140. Peters, B., Aidley, J., Cadzow, M., Twell, D. and Brownfield, L. (2017) Identification of cis-regulatory modules that function in the male germline of flowering plants. Methods in Molecular Biology (in press).

139. Twell, D. and Brownfield, L. (2017) Analysis of fluorescent reporter activity in the male germline during pollen development by confocal microscopy. Methods in Molecular Biology (in press).

138. Rieu, I., Twell, D. and Firon, N. (2017) Pollen development at high temperature: from acclimation to collapse. Plant Physiology 173, 1967-1976.

137. Peters, B., Casey, J., Aidley, J., Zohrab, S., Borg, M., Twell, D.*, Lynette Brownfield, L.* (2017) A conserved cis-regulatory module determines germline fate through activation of the transcription factor DUO1. Plant Physiology 173, 280-293. * Co-senior authors.


136. Yue, L., Twell, D., Kuang, Y., Liao, J. and Zhou, X. (2016). Transcriptome analysis of Hamelia patens (Rubiaceae) anthers reveals candidate genes for tapetum and pollen wall development. Frontiers Plant Sci. doi: 10.3389/fpls.2016.01991

135. Grover, A, Twell, D. Schleiff, E (2016) Pollen as a target of environmental changes (editorial). Plant Reproduction 29, 1-2.

134. Brownfield, L. and Twell, D. (2016) Plant Reproduction. In: eLS (Encyclopedia of Life Sciences). John Wiley & Sons, Ltd, Chichester. [doi: 10.1002/9780470015902.a0002046.pub2]

133. Oh, S-A, Jeon, J., Park, H-J., Grini, P. Twell, D. and Park, S-K. (2016) Analysis of gemini pollen 3 mutant suggests a broad function of AUGMIN in microtubule organization during sexual reproduction in Arabidopsis. Plant Journal 87, 188-201.


132. Rutley, N. and Twell, D. (2015). A decade of pollen transcriptomics. Plant Reproduction 28, 73-89.

131. Harrison, M., Mallon, E., Twell, D. and Hammond, R. (2015) Deleterious mutation accumulation in Arabidopsis thaliana pollen genes: a role for a recent relaxation of selection. BioRxiv. Doi:

130. Brownfield, L., Yi, J., Jiang, H., Minina, E.A., Twell, D., Köhler, C. (2015). Organelles maintain spindle position in plant meiosis.  Nature Communications 6:6492.

129. Schoft_V., Chumak, N., Slusarz, L., Kohler, C., Twell, D. and Tamaru, H. (2015). SYBR green activated Arabidopsis pollen nucleus sorting based on different DNA/RNA content. Plant Reproduction 28:61-72.

128. Cui, X., Lv, Y., Chen, M., Nikoloski, Z. Twell, D. and Zhang, D. (2015). Young genes out of the male: an insight from evolutionary age analysis of the pollen transcriptome. Molecular Plant 8, 935-945.


127. Park, G.T., Frost, J. M., Park, J-S., Kim, T.H., Lee, J.S., Oh, S-A., Twell, D., Brooks, J.S., Fischer, R. L. and Choi, Y. (2014). Nucleoporin MOS7/Nup88 is required for mitosis in gametogenesis and seed development in Arabidopsis. Proc Natl Acad. Sci. USA. 111:18393-8.

126. Hoedemaekers, K., Derksen, J., Hoogstrate, S., Wolters-Arts, M., Oh, S-A., Twell, D., Mariani, C., Rieu, I. (2014). The Arabidopsis glycosyltransferase BURSTING POLLEN (BUP) is required to organise the pollen germination plaque and pollen tube tip. New Phytologist 206:255-67.

125. Riddell, C.E., Lobaton-Garces, J.D., Adams, S., Barribeau, S.M., Twell, D. and Mallon, E.B. (2014). Differential gene expression and alternative splicing in insect immune specificity. BMC genomics 15:1031.

 124. Rennie, E.A, Ebert, B., Miles, G.P., Christiansen, K., Stonebloom, S., Khatab, H., Twell, D., Petzold, C.J., Adams, P.D., Dupree, P., Heazlewood, J-L. and Scheller, H.V. (2014) Identification of a sphingolipid α-glucuronosyltransferase that is essential for pollen function in Arabidopsis. Plant Cell 26: 3314-25.

