This review paper highlights about the role of salicyclic acid on somatic embryogenesis and also recent updates on cloning mature trees ofÃ‚Â  conifers using salicyclic acid were discussed. Salicyclic acid (SA) is a mobile molecule, which is capable of acting as a cell signal that senses, amplifies, and transmit information from a cell and might help in programmingÃ‚Â  towards embryogenesis during cloning. Very recently incorporation of 1.0 mg-1 SA in the DCR induction medium was found to be optimum for allÃ‚Â  the genotypes of P. roxburghii in terms of increasing the percentage of somatic embryogenesis compared against control. Therefore, salicyclic acid can be used as growth regulator in conifer somatic embryogenesis and its use might help to solve the low initiation frequencies of many other recalcitrant pines. However, the mechanism of salicylate-inducedÃ‚Â  differentiation in plants is not known, although salicylate is a signal molecule implicated in eliciting many physiological functions in plants

Kumar, S. Vijaya

Malabadi, Ravindra B.

Mulgund, Gangadhar S.

Nataraja, K.

T. Meti, Neelambika

English

Update Publishing (E-Journals)

 Research in Biotechnology, 3(2): 57-61, 2012                                                   ISSN: 2229-791X      www.researchinbiotechnology.com                                    Short Communication Role of salicyclic acid on conifer somatic embryogenesis    Gangadhar S. Mulgund1, Neelambika T. Meti2, Ravindra B. Malabadi*, K. Nataraja1,               S. Vijaya Kumar3  1Department of Botany, Karnatak University, Pavate nagar, Dharwad-580003,  Karnataka, India 2Department of Agricultural Biotechnology, Bharati Vidyapeeth University, Pune-Satara Road,Katraj, Pune - 411046, Maharashtra, India 3Department of Biotechnology, Madanapalle Institute of Technology and Science, Madanapalle-517325, Chitoor, Andhra Pradesh, India *Corresponding author e-mail: mlbd712@rediffmail.com      This review paper highlights about the role of salicyclic acid on somatic embryogenesis and also recent updates on cloning mature trees of conifers using salicyclic acid were discussed. Salicyclic acid (SA) is a mobile molecule, which is capable of acting as a cell signal that senses, amplifies, and transmit information from a cell and might help in programming towards embryogenesis during cloning. Very recently  incorporation of   1.0 mg-1 SA in the DCR induction medium was found to be optimum for all the genotypes of P. roxburghii in terms of increasing the percentage of somatic embryogenesis compared against control.   Therefore, salicyclic acid can be used as growth regulator in conifer somatic embryogenesis and its use might help to solve the low initiation frequencies of many other recalcitrant pines. However, the mechanism of salicylate-induced differentiation in plants is not known, although salicylate is a signal molecule implicated in eliciting many physiological functions in plants.     Key words: Cloning, conifers, mature trees, forestry, India, signaling  Abbreviations: ASA- acetylsalicyclic acid; DCR- Durzan and Gupta medium; H2O2- Hydrogen peroxide; PGR- Plant growth regulators; SA-salicyclic acid; TDZ-  thidiazuron;  tTCLs-transverse thin cell layers;  Overview The plant growth regulator, salicyclic acid (SA), when applied to plants, affects diverse physiological processes (Malabadi et al. 2008a, 2008b; Cleland and Ajami, 1974; Cleland et al. 1982; Conrath et al. 1995; Dean and Delaney, 2008; Durner and Klessig, 1995; Gutierrez-Coronado et al. 1998; Hao et al. 2006; Hutchinson and Saxena, 1996; Komaraiah et al. 2004; Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001).  In African violet, the application of SA at low concentrations affects plant size, and the number of leaves and flowers. Moreover, application of SA to the growth medium induced flowering in several species of Lemnaceae.  SA has also been reported to increase the activity of superoxide dismutase, and inhibit activities of ascorbate peroxidase, and catalase, thus leading to endogenous Mulgund et al. /Research in Biotechnology, 3(2): 57-61, 2012  58  H2O2 accumulation (Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001).  Thus, SA inhibits the decomposition of H2O2 produced in plants. SA is an important signaling molecule involved in plant defense responses to pathogens and abiotic stress, as well as in plant growth and development. There are several reports describing exogenous SA and acetylsalicyclic acid (ASA) enhancing somatic embryogenesis in plants viz. carrot (Daucus carota), pearl millet (Pennisetum americanum), geranium (Pelargonium × hortorum Bailey), Astragalus adsurgens Pall, Plumbago rosea L., naked oat (Avena nuda), and Pinus roxburghii. On the other hand salicyclic acid also promotes induction of flowering in duckweeds, increased cell division, and growth during somatic embryogenesis in tissue culture of Coffea arabica, stomatal closure in Phaseolus vulgaris, increase in the accumulation of nitrates in roots of Pinus patula, inhibition of the biosynthesis of ethylene in cell suspension cultures of pears and seed germination in species, and root growth stimulation of soybean (Malabadi et al. 2008a, 2008b; Cleland and Ajami, 1974; Cleland et al. 1982; Conrath et al. 1995; Dean and Delaney, 2008; Durner and Klessig, 1995; Gutierrez-Coronado et al. 1998; Hao et al. 2006; Hutchinson and Saxena, 1996; Komaraiah et al. 2004; Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001; Martin-Max et al. 2005; San Miguel et al. 2002; Raskin, 1992; Quiroz et al. 2001; Ping et al. 2001).  