Wim van Ieperen

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 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.

Photosynthetic Quantum Yield Dynamics: From Photosystems to Leaves

The quantum yield for CO2 fixation is wavelength dependent due to (1) light absorption by nonphotosynthetic pigments, (2) inefficient energy transfer, and (3) the excitation balance between the two photosystems. The growth-light spectrum alters the excitation balance by altering the photosystem composition, as shown both in vivo and in vitro. Enhancement effects can increase the quantum yield. The mechanisms underlying the wavelength dependence of the quantum yield for CO2 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 CO2 fixation. This also proves that combining different wavelengths can enhance quantum yields substantially.

Plants under continuous light

Continuous light is an essential tool for understanding the plant circadian clock. Additionally, continuous light might increase greenhouse food production. However, using continuous light in research and practice has its challenges. For instance, most of the circadian clock-oriented experiments were performed under continuous light; consequently, interactions between the circadian clock and the light signaling pathway were overlooked. Furthermore, in some plant species continuous light induces severe injury, which is only poorly understood so far. In this review paper, we aim to combine the current knowledge with a modern conceptual framework. Modern genomic tools and rediscovered continuous light-tolerant tomato species (Solanum spp.) could boost the understanding of the physiology of plants under continuous light.

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

Long-term effects of light quality on leaf hydraulic conductance (Kleaf) and stomatal conductance (gs) 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. Kleaf and gs were much lower in leaves that developed without blue light. Differences in gs 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 (AN) was lowest in R-, intermediate in B-, and highest in RB- grown leaves. The low AN in R-grown leaves correlated with a low leaf internal CO2 concentration and reduced PSII operating efficiency. In response to osmotic stress, all leaves showed similar degrees of stomatal closure, but the reduction in AN 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 Kleaf and gs across different light qualities, while the presence of blue in the light spectrum seems to drive both Kleaf and gs 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.

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.

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.

Establishing in vitro Zinnia elegans cell suspension culture with high tracheary element differentiation.

The Zinnia elegans mesophyll cell culture is a useful system for xylogenesis studies. The system is associated with highly synchronous tracheary element (TE) differentiation, making it more suitable for molecular studies requiring larger amounts of molecular isolates, such as mRNA and proteins and for studying cellulose synthesis. There is, however, the problem of non‐uniformity and significant variations in the yields of TEs (%TE). One possible cause for this variability in the%TE could be the lack of a standardized experimental protocol in various research laboratories for establishing the Zinnia culture. Mesophyll cells isolated from the first true leaves of Z. elegans var Envy seedlings of approximately 14 days old were cultured in vitro and differentiated into TEs. The xylogenic culture medium was supplied with 1 mg/l each of benzylaminopurine (BA) and α‐naphthalene acetic acid (NAA). Application of this improved culture method resulted in stable and reproducible amounts of TE as high as 76% in the Zinnia culture. The increase was mainly due to conditioning of the mesophyll cell culture and adjustments of the phytohormonal balance in the cultures. Also, certain biochemical and cytological methods have been shown to reliably monitor progress of TE differentiation. We conclude that, with the adoption of current improvement in the xylogenic Z. elegans culture, higher amounts of tracheary elements can be produced. This successful outcome raises the potential of the Zinnia system as a suitable model for cellulose and xylogenesis research.

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.

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.

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.