Far-red light effects on plant photosynthesis: from short-term enhancements to long-term effects of artificial solar light.

BACKGROUND AND AIMS: Long-term exposure over several days to Far-Red (FR) increases leaf expansion, while short-term exposure (minutes) may enhance the PSII operating efficiency (ϕPSII). The interaction between these responses at different time scales, and their impact on photosynthesis at whole-plant level is not well understood. Our study aimed to assess the effects of FR in an irradiance mimicking the spectrum of sunlight (referred to as artificial solar irradiance) both in the long and short-term, on whole-plant CO2 assimilation rates and in leaves at different positions in the plant. METHODS: Tomato (Solanum lycopersicum) plants were grown under artificial solar irradiance conditions with either a severely reduced or normal fraction of FR(SUN(FR-) vs. SUN). To elucidate the interplay between the growth light treatment and the short-term reduction of FR, we investigated this interaction at both the whole-plant and leaf level. At whole-plant level, CO2 assimilation rates were assessed under artificial solar irradiance with a normal and a reduced fraction of FR. At the leaf level, the effects of removal and presence of FR (0FR and 60FR) during transition from high to low light on CO2 assimilation rates and chlorophyll fluorescence were evaluated in upper and lower leaves. KEY RESULTS: SUN(FR-) plants had lower leaf area, shorter stems, and darker leaves than SUN plants. While reducing FR during growth did not affect whole-plant photosynthesis under high light intensity, it had a negative impact at low light intensity. Short-term FR removal reduced both plant and leaf CO2 assimilation rates, but only at low light intensity and irrespective of the growth light treatment and leaf position. Interestingly, the kinetics of ϕPSII from high to low light were accelerated by 60FR, with a larger effect in lower leaves of SUN than in SUN(FR-) plants. CONCLUSIONS: Growing plants with a reduced amount of FR light lowers whole-plant CO2 assimilation rates at low light intensity through reduced leaf area, despite maintaining similar leaf-level CO2 assimilation to leaves grown with a normal amount of FR. The short-term removal of FR brings about significant but marginal reductions in photosynthetic efficiency at the leaf level, regardless of the long-term growth light treatment.

Shining light on diurnal variation of non-photochemical quenching: Impact of gradual light intensity patterns on short-term NPQ over a day.

Maximal sunlight intensity varies diurnally due to the earth's rotation. Whether this slow diurnal pattern influences the photoprotective capacity of plants throughout the day is unknown. We investigated diurnal variation in NPQ, along with NPQ capacity, induction, and relaxation kinetics after transitions to high light, in tomato plants grown under diurnal parabolic (DP) or constant (DC) light intensity regimes. DP light intensity peaked at midday (470 µmol m-2 s-1) while DC stayed constant at 300 µmol m-2 s-1 at a similar 12-hour photoperiod and daily light integral. NPQs were higher in the morning and afternoon at lower light intensities in DP compared to DC, except shortly after dawn. NPQ capacity increased from midday to the end of the day, with higher values in DP than in DC. At high light ΦPSII did not vary throughout the day, while ΦNPQ varied consistently with NPQ capacity. Reduced ΦNO suggested less susceptibility to photodamage at the end of the day. NPQ induction was faster at midday than at the start of the day and in DC than in DP, with overshoot occurring in the morning and midday but not at the end of the day. NPQ relaxation was faster in DP than in DC. The xanthophyll de-epoxidation state and reduced demand for photochemistry could not explain the observed diurnal variations in photoprotective capacity. In conclusion, this study showed diurnal variation in regulated photoprotective capacity at moderate growth light intensity, which was not explained by instantaneous light intensity or increasing photoinhibition over the day and was influenced by acclimation to constant light intensity.

Crassulacean acid metabolism species differ in the contribution of C3 and C4 carboxylation to end of day CO2 fixation.

