UV radiation: plant responses and an in-depth mechanism of sustainability under climatic extremities

doi:10.21203/rs.3.rs-2219238/v1

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UV radiation: plant responses and an in-depth mechanism of sustainability under climatic extremities

https://doi.org/10.21203/rs.3.rs-2219238/v1

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Light is the primary element that plants need to grow, and UV rays, the harmful part of solar radiation, can decrease plant growth by slowing down the metabolic rate of photosynthesis. There are three different types of ultraviolet (UV) radiation, each with a distinct waveband: UV-A (315–400 nm), UV-B (280–315 nm), and UV-C (280 nm). Wavelength, intensity, and exposure to light have a big impact on plant quality and growth. Photoreceptors such as UV Resistance Locus 8 (UVR8), which promotes adaptation to UV-B stress, control how plants interact with UV radiation. While it is well established that UV in large amounts can harm production and quality metrics, some studies suggest that UV in moderate doses may promote biomass growth and the creation of beneficial chemicals that primarily absorb UV. It is well known that UV radiation causes differences in plant architecture, which are significant in ornamental crops and raise their economic value. Increased resistance to insects and diseases and a reduction in postharvest quality depletion are two effects of abiotic stress brought on by UV exposure. This review emphasises how UV may affect plant quality, growth, photomorphogenesis, and resistance to abiotic and biotic stress.

UV-A

UV-B

UV-C

ROS

UV radiation

Absorbing compounds

Photomorphogenesis

Plants, which are sessile organisms, are inevitably subjected to ecological changes that have a negative impact on their development and function (Singh et al. 2019). Plant life depends on solar radiation is essential for plant life, and UV radiation is a component of the sunlight that reaches the Earth's surface and affects living organisms (Yin and Ulm 2017; Zlatev et al. 2012). Based on their wavelength ranges (100–280 nm, 280–315 nm, and UVA 315–400 nm, respectively), the UV radiation range is divided into UVC, UVB, and UVA (Dawood et al. 2020; Müller-Xing et al. 2014). Despite making only a small portion of the sunlight that reaches Earth's surface, UV radiation has a substantial biological impact on organisms, which can directly alter plant-phyllosphere interactions between plants and their rhizosphere, with most research focusing on UV-B (Carvalho and Castillo 2018; Robson et al. 2015). In terms of wavelength, intensity, and exposure, UV radiation has a significant effect on the quality and growth of plants. Numerous studies have demonstrated that high UV-A doses have a detrimental effect on the elaboration of PSII, reducing maximum quantum efficiency, electron transport rate, and photosynthesis because they result in a decrease in ribulose-1,5-bisphosphate carboxylase-oxygenase (RuBisCO) (C3 species) or PEP carboxylase (C4 species), as well as an increase in stomatal resistance and the production of reactive oxygen species (ROS) (Nawkar et al. 2013; Prado et al. 2012; Takahashi and Badger 2011; Vass et al. 2002). Abiotic stress and photomorphogenic alterations can be brought on by UV-B. High UV-B exposure results in physiological stress that impairs cell functions, produces ROS and degrades and damages DNA (Heijde and Ulm 2012; Shi and Liu 2021). Low UV-B exposure encourages the development of photomorphogenic traits like cotyledon expansion, anthocyanin and flavonoid production, inhibition of hypocotyl growth, and stomatal opening (Jenkins 2009). UV Resistance Locus 8 (UVR8), a photoreceptor that enables adaptation to UV-B stress, controls how plants interact with UV light (Favory et al. 2009; Rizzini et al. 2011). The promoter region of the gene elongated hypocotyl 5 (HY5) is connected to the chromatin of UV-B responsive genes (UVR8). The E3 ubiquitin ligase, also known as constitutively photomorphogenic 1, and UVR8 activity are linked (COP1). Together, UVR8 and COP1 interact to cause a photomorphogenic response (Favory et al. 2009; Liao and Jenkins 2022). UVC, the most harmful Ultraviolet spectrum, is virtually ever absorbed by living beings on our planet due to the ozone layer in the stratosphere (Hornung and Downham 2018). The harmful effects of UVC radiation are now attributed to issues with ozone depletion in the stratosphere that are gradually getting worse and are considered stressors (Ballaré et al. 2011; Biever et al. 2014; Rai and Agrawal 2020). Despite effectively blocking UVC light, the ozone layer in the stratosphere reduces the amount that reaches the Earth's surface, releasing chlorofluorocarbons into the atmosphere and thinning the ozone layer. Thus, the future trend of growing UVC radiation reaching the soil brings about the ecological effects of increased UVC on the environment and on the agricultural system (Kataria and Guruprasad 2012). High UVB and UVC exposure led to dimerization and ionisation of DNA's pyrimidines, damaging it and interfering with protein synthesis, which caused a significant change in plants' typical cellular activities (Nawkar et al. 2013; Sharma et al. 2017; Tossi et al. 2019). When exposed to high-energy UVC radiation, leaf area, chlorophyll content, photosynthesis, DNA, membrane lipids, and proteins undergo morphological, developmental, and physiological damage (Geras’kin et al. 2016; Müller-Xing et al. 2014). Chloroplasts appear to be the most sensitive structures in plant cells or organelles where UVC radiation generates oxidative stress, thereby decreasing photosynthetic output and reducing crop productivity (Acuña-Rodríguez et al. 2021; Czégény et al. 2016; Nawkar et al. 2013; Vanhaelewyn et al. 2020a).

