Interspecific prediction of photosynthetic light response curves using specific leaf mass and leaf nitrogen content: effects of differences in soil fertility and growth irradiance

Background and Aims

Previous work has shown that the entire photosynthetic light response curve, based on both Mitscherlich and Michaelis–Menten functions, could be predicted in an interspecific context through allometric relations linking the parameters of these functions to two static leaf traits: leaf nitrogen (N) content and leaf mass per area (LMA). This paper describes to what extent these allometric relations are robust to changes in soil fertility and the growth irradiance of the plants.

Methods

Plants of 25 herbaceous species were grown under controlled conditions in factorial combinations of low/high soil fertility and low/high growth irradiance. Net photosynthetic rates per unit dry mass were measured at light intensities ranging from 0 to 700 µmol m⁻² s⁻¹ photosynthetically active radiation (PAR).

Key Results

The differing growth environments induced large changes in N, LMA and in each of the parameter estimates of the Mitscherlich and Michaelis–Menten functions. However, the differing growth environments induced only small (although significant) changes in the allometric relationships linking N and LMA to the parameters of the two functions. As a result, 88 % (Mitcherlich) and 89 % (Michaelis–Menten) of the observed net photosynthetic rates over the full range of light intensities (0–700 µmol m⁻² s⁻¹ PAR) and across all four growth environments could be predicted using only N and LMA using the same allometric relations.

Conclusions

These results suggest the possibility of predicting net photosynthetic rates in nature across species over the full range of light intensities using readily available data.     

Hysteresis light curves: a protocol for characterizing the time dependence of the light response of photosynthesis

Photosynthesis vs. light curves (LCs) have played a central role in photosynthesis research for decades. They are the commonest form of describing how photosynthesis responds to changes in light, being frequently used for characterizing photoacclimation. However, LCs are often interpreted exclusively regarding the response to light intensity, the effects of time of exposure not being explicitly considered. This study proposes the use of ‘hysteresis light curves’ (HLC), an experimental protocol focused on the cumulative effects of light exposure to obtain information on the time dependence of photosynthetic light responses. HLC are generated by exposing samples to a symmetrical sequence of increasing and decreasing light levels. The comparison of the light-increasing and the light-decreasing phases allows the quantification of the hysteresis caused by high-light exposure, the magnitude and direction of which inform on the activation, and subsequent relaxation of high-light-induced photosynthetic processes. HLCs of the chlorophyll fluorescence indices rETR (relative electron transport rate of photosystem II) and Y(NPQ) (index of non-photochemical quenching) were measured on cyanobacteria, algae, and plants, with the aim of identifying main patterns of hysteresis and their diversity. A non-parametric index is proposed to quantify the magnitude and direction of hysteresis in HLCs of rETR and Y(NPQ). The results of this study show that HLCs can provide additional relevant information on the time dependence of the light response of photosynthetic samples, not obtainable from conventional LCs, useful for phenotyping photosynthetic traits, including photoacclimation state and kinetics of light activation and relaxation of electron flow and energy dissipation processes.

     

Geometrical similarity analysis of photosynthetic light response curves, light saturation and light use efficiency

Light absorption and use efficiency (LAUE mol mol⁻¹, daily gross photosynthesis per daily incident light) of each leaf depends on several factors, including the degree of light saturation. It is often discussed that upper canopy leaves exposed to direct sunlight are fully light-saturated. However, we found that upper leaves of three temperate species, a heliophytic perennial herb Helianthus tuberosus, a pioneer tree Alnus japonica, and a late-successional tree Fagus crenata, were not fully light-saturated even under full sunlight. Geometrical analysis of the photosynthetic light response curves revealed that all the curves of the leaves from different canopy positions, as well as from the different species, can be considered as different parts of a single non-rectangular hyperbola. The analysis consistently explained how those leaves were not fully light-saturated. Light use optimization models, called big leaf models, predicted that the degree of light saturation and LAUE are both independent of light environment. From these, we hypothesized that the upper leaves should not be fully light-saturated even under direct sunlight, but instead should share the light limitation with the shaded lower-canopy leaves, so as to utilize strong sunlight efficiently. Supporting this prediction, within a canopy of H. tuberosus, both the degree of light saturation and LAUE were independent of light environment within a canopy, resulting in proportionality between the daily photosynthesis and the daily incident light among the leaves.     

