)がくみ出される。そしてこのプロトンはATP生産の動力源として用いられる。シトクロムcは、全てのエンジンを滑らかに動かし続けることで、一つの複合体から別の複合体へ必要に応じて電子を輸送している。

構造を見る

シトクロムc(上のピンク色の管で示された分子)とシトクロムbc 1 複合体(下の黄色い管で示された分子) 電子を輸送するヘム基の各原子は球で、その中の鉄原子は黄色い球で示す。PDB:1kyo

PDBエントリー 1kyo では、細胞内で電子がどのように輸送されるかを拡大して見ることができる。電子は、身近で見かける電気器具のように連続的な線を通って流れる訳ではない。電子はこの短い距離を、輸送体から隣の輸送体へと直接移動する。上図にはシトクロムc(上)と大きなシトクロムbc 1 複合体(下)との複合体を示している。シトクロムcのタンパク質鎖はピンク色の管で、bc1複合体のタンパク質鎖は黄色い管で示す。ヘムは各原子とも球で、鉄原子は黄色の球で示す。シトクロムcのヘム基が押し出され、bc1複合体のヘム基に近づいていることに注目して欲しい。電子はこの距離を、bc1複合体のヘムからシトクロムcのヘムへ、百万分の1秒未満で移動する。

2002年12月時点でPDBに登録されているエントリーリストはこちらのリスト

2002年12月 のキーワード検索による関連PDBエントリー一覧
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PDB ID Title Authors Publication Year Journal Name Volume No First Page Pubmed ID
155c The structure of Paracoccus denitrificans cytochrome c550. R.Timkovich, R.E.Dickerson 1976 J Biol Chem 251 4033 180013
19hc The Primary and Three-Dimensional Structures of a Nine-Haem Cytochrome C from Desulfovibrio Desulfuricans Atcc 27774 Reveal a New Member of the Hmc Family P.M.Matias, R.Coelho, I.A.Pereira, A.V.Coelho, A.W.Thompson, L.C.Sieker, J.L.Gall, M.A.Carrondo 1999 Structure (London) 7 119 10368280
1a2s Solution structure of oxidized cytochrome c6 from the green alga Monoraphidium braunii. L.Banci, I.Bertini, M.A.De la Rosa, D.Koulougliotis, J.A.Navarro, O.Walter 1998 Biochemistry 37 4831 9538000
1a56 Primary sequence and solution conformation of ferrocytochrome c-552 from Nitrosomonas europaea. R.Timkovich, D.Bergmann, D.M.Arciero, A.B.Hooper 1998 Biophys J 75 1964 9746537
1a7v Basis for monomer stabilization in Rhodopseudomonas palustris cytochrome c' derived from the crystal structure. N.Shibata, S.Iba, S.Misaki, T.E.Meyer, R.G.Bartsch, M.A.Cusanovich, Y.Morimoto, Y.Higuchi, N.Yasuoka 1998 J Mol Biol 284 751 9826513
1akk Solution structure of oxidized horse heart cytochrome c. L.Banci, I.Bertini, H.B.Gray, C.Luchinat, T.Reddig, A.Rosato, P.Turano 1997 Biochemistry 36 9867 9245419
1aqe Structural and kinetic studies of the Y73E mutant of octaheme cytochrome c3 (Mr = 26 000) from Desulfovibrio desulfuricans Norway. C.Aubert, M.T.Giudici-Orticoni, M.Czjzek, R.Haser, M.Bruschi, A.Dolla 1998 Biochemistry 37 2120 9485359
1ayg Solution structure of thermostable cytochrome c-552 from Hydrogenobacter thermophilus determined by 1H-NMR spectroscopy. J.Hasegawa, T.Yoshida, T.Yamazaki, Y.Sambongi, Y.Yu, Y.Igarashi, T.Kodama, K.Yamazaki, Y.Kyogoku, Y.Kobayashi 1998 Biochemistry 37 9641 9657676
1b7v Crystal Structure of Oxidized Bacillus Pasteurii Cytochrome C(553) at 0.97-A Resolution S.Benini, A.Gonzalez, W.R.Rypniewski, K.S.Wilson, J.J.Van-Beeumen, S.Ciurli 2000 Biochemistry 39 13115 11052663
1bbh Atomic structure of a cytochrome c' with an unusual ligand-controlled dimer dissociation at 1.8 A resolution. Z.Ren, T.Meyer, D.E.McRee 1993 J Mol Biol 234 433 8230224
1bvb Heme packing motifs revealed by the crystal structure of the tetra-heme cytochrome c554 from Nitrosomonas europaea. T.M.Iverson, D.M.Arciero, B.T.Hsu, M.S.Logan, A.B.Hooper, D.C.Rees 1998 Nat Struct Biol 5 1005 9808046
1c2n Solution structure, rotational diffusion anisotropy and local backbone dynamics of Rhodobacter capsulatus cytochrome c2. F.Cordier, M.Caffrey, B.Brutscher, M.A.Cusanovich, D.Marion, M.Blackledge 1998 J Mol Biol 281 341 9698552