The non-ionizing part of the electromagnetic spectrum, which includes ultraviolet light (UV), accounts for 8–9% of all solar radiation. Three classes comprise the ultraviolet (UV) component of the electromagnetic spectrum: UV-A (315–400 nm), UV-B (280–315 nm), and UV-C (200–280 nm) (Kim et al. 2016). The only two of the three that reach the earth's surface are UV-B and UV-A. Even while UV radiation is only a small portion of the sunlight that reaches the Earth's surface, it has a substantial biological impact on organisms, with most research focused on UV-B, which can have an impact on interactions between plants and their environment (Balcerowicz 2022; Carvalho and Castillo 2018; Robson et al. 2015). While UV-B and UV-C greatly stimulate the formation of secondary metabolites, UV-A and UV-B primarily influence morphogenesis and phototropism. Plant diseases are negatively impacted by short-wave UV radiation (350 nm) both directly and indirectly. DNA damage, protein polymerization, enzyme inactivation, and increased cell membrane permeability all have direct consequences. Since UV-C is the most intense radiation, it kills germs more effectively at lower concentrations but, as a result, frequently damages plants (Vanhaelewyn et al. 2020a). Indirect effects can be attributed to UV-B-specific pathways, such as the upregulated plant defence responses that are UVR8 dependent, the upregulated ROS accumulation caused by UV-B and UV-C, and the formation of secondary metabolites such as phenolic compounds (Vanhaelewyn et al. 2019).

Without photosynthesis, there wouldn't be any life on Earth, which makes it the most crucial process in the living world. Furthermore, biomass- one of the results of photosynthesis—serves as the basis for numerous technologies and the main food supply for a range of animals as well as humans. The ozone layer's depletion, on the other hand, increases UV radiation, which has an impact on plants' ability to photosynthesize. Being immobile, plants are constantly subjected to UV radiation and must thus adapt and respond to the changing environment. These adaptations take the form of altered phenotypic traits, changes in plant architecture (acclimation responses), physiological attributes (Takshak and Agrawal 2019), and modifications in biochemical and genetic levels (Kumari et al. 2010; Tripathi et al. 2011). UV radiation, which intracellular membranes can readily absorb when exposed in high doses, and molecules like amino acids, nucleic acids, and membrane polypeptides damage living organisms (De Jager et al. 2017; Roy 2017). Due to its mutagenicity and non-selective harmful characteristics, it is classified as a "complete carcinogen." As a replication-, transcription-, and translation-preventive agent, it is a certain tumour starter and promoter in addition to its role in oxidative damage (D’Orazio et al. 2013; Roy 2017). Severe stress circumstances have been shown to result in metabolic dysfunction. Plants respond to UV-B radiation depending on variability, intensity, fluency rate, duration of exposure and wavelength. According to a meta-analysis of studies on the effects of UVB radiation on crops, agricultural yield declines by 1% for every 3% rise in weighted UVB irradiation (Ballaré et al. 2011). Studies on plants have revealed that UVB decreases the activity of Rubisco, the level of carotenoids and chlorophyll, as well as the amount of leaf area and leaf thickness in field crops (Martel and Qaderi 2016). Plant metabolic activities are significantly harmed by UV-B, which increases defensive chemicals such as flavonoids (Takshak and Agrawal 2019). UV-B radiation enhanced lipid peroxidation and altered the activities of antioxidant enzymes that scavenge reactive oxygen species. With the increasing UV-B radiation intensity, a drop has been observed in superoxide dismutase, peroxidase, and catalase activity but ascorbate peroxidase activity was increased (Chen et al. 2019). There are hazards associated with applying or exposing organisms to UV light. Some UV-induced morphological responses can be attributed to the interaction of the highly energetic UV radiation with molecules, in addition to active detection and signal induction to UV radiation by photoreceptors. Because UV-A is not absorbed by DNA, it has been demonstrated that UV-A irradiation of plant tissues has little effect on DNA damage (Vanhaelewyn et al. 2020b). However, ROS production resulting from photosensitizing reactions, particularly by highly reactive singlet oxygen, can still lead to indirect DNA damage (Alscher et al. 1997). Higher energy UV irradiation, such as UV-B and UV-C, on the other hand, is significantly absorbed by nuclear, mitochondrial, and chloroplast DNA and results in a similar mutational process (Vanhaelewyn et al. 2020b).

