Photosynthetica, 2011 (vol. 49), issue 3

Photosynthetica 2011, 49(3):448 | DOI: 10.1007/s11099-011-0057-4

Responses in the physiology and biochemistry of Korean pine (Pinus koraiensis) under supplementary UV-B radiation

Y. G. Zu1, X. X. Wei2, J. H. Yu1, D. W. Li1,*, H. H. Pang1, L. Tong1
1 Key Laboratory of Forest Plant Ecology, Ministry of Education, Northeast Forestry University, Harbin, China
2 Institute of Volcano and Spring, Heilongjiang Academy of Sciences, Wudalianchi, China

The effect of supplementary UV-B radiation on Korean pine (Pinus koraiensis Sieb. et Zucc) was investigated. Compared with the control, the T1, T2, and T3 UV-B treatments increased by 1.40, 2.81, and 4.22 kJ m-2 d-1, respectively. Gas-exchange parameters, photosynthetic pigment concentrations, contents of secondary metabolites, epicuticular wax, free radical, malondialdehyde (MDA), and the activities of antioxidant enzymes were determined after 40 d of exposure. The concentrations of chlorophyll (Chl) a, Chl b, total Chl, carotenoid (Car), and the ratio Chl a/b in the pine needles were in the following order: T1 > T2 > T3. Compared with the control, the contents of flavonoids and epicuticular wax significantly decreased in all levels of supplementary UV-B radiations (p<0.05). Moreover, the contents of hydrogen peroxide (H2O2) and MDA significantly increased with the enhanced UV-B radiations (p<0.05). Korean pine can increase the catalase, ascorbate peroxidase, and superoxide dismutase activities to prevent oxidative stress by supplementary UV-B radiation. However, its defence mechanism is not efficient enough to prevent UV-Binduced damage.

Keywords: epicuticular wax; Korean pine; photosynthesis; UV-B radiation

Received: August 17, 2010; Accepted: June 12, 2011; Published: September 1, 2011Show citation

ACS AIP APA ASA Harvard Chicago IEEE ISO690 MLA NLM Turabian Vancouver
Zu, Y.G., Wei, X.X., Yu, J.H., Li, D.W., Pang, H.H., & Tong, L. (2011). Responses in the physiology and biochemistry of Korean pine (Pinus koraiensis) under supplementary UV-B radiation. Photosynthetica, 49(3), 448. doi: 10.1007/s11099-011-0057-4.
Download citation
  • BibTeX (.bib)
  • Bookends (.ris)
  • EasyBib (.ris)
  • EndNote (.enw)
  • EndNote 8 (.xml)
  • ISI WoS (.isi)
  • Medlars (.medlars)
  • Mendeley (.ris)
  • MODS (.xml)
  • Papers (.ris)
  • RefWorks (.txt)
  • RefManager (.ris)
  • RIS (.ris)
  • MS Word (.xml)
  • Zotero (.ris)
    • Open full article

    References

    1. Ahmad, P., Sarwat, M., Sharma, S.: Reactive oxygen species, antioxidants and signaling in plants. - J. Plant. Biol. 51: 167-173, 2008. Go to original source...
    2. Asada, K.: Ascorbate peroxidase: a hydrogen peroxide-scavenging enzyme in plants. - Physiol. Plant. 85: 235-241, 1992. Go to original source...
    3. Barnes, J.D., Percy, K.E., Paul, N.D., Jones, P., McLaughlin, C.K., Mullineaux, P.M., Creissen, G., Wellburn, A.R.: The influence of UV-B radiation on the physicochemical nature of tobacco (Nicotiana tabacum L.) leaf surfaces. - J. Exp. Bot. 47: 99-109, 1996. Go to original source...
    4. Baumbusch, L.O., Eiblmeier, M., Schnitzler, J.-P., Heller, W., Sandermann, H., Polle, A.: Interactive effects of ozone and low UV-B radiation on antioxidants in spruce (Picea abies) and pine (Pinus sylvestris) needles. - Physiol. Plant. 104: 248-254, 1998. Go to original source...
    5. Bornman, J.F., Reuber, S., Cen, Y.-P., Weissenbock, G.: Ultraviolet radiation as a stress factor and the role of protective pigments. - In: Lumbsden, P.J. (ed.): Plants and UV-B: Responses to Environmental Change. Pp. 157-171. Cambridge Univ. Press, Cambridge 1997. Go to original source...