Crassulacean acid metabolism (CAM) is a photosynthetic pathway that temporally separates the nocturnal CO2 uptake, via phosphoenolpyruvate carboxylase (PEPC, C4 carboxylation), from the diurnal refixation by Rubisco (C3 carboxylation). At the end of the day (CAM-Phase IV), when nocturnally stored CO2 has depleted, stomata reopen and allow additional CO2 uptake, which can be fixed by Rubisco or by PEPC. This work examined the CO2 uptake via C3 and C4 carboxylation in phase IV in the CAM species Phalaenopsis "Sacramento" and Kalanchoe blossfeldiana "Saja." Short blackout periods during phase IV caused a sharp drop in CO2 uptake in K. blossfeldiana but not in Phalaenopsis, indicating strong Rubisco activity only in K. blossfeldiana. Chlorophyll fluorescence revealed a progressive decrease in ΦPSII in Phalaenopsis, implying decreasing Rubisco activity, while ΦPSII remained constant in phase IV in K. blossfeldiana. However, short switching to 2% O2 indicated the presence of photorespiration and thus Rubisco activity in both species throughout phase IV. Lastly, in Phalaenopsis, accumulation of starch in phase IV occurred. These results indicate that in Phalaenopsis, PEPC was the main carboxylase in phase IV, although Rubisco remained active throughout the whole phase. This will lead to double carboxylation (futile cycling) but may help to avoid photoinhibition.

Demonstration of a relationship between state transitions and photosynthetic efficiency in a higher plant.

A consequence of the series configuration of PSI and PSII is that imbalanced excitation of the photosystems leads to a reduction in linear electron transport and a drop in photosynthetic efficiency. Achieving balanced excitation is complicated by the distinct nature of the photosystems, which differ in composition, absorption spectra, and intrinsic efficiency, and by a spectrally variable natural environment. The existence of long- and short-term mechanisms that tune the photosynthetic apparatus and redistribute excitation energy between the photosystems highlights the importance of maintaining balanced excitation. In the short term, state transitions help restore balance through adjustments which, though not fully characterised, are observable using fluorescence techniques. Upon initiation of a state transition in algae and cyanobacteria, increases in photosynthetic efficiency are observable. However, while higher plants show fluorescence signatures associated with state transitions, no correlation between a state transition and photosynthetic efficiency has been demonstrated. In the present study, state 1 and state 2 were alternately induced in tomato leaves by illuminating leaves produced under artificial sun and shade spectra with a sequence of irradiances extreme in terms of PSI or PSII overexcitation. Light-use efficiency increased in both leaf types during transition from one state to the other with remarkably similar kinetics to that of F'm/Fm, F'o/Fo, and, during the PSII-overexciting irradiance, ΦPSII and qP. We have provided compelling evidence for the first time of a correlation between photosynthetic efficiency and state transitions in a higher plant. The importance of this relationship in natural ecophysiological contexts remains to be elucidated.

Sucrose and Starch Content Negatively Correlates with PSII Maximum Quantum Efficiency in Tomato (Solanum lycopersicum) Exposed to Abnormal Light/Dark Cycles and Continuous Light.

Light is most important to plants as it fuels photosynthesis and provides clues about the environment. If provided in unnatural long photoperiods, however, it can be harmful and even lethal. Tomato (Solanum lycopersicum), for example, develops mottled chlorosis and necrosis when exposed to continuous light. Understanding the mechanism of these injuries is valuable, as important pathways regulating photosynthesis, such as circadian, retrograde and light signaling pathways are probably involved. Here, we use non-targeted metabolomics and transcriptomics analysis as well as hypothesis-driven experiments with continuous light-tolerant and -sensitive tomato lines to explore the long-standing proposed role of carbohydrate accumulation in this disorder. Analysis of metabolomics and transcriptomics data reveals a clear effect of continuous light on sugar metabolism and photosynthesis. A strong negative correlation between sucrose and starch content with the severity of continuous light-induced damage quantified as the maximum quantum efficiency of PSII (Fv/Fm) was found across several abnormal light/dark cycles, supporting the hypothesis that carbohydrates play an important role in the continuous light-induced injury. We postulate that the continuous light-induced injury in tomato is caused by down-regulation of photosynthesis, showing characteristics of both cytokinin-regulated senescence and light-modulated retrograde signaling. Molecular mechanisms linking carbohydrate accumulation with down-regulation of carbon-fixing enzymes are discussed.

