Environmental and Experimental Botany

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EEB 2954 No. of Pages 17
Environmental and Experimental Botany xxx (2015) xxx–xxx
Contents lists available at ScienceDirect
Environmental and Experimental Botany

journal homepage: www.elsevier.com/locate/envexpbot
Light signaling and plant responses to blue and UV radiations—

Perspectives for applications in horticulture
Lydie Huché-Théliera,b,* , Laurent Crespelb,c,1, José Le Gourrierecb,d,1, Philippe Morela,b,1,
Soulaiman Sakrb,c , Nathalie Leducb,d
a
INRA (Institut National de la Recherche Agronomique), Unité Mixte de Recherche 1345 IRHS (Institute of Research on Horticulture and Seeds),
Beaucouzé F-49070, France
b
SFR (Federal Research Structure) 4207 Qualité de la Santé des Plantes (Plant Quality and Health), Angers F-49000, France
c
Agrocampus-Ouest, Unité Mixte de Recherche 1345 IRHS (Institute of Research on Horticulture and Seeds), Angers F-49000, France
d
University of Angers, L'Université Nantes Angers Le Mans, Unité Mixte de Recherche 1345 IRHS (Institute of Research on Horticulture and Seeds), Angers F-
49000, France
A R T I C L E I N F O A B S T R A C T
Article history: Ultra-violet (UV) and blue radiations are perceived by plants through several photoreceptors. They
Received 18 February 2015 regulate a large range of processes throughout plant life. Along with red radiations, they are involved in
Received in revised form 8 June 2015 diverse photomorphogenic responses, e.g., seedling development, branching or flowering. In this paper,
Accepted 9 June 2015
we present an overview of UV- and blue-radiations signaling pathways in some key physiological
Available online xxx
processes and describe effects of plant exposure to these wavelengths on phenotype as well as on
contents in useful metabolites and resistance to bio aggressors. Taking these knowledge into account, we
Keywords:
finally discuss possible applications of the use of such radiations to improve plant production in
Biotic resistance
Flowering
horticulture.
Growth ã 2015 Elsevier B.V. All rights reserved.
Photomorphogenesis
Photoreceptors
Secondary metabolites
Introduction resistance to pests and diseases, product quality), through (1)

better management of plant density or architecture for light to
Light is one of the most important environmental factors that better penetrate into the crop or into the tree canopy, and (2)
regulates plant growth and development (Smith, 1982). Plants modification of the light environment under greenhouse con-
need light not only for photosynthesis but also for fine-tuned ditions. The need for artificial and efficient growing systems is
regulation of their development. UV, blue and red radiations are indeed essential to feed a constantly expanding human population
involved in diverse photomorphogenic responses such as seedling (Darko et al., 2014). Furthermore, as far as environmental and
development, vegetative growth, flowering and branching. A large health issues are concerned, growers are incited to find cultural
amount of available literature is on the perception and the plant alternatives to the use of pesticides and growth retardants on
response to red (R), far-red (FR) and R/FR ratio, mediated by crops. The use of sustainable techniques such as modulation of
phytochromes (Demotes-Mainard et al., 2015). UV and blue incoming light is one possibility.
radiations are also highly involved in the photomorphogenetic Even if morphological responses to blue radiations are known
effects. Understanding the mechanisms involved in these to be species-dependent, increasing the part of blue rays in the
responses can contribute to improving crop production (yield, light often reduces plant height and could be used as an alternative
to the growth retardants. Moreover, high amount of blue light
triggers specific biochemical and physiological processes resulting
* Corresponding author. in a higher accumulation of epidermal flavonols that allows a
E-mail addresses: lydie.thelier@angers.inra.fr (L. Huché-Thélier), better acclimation of pepper to UV stress (Hoffmann et al., 2015).
laurent.crespel@agrocampus-ouest.fr (L. Crespel), Indeed, exposure to UV-rich radiations leads to generation of free
jose.gentilhomme@univ-angers.fr (J.L. Gourrierec), philippe.morel@angers.inra.fr
radicals that damage DNA, proteins, lipids, chloroplasts and
(P. Morel), soulaiman.sakr@agrocampus-ouest.fr (S. Sakr),
nathalie.leduc@univ-angers.fr (N. Leduc). photosynthetic pigments (Hideg et al., 2013), but applied at lower
1
These authors contributed equally to this work. level and during specific and short periods in plant lighting, UV
http://dx.doi.org/10.1016/j.envexpbot.2015.06.009
0098-8472/ ã 2015 Elsevier B.V. All rights reserved.
Please cite this article in press as: L. Huché-Thélier, et al., Light signaling and plant responses to blue and UV radiations—Perspectives for
applications in horticulture, Environ. Exp. Bot. (2015), http://dx.doi.org/10.1016/j.envexpbot.2015.06.009
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2 L. Huché-Thélier et al. / Environmental and Experimental Botany xxx (2015) xxx–xxx
radiations enhance plant resistance to pathogens and herbivores 2.1. UVR8:

