Phytochemical changes and growth performance of
Thymus daenensis Celak. essential oil under the influence of light and salicylic acid
Subject Areas :
Medicinal Plants
Leila Abdi
1
,
Hamid reza Asghari
2
,
majid Tolyat Abolhassan
3
,
Mohammad Rea Amerian
4
,
Hassanali Naghdi Badi
5
1 - phD student, Faculty of Agriculture, Shahrood University of Technology, Shahrood, Iran,
2 - Associate professor, Faculty of Agriculture, Shahrood University of Technology, Shahrood, Iran
3 - Assistant professor, Medicinal Plants Research Center, Institute of Medicinal Plants, ACECR, Karaj, Iran
4 - Associate professor, Faculty of Agriculture, Shahrood University of Technology, Shahrood, Iran
5 - Associate professor, Iranian Biolocal Resource Center, Institute of Medicinal Plants, ACECR, Karaj, Iran
Received: 2021-11-02
Accepted : 2022-03-22
Published : 2022-11-21
Keywords:
Thymus daenensis,
Salicylic acid,
Carvacrol,
Thymol,
Light,
Essential oil,
Abstract :
In this study, to investigate the phytochemical changes of essential oil and growth performance of Thymus daenensis Celak. affected by light and salicylic acid, an experiment was conducted as a split plot in a completely randomized block design with three replications. The tested factors included salicylic acid foliar spraying at three levels (0-0.2 M) and two light levels (50-100%). Special nets were used to create 50% shading. The amount of light reduction compared to the control treatment was measured by a lux meter. Foliar spraying was done before flowering. Plant essential oil was obtained from flowering aerial branches by Clevenger apparatus and analyzed by GC-MS. The highest yield of flowering branches and height were from 0.1 M salicylic acid (33.35 kg/ha and 24.33 cm, respectively) and full light treatments (32.50 kg/ha and 22.47cm respectively). The highest amount of chlorophyll a (5.2 mg/g), chlorophyll b (1.98 mg/g) and the number of lateral branches (180/35) due to the application of 0.1 M salicylic acid × 50% light treatment significantly increased compared to the control treatment. The amount of essential oil decreased with increasing light intensity and the use of salicylic acid improved this attribute. So that with full light intensity it reached the lowest level of 1.95% and the application of 0.1 M salicylic acid increased the percentage of essential oil (3.1) compared to the control treatment. The most components of thyme essential oil with the use of 0.1 M salicylic acid× 50% light treatment include carvacrol (4.4%), paracetamol (14.6%), beta-caryophylline (5.95%). Also, the highest amount of thymol was related to 0.1 M salicylic acid treatment (59.66%) and full light (57.8%). In general, the results showed that the use of salicylic acid has a positive effect on improving the qualitative and quantitative characteristics of the essential oil components of thyme. (5.95%). Also, the highest amount of thymol was related to 0.1 M salicylic acid (59.66%) and full light (57.8%). In general, the results showed that the application of salicylic acid had a positive effect on improving the qualitative and quantitative characteristics of the essential oil components of thyme.
References:
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Al-Mariri, A., Swied, G., Oda, A. and Al Hallab, L. 2013. Antibacterial activity of Thymus syriacus essential oil and its components against some Syrian Gram-negative bacteria isolates. Iran J. Med. Sci., 38(2): 180-186.
Arnon, A.N. 1967. Method of extraction of chlorophyll in the plants. Agron. J, 23:112-121.
Bakry, B.A., El Hariri, D.M., Mervat. S.S. and El Bassiouny, H.M.S. 2012. Drought stress mitigation by foliar application of salicylic acid in two linseed varieties grown under newly reclaimed sandy Aoil. Journal of applied sciences research, 7: 3503-3514.
Baj, T., Baryluk, A., and Sieniawska, E. 2018. Application of mixture design for optimum antioxidant activity of mixtures of essential oils from Ocimum basilicum, Origanum majorana L. and Rosmarinus officinalis L. Ind. Crops Prod, 115: 52–61.
Chang, X., Alderson, P.G. and Wright, C.J. 2007. Solar irradiance level alters the growth of Basil (Ocimum basilicum ) and its content of volatile oils. Environ. Exp. Bot, 63: 216–223.
