The Effect of Different Qualities of LED Light on the Morphophysiological Indicators of Cucumis sativus L. var. Officer

علوم باغبانی

Article 10, Volume 37, Issue 4 - Serial Number 60, January 2024, Pages 1029-1041   

Document Type: Research Article

DOI: 10.22067/jhs.2023.80257.1222

Authors

M. Zare Mehrjerdi¹; N. Safari²; S. M. Kharrazi³; A. Khadem³; A. Sharifi^{* 3}

¹Agriculture Faculty of Shirvan, University of Bojnord, Bojnord, Iran

²Department of Horticultural Science and Landscape, Faculty of Agriculture, Ferdowsi University of Mashhad, Mashhad, Iran

³Horticultural Plants Biotechnology Department, Research Institute for Industrial Biotechnology, Iranian Academic Centre for Education, Culture and Research (ACECR), Khorasan Razavi Branch, Mashhad, Iran

Abstract

Introduction
 In addition to providing the necessary energy for photosynthesis, light controls many plant metabolic processes. Nowadays, the use of supplemental light significantly improves the quality of the food product in the conditions of lack of natural light in the autumn and winter seasons. Light-emitting diodes have been proposed as alternative light sources in controlled agricultural environments. These lamps are the first light sources with the ability to control the light spectrum. Therefore, by controlling the light spectrum and matching the wavelength of LED lamps with the photoreceptors of the plant, the performance and quality of the plant is improved. Cucumber is one of the most important greenhouse plants and its seedlings are generally produced during the autumn and winter seasons, when natural light is relatively low; therefore, the use of supplementary light is required. The use of artificial light sources in fully controlled conditions can change and improve the growth conditions of the plant and also improve its quantitative and qualitative traits. In this research, the aim was to investigate the effect of different qualities of light on the morphophysiological indicators of cucumber seedlings.
 
Materials and Methods
 This research was performed at the Horticultural Plants Biotechnology Department, Industrial Biotechnology Research Institute of Khorasan Razavi. First, the seeds of the cucumber plant (Cucumis sativus L. var. Officer) were planted in a planting tray, and placed under the light panel with different light treatments. The experimental treatments included four light qualities including white light (6000-6500K) blue light (460-470 nm), red light (625 nm) and combined light (blue + red + white). The amount of photosynthetic photon flux was considered the same in all light treatments, which was equal to 2.75 µol m-2 s-1 CO2 assimilation. Thirty days after planting of seeds, the growth of plants were measured in three stages every 10 days. In each stage, 4 plant samples were selected and then plant height, leaf area, fresh and dry weight of shoots and roots, plant dry matter index and leaf chlorophyll content were measured. Data preparation was done in Excel software, data analysis was done using JMP-8 software and treatment averages were compared using LSD test at 5% probability level.
 
Results and Discussion
The results of the experiments showed that the quality of light was significantly effective on the growth indicators of Cucumber plants. Plant height in red light and blue light treatments compared to the white light and combined light (red+blue+white with the same intensity) treatments, increased by more than 130% and 60%, respectively. The lowest height was observed in plants grown under combined light, 9 and 14.25 cm after 40 and 50 days of cultivation respectively. The exposure of cucumber seedlings to the red and combined lights recorded the highest and the lowest leaf area in the plants, 618.65 and 377.26 cm² respectively. Also white light significantly improved the dry weight of the roots. It is worth to mention that the highest plant fresh weight was observed in blue and white lights, 20.9 and 19.5 g respectively. For parameters such as dry weight, dry matter index, and pigment content, the light treatments did not exert a significant effect. However, the utilization of red and blue light, both individually and in combination, positively impacted plant growth. Notably, exposure to red light alone led to a significant increase in leaf surface area, root dry weight, and plant height compared to other light conditions. Previous studies have indicated that red light enhances leaf area, stem length, and fresh weight of plants. Leaves play a critical role in plant photosynthesis and overall growth. Therefore, increasing the leaf area in the plant increases the amount of photosynthesis, growth and development of the plant. In this study, with the increases of the leaf area in the plant and the subsequent increases in the amount of photosynthesis, allocation of dry mater to root increased. The results of the research has been shown that the combination of blue and red wavelengths in pepper, tomato and cucumber seedlings is effective in stimulating plant growth and improving the morphological characteristics under controlled conditions. Blue and red lights can increase the proton flow rate of epidermal cells through the separation mechanism and thus affect leaf development. Blue light directly through the interaction with proton pumps and indirectly through receptors, affects proton pumps by modulating passive ion conduction of potassium and calcium channels.
 