(Ballaré, 2014). The numerous studies on the consequences on
plants of the depletion of the ozone layer, which results in local UV-B radiation is perceived by UV RESISTANCE LOCUS 8
elevations in UV-B radiations (Caldwell et al., 2007; Kakani et al., (UVR8) (Fig. 1), a symmetrical homodimer made up of seven-
2003), provide interesting information for further applications to bladed b-propeller protein homodimers bearing tryptophans that
horticultural crops. may act as a chromophore (Wu et al., 2012; Heijde and Ulm,
Moreover, greenhouses were commonly equipped with high 2012). UV-B irradiation disrupts hydrogen bonds and thus
pressure sodium (HPS) lamps with a spectrum of lighting dissociates the homodimer into two monomers.
containing small amounts of blue and high amounts of red and UVR8 monomers then migrate from the cytosol into the nucleus
green lights. Now, these structures are gradually being equipped where they interact with a molecular partner involved in the UV-
with new lighting technologies such as photo-selective films or B signal transduction pathway. Only few of these molecular
light emitting diodes (LEDs). LEDs can cover the fluence and players have been identified to date (Wu et al., 2012) and acts
wavelength requirements of plants for different growth phases upstream of CONSTITUTIVELY PHOTOMORPHOGENIC1 (COP1, an
(Darko et al., 2014), and will contribute to quantitative and E3-ubiquitin ligase), and LONG HYPOCOTYL5 (HY5), two well-
qualitative increases in crop yields. known central regulators of other light signaling pathways.
In this context, this review focuses on plant responses to UV and UVR8 associates with COP1 to activate UV-B responsive genes
blue lights, with an overview of the photoreceptors involved in the (Wu et al., 2012; Rizzini et al., 2011).
perception and signaling pathways of these wavelengths in main
key physiological processes. The consequences on the phenotype 2.2. Phototropins
of plants exposed to these wavelengths are described and this
analysis leads to a discussion on the potential use of UV and blue Phototropins (phots) are photoreceptors that perceive UV-A
radiations in the improvement of plant production in horticulture. and blue lights (Fig. 1). They are serine/threonine protein kinases
that bear flavin mononucleotide (FMN) chromophores bound to
1. The light spectrum controls plant growth and development the protein at two light-oxygen-voltage (LOV) domains (Christie,
2007; Demarsy and Fankhauser, 2009). Two phototropins are
The sensitivity range of plants to light extends from UV through present in Arabidopsis: phot1 and phot2. Phot1 responds to low-
the visible spectrum to far-red radiation (700–800 nm). Regarding fluence blue light (<1 mmol m2 s1), while phot2 mediates the
ultraviolet radiation, UV-C (100–280 nm) radiation is thoroughly high-fluence response together with phot1. Blue light perception
absorbed by the stratospheric ozone layer and the atmosphere; triggers conformational changes of these photoreceptors, leading
only UV-A (315–400 nm) and UV-B (280–315 nm) radiations reach to activation of the kinase domain and then autophosphorylation
plants (Fig. 1). In the visible light spectrum (400–700 nm), the of the serine residues (Briggs and Christie, 2002; Inoue et al.,
major wavelengths perceived by plant photoreceptors and pig- 2008a). This triggers phot signaling, involving NONPHOTOTROPIC
ments are those corresponding to blue (400–500 nm) and red HYPOCOTYL 3 (NPH3) and ROOT PHOTOTROPISM2 (RPT2), two
(600–700 nm) and, to a lesser extent, green (500–600 nm) (Fig. 1). partner proteins that bind to PHOT1 (Inada et al., 2004;
A small fraction of near-infrared radiation, i.e., far-red light Motchoulski and Liscum, 1999). Phots localize at the plasma
perceived by phytochromes with a sensitivity peak at 730 nm, is membrane and dissociate from it once activated by blue and UV-A
also essential to plant development. photons. PHOT expression is regulated by blue light, with blue light
stimulating PHOT2 expression and downregulating PHOT1 expres-
2. UV and blue radiation photoreceptors sion. Mutant studies in Arabidopsis reveal that CRYPTOCHROME 1
(CRY1) and PHYTOCHROME B (PHYB) regulate PHOT1 expression,
UV and blue radiations are perceived by plants through a range while CRY1, CRYPTOCHROME 2 (CRY2) and PHYTOCHROME A
of photoreceptors: (PHYA) control PHOT2 expression (Łabuz et al., 2012). Phots
contribute to several developmental responses to light, and
phot1 and phot2 have most often overlapping functions. In
Arabidopsis, both are responsible for phototropism, including
hypocotyl and stem bending as well as leaf positioning (Esmon
Fig. 1. The active light spectrum in plants ranges from ultra-violet (UV) to infrared, and plant photoreceptors absorb specific wavelength regions (bordered by dotted lines):
UVR8, UV RESISTANCE LOCUS 8; PHOT, phototropins; CRY, cryptochromes; ZTL/FKF1/LKP2, Zeitlupe/Flavin-binding Kelch/LOV Kelch Protein; PHY, phytochromes.
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et al., 2006; Inoue et al., 2008b), chloroplast accumulation and PIF5 that take part in signaling of the R/FR ratio in SAS response,
stomatal opening (Ma et al., 2001). However, phot2 is the only phot may also be required for full expression of the hyponastic
involved in the chloroplast avoidance response (Jarillo et al., 2001; response of Arabidopsis leaves to blue light attenuation (Keller
Łabuz et al., 2012). et al., 2011), but so far the mechanisms of interaction between
CRY1 and PIF have not been elucidated.
2.3. Cryptochromes
2.4. ZTL/FKF1/LKP2
Cryptochromes (crys) can perceive UV-A and blue radiations
(Fig. 1) with two wavelengths optima (370 and 450 nm). Most Zeitlupe (ZTL), Flavin-binding Kelch (FKF1) and LOV Kelch
plants have several crys: in Arabidopsis thaliana, two crys, Proteins (LKP2/FKL1) form a family of UV-A and blue light photo-
AtCRY1 and AtCRY2, the latter being light-labile, were clearly receptors (Fig. 1) that bear flavin mononucleotide (FMN) as a
identified based on their C-terminal extensions (Ahmad and chromophore, as phototropins do. These proteins are composed of
Cashmore, 1993; Hoffman et al., 1996; Lin et al., 1996), while three domains: a LOV, an F-box on the N-terminal side and a Kelch
CRY3 function is not clearly established yet (Brudler et al., 2003; domain on the C-terminal end. The LOV domain links in a non-
Kleine et al., 2003). The CRY gene family varies among plants: some covalent manner to the FMN chromophore, while the F-box domain
bear two CRY1-like genes (tomato, rice, barley: Matsumoto et al., interacts with the ubiquitination complex E2-E3 that leads to protein
2003; Perrotta et al., 2000), while others bear two differentCRY2 degradation. The Kelch motif contains 6 repeated tandem sequences
genes (Pisum sativum L., Medicago: Platten et al., 2005a,b). Blue forming a b-helix. Blue light irradiation induces a conformational
light acts differently on CRY gene expression depending on species. change leading to photoreceptor autophosphorylation that triggers
For example, it strongly downregulates PsCRY2b expression in P. signaling (Imaizumi et al., 2003).
sativum L. (Platten et al., 2005b), while it enhances CRY1
expression in Brassica napus (Chatterjee et al., 2006). 2.5. Phytochromes
Subcellular localization of crys has been described in some
species. In Arabidopsis, AtCRY1 and AtCRY2 are both present in the Phytochromes (phys) are proteins that bear a phytochromobi-
nucleus (Cashmore et al., 1999; Guo et al., 1999), and blue light lin, a linear tetrapyrrole, as a chromophore. The chromophore is
triggers CRY1 transfer to the cytosol while CRY2 remains in the synthesized in the chloroplast, and migrates into the cytoplasm
nucleus (Yu et al., 2007). In wheat (Triticum aestivum), similar where it binds to the protein part of the phy. In Arabidopsis, 5 phys
translocation of TaCRY1a from nucleus to cytoplasm upon are present, named phyA, phyB, phyC, phyD, and phyE (Lau and
exposure to blue light has been observed, as well as degradation Deng, 2010; Leivar and Quail, 2011). They each exist under two
of TaCRY2 in the nucleus under blue light (Xu et al., 2009). In rice, forms: the Pr inactive form in which they are synthesized, and the
on the contrary, CRY1 has been found both in the cytosol and in the Pfr active form. In the nucleus, light-activated phys interact with
nucleus independently of light conditions and functions as blue PIFs and COP1 to regulate downstream photomorphogenic
light receptor both in the nucleus and the cytoplasm (Matsumoto molecular actors (Li et al., 2011).
et al., 2003). Photo-transformation between the Pr inactive and Pfr active
CRYs have structural homology with photolyases although they forms is efficiently achieved by red light (around 660 nm), but also
do not have such enzymatic activity, and unlike photolyases they by any other wavelength ranging from UV (300 nm) and blue to far-
display a C-terminal domain (Lin and Todo, 2005). They bear a flavin red (800 nm) even though at a far less efficient rate (Fig. 1,
adenine dinucleotide (FAD) and methenyltetrahydrofolate (MTHF) Shinomura et al., 1996). Under wavelengths other than red light,
as chromophores (Heijde and Ulm, 2012; Möglich et al., 2010). Upon low quantities of Pfr accumulate. Still, this allows plants to use phy
absorption of blue light energy by MTHFand transfer to oxidized FAD, signaling pathways in response to particular light conditions such
