Cited by 866Light effects on plantsPlant Parasitism and ResistancePlant Molecular Biology Research

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Photoreceptor Signaling Networks in Plant Responses to Shade

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Casal, Jorge J.·2013·Annual Review of Plant Biology

DOI 10.1146/annurev-arplant-050312-120221

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Abstract

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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“On the other hand, the initial difference in leaf growth between control and thinned plants can be attributed to neighboring plants sensing and thus at least in part phytochrome‑mediated signaling (Casal 2013) may be involved.”
Improving phytochemicals and antioxidant activity of clove basil (Ocimum gratissimum L.) with far-red and UV-A light supplementation in late production stages
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Brassinosteroids' regulation of plant architecture
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Plant growth: Sentinel leaf tip communicates the shade threat to its base
Casal, J. J. (2023) · Current Biology · doi:10.1016/j.cub.2022.10.057 · Cited by 2
Self-citation
Approaches to Study Light Effects on Brassinosteroid Sensitivity
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“According to Casal (2013), the expression of genes for biosynthesis of gibberellin was sensitive against light alteration.”
Background
The effect of red light (660 nm) on proliferative activity and growth reactions in seedlings of plants with contrast photoperiodic reaction
Авксентьева, О. А., & Batueva, E. D. (2020) · The Journal of V N Karazin Kharkiv National University Series Biology · doi:10.26565/2075-5457-2020-34-16 · Cited by 2
Information Processing and Survival Strategies
Heilmann, I. (2014) · doi:10.1002/9783527686063.ch7 · Cited by 2
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“…avoidance, phytochromes are also involved in regulation of seed germination, photomorphogenesis, flowering time, the circadian clock and gravitropism (Casal, 2013; Correll et al., 2003; Franklin & Whitelam, 2007b; Lin, 2000; Nagy & Schafer, 2002; Pierik & de Wit, 2014; Somers, Devlin, & Kay, 1998).”
Background
Molecular Breeding Strategies for Enhancing Rice Yields Under Low Light Intensity
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“Plant PHYs are also capable of perceiving changes in the R/FR ratio of ambient light caused by neighboring vegetation, initiating shade avoidance and acclimation reactions ( 7 ).”
Background
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Belkov, V. I., Belogub, K. E., Garnik, E. Y., Тарасенко, В. И., & Konstantinov, Y. M. (2021) · Proceedings of universities Applied chemistry and biotechnology · doi:10.21285/2227-2925-2021-11-2-251-259 · Cited by 1
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“SAS involves responses such as erect leaf angles and stem elongation, which are well studied in Arabidopsis and tomatoes (Casal 2013).”
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“Many studies reported that red color light stimulated shoot elongation in plants (Casal, 2013; dos Reis Oliveira et al., 2020).”
Identification of Novel Quantitative Trait Loci and Candidate Genes Associated with Grain Yield and Related Traits Under Low-Light Stress Conditions in Rice
Mohanty, S., Das, S., Panda, D., Choudhury, N., Mishra, B., Jena, R. K., Sah, R. P., Anilkumar, C., Devanna, B. N., R., R. R. K., et al. (2025) · Biomolecules · doi:10.3390/biom15101388 · Cited by 1
“This compromised physiological state weakens the plant’s resilience against stressors, further emphasizing the impact of low-light conditions [56].”
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Water shortage reduces PHYTOCHROME INTERACTING FACTOR 4 , 5 and 3 expression and shade avoidance in Arabidopsis
Semmoloni, M., Rojas, C. C., Escudero, C. M., Yan, Y., Sánchez, D. H., & Casal, J. J. (2025) · The Plant Journal · doi:10.1111/tpj.70610 · Cited by 4
Self-citation
Avoiding the shadow: how plants perceive neighbors and reshape the crop light environment
Pereyra, M. E., Sadras, V. O., & Casal, J. J. (2026) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiag034 · Cited by 1
Self-citation
The effect of discontinuous blue light intervention on red-light-grown tomato fruit coloration
Chen, X., Liu, Y., Gao, X., Zhai, Y., Wei, X., & Wang, L. (2026) · Scientia Horticulturae · doi:10.1016/j.scienta.2026.114688 · Cited by 1
Light Mediated Flowering in Plants: Mechanisms and Molecular Insights
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“Far-red light (700–750 nm) enhances shade-avoidance responses through PHYB inactivation and PIF (Phytochrome Interacting Factors) signalling, often accelerating floral initiation, particularly when applied as end-of-day far-red treatments (Casal 2013; Smith 2000).”
Integrated phenotype, endogenous hormones and transcriptome analysis revealed the mechanism of response of Phoebe bournei seedlings to shade signals
Huo, H., Wei, X., Jiang, F., Feng, S., & An, J. (2026) · BMC Genomics · doi:10.1186/s12864-025-12289-z · Cited by 1
“It is shown that the R: FR ratio ranges from 1 to 1.5 in natural light and drops to below 1.0 in the presence of neighbors or under the canopy [3, 4].”
Influential
Spatial Proximity to Perennial Groundcover Triggers Shade Avoidance Responses in Corn
Moro, A., Goggi, A. S., Moore, K. J., Fei, S., & Kaleita, A. L. (2026) · Agronomy · doi:10.3390/agronomy16070729 · Cited by 1
Phytochrome‐mediated shade‐avoidance responses and its impact on growth and physiology in crops—A review
Surya, K., Krishnan, R., Saitheja, V., Janaki, P., Parameswari, E., Govindan, S. K., Gnanachitra, M., Suresh, K., Paul, R., & Ammaiyappan, A. (2026) · Photochemistry and Photobiology · doi:10.1111/php.70098 · Cited by 1
UV wavelength- and dose-dependent changes in the biosynthesis of phenolic compounds in kale (Brassica oleracea var. acephala) plants
Yoon, H. I., Choi, Y.-S., & Oh, M. (2026) · Photochemical & Photobiological Sciences · doi:10.1007/s43630-026-00919-8 · Cited by 1
“UV-A radiation is partially absorbed in blue light photoreceptors, such as cryptochromes and phototropins [ 11, 12].”

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
DOI: 10.1146/annurev-arplant-050312-120221
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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Photoreceptor Signaling Networks in Plant Responses to Shade

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