ヘモグロビン

◆チューブワーム由来ヘモグロビンとヒトヘモグロビンの大きさ比べ
チューブワーム由来1YHU　　同Chain A　同PDBsum1yhu_HEM-OXY(A鎖) $
チューブワーム由来1YHU（左）とヒトヘモグロビン1HHO（4量体；右）
　　→ チューブワーム関連データのSITE例

◆サツマハオリムシ由来ヘモグロビン
チューブワームの一種サツマハオリムシ由来3WCT（24量体；A-H鎖×3）

◆マシコヒゲムシ由来ヘモグロビン
マシコヒゲムシ由来2D2N（24量体；A-D鎖×6）　同（4量体；A-D鎖）
PDBsum2d2n_MMC(メチル水銀，A鎖；A-D鎖で近傍アミノ酸に違い) $
　　→ SPring-8プレスリリース（2005/10/04）

◆アカムシユスリカ由来ヘモグロビン（ヘモグロビンV；Hb V）
アカムシユスリカ由来3ARK

◆シロイヌナズナ由来ヘモグロビン
シロイヌナズナ由来3ZHW　　同Chain A　同PDBsum3zhw_HEM(A鎖) $

◆ダイズ由来ヘモグロビン（レグヘモグロビン）
ダイズ由来1BINのChain A（レグヘモグロビン）　→ Jmolトピック

◆細菌Vitreoscilla stercoraria由来ヘモグロビン
Vitreoscilla stercoraria由来2VHB（アジ化物イオンが結合）　　同Chain A　同PDBsum2vhb_AZI-HEM(A鎖) $

【参考】
■“生物界で見られる4種類のテトラピロール”（岩波「科学」2013年7月号コラム，p.723図を参考に）
(a) クロロフィルa PDBデータ 3arcのChain A（光合成光化学系IIの部分構造）　同PDBsumデータ 3arc_CLA607s（多数のCLAの中から607番；CLA607以外のヘテロ分子は削除）　CLA（クロロフィルa）選択　［Chlorophyll a - Wikipedia，光合成に関係する生体分子PDB 3ARCの部分構造例（本サイト）］
(b) ヘム PDBデータ 2hhb（デオキシヘモグロビン，4量体）　同PDBsumデータ 2hhb_HEM（A鎖）$　HEM（ヘム）選択　［Heme - Wikipedia，ヘモグロビン（本サイト）］
(c) ビタミンB₁₂（シアノコバラミン） PDBデータ 1n2zのChain A（ビタミンE輸送タンパク質）　同PDBsumデータ 1n2z_CNC$　CNC（ビタミンB₁₂）選択　［Vitamin B₁₂ - Wikipedia，水溶性ビタミンと脂溶性ビタミン（本サイト）］
(d) F430 PDBデータ 3pot（メチル補酵素Mレダクターゼ）　同PDBsumデータ 3pot_F43（COM含む；06Cは削除）$　F43（F430）選択　［Coenzyme-B sulfoethylthiotransferase - Wikipedia，Cofactor F430 - Wikipedia］
※上記4種のテトラピロール同時表示 4種類のテトラピロール（左上から時計回りでクロロフィルa，ヘム，F430，ビタミンB₁₂）　CLA（クロロフィルa）選択　HEM（ヘム）選択　CNC（ビタミンB₁₂）選択　F43（F430）選択
※参考（テトラピロール関連分子）ポルフィリン

［左］ヒトヘモグロビン2HHBとチューブワーム由来ヘモグロビン1YHUのPyMOLによる重ね合わせ表示（何れもChain A）
［右］ヒトヘモグロビン2HHB（Chain A）のPDBsumデータ

［左］3Dプリンタによるデオキシヘモグロビン2HHB（4量体）の分子模型　［右］4量体模型と「鍵と鍵穴」模型
※新分子模型Kawakami Modelに関する資料はこちらもご覧ください。
★本分子模型を使った出張講座・土曜講座の受け付けをしています。

〔左〕ヘモグロビンS2hbs B鎖，〔右〕ヘモグロビン4hhb B鎖

オキシヘモグロビン1HHO A鎖

同4量体（A・B鎖×2）

同4量体のPyMOLによる表示例

ヒトのデオキシヘモグロビン2HHBの4量体；右端は必須・準必須・非必須アミノ酸区別着色

デオキシヘモグロビン2HHB Chain Bのヘム（HEM）近傍

デオキシヘモグロビン2HHBのPDBsumデータ（HEM）

デオキシヘモグロビン2HHBのHEM近傍His（〇印）

2HHB A鎖（ヘモグロビンα；赤字がFe近傍His）

001-025	Val	Leu	Ser	Pro	Ala	Asp	Lys	Thr	Asn	Val	Lys	Ala	Ala	Trp	Gly	Lys	Val	Gly	Ala	His	Ala	Gly	Glu	Tyr	Gly
026-050	Ala	Glu	Ala	Leu	Glu	Arg	Met	Phe	Leu	Ser	Phe	Pro	Thr	Thr	Lys	Thr	Tyr	Phe	Pro	His	Phe	Asp	Leu	Ser	His
051-075	Gly	Ser	Ala	Gln	Val	Lys	Gly	His	Gly	Lys	Lys	Val	Ala	Asp	Ala	Leu	Thr	Asn	Ala	Val	Ala	His	Val	Asp	Asp
076-100	Met	Pro	Asn	Ala	Leu	Ser	Ala	Leu	Ser	Asp	Leu	His	Ala	His	Lys	Leu	Arg	Val	Asp	Pro	Val	Asn	Phe	Lys	Leu
101-125	Leu	Ser	His	Cys	Leu	Leu	Val	Thr	Leu	Ala	Ala	His	Leu	Pro	Ala	Glu	Phe	Thr	Pro	Ala	Val	His	Ala	Ser	Leu
126-	Asp	Lys	Phe	Leu	Ala	Ser	Val	Ser	Thr	Val	Leu	Thr	Ser	Lys	Tyr	Arg

Ile = 0
Val = 13
Leu = 18
　BCAA = 31（22.0％）
Cys = 1
His = 10
Total = 141

2HHB B鎖（ヘモグロビンβ；赤字がFe近傍His）

001-025 Val His Leu Thr Pro Glu Glu Lys Ser Ala Val Thr Ala Leu Trp Gly Lys Val Asn Val Asp Glu Val Gly Gly