Phenotypic plasticity to altered apical bud temperature in Cucumis sativus: more leaves-smaller leaves and vice versa.

Many studies investigated temperature effects on leaf initiation and expansion by relating these processes to air temperature or the temperature of a specific organ (e.g. leaf temperature). In reality plant temperature is hardly ever equal to air temperature or spatially uniform. Apical bud temperature (Tbud ), for example, may greatly differ from the temperature of the rest of the plant (Tplant ) dependent on the environment. Recent research in Cucumis sativus showed that Tbud influences leaf initiation independent of Tplant . These findings trigger the question if such spatial temperature differences also influence leaf expansion and plant phenotype. In a 28 day study, we maintained temperature differences between Tbud and Tplant ranging from -7 to +8 °C using a custom-made bud temperature control system. Leaf expansion did not only depend on leaf temperature but also on the difference between bud and leaf temperature. Differences between Tbud and Tplant considerably influenced vertical leaf area distribution over the shoot: increasing Tbud beyond Tplant resulted in more and smaller leaves, while decreasing Tbud below Tplant resulted in less and larger leaves. The trade-off between leaf number and leaf area resulted in phenotypic alterations that cannot be predicted, for example, by crop models, when assuming plant temperature uniformity.

A unique approach to demonstrating that apical bud temperature specifically determines leaf initiation rate in the dicot Cucumis sativus.

MAIN CONCLUSION: Leaf initiation rate is largely determined by the apical bud temperature even when apical bud temperature largely deviates from the temperature of other plant organs. We have long known that the rate of leaf initiation (LIR) is highly sensitive to temperature, but previous studies in dicots have not rigorously demonstrated that apical bud temperature controls LIR independent of other plant organs temperature. Many models assume that apical bud and leaf temperature are the same. In some environments, the temperature of the apical bud, where leaf initiation occurs, may differ by several degrees Celsius from the temperature of other plant organs. In a 28-days study, we maintained temperature differences between the apical bud and the rest of the individual Cucumis sativus plants from -7 to +8 °C by enclosing the apical buds in transparent, temperature-controlled, flow-through, spheres. Our results demonstrate that LIR was completely determined by apical bud temperature independent of other plant organs temperature. These results emphasize the need to measure or model apical bud temperatures in dicots to improve the prediction of crop development rates in simulation models.

Continuous-light tolerance in tomato is graft-transferable.

Continuous light induces a potentially lethal injury in domesticated tomato (Solanum lycopersicum) plants. Recently, continuous-light tolerance was reported in several wild tomato species, yet the molecular mechanisms underpinning tolerance/sensitivity are still elusive. Here, we investigated from which part of the plant continuous-light tolerance originates and whether this trait acts systemically within the plant. By exposing grafted plants bearing both tolerant and sensitive shoots, the trait was functionally located in the shoot rather than the roots. Additionally, an increase in continuous-light tolerance was observed in sensitive plants when a continuous-light-tolerant shoot was grafted on it. Cultivation of greenhouse tomatoes under continuous light promises high yield increases. Our results show that to pursuit this, the trait should be bred into scion rather than rootstock lines. In addition, identifying the nature of the signal/molecule(s) and/or the mechanism of graft-induced, continuous-light tolerance can potentially result in a better understanding of important physiological processes like long-distance signaling.

Photosynthetic quantum yield dynamics: from photosystems to leaves.

