JP2009213481A - ペントース糖の発酵 - Google Patents
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Abstract
【解決手段】発現したときにキシロース異性化酵素をコードする配列は、キシロースをキシルロースに変換する能力を宿主細胞に付与し、それは宿主細胞により更に代謝される。従って、宿主細胞は、炭素源としてのキシロースに対して増殖することができる。宿主細胞は酵母または糸状菌のような真核性微生物であることが望ましい。さらにエタノールのような発酵産物の生産プロセスに関し、そのプロセスにおいて、宿主細胞は増殖及び発酵産物の生産のためにキシロースを利用する。さらに嫌気性菌から得られる真核生物性キシロース異性化酵素及びキシルロースキナーゼをコードする核酸配列。
【選択図】なし
Description
(定義)
キシロース異性化酵素
本明細書において、酵素「キシロース異性化酵素」(EC5.3.1.5)は、D−キシロースのD−キシルロースへのの直接の異性化、及びその逆の直接の異性化を触媒する酵素として定義される。この酵素はまた、D−キシロースケト異性化酵素としても知られている。一部のキシロース異性化酵素は、D−グルコース及びD−フルクトースの間の変換を触媒することもできるので、時にはグルコース異性化酵素とも呼ばれる。キシロース異性化酵素は補因子としてマグネシウムを必要とする。本発明のキシロース異性化酵素はさらに後述するアミノ酸配列によって定義される。同様に、キシロース異性化酵素はこの酵素をコードするヌクレオチド配列並びに後述されるキシロース異性化酵素をコードする対照ヌクレオチド配列にハイブリダイズするヌクレオチド配列によって定義することができる。
本明細書において配列同一性(sequence identity)とは、配列比較により決定される、二以上のアミノ酸(ポリペプチドまたはタンパク)配列または二以上の核酸(ポリヌクレオチド)配列の関係として定義される。この分野で、「同一性」は、場合によって、アミノ酸配列又は核酸配列の配列間の一致により決定される、アミノ酸または核酸配列間の配列関連性(sequence relatedness)の程度も意味している。二つのアミノ酸配列の「類似性(similarity)」は、あるポリペプチドの他のポリペプチドに対する、アミノ酸配列及び保存的置換アミノ酸を比較して決定される。「同一性」及び「類似性」は、限定はしないが以下に記述されている方法を含む既知方法により容易に計算することができる(Computational Molecular Biology, Lesk, A. M., ed., Oxford University Press, New York, 1988; Biocomputing:Informatics and Genome Projects, Smith, D. W., ed., Academic Press, New York, 1993; Computer Analysis of Sequence Data, Part I, Griffin, A. M., and Griffin, H. G., eds., Humana Press, New Jersey, 1994;Sequence Analysis in Molecular Biology, von Heine, g., Academic Press, 1987; and Sequence Analysis Primer, Gribskov, M. and Devereux, J., eds., M Stockton Press, New York, 1991; and Carillo, H., and Lipman, D., SIAM J. Applied Math., 48: 1073(1988)。
本発明のキシロース異性化酵素またはキシルロースキナーゼをコードする核酸配列は、中等度のまたは望ましくは厳密なハイブリッド形成条件下に、SEQ ID NO.2またはSEQ ID NO.4のヌクレオチド配列それぞれとハイブリッド形成する能力によって規定することもできる。厳密なハイブリッド形成条件とは、少なくとも約25、望ましくは約50ヌクレオチド、75または100及び最も望ましくは約200またはそれ以上のヌクレオチドの核酸配列を、約65℃の温度において、1Mの塩、望ましくは6xSSCまたは同等のイオン強度を持つその他の溶液中でハイブリッド形成させることができ、そして0.1Mの塩、またはそれ以下、望ましくは0.2xSSCまたは同等のイオン強度を持つその他の溶液中で65℃において洗浄する条件とここでは定義する。望ましくは、ハイブリッド形成は終夜、すなわち少なくとも10時間行い、望ましくは、洗浄は洗浄液を少なくとも2回交換して少なくとも1時間行う。この条件により、通常約90%またはそれ以上の配列同一性を持つ配列との特異的ハイブリッド形成が可能であろう。
