Rhizobium radiobacter(
Agrobacterium tumefaciens)
アグロバクテリウム (Agrobacterium) とはグラム陰性菌に属する土壌細菌であるリゾビウム属 (Rhizobium) の内、植物に対する病原性を持つものの総称。特にその中で根頭癌腫病に関連するAgrobacterium tumefaciens（Rhizobium radiobacterの異名）[1]を指すことが多い。かつてアグロバクテリウム属という独立の属が与えられていたが、系統解析の結果多くはリゾビウム属に含まれることがわかり、その他も新設されたルエゲリア属（英語版） (Ruegeria)、シュードロドバクター属（英語版） (Pseudorhodobacter)、スタッピア属（英語版） (Stappia) に分類され、学名としては廃された。このため使用には注意が必要である。しかしながら、アグロバクテリウムという分類は便利なため、分野や用途によってはこの呼称も広く使われている。
アグロバクテリウムは、植物細胞に感染してDNAを送り込む（形質転換）性質があるため、植物のバイオテクノロジーでよく利用される。
目次
- 1 植物に対する病原性
- 2 バイオテクノロジーへの利用
- 3 その他の性質
- 4 脚注
- 5 関連項目
植物に対する病原性
根の大きく成長した部位は
Agrobacterium sp.によって誘導された虫こぶである。
A. tumefaciensは多くの双子葉植物および一部の裸子植物・単子葉植物に虫こぶ様の腫瘍（根元などに生じ、根頭癌腫、クラウンゴール crown gall と呼ばれる）を起こす。この菌はTiプラスミド（pTi: Tiはtumor-inducingの略）と呼ばれる巨大なプラスミドを有しており、その一部であるT-DNA（transfer DNAの意）と呼ばれるDNA断片を植物細胞に注入し、T-DNAは相同組換えにより植物細胞のゲノムに挿入される。ただし、T-DNAの両末端の極めて短い配列とゲノムの配列との相同組換えであるため挿入位置はかなりランダムであり、実質的にはほとんどが非相同組換えによってゲノムに挿入されるといってよい。T-DNAは植物ホルモン（オーキシンとサイトカイニン）を生成する酵素の遺伝子であるiaaM(tms1), iaaH(tms2), iptZ(tmr)を含み、これらによって生産される大量のオーキシンやサイトカイニンにより腫瘍（A. tumefaciensに特有）が形成される。またT-DNAはオパイン (オピン: Opine) と総称される特殊なイミノ酸（アグロバクテリウムは炭素源や窒素源として代謝できるが、他の細菌はほとんど利用できない）を植物に作らせる酵素をコードしている[2]。根粒菌などの窒素固定細菌とは異なりA. tumefaciensは寄生細菌であって、植物にとって利益はない[3]。このA. tumefaciensの性質は「植物に対する遺伝的植民地化」とも喩えられる。
Tiプラスミドは大きい（20万塩基対前後）プラスミドで、T-DNAの他にT-DNAを植物細胞に輸送するのに働く遺伝子群(vir region)やオパインを分解消費するための遺伝子などを持っている。vir regionはvirA, B, G, C, D, Eの６つのオペロンから形成されており、それぞれのオペロンは複数の構造遺伝子を含んでいる。なお、virCオペロンのみ転写方向が他のvirオペロンとは異なる。サクラの木とサクランボの木のクラウンゴールから分離されたTiプラスミド（pTi-SAKURAとpTiC58）の全塩基配列が明らかにされている。最もよく研究されている細胞株はA. tumefaciens C58（サクランボの木のクラウンゴールから分離された）で、Goodnerら[4]とWoodら[5]により同時にゲノムの完全配列が明らかにされた。A. tumefaciens C58のゲノムは環状の染色体、2個の環状プラスミド、および1本の直線状染色体からなる。環状染色体を有する細菌はごく普通だが、それに加え直線状染色体を持つのはアグロバクテリウム属の一部のグループに特有である。2つの環状プラスミドはpTiC58（病原性に関与する[4]）とpAtC58[5]である。pAtC58はオパイン（A. tumefaciens C58が生成するオパインはノパリン〔Nopalin〕と呼ばれる）の代謝に関与し、これはpTiC58がない場合には他の細菌にも転移する[6]。