123. Borg, M., Rutley, N., Kagale, S. Hamamura, Y., Gherghinoiu, M., Kumar, S., Sari, U., Esparza-Franco, MA., Sakamoto, W., Rozwadowski, K., Higashiyama, T. and Twell, D. (2014). An EAR-dependent regulatory module promotes male germ cell division and sperm fertility in Arabidopsis. Plant Cell 26:1-17.

122. Oh, S.A., Bourdon, V. Dickinson, H.G., Twell, D.* and Park, S.K.* (2014) Arabidopsis Fused kinase TWO-IN-ONE dominantly inhibits male meiotic cytokinesis. Plant Reproduction. 27(1):7-17 .* Co-senior authors.


121. Grant-Downton, R., Kourmpetli, S., Hafidh, S., Khatab, H., LeTrionnaire, G., Dickinson, H.G. and Twell, D. (2013). Artificial microRNAs reveal cell-specific differences in small RNA activity in pollen. Curr Biol. 23 (14), R599-R601.


120. Oh, S.A., Allen, T., Kim, G.J., Sidorova, A., Michael Borg, M., Park, S.K.* and Twell, D.* (2012) Arabidopsis Fused kinase and the Kinesin-12 subfamily constitute a signalling module required for phragmoplast expansion. Plant J. 72, 308-319.* Co-senior authors.

119. Kasahara, R.D., Maruyama, D., Hamamura, Y., Sakakibara, T., Twell, D. and Higashiyama, T. (2012). Fertilization recovery after defective sperm cell release in Arabidopsis. Curr Biol. 22, 1084-1089. 


118. Berger, F. and Twell, D. (2011) Germline specification and function in plants. Annu Rev Plant Biol 62, 461-484.

117. Borg, M. and Twell, D. (2011) Pollen: Structure and Development. In: eLS (Encyclopedia of Life Sciences). John Wiley & Sons, Ltd: Chichester, Nature Publishing Group ( [DOI: 10.1002/9780470015902.a0002039.pub2]

116. Oh, SA, Twell, D. and Park, SK (2011) SIDECAR POLLEN suggests a plant-specific regulatory network underlying asymmetric microspore division in Arabidopsis. Plant Signal Behav 6, 416-419.

115. Borg, M., Brownfield, L., Khatab, H., Sidorova, A., Lingaya, M. and Twell, D. (2011) The R2R3 MYB transcription factor DUO1 activates a male germline-specific regulon essential for sperm cell differentiation in Arabidopsis. Plant Cell 23, 1-16.

114. Twell, D. (2011) Male gametogenesis and germline specification in flowering plants. Sex Plant Reprod. 24:149-160.

113. Le Trionnaire, G. Grant-Downton, R. Kourmpetli, S. Dickinson H. and Twell, D. (2011) Small RNA activity and function in angiosperm gametophytes. J Exp Bot 62, 1601-1610.

112. Matsushima, R., Tang, L.Y., Zhang, L., Yamada, H., Twell, D. and Sakamoto, W. (2011) A conserved, Mg2+-dependent exonuclease degrades organelle DNA during Arabidopsis pollen development. Plant Cell 23, 1608-1624.

111. Kim, H.J., Ok, S.H., Bahn, S.C., Jang, J., Oh, S.A., Park, S.K., Twell, D., Ryu, S.B. and Shin, J.S. (2011) Endoplasmic Reticulum– and Golgi-Localized Phospholipase A2Plays Critical Roles in Arabidopsis Pollen Development and Germination. Plant Cell 23, 1608-1624.


110. Li, S-L.,  Zong, J.,   Zhou, Z-G., Twell, D. and Zhang, D-B. (2010) The rice MYB transcription factor OsDUO1, cDNA cloning and preliminary analysis of its expression pattern. Plant Physiol. Comm. 10, 1033-1039.

109. Oh, S-A, Park, K.S., Twell, D.*, Park, S.K.* (2011) The SIDECAR POLLEN gene encodes a microspore-specific LOB/AS2 domain protein required for the correct timing and orientation of asymmetric cell division. Plant J 64:839-850.*Co-senior authors.

