Transgene expression variability (position effect) of CAT and GUS reporter genes driven by linked divergent T-DNA promoters

Amasino RM, Powell ALT, Gordon MP: Changes in T-DNA methylation and expression are associated with phenotypic variation and plant regeneration in a crown gall tumor line. Mol Gen Genet 297: 437?446 (1984).

An G, Ebert PR, Mitra A, Ha JB: Binary vectors. In: Gelvin SB, Schilperoort RA (eds) Plant Molecular Biology Manual, pp A3/1-A3/19. Kluwer Academic Publishers, Dordrecht (1988).

An G, Watson BD, Stachel S, Gordon MP, Nester EW: New cloning vehicles for transformation of higher plants. EMBO J 4: 277?284 (1985).

An G: Development of plant promoter expression vectors and their use for analysis of differential activity of nopaline synthase promoter in transformed tobacco cells. Plant Physiol 81: 86?91 (1986).

Bassiri A, Carlson PS: Isozyme patterns and differences in plant parts and their callus cultures in common bean. Crop Sci 19: 955?958 (1978).

Bennetzen JL: Covalent DNA modification and the regulation of Mutator element transposition in maize. Mol Gen Genet 208: 45?51 (1987).

Bradford MM: A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 72: 248?254 (1976).

Chandler VL, Talbert LE, Raymond F: Sequence, genomic distribution and DNA modification of a Mu1 element from non-mutator maize stocks. Genetics 119: 951?958 (1988).

Chandler VL, Walbot V: DNA modification of a maize transposable element correlates with loss of activity. Proc Natl Acad Sci USA 83: 1767?1771 (1986).

Chomet PS, Wessler S, Dellaporta SL: Inactivation of the maize transposable element Activater (Ac) is associated with its DNA methylation. EMBO J 6: 295?302 (1987).

Coates D, Taliercio EW, Gelvin SB: Chromatin structure of integrated T-DNA in crown gall tumors. Plant Mol Biol 8: 159?168 (1987).

Czernilofsky AP, Hain R, Herrera-Estrella L, Horst L, Goyvaerts E, Baker BJ, Schell J: Fate of selectable marker DNA integrated into the genome ofNicotiana tabacum. DNA 5: 101?113 (1986).

De Almeida ERP, Gossele V, Muller CG, Dockx Jan, Reynaerts A, Botterman J, Krebbers E, Timko MP: Transgenic expression of two marker genes under the control of anArabidopsis rbcS promoter: Sequences encoding the rubisco transit peptide increase expression levels. Mol Gen Genet 218: 78?86 (1989).

Dean C, Favreau M, Tamaki S, Bond-Nutter D, Dunsmuir P, Bedbrook J: Expression of tandem gene fusions in transgenic tobacco plants. Nucl Acids Res 16: 7601?7617 (1988).

Dean C, Jones J, Favreau M, Dunsmuir P, Bedbrook J: Influence of flanking sequences on variability in expression levels of an introduced gene in transgenic tobacco plants. Nucl Acid Res 16: 9267?9282 (1988).

DeRita VJ, Gelvin SB: Deletion analysis of the mannopine synthase gene promoter in sunflower crown gall tumors andAgrobacterium tumefaciens. Mol Gen Genet 207: 233?241 (1987).

Doyle JJ, Doyle JL: Isolation of plant DNA from fresh tissue. Focus 12(1) 13?15 (1990).

Dunsmuir P, Bedbrook J, Bond-Nuttler D, Dean C, Gidoni D, Jones J: The expression of introduced genes in regenerated plants. In: Setlow J, Hollaender A (eds) Genetic Engineering: Principles and Methods, vol. 1 pp. 45?58. Plenum Press, New York (1987).

Fang R-X, Nagy F, Sivasubramaniam S, Chua N-H: Multiplecis regulatory elements for maximal expression of the cauliflower mosaic virus 35S promoter in transgenic plants. Plant Cell 1: 141?150 (1989).

Fluhr R, Kuhlemeier CF, Nagy F, Chua N-H: Organspecific and light-induced expression of plant genes. Science 232: 1106?1112 (1986).

Forrester WC, Takegawa S, Papayannopoulou T, Stamatoyannopoulos G, Groudine M: Evidence for a locus activation region: the formation of developmentally stable hypersensitive sites in globin-expressing hybrids. Nucl Acids Res 15: 10159?10177 (1987).

Gendloff EH, Bowen B, Buchholz WG: Quantitation of chloramphenicol acetyl transferase in transgenic tobacco by ELISA and correlation with gene copy number. Plant Mol Biol 14: 575?583 (1990).

