JP2021513846A - キメラテルペンシンターゼ - Google Patents
キメラテルペンシンターゼ Download PDFInfo
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- JP2021513846A JP2021513846A JP2020543367A JP2020543367A JP2021513846A JP 2021513846 A JP2021513846 A JP 2021513846A JP 2020543367 A JP2020543367 A JP 2020543367A JP 2020543367 A JP2020543367 A JP 2020543367A JP 2021513846 A JP2021513846 A JP 2021513846A
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- nucleic acid
- host cell
- synthase
- chimeric
- acid molecule
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Abstract
Description
本出願は、米国特許法第119条(e)の定めにより、2018年2月14日出願の米国仮特許出願第62/630,640号、表題“キメラテルペンシンターゼ”に基づく優先権の利益を主張し、その内容全体を引用により本明細書中に包含させる。
本発明は、キメラテルペンシンターゼ、キメラテルペンシンターゼの製造方法およびキメラテルペンシンターゼを用いるテルペン類の製造方法に関する。
テルペン類は、五炭素を構成要素とする有機化合物の多様なクラスであり、少なくとも400の異なる構造的ファミリーを包含する。テルペン類は、その構造的多様性により、フェロモン、抗酸化剤および抗微生物剤などの多くの役割を有している。テルペンシンターゼは原核生物および真核生物の両方でテルペンを生成するが、その多様なテルペン異性体のために、天然源から高収率で抽出することができないことが多い。さらに、テルペンは構造が複雑なため、新規の化学合成には限界がある。
本発明の一面は、配列番号1−52からなる群より選択されるアミノ酸配列と少なくとも90%同一のアミノ酸配列を含むキメラテルペンシンターゼに関する。ある態様において、キメラテルペンシンターゼは、配列番号1−52からなる群より選択されるアミノ酸配列と少なくとも91%、少なくとも92%、少なくとも93%、少なくとも94%、少なくとも95%、少なくとも96%、少なくとも97%、少なくとも98%または少なくとも99%同一のアミノ酸配列を含む。ある態様において、キメラテルペンシンターゼは、配列番号1−52からなる群より選択されるアミノ酸配列と同一のアミノ酸配列を含む。
テルペン類は、香料産業において広く用いられているが、天然源からのテルペン類の精製および新規の化学合成は、高い製造コストおよび低い収率であることが多い。本発明は、少なくとも1つの希少植物または絶滅した植物由来のテルペンシンターゼ配列の一部を含むキメラテルペンシンターゼが、多様なセスキテルペン類を製造できるという予期しない発見を前提としている。従って、本発明は、キメラテルペンシンターゼ、キメラテルペンシンターゼの製造方法および記載のキメラテルペンシンターゼを用いたテルペン類の製造方法を提供する。ある態様において、キメラテルペンシンターゼは、キメラセスキテルペンシンターゼである。
本発明の一面は、少なくとも2つのテルペンシンターゼに由来するフラグメント(例えば、配列)を含むキメラテルペンシンターゼに関し、ここで、2つまたはそれ以上のテルペンシンターゼの少なくとも1つは、希少植物または絶滅した植物に由来する。例えば、キメラテルペンシンターゼの配列は、少なくとも2個、少なくとも3個、少なくとも4個、少なくとも5個、少なくとも6個、少なくとも7個、少なくとも8個、少なくとも9個または少なくとも10個のテルペンシンターゼに由来する1以上のフラグメント(例えば、全配列の1以上の部分)を含み得る。本明細書に記載のキメラテルペンシンターゼは合成であり得ることが理解されるべきである。従って、本明細書に記載の、合成キメラテルペンシンターゼを含むキメラテルペンシンターゼは、2個以上のテルペンシンターゼに由来する配列を含み、ここで、テルペンシンターゼの少なくとも1個は、希少植物または絶滅した植物に由来する。ある態様において、キメラテルペンシンターゼは、キメラセスキテルペンシンターゼである。
