The DOI matches its Crossref record; 144 of the 144 references it cites were found in a registry or an index.
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.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“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
Citations
Cites
Phytochromes and light signal perception by plants—an emerging synthesis
Smith, H. (2000) · Nature · doi:10.1038/35036500 · Cited by 1,385
A molecular framework for light and gibberellin control of cell elongation
de Lucas, M., Davière, J., Rodríguez-Falcón, M., Pontín, M., Iglesias‐Pedraz, J. M., Lorrain, S., Fankhauser, C., Blázquez, M. A., Titarenko, E., & Prat, S. (2008) · Nature · doi:10.1038/nature06520 · Cited by 1,320
Light Quality, Photoperception, and Plant Strategy
Smith, H. (1982) · Annual Review of Plant Physiology · doi:10.1146/annurev.pp.33.060182.002405 · Cited by 1,253
Coordinated regulation of Arabidopsis thaliana development by light and gibberellins
Feng, S., Martínez, C., Gusmaroli, G., Wang, Y., Zhou, J., Wang, F., Chen, L., Yu, L., Iglesias‐Pedraz, J. M., Kircher, S., et al. (2008) · Nature · doi:10.1038/nature06448 · Cited by 1,118
Tao, Y., Ferrer, J.-L., Ljung, K., Pojer, F., Hong, F., Long, J. A., Li, L., Moreno, J. E., Bowman, M. E., Ivans, L. J., et al. (2008) · Cell · doi:10.1016/j.cell.2008.01.049 · Cited by 1,062
PIFs: pivotal components in a cellular signaling hub
Leivar, P., & Quail, P. H. (2010) · Trends in Plant Science · doi:10.1016/j.tplants.2010.08.003 · Cited by 977
Phototropin Blue-Light Receptors
Christie, J. M. (2007) · Annual Review of Plant Biology · doi:10.1146/annurev.arplant.58.032806.103951 · Cited by 937
The ELF4–ELF3–LUX complex links the circadian clock to diurnal control of hypocotyl growth
Nusinow, D. A., Helfer, A., Hamilton, E. E., King, J. J., Imaizumi, T., Schultz, T. F., Farré, E. M., & Kay, S. A. (2011) · Nature · doi:10.1038/nature10182 · Cited by 908
phot1 and phot2 mediate blue light regulation of stomatal opening
Kinoshita, T., Doi, M., Suetsugu, N., Kagawa, T., Wada, M., & Shimazaki, K.-I. (2001) · Nature · doi:10.1038/414656a · Cited by 899
Interaction between BZR1 and PIF4 integrates brassinosteroid and environmental responses
Oh, E., Zhu, J.-Y., & Wang, Z. (2012) · Nature Cell Biology · doi:10.1038/ncb2545 · Cited by 886
Research works printed but not linked: 2
- Jang IC, 2010, Arabidopsis, 22, 2370
- 10.1111/j.1399-3054.1996.tb00205.x · DOI
Cited by
Sorted by citations
Items 801–866 of 866
Monteagudo, A., Kiss, T., Mayer, M., Casas, A. M., Igartua, E., & Karsaï, I. (2020) · Figshare · doi:10.6084/m9.figshare.12291452
Petrella, D. P., Breuillin‐Sessoms, F., & Watkins, E. (2022) · Figshare · doi:10.6084/m9.figshare.19143116
Sun, T., Wang, Y., Wu, X., Ye, J., & Cheng, F. (2023) · Figshare · doi:10.6084/m9.figshare.22622344
Zheng, L.-W., Ma, S.-J., Zhou, T., Yue, C.-P., Hua, Y., & Huang, J.-Y. (2021) · Figshare · doi:10.6084/m9.figshare.14843209
Far-Red Light Regulates the Circadian Rhythm Pathway to Accelerate Rice Flowering
Li, Z., Zhou, C., Hu, J., Xie, J., Yuan, Q., Wang, F., Wang, S., & Yang, Q. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27041683
“This can be attributed to the role of GA in mediating the shade avoidance syndrome, specifically in promoting internode elongation and accelerating the transition to flowering [31].”
Liu, Y., Liu, G., Dong, R., Zhang, Y., Huan, H., Yu, D., Wang, W., & Wang, Z. (2026) · Grass Research · doi:10.48130/grares-0025-0033
Li, F., Zeng, Y., Wang, J., & Lin, L. H. (2026) · Plant Communications · doi:10.1016/j.xplc.2026.101789
“In agricultural systems, SAS in shade-intolerant crops restricts planting density and ultimately limits yield (Ballare´ and Pierik, 2017).”
InfluentialPlant movements: navigating the light environment
Matton, S. E. A., Oskam, L., & Pierik, R. (2026) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiag110
“Active phytochromes repress phytochrome interacting factors (PIFs) (Casal 2013), a subgroup of bHLH transcription factors (Leivar and Monte 2014), through direct interaction.”