    6. Caldwell, M.M., Björn, L.O., Bornman, J.F., Flinta, S.D., Kulandaivelu, G., Teramura, A.H., Tevini, M.: Effects of increased solar ultraviolet radiation on terrestrial ecosystem. - J. Photochem. Photobiol. B: Biol. 46: 40-52, 1998. Go to original source...
    7. Caldwell, M.M.: Solar UV irradiation and the growth and development of higher plants. - In: Giese, A.C. (ed.): Photobiology. Pp. 131-177. Academic Press, New York 1971. Go to original source...
    8. Clark, J.B., Lister, G.R.: Photosynthetic action spectra of trees: II. The relationship of cuticle structure to the visible and ultraviolet spectral properties of needles from four coniferous species. - Plant Physiol. 5: 407-413, 1975. Go to original source...
    9. Creelman, R.A., Mulletl, J.E.: Oligosaccharins, brassinolides, and jasmonates: nontraditional regulators of plant growth, development, and gene expression. - Plant Cell. 9: 1211-1223, 1997. Go to original source...
    10. Dai, Q.-J., Yan, B., Huang, S.-B., Liu, X.-Z., Peng, S.-B., Miranda, M.L.L., Chavez, A.Q., Vergara, B.S., Olszyk, D.M.: Response of oxidative stress in rice (Oryza sativa) leaves with supplemental UV-B radiation. - Physiol. Plant. 101: 301-308, 1997. Go to original source...
    11. Day, T.A., Howells, B.W., Ruhland, C.T.: Changes in growth and pigment concentrations with leaf age in pea under modulated UV-B radiation field treatments. - Plant Cell Environ. 19: 101-108, 1996. Go to original source...
    12. Day, T.A., Martin, G., Vogelmann, T.C.: Penetration of UV-B radiation in foliage: evidence that the epidermis behaves as a non-uniform filter. - Plant Cell Environ. 16: 735-741, 1993. Go to original source...
    13. Day, T.A., Ruhland, C.T., Grobe, C.W., Xiong, F.: Growth and reproduction of Antarctic vascular plants in response to warming and UV radiation reductions in the field. - Oecologia 119: 24-35, 1999. Go to original source...
    14. Day, T.A.: Relating UV-B radiation screening effectiveness of foliage to absorbing-compound concentration and anatomical characteristics in a diverse group of plants. - Oecologia 95: 542-550, 1993. Go to original source...
    15. De Zwart, L.L., Meerman, J.H.N., Commandeur, J.N.M., Vermeulen, N.P.E.: Biomarkers of free radical damage applications in experimental animals and in humans. - Free Radic. Biol. Med. 26: 202-226, 1999. Go to original source...
    16. Deckmyn, G., Martens, C., Impens, I.: The importance of the ratio UV-B/photosynthetic active radiation (PAR) during leaf development as determining factor of plant sensitivity to increased UV-B irradiance: effects on growth, gas exchange and pigmentation of bean plants (Phaseolus vulgaris cv. Label). - Plant Cell Environ. 17: 295-301, 1994. Go to original source...
    17. DeLucia, E.H., Day, T.A., Vogelmann, T.C.: Ultraviolet-B and visible light penetration into needles of two species of subalpine conifers during foliar development. - Plant Cell Environ. 15: 921-929, 1992. Go to original source...
    18. Demmig-Adams B., Adams, W.W., III: Photoprotection and other responses of plants to high light stress. - Annu. Rev. Plant Physiol. Plant Mol. Biol. 43: 599-626, 1992. Go to original source...
    19. Esch, A., Mengel, K.: Combined effects of acid mist and frost drought on the water status of young spruce trees (Picea abies). - Environ. Exp. Bot. 39: 57-65, 1998. Go to original source...
    20. Flint, S.D., Jordan, P.W., Caldwell, M.M.: Plant protective response to enhanced UV-B radiation under field conditions: leaf optical properties and photosynthesis. - Photochem. Photobiol. 41: 95-99, 1985. Go to original source...
