JP7019580B2 - 単為結実植物およびその製造方法 - Google Patents
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Description
米国特許出願第20100146656号明細書
米国特許出願第20090089902号明細書
米国特許出願第20030217391号明細書
米国特許出願第20070250945号明細書
米国特許第6,268,552号明細書。
(i)非単為結実トマトの受精可能条件下において、植物当たりの果実収量が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの果実収量とほぼ同一またはそれ以上である、
(ii)非単為結実トマトの受精可能条件下において、植物当たりの平均果実重量が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの平均果実重量とほぼ同一またはそれ以上である、
(iii)前記ナス科植物がトマトであるとき、ゼリー部を含む、
(iv)前記果実収量の少なくとも80%にホメオティック遺伝子異常がない、
(v)種なし果実内の肥大した胚珠が非受精子房から発生したものである。
(i)非単為結実トマトの受精可能条件下において、植物当たりの果実収量(fruit yield/plant)が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの果実収量とほぼ同一(例えば、80%、90%、100%、110%)またはそれ以上である、
(ii)非単為結実トマトの受精可能条件下において、植物当たりの平均果実重量(fruit weight/plant)が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの平均果実重量とほぼ同一またはそれ以上である、
(iii)植物がトマトであるとき、ゼリー部を含む、
(iv)果実収量の少なくとも80%にホメオティック遺伝子異常がない、
(v)種なし果実内の肥大した胚珠が非受精子房から発生したものである。
(i)アグロバクテリウム仲介性遺伝子導入:Klee et al. (1987) Annu. Rev. Plant Physiol. 38:467-486、Klee and Rogers in Cell Culture and Somatic Cell Genetics of Plants, Vol. 6, Molecular Biology of Plant Nuclear Genes, eds. Schell, J., and Vasil, L. K., Academic Publishers, San Diego, Calif. (1989) p. 2-25、Gatenby, in Plant Biotechnology, eds. Kung, S. and Arntzen, C. J., Butterworth Publishers, Boston, Mass. (1989) p. 93-112。
(ii)直接DNA取り込み:Paszkowski et al., in Cell Culture and Somatic Cell Genetics of Plants, Vol. 6, Molecular Biology of Plant Nuclear Genes eds. Schell, J., and Vasil, L. K., Academic Publishers, San Diego, Calif. (1989) p. 52-68。この方法には、プロトプラストに直接DNAを取り込ませる方法も含まれる、Toriyama, K. et al. (1988) Bio/Technology 6:1072-1074。植物細胞の短時間の電気ショックによって誘導されるDNA取り込み:Zhang et al. Plant Cell Rep. (1988) 7:379-384.Fromm et al. Nature (1986) 319:791-793。微粒子銃による植物の細胞または組織へのDNA注入、Klein et al. Bio/Technology (1988) 6:559-563、McCabe et al. Bio/Technology (1988) 6:923-926、Sanford, Physiol. Plant. (1990) 79:206-209。マイクロピペットシステムの使用による方法:Neuhaus et al., Theor. Appl. Genet. (1987) 75:30-36、Neuhaus and Spangenberg, Physiol. Plant. (1990) 79:213-217。
EMS誘導M2集団の、極端な温度下での収量についてのスクリーニング: 約1000M2 EMSで変異誘発されたファミリーについて、1ファミリー12株の植物を圃場に植え付けた。当該12株は、2つの複製となる群に分けられた、即ち、各々6株ずつを別個のブロックに植えた。対照となる親系統M82を同様に、1群6株が複製となるように圃場に植えた。このとき、22組の2重列の各組に対して、対照が少なくとも1群となるようにしたに。実生を晩夏(2009年7月7日)に圃場に植え、8月~9月の極めて暑い月の果実結実および発生について、8月12日を開始日とし、スクリーニングを繰り返した。選択した変異2012と、大きな単為結実果実であるが、その遠位(花柱)末端が幾分かゆがんだ形状である別の1種とを除いては、いくつかのファミリーから1株のみが単為結実果実を結実することがわかった。その他のすべてが比較的小さい(35g未満)種なし果実を結実し、これらでは、果実形成能を最大限に引き出すという種子の貢献が部分的にしか補償されていないと考えられる。
Solyc01g093960 gRNA標的部位を、PAM上流の3bpと重複する、Cas9ヌクレアーゼの予測される切断部位であるAclI制限酵素部位(AACGTT、配列番号12)を含むように設計することで(図2Bを参照)、DNA二本鎖切断(DSB)修復が、制限部位を破壊し得るようにした。R0植物を、変異標的部位を保持するキメラセクションの存在についてスクリーニングした。再生R0植物の後期発生期で生じる、Cas9によって作製された変異が検出される可能性を高めるために、主茎上の10~12番目の葉よりも若い葉からゲノムDNAを抽出した。以下の特異的プライマー:2012-F: 5’-GCCTTGAAATCAGTAAGAGTATTGG-3’(配列番号13)、および2012-R: 5’-GTTCGTTGAAGTGCTTCAAACTTGG-3’(配列番号14)を使用してDNAを増幅し、sgRNA標的配列に隣接する354bpの断片を得た。変異されていない限り、そのAclI消化産物は、類似した大きさの2本のバンド(170bp/184bp、図2Bを参照)をもたらす。消化しれたアンプリコンをゲル電気泳動に供した。UV光下で明瞭なバンドが可視化されない場合であっても、切断されていないバンドの位置のゲルからDNAを抽出した。抽出されたDNAを、-20℃で一晩沈殿させ(0.3M酢酸Na、pH=5.2、2.2容量エタノールおよび1μgグリコーゲン)、10μLの水に再懸濁し、それから1μLを、同じプライマーを使用する増幅のための鋳型として用いた。制限部位の喪失および作製された変異の性質を試験するために、増幅されたバンドの両方をAclIを用いて制限処理し、フォワードおよびリバースプライマーの両方を使用してシーケンシングした。変異型を含有するとわかったR0植物のほとんどにおいて、AclI耐性PCRバンドは、制限処理された二本鎖よりもかなり少量であり、変異型が、R0植物の再生初期ではなく後期に作製されたことを強力に示唆した。同一手順をR1後代の遺伝子型解析に適用した。
