Cent. Eur. J. Biol.• 6(5) • 2011 • 821-828 DOI: 10.2478/s11535-011-0059-z Central European Journal of Biology The impact of red and blue light-emitting diode illumination on radish physiological indices Research Article Giedrė Samuolienė*, Ramūnas Sirtautas, Aušra Brazaitytė, Jurga Sakalauskaitė, Sandra Sakalauskienė, Pavelas Duchovskis Institute of Horticulture, Lithuanian Research Centre for Agriculture and Forestry, Lt-54333 Babtai, Kaunas, Lithuania Received 20 January 2011; Accepted 19 May 2011 Abstract: The objective was to evaluate the effect of different combinations of red (638 nm) and blue (455 nm) light produced by solid-state light-emitting diodes (LEDs) on physiological indices (net assimilation rate, hypocotyl-to-leaf ratio, leaf area, leaf dry weight, hypocotyl length and diameter, plant length, developed leaves), variation of photosynthetic pigments and non-structural carbohydrates in radish (Raphanus sativus L., var. ‘Faraon’). Lighting experiments were performed under controlled conditions (total PPFD - 200 µmol m-2 s-1; 16 h photoperiod; 14/18°C night/day temperature). The LED conditions: 638 nm; 638 + 5% 455 nm; 638 + 10% 455 nm; 638 + 10% 455 + 731 nm; 638 + 10% 455 + 731 + 669 nm. Our results showed that radishes grown under red (638 nm) alone were elongated, and the formation of hypocotyl was weak. The net assimilation rate, hypocotyl–to–leaf ratio, and leaf dry weight also were low due to the low accumulation of photosynthetic pigments and non-structural carbohydrates in leaves. The supplemented blue (455 nm) light was necessary for the non–structural carbohydrates distribution between radish storage organs and leaves which resulted in hypocotyl thickening. Red alone (638 nm) or in combination with far–red (731 nm), or red669 for radish generative development was required. Keywords: Carbohydrates • Chlorophyll • Hypocotyl • Leaf dry weigh • Root:shoot ratio © Versita Sp. z o.o. 1. Introduction Light quality and quantity in the growth environment through photosynthesis and photomorphogenetic photoreceptors strongly influences plant development and physiology [1-4]. One of the problems of controlled environment crop production is the distribution of adequate light for plant growth. Some electrically light sources lack specific wavelengths required for natural plant photomorphogenesis. Generally fluorescent, highpressure sodium, and metal halide lamps are used for illumination in greenhouses or in other controlled growth environment [5]. These lamps vary in terms of spectral quality, which can result in differences in plant growth and morphology [6]. Recently most studies are based on solid-state light-emitting diodes (LEDs) lighting [7] as a potential light source for successful growth [8] and to determine if the requirements for plant photosynthesis and photomorphogenesis are met [9,10]. It is known that the main role in photomorphogenesis, through the photoreceptors, needs blue light and thus it is essential for the growth and development of higher plants [8,10]. In photomorphogenesis the attention falls on the red far-red light-absorbing phytochromes, the blue, UV-A light-absorbing cryptochromes and blue light photoreceptors phototropins [4,11]. Photosynthetic receptors, chlorophylls and carotenoids absorb at red and blue light [8]. It is important to understand the light competition between photosynthetic and photomorphogenetic pigments and related processes. According to literature data [9], plants can complete a life cycle under red LEDs alone, but supplementing red light with blue light promotes dry matter production in several plant species [2,10], including radish [1]. The trend of higher biomass production and photosynthetic capacity due to blue and red light combinations was observed in many papers [2,9,10]. The effect of light quantity on plant development and physiological indices is also important. Goins and co–workers [2] ascertained that wheat grown under red LEDs supplemented with blue light from fluorescent lamps (BF) were larger and greater amounts of seed were produced. According to * E-mail: [email protected] 821 Unauthenticated Download Date | 6/17/17 11:49 PM The impact of red and blue light-emitting diode illumination on radish physiological indices Yorio et al. [1] data, 10% of blue light from BF lamps and red LEDs were still insufficient for achieving maximal growth for radish, lettuce and spinach. Hogewoning