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Primary structure of A B1 hordein gene from barley


The 873 base pair coding region of a Hor-2 gene of barley and the adjacent 550 base pair upstream and 413 base pair downstream regions were sequenced. The gene is devoid of introns and encodes a 271 amino acid long B1 hordein polypeptide containing a putative 19 amino acid signal peptide. The remaining part of the coding sequence can be divided into three parts. In the 53 residue amino-terminal region there are 9 glutamine-proline blocks with a preferred core sequence PQQP, separated by one or two other residues giving a glutamine proline content of 78%. The second part encodes 164 amino acids, 41% of which are glutamine+proline organised in scattered blocks. Seven cysteine residues are coded for by this portion of the gene. The last part encodes the carboxyterminal 35 amino acids none of which is glutamine.

In the 550 base pair upstream region the sequence TATAAATA is found at- 71 base pairs from the initiator methionine. In the 3′ non-coding region three putative polyadenylation signals, AATAAA, are present.

Comparison of the gene with 3 partial cDNA clones indicates that the charge polymorphism in the B1 polypeptide group is due to point mutations in the part of the gene corresponding to the carboxy terminal half of the polypeptide. Comparison with the sequence of a second B hordein gene suggests that insertions or deletions of glutamine-proline blocks in the amino-terminal domain are a major source of size polymorphisms in the B hordein family.



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  1. 1.

    Anderson, O. D., J. C. Litts, M.-F. Gautier &F. C. Greene: Nucleic acid squence and chromosome assignment of a wheat storage protein gene. Nucl. Acids Res. 12, 8129–8144 (1984)

  2. 2.

    Benton, W. D. &R. W. Davis: Screening λ gt recombinant clones by hydridization to single plaques in situ. Science 196, 180–182 (1977)

  3. 3.

    Brandt, A. &J. Ingvesen: Isolation and translation of hordein messenger RNA from wild type and mutant endosperm in barley. Carlsberg Res. Commun. 43, 451–469 (1978)

  4. 4.

    Broglie, R., G. Coruzzi, R. T. Fraley, S. G. Rogers, R. B. Horsch, J. G. Niedermeyer, C. L. Fink, J. S. Flick &N.-H. Chua: Light-regulated expression of a pea ribulose-1,5-bisphosphate carboxylase, small subunit gene in transformed plant cells. Science 224, 838–843 (1984)

  5. 5.

    Cameron-Mills, V.: The structure and composition of protein bodies purified from barley endosperms by silica sol density gradients. Carlsberg Res. Commun. 45, 557–576 (1980)

  6. 6.

    Cameron-Mills, V. &J. Ingversen: In vitro synthesis and transport of barley endosperm proteins: Reconstitution of functional rough microsomes from polyribosomes and stripped microsomes. Carlsberg Res. Commun. 43, 471–489 (1978)

  7. 7.

    Cameron-Mills, V. &D. von Wettstein: Protein body formation in the developing barley endosperm. Carlsberg Res. Commun. 45, 577–594 (1980)

  8. 8.

    Covarrubias, L., L. Cervantes, A. Covarrubias, X. Soberon, I. Vichido, A. Blanco, Y. M. Kupersztoch-Portnoy &F. Bolivar: Construction and characterization of new cloning vehicles. V. Mobilization and coding properties of pBR322 and several deletion derivatives including pBR327 and pBR328. Gene 13, 25–35 (1981)

  9. 9.

    Faulks, A. J., P. R. Shewry &B. J. Miflin: The polymorphism and structural homology of storage polypeptides (hordein) coded by the Hor 2 locus in barley (Hordeum vulgare L.). Biochem. Genet. 19, 841–858 (1981)

  10. 10.

    Fincher, G.-B., P. A. Lock, M. M. Morgan, K., Lingelbach, R. E. H. Wettenhall, J. F. B. Mercer, A. Brandt & K. K. Thomsen: Primary structure of the (1→3,1→4)-β-glucanase from barley aleurone. Proc. Natl. Acad. Sci. USA (in press)

  11. 11.

