The role of far-red light (FR) in photomorphogenesis and its use in greenhouse plant production

Bojka KUMP


Light energy is one of the most important factors regulating the growth and development of plants. In greenhouses and other controlled- environments in which the natural radiation intensities are often low, plant production relies on supplementary lighting to optimize the photosynthesis, increase production levels, and enable year-round production. For a long time, the research related to artificial lighting sources focused on the optimization of the efficiency of use for photosynthesis. The quality of light in plant production has been widely addressed only recently with the development of advanced LED technology that is energy efficient and enables the control of the spectral composition of light. Red and far-red light are sensed by the phytochromes that trigger several morphological and developmental processes that impact productivity and yield quality. Thus, to efficiently exploit all the advantages of LEDs and to develop LED arrays for specific plant applications, it is essential to understand thoroughly how light quality influences plant growth and development. This paper presents an overview of the recent developments in light quality manipulation, focusing on far-red light and the R: FR ratio, to improve yield and quality of products and to manage plant architecture and flowering in vegetable and ornamental horticulture.


far-red light; greenhouse; LEDs, light quality; photomorphogenesis; R: FR ratio

Full Text:



Ahmad, M., Jarillo, J.A., Smirnova, O., & Cashmore, A.R. (1998). The CRY1 blue light photoreceptor of Arabidopsis interacts with phytochrome A in vitro. Molecular Cell, 1, 939–948.

Akutsu, M., Izena, J., & Takakura, T. (2017). Effect of EOD-FR Treatment on the Growth and Morphology of Japanese Mustard Spinach and Pak-choi. Hort. Res. (Japan), 16 (4), 449-454.

Alrifai, O., Hao, X., Marcone, M.F., & Tsao, R. (2019). Current review of the modulatory effects of LED lights on photosynthesis of secondary metabolites and future perspectives of microgreen vegetables. J. Agric. Food Chem., 67, 6075-6090.

Ballaré, C. L. (2017). Phytochrome Responses: Think Globally, Act Locally. Trends in Plant Science, 22 (11), 909-911.

Ballaré, C.L. (2014). Light regulation of plant defense. Annu. Rev. Plant Biol., 65, 335–363.

Bantis, F., Smirnakou, S., Ouzounis, T., Koukounaras, A., Ntagkas, N., & Radoglou, K. (2018). Current status and recent achievements in the field of horticulture with the use of light-emitting diodes (LEDs). Sci. Hort., 235, 437-451.

Bilodeau, S.E., Wu, B., Rufyikiri, A., MacPherson, S., & Lefsrud, M. (2019). An update on plant photobiology and implications for Cannabis production. Front. Plant. Sci., 10, 296.

Bradbeer, J.W. (1971). Plastid Development in Primary Leaves of Phaseolus vulgaris: the effect of short blue, red, far-red, and white light treatments on dark-grown plants. Journal of Experimental Botany, 22 (2), 382–390.

Cao, K., Yu, J., Xu, D., Ai, K., Bao, E., & Zou, Z. (2018). Exposure to lower red to far-red light ratios improve tomato tolerance to salt stress. Plant Biol., 18, 92.

Cargnel, M.D., Demkura, P.V., & Ballaré, C.L. (2014). Linking phytochrome to plant im-munity: low red: far-red ratios increase Arabidopsis susceptibility to Botrytis cinerea by reducing the biosynthesis of indolic glucosinolates and camalexin. New Phytol. 204, 342–354.

Casal, J.J., & Smith, H. (1989). The function, action and adaptive significance of phytochrome in light-grown plants. Plant Cell Environ, 12, 855-862.

Casal, J.J., Sanchez, R.A., & Yanovsky, M.J. (1997). The function of phytochrome A. Plant cell environ, 20 (6), 313-819.

Cerrudo, I., Keller, M.M., Cargnel, M.D., et al. (2012). Low red/far-red ratios reduceArabidopsis resistance toBotrytis cinereaand jasmonate responses via a COI1-JAZ10-dependent, salicylic acid-independent mechanism. Plant Physiol. 158, 2042–2052.

Chang, M.-H., Das, D., Varde, P., & Pecht, M. (2012). Light emitting diodes reliability review. Microelectron. Reliab. 52, 762–782.

Chia, P., & Kubota, C. (2010). End-of-day far-red light quality and dose requirements for tomato rootstock hypocotyl elongation. HortScience, 45 (19), 1501-1506.