    21. Flint, S.D., Searles, P.S., Caldwell, M.M.: Field testing of biological spectral weighting functions for induction of UV-absorbing compounds in higher plants. - Photochem. Photobiol. 79: 399-403, 2004. Go to original source...
    22. Forman, H.J., Fisher, A.B.: Antioxidant defenses. - In: Gilbert, D.L. (ed.): Oxygen and living processes: an interdiscriplinary approach. Pp. 235-249. Springer-Verlag, Berlin 1981. Go to original source...
    23. Foyer, C.H., Lopez-Delgado, H., Dat, J.F., Scott, I.M.: Hydrogen peroxide and glutathione-associated mechanism of acclimatory stress tolerance and signaling. - Physiol. Plant. 100: 241-254, 1997. Go to original source...
    24. Germ, M., Kreft, I., Osvald, J.: Influence of UV-B exclusion and selenium treatment on photochemical efficiency of photosystem II, yield and respiratory potential in pumpkins (Cucurbita pepo L.). - Plant Physiol. Biochem. 43: 445-448, 2005. Go to original source...
    25. Gómez, I., Pérez-Rodríguez, E., Viñegla, B., Figueroa, F.L., Karsten, U.: Effects of solar radiation on photosynthesis, UV-absorbing compounds and enzyme activities of the green alga Dasycladus vermicularis from southern Spain. - J. Photochem. Photobiol. B: Biol. 47: 46-57, 1998. Go to original source...
    26. Gu, J., Chen, Z.-Y., Zi, X.-N., Luo, L.-Q.: [Response mechanism of plant enzymatic system to UV-B radiation.] - J. Ecol. 25: 1269-1274, 2006. [In Chin.]
    27. He, J.-M., Xu, H., She, X.-P., Song, X.-G., Zhao, W.-M.: The role and the interrelationship of hydrogen peroxide and nitric oxide in the UV-B-induced stomatal closure in broad bean. - Funct. Plant Biol. 32: 237-247, 2005. Go to original source...
    28. Heath, R.L., Packer, L.: Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation. - Arch. Biochem. Biophys. 125: 189-198, 1968. Go to original source...
    29. Heinonen, I.M., Meyer, A.S., Frankel, E.N.: Antioxidant activity of berry phenolics on human low-density lipoprotein and liposome oxidation. - J. Agric. Food Chem. 46: 4107-4112, 1998. Go to original source...
    30. Hideg, E., Vass, I.: UV-B induced free radical production in plant leaves and isolated thylakoids. - Plant Sci. 115: 251-260, 1996. Go to original source...
    31. Hodges, D.M., Delong, J.M., Foney C.F., Prange, R.K.: Improving the thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds. - Planta 207: 604-611, 1999. Go to original source...
    32. Ibañez, S., Rosa, M., Hilal, M., Gonzalez, J.A., Prado, F.E.: Leaves of Citrus aurantifolia exhibit a different sensibility to solar UV-B radiation according to development stage in relation to photosynthetic pigments and UV-B absorbing compounds production. - J. Photochem. Photobiol. B: Biol. 90: 163-169, 2008.
    33. Jain, K., Kataria, S., Guruprasad, K.N.: Effect of UV-B radiation on antioxidant enzymes and its modulation by benzoquinone and α-tocopherol in cucumber cotyledons. - Curr. Sci. India. 87: 87-90, 2004.
    34. Jenks, M.A., Tuttle, H.A., Eigenbrode, S.D., Feldmann, K.A.: Leaf epicuticular waxes of the eceriferum mutants in Arabidopsis. - Plant Physiol. 108: 369-377, 1995. Go to original source...
    35. Jeremy, D. B., Kevin, E. P., Nigel, D. P., Pam, J., Chris, K. M., Phil, M. M., Gary, C., Alan, R. W.: The influence of UV-B radiation on the physicochemical nature of tobacco (Nicotiana tabacum L.) leaf surfaces. - J. Exp. Bot., 47: 99-109, 1996.
    36. Jiang, M.-Y., Zhang, J.-H.: Water stress-induced abscisic acid accumulation triggers the increased generation of reactive oxygen species and up-regulates the activities of antioxidant enzymes in maize leaves. - J. Exp. Bot. 53: 2401-2410, 2002. Go to original source...