Gidoni D, Bond-Nutter D, Brosio P, Jones J, Bedbrook J, Dunsmuir P: Coordinated expression between two photosynthetic petunia genes in transgenic plants. Mol Gen Genet 211: 507?514 (1988).

Grosvelt F, van Assendelft GB, Greaves DR, Kollias G: Position-independent, high-level expression of the human ?-globin gene in transgenic mice. Cell 51: 975?985 (1987).

Gruenbaum Y, Naveh-Many T, Ceder H, Razin A: Sequence specificity of methylation in higher plant DNA. Nature 292: 860?862 (1981).

Hepburn AG, Clarke LE, Pearson L, White J: The role of cytosine methylation in the control of nopaline synthase gene expression in a plant tumor. J Mol Appl Genet 2: 315?329 (1983).

Jefferson RA, Burgess SM, Hirsh D: ?-glucuronidase fromEscherichia coli as a gene-fusion marker. Proc Natl Acad Sci USA 83: 8447?8451 (1986).

Jefferson RA: Assaying chimeric genes in plants: the GUS fusion system. Plant Mol Biol Rep 5: 387?405 (1987).

Jones JDG, Dunsmuir P, Bedbrook J: High level expression of introduced chimaeric genes in regenerated transformed plants. EMBO J 4: 2411?2418 (1985).

Jones JDG, Gilbert DE, Grady KL, Jorgensen RA: T-DNA structure and gene expression in petunia plants transformed byAgrobacterium tumefaciens C58 derivatives. Mol Gen Genet 207: 478?485 (1987).

Junghans H, Metzlaff M: A simple and rapid method for the preparation of total plant DNA. Biotechniques 8: 176 (1990).

Kozak M: Point mutations define as sequence flanking the AUG initiator codon that modulates translation by eukaryotic ribosomes. Cell 44: 283?292 (1986).

Langridge WH, Fitzgerald KJ, Koncz C, Schell J, Szalay AA: Dual promoter ofAgrobacterium tumefaciens mannopine synthase genes in regulated by plant growth hormones. Proc Natl Acad Sci USA 86: 3219?3223 (1989).

Maliga P, Sz.-Brenznovits A, Marton L: Streptomycin-resistant plants from callus culture of haploid tobacco. Nature New Biol 244: 28?30 (1973).

Maniatis T, Fritsch EF, Sambrook J: Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York (1982).

Martin C, Prescott A, Lister C, MacKay S: Activity of the transposon Tam3 inAntirrhinum and tobacco: possible role of DNA methylation. EMBO J 8: 997?1004 (1989).

Marton L, Wullems GJ, Molendijk L, Schilperoort RA:In vitro transformation of cultured cells fromNicotiana tabacum byAgrobacterium tumefaciens. Nature 277: 129?131 (1979).

Matzke MA, Primig M, Trnovsky J, Matzke AJM: Reversible methylation and inactivation of marker genes in sequentially transformed tobacco plants. EMBO J 8: 643?649 (1989).

Monk M: Memories of mother and father. Nature 328: 203?205 (1987).

Murashigi T, Skoog F: A revised medium for rapid growth and bio-assay with tobacco tissue cultures. Physiol Plant 15: 473?497 (1962).

Nagy F, Morelli G, Fraley RT, Rogers SG, Chua N-H: Photoregulated expression of a pearbcS gene in leaves of transgenic plants. EMBO J 12: 3063?3068 (1985).

Nagy JI, Maliga P: Callus induction and plant regeneration from mesophyll protoplasts ofNicotiana sylvestris. Z Pflanzenphysiol 78: 453?455 (1976).

Napoli C, Lemieux C, Jorgensen R: Introduction of a chimeric chalcone synthase gene into petunia results in reversible co-suppression of hormologous genes intrans. Plant Cell 2: 279?289 (1990).

Neumann JR, Morency CA, Russian KO: A novel rapid assay for chloramphenicol acetyltransferase gene expression. Biotechniques 5: 444?448 (1987).

Odell JT, Nagy F, Chua N-H: Variability in 35S promoter expression between independent transformants. In: Key J, McIntosh L (eds) Plant Gene Systems and their Biology, pp. 289?299. A.R. Liss, New York (1987).

Peach C, Velten J: Application of the chloramphenicol acetyltransferase (CAT) diffusion assay to transgenic plant tissues. Bio Techniques (submitted) (1991).

Peach C, Velten J: Simultaneous analysis of the expression characteristics of two linked, divergent T-DNA promoters in transgenic tobacco. Plant Mol Biol (in preparation) (1991).

Peerbolte R, Leenhouts K, Hooykaas-van Slogteren GMS, Wullems GJ, Schilperoort RA: Clones from a shooty tobacco crown gall tumor II: irregular T-DNA structures and organization, T-DNA methylation and conditional expression of opine genes. Plant Mol Biol 7: 285?299 (1986).