実験を4回の反復で実施し、各々は、遺伝子型あたり17~27植物体からなる。植物を2016年4月20日にネットハウス内に植え付け、2016年6月26日に最初の収穫を実施した(図13を参照のこと)。各反復実験について、すべての赤色果実を収穫し、秤量した。各反復実験について、30個の果実を6~8バッチ秤量した結果から、植物あたりの果実重量および果実数を算出した。図13に示されるデータは、植物の数が反復間で変動したため、植物あたりで算出した。Brixは、各反復実験について、それぞれが3個の赤く熟した果実からなる2つのプールから絞られたジュースで測定した(Carmi et al.,2003に従って)。pHは、各反復実験について、それぞれが6~8個の赤く熟した果実からなる2つのプールで測定した。
EMS変異誘発されたM82集団由来の、単為結実について新規な一遺伝子性劣性変異体としての系統2012の同定
2012変異のNGS補助マッピング
変異をマッピングするために、バルク分離アプローチ(Michelmore et al 1991)を選択した。ディープシーケンシングを2つのゲノムライブラリーで実施した。当該ライブラリーは、一方は2012 BC1F2単為結実同胞のバルクに由来し、もう一方は、その非単為結実同胞に由来する。
2012変異のマーカー補助微細マッピング
2012変異体の根底にある変異の位置を限定する試みとして、変異位置として示唆される染色体区間に沿って分散された6個のSNP(表1、第2列)と、単為結実表現型との共分離を、(材料および方法に記載のように作製された)試験交雑種(TC)集団において調べた。この集団を、2013年の夏の終わりに熱ストレス下で果実結実させた。表1、第2列に特定した6つの候補SNPの各々について、TC集団の遺伝子型解析を、イスラエル国のDYN R&D社に委託して実施した(材料および方法を参照のこと)。
2012単為結実の遺伝子としての、変異SlAGL6のCRISPRに基づく検証
変異AGL6が、2012単為結実の根底にある遺伝子であることを確認するために、CRISPR技術を利用して、SlAGL6遺伝子Solyc01g093960をノックアウトした。
CRISPR誘導性AGL6変異R1後代は、単為結実である
性質は異なっているが、すべて未成熟停止コドンにつながる変異を有する、sg1、sg4およびsg5と呼ばれる3株のR0植物の後代(表4)を、単為結実果実を結実するその能力の解析のために選択した。これらの変異は、未成熟停止コドンをもたらすので、これらの変異遺伝子から転写された異常なmRNAは、ナンセンス変異依存性分解機構(Nonsense-Mediated Decay)(NMD)RNAサーベイランス機構を介して破壊されると予測される(例えば、Chang et al. 2007、Drechsel et al. 2013、Degtiar et al 2015)。
収量パラメータに対する変異SlAGL6の影響の推定は、副作用を示唆しない
AGL6変異が収穫パラメータに与え得るペナルティーを評価するための第1のステップとして、以下の3つの異なる集団における果実重量に対するその効果を調べた: 系統M82をバックグラウンドとする、分離有限成長2012 BC2F2集団、大果実、半有限成長栽培品種マルマンデと単為結実2012との間の分離F2集団、および無限成長MP-1系統をバックグラウンドとする、CRISPR/Cas9系統sg1のR1後代。2012 BC2F2集団では、総収量に対する効果も推定した。
一般に、2012 BC2F2集団のホモ接合性変異(m/m)植物で発生した果実のほとんどが、周囲条件下で発生させた場合であっても、種なしであった。しかし、変異は、種子から繁殖されるトマトのような作物にとって重篤な弱点となる、不可避な単為結実のためのものではない。受精可能条件下での変異植物による種子結実に好都合な正確な条件は、解明されるべきもののままである。活発な花序震盪は、それなしでは主に種なしのものを結実する(データは示さず)植物から多数の種あり果実をもたらした。意図的な花序震盪の不在下では、増強された傾向を2つのパラメータと明確に関連させることができた。第1に、周囲条件下で古い植物で発生した小さい果実は、頻繁に種子を有していた。種あり果実が、同一植物で発生した種子が不十分なものまたは単為結実のものよりも大きいということは、広く受け入れられているので、これは例外的な現象である(例えば、Imanshi and Hiura 1975、Varga and Bruinsma. 1976、Carmi et al 2003)。第2に、最適温度よりも少し低い温度で結実した果実は、種子を含有することが多かった。どちらの場合も、種子の産生は、おそらくは、果実への子房拡大の速度を減速させる条件を反映し、したがって、そうでなければ急速に拡大する胚珠/果実から花柱が脱離する前の、花粉粒の完全な発芽、伸長および胚珠の受精を可能にする。多数の種あり果実も、2012×マルマンデ交雑種の(m/m)F2後代でも発生した(図9A~B)。同様に、周囲条件下で、MP-1をバックグラウンドとするde novo変異AGL6も、種なし果実および種あり果実の両方の形成を可能にした(表5、図12)。
Slagl6は、熱ストレス下での収量を改善する
MP-1と、Slagl6についてホモ接合性であり、(R2が)Cas9カセットを欠くsg1系統とを比較することによって、天然の熱ストレス条件下での収量を調べた。2016年4月20日にネットハウス中に植物を植え、67日後に最初の収穫を実施した。5月および6月の間の日中温度は非常に高く、(2016年5月14日から16日の)3日連続した極めて高い温度(最大日中温度38℃以上、図13H)も含まれる。これらの天然に存在する熱ストレス条件は、小胞子形成、したがって、受精依存性果実結実を妨害すると知られている「慢性の軽度熱ストレス」の定義に当てはまる(Mesihovic et al., 2016およびその中の参考文献を参照)。図13Aによって示されるように、これらの気候条件下で、親系統MP-1の赤色果実収量は、系統sg1のものよりも有意に低かった(約85%低い)。この違いは、系統MP-1において83%低かった、産生果実数における著しい相違(図13B、F、G)と、また13%だけではあるが、後者において有意に低い果実重量(図13C)を主に反映している。その他の単為結実変異体(例えば、Carmi et al., 2003、Casas Diaz et al., 1987)と同様に、赤く熟した種なし果実のBrixとして表される総可溶性固形物(TSS)含量が、MP-1の種あり果実のものよりも有意に高かったが(図13D)、果実の酸度(pH)は、同様のままであった(図13E)。
条件的単為結実ナスおよびトウガラシの作製