Trasser, M., Robson, T. M., & Durand, M. (2026) · BMC Plant Biology · doi:10.1186/s12870-026-08455-4
“…under controlled conditions have been well described, but most studies have relied on plants grown under steady light and temperature conditions in growth rooms or chambers, and subsequently exposed to the spectral regions of interest only for a relatively short period of time [11, 16, 29].”
Deslous, P. (2018) · theses.fr (ABES) · doi:10.70675/01735f17z51cez4bf0zba24zc47edf4ac548 · Cited by 1
Basso, V. (2019) · theses.fr (ABES) · doi:10.70675/e78df998z0331z447dz8c26z92f0b9a6942e · Cited by 1
Chou, C. T., Christabel, V., Le, M. A., Tsai, M., & Wang, S. (2026) · Agronomy · doi:10.3390/agronomy16080770
Decoding light perception and fine-tuning photomorphogenesis through posttranslational modifications
Gopalan, S. M., Priyanka Nair, T., Regy, A., T, S. K., & Srivastava, M. (2026) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiag189
Tao, Q., Wang, G., Wang, Z., An, Y., Ma, L., & Li, G. (2026) · Plant Communications · doi:10.1016/j.xplc.2026.101848
“Prolonged shade inhibits axillary bud outgrowth, reducing branching and promoting the elongation of stems or stalks (Casal, 2013; Schneider et al., 2019).”
Regulation of Phenolics Accumulation by Blue Light and Salicylic Acid in Mung Bean Sprouts
Kang, H., Cheng, Y., & Guo, X. (2026) · Journal of Plant Growth Regulation · doi:10.1007/s00344-026-12172-5
“BL affects certain physiological responses to plant growth and metabolite biosynthesis mainly by modulating CRY and phototropin levels (Landi et al. 2020).”
Liu, Z., Yu, H., Zhao, X., Nan, Z., Zhang, J., Guo, F., Zhang, Z., & Wan, S. (2026) · Journal of Integrative Agriculture · doi:10.1016/j.jia.2026.04.035
A plant-centered perspective on the radiation spectrum
Jan, S., & Bhat, G. H. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-21727-2.00006-x
Control and coordination between light spectrum and plant metabolism
Jan, S., & Bhat, G. H. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-21727-2.00005-8
Optical properties of leaf structural elements
Jan, S., & Bhat, G. H. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-21727-2.00002-2
Han, Y., Yu, J., Jun, S.-H., Bang, K.-H., & Kim, Y.-G. (2026) · Korean Journal of Medicinal Crop Science · doi:10.7783/kjmcs.2026.34.2.95
Sousa, A. L. B. D., & Nazaré, K. C. D. (2026) · Revista Multidisciplinar do Nordeste Mineiro · doi:10.66104/v908th38
Decoding darkness by seeking photoreceptor functions with and without light
Zeng, D., & Liu, H. (2026) · Nature Plants · doi:10.1038/s41477-026-02307-7 · Cited by 1
“The perceived light environment is sculpted by multi-scale interactions—from planetary rhythms to leaf shading—creating an inter-woven continuum of light and darkness that varies across time, space, direction, spectrum and intensity 4 .”
Bogino, M. F., Senz, J. M. L., Tamagnone, N., Kourdova, L. T., Romanowski, A., Wirthmueller, L., & Fabro, G. (2026) · Plant Physiology and Biochemistry · doi:10.1016/j.plaphy.2026.111418
Le, Q. V., Đào, T. M. C., Nguyen, A. D., Nguyen, T. T., & Nguyen, T. B. L. (2026) · Forests · doi:10.3390/f17060643
Han, J., Tian, Y., Liu, J., Deng, H., Han, K., Si, H., Shasha, Ling, Yu, L., El-Kassaby, Y. A., et al. (2026) · Horticulture Research · doi:10.1093/hr/uhag202
“PHYA is a key photoreceptor that senses low-light environments [ 45– 47].”
Xiang, M., Jin, D., Zeng, H., Yan, Z., Fang, L., Sun, H., Zheng, G., Liao, J., Si, X., Zhu, Y., et al. (2026) · The Crop Journal · doi:10.1016/j.cj.2026.05.007
Zhao, F., Yang, R., Song, Z., Bian, Y., Xiao, Y., & Xu, D. (2026) · PLoS Genetics · doi:10.1371/journal.pgen.1012177
“Shade-intolerant plants undergo SAS when grown in crowded environments [4,5].”
