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Photoreceptor Signaling Networks in Plant Responses to Shade
Open accessAbstract
The dynamic light environment of vegetation canopies is perceived by phytochromes, cryptochromes, phototropins, and UV RESISTANCE LOCUS 8 (UVR8). These receptors control avoidance responses to preclude exposure to limiting or excessive light and acclimation responses to cope with conditions that cannot be avoided. The low red/far-red ratios of shade light reduce phytochrome B activity, which allows PHYTOCHROME INTERACTING FACTORS (PIFs) to directly activate the transcription of auxin-synthesis genes, leading to shade-avoidance responses. Direct PIF interaction with DELLA proteins links gibberellin and brassinosteroid signaling to shade avoidance. Shade avoidance also requires CONSTITUTIVE PHOTOMORPHOGENESIS 1 (COP1), a target of cryptochromes, phytochromes, and UVR8. Multiple regulatory loops and the input of the circadian clock create a complex network able to respond even to subtle threats of competition with neighbors while still compensating for major environmental fluctuations such as the day-night cycles.
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“Plants quantify this ratio through the phytochrome system and react with the shade avoidance syndrome that enables plants to outgrow the competitors for light (Casal, 2013; Pierik and de Wit, 2014).”
From Shaping plant architecture (2015) · cites this paper as Casal, 2013
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“Cryptochromes ( CRY ) such as CRY1 , CRY2 , and phototropins ( PHOT ) like PHOT1 , PHOT2 are the UV-A/ blue light photoreceptors, and UV RESISTENCE LOCUS8 ( UVR8 ) is the UV-B absorbing photoreceptor that has been discovered lately (Favory et al. 2009; Rizzini et al. 2011; Casal 2013).”
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“Light is crucial for plant growth [1,2], significantly affecting development under canopy shade.”
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Dephosphorylation of bZIP59 by PP2A ensures appropriate shade avoidance response in Arabidopsis
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“PhyA can make a direct contribution to shade responses, which becomes obvious in the phyB mutant background [16].”
BackgroundRockwell, N. C., & Lagarias, J. C. (2023) · Journal of Molecular Biology · doi:10.1016/j.jmb.2023.168313 · Cited by 13
GA signalling and cross-talk with other signalling pathways
Lor, V. S., & Olszewski, N. E. (2015) · Essays in Biochemistry · doi:10.1042/bse0580049 · Cited by 13
“Light signalling has recently been reviewed [22].”
Auxin Contributes to the Intraorgan Regulation of Gene Expression in Response to Shade
Kim, S.-J., Mochizuki, N., Deguchi, A., Nagano, A. J., Suzuki, T., & Nagatani, A. (2018) · PLANT PHYSIOLOGY · doi:10.1104/pp.17.01259 · Cited by 13
Kebrom, T. H., McKinley, B. A., & Mullet, J. E. (2020) · Plant Direct · doi:10.1002/pld3.235 · Cited by 13
“Noticeable differential expression of circadian clock genes has not been reported in transcriptome studies of SAS in Arabidopsis seedlings and in the stem of tomato and other species (Cagnola, Ploschuk, Benech-Arnold, Finlayson, & Casal, 2012; Devlin et al., 2003).”
BackgroundInfluentialBernado, W. D. P., Baroni, D. F., Ruas, K. F., Santos, A. R., de Souza, S. B., Passos, L. C., Façanha, A. R., Ramalho, J. C., Campostrini, E., Rakočević, M., et al. (2022) · Scientia Horticulturae · doi:10.1016/j.scienta.2022.110881 · Cited by 13
Sher, J., Zheng, Y., Burns, J. H., Jan, G., & Zhang, J. (2025) · Journal of Plant Interactions · doi:10.1080/17429145.2025.2548579 · Cited by 12
“Light plays a crucial role in photosynthesis, plant growth, and overall health its availability can lead to strong competition among plant species (De Wit et al. 2012; Casal 2013; Simkin et al. 2022).”
Chen, B., Shi, Y. Y., Sun, Y., Lu, L., Wang, L., Liu, Z., & Cheng, S. (2024) · aBIOTECH · doi:10.1007/s42994-023-00129-1 · Cited by 13
“Plants utilize a range of photoreceptors to sense light, among which cryptochromes and phytochromes are notable (Mo¨glich et al. 2010; Casal 2013).”