CaAGL6(トウガラシ)またはSmAGL6(ナス)を、トマトについて記載したように(本明細書の実施例1)、種子の化学的変異誘発によって変異させる。変異誘発処理した種子から栽培し、自家受粉された植物から生じたM1変異誘発ファミリーを、例えば、TILLING技術(Till et al 2003)に従って、標的遺伝子中の変異についてスクリーニングする。標的遺伝子中に変異を保持すると同定されたファミリーを、熱ストレスまたは花の雄蕊除去のいずれかである受精制限条件下で繁殖させ、単為結実果実結実について試験する。上記の代わりに、安定な形質転換を介する、トマトについて行ったような、例えば、CRISPR/Cas9技術を利用する遺伝子編集によって、標的遺伝子中に欠失を作製する。
>Chr01-44480416番ヌクレオチド~44480435番ヌクレオチドの間のCapana01g00134遺伝子標的
GTATCACCAAAACCCTTGAG(配列番号17)。
182番~201番ヌクレオチドの間のCapana01g00134 cDNAを標的と、切断は、198番~199番ヌクレオチドの間で起こると予測される。
>29543番~29562番ヌクレオチド(マイナス鎖)の間のSme2.5_06058.1標的
GAGGATTAAGGCAACAACGT(配列番号18)。
204番~229番ヌクレオチドの間のSME2.5 06058.1 cDNAを標的とし、切断は、212番~213番ヌクレオチドの間で起こると予測される。
1. 100 Tomato Genome Sequencing Consortium, Aflitos S, Schijlen E, de Jong H, de Ridder D, Smit S, Finkers R, Wang J, Zhang G, Li N, Mao L, Bakker F, Dirks R, Breit T, Gravendeel B, Huits H, Struss D, Swanson-Wagner R, van Leeuwen H, van Ham RC, Fito L, Guignier L, Sevilla M, Ellul P, Ganko E, Kapur A, Reclus E, de Geus B, van de Geest H, Te Lintel Hekkert B, van Haarst J, Smits L, Koops A, Sanchez-Perez G, van Heusden AW, Visser R, Quan Z, Min J, Liao L, Wang X, Wang G, Yue Z, Yang X, Xu N, Schranz E, Smets E, Vos R, Rauwerda J, Ursem R, Schuit C, Kerns M, van den Berg J, Vriezen W, Janssen A, Datema E, Jahrman T, Moquet F, Bonnet J, Peters S. (2014) Exploring genetic variation in the tomato (Solanum section Lycopersicon) clade by whole-genome sequencing. Plant J 80(1):136-48.
2. Adelberg JW; Xhang XP, Rhodes BB. (1997) Micropropagation of Citrullus lanatus (Thunb.) Matsum. and Nakai (watermelon). In: Bajaj, Y. P. S., ed. Biotechnology in agriculture and forestry, Vol. 39: High-tech micropropagation. Berlin: Springer-Verlag; pp.:60-76.
3. Ariizumi T, Shinozaki Y, Ezura H. (2013) Genes that influence yield in tomato. Breed Sci. 63(1):3-13.
4. Barg R, Pilowsky M, Shabtai S, Carmi N, Szechtman AD, Dedicova B, Salts Y. (1997) The TYLCV-tolerant tomato line MP-1 is characterized by superior transformation competence J. Exp. Bot 48: 1919-1923.
5. Beraldi D, Picarella ME, Soressi GP, Mazzucato A. Fine mapping of the parthenocarpic fruit (pat) mutation in tomato. Theor Appl Genet. 2004.
6. Bohner J, Bangerth F (1988) Cell number, cell size and hormone levels in semi-isogenic mutants of Lycopersicon pimpinellifolium differing in fruit size. Physiologia Plant. 72: 316-320.
7. Bolger A, Scossa F, Bolger ME, Lanz C, Maumus F, Tohge T, Quesneville H, Alseekh S, Sorensen I, Lichtenstein G, Fich EA, Conte M, Keller H, Schneeberger K, Schwacke R, Ofner I, Vrebalov J, Xu Y, Osorio S, Aflitos SA, Schijlen E, Jimenez-Gomez JM, Ryngajllo M, Kimura S, Kumar R, Koenig D, Headland LR, Maloof JN, Sinha N, van Ham RC, Lankhorst RK, Mao L, Vogel A, Arsova B, Panstruga R, Fei Z, Rose JK, Zamir D, Carrari F, Giovannoni JJ, Weigel D, Usadel B, Fernie AR. (2014) The genome of the stress-tolerant wild tomato species Solanum pennellii. Nat Genet. 46(9):1034-1038.
8. Bolger AM, Lohse M, Usadel B. (2014) Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics. 30(15):2114-2120.
9. Carmi N, Salts Y, Dedicova B, Shabtai S, Barg R (2003) Induction of parthenocarpy in tomato via specific expression of the rolB gene in the ovary. Planta 217: 726-735.
10. Casas Diaz AV, Hewitt JD, Lapushner D (1987) Effects of parthenocarpy on fruit quality in tomato. J Am Soc Hort Sci 112:634-637.