The mechanisms underlying the wavelength dependence of the quantum yield for CO(2) fixation (α) and its acclimation to the growth-light spectrum are quantitatively addressed, combining in vivo physiological and in vitro molecular methods. Cucumber (Cucumis sativus) was grown under an artificial sunlight spectrum, shade light spectrum, and blue light, and the quantum yield for photosystem I (PSI) and photosystem II (PSII) electron transport and α were simultaneously measured in vivo at 20 different wavelengths. The wavelength dependence of the photosystem excitation balance was calculated from both these in vivo data and in vitro from the photosystem composition and spectroscopic properties. Measuring wavelengths overexciting PSI produced a higher α for leaves grown under the shade light spectrum (i.e., PSI light), whereas wavelengths overexciting PSII produced a higher α for the sun and blue leaves. The shade spectrum produced the lowest PSI:PSII ratio. The photosystem excitation balance calculated from both in vivo and in vitro data was substantially similar and was shown to determine α at those wavelengths where absorption by carotenoids and nonphotosynthetic pigments is insignificant (i.e., >580 nm). We show quantitatively that leaves acclimate their photosystem composition to their growth light spectrum and how this changes the wavelength dependence of the photosystem excitation balance and quantum yield for CO(2) fixation. This also proves that combining different wavelengths can enhance quantum yields substantially.

Meristem temperature substantially deviates from air temperature even in moderate environments: is the magnitude of this deviation species-specific?

Meristem temperature (Tmeristem ) drives plant development but is hardly ever quantified. Instead, air temperature (Tair ) is usually used as its approximation. Meristems are enclosed within apical buds. Bud structure and function may differ across species. Therefore, Tmeristem may deviate from Tair in a species-specific way. Environmental variables (air temperature, vapour pressure deficit, radiation, and wind speed) were systematically varied to quantify the response of Tmeristem . This response was related to observations of bud structure and transpiration. Tomato and cucumber plants were used as model plants as they are morphologically distinct and usually growing in similar environments. Tmeristem substantially deviated from Tair in a species-specific manner under moderate environments. This deviation ranged between -2.6 and 3.8 °C in tomato and between -4.1 and 3.0 °C in cucumber. The lower Tmeristem observed in cucumber was linked with the higher transpiration of the bud foliage sheltering the meristem when compared with tomato plants. We here indicate that for properly linking growth and development of plants to temperature in future applications, for instance in climate change scenarios studies, Tmeristem should be used instead of Tair , as a species-specific trait highly reliant on various environmental factors.

Effect of preharvest conditions on cut-flower quality.

The cut flower industry has a global reach as flowers are often produced in countries around the equator and transported by plane or ship (reefer) mostly to the global north. Vase-life issues are often regarded as linked to only postharvest conditions while cultivation factors are just as important. Here, we review the main causes for quality reduction in cut flowers with the emphasis on the importance of preharvest conditions. Cut flower quality is characterised by a wide range of features, such as flower number, size, shape, colour (patterns), fragrance, uniformity of blooming, leaf and stem colour, plant shape and developmental stage, and absence of pests and diseases. Postharvest performance involves improving and preserving most of these characteristics for as long as possible. The main causes for cut flower quality loss are reduced water balance or carbohydrate availability, senescence and pest and diseases. Although there is a clear role for genotype, cultivation conditions are just as important to improve vase life. The role of growth conditions has been shown to be essential; irrigation, air humidity, and light quantity and quality can be used to increase quality. For example, xylem architecture is affected by the irrigation scheme, and the relative humidity in the greenhouse affects stomatal function. Both features determine the water balance of the flowering stem. Light quality and period drives photosynthesis, which is directly responsible for accumulation of carbohydrates. The carbohydrate status is important for respiration, and many senescence related processes. High carbohydrates can lead to sugar loss into the vase water, leading to bacterial growth and potential xylem blockage. Finally, inferior hygiene during cultivation and temperature and humidity control during postharvest can lead to pathogen contamination. At the end of the review, we will discuss the future outlook focussing on new phenotyping tools necessary to quantify the complex interactions between cultivation factors and postharvest performance of cut flowers.

Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves.