BackgroundOskam, L., Snoek, B. L., Pantazopoulou, C. K., van Veen, H., Matton, S. E. A., Dijkhuizen, R., & Pierik, R. (2024) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiae097 · Cited by 12
“This is particularly challenging in dense vegetation, both in natural and agricultural fields, where plants compete for access to this resource because they mutually shade each other (Casal 2013; Huber et al. 2021).”
BackgroundProtein design accelerates the development and application of optogenetic tools
Zhu, L., Wang, Y., Wu, X., Wu, G., Zhang, G., Liu, C., & Zhang, S. (2025) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2025.02.014 · Cited by 13
“Most photosensitive proteins are found in plants, where they play crucial roles in mediating cellular responses to light, including phytochromes, cryptochromes, phototropins, and UV Resistance Locus 8 (UVR8) [7,8].”
Yang, X., Wang, S., Liu, W., Huang, S., Xie, Y., Meng, X., Li, Z., Jin, N., Jin, L., Lyu, J., et al. (2025) · Plants · doi:10.3390/plants14091369 · Cited by 12
“Endogenous hormones play an indispensable role in the growth and development of plants [58,59].”
Plantenga, F. D. M., Heuvelink, E., Rienstra, J. A., Visser, R. G. F., Bachem, C., & Marcelis, L. F. M. (2018) · Physiologia Plantarum · doi:10.1111/ppl.12885 · Cited by 12
Krahmer, J., Abbas, A., Mengin, V., Ishihara, H., Romanowski, A., Furniss, J. J., Moraes, T. A., Krohn, N., Annunziata, M. G., Feil, R., et al. (2021) · Journal of Experimental Botany · doi:10.1093/jxb/erab038 · Cited by 11
Chen, Y., Yang, H., Tang, B., Li, F., Xie, Q., Chen, G., & Hu, Z. (2022) · Plant Cell Reports · doi:10.1007/s00299-022-02963-x · Cited by 12
“Light signals can be perceived by families of photoreceptors, including phytochromes, cryptochromes, phototropins, and UV resistance locus 8 (UVR8) (Casal 2013).”
BackgroundRanade, S. S., & García‐Gil, M. R. (2015) · Forest Ecology and Management · doi:10.1016/j.foreco.2015.11.034 · Cited by 12
Shade avoidance cues reduceBeta vulgarisgrowth
Schambow, T. J., Adjesiwor, A. T., Lorent, L., & Kniss, A. R. (2019) · Weed Science · doi:10.1017/wsc.2019.2 · Cited by 12
“Plants have evolved the ability to sense and respond to changes in the quality of light using a family of photoreceptors, particularly cryptochromes and phytochromes, which act as molecular switches in response to blue-violet light and R:FR ratio, respectively (Casal 2013).”
BackgroundKalmatskaya, O. А., Karavaev, V. A., & Tikhonov, A. N. (2019) · Photosynthesis Research · doi:10.1007/s11120-019-00663-4 · Cited by 12
Quantification of Spectral Perception of Plants with Light Absorption of Photoreceptors
Kang, W. H., Kim, J.-W., Yoon, H. I., & Son, J. E. (2020) · Plants · doi:10.3390/plants9050556 · Cited by 12
“High fractions of green and far-red light induce shade-avoidance syndrome, represented by an elongated hypocotyl and petioles to enable the plant to move out of shade [8,9] because those conditions occur when the red and blue light within solar radiation is filtered out by dense foliage above or nearby [10–12].”
BackgroundFunctional specialization of Nicotiana attenuata phytochromes in leaf development and flowering time
Fragoso, V., Oh, Y., Kim, S., Gase, K., & Baldwin, I. T. (2016) · Journal of Integrative Plant Biology · doi:10.1111/jipb.12516 · Cited by 12
Zhao, J. C., Shi, X., Chen, L., Chen, Q., Tian, X., Ai, L., Zhao, H., Yang, C., Yan, L., & Zhang, M. (2022) · Frontiers in Plant Science · doi:10.3389/fpls.2022.973643 · Cited by 12
“Phototropins may play a role in shade avoidance by controlling phototropic growth in response to gradients of blue light (Galen et al., 2004; Casal, 2013; Fraser et al., 2016; Pierik and Ballare, 2021).”