Conclusion
 According to the results of this research, it was found that exposing the plant to different light quality had different responses in the cucumber plants. Although variables such as dry weight and the amount of photosynthetic pigments were not significantly affected by light quality, however, traits such as plant height, leaf area, root dry weight, and plant fresh weight were affected by light quality. The affected parameters are among the traits that are influenced by the gibberellin hormone and according to the reports related to the effect of light quality on the gibberellin biosynthesis and response to this hormone. It seems that plant action to the quality of light can be attributed to the regulation of this hormone. So it is possible to choose the appropriate light quality in fully controlled conditions according to the production goal and results. In this research, according to the plant leaf rea, root dry weight and plant height, it was determined that white light can be used in the seedling production stage.

Keywords

Blue light; Growth characteristics; Light emitting diodes; Red light; White light

References

1. Adams, S.R., Valdes, V.M., & Langton, F.A. (2008). Why does low intensity, long-day lighting promote growth in petunia, impatiens, and tomato? The Journal of Horticultural Science and Biotechnology, 83(5), 609-615. https://doi.org/10.1080/14620316.2008.11512431.
2. Astolfi, S., de Biasi, M.G., & Passera, C. (2001). Effect of irradiance-sulphur interaction on enzyme of carbon, nitrogen, and Sulphur metabolism in maize plant. Photosynthetica, 39, 177-181. https://doi.org/10.1023/A:1013762605766.
3. Bernie, G., & Perilleux, C. (2005). A physiological overview of the genetics of flowering time control. Plant Biotechnology Journal, 3, 3-16. https://doi.org/1111/j.1467-7652.2004.00114.x.
4. Bian, Z.H., Cheng, R.F., Yang, Q.C., Wang, J., & Lu, C. (2016). Continuous light from red, blue, and green light-emitting diodes reduces nitrate content and enhances phytochemical concentrations and antioxidant capacity in lettuce. Journal of the American Society for Horticultural Science, 141, 186–195. https://doi.org/10.21273/JASHS.141.2.186.
5. Bian, Z.H., Yang, Q.C., & Liu, W.K. (2015). Effects of light quality on the accumulation of phytochemicals in vegetables produced in controlled environments: a review. Journal of the Science of Food and Agriculture, 95(5), 869–877. https://doi.org/1002/jsfa.6789
6. Biswal, A.K., Pattanayak, G.K., Pandey, S.S., Leelavathi, S., Reddy, V.S., & Tripathy, B.C. (2012). Light intensitydependent modulation of chlorophyll b biosynthesis and photosynthesis by overexpression of chlorophyllide a oxygenase in tobacco. Plant Physiology, 159, 433-449. https://doi.org/10.1104/pp.112.195859.
7. Bourget, C.M. (2008) An introduction to light-emitting diodes. HortScience, 43, 1944-1946. https://doi.org/10.21273/HORTSCI.43.7.1944.
8. Cao, G., Zhang, G., Yu, J., & Ma, Y. (2013). Effects of different led light qualities on cucumber seedling growth and chlorophyll fluorescence parameters. Scientia Agricultura Sinica, 46(6), 1297–1304.
9. Cerdan, P.D., & Chory, J. (2003). Regulation of flowering time by light quality. Nat, 423, 881-885. https://doi.org/10.1038/nature01636
10. Chen, M., Chory, J., & Fankhauser, C. (2004). Light signal transduction in higher plants. Annual Review of Genetics, 38(1), 87–117. https://doi.org/10.1146/annurev.genet.38.072902.092259
11. Cosgrove, D.J. (1981). Rapid suppression of growth by blue light. Plant Physiology, 67, 584-590. https://doi.org/10.1104/pp.67.3.584