1c53 S-class cytochromes c have a variety of folding patterns: structure of cytochrome c-553 from Desulfovibrio vulgaris determined by the multi-wavelength anomalous dispersion method. A.Nakagawa, Y.Higuchi, N.Yasuoka, Y.Katsube, T.Yagi 1990 J Biochem (Tokyo) 108 701 1964450
1c6o Amino acid sequence, crystallization and structure determination of reduced and oxidized cytochrome c6 from the green alga Scenedesmus obliquus. J.Schnackenberg, M.E.Than, K.Mann, G.Wiegand, R.Huber, W.Reuter 1999 J.Mol.Biol. 290 1019 10438600
1c6r
1c6s Solution structure of cytochrome c6 from the thermophilic cyanobacterium Synechococcus elongatus. M.Beissinger, H.Sticht, M.Sutter, A.Ejchart, W.Haehnel, P.Rosch 1998 EMBO J 17 27 9427738
1c75 Crystal Structure of Oxidized Bacillus Pasteuriicytochrome C(553) at 0.97-A Resolution S.Benini, A.Gonzalez, W.R.Rypniewski, K.S.Wilson, J.J.Van-Beeumen, S.Ciurli 2000 Biochemistry 39 13115 11052663
1c7m Solution Structure of the Functional Domain of Paracoccus Denitrificans Cytochrome C552 in the Reduced State P.Pristovsek, C.Luecke, B.Reincke, B.Ludwig, H.Rueterjans 2000 Eur. J. Biochem. 267 4205 10866825
1cc5 Crystal structure of Azotobacter cytochrome c5 at 2.5 A resolution. D.C.Carter, K.A.Melis, S.E.O'Donnell, B.K.Burgess, W.R.Furey Jr., B.C.Wang, C.D.Stout 1985 J Mol Biol 184 279 2993632
1cch Investigation of the solution conformation of cytochrome c-551 from Pseudomonas stutzeri. M.Cai, E.G.Bradford, R.Timkovich 1992 Biochemistry 31 8603 1327105
1ced The solution structure of cytochrome c(6) from the green alga Monoraphidium braunii. L.Banci, I.Bertini, G.Quacquarini, O.Walter, A.Diaz, M.Hervas, M.A.delaRosa 1996 J Biol Inorg Chem 1 330 n/a
1cgn Three-dimensional structure of cytochrome c' from two Alcaligenes species and the implications for four-helix bundle structures. A.J.Dobbs, B.F.Anderson, H.R.Faber, E.N.Baker 1996 Acta Crystallogr., Sect.D 52 356 15299707
1cgo
1chh Structural studies of the roles of residues 82 and 85 at the interactive face of cytochrome c. T.P.Lo, J.G.Guillemette, G.V.Louie, M.Smith, G.D.Brayer 1995 Biochemistry 34 163 7819192
1chi
1chj
1cie Structural and functional effects of multiple mutations at distal sites in cytochrome c. T.P.Lo, S.Komar-Panicucci, F.Sherman, G.McLendon, G.D.Brayer 1995 Biochemistry 34 5259 7711047
1cif
1cig
1cih
1cno MAD Structure of Psueudomonas Nautica Dimeric Cytochrome C552 Mimics the C4 Dihemic Cytochrome Domain Association K.Brown, D.Nurizzo, S.Besson, W.Sheppard, J.Moura, I.Moura, M.Tegoni, C.Cambillau 1999 J. Mol. Biol. 289 1017 10369779
1co6 Refined Crystal Structure of Ferrocytochrome C2 from Rhodopseudomonas Viridis at 1.6 A Resolution S.Sogabe, K.Miki 1995 J.Mol.Biol. 252 235 7674304
1cot X-Ray structure of the cytochrome c2 isolated from Paracoccus denitrificans refined to 1.7-A resolution. M.M.Benning, T.E.Meyer, H.M.Holden 1994 Arch Biochem Biophys 310 460 8179333
1cpq High-resolution crystal structures of two polymorphs of cytochrome c' from the purple phototrophic bacterium rhodobacter capsulatus. T.H.Tahirov, S.Misaki, T.E.Meyer, M.A.Cusanovich, Y.Higuchi, N.Yasuoka 1996 J Mol Biol 259 467 8676382
1rcp
1cpr Structure of cytochrome c' from Rhodobacter capsulatus strain St Louis: an unusual molecular association induced by bridging Zn ions. T.H.Tahirov, S.Misaki, T.E.Meyer, M.A.Cusanovich, N.Yasuoka 1997 Acta Crystallogr., Sect.D 53 658 15299853
