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Blue light dose–responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light
Abstract
The blue part of the light spectrum has been associated with leaf characteristics which also develop under high irradiances. In this study blue light dose–response curves were made for the photosynthetic properties and related developmental characteristics of cucumber leaves that were grown at an equal irradiance under seven different combinations of red and blue light provided by light-emitting diodes. Only the leaves developed under red light alone (0% blue) displayed dysfunctional photosynthetic operation, characterized by a suboptimal and heterogeneously distributed dark-adapted Fv/Fm, a stomatal conductance unresponsive to irradiance, and a relatively low light-limited quantum yield for CO2 fixation. Only 7% blue light was sufficient to prevent any overt dysfunctional photosynthesis, which can be considered a qualitatively blue light effect. The photosynthetic capacity (Amax) was twice as high for leaves grown at 7% blue compared with 0% blue, and continued to increase with increasing blue percentage during growth measured up to 50% blue. At 100% blue, Amax was lower but photosynthetic functioning was normal. The increase in Amax with blue percentage (0–50%) was associated with an increase in leaf mass per unit leaf area (LMA), nitrogen (N) content per area, chlorophyll (Chl) content per area, and stomatal conductance. Above 15% blue, the parameters Amax, LMA, Chl content, photosynthetic N use efficiency, and the Chl:N ratio had a comparable relationship as reported for leaf responses to irradiance intensity. It is concluded that blue light during growth is qualitatively required for normal photosynthetic functioning and quantitatively mediates leaf responses resembling those to irradiance intensity.
Introduction
Plant development and physiology are strongly influenced by the light spectrum of the growth environment. The underlying mechanisms of the effect of different growth spectra on plant development are not known in detail, although the involvement of photoreceptors has been demonstrated for a wide range of spectrum-dependent plant responses. Cryptochromes and phototropins are specifically blue light sensitive, whereas phytochromes are more sensitive to red than to blue (Whitelam and Halliday, 2007). Blue light is involved in a wide range of plant processes such as phototropism, photomorphogenesis, stomatal opening, and leaf photosynthetic functioning (Whitelam and Halliday, 2007). At the chloroplast level, blue light has been associated with the expression of ‘sun-type’ characteristics such as a high photosynthetic capacity (Lichtenthaler et al., 1980). Most studies assessing blue light effects on the leaf- or whole-plant level have either compared responses to a broad-band light source with responses to blue-deficient light (e.g. Britz and Sager, 1990; Matsuda et al., 2008), or compared plants grown under blue or a combination of red and blue light with plants grown under red light alone (e.g. Brown et al., 1995; Bukhov et al., 1995; Yorio, 2001; Matsuda et al., 2004; Ohashi et al, 2006). Overall there is a trend to higher biomass production and photosynthetic capacity in a blue light-containing irradiance. Before the development of light-emitting diodes (LEDs) that were intense enough to be used for experimental plant cultivation (Tennessen et al., 1994), light sources emitting wavelengths in a broader range than strictly the red (i.e. 600–700 nm) or blue (i.e. 400–500 nm) region were often used (e.g. Voskresenskaya et al., 1977). Other wavelengths can interact with blue light responses. For example, green light has been reported to antagonize some blue light responses, such as stomatal opening and inhibition of hypocotyl elongation in seedlings (Folta and Maruhnich, 2007). The blue light enhancement effect on photosynthetic capacity appears to be greater when using combinations of red and blue light produced by LEDs than when broad-band light is made deficient in blue by a filter (e.g. for spinach compare Matsuda et al., 2007 and 2008). This raises the question of whether plants exposed to red light alone suffer a spectral ‘deficiency’ syndrome, which may be reversed by blue light as well as by longer wavelengths.
Poorter et al. (2010) stress the importance of dose–response curves for quantitative analysis of the effects of environmental factors on plant phenotypes, allowing a better understanding of plant–environment interactions than the comparison of two treatments only. It is not clear whether the enhancement effect of blue light on leaf photosynthetic capacity is a qualitative threshold response or a quantitative progressive response, or a combination of both. Only few specific processes in leaves have been identified as quantitative blue light responses, such as chloroplast movement (Jarillo et al., 2001) and stomatal conductance (Sharkey and Raschke, 1981). Matsuda et al. (2007) found a higher photosynthetic capacity for spinach leaves grown under 300 μmol m−2 s−1 mixed red/blue irradiance containing 30 μmol m−2 s−1 blue than for leaves grown under red alone. A higher blue light fraction did not yield a significant further enhancement in light-saturated assimilation (Amax), which may be interpreted as a qualitative blue light effect. However, a quantitative blue light effect at quantum fluxes <30 μmol m−2 s−1 cannot be excluded.
A diverse choice of LEDs powerful enough for use as a growth irradiance source in controlled environments has recently become available (e.g. Massa et al., 2008). These LEDs allow the effect of light quality to be investigated independently of the amount of photosynthetic irradiance. LED illumination has been used here to study the response curves of a range of parameters related to leaf photosynthesis of plants that were grown at an irradiance with a proportion of blue light ranging from 0% to 100%. A range of other leaf characteristics important for the functioning of photosynthesis, such as stomatal development and behaviour, leaf mass per area (LMA), and the content of N, pigments, and carbohydrates, were also determined. The spectra and the extent of variation in the ratio of red and blue irradiance that can be achieved with LED lighting are dissimilar to field conditions. However, the responses of leaves to these unnatural environments provides the possibility to unravel the complex developmental and functional interactions that normally occur in the natural light environment.