026-050 Glu Ala Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Glu Ser Phe Gly Asp Leu Ser Thr

051-075 Pro Asp Ala Val Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Gly Ala Phe Ser Asp Gly Leu

076-100 Ala His Leu Asp Asn Leu Lys Gly Thr Phe Ala Thr Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

101-125 Glu Asn Phe Arg Leu Leu Gly Asn Val Leu Val Cys Val Leu Ala His His Phe Gly Lys Glu Phe Thr Pro Pro

126- Val Gln Ala Ala Tyr Gln Lys Val Val Ala Gly Val Ala Asn Ala Leu Ala His Lys Tyr His

Ile = 0
Val = 18
Leu = 18
　BCAA = 36（24.7％）
Cys = 2
His = 9
Total = 146

4MQJ F鎖（胎児ヘモグロビンHbF，ヘモグロビンγ；赤字がFe近傍His）

001-025 Val His Phe Thr Glu Glu Asp Lys Ala Thr Ile Thr Ser Leu Trp Gly Lys Val Asn Val Glu Asp Ala Gly Gly

026-050 Glu Thr Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Asp Ser Phe Gly Asn Leu Ser Ser

051-075 Ala Ser Ala Ile Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Thr Ser Leu Gly Asp Ala Ile

076-100 Lys His Leu Asp Asp Leu Lys Gly Thr Phe Ala Gln Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

101-125 Glu Asn Phe Lys Leu Leu Gly Asn Val Leu Val Thr Val Leu Ala Ile His Phe Gly Lys Glu Phe Thr Pro Glu

126- Val Gln Ala Ser Trp Gln Lys Met Val Thr Gly Val Ala Ser Ala Leu Ser Ser Arg Tyr His

Ile = 4
Val = 14
Leu = 17
　BCAA = 35（24.0％）
Cys = 1
His = 7
Total = 146

［左］成人のヘモグロビンHbA（1HHO；α₂β₂）と胎児ヘモグロビンHbF（4MQJ；α₂γ₂）の比較
［右］胎児ヘモグロビンHbF（1FDH）　→ PDBj今月の分子 257: 胎児ヘモグロビン｜Jmolトピック

1HHO B鎖（ヘモグロビンβ，I/O値順着色）

Val

His

Leu

Thr

Pro

Glu

Lys

Ser

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Val

Thr

Ala

Leu

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Glu

Val

Gly

Glu

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Leu

Gly

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Leu

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Trp

Thr

Gln

Arg

Phe

Glu

Ser

Phe

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Leu

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Ala

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Gly

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Gly

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Gly

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Asn

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His

4MQJ F鎖（ヘモグロビンγ，I/O値順着色）

Val

His

Phe

Thr

Glu

Asp

Lys

Ala

Thr

Ile

Thr

Ser

Leu

Trp

Gly

Lys

Val

Asn

Val

Glu

Asp

Ala

Gly

Glu

Thr

Leu

Gly

Arg

Leu

Val

Tyr

Pro

Trp

Thr

Gln

Arg

Phe

Asp

Ser

Phe

Gly

Asn

Leu

Ser

Ala

Ser

Ala

Ile

Met

Gly

Asn

Pro

Lys

Val

Lys

Ala

His

Gly

Lys

Val

Leu

Thr

Ser

Leu

Gly

Asp

Ala

Ile

Lys

His

Leu

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Leu

Lys

Gly

Thr

Phe

Ala

Gln

Leu

Ser

Glu

Leu

His

Cys

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Lys

Leu

His

Val

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Pro

Glu

Asn

Phe

Lys

Leu

Gly

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Thr

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Leu

Ala

Ile

His

Phe

Gly

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Phe

Thr

Pro

Glu

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Gln

Ala

Ser

Trp

Gln

Lys

Met

Val

Thr

Gly

Val

Ala

Ser

Ala

Leu

Ser

Arg

Tyr

His

ヘム近傍PDBsumデータの比較（何れもO₂結合）

CO結合カルボモノオキシヘモグロビンのヘム近傍PDBsumデータの比較（1AJ9と4MQC，何れもA鎖）
※V62M変異の4MQCは自己免疫性溶血性貧血のEvans症候群に関係すると考えられる。

一酸化炭素が結合した Ni(II)-Fe(II)混成ヘモグロビン6KAEのChain A-DとそのPDBsumデータ（Chain A〈CO結合〉・B）　→ Jmolトピック
※ヘモグロビンによるガス分子運搬の直接観察に成功（横浜市立大学，2020/02/18）　…6KAE以外に6KA9・6KAH・6KAI・6KAO～6KAV・6L5V～6L5Y・6LCW・6LCX公開

ヒトヘモグロビン2HHBとマンモスヘモグロビン3VRFのPyMOLによる表示（何れもChain A）

ヘムのPDBsumデータの構成アミノ酸の違い（右はPDBsumデータ2hhb_HEM）

デオキシヘモグロビン3KMFのPDBsumデータ（A鎖HEM，PDBsumデータにH・D追加；中性子回折による座標)

参考：What is a Protein? - YouTube

参考：The flyer What is a Protein? (RCSB PDB) [PDF]
※数研出版「フォトサイエンス生物図録」，同「チャート式新化学」，講談社ブルーバックス「動く分子事典」p.256なども参照。

★水素原子は省略
消去　針金[2]　スティック[3]　球棒[4]　空間充填[5]
Jmol色[1]　原子球白色表示
｜｜
背景・黒　灰　白　紺

★ [ ] の番号順にボタンをクリックしてみましょう！
あとは以下をいろいろ押してみよう。
鍵と鍵穴区別表示1（1次構造；下の“鍵穴構成部品色別表示1”も押して）
鍵と鍵穴区別表示2（2次構造）
全選択　鍵穴を選択　鍵を選択
鍵穴構成部品色別表示1　2　3
鍵穴構造を維持する結合表示