本明細書で使用される「作動的に連結した」とは、機能的に関連するポリヌクレオチド配列の連結のことである。他の核酸配列との機能的な関連を持って配置されている場合に、核酸は「作動的に連結している」。例えば、プロモーターまたはエンハンサーは、それがコード配列の転写に影響するならば、コード配列と作動的に連結している。作動的に連結したとは、典型的には、連結したDNA配列同士が隣接しており、二個のタンパクコード領域を結合する必要がある場合には、連続し、読み枠内にあることを意味する。
本明細書で使用される「プロモーター」とは、転写の方向の点で遺伝子の転写開始部位の上流に存在する1またはそれ以上の遺伝子の転写を調節する機能を持つ核酸フラグメントであって、DNA依存RNAポリメラーゼ、転写開始部位、及び限定はしないが、転写因子結合部位、リプレッサー及び活性化タンパク結合部位を含むその他のDNA配列、並びにプロモーターの転写の量を直接的若しくは間接的に調節することが当業者には既知のその他のヌクレオチドの配列によって構造的に同定される。「構成的」プロモーターはほとんどの環境及び発生条件において活性であるプロモーターである。「誘導的」プロモーターは環境及び発生が調節されたときに活性となるプロモーターである。
本発明の最初の態様は、キシロースからキシルロースへ異性化する能力を持つ形質転換宿主細胞に関するものである。キシロースからキシルロースへ異性化する能力は、キシロース異性化酵素をコードする核酸配列を含む核酸構築物で宿主細胞を形質転換することにより付与される。キシロースからキシルロースへ異性化する形質転換宿主細胞の能力は、キシロースからキシルロースへの直接の異性化である。これは、それぞれキシロース還元酵素およびキシリトール脱水素酵素に触媒されてキシリトール中間体を経由してキシロースからキシルロースへ変換する2ステップ反応に対して、キシロース異性化酵素により触媒される1ステップ反応によりキシロースがキシルロースへ異性化したことを意味すると理解される。
(a)SEQ ID NO.1のアミノ酸配列と少なくとも40,45,49,50,53,55,60,70,80,90,95,97,98,または99%の配列同一性を有するアミノ酸配列を含むポリペプチドをコードするヌクレオチド配列;
(b)SEQ ID NO.2のヌクレオチド配列と少なくとも40,50,55,56,57,60,70,80,90,95,97,98,または99%の配列同一性を有するヌクレオチド配列を含むヌクレオチド配列;
(c)その相補鎖が、(a)または(b)の核酸分子配列にハイブリダイズするヌクレオチド配列;
(d)遺伝子コードの縮重による、(c)の核酸分子の配列と異なる配列からなるヌクレオチド配列:
からなる群から選択することができる。
(a)SEQ ID NO.1のアミノ酸配列と少なくとも50,53,54,55,60,70,80,90,95,97,98,または99%の配列同一性を有するアミノ酸配列を含むポリペプチドをコードする核酸分子;
(b)SEQ ID NO.2のヌクレオチド配列と少なくとも50,56,57,58,60,70,80,90,95,97,98,または99%の配列同一性を有するヌクレオチド配列を含む核酸分子;
(c)その相補鎖が、(a)または(b)の核酸分子配列にハイブリダイズする核酸分子;及び
(d)遺伝子コードの縮重により(c)の核酸分子配列と異なる配列からなる核酸分子;
からなる群から選択されることが望ましい。
(a)SEQ ID NO.3のアミノ酸配列と少なくとも45,47,48,49,50,55,60,70,80,90,95,97,98,または99%の配列同一性を有するアミノ酸配列を含むポリペプチドをコードする核酸分子;
(b)SEQ ID NO.4のヌクレオチド配列と少なくとも30,37,38,39,40,50,60,70,80,90,95,97,98,または99%の配列同一性を有するヌクレオチド配列からなる核酸分子;