11. El Ahmadi AB, Stevens MA (1979) - Reproductive responses of heat-tolerant tomatoes to high temperatures. J. Amer. Soc. Hort. Sci. 104:686-691.
12. Falavigna A, Badino M, Soressi GP. - (1978) Potential of the monomendelian factor pat in the tomato breeding for industry. Genetica Agraria, 32: 159-160 (Abstr.).
13. Ficcadenti N, Sestili S, Pandolfini T, Cirillo C, Rotino GL, Spena A (1999) Genetic engineering of parthenocarpic fruit development in tomato, Mol. Breed. 5:463-470.
14. Galpaz N, Burger Y, Lavee T, Tzuri G, Sherman A, Melamed T, Eshed R, Meir A, Portnoy V, Bar E, Shimoni-Shor E, Feder A, Saar Y, Saar U, Baumkoler F, Lewinsohn E, Schaffer AA, Katzir N, Tadmor Y.(2013) Genetic and chemical characterization of an EMS induced mutation in Cucumis melo CRTISO gene. Arch Biochem Biophys. 539(2):117-125.
15. Ghio C, Ramos ML, Altieri E, Bulos M, Sala CA. (2013) Molecular characterization of Als1, an acetohydroxyacid synthase mutation conferring resistance to sulfonylurea herbicides in soybean. Theor Appl Genet. 126(12):2957-2968.
16. Gillaspy G, Ben-David H, Gruissem W (1993) Fruits: a developmental perspective. Plant Cell 5: 1439-1451.
17. Gorguet B, Eggink PM, Ocana J, Tiwari A, Schipper D, Finkers R, Visser RGF, van Heusden AW (2008) Mapping and characterization of novel parthenocarpy QTLs in tomato. Theor Appl Genet 116:755-767.
18. Gorguet B, van Heusden AW, Lindhout P. (2005) Parthenocarpic fruit development in tomato. Plant Biol (Stuttg). 7:131-139.
19. Hazra P, Dutta AK. (2010). Inheritance of parthenocarpy in tomato (Solanum lycopersicum) and its association with two marker characters. Int J Plant Sci, 1: 144-149.
20. Imanshi S, Hiura I (1975) Relationship between fruit weight and seed content in the tomato. J Jpn Soc Hort Sci 44:33-40.
21. Joubes J, Chevalier C. (2000) Endoreduplication in higher plants. Plant Mol Biol. 43:735-745. Review.
22. Koo, S.C., Bracko, O., Park, M.S., Schwab, R., Chun, H.J., Park, K.M., Seo, J.S., Grbic, V., Balasubramanian, S., Schmid, M., Godard, F., Yun, D.J., Lee, S.Y., Cho, M.J., Weigel, D. and Kim, M.C. (2010) Control of lateral organ development and flowering time by the Arabidopsis thaliana MADS-box Gene AGAMOUS-LIKE6. Plant J. 62,807-816.
23. Li H, Durbin R. (2009) Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics. 25(14):1754-1760.
24. Li H, Handsaker B, Wysoker A, Fennell T, Ruan J, Homer N, Marth G, Abecasis G, Durbin R; 1000 Genome Project Data Processing Subgroup.(2009) The Sequence Alignment/Map format and SAMtools. Bioinformatics. 25(16):2078-2079.
25. Li JF, Norville JE, Aach J, McCormack M, Zhang D, Bush J, Church GM, Sheen J. (2013). Multiplex and homologous recombination-mediated genome editing in Arabidopsis and Nicotiana benthamiana using guide RNA and Cas9. Nat. Biotechnol. 31: 688-691.
26. Lieber MR. (2010) The mechanism of double-strand DNA break repair by the nonhomologous DNA end-joining pathway. Annu Rev Biochem. 79:181-211.
27. Lin S, George WL, Splittstoesser WE (1984) Expression and inheritance of parthenocarpy in ‘Severianin’ tomato. J Hered 75 (1): 62-66.
28. Lippman Z, Tanksley SD. (2001) Dissecting the genetic pathway to extreme fruit size in tomato using a cross between the small-fruited wild species Lycopersicon pimpinellifolium and L. esculentum var. Giant Heirloom. Genetics. 158: 413-422.
29. Mazzucato A, Taddei AR, Soressi GP (1998) The parthenocarpic fruit (pat) mutant of tomato (Lycopersicon esculentum Mill.) sets seedless fruits and has aberrant anther and ovule development. Development 125:107-114.
30. Michelmore RW, Paran I, Kesseli RV. (1991) Identification of markers linked to disease-resistance genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by using segregating populations. Proc Natl Acad Sci U S A. 88(21):9828-9832.
31. Nuez F, Costa J, Cuartero J (1986) Genetics of the parthenocarpy for tomato varieties “Sub-Artic Plenty,” “75/59” and “Severianin.”. Z Pflanzenzucht 96:200-206.
32. Pan, I.L., McQuinn, R., Giovannoni, J.J. and Irish, V.F. (2010) Functional diversification of AGAMOUS lineage genes in regulating tomato flower and fruit development. J. Exp. Bot. 61, 1795-1806.
33. Pandolfini T, Rotino GL, Camerini S, Defez R, Spena A. (2002) Optimisation of transgene action at the post-transcriptional level: high quality parthenocarpic fruits in industrial tomatoes. BMC Biotechnol. 2:1
34. Anderson P.(1995) Mutagenesis. Methods Cell Biol.;48:31-58. Review.
35. Pattison RJ, Csukasi F, Zheng Y, Fei Z, van der Knaap E, Catala C. (2015) Comprehensive tissue-specific transcriptome analysis reveals distinct regulatory programs during early tomato fruit development. Plant Physiol. 168:1684-1701.
36. Perry FE. (1915) The inheritance of size in tomatoes. The Ohio Naturalist 15: 473-497.
37. Philouze J, Maisonneuve B (1978b) Heredity of the natural ability to set parthenocarpic fruits in a German line. Tomato Genet. Coop. 28: 12.