Plant sensitivity to UV exposure may be assessed through direct UV damage to cell structure and functional components or inadequate acclimation (Smith et al. 2000). By altering the structure of the leaves, UV-B radiation can cause the processes of leaf differentiation and senescence. Exposure to UV-B increased auxiliary branching and leaf curling while decreasing plant height, leaf area, and dry weight (Zhao et al. 2003). According to Dai et al. (1995), rice plants' leaf area and dry weight dramatically decreased after a few weeks of UV-B exposure. Increased UV-A reduced leaf area and improved biomass productivity per unit of leaf area and nitrogen, as well as per unit of plant biomass (Weih et al. 1998). As the leaf area ratio, leaf area productivity, and leaf nitrogen productivity were all decreased by high UV-B levels, it was evident that these factors decreased the relative growth rate and nitrogen productivity. Avena fatua and Setaria viridis species produced in greenhouses under UV-B radiation displayed varied development habits and responses to UV-B radiation. While tolerance levels depend on the type of plant, its age, how long it has been exposed to the radiation, and other factors, plants are capable of adapting to certain UV-B radiation as well as to light intensity. Plant leaf anatomy is altered and biomass is reduced if the UV-B dosage exceeds the tolerance limits (Coleman and Day 2004; Kakani et al. 2003a; Parihar et al. 2021; Zhao et al. 2003). UV B radiation was discovered to have a favourable effect on seed germination rate. After being exposed to UV B radiation, the germination rate of V. mungo seeds increased by 33% (Shaukat et al. 2013). The germination rate of Phaseolus vulgaris (black bean) seeds was likewise boosted by UV-C exposure. Longer periods of exposure to low levels of UV (460 W/cm² for 60 min) accelerated kidney bean development and biomass buildup. Biennial crops, however, responded favourably to shorter exposure times. In kidney beans, exposure to UV light increases plant height by 11–39% (Kacharava et al. 2009). After 3 hours at an irradiance of 1.3 mW/cm2, the seed coat thickness of the soft, macha, and red dika kinds of wheat rose by 28%, 25–36%, and 66–77%, respectively. In comparison to the other two wheat varieties, the red dika wheat had the thickest seed coat (Badridze et al. 2016). Exposure to UV A enhanced the germination of mung bean (Vigna radiata L.) seeds and improved seedling growth in terms of leaf area, root and shoot length, and dry weight (Hamid and Jawaid 2011). The UV C (280 nm) treatment of ground nut (Arachis hypogaea) and mung bean (Vigna radiata) seeds increased the germination rate and seedling vigour over a range of periods (5–60 min). Mung bean seed germination was 100% after 30 minutes, and ground nut seed germination was 87% after 60 minutes of UV-C exposure. In mung beans, the maximum shoot length and weight were reached after 15 minutes of UV C exposure, but the maximum root length and weight were reached after 30 minutes. But in the case of ground nuts, all the above metrics obtained a maximum of 60 min UV exposure (Siddiqui et al. 2011). In comparison to controls, the ground nut seeds' seedling vigour and bloom production increased when exposed to UV C for periods ranging from 5 to 60 minutes. Similarly, all UV C treatments increased pod biomass (Neelamegam and Sutha 2015). Buckwheat seeds were exposed to UV B radiation in the field using filtered UV fluorescent lights (275–380 nm), and it was discovered that this had a good impact on the plant's height and production (Yao et al. 2007). Similar to this, peppermint showed accelerated vegetative growth and an increase in the number of peltate glandular trichomes per leaf when exposed to UV B and photosynthetically active light together (PAR) (Behn et al. 2010).

The procedure by which CO₂ and water are transformed into carbohydrates in the presence of photosynthesis is the process of using sunlight. Two components make up the photosynthetic apparatus: PS-I and PS-II photosystems. Even though UV-B affects most elements of damage to the ultrastructure of the chloroplast and light-harvesting complex, with a loss in Rubisco activity, a drop in oxygen evolution, and a rise in CO₂ fixation, a decline in chlorophyll content, and a rise in starch levels. Moreover, the photosynthetic plant species, cultivars, and other factors influence how plants respond to UV-B radiation, the environment, UV-B radiation dosage, and the proportion of PAR to UV-B radiation (Rai and Agrawal 2017).

There have been numerous studies on the function of photosynthetically active radiation (PAR) between 400 and 700 nm, plant sensitivity to UV-B and photomorphogenic reactions radiation and its opposite. Direct improvements in photo repair, levels of photoprotective chemicals and photo-reactivation as well as indirect mechanisms, such as alterations in leaf anatomy, have been proposed to account for the UV-protective benefits of high PAR light circumstances (Escobar-Bravo et al. 2021; Hoffmann et al. 2015).