(c)その相補鎖が、(a)または(b)の核酸分子配列にハイブリダイズする核酸分子;及び
(d)遺伝子コードの縮重による、(c)の核酸分子配列と異なる配列からなる核酸分子:
からなる群から選択されることが望ましい。
(生物及び増殖条件)
インド象の糞から単離した嫌気性菌Piromyces sp.E2(ATCC76762)を、N2/CO2(80%/20%)中、39℃で、種々の炭素源を添加したM2培地で嫌気的に増殖した(24)。使用した炭素源は、アビセル(微結晶セルロース、タイプPH105、Serva,ドイツ)、フルクトースまたはキシロース(全て0.5%、w/v)であった。増殖が止まった後(水素の発生により判断した)、細胞を遠心分離(15,000xg,4℃,15分間)またはナイロンガーゼ(30μm孔径)による濾過により回収した。
菌細胞を脱イオン水で洗い、培地成分を除去した。細胞を液体窒素中で凍結し、次いで、乳鉢中で、硝子ビーズ(0.10−0.11mm径)ですりつぶして、無細胞抽出物を調製した。Tris/HCl緩衝液(100mM,pH7.0)を粉末に加え(1:1,w/v)そして15分間解凍した後、懸濁液を遠心分離した(18,000xg,4℃,15分間)。透明な上清を細胞内酵素の原料として使用した。
キシロース異性化酵素活性は、50mMリン酸緩衝液(pH7.0),10mMキシロース,10mM MgCl2及び適当量の無細胞抽出物を含む反応混合物中37℃で検定した。生成したキシルロースの量は、システイン−カルバゾール法(9)により測定した。キシルロースキナーゼ及びキシロース還元酵素活性はWitteveen et al.(28)による記述にしたがって検定した。活性の1単位は、検定条件の下に、毎分1 nmolのキシルロースを生産する酵素量と定義される。生成したキシルロースは、Dische and Borenfreund(Dische and Borenfreund, 1951, J. Biol. Chem. 192: 583-587)の方法によるか、または80℃のBiorad HPX−87Nカラムを使用し、溶出液として0.01M Na2HPO4を使用して0.6ml/minで溶出するHPLCにより測定した。キシロース及びキシルロースは、内部温度60℃で屈折計により測定した。
既に記述されている(2)ベクターラムダZAPII中に構築したcDNAライブラリーを使用した。このライブラリーの一部をExAssistヘルパーファージ(Stratagene,La Jolla,CA,米国)による大量切除(mass excission)によりpBluescript SK−クローンに変換した。無作為に取り出したクローンをM13逆プライマーを使用して配列解析し、5’部分の配列を得た。不完全cDNAを使用してプローブを合成し、ライブラリーを再スクリーニングするために使用した。全長配列を得るためにpUC18にサブクローニングを生成さた。配列分析は、dRhodamineターミネーターサイクルシークエンス即時反応DNAシークエンスキット(Perkin-Elmer Applied Biosystems)を使用してABIプリズム310自動化シークエンサーで実施した。
嫌気性菌Piromyces sp.E2のcDNAライブラリーから無作為に選択したクローンを配列解析し、その結果キシロース異性化酵素及びD−キシルロキナーゼ遺伝子にそれぞれ高い相同性を示す二つのクローン(pH97及びpAK44)を得た。これらのクローンを詳細に解析した。
Piromyces sp.E2のキシロース異性化酵素のSaccharomyces cerevisiae中の発現
Piromyces sp.E2のcDNAをpfuポリメラーゼ(Stratgene)を使用するPCR反応に使用した。プライマーはキシロース異性化酵素の5’及び3’末端の配列を使用して設計し、SfiI及びXbaI制限部位を含めた。PCR産物をpPICZαベクター(Invitrogen,Carlsbad,CA,米国)中にクローニングした。キシロース異性化酵素を取り出すために、pPICZαベクターをEcoRI及びXbaIで消化した。消化産物をpYes2ベクター(Invitrogen)に連結した。キシロース異性化酵素を含むpYes2プラスミドをSaccharomyces cerevisiae(株BJ1991,Beth Johns,UvAから供与)に導入した。この株の遺伝型は:matα,leu2,trp1,ura3−251,prb1−1122及びpep4−3、である。形質転換細胞をSCプレート(0.67% YNB培地+0.05% L−Leu+0.05% L−Trp+2%グルコース+2%アガロース)に接種した。