本明細書に記載のキメラテルペンシンターゼの配列の少なくとも一部は、希少植物または絶滅した植物に由来する。本明細書で用いる用語“希少植物”または“希少植物類”は、珍しい、希少な、まれに遭遇する、絶滅危惧種(例えば、絶滅の危機に曝されている)、脆弱であり、個人蒐集地でのみ入手可能で、特定の地域(endemic location)では見つからず、栽培でしか入手できず、および/または絶滅した、植物を包含する。ある態様において、希少植物は、まれに遭遇する(例えば、1、2、3、4または5箇所などのいくつかの箇所でのみ遭遇する)植物である。ある態様において、希少植物は絶滅した植物である。本明細書で用いる、絶滅した植物とは、生存種を有しない、生存種を有しないと分類された、または当業者によって生存種を有しないと推測された、植物種を意味する。限定的でない例として、絶滅危惧種の国際自然保護連合(IUCN)のレッドリストを用いて、植物の保全状況を決定し、希少植物を特定できる。例えば、絶滅した、野生で絶滅した、絶滅の危機に瀕している、絶滅の危機にある、および絶滅が近い、IUCNレッドリストで絶滅危惧種に分類されている植物は、希少植物と見なされ得る。
本発明はまた、キメラテルペンシンターゼをコードする核酸分子を記載する。ある態様において、かかるキメラテルペンシンターゼをコードする核酸分子の少なくとも10%(例えば、少なくとも11%、12%、13%、14%、15%、16%、17%、18%、19%、20%、21%、22%、23%、24%、25%、26%、27%、28%、29%、30%、31%、32%、33%、34%、35%、36%、37%、38%、39%、40%、41%、42%、43%、44%、45%、46%、47%、48%、49%、50%、51%、52%、53%、54%、55%、56%、57%、58%、59%、60%、61%、62%、63%、64%、65%、66%、67%、68%、69%、70%、71%、72%、73%、74%、75%、76%、77%、78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%、99%、または99%を超える、その間の全ての値を含む)は、希少植物または絶滅した植物に由来し得る。
実施例1.キメラテルペンシンターゼの機能的特徴付け
12種の絶滅した植物サンプルからのゲノムDNAを配列決定した(表2)。セスキテルペンシンターゼ(SQTS)フラグメントは、7つの植物から取得されたが(表11)、配列のギャップにより完全長遺伝子の再構築は不可能であった。2,738のテルペンシンターゼキメラ(配列のギャップを埋めるために、現存する植物のセスキテルペンシンターゼの配列を含み、配列のギャップを埋める)を含むライブラリーをスクリーニングした。6つの希少植物(表2)からの52のSQTSキメラ(表10に示す配列)の発現により、スクリーニング株でセスキテルペンの産生がもたらされた。本実施例に記載された各操作の方法および材料は、実施例2に見出され得る。
Capture-seqおよびキメラ足場用のテルペンシンターゼ
セスキテルペンシンターゼ(SQTS)の候補を、希少花ゲノム(rare flower genome)からの配列フラグメント(表11)とUniProtおよびGenBankを含む供給源からの“足場”(scaffold)SQTSを組み合わせることにより設計した。
キメラSQTSを構築するために2つの方法を用いた:1)blastn-mapDamageパイプラインおよびb)tblastnパイプライン。
一般的に、blastn-mapDamageパイプラインを、足場と類似性の高いヌクレオチドを有するフラグメントを注意深く(conservatively)に検出し、結果として、希少花の元の酵素配列と非常に近いと思われるキメラテルペンシンターゼ(例えば、キメラセスキテルペンシンターゼ)を得た。希少花DNAのステレオタイプの損傷による人工物(artifact)であり得る変異体を検出するために、bam形式のイルミナの読み取りアラインメントをmapDamageソフトウェアに入力した。
1.ゲノム capture-seq ランからのIllumina読み取り(fastqファイル)を合わせ、SPADESによってより長いコンティグ(contig)にアセンブルした。
2.1521セットのSQTS足場を、SPADESコンティグに対するデフォルトパラメーターを用いたblastn検索でクエリーとして用いた。比較的少数の骨格にヒットがあったため、次のステップでの読み取りアライメントの対照として、ヒットのある骨格をすべて選択した。
3.配列決定ランからの読み取りの組み合わせは、質をトリミングし(quality-timmed)(bbdukを使用)、ペアを合わせ(bbmergeを使用)、そしてbwamemを用いて選択したSQTS対照配列に整列させた。結果は、BAMに再フォーマットされ、ソートされ、索引付けされた。
4.mapDamageを、整列された読み取りに対して実行した。これにより、DNA損傷に似たSNPに低い質スコアが割り当てられた読み取りアライメントが得られた。