the reduced and active form FADH accumulates. Further reduction as very low irradiance, and therefore best adapt to their
into FADH may also occur under green light exposure and environment (Shinomura et al., 1996). PhyA is the only phy able
contribute to stop blue light signaling (Bouly et al., 2007), while to produce such a very low fluence response (VLFR) (Casal, 2013),
under darkness, reoxidation of FADH into FAD replenishes the pool due to its highest sensitivity (Shinomura et al., 1996). Some
of inactivated CRY (Möglich et al., 2010). Upon activation, responses to blue light through phyA have been identified: in
conformational changes of the CRY protein allow the effector Arabidopsis, repression of hypocotyl gravitropism in response to
domain of CRY, likely represented by the C-terminal extension, to very low irradiance (0.1–0.7 mmol m2 s1) blue light is under the
directly interact with effector proteins. Homodimerization of control of phyA (Lariguet and Fankhauser, 2004); it also irrevers-
CRY1 and CRY2 contributes to the regulation of their signaling. ibly triggers the photoinduction of seed germination upon
Up to now, two interacting partners of CRYs have been irradiation under extremely low-fluence UV-A radiation and blue
identified (Liu et al., 2011). The Cryptochrome-Interacting Basic- light (Shinomura et al., 1996). Some responses so far attributed to
helix-loop helix (CIB1) protein interacts with CRY2 under blue other blue light photoreceptors may in fact involve phyA or phyB
light and binds to the promoter of certain genes to activate their (Shinomura et al., 1996).
transcription (Liu et al., 2008). This interaction is notably involved
in floral initiation (see below). The second CRY-interacting 3. Impact of blue and UV radiations on plant vegetative growth
partner is the SUPRESSOR OF PHYTOCHROME A-105 1 (SPA1),
which is involved in post-transcriptional regulation of gene Plant vegetative growth is the result of key processes such as
expression under blue light. In this system, SPA1 is a positive cell division and elongation, directional growth and branching.
regulator of E3 ubiquitin ligase COP1 (Lian et al., 2011; Liu et al., These processes are dependent on light spectrum, intensity and
2011; Zuo et al., 2011). Interaction of SPA1 with CRY leads to direction that modulate photosynthetic activity as well as
suppression of COP1-dependent degradation of transcriptional photomorphogenic signals. In this review, we focus on perception
regulators such as bZIP transcription factors HY5, HY5HOMO- and signaling pathways of blue and UV radiations in some of these
LOGUE (HYH) and bHLH transcription factor Long Hypocotyl in processes before investigating the main consequences on plant
Far-Red1 (HFR1) involved in shade avoidance syndrome (SAS) morphology. We resume these physiological processes and
responses (Liu et al., 2011). The basic-Helix-Loop Helix (bHLH) developmental phases of a plant on two schemes (Fig. 2 for blue
transcription factors Phytochrome-Interacting Factors PIF4 and radiations and Fig. 3 for UV radiations).
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Fig. 2. Physiological processes (blue square) and developmental phases of a plant (grey square) that may be targeted to improve crop characters (deep Blue squares) after
manipulation of the blue light environment during culture. Plant photoreceptors absorbing blue radiations: PHOT, phototropins; CRY, cryptochromes; FKF1, Flavin-binding
Kelch, PHY, phytochromes. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
3.1. Cell elongation expression early (24 h) upon exposure of etiolated pea seedlings
to blue light, illustrating the complex interaction among photo-
Blue light is an essential signal that helps plants to adjust their receptors in the regulation of this process (Platten et al., 2005b).
growth to their environment at different key stages of their Blue light perception and signaling through these photo-
development, e.g., hypocotyl de-etiolation upon above-ground receptors is believed to lead (1) to increased rigidity of the
growth, increased vertical shoot elongation to avoid shade from hypocotyl caused by increased cell turgor pressure (Cosgrove and
neighboring plants, or directional growth (phototropism) to reach Green, 1981), and (2) to reduced cell wall extensibility and thus
light in gaps left by higher plant layers, or to grow away from too repression of hypocotyl etiolation upon aboveground light
high irradiance conditions. In all these responses, blue light exposure, as shown in cucumber (Cosgrove, 1981). In Arabidopsis,
controls cell elongation. blue light acts through CRY1 and represses gibberellin (GA) and
Hypocotyl de-etiolation under blue light is under the control of auxin synthesis and/or sensitivity, in turn affecting the expression
phot, cry and phy photoreceptors. PHOT1 mediates a very early 30 s of several genes involved in hypocotyl growth repression (Folta
after blue light exposure—initial growth inhibition response in et al., 2003). Conversely, when blue light intensity drops as in SAS
hypocotyls (Folta and Spalding, 2001; Parks et al., 2001), then further conditions, hypocotyl cell wall extensibility is increased due to
inhibition is taken over by CRYs and PHYs. In A. thaliana, CRY2 (Lin stimulation of the transcriptional activity of cell-expansion-related
et al., 1998) and CRY1 (Ahmad and Cashmore, 1993) contribute to genes such as expansin genes in Stellaria (Sasidharan et al., 2010)
de-etiolation, together with PHYA (Ahmad and Cashmore, 1997; and of two groups of XYLOGLUCAN ENDOTRANSGLUCOSYLASE/
Neff and Chory,1998). In this process, PHYB is thought to antagonize HYDROLASE (XTH) genes in Arabidopsis (Keuskamp et al., 2011).
CRY1 and PHYA (Folta and Spalding, 2001). Photoreceptors respond Further investigations revealed that auxin and brassinosteroids
to different light levels for a fine tuning of the de-etiolation process: both controlled a subset of these XTH genes in hypocotyl de-
in pea, hypocotyl de-etiolation is triggered by PHYA, PHYB and etiolation in a non-redundant and non-synergistic manner
CRY1 perception of high-irradiance (from 4 to 20 mmol m2 s1) (Keuskamp et al., 2011). Interestingly, blue light depletion and
blue light, while only PHYA is involved in the response to low- CRY1 perception triggered only brassinosteroid signaling in two
irradiance (from 0.2 to 2 mmol m2 s1) blue light (Platten et al., other SAS-related responses in Arabidopsis seedlings: leaf
2005a,b). Interestingly, PHYA and CRY1 can repress CRY2b hyponasty and petiole elongation (Keller et al., 2011).
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Fig. 3. Physiological processes (orange square) and developmental phases of a plant (grey square) that may be targeted to improve crop characters (deep orange squares) after
manipulation of the UV light environment during culture. Plant photoreceptors absorbing UV radiations: PHOT, phototropins; CRY, cryptochromes; PHY, phytochromes;
UVR8, UV RESISTANCE LOCUS 8. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
In the stem of older plants, photoreceptors have an opposite The downstream sequence of events following blue light
effect on elongation upon blue light perception. In pea, PHYA is a perception by phot1 is not clearly established. Several mechanisms
promoter of internode growth, while CRY1 is a repressor (Platten seem plausible. According to Christie et al. (2011), phot1 physically
et al., 2005a; Weller et al., 2001). CRY1 is also a repressor in tomato interacts with the auxin efflux carrier P-GLYCOPROTEIN 19
grown at low density (Casal, 2013). According to Foo et al. (2006), (PGP19), possibly causing its phosphorylation as observed in vitro,
blue light perception by CRY1 and PHYA in pea stems exposed to and leading to a change in auxin fluxes. In an alternative
seven days of darkness represses stem cell elongation, and thus mechanism (Huang et al., 2010; Zhang et al., 2010),
etiolation, by redundantly reducing levels of bioactive gibberellins phot1 controls the orientation of auxin fluxes through the control
GA1 through repression of PsGA3ox1 and stimulation of PsGA-2ox of PINOID expression. This gene encodes a PID kinase that
transcription, and regulation of GA sensitivity. phosphorylates the auxin efflux carrier PIN. Since the
phosphorylation state of PIN drives its cellular localization,
3.2. Directional growth phot1-controlled PID transcript accumulation may influence PIN
distribution within the cell and thus orientation of the auxin flux.
In order to optimize photosynthetic activity, plant direct their Indirect control of phots on auxin efflux carriers may take place
growth towards incoming light to best position their leaves. In this through control of the Ca2+ influx (Stoelzle et al., 2003; Zhao et al.,
reaction, direction of blue and UV-A radiations in the incoming 2013), as PID is also regulated by intracellular Ca2+ (Benjamins
light constitutes the signal perceived by plants. In the case of et al., 2003). Finally, recent work by Sun et al. (2013) brought
hypocotyl phototropism, the signal will lead to asymmetrical cell evidence for another mechanism involving PIF4 and PIF5 as
growth upon formation of a lateral gradient of auxin distribution, negative regulators of hypocotyl phototropism. In their study,
and auxin transfer from the lit side to the shaded side of the organ directional blue light increases expression of these two PIF genes
