一些海洋浮游植物量子产值的研究

现场实验以及用硅藻、金藻和绿藻所做的实验表明,在光饱和深度以下,随着深度的增加,浮游植物的量子产懂具有不变、增加和下降3种垂直变化趋势。偏离理论模式的后两种结果可能是由浮游植物的光强适应(Light-shade adaptation)等原因引起。还探讨了量子产值作为初级生产力模型和光利用效率模型中的参数的问题。     

USE OF PULSE‐AMPLITUDE‐MODULATED FLUORESCENCE TO ASSESS THE PHYSIOLOGICAL STATUS OF CLADOPHORA SP. ALONG A WATER QUALITY GRADIENT1

This study investigated the application of pulse‐amplitude‐modulated (PAM) fluorometry as a rapid assessment of benthic macroalgal physiological status. Maximum quantum efficiency (F_v/F_m), dark–light induction curves, and rapid fluorescence light‐response curves (RLC) were measured on the filamentous macroalgal Cladophora sp. from Lake Ontario on 5 d at 16 sites spanning a gradient of light and nutrient supply. For Cladophora sp. growing in situ, light limitation was assessed by comparing average daily irradiance with the light utilization efficiency parameter (α) derived from RLCs. In this study, there was a nonlinear relationship between F_v/F_m and the degree of P limitation in macroalgae. However, only light‐saturated Cladophora sp. showed a significant positive linear relationship between F_v/F_m and P nutrient status. The absence of this relationship among light‐limited algae indicates that their photosynthetic rate would be stimulated by increased water clarity, and not by increased P supply. PAM fluorescence measures were successfully able to identify light‐saturated macroalgae and, among these, assess the degree to which they were nutrient limited. These results enable us to test hypotheses arising from numeric models predicting the impact of changes in light penetration and nutrient supply on benthic primary production.     

EFFECTS OF TEMPERATURE ON GROWTH,LIGHT ABSORPTION,AND QUANTUM YIELD IN DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

The effects of growth temperature on the marine chlorophyte Dunaliella tertiolecta Butcher were studied to provide a more mechanistic understanding of the role of environmental factors in regulating bio-optical properties of phytoplankton. Specific attention was focused on quantities that are relevant for modeling of growth and photosynthesis. Characteristics including chlorophyll a (chl z)-specific light absorption (a*_ph(λ)), C:chl a ratio, and quantum yield for growth (φ_μ) varied as functions of temperature under conditions of excess light and nutrients. As temperature increased over the range examined (12°-28°C), intracellular concentrations of chl a increased by a factor of 2 and a*_ph(λ) values decreased by more than 50% at blue to green wavelengths. The lower values of a*_ph(λ) were due to both a decrease in the abundance of accessory pigments relative to chl a and an increase in pigment package effects arising from higher intracellular pigment concentrations. Intracellular pigment concentration increased as a consequence of higher cellular pigment quotas combined with lower cell volume. At high growth temperatures, slightly more light was absorbed on a per-cell-C basis, but the dramatic increases in growth rate from μ= 0.5 d^?1 at 12° C to μ= 2.2 d^?1 at 28°C were primarily due to an increase in φ_μ (0.015–0.041 mol C (mol quanta)^?1). By comparison with previous work on this species, we conclude the effects of temperature on a*_ph(λ) and φ_μ are comparable to those observed for light and nutrient limitation. Patterns of variability in a*_ph(λ)and φ_μ as a function of growth rate at different temperatures are similar to those previously documented for this species grown at the same irradiance but under a range of nitrogen-limited conditions. These results are discussed in the context of implications for bio-optical modeling of aquatic primary production by phytoplankton.     

风力胁迫对具鞘微鞘藻结皮光合活性的影响

抵抗风力胁迫是荒漠藻类适应干旱区环境的重要生物学机制,也是藻结皮能够拓殖流沙的必要条件之一,但有关藻类对风力胁迫的响应机理国内外尚无研究报道。以具鞘微鞘藻(Microcoleus vaginatus Gom.)人工结皮为实验对象,研究了不同强度风力吹蚀对结皮含水量、藻类活力、生物量、及其光合活性的影响。结果表明:不论低于当地起沙风(3m/s)还是高于起沙风(5m/s和7m/s)的风力吹蚀,结皮中藻类生物量均明显下降,而且结皮生物量的变化与风速大小和吹蚀时间呈线性关系(y=14.78+0.035a-1.48b,a风速,b时间,r2=0.79)。进一步分析发现,风力吹蚀后结皮中藻类活力并没有降低但主要光合色素和天线色素的含量普遍降低,叶绿素荧光(Fv/Fm)、表观电子传递速率(ETR)和净光合速率(Pn)明显下降,并且风速越大,降幅越大。这些结果说明风力胁迫对藻结皮生长和光合活性的影响主要是通过影响光合色素代谢合成和电子传递速率引起的,对其生命力没有明显影响。       