1crc The low ionic strength crystal structure of horse cytochrome c at 2.1 A resolution and comparison with its high ionic strength counterpart. R.Sanishvili, K.W.Volz, E.M.Westbrook, E.Margoliash 1995 Structure 3 707 8591047
1crg The role of a conserved internal water molecule and its associated hydrogen bond network in cytochrome c. A.M.Berghuis, J.G.Guillemette, G.McLendon, F.Sherman, M.Smith, G.D.Brayer 1994 J Mol Biol 236 786 8114094
1crh
1cri
1crj
1cry Application of an automatic molecular-replacement procedure to crystal structure analysis of cytochrome c2 from Rhodopseudomonas viridis. K.Miki, S.Sogabe, A.Uno, T.Ezoe, N.Kasai, M.Saeda, Y.Matsuura, M.Miki 1994 Acta Crystallogr., Sect.D 50 271 15299438
1csu Replacements in a conserved leucine cluster in the hydrophobic heme pocket of cytochrome c. T.P.Lo, M.E.Murphy, J.G.Guillemette, M.Smith, G.D.Brayer 1995 Protein Sci 4 198 7757009
1csv
1csw
1csx
1ctj Ab initio determination of the crystal structure of cytochrome c6 and comparison with plastocyanin. C.Frazao, C.M.Soares, M.A.Carrondo, E.Pohl, Z.Dauter, K.S.Wilson, M.Hervas, J.A.Navarro, M.A.De la Rosa, G.M.Sheldrick 1995 Structure 3 1159 8591027
1cxa Crystallization and X-ray structure determination of cytochrome c2 from Rhodobacter sphaeroides in three crystal forms. H.L.Axelrod, G.Feher, J.P.Allen, A.J.Chirino, M.W.Day, B.T.Hsu, D.C.Rees 1994 Acta Crystallogr., Sect.D 50 596 15299423
1cxc
2cxb
1cyi The structure of chloroplast cytochrome c6 at 1.9 A resolution: evidence for functional oligomerization. C.A.Kerfeld, H.P.Anwar, R.Interrante, S.Merchant, T.O.Yeates 1995 J Mol Biol 250 627 7623381
1cyj
1czj Crystal structure of a dimeric octaheme cytochrome c3 (M(r) 26,000) from Desulfovibrio desulfuricans Norway. M.Czjzek, F.Guerlesquin, M.Bruschi, R.Haser 1996 Structure 4 395 8740362
1h21 A novel iron centre in the split-Soret cytochrome c from Desulfovibrio desulfuricans ATCC 27774. I.A. Abreu, A.I. Lourenco, A.V. Xavier, J. Legall, A.V. Coelho, P.M. Matias, D.M. Pinto, M.A. Carrondo, M.Teixeira, L.M. Saraiva 2003 J. Biol. Inorg. Chem. 8 360 12589573
1dt1 Integrity of thermus thermophilus cytochrome c552 synthesized by Escherichia coli cells expressing the host-specific cytochrome c maturation genes, ccmABCDEFGH: biochemical, spectral, and structural characterization of the recombinant protein. J.A.Fee, Y.Chen, T.R.Todaro, K.L.Bren, K.M.Patel, M.G.Hill, E.Gomez-Moran, T.M.Loehr, J.Ai, L.Thony-Meyer, P.A.Williams, E.Stura, V.Sridhar, D.E.Mcree 2000 Protein Sci. 9 2074 11152119
1duw Three-dimensional structure of the nonaheme cytochrome c from Desulfovibrio desulfuricans Essex in the Fe(III) state at 1.89 Angstroms resolution. S.Umhau, G.Fritz, K.Diederichs, J.Breed, W.Welte, P.M.Kroneck 2001 Biochemistry 40 1308 11170457
1dvh Structure and dynamics of ferrocytochrome c553 from Desulfovibrio vulgaris studied by NMR spectroscopy and restrained molecular dynamics. M.J.Blackledge, S.Medvedeva, M.Poncin, F.Guerlesquin, M.Bruschi, D.Marion 1995 J Mol Biol 245 661 7844834
2dvh
1dvv Selected mutations in a mesophilic cytochrome c confer the stability of a thermophilic counterpart. J.Hasegawa, S.Uchiyama, Y.Tanimoto, M.Mizutani, Y.Kobayashi, Y.Sambongi, Y.Igarashi 2000 J.Biol.Chem. 275 37824 10918067