The Paramecium circadian behavioral rhythm: light phase response curves and entrainment

The population of a ciliate protozoan, Paramecium multimicronucleatum, exhibits a circadian rhythm as measured by the number of the cells traversing an observation point ("traverse frequency," or TF). The present study examined phase shifting of the TF rhythm by administering 2-hr light pulses at different phases of the circadian cycle to cultures free-running in constant darkness (DD). The results were summarized in a phase response curve (PRC), categorized as Type 1. This PRC indicated a relatively narrow phase zone insensitive to the light pulse ("dead zone"). Entrainment of the rhythm to light pulses repeated at 24-hr intervals was also examined, and it was found that the rhythm gradually reached a steady state, following several transient cycles, with the pulses falling at a phase corresponding to the narrow dead zone. Such a steady-state rhythm, with a minimum at approximately 3 hr after the pulse and a maximum at approximately 12 hr after the pulse, was mathematically simulated by superimposing a response function to the pulse on a sinusoidal function representative of the free-running rhythm in DD.     

Photosynthetic apparatus of pea thylakoid membranes : response to growth light intensity

We investigated the effect of growth light intensity on the photosynthetic apparatus of pea (Pisum sativum) thylakoid membranes. Plants were grown either in a growth chamber at light intensities that ranged from 8 to 1050 microeinsteins per square meter per second, or outside under natural sunlight. In thylakoid membranes we determined: the amounts of active and inactive photosystem II, photosystem I, cytochrome b/f, and high potential cytochrome b₅₅₉, the rate of uncoupled electron transport, and the ratio of chlorophyll a to b. In leaves we determined: the amounts of the photosynthetic components per leaf area, the fresh weight per leaf area, the rate of electron transport, and the light compensation point. To minimize factors other than growth light intensity that may alter the photosynthetic apparatus, we focused on peas grown above the light compensation point (20-40 microeinsteins per square meter per second), and harvested only the unshaded leaves at the top of the plant. The maximum difference in the concentrations of the photosynthetic components was about 30% in thylakoids isolated from plants grown over a 10-fold range in light intensity, 100 to 1050 microeinsteins per square meter per second. Plants grown under natural sunlight were virtually indistinguishable from plants grown in growth chambers at the higher light intensities. On a leaf area basis, over the same growth light regime, the maximum difference in the concentration of the photosynthetic components was also about 30%. For peas grown at 1050 microeinsteins per square meter per second we found the concentrations of active photosystem II, photosystem I, and cytochrome b/f were about 2.1 millimoles per mol chlorophyll. There were an additional 20 to 33% of photosystem II complexes that were inactive. Over 90% of the heme-containing cytochrome f detected in the thylakoid membranes was active in linear electron transport. Based on these data, we do not find convincing evidence that the stoichiometries of the electron transport components in the thylakoid membrane, the size of the light-harvesting system serving the reaction centers, or the concentration of the photosynthetic components per leaf area, are regulated in response to different growth light intensities. The concept that emerges from this work is of a relatively fixed photosynthetic apparatus in thylakoid membranes of peas grown above the light compensation point.     

Photosynthetic efficiency and oxygen evolution of Chlamydomonas reinhardtii under continuous and flashing light