108. LeTrionnaire, G. and Twell, D. (2010) Small RNAs in angiosperm gametophytes: from epigenetics to gamete development. Genes Dev. 24, 1081-1085.

107. Oh, S-A, Allen, T. and Twell, D. (2010) A ticket for the live show: Microtubules in male gametophyte development. Plant Signal. Behav. 5, 1-4.

106. Oh, S-A., Das Pal, M., Park, S-K, Johnson, J.A. and Twell, D. (2010) The MAP215/Dis1-family protein TMBP200 is required for the organization and function of gametophytic microtubule arrays associated with asymmetric division and male germline establishment in tobacco. J Exp Bot 61, 969-981.

105. Borg, M. and Twell, D. (2010) Life after meiosis: Patterning the angiosperm male gametophyte. Biochem Soc Trans 38, 577-582.


103. Brownfield, L, Twell, D (2009) A dynamic DUO of regulatory proteins coordinates gamete  specification and germ cell mitosis in the angiosperm male germline. Plant Signaling & Behavior 4(12):1-4.

102. Erilova, A, Brownfield, L, Exner, V, Rosa, M, Twell, D, Mittelsten Scheid, O, Hennig, L Köhler, C (2009) Imprinting of the Polycomb Group Gene MEDEA Serves as a Ploidy Sensor in Arabidopsis. PLoS Genet. 5(9):e1000663

101. Schoft, VK, Chumak, N, Mosiolek, M, Slusarz, L, Komnenovic, V, Brownfield, L, Twell, D., Kakutani, T, Tamaru, H (2009) Induction of RNA-directed DNA methylation upon decondensation of constitutive heterochromatin. EMBO Rep 10, 1015-21.

100. Brownfield, L, Hafidh, S, Durbarry, A, Khatab, H, Sidorova, A, Doerner, P, Twell D (2009) Arabidopsis DUO POLLEN 3 is a key regulator of male germline development and embryogenesis. Plant Cell 21, 1940-56.  

99. Gibalová, A, Reňák, D, Matczuk, K, Dupľáková, N, Cháb, D, Twell, D, Honys, D (2009) AtbZIP34 is required for cell wall patterning and the control of lipid metabolism and cellular transport genes in Arabidopsis pollen. Plant Mol Biol 70, 581-601.

98. Zhong, C, Hafidh, S, Poh, SH, Twell, D, Berger, F (2009) Hyperproliferation caused by loss of Retinoblastoma prevents cell fate establishment during Arabidopsis male gametogenesis. Proc Natl Acad Sci USA 106, 7257-62.

97. Brownfield, L, Hafidh, S, Borg, M, Sidorova, A, Mori, T, Twell, D (2009) A plant germ cell-specific integrator of cell cycle progression and sperm specification PLoS Genet. 5: e1000430.

96. Twell, D (2009) Male Gametophyte Development. In E. C. Chong & M. R. Davies (Eds.) Plant Developmental Biology-Biotechnological Perspectives. Springer, Germany. (in press)

95. Borg, M, Brownfield, L, Twell, D, (2009) Male gametophyte development: A molecular perspective. J Exp Bot 60, 1465-78.

94. Grant-Downton, R, Hafidh, S, Twell, D, Dickinson, HG (2009) Small RNA pathways are present and functional in the angiosperm male gametophyte. Mol. Plant 2, 500-12.


93. Moll C, von Lyncker L, Kägi C, Baumann N, Twell D, Grossniklaus U, Groß-Hardt R (2008) CLOTHO is a novel regulator of gametic cell fate in plants and acts in a non-cell autonomous manner Plant J 56, 913-921.

92. Kim HJ, Oh S-A, Brownfield L, Ryu H, Hwang I, Twell D, Nam H-G (2008) Control of plant male germline proliferation by SCFFBL17 degradation of cell cycle inhibitors. Nature 455, 1134-1137.

91. Oh S-A, Bourdon V, Das Pal M, Dickinson HG, Twell D (2008) Arabidopsis kinesins HINKEL and TETRASPORE act redundantly to control cell plate expansion during cytokinesis in the male gametophyte. Mol Plant 1, 794-799.