LIGHT ABSORPTION AND QUANTUM YIELD FOR GROWTH IN FIVE SPECIES OF MARINE MACROALGA1

Light utilization efficiency in five species of marine macroalgae was measured in laboratory growth experiments (13–41 days duration) at different irradiances at 7°C. All species acclimated to irradiance by changing their light absorption, resulting in a peak in light absorption between 2 and 15 μmol·m^?2.s^?1. Light absorption increased with thallus-specific chlorophyll and carbon content according to linear inverse relationships between chlorophyll content (chlarea^?1) and log[transmission] and between log[carbon content, Carea^?1] and log[transmission]. Quantum yields for light-limited growth and estimated gross photosynthesis were calculated based on incident and absorbed light. Quantum yield for photosynthesis based on light absorbed by pigments was high (mean = 114 mmol C·mol^?1 photons) and similar among the species. Quantum yield for net growth based on incident light was also high but more variable, between 22 and 75 mmol C·mol^?1 photons. Differences among species were mainly due to differences in light absorption. In conclusion, all species acclimated to low light by increasing light absorption to the maximum attainable, and growth efficiencies based on absorbed light were close to the maximum theoretically possible.     

PHYSIOLOGICAL RESPONSES TO PHOSPHORUS LIMITATION IN BATCH AND STEADY-STATE CULTURES OF DUNALIELLA TERTIOLECTA (CHLOROPHYTA): A UNIQUE STRESS PROTEIN AS AN INDICATOR OF PHOSPHATE DEFICIENCY1

A protein unique to phosphorus stress observed in Dunaliella tertiolecta Butcher was studied in the context of phosphate-limited cell physiology and is a potential diagnostic indicator of phosphate deficiency in this alga. Cells were grown over a range of limited, steady-state growth rates and at maximum (replete) and zero (phosphate-starved) growth rates. The stress protein, absent in nutrient-replete cells, was produced under all steady-state phosphate-limited conditions and increased in abundance with increasing limitation (decreasing growth rate). Cellular carbon: phosphorus ratios and the maximum uptake rate of phosphate (V_m) increased with increasing limitation, whereas the ratio of chlorophyll a: carbon decreased. Alkaline phosphatase activity did not respond to limitation but was measurable in starved, stationary-phase cells. F_v/F_m, a measure of photochemical efficiency, was a nonlinear, saturating function of p, as commonly observed under N limitation. The maximum F_v/F_m of 0.64 was measured in nutrient-replete cells growing at μ_max, and a value of zero was measured in stationary-phase starved cells. When physiological parameters were compared, the P-stress protein abundance and F_v/F_m were the most sensitive indicators of the level of deficiency. The stress protein was not produced under N- or Fe-limited conditions. It is of high molecular weight (>200) and is associated with internal cell membranes. The stress protein has several characteristics that make it a potential diagnostic indicator: it is 1) unique to phosphorus limitation (i.e. absent under all other conditions), 2) present under limiting as well as starved conditions, 3) sensitive to the level of limitation, and 4) observable without time-course incubation of live samples.     

WAVELENGTH DEPENDENCY OF THE MAXIMUM QUANTUM YIELD OF CARBON FIXATION FOR TWO RED TIDE DINOFLAGELLATES,HETEROCAPSA PYGMAEA AND PROROCENTRUM MINIMUM (PYRROPHYTA): IMPLICATIONS FOR MEASURING PHOTOSYNTHETIC RATES1

The influence of photoadaptive state on the spectral dependency of the maximum quantum yield for carbon fixation was determined for two red tide dinoflagellates, Heterocapsa pygmaea Loeblich, Schmidt, et Sherley and Prorocentrum minimum Pavillard. Cultures were acclimated to green, blue, red, and white light. The spectral dependency in the light-limited slope of the photosynthesis–irradiance curves (α) was measured with carbon action spectra that, when divided by the spectrally weighted absorption coefficient, provided estimates of the maximum quantum yield (φ_max) for carbon fixation. Values of φ_max varied with wavelength within each culture condition as well as between different culture conditions. The degree to which the spectral dependency in φ_max was influenced by the presence of photoprotective carotenoids and/or energy imbalances between photosystems I and II was assessed for both dinoflagellates. The impact of photoprotective pigmentation on the spectral dependency of φ_max was most significant for cells grown under high light conditions reflecting the enrichment of diadinoxanthin. Energy imbalances between the photosystems was assessed by quantifying enhancement effects on spectral φ_max in the presence of background illumination. Under our experimental conditions, enhancement effects on carbon action spectra were evident for H. pygmaea under nearly all growth conditions but were not detectable for P. minimum under any growth condition. We hypothesize that sensitivity to enhancement effects reflected differences in the structure of the photosynthetic machinery of these two peridinin-containing dinoflagellates. While measurements of φ_max are sensitive to the color of the light within an incubator, the relative impact on the spectral dependency of a was less than the wavelength dependency associated with the cellular absorption properties. Finally we used our data to validate an approach proposed by others to aid in the correction of photosynthetic measurements where the in situ spectral light field cannot be easily mimicked. The average error using this approach was 8%, which was significantly less than the error associated with ignoring the spectral dependency in α.     