Cho, J., Park, J. H., Kim, J. K., & Schubert, E. F. (2017). White light-emitting diodes: history, progress, and future. Laser Photonics Rev, 11, 1600147.

Craig, D.S., & Runkle, E.S. (2013). A Moderate to high red to far-red light ratio from light-emitting diodes controls flowering of short-day plants. J. Amer. Soc. Hort. Sci., 138 (3), 167-172.

De Simone, S., Oka, Y., Inoue, Y., Nishioka, N., Tadano, S., & Inoue, Y. (2000). Evidence of phytochrome mediation in the low-pH-induced root hair formation process in lettuce (Lactuca sativa L. cv. Grand Rapids) seedlings. J. Plant Res., 113, 45-53.

De Wit, M., Spoel, S.H., Sanchez-Perez, G.F., et al. (2013). Perception of low red: far-redratio compromises both salicylic acid- and jasmonic acid-dependent pathogen de-fences in Arabidopsis. Plant J. 75, 90–103.

Demotes-Mainard, S., Péron, T., Corot, A. et al. (2016). Plant responses to red and far-red lights, applications in horticulture. Environ. Exp. Bot., 121, 4-21.

Downs, R.J., & Thomas, J.F. (1982) Phytochrome regulation of flowering in the long-day plant, Hyoscyamus niger. Plant Physiol., 70 (3), 898-900.

Elkins, C.A., Martin, M., & van Iersel, M.W. (2019). Growth and morphological responsesof Digitalis and Rudbeckia seedlings to supplemental far-red LED light. HortScience, 54(9) Supplement, S89.

Fanwoua, J., Vercambre, G., Buck-Sorlin, G., Dieleman, J.A., de Visser, P., & Génard, P. (2019). Supplemental LED lighting affects the dynamics of tomato fruit growth and composition. Sci. Hort., 256, 109402.

Favre, N., Bárcena, A., Bahima, J.V., Martinez, G., & Costa., L. (2018). Pulses of low intensity light as promising technology to delay postharvest senescence of broccoli. Postharvest Biol Technol, 142, 107-114.

Franklin, K.A., Toledo-Ortiz, G., Pyott, D.E., & Halliday, K.J. (2014). Interaction of light and temperature signalling. J. Exp. Bot., 65 (11), 2859-2871.

Furuya, M. (1993). Phytochromes: their molecular species, gene families and functions. Annu. Rev. Plant Physiol. Plant Mol. Biol., 44, 617–645.

González, C.V., Ibbara, S.E., Piccoli, P.N., Botto, J.F., & Boccalandro H.E. (2012). Phytochrome B increases drought tolerance by enhancing ABA sensitivity in Arabidopsis thaliana. Plant Cell Environ., 35, 1958-1968.

Gundel, P. E., Pierik, R., Mommer, L., & Ballaré, C. L. (2014). Competing neighbors: light perception and root function. Oecologia, 176, 1–10.

Hao, X., Zheng, J., Little, C., & Khosla S. (2012). Led inter-lighting in year-round greenhouse mini-cucumber production. Acta Hortic., 956, 335-340.

Holmes, M.G., & Smith, H. (1977). Function of phytochrome in natural environment 1, Characterization of daylight for studies in photomorphogenesis and photoperiodism. Photochem. Photobiol., 25, 533-538.

Ilias, I.F., & Rajapakse, N. (2005). The effects of end-of-the-day red and far-red light on growth and flowering of Petunia xhybrida ‘Countdown Burgundy’ grown under photoselective films. HortScience, 40 (1), 131-133.

Islam, M.A., Gislerod, H.R., Torre, S., & Olsen, J.E. (2015). Control of shoot elongation and hormone physiologyin poinsettia by light quality provided by light emitting diodes – a minireview. Acta Horticulturae, 1104, 131-136.

Islam, M.A., Tarkowska, D., Clarke, J.L., Blystad, D.R., Gislerod, H.R., Torre, S., & Olsen, J.E. (2014). Impact of end-of-day red and far-red light on plant morphology and hormone physiology of poinsettia. Sci. Hort., 174, 77.86.

Jung, J.H. (2016). Phytochromes function as thermosensors in Arabidopsis. Science, 354, 886-889.

Kalaitzoglou, P., van Ieperen, W., Harbinson, J., van der Meer, M., Martinakos, S., Weerheim, K., Nicole, C.C.S., & Marcelis, F.M. (2019). Effects of continuos or end-of-day far-red light on tomato plant growth, morphology, light absorbtion, and fruit production. Front. Plant. Sci., 10, 322.