38. Philouze J. (1989) Natural parthenocarpy in tomato. IV. A study of the polygenic control of parthenocarpy in line 75/59. Agronomie 9:63-75.
39. Picken AJF (1984) A review of pollination and fruit set in the tomato (Lycopersicon esculentum Mill.). J Hort Sci 59:1- 13.
40. Ruan YL, Patrick JW, Bouzayen M, Osorio S, Fernie AR. (2012) Molecular regulation of seed and fruit set. Trends Plant Sci. 17:656-665.
41. Sato S, Kamiyama M, Iwata T, Makita N, Furukawa H, Ikeda H. (2006) Moderate increase of mean daily temperature adversely affects fruit set of Lycopersicon esculentum by disrupting specific physiological processes in male reproductive development. Ann Bot (Lond). 97:731-738.
42. Shan Q, Wang Y, Li J, Zhang Y, Chen K, Liang Z, Zhang K, Liu J, Xi JJ, Qiu JL , Gao C. (2013) Targeted genome modification of crop plants using a CRISPR-Cas system. Nat Biotechnol. 31(8):686-688.
43. Shinozaki Y, Ezura K. (2016) Tomato fruit set and its modification using molecular breeding techniques. In: Functional Genomics and Biotechnology in Solanaceae and Cucurbitaceae Crops (H. Ezura et al. eds.), Biotechnology in Agriculture and Forestry 70. Springer-Verlag Berlin Heidelberg. pp.93-112.
44. Smaczniak C, Immink RG, Angenent GC, Kaufmann K. (2012) Developmental and evolutionary diversity of plant MADS-domain factors: insights from recent studies. Development. 139:3081-3098.
45. Tang N, Deng W, Hu G, Hu N, Li Z. (. 2015) Transcriptome profiling reveals the regulatory mechanism underlying pollination dependent and parthenocarpic fruit set mainly mediated by auxin and gibberellin. PLoS One 24;10(4):e0125355.
46. Vardy E, Lapushner D, Genizi D, Hewitt J (1989a) Genetics of parthenocarpy in tomato under a low temperature regime: I. Line RP75/59. Euphytica 41:1-8.
47. Vardy E, Lapushner D, Genizi D, Hewitt J (1989b) Genetics of parthenocarpy in tomato under a low temperature regime: II. Cultivar "Severianin". Euphytica 41:
9-15.
48. Varga A, Bruinsma J (1986) Tomato. In: CRC Handbook of Fruit Set and Development (Ed. Monselise SP) CRC Press Inc. Boca Raton, FL. pp. 461-481.
49. Varga A, Bruinsma J. (1976) Roles of seeds and auxins in tomato fruit growth. Zeitschrift fur Pflanzenphysiologie 1976;80:95-104.
50. Varoquaux F, Blanvillain R, Delseny M, Gallois P. (2000) Less is better: new approaches for seedless fruit production. Trends Biotechnol. 18(6):233-242. Review.
51. Waibel F. Filipowicz W. (1990) U6 snRNA genes of Arabidopsis are transcribed by RNA polymerase III but contain the same two upstream promoter elements as RNA polymerase II-transcribed U-snRNA genes. Nucleic Acids Res., 18:3451-3458.
52. Wang H, Jones B, Li Z, Frasse P, Delalande C, Regad F, Chaabouni S, Latche A, Pech JC, Bouzayen M. (2005) The tomato Aux/IAA transcription factor IAA9 is involved in fruit development and leaf morphogenesis. Plant Cell. 17(10):2676-2692.
53. Wang H, Schauer N, Usadel B, Frasse P, Zouine M, Hernould M, Latche A,
54. Wang, P., Liao, H., Zhang, W., Yu, X., Zhang, R., Shan, H., Duan, X., Yao, X. and Kong, H. (2015) Flexibility in the structure of spiral flowers and its underlying mechanisms. Nat. Plants, 2, 15188.
55. Pech JC, Fernie AR, Bouzayen M. (2009) Regulatory features underlying pollination-dependent and -independent tomato fruit set revealed by transcript and primary metabolite profiling. Plant Cell. 21:1428-1452.
56. Ye J, Parra EJ, Sosnoski DM, Hiester K, Underhill PA, Shriver MD. (2002) Melting curve SNP (McSNP) genotyping: a useful approach for diallelic genotyping in forensic science. J Forensic Sciences 47: 593-600.
57. Erickson AN,. Markhart AH (2002) Flower developmental stage and organ sensitivity of bell pepper (Capsicum annuum L.) to elevated temperature. Plant Cell Environment 25(1): 123-130
58. Kawasaki Y (2015) Fruit set and temperature stress. In: Abiotic stress biology in horticultural plants. (Y. Kanayama, A. Kochetov, eds.), Springer Japan. Pp. 87-99.
59. Karapanos IC, Mahmood S, Thanopoulos C (2008) Fruit Set in Solanaceous Vegetable Crops as Affected by Floral and Environmental Factors. The European Journal of Plant Science and Biotechnology 2 (Special Issue 1), 88-105.
60. Till BJ, Reynolds SH, Greene EA, Codomo CA, Enns LC, Johnson JE, Burtner C, Odden AR, Young K, Taylor NE, Henikoff JG, Comai L, Henikoff S. (2003) Large-scale discovery of induced point mutations with high-throughput TILLING. Genome Res. 13(3):524-530
62. Selleri, L. (2011) Ph. D. Thesis www.hdl.handle.net/2067/2501
Bassel, G.W., Mullen, R.T. and Bewley, J.D. (2008) procera is a putative DELLA mutant in tomato (Solanum lycopersicum): effects on the seed and vegetative plant. J. Exp. Bot. 59, 585-593.