et al. [8] found that 7% of blue light during cucumber growth is qualitatively required for normal photosynthetic functioning and quantitatively mediates leaf responses resembling those to irradiance intensity. Several studies have shown that biochemical properties of photosynthesis (chlorophyll a, b contents and ratios) are positively influenced by blue light [8]. Dougher and Bugbee [12] reported that with regard to morphogenesis, dicotyledonous species tend to be more sensitive to blue light than monocotyledonous species. The regulation of photosynthesis is associated with carbohydrate accumulation in leaves, [13] but there still is not much literature data associated with light treatments. With concern to its economical importance, radish (Raphanus sativus L.) has attracted the attention of researchers studying whole plant responses to environmental conditions [14]. A short growth cycle, generally day–length neutral, regularly shaped leaves, distribution of biomass and a division between leaves and a below–ground storage organ (hypocotyl) make radish a convenient experimental plant. It is useful as a model crop for studying developmental characteristics and for applications between major plant organs. The aim of this study was to evaluate the effect of different combinations of red and blue light produced by LEDs on physiological indices, variation of photosynthetic pigments and non-structural carbohydrates in radish (Raphanus sativus L., ‘Faraon’). 2. Experimental Procedures 2.1 Chemicals Raffinose, sucrose, fructose, mannose and glucose were obtained from Sigma-Aldrich (Germany). Acetone was obtained from Merck (Germany). Calcium carbonate (CaCO3) was obtained from Lach-Ner (Czech Republic). All standards and samples for HPLC were filtered through 0.25 µm syringe filters (Albet®, Germany). 2.2 Growth conditions and plant material Lighting experiments were performed in the phytotron chambers, under controlled environmental conditions. The originally designed [15] light emitting diode based lighting units, consisted of commercially available wavelengths: red component (638 nm, delivered by AlGaInP LEDs LuxeonTM type LXHL-MD1D, Lumileds Lighting, USA) and additional blue (445 nm, LuxeonTM type LXHL-LR5C, Lumileds Lighting, USA), red (669 nm, L670-66-60, Epitex, Japan), and far red (731 nm, L735-05-AU, Epitex, Japan) LEDs were used. The LED combinations are presented in Table 1, red (638 nm) light treatment was used as a control. Total photosynthetic photon flux density of about 200 µmol m-2 s-1 and 16 h photoperiod were maintained in LED treatments. The night/day temperature was maintained at 14/18°C. Radishes (Raphanus sativus L., ‘Faraon’) were grown in peat (pH ≈ 7, accuracy ±0.01 pH units) substrate under the light conditions described in Table 1. The amount of nutrients (mg/l) in the substrate was as follows: N 60-80, P 30, K 140-180, Ca 200-300, Mg 40-60. Electrical conductivity varied between 1.0 and 2.5 mS cm-1 (±0.03 mS cm-1). Plants were fertilized with 0.2% ammonium nitrate solution once a week. Analyses were performed 25 days after germination when rhizocarp was formed (vegetative state); and 50 days after germination when the formation of generative organs was observed (the differentiation of apical meristems started). 2.3 Determination carbohydrates of non–structural Raffinose, sucrose, fructose, mannose and glucose were measured by high performance liquid chromatography (HPLC) method. About 1 g of fresh plant tissue (leaves or hypocotyl) was ground and diluted with +70ºC 4 ml double distilled water. The extraction was carried out for 24 h. The samples were filtered using cellulose acetate (pore diameter 0.25 µm) syringe filters. The analyses were performed on Shimadzu HPLC (Japan) chromatograph with refractive index detector (RID 10A), oven temperature was maintained at +80ºC. Separation of carbohydrates was performed with a Shodex SC-1011 column (300 x 4.6 mm) (Japan), mobile phase – double distilled water. The sensitivity of the HPLC method was established using a method validation protocol [16]. Photosynthetically active flux density, µmol m-2 s-1 Treatments Blue 445 nm R Red 638 nm Far–red 731 nm 10 4 200 R+5B 5% 10 190 R+10B 10% 20 180 R+10B+FR 10% 20 176 R+10B+FR+R669 10% 20 166 Table 1. Red 669 nm 4 The light-emitting diode combinations and photon flux densities 822 Unauthenticated Download Date | 6/17/17 11:49 PM G. Samuolienė et al. 2.4 Determination of photosynthetic pigments About 0.2 g of fresh leaf tissue was ground with 0.5 g CaCO3, washed with pure acetone, and filtered through a cellulose filter. The sample was diluted to 50 ml with 100% acetone. Chlorophyll a, b and carotinoids were measured by the specrophotometric method of Wetshtein [17]. The spectrum of photosynthetic pigments was measured at 440.5 nm, 662 nm and 644 nm respectively. 