90. Töller A, Brownfield L, Neu C, Twell D, Schulze-Lefert P (2008). Dual function of Arabidopsis Glucan Synthase-Like genes GSL8 and GSL10 in male gametophyte development and plant growth. Plant J 54, 911-923.

89. Quan L, Xiao R, Li W, Oh S-A, Kong H, Ambrose J, Malcos J, Cyr R, Twell D, Ma H (2008) Functional divergence of the duplicated AtKIN14b and AtKIN14b genes: critical roles in Arabidopsis meiosis and gametophyte development. Plant J 53, 1013-1026.


88. Andersen SU, Algreen-Petersen RG, Hoedl M, Jurkiewicz A, Cvitanich C, Braunschweig U, Schauser L, Oh S-A, Twell D and Jensen EO (2007) The conserved cysteine-rich domain of a tesmin/TSO1-like protein binds zinc in vitro and TSO1 is required for both male and female fertility in Arabidopsis thaliana. J Exp Bot 58, 3657-3670.

87. Verelst W, Twell D, de Folter S, Immink R, Saedler H and Münster T (2007) MADS-complexes regulate transcriptome dynamics during pollen maturation. Genome Biol 8:R249

86. Dupľáková N, Reňák D, Hovanec P, Honysová B, Twell D, and Honys D (2007) Arabidopsis Gene Family Profiler (aGFP) - user-oriented transcriptomic database with easy-to-use graphic interface. BMC Plant Biology 7, 39.


85. Honys D, Oh S-A, Renak D, Donders M, Solcova B, Johnson JA, Boudova R and Twell D (2006) Identification of microspore-active promoters that allow targeted manipulation of gene expression at early stages of microgametogenesis in Arabidopsis. BMC Plant Biol 6:31.

84. Novak JP, Kim S-Y, Xu J, Modlich O, Volsky DJ, Honys D, Slonczewski JL, Bell DA, Blattner FR, Blumwald E, Boerma M, Cosio M, Gatalica Z, Hajduch M, Hidalgo J, McInnes RR, Miller MC III, Penkowa M, Rolph MS, Sottosanto J, St-Arnaud R, Szego MJ, Twell D and Wang C (2006) Generalization of DNA microarray dispersion properties: microarray equivalent of t-distribution. Biology Direct 1:27.

83. Bock KW, Honys D, Ward JM, Padmanaban S, Nawrocki EP, Hirschi KD, Twell D and Sze H (2006) Integrating membrane transport with male gametophyte development and function through transcriptomics. Plant Physiol 140, 1151-68.

82. Honys D, Renak D and Twell D (2006) Male Gametophyte Development and Function. In Teixera da Silva JA (Ed.), Floriculture, Ornamental and Plant Biotechnology: Advances and Topical Issues (1st Edition). Global Science Books, London, UK. pp. 76-87.

81. Twell D, Oh S-A and Honys D (2006) Pollen development, a genetic and transcriptomic view. In Malho R (ed.) The Pollen Tube: A Cellular and Molecular Perspective. Plant Cell Monographs Volume 3. Springer-Verlag, Berlin. pp.15-45.

80. Ehsanpour AA and Twell D (2006) Yeast two-hybrid screening of Arabidopsis thaliana for SETH4 protein interaction. J Sci Is Rep Iran 17, 5-10.

79. Twell D (2006) A blossoming romance: gamete interactions in flowering plants. Nature Cell Biol 8, 14-16.

78. Oh S-A, Johnson JA, Smertenko A, Rahman D, Park S-K, Hussey PJ and Twell D (2006) A divergent cellular role for the FUSED kinase family in the plant-specific cytokinetic phragmoplast. Curr Biol 15, 2107-2111.


77. Baldwin A, Wardle A, Patel R, Dudley P, Park SK, Twell D, Inoue K and Jarvis P (2005) A molecular-genetic study of the Arabidopsis Toc75 gene family. Plant Physiol 138, 715-733.

76. Rotman N, Durbarry A, Wardle A, Yang W-C, Chaboud A, Faure J-E, Berger F and Twell D (2005) A novel class of MYB factors controls sperm cell formation in plants. Curr Biol 15, 244-248.

75. Durbarry A, Vizir I and Twell D (2005) Male germ line development in Arabidopsis: duo pollen mutants reveal gametophytic regulators of generative cell cycle progression. Plant Physiol 137, 297-307.