Nevertheless, recent experimental data indicate the possibility that high PAR and UV-B could work in harmony and favourably impact plant photo-defence. UV-B exposure has been demonstrated to boost net photosynthesis following exposure to lettuce (Lactuca sativa) under situations of high light intensity (Wargent et al. 2015). Similarly, UV-B induced photosynthetic rates in Swedish ivy (Plectranthus coleoides) by increasing stomatal conductance, the rate of CO₂ assimilation, and internal CO₂ content in the presence of both high and low PAR circumstances (Vidović et al. 2015), particularly under direct sunshine photo-inhibition (light-induced deactivation of conditions).

There was an important gap in how plants perceive UV radiation, although in recent times our understanding has been improved on this matter. Before this, it was thought that photomorphogenic responses to UV-A radiation were mediated by UV-A/blue photoreceptors, cryptochromes and phototropin, whereas UV-B radiation mediated by "UV-B photoreceptor" UV RESISTANCE LOCUS 8 (UVR8). Several experimental facts now settle the fact that UVR8 can alone play a major role in the perception of UV-A (315–350 nm) and UV-B or it may jointly regulate the genetic expression with cryptochromes through various interactions (Rai et al. 2021).

Cryptochromes (cry1 and cry2), phototropins (phot1 and phot2) and zeitlupe proteins are labelled as UV-A photoreceptors or blue photoreceptors (Christie et al. 2015) whereas UVR8 known as UV-B photoreceptors (Rizzini et al. 2011). Based on the responses that UVR8, CRYs and PHOTs mediate towards monochromatic radiations (Robson et al. 2019) and their strong photon absorbing capability in particular regions (Christie et al. 2012) they have been attributed to these roles.

While perceiving UV radiation plants also undergo some changes in response to different UV radiations. Blue light-induced gene expression, expansion of cotyledons, accumulation of phenolics, and alteration in flowering time are mostly mediated by cryptochromes (CRYs) (Wang et al. 2017). Phototropism and chloroplastic movement are mainly mediated through phototropins (PHOTs) (Christie et al. 2015), however under blue light CRYs and PHOTs are both involved in the regulation of stomatal openings and inhibition of hypocotyl growth (Wang et al. 2020). Likewise, UVR8 also led to various changes both inhibitory and inductive. Inhibitory effects were found on hypocotyl length, and expansion of cotyledons whereas induction of biosynthesis of flavonoids and accumulation of flavonoid are been found (Rai et al. 2019; Yin and Ulm 2017).

The signalling and molecular perspective of this discussion could be proved beneficial towards understanding the mechanism of UV perception by photoreceptors. Absorbing a photon by a photoreceptor is a similar process irrespective of wavelengths (Rai et al. 2021). Activation photoreceptors are simultaneous under both sunlight and shade light and their downstream interactions are very important for various responses exhibited by plants (Ballaré and Pierik 2017). UVR8 occurs as a homodimer and is mostly found in the cytosol in the absence of UV-B, but UV-B exposure causes UVR8 to monomerize and quickly accumulate in the nucleus (Rizzini et al. 2011). The same has been followed i.e., upon blue light exposure conversion of CRYs monomers to dimers has occurred but there are differences too and that is cry2 is mostly found in the nucleus, cry1 is found in both the cytosol and nucleus (Wang et al. 2016).

On a contradictory note, UVR8 can also monomerize under UV-A exposure but up to 335 nm (Rai et al. 2020) and this can be explained by the excitation energy transfer between three groups of tryptophan present in UVR8 protein (Li et al. 2020). After exposure to UV-A and UV-B, CRYs and UVR8 bind with E3 ubiquitin ligase COP1 (CONSTITUTIVELY PHOTOMORPHOGENIC 1) and inhibit the E3 ubiquitin ligase and stabilize the HY5 transcription factor (Holtkotte et al. 2017; Lau et al. 2019; Podolec and Ulm 2018; Qian et al. 2020).