5.読み取りアライメントを以下のように処理した:質が25未満の塩基をマスキングした(対照、に変更した);アラインメントをfastaに再フォーマットした;カウント数6未満のSNPをマスキングした;重複した読み取りを除いた;頻度が0.1未満のSNPをマスキングした;他の読み取りの正確な部分配列(subsequences)の読み取りを除いた;読み取りを対照のフレームで翻訳した;そして、部分配列を再び除いた;アライメントのマスキングに用いる質およびSNP頻度の閾値を、質およびSNPの頻度の分布を調べることにより、実験的に決定した。
6.読み取りアライメントおよびSPADEコンティグアライメント(対照フレーム翻訳後)を合わせ、Clustal Omegaを用いて再配置した。これは、一部のコンティグが、読み取りされなかった足場の領域にまたがっていたために行われた。
1.アラインメントを、各サブリージョンには、別のサブリージョン(サブリージョン間で同一の重複が許可された)のフラグメントと重複するおよび異なるサブリージョンからのフラグメント(アラインメントされた読み取り)が含まれていないように、“独立したサブリージョン”に分割した。
2.各サブリージョンで、“互換性のある(compatible)フラグメント”のすべての可能な組み合わせを列挙した。互換性のあるフラグメントは、同一としてオーバーラップする(従って、より長いフラグメントに融合できる)か、まったくオーバーラップしない(例えば、同じハプロタイプに由来すると推定された)フラグメントとして定義された。フラグメントの組み合わせは“最大カバレッジ(coverage)”であった。すなわち、可能な限り多くの互換性のあるフラグメントが含まれた。各最大カバレッジフラグメントの組み合わせを、アライメントのその領域の可能な再構築であると見なし、スーパーフラグメント(ギャップが含まれていてよい)に融合させて(merged)、保存した。
3.各サブリージョンのスーパーフラグメントを、通常の反復アルゴリズムを用いて90%または95%の同一性にダウンサンプリングし、異なるサブリージョンのダウンサンプリングされたスーパーフラグメントのすべての可能な組み合わせを組合せた。特定の閾値より短い領域を、単一の配列にダウンサンプリングする。スーパーフラグメントの各組み合わせを足場にマージして、キメラ配列を作成した。ダウンサンプリングパラメータを、いずれの場合も1を超えるが100未満のキメラを構築できるように、サンプルおよび足場に応じてわずかに変化させた。
一般に、tblastnパイプラインは、SQTS足場に相同なフラグメントを検出する感度を最大化したため、使用可能な配列に対して広いネットを形成した。
1.1521セットのSQTS足場を、SPADESコンティグ(上記)のすべてのフレームの翻訳を検索するためのtblastnへのタンパク質クエリーとして用いた。
2.ヒット(整列されたコンティグ)を、足場に対して最低40%の同一性、およびヒューリスティック関数によってヒットの同一性に依存する最小の長さにフィルタリングした。フィルタリング基準は、すべてのサンプルにわたる同一性とヒット長プロットの検定によって選択した。
3.足場のダウンサンプリングを、各ヒットと同一の残基の数によって足場を階層的にクラスター化することによって実行した。キメラの再構築のために、すべてのヒットに対して同一性の数が最も多い骨格を各クラスターにおいて保持した。ダウンサンプリングにより、骨格の数が20分の1に減少した。ヒットが10未満の骨格の場合、この手順はスキップされた。
4.特定の骨格が、以前に活性を有していると識別(同定)されていたり、および/または文献で知られていたりしたため(別の配列がヒットに対してより多くの同一性を有していたとしても)、常にクラスターの代表として選択された。これらの好ましい足場はダウンサンプリングされず、キメラ構築のためにtblastnヒットを保持した。
5.足場にヒットするすべてのコンティグの整列部分を、Clustal Omegaを用いて足場に再整列した。コンティグの整列されていない部分は、イントロンを表す可能性が高いとして廃棄した。この整列(アライメント)を、キメラ構築に用いた。
6.キメラを、サブ領域でダウンサンプリングすることなく、上記のコンビナトリアル互換フラグメント法を用いて、整列されたtblastnヒットから構築した。“最大カバレッジ”(各セットで可能な限り互換性のあるフラグメント)と“最小カバレッジ”(各セットでただ1つの互換性のあるフラグメント)の両方のキメラを作成した。最小カバレッジキメラは、無関係な配列からのフラグメントの結合を回避し得る。
各酵素を2回コドン最適化した。1回目は酵母発現加重コドン表を用い、もう1回は、10%未満の頻度でコドンを除去した後に酵母発現加重コドン表を用いた。異なる乱数(random number)を各エンコーディングのシードとして用いた。異なる酵素のエンコーディングは完全に独立しており、キメラが足場から継承した残基のコドンを保存するために特定の操作は用いなかった。