(培地組成)
Saccharomyces cerevisiae株を下記組成のSC−培地上で増殖した:0.67%(w/v)酵母窒素塩基;0.01%(w/v)L−トリプトファン;0.01%(w/v)L−ロイシン及びグルコース、ガラクトースまたはキシロースまたはこれら基質の組合せのいずれか(下記参照)。寒天培地用に培地に2%(w/v)細菌用寒天を加えた。
挿入のないpYes2で形質転換されたSaccharomyces cerevisiae株BJ1991(遺伝型:matα,leu2,trp1,ura3−251,prb1−1122,pep4−3)及びPiromyces sp.E2キシロース異性化酵素遺伝子を持つpYes2を含む形質転換細胞(16.2.1;16.2.2及び14.3)を、炭素源として10mMグルコースを含むSC−寒天プレート上で増殖した。コロニーが見えるようになったとき、一つのコロニーを使用して、炭素源として100mMキシロース及び25mMガラクトースを含むSVC液体培地に接種した。LKB Ultrospec K分光光度計を使用して600nmの光学密度の増加を測定することにより増殖を監視した。
増殖実験の結果を図1にまとめた。挿入のないpYes2で形質転換したBJ1991株の培養は80時間までOD600の増加を示した。この後徐々に減少が観察された。これは増殖の末期にしばしば観察される酵母細胞の凝集によるものである。3種の形質転換細胞は、80時間後も増殖を止めず、少なくとも150時間までさらに増加を示した。
キシロース異性化酵素をコードするPiromyces sp.E2遺伝子を含むpPICZαベクターを、VentR DNAポリメラーゼ(New England Biolabs)によるPCRの鋳型として使用した。プライマーはキシロース異性化酵素をコードする遺伝子の5’及び3’配列を使用して設計し、EcoRI及びSpeI部位を含めた。さらに、プライマーはpPICZα構築中に認められるXbaI部位を除去し、その代わりに終結コドン(TAA)を入れて設計した。最終産物は、pPICZα構築に認められる追加のアミノ酸(his及びc−Mycタグ)のない、元のオープンリードフレームを復元するように設計した。PCR産物をEcoRI及びSpeIで切り出した。最終産物をpYES2(Invitrogen)由来のベクター中にクローニングした。このベクター中において、キシロース異性化酵素の構成的発現を確実にし、それにより培地にガラクトースを加える必要をなくするために、pYES2中にあるGAL1プロモーターをTPI1プロモーターに置換した。TPI1プロモーターはプラスミドpYX012(R&D systems)の修飾型からクローニングした。このプロモーターをNheI−EcoRIフラグメントとして切り出した。
Claims (17)
- SEQ ID NO.1のアミノ酸配列と少なくとも70%配列同一性(sequence identity)を有するアミノ酸配列(配列同一性は、Needleman and Wunsch,J.Mol.Biol.48:443-453 (1970)に記載のアルゴリズムを使用し、Hentikoff and Hentikoff,Proc.Natl.Acad.Sci.USA 89:10915-10919(1992)に記載のBLOSSUM62 Comparison matrixを使用し、Gap Penalty:12及びGap Length Penalty:4で同定した)を含むキシロース異性化酵素をコードするヌクレオチド配列を含む核酸構築物で形質転換した真菌宿主細胞であって、
該宿主細胞の形質転換で、該核酸構築物により、キシロースをキシルロースに異性化する能力を付与された、真菌宿主細胞。 - 前記宿主細胞が、Saccharomyces,Kluyveromyces,Candida,Pichia,Schizosaccharmyces,Hansenula,Kloeckera,Schwanniomyces,及びYarrowia属の何れか一つに属する酵母である、請求項1に記載の形質転換宿主細胞。