Kang, C., Lian, H., Wang, F., Huang, J., & Yang, H. (2009). Cryptochromes, Phytochromes, and COP1 Regulate Light-Controlled Stomatal Development in Arabidopsis. Plant Cell, 21, 2624-2641.

Kim, H., Lin, M., & Mitchell, C.A. (2019). Light spectral and thermal properties govern biomass allocation in tomato through morphological and physiological changes. Environ. Exp. Bot., 157, 228-240.

Klem, K., Gargallo-Garriga A., Rattanapichai, W., Oravec, M., Holub, P., Veselá, B., Sardans, J., Peñuelas, J., & Urban, O. (2019). Distinct morphological, physiological, and biological responses ti light quality in barley leaves and roots. Front. Plant Sci., 10 (1026), 1-18.

Kono, M., Kawaguchi, H., Mizusawa, N.,Yamori, W., Suzuki , Y., & Terashima, I. (2020). Far-Red Light Accelerates Photosynthesis in the Low-Light Phases of Fluctuating Light, Plant and Cell Physiology, 61 (1), 192–202.

Kopsell, D.A., & Sams, C.E. (2013). Increases in shoot tissue pigments, glucosinolates, and mineral elements in sprouting broccoli after exposure to short-duration blue light from light emitting diodes. J. Am. Soc. Hortic. Sci., 138, 31-37.

Kurepin, L.V. Emery, R.J., Pharis, R.P., & Reid, D.M. (2007). Uncoupling light quality from light irradiance effects in Helianthus annuus shoots. Putative roles for plant hormones in leaf and internode growth. J. Exp. Bot., 58, 2145-2157.

Lagarias, J.C., & Rapoport, H. (1980). Chromopeptides from phytochrome: the structure and linkage of the Pfr form of the phytochrome chromophore. Journal of the American Chemical Society, 102, 4821-4828.

Larcher, W. (2003). Physiological plant ecology. Berlin Heidelberg: Springer-Verlag.

Legris, M., Ince, Y.Ç., & Fankhauser, C. (2019). Molecular mechanisms underlying phytochrome-controlled morphogenesis in plants. Nat Commun, 10, 5219.

Li, Q., & Kubota, C. (2009). Effects of supplemental light quality on growth and phytochemicals of baby leaf lettuce. Environ. Exp. Bot., 67, 59- 64.

Liscum, E.; Young, J.C.; Poff, K.L., & Hangarter, R.P. (1992). Genetic separation of pgototropism ad blue light inhibition of stem elongation. Plant Physiol., 100, 267-271.

Lorenzo, C.D., Sanchez-Lamas, M., Antonietti, M.S., & Cerdan, P.D. (2016). Emerging hubs in plant light and temperature signaling. Photochem. Photobiol. 92, 3–13.

Más, P., Devlin, P.F., Panda, S., & Kay, S.A. (2000). Functional interaction of phytochrome B and cryptochrome 2. Nature, 408, 207–211.

Massa, G. D., Emmerich, J. C., Mick, M. E., Kennedy, R., Morrow, R. C., & Mitchell, C. A. (2005). Development and testing of an efficient LED intracanopy lighting design for minimizing Equivalent System Mass in an advanced life-support system. Gravit. Space Bio Bull.,18, 87–88.

Massa, G.D., Kim, H.-H., Wheeler, R.M., & Mitchell, C.A. (2008). Plant productivity in response to LED lighting. HortScience, 43, 1951–1956.

McCree, K.J. (1972). The action spectrum, absorptance and quantum yield of photosynthesis in crop plants. Agric. Meteorol., 9, 191–216.

McGuire, R., & Agrawal, A.A. (2005). Trade-offs between the shade-avoidance response andplant resistance to herbivores? Tests with mutantCucumis sativus. Funct. Ecol., 19, 1025–1031.

Meng, Q., & Runkle, E.S. (2019). Far-red radiation interacts with relative and absolute blue and red photon flux densities to regulate growth, morphology, and pigmentation of lettuce and basil seedlings. Sci. Hort., 255, 269-280.

Meng, Q., Kelly, N., & Runkle, E.S. (2019). Substituting green or far-red radiation for blue radiation induces shade avoidance and promotes growth in lettuce and kale. Environ. Exp. Bot., 162, 383-391.