63. Mazzucato, A., Cellini,F., Bouzayen, M., Zouine, M., Mila, I., Minoia, S., Petrozza, A., Maurizio, E. Picarella, M.E., Ruiu, F. and. Carriero, F. (2015) A TILLING allele of the tomato Aux/IAA9 gene offers new insights into fruit set mechanisms and perspectives for breeding seedless tomatoes. Mol. Breeding, 35, 22.
64. Saito, T., Ariizumi, T., Okabe, Y., Asamizu, E., Hiwasa-Tanase, K., Fukuda, N., Mizoguchi, T., Yamazaki, Y., Aoki, K. and Ezura, H. (2011) TOMATOMA: a novel tomato mutant database distributing Micro-Tom mutant collections. Plant Cell Physiol. 52, 283-896.
65. Carrera, E., Ruiz-Rivero, O., Peres, L/E., Atares, A. and Garcia-Martinez, J.L. (2012) Characterization of the procera tomato mutant shows novel functions of the SlDELLAprotein in the control of flower morphology, cell division and expansion, and the auxin-signaling pathway during fruit-set and development. Plant Physiol. 160, 1581-1596.
66. Zhang, J., Nallamilli, B.R., Mujahid, H. and Peng, Z. (2010) OsMADS6 plays an essential role in endosperm nutrient accumulation and is subject to epigenetic regulation in rice (Oryza sativa). Plant J. 64, 604-617
67. Duan, Y., Xing, Z., Diao, Z., Xu, W., Li, S., Du, X., Wu, G., Wang, C., Lan, T., Meng, Z., Liu, H., Wang, F., Wu, W. and Xue, Y. (2012) Characterization of Osmads6-5, a null allele, reveals that OsMADS6 is a critical regulator for early flower development in rice (Oryza sativaL.). Plant Mol. Biol. 80, 429-442.
配列番号2: SlAG6 mRNA配列
配列番号3: SlAG6アミノ酸配列
配列番号5: AGL6コード配列
配列番号6: AGL6タンパク質のアミノ酸配列
配列番号7: AGL6遺伝子配列
配列番号8: AGL6コード配列
配列番号9: AGL6タンパク質のアミノ酸配列
配列番号10: 1本鎖DNAオリゴヌクレオチド
配列番号11: 1本鎖DNAオリゴヌクレオチド
配列番号12: AclI制限酵素部位
配列番号13: 1本鎖DNAオリゴヌクレオチド
配列番号14: 1本鎖DNAオリゴヌクレオチド
配列番号15: 1本鎖DNAオリゴヌクレオチド
配列番号16: 1本鎖DNAオリゴヌクレオチド
配列番号17: Capana01g00134遺伝子のCRISPRに基づく遺伝子編集の標的として使用可能な例示的な配列
配列番号18: Sme2.5_06058.1遺伝子のCRISPRに基づく遺伝子編集の標的として使用可能な例示的な配列
配列番号19: 1本鎖DNAオリゴヌクレオチド
配列番号20: 1本鎖DNAオリゴヌクレオチド
Claims (25)
- 条件的単為結実を示し、果実収量の少なくとも80%にホメオティック遺伝子異常がなく、且つAGL6遺伝子のコード配列中に機能喪失変異を含む、トマト植物。
- 下記特性の少なくとも1種をさらに示す、請求項1に記載のトマト植物。
(i)非単為結実トマトの受精可能条件下において、植物当たりの果実収量が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの果実収量と同一またはそれ以上である、
(ii)非単為結実トマトの受精可能条件下において、植物当たりの平均果実重量が、同じ遺伝的バックグラウンドの非単為結実トマトの植物当たりの平均果実重量と同一またはそれ以上である、
(iii)ゼリー部を含む、
(iv)種なし果実内の肥大した胚珠が非受精子房から発生したものである。 - 前記果実収量が、成熟果実収量である、請求項1または2に記載のトマト植物。
- 加工用のトマトである、請求項1または2に記載のトマト植物。
- 有限成長型のトマトである、請求項1または2に記載のトマト植物。
- 無限成長型のトマトである、請求項1または2に記載のトマト植物。
- 半有限成長型のトマトである、請求項1または2に記載のトマト植物。
- 選良品種である、請求項1~7のいずれか一項に記載のトマト植物。
- トランスジェニックである、請求項1~8のいずれか一項に記載のトマト植物。
- 前記トマト植物が、リコペルシコン・エスカレンタム、リコペルシコン・セラシフォルメ、リコペルシコン・ピムピネリフォリウム、リコペルシコン・チーズマニイ、リコペルシコン・パルビフロルム、リコペルシコン・クミエレウスキィ、リコペルシコン・ヒルストゥム、リコペルシコン・ペネリィ、リコペルシコン・ペルビアヌム、リコペルシコン・チレンセおよびソラヌム・リコペルシコイデスからなる群から選択される種である、請求項4~8のいずれか一項に記載のトマト植物。
- 前記トマト植物が、1果房1果のトマト、分枝状のトマトおよびチェリートマトからなる群から選択される、請求項4~8および10のいずれか一項に記載のトマト植物。
- 前記条件的単為結実が、熱ストレス下または冷ストレス下で生じる、請求項1~11のいずれか一項に記載のトマト植物。
- 自殖系統である、請求項1~12のいずれか一項に記載のトマト植物。
- 前記機能喪失変異が、ホモ接合型である、請求項1~13のいずれか一項に記載のトマト植物。
- AGL6遺伝子の発現を抑制するためのサイレンシング作用因子を含む、請求項1~13のいずれか一項に記載のトマト植物。
- メガヌクレアーゼ、RNAガイドDNAエンドヌクレアーゼ、ジンクフィンガーヌクレアーゼおよびTALENからなる群から選択されるヌクレアーゼを外因的に発現する、請求項14に記載のトマト植物。
- 請求項1~16のいずれか一項に記載のトマト植物の果実。
- 請求項1~16のいずれか一項に記載のトマト植物の種子。
- 請求項1に記載のトマト植物の交雑種子。
- 請求項1~16のいずれか一項に記載のトマト植物またはその果実の、食用加工品。
- トマトペースト、ケチャップ、トマトソース、トマトスープ、トマトジュース、トマトパウダー、ダイストマト、クラッシュトマト、チョップドトマトおよびトマト濃縮物からなる群から選択される、請求項20に記載の加工品。
- 請求項1~16のいずれか一項に記載のトマト植物の製造方法であって、前記トマト植物のAGL6遺伝子の発現または活性を下方制御し、前記トマト植物のAGL6遺伝子内の変異を検出することを含む、方法。
- 前記下方制御を、前記トマト植物のRNAサイレンシング作用因子を用いた処理によって実施する、請求項22に記載の方法。
- 前記下方制御を、前記トマト植物のDNA編集因子を用いた処理によって実施する、請求項22に記載の方法。
- 請求項1~16のいずれか一項に記載のトマト植物を自家受粉または交雑させることを含む、育種方法。
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Family Cites Families (59)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
NL154600B (nl) | 1971-02-10 | 1977-09-15 | Organon Nv | Werkwijze voor het aantonen en bepalen van specifiek bindende eiwitten en hun corresponderende bindbare stoffen. |
NL154598B (nl) | 1970-11-10 | 1977-09-15 | Organon Nv | Werkwijze voor het aantonen en bepalen van laagmoleculire verbindingen en van eiwitten die deze verbindingen specifiek kunnen binden, alsmede testverpakking. |