74. Combe JP, Petracek ME, van Eldik G, Meulewaeter F and Twell D (2005) Translation initiation factors eIF4E and eIFiso4E are required for polysome formation and regulate plant growth in tobacco. Plant Mol Biol 57, 749-760.

73. Ehsanpour AA and Twell D (2006) Analysis of SFL1 and SFL2 promoter region in Arabidopsis thaliana using Gateway cloning system. J Sci Is Rep Iran 16, 303-309.

72. Hrubá P, Honys D, Twell D, Capková V and Tupy J (2005) Expression of beta-galactosidase and beta-xylosidase genes during microspore and pollen development. Planta 220, 931-940.

71. Kovacheva S, Bedard J, Patel R, Dudley P, Twell D, Rios G, Koncz C and Jarvis P (2005) In vivo studies on the roles of Tic110, Tic40 and Hsp93 during chloroplast protein import. Plant J 41, 412-428.


70. Honys D and Twell D (2004) Transcriptome analysis of haploid male gametophyte development in Arabidopsis. Genome Biology 5/11/R85.

69. Lalanne E, Michaelidis C, Moore JM, Gagliano W, Johnson A, Patel R, Howden R, Vielle-Calzada J-P, Grossniklaus U and Twell D (2004 ) Analysis of transposon insertion mutants highlights the diversity of mechanisms underlying male progamic development in Arabidopsis. Genetics 167, 1975-1986.

68. Lobstein E, Guyon A, Férault M, Twell D, Pelletier G and Bonhomme S (2004) The putative Arabidopsis homolog of yeast Vps52p is required for pollen tube elongation, localizes to Golgi and might be involved in vesicle trafficking. Plant Physiol 135, 1480-1490.

67. Park SK, Rahman D, Oh SA and Twell D (2004) gemini pollen 2, a male and female gametophytic cytokinesisdefectivemutation. Sex Plant Reprod 17, 63-70.

66. Sze H, Padmanaban S, Cellier F, Honys D, Cheng N-H, Bock KW, Conéjéro G, Li X, Twell D, Ward JM and Hirschi K (2004) Expression patterns of a novel AtCHX gene family highlight potential roles in osmotic adjustment and K+ homeostasis in pollen development. Plant Physiol 136, 2532-2547.

65. Honys D and Twell D (2004) Male gametogenesis. In Goodman RM (ed.) Encyclopedia of Plant and Crop Science. Marcel Dekker, Inc., New York. pp. 663-668.

64. Lalanne E, Honys D, Johnson A, Borner GHH, Lilley KS, Dupree P, Grossniklaus U and Twell D (2004) Seth1 and Seth2, two components of the glycosylphosphatidylinositol anchor biosynthetic pathway, are required for pollen germination and tube growth in Arabidopsis. Plant Cell 16, 229-240.


63. Cheung AY, Chen CY-H, Tao L-Z, Andreyeva T, Twell D and Wu H-M (2003). Regulation of Pollen Tube Growth by Rac-like GTPases J Exp Bot 54, 73-81.

62. Honys D and Twell D (2003) Comparative analysis of the Arabidopsis pollen transcriptome. Plant Physiol 132, 640-652.

61. Goubet F, Misrahi A, Park S-K, Zhang Z, Twell D and Dupree P (2003) AtCSLA7, a cellulose-synthase-like putative glycosyltransferase, is important for pollen tube growth and embryogenesis in Arabidopsis. Plant Physiol 131, 547-557.

60. O, S-A, Park S-K, Jang I, Howden R, Moore JM, Grossniklaus U and Twell D (2003) halfman, an Arabidopsis male gametophytic mutant associated with a 150 kb chromosomal deletion at the site of transposon insertion. Sex Plant Reprod 16, 99-102.


59. Hulzink RJM, de Groot PFM, Croes AF, Quaedvlieg W, Twell D, Wullems GJ and van Herpen MMA (2002) The 5’ UTR of the ntp303 gene strongly enhances translation during pollen tube growth, but not during pollen maturation. Plant Physiol 129, 342-353.

58. Lalanne E and Twell D (2002) Genetic control of male germ unit organization in Arabidopsis Plant Physiol 129, 865-875.