Long-term effects of light quality on leaf hydraulic conductance (K(leaf)) and stomatal conductance (g(s)) were studied in cucumber, and their joint impact on leaf photosynthesis in response to osmotic-induced water stress was assessed. Plants were grown under low intensity monochromatic red (R, 640 nm), blue (B, 420 nm) or combined red and blue (R:B, 70:30) light. K(leaf) and g(s) were much lower in leaves that developed without blue light. Differences in g(s) were caused by differences in stomatal aperture and stomatal density, of which the latter was largely due to differences in epidermal cell size and hardly due to stomatal development. Net photosynthesis (A(N)) was lowest in R-, intermediate in B-, and highest in RB- grown leaves. The low A(N) in R-grown leaves correlated with a low leaf internal CO(2) concentration and reduced PSII operating efficiency. In response to osmotic stress, all leaves showed similar degrees of stomatal closure, but the reduction in A(N) was larger in R- than in B- and RB-grown leaves. This was probably due to damage of the photosynthetic apparatus, which only occurred in R-grown leaves. The present study shows the co-ordination of K(leaf) and g(s) across different light qualities, while the presence of blue in the light spectrum seems to drive both K(leaf) and g(s) towards high, sun-type leaf values, as was previously reported for maximal photosynthetic capacity and leaf morphology. The present results suggest the involvement of blue light receptors in the usually harmonized development of leaf characteristics related to water relations and photosynthesis under different light environments.

Using a biochemical C4 photosynthesis model and combined gas exchange and chlorophyll fluorescence measurements to estimate bundle-sheath conductance of maize leaves differing in age and nitrogen content.

Bundle-sheath conductance (g(bs) ) affects CO(2) leakiness, and, therefore, the efficiency of the CO(2) -concentrating mechanism (CCM) in C(4) photosynthesis. Whether and how g(bs) varies with leaf age and nitrogen status is virtually unknown. We used a C(4) -photosynthesis model to estimate g(bs) , based on combined measurements of gas exchange and chlorophyll fluorescence on fully expanded leaves of three different ages of maize (Zea mays L.) plants grown under two contrasting nitrogen levels. Nitrogen was replenished weekly to maintain leaf nitrogen content (LNC) at a similar level across the three leaf ages. The estimated g(bs) values on leaf-area basis ranged from 1.4 to 10.3 mmol m(-2) s(-1) and were affected more by LNC than by leaf age, although g(bs) tended to decrease as leaves became older. When converted to resistance (r(bs) = 1/g(bs)), r(bs) decreased monotonically with LNC. The correlation was presumably associated with nitrogen effects on leaf anatomy such as on wall thickness of bundle-sheath cells. Despite higher g(bs), meaning less efficient CCM, the calculated loss due to photorespiration was still low for high-nitrogen leaves. Under the condition of ambient CO(2) and saturating irradiance, photorespiratory loss accounted for 3-5% of fixed carbon for the high-nitrogen, versus 1-2% for the low-nitrogen, leaves.

Photosynthetic acclimation in relation to nitrogen allocation in cucumber leaves in response to changes in irradiance.

Leaves deep in canopies can suddenly be exposed to increased irradiances following e.g. gap formation in forests or pruning in crops. Studies on the acclimation of photosynthesis to increased irradiance have mainly focused on the changes in photosynthetic capacity (A(max)), although actual irradiance often remains below saturating level. We investigated the effect of changes in irradiance on the photosynthesis irradiance response and on nitrogen allocation in fully grown leaves of Cucumis sativus. Leaves that fully developed under low (50 µmol m⁻² s⁻¹) or moderate (200 µmol m⁻² s⁻¹) irradiance were subsequently exposed to, respectively, moderate (LM-leaves) or low (ML-leaves) irradiance or kept at constant irradiance level (LL- and MM-leaves). Acclimation of photosynthesis occurred within 7 days with final A(max) highest in MM-leaves, lowest in LL-leaves and intermediate in ML- and LM-leaves, whereas full acclimation of thylakoid processes underlying photosystem II (PSII) efficiency and non-photochemical quenching occurred in ML- and LM-leaves. Dark respiration correlated with irradiance level, but not with A(max). Light-limited quantum efficiency was similar in all leaves. The increase in photosynthesis at moderate irradiance in LM-leaves was primarily driven by nitrogen import, and nitrogen remained allocated in a similar ratio to Rubisco and bioenergetics, while allocation to light harvesting relatively decreased. A contrary response of nitrogen was associated with the decrease in photosynthesis in ML-leaves. Net assimilation of LM-leaves under moderate irradiance remained lower than in MM-leaves, revealing the importance of photosynthetic acclimation during the leaf developmental phase for crop productivity in scenarios with realistic, moderate fluctuations in irradiance that leaves can be exposed to.