- 前記酵母が、S.cerevisiae,S.bulderi,S.barnetti,S.exiguus,S.uvarum,S.diastaticus,K.lactis,K.marxianus,及びK.fragilis種の何れか一つに属する、請求項2に記載の形質転換宿主細胞。
- 前記宿主細胞が、Aspergillus,Trichoderma,Humicola,Acremonium,Fusarium,及びPenicillium属の何れか一つに属する糸状菌である、請求項1に記載の形質転換宿主細胞。
- キシロースをキシルロースに異性化する能力を宿主細胞に付与するために、キシロース異性化酵素をコードするヌクレオチド配列を、宿主細胞中でキシロース異性化酵素を充分に発現させるプロモーターに作動的に連結している、請求項1から4の何れか1項に記載の形質転換宿主細胞。
- 前記宿主細胞中で、該プロモーターがカタボライト抑制に非感受性である、請求項5に記載の形質転換宿主細胞。
- 前記宿主細胞が
(a)宿主細胞中へキシロース輸送の増加;
(b)キシルロースキナーゼ活性の増加;
(c)ペントースリン酸経路の流量の増加;
(d)カタボライト抑制に対する感受性の減少;
(e)エタノール、浸透圧または有機酸に対する耐性の増加;及び
(f)副産物生産の減少、
からなる群から選択された特徴を生じる遺伝子修飾を含む、請求項1から6の何れか1項に記載の形質転換宿主細胞。 - 遺伝子修飾が、内在性遺伝子の過剰発現、異種遺伝子の発現、またはそれらの組合せからなり、それらによって、該遺伝子が;ヘキソースまたはペントーストランスポーター、キシルロースキナーゼ;ペントースリン酸経路系酵素、解糖系酵素、及びエタノール代謝酵素をコードする遺伝子からなる群から選択される、請求項7に記載の形質転換宿主細胞。
- 前記遺伝子修飾が内在性遺伝子の不活化からなり、それによって、該遺伝子がヘキソースキナーゼ遺伝子、Saccharomyces MIG1及びMIG2遺伝子、並びにそれらの遺伝子の塩基配列に対する相補配列を有するポリヌクレオチドとストリジェントな条件でハイブリダイズし、ヘキソキナーゼ、MIG1又はMIG2活性を有するタンパク質をコードする遺伝子ホモログからなる群から選択される、請求項7に記載の形質転換宿主細胞。
- 前記宿主細胞が、該宿主細胞に、乳酸、酢酸、コハク酸、アミノ酸、1,3−プロパン−ジオール、エチレン、グリセロール、β−ラクタム抗生物質及びセファロスポリンを生産する能力を付与する一つまたはそれ以上の酵素を発現している、請求項1から9の何れか1項に記載の形質転換宿主細胞。
- 前記宿主細胞が、アルコール脱水素酵素活性を減少させる遺伝子修飾を含む、請求項10に記載の形質転換宿主細胞。
- (a)請求項1から9の何れか一項に記載の形質転換真菌宿主細胞であって、アルコール発酵能力を有する細胞と共にキシロース資源を含有する培地を発酵することによって、該宿主細胞によってキシロースをエタノールに発酵させ、任意に
(b)エタノールを回収する、
ステップを含む、エタノールの生産プロセス。 - 前記培地がグルコース資源も含む、請求項12に記載のプロセス。
- エタノールの容積生産性が、少なくとも0.5g/リットル/時間である、請求項12または13に記載のプロセス。
- エタノール収率が少なくとも50%である、請求項12から14の何れか1項に記載のプロセス。
- (a)請求項10又は11に記載の形質転換真菌宿主細胞と共にキシロース資源を含む培地を発酵することによって、該真菌宿主細胞によりキシロースを発酵産物に発酵させ、任意に
(b)発酵産物を回収する、
ステップを含む、乳酸、酢酸、コハク酸、1,3−プロパン−ジオール、エチレン、グリセロール、β−ラクタム抗生物質及びセファロスポリンからなる群から選択される発酵産物を生産するプロセス。 - 該培地がグルコース資源も含む、請求項16に記載のプロセス。
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R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
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EXPY | Cancellation because of completion of term |