なお、植物の腫瘍はアグロバクテリウムだけでなくむしろ昆虫（虫こぶ）などによるものが多い。根にこぶを作る病原体には根こぶ病菌（原生生物ネコブカビ）やネコブセンチュウ（線虫）がある。
バイオテクノロジーへの利用
A: Agrobacterium tumefaciens
B: Agrobacteriumゲノム
C: Tiプラスミド : a: T-DNA , b: vir遺伝子群 , c: 複製起点 , d: オパイン異化遺伝子
D: 植物細胞
E: ミトコンドリア
F: 葉緑体
G: 核
T-DNAは植物遺伝子工学の特に有用なベクターであり、アグロバクテリウムのDNA転移能力は植物の核ゲノムに外来遺伝子を導入する（トランスジェニック植物の作出）手段として盛んに利用されている[7]。
具体的には、目的の遺伝子配列と植物での選択マーカー遺伝子をT-DNA内に挿入し、それを植物細胞の核ゲノムに挿入させる。T-DNAの中で植物ゲノムへの挿入に必須なのはT-DNA両端に存在するRB(right border:右境界配列)とLB(left border:左境界配列)とよばれる25塩基対(コンセンサス配列: 5'-TGGCAGGATATATN(C/G)N(G/A)(T/G)TGTAA(A/T)(T/C)-3', NはACGTのいずれでも構わない) )である。RBとLBに挟まれた内側の配列には特異性はなく、どのような配列でも構わない。また、野生型のT-DNA中に存在する腫瘍形成遺伝子群は挿入には必要ないだけでなく植物体再生に悪影響を及ぼす。そこで、腫瘍形成遺伝子群を目的の遺伝子と置き換えれば、目的の遺伝子を植物細胞に導入できるうえに増殖した形質転換細胞が腫瘍を形成することもない。
形質転換植物体を得る方法としては、まず組織または細胞にアグロバクテリウムを感染させ、これを培養して植物体に再生させる方法がある。もう1つの方法としては、花にアグロバクテリウムを感染させ、種子を形成させる方法(フローラル・ディップ(floral dip)法やフローラル・スプレー(floral spray)法)がある。
形質転換植物の作製法の詳細については、遺伝子組換え作物の作製法を参照。
一例としてホタルのルシフェラーゼを用いた「光る植物」の作出にも用いられ、この方法は植物の葉緑体機能の研究やレポーター遺伝子（遺伝子の調節領域の研究用）としての利用に有用である[8]。アグロバクテリウムは自然には双子葉植物などにしか感染しないが、現在ではT-DNAはイネなどの単子葉植物や真菌などでの利用も可能になっている[9]。さらにT-DNAをヒト細胞に転移することも実験的には可能である[10]。
その他の性質
アグロバクテリウムが植物細胞にT-DNAを送り込むメカニズムは、タイプIV分泌系といわれ、ヒト病原菌の多くが細胞にタンパク質などを送り込むメカニズム（タイプIII分泌系）に類似している[11]。また多くのグラム陰性細菌に見られるクオラムセンシング（他の同種菌の分泌物質を感知して同調行動を取るためのシグナル伝達系）を有する。
同じアグロバクテリウム属に属するアグロバクテリウム・リゾゲネス（Agrobacterium rhizogenes、現在の正式な学名はRhizobium rhizogenes）もpTiに相当するプラスミドpRi内にT-DNAをもつが、これは植物に腫瘍でなく不定根を発生させる性質がある。この不定根の形成はひげ毛病とよばれ、高密度に枝分かれした根が大量に増殖するというものである。
脚注
- ^ 現在の正式な学名はRhizobium radiobacterであるが、この中には旧来Agrobacterium radiobacterと呼ばれていた感染性のないものも含まれている。また、A. tumefaciensとされていた株の中には別の種に分類されたものもある。
- ^ Zupan J, Muth TR, Draper O, Zambryski P. (2000). “The transfer of DNA from Agrobacterium tumefaciens into plants: a feast of fundamental insights”. Plant J. 23: 11-28. doi:10.1046/j.1365-313x.2000.00808.x. PMID 10929098.