Phillips RL, Kaeppler SM, Peschke VM: Do we understand somaclonal variation? In: Nijkamp HJJ, van der Plas KHW, van Aartrijk J (eds) Current Plant Science and Biotechnology in Agriculture, vol. 9, pp. 131?141. Kluwer Academic Publishers, Dordrecht (1990).

Reik W, Collick A, Norris ML, Barton SC, Surani MA: Genetic imprinting determines methylation of parental alleles in transgenic mice. Nature 328: 248?251 (1987).

Reitman M, Lee E, Westphal H, Felsenfeld G: Site-independent expression of the chicken 60?1 gene in transgenic mice. Nature 348: 749?752 (1990).

Rosahl S, Schell J, Willmitzer L: Expression of a tuberspecific storage protein in transgenic tobacco plants: demonstration of an esterase activity. EMBO J 6: 1155?1159 (1987).

Sanger M, Daubert S, Goodman RM: Characteristics of a strong promoter from figwort mosaic virus: comparison with the analogous 35S promoter from cauliflower mosaic virus and the regulated mannopine synthase promoter. Plant Mol Biol 14: 433?443 (1990).

Sapienza C, Peterson AC, Rossant J, Balling R: Degree of methyaltion of transgenes is dependent on gamete of origin. Nature 328: 251?254 (1987).

Saul MW, Shillito RD, Negrutiu I: Direct DNA transfer to protoplasts with and without electroporation. In: Gelvin SB, Schilperoort RA (eds) Plant Molecular Biology Manual, pp. A1/1-A1/16. Kluwer Academic Publishers, Dordrecht (1988).

Schwartz D, dennis E: Transposase activity of the Ac controlling element in maize is regulated by its degree of methylation. Mol Gen Genet 205: 476?482 (1986).

Shirsat AH, Wilford N, Croy RRD: Gene copy number and levels of expression in transgenic plants of a seed specific promoter. Plant Sci 61: 75?80 (1989).

Southern E: Detection of specific sequences among DNA fragments separated by gel electrophoresis. J Mol Biol 98: 503?517 (1975).

Spiker S: Chromatin structure and gene regulation in higher plants. Adv Genet 22: 145?207 (1984).

Teeri TH, Lehväslaiho H, Franck M, Uotila J, Heino P, Palva ET, Van Montagu M, Herrera-Estrella L: Gene fusions tolacZ reveal new expression patterns of chimeric genes in transgenic plants. EMBO J 8: 343?350 (1989).

Tuan D, Solomon W, Qiliang L, London IM: The ??-globin? gene domain in human erythroid cells. Proc Natl Acad Sci USA 82: 6384?6388 (1985).

Uchimiya H, Hirochika H, Hashimoto T, Hara A, Masuda T, Kasumimto T, Harada H, Ikeda J-E, Yoshioka M: Co-expression and inheritance of foreign genes in transformants obtained by direct DNA transformation of tobacco protoplasts. Mol Gen Genet 205: 1?8 (1986).

van der Krol AR, Mur LA, Beld M, Mol JNM, Stuitje AR: Flavonoid genes in petunia: addition of a limited number of gene copies may lead to a suppression of gene expression. Plant Cell 2: 291?299 (1990).

van Slogteren GMS, Hooykaas PJJ, Schilperoort RA: Silent T-DNA genes in plant lines transformed byAgrobacterium tumefaciens are activated by grafting and by 5-azacytidine treatment. Plant Mol Biol 3: 333?336 (1984).

Velten J, Schell J: Selection-expression plasmid vectors for use in genetic transformation of higher plants. Nucl Acids Res 13: 6981?6998 (1985).

Velten J, Velten L, Hain R, Schell J: Isolation of a dual plant promoter fragment from the Ti plasmid ofAgrobacterium tumefaciens. EMBO J 3: 2723?2730 (1984).

Winter JA, Wright RL, Gurley WB: Map locations of five transcripts homologous to TR-DNA in tobacco and sunflower gall tumors. Nucl Acids Res 9: 4801?4812 (1984)

Wullems GJ, Krens FA, Schilperoort RA: Transformation of protoplasts via cocultivation with Agrobacterium and via Ti plasmids. In: Evans DA, Sharp WR, Ammirato PV (eds) Handbook of Plant Cell Culture, vol 4: Techniques and Applications, pp. 197?220. Macmillan, New York (1986).

Zambryski P, Joos H, Genetello C, Leemans J, Van Montagu M, Schell J: Ti plasmid vector for the introduction of DNA into plants cells without alteration of their normal regeneration capacity. EMBO J 2: 2143?2150 (1983).