Chen, J.W., Kuang, S.B., Long, G.Q., Yang, S.C., Meng, Z.G., Li, L.G., Chen, Z.J. and Zhang, G.H. 2016. Photosynthesis light energy partitioning and photoprotection in the shade-demanding species Panax notoginseng under high and low level of growth irradiance. Functional Plant Biology, 43: 479-491.
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Ding, Y., Sun, T., Ao, K., Peng, Y., Zhang, Y., Li, X. and Zhang, Y. 2018. Opposite roles of salicylic acid receptors NPR1 and NPR3 / NPR4 in transcriptional regulation of plant immunity. Cell 173, this issue, 1454–1467.
Djennane, S., Oyant, L.H., Kawamura, K., Lalanne, D., Laffaire, M., Thouroude, T., Chalain, S., Sakr, S., Boumaza, R. and Foucher, F. 2014. Impacts of light and temperature on shoot branching gradient and expression of strigolactone synthesis and signalling genes in rose. Plant Cell Environ, 37: 742–757.
Elizabeth Abreu, M. and Munne Bosch, S. 2014. Salicylic acid may be involved in the regulation of drought induced leaf senescence in perennials. A case study in field-grown Salvia officinalis plants. Environmental and Experimental Botany, 64:105–112.
Emarat Pardaz, J., Shakiba, M.R., Toorchi, M. and Mohammadinasab, A.D. 2013. The influence of light intensities and nitrogen on growth of Hypericum perforatum International Journal of Agriculture, 3: 77-781.
Fernandez, M.B., Tossi, V., Lamattina, L. and Cassia, R. 2016. A comprehensive phylogeny reveals functional conservation of the UV-B photoreceptor UVR8 from green algae to higher plants. Front. Plant Sci, 7:1698.
Fiorucci, A.S., and Fankhauser, C. 2017. Plant strategies for enhancing access to sunlight. Cur. Biol, 27: 931–940.
Figurera, P., Marely, G., Rocha, N.E. and Reynosa, R. 2014. Effect of chemical elicitors on peppermint (Mentha piperita) plants and their impact on the metabolite profil and antioxidant capacity of resulting infusion. Food Chemistry, 156: 273-278.
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Ghorbanli, M., Kiapur, A. 2012. Investigation of the effect of different concentrations of copper on pigments and the activity of defense systems
Non-enzymatic and enzymatic in the portulaca oleracea plant (Portulaca oleracea L.). Journal of Medicinal and Aromatic Plants Research, 28(2): 235-247.
Goss, R. and Lepetit, B. 2015. Biodiversity of NPQ. J. Plant Physiol, 172: 13-32.
Heidari, Z., Salehzadeh, A., Sadat Shandiz, S.A. 2018. Anti-cancer and anti-oxidant properties of ethanolic leaf extract of Thymus vulgaris and its bio-functionalized silver nanoparticles, 3 Biotech, 8: 1–14.
Horvath, E., Szalai, G. and Janda, T. 2007. Induction of abiotic stress tolerance by salicylic acid signaling. Journal of plant growth regulation, 26: 290-300.
Idrees, M.M., Khan, A., Aftab, T., Naeem, M. and Hashmi, N. 2010. Salicylic acid induced physiological and biochemical changes in lemongrass varieties under water stress. J Plant Interact, 5: 293–303.
Klessig, D.F., Choi, H.W. and Dempsey, D.M.A. 2018. Systemic acquired resistance and salicylic acid: past, present, and future. Mol. Plant Microbe Interact, 31: 871–888.
Kshavrz, H., ModaresSanavi, S.A.M., ZarinKamr, A., Dolatabadian, F., Panahi. M. and Sadaj Asilan, K. 2012. Evolution effect salicylic acid foliar on same traits biochemical two Brassica napus under cool stress. Iran. J. of Agri. Sci., 42: 723-734.
Kuete, V. 2017. Thymous vulgaris, in: V. Kuete (Ed.), Medicinal spices and vegetables from Africa, first ed. Elsevier Inc, pp. 599–609.
Letchamo, W., Xu, H.L. and Gosselin, A. 2015. Variations in photosynthesis and essential oil in thyme. J of Plant Physiol, 147(1): 29-37.
Li, H., Xu, H., Zhang, P., Gao, M., Wang, D., and Zhao, H. 2017. High temperature effects on D1 protein turnover in three wheat varieties with different heat susceptibility. Plant Growth Regulation, 81: 1-9.