2.5 Physiological indices The development levels and growth parameters of plants were evaluated. The net assimilation rate (NAR) of a plant is defined as its growth rate per unit leaf area for any given time period (day). It can be calculated as: NAR (g cm-2 d-1)=(1/ LA)(dW/dt), where LA is leaf area (cm2) and dW/dt is the change in plant dry mass per unit time. Root–to–shoot ratio - is the distribution of biomass between the below-ground storage organ (hypocotyl) and the shoot (leaves). In this case it was expressed as hypocotyl–to–leaf ratio. The leaf area of the radish was measured by “WinDias” leaf area meter (Delta-T Devices Lts, UK). Leaves and hypocotyls were dried in a drying oven at 105°C for 24 h to determine the dry mass. Development, % 2.6 Statistical analysis The analyses were performed in seven (biometrical measurements) or five (analytical measurements) replications and data analysis was processed using oneway analysis of variance Anova, the Fisher’s LSD test to trial mean at the confidence level P = 0.05. The standard deviation of mean to express some carbohydrate values was used. 3. Results After 25 days, radishes grown under red (R) light alone were elongated (Table 2). The length of the hypocotyl and the total plant length differed significantly from the mean, 33% and 19% respectively. Increasing the blue (B) light fraction from 0% to 10% resulted in significantly decreased plant length (both hypocotyl and leaves). The addition of far–red (FR) or FR together with red 669 nm (R669) light component resulted in significantly increased hypocotyl diameter. 50 days growth duration and LED treatment resulted in developmental differences. The differentiation of apical meristems and the formation of generative structures were observed in radishes grown under R alone (36.7%) or under R in combination with FR The length of hypocotyl, cm The diameter of hypocotyl, cm R 2.99B 0.50A 15.6B 3.9 R+5B 2.27 1.79 14.7 4.0 R+10B 1.74 1.45 10.7 3.3A R+10B+FR 1.86 1.76B 12.4 4.0 R+10B+FR+R669 1.94 2.12 12.9 4.1B 0.38 0.22 0.9 0.2 Treatments Total plant length, cm The number of developed leaves 25 days after germination Vegetative state, 100 % LSD 05 A B B B A 50 days after germination Vegetative, 63.3 % R 3.00 2.06A 15.0 4.3 Vegetative, 100.0 % R+5B 2.71 3.13 16.8 5.3B Vegetative, 100.0 % R+10B 2.69 2.54 14.0 4.6 Vegetative, 62.5 % R+10B+FR 2.83 2.91 15.0 4.7 Vegetative, 70.0 % R+10B+FR+R669 2.76 2.73 13.2A 5.0 0.22 0.26 1.1 0.5 LSD 05 B B Generative, 36.7 % R 2.78 0.64 11.5 4.8 Generative, 37.5 % R+10B+FR 1.82 1.42 12.4 5.6 Generative, 30.0 % R+10B+FR+R669 2.38 1.41 11.9 5.2 0.60 0.42 1.8 1.1 LSD 05 Table 2. A Growth parameters of radish plants. The values with the same letters are not significantly different with P≤0.05 (n=7). 823 Unauthenticated Download Date | 6/17/17 11:49 PM The impact of red and blue light-emitting diode illumination on radish physiological indices (37.5%), or with FR and R669 (30.0%). Radishes grown under R with 5% or 10% of B remained in a vegetative state. Plants grown under R in combination with 5% B were still elongated. However, these conditions were sufficient for the formation of hypocotyl (Table 2). Significantly thinner hypocotyl in both vegetative and generative stages were formed under R lighting alone. As there were no significant differences in the length of hypocotyl, the significant decrease of total plant length in a vegetative stage was observed under lighting combination with R669, due to shorter leaves. A significantly negative effect of R alone was observed on assimilative indices of radish (Table 3). Under these lighting conditions, after 25 days of treatment, NAR and hypocotyl–to–leaf ratio was significantly lower. Moreover after 50 days of treatment, a significant decrease in leaf area was observed. The significant decrease in leaf area under R in combination with 10% of B, after both 25 and 50 days duration was observed, too. A significant positive effect of R with 5% of