VARIABILITY OF PHOTOSYSTEM II QUANTUM YIELD AND RELATED PROCESSES IN CHLORELLA PYRENOIDOSA (CHLOROPHYTA) ACCLIMATED TO AN OSCILLATING LIGHT REGIME SIMULATING A MIXED PHOTIC ZONE1

The regulation of photosystem II (PSII)-related parameters was characterized for Chlorella pyrenoidosa Chick grown in continuous nutrient-replete culture and acclimated to a light field simulating fluctuations of a natural environment. The maximum quantum yield for charge separation at PSII showed a light-driven decrease during the first half of the photoperiod when incident irradiance (integrated hourly) was increasing to a midday maximum, This change was accompanied by a decrease in the effective cross-section for all PSIIs and the connectivity between PSIIs, while the cross-section for inactive centers increased. Furthermore, a decrease in the number of active PSII centers and the amount of Q_B bound to D1 became evident. During the second half of the day, as irradiance declined toward the end of the photoperiod, all the observed changes in PSII photochemistry were reversed. The low quantum yield values at the middle of the photoperiod returned to 85% of the value at the end of the photoperiod by the addition of far-red light, suggesting that a State I–II–I transition took place during the day. Measured rates of oxygen production correlated well with fluorescence-derived estimates of photosynthesis. The overall results suggest that PSII photochemistry is optimized to minimize the susceptibility to photoinhibition and to allow balanced growth under fluctuating irradiation and that the impact of these alterations on primary production can be adequately monitored through bio-optical measurements alone.     

CHROMATIC REGULATION OF QUANTUM YIELDS FOR PHOTOSYSTEM II CHARGE SEPARATION,OXYGEN EVOLUTION,AND CARBON FIXATION IN HETEROCAPSA PYGMAEA (PYRROPHYTA)1

Operational and maximum quantum yields for system (PSII) charge separation, oxygen evolution, and carbon fixation were quantified and compared for Heterocapsa pygmaea Loeblich, Schmidt et Sherley populations chromatically adapted to white, green, blue, and red light. Significant variability in quantum yields was induced by chromatic adaptation alone or when chromatically adapted cells were suddenly exposed to biased light fields (i.e. white light). Results indicated a close coupling between the variability in quantum yields for PSII charge separation and oxygen evolution, but not between quantum yields for oxygen evolution and carbon fixation. But not between quantum yields for oxygen evolution and carbon fixation. The ability to regrlate and optimize light energy distribution between PSII and photosystem I (PSI) appears to be the mechanism underlying chromatic adaptation for PSII charge separation and oxygen evolution. Conceptually, the resulting impacts on PSI cyclic electron transport rates could account for observed variability in quantum yields for oxygen evolution and some variability in quantum yields for carbon fixation. Similarly, enzymatic processes associated with organic carbon synthesis appeared to be variably dependent on spectral growth irradiances and contributing to the observed variability in quantum yields for carbon fixation. The relevance of these findings to the in situ primary production is discussed.     

Phosphorylation of PSII proteins in maize thylakoids in the presence of Pb ions

Lead is potentially toxic to all organisms including plants. Many physiological studies suggest that plants have developed various mechanisms to contend with heavy metals, however the molecular mechanisms remain unclear. We studied maize plants in which lead was introduced into detached leaves through the transpiration stream. The photochemical efficiency of PSII, measured as an Fv/Fm ratio, in the maize leaves treated with Pb was only 10% lower than in control leaves. The PSII activity was not affected by Pb ions in mesophyll thylakoids, whereas in bundle sheath it was reduced. Protein phosphorylation in mesophyll and bundle sheath thylakoids was analyzed using mass spectrometry and protein blotting before and after lead treatment. Both methods clearly demonstrated increase in phosphorylation of the PSII proteins upon treatment with Pb²⁺, however, the extent of D1, D2 and CP43 phosphorylation in the mesophyll chloroplasts was clearly higher than in bundle sheath cells. We found that in the presence of Pb ions there was no detectable dephosphorylation of the strongly phosphorylated D1 and PsbH proteins of PSII complex in darkness or under far red light. These results suggest that Pb²⁺ stimulates phosphorylation of PSII core proteins, which can affect stability of the PSII complexes and the rate of D1 protein degradation. Increased phosphorylation of the PSII core proteins induced by Pb ions may be a crucial protection mechanism stabilizing optimal composition of the PSII complexes under metal stress conditions. Our results show that acclimation to Pb ions was achieved in both types of maize chloroplasts in the same way. However, these processes are obviously more complex because of different metabolic status in mesophyll and bundle sheath chloroplasts.     