As a result of mixing and light attenuation in a photobioreactor (PBR), microalgae experience light/dark (L/D) cycles that can enhance PBR efficiency. One parameter which characterizes L/D cycles is the duty cycle; it determines the time fraction algae spend in the light. The objective of this study was to determine the influence of different duty cycles on oxygen yield on absorbed light energy and photosynthetic oxygen evolution. Net oxygen evolution of Chlamydomonas reinhardtii was measured for four duty cycles (0.05, 0.1, 0.2, and 0.5) in a biological oxygen monitor (BOM). Oversaturating light flashes were applied in a square-wave fashion with four flash frequencies (5, 10, 50, and 100 Hz). Algae were precultivated in a turbidostat and acclimated to a low photon flux density (PFD). A photosynthesis–irradiance (PI) curve was measured under continuous illumination and used to calculate the net oxygen yield, which was maximal between a PFD of 100 and 200 μmol m^?2?s^?1. Net oxygen yield under flashing light was duty cycle-dependent: the highest yield was observed at a duty cycle of 0.1 (i.e., time-averaged PFD of 115 μmol m^?2?s^?1). At lower duty cycles, maintenance respiration reduced net oxygen yield. At higher duty cycles, photon absorption rate exceeded the maximal photon utilization rate, and, as a result, surplus light energy was dissipated which led to a reduction in net oxygen yield. This behavior was identical with the observation under continuous light. Based on these data, the optimal balance between oxygen yield and production rate can be determined to maximize PBR productivity.     

Photosynthetic performance and light response of two olive cultivars under different water and light regimes

The olive tree (Olea europaea L.) is commonly grown in the Mediterranean area, where it is adapted to resist periods characterized by severe drought and high irradiance levels. Photosynthetic efficiency (in terms of F_v/F_m and Φ_PSII), photochemical (q_P) and nonphotochemical quenching (NPQ) were determined in two-year-old olive plants (cultivars Coratina and Biancolilla) grown under two different light levels (exposed plants, EP, and shaded plants, SP) during a 21-day controlled water deficit. After reaching the maximum level of drought stress, plants were rewatered for 23 days. During the experimental period, measurements of gas exchange and chlorophyll (Chl) fluorescence were carried out to study the photosynthetic performance of olive plants. The synergical effect of drought stress and high irradiance levels caused a reduction of gas exchange and photosynthetic efficiency and these decreases were more marked in EP. EP showed a higher degree of photoinhibition, a higher NPQ and a lower q_P if compared to SP. Coratina was more sensitive to high light and drought stress but also showed a slower recovery during rewatering, whereas Biancolilla showed a less marked photosynthesis depression during drought and a considerable resilience during rewatering. The results confirm that photoinhibition due to high light intensity and water deficit can be an important factor that affects photosynthetic productivity in this species.     

四种水稻品种的光合光响应曲线及其模型拟合

应用4种典型的光响应模型对4个水稻品种“02428”、“淦鑫688”、“JR8892-1”和“JR8892-2”叶片的光合光响应曲线进行了拟合.结果表明:直角双曲线模型、非直角双曲线模型、指数函数以及修正的直角双曲线模型对4个水稻品种的光合光响应曲线都可以进行拟合,决定系数均为R2 ＞0.99.但只有修正的直角双曲线模型可以拟合发生光抑制部分的光响应曲线,且可直接计算其饱和光强(Isat);由各模型计算的光合参数可知,直角双曲线模型、非直角双曲线模型求得的最大净光合速率(Pnmax)远高于实测值,指数函数与修正的直角双曲线模型求得的Pnm.与实测值最为接近;直角双曲线、非直角双曲线模型以及指数函数求得的Isat均与实测值相差较大,除了“JR8892-1”采用指数函数求得的Isat大于实测值外,其余3个品种用上述3种模型拟合求得的Isat均小于实测值,唯有修正的直角双曲线模型求得的Isat与实测值最为相符;对于低光强部分的参数光补偿点(Ic)与暗呼吸速率(Rd),修正的直角双曲线模型最接近其测量值;不同水稻品种的光合特性,“JR8892-1”饱和光强与最大净光合速率明显高于高光效品种“02428”;处于分蘖盛期的“JR8892-2”也表现出较高的饱和光强和最大净光合速率,而超级稻“淦鑫688”相对于其他品种,则具有较低的暗呼吸速率,光合产物消耗少.     

