PHOTOSYNTHESIS IN AN EXTREME SHADE ENVIRONMENT: BENTHIC MICROBIAL MATS FROM LAKE HOARE, A PERMANENTLY ICE-COVERED ANTARCTIC LAKE

We investigated the composition of benthic microbial mats in permanently ice-covered Lake Hoare, Antarctica, and their irradiance vs. photosynthetic oxygen exchange relationships. Mats could be subdivided into three distinct depth zones: a seasonally ice-free “moat” zone and two under-ice zones. The upper under-ice zone extended from below the 3.5 m thick ice to approximately 13 m and the lower from below 13 m to 22 m. Moat mats were acclimated to the high irradiance they experienced during summer. They contained photoprotective pigments, predominantly those characteristic of cyanobacteria, and had high compensation and saturating irradiances (E_c and E_k) of 75 and 130 μmol photons·m⁻²·s⁻¹, respectively. The moat mats used light inefficiently. The upper under-ice community contained both cyanobacteria and diatoms. Within this zone, biomass (as pigments) increased with increasing depth, reaching a maximum at 10 m. Phycoerythrin was abundant in this zone, with shade acclimation and efficiency of utilization of incident light increasing with depth to a maximum of 0.06 mol C fixed·mol⁻¹ incident photons under light-limiting conditions. Precipitation of inorganic carbon as calcite was associated with this community, representing up to 50% of the carbon sequestered into the sediment. The lower under-ice zone was characterized by a decline in pigment concentrations with depth and an increasing prevalence of diatoms. Photosynthesis in this community was highly shade acclimated and efficient, with E_c and E_k below 0.5 μmol·m⁻²·s⁻¹ and 2 μmol·m⁻²·s⁻¹, respectively, and maximum yields of 0.04 mol C fixed·mol⁻¹ incident quanta. Carbon uptake in situ by both under-ice and moat mats was estimated at up to 100 and 140 mg·m⁻²·day⁻¹, based on the photosynthesis–irradiance curves, incident irradiance, and light attenuation by ice and the water column.     

Bio-optical Characteristics and the Vertical Distribution of Photosynthetic Pigments and Photosynthesis in an Artificial Cyanobacterial Mat

Abstract Zonations of photosynthesis and photopigments in artificial cyanobacterial mats were studied with (i) oxygen and pH microsensors, (ii) fiber-optic microprobes for field radiance, scalar irradiance, and PSII fluorescence, and (iii) a light microscope equipped with a spectrometer for spectral absorbance and fluorescence measurements. Our analysis revealed the presence of several distinct 1–2 mm thick cyanobacterial layers mixed with patches of anoxygenic photosynthetic bacteria. Strong attenuation of visible light confined the euphotic zone to the uppermost 3 mm of the mat, where oxygen levels of 3–4 times air saturation and a pH peak of up to pH 8.8 were observed under saturating irradiance (413 μmol photon m⁻² s⁻¹). Oxygen penetration was 5 mm in light and decreased to 1 mm in darkness. Volumetric oxygen consumption in the photic and aphotic zones of illuminated mat was 5.5 and 2.9 times higher, respectively, than oxygen consumption in dark incubated mats. Scalar irradiance reached 100–150% of incident irradiance in the upper 0.5 mm of the mat due to intense scattering in the matrix of cells, exopolymers, and carbonate precipitates. In deeper mat layers scalar irradiance decreased nearly exponentially, and highest attenuation coefficients of 6–7 mm⁻¹ were found in cyanobacterial layers, where photosynthesis and photopigment fluorescence also peaked. Visible light was attenuated >100 times more strongly than near infrared light. Microscope spectrometry on thin sections of mats allowed detailed spectral absorbance and fluorescence measurements at defined positions relative to the mat surface. Besides strong spectral signals of cyanobacterial photopigments (Chl a and phycobiliproteins), the presence of both green and purple photosynthetic bacteria was evident from spectral signals of Bchl a and Bchl c. Microprofiles of photopigment absorbance correlated well with microdistributions of phototrophs determined in an accompanying study. Received: 20 December 1999; Accepted: 10 June 2000; Online Publication: 28 August 2000     