NL154599B (nl) | 1970-12-28 | 1977-09-15 | Organon Nv | Werkwijze voor het aantonen en bepalen van specifiek bindende eiwitten en hun corresponderende bindbare stoffen, alsmede testverpakking. |
US3901654A (en) | 1971-06-21 | 1975-08-26 | Biological Developments | Receptor assays of biologically active compounds employing biologically specific receptors |
US3853987A (en) | 1971-09-01 | 1974-12-10 | W Dreyer | Immunological reagent and radioimmuno assay |
US3867517A (en) | 1971-12-21 | 1975-02-18 | Abbott Lab | Direct radioimmunoassay for antigens and their antibodies |
NL171930C (nl) | 1972-05-11 | 1983-06-01 | Akzo Nv | Werkwijze voor het aantonen en bepalen van haptenen, alsmede testverpakkingen. |
US3850578A (en) | 1973-03-12 | 1974-11-26 | H Mcconnell | Process for assaying for biologically active molecules |
US3935074A (en) | 1973-12-17 | 1976-01-27 | Syva Company | Antibody steric hindrance immunoassay with two antibodies |
US3996345A (en) | 1974-08-12 | 1976-12-07 | Syva Company | Fluorescence quenching with immunological pairs in immunoassays |
US4034074A (en) | 1974-09-19 | 1977-07-05 | The Board Of Trustees Of Leland Stanford Junior University | Universal reagent 2-site immunoradiometric assay using labelled anti (IgG) |
US3984533A (en) | 1975-11-13 | 1976-10-05 | General Electric Company | Electrophoretic method of detecting antigen-antibody reaction |
US4038424A (en) | 1976-01-22 | 1977-07-26 | Swift & Company | Vegetable sauce composition |
US4098876A (en) | 1976-10-26 | 1978-07-04 | Corning Glass Works | Reverse sandwich immunoassay |
CA1100351A (en) | 1978-01-31 | 1981-05-05 | House Of Paris Pate Inc.(The)/La Maison Paris Pate Inc. | Food spread product and process for its preparation |
US4879219A (en) | 1980-09-19 | 1989-11-07 | General Hospital Corporation | Immunoassay utilizing monoclonal high affinity IgM antibodies |
CA1192510A (en) | 1981-05-27 | 1985-08-27 | Lawrence E. Pelcher | Rna plant virus vector or portion thereof, a method of construction thereof, and a method of producing a gene derived product therefrom |
JPS6054684A (ja) | 1983-09-05 | 1985-03-29 | Teijin Ltd | 新規dνa及びハイブリツドdνa |
US4556576A (en) | 1984-03-02 | 1985-12-03 | Campbell Soup Company | Process for preparing tomato products of increased consistency |
US5011771A (en) | 1984-04-12 | 1991-04-30 | The General Hospital Corporation | Multiepitopic immunometric assay |
US4666828A (en) | 1984-08-15 | 1987-05-19 | The General Hospital Corporation | Test for Huntington's disease |
CA1288073C (en) | 1985-03-07 | 1991-08-27 | Paul G. Ahlquist | Rna transformation vector |
US4683202A (en) | 1985-03-28 | 1987-07-28 | Cetus Corporation | Process for amplifying nucleic acid sequences |
US4801531A (en) | 1985-04-17 | 1989-01-31 | Biotechnology Research Partners, Ltd. | Apo AI/CIII genomic polymorphisms predictive of atherosclerosis |
GB8608850D0 (en) | 1986-04-11 | 1986-05-14 | Diatech Ltd | Packaging system |
JPS6314693A (ja) | 1986-07-04 | 1988-01-21 | Sumitomo Chem Co Ltd | 植物ウイルスrnaベクタ− |
DE3850683T2 (de) | 1987-02-09 | 1994-10-27 | Lubrizol Genetics Inc | Hybrides RNS-Virus. |
US5272057A (en) | 1988-10-14 | 1993-12-21 | Georgetown University | Method of detecting a predisposition to cancer by the use of restriction fragment length polymorphism of the gene for human poly (ADP-ribose) polymerase |
US5302523A (en) | 1989-06-21 | 1994-04-12 | Zeneca Limited | Transformation of plant cells |
US5192659A (en) | 1989-08-25 | 1993-03-09 | Genetype Ag | Intron sequence analysis method for detection of adjacent and remote locus alleles as haplotypes |
US5035909A (en) | 1990-08-07 | 1991-07-30 | Imit, A.C. | Process for dehydrating tomatoes |
US5281521A (en) | 1992-07-20 | 1994-01-25 | The Trustees Of The University Of Pennsylvania | Modified avidin-biotin technique |
US20030217391A1 (en) | 1996-12-06 | 2003-11-20 | Western Seed Espana | Seedless tomato |
WO1998049888A1 (en) | 1997-05-06 | 1998-11-12 | Kansas State University Research Foundation | Transgenic seedless fruit and methods |
US6774279B2 (en) | 1997-05-30 | 2004-08-10 | Carnegie Institution Of Washington | Use of FLP recombinase in mice |
US5914146A (en) | 1997-10-31 | 1999-06-22 | Vallejo; Bonnie Kay | Method of preparing a salsa product |