Vegetative traits can predict flowering quality in Phalaenopsis orchids despite large genotypic variation in response to light and temperature.

Phalaenopsis is an economically important horticultural ornamental, but its growth is slow and costly. The vegetative cultivation phase is long and required to ensure sufficient plant size. This is needed to develop high quality flowering plants. We studied the effects of temperature (27 or 31 °C) and light intensity (60 or 140 µmol m-2 s-1) on plant growth and development during the vegetative cultivation phase in two experiments, with respectively 19 and 14 genotypes. Furthermore, the after-effects of treatments applied during vegetative growth on flowering traits were determined. Increasing light intensity in the vegetative phase accelerated both vegetative plant growth and development. Increasing temperature accelerated vegetative leaf appearance rate, but strongly reduced plant and root biomass accumulation when temperatures were too high. Flowering was greatly affected by treatments applied during vegetative growth, and increased light and temperature increased number of flower spikes, and number of flowers and buds. Genotypic variation was large in Phalaenopsis, especially in traits related to flowering, thus care is needed when generalising results based on a limited number of cultivars. Plant biomass and number of leaves during vegetative growth were positively correlated with flowering quality. These traits can be used as an early predictor for flowering capacity and quality of the final product. Additionally, this knowledge can be used to improve selection of new cultivars.

Red light imaging for programmed cell death visualization and quantification in plant-pathogen interactions.

Studies on plant-pathogen interactions often involve monitoring disease symptoms or responses of the host plant to pathogen-derived immunogenic patterns, either visually or by staining the plant tissue. Both these methods have limitations with respect to resolution, reproducibility, and the ability to quantify the results. In this study we show that red light detection by the red fluorescent protein (RFP) channel of a multipurpose fluorescence imaging system that is probably available in many laboratories can be used to visualize plant tissue undergoing cell death. Red light emission is the result of chlorophyll fluorescence on thylakoid membrane disassembly during the development of a programmed cell death process. The activation of programmed cell death can occur during either a hypersensitive response to a biotrophic pathogen or an apoptotic cell death triggered by a necrotrophic pathogen. Quantifying the intensity of the red light signal enables the magnitude of programmed cell death to be evaluated and provides a readout of the plant immune response in a faster, safer, and nondestructive manner when compared to previously developed chemical staining methodologies. This application can be implemented to screen for differences in symptom severity in plant-pathogen interactions, and to visualize and quantify in a more sensitive and objective manner the intensity of the plant response on perception of a given immunological pattern. We illustrate the utility and versatility of the method using diverse immunogenic patterns and pathogens.

Caspase inhibitors affect the kinetics and dimensions of tracheary elements in xylogenic Zinnia (Zinnia elegans) cell cultures.

BACKGROUND: The xylem vascular system is composed of fused dead, hollow cells called tracheary elements (TEs) that originate through trans-differentiation of root and shoot cambium cells. TEs undergo autolysis as they differentiate and mature. The final stage of the formation of TEs in plants is the death of the involved cells, a process showing some similarities to programmed cell death (PCD) in animal systems. Plant proteases with functional similarity to proteases involved in mammalian apoptotic cell death (caspases) are suggested as an integral part of the core mechanism of most PCD responses in plants, but participation of plant caspase-like proteases in TE PCD has not yet been documented. RESULTS: Confocal microscopic images revealed the consecutive stages of TE formation in Zinnia cells during trans-differentiation. Application of the caspase inhibitors Z-Asp-CH2-DCB, Ac-YVAD-CMK and Ac-DEVD-CHO affected the kinetics of formation and the dimensions of the TEs resulting in a significant delay of TE formation, production of larger TEs and in elimination of the 'two-wave' pattern of TE production. DNA breakdown and appearance of TUNEL-positive nuclei was observed in xylogenic cultures and this was suppressed in the presence of caspase inhibitors. CONCLUSIONS: To the best of our knowledge this is the first report showing that caspase inhibitors can modulate the process of trans-differentiation in Zinnia xylogenic cell cultures. As caspase inhibitors are closely associated with cell death inhibition in a variety of plant systems, this suggests that the altered TE formation results from suppression of PCD. The findings presented here are a first step towards the use of appropriate PCD signalling modulators or related molecular genetic strategies to improve the hydraulic properties of xylem vessels in favour of the quality and shelf life of plants or plant parts.