- ^ Moore LW, Chilton WS, Canfield ML (1997). “Diversity of Opines and Opine-Catabolizing Bacteria Isolated from Naturally Occurring Crown Gall Tumors”. App. Environ. Microbiol. 63: 201-207. PMC 1389099. PMID 16535484. http://aem.asm.org/content/63/1/201.abstract.
- ^ a b Goodner B, Hinkle G, Gattung S, Miller N, et al. (2001). “Genome Sequence of the Plant Pathogen and Biotechnology Agent Agrobacterium tumefaciens C58”. Science 294: 2323-2328. doi:10.1126/science.1066803. PMID 11743194.
- ^ a b Wood DW, Setubal JC, Kaul R, Monks DE, et al. (2001). “The Genome of the Natural Genetic Engineer Agrobacterium tumefaciens C58”. Science 294: 2317-2323. doi:10.1126/science.1066803. PMID 11743193.
- ^ Vaudequin-Dransart V, Petit A, Chilton WS, Dessaux Y (1998). “The cryptic plasmid of Agrobacterium tumefaciens cointegrates with the Ti plasmid and cooperates for opine degradation”. Mol. Plant-microbe Interact. 11: 583-591. doi:10.1094/MPMI.1998.11.7.583.
- ^ Zambryski P, Joos H, Genetello C, Leemans J, Montagu MV, Schell J (1983). “Ti plasmid vector for introduction of DNA into plant cells without alteration of their normal regeneration capacity”. EMBO J. 2: 2143-2150. PMC 555426. PMID 16453482. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=555426.
- ^ Root, M. (1988). “Glow in the dark biotechnology”. Bioscience 38: 745-747. http://www.jstor.org/stable/1310781.
- ^ Hiei Y, Ohta S, Komari T, Kumashiro T (1994). “Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA”. Plant J. 6: 271-82. doi:10.1046/j.1365-313X.1994.6020271.x. PMID 7920717.
- ^ Kunik T, Tzfira T, Kapulnik Y, Gafni Y, Dingwall C, Citovsky V (2001). “Genetic transformation of HeLa cells by Agrobacterium”. Proc. Natl. Acad. Sci, U. S. A. 98: 1871-1876. doi:10.1073/pnas.98.4.1871.
- ^ Li PL, Everhart DM, Farrand SK (1998). “Genetic and sequence analysis of the pTiC58 trb locus, encoding a mating-pair formation system related to members of the type IV secretion family”. J Bacteriol. 180: 6164-6172. PMC 107700. PMID 9829924. http://jb.asm.org/content/180/23/6164.abstract.
関連項目
| Agrobacterium |
|
| Scientific classification |
| Kingdom: |
Bacteria |
| Phylum: |
Proteobacteria |
| Class: |
Alpha Proteobacteria |
| Order: |
Rhizobiales |
| Family: |
Rhizobiaceae |
| Genus: |
Agrobacterium |
| Type species |
Agrobacterium tumefaciens
(Smith and Townsend 1907) Conn 1942 |
| Species |
- ‘Agrobacterium albertimagni’ Salmassi et al. 2002
- Agrobacterium larrymoorei Bouzar and Jones 2001
- Agrobacterium nepotum (Puławska et al. 2012) Mousavi et al. 2015
- Agrobacterium pusense (Panday et al. 2011) Mousavi et al. 2015
- Agrobacterium radiobacter (Beijerinck and van Delden 1902) Conn 1942
- Agrobacterium rubi (Hildebrand 1940) Starr and Weiss 1943
- Agrobacterium skierniewicense (Puławska et al. 2012) Mousavi et al. 2015
- Agrobacterium tumefaciens (Smith and Townsend 1907) Conn 1942
|
| Synonyms |
|
|
Agrobacterium is a genus of Gram-negative bacteria established by H. J. Conn that uses horizontal gene transfer to cause tumors in plants. Agrobacterium tumefaciens is the most commonly studied species in this genus. Agrobacterium is well known for its ability to transfer DNA between itself and plants, and for this reason it has become an important tool for genetic engineering.