Since UVR8 and CRYs downstream signalling shares a lot of components, multiple points of interaction are possible. According to Rai et al. (2021), there are three main levels of interaction that took place. At the first level of interaction, the binding of the VP-peptide motif of UVR8 and CRYs occurs at the WD40 domain of COP1 (Lau et al. 2019; Ponnu et al. 2019). The second stage of interaction involves the TF-specific regulation of gene expression, like HY5 (HYPOCOTYL 5), BES1 (BRI1-EMS-SUPPRESSOR1), BIM1 (BES1-INTERACTING MYC-LIKE 1), and PIFs (PHYTOCHROME INTERACTING FACTORs (Liang et al. 2018; Wang et al. 2018). Regulation of negative feedback happened at the third stage of interaction where REPRESSOR OF UV-B PHOTOMORPHOGENESIS (RUP1 and RUP2), and BLUE-LIGHT INHIBITOR OF CRYPTOCHROMES 1 (BIC1 and BIC2) are involved (Findlay and Jenkins 2016; Wang et al. 2017). As UV-B radiation occurs, it induces the expression of BIC1 and BIC2 thus activating UVR8 signalling, but overexpression of BIC1 and BIC2 masks the process of UVR8 redimerization which is mediated by CRYs (Tissot and Ulm 2020), reciprocally, expression of RUP1 and RUP2 under blue light exposure activate CRYs signalling and promoted UVR8 redimerization (Tissot and Ulm 2020). Redimerization of monomeric UVR8 RUPS negatively regulate UVR8 signalling (Findlay and Jenkins 2016), whereas CRYs dimerization is inhibited by BICs as negatively regulated (Wang et al. 2017). Further downstream interactions contributed by different other TFs mediated regulation of genes (Rai et al. 2020).

For a long time, UV radiation is considered to be hazardous and harmful to plants, but recent studies suggested that supplemental UV-B exposure can act as eustress (i.e., positive stress) and established the data that stated that a range of positive responses has been found to various plant process such as synthesis of UV shielding compounds (anthocyanin, flavonoids and phenolic acids), various antioxidants such as ascorbate, carotenoid and glucosinolates (Castagna et al. 2014; Moreira-Rodríguez et al. 2017; Sun et al. 2012). Yet UV possesses a small portion of the spectrum, but it is involved in many positive and negative effects of the plan’s growth (Yadav et al. 2020). The negative effects of UV are mainly exhibited by the morphology of the plant, as the plant adopts various changes in its vegetative structure to escape UV-mediated damage. According to Robson et al. (2019), UV-B-induced plants develop thicker leaves, shortening of petioles and curling of leaves. Liu et al. (2017) found that UV-B exposure at 3.33 µmol. m^− 2. s^− 1 exhibits a positive effect with increased stem diameter, total soluble sugar, total ascorbic acid, and enhanced activities of POX, SOD, and CAT in cucumber leaves while not compromising the net photosynthesis rate. Exposure of UV-C to Tomato plants at 1.0 kJ.m^− 2 was also found beneficial response on fruit number, fruit weight and net photosynthesis (Darras et al. 2020). Whereas according to Kaiserli (2018), UV-B radiation negatively regulates the expression of most of the cell-wall elongation genes that were induced by BRI1-EMS-SUPPRESSOR 1 (BES1). Reduction in plant stem elongation process by suppressing the biosynthesis and accumulation of auxin also found in Arabidopsis and coriander plants under high UV-B doses (Fraser et al. 2017). Irradiation of UV-A and UV-B was found to be beneficial in fruit quality enhancement (Ghasemi et al. 2019; Mariz-Ponte et al. 2019).

Plant development and environmental acclimation responses are significantly influenced by ultraviolet-B (UV-B, 280–315 nm) light, which is a natural component of sunlight. Many studies have shown that UV-B enhances plant photomorphogenesis, inhibiting hypocotyl elongation, increasing flavonoid and anthocyanin levels, and increasing the expression of UV-B-responsive genes (Fina et al. 2017; Henry-Kirk et al. 2018). Utilizing the UV-B signal, plants can UVR8. When UVR8 (UV RESISTANCE LOCUS 8) receives the UV-B signal, it depolymerizes an active monomer form from the inactive dimer, which it then transports to the nucleus in a dynamic, monomeric form (Heijde and Ulm 2013; Moriconi et al. 2018). RUPs (REPRESSOR OF UV-B PHOTOMORPHOGENESIS) promote the re-dimerization of UVR8.RUPs may physically interact with UVR8 and UVR8 re-dimerization, negatively affecting the transduction of UV-B signals (Yin and Ulm 2017). In horticulture, plant architecture is essential to quality and output (Loconsole and Santamaria 2021). In higher latitudes with limited light quality and quantity, when plants tend to elongate and the quality of production is poor, dwarfed plant design is crucial for aesthetic plants, boosting their economic worth. Dwarfed architecture is less prone to harm and provides advantages during production, transplant, and transportation (Mata and Botto 2009; Qian et al. 2020). Plants that are smaller in stature require less space to travel and grow in transplant nurseries. The cucumber is one example of a dwarfed plant that bears the same quantity and size of fruit as non-dwarfed plants. More sustainable production is made possible by saving space and minimising damage (Qian et al. 2020). One of the most significant factors in dwarfism is light. There has been extensive research on how UV radiation affects plant morphology. Plant responses vary considerably depending on the species (Kakani et al. 2003b). In some species of plants, exposure to UV-B has been reported to result in a decrease in foliar area, shoot dry mass, and shoot length (Liu et al. 2013; Torre et al. 2012; Zhang et al. 2014), however in other species, it has been found to result in an increase in leaf area, fresh weight, and dry weight, even though the impact of UV-B varies and depends on the species, exposure often increases the number of stems per tiller (Sakalauskaitė et al. 2013). Studies comparing the effects of UV-A and UV-B exposure on cucumbers reveal that while both UV-A and UV-B-exposed plants are dwarfed, UV-B-exposed plants lack a strong stem (Qian et al. 2020). In Arabidopsis, UV-A and UV-B respectively had stimulating and inhibiting effects. In eight accessions of A. thaliana grown indoors under low PAR conditions, further exposure to UV-A resulted in a 30–150% increase in rosette diameter. UV-B supplementation was added to improve this phenotype. The same result was seen in outdoor environments. More research revealed that the UV-B dwarfing effect outweighed the leaf size stimulatory effects of UV-A (Biswas and Jansen 2012).