タンパク質の1次構造～3次構造を知るために（4HHB B鎖を例に；4次構造は冒頭の“4量体”が例）。
→ 分子とは？から順にたどってみよう。

※注：本アニメgif中の2次構造画像にはα-ヘリックスのみでβ-シートは含まれていません。他のJmolコンテンツをご参照ください。

【参考】いろいろな生物のヘモグロビンのアミノ酸配列比較
ヘモグロビンのアミノ酸配列比較例（付記がないものは何れもA鎖）；ヒト（2hhb），ヒト胎児HbF（4mqj，B鎖；A鎖はヒトと同じ），マンモス（3vrf），ハツカネズミ（3hrw），ウズラ（3mjp），クロマグロ（1v4w），シロザメ（1gcw），ヤツメウナギ（2lhb），ヌタウナギ（1it3），チューブワーム（1yhu），マシコヒゲムシ（2d2n）　　→ その他参考：Glossoscolex paulistusの巨大ヘモグロビン

ヘモグロビン構造の比較 [1]（何れもA鎖）：ヒト（2HHB，デオキシヘモグロビン），マンモス（3VRF，CO結合），シロザメ（1GCW，CO結合），チューブワーム（1YHU）

同 [2]（何れもB鎖）：ヒト（4HHB，デオキシヘモグロビン），ワニ（8WIZ，重炭酸イオン〈HCO₃^-〉結合）

同 [3]（何れもA鎖）：ヒト（2HHB，デオキシヘモグロビン），ヤツメウナギ（2LHB，CN^-結合），ヌタウナギ（1IT3，CO結合）

同 [4]：ヒト（2HHB），ウマ（6R2O）

同 [5]：ヒト（2HHB）・ウマ（6R2O）・オオカミ（タテガミオオカミ，1FHJ）・イヌ（グレイハウンド，3PEL）；何れもA鎖
→ アミノ酸配列比較｜ Jmolトピック（明石書店「飼いならす」）

同 [6]：サツマハオリムシ（3WCT；24量体，Chain A-H × 3）
【参考（サツマハオリムシ研究例）】鹿児島湾の奇妙な生き物サツマハオリムシの全ゲノムを解読 ―化学合成生態系への適応メカニズムに迫る―（東京大学，2023/06/26） [NEW!]
※プレスリリース中の“Toll様受容体（TLR）”についてはToll様受容体参照

同 [7]：細菌Vitreoscilla stercoraria由来ヘモグロビン（2VHBのA鎖，アジ化物イオンが結合）とそのPDBsumデータ

I/O値順による着色
001-050

ヒト Val Leu Ser Pro Ala Asp Lys Thr Asn Val Lys Ala Ala Trp Gly Lys Val Gly Ala His Ala Gly Glu Tyr Gly Ala Glu Ala Leu Glu Arg Met Phe Leu Ser Phe Pro Thr Thr Lys Thr Tyr Phe Pro His Phe Asp Leu Ser His

ヒト　B鎖 Val His Leu Thr Pro Glu Glu Lys Ser Ala Val Thr Ala Leu Trp Gly Lys Val Asn Val Asp Glu Val Gly Gly Glu Ala Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Glu Ser Phe Gly Asp Leu Ser Thr

ヒト胎児HbF　B鎖 ** Val His Phe Thr Glu Glu Asp Lys Ala Thr Ile Thr Ser Leu Trp Gly Lys Val Asn Val Glu Asp Ala Gly Gly Glu Thr Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Asp Ser Phe Gly Asn Leu Ser Ser

マンモス Val Leu Ser Asp Asn Asp Lys Thr Asn Val Lys Ala Thr Trp Ser Lys Val Gly Asp His Ala Ser Asp Tyr Val Ala Glu Ala Leu Glu Arg Met Phe Phe Ser Phe Pro Thr Thr Lys Thr Tyr Phe Pro His Phe Asp Leu Ser His

ハツカネズミ Val Leu Ser Gly Glu Asp Lys Ser Asn Ile Lys Ala Ala Trp Gly Lys Ile Gly Gly His Gly Ala Glu Tyr Gly Ala Glu Ala Leu Glu Arg Met Phe Ala Ser Phe Pro Thr Thr Lys Thr Tyr Phe Pro His Phe Asp Val Ser His

ウズラ Val Leu Ser Ala Ala Asp Lys Thr Asn Val Lys Gly Ile Phe Ala Lys Ile Ala Gly His Ala Glu Glu Tyr Gly Ala Glu Ala Leu Asp Arg Met Phe Thr Thr Tyr Pro Gln Thr Lys Thr Tyr Phe Pro His Phe Asp Val Ser His

クロマグロ (X) Thr Thr Leu Ser Asp Lys Asp Lys Ser Thr Val Lys Ala Leu Trp Gly Lys Ile Ser Lys Ser Ala Asp Ala Ile Gly Ala Asp Ala Leu Gly Arg Met Leu Ala Val Tyr Pro Gln Thr Lys Thr Tyr Phe Ser His Trp Pro Asp

シロザメ Ala Phe Thr Ala Cys Glu Lys Gln Thr Ile Gly Lys Ile Ala Gln Val Leu Ala Lys Ser Pro Glu Ala Tyr Gly Ala Glu Cys Leu Ala Arg Leu Phe Val Thr His Pro Gly Ser Lys Ser Tyr Phe Glu Tyr Lys Asp Tyr Ser Ala

ヤツメウナギ Pro Ile Val Asp Thr Gly Ser Val Ala Pro Leu Ser Ala Ala Glu Lys Thr Lys Ile Arg Ser Ala Trp Ala Pro Val Tyr Ser Thr Tyr Glu Thr Ser Gly Val Asp Ile Leu Val Lys Phe Phe Thr Ser Thr Pro Ala Ala Gln Glu

ヌタウナギ Pro Ile Ile Asp Gln Gly Pro Leu Pro Thr Leu Thr Asp Gly Asp Lys Lys Ala Ile Asn Lys Ile Trp Pro Lys Ile Tyr Lys Glu Tyr Glu Gln Tyr Ser Leu Asn Ile Leu Leu Arg Phe Leu Lys Cys Phe Pro Gln Ala Gln Ala

チューブワーム Ala Cys Ala Met Leu Glu Arg Ala Lys Val Lys Asp Glu Trp Ala Lys Ala Tyr Gly Ile Gly Ala Ala Arg Ser Lys Phe Gly Asp Ala Leu Trp Arg Asn Val Phe Asn Tyr Ala Pro Asn Ala Arg Asp Ile Phe Glu Ser Val Asn