絶滅した花種の1つであるShorea cuspidataについては、現存する近縁種であるShorea beccarianaのトランスクリプトーム配列データが利用可能であった。これにより、関連する花からSQTS足場を用いてキメラを構築することが可能になった。これは2段階のプロセスで行われた:
1.S. beccariana (Sb)トランスクリプトームデータを組み立て、SQTS相同体のために採用した。データを、NCBI SRAデータベースのデータセットSRR687302からダウンロードした。組み立てはTrinityを用いて行い、ORFはTransdecoderにより予測された。BLASTを用いて、1,500のキュレートされたSQTS配列のセットに相同なフラグメントを同定した。
2.同定されたSb SQTSまたはSQTSフラグメントを、tblastnまたはblastn-mapDamageパイプラインで足場配列として用いて、キメラを再構築した。足場自体がフラグメントだった場合は、次に、最も近いUniprotをソースとするSQTS配列にマージして、完全長キメラを生成した。
キメラセスキテルペンシンターゼを、ガラクトース誘導性P(gal1)プロモーターの制御下で高コピーpESC-URA3-由来の発現ベクターに形質転換した(Sikorski et al., A system of shuttle vectors and yeast host strains designed for efficient manipulation of DNA in Saccharomyces cerevisiae. Genetics. 1989 May;122(1):19-27、この目的のためにその内容全体を引用により本明細書中に包含させる)。
形質転換コロニーを、96ディープウェルプレート中、300μlのSC-ura培地(2%デキストロースを含む合成完全培地、ウラシル無添加)に接種した。プレートを、Excel Scientific AeroSeal膜(BS-25)で覆い、振とうインキュベーター中で30℃にて48時間インキュベートした。30μlの培養物(1:15希釈)を、炭素源として、1.8%ガラクトースおよび0.2%ラフィノースを含む420μlのSC-ura 誘導培地に混合し、600nm(OD600)での開始光学密度が約0.1−0.2となるようにした。0.88%ドデカンオーバーレイ(4μl)を各ウェルに添加し、プレートをAeroSeal膜で覆い、振とうインキュベーター中で30℃にて4日間培養した。各培養物15μlを取りだし、4日間の終了時にOD600を測定した。350μlの酢酸エチル(250μMトリデカン内部標準)を各ウェルに直接添加し、混合した(1:1抽出)。次いで、96ウェルプレートを遠心分離し、酢酸エチル抽出物を、GC−MSによる分析までガラスバイアル中で−80℃にて保存した。
酢酸エチルサンプル(1.0uL)を、Agilent/Gerstel 7890B GCシステムに注入し、GCインレットを250℃に設定し、スプリット比を2:1にした。キャピラリーカラムは、担体ガス(ヘリウム)の流量を1.5ml/分に設定してAgilent DB-5MS (20m×0.18mm×0.18μm)を用いた。GCオーブン温度を、40℃/分の上昇で100℃(0.10分間保持)に設定し、155℃まで上昇させ、そこから15℃/分の上昇で190℃まで上昇させ、最後に75℃/分の上昇で280℃まで上昇させた(5分間法)。標的のより包括的な分析のために、GCオーブン温度を、10℃/分の上昇で100℃(2.0分間保持)に設定し、250℃まで上昇(2.0分間保持)させる方法を利用した(20分間法)。両法とも、Agilent 5977B MSD (Etune)上のMS源および四重極は、それぞれ230℃および180℃に設定した。マススキャン範囲を40−250mzに設定し、スペクトルおよび線形保持指数の計算を、利用可能な標準物質および精油に加えて、NIST MS データベース (2008年版)と照合した。
当業者は、本明細書に記載される本発明の特定の態様に対する多くの均等物を認識し、または常套の実験のみを用いて確認することができるであろう。そのような均等物は、以下の特許請求の範囲に含まれることが意図される。
Claims (41)
- 配列番号1−52からなる群より選択されるアミノ酸配列と少なくとも90%の同一性を有するアミノ酸配列を含む、キメラテルペンシンターゼ。
- 配列番号1−52からなる群より選択されるアミノ酸配列と同一のアミノ酸配列を含む、請求項1記載のキメラテルペンシンターゼ。
- 請求項2記載のキメラテルペンシンターゼをコードする核酸分子。
- 請求項3記載の核酸分子を含むベクター。
- ウイルスベクター、一過性発現用ベクターまたは発現誘導用ベクターである、請求項4記載のベクター。