Li, G., Zhang, W., Benoit, F. and Ceustermans, N. Effects of environment factors on the growth and incidence of blossom-end rot in soilless Capsicum fructescens var. grossum. Acta Hort, 633: 382–389.
Li, Y., Yang, Y., Hu, Y., Liu, H., He, M., Yang, Z., Kong, F., Liu, X., and Hou, X. 2019. DELLA and EDS1 form a feedback regulatory module to fine-tune plant growth–defense tradeoff in Arabidopsis. Mol. Plant, 12: 1485-1498.
Li, F.W. and Mathews, S. 2016. Evolutionary aspects of plant photoreceptors. J. Plant Res, 129:115–122.
Li, P., Huang, Z., Li, P., Fang, B., Chu, S. and Guo, H. 2017. A tripartite amplification loop involving the transcription factor WRKY75, salicylic acid and reactive oxygen species accelerates leaf senescence. Plant Cell, 29:2854–2870.
Najafian, S.H, Khushkhui, M., Tavalliali, V. and Saharkhiz. M.J. 2009. Effect of salicylic acid and salinity in Thyme (Thymus vulgaris L.). Journal of Basic and Applied Science, 3(3): 2620-2626.
Nazar, R., Umar, S., Khan, N.A. and Sareer, O. 2015. Salicylic acid supplementation improves photosynthesis and growth in mustard through changes in proline accumulation and ethylene formation under drought stress. South African Journal of Botany, 98:84-94.
Nazar, R.., Umar, S., Khan, N.A., and Sareer, O. 2015. Salicylic acid supplementation improves photosynthesis and growth in mustard through changes in proline accumulation and ethylene formation under drought stress. South Afr. J. Bot, 98: 84–94.
Nazar, R., Iqbal, N., Syeed, S., and Khan, N.A. 2011. Salicylic acid alleviates decreases in photosynthesis under salt stress by enhancing nitrogen and sulfur assimilation and antioxidant metabolism differentially in two mung bean cultivars.Plant Physiol, 168: 807–815.
Nourafcan, H. 2014. Effect of salicylic acid on salinity stress tolerance improvement of peppermint (Mentha piperita L.) in greenhouse conditions. Agroecology Journal (Modern Science of Sustainable Agriculture Journal), 10 (2): 85-95.
Nourruzi Shahri, F., Pouryousef, M., Tavakkoli, A., Saba, J. and Yazdinezhad, A.R. 2015. Evaluation of the function of some native fennel masses (Foeniculum vulgare Mill.) of Iran under drought stress. Journal of Iranian Crop Science, 46(1): 49-56.
Odabas, M.S., Temizel, K.E., Caliskan, O., Senyer, N., Kayhan, G. and Ergun, E. 2014. Determination of reflectance values of Hypericum's leaves under stress conditions using adaptive network based fuzzy inference system. Neural Network World, 24: 79-87.
Parvaiz, A. and Satyawati, S. 2008. Salt stress and phyto-biochemical responses of plants. Plant Soil Environment, 54: 89-99.
Pessarkkli, M. 2019. Hand book of Plant hand Crop Stress. Maecel Dekker Inc. New York.
Ram, m., Singh, R., Naqvi, A.A., Lohia, R.S., Bansal, R.P., and Kumar, S. 1997. Effect of salisylic acid on the yield and quality of essential oil in aromatic crops. J. Med. Aromatic Pl. Sci, 19: 24-27.
Rahim, M., and Yadegari, M. 2012. Effects of jasmonic and salicylic acid on the phytochemical properties of sage leaves, herbal medicines, 3 (2): 89- 94.
Rezai, S., Etemadi, N., Nikbakht, A., Yousefi, M., and Majidi, M.M. 2018. Effect of light intensity on leaf morphology, photosynthetic capacity and chlorophyll content in Sage (Salvia officinalis ). Hort. Sci. Technol, 36 (1): 46–57.
Ruban, A.V. 2016. Nonphotochemical chlorophyll fluorescence quenching: mechanism and effectiveness in protecting plants from photodamage. Plant Physiol, 170: 1903-1916.
Setiawati, T., Ayalla, A., Nurzaman, M.A., and Mutaqin, Z. 2018. Influence of light intensity on leaf photosynthetic traits and alkaloid content of Kiasahan (Tetracera scandens L.). IOP Conf. Ser. Earth Environ. Sci, 166: 12–25.