サツマハオリムシ Asp Cys Asn Ile Leu Gln Arg Leu Lys Val Lys Met Gln Trp Ala Lys Ala Tyr Gly Phe Gly Thr Glu Arg Ala Lys Phe Gly Asn Ser Leu Trp Thr Ser Ile Phe Asn Tyr Ala Pro Asp Ala Arg Asp Leu Phe Lys Ser Val Lys

マシコヒゲムシ Val Cys Asn Arg Leu Glu Gln Ile Leu Val Lys Thr Gln Trp Ala Gln Ser Tyr Gly Glu Ala Glu Asn Arg Ala Ala Phe Ser Arg Asp Leu Phe Ser Glu Leu Phe Asn Ile Gln Gly Ser Ser Arg Ala Leu Phe Ser Gly Val Gly

051-100

ヒト Gly Ser Ala Gln Val Lys Gly His Gly Lys Lys Val Ala Asp Ala Leu Thr Asn Ala Val Ala His Val Asp Asp Met Pro Asn Ala Leu Ser Ala Leu Ser Asp Leu His Ala His Lys Leu Arg Val Asp Pro Val Asn Phe Lys Leu

ヒト　B鎖 Pro Asp Ala Val Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Gly Ala Phe Ser Asp Gly Leu Ala His Leu Asp Asn Leu Lys Gly Thr Phe Ala Thr Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

ヒト胎児HbF　B鎖 ** Ala Ser Ala Ile Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Thr Ser Leu Gly Asp Ala Ile Lys His Leu Asp Asp Leu Lys Gly Thr Phe Ala Gln Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

マンモス Gly Ser Gly Gln Val Lys Gly His Gly Lys Lys Val Gly Glu Ala Leu Thr Gln Ala Val Gly His Leu Asp Asp Leu Pro Ser Ala Leu Ser Ala Leu Ser Asp Leu His Ala His Lys Leu Arg Val Asp Pro Val Asn Phe Lys Leu

ハツカネズミ Gly Ser Ala Gln Val Lys Gly His Gly Lys Lys Val Ala Asp Ala Leu Ala Ser Ala Ala Gly His Leu Asp Asp Leu Pro Gly Ala Leu Ser Ala Leu Ser Asp Leu His Ala His Lys Leu Arg Val Asp Pro Val Asn Phe Lys Leu

ウズラ Gly Ser Ala Gln Ile Lys Gly His Gly Lys Lys Val Ala Ala Ala Leu Val Glu Ala Ala Asn His Ile Asp Asp Ile Ala Gly Thr Leu Ser Lys Leu Ser Asp Leu His Ala Gln Lys Leu Arg Val Asp Pro Val Asn Phe Lys Leu

クロマグロ Met Ser Pro Gly Ser Gly Pro Val Lys Ala His Gly Lys Lys Val Met Gly Gly Val Ala Leu Ala Val Ser Lys Ile Asp Asp Leu Thr Thr Gly Leu Gly Asp Leu Ser Glu Leu His Ala Phe Lys Met Arg Val Asp Pro Ser Asn

シロザメ Ala Gly Ala Lys Val Gln Val His Gly Gly Lys Val Ile Arg Ala Val Val Lys Ala Ala Glu His Val Asp Asp Leu His Ser His Leu Glu Thr Leu Ala Leu Thr His Gly Lys Lys Leu Leu Val Asp Pro Gln Asn Phe Pro Met

ヤツメウナギ Phe Phe Pro Lys Phe Lys Gly Leu Thr Thr Ala Asp Glu Leu Lys Lys Ser Ala Asp Val Arg Trp His Ala Glu Arg Ile Ile Asn Ala Val Asp Asp Ala Val Ala Ser Met Asp Asp Thr Glu Lys Met Ser Met Lys Leu Arg Asn

ヌタウナギ Ser Phe Pro Lys Phe Ser Thr Lys Lys Ser Asn Leu Glu Gln Asp Pro Glu Val Lys His Gln Ala Val Val Ile Phe Asn Lys Val Asn Glu Ile Ile Asn Ser Met Asp Asn Gln Glu Glu Ile Ile Lys Ser Leu Lys Asp Leu Ser

チューブワーム Ser Lys Asp Met Ala Ser Pro Glu Phe Lys Ala His Ile Ala Arg Val Leu Gly Gly Leu Asp Arg Val Ile Ser Met Leu Asp Asn Gln Ala Thr Leu Asp Ala Asp Leu Ala His Leu Lys Ser Gln His Asp Pro Arg Thr Ile Asp

サツマハオリムシ Ser Glu Asp Met Arg Ser Pro Gln Phe Lys Ala His Ile Ala Arg Val Ile Gly Gly Leu Asp Arg Val Ile Ser Met Phe Asp Asn Glu Asp Ala Leu Asn Ala Asp Leu Glu His Leu Lys Ser Gln His Asp Pro Arg Gly Leu Asp

マシコヒゲムシ Val Asp Asp Met Asn Ser Ala Ala Phe Thr Ala His Cys Leu Arg Val Thr Gly Ala Leu Asn Arg Leu Ile Ser Gln Leu Asp Gln Gln Ala Thr Ile Asn Ala Asp Leu Ala His Leu Ala Gly Gln His Ala Ser Arg Asn Leu Asp

101-

ヒト Leu Ser His Cys Leu Leu Val Thr Leu Ala Ala His Leu Pro Ala Glu Phe Thr Pro Ala Val His Ala Ser Leu Asp Lys Phe Leu Ala Ser Val Ser Thr Val Leu Thr Ser Lys Tyr Arg

ヒト　B鎖 Glu Asn Phe Arg Leu Leu Gly Asn Val Leu Val Cys Val Leu Ala His His Phe Gly Lys Glu Phe Thr Pro Pro Val Gln Ala Ala Tyr Gln Lys Val Val Ala Gly Val Ala Asn Ala Leu Ala His Lys Tyr His

ヒト胎児HbF　B鎖 ** Glu Asn Phe Lys Leu Leu Gly Asn Val Leu Val Thr Val Leu Ala Ile His Phe Gly Lys Glu Phe Thr Pro Glu Val Gln Ala Ser Trp Gln Lys Met Val Thr Gly Val Ala Ser Ala Leu Ser Ser Arg Tyr His