1dxr Structural basis of the drastically increased initial electron transfer rate in the reaction center from a Rhodopseudomonas viridis mutant described at 2.00-A resolution. C.R.D.Lancaster, M.Bibikova, P.Sabatino, D.Oesterhelt, H.Michel 2000 J.Biol.Chem. 275 39364 11005826
1e29 Crystal structure of low-potential cytochrome c549 from Synechocystis sp. PCC 6803 at 1.21 angstroms resolution. C.Frazao, F.J.Enguita, R.Coelho, G.M.Sheldrick, J.A.Navarro, M.Hervas, M.A.De La Rosa, M.A.Carrondo 2001 J. Biol. Inorg. Chem. 6 324 11315568
1e2r X-ray crystallographic study of cyanide binding provides insights into the structure-function relationship for cytochrome cd1 nitrite reductase from Paracoccus pantotrophus. A.Jafferji, J.W.A.Allen, S.J.Ferguson, V.Fulop 2000 J.Biol.Chem. 275 25089 10827177
1e39 Identification of the active site acid/base catalyst in a bacterial fumarate reductase: a kinetic and crystallographic study. M.K.Doherty, S.L.Pealing, C.S.Miles, R.Moysey, P.Taylor, M.D.Walkinshaw, G.A.Reid, S.K.Chapman 2000 Biochemistry 39 10695 10978153
1e83 Unprecedented proximal binding of nitric oxide to heme: implications for guanylate cyclase. D.M.Lawson, C.E.Stevenson, C.R.Andrew, R.R.Eady 2000 Embo J. 19 5661 11060017
1e84
1e85
1e86
1e8e Solution structure of Methylophilus methylotrophus cytochrome c": insights into the structural basis of haem-ligand detachment. L.Brennan, D.L.Turner, P.Fareleira, H.Santos 2001 J.Mol.Biol. 308 353 11327772
1ehj A proton-NMR investigation of the fully reduced cytochrome c7 from Desulfuromonas acetoxidans. Comparison between the reduced and the oxidized forms. M.Assfalg, L.Banci, I.Bertini, M.Bruschi, M.T.Giudici-Orticoni 1999 Eur. J. Biochem. 266 634 10561607
1f22
1eky De novo determination of protein structure by NMR using orientational and long-range order restraints. J.C.Hus, D.Marion, M.Blackledge 2000 J.Mol.Biol. 298 927 10801359
1etp Crystal structure of the dihaem cytochrome c4 from Pseudomonas stutzeri determined at 2.2A resolution. A.Kadziola, S.Larsen 1997 Structure 5 203 9032080
1f1c Structures of cytochrome c-549 and cytochrome c6 from the cyanobacterium Arthrospira maxima. M.R.Sawaya, D.W.Krogmann, A.Serag, K.K.Ho, T.O.Yeates, C.A.Kerfeld 2001 Biochemistry 40 9215 11478889
1f1f
1fhb Three-dimensional solution structure of the cyanide adduct of a Met80Ala variant of Saccharomyces cerevisiae iso-1-cytochrome c. Identification of ligand-residue interactions in the distal heme cavity. L.Banci, I.Bertini, K.L.Bren, H.B.Gray, P.Sompornpisut, P.Turano 1995 Biochemistry 34 11385 7547866
1fi3 Solution conformation of the Met 61 to His 61 mutant of Pseudomonas stutzeri ZoBell ferrocytochrome c-551. G.T.Miller, J.K.Hardman, R.Timkovich 2001 Biophys.J. 80 2928 11371465
1fi7 Effects of extrinsic imidazole ligation on the molecular and electronic structure of cytochrome c L.Banci, I.Bertini, G.Liu, J.Lu, T.Reddig, W.Tang, Y.Wu, D.Zhu 2001 J. Biol. Inorg. Chem. 6 628 11472026
1fi9
1fj0 Cleavage of the iron-methionine bond in c-type cytochromes: Crystal structure of oxidized and reduced cytochrome c(2) from Rhodopseudomonas palustris and its ammonia complex. S.Geremia, G.Garau, L.Vaccari, R.Sgarra, M.S.Viezzoli, M.Calligaris, L.Randaccio 2002 Protein Sci. 11 6 11742117
