Ukr.Biochem.J. 2014; Volume 86, Issue 5, Sep-Oct, pp. 26-36


Functions of tocopherols in the cells of plants and other photosynthetic organisms

V. M. Mokrosnop

M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kyiv;

Tocopherol synthesis has only been observed in photosynthetic organisms (plants, algae and some cyanobacteria). Tocopherol is synthesized in the inner membrane of chloroplasts and distributed between chloroplast membranes, thylakoids and plastoglobules. Physiological significance of tocopherols for human and animal is well-studied, but relatively little is known about their function in plant organisms. Among the best characterized functions of tocopherols in cells is their ability to scavenge and quench reactive oxygen species and fat-soluble by-products of oxidative stress. There are the data on the participation of different mechanisms of α-tocopherol action in protecting photosystem II (PS II) from photoinhibition both by deactivation of singlet oxygen produced by PS II and by reduction of proton permeability of thylakoid membranes, leading to acidification of lumen under high light conditions and activation of violaxanthin de-epoxidase. Additional biological activity of tocopherols, independent of its antioxidant functions have been demonstrated. Basic mechanisms for these effects are connected with the modulation of signal transduction pathways by specific tocopherols and, in some instances, by transcriptional activation of gene expression.

Keywords: , , , , , , , , ,


  1. Li Y, Hussain N, Zhang L, Chen X, Ali E, Jiang L. Correlations between tocopherol and fatty acid components in germplasm collections of Brassica oilseeds. J Agric Food Chem. 2013 Jan 9;61(1):34-40. PubMed, CrossRef
  2. Bellock O. New topics in vitamin E research. New York: Nova science publishers, 2007. 167 p.
  3. Preedy V, Watson R. The encyclopedia of vitamin E. London: CABI Publishing, 2007. 945 p.
  4. Munné-Bosch S. Alpha-tocopherol: a multifaceted molecule in plants. Vitam Horm. 2007;76:375-92. Review. PubMed, CrossRef
  5. Lichtenthaler HK. Biosynthesis, accumulation and emission of carotenoids, alpha-tocopherol, plastoquinone, and isoprene in leaves under high photosynthetic irradiance. Photosynth Res. 2007 May;92(2):163-79. Review. PubMed, CrossRef
  6. Szymańska R, Kruk J. Plastoquinol is the main prenyllipid synthesized during acclimation to high light conditions in Arabidopsis and is converted to plastochromanol by tocopherol cyclase. Plant Cell Physiol. 2010 Apr;51(4):537-45. Epub 2010 Feb 17. PubMed, CrossRef
  7. Havaux M, Eymery F, Porfirova S, Rey P, Dörmann P. Vitamin E protects against photoinhibition and photooxidative stress in Arabidopsis thaliana. Plant Cell. 2005 Dec;17(12):3451-69. Epub 2005 Oct 28. PubMed, PubMedCentral, CrossRef
  8. Munne-Bosch S, Alegre L. The function of tocopherols and tocotrienols in plants. Crit Rev Plant Sci. 2002;21(1):31-57. CrossRef
  9. Lichtenthaler HK, Prenzel U, Douce R, Joyard J. Localization of prenylquinones in the envelope of spinach chloroplasts. Biochim Biophys Acta. 1981 Feb 20;641(1):99-105. PubMed, CrossRef
  10. Szarka A, Tomasskovics B, Bánhegyi G. The ascorbate-glutathione-α-tocopherol triad in abiotic stress response. Int J Mol Sci. 2012;13(4):4458-83. PubMed, PubMedCentral, CrossRef