    Forde, B. G., M. Kreiis, M. S. Williamson, R. P. Fry, J. Pywell, P. R. Shewry, N. Brunce &B. J. Miflin: Short tandem repeats shared by B- and C-hordein cDNAs suggest a common evolutionary origin for two groups of cereal storage protein genes. The EMBO J. 4, 9–15 (1985)

  12. 12.

    Forde, B. G., A. Heyworth, J. Pywell &M. Kreis: Nucleotide sequence of aB1 hordein gene and the identification of possible upstream regulatory elements in endosperm storage protein genes from barley, wheat and maize. Nucl. Acids Res. 13, 7327–7339 (1985)

  13. 13.

    Forde, J., J.-M. Malpica, N. G. Halford, P. R. Shewry, O. D. Anderson, F. C. Greene &B. J. Miflin: The nucleotide sequence of a HMW glutenin subunit gene located on chromosome 1A of wheat (Triticum aestivum L.) Nucl. Acids Res. 13, 6817–6832 (1985)

  14. 14.

    Giese, H., B. Andersen &H. Doll: Synthesis of the major storage protein, hordein, in barley. Pulse-labeling study of grain filling in liquid-cultured detached spikes. Planta 159, 60–65 (1983)

  15. 15.

    Giese, H. &H. E. Hopp: Influence of nitrogen nutrition on the amount of hordein, protein Z and β-amylase messenger RNA in developing endosperms of barley. Carlsberg Res. Commun. 49, 365–383 (1984)

  16. 16.

    Heijne, G. von: On the hydrophobic nature of signal sequences. Eur. J. Biochem. 116, 419–422 (1981)

  17. 17.

    Hohn, B. &K. Murray: Packaging recombinant DNA molecules into bacteriophage particles in vitro. Proc. Nat. Acad. Sci. 74, 3259–3263 (1977)

  18. 18.

    Holder, A. A. &J. Ingversen: Peptide mapping of the major components of in vitro synthesized barley hordein: Evidence for structural homology. Carlsberg Res. Commun. 43, 177–184 (1978)

  19. 19.

    Hopp, H. E., S. K. Rasmussen &A. Brandt: Organization and transcription ofB1 hordein genes in high lysine mutants of barley. Carlsberg Res. Commun. 48, 201–216 (1983)

  20. 20.

    Hu, N.-T., M. A. Peifer, G. Heidecker, I. Messing &I. Rubenstein: Primary structure of a genomic zein sequence of maize. The EMBO J. 1, 1337–1342 (1982)

  21. 21.

    Jensen, I. H., J. Jørgensen, H. P. Jensen, H. Giese &H. Doll: Linkage of the hordein loci Horl and Hor2 with the powdery mildew resistance loci Ml-k and Ml-a on barley chromosome 5. Theor. Appl. Genet. 58, 27–31 (1980)

  22. 22.

    Kreis, M., P. R. Shewry, B. G. Forde, S. Rahman &B. J. Miflin: Molecular analysis of a mutation conferring the high-lysine phenotype on the grain of barley (Hordeum vulgare). Cell 34, 161–167 (1983)

  23. 23.

    Loenen, W. A. M. &W. J. Brammer: A bacteriophage lambda vector for cloning large DNA fragments made with several restriction enzymes. Gene 10, 249–259 (1980)

  24. 24.

    Lycett, G. W., R. R. D. Croy, A. H. Shirsat &D. Boulter: The complete nucleotide sequence of a legumin gene from pea (Pisum sativum L.). Nucl. Acids Res. 12, 4493–4506 (1984)

  25. 25.

    Lycett, G. W., R. R. D. Croy, A. H. Shirsat, D. M. Richards &D. Boulter: The 5′-flanking regions of three pea legumin genes: comparison of the DNA sequences. Nucl. Acids Res. 13, 6733–6743 (1985).

  26. 26.