The Agrobacterium genus is quite heterogeneous. Recent taxonomic studies have reclassified all of the Agrobacterium species into new genera, such as Ahrensia, Pseudorhodobacter, Ruegeria, and Stappia,[1][2] but most species have been controversially reclassified as Rhizobium species.[3][4][5]
Contents
- 1 Plant pathogen
- 2 In humans
- 3 Uses in biotechnology
- 4 Genomics
- 5 History
- 6 See also
- 7 References
- 8 External links
Plant pathogen
The large growths on these roots are galls induced by
Agrobacterium sp.
A. tumefaciens causes crown-gall disease in plants. The disease is characterised by a tumour-like growth or gall on the infected plant, often at the junction between the root and the shoot. Tumors are incited by the conjugative transfer of a DNA segment (T-DNA) from the bacterial tumour-inducing (Ti) plasmid. The closely related species, A. rhizogenes, induces root tumors, and carries the distinct Ri (root-inducing) plasmid. Although the taxonomy of Agrobacterium is currently under revision it can be generalised that 3 biovars exist within the genus, A. tumefaciens, A. rhizogenes, and A. vitis. Strains within A. tumefaciens and A. rhizogenes are known to be able to harbour either a Ti or Ri-plasmid, whilst strains of A. vitis, generally restricted to grapevines, can harbour a Ti-plasmid. Non-Agrobacterium strains have been isolated from environmental samples which harbour a Ri-plasmid whilst laboratory studies have shown that non-Agrobacterium strains can also harbour a Ti-plasmid. Some environmental strains of Agrobacterium possess neither a Ti nor Ri-plasmid. These strains are avirulent.[6]
The plasmid T-DNA is integrated semi-randomly into the genome of the host cell,[7] and the tumor morphology genes on the T-DNA are expressed, causing the formation of a gall. The T-DNA carries genes for the biosynthetic enzymes for the production of unusual amino acids, typically octopine or nopaline. It also carries genes for the biosynthesis of the plant hormones, auxin and cytokinins, and for the biosynthesis of opines, providing a carbon and nitrogen source for the bacteria that most other micro-organisms can't use, giving Agrobacterium a selective advantage.[8] By altering the hormone balance in the plant cell, the division of those cells cannot be controlled by the plant, and tumors form. The ratio of auxin to cytokinin produced by the tumor genes determines the morphology of the tumor (root-like, disorganized or shoot-like).
In humans
Although generally seen as an infection in plants, Agrobacterium can be responsible for opportunistic infections in humans with weakened immune systems,[9][10] but has not been shown to be a primary pathogen in otherwise healthy individuals. One of the earliest associations of human disease caused by Agrobacterium radiobacter was reported by Dr. J. R. Cain in Scotland (1988).[11] A later study suggested that Agrobacterium attaches to and genetically transforms several types of human cells by integrating its T-DNA into the human cell genome. The study was conducted using cultured human tissue and did not draw any conclusions regarding related biological activity in nature.[12]
Uses in biotechnology
See also: horizontal gene transfer
The ability of Agrobacterium to transfer genes to plants and fungi is used in biotechnology, in particular, genetic engineering for plant improvement. A modified Ti or Ri plasmid can be used. The plasmid is 'disarmed' by deletion of the tumor inducing genes; the only essential parts of the T-DNA are its two small (25 base pair) border repeats, at least one of which is needed for plant transformation.[13][14] The genes to be introduced into the plant are cloned into a plant transformation vector that contains the T-DNA region of the disarmed plasmid, together with a selectable marker (such as antibiotic resistance) to enable selection for plants that have been successfully transformed. Plants are grown on media containing antibiotic following transformation, and those that do not have the T-DNA integrated into their genome will die. An alternative method is agroinfiltration.[15][16]
Plant (
S. chacoense) transformed using
Agrobacterium. Transformed cells start forming calluses on the side of the leaf pieces
Transformation with Agrobacterium can be achieved in two ways. Protoplasts or alternatively leaf-discs can be incubated with the Agrobacterium and whole plants regenerated using plant tissue culture. A common transformation protocol for Arabidopsis is the floral-dip method:[17] inflorescence are dipped in a suspension of Agrobacterium, and the bacterium transforms the germline cells that make the female gametes. The seeds can then be screened for antibiotic resistance (or another marker of interest), and plants that have not integrated the plasmid DNA will die when exposed to the correct condition of antibiotic.[15]
Agrobacterium does not infect all plant species, but there are several other effective techniques for plant transformation including the gene gun.