Morrow, R.C. (2008). LED lighting in horticulture. HortScience 43, 1947-1950.

Myers, J. (1971). Enhancement studies in photosynthesis. Annu. Rev. Plant Physiol., 22, 289–312.

Nagata, M., Yamamoto, N., Shigeyama, T. et al. (2015). Red/far red light controls arbuscular mycorrhizal colonization via jasmonic acid and strigolactone signalling. Plant Cell Physiol., 56 (11), 2100-2109.

Nájera, C., Guil-Guerrero, J.L., Enriquez, L.J., & Álvaro, J.E. (2018). LED-enhanced dietary and organoleptic qualities in postharvest tomato fruit. Postharvest Biol Technol, 145, 151-156.

Nelson, J. A., & Bugbee, B. (2014). Economic analysis of greenhouse lighting: light emitting diodes vs. high intensity discharge fixtures. PLOS ONE, 9(6), e99010.

Nishimura, Y., Wada, E., Fukumoto, Y., Aruga, H., & Shimoi, Y. (2012). The effect of spectrum conversion covering film on cucumber in soilless culture. Acta Hortic., 956, 481-487.

Olle, M., & Viršile, A. (2013). The effects of light-emitting diode lighting on greenhouse plant growth and quality. Agric. Food Sci. 22, 223–234.

Park, Y., & Runkle, E.S. (2017). Far-red radiation promotes growth of seedlings by increasing leaf expansion and whole-plant net assimilation. Environ. Exp. Bot., 136, 41-49.

Park, Y., & Runkle, E.S. (2019). Blue radiation attenuates the effects of the red to far-red ratio on extension growth but not on flowering. Environ. Exp. Bot., 168, 103871.

Parks, B.M., Quail, P.H., & Hangarter, R.P. (1996). Phytochrome A regulates red light induction of phototropic enhancement in Arabidopsis. Plant Physiology, 110, 155–162.

Patel, D., Basu, M., Hayes, S., Majlath, I., Hetherington, F.M., Tschaplinski,T.J., & Franklin, K.A. (2013). Temperature-dependent shade avoidanceinvolves the receptor-like kinase ERECTA. Plant J., 73, 980–992.

Pinho, P., Jokinen, K., & Halonen, L. (2017). The influence of the LED light spectrum on the growth and nutrient uptake of hydroponically grown lettuce. Lighting Res. Technol., 49 (7), 866-881.

Rajapakse, N.C., Young, R.E., McMahon, M.J., & Oi, R. (1999). Plant height control by photoselective filters: current status and future prospects. Hort Technology, 9 (4), 618-624.

Roberts, M.R., & Paul, N.D. (2006). Seduced by the dark side: integrating molecular and ecological perspectives on the influence of light on plant defence against pests and pathogens. New Phytol., 170, 677–699.

Runkle, E. S. (2013). Manipulating light quality to elicit desirable plant growth and flowering responses. IFAC Proceedings Volumes, 46 (4), 196-200.

Runkle, E.S., & Heins, R.D. (2001). Specific Functions of red, far red, and blue light in flowering and stem extension of long-day plants. J. Amer. Soc. Hort. Sci., 126 (3), 275-282.

Sadras, V. O., Hall, A. J., Trapani, N., & Vilella, F. (1989). Dynamics of rooting and root-length: leaf-area relationships as affected by plant population in sunflower crops. Field Crops Res., 22, 45–57.

Sánchez-Lamas, M., Lorenzo, C.D., & Cerdán, P.D. (2016). Bottom-up assembly of the pgytochrome network. PLoS Genet., 12 (11), e1006413.

Shibuya, T., Endo, R., Kitaya, Y., & Hayashi, S. (2016). Growth analysis and photosynthesis measurements of cucumber seedlings grown under light with different red to far-red ratios. HortSci. 51 (7), 843-846.

Shibuya, T., Itagaki, K., Tojo, M., Endo, R., & Kitaya, Y. (2011). Fluorescent illumination with high red-to-far-red ratio improves resistance of cucumber seedlings to powdery mildew. HortScience, 46 (3), 429-431.

Singh, D., Basu, C., Meinhardt-Wollweber, M., & Roth, B. (2015). LEDs for energy efficient greenhouse lighting. Renew. Sustain. Energy Rev., 49, 139-147.

Srikanth, A., & Schmid, M. (2011). Regulation of flowering time: all roads lead to Rome. Cell. Mol. Life Sci. CMLS, 68, 2013-2037.