Photosynthetic response of seagrasses to ultraviolet-a radiation and the influence of visible light intensity

Inhibition of photosynthesis by ultraviolet-A radiation (UV-A, 315-380 nanometers) was examined in three marine angiosperms: Halophila engelmannii Aschers, Halodule wrightii Aschers, and Syringodium filiforme Kütz. Sensitivity to UV-A and photosensitization to UV-A by photosynthetically active radiation (PAR, 380-700 nanometers) were characterized.     

Photosynthetic gas exchange response of poplars to steady-state and dynamic light environments

The steady-state and dynamic photosynthetic response of two poplar species (Populus tremuloides and P. fremontii) to variations in photon flux density (PFD) were observed with a field portable gas exchange system. These poplars were shown to be very shade intolerant with high light saturation (800 to 1300 mol photons m^–2 s^–1) and light compensation (70 to 100 mol m^–2 s^–1) points. Understory poplar leaves showed no physiological acclimation to understory light environments. These plants become photosynthetically induced quickly (10 min). Activation of Rubisco was the primary limitation for induction, with stomatal opening playing only a minor role. Leaves maintained high stomatal conductances and stomata were unresponsive to variations in PFD. Leaves were very efficient at utilizing rapidly fluctuating light environments similar to those naturally occurring in canopies. Post-illumination CO₂ fixation contributed proportionally more to the carbon gain of leaves during short frequent lightflecks than longer less frequent ones. The benefits of a more dynamic understory light environment for the carbon economy of these species are discussed.     

Blue light induced response in the absence of free oxygen

The absence of oxygen had no effect on the photoinduced conidiationof Trichoderma, a system (unlike previously used systems) wherethe transient photoinduction and the prolonged respiratory oxygenrequirements are separable. The nature of the presumably ubiquitous blue-light absorbingpigment is not restricted to those requiring oxygen. ¹ Irene and David Schwartz Professor of Plant Genetics. (Received September 19, 1974; )     

The scaling of leaf area and mass: the cost of light interception increases with leaf size

For leaves, the light-capturing surface area per unit dry mass investment (specific leaf area, SLA) is a key trait from physiological, ecological and biophysical perspectives. To address whether SLA declines with leaf size, as hypothesized due to increasing costs of support in larger leaves, we compiled data on intraspecific variation in leaf dry mass (LM) and leaf surface area (LA) for 6334 leaves of 157 species. We used the power function LM=alpha LAbeta to test whether, within each species, large leaves deploy less surface area per unit dry mass than small leaves. Comparing scaling exponents (beta) showed that more species had a statistically significant decrease in SLA as leaf size increased (61) than the opposite (7) and the average beta was significantly greater than 1 (betamean=1.10, 95% CI 1.08-1.13). However, scaling exponents varied markedly from the few species that decreased to the many that increased SLA disproportionately fast as leaf size increased. This variation was unrelated to growth form, ecosystem of origin or climate. The average within-species tendency found here (allometric decrease of SLA with leaf size, averaging 13%) is in accord with concurrent findings on global-scale trends among species, although the substantial scatter around the central tendency suggests that the leaf size dependency does not obligately shape SLA. Nonetheless, the generally greater mass per unit leaf area of larger than smaller leaves directly translates into a greater cost to build and maintain a unit of leaf area, which, all else being equal, should constrain the maximum leaf size displayed.     

A more sensitive modification of the catalase assay with the Clark oxygen electrode : Application to the kinetic study of the pea leaf enzyme

A modification of the method of catalase determination by means of the Clark oxygen electrode is described. The assay is based on measurement of the initial rate at which oxygen is released by catalase in an oxygen-free buffer. Displacement of oxygen was brought about by flushing with nitrogen, and the substrate used was hydrogen peroxide at a 33.5 m final concentration. The method is rapid and can be used with crude catalase preparations. Its sensitivity is at least 20 times higher than that of previous methods; it has an interval of measurable activity of about 0.01–8.4 μmol of O₂/min and, therefore, is applicable to an 840-fold range of catalase concentrations. This modification was applied to the kinetic study of crude extracts of pea leaf catalase. An apparent K_m of 0.190 was calculated.     