Oxidative stress in plants under UV radiation is very closely related to drought and high light/temperature stress as UV rays are an integral part of solar light and excessive exposure to solar light could lead to UV stress associated with the other stress described earlier. Reactive oxygen species levels rise in cells after excessive exposure to UV-B radiation, as they do under most stressful circumstances (ROS) (Qamer et al. 2021). Particularly, on UV-B exposure superoxides accumulates in apoplastic space as a result of NADPH oxidases AtRBOHD and AtRBOHF, and in the chloroplastic region as UV-B alters PS-II function (Qamer et al. 2021). Plants have activated their defence against UV-B stress through the activation of various antioxidant enzymes such as catalase, peroxidase, superoxide dismutase, ascorbate peroxidase and glutathione reductase (Ferreyra et al. 2021).

Over-accumulation of ROS due to UV-B exposure can cause harmful effects on the expression of genes and enzymatic activities, thus disturbing various biochemical pathways, and leading to cellular damage and programmed cell death (Qamer et al. 2021). It was found that class II TGA factors involve in the control of H₂O₂ accumulation and degree of oxidative damage in the context of tolerance response towards UV-B induced damage, as well as expression of genes that code for enzymes that scavenge H₂O₂ such as GSTU (Herrera-Vásquez et al. 2021). Flavonoids are also involved in the scavenging of ROS and lowering the risk of oxidative damage (Fig. 1) as they contain phenolic hydroxyl groups in their structure (Jan et al. 2022).

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A major and well-known abiotic stress to the plant is UV radiation which negatively affects plant growth and productivity, but on other hand lower dose of UV specifically, UV-B radiation applied to the seed as a priming agent could have a positive effect on various aspects of the plant (Thomas and Puthur 2021). It was accepted by various researchers that solar UV radiation mostly plays as a regulator of growth and development but only in exceptional it acts as a stressor (Jenkins 2017; Verdaguer et al. 2017). Low dosage UV-B exposure can lead to changes in the antioxidant status, such as the regulation of the glutathione, phenylpropanoids, cinnamates, or flavonoids, and pyridoxine biosynthesis pathways (Hideg et al. 2013), and create an imprint of stress memory and trigger early adaptation when the plant gets the stressful condition later thus acclimatize plants to protect (Müller-Xing et al. 2014).

According to Thomas and Puthur (2020), phenylalanine ammonia-lyase (PAL), flavonoids and anthocyanins were accumulated higher when seeds primed with UV-B radiation. 4 kJm^− 2 of UV-B irradiation help plant by reducing the harmful effects of drought while enhances the accumulation of proline, soluble sugars, amino acids etc. also increased ROS scavenging capability of plants (Sen and Puthur 2021). Rice seed primed UV-B promoted innate tolerance potential of the seedling against NaCl and PEG stresses leading to reduce oxidative stress in seedlings of rice (Sen and Puthur 2021). UV-C also proved as a potential inducer of ROS-scavenging enzymes in plants under salt stress with higher antioxidant activity (Ouhibi et al. 2014). In 40 days old strawberry plant, UV-C exposure enhances defence against leaf spot disease (Xu et al. 2019). It was suggested that UV-C induce SAR in plants and regulates several transduction pathways and is involved in the activation of TFs such as FvPINS, PINS1 and FaNES (Bera et al. 2022b).

The main defence techniques used by plants against possibly harmful ultraviolet rays are the accumulation of UV-absorbing substances (flavonoids and relevant phenylpropanoids) and the resulting decrease in epidermal UV transmittance (TUV). These processes are essential parts of the general acclimatization response of plants to adapt to UV environments (Barnes et al. 2016). In plants, the epidermis acts as the main primary filter of sunlight or solar UV rays while transmitting visible light necessary for photosynthesis. Under UV environments reducing UV transmission is the very early stage of defence which started with an accumulation of UV-absorbing substances in epidermal tissues (Barnes et al. 2016; Bornman et al. 2015).