12. Fan, X., Zang, J., Xu, Z., Guo, S., Jiao, X., Liu, X., & Gao, Y. (2013). Effects of different light quality on growth, chlorophyll concentration and chlorophyll biosynthesis precursors of non-heading Chinese cabbage (Brassica campestris). Acta Physiologiae Plantarum, 35, 2721-2726. https://doi.org/10.1007/s11738-013-1304-z
13. Fukuda, N., Ajima, C., Yukawa, T., & Olsen, J. (2016). Antagonistic action of blue and red light on shoot elongation in petunia depends on gibberellin, but the effects on flowering are not generally linked to gibberellin. Environmental and Experimental Botany, 121, 102-111. https://doi.org/10.1016/j.envexpbot.2015.06.014
14. Goto, E. (2003). Effect of light quality on growth of crop plants under artificial lighting. Environmental Control in Biology, 41, 121-132. https://doi.org/10.2525/ecb1963.41.12
15. Hanyu, H., & Shoji, K. (2000) Effect of blue light and red light on kidney bean plants grown under combined radiation from narrow-band light source. Environmental Control in Biology, 38, 13-24. http://doi.org/10.2525/ecb1963.38.13
16. Hemming, S., Mohammadkhani, V., & Dueck, T. (2008). Diffuse greenhouse coveringmaterials–material technology, measurements and evaluation of optical properties. Acta Horticulturae, 469–475. http://doi.org/17660/ActaHortic.2008.797.68
17. Heo, J.W., Kang, D.H., Bang, H.S., Hong, S.G., Chun, C.H., & Kang, K.K. (2012). Early growth, pigmentation, protein content, and phenylalanine ammonia-lyase activity of red curled lettuces grown under different lighting conditions. Korean Journal of Horticultural Science and Technology, 30(1), 6–12. http://doi.org/10.7235/hort.2012.11118
18. Hernández, R., & Kubota, C. (2016). Physiological responses of cucumber seedlings under different blue and red photon flux ratios using LEDs. Environmental and Experimental Botany, 121, 66-74.
19. Hernández, R., & Kubota, C. (2015). Physiological, morphological, and energy-use efficiency comparisons of LED and HPS supplemental lighting for cucumber transplant production. HortScience, 50, 351-357.
20. Hernández, R., & Kubota, C. (2012). Tomato seedling growth and morphologicalresponses to supplemental LED lighting Red:Blue rations under varied daily solar light integrals. Acta Horticulturae, 956, 187–194. http://doi.org/10.17660/ActaHortic.2012.956.19
21. Hernandez, R., Eguchi, T., & Kubota, C. (2016). Growth and morphology of vegetable seedlings under different blue and red photon flux ratios using light-emitting diodes as solesource lighting. Acta Horticulturae (ISHS), 1134, 195–200. http://doi.org/10.17660/ActaHortic.2016.1134.26
22. Heuvelink, E., Bakker, M.J., Hogendonk, L., Janse, J., Kaarsemaker, R.C., & Maaswinkel, R.H.M. (2006). Horticultural lightening in the Neetherlands: new developments, Acta Horticulturae, 711, 25-33. https://doi.org/10.17660/ActaHortic.2006.711.1
23. Hoagland, D.R., & Arnon, D.I. (1938). The water culture method for growing plants without soil. California Agricultural Experiment Station Circulation, 347, 32.
24. Hoffmann, A.M., Noga, G., & Hunsche, M. (2015). High blue light improves acclimation and photosynthetic recovery of pepper plants exposed to UV stress. Environmental and Experimental Botany, 109(2), 254–63. https://doi.org/10.1016/j.envexpbot.2014.06.017
25. Hogewoning, S.W., Trouwborst, G., Maljaars, H., Poorter, H., van Ieperen, W., & Harbinson, J. (2010). Blue light dose–responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light. Journal of Experimental Botany, 61, 3107-3117. https://doi.org/10.1093/jxb/erq132
26. Hogewoning, S.W., Wienties, E., Douwatra, P., Trouwborst, G., van Irperen, W., Croce, R., & Harbinson, J. (2012). Photosynthetic quantum yeild dynamic: From photosysytems to leaves. The Plant Cell, 24, 1921-1935. https://doi.org/1105/tpc.112.097972