where it accumulates (Ding et al., 2011; Friml et al., 2002). This by bending hypocotyls of 3-day-old seedlings, and promotes
ultimately causes phototropic growth of the shoot towards the accumulation of PIF proteins to the hypocotyl apex where
light source through auxin-mediated increased cell growth on the phototropic perception takes place. Using photoreceptor mutants,
shaded side, as recently reviewed by Christie and Murphy (2013) they established that the blue light photoreceptors involved in this
and by Hohm et al. (2013). In this case, phots are the main response were PHYA acting as a positive regulator of PIF4 and
photoreceptors involved in perception of directional light, with PIF5 expression, and PHOT1 and PHOT2 acting as repressors. In the
CRY and PHYA contributing too. PHOT1 acts as the main photo- proposed model, blue light perception by PHOT1 and
tropin in this response. PHOT2 represses PIF4 and PIF5 transcription, and this reduces
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cell auxin response on the lit side of the organ. PIF4 is thought to crys might regulate stomatal opening by regulating MYB60
act through direct binding to the G-boxes of the promoters of the expression (Wang et al., 2010a). Downstream of crys and phots,
IAA19 and IAA29 auxin response repressor genes acting as negative COP1 regulates the stomatal response to blue light (Mao et al., 2005;
regulators of the auxin-responsive transcriptional activator ARF7. Wang et al., 2010a), but how it participates in this regulation
remains unknown (Chen et al., 2012). Phytochromes enhance cry
3.3. Photosynthesis and phot activity (Casal, 2000). In Arabidopsis, stomata of the triple
mutants cry1 cry2 phyB, phot1 phot2 phyB or cry1 phyA phyB open
The effects of blue and UV-B radiations on photosynthesis are less than those of the single or double mutants under white light.
the consequences of the specific absorption spectrum of photo- This implies that phyB acts in concert with phyA, cry, and phot in
synthetic pigments (chlorophylls, carotenoids) or of higher or light regulation of stomatal opening (Wang et al., 2010a). PhyB-
lower absorption of micronutrients essential to the photosynthetic mediated stomatal opening responds not only to red light but also to
electron transport chain (Kopsell et al., 2014; O’Carrigan et al., blue light, probably due to a crosstalk between the different
2014). Chlorophylls a and b absorb strongly in the red (maximum photoreceptors at a common intermediate signaling step (Wang
absorption at 663 and 642 nm, respectively) and blue (maximum et al., 2010a). Even if transduction of the light signal from the
absorption at 430 and 453 nm, respectively) regions (Wright and different photoreceptors is not completely deciphered, several
Shearer, 1984). The carotenoid pigments lutein and b-carotene transcription factors have been identified as regulators of stomatal
absorb strongly in the blue region, with maximum absorption at opening in response to blue light: (1) COP1, (2) MYB proteins
448 and 452 nm, respectively (Wright and Shearer, 1984). Plants (AtMYB60 as a positive regulator (Wang et al., 2010a), as mentioned
tend to adapt their content of chlorophyll pigments to the light above, or AtMYB61 as a negative regulator (Liang et al., 2005), or (3)
spectrum. Under blue light alone compared to white light, total EARLY FLOWERING 3 (EFL3) and FLOWERING LOCUS T (FT). EFL3 and
chlorophyll (chl) content decreases in some species (Wang et al., FT may be implied in the photoperiodic regulation of stomatal
2014) but is not affected in others (Abidi et al., 2013). Nevertheless, sensitivity to blue light (Kinoshita et al., 2011).
when blue light is present with other wavelengths, chl content Blue wavelengths also act on chloroplast relocation within the
tends to increase with the proportion of blue light in plant lighting cell. Under low fluence rates, chloroplasts accumulate at the cell
(Hernández and Kubota, 2014; Hogewoning et al., 2010; Matsuda surface to increase their photosynthetic ability, while under higher
et al., 2007; Yorio et al., 2001). Independently of species, both blue fluence rates, they move to the opposite side to light
monochromatic blue light and higher proportion of blue light in (avoidance response) to avoid damage (Kasahara et al., 2002).
plant lighting tend to increase the chl a/b ratio (Abidi et al., 2013; These movements are mediated by phots (Kong et al., 2013a). In
Hogewoning et al., 2010; Johkan et al., 2010; Kopsell and Sams, Arabidopsis, both phot1 and phot2 regulate chloroplast accumula-
2013; Matsuda et al., 2007; Ohashi-Kaneko et al., 2007; Wang et al., tion at the cell surface through long-distance signaling (Kong and
2014). When plants are exposed to UV-B radiations, chl content Wada, 2011; Kong et al., 2013b). The avoidance response is mainly
decreases (Nedunchezhian and Kulandaivelu, 1997; Ziska et al., regulated by phot2 (Kong et al., 2013b), though phot1 is also
1993) due to high decrease in chlorophyll a content and, involved, but in a minor capacity (Luesse et al., 2010). Chloroplast
consequently, in chl a/b ratio (Lidon and Ramalho, 2011). movement is mediated through the rapid reorganization of
Photosynthetic activity increases proportionally with the level chloroplast actin (cp-actin) filaments at the interface between
of blue light in plant lighting (Hernández and Kubota, 2014; the plasma membrane and chloroplast outer membrane (Kadota
Hogewoning et al., 2010; Matsuda et al., 2007; Yorio et al., 2001), et al., 2009). The rate of cp-actin filament disappearance depends
but only when blue light is present together with other wave- on blue light intensity (Kadota et al., 2009) and is solely regulated
lengths. Photosynthetic activity is reduced under blue light alone by phot2 (Kong and Wada, 2011).
compared to white light (Abidi et al., 2013; Hogewoning et al., The responses of stomata to UV-B radiations are less constant
2010; O’Carrigan et al., 2014), or when blue wavelengths are than those to blue light. UV-B either stimulate stomatal opening or
missing in the light spectrum (Goins et al., 1997; Hogewoning et al., stomatal closing and neither of these responses is readily reversible
2010; Matsuda et al., 2007; Yorio et al., 2001). Likewise, exposure (Jansen and van den Noort, 2000). These responses depend on the
to supplemental UV-B radiation decreases plant photosynthesis metabolic state of guard cells (Jansen and van den Noort, 2000) but so
efficiency by 3–90% according to growth conditions (see Lidon far, the implicated mechanisms have not been elucidated.
et al., 2012; for a review). Long term exposure to blue light modifies leaf and chloroplast
Blue and UV-B radiations act on photosynthetic activity both at characteristics. Leaf thickness, stomata number and length of
short- and long-term scales. On the short term, blue light has an palisade tissue cells are increased under blue light as compared to
effect on photosynthesis by inducing stomatal opening (Lawson, plants grown under red or green light (O’Carrigan et al., 2014). The
2009; Zeiger et al., 2002). Both blue and red lights are able to promote epidermal cell area of birch leaves was larger and the functional
stomatal opening, and three families of blue light photoreceptors viz. area of chloroplasts (starch-free part of the chloroplast) is greater
phots, crys, and phys regulate this response (Kinoshita et al., 2001; in plantlets grown under blue light than in plantlets grown under
Mao et al., 2005). In Arabidopsis, autophosphorylation of phot1 and white or red light (Sæbø et al., 1995). In cucumber grown under low
phot2, which are blue-light induced, activates signal transduction, radiations, chloroplasts under blue light have a higher number of
while their dephosphorylation halts the signaling process (Inoue grana lamellae and more stacked thylakoid membranes than under
et al., 2008a). Phosphorylation of phot1 and phot2 activates a white or red light (Wang et al., 2014). Moreover, the lower
plasma membrane H+-ATPase and leads to proton extrusion, accumulation of starch grains in blue-light-grown leaves reduces
membrane hyperpolarization and K+ influx into stomatal guard the risk for thylakoids to be pressed close together, which can
cells (Assmann and Shimazaki, 1999; Mott, 2009; Takemiya et al., physically prevent light from reaching them (Bondada and
2005). As a result, guard cells swell and bend, thereby pushing the Oosterhuis, 2003).
stomatal pore open and enabling gas exchanges between the leaf
inside and the atmosphere (Roelfsema and Hedrich, 2005). MYB60, 3.4. Consequences of UV and blue light perception on plant vegetative
an R2R3-MYB gene of Arabidopsis, is the first transcription factor growth
ever identified in the regulation of stomatal movements (Cominelli
et al., 2005). Its expression is reduced in cry1 cry2 mutants under Numerous studies mention morphological effects of UV and
blue light, but increased in CRY1-overexpressing plants. Therefore blue radiations on plants, which often depend on species and
G Model
Table 1
Effects of blue light on vegetative and reproductive development, pathogen – pest resistance and production of metabolites in crops. ": significant increase; #: significant
decrease; =: no significant effect. PPFD: photosynthetic flux density.
Process Variables Blue Quantification Species/disease Light conditions References