57. Twell D, Park SK, Hawkins TJ, Schubert D, Schmidt R, Smertenko A and Hussey PJ (2002) MOR1/GEM1 plays an essential role in the plant-specific cytokinetic phragmoplast. Nat Cell Biol 4, 711-714.

56. Twell D (2002) Pollen developmental biology. In O'Neill SD and Roberts JA (eds.) Plant Reproduction. Annual Plant Reviews, Volume 6. Sheffield Academic Press, Sheffield. pp. 86-153.


55. Yang S, Sweetman J, Amirsadeghi S, Barghchi M, Huttly AK, Chung W-I, Twell D (2001) Novel anther-specific myb genes from tobacco as putative regulators of phenylalanine ammonia-lyase expression. Plant Physiol 126, 1738-1753.

54. Park SK and Twell D (2001) Novel patterns of ectopic cell plate growth and lipid body distribution in the Arabidopsis gemini pollen1 mutant. Plant Physiol 126, 899-909.

53. Rogers HJ, Bate N, Combe J, Sullivan J, Sweetman JS, Swan C, Lonsdale DM and Twell D (2001) Functional analysis of cis-regulatory elements within the promoter of the tobacco late pollen gene g10. Plant Mol Biol 45, 577-585.


52. Honys D, Combe JP, Twell D and Capková V (2000) The translationally repressed pollen-specific ntp303 mRNA is stored in non-polysomal mRNPs during pollen maturation. Sex Plant Reprod 13,135-144.

51. Twell D (2000) Pollen structure development and function. Encyclopedia of Life Sciences. Macmillan Reference Ltd. Nature, Scientific American Publishing Group.

50. Sweetman J, Spurr C, Eliasson A, Gass N, Steinmetz A and Twell D (2000) Isolation and characterisation of two pollen-specific LIM domain protein cDNAs from Nicotiana tabacum. Sex Plant Reprod 12, 339-345.

49. Wittink FRA, Knuiman B, Derksen J, Capková V, Twell D, Schrauwen JAM and Wullems GJ (2000) The pollen-specific gene Ntp303 encodes a 69kD glycoprotein associated with the vegetative membranes and the cell wall. Sex Plant Reprod 12, 276-284.


48. Turner RL, Glynn M, Taylor SC, Cheung M-K, Spurr C, Twell D and Foster GD (1999) Analysis of a translational enhancer present within the 5-terminal sequence of the genomic RNA of potato virus S Arch Virol 144, 1451-1461.

47. Twell D (1999). Mechanisms of microspore polarity and differential cell fate determination in developing pollen. In Cresti M, Cai G and Moscatelli A (eds.). Fertilization in Higher Plants. Molecular and Cytological Aspects. Springer-Verlag, Berlin. pp. 201-215.


46. Amirsadeghi S, Sweetman J, Barghchi M and Twell D (1998) Physiological analysis of plant dwarfing. In: Davey MR, Alderson PG, Lowe KC, Power JB (eds.) Tree Biotechnology: Towards the Millenium. Nottingham University Press, Nottingham. pp.267-275.

45. Bate N and Twell D (1998) Functional architecture of a late pollen promoter: pollen-specific transcription is developmentally regulated by multiple stage-specific and co-dependent activator elements. Plant Mol Biol 37, 859-869.

44. Howden R, Park SK, Moore JM, Orme J, Grossniklaus U and Twell D (1998) Selection of T-DNA-tagged male and female gametophytic mutants by segregation distortion in Arabidopsis. Genetics 149, 621-631.

43. Park SK, Howden R and Twell D (1998) The Arabidopsis thaliana gametophytic mutation gemini pollen 1 disrupts microspore polarity, division asymmetry and pollen cell fate. Development 125, 3789-3799.

42. Tichtinsky G, Tavares R, Takvorian A, Schwebel-Dugué N, Twell D, Kreis M (1998) An evolutionary conserved group of plant GSK-3/shaggy like protein kinase genes preferentially expressed in developing pollen. Bioch Biophys Acta - Gene Structure and Expression. 1442, 261-273.

41. Twell D, Park SK and Lalanne E (1998) Asymmetric division and cell fate determination in developing pollen. Trends Pl Sci 3, 305-310.