マンモス Leu Ser His Cys Leu Leu Val Thr Leu Ser Ser His Gln Pro Thr Glu Phe Thr Pro Glu Val His Ala Ser Leu Asp Lys Phe Leu Ser Asn Val Ser Thr Val Leu Thr Ser Lys Tyr Arg

ハツカネズミ Leu Ser His Cys Leu Leu Val Thr Leu Ala Ser His His Pro Ala Asp Phe Thr Pro Ala Val His Ala Ser Leu Asp Lys Phe Leu Ala Ser Val Ser Thr Val Leu Thr Ser Lys Tyr Arg

ウズラ Leu Gly Gln Cys Phe Leu Val Val Val Ala Ile His His Pro Ala Ala Leu Thr Pro Glu Val His Ala Ser Leu Asp Lys Phe Leu Cys Ala Val Gly Thr Val Leu Thr Ala Lys Tyr Arg

クロマグロ Phe Lys Ile Leu Ser His Cys Ile Leu Val Val Val Ala Lys Met Phe Pro Lys Glu Phe Thr Pro Asp Ala His Val Ser Leu Asp Lys Phe Leu Ala Ser Val Ala Leu Ala Leu Ala Glu Arg Tyr Arg

シロザメ Leu Ser Glu Cys Ile Ile Val Thr Leu Ala Thr His Leu Thr Glu Phe Ser Pro Asp Thr His Cys Ala Val Asp Lys Leu Leu Ser Ala Ile Cys Gln Glu Leu Ser Ser Arg Tyr Arg

ヤツメウナギ Leu Ser Gly Lys His Ala Lys Ser Phe Gln Val Asp Pro Glu Tyr Phe Lys Val Leu Ala Ala Val Ile Ala Asp Thr Val Ala Ala Gly Asp Ala Gly Phe Glu Lys Leu Met Ser Met Ile Cys Ile Leu Leu Arg Ser Ala Tyr

ヌタウナギ Gln Lys His Lys Thr Val Phe Lys Val Asp Ser Ile Trp Phe Lys Glu Leu Ser Ser Ile Phe Val Ser Thr Ile Asp Gly Gly Ala Glu Phe Glu Lys Leu Phe Ser Ile Ile Cys Ile Leu Leu Arg Ser Ala Tyr

チューブワーム Pro Val Asn Phe Val Val Phe Arg Lys Ala Leu Ile Ala Thr Val Ala Gly Thr Phe Gly Val Cys Phe Asp Val Pro Ala Trp Gln Gly Cys Tyr Asn Ile Ile Ala Lys Gly Ile Thr Gly Ser Asp Ala Ala

サツマハオリムシ Ala Leu Asn Phe Val Val Phe Gly Lys Ala Leu Phe Ala Thr Val Gly Gly Gln Phe Gly Val Cys Phe Asp Leu Pro Ala Trp Glu Ser Cys Tyr Lys Val Ile Ala Met Gly Ile Thr Gly Asn Asp Met Phe Ser

マシコヒゲムシ Ala Ser Asn Phe Ala Ala Met Gly Gln Ala Val Met Ser Val Val Pro Thr His Leu Asp Cys Phe Asn Gln His Ala Trp Gly Glu Cys Tyr Glu Arg Ile Ala Ser Gly Ile Ser Gly

** A鎖はヒトと同配列。

ヒトの必須・準必須・非必須による着色
001-050

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Val

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** A鎖はヒトと同配列。
※各表は１文字アミノ酸配列から各種HTMLデータを作成する秀丸マクロにより作成。

<参考

ヒト（2hhb）・ウマ（6r2o）・オオカミ（タテガミオオカミ，1fhj）・イヌ（グレイハウンド，3pel）の比較；何れもA鎖　→ Jmolトピック（明石書店「飼いならす」）