B, and R with 10% of B supplemented by FR on assimilative indices was noticed. After 50 days of treatment these spectral conditions also had a significant positive effect on leaf dry weight accumulation. However the addition of R669 resulted in significant decrease in leaf area and leaf dry Treatments Development Net assimilation rate, g cm-2 d-1 weight of radishes which remained in a vegetative state (Table 3). After 50 days of treatment in the generative stage, the NAR decreased 4.5 and 3.6 times under R alone or in combination with 10% B, FR and R669, and under R with 10% B, FR, respectively. However under R with 10% B, FR treatment, the NAR in both the vegetative and the generative stages were significantly higher due to significantly bigger leaf area. The significant decrease of chlorophyll (Chl) a to b ratio due to significant decrease of Chl a and increase of Chl b content after 25 days of treatment under R alone in radish leaves was observed (Table 4). Increasing the blue light fraction from 0% to 10% and the addition of far–red light normalized the Chl a to b ratio (3.02 – 3.37). However, only R+10B+FR+R669 LED spectral combination lead to the significant increase of photosynthetic pigment ratio and content. Such affect lasted for 50 days under the above-mentioned spectral combination treatment for radishes which remained in a vegetative state. The addition of FR light had a positive effect on radish development rate (37.5%) (Table 2) and lead to significant increase of Chl a to b ratio, Chl a and carotenoid content (Table 4). Radishes grown under R alone or R supplemented with 5% of blue light for 25 days accumulated a Hypocotyl–to–leaf ratio Leaf area, cm-2 Leaf dry weight, g 25 days after germination R 0.009 0.22A 90.5 0.19 R+5B 0.020 1.04 112.1B 0.25 0.018 0.83 65.9A 0.30B R+10B+FR 0.015 1.04 92.1 0.19 100.0 0.20 11.2 0.09 A Vegetative state R+10B R+10B+FR+R669 0.019 1.54 LSD 05 0.004 0.27 R 0.027A 2.34A 83.0A 0.28 R+5B 0.050 2.87 148.7 0.46B 0.033 3.66 72.2A 0.27A R+10B+FR 0.058 B 3.73 112.4 0.38B R+10B+FR+R669 0.026A 3.78 73.2A 0.25A LSD 05 0.011 0.73 15.0 0.06 R A B 50 days after germination R+10B Vegetative state B B B 0.006 0.96 38.5 0.21 0.016B 1.71 76.1B 0.26 R+10B+FR+R669 0.006 2.67 52.2 0.17 LSD 05 0.006 0.73 16.9 0.07 R+10B+FR Table 3. Generative state A B Assimilative indices of radish. The values with the same letters are not significantly different with P≤0.05 (n=7). 824 Unauthenticated Download Date | 6/17/17 11:49 PM G. Samuolienė et al. Content of chlorophyll b, mg g-1 FM Content of carotenoids, mg g-1 FM The chlorophyll a to b ratio Content of chlorophyll a, mg g-1 FM R 1.97A 0.54A 0.28B 0.17A R+5B 3.37 0.74 0.22 0.22 Treatment Development 25 days after germination 3.33 0.64 0.18 0.22 R+10B+FR 3.02 0.66 0.22 0.19A R+10B+FR+R669 3.60B 0.90B 0.25 0.26B LSD 05 0.34 0.05 0.03 0.02 R 2.47B 0.35 0.15 0.13 1.91 0.29 0.14 0.15 1.71 0.29 0.19 0.14 R+10B+FR 1.44 0.24 0.18 0.13A R+10B+FR+R669 1.79 0.34B 0.23B 0.20B LSD 05 0.23 0.04 0.05 0.02 R 2.25 A 0.33 0.16 0.16A 2.44B 0.50B 0.15 0.21B R+10B+FR+R669 2.30 0.40 0.17 0.19 LSD 05 0.04 0.02 0.08 0.01 Vegetative state R+10B A A 50 days after germination R+5B Vegetative state R+10B A A R+10B+FR Table 4. Generative state A The distribution of photosynthetic pigments in radish leaves. The values with the same letters are not significantly different with P≤0.05 (n=5). significant lower amount of total carbohydrates in both hypocotyl and leaves (Table 5). 10% of blue light lead to an increased fructose content in the whole plant. The significant biggest content (about 2 times) of total non– structural carbohydrates in both hypocotyl and leaves was detected under R+10B LED spectral combination. In contrast to leaves, a significant higher content of monocarbohydrates and a lower amount of sucrose in hypocotyl were detected. The addition of FR spectral component influenced better accumulation of total carbohydrates in leaves, but the total amount in the hypocotyl was significantly lower. The R+10B+FR+R669 spectral composition conditioned the opposite distribution of carbohydrates, and the statistically bigger amounts of glucose and fructose were determined to be in the hypocotyl. The same effect was found with different LED lighting sustained for 50 days of treatment for radishes which remained in a vegetative state. Also, the total amount of hypocotyl carbohydrates increased from 27.0% to 49.8%, especially due to bigger contents of monocarbohydrates. The increase of total leaf carbohydrates was due to an increased accumulation of both hexoses, especially of sucrose. The distribution of non–structural carbohydrates changed drastically in plants grown under R alone, which started to form generative organs. A significant increase of total content of carbohydrates, especially of sucrose, in the hypocotyl and leaves was observed, 1.7 and 2 times respectively (Table 5). 