    Maniatis T., E. F. Fritsch & J. Sambrook: Molecular cloning, a laboratory manual. Cold Spring Harbor Laboratory (1982)

  27. 27.

    Messing, J. &J. Vieira: A new pair of M13 vectors for selecting either DNA strand of doubie-digest restriction fragments. Gene 19, 269–276 (1982)

  28. 28.

    Pedersen, K., J. Devereux, D. R. Wilson, E. Sheldon &B. A. Larkins: Cloning and sequence analysis reveal structural variation among related zein genes in maize. Cell 29, 1015–1026 (1982)

  29. 29.

    Rafalski, J. A., K. Scheets, M. Metzler, D. M. Peterson, C. Hedgcoth &D. Soll: Developmentally regulated plant genes: The nucleotide sequence of a wheat gliadin genomic clone. The EMBO J. 3, 1409–1415 (1984)

  30. 30.

    Rasmussen, S. K., H. E. Hopp &A. Brandt: Nucleotide sequences of cDNA clones for Bl hordein polypeptides. Carlsberg Res. Commun. 48, 187–199 (1983)

  31. 31.

    Rogers, J. C. &C. Milliman: Isolation and sequence analysis of a barley α-amylase cDNA clone. J. Biol. Chem. 258, 8169–8174 (1983)

  32. 32.

    Sanger, F., S. Nicklen &A. R. Coulson: DNA sequencing with chain-terminating inhibitors. Proc. Nat. Acad. Sci. USA 74, 5463–5467 (1977)

  33. 33.

    Schmitt, J. M.: Purification of hordein polypeptides by column chromatography using volatile solvents. Carlsberg Res. Commun. 44, 431–438 (1979)

  34. 34.

    Schmitt, J. M. &I. Svendsen: Partial amino acid sequence from hordein polypeptide B1. Carlsberg Res. Commun. 45, 549–555 (1980)

  35. 35.

    Sengupta-Gopalan, C., N. A. Reichert, R. F. Barker T. C. Hall &J. D. Hemp: Developmentally regulated expression of the bean β-phaseolin gene in tobacco seed. Proc. Natl. Acad. Sci. USA 82, 3320–3324 (1985)

  36. 36.

    Shewry, P. R., A. J. Faulks, R. A. Pickering, I. T. Jones, R. A. Finch &B. J. Miflin: The genetic analysis of barley storage proteins. Heredity 44, 383–389 (1980)

  37. 37.

    Souciet, J.-L., J.-C. Hubert &F. Lacroute: Cloning and restriction mapping of the yeast URA2 gene coding for the carbamyl phosphate synthetase aspartate-transcarbamylase complex. Mol. Gen. Genet. 186, 385–390 (1985)

  38. 38.

    Sumner-Smith, M., J. A. Rafalski, T. Sugiyama, M. Stoll &D. Soll: Conservation and variability of wheat α/β-gliadin genes. Nucl. Acids Res. 13, 3905–3916 (1985)

  39. 39.

    Thompson, R. D., D. Bartels &N. P. Harberd: Nucleotide sequence of a gene from chromosome 1D of wheat encoding a HMW-glutenin subunit. Nucl. Acids Res. 13, 6833–6846 (1985)

  40. 40.

    Weber, A. &A. Brandt: Species specific signal peptide cleavage of plant storage protein precursors in the endoplasmic reticulum. Carlsberg Res. Commun. 50, 229–308 (1985)

  41. 41.

    Weiher, H., M. Konig &P. Gruss: Multiple point mutations affecting the simian virus 40 enhancer. Science 219, 626–631 (1983)

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Brandt, A., Montembault, A., Cameron-Mills, V. et al. Primary structure of A B1 hordein gene from barley. Carlsberg Res. Commun. 50, 333 (1985). https://doi.org/10.1007/BF02907156

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  • Endosperm
  • storage protein
  • Hordeum vulgare
  • gene family
  • heterogeneity
  • molecular cloning