- レンチウイルスベクター、レトロウイルスベクター、アデノウイルスベクター、アデノ随伴ウイルスベクター、ガラクトース誘導性ベクターまたはドキシサイクリン誘導性ベクターである、請求項4または5記載のベクター。
- 請求項3記載の核酸分子を含む宿主細胞。
- 請求項4から6のいずれか一項記載のベクターで形質転換されている宿主細胞。
- 真菌細胞である、請求項7または8記載の宿主細胞。
- 真菌細胞が酵母細胞である、請求項9記載の宿主細胞。
- 酵母細胞が出芽酵母、ピキア酵母、クリベロマイセス酵母、ハンゼヌラ属酵母またはヤロウイア属酵母の細胞である、請求項10記載の宿主細胞。
- 出芽酵母細胞である、請求項11記載の宿主細胞。
- 植物細胞である、請求項7または8記載の宿主細胞。
- 細菌細胞である、請求項7または8記載の宿主細胞。
- キメラテルペンシンターゼをコードする核酸分子であって、ここで、該核酸分子が、配列番号67−118からなる群より選択されるヌクレオチド配列と少なくとも90%の同一性を有するヌクレオチド配列を含む、核酸分子。
- 配列番号67−118に記載の配列セットを含む、請求項15記載の核酸分子。
- TATAボックス配列をさらに含む、請求項15または16記載の核酸分子。
- 核酸分子配列の少なくとも40%が絶滅した植物に由来する、請求項3記載の核酸分子。
- キメラテルペンシンターゼが、キメラセスキテルペンシンターゼである、請求項1から18のいずれか一項記載のキメラテルペンシンターゼ、核酸分子または宿主細胞。
- キメラテルペンシンターゼが、アルファ−グアイエンシンターゼである、請求項19記載のキメラテルペンシンターゼ、核酸分子または宿主細胞。
- アルファ−グアイエンシンターゼが、配列番号17、22または29と少なくとも90%同一の配列を含む、請求項20記載のキメラテルペンシンターゼ、核酸分子または宿主細胞。
- アルファ−グアイエンシンターゼが、配列番号17、22または29を含む、請求項21記載のキメラテルペンシンターゼ、核酸分子または宿主細胞。
- アルファ−グアイエンシンターゼが、配列番号17、22または29からなる、請求項18記載のキメラテルペンシンターゼ、核酸分子または宿主細胞。
- 絶滅した植物が、Hibiscadelphus wilderianus、Leucadendron grandiflorum、Macrostylis villosa、Orbexilum stipulatum、Shorea cuspidateおよびWendlandia angustifoliaからなる群より選択される、請求項18記載の核酸分子。
- 1以上のセスキテルペンを製造する方法であって、ここで、該方法が、請求項3、15から18および24のいずれか一項記載の核酸を含む宿主細胞を、1以上のセスキテルペンを製造するのに適する条件下で培養することを含む、方法。
- 請求項25記載の方法により製造された1以上のセスキテルペンを含む組成物。
- 1以上のセスキテルペンの少なくとも1つが芳香化合物である、請求項26記載の組成物。
- 香料を製造する方法であって、該方法が、
請求項3、15から18および24のいずれか一項記載の核酸を含む宿主細胞を、1以上のセスキテルペンを製造するのに適する条件下で培養すること;および
1以上のセスキテルペンを抽出すること
を含む、方法。 - アルファ−グアイエンを製造可能なキメラテルペンシンターゼであって、ここで、該キメラテルペンシンターゼのアミノ酸配列の少なくとも40%が、絶滅した植物に由来する、キメラテルペンシンターゼ。
- アルファ−グアイエンを製造可能なキメラテルペンシンターゼをコードする核酸分子であって、ここで、該核酸分子配列の少なくとも40%が、絶滅した植物に由来する、核酸分子。
- 請求項30記載の核酸を発現する宿主細胞。
- アルファ−グアイエンを製造可能なキメラテルペンシンターゼを発現する宿主細胞。
- 請求項31または32記載の宿主細胞を培養することを含む、アルファ−グアイエンの製造方法。
- 宿主細胞により産生されたアルファ−グアイエンを取得することをさらに含む、請求項33記載の方法。
- 真菌細胞である、請求項31または32記載の宿主細胞。
- 植物細胞である、請求項31または32記載の宿主細胞。
- 細菌細胞である、請求項31または32記載の宿主細胞。
- 酵母細胞である、請求項31、32または35記載の宿主細胞
- 酵母細胞が、出芽酵母、ピキア酵母、クリベロマイセス酵母、ハンゼヌラ属酵母またはヤロウイア属酵母である、請求項38記載の宿主細胞。
- 酵母細胞が出芽酵母細胞である、請求項39記載の宿主細胞。
- 配列番号119−357からなる群より選択される配列を含むキメラテルペンシンターゼ。
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