4. Discussion Some plants systems quite easy accept light-driven manipulation [3]. As photosynthesis is the basis for primary metabolite production, light also modulates several metabolic pathways altering gene expression which generates a physiological response [4,11]. Pursuant to literature data [2], radishes grown under R alone were elongated, the formation of hypocotyl was weak (Table 2), the net assimilation rate and hypocotyl– to–leaf ratio was low (Table 3), leaf dry weight also was low due to the low accumulation of photosynthetic pigments (Table 4) and non–structural carbohydrates in leaves (Table 5). A wide variety of plant responses, such as procession of flowering [18] or morphological alterations [3], is the result of light activated phytochromes or cryptochromes action. Besides, changes in the 825 Unauthenticated Download Date | 6/17/17 11:49 PM Treatment Organs Development The impact of red and blue light-emitting diode illumination on radish physiological indices Raffinose, mg g-1, FM Sucrose, mg g-1, FM Glucose, mg g-1, FM Mannose, mg g-1, FM Fructose, mg g-1, FM Total, mg g-1, FM 1.78A 10.33A 25 days after germination 5.71B 2.57A R+5B 1.25A 2.21A 6.71B 10.18A 1.80A 12.14B 7.04B 20.97B 1.21A 8.01B 4.63A 13.85A 2.20 9.54 5.90 17.64B 0.02 0.01 0.01 0.02 2.57 1.64 A 0.29 4.51A 3.09A 1.05A 0.15 0.50B 8.20A 15.68B 2.38B 0.16B 0.53B 18.75B 8.84B 5.34A 3.03B 0.12A 0.63B 17.96B R+10B+FR+R669 4.34A 2.67A 1.35A 0.16B 0.43A 8.96A LSD 05 0.03 0.03 0.01 0.00 0.00 0.04 hypocotyl R R+10B R+10B+FR+R669 LSD 05 R R+5B A Vegetative state R+10B+FR 3.41A leaves R+10B A R+10B+FR 0.27±0.00 B B A 50 days after germination 2.46A 11.69A 4.06A 18.21A R+5B 4.27 10.59 A 2.95 17.81A 8.72B 14.94B 5.07B 28.73B 3.62A 9.20A 6.10B 22.73A 9.30B 21.70B 4.12A 35.12B 0.01 0.04 0.01 0.05 1.77 B 2.04 17.08A 0.31A 0.87A 14.61A 0.55A 21.43B hypocotyl R R+10B R+10B+FR+R669 LSD 05 R R+5B 3.72±0.00 Vegetative state R+10B+FR 10.92 A R+10B+FR R+10B+FR+R669 LSD 05 R R+10B+FR R+10B+FR+R669 LSD 05 Table 5. leaves LSD 05 R Generative state R+10B+FR+R669 hypocotyl 1.87±0.01 R+10B+FR A 2.35 A 13.43A leaves R+10B A 0.08±0.00 B 19.49B 1.39A 24.47B 3.14B 0.23A 1.48B 29.32B 13.24A 3.03B 0.79B 1.32 18.38A 0.04 0.00 0.00 0.01 0.04 15.84 9.85 0.18±0.00 A 2.63 30.36B 4.45A 14.44B 6.79B 25.67A 4.89A 13.80B 6.30B 24.99A B A 0.02 0.01 26.86B 5.22±0.01 0.02 0.02 0.13A 1.97B 34.18B 14.82A 2.86B 1.85A 19.54A 14.13A 1.70B 1.84A 17.67A 0.09 0.01 0.01 0.03 The distribution of non–structural carbohydrates in radish hypocotyl and leaves. The values with the same letters are not significantly different with P≤0.05 (mean ± SD; n=5). 826 Unauthenticated Download Date | 6/17/17 11:49 PM G. Samuolienė et al. partitioning of photoassimilates may be involved by photomorphogenetic responses [4]. Acording to our data, R light was sufficient for flowering induction, 36.7% of plants formed the generative organs (Table 2). Acording to Goins and co–workers [2] data, wheat grown under red LEDs alone also were elongated, shoot dry matter and net leaf photosynthesis rate were decreased compared to plants grown under supplemental blue fluorescent light (from 1% to 10%). Yorio and co–workers [1] also stated that radish, lettuce, and spinach grown under red LEDs alone accumulated significantly less total dry weight than when grown under cool-white fluorescent light or R LEDs supplemented with 10%B fluorescent light. According to our data, the addition of 5% blue light to red LEDs improved the hypocotyl thickening (Table 2), and leaf area significantly increased (Table 3), but plants were still elongated and there was no positive effects on photosynthetic pigment (Table 4) and non–structural carbohydrate (Table 5) accumulation. In this case, B and R light radiation was insufficient for photosynthetic and photomorphogenetic responses, although chlorophylls and carotenoids accumulated abundantly in plant tissue as compared to photomorphogenetic