1-50	ヒト（2HHB）	Val	Leu	Ser	Pro	Ala	Asp	Lys	Thr	Asn	Val	Lys	Ala	Ala	Trp	Gly	Lys	Val	Gly	Ala	His	Ala	Gly	Glu	Tyr	Gly	Ala	Glu	Ala	Leu	Glu	Arg	Met	Phe	Leu	Ser	Phe	Pro	Thr	Thr	Lys	Thr	Tyr	Phe	Pro	His	Phe	Asp	Leu	Ser	His
	ウマ（6R2O）	Val	Leu	Ser	Ala	Ala	Asp	Lys	Thr	Asn	Val	Lys	Ala	Ala	Trp	Ser	Lys	Val	Gly	Gly	His	Ala	Gly	Glu	Tyr	Gly	Ala	Glu	Ala	Leu	Glu	Arg	Met	Phe	Leu	Gly	Phe	Pro	Thr	Thr	Lys	Thr	Tyr	Phe	Pro	His	Phe	Asp	Leu	Ser	His
	オオカミ（1FHJ）	Val	Leu	Ser	Pro	Ala	Asp	Lys	Thr	Asn	Ile	Lys	Ser	Thr	Trp	Asp	Lys	Ile	Gly	Gly	His	Ala	Gly	Asp	Tyr	Gly	Gly	Glu	Ala	Leu	Asp	Arg	Thr	Phe	Gln	Ser	Phe	Pro	Thr	Thr	Lys	Thr	Tyr	Phe	Pro	His	Phe	Asp	Leu	Ser	Pro
	イヌ（3PEL）	Val	Leu	Ser	Pro	Ala	Asp	Lys	Thr	Asn	Ile	Lys	Ser	Thr	Trp	Asp	Lys	Ile	Gly	Gly	His	Ala	Gly	Asp	Tyr	Gly	Gly	Glu	Ala	Leu	Asp	Arg	Thr	Phe	Gln	Ser	Phe	Pro	Thr	Thr	Lys	Thr	Tyr	Phe	Pro	His	Phe	Asp	Leu	Ser	Pro
50-100	ヒト（2HHB）	Gly	Ser	Ala	Gln	Val	Lys	Gly	His	Gly	Lys	Lys	Val	Ala	Asp	Ala	Leu	Thr	Asn	Ala	Val	Ala	His	Val	Asp	Asp	Met	Pro	Asn	Ala	Leu	Ser	Ala	Leu	Ser	Asp	Leu	His	Ala	His	Lys	Leu	Arg	Val	Asp	Pro	Val	Asn	Phe	Lys	Leu
	ウマ（6R2O）	Gly	Ser	Ala	Gln	Val	Lys	Ala	His	Gly	Lys	Lys	Val	Gly	Asp	Ala	Leu	Thr	Leu	Ala	Val	Gly	His	Leu	Asp	Asp	Leu	Pro	Gly	Ala	Leu	Ser	Asn	Leu	Ser	Asp	Leu	His	Ala	His	Lys	Leu	Arg	Val	Asp	Pro	Val	Asn	Phe	Lys	Leu
	オオカミ（1FHJ）	Gly	Ser	Ala	Gln	Val	Lys	Ala	His	Gly	Lys	Lys	Val	Ala	Asp	Ala	Leu	Thr	Thr	Ala	Val	Ala	His	Leu	Asp	Asp	Leu	Pro	Gly	Ala	Leu	Ser	Ala	Leu	Ser	Asp	Leu	His	Ala	Tyr	Lys	Leu	Arg	Val	Asp	Pro	Val	Asn	Phe	Lys	Leu
	イヌ（3PEL）	Gly	Ser	Ala	Gln	Val	Lys	Ala	His	Gly	Lys	Lys	Val	Ala	Asp	Ala	Leu	Thr	Thr	Ala	Val	Ala	His	Leu	Asp	Asp	Leu	Pro	Gly	Ala	Leu	Ser	Ala	Leu	Ser	Asp	Leu	His	Ala	Tyr	Lys	Leu	Arg	Val	Asp	Pro	Val	Asn	Phe	Lys	Leu
100-	ヒト（2HHB）	Leu	Ser	His	Cys	Leu	Leu	Val	Thr	Leu	Ala	Ala	His	Leu	Pro	Ala	Glu	Phe	Thr	Pro	Ala	Val	His	Ala	Ser	Leu	Asp	Lys	Phe	Leu	Ala	Ser	Val	Ser	Thr	Val	Leu	Thr	Ser	Lys	Tyr	Arg
	ウマ（6R2O）	Leu	Ser	His	Cys	Leu	Leu	Ser	Thr	Leu	Ala	Val	His	Leu	Pro	Asn	Asp	Phe	Thr	Pro	Ala	Val	His	Ala	Ser	Leu	Asp	Lys	Phe	Leu	Ser	Ser	Val	Ser	Thr	Val	Leu	Thr	Ser	Lys	Tyr	Arg
	オオカミ（1FHJ）	Leu	Ser	His	Cys	Leu	Leu	Val	Thr	Leu	Ala	Cys	His	His	Pro	Thr	Glu	Phe	Thr	Pro	Ala	Val	His	Ala	Ser	Leu	Asp	Lys	Phe	Phe	Thr	Ala	Val	Ser	Thr	Val	Leu	Thr	Ser	Lys	Tyr	Arg
	イヌ（3PEL）	Leu	Ser	His	Cys	Leu	Leu	Val	Thr	Leu	Ala	Cys	His	His	Pro	Thr	Glu	Phe	Thr	Pro	Ala	Val	His	Ala	Ser	Leu	Asp	Lys	Phe	Phe	Ala	Ala	Val	Ser	Thr	Val	Leu	Thr	Ser	Lys	Tyr	Arg

シロイヌナズナ（3zhw A鎖）

Met

Glu

Ser

Glu

Gly

Lys

Ile

Val

Phe

Thr

Glu

Gln

Glu

Ala

Leu

Val

Lys

Ser

Trp

Ser

Val

Met

Lys

Asn

Ser

Ala

Glu

Leu

Gly

Leu

Lys

Leu

Phe

Ile

Lys

Ile

Phe

Glu

Ile

Ala

Pro

Thr

Lys

Met

Phe

Ser

Phe

Leu

Arg

Asp

Ser

Pro

Ile

Pro

Ala

Glu

Gln

Asn

Pro

Lys

Leu

Lys

Pro

His

Ala

Met

Ser

Val

Phe

Val

Met

Cys

Glu

Ser

Ala

Val

Gln

Leu

Arg

Lys

Thr

Gly

Lys

Val

Thr

Val

Arg

Glu

Thr

Leu

Lys

Arg

Leu

Gly

Ala

Ser

His

Ser

Lys

Tyr

Gly

Val

Asp

Glu

His

Phe

Glu

Val

Ala

Lys

Tyr

Ala

Leu

Glu

Thr

Ile

Lys

Glu

Ala

Val

Pro

Glu

Met

Trp

Ser

Pro

Glu

Met

Lys

Val

Ala

Trp

Gly

Gln

Ala

Tyr

Asp

His

Leu

Val

Ala

Ile

Lys

Ala

Glu

Met

Asn

Leu

Ser

Asn

アカムシユスリカ（3ark A鎖〈単量体〉）　…ヘモグロビンV（Hb V）　→ Jmolトピック

Ala

Phe

Val

Gly

Leu

Ser

Asp

Ser

Glu

Lys

Leu

Val

Arg

Asp

Ala

Trp

Ala

Pro

Ile

His

Gly

Asp

Leu

Gln

Gly

Thr

Ala

Asn

Thr

Val

Phe

Tyr

Asn

Tyr

Leu

Lys

Tyr

Pro

Ser

Asn

Gln

Asp

Lys

Phe

Glu

Thr

Leu

Lys

Gly

His

Pro

Leu

Asp

Glu

Val

Lys

Asp

Thr

Ala

Asn

Phe

Lys

Leu

Ile

Ala

Gly

Arg

Ile

Phe

Thr

Ile

Phe

Asp

Asn

Cys

Val

Lys

Asn

Val

Gly

Asn

Asp

Lys

Gly

Phe

Gln

Lys

Val

Ile

Ala

Asp

Met

Ser

Gly

Pro

His

Val

Ala

Arg

Pro

Ile

Thr

His

Gly

Ser

Tyr

Asn

Asp

Leu

Arg

Gly

Val

Ile

Tyr

Asp

Ser

Met

His

Leu

Asp

Ser

Thr

His

Gly

Ala

Trp

Asn

Lys

Met

Asp

Asn

Phe

Tyr

Val

Phe

Tyr

Glu

Cys

Leu

Asp

Gly

Arg

Cys

Ser

Gln

Phe

Ser

【参考】ヘモグロビンとヘモグロビンS
4hhbと2hbsのアミノ酸配列の違い（FASTA Sequence，何れもB鎖について）
※赤字が最初の共通開始部分トップのMet