Schottler, M.A. and Toth, S.Z. 2014. Photosynthetic complex stoichiometry dynamics in higher plants: environmental acclimation and photosynthetic flux control. Front. Plant Sci, 5: 188.
Setiawati, T., Ayalla, A., Nurzaman, M.A., and Mutaqin, Z. 2018. Influence of light intensity on leaf photosynthetic traits and alkaloid content of Kiasahan (Tetracera scandens L.). IOP Conf. Ser. Earth Environ. Sci. 166: 12–25.
Shilpa, K., Varun, K., and Lakshmi, B. 2010. An alternate method of natural drug production: eliciting secondary metabolite production using plant cell culture. J. Plant Sci., 5: 222-247.
Silvestrini, M., Valio, I.F.M. and Mattos, E.A.D. 2007. Photosynthesis and carbon gain under contrasting light levels in seedlings of a pioneer and a climax tree from a Brazilian Semideciduous Tropical Forest. Brazilian Journal of Botany. 30: 463-474.
Temizel, K.E. 2015. Estimation of the phenolics content of St. John’s wort (Hyperıcum perforatum ) grown under different water and salt levels based on reflectance spectroscopy. Kuwait Journal of Science, 42: 104-114.
Tolit Abolhassani, M., Tavakol Afshari, R., Jahansuz, M.R., Najafi, F., and Naqdi Badi, H.A. 2015. The effect of different temperatures on the germination behavior of thyme seeds of Elam Thy ecotype. Iranian Journal of Seed Science and Technology,1:97107
Uzunova, A.N., Popova, L.P. 2000. Effect of salicylic acid on leaf anatomy and chloroplast ultrastructure of barley plants Photosynthetica, 38: 243-250.
Victor Perez, T., Antonio Lopez, O., Asception Martinez, P. and Antonio, A.C. 2012. Antioxidant activity and rosmarinic acid changes in salicylic acid tread Thymus membranceus L. shoots. Food Chemistry, 130: 362- 369.
Wang, H. and Wang, H. 2015. Phytochrome signaling: time to tighten up the loose ends. Mol. Plant, 8:540–551.
Wang, Y.Y., Wang, Y., Li, G.Z., and Hao, L. 2019. Salicylic acid-altering Arabidopsis plant response to cadmium exposure: underlying mechanisms affecting antioxidation and photosynthesis-related processes. Ecotoxicol. Environ. Saf, 169: 645–653.
Yamori, W. and Shikanai, T. 2016. Physiological functions of cyclic electron transport around photosystem I in sustaining photosynthesis and plant growth. Annual Review of Plant Biology, 67: 81-106.
Yuan, M., Zhao, Y.Q., Zhang, Z.W., Chen, Y.E., Ding, C.B., and Yuan, S. 2017. Light regulates transcription of chlorophyll biosynthetic genes during chloroplast biogenesis. Crit. Rev. Plant Sci, 36: 35–54.
Zhu, H., Li, X., Zhai, W., Liu, Y., Gao, Q., Liu, J., and Zhu, Y. 2017. Effects of low light on photosynthetic properties, antioxidant enzyme activity, and anthocyanin accumulation in purple pak-choi (Brassica campestris ssp. Chinensis Makino). PloS One, 12 (6).
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Adams, R.P. 2001. Identification of essential oil components by gas chromatograph mass spectrometry. Allured Publishing Corporation Carol Stream. IL.
Ade–Ademilua, E.O., Obi, H.O. and Craker, L.E. 2013. Growth and essential oil yield of African Basil. Ocimum gratissimum. under light and water stress. Journl of Medicinally active plants, 1 (4): 142-149.
Al-Mariri, A., Swied, G., Oda, A. and Al Hallab, L. 2013. Antibacterial activity of Thymus syriacus essential oil and its components against some Syrian Gram-negative bacteria isolates. Iran J. Med. Sci., 38(2): 180-186.
Arnon, A.N. 1967. Method of extraction of chlorophyll in the plants. Agron. J, 23:112-121.
Bakry, B.A., El Hariri, D.M., Mervat. S.S. and El Bassiouny, H.M.S. 2012. Drought stress mitigation by foliar application of salicylic acid in two linseed varieties grown under newly reclaimed sandy Aoil. Journal of applied sciences research, 7: 3503-3514.