BackgroundInfluentialThe role of plant-derived smoke and karrikinolide in abiotic stress mitigation: An Omic approach
Singh, S., Uddin, M., Khan, M. M. A., Chishti, A. S., Singh, S., & Bhat, U. H. (2023) · Plant Stress · doi:10.1016/j.stress.2023.100147 · Cited by 14
Lei, K., Hu, H., Chang, M., Sun, C., Ullah, A., Yu, J., Dong, C., Gao, Q., Jiang, D., Cao, W., et al. (2023) · Plant Physiology and Biochemistry · doi:10.1016/j.plaphy.2023.107850 · Cited by 11
Rosati, A., Proctor, K., Dazaea, A., Graham, M., Ateş, S., Kirschten, H. M., & Higgins, C. W. (2023) · Agroforestry Systems · doi:10.1007/s10457-023-00914-3 · Cited by 11
“The red/far-red ratio (R:FR), for example, decreases with increasing canopy shade in forests and dense canopies (De Castro 2000; Kurepin et al. 2006; Casal 2013; Park and Runkle 2017), because R radiation is absorbed much more than FR (Woolley 1971; Kasperbauer 1987; Ruberti et al. 2012).”
Zhang, Y., Li, J., Xu, Q., Chen, C., Nie, S., Lei, J., & Duan, L. (2024) · Frontiers in Plant Science · doi:10.3389/fpls.2024.1498123 · Cited by 11
Mu, D., Shui, Z., Guo, H., Jiang, H., Liu, Y., Luo, L., Zhang, Y., Zhang, X., Yu, J., Liao, S., et al. (2024) · BMC Plant Biology · doi:10.1186/s12870-024-05950-4 · Cited by 11
Makinen, A. S., Ishihara, H., Poque, S., Sipari, N., Himanen, K., Varjus, I., Heininen, J., Pastell, M., Elomaa, P., Shapiguzov, A., et al. (2025) · Physiologia Plantarum · doi:10.1111/ppl.70405 · Cited by 11
Layering contrasting photoselective filters improves the simulation of foliar shade
Petrella, D. P., Breuillin‐Sessoms, F., & Watkins, E. (2022) · Plant Methods · doi:10.1186/s13007-022-00844-8 · Cited by 11
Vodnik, D., Vogrin, Ž., Šircelj, H., Grohar, M. C., Medič, A., Carović‐Stanko, K., Safner, T., & Lazarević, B. (2022) · Scientia Horticulturae · doi:10.1016/j.scienta.2022.111638 · Cited by 11
Metadata
- Authors
- Casal, J. J.
- Affiliations
- Consejo Nacional de Investigaciones Científicas y Técnicas (AR)
- Corresponding author
- Casal, Jorge J.
- ORCID
- Casal, Jorge J.
- Venue
- Annual Review of Plant Biology
- Publisher
- Annual Reviews
- Published
- 2013-04-29
- Volume
- 64
- Issue
- 1
- Pages
- 403-427
- ISSN
- 1543-5008 (ISSN-L)
- 1545-2123
- OpenAlex
- W2107201571
- Cited by
- 865
- References
- 144
- Citations by year
- 2026: 81 · 2025: 66 · 2024: 72 · 2023: 61 · 2022: 79 · 2021: 91 · 2020: 67 · 2019: 60 · 2018: 62 · 2017: 60 · 2016: 58 · 2015: 44 · 2014: 51 · 2013: 13
- FWCI
- 5.94
- Type
- journal article
- Language
- en
- Open access
- green
- hdl.handle.net
- Licence
- cc-by-nc-nd
- Locations
- Annual Review of Plant Biology (publishedVersion)
- PubMed (publishedVersion)
- CONICET Digital (CONICET) (submittedVersion, cc-by-nc-nd)
- Funders
- Consejo Nacional de Investigaciones Científicas y Técnicas
- Universidad de Buenos Aires (CONICET)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Light effects on plants
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Parasitism and Resistance
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Molecular Biology Research
- Keywords
- shade avoidance, photoreceptors, phytochrome, cryptochrome, phototropin, UVR8, phytochrome-interacting factors, DELLA proteins, gibberellin signaling, brassinosteroid signaling, auxin biosynthesis, COP1, circadian clock, red:far-red ratio, plant light signaling, canopy light environment
- MeSH terms
- Gibberellins, Light, Phytochrome, Plant Growth Regulators, Plants, Signal Transduction, Ecosystem, Photoreceptors, Plant
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