VHLTPVEKSAVTALWGKVNVDEVGGEALGRLLVVYPWTQRFFESFGDLST PDAVMGNPKVKAHGKKVLGAFSDGLAHLDNLKGTFATLSELHCDKLHVDP ENFRLLGNVLVCVLAHHFGKEFTPPVQAAYQKVVAGVANALAHKYH

※以下にこの配列について，１文字アミノ酸配列から各種HTMLデータを作成する秀丸マクロにより処理したデータを示す。

4HHB（アミノ酸別着色）

Val His Leu Thr Pro Glu Glu Lys Ser Ala Val Thr Ala Leu Trp Gly Lys Val Asn Val Asp Glu Val Gly Gly Glu Ala Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Glu Ser Phe Gly Asp Leu Ser Thr

Pro Asp Ala Val Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Gly Ala Phe Ser Asp Gly Leu Ala His Leu Asp Asn Leu Lys Gly Thr Phe Ala Thr Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

Glu Asn Phe Arg Leu Leu Gly Asn Val Leu Val Cys Val Leu Ala His His Phe Gly Lys Glu Phe Thr Pro Pro Val Gln Ala Ala Tyr Gln Lys Val Val Ala Gly Val Ala Asn Ala Leu Ala His Lys Tyr His

2HBS（アミノ酸別着色）

Val

His

Leu

Thr

Pro

Val

Glu

Lys

Ser

Ala

Val

Thr

Ala

Leu

Trp

Gly

Lys

Val

Asn

Val

Asp

Glu

Val

Gly

Glu

Ala

Leu

Gly

Arg

Leu

Val

Tyr

Pro

Trp

Thr

Gln

Arg

Phe

Glu

Ser

Phe

Gly

Asp

Leu

Ser

Thr

Pro

Asp

Ala

Val

Met

Gly

Asn

Pro

Lys

Val

Lys

Ala

His

Gly

Lys

Val

Leu

Gly

Ala

Phe

Ser

Asp

Gly

Leu

Ala

His

Leu

Asp

Asn

Leu

Lys

Gly

Thr

Phe

Ala

Thr

Leu

Ser

Glu

Leu

His

Cys

Asp

Lys

Leu

His

Val

Asp

Pro

Glu

Asn

Phe

Arg

Leu

Gly

Asn

Val

Leu

Val

Cys

Val

Leu

Ala

His

Phe

Gly

Lys

Glu

Phe

Thr

Pro

Val

Gln

Ala

Tyr

Gln

Lys

Val

Ala

Gly

Val

Ala

Asn

Ala

Leu

Ala

His

Lys

Tyr

His

4HHB（極性・非極性別）

Val

His

Leu

Thr

Pro

Glu

Lys

Ser

Ala

Val

Thr

Ala

Leu

Trp

Gly

Lys

Val

Asn

Val

Asp

Glu

Val

Gly

Glu

Ala

Leu

Gly

Arg

Leu

Val

Tyr

Pro

Trp

Thr

Gln

Arg

Phe

Glu

Ser

Phe

Gly

Asp

Leu

Ser

Thr

Pro

Asp

Ala

Val

Met

Gly

Asn

Pro

Lys

Val

Lys

Ala

His

Gly

Lys

Val

Leu

Gly

Ala

Phe

Ser

Asp

Gly

Leu

Ala

His

Leu

Asp

Asn

Leu

Lys

Gly

Thr

Phe

Ala

Thr

Leu

Ser

Glu

Leu

His

Cys

Asp

Lys

Leu

His

Val

Asp

Pro

Glu

Asn

Phe

Arg

Leu

Gly

Asn

Val

Leu

Val

Cys

Val

Leu

Ala

His

Phe

Gly

Lys

Glu

Phe

Thr

Pro

Val

Gln

Ala

Tyr

Gln

Lys

Val

Ala

Gly

Val

Ala

Asn

Ala

Leu

Ala

His

Lys

Tyr

His

2HBS（極性・非極性別）

Val

His

Leu

Thr

Pro

Val

Glu

Lys

Ser

Ala

Val

Thr

Ala

Leu

Trp

Gly

Lys

Val

Asn

Val

Asp

Glu

Val

Gly

Glu

Ala

Leu

Gly

Arg

Leu

Val

Tyr

Pro

Trp

Thr

Gln

Arg

Phe

Glu

Ser

Phe

Gly

Asp

Leu

Ser

Thr

Pro

Asp

Ala

Val

Met

Gly

Asn

Pro

Lys

Val

Lys

Ala

His

Gly

Lys

Val

Leu

Gly

Ala

Phe

Ser

Asp

Gly

Leu

Ala

His

Leu

Asp

Asn

Leu

Lys

Gly

Thr

Phe

Ala

Thr

Leu

Ser

Glu

Leu

His

Cys

Asp

Lys

Leu

His

Val

Asp

Pro

Glu

Asn

Phe

Arg

Leu

Gly

Asn

Val

Leu

Val

Cys

Val

Leu

Ala

His

Phe

Gly

Lys

Glu

Phe

Thr

Pro

Val

Gln

Ala

Tyr

Gln

Lys

Val

Ala

Gly

Val

Ala

Asn

Ala

Leu

Ala

His

Lys

Tyr

His

4HHB（I/O値順）

Val His Leu Thr Pro Glu Glu Lys Ser Ala Val Thr Ala Leu Trp Gly Lys Val Asn Val Asp Glu Val Gly Gly Glu Ala Leu Gly Arg Leu Leu Val Val Tyr Pro Trp Thr Gln Arg Phe Phe Glu Ser Phe Gly Asp Leu Ser Thr

Pro Asp Ala Val Met Gly Asn Pro Lys Val Lys Ala His Gly Lys Lys Val Leu Gly Ala Phe Ser Asp Gly Leu Ala His Leu Asp Asn Leu Lys Gly Thr Phe Ala Thr Leu Ser Glu Leu His Cys Asp Lys Leu His Val Asp Pro