Baj, T., Baryluk, A., and Sieniawska, E. 2018. Application of mixture design for optimum antioxidant activity of mixtures of essential oils from Ocimum basilicum, Origanum majorana L. and Rosmarinus officinalis L. Ind. Crops Prod, 115: 52–61.
Chang, X., Alderson, P.G. and Wright, C.J. 2007. Solar irradiance level alters the growth of Basil (Ocimum basilicum ) and its content of volatile oils. Environ. Exp. Bot, 63: 216–223.
Chen, J.W., Kuang, S.B., Long, G.Q., Yang, S.C., Meng, Z.G., Li, L.G., Chen, Z.J. and Zhang, G.H. 2016. Photosynthesis light energy partitioning and photoprotection in the shade-demanding species Panax notoginseng under high and low level of growth irradiance. Functional Plant Biology, 43: 479-491.
DebMandal, S.M. 2016. Thyme (Thymus vulgaris L.) oils, in: V. Preedy (Ed.), Essential Oils in Food Preservation, Flavor and Safety, Academic Press, London, UK, pp, 825–834.
Ding, Y., Sun, T., Ao, K., Peng, Y., Zhang, Y., Li, X. and Zhang, Y. 2018. Opposite roles of salicylic acid receptors NPR1 and NPR3 / NPR4 in transcriptional regulation of plant immunity. Cell 173, this issue, 1454–1467.
Djennane, S., Oyant, L.H., Kawamura, K., Lalanne, D., Laffaire, M., Thouroude, T., Chalain, S., Sakr, S., Boumaza, R. and Foucher, F. 2014. Impacts of light and temperature on shoot branching gradient and expression of strigolactone synthesis and signalling genes in rose. Plant Cell Environ, 37: 742–757.
Elizabeth Abreu, M. and Munne Bosch, S. 2014. Salicylic acid may be involved in the regulation of drought induced leaf senescence in perennials. A case study in field-grown Salvia officinalis plants. Environmental and Experimental Botany, 64:105–112.
Emarat Pardaz, J., Shakiba, M.R., Toorchi, M. and Mohammadinasab, A.D. 2013. The influence of light intensities and nitrogen on growth of Hypericum perforatum International Journal of Agriculture, 3: 77-781.
Fernandez, M.B., Tossi, V., Lamattina, L. and Cassia, R. 2016. A comprehensive phylogeny reveals functional conservation of the UV-B photoreceptor UVR8 from green algae to higher plants. Front. Plant Sci, 7:1698.
Fiorucci, A.S., and Fankhauser, C. 2017. Plant strategies for enhancing access to sunlight. Cur. Biol, 27: 931–940.
Figurera, P., Marely, G., Rocha, N.E. and Reynosa, R. 2014. Effect of chemical elicitors on peppermint (Mentha piperita) plants and their impact on the metabolite profil and antioxidant capacity of resulting infusion. Food Chemistry, 156: 273-278.
Golparvar, A.R. 2011. Effects of phenological stages on quality and quantity of essential oil in Kermanian Thyme (Thymus caramanicus ). Electronic J of Bio, 7(4): 70-73.
Ghorbanli, M., Kiapur, A. 2012. Investigation of the effect of different concentrations of copper on pigments and the activity of defense systems
Non-enzymatic and enzymatic in the portulaca oleracea plant (Portulaca oleracea L.). Journal of Medicinal and Aromatic Plants Research, 28(2): 235-247.
Goss, R. and Lepetit, B. 2015. Biodiversity of NPQ. J. Plant Physiol, 172: 13-32.
Heidari, Z., Salehzadeh, A., Sadat Shandiz, S.A. 2018. Anti-cancer and anti-oxidant properties of ethanolic leaf extract of Thymus vulgaris and its bio-functionalized silver nanoparticles, 3 Biotech, 8: 1–14.
Horvath, E., Szalai, G. and Janda, T. 2007. Induction of abiotic stress tolerance by salicylic acid signaling. Journal of plant growth regulation, 26: 290-300.
Idrees, M.M., Khan, A., Aftab, T., Naeem, M. and Hashmi, N. 2010. Salicylic acid induced physiological and biochemical changes in lemongrass varieties under water stress. J Plant Interact, 5: 293–303.