  11. Collin VC, Eymery F, Genty B, Rey P, Havaux M. Vitamin E is essential for the tolerance of Arabidopsis thaliana to metal-induced oxidative stress. Plant Cell Environ. 2008 Feb;31(2):244-57. PubMed, CrossRef
  12. Li Y, Wang Z, Sun X, Tang K. Current opinions on the functions of tocopherol based on the genetic manipulation of tocopherol biosynthesis in plants. J Integr Plant Biol. 2008 Sep;50(9):1057-69. Review. PubMed, CrossRef
  13.  Niki E, Noguchi N. Dynamics of antioxidant action of vitamin E. Acc Chem Res. 2004 Jan;37(1):45-51. PubMed, CrossRef
  14. Mène-Saffrané L, Jones AD, DellaPenna D. Plastochromanol-8 and tocopherols are essential lipid-soluble antioxidants during seed desiccation and quiescence in Arabidopsis. Proc Natl Acad Sci USA. 2010 Oct 12;107(41):17815-20. PubMed, PubMedCentral, CrossRef
  15. Gill SS, Tuteja N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiol Biochem. 2010 Dec;48(12):909-30. Review. PubMed, CrossRef
  16. Abbasi A. Functional analysis of tocopherol biosynthesis in plants. Aus Tehran: IRI, 2007. 171 p.
  17. Sattler SE, Gilliland LU, Magallanes-Lundback M, Pollard M, DellaPenna D. Vitamin E is essential for seed longevity and for preventing lipid peroxidation during germination. Plant Cell. 2004 Jun;16(6):1419-32. PubMed, PubMedCentral, CrossRef
  18. Kruk J, Holländer-Czytko H, Oettmeier W, Trebst A. Tocopherol as singlet oxygen scavenger in photosystem II. J Plant Physiol. 2005 Jul;162(7):749-57. Review. PubMed, CrossRef
  19. Krieger-Liszkay A, Trebst A. Tocopherol is the scavenger of singlet oxygen produced by the triplet states of chlorophyll in the PSII reaction centre. J Exp Bot. 2006;57(8):1677-84. Epub 2006 May 19. Review. PubMed, CrossRef
  20. Kruk J., Strzalka K. Occurrence and function of α-tocopherol quinine in plants. J Plant Physiol. 1995;145(4):405-409. CrossRef
  21. Li Z, Keasling JD, Niyogi KK. Overlapping photoprotective function of vitamin E and carotenoids in Chlamydomonas. Plant Physiol. 2012 Jan;158(1):313-23. PubMed, PubMedCentral, CrossRef
  22. Inoue S, Ejima K, Iwai E, Hayashi H, Appel J, Tyystjärvi E, Murata N, Nishiyama Y. Protection by α-tocopherol of the repair of photosystem II during photoinhibition in Synechocystis sp. PCC 6803. Biochim Biophys Acta. 2011 Feb;1807(2):236-41. PubMed, CrossRef
  23. Trebst A, Depka B, Holländer-Czytko H. A specific role for tocopherol and of chemical singlet oxygen quenchers in the maintenance of photosystem II structure and function in Chlamydomonas reinhardtii. FEBS Lett. 2002 Apr 10;516(1-3):156-60. PubMed, CrossRef
  24. Sakuragi Y, Maeda H, Dellapenna D, Bryant DA. alpha-Tocopherol plays a role in photosynthesis and macronutrient homeostasis of the cyanobacterium Synechocystis sp. PCC 6803 that is independent of its antioxidant function. Plant Physiol. 2006 Jun;141(2):508-21. PubMedPubMedCentral, CrossRef
  25. Maeda H, DellaPenna D. Tocopherol functions in photosynthetic organisms. Curr Opin Plant Biol. 2007 Jun;10(3):260-5. Review. PubMed, CrossRef
  26. Quadrana L, Almeida J, Otaiza S, Duffy T, Correa da Silva J, de Godoy F, Asis R, Bermudez L, Fernie A, Carrari F, Rossi M. Transcriptional regulation of tocopherol biosynthesis in tomato.  Plant Mol Biol. 2013 Feb;81(3):309-25. PubMed, CrossRef
  27. Mène-Saffrané L, DellaPenna D. Biosynthesis, regulation and functions of tocochromanols in plants. Plant Physiol Biochem. 2010 May;48(5):301-9. Review. PubMed, CrossRef