light
effect
PPFD Blue Growth conditions
(mmol m2 s1) wavelengths
(nm)
Growth Shoot length " +28% Cucumber From 50 to 60 NAa Fluorescent lamps:blue vs. daylight Piszczek and
Głowacka
(2008)
# 68% Potato 100 NA Blue fluorescent lamps vs. low-pressure Wilson et al.,
sodium (LPS) lamps. 4 cultivars, in vitro (1993)
30% Tomato 67 NA Fluorescent lamps:blue vs. daylight. Glowacka,
3 cultivars (2004)
= Rose bush 110 From 400 to fluorescent lamps : blue vs. white, Abidi et al.,
500 max:450 2 cultivars (2013)
Internode " +29% Cucumber From 50 to 60 NA Fluorescent lamps : blue vs. daylight Piszczek and
length Głowacka,
(2008)
# 27% Tomato 110 NA Fluorescent lamps : blue vs. daylight. Glowacka,
3 cultivars (2004)
= Rose bush 110 From 400 to Fluorescent lamps : blue vs. white, Abidi et al.,
500 2 cultivars (2013)
max:450
Leaf area " +26% Pepper 300 From 400 to Red LEDs with supplemental blue Brown et al.,
500 max:440 (fluorescent lamps) vs. red LEDs alone. (1995)
# 9% Cucumber 250 Max:455 High solar irradiance, supplemental red Hernández
FWHMa :15 LEDS with or without supplemental blue and Kubota,
LEDs (2014)
38% Lettuce 300 From 400 to Fluorescent lamps : blue vs. white Ohashi-
500 max: Kaneko et al.,
62% Spinach
450 (2007)
= Komatsuna
Rose Bush 110 From 400 to Fluorescent lamps : blue vs. white. Abidi et al.,
max: 450
Leaf mass area " Maximum Cucumber 100 450 Different light qualities with red and Hogewoning
with 50% blue blue LEDs. Proportions of PAR provided et al., (2010)
by blue light:0, 15, 30, 50 or 100%
+20% Rose bush 110 From 400 to Fluorescent lamps:blue vs. white. Abidi et al.,
500 cultivar ‘Radrazz” (2013)
max: 450
=
Stem " +20% Cucumber From 50 to 60 NA Fluorescent lamps:blue vs. daylight Piszczek and
thickness Glowacka
(2008)
+8% Tomato 67 NA Fluorescent lamps:blue vs. daylight. Glowacka,
3 cultivars (2004)
Shoot dry " +15% 200 max:450 Blue (LEDs) + red + far-red vs. HPS lamps Hogewoning
mass et al., (2012)
+21% Cucumber From 50 to 60 NA Fluorescent lamps : blue vs. daylight Piszczek and
Głowacka,
+10%
(2008)
# 24% 250 Max:455 High solar irradiance, supplemental red Hernández
FWHMa :15 LEDS with or without supplemental blue and Kubota,
LEDs (2014)
39% Lettuce 300 From 400 to Fluorescent lamps : blue vs. white Ohashi-
500 max: Kaneko et al.,
58% Spinach
450 (2007)
= Komatsuna
Potato 100 NA Blue fluorescent lamps vs. LPS lamps. Wilson et al.,
4 cultivars (1993)
Tomato 67 NA Fluorescent lamps : blue vs. daylight). 3 Glowacka,
cultivars (2004)
Blue (LED) s + red vs. HPS lamps Hogewoning
et al., (2012)
G Model
Table 1 (Continued)
light
effect
(nm)
Cucumber
Branching Stem or tiller " +22% Chrysanthemum 93–146–237 NA Blue vs. white light. three levels of PAR Mortensen
number and
Strømme,
(1987)
+58% Plum 40 Blue + red vs. red light Muleo et al.,
(2001)
+31% Tomato 93–146–237 NA Blue vs. white light. three levels of PAR Mortensen
and
Strømme,
(1987)
+25% Wheat 200 NA High (25%) vs. low (1%) part of blue light Barnes and
provided by metal halide lamps Bugbee,
(1992)
= Rose bush 110 From 400 to Fluorescent lamps : blue vs. white. Abidi et al.,
max: 450
Bud burst rate " +93% Potato 100 NA Blue fluorescent lamps vs. LPS lamps. Wilson et al.,
4 cultivars, in vitro (1993)
Dark: 0% Rose bush 200 NA Fluorescent lamps:blue light vs. Girault et al.,
White: 100% darkness (2008)
= Blue:100% Fluorescent lamps: blue vs. white light
Flowering Flowering = 110 From 400 to Fluorescent lamps:blue vs. white light. Abidi et al.,
rate 500 2 cultivars (2013)
max: 450
Delay before " +5.5 days Chrysanthemum 3.05 MJ m2 day1 NA Increasing% of blue in PAR from 30 to Khattak and
flowering 40% Pearson,
(2006)
# 9.1 days Increasing% of blue in PAR from 14 to
30%
Flower " +7%
diameter
= Increasing% of blue in PAR from 30 to
40%
Rose bush 110 From 400 to Fluorescent lamps:blue vs. white. Abidi et al.,
max: 450
Pathogen % Wilted # -21% Pepper/tomato 300 From 400 to Red LEDs + blue fluorescent lamps vs. red Schuerger
resistance plants mosaic virus 500 LEDs and Brown,
(1997)
Colony " +31% Cucumber/ 350 Max:452.5 Incandescent reflector lamps:blue vs. Wang et al.,
number/leaf Sphaerotheca white light (2010b)
fuliginea
Concentration b-carotene " +92% Broccoli 250 470 LEDs (20% blue – 80 % red) vs. white light Kopsell et al.,
in (fluorescent lamps 5% blue) (2014)
metabolites +8% Kale 275 440 Specific LED wavelengths blue vs. red Lefsrud et al.,
(660). (2008)
+8% Lettuce 110 Blue Fluorescent lamps: white light added by Li and
LEDs:476 supplemental blue LEDs vs. white light Kubota,
only (2009)
Total " +32% Spinach 300 From 400 to Fluorescent lamps : blue vs. white Ohashi-
carotenoids 500 Kaneko et al.,
max: 450 (2007)
Anthocyanins " +36% Lettuce 110 Blue White light (Fluorescent lamps) + Li and
LEDs:476 supplemental blue LEDs vs. white light Kubota,
only (2009)
Aliphatic " +205% Broccoli 250 Max:470 LEDs (20% blue 80 % red) vs. white light Kopsell et al.,
glucosinolates (fluorescent lamps 5% blue) (2014)
Chlorogenic " +662% Basil 300 From 410 to HPS + supplemental blue LEDs vs. HPS Taulavuori
acid 460. Max: alone et al., (2013)
450
+96% Tomato
Nitrate # 35% Lettuce 300 Fluorescent lamps:blue vs. white
G Model
Table 1 (Continued)
light
effect
(nm)
From 400 to Ohashi-
500 Kaneko et al.,
max:450 (2007)
= Spinach 300 From 400 to Fluorescent lamps:blue vs. white Ohashi-
500 Kaneko et al.,
max:450 (2007)
Komatsuna
a
NA: not available information, FWHM: full width at half maximum.
growth conditions (light intensity, relative shares of these wave- and internode lengths, shoot dry matter, as well as leaf area (Abidi
lengths in the light spectrum). Even within a single species, plant et al., 2013; Brown et al., 1995; Glowacka, 2004; Hernández and
responses to blue light can differ among genotypes, as shown in Kubota, 2014; Ohashi-Kaneko et al., 2007; Piszczek and Głowacka,
tomato (Glowacka, 2006). Blue light can increase or decrease shoot 2008; Wilson et al., 1993) (Table 1). In some species, it has no
Table 2
Effects of UV-radiations on vegetative and reproductive developments, pathogen - pest resistance and production of metabolites in crops. ": significant increase; #: significant
decrease; =: no significant effect.
Process Variables UV Quantification Species/ Light conditions References