27. Jeong, H.W., Lee, H.R., Kim, H.M., Kim, H.M., Hwang H.S., & Hwang, S.J. (2020). Using light quality for growth control of cucumber seedlings in closed-type plant production system. Plants, 639, 1-12. https://doi.org/10.3390/plants9050639
28. Johkan, M., Shoji, K., Goto, F., Hashida, S., & Yoshihara, T. (2010). Blue light-emitting diode light irradiation of seedlings improves seedling quality and growth after transplanting in red leaf lettuce. HortScience, 45(12), 1809–1814. https://doi.org/10.21273/HORTSCI.45.12.1809
29. Kaiser, E., Ouzounis, T., Giday, H., Schipper, R., Heuvelink, E., & Marcelis, L.F.M. (2019). Adding blue to red supplemental light increases biomass and yield of greenhouse-grown tomatoes, but only to an optimum. Frontiers in Plant Science, 9, 1–11. https://doi.org/10.3389/fpls.2018.02002
30. Kim, H.H., Goins, G.D., Wheeler, R.M., & Sager, J.C. (2004). Green-light supplementation for enhanced lettuce growth under red- and bluelight- emitting diodes. HortScience, 39, 1617–1622.
31. Kim, H.-J., Lin, M.-Y., & Mitchell, C.A. (2019). Light spectral and thermal properties govern biomass allocation in tomato through morphological and physiological changes, Environmental and Experimental Botany, 157, 228–240. https://doi.org/10.1016/j.envexpbot.2018.10.019
32. Klein, R.M. (1992). Effect of green light on biological systems. Biological Reviews, 67, 199-284. https://doi.org/10.1111/j.1469-185x.1992.tb01019.x
33. Kong, Y., Kamath, D., & Zheng, Y. (2019). Blue versus red light can promote elongation growth independent of photoperiod: a study in four Brassica microgreens species. Hortscience, 54(11), 1955–1961. https://doi.org/10.21273/HORTSCI14286-19
34. Kopsell, D.A., Sams, C.E., & Barickman, T.C. (2014). Sprouting broccoli accumulate higher concentrations of nutritionally important metabolites under narrow-band light-emitting diode lighting. Horticultural Science, 139, 469–477. https://doi.org/10.21273/JASHS.139.4.469
35. Kreslavski, V.D., Lyubimov, V.Y., Shirshikova, G.N., Shmarev, A.N., Kosobryukhov, A.A., Schmitt, F.J., Friedrich, T., & Allakhverdiev, S.I. (2013). Preillumination of lettuce seedlings with red light enhances the resistance of photosynthetic apparatus to UV-A. Journal of Photochemistry Photobiology B: Biology, 122(1), 1–6. https://doi.org/10.1016/j.jphotobiol.2013.02.016
36. Lanoue, J., Leonardos, E.D., & Grodzinski, B. (2018). Effects of light quality and intensity on diurnal patterns and rates of photo-assimilate translocation and transpiration in tomato leaves. Frontiers in Plant Science, 9, 756. https://doi.org/10.3389/fpls.2018.00756
37. Li, H., Tang, C., & Xu, Z. (2013). The effects of different light qualities on rapeseed (Brassica napus ) plantlet growth and morphogenesis in vitro. Scientia Horticulturae, 150, 117–124. https://doi.org/10.1016/j.scienta.2012.10.009.
38. Lichtenthaler, H.K., & Buschmann, C. (2001). Chlorophylls and carotenoids: measurement and characterization by UV-vis spectroscopy. Current Protocols in Food Analytical Chemistry F4.3.1-F4.3.8. https://doi.org/10.1002/0471142913.faf0403s01
39. Liu, N., Ji, F., Xu, L.J., & He, D.X. (2019). Effects of LED light quality on the growth of pepper seedling in plant factory. International Journal of Agricultural and Biological Engineering, 12(5), 44–50. https://doi.org/25165/j.ijabe.20191205.4847
40. Liu, X.Y., Chang, T.T., Guo, S.R., Xu, Z.G., & Li, J., (2011). Effects of different light qualityof LED on growth of LED on growth and photosynthetic characters in cherry tomato seedlings. Acta Horticulturae, 907, 325–330. https://doi.org/17660/ActaHortic.2011.907.53
41. Lu, N., Maruo, T., Johkan, M., Hohjo, M., Tsukagoshi, S., Ito, Y., Ichimura, T., & Shinohara, Y. (2012). Effects of supplemental lighting with light-emitting diodes (LEDs) on tomato yield and quality of single-truss tomato plants grown at high planting density. Environmental Control in Biology, 50(1), 63–74. https://doi.org/10.2525/ecb.50.63