Blue light dose-responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light.

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 F(v)/F(m), a stomatal conductance unresponsive to irradiance, and a relatively low light-limited quantum yield for CO(2) 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 (A(max)) 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, A(max) was lower but photosynthetic functioning was normal. The increase in A(max) 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 A(max), 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.

An artificial solar spectrum substantially alters plant development compared with usual climate room irradiance spectra.

Plant responses to the light spectrum under which plants are grown affect their developmental characteristics in a complicated manner. Lamps widely used to provide growth irradiance emit spectra which are very different from natural daylight spectra. Whereas specific responses of plants to a spectrum differing from natural daylight may sometimes be predictable, the overall plant response is generally difficult to predict due to the complicated interaction of the many different responses. So far studies on plant responses to spectra either use no daylight control or, if a natural daylight control is used, it will fluctuate in intensity and spectrum. An artificial solar (AS) spectrum which closely resembles a sunlight spectrum has been engineered, and growth, morphogenesis, and photosynthetic characteristics of cucumber plants grown for 13 d under this spectrum have been compared with their performance under fluorescent tubes (FTs) and a high pressure sodium lamp (HPS). The total dry weight of the AS-grown plants was 2.3 and 1.6 times greater than that of the FT and HPS plants, respectively, and the height of the AS plants was 4-5 times greater. This striking difference appeared to be related to a more efficient light interception by the AS plants, characterized by longer petioles, a greater leaf unfolding rate, and a lower investment in leaf mass relative to leaf area. Photosynthesis per leaf area was not greater for the AS plants. The extreme differences in plant response to the AS spectrum compared with the widely used protected cultivation light sources tested highlights the importance of a more natural spectrum, such as the AS spectrum, if the aim is to produce plants representative of field conditions.

The responses of light interception, photosynthesis and fruit yield of cucumber to LED-lighting within the canopy.

Mathematical models of light attenuation and canopy photosynthesis suggest that crop photosynthesis increases by more uniform vertical irradiance within crops. This would result when a larger proportion of total irradiance is applied within canopies (interlighting) instead of from above (top lighting). These irradiance profiles can be generated by Light Emitting Diodes (LEDs). We investigated the effects of interlighting with LEDs on light interception, on vertical gradients of leaf photosynthetic characteristics and on crop production and development of a greenhouse-grown Cucumis sativus'Samona' crop and analysed the interaction between them. Plants were grown in a greenhouse under low natural irradiance (winter) with supplemental irradiance of 221 micromol photosynthetic photon flux m(-2) s(-1) (20 h per day). In the interlighting treatment, LEDs (80% Red, 20% Blue) supplied 38% of the supplemental irradiance within the canopy with 62% as top lighting by High-Pressure Sodium (HPS)-lamps. The control was 100% top lighting (HPS lamps). We measured horizontal and vertical light extinction as well as leaf photosynthetic characteristics at different leaf layers, and determined total plant production. Leaf mass per area and dry mass allocation to leaves were significantly greater but leaf appearance rate and plant length were smaller in the interlighting treatment. Although leaf photosynthetic characteristics were significantly increased in the lower leaf layers, interlighting did not increase total biomass or fruit production, partly because of a significantly reduced vertical and horizontal light interception caused by extreme leaf curling, likely because of the LED-light spectrum used, and partly because of the relatively low irradiances from above.