effect disease
UV-irradiance UV Growth conditions
(kJ m2 d1) wavelengths
(nm)
Growth Shoot or " +22% Cucumber NA NA Solar UV-A without UV-B radiations vs. no solar Krizek et al.,
stem length UV radiations (UV exclusion chambers) (1997)
# 15% Soya 13 NA UV-B Supplementation (7 h.day1, except rainy Liu et al.,
days) simulating a 21% depletion of the ozone (2013)
layer vs. no UV-B radiation
31% NA From 280 to Natural lighting vs. glasshouse with complete Zhang et al.,
320 UV-B absorption (2014)
20% Rose bush UV-A:12.0 vs. UV-A:315- High altitude (2800 m) natural light vs. crop Terfa et al.,
1.7 400 under plastic film absorbing 96% of UV (2014)
UV- UV-B: 280- radiation
B:0.88 vs.0.036 315
(V m2 s1)
= Poinsettia 0.36 or 0.72 NA Irradiation (1h) with UV-B at the middle of the Torre et al.,
night period (2012)
Internode # 9%
length
Shoot or # 17% Soya 13 NA UV-B supplementation (7h day1, except rainy Liu et al.,
stem dry days) simulating a 21% depletion of the ozone (2013)
mass layer vs. no UV-B radiation
16% NA From 280 to Natural lighting vs. glasshouse with complete Zhang et al.,
Leaf area " +19% Cucumber NA NA Solar UV-A without UV-B radiation vs. no solar Krizek et al.,
UV radiation (1997)
" 40% Lettuce 12 NA UV-B supplementation vs. no supplementation Wargent et al.,
(2006)
22% Soya NA From 280 to Natural lighting vs. glasshouse with complete Zhang et al.,
46% Rose bush UV-A: 12.0 vs. UV-A: 315- High altitude (2800 m) natural light vs. crop Terfa et al.,
1.7 400 under plastic film absorbing 96% of UV (2014)
UV-B: UV-B: 280- radiation
0.88 vs.0.036 315
(V m2 s1)
Leaf " +20% Lettuce 12 From 280 to UV-B supplementation vs. no UV-B radiation Wargent et al.,
thickness 320 (2006)
Branching Stem or " +8% Bean 9.6 NA UV-B supplementation simulating a 21% Barnes et al.,
tiller depletion of the ozone layer vs. no UV-B (1990)
+40% Kochia
number radiation
scoparia
+15% Oat
+15% Wheat
+13% Poinsettia 0.36 or 0.72 NA Irradiation (1 h) with UV-B at the middle of the Torre et al.,
night period (2012)
G Model
Table 2 (Continued)
effect disease
(nm)
NQa Broad bean 9.6 NA UV-B supplementation simulating a 21% Barnes et al.,
depletion of the ozone layer vs. no UV-B (1990)
Maize
radiation
Sunflower
Flowering Flowering # NQa Limnanthes 3, 6, 9, 12 or 15 From 280 to Plants grown under glasshouse, with 5 dosages Sampson and
rate alba 320 of UV-B supplementation (5h d1) Cane, (1999)
= Malcolmia NA From 280 to UV-B supplementation simulating a 15% Petropoulou
maritima 320 depletion of the ozone layer vs. no UV-B et al., (2001)
radiation
Phacelia 3, 6, 9, 12 or 15 From 280 to Plants grown under glasshouse, with 5 dosages Sampson and
campanularia 320 of UV-B supplementation (5h d1) Cane, (1999)
Delay before " NQa
flowering
= Limnanthes
alba
Flowering = Malcolmia NA From 280 to UV-B supplementation simulating a 15% Petropoulou
duration maritima 320 depletion of the ozone layer vs. no UV-B et al., (2001)
radiation
Limnanthes 3, 6, 9, 12 or 15 From 280 to Plants grown under glasshouse, with 5 dosages Sampson and
alba 320 of UV-B supplementation (5h.d1) Cane, (1999)
Flower " NQa Malcolmia NA From 280 to UV-B supplementation simulating a 15% Petropoulou
diameter maritima 320 depletion of the ozone layer vs. no UV-B et al., (2001)
radiation
Pathogen and % leaves # 51% soybean/ NA 315 Field plots receiving solar UV-B vs. plots Zavala et al.,
herbivore with lesions Anticarsia covered with polyester film excluding the UV-B (2001)
resistance gemmatalis component of sunlight
63% Grapevine/ from 3 to 5 UV-A:315- Field plots receiving solar UV vs. plots covered Keller et al.,
Botrytis 400 with a diacetate film blocking 98% of UV-A and (2003)
cinerea UV-B:280- UV-B radiation
315
Lesion # 240%
number
/leaf
Area lesions " 68% Arabidospis/ NA 310 Field conditions with ACLAR film which Caputo et al.,
/plant Plutella transmit >90% of all UV radiations vs. polyester (2006)
xylostella L. film blocking all UV radiations below 310 nm
-52% Arabidopsis/ 5.5 NA greenhouse. Plants irradiated 4h d1. Polyester Demkura and
Botrytis film transparent to UV-B vs. polyester film Ballaré, (2012)
cinerea blocking UV-B radiations
From 63% to Cucumber/ 0.9 NA greenhouse. Plants irradiated 15min d1 with Suthaparan
80% Podosphaera UV-B radiations vs. non-irradiated plants et al., (2014)
xanthii
Concentration b-carotene = Lettuce 18 mmol m2 UV-A white light (Fluorescent lamps–110 mmol. Li and Kubota,
in s1 peak:373 m2 s1), supplemented by UV-A LEDs vs. white (2009)
metabolites light only
Carotenoids # 11% Pepper 5.53 (UV-B: UV-C: 200- LEDs providing 30% (low) blue light (100 mmol. Hoffmann
4.5, UV-C: 1.0, 280 m2 s1) with or without exposure to UV et al., (2015)
UV-A:0.03) UV-B: 280-
320 peak at
290
UV-A: 320-
400
6% LEDs providing 62% (high) blue light
((100 mmol. m2.s1) with or without exposure
to UV
Anthocyanin " +800% Lettuce NA above 320, tunnels covered with a UV-transparent Tsormpatsidis
350, 370 or polyethylene film vs. UV-B-blocking et al., (2008)
380 polyethylene film
+11% 18 mmol m2 UV-A white light (Fluorescent lamps— Li and Kubota,
s1 peak:373 110 mmol m2 s1), supplemented by UV-A (2009)
LEDs vs. white light only
+675% NA from 280 to tunnel covered with a UV-transparent Tsormpatsidis
400 polyethylene film (81% of transmission) vs. UV- et al., (2010)
B-blocking polyethylene film (3.5% of
transmission). Plants transferred from UV-
G Model
Table 2 (Continued)
effect disease
(nm)
blocking to UV-transparent film at
transplantation stage, vs. no transferred plants.
Measurements at harvest stage, 14 days later
+367% plants grown under UV-blocking film and
transferred under UV-transparent film 6 days
before harvest vs. no transferred plants.
Measurements at harvest stage
= Pepper 5.53 (UV-B: UV-C: 200– LEDs providing 30% (low) or 62% (high) blue Hoffmann
4.5, UV-C: 1.0, 280 light (100 mmol. m2 s1) with or without et al., (2015)
UV-A:0.03) UV-B: 280– exposure to UV
320 peak at
290
UV-A: 320-
400
Flavonoids " NQa Lettuce NA Above 320, Tunnel covered with a UV-transparent Tsormpatsidis
350, 370 ou polyethylene film vs. UV-B-blocking et al., (2008)
380 polyethylene film
# 27% Alnus incana NA NA Field conditions. Branchs covered with a UV- Kotilainen
transparent plastic film vs. UV-B-blocking et al., (2008)
10% Betula
plastic film
pubescens
= Pepper 5.53 (UV-B: UV-C: 200- LEDs providing 30% (low) blue light (100 mmol. Hoffmann
4.5, UV-C: 1.0, 280 m2 s1) with or without exposure to UV et al., (2015)
UV-A:0.03) UV-B: 280-
320 peak at
290
UV-A: 320-
400
LEDs providing 62% (high) blue light (100 mmol.
m2 s1) with or without exposure to UV
Phenolics or " NQa Lettuce NA Above 320, Tunnels covered with a UV-transparent Tsormpatsidis
phenolic 350, 370 ou polyethylene film vs. UV-B-blocking et al., (2008)
acids 380 polyethylene film
= Betula From 1.3 to 13 From 240 to Greenhouse, UV-B supplementation with De La Rosa
pendula 400 fluorescent lamps et al., (2001)
Alnus incana NA NA Field conditions. Branches covered with a UV- Kotilainen
transparent plastic film vs. UV-B-blocking et al., (2008)
Phenolics " +51%
plastic film
glycosides
= Betula From 1.3 to 13 From 240 to Greenhouse, UV-B supplementation with De La Rosa
pendula 400 fluorescent lamps et al., (2001)
tannins =
NA NA Field conditions. Branches covered with a UV- Kotilainen
Alnus incana transparent plastic film vs. UV-B-blocking et al., (2008)
plastic film
Chlorogenic # 19%
acids = Betula
pendula
a
NA: not available information; NQ: not quantifiable (graphics).
impact on these parameters (Table 1). A recent work using specific repression (Wilson et al., 1993), or no effect observed (Abidi
bandwidths of 5-nm intervals in the 430–470-nm spectrum et al., 2013; Sarala et al., 2007) (Table 1). In rose, blue light alone is
revealed that wavelengths shorter than 450 nm promoted more able to induce both shoot meristem organogenic activity and leaf
leaf and stem growth in green Perilla (Perilla frutescens) than longer expansion, whereas these processes are completely inhibited in
wavelengths (Lee et al., 2014). This could partly explain the the absence of light (Girault et al., 2008). This indicates that blue
diversity of morphological plant responses to blue light according light can drive all the morphogenic processes induced by natural
to wavelengths. The effects of blue light on leaf dry mass (LMA) are white light (Abidi et al., 2013).
more stable across species, with greater LMA due to greater leaf Exposure to UV also modifies plant growth. Morphological plant
thickness (Brown et al., 1995; Hogewoning et al., 2010) and to responses to UV-B exposure are also species-dependent (Kakani
increased epidermal cell area (Sæbø et al., 1995) when blue light in et al., 2003). In numerous species, UV-B exposure decreases shoot
the spectrum is more intense. length, shoot dry mass and foliar area (Kuhlmann and Müller, 2009a,
The effects of blue light on branching are also species- b; Liu et al., 2013; Singh et al., 2011; Torre et al., 2012; Wargent et al.,
dependent, with bud burst stimulation (Barnes and Bugbee, 2006; Zhang et al., 2014) (Table 2). In some other species, exposure to
1992; Mortensen and Strømme, 1987; Muleo et al., 2001), supplementary UV-B light significantly increases leaf area as well as
G Model