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Abstract
The blue part of the light spectrum has been associated with leaf characteristics which also develop under high irradiances. In this study blue light dose–response curves were made for the photosynthetic properties and related developmental characteristics of cucumber leaves that were grown at an equal irradiance under seven different combinations of red and blue light provided by light-emitting diodes. Only the leaves developed under red light alone (0% blue) displayed dysfunctional photosynthetic operation, characterized by a suboptimal and heterogeneously distributed dark-adapted Fv/Fm, a stomatal conductance unresponsive to irradiance, and a relatively low light-limited quantum yield for CO2 fixation. Only 7% blue light was sufficient to prevent any overt dysfunctional photosynthesis, which can be considered a qualitatively blue light effect. The photosynthetic capacity (Amax) was twice as high for leaves grown at 7% blue compared with 0% blue, and continued to increase with increasing blue percentage during growth measured up to 50% blue. At 100% blue, Amax was lower but photosynthetic functioning was normal. The increase in Amax with blue percentage (0–50%) was associated with an increase in leaf mass per unit leaf area (LMA), nitrogen (N) content per area, chlorophyll (Chl) content per area, and stomatal conductance. Above 15% blue, the parameters Amax, LMA, Chl content, photosynthetic N use efficiency, and the Chl:N ratio had a comparable relationship as reported for leaf responses to irradiance intensity. It is concluded that blue light during growth is qualitatively required for normal photosynthetic functioning and quantitatively mediates leaf responses resembling those to irradiance intensity.
Introduction
Plant development and physiology are strongly influenced by the light spectrum of the growth environment. The underlying mechanisms of the effect of different growth spectra on plant development are not known in detail, although the involvement of photoreceptors has been demonstrated for a wide range of spectrum-dependent plant responses. Cryptochromes and phototropins are specifically blue light sensitive, whereas phytochromes are more sensitive to red than to blue (Whitelam and Halliday, 2007). Blue light is involved in a wide range of plant processes such as phototropism, photomorphogenesis, stomatal opening, and leaf photosynthetic functioning (Whitelam and Halliday, 2007). At the chloroplast level, blue light has been associated with the expression of ‘sun-type’ characteristics such as a high photosynthetic capacity (Lichtenthaler et al., 1980). Most studies assessing blue light effects on the leaf- or whole-plant level have either compared responses to a broad-band light source with responses to blue-deficient light (e.g. Britz and Sager, 1990; Matsuda et al., 2008), or compared plants grown under blue or a combination of red and blue light with plants grown under red light alone (e.g. Brown et al., 1995; Bukhov et al., 1995; Yorio, 2001; Matsuda et al., 2004; Ohashi et al, 2006). Overall there is a trend to higher biomass production and photosynthetic capacity in a blue light-containing irradiance. Before the development of light-emitting diodes (LEDs) that were intense enough to be used for experimental plant cultivation (Tennessen et al., 1994), light sources emitting wavelengths in a broader range than strictly the red (i.e. 600–700 nm) or blue (i.e. 400–500 nm) region were often used (e.g. Voskresenskaya et al., 1977). Other wavelengths can interact with blue light responses. For example, green light has been reported to antagonize some blue light responses, such as stomatal opening and inhibition of hypocotyl elongation in seedlings (Folta and Maruhnich, 2007). The blue light enhancement effect on photosynthetic capacity appears to be greater when using combinations of red and blue light produced by LEDs than when broad-band light is made deficient in blue by a filter (e.g. for spinach compare Matsuda et al., 2007 and 2008). This raises the question of whether plants exposed to red light alone suffer a spectral ‘deficiency’ syndrome, which may be reversed by blue light as well as by longer wavelengths.
Poorter et al. (2010) stress the importance of dose–response curves for quantitative analysis of the effects of environmental factors on plant phenotypes, allowing a better understanding of plant–environment interactions than the comparison of two treatments only. It is not clear whether the enhancement effect of blue light on leaf photosynthetic capacity is a qualitative threshold response or a quantitative progressive response, or a combination of both. Only few specific processes in leaves have been identified as quantitative blue light responses, such as chloroplast movement (Jarillo et al., 2001) and stomatal conductance (Sharkey and Raschke, 1981). Matsuda et al. (2007) found a higher photosynthetic capacity for spinach leaves grown under 300 μmol m−2 s−1 mixed red/blue irradiance containing 30 μmol m−2 s−1 blue than for leaves grown under red alone. A higher blue light fraction did not yield a significant further enhancement in light-saturated assimilation (Amax), which may be interpreted as a qualitative blue light effect. However, a quantitative blue light effect at quantum fluxes <30 μmol m−2 s−1 cannot be excluded.
A diverse choice of LEDs powerful enough for use as a growth irradiance source in controlled environments has recently become available (e.g. Massa et al., 2008). These LEDs allow the effect of light quality to be investigated independently of the amount of photosynthetic irradiance. LED illumination has been used here to study the response curves of a range of parameters related to leaf photosynthesis of plants that were grown at an irradiance with a proportion of blue light ranging from 0% to 100%. A range of other leaf characteristics important for the functioning of photosynthesis, such as stomatal development and behaviour, leaf mass per area (LMA), and the content of N, pigments, and carbohydrates, were also determined. The spectra and the extent of variation in the ratio of red and blue irradiance that can be achieved with LED lighting are dissimilar to field conditions. However, the responses of leaves to these unnatural environments provides the possibility to unravel the complex developmental and functional interactions that normally occur in the natural light environment.
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