Klessig, D.F., Choi, H.W. and Dempsey, D.M.A. 2018. Systemic acquired resistance and salicylic acid: past, present, and future. Mol. Plant Microbe Interact, 31: 871–888.
Kshavrz, H., ModaresSanavi, S.A.M., ZarinKamr, A., Dolatabadian, F., Panahi. M. and Sadaj Asilan, K. 2012. Evolution effect salicylic acid foliar on same traits biochemical two Brassica napus under cool stress. Iran. J. of Agri. Sci., 42: 723-734.
Kuete, V. 2017. Thymous vulgaris, in: V. Kuete (Ed.), Medicinal spices and vegetables from Africa, first ed. Elsevier Inc, pp. 599–609.
Letchamo, W., Xu, H.L. and Gosselin, A. 2015. Variations in photosynthesis and essential oil in thyme. J of Plant Physiol, 147(1): 29-37.
Li, H., Xu, H., Zhang, P., Gao, M., Wang, D., and Zhao, H. 2017. High temperature effects on D1 protein turnover in three wheat varieties with different heat susceptibility. Plant Growth Regulation, 81: 1-9.
Li, G., Zhang, W., Benoit, F. and Ceustermans, N. Effects of environment factors on the growth and incidence of blossom-end rot in soilless Capsicum fructescens var. grossum. Acta Hort, 633: 382–389.
Li, Y., Yang, Y., Hu, Y., Liu, H., He, M., Yang, Z., Kong, F., Liu, X., and Hou, X. 2019. DELLA and EDS1 form a feedback regulatory module to fine-tune plant growth–defense tradeoff in Arabidopsis. Mol. Plant, 12: 1485-1498.
Li, F.W. and Mathews, S. 2016. Evolutionary aspects of plant photoreceptors. J. Plant Res, 129:115–122.
Li, P., Huang, Z., Li, P., Fang, B., Chu, S. and Guo, H. 2017. A tripartite amplification loop involving the transcription factor WRKY75, salicylic acid and reactive oxygen species accelerates leaf senescence. Plant Cell, 29:2854–2870.
Najafian, S.H, Khushkhui, M., Tavalliali, V. and Saharkhiz. M.J. 2009. Effect of salicylic acid and salinity in Thyme (Thymus vulgaris L.). Journal of Basic and Applied Science, 3(3): 2620-2626.
Nazar, R., Umar, S., Khan, N.A. and Sareer, O. 2015. Salicylic acid supplementation improves photosynthesis and growth in mustard through changes in proline accumulation and ethylene formation under drought stress. South African Journal of Botany, 98:84-94.
Nazar, R.., Umar, S., Khan, N.A., and Sareer, O. 2015. Salicylic acid supplementation improves photosynthesis and growth in mustard through changes in proline accumulation and ethylene formation under drought stress. South Afr. J. Bot, 98: 84–94.
Nazar, R., Iqbal, N., Syeed, S., and Khan, N.A. 2011. Salicylic acid alleviates decreases in photosynthesis under salt stress by enhancing nitrogen and sulfur assimilation and antioxidant metabolism differentially in two mung bean cultivars.Plant Physiol, 168: 807–815.
Nourafcan, H. 2014. Effect of salicylic acid on salinity stress tolerance improvement of peppermint (Mentha piperita L.) in greenhouse conditions. Agroecology Journal (Modern Science of Sustainable Agriculture Journal), 10 (2): 85-95.
Nourruzi Shahri, F., Pouryousef, M., Tavakkoli, A., Saba, J. and Yazdinezhad, A.R. 2015. Evaluation of the function of some native fennel masses (Foeniculum vulgare Mill.) of Iran under drought stress. Journal of Iranian Crop Science, 46(1): 49-56.
Odabas, M.S., Temizel, K.E., Caliskan, O., Senyer, N., Kayhan, G. and Ergun, E. 2014. Determination of reflectance values of Hypericum's leaves under stress conditions using adaptive network based fuzzy inference system. Neural Network World, 24: 79-87.
Parvaiz, A. and Satyawati, S. 2008. Salt stress and phyto-biochemical responses of plants. Plant Soil Environment, 54: 89-99.
Pessarkkli, M. 2019. Hand book of Plant hand Crop Stress. Maecel Dekker Inc. New York.
Ram, m., Singh, R., Naqvi, A.A., Lohia, R.S., Bansal, R.P., and Kumar, S. 1997. Effect of salisylic acid on the yield and quality of essential oil in aromatic crops. J. Med. Aromatic Pl. Sci, 19: 24-27.