fresh and dry biomass (Sakalauskaite_ et al., 2013). The effects of UV-A by blue light which reduced flower diameter (Khattak and Pearson,
on plant morphology have been less studied, but Krizek et al. (1997) 2006) or did not affect it but reduced flower peduncle length (Abidi
noted that exposure to UV-A without UV-B radiation decreased plant et al., 2013) (Table 1).
height and leaf area (Table 2). Whereas the effects of UV-B on flowering induction, precocity
Even if the effects of UV-B on branching vary in range according and duration depend on species (Petropoulou et al., 2001;
to species, UV-B exposure tends to increase the number of stems/ Sampson and Cane, 1999) plant exposure to UV-B can increase
tiller (Barnes et al., 1990; Torre et al., 2012) (Table 2). flower size (Petropoulou et al., 2001) (Table 2).
4. Blue and UV-B radiations also modulate plant sexual 5. Impact of UV and blue radiations on processes involved on
reproduction resistance to some biotic and abiotic stress
4.1. Impact on floral induction UV, and blue radiations to a lesser extent, can induce the
production of secondary metabolites or mechanisms of resistance
Flowering time, more specifically floral induction, determine to pathogens or herbivores that confer interesting qualities to
flower and fruit production ensuring high seed yield. Tight plants exposed to these radiations.
regulation of this latter is greatly influenced by environmental
light conditions. It is of crucial importance to increase our 5.1. UV-protective response
understanding of how blue and UV radiations modulate this step.
In plants, flowering is under the control of internal (autonomous, UV-B causes DNA damage and generates reactive oxygen
aging, gibberellins) and external (ambient temperature, vernaliza- species against which plants need to protect themselves. Protec-
tion and daylength) signals. All these cues converge toward two floral tion involves perception and signal transduction through photo-
integrator genes FLOWERING LOCUS T (FT) and SUPPRESSOR OF receptor UVR8. UV-B perception by this photoreceptor induces its
OVEREXPRESSION OF CONSTANS 1 (SOC1) which govern flowering activation through its monomerization. In its active form,
time (Parcy, 2005). The photoperiod pathway controls the response UVR8 binds to COP1, causing stabilization and accumulation of
to daylength through a signaling cascade involving GIGANTEA (GI) COP1 in the nucleus (Oravecz et al., 2006). This UV-B-dependent
and the transcriptional regulator CONSTANS (CO) (Mouradov et al., interaction between UVR8 and COP1 then leads to HY5 activation
2002). Protein CO in turn upregulates FT which encodes the mobile and stabilization (Favory et al., 2009). HY5 in turn is believed to
florigen in leaves, and protein FT then moves to the meristem promote the expression of several genes involved in UV-B
through the phloem to initiate flowering together with FLOWERING protection such as flavonoid synthesis and chalcone synthase
LOCUS D (FD) and 14-3-3 proteins (Abe et al., 2005; Corbesier et al., genes, as well as genes involved in DNA repair such as photolyase
2007). In the photoperiodic pathway, blue light perception is genes (Heijde and Ulm, 2012; Stracke et al., 2010; Ulm and Nagy,
mediated first by CRY1 and CRY2, and secondly by the blue light 2005). Interestingly, cry1 perceives UV-A–besides blue light –, and
receptor FLAVIN-BINDING KELCH REPEAT F-BOX 1 (FKF1). also promotes flavonoid synthesis under high irradiance through
CRY2 promotes flowering by mediating stabilization of CO (Valve- HY5. The cry1 signaling pathway of UV-A blue light under high
rde et al., 2004). At the end of the day in response to blue light, irradiance also contributes to the expression of genes involved in
CRY2 interacts with SUPPRESSOR OF PHYA-105 (SPA1). As a result, photooxidative stress tolerance, similarly to UVR8. In this case,
COP1 activity is suppressed, which decreases CO degradation and cry1 acts through ZINC FINGER PROTEIN EXPRESSED IN INFLORES-
promotes floral initiation in vascular bundles (Endo et al., 2007; Zuo CENCE MERISTEM LIKE1 (ZML1) and ZML2 (Shaikhali et al., 2012)
et al., 2011). When sensing blue light in long days (LD), to enhance the expression of antioxidative enzymes.
FKF1 interacts with GIGANTEA (GI) through its LOV domain. The
resulting FKF1–GI complex mediates blue-light-ubiquitin-depen- 5.2. Pathogen and herbivore resistance
dent degradation of CYCLING DOF FACTOR (CDF) proteins that
repress transcription of CO and FT genes (Fornara et al., 2009; Plant exposure to solar UV-B radiation tends to increase both
Imaizumi et al., 2005; Song et al., 2012). FKF1 also interacts resistance to herbivorous insects (see Caldwell et al., 2007 for a
physically with CO in a blue-light dependent manner, and the FKF1- review) and to microbial pathogens (Demkura and Ballaré, 2012;
CO complex increases CO stability in LD conditions (Song et al., Gunasekera and Paul, 2007; Kunz et al., 2008). The effects of blue
2012). Liu et al. (2013, 2008) recently showed that CRY2-interacting and UV-A radiations on plant diseases are more variable and differ
bHLHs (CIBs) were positive regulators of CRY2 blue-light-mediated according to the pathosystem. For example, without UV-A or
flowering through FT activation. UV-B is perceived by UVR-8 and without blue wavelengths, light sources favor the development of
acts as an input signals of the circadian clock involved in the the tomato mosaic virus disease in pepper but reduce the
photoperiod pathway (Fehér et al., 2011). development of powdery mildew in cucumber (Schuerger and
Brown, 1997; Wang et al., 2010b).
4.2. Consequences on flowering time and morphology An analysis of these responses needs to consider the effects of
specific wavelengths on all organisms involved: higher or lower
In A. thaliana, blue light accelerated flowering by 15 days when plant resistance, but also bioaggressor preferences for specific
plants were grown under continuous blue light as compared to wavelengths or direct damage to pathogens by specific radiations.
those grown under red light at a similar intensity (Guo et al., 1998). Just after the detection of a consumer, various hormones, mainly
Recently, blue light alone induced full floral development in rose jasmonate (JA) and salicylic acid (SA), coordinate the plant immune
(Abidi et al., 2013). Compared to white light, both a reduction and response (see Ballaré, 2014 for a review; Cerrudo et al., 2012;
an increase of the share of blue light in incident light delayed McGuire and Agrawal, 2005; De Wit et al., 2013). The SA pathway is
flowering in chrysanthemum (Khattak and Pearson, 2006), activated predominantly to fend off biotrophic microbial patho-
whereas blue light alone did not affect or delay flowering in gens (Glazebrook, 2005), whereas the JA pathway is mainly
two rose bush cultivars (Abidi et al., 2013). While the impact of activated in response to attacks by necrotrophic pathogens or
blue light on the timing of floral induction is well studied, less is chewing insects.
known about the processes involved in flower development. The role of blue photoreceptors in the regulation of defense
However, morphological flower traits were only slightly modified responses is not entirely established. CRY1 positively regulates SA-
G Model
induced PATHOGENESIS-RELATED (PR) gene expression, increasing human health can also consist of decreased levels of unwanted
Arabidopsis resistance to Pseudomonas syringae but only under components such as oxalate or nitrate, which decrease when
continuous blue light conditions (Wu and Yang, 2010). It has no plants are exposed to blue light (Ohashi-Kaneko et al., 2007)
role in induced defense responses under short-day blue light (Table 1).
conditions (Griebel and Zeier, 2008). CRY2 and PHOT2 are
specifically required for PR-mediated Arabidopsis defense against 7. Use of blue and UV-B lights in ornamental and vegetable
turnip crinkle virus (Jeong et al., 2010). UV-B radiation is a positive horticulture
regulator of plant defence through both JA-dependent and JA-
independent pathways. Positive effects of UV-B radiation on plant The irradiance of the ornamental and vegetable cultures,
defense against Botrytis cinerea are conserved in Arabidopsis complementary to the natural light, has been used for many years
mutants deficient in JA-signaling or insensitive to JA (Demkura in northern countries. It is usually provided by HPS lamps which
and Ballaré, 2012), and plant defenses are lost in some mutants have the advantage of being easily available and cheap, emitting a
disrupted in the JA signaling pathway (Caputo et al., 2006 in strong light intensity. However these lamps consume a lot of
Arabidopsis; Demkura et al., 2010 in a Nicotiana attenuata mutant electrical energy, have a relatively weak light output and present a
silenced for a lipoxygenase gene (NaLOX3) involved in JA broad light spectrum, but unbalanced with a weak emission in
biosynthesis). Solar UV-B radiation does not affect JA accumulation blue. Recently, a new type of lighting, based on LEDs, appeared on
in Nicotiana attenuata, but increases JA sensitivity (Demkura et al., the market. These lamps consume little electrical energy (20–30%