1fs7 Cytochrome c nitrite reductase from Wolinella succinogenes. Structure at 1.6 A resolution, inhibitor binding, and heme-packing motifs. O.Einsle, P.Stach, A.Messerschmidt, J.Simon, A.Kroger, R.Huber, P.M.H.Kroneck 2000 J.Biol.Chem. 275 39608 10984487
1fs8
1fs9
1ft5 High-resolution structures of the oxidized and reduced states of cytochrome c554 from Nitrosomonas europaea. T.M.Iverson, D.M.Arciero, A.B.Hooper, D.C.Rees 2001 J. Biol. Inorg. Chem. 6 390 11372197
1ft6
1gdv Structure of cytochrome c6 from the red alga Porphyra yezoensis at 1.57 Angstroms resolution. S.Yamada, S.-Y.Park, H.Shimizu, Y.Koshizuka, K.Kadokura, T.Satoh, K.Suruga, M.Ogawa, Y.Isogai, T.Nishio, Y.Shiro, T.Oku 2000 Acta Crystallogr., Sect.D 56 1577 11092924
1giw Solution structure of reduced horse heart cytochrome c. L.Banci, I.Bertini, J.G.Huber, G.A.Spyroulias, P.Turano 1999 J Biol Inorg Chem 4 21 10499099
1gks Ectothiorhodospira halophila ferrocytochrome c551: solution structure and comparison with bacterial cytochromes c. B.Bersch, M.J.Blackledge, T.E.Meyer, D.Marion 1996 J Mol Biol 264 567 8969306
1gm4 Conformational component in the coupled transfer of multiple electrons and protons in a monomeric tetraheme cytochrome. R.Louro, I.Bento, P.Matias, T.Catarino, A.Baptista, C.Soares, M.Carrondo, D.Turner, A.Xavier 2001 J.Biol.Chem. 276 44044 11551953
1gmb
1gu6 Structure and spectroscopy of the periplasmic cytochrome c nitrite reductase from Escherichia coli. V.A.Bamford, H.C.Angove, H.E.Seward, A.J.Thomson, J.A.Cole, J.N.Butt, A.M.Hemmings, D.J.Richardson 2002 Biochemistry 41 2921 11863430
1gyo Structure of dimeric cytochrome c3 from Desulfovibrio gigas at 1.2 Å resolution D.Aragao, C.Frazao, L.Sieker, G.M.Sheldrick, J.Legall, M.A.Carrondo 2003 Acta Crystallogr., Sect.D 59 644 12657783
1h9x The Structure of an alternative form of Paracoccus pantotrophus cytochrome cd(1) nitrite reductase. T.Sjogren, J.Hajdu 2001 J.Biol.Chem. 276 29450 11373294
1h9y
1hcm
1hh5 Structure of cytochrome c7 from Desulfuromonas acetoxidans at 1.9 angstroms resolution. M.Czjzek, P.Arnoux, R.Haser, W.Shepard 2001 Acta Crystallogr., Sect.D 57 670 11320307
1hh7 Crystallization and preliminary X-ray analysis of two pH-dependent forms of cytochrome c2 from Rhodopseudomonas palustris. G.Garau, S.Geremia, L.Randaccio, L.Vaccari, M.S.Viezzoli 2000 Acta Crystallogr., Sect.D 56 1699 11092951
1hrc High-resolution three-dimensional structure of horse heart cytochrome c. G.W.Bushnell, G.V.Louie, G.D.Brayer 1990 J Mol Biol 214 585 2166170
1hro Molecular structure of a high potential cytochrome c2 isolated from Rhodopila globiformis. M.M.Benning, T.E.Meyer, H.M.Holden 1996 Arch Biochem Biophys 333 338 8809072
1i54 Electron tunneling in protein crystals. F.A.Tezcan, B.R.Crane, J.R.Winkler, H.B.Gray 2001 Proc. Nat. Acad. Sci. USA 98 5002 11296248
1i55
1i5t Solution structure of cyanoferricytochrome c: ligand-controlled conformational flexibility and electronic structure of the heme moiety. Y.Yao, C.Qian, K.Ye, J.Wang, Z.Bai, W.Tang 2002 J. Biol. Inorg. Chem. 7 539 11941512
1i6d Solution structure and dynamics of the functional domain of Paracoccus denitrificans cytochrome c(552) in both redox states. B.Reincke, C.Perez, P.Pristovsek, C.Luecke, C.Ludwig, F.Loehr, V.V.Rogov, B.Ludwig, H.Rueterjans 2001 Biochemistry 40 12312 11591150
1i6e