The causes of altered chlorophyll fluorescence quenching induction in the Arabidopsis mutant lacking all minor antenna complexes

Non-photochemical quenching (NPQ) of chlorophyll fluorescence is the process by which excess light energy is harmlessly dissipated within the photosynthetic membrane. The fastest component of NPQ, known as energy-dependent quenching (qE), occurs within minutes, but the site and mechanism of qE remain of great debate. Here, the chlorophyll fluorescence of Arabidopsis thaliana wild type (WT) plants was compared to mutants lacking all minor antenna complexes (NoM). Upon illumination, NoM exhibits altered chlorophyll fluorescence quenching induction (i.e. from the dark-adapted state) characterised by three different stages: (i) a fast quenching component, (ii) transient fluorescence recovery and (iii) a second quenching component. The initial fast quenching component originates in light harvesting complex II (LHCII) trimers and is dependent upon PsbS and the formation of a proton gradient across the thylakoid membrane (ΔpH). Transient fluorescence recovery is likely to occur in both WT and NoM plants, but it cannot be overcome in NoM due to impaired ΔpH formation and a reduced zeaxanthin synthesis rate. Moreover, an enhanced fluorescence emission peak at ~679?nm in NoM plants indicates detachment of LHCII trimers from the bulk antenna system, which could also contribute to the transient fluorescence recovery. Finally, the second quenching component is triggered by both ΔpH and PsbS and enhanced by zeaxanthin synthesis. This study indicates that minor antenna complexes are not essential for qE, but reveals their importance in electron stransport, ΔpH formation and zeaxanthin synthesis.     

Ambient UV-B radiation reduces PSII performance and net photosynthesis in high Arctic Salix arctica

Ambient ultraviolet-B (UV-B) radiation potentially impacts the photosynthetic performance of high Arctic plants. We conducted an UV-B exclusion experiment in a dwarf shrub heath in NE Greenland (74°N), with open control, filter control, UV-B filtering and UV-AB filtering, all in combination with leaf angle control. Two sites with natural leaf positions had ground angles of 0° (‘level site’) and 45° (‘sloping site’), while at a third site the leaves were fixed in an angle of 45° to homogenize the irradiance dose (‘fixed leaf angle site’). The photosynthetic performance of the leaves was characterized by simultaneous gas exchange and chlorophyll fluorescence measurements and the PSII performance through the growing season was investigated with fluorescence measurements. Leaf harvest towards the end of the growing season was done to determine the specific leaf area and the content of carbon, nitrogen and UV-B absorbing compounds. Compared to a 60% reduced UV-B irradiance, the ambient solar UV-B reduced net photosynthesis in Salix arctica leaves fixed in the 45° position which exposed leaves to maximum natural irradiance. Also a reduced Calvin Cycle capacity was found, i.e. the maximum rate of electron transport (J_max) and the maximum carboxylation rate of Rubisco (V_cmax), and the PSII performance showed a decreased quantum yield and increased energy dissipation. A parallel response pattern and reduced PSII performance at all three sites indicate that these responses take place in all leaves across position in the vegetation. These findings add to the evidence that the ambient solar UV-B currently is a significant stress factor for plants in high Arctic Greenland.     

A theoretical study on effect of the initial redox state of cytochrome b559 on maximal chlorophyll fluorescence level (F(M)): implications for photoinhibition of photosystem II

In this work, we extended the reversible radical pair model which describes energy utilization and electron transfer up to the first quinone electron acceptor (Q(A)) in photosystem II (PSII), by redox reactions involving cytochrome (cyt) b559. In the model, cyt b559 accepts electrons from the reduced primary electron acceptor in PSII, pheophytin, and donates electrons to the oxidized primary electron donor in PSII (P680+). Theoretical simulations of chlorophyll fluorescence rise based on the model show that the maximal fluorescence, F(M), increases with an increasing amount of initially reduced cyt b559. In this work we applied, the first to our knowledge, metabolic control analysis (MCA) to a model of reactions in PSII. The MCA was used to determine to what extent the reactions occurring in the model control the F(M) level and how this control depends on the initial redox state of cyt b559. The simulations also revealed that increasing the amount of initially reduced cyt b559 could protect PSII against photoinhibition. Also experimental data, which might be used to validate our theory, are presented and discussed.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.