Redimerization of UVR8 leads to the activation of specific TFs responsible activation of genes for biosynthesis and localization of flavonoids (Qian et al. 2020; Tossi et al. 2019). Pathways involve in flavonoid synthesis are chalcone synthase (CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), and flavonol synthase (FLS) pathways (Hideg et al. 2013; Jan et al. 2022). Some HY5 and HYH are two homologues that regulate UV-B signalling and are also activated by blue and red light (Tissot and Ulm 2020). Cryptochromes (CRYs) and phytochromes (PHYs) are perceived as blue/UV-A and red light respectively through the HY5/HYH activation thus regulating the synthesis of flavonoids. According to Tissot and Ulm (2020), cross-regulation of photoreceptor pathways exists under a polychromatic light environment, and CRY1 and PHYB together with UVR8 contribute to UV-B responses plant exhibit under UV-B environments.

Besides the flavonoid-based protective mechanisms, there are some changes in plant morphology such as leaf thickness increases, an abundance of trichomes, vertical leaf orientation, protective basal sheath, and concealed apical meristem etc. (Thomas and Puthur 2017).

Table 1: Positive effects of UV exposure on different plants

Plant	UV dose	UV induced changes	Reference
Trigonella foenum-graecum	UV lamp exposure at 15 and 30 W for 1, 2, 3 and 4 h	Increased activity of SOD, GPX, GR and APX	(El-Shora et al. 2015)
Triticum aestivum	DRT-400 lamp Exposure of UV-A, B, and C at 1.19, 1.3 and 1.84 mW/cm² for 3 hours	Significantly increased proline content, ascorbate, and phenolic contents induce the activity of CAT, POD	(Badridze et al. 2015; Badridze et al. 2016)
Cucumis sativus	Photosynthetic active radiation at 200–212 µmol.m^− 2.s^− 1	Increased concentration of sugars, amino acids, phenolics, and hormones	(Palma et al. 2022)
Oryza sativa	Exposure of UV-B lamps from Q-Panel at 313, 40 W at 2.5, 5, 7.5 kJm^− 2 for 6, 24, 48 and 72 hours	Enhanced activity of SOD, CAT, POD	(Inostroza-Blancheteau et al. 2016)
Triticum aestivum	UV-C for 30–180 seconds	Increased germination chlorophylls and carotenoids content	(Sharlaeva et al. 2022)
Allium cepa L.	UV-C (254nm) and UV-A (365nm) were irradiated with a UVGL-58 Handheld UV Lamp for 72 hours	Antioxidant enzyme activities were stimulated significantly	(Çavuşoğlu et al. 2022)
Vigna mungo	UV B (312 nm) with fluorescent UV-B tube (TL e 40 W/12	Promoted activities of SOD, CAT, GPX	(Dwivedi et al. 2015)
Fragaria ananassa	UV (270–380 nm) irradiation for 3 hr at 0.17–0.96 kJ.m^− 2.day^− 1	Significantly reduced the incidence of powdery mildew	(Ota et al. 2021)
Nicotiana benthamiana	6.9 kJ.m^− 2.d^− 1 for 4 days	Positive effects on antioxidant capacity and flavonoid index were observed	(Mátai et al. 2019)
Fragaria ananassa	UV exposure (270–380 nm) for 3 hr (0.17–0.96 kJ.m^− 2.day^− 1)	Significantly reduced the incidence of powdery mildew on leaves	(Ota et al. 2021)
Triticosecale wittmack	UV-B	Higher maximum and minimum chlorophyll fluorescence, chlorophylls, carotenoids, starch and concentrations of UV-B absorbing compounds and intrinsic water use efficiency	(Bacelar et al. 2015)

Optimum germination of seeds is a prime condition for the good stand establishment which is a basic factor in crop production but due to various abiotic and biotic factors, different primary aspects of crop growth such as germination, field emergence, and seedling vigour are badly affected thus finally result in poor yield (Bera et al. 2022a; Rhaman et al. 2020). To encounter the limiting factors of the environment, plants adopt various modifications and changes in metabolic processes. In this context of protecting plants from various limiting factors “seed priming” has been developed as a cost-effective technique with controlled hydration and drying that initiate and enhances the pre-germinative metabolic process thus improving the germination process and acclimatising for various stress tolerance (Rhaman et al. 2020).

As time passes the area of priming was widen from hydropriming to chemical priming and now non-chemical physical priming that is priming with various natural factors such as magnetic field, UV radiation, solar radiation and others such as plasma, ultrasonic sound, microwaves, tele waves, x-ray and gamma radiation etc. (Bera et al. 2022b).