The rapid light response of leaf hydraulic conductance: new evidence from two experimental methods

Previous studies have shown a rapid enhancement in leaf hydraulic conductance (K(leaf)) from low to high irradiance (from <10 to >1000 micromol photons m(-2) s(-1)), using the high-pressure flow meter (HPFM), for 7 of 14 tested woody species. However, theoretical suggestions have been made that this response might arise as an artifact of the HPFM. We tested the K(leaf) light response for six evergreen species using refined versions of the rehydration kinetics method (RKM) and the evaporative flux method (EFM). We found new evidence for a rapid, 60% to 100% increase in K(leaf) from low to high irradiance for three species. In the RKM, the leaf rehydration time constant declined by up to 70% under high irradiance relative to darkness. In the EFM, under higher irradiance, the flow rate increased disproportionately to the water potential gradient. Combining our data with those of previous studies, we found that heterobaric species, i.e. those with bundle sheath extensions (BSEs) showed a twofold greater K(leaf) light response on average than homobaric species, i.e. those without BSEs. We suggest further research to characterize this substantial dynamic at the nexus of plant light- and water-relations.     

Photosynthetic energy conversion under extreme conditions--II: the significance of lipids under light limited growth in Antarctic sea ice diatoms

Low photosynthetic active radiation is a strong determinant in the development and growth of sea ice algae. The algae appear to have universal mechanisms to overcome light limitation. One important process, which is induced under light limitation, is the desaturation of chloroplast membrane lipids. In order to discover whether this process is universally valid in sea ice diatoms, we investigated three species coexisting in chemostats illuminated with 15 and 2 micromol photons m(-2) s(-1) at -1 degrees C. Growth under 2 micromol photons m(-2) s(-1) caused a 50% increase in monogalactosyldiacylglycerols (MGDG) thylakoid membrane related 20:5 n-3 fatty acids. This fatty acid supports the fluidity of the thylakoid membrane and therefore the velocity of electron flow, which is indicated by increasing rate constants for the electron transport between Q(A) (first stable electron acceptor) and bound Q(B) (second stable electron acceptor) (11.16 +/- 1.34 to 23.24 +/- 1.35 relative units). Two micromol photons m(-2) s(-1) furthermore resulted in higher amounts of non-lipid bilayer forming MGDG in relation to other bilayer forming lipids, especially digalactosydiacylglycerol (DGDG). The ratio of MGDG:DGDG increased from 3.4 +/- 0.3 to 5.7 +/- 0.3. The existence of bilayer thylakoid membranes with high proportions of non. bilayer forming lipids is only possible when sufficient thylakoid pigment-protein complexes are present. If more thylakoid pigment-protein complexes are present in membranes, as found under extreme light limitation, less bilayer forming lipids such as DGDG are required to stabilize the bilayer structure. Differences in protein contents between both light intensities were not found. Consequently pigment contents which nearly doubled under 2 micromol photons m(-2) s(-1) must be responsible in balancing the potential stability loss resulting from an increase in MGDG:DGDG ratio.     

Effect of light intensity on the phase and period response curves in the nocturnal field mouse Mus booduga

The effect of light intensity on the phase response curve (PRC) and the period response curve (τRC) of the nocturnal field mouse Mus booduga was studied. PRCs and τRCs were constructed by exposing animals free-running in constant darkness (DD), to fluorescent light pulses (LPs) of 100 lux and 1000 lux intensities for 15min duration. The waveform of the PRCs and τRCs evoked by high light intensity (1000 lux) stimuli was significantly different compared to those constructed using low light intensity (100 lux). Moreover, a weak but significant correlation was observed between phase shifts and period changes when light stimuli of 1000 lux intensity were used; however, the phase shifts and period changes in the 100 lux PRC and τRC were not correlated. This suggests that the intensity of light stimuli affects both phase and period responses in the locomotor activity rhythm of the nocturnal field mouse M. booduga. These results indicate that complex mechanisms are involved in entrainment of circadian clocks, even in nocturnal rodents, in which PRC, τRC, and dose responses play a significant role.