Glu Asn Phe Arg Leu Leu Gly Asn Val Leu Val Cys Val Leu Ala His His Phe Gly Lys Glu Phe Thr Pro Pro Val Gln Ala Ala Tyr Gln Lys Val Val Ala Gly Val Ala Asn Ala Leu Ala His Lys Tyr His

2HBS（I/O値順）

Val

His

Leu

Thr

Pro

Val

Glu

Lys

Ser

Ala

Val

Thr

Ala

Leu

Trp

Gly

Lys

Val

Asn

Val

Asp

Glu

Val

Gly

Glu

Ala

Leu

Gly

Arg

Leu

Val

Tyr

Pro

Trp

Thr

Gln

Arg

Phe

Glu

Ser

Phe

Gly

Asp

Leu

Ser

Thr

Pro

Asp

Ala

Val

Met

Gly

Asn

Pro

Lys

Val

Lys

Ala

His

Gly

Lys

Val

Leu

Gly

Ala

Phe

Ser

Asp

Gly

Leu

Ala

His

Leu

Asp

Asn

Leu

Lys

Gly

Thr

Phe

Ala

Thr

Leu

Ser

Glu

Leu

His

Cys

Asp

Lys

Leu

His

Val

Asp

Pro

Glu

Asn

Phe

Arg

Leu

Gly

Asn

Val

Leu

Val

Cys

Val

Leu

Ala

His

Phe

Gly

Lys

Glu

Phe

Thr

Pro

Val

Gln

Ala

Tyr

Gln

Lys

Val

Ala

Gly

Val

Ala

Asn

Ala

Leu

Ala

His

Lys

Tyr

His

《話題》オキシヘモグロビン・デオキシヘモグロビンとfMRI
　fMRI（機能的磁気共鳴画像）は脳の活動を調べる神経活動計測法の一つであるが，その測定原理となっているのが，オキシヘモグロビンとデオキシヘモグロビンの磁性体としての性質の差である。
　デオキシヘモグロビンの方が常磁性体としての性質をもつので，MRIの強い磁場の中において磁場を乱してMRI信号を減弱させる（Blood oxygen level dependent効果，BOLD効果）。ところが脳の活動が活発化して，網細血管にオキシヘモグロビンが流入するとMRI信号を弱めていたデオキシヘモグロビンが押し出されて信号強度が回復する。この現象による局所的なシグナルの強弱を測定して脳の活動状態を調べるのである。

参考

松岡香弥子，『fMRIで心のしくみを探る』，バイオニクス 2005年2月号

分子科学研究所編，「分子科学者がいどむ12の謎」／9章『生体中でのタンパク質の機能をさぐる』，化学同人(2005)　※ミオグロビン中のヘムの構造変化について詳細に解説

2003年日本国際賞，医学における視覚化技術分野（磁気共鳴機能画像法の基礎原理の発見）『小川誠二博士　脳機能を知る一つの方法』

矢野良治，『ブレインイメージング：神経疾患の診断・治療に向けた非侵襲技術応用研究の体制強化の必要性』

牧敦，『光で見たこどもの脳』，科学 2001年6月号（特集-1＝こどもの脳で何がおこっているのか），岩波書店　※fMRIではなく光トポグラフィ法

（タイムスリップ）偶然見えた脳の動き　fMRI開発、マウス酸欠きっかけ（朝日，2013/09/16）

生化学 : ヘモグロビンを安定化する（Nature，2005/06/02）

進化医学 - Wikipedia

参考文献・Webページ

C.K.Mathews ほか著，清水孝雄ほか監訳，「カラー生化学」，pp.164-174，西村書店(2003)

Bruce Alberts ほか著，中村桂子・松原謙一　監訳，「細胞の分子生物学第4版」，ニュートンプレス(2004)
桜井弘編，「生命元素事典」，オーム社(2006)　※p.80の，O₂の両O原子がFeにside-onで配位する構造（下図はその近傍）については，上掲1o7m閲覧（生体分子の構成元素も参照）
ヘモグロビン - 今月の分子（PDBj）｜ PDB Molecule of the Month - Hemoglobin　※Cooperation Makes It Easierにオキシヘモグロビン・デオキシヘモグロビンのアニメーション
今月の分子 206: グロビンの進化（Globin Evolution）（PDBj，2017/02）｜ Molecule of the Month（PDB）　→ Jmolトピック
今月の分子 257: 胎児ヘモグロビン（Fetal Hemoglobin）（PDBj，2021/05）｜ Molecule of the Month（PDB）
Chime Resources at UMass ｜ Protein Explorer FrontDoor ｜ HEMOGLOBIN
Chemistry 227 / Intro To Biochem (Wellesley College) ｜ HEMOGLOBIN
PPS 96' - Protein Interactions: Haemoglobin 2
Google検索結果：　“オキシヘモグロビン” ｜ “haemoglobin OR hemoglobin + Jmol” ∥ イメージ検索： “haemoglobin OR hemoglobin”
タンパク質の「深呼吸」（KEK，2009/02/12）
巨大ヘモグロビンの結晶構造の解明に成功（SPring-8，2005/10/04）　※PDB 2D2Nを上掲。
ヘモグロビンへの配位子結合過程を直接観測（理研，2003/06/09）
有鬚（ゆうしゅ）動物巨大ヘモグロビンの構造生物学（金沢大学動物・微生物生理化学研究室）
ヘモグロビン - Wikipedia ｜鎌状赤血球症｜進化医学

3D印刷技術を応用した新たな分子模型の作製方法を開発（JAIST，2012/08/06）
※同分子模型に関する資料はこちらをご覧ください。
本間善夫・川端潤，「パソコンで見る動く分子事典」，講談社ブルーバックス(2007)　※p.92“ヘム色素”，p.256“ヘモグロビン”など
野村祐次郎・辰巳敬・本間善夫，「チャート式シリーズ　新化学」，p.408，数研出版(2009)
「視覚でとらえるフォトサイエンス化学図録」，p.202，数研出版(2012)
「ダイナミックワイド図説化学」，p.175，東京書籍(2011)
「サイエンスビュー化学総合資料」，p.272，実教出版(2012)
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