ES2239482T3 (es) | 1998-12-01 | 2005-09-16 | Sudzucker Aktiengesellschaft Mannheim/Ochsenfurt | Procedimiento para reforzar la textura pulposa o granulosa de alimentos. |
US6689279B1 (en) | 2000-09-05 | 2004-02-10 | Elizabeth F. Train | Device for separating and dispensing high viscosity fluid from low viscosity fluids |
IL138515A (en) | 2000-09-17 | 2003-06-24 | Lycored Natural Prod Ind Ltd | Tomato fiber composition, process for its preparation and uses thereof |
US6576284B1 (en) | 2001-04-27 | 2003-06-10 | Mcpeek Ruth Ann | Tomato-based caramelized sauce and process for making same |
US6586662B2 (en) | 2001-06-13 | 2003-07-01 | California Hybrids | Yellow, determinate tomatoes suitable for processing |
US6869634B2 (en) | 2001-06-18 | 2005-03-22 | Naomi H. Lincoln | Barbecue sauce and process for making same |
IL160906A0 (en) | 2001-09-19 | 2004-08-31 | Conagra Grocery Prod Co | System and method for producing concentrated food products with fractionation concentration |
US20030232410A1 (en) | 2002-03-21 | 2003-12-18 | Monika Liljedahl | Methods and compositions for using zinc finger endonucleases to enhance homologous recombination |
CA2497913C (en) | 2002-09-05 | 2014-06-03 | California Institute Of Technology | Use of chimeric nucleases to stimulate gene targeting |
WO2005080571A1 (en) | 2004-02-02 | 2005-09-01 | Dongbu Hannong Chemical Co., Ltd. | The usage of mad-box genes in fruit & seed development by regulating active gibberellin synthesis |
US20060014264A1 (en) | 2004-07-13 | 2006-01-19 | Stowers Institute For Medical Research | Cre/lox system with lox sites having an extended spacer region |
EP2360244B1 (en) | 2005-10-18 | 2014-12-24 | Precision Biosciences | Rationally-designed meganucleases with altered sequence specificity and DNA-binding affinity |
EP1992218A1 (en) | 2007-05-18 | 2008-11-19 | De Ruiter Seeds R&D B.V. | Parthenocarpic genetic elements derived from S. habrochaites |
EP2067402A1 (en) | 2007-12-07 | 2009-06-10 | Max Delbrück Centrum für Molekulare Medizin (MDC) Berlin-Buch; | Transponson-mediated mutagenesis in spermatogonial stem cells |
USPP21199P3 (en) | 2008-09-02 | 2010-08-17 | Keisei Rose Nurseries, Inc. | Rose plant named ‘Keishisutoki’ |
JP6208580B2 (ja) | 2010-05-17 | 2017-10-04 | サンガモ セラピューティクス, インコーポレイテッド | 新規のdna結合タンパク質及びその使用 |
NL2005908C2 (en) * | 2010-12-22 | 2012-06-28 | Rijk Zwaan Zaadteelt En Zaadhandel Bv | Fruit formation in the absence of fertilisation. |
WO2013126963A1 (en) | 2012-02-29 | 2013-09-06 | Benitec Biopharma Limited | Pain treatment |
PT3502240T (pt) | 2012-11-27 | 2021-08-11 | Childrens Medical Ct Corp | Elementos reguladores distais de bcl11a como alvo para a reindução de hemoglobina fetal |
US8697359B1 (en) | 2012-12-12 | 2014-04-15 | The Broad Institute, Inc. | CRISPR-Cas systems and methods for altering expression of gene products |
KR20150103280A (ko) | 2013-01-08 | 2015-09-09 | 베니텍 바이오파마 리미티드 | 연령-관련 황반 변성 치료 |
EP3690042A1 (en) | 2014-01-17 | 2020-08-05 | National Research and Development Agency National Agriculture and Food Research Organization | Parthenocarpy regulation gene and use thereof |
CN109068605B (zh) | 2016-01-21 | 2022-11-11 | 以色列国家农业和农村发展农业研究组织沃尔坎尼中心 | 单性结实植物及其产生方法 |
-
2017
- 2017-01-19 CN CN201780018857.8A patent/CN109068605B/zh active Active
- 2017-01-19 US US16/071,097 patent/US11839195B2/en active Active
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- 2018-07-16 IL IL260610A patent/IL260610B/en unknown
-
2021
- 2021-10-07 JP JP2021165483A patent/JP7273919B2/ja active Active
Non-Patent Citations (2)
Title |
---|
Takisawa et al., BMC Plant Biology, 2018, 18:72 |
農業生産技術管理学会誌, 2011, vol.18, no.2, p.67-73 |
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JP2019502392A (ja) | 2019-01-31 |
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CN109068605A (zh) | 2018-12-21 |
MX2018008910A (es) | 2019-03-28 |
BR112018014902A2 (pt) | 2018-12-18 |
JP7273919B2 (ja) | 2023-05-15 |
EP3405024A1 (en) | 2018-11-28 |
IL260610B (en) | 2022-04-01 |
WO2017125931A1 (en) | 2017-07-27 |
US11839195B2 (en) | 2023-12-12 |
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CN109068605B (zh) | 2022-11-11 |
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