Stutte, G.W., Edney, S., & Skerritt, T. (2009). Photoregulation of bioprotectant content of red leaf lettuce with light-emitting diodes. HortScience, 44 (1), 79-82.

Suzuki, A., Suriyagoda, L., Shigeyama, T. et al. (2011). Lotus japonicus nodulation is photomorphogenetically controlled by sensing the red/far red (R/FR) ratio through jasmonic acid (JA) signalling. Proc. Natl. Acad. Sci. USA, 108, 16837-16842.

Taiz, L., & Zeiger, E. (2014). Plant physiology. Sunderland, MA: Sinauer Associates Inc.

Thiele, A., Herold, M., Lenk, I., Quail, P.H., & Gatz, C. (1999). Heterologous expression of Arabidopsis phytochrome B in transgenic potato influences photosynthetic performance and tuber development. Plant Physiol., 120, 73-82.

Tokuhisha, J.G., Daniels, S.M., & Quail, P.H. (1985). Phytochrome in green tissue: Spectral and immunochemical evidence for two distinct molecular species of phytochrome in light-grown Avena sativa L. Planta, 164, 321-332.

Turnbull, M.H., & Yates, D.J. (1993). Seasonal variation in the red/far-red ratio and photon flux density in an Australian sub-tropical rainforest. Agr. For. Met.,64 (1-2), 111-127.

Viczián, A., Klose, C., Adam, E., & Nagy, F. (2017). New insights of red light-induced development. Plant Cell Environ., 40, 2457-2468.

Viršilė, A., Olle, M., & Duchovskis, P. (2017). “LED lighting in horticulture” in Light Emitting Diodes for Agriculture. ed. S.D. Gupta. (Singapore: Springer), 113–147.

Yang, F., Feng, L., Liu, Q., et al. (2018). Effect of interactions between light intensity and red-to far-red ratio on the photosynthesis of soybean leaves under shade condition. Environ. Exp. Bot., 150, 79-87.

Yang, Z., Kubota, C., Chia, P., & Kacira, M. (2012). Effect of end-of-day far-red light from a movable LED fixture on squash rootstock hypocotyl elongation. Sci. Hortic., 136, 81-86.

Yeh, N., & Chung, J.-P. (2009). High-brightness Leds-Energy efficient lighting sources and their potential in indoor plant cultivation. Renew. Sust. Energ. Rev. 13, 2175–2180.

Zahedi, S.M., & Sarikhani, H. (2016). Effect of far-red light, temperature, and plant age on morphological changes and induction of flowering of a ’June-bearing’ strawberry. Hortic. Environ. Biotechnol., 57 (4), 340-347.

Zhang, M., & Runkle, E.S. (2019). Regulating Flowering and Extension Growth of Poinsettia Using Red and Far-red Light-emitting Diodes for End-of-day Lighting. HortScience, 54 (2), 323-327.

Zhang, M., Whitman, C.M., & Runkle, E.S. (2019a). Manipulating growth, color, and taste attributes of fresh cut lettuce by greenhouse supplemental lighting. Scientia Hort., 252, 274-282.

Zhang, Y., Zhang, Yu., Yang, Q., & Tao, L. (2019b). Overhead supplemental far-red light stimulates tomato growth under intra-canopy lighting with LEDs. J. Integr. Agric., 17, 62-69.

Zhen, S., & Bugbee, B. (2019). Far-Red Photons Are Necessary for Efficient Photosynthesis: Whole-Canopy Photosynthesis and Radiation Capture. HortScience, 54 (9), S178.

Zhen, S., & van Iersel, M.W. (2017). Far-red light is needed for efficient photochemistry and photosynthesis. J. Plant Physiol, 209, 115-122.

Zhen, S., Haidekker, M., & van Iersel, M.W. (2019). Far-red light enhances photochemical efficiency in a wavelength-dependent manner. Physiologia plantarum, 167 (1), 21-33.

Zou, J., Zhang, Y., Zhang, Yu., Bian, Z., Fanourakis, D., Yang, Q., & Li, T. (2019). Morphological and physiological properties of indoor cultivated lettuce in response to additional far-red light. Scientia Hort., 257, 108725.



  • There are currently no refbacks.

Copyright (c) 2020 Bojka Kump

Creative Commons License
This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.


Acta agriculturae Slovenica is an Open Access journal published under the terms of the Creative Commons CC BY License.


eISSN 1854-1941