Role of salicyclic acid on somatic embryogenesis       SA is an important signaling molecule not only involved in defense responses but also in somatic embryogenesis of many plant species. SA is a mobile molecule, which is capable of acting as a cell signal that senses, amplifies, and transmit information from a cell and might help in programming towards embryogenesis during cloning. Secondly, SA is involved (together with nitrogen oxide, hydrogen peroxide, and other metabolites) in the function of several signal systems, unifying them into an intricate network of regulatory interactions. Perhaps embryo differentiation may sharing some of the intermediates in the salicylate signal pathway(Malabadi et al. 2008a, 2008b; Cleland and Ajami, 1974; Cleland et al. 1982; Conrath et al. 1995; Dean and Delaney, 2008; Durner and Klessig, 1995; Gutierrez-Coronado et al. 1998; Hao et al. 2006; Hutchinson and Saxena, 1996; Komaraiah et al. 2004; Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001; Martin-Max et al. 2005; San Miguel et al. 2002; Raskin, 1992; Quiroz et al. 2001; Ping et al. 2001). In a recent study, it was observed that SA acts as a signaling molecule during cloning mature trees of P. roxburghii. Therefore, SA can be used as growth regulator in conifer somatic embryogenesis and its use might help to solve the low initiation frequencies of many other recalcitrant pines.  Embryogenic cultures have been generated for most of the conifers. For the most part, however, even the best of these systems lack commercial viability for two reasons: first, a low frequency of regeneration for many of the most desirable clones; and secondly, unproven genotypes, as starting material for the cultures is derived from seeds or seedlings. Therefore, the current approach of cloning mature trees of conifers using SA has many practical applications particularly in clonal forestry schemes. Use of SA might be helpful in solving many problems of conifer somatic embryogenesis.  The pre-treatment of tTCLs from any of the 10 genotypes of P. roxburghii with 0.1, 0.2 and 0.4 mg-1 SA could not effectively increasing the percentage of somatic embryogenesis when compared to the control (Malabadi et al. 2008a, 2008b; Malabadi and Teixeira da Silva, 2011). Pretreatment of explants with higher concentrations (2.0-5.0 mg-1) of SA might have had a toxic effect and resulted in the browning of explants without callus Mulgund et al. /Research in Biotechnology, 3(2): 57-61, 2012  59  formation in all 10 genotypes.  Incorporation of 1.0 mg-1 SA in the induction medium was optimum for all 10 genotypes by increasing the percentage of somatic embryogenesis compared to the control. The addition of 1.0 mg-1 SA to the induction medium was very beneficial since in the control this genotype failed to induce somatic embryogenesis. This clearly indicates the positive role of SA as a signaling molecule during cloning of mature P. roxburghii trees. In this study SA alone (i.e. without PGRs) did not induce somatic embryogenesis and resulted in the browning of explants and callus. Therefore, SA when combined with 22.6 µM 2, 4-D, 26.8 µM NAA, 8.9 µM BAP in the induction medium, improved the percentage of somatic embryogenesis.  Hence, the combination of SA with other PGRs such as 2,4-D/NAA/BA might be beneficial in inducing somatic embryogenesis in Chir pine. In geranium (Pelargonium x hortorum Bailey), thidiazuron (TDZ) effectively induced somatic embryo-genesis in cultured hypocotyls explants during only a 3-day period of induction.  The presence of acetylsalicyclic acid (ASA) during this period caused a two-fold increase in the number of somatic embryos an enhanced synchronization of embryo development compared to the TDZ treatment alone. However, in the same study, SA was ineffective in modulating similar embryogenic responses as ASA in geranium. Enhanced somatic embryogenesis and plant regeneration have been obtained using young leaf bases of naked oat (Avena nuda) as explants by including 0.5 mM SA and carrot embryogenic callus extracts in MS media. An improvement was achieved in somatic embryogenesis and plant regeneration on the corresponding media supplemented with 0.5 mM SA and carrot embryogenic callus extracts as compared to control. Somatic embryogenesis was induced from suspension cultures derived from leaf callus of an important medicinal plant, Plumbago rosea L. in which 8.32 µM ASA alone induced embryogenesis, but IAA, NAA or IBA alone failed to elicit a similar response. Optimal embryogenic response per culture (216 embryos per culture) was observed in MS medium containing a combination of ASA (8.32 µM) and IAA (5.06 µM), i.e. a similar synergistic response as observed in our study between SA/ASA and other PGR(s) in the medium. It was also observed that by increasing the concentration of ASA alone (without auxin) in the medium (up to 11.09 µM) the number of somatic embryos formed per culture increased. The interactive effect of ASA and IAA appears to be essential for enhanced production of embryos per culture since no embryogenesis was noticed when IAA alone was added in Plumbago rosea L.  