1i77 Spectroscopic investigation and determination of reactivity and structure of the tetraheme cytochrome c3 from Desulfovibrio desulfuricans Essex 6. O.Einsle, S.Foerster, K.H.Mann, G.Fritz, A.Messerschmidt, P.M.H.Kroneck 2001 Eur. J. Biochem. 268 3028 11358521
1i8o Cleavage of the iron-methionine bond in c-type cytochromes: crystal structure of oxidized and reduced cytochrome c(2) from Rhodopseudomonas palustris and its ammonia complex. S.Geremia, G.Garau, L.Vaccari, R.Sgarra, M.S.Viezzoli, M.Calligaris, L.Randaccio 2002 Protein Sci. 11 6 11742117
1i8p
1io3 Crystal structure of the oxidized cytochrome c(2) from Blastochloris viridis. S.Sogabe, K.Miki 2001 FEBS Lett. 491 174 11240122
1irv Mechanistic and structural contributions of critical surface and internal residues to cytochrome c electron transfer reactivity. S.P.Rafferty, J.G.Guillemette, A.M.Berghuis, M.Smith, G.D.Brayer, A.G.Mauk 1996 Biochemistry 35 10784 8718869
1irw
1it1 Redox-coupled conformational alternations in cytochrome c(3) from D. vulgaris Miyazaki F on the basis of its reduced solution structure. E.Harada, Y.Fukuoka, T.Ohmura, A.Fukunishi, G.Kawai, T.Fujiwara, H.Akutsu 2002 J.Mol.Biol. 319 767 12054869
1jaf Crystal structure of cytochrome c' from Rhodocyclus gelatinosus and comparison with other cytochromes c'. M.Archer, L.Banci, E.Dikaya, M.J.Romao 1997 J Biol Inorg Chem 2 611 n/a
1jdl Structure of cytochrome c2 from Rhodospirillum centenum. A.Camara-Artigas, J.C.Williams, J.P.Allen 2001 Acta Crystallogr., Sect.D 57 1498 11679712
1jni The 1.25 Å resolution structure of the diheme NapB subunit of soluble nitrate reductase reveals a novel cytochrome c fold with a stacked heme arrangement. A.Brige, D.Leys, T.E.Meyer, M.A.Cusanovich, J.J.Van Beeumen 2002 Biochemistry 41 4827 11939777
1k3g NMR solution structure, backbone mobility, and homology modeling of c-type cytochromes from gram-positive bacteria. L.Banci, I.Bertini, S.Ciurli, A.Dikiy, J.Dittmer, A.Rosato, G.Sciara, A.R.Thompsett 2002 Chembiochem Europ. J. Chem. Biol. 3 299 11933230
1k3h
1kib Structure of Cytochrome C6 from Arthrospira Maxima: An Assembly of 24 Subunits in the Form of a Nearly Symmetric Shell C.A.Kerfeld, M.R.Sawaya, D.Krogmann, T.O.Yeates 2002 Acta Crystallogr., Sect.D 58 1104 n/a
1kt2 Structural Basis of Cytochrome C Presentation by Ie(K) D.H.Fremont, S.Dai, H.Chiang, F.Crawford, P.Marrack, J.Kappler 2002 J.Exp.Med. 195 1043 n/a
1ktd
1kwj A Quick Solution Structure Determination of the Fully Oxidized Double Mutant K9-10A Cytochrome C7 from Desulfuromonas Acetoxidans and Mechanistic Implications M.Assfalg, I.Bertini, P.Turano, M.Bruschi, M.C.Durand, M.T.Giudici-Orticoni, A.Dolla 2002 J.Biomol.NMR 22 107 n/a
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1kyo Crystal Structure of the Yeast Cytochrome Bc1 Complex with its Bound Substrate Cytochrome C C.Lange, C.Hunte 2002 Proc. Nat. Acad. Sci. USA 99 2800 n/a
1l9b X-Ray Structure Determination of the Cytochrome-C(2):Reaction Center Electron Transfer Complex from Rhodobacter Sphaeroides H.L.Axelrod, E.C.Abresch, M.Y.Okamura, A.P.Yeh, D.C.Rees, G.Feher 2002 J.Mol.Biol. 319 501 n/a
1l9j
1lfm Using Deeply Trapped Intermediates to Map the Cytochrome C Folding Landscape F.A.Tezcan, W.M.Findley, B.R.Crane, S.A.Ross, J.G.Lyubovitsky, H.B.Gray, J.R.Winkler 2002 Proc. Nat. Acad. Sci. USA 99 8626 n/a