Agrobacterium is listed as being the vector of genetic material that was transferred to these USA GMOs:[18]
- Soybean
- Cotton
- Corn
- Sugar Beet
- Alfalfa
- Wheat
- Rapeseed Oil (Canola)
- Creeping bentgrass (for animal feed)
- Rice (Golden Rice)
Genomics
The sequencing of the genomes of several species of Agrobacterium has permitted the study of the evolutionary history of these organisms and has provided information on the genes and systems involved in pathogenesis, biological control and symbiosis. One important finding is the possibility that chromosomes are evolving from plasmids in many of these bacteria. Another discovery is that the diverse chromosomal structures in this group appear to be capable of supporting both symbiotic and pathogenic lifestyles. The availability of the genome sequences of Agrobacterium species will continue to increase, resulting in substantial insights into the function and evolutionary history of this group of plant-associated microbes.[19]
History
Marc Van Montagu and Jozef Schell at the University of Ghent (Belgium) discovered the gene transfer mechanism between Agrobacterium and plants, which resulted in the development of methods to alter Agrobacterium into an efficient delivery system for gene engineering in plants.[13][14] A team of researchers led by Dr Mary-Dell Chilton were the first to demonstrate that the virulence genes could be removed without adversely affecting the ability of Agrobacterium to insert its own DNA into the plant genome (1983).
See also
- Agroinfiltration
- Marc Van Montagu
- Rhizobium rhizogenes (formerly Agrobacterium rhizogenes)
References
- ^ Uchino, Yoshihito; Yokota, Akira; Sugiyama, Junta (1997). "Phylogenetic position of the marine subdivision of Agrobacterium species based on 16S rRNA sequence analysis". The Journal of General and Applied Microbiology. 43 (4): 243–247. doi:10.2323/jgam.43.243. PMID 12501326.
- ^ Uchino, Yoshihito; Hirata, Aiko; Yokota, Akira; Sugiyama, Junta (1998). "Reclassification of marine Agrobacterium species: Proposals of Stappia stellulata gen. nov., comb. Nov., Stappia aggregata sp. nov., nom. Rev., Ruegeria atlantica gen. nov., comb. Nov., Ruegeria gelatinovora comb. Nov., Ruegeria algicola comb. Nov., and Ahrensia kieliense gen. nov., sp. nov., nom. Rev". The Journal of General and Applied Microbiology. 44 (3): 201–210. doi:10.2323/jgam.44.201. PMID 12501429.
- ^ Young, J. M.; Kuykendall, L. D.; Martínez-Romero, E; Kerr, A; Sawada, H (2001). "A revision of Rhizobium Frank 1889, with an emended description of the genus, and the inclusion of all species of Agrobacterium Conn 1942 and Allorhizobium undicola de Lajudie et al. 1998 as new combinations: Rhizobium radiobacter, R. rhizogenes, R. rubi, R. undicola, and R. vitis". Int J Syst Evol Microbiol. 51 (Pt 1): 89–103. doi:10.1099/00207713-51-1-89. PMID 11211278.
- ^ Farrand, S. K.; Van Berkum, P. B.; Oger, P (2003). "Agrobacterium is a definable genus of the family Rhizobiaceae". International Journal of Systematic and Evolutionary Microbiology. 53 (5): 1681–7. doi:10.1099/ijs.0.02445-0. PMID 13130068.