  28. Hofius D, Hajirezaei MR, Geiger M, Tschiersch H, Melzer M, Sonnewald U. RNAi-mediated tocopherol deficiency impairs photoassimilate export in transgenic potato plants. Plant Physiol. 2004 Jul;135(3):1256-68. PubMed, PubMedCentral, CrossRef
  29. Botha CEJ, Cross RHM, van Bel AJE, Peter CI. Phloem loading in the sucrose-export-defective (SXD-1) mutant maize is limited by callose deposition at plasmodesmata in bundle sheath-vascular parenchyma interface. Protoplasma. 2000 Mar;214(1-2):65-72. CrossRef
  30. Sattler SE, Cahoon EB, Coughlan SJ, DellaPenna D. Characterization of tocopherol cyclases from higher plants and cyanobacteria. Evolutionary implications for tocopherol synthesis and function. Plant Physiol. 2003 Aug;132(4):2184-95. PubMed, PubMedCentral, CrossRef
  31. Bergmüller E, Porfirova S, Dörmann P. Characterization of an Arabidopsis mutant deficient in gamma-tocopherol methyltransferase. Plant Mol Biol. 2003 Aug;52(6):1181-90. PubMed, CrossRef
  32. Havaux M, Eymery F, Porfirova S, Rey P, Dörmann P. Vitamin E protects against photoinhibition and photooxidative stress in Arabidopsis thaliana. Plant Cell. 2005 Dec;17(12):3451-69. PubMed, PubMedCentral, CrossRef
  33. Maeda H, Song W, Sage TL, DellaPenna D. Tocopherols play a crucial role in low-temperature adaptation and Phloem loading in Arabidopsis. Plant Cell. 2006 Oct;18(10):2710-32. PubMed, PubMedCentral, CrossRef
  34. Munné-Bosch S. Alpha-tocopherol: a multifaceted molecule in plants. Vitam Horm. 2007;76:375-92. Review. PubMed, CrossRef
  35. Sattler SE, Mène-Saffrané L, Farmer EE, Krischke M, Mueller MJ, DellaPenna D. Nonenzymatic lipid peroxidation reprograms gene expression and activates defense markers in Arabidopsis tocopherol-deficient mutants. Plant Cell. 2006 Dec;18(12):3706-20. PubMed, PubMedCentral, CrossRef
  36. Maeda H, Sakuragi Y, Bryant DA, Dellapenna D. Tocopherols protect Synechocystis sp. strain PCC 6803 from lipid peroxidation. Plant Physiol. 2005 Jul;138(3):1422-35.  PubMed, PubMedCentral, CrossRef
  37. Munne-Bosch S. Linking tocopherols with cellular signaling in plants. New Phytol. 2005 May;166(2):363-6. PubMed, CrossRef
  38. Munné-Bosch S, Falk J. New insights into the function of tocopherols in plants. Planta. 2004 Jan;218(3):323-6. Review. PubMed, CrossRef
  39. Lushchak VI, Semchuk NM. Tocopherol biosynthesis: chemistry, regulation and effects of environmental factors. Acta Physiol Plant. 2012;34(5): 1607-28.  CrossRef
  40. Azzi A, Stocker A. Vitamin E: non-antioxidant roles. Prog Lipid Res. 2000 May;39(3):231-55. Review. PubMed, CrossRef
  41. Sandorf I, Holländer-Czytko H. Jasmonate is involved in the induction of tyrosine aminotransferase and tocopherol biosynthesis in Arabidopsis thaliana. Planta. 2002 Nov;216(1):173-9.  PubMed, CrossRef
  42. Schaller F. Enzymes of the biosynthesis of octadecanoid-derived signalling molecules. J Exp Bot. 2001 Jan;52(354):11-23. Review. PubMed, CrossRef
  43. Badrhadad A., Piri K., Ghiasvand T. Increasing of alpha-tocopherol in cell suspension cultures Elaeagnus angustifolia L. Intl  J Agri Crop Sci. 2013;5(12):1328-1331.
  44. Kayal W, Keller G, Debayles C, Kumar R, Weier D, Teulieres C, Marque C. Regulation of tocopherol biosynthesis through transcriptional control of tocopherol cyclase during cold hardening in Eucalyptus gunnii. Physiol. Plantarum. 2006;126:212-223. CrossRef

Creative CommonsThis work is licensed under a Creative Commons Attribution 4.0 International License.