SA is endogenously produced in many plants in normal as well as during stress conditions.   However, the mechanism of salicylate-induced differentiation in plants is not known, although salicylate is a signal molecule implicated in eliciting many physiological functions in plants. It was also reported that the inclusion of SA to differentiation medium below 200 µmol/L significantly enhanced somatic embryo-genesis in Astragalus adsurgens Pall., callus cultures, the highest frequency of somatic embryogenesis occurring at 150 µmol/L SA. They also reported that enhanced somatic embryogenesis by SA was accompanied by an increase in the endogenous hydrogen peroxide (H2O2) level compared to controls (Malabadi et al. 2008a, 2008b; Cleland and Ajami, 1974; Cleland et al. 1982; Conrath et al. 1995; Dean and Delaney, 2008; Durner and Klessig, 1995; Gutierrez-Coronado et al. 1998; Hao et al. 2006; Hutchinson and Saxena, 1996;  Komaraiah et al. 2004; Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001; Martin-Max et al. 2005; San Miguel et al. 2002; Raskin, 1992; Quiroz et al. 2001; Ping et al. 2001).   This increased endogenous H2O2 level was related to the inhibition of ascorbate peroxidase and catalase activities. Although Mulgund et al. /Research in Biotechnology, 3(2): 57-61, 2012  60  the promoting effect of exogenous H2O2 was significantly lower than that of exogenous SA on the development of somatic embryos, the pre-treatment of callus cultures of A. adsurgens with dimethylurea (a trap for H2O2) significantly inhibited somatic embryo-genesis, even if callus was cultured on the differentiation medium supplemented with 150 µmol/L SA, suggesting that endogenous H2O2 was required for SA-enhanced somatic embryogenesis in A. adsurgens (Malabadi et al. 2008a, 2008b; Cleland and Ajami, 1974; Cleland et al. 1982; Conrath et al. 1995; Dean and Delaney, 2008;  Durner and Klessig, 1995; Gutierrez-Coronado et al. 1998; Hao et al. 2006; Hutchinson and Saxena, 1996; Komaraiah et al. 2004; Larque-Saavedra, 1978, 1979; Leslie and Ramani, 1988; Luo et al. 2001; Martin-Max et al. 2005; San Miguel et al. 2002; Raskin, 1992; Quiroz et al. 2001; Ping et al. 2001).  Therefore, one possible link between oxidative stress and plant regeneration in tissue culture could be H2O2. SA is endogenously produced in many plants in normal as well as during stress conditions. SA also inhibited ethylene biosynthesis in cell suspension cultures of carrot.   It is well known that ethylene inhibits differentiation in plants. Therefore, SA may be promoting embryo development by inhibiting ethylene biosynthesis. Another hypothesis is that SA has been reported to increase the activity of superoxide dismutase, and inhibits the activities of ascorbate peroxidase and catalases, thus leading to endogenous H2O2 accumulation in Arabidopsis thaliana.   References Cleland CF, Ajami A (1974) Identification of the flower-inducing factor isolated from aphid honeydew as being salicyclic acid. Plant Physiology 54, 909-906 Cleland CF, Tanaka O, Feldman IJ (1982) Influence of plant growth substances and salicyclic acid on flowering and growth in the Lemnaceae (duckweeds). Aquatic Botany 13, 3-20 Conrath U, Chen Z, Ricigliano JR, Klessig DF  (1995) Two inducers of plant defense responses, 2, 6-dichloroisonicotinic acid and salicyclic acid inhibit catalase activities in tobacco. Proceedings of  National Academy of  Sciences USA 92, 7143-7147 Dean JV, Delaney SP (2008) Metabolism of salicyclic acid in wild-type, ugt74f1 and ugt74f2 glucosyltransferase mutants of Arabidopsis thaliana. Physiologia Plantarum 132, 417-425 Durner J, Klessig DF  (1995) Inhibition of ascorbate peroxidase by salicyclic acid and 2, 6-dichloroisonicotinic acid, two inducers of plant defense responses. Proceedings of National Academy of  Sciences USA 92, 11312-11316 Gutierrez-Coronado M, Trejo CL, Larque-Saavedra A  (1998) Effects of salicyclic acid on the growth of roots and shoots in soybean. 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Role of salicyclic acid on conifer somatic embryogenesis

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Role of salicyclic acid on conifer somatic embryogenesis

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