- ^ Young, J. M.; Kuykendall, L. D.; Martínez-Romero, E; Kerr, A; Sawada, H (2003). "Classification and nomenclature of Agrobacterium and Rhizobium—a reply to Farrand et al. (2003)". International Journal of Systematic and Evolutionary Microbiology. 53 (5): 1689–95. doi:10.1099/ijs.0.02762-0. PMID 13130069.
- ^ Sawada, H.; Ieki, H.; Oyaizu, H.; Matsumoto, S. (1993). "Proposal for Rejection of Agrobacterium tumefaciens and Revised Descriptions for the Genus Agrobacterium and for Agrobacterium radiobacter and Agrobacterium rhizogenes". International Journal of Systematic Bacteriology. 43 (4): 694–702. doi:10.1099/00207713-43-4-694. PMID 8240952.
- ^ Francis, Kirk E.; Spiker, Steven (2004). "Identification of Arabidopsis thaliana transformants without selection reveals a high occurrence of silenced T-DNA integrations". The Plant Journal. 41 (3): 464–77. doi:10.1111/j.1365-313X.2004.02312.x. PMID 15659104.
- ^ Pitzschke, Andrea; Hirt, Heribert (2010). "New insights into an old story: Agrobacterium-induced tumour formation in plants by plant transformation". The EMBO Journal. 29 (6): 1021–32. doi:10.1038/emboj.2010.8. PMC 2845280 . PMID 20150897.
- ^ Hulse, M.; Johnson, S.; Ferrieri, P. (1993). "Agrobacterium Infections in Humans: Experience at One Hospital and Review". Clinical Infectious Diseases. 16 (1): 112–7. doi:10.1093/clinids/16.1.112. PMID 8448285.
- ^ Dunne Jr, W. M.; Tillman, J; Murray, J. C. (1993). "Recovery of a strain of Agrobacterium radiobacter with a mucoid phenotype from an immunocompromised child with bacteremia". Journal of clinical microbiology. 31 (9): 2541–3. PMC 265809 . PMID 8408587.
- ^ Cain, John Raymond (1988). "A case of septicaemia caused by Agrobacterium radiobacter". Journal of Infection. 16 (2): 205–6. doi:10.1016/s0163-4453(88)94272-7. PMID 3351321.
- ^ Kunik, T.; Tzfira, T; Kapulnik, Y; Gafni, Y; Dingwall, C; Citovsky, V (2001). "Genetic transformation of HeLa cells by Agrobacterium". Proceedings of the National Academy of Sciences. 98 (4): 1871–6. Bibcode:2001PNAS...98.1871K. doi:10.1073/pnas.041327598. JSTOR 3054968. PMC 29349 . PMID 11172043.
- ^ a b Schell, J.; Van Montagu, M. (1977). "The Ti-Plasmid of Agrobacterium Tumefaciens, A Natural Vector for the Introduction of NIF Genes in Plants?". In Hollaender, Alexander; Burris, R. H.; Day, P. R.; Hardy, R. W. F.; Helinski, D. R.; Lamborg, M. R.; Owens, L.; Valentine, R. C. Genetic Engineering for Nitrogen Fixation. Basic Life Sciences. 9. pp. 159–79. doi:10.1007/978-1-4684-0880-5_12. ISBN 978-1-4684-0882-9. PMID 336023.
- ^ a b Joos, H; Timmerman, B; Montagu, M. V.; Schell, J (1983). "Genetic analysis of transfer and stabilization of Agrobacterium DNA in plant cells". The EMBO Journal. 2 (12): 2151–60. PMC 555427 . PMID 16453483.
- ^ a b Thomson JA. "Genetic Engineering of Plants" (PDF). Biotechnology. Encyclopedia of Life Support Systems. 3. Retrieved 17 July 2016.