Rahim, M., and Yadegari, M. 2012. Effects of jasmonic and salicylic acid on the phytochemical properties of sage leaves, herbal medicines, 3 (2): 89- 94.
Rezai, S., Etemadi, N., Nikbakht, A., Yousefi, M., and Majidi, M.M. 2018. Effect of light intensity on leaf morphology, photosynthetic capacity and chlorophyll content in Sage (Salvia officinalis ). Hort. Sci. Technol, 36 (1): 46–57.
Ruban, A.V. 2016. Nonphotochemical chlorophyll fluorescence quenching: mechanism and effectiveness in protecting plants from photodamage. Plant Physiol, 170: 1903-1916.
Setiawati, T., Ayalla, A., Nurzaman, M.A., and Mutaqin, Z. 2018. Influence of light intensity on leaf photosynthetic traits and alkaloid content of Kiasahan (Tetracera scandens L.). IOP Conf. Ser. Earth Environ. Sci, 166: 12–25.
Schottler, M.A. and Toth, S.Z. 2014. Photosynthetic complex stoichiometry dynamics in higher plants: environmental acclimation and photosynthetic flux control. Front. Plant Sci, 5: 188.
Setiawati, T., Ayalla, A., Nurzaman, M.A., and Mutaqin, Z. 2018. Influence of light intensity on leaf photosynthetic traits and alkaloid content of Kiasahan (Tetracera scandens L.). IOP Conf. Ser. Earth Environ. Sci. 166: 12–25.
Shilpa, K., Varun, K., and Lakshmi, B. 2010. An alternate method of natural drug production: eliciting secondary metabolite production using plant cell culture. J. Plant Sci., 5: 222-247.
Silvestrini, M., Valio, I.F.M. and Mattos, E.A.D. 2007. Photosynthesis and carbon gain under contrasting light levels in seedlings of a pioneer and a climax tree from a Brazilian Semideciduous Tropical Forest. Brazilian Journal of Botany. 30: 463-474.
Temizel, K.E. 2015. Estimation of the phenolics content of St. John’s wort (Hyperıcum perforatum ) grown under different water and salt levels based on reflectance spectroscopy. Kuwait Journal of Science, 42: 104-114.
Tolit Abolhassani, M., Tavakol Afshari, R., Jahansuz, M.R., Najafi, F., and Naqdi Badi, H.A. 2015. The effect of different temperatures on the germination behavior of thyme seeds of Elam Thy ecotype. Iranian Journal of Seed Science and Technology,1:97107
Uzunova, A.N., Popova, L.P. 2000. Effect of salicylic acid on leaf anatomy and chloroplast ultrastructure of barley plants Photosynthetica, 38: 243-250.
Victor Perez, T., Antonio Lopez, O., Asception Martinez, P. and Antonio, A.C. 2012. Antioxidant activity and rosmarinic acid changes in salicylic acid tread Thymus membranceus L. shoots. Food Chemistry, 130: 362- 369.
Wang, H. and Wang, H. 2015. Phytochrome signaling: time to tighten up the loose ends. Mol. Plant, 8:540–551.
Wang, Y.Y., Wang, Y., Li, G.Z., and Hao, L. 2019. Salicylic acid-altering Arabidopsis plant response to cadmium exposure: underlying mechanisms affecting antioxidation and photosynthesis-related processes. Ecotoxicol. Environ. Saf, 169: 645–653.
Yamori, W. and Shikanai, T. 2016. Physiological functions of cyclic electron transport around photosystem I in sustaining photosynthesis and plant growth. Annual Review of Plant Biology, 67: 81-106.
Yuan, M., Zhao, Y.Q., Zhang, Z.W., Chen, Y.E., Ding, C.B., and Yuan, S. 2017. Light regulates transcription of chlorophyll biosynthetic genes during chloroplast biogenesis. Crit. Rev. Plant Sci, 36: 35–54.
Zhu, H., Li, X., Zhai, W., Liu, Y., Gao, Q., Liu, J., and Zhu, Y. 2017. Effects of low light on photosynthetic properties, antioxidant enzyme activity, and anthocyanin accumulation in purple pak-choi (Brassica campestris ssp. Chinensis Makino). PloS One, 12 (6).