1lm2 The Metal Reductase Activity of Some Multiheme Cytochromes C: NMR Structural Characterization of the Reduction of Chromium(Vi) to Chromium(III) by Cytochrome C7 M.Assfalg, I.Bertini, M.Bruschi, C.Michel, P.Turano 2002 Proc. Nat. Acad. Sci. USA 99 9750 n/a
1m1p Crystal Structures at Atomic Resolution Reveal the Novel Concept of 'Chi Electron Harvesting' as a Role for the Small Tetraheme Cytochrome C. D.Leys, T.E.Meyer, A.I.Tsapin, K.H.Nealson, M.A.Cusanovich, J.J.Van Beeumen n/a To be Published n/a n/a n/a
1m1q
1m1r
1mdv Key role of phenylalanine 20 in cytochrome c3: structure, stability, and function studies. A.Dolla, P.Arnoux, I.Protasevich, V.Lobachov, M.Brugna, M.T.Giudici-Orticoni, R.Haser, M.Czjzek, A.Makarov, M.Bruschi 1999 Biochemistry 38 33 9890880
1nbb Concerted movement of side chains in the haem vicinity observed on ligand binding in cytochrome c' from rhodobacter capsulatus. T.H.Tahirov, S.Misaki, T.E.Meyer, M.A.Cusanovich, Y.Higuchi, N.Yasuoka 1996 Nat Struct Biol 3 459 8612077
1new 800 MHz 1H NMR solution structure refinement of oxidized cytochrome c7 from Desulfuromonas acetoxidans. M.Assfalg, L.Banci, I.Bertini, M.Bruschi, P.Turano 1998 Eur J Biochem 256 261 9760163
1ocd Solution structure of horse heart ferricytochrome c and detection of redox-related structural changes by high-resolution 1H NMR. P.X.Qi, R.A.Beckman, A.J.Wand 1996 Biochemistry 35 12275 8823161
2frc
1qdb Structure of Cytochrome C Nitrite Reductase O.Einsle, A.Messerschmidt, P.Stach, G.P.Bourenkov, H.D.Bartunik, R.Huber, P.M.H.Kroneck 1999 Nature 400 476 n/a
1qjd Structural and Mechanistic Mapping of a Unique Fumarate Reductase P.Taylor, S.L.Pealing, G.A.Reid, S.K.Chapman, M.D.Walkinshaw 1999 Nat. Struct. Biol. 6 1108 n/a
1qks The Anatomy of a Bifunctional Enzyme: Structural Basis for Reduction of Oxygen to Water and Synthesys of Nitric Oxide by Cytochrome Cd1 V.Fulop, J.W.B.Moir, S.J.Ferguson, J.Hajdu 1995 Cell (Cambridge, Mass.) 81 369 n/a
1ql3 Structure of the Soluble Domain of Cytochrome C552 from Paracoccus Denitrificans in the Oxidized and Reduced States A.Harrenga, B.Reincke, H.Rueterjans, B.Ludwig, H.Michel 2000 J.Mol.Biol. 295 667 n/a
1ql4
1qn0 Solution Structure of Desulfovibrio Vulgaris (Hildenborough) Ferrocytochrome C3: Structural Basis for Functional Cooperativity A.C.Messias, M.L.Teodoro, L.Brennan, J.Legall, H.Santos, A.V.Xavier, D.L.Turner n/a n/a n/a n/a n/a
1qn1 Structural Basis for the Network of Functional Cooperativities in Cytochromes C3 from Desulfovibrio Gigas: Solution Structures of the Oxidised and Reduced States L.Brennan, D.L.Turner, A.C.Messias, M.L.Teodoro, J.Legall, H.Santos, A.V.Xavier 2000 J.Mol.Biol. 298 61 n/a
1qn2 The Molecular Structure of an Unusual Cytochrome C2 Determined at 2.0A; the Cytochrome Ch from Methylobacterium Extorquens J.Read, R.Gill, S.L.Dales, J.B.Cooper, S.P.Wood, C.Anthony 1999 Protein Sci. 8 1232 n/a
1rap The structure and function of omega loop A replacements in cytochrome c. M.E.Murphy, J.S.Fetrow, R.E.Burton, G.D.Brayer 1993 Protein Sci 2 1429 8401228
1raq
1wad Cytochrome c3 from Desulfovibrio gigas: crystal structure at 1.8 A resolution and evidence for a specific calcium-binding site. P.M.Matias, J.Morais, R.Coelho, M.A.Carrondo, K.Wilson, Z.Dauter, L.Sieker 1996 Protein Sci 5 1342 8819167