- ^ Leuzinger K, Dent M, Hurtado J, Stahnke J, Lai H, Zhou X, Chen Q (2013). "Efficient Agroinfiltration of Plants for High-level Transient Expression of Recombinant Proteins". Journal of Visualized Experiments. 77 (50521). doi:10.3791/50521. PMC 3846102 . PMID 23913006.
- ^ Clough, Steven J.; Bent, Andrew F. (1998-12-01). "Floral dip: a simplified method forAgrobacterium-mediated transformation ofArabidopsis thaliana". The Plant Journal. 16 (6): 735–743. doi:10.1046/j.1365-313x.1998.00343.x. ISSN 1365-313X. PMID 10069079.
- ^ The FDA List of Completed Consultations on Bioengineered Foods Archived May 13, 2008, at the Wayback Machine.
- ^ Setubal, Joao C.; Wood, Derek; Burr, Thomas; Farrand, Stephen K.; Goldman, Barry S.; Goodner, Brad; Otten, Leon; Slater, Steven (2009). "The Genomics of Agrobacterium: Insights into its Pathogenicity, Biocontrol, and Evolution". In Jackson, Robert W. Plant Pathogenic Bacteria: Genomics and Molecular Biology. Caister Academic Press. pp. 91–112. ISBN 978-1-904455-37-0.
External links
- Kyndt, Tina; Quispe, Dora; Zhai, Hong; Jarret, Robert; Ghislain, Marc; Liu, Qingchang; Gheysen, Godelieve; Kreuze, Jan F. (2015). "The genome of cultivated sweet potato contains Agrobacterium T-DNAs with expressed genes: An example of a naturally transgenic food crop". Proceedings of the National Academy of Sciences. 112: 201419685. doi:10.1073/pnas.1419685112. PMC 4426443 . PMID 25902487. Lay summary – Phys.org (April 21, 2015).
- Current taxonomy of Agrobacterium species, and new Rhizobium names
- Agrobacteria is used as gene ferry - Plant transformation with Agrobacterium]
|
Genetic engineering
|
|
Genetically
modified
organisms |
|
Crops
|
|
Maize
|
- MON 810
- MON 863
- StarLink
- List of varieties of genetically modified maize
|
|
|
Potato
|
|
|
|
Rice
|
|
|
|
Soybean
|
- Roundup ready soybean
- Vistive Gold
|
|
|
Tomato
|
|
|
|
Cotton
|
|
|
|
Other
|
- Apple
- Arabidopsis
- Brinjal
- Canola
- Papaya
- Rose
- SmartStax
- Sugar beet
- Tobacco
- Trees
- Wheat
|
|
|
|
Mammals
|
- Mouse
- Enviropig
- Herman the Bull
- Knockout rat
|
|
|
Other animals
|
|
|
|
Bacteria
and viruses
|
- Ice-minus bacteria
- Hepatitis B vaccine
- Oncolytic virus
|
|
|
| Processes |
|
Inserting DNA
|
- Agrobacteria
- Biolistics
- Electroporation
- Genetic transduction
- Lipofection
- Microinjection
- Transfection
|
|
|
Types
|
- Recombinant DNA
- Transgenesis
- Cisgenesis
|
|
|
| Uses |
|
In agriculture
|
- Genetically modified food
- Pharming
- Companies
- BASF
- Bayer
- Dow AgroSciences
- DuPont Pioneer
- Monsanto
- Syngenta
|
|
|
In humans and
diagnostics
|
- Gene therapy
- Genetic enhancement
|
|
|
In research
|
- Gene knockout
- Gene knockdown
- Gene targeting
|
|
|
Related
articles |
- Transgene
- Detection of genetically modified organisms
- Genetic pollution
- Genetic engineering in fiction
- Human enhancement
- Reverse transfection
- Transhumanism
|
|
| Regulation |
- Regulation of the release of GMOs
- Regulation of GMOs in the European Union
- Regulation of GMOs in Switzerland
- Cartagena Protocol on Biosafety
|
|
| Geography |
|
|
| Similar fields |
- Synthetic biology
- Cloning
- Stem cell research
|
|
- Biology
- Genetics
- Biotechnology
- Bioethics
|