ANTARCTIC CYANOBACTERIA: LIGHT,NUTRIENTS, AND PHOTOSYNTHESIS IN THE MICROBIAL MAT ENVIRONMENT1

The microenvironmental and photosynthetic characteristics of Antarctic microbial mats were measured in a series of ponds near McMurdo Sound. As elsewhere in Antarctica, these cold-water benthic communities were dominated by oscillatoriacean cyanobacteria. Despite large variations in mat thickness, surface morphology, and color, all of the communities had a similar pigment organization, with a surface carotenoid-rich layer that overlaid a deep chlorophyll maximum (DCM) enriched in phycocyanin as well as chlorophyll a. Spectroradiometric analyses showed that the DCM population inhabited an orange-red shade environment. In several of the mats, the deep-living trichomes migrated up to the surface of the mat within 2 h in response to a 10-fold decrease in surface irradiance. The euphotic layer of the mats was supersaturated in oxygen and contained ammonium and dissolved reactive phosphorus concentrations in excess of 100 mg N·m^?3 or P·m^?3. Integral photosynthesis by core samples was saturated at low irradiances and varied two- to threefold throughout the continuous 24-h radiation cycle. Oxygen microelectrode analyses showed that the photosynthetic rates were slow to negligible near the surface and maximal in the DCM. These compressed, nutrient-rich euphotic zones have some properties analogous to planktonic systems, but the integrated photosynthetic responses of the community reflect the strong self-shading within the mat and physiological dominance by the motile, DCM populations.     

The interaction of components controlling net phytoplankton photosynthesis in a well-mixed lake (Lough Neagh, Northern Ireland)

The homogeneous distribution of the phytoplankton in a shallow (mean depth 8·6 m) unstratified lake, L. Neagh, Northern Ireland, facilitated the study of the interaction of components controlling gross photosynthesis per unit area. These included the photosynthetic capacity, the phytoplankton content of the euphotic zone, and a logarithmic function describing the effective radiation input. These factors were analysed for two sites, the open lake and Kinnego Bay, which respectively had standing crops of up to 90 and 300 mg chlorophyll a m^?3 and maximum daily rates of gross integral photosynthesis of 11·7 and 15·6 g O₂ m^?2 day^?1. Values are reduced by the high contribution to light attenuation by non-algal sources, which increases at low standing crops particularly in winter, when values of integral photosynthesis decrease to 0·5 g O₂ m^?2 day^?1. This relative change is the result of self-shading behaviour of the phytoplankton altering the crop content of the euphotic zone at different population densities. Changes in the irradiance function, incorporating day length, are largely responsible for the changes in daily rates of integral gross photosynthesis; as daily irradiance is also a determinant of water temperature, it exerts further influence through the photosynthetic capacity which was strongly correlated with temperature. Much of the gain in gross photosynthesis resulting from higher photosynthetic capacity may not be reflected in a higher net column photosynthesis, because of the greater proportional rise in respiration with temperature. The balance in the water column between respiration losses and photosynthetic input may frequently alter since the ratio of illuminated to dark zones is between 1/4 to 1/5 in the open lake, and small shifts in any of the controlling features may result in conditions unfavourable for growth. This is analysed especially for the increase of diatoms in spring, when small modifications of the underwater light field can delay growth.     

MICROENVIRONMENTAL CONTROL OF PHOTOSYNTHESIS AND PHOTOSYNTHESIS-COUPLED RESPIRATION IN AN EPILITHIC CYANOBACTERIAL BIOFILM1

The photosynthetic performance of an epilithic cyano-bacterial biofilm was studied in relation to the in situ light field by the use of combined microsensor measurements of O₂, photosynthesis, and spectral scalar irradiance. The high density of the dominant filamentous cyanobacteria (Oscillatoria sp.) embedded in a matrix of exopolymers and bacteria resulted in a photic zone of < 0.7 mm. At the biofilm surface, the prevailing irradiance and spectral composition were significantly different from the incident light. Multiple scattering led to an intensity maximum for photic light (400–700 nm) of ca. 120% of incident quantum irradiance at the biofilm surface. At the bottom