Ren, R., Juan, Y. A. N. G., Hongchao, L. I., Kong, D., Liu, Q., Haiyang, W. A. N. G., & Wang, S. (2026) · Maize Sciences · doi:10.2738/ms.2026.0006
Redundant in the light, indifferent to the heat: the role of the PIF family in mature tomato
Thomas, H. R. (2026) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiag380
“Shade has a low R:FR, which inactivates PhyB (Casal 2013).”
Light Signaling: Phytochrome and Photoreceptors
Luo, F., Mickles, A. J., & Fu, W. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-40403-0.00011-1
Mendes, F. M., Leite, D. M., de Sousa, K. I. R., de Andrade, C. M. L., Miranda, D. P., & Brondani, G. E. (2026) · Plant Cell Tissue and Organ Culture (PCTOC) · doi:10.1007/s11240-026-03516-w
“…activated by red light, triggering physiological responses similar to shade conditions, inducing auxin and gibberellin biosynthesis, which promote cell elongation and division, resulting in stem and petiole elongation, reduced leaf mass, and lower chlorophyll content (Casal 2013; Wei et al. 2023).”
Rosati, A., Kirschten, H. M., Graham, M., Maxwell, T. M. R., Dazaea, A., & Ateş, S. (2026) · Agroforestry Systems · doi:10.1007/s10457-026-01606-4
“In particular, green canopies absorb much more R radiation while transmitting more FR (Woolley 1971; Kasperbauer 1987; Ruberti et al. 2012), thus R/FR decreases with increasing canopy shade (De Castro 2000; Kurepin et al. 2006; Casal 2013; Park and Runkle 2017).”
He, Y. U. E. J. I., Wei, Y., Chen, Y., Feng, J., Zhou, D., Zhang, M., Qi, D., Wang, W. M., Wang, P. Y., Xie, J., et al. (2026) · Horticulturae · doi:10.3390/horticulturae12080908
QIN, W., Myint, H. S., Chai, J., Deng, J., Guo, J., Zhao, G., & Liu, J. (2026) · Research Square · doi:10.21203/rs.3.rs-10522073/v1
Zha, L., Zheng, X., He, X., Liu, L., Chen, Z., & Zhang, J. (2026) · Horticulturae · doi:10.3390/horticulturae12080933
Magnin, L., Hiltpold, I., Baux, A., Fortineau, A., & Jullien, A. (2026) · Frontiers in Plant Science · doi:10.3389/fpls.2026.1878300
“…environments and are generally associated with changes in resource availability and plant perception of neighboring vegetation (Ballare ́ al., 1987; Casal, 2013). maintaining or improving yield performance in Among the environmental factors modi fi ed by the presence of FB, temperature appeared…”
Kreslavski, V. D., Shmarev, A. N., Khudyakova, A., Strokina, V. V., Elizarov, D. O., & Kosobryukhov, A. A. (2026) · Russian Journal of Plant Physiology · doi:10.1134/s1021443726603447
“…indicator of plant canopy density: a decrease in RL/FRL 1 significantly signals shading, causing the so-called shade avoidance syndrome (SAS)—hypocotyl elongation, changes in leaf area, acceleration of flowering, and redistribution of resources to the stem at the expense of the root system [3, 4].”
Huang, Y., Ding, M. B., Bai, F. H., Zhang, Y. B., Jiang, F. J., Yan, X. Y., Li, Q. F., & Xu, T. L. (2026) · Russian Journal of Plant Physiology · doi:10.1134/s1021443726603320
“This aboveground increase likely results from stem elongation and leaf expansion—typical responses to deep shade that serve to position leaves in higher light layers [18].”
Liu, Z. (2026) · Springer eBooks · doi:10.1007/978-981-92-2960-4_6
Pourkarimi, M., & Moghaddam, A. H. (2026) · Scientia Horticulturae · doi:10.1016/j.scienta.2026.115075
ZHANG, Y., Yang, Y., Chen, Z., Wu, L., Peng, H., Wang, Q., Gao, J., Wang, J., Liao, Y., Tan, X., et al. (2026) · Research Square · doi:10.21203/rs.3.rs-10610097/v1
Off-season seed multiplication in spring barley: a scale-up step for speed breeding
Marenkova, A. G., Николаевич, И. В., Kocheshkova, A. A., Yu, K. P., & Divashuk, M. G. (2026) · Frontiers in Plant Science · doi:10.3389/fpls.2026.1912336
“A low red-to-far-red ratio suppresses tiller appearance in barley (Skinner and Simmons, 1993; Davis and Simmons, 1994) as part of the shade-avoidance syndrome (Casal, 2013); the lower far-red content of the LED spectrum may therefore have contributed to its higher tillering.”