    37. Kinnunen, H., Huttunen, S., Laakso, K.: UV-absorbing compounds and waxes of Scots pine needles during a third growing season of supplemental UV-B. - Environ. Pollut. 112: 215-220, 2001. Go to original source...
    38. Kinnunen, H., Laakso, K., Huttunen, S.: Methanol-extractable UV-B-absorbing compounds in Scots pine needles. - Chemosphere Global Change Sci. 1: 455-460, 1999. Go to original source...
    39. Kolattukudy, P.E.: Cutin, suberin and waxes. - In: Stumpf, P.K., Conn, E.E. (ed.): The Biochemistry of Plants. Pp. 571-645. Academic Press, New York 1980. Go to original source...
    40. Koti, S., Reddy, K.R., Kakani, V.G., Zhao, D.-L., Gao, W.: Effects of carbon dioxide temperature and ultraviolet-B radiation and their interactions on soybean (Glycine max L.) growth and development. - Environ. Exp. Bot. 60: 1-10, 2007. Go to original source...
    41. Kunst, L., Samuels, A.L.: Biosynthesis and secretion of plant cuticular wax. - Prog. Lipid Res. 42: 51-80, 2003. Go to original source...
    42. Laakso, K., Sullivan, J.H., Huttunen, S.: The effects of UV-B radiation on epidermal anatomy in loblolly pine (Pinus taeda L.) and Scots pine (Pinus sylvestris L.). - Plant Cell Environ. 23: 461-472, 2000. Go to original source...
    43. Lavola, A., Aphalo, P.J., Lahti, M., Julkunen-Tiitto, R.: Nutrient availability and the effect of increasing UV-B radiation on secondary plant compounds in Scots pine. - Environ. Exp. Bot. 49: 49-60, 2003. Go to original source...
    44. Li, J.-W., Ge, J.-P., Ma, J.-L., Chen, D.: The ecology and management of Korean pine mixed forest in the Northeast China Harbin. - The Press of North-East Forestry University. Harbin 1997. [In Chin.]
    45. Li, P., Li, Q.-F., Huang, Y.-Y., Huang, R.-D.: [The advance of plant flavonoids of anti-UV-B radiation research.] - J. Ecol. 20: 36-40, 2001. [In Chin.]
    46. Lu, Y.-W., Duan, B.-L., Zhang, X.-L., Korpelainen, H., Li, C.- Y.: Differences in growth and physiological traits of Populus cathayana populations as affected by enhanced UV-B radiation and exogenous ABA. - Environ. Exp. Bot. 66: 100-109, 2009. Go to original source...
    47. Mackerness, S.A.-H., John, C.F., Jordan, B.R., Thomas. B.: Early signaling components in ultraviolet-B response: Distinct roles for different reactive oxygen species and nitric oxide. - FEBS Lett. 489: 237-242, 2001. Go to original source...
    48. Malanga, G., Puntarulo, S.: Oxidative stress and antioxidant content in Chlorella vulgaris after exposure to ultraviolet-B radiation. - Physiol. Plant. 94: 672-679, 1995. Go to original source...
    49. Manetas, Y., Petropoulou, Y., Stamatakis, K., Nikolopoulos, D., Levizou, E., Psaras, G., Karabourniotis, G.: Beneficial effects of enhanced UV-B radiation under field conditions: improvement of needle water relations and survival capacity of Pinus pinea L. seedlings during the dry Mediterranean summer. - Plant Ecol. 128: 101-108, 1997. Go to original source...
    50. Mishra, V., Srivastava, G., Prasad, S.M.: Antioxidant response of bitter gourd (Momordica charantia L.) seedlings to interactive effect of dimethoate and UV-B irradiation. - Sci. Hortic-Amsterdam 120: 373-378, 2009. Go to original source...
    51. Patterson, B.D., Macrae, E.A., Ferguson, I.B.: Estimation of hydrogen peroxide in plant extracts using titanium (IV). - Anal. Biochem. 139: 487-492, 1984. Go to original source...
    52. Percy, K.E., McQuattie, C.J., Rebbeck, J.A.: Effects of air pollutants on epicuticular wax chemical composition. - In: Percy, K.E., Cape, J.N., Jagels, R., Simpson, C.J. (ed.): Air Pollutants and the Leaf Cuticle. Pp. 67-79. NATO ASI Series, Springer-Verlag, Berlin 1994. Go to original source...