40. Twell D and Howden R (1998) Mechanisms of asymmetric division and cell fate determination in developing pollen. In Chupeau Y, Caboche M, Henry Y (eds,) Androgenesis and haploid plants, 1967-1994. In memory of Jean-Pierre Bourgin. INRA-Springer-Verlag. pp. 69-103.

39. Twell D and Howden R (1998). Cell polarity, asymmetric division and cell fate determination in developing pollen. In Owens SJ and Rudall PJ (eds.) Reproductive Biology. Royal Botanic Gardens, Kew. pp. 197-218.

38. Wilkinson JE, Lindsey K and Twell D (1998) Antisense-mediated suppression of transgene expression targeted specifically to pollen. J Exp Bot 49, 1481-1490.


37. Wei W, Twell D and Lindsey K (1997) A novel nucleic acid helicase gene identified in Arabidopsis by promoter trapping. Plant J 11, 1307-1314.

36. Wilkinson JE, Twell D and Lindsey K (1997) Activity of CaMV35S and nos promoters in pollen: implications for field release of transgenic plants. J Exp Bot 48, 265-275.


35. Bate N, Spurr C, Foster GD and Twell D (1996) Maturation-specific translational enhancement mediated by the 5’-UTR of a late pollen transcript. Plant J 10, 101-111.

34. Foster G and Twell D (eds.) Plant Gene Isolation: principles and practice John Wiley & Sons Limited, Chichester. (1996).

33. Hull GA, Bies N, Twell D, and Delseny M (1996) Analysis of the promoter of an ABAresponsive late embryogenesis abundant gene of Arabidopsis thaliana. Plant Sci 114, 181-192.

32. Worrall D, Patel S, Lindsey K and Twell D (1996) A novel transient assay system demonstrates that DT-Atsm is a temperature-sensitive toxin in plant tissues. Plant Sci 113, 59-65.


31. Eady C, Lindsey K and Twell D (1995) The significance of microspore division and division symmetry for vegetative cell-specific transcription and generative cell differentiation. Plant Cell 7, 65-74.

30. Eady C, Twell D and Lindsey K (1995) Pollen viability and transgene expression following storage in honey. Transgenic Res 4, 226-231.

29. Hehn A, Bouzoubaa S, Marbach J, Bate N, Twell D, Richards K, Guilley H and Jonard G (1995) The small cysteine-rich protein P14 of beet necrotic yellow vein virus regulates accumulation of RNA 2 in cis and coat protein in trans. Virology 210, 73-81.

28. Martinussen I, Bate N, Weterings KAP, Junttila O and Twell D (1995). Analysis of gene regulation in growing pollen tubes of angiosperm and gymnosperm species using microprojectile bombardment. Physiol Plant 93, 445-450.

27. Twell D (1995) Diphtheria toxin-mediated cell ablation in developing pollen: vegetative cell ablation blocks generative cell migration. Protoplasma 187, 144-154.

26. Weterings K, Schrauwen J, Wullems G and Twell D (1995) Functional dissection of the promoter of the pollen-specific gene NTP303 reveals a novel pollen-specific, and conserved cis-regulatory element. Plant J 8, 55-63.


25. Eady C, Lindsey K and Twell D (1994) Differential activation and conserved vegetative-cell-specific activity of a late pollen promoter in species with bi- and tricellular pollen. Plant J 5, 543-550.

24. Martinussen I, Junttila O and Twell D (1994). Optimization of transient gene expression in pollen of Norway spruce (Picea abies) by particle acceleration. Physiol Plant 92, 412-416.

23. Turner R, Bate N, Twell D and Foster G (1994) Analysis of a translational enhancer upstream from the coat protein open reading frame of potato virus S Arch Virol 134, 321-333.

22. Turner R, Bate N, Twell D and Foster G (1994). In vivo characterisation of a translational enhancer upstream from the coat protein open reading frame of potato virus S  Arch Virol 137, 123-132.

21. Twell D (1994) The diversity and regulation of gene expression in the pathway of male gametophyte development. In: Scott RJ and Stead AD (eds.) Molecular and cellular aspects of plant reproduction. Seminar series; 55 (Society for Experimental Biology). Cambridge University Press, Cambridge. pp. 83-135.