Samarina, L. S., Turbekova, A., Jantassov, S. K., Demir, H., Kozhakhmetova, F., Begalina, A., Akzhunis, R., & Aisakulova, K. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27167408
Photoreceptors as central hubs of developmental plasticity and environmental adaptation
Hwang, H., Han, S., & Cho, H. (2026) · Molecules and Cells · doi:10.1016/j.mocell.2026.100390
Bian, Y., Zhao, F., Song, Z., Zhang, L., & Xu, D. (2026) · PLANT PHYSIOLOGY · doi:10.1093/plphys/kiag636
Kim, K. Y., Kang, J., Jeon, W.-T., Lee, J., Woo, J., Je, S., Yamaoka, Y., & Lee, Y. (2026) · Journal of Experimental Botany · doi:10.1093/jxb/erag360
Salinas, D., Lindström, M., Ibáñez, C., Eriksson, M. E., Bascuñán‐Godoy, L., Aronsson, H., González‐Teuber, M., Ojeda-May, P., Domínguez, P. G., & Vergara, A. (2026) · SSRN Electronic Journal · doi:10.2139/ssrn.7386949
Haile, A., Degu, H. D., Roro, A. G., & Tesfaye, B. (2026) · Plant Physiology and Biochemistry · doi:10.1016/j.plaphy.2026.111718
Zakaria, T. N. A. A. T., Teoh, S. C., Chear, N. J., Subramanıam, S., Murugaiyah, V., Hassan, Z., & Chew, B. L. (2026) · Industrial Crops and Products · doi:10.1016/j.indcrop.2026.124344
Beyene, A., & Ali, U. M. (2026) · Scientific African · doi:10.1016/j.sciaf.2026.e03643
Roh, Y. S., & Yoo, Y. K. (2026) · Horticultural Science and Technology · doi:10.7235/hort.20260032
da-Silva, C. J., Collado, C. E., van Butsele, J., & Hernández, R. (2026) · Annals of Applied Biology · doi:10.1111/aab.70158
Patandean, B., Leki, M. P. K. B., Pa, B. Y., Ndandarmana, N., Abadi, E., Christian, R., Masela, E. S., Navaro, A., Yusuf, A. R., Purba, M. M., et al. (2026) · F1000Research · doi:10.12688/f1000research.189197.1
Feng, X., An, S., Guo, W., Song, Y., Wang, G., & Li, L. (2026) · Horticulturae · doi:10.3390/horticulturae12091169
Photomorphogenesis: A molecular brake on PIF4
Mitra, O., & Huq, E. (2026) · Current Biology · doi:10.1016/j.cub.2026.07.068
Chen, R., Feng, J., Xi, S., Ji, Y., Fan, S., Wang, J., Kong, W., Guo, H., Chen, J., & Li, D. (2026) · Plant Physiology and Biochemistry · doi:10.1016/j.plaphy.2026.111760
Yang, H., Zhang, T., Liu, Y., Shan, Y., Yang, L., Liu, Y., Yang, Z., Li, F., Peng, J., & Wang, Z. (2026) · Planta · doi:10.1007/s00425-026-05179-5
“Early physiological studies indicated that plants exhibit differential sensitivity to spectral composition, and that photoperiodic responses differ among plant species, suggesting potential evolutionary divergence in photoreceptor systems among plant lineages (Casal 2013; Song et al. 2013).”
Xu, J., Wang, J., Long, M., Lang, C., Wang, Y., Zhang, J., & Xu, K. (2026) · Scientia Horticulturae · doi:10.1016/j.scienta.2026.115182
Phytochrome Signaling in Rice Immunity: Mechanisms, Trade-Offs, and Translational Opportunities
Rahman, M. M., Chen, H., Kwon, Y.-S., Rahman, M. M., Afrin, W., Choi, S., Yuanhu, X., & Jeon, J. (2026) · Rice Science · doi:10.1016/j.rsci.2026.09.010
Mooney, B. (2026) · The Plant Journal · doi:10.1111/tpj.71158
“These responses enable plants to grow beyond their neighbour’s shadow and regain access to light (Casal, 2013).”
Thangaraj, A., Kaul, R., Verma, R., Idayaraja, E., Ibrahim, S. M., Sharda, S., & Kaul, T. (2026) · Plant Physiology Reports · doi:10.1007/s40502-026-01003-0
Plant biointelligence: A genetic, biochemical, molecular, and cellular perspective
Lin, Y. (2026) · Current Plant Biology · doi:10.1016/j.cpb.2026.100686
한민희, Kim, J., Bang, J., 이재한, Kim, J. H., & Kim, D. (2026) · Horticulture Environment and Biotechnology · doi:10.1007/s13580-026-00860-w
Li, R., Cui, Y., Naser, M., Wu, Z., Xu, Y., Chen, X., Zhu, Y., Song, W., Xu, C., & 吴存祥 (2026) · Field Crops Research · doi:10.1016/j.fcr.2026.110769
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
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review