    53. Petropoulou, Y., Kyparissis, A., Nikopoulos, D., Manetas, Y.: Enhanced UV-B radiation alleviates the adverse effects of summer drought in two Mediterranean pines under field conditions. - Physiol. Plant. 94: 37-44, 1995. Go to original source...
    54. Pilon, J.J., Lambers, H., Baas, W., Tosserams, M., Rozema, J., Atkin, O.K.: Leaf waxes of slow-growing alpine and fastgrowing lowland Poa species: inherent differences and responses to UV-B radiation. - Phytochem. 50: 571-580, 1999. Go to original source...
    55. Prasad, S.M., Kumar, D., Zeeshan, M.: Growth, photosynthesis, active oxygen species and antioxidants responses of paddy field cyanobacterium Plectonema boryanum to endosulfan stress. - J. Gen. Appl. Microbiol. 51: 115-123, 2005. Go to original source...
    56. Pyle, J.A.: Global ozone depletion: observations and theory. - In: Lumbsden, P.J. (ed.): Plants and UV-B: Responses to Environmental Change. Pp. 3-12. Cambridge Univ. Press, Cambridge 1997. Go to original source...
    57. Rao, M.V., Ormrod, D.P.: Impact of UV-B and O3 on the oxygen free radical scavenging system in Arabidopsis thaliana genotypes differing in flavonoid biosynthesis. - Photochem. Photobiol. 62: 719-726, 1995. Go to original source...
    58. Saile-Mark, M., Tevini, M.: Effects of solar UV-B radiation on growth, flowering and yield of central and southern European bush bean cultivars (Phaseolus vulgaris L.). - Plant Ecol. 128: 115-125, 1997. Go to original source...
    59. Sakaki, T., Kondo, N., Sugahara, K.: Breakdown of photosynthetic pigments and lipid in spinach leaves with ozone fumigation, role of active oxygen. - Physiol. Plant. 59: 28-34, 1983. Go to original source...
    60. Schnitzler, J.-P., Jungblut, T.P., Feicht, C., Köfferlein, M., Langebartels, C., Heller, W., Sandermann, H.: UV-B induction of flavonoid biosynthesis in Scots pine (Pinus sylvestris L.) seedlings. - Trees-Struct. Funct. 11: 162-168, 1997.
    61. Schuck, H.J.: [Quantitative variation der wachsauflage und der cuticularen kohlenwasserstoffe bei Picea abies Nadeln.] - Flora (Jena) 165: 303-314, 1976. [In German.] Go to original source...
    62. Schulze, E.D., Eller, B.M., Thomas, D.A., Willert, D.J., Brinckmann, E.: Leaf temperature and energy balance of Welwitschia mirabilis in its natural habitat. - Oecologia 44: 258-262, 1980. Go to original source...
    63. Song, Y.-X., Guo, S.-H., Ma, H.-A.: [Observation of leaf epidermis of 15 species shrubsin Helanshan mountain by SEM.] - Acta Bot. Boreal. Occident. Sin. 23: 1283-1287, 2003. [In Chin.]
    64. Šprtová, M., Marek, M.V., Nedbal, L., Prášil, O., Kalina, J.: Seasonal changes of photosynthetic assimilation of Norway spruce under the impact of enhanced UV-B radiation. - Plant Sci. 142: 37-45, 1999. Go to original source...
    65. Steiner, M.G., Babbs, C.F.: Quantitation of the hydroxyl radical by reaction with dimethyl sulfoxide. - Arch. Biochem. Biophys. 278: 478-481, 1990. Go to original source...
    66. Stewart, J.D., Hoddinott, J.: Photosynthetic acclimation to elevated atmospheric carbon dioxide and UV irradiation in Pinus banksiana. - Physiol. Plant. 88: 493-500, 1993. Go to original source...
    67. Sullivan, J.H., Teramura, A.H.: Effects of ultraviolet-B radiation on seedling growth in the Pinaceae. - Am. J. Bot. 75: 225-230, 1988. Go to original source...
    68. Surabhi, G.K., Reddy, K.R., Singh, S.K.: Photosynthesis, fluorescence, shoot biomass and seed weight responses of three cowpea (Vigna unguiculata (L.) Walp.) cultivars with contrasting sensitivity to UV-B radiation. - Environ. Exp. Bot. 66: 160-171, 2009. Go to original source...
