CAM-physiology and carbon gain of the orchid Phalaenopsis in response to light intensity, light integral and CO2.

The regulation of photosynthesis and carbon gain of crassulacean acid metabolism (CAM) plants has not yet been disclosed to the extent of C3-plants. In this study, the tropical epiphyte Phalaenopsis cv. "Sacramento" was subjected to different lighting regimes. Photosynthesis and biochemical measuring techniques were used to address four specific questions: (1) the response of malate decarboxylation to light intensity, (2) the malate carboxylation pathway in phase IV, (3) the response of diel carbon gain to the light integral and (4) the response of diel carbon gain to CO2 . The four CAM-phases were clearly discernable. The length of phase III and the malate decarboxylation rate responded directly to light intensity. In phase IV, CO2 was initially mainly carboxylated via Rubisco. However, at daylength of 16 h, specifically beyond ±12 h, it was mainly phosphoenolpyruvate carboxylase (PEP-C) carboxylating CO2 . Diel carbon gain appeared to be controlled by the light integral during phase III rather than the total daily light integral. Elevated CO2 further enhanced carbon gain both in phase IV and phase I. This establishes that neither malate storage capacity, nor availability of PEP as substrate for nocturnal CO2 carboxylation were limiting factors for carbon gain enhancement. These results advance our understanding of CAM-plants and are also of practical importance for growers.

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.

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.

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.

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.