In this review, our focus will be on the priming potentiality of different UV radiations. In especially, how plants have developed homeostasis between the negative effects of UV radiation and the beneficial effects of UV priming on plants. This can be explained as there are certain signalling molecules activated when plants get radiation at high doses and these molecules are also activated during exposure at very low doses or dose lower than potentially harmful to the plant. Priming with low doses of UV makes a stress imprint or epigenetic memory in the seed or seedling or different plant parts, later when the stress intensity became higher than that of the threat range plants start to acclimatize and associated changes have been exhibited by the plants (Bera et al. 2022b; Hossain et al. 2018). These changes in plants in the context of morphological, physiological, biochemical and molecular characteristics assist them to uphold protection and tolerate unfavourable conditions during stress exposure (Song et al. 2019). Some of the positive effects of UV exposure on different plants are highlighted in Table 1. According to Ansari et al. (2021), in adverse environmental conditions plants modulate their metabolic processes and plants growing in these adverse conditions modulate to sustain their normal cellular functions.

<<<<<<<<<<<<<<<<<<<<<<<<<<<Table 1>>>>>>>>>>>>>>>>>>>>>>>>>>>>>

Germination of seed, photosynthesis and crop growth mostly depends on the environmental condition and plant genotype (Verdaguer et al. 2017), but UV-A and B radiation evidenced advantages in these parameters (Mariz-Ponte et al. 2018). According to Coffey et al. (2017), in plants, UV-B radiation can prompt the accumulation of phenolics which helps to decrease oxidative stress-mediated damage and inhibit the penetration of UV-B to the inner photosynthetic layers of leaves and other plant parts. Supporting this fact, Yang et al. (2018) described that in chrysanthemum leaves, 12-hour exposure to UV-B significantly increases total and individual flavonoids and phenylpropanoids content thus protecting the plant from the harmful effects of UV-B. Plants grown in a protected environment or within greenhouses reported improving crop protection against different pests and diseases after exposure to UV radiation controlled artificially (Escobar-Bravo et al. 2019a; Escobar-Bravo et al. 2019b). Without the use of pesticides, UV-B radiation has the ability to increase productivity and protect crops from pests (Escobar-Bravo et al. 2019b; Mazza et al. 2013). Seed priming with UV-B tempts mild stress in plants thus triggering the expression of stress-responsive genes and proteins such as LEAs (late embryogenesis abundant) which enhance drought tolerance levels in plants (Sen et al. 2021; Thomas et al. 2020). Positive effects of UV-C priming on Vicia villosa were found when UV-C was applied to Vicia seeds at a dose of 1000 J/m² (Semenov et al. 2021). At this dose seed hydration rate, germination, seed vigour, chlorophyll-a and b content, and carotenoid production are significantly enhanced. From the experiment of Hamid and Jawaid (2011), it was established that UV-A priming on mung bean seeds proved very promising in terms of seed germination rate, leaf area, vigour index, and biomass accumulation at the seedling stage. In lettuce seeds, UV-C at 0.82 and 3.42 kJm^− 2 for 1 and 4 minutes exposure reported higher antioxidant activity under salt stress thus helping the plant to withstand oxidative damages caused by the stress in the leaf of seedlings (Ouhibi et al. 2014). Another experiment with UV-C priming was proven beneficial to Phaseolus vulgaris as exposure to UV-C at 460 µW.cm^− 2 for 60 minutes enhanced seed germination and accumulation of bioactive molecules in seed coats (Guajardo-Flores et al. 2014).

Although ultraviolet rays reaching Earth's surface comprises a very minute portion of the total solar light spectrum, its presence especially UV-B has mostly deleterious effects on plants' growth and development. In recent times scientists are keen to work on how these harmful UV rays could use as a tool to replace chemical-based agriculture. As crops in the field are sessile and bare to the environment, it is difficult to protect them from solar radiation. Instead, UV rays at low doses or in a short period as supplementary exposure could use to preprint the stress memory in the early seedling stage or as seed priming agents, and when they got exposed to nature and radiation, the adaptive mechanisms can initiate and protect the plant. We have walked a little on this road and established a few positive impacts on various aspects of plants such as abiotic and biotic stress tolerance, yield attributes, vegetative growth etc. Although much research has been done in this particular matter and some mechanisms, and pathways have also been discovered but there is a lot more to unreveal like the exact mode of action of priming mechanisms, molecular explanation of various pathways involved in different acclimatisation of plants at a different intensity or UV types.

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UV radiation: plant responses and an in-depth mechanism of sustainability under climatic extremities

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Abstract

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Introduction

Ultraviolet Radiation

Plant Damage By Uv Radiation

Effect On Plant Morphology

Effect Of Uv On Photosystems Of Plants

Perceiving Uv In Plants Through Photoreceptors

Influence Of Uv Radiation On Plant Growth

Uv Signaling For Photomorphogenesis

Uv Radiation And Oxidative Stress

Uv Irradiation And Stress Tolerance

Protective Adaptation By Plants To Escape Uv Radiation

Priming With Uv Radiations And Their Positive Responses

Conclusion

References

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