1wej Structural basis for the binding of an anti-cytochrome c antibody to its antigen: crystal structures of FabE8-cytochrome c complex to 1.8 A resolution and FabE8 to 2.26 A resolution. S.E.Mylvaganam, Y.Paterson, E.D.Getzoff 1998 J Mol Biol 281 301 9698550
1ycc High-resolution refinement of yeast iso-1-cytochrome c and comparisons with other eukaryotic cytochromes c. G.V.Louie, G.D.Brayer 1990 J Mol Biol 214 527 2166169
1yea Structure determination and analysis of yeast iso-2-cytochrome c and a composite mutant protein. M.E.Murphy, B.T.Nall, G.D.Brayer 1992 J Mol Biol 227 160 1326054
1yeb
1yic Solution structure of oxidized Saccharomyces cerevisiae iso-1- cytochrome c. L.Banci, I.Bertini, K.L.Bren, H.B.Gray, P.Sompornpisut, P.Turano 1997 Biochemistry 36 8992 9220987
1ytc Thermodynamic cycles as probes of structure in unfolded proteins. W.A.McGee, F.I.Rosell, J.R.Liggins, S.Rodriguez-Ghidarpour, Y.Luo, J.Chen, G.D.Brayer, A.G.Mauk, B.T.Nall 1996 Biochemistry 35 1995 8639684
2cth Refinement of the Three-Dimensional Structures of Cytochromes C3 from Desulfovibrio Vulgaris Hildenborough at 1.67 Angstrom Resolution and from Desulfovibrio Desulfuricans Atcc 27774 at 1.6 Angstrom Resolution P.Simoes, P.M.Matias, J.Morais, K.Wilson, Z.Dauter, M.A.Carrondo n/a To be Published n/a n/a n/a
2cy3 Crystal structure of cytochrome c3 from Desulfovibrio desulfuricans Norway at 1.7 A resolution. M.Czjzek, F.Payan, F.Guerlesquin, M.Bruschi, R.Haser 1994 J Mol Biol 243 653 7966289
2cym Effects of amino acid substitution on three-dimensional structure: an X-ray analysis of cytochrome c3 from Desulfovibrio vulgaris Hildenborough at 2 A resolution. Y.Morimoto, T.Tani, H.Okumura, Y.Higuchi, N.Yasuoka 1991 J Biochem (Tokyo) 110 532 1663945
2giw Solution Structure of Reduced Horse Heart Cytochrome C L.Banci, I.Bertini, J.G.Huber, G.A.Spyroulias, P.Turano n/a To be Published n/a n/a n/a
2new 800 Mhz 1H-NMR Solution Structure Refinement of Oxidized Cytochrome C7 from Desulfuromonas Acetoxidans M.Assfalg, L.Banci, I.Bertini, M.Bruschi, P.Turano n/a To be Published n/a n/a n/a
2pac Solution structure of Fe(II) cytochrome c551 from Pseudomonas aeruginosa as determined by two-dimensional 1H NMR. D.J.Detlefsen, V.Thanabal, V.L.Pecoraro, G.Wagner 1991 Biochemistry 30 9040 1654086
3cao Crystal Structure of the Oxidised and Reduced Acidic Cytochrome C3 from Desulfovibrio Africanus S.Norager, P.Legrand, L.Pieulle, C.Hatchikian, M.Roth 1999 J.Mol.Biol. 290 881 n/a
3car
3cyr Refinement of the Three-Dimensional Structures of Cytochrome C3 from Desulfovibrio Vulgaris Hildenborough at 1.67 Angstroms Resolution and from Desulfovibrio Desulfuricans Atcc 27774 at 1.6 Angstroms Resolution P.Simoes, P.M.Matias, J.Morais, K.Wilson, Z.Dauter, M.A.Carrondo n/a To be Published n/a n/a n/a
3cyt Redox conformation changes in refined tuna cytochrome c. T.Takano, R.E.Dickerson 1980 Proc. Nat. Acad. Sci. USA 77 6371 6256733

参考文献

当記事に関係する別の文献を以下に示します。

  • Richard E. Dickerson (1980): Cytochrome c and the evolution of energy metabolism. Scientific American 242 (3), pp. 136-153.
  • Christian Lange and Carola Hunte (2002): Crystal structure of the yeast cytochrome bc1 complex with its bound substrate cytochrome c. Proceedings of the National Academy of Science USA 99



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