2010). Thus, conversely to low R/FR ratios that limit the investment of the electricity of HPS lamps) (Taulavuori et al., 2013), with a good
of plant resources in anti-herbivore defence to benefit growth, UV- light output and offer the possibility of a precise spectral band (van
B enhances jasmonate sensitivity and acts as a signal of abundant Ieperen, 2012). Thus the use of LEDs makes it possible to modulate
light and, therefore, do not limit the investment of resources in the wavelengths implied in photosynthesis and the photo-
antiherbivore defense (Demkura et al., 2010). The photoreceptor morphogenesis, such as B and R (B light is seldom applied alone)
UVR8 is implicated in the positive effects of UV-B on plant defense or in plant health such as UV. The launch of monochromatic LEDs
(Ballaré et al., 2012). It is required for UV-B-induced accumulation will certainly allow a more precise control of plant development
of sinapates (Demkura and Ballaré, 2012), important precursors in and health and a fast development of the LEDs techniques.
the synthesis of syringyl-type (“defense”) lignin which strengthens For ornamental potted plants cultivated for the interior
plant cell walls and prevents penetration of fungal hyphae decoration or the garden, plant shape is one of the essential
(Jenkins, 2009; Lloyd et al., 2011). Moreover, UV-B induces components of visual quality (Boumaza et al., 2010), therefore of
flavonoid accumulation controlled by UVR8 (Jenkins, 2009) and the commercial value of the product. Shape is the result of
flavonoids increase plant defense against herbivorous insects architectural construction, which in turn is the result of the growth
(Hoffmann-Campo et al., 2001). and branching processes. It is thus necessary for horticulturists to
control growth and branching. Thus, under greenhouse conditions,
6. Impact of UV and blue radiations on metabolites production. supplementary R (80%) and B (20%) LED lighting reduced stem
length by 34% in poinsettia (Euphorbia pulcherrima) compared to
Exposure to blue or UV radiations also tends to increase HPS lamp lighting (Islam et al., 2012). Under natural light and a B-
production of bioactive compounds, which, for vegetable and fruit light-transparent filter, chrysanthemum (Dendranthema grandi-
products, can carry out beneficial functions for human health. flora) plant height was reduced by 23% and branching was
Among photosynthesis pigments, blue light can increase total increased by 22% (Mortensen and Strømme, 1987). In plants
carotenoid (Johkan et al., 2010; Kopsell and Sams, 2013; Li and marketed for flowers, the horticulturist must also control the
Kubota, 2009; Ohashi-Kaneko et al., 2007; Wang et al., 2014), flowering process, more particularly floral induction. B light alone
(Table 1). By contrast, exposure to UV-A tends to decrease does not seem to sufficiently modify the flowering process to be of
carotenoid levels, but a higher proportion of blue light in the practical value in horticulture, e.g., by shifting the flowering period.
spectrum limits this decrease (Hoffmann et al., 2015) (Table 2). However, in combination with R light, B light can delay flowering,
In order to protect cells and prevent damage by UV radiations, as observed in fuchsia (Fuchsia hybrida) cultivated in growth
plants accumulate UV-absorbing compounds in their cell epider- chamber conditions (Moe et al., 2002). This response appears to be
mis. Accumulation of flavonoids, suspected to offer UV protection species-dependent since it was not observed in azalea (Rhodo-
(Treutter, 2005), is induced by UV-B. This accumulation under UV- dendron simsii) (Schamp et al., 2012).
B exposure is mentioned for several species (Rozema et al., 1997; In vegetable horticulture, the essential components of the
Tsormpatsidis et al., 2008). Nevertheless, UV-B exposure does not commercial yield of a crop are biomass production and product
induce accumulation of flavonoids in pepper (Hoffmann et al., quality, i.e., its visual aspect and nutritional value. These processes
2015), and even strongly decreases (27%) flavonoid content in directly depend on the quantity and quality of photosynthetically
Alnus incana (Kotilainen et al., 2008) (Table 2). Accumulation of active radiations, especially B light and the R/FR ratio, received by
anthocyanins is promoted by supplementation with UV-A or UV-B plant leaves. Blue light alone caused a 39% reduction in lettuce
or blue wavelengths (Carvalho and Folta, 2014; Li and Kubota, (Lactuca sativa) biomass, and 58% in spinach (Spinacia oleracea), in
2009Tsormpatsidis et al., 2008; Tsormpatsidis et al., 2010), even growth chamber conditions, as compared to plants cultivated
when UV-B irradiation is only applied over a short period (6 days) under fluorescent lighting (Ohashi-Kaneko et al., 2007). A
before harvest to avoid a depressive effect of UV-B on growth combination of B, R and FR lights under greenhouse conditions
(Tsormpatsidis et al., 2010). increased dry matter weights in cucumber (Cucumis sativus) and
As for effects on human health, the concentration in total tomato (Solanum lycopersicon), by 21% and 15%, respectively, as
aliphatic glucosinolate, an anti-cancer compound, increases when compared to plants cultivated under HPS lamps (Hogewoning
the part of blue light in Brassica oleacea lighting increases (Kopsell et al., 2012). This positive effect on biomass seems to require a low
et al., 2014) (Table 1) or after a short UV-B exposure before harvest B/R ratio (Nanya et al., 2012). Such a positive effect of B light is also
(Mewis et al., 2012). The concentration of chlorogenic acid, a commonly observed on the nutritional value of products. Thus, in
compound which decreases the risk of diabetes type 2, highly broccoli (Brassica oleacea var. italic) cultivated in growth chamber
increases in tomato and basil under supplemental blue light conditions with a 20/80 B/R ratio, concentrations in total
(Taulavuori et al., 2013). Favourable effects of light quality on carotenoids, macro-elements such as magnesium, micro-elements
G Model
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of bioactive compounds to improve nutritional and health values of receptors for plants and animals. Science 284, 760–765.
vegetables and fruits. Other fields of investigation are possible, Cerrudo, I., Keller, M.M., Cargnel, M.D., Demkura, P.V., Wit, M., de, Patitucci, M.S.,
Pierik, R., Pieterse, C.M.J., Ballaré, C.L., 2012. Low red/far-red ratios reduce
notably genetic improvement, with the selection of plants more Arabidopsis resistance to Botrytis cinerea and jasmonate responses via a COI1-
sensitive to specific B/R ratio, for example to obtain more compact JAZ10-dependent, salicylic acid-independent mechanism. Plant Physiol. 158,
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up-regulated by blue light and controls hypocotyl/stem growth and
Acknowledgments anthocyanin accumulation. Plant Physiol. 141, 61–74.
Chen, C., Xiao, Y.-G., Li, X., Ni, M., 2012. Light-Regulated Stomatal Aperture in
Arabidopsis. Mol. Plant 5, 566–572.
We thank our colleagues from ARCH-E (Architecture and Christie, J.M., Murphy, A.S., 2013. Shoot phototropism in higher plants: new light
Environment) team from IRHS for fruitful discussions and through old concepts. Am. J. Bot. 100, 35–46.
collaboration in research work on plant response to environment, Christie, J.M., Yang, H., Richter, G.L., Sullivan, S., Thomson, C.E., Lin, J.,
Titapiwatanakun, B., Ennis, M., Kaiserli, E., Lee, O.R., Adamec, J., Peer, W.A.,
and light in particular, over the last years. We also thank Sabine
Murphy, A.S., 2011. Phot1 inhibition of ABCB19 primes lateral auxin fluxes in the
Demotes-Mainard and Vincent Guerin (UMR IRHS) for useful shoot apex required for phototropism. PLoS Biol. 9, e1001076.
suggestions on the manuscript and Annie Buchwalter (scientific Christie, J.M., 2007. Phototropin blue-light receptors. Annu. Rev. Plant Biol. 58, 21–
translator) for reviewing the paper for English usage. 45.
Cominelli, E., Galbiati, M., Vavasseur, A., Conti, L., Sala, T., Vuylsteke, M., Leonhardt,
N., Dellaporta, S.L., Tonelli, C., 2005. A guard-cell-specific MYB transcription
factor regulates stomatal movements and plant drought tolerance. Curr. Biol. 15,
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EEB 2954 No. of Pages 17

Environmental and Experimental Botany xxx (2015) xxx–xxx

Contents lists available at ScienceDirect

Environmental and Experimental Botany

Light signaling and plant responses to blue and UV radiations—

Introduction resistance to pests and diseases, product quality), through (1)

2 L. Huché-Thélier et al. / Environmental and Experimental Botany xxx (2015) xxx–xxx

radiations enhance plant resistance to pathogens and herbivores 2.1. UVR8:

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6 L. Huché-Thélier et al. / Environmental and Experimental Botany xxx (2015) xxx–xxx

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Process Variables Blue Quantiﬁcation Species/disease Light conditions References

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L. Huché-Thélier et al. / Environmental and Experimental Botany xxx (2015) xxx–xxx 9

Process Variables UV Quantiﬁcation Species/ Light conditions References

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