photoreceptors [11]. Hogewoning and co–workers [8] stated that 7% of B light to R LEDs was sufficient enough to prevent any overt dysfunctional photosynthesis. The authors noticed that an increase of photosynthetic capacity associated with an increase in leaf mass per unit leaf area, Chl content per area continued to increase with increasing B percentage up to 50%. In agreement with literature data [8], our results demonstrated that Chl content on leaf dry weight basis was not affected by blue light percentage (Tables 2 and 3). In contrary to Yorio and co– workers [1], radish grown under R LEDs supplemented with 10% B LEDs accumulated significantly higher leaf dry weight (Table 3) due to significantly increased leaf carbohydrates contents (Table 5). Moreover, in contrary to wheat development [2], the blue light did not influence the generative growth of radish and all plants remained in a vegetative state (Table 2). According to our data, supplementation of R and B LEDs with FR or R669 components had a positive effect on radish development, and the significant hypocotyl thickening was observed (Table 2). Such transition to flowering might be influenced by integrated phytochromes and cryptochromes action. As leaves filter out B and R light and transmit far-red, there is a reduction in B and R light leading to the decrease of photosynthetically active radiation and light competitiveness [19]. So, due to light quality and interaction between photomorphogenetic and photosynthetic receptors, plants can change the pattern of development during vegetation. FR light significantly positively affected the assimilative indices of radish (Table 3). Parks et al. [20] stated that a decrease in stem elongation and an increase in leaf expansion light activated phytochrome action. Dillard and co–workers [21] also noticed that radish grown under red, green, and blue supplemented by FR light partitioned significantly more biomass than plants grown without FR light. The significant hypocotyl–to–leaf ratio under R+10%B+FR+R669 (Table 3) was observed and it correlated with carbohydrates distribution in the hypocotyl and leaves of radish (Table 5). Thus the response of radish growth and biomass partitioning was linked to patterns of non–structural carbohydrate distribution between plant organs. As it is known, the sufficient doses of photosynthetically active radiation are crucial for the formation of assimilates and thus the accumulation of biomass [9,22]. But spectral composition of light controls formative processes and may initiate developmental processes during plant growth [2,23]. Moreover, as photomorphogenesis is the process by which light regulates many plant development aspects, so some metabolites which are produced by photosynthesis are used in photomorphogenesis [11]. Our results showed that for radish generative development, red alone (638 nm) or combination with FR, or R669 was required (Table 2). However, for hypocotyl formation a minimum 5% of blue light was necessary (Table 2). Besides, the indirect effect of light spectral composition on hypocotyl growth via formation and distribution of assimilates was observed. The discussed data is in agreement with Ballaré and Casal [4], where they state that stem growth and dry matter partitioning are correlated, notwithstanding changes in dry matter allocation are not entirely the result of changes in stem elongation. This indicates that photomorphogenetic responses, depend on the interaction of photosynthetic and photomorphogenetic photoreceptors. The co-operation of both signals, chlorophyll and phytochrome, cryptochrome action, control the morphology, development and distribution of photoassimilates. Also an intense accumulation of mannose in radish leaves inhibited sucrose synthesis (Table 5). In agreement with Harris et al. [24], mannose perturbs normal metabolic processes of glucose, inhibits sucrose synthesis, moreover it distinguishes in photosynthetic toxicity. Whereas high levels of raffinose, synthesized from sucrose, are present under normal and/or stressful conditions act as an antioxidant and together with glucose, fructose and sucrose efficiently scavenge hydroxyl radicals [25]. In summary, blue light has been shown to be necessary for non–structural carbohydrates distribution between radish storage organs and leaves resulting in hypocotyl thickening. 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