20. Wilkinson JE, Twell D and Lindsey K (1994). Methanol does not specifically inhibit endogenous ß-glucuronidase (GUS) activity. Plant Sci 97, 61-67.

19. Worrall D and Twell D (1994). Pollen: Where ubiquitin is not required?. Bioessays 16, 873-875.


18. Twell D (1993) Gene regulation during pollen development. Lin Soc Palynol Spec Grp Newslet pp. 2-3.

17. Twell D, Patel S, Sorensen A, Roberts M, Scott R, Draper J and Foster G (1993). Activation and developmental regulation of an Arabidopsis anther-specific promoter in microspores and pollen of Nicotiana tabacum. Sex Plant Reprod 6, 217-224.


16. Twell D (1992). Use of a nuclear-targeted ß-glucuronidase fusion protein to demonstrate vegetative cell-specific gene expresion in developing pollen. Plant J 2, 887-892.


15. McCormick S, Twell D, Vancanneyt G and Yamaguchi J (1991). Molecular analysis of gene regulation and function during male gametophyte development. In: Jenkins G and Schuch W (eds.) Molecular Biology of Plant Development. Proceedings of SEB Symposium No. 45. Company of Biologists, Cambridge. pp. 229-244.

14. McCormick S, Yamaguchi J and Twell D (1991). Deletion analysis of pollen-expressed promoters. In Vitro Cell Dev Biol 27,  15-20.

13. Twell D, Klein TM and McCormick S (1991). Transformation of pollen by particle bombardment. In: Lindsey K (ed.) Plant Tissue Culture Manual: Fundamentals and Applications, Kluwer Academic Publishers, Dordrecht, Netherlands. pp. D2: 1-14.

12. Twell, D., Yamaguchi, J., Wing, R.A., Ushiba, J. and McCormick, S. (1991). Promoter analysis of three genes that are coordinately expressed during pollen development reveals pollen-specific enhancer sequences and shared regulatory elements. Genes Dev 5, 496-507.


11. Twell D, Yamaguchi J and McCormick S (1990). Pollen-specific expression in transgenic plants: coordinate regulation of two different promoters during microsporogenesis. Development 109, 705-713.


10. McCormick S, Twell D, Wing R, Ursin V, Yamaguchi J and Larabell S (1989). Anther-specific genes: molecular characterization and promoter analysis in transgenicplants. In: Lord E and Bernier G (eds.), Plant Reproduction: from Floral Induction to Pollination. Proceedings from the 12th Annual Symposium in Plant Physiology. pp. 128-136.

9. Twell D, Wing RA, Yamaguchi J and McCormick S (1989). Isolation and expression of an anther-specific gene from tomato. Mol Gen Genet 217, 240-245.

8. Twell D, Klein T, Fromm ME and McCormick S (1989).Transient expression of chimeric genes delivered into pollen by microprojectile bombardment. Plant Physiol 91, 1270-1274.


7. Twell D (1988). Genetic engineering of potato. Plants Today 1:191-196.

6. Twell D and Ooms G (1988) Structural diversity of the patatin gene family in potato cv. Desiree. Mol Gen Genet 212, 325-336.


5. Twell D and Ooms G (1987). The 5' flanking DNA of a patatin gene directs tuber-specific expression of a chimaeric gene in potato. Plant Mol Biol 9:345-375.


4. Ooms G, Twell D, Bossen ME, Hoge JH and Burrell MM (1986). Developmental regulation of Ri T-DNA gene expression in roots, shoots and tubers of transformed potato (Solanum tuberosum cv. Desiree). Plant Mol Biol 6, 321-330.


3. Burrell MM, Twell D, Karp A and Ooms G (1985). Expression of shoot-inducing Ti T-DNA in differentiated tissues of potato (Solanum tuberosum cv. Maris bard). Plant Mol Biol 5, 231-222.

2. Ooms G, Bains A, Burrell MM, Karp A, Twell D and Wilcox E (1985). Genetic manipulation in cultivars of oilseed rape (Brassica napus) using Agrobacterium. Theor Appl Genet 71, 325-329.

1. Ooms G, Karp A, Burrell MM, Twell D and Roberts J (1985). Genetic modification of potato development using Ri T-DNA. Theor Appl Genet 70, 440-446.

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