    69. Swanson, A., Fahselt, D.: Effects of ultraviolet on polyphenolics of Umbilicaria americana. - Can. J. Bot. 75: 284-289, 1997. Go to original source...
    70. Teramura, A.H.: Effects of ultraviolet-B radiation on the growth and yield of crop plants. - Physiol. Plant. 58: 415-427, 1983. Go to original source...
    71. Tevini, M., Steinmueller, D.: Influence of light, UV-B radiation, and herbicides on wax biosynthesis of cucumber seedlings. - J. Plant. Physiol. 131: 111-121, 1987. Go to original source...
    72. Turtol, S., Sallas, L., Holopainen, J.K., Julkunen-Tiitto, R., Kainulainen, P.: Long-term exposure to enhanced UV-B radiation has no significant effects on growth or secondary compounds of outdoor-grown Scots pine and Norway spruce seedlings. - Environ. Pollut. 144: 166-171, 2006. Go to original source...
    73. Vagish, M., Garima, S., Sheo, M.P., Gerard, A.: Growth, photosynthetic pigments and photosynthetic activity during seedling stage of cowpea (Vigna unguiculata) in response to UV-B and dimethoate. - Pestic. Biochem. Physiol. 92: 30-37, 2008.
    74. Vogt, T., Gul, P.G.: Accumulation of avonoids during leaf development in Cistus laurifolius. - Phytochem. 36: 591-597, 1994. Go to original source...
    75. Wang, S.-W., Xie, B.-T., Yin, L.-N., Duan, L.-S., Li, Z.-H., Eneji, A.E., Tsuji, W., Tsunekawa, A.: Increased UV-B radiation affects the viability, reactive oxygen species accumulation and antioxidant enzyme activities in Maize (Zea mays L.) pollen. - Photochem. Photobiol. 86: 110-116, 2010. Go to original source...
    76. Wellburn, A.R.: The spectral determination of chlorophylls a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resolution. - J. Plant. Physiol. 144: 307-313, 1994. Go to original source...
    77. Wu, Z.-L.: [The classification and distribution of Pinus.] - J. Syst. Evol. 5: 131-163, 1956. [In Chin.]
    78. Xu, D.-Y., Yan, H.: A study of the impacts of climate change on the geographic distribution of Pinus koraiensis in China. - Environ. Int. 27: 201-205, 2001. Go to original source...
    79. Yamamoto, H.Y., Bassi, R.: Carotenoids: localization and function. - In: Ort, R.R., Yokum, C.F. (ed.): Oxygenic Photosynthesis. The Light Reactions. Pp. 539-563. Kluwer Academic Publishers, Dordrecht 1996. Go to original source...
    80. Yazici, I., Türkan, I., Sekmen, A.H., Demiral, T.: Salinity tolerance of purslane (Portulaca oleracea L.) is achieved by enhanced antioxidative system, lower level of lipid peroxidation and proline accumulation. - Environ. Exp. Bot. 61: 49-57, 2007. Go to original source...
    81. Ye, Z.-P.: A new model for relationship between irradiance and the rate of photosynthesis in Oryza sativa. - Photosynthetica 45: 637-640, 2007. Go to original source...
    82. Zancan, S., Suglia, I., Rocca, N.L., Ghisi, R.: Effects of UV-B radiation on antioxidant parameters of iron-deficient barley plants. - Environ. Exp. Bot. 63: 71-79, 2008. Go to original source...
    83. Zhao, D.-L., Reddy, K.R., Kakani, V.G., Mohammed, A.R., Read, J.J., Gao, W.: Leaf and canopy photosynthetic characteristics of cotton (Gossypium hirsutum) under elevated CO2 concentration and UV-B radiation. - J. Plant. Physiol. 161: 581-590, 2004. Go to original source...
    84. Zu, Y.-G., Pang, H.-H., Yu, J.-H., Li, D.-W., Wei, X.-X., Gao, Y.-X., Tong, L.: Responses in the morphology, physiology and biochemistry of Taxus chinensis var. mairei grown under supplementary UV-B radiation. - J. Photochem. Photobiol. B: Biol. 98: 152-158, 2010. Go to original source...

    Return to the content