42. Massa, G.D., Kim H.H., Wheeler, R.M., & Mitchell, C.A. (2008). Plant productivity in response to LED lighting. Hortscience, 43, 1951- 1956. https://doi.org/10.21273/HORTSCI.43.7.1951
43. McCree, K.J. (1971). The action spectrum, absorptance and quantum yield of photosynthesis in crop plants. Agricultural Meteorology, 9, 191-216. https://doi.org/10.1016/0002-1571(71)90022-7
44. Miao, Y.X., Wang, X.Z., Gao, L.H., Chen, Q.Y., & Qu, M. (2016). Blue light is more essential than red light for maintaining the activities of photosystem II and I and photosynthetic electron transport capacity in cucumber leaves. Journal of Integrative Agriculture, 15(1), 87-100. https://doi.org/10.1016/S2095-3119(15)61202-3
45. Moradi, M., Abedi, B., Arouiee, H., Aliniaeifard, S., & Ghasemi Bezdi, K. (2023). Effect of different light spectral on photosynthetic performance, growth indicators and essential oil content of Salvia officinalis. Journal of Horticultural Science, 37(3), 821-841. (In Persian with English abstract). https://doi.org/10.22067/jhs.2023.78806.1211
46. Morrow, R.C. (2008). LED lighting in horticulture. HortScience, 43(7), 1947-1950. https://doi.org/10.21273/HORTSCI.43.7.1947
47. Naznin, M.T., Lefsrud, M., Gravel, V., & Azad, M.O.K. (2019). Blue light added with red LEDs enhance growth characteristics, pigments content, and antioxidant capacity in lettuce, spinach, kale, basil, and sweet pepper in a controlled environment. Plants, 8(4). https://doi.org/10.3390/plants8040093
48. Netto, A.T., Campostrini, E., & de Oliveira J.G. (2005). Photosynthetic pigments, nitrogen, chlorophyll a fluorescence and SPAD-502 readings in coffee leaves. Scientia Horticulturae, 104, 199–209. https://doi.org/10.1016/j.scienta.2004.08.013
49. Olle, , & Viršile, A. (2013). The effects of light-emitting diodes on greenhouse plant growth and quality. Agricultural and Food Science, 22, 223–234. https://doi.org/10.23986/afsci.7897
50. Randall, W.C., & Lopez, R.G. (2014). Comparison of supplemental lighting from high-pressure sodium lamps and light-emitting diodes during bedding plant seedling production. HortScience, 49, 589–595. https://doi.org/10.21273/HORTSCI.49.5.589
51. Savvides, A., Fanourakis, D., & van Ieperen, W. (2012). Co-ordination of hydraulic andstomatal conductances across light qualities in cucumber leaves. Journal of Experimental Botany, 63, 1135–1143. https://doi.org/10.1093/jxb/err348
52. Singh, D., Basu, C., Meinhardt-Wollweber, M., & Roth, B. (2015). LEDs for energy efficient greenhouse lighting. Renew. Renewable and Sustainable Energy Reviews, 49, 139–147. https://doi.org/10.1016/j.rser.2015.04.117
53. Son, K.H., & Oh, M.M. (2013). Leaf shape, growth, and antioxidant phenolic compounds of two lettuce cultivars grown under various combinations of blue and red light-emitting diodes. Horticulture Science, 48(8), 988–995. https://doi.org/10.21273/HORTSCI.48.8.988
54. Song, J.X., Meng, Q.W., Du, W.F., & He, D.X. (2017). Effects of light quality on growth and development of cucumber seedlings in controlled environment. International Journal of Agricultural and Biological Engineering, 10(3), 312–318. https://doi.org/3965/j.ijabe.20171003.2299
55. Spaargaren, J.J. (2001) Supplemental lightening for green house crop. Hortilux Schreder B.V. and P.L light System, Inc.-Ortario, Canada, p. 26-29.
56. Tabaka, P., & Wtorkiewicz, J. (2022). Analysis of the spectral sensitivity of luxmeters and light sensors of smartphones in terms of their influence on the results of illuminance measurements-example cases. Energies, 15, https://doi.org/10.3390/en15165847
57. Tamulaitis, G., Duchovskis, P., Bliznikas, Z., Breivė, K., Ulinskaite, R., Brazaityte, A., Novičkovas, A., & Žukauskas, A. (2005). High-power light wmmitting dioded based facility for plant cultivation. Journal of Physics. D: Applied Physics, 44, 261-269. https://doi.org/1088/0022-3727/38/17/S20
58. Tibbitts, T., Morgan, D., & Warrington, I. (1983). Growth of lettuce, spinach, mustard, and wheat plants under four combinations of high-pressure sodium, metal halide, and tungsten halogen lamps at equal PPFD. Journal of the American Society for Horticultural Science, 108, 622–630. https://doi.org/10.21273/JASHS.108.4.622
59. Trouwborst, G., Oosterkamp, J., Hogewoning, S.W., Harbinson, J., & van leperen, W., (2010). The responses of light interception, photosynthesis and fruit yield of cucumber to LED-lighting within the canopy. Physiologia Plantarum, 138, 289–300. https://doi.org/1111/j.1399-3054.2009.01333.x
60. Urbonaviciute, A., Pinho, P., Samuoliene, G., Duchovskis, P., Vitta P., Stonkus, A., Tamulaitis, G., Zukauskas, A., & Halonen, L. (2007). Effect of short wavelength light lettuce growth and nutritional quality. Sodininkyste ir Darzininkyste, 26(1), 157-165.
61. VanIeperen, W., Savvides, A., & Fanourakis, D. (2012). Red and blue light effects during growth on hydraulic and stomatal conductance in leaves of young cucumber plants. Acta Horticulturae, 956, 223–230. https://doi.org/17660/ActaHortic.2012.956.24
62. Volkenburgh, E.V. (1999). Leaf expansion–an integrating plant behaviour. Plant Cell and Environment, 22(12), 1463–1473. https://doi.org/1046/j.1365-3040.1999.00514.x
63. Wang, H., Gu, M., Cui, J., Shi, K., Zhou, Y., & Yu, J. (2009). Effects of light quality on CO₂ assimilation, chlorophyll-fluorescence quenching, expression of Calvin cycle genes and carbohydrate accumulation in Cucumis sativus. Journal of Photochemistry and Photobiology B: Biology, 96(1), 30–37. https://doi.org/1016/j.jphotobiol.2009.03.010
64. Watjanatepin, N. (2019). Effect of three specific spectra of LED light on the growth, yield, and fruit quality of Sida tomato. International Journal of Advanced and Applied Sciences, 6(6), 15-21. https://doi.org/10.21833/ijaas.2019.06.003
65. Wei, B., Song, C.Y., Wang, S.J., Sang, S.P., Li, F.T., Wang, Z.Z., & Yang, J.Z. (2018). Nitrogen application time and R/Fr ratio: Effect on growth physiological characters and yield of maize. Chinese Agricultural Science Bulletin, 10–18. (In Chinese with English abstract)
66. Yang, Q.C. (2008). Application and prospect of light emitting diode (LED) in agriculture and bio-industry. Journal of Agricultural Science and Technology, 6, 42–47. https://doi.org/3969/j.issn.1008-0864.2011.05.06
67. Yang, X., Xu, H., Shao, L., Li, T., Wang, Y., & Wang, R. (2018). Response of photosynthetic capacity of tomato leaves to different LED light wavelength. Environmental and Experimental Botany, 150(3), 161–71. https://doi.org/10.1016/J.ENVEXPBOT.2018.03.013
68. Yorio, N.C., Goins, G.D., Kagie, H.R., Wheeler, R.M., & Sager, J.C. (2001). Improving spinach, radish, and lettuce growth under red light-emitting diodes (LEDs) with blue light supplementation. HortScience, 36(2), 380–383. https://doi.org/10.21273/HORTSCI.36.2.380
69. Zandavifard, Z., & Azizi, M. (2021). Influence of different light spectra on morphological traits and hypericin content in St. Johnʼs Wort (Hypericum perforatum). Journal Of Horticultural Science, 35(3), 331-339. (In Persian with English abstract). https://doi.org/ 10.22067/JHS.2021.57256.0
70. Zheng, L., & Van Labeke, M.C. (2017) Long-term effects of red-and blue-light emitting diodes on leaf anatomy and photosynthetic efficiency of three ornamental pot plants. Frontiers in Plant Science, 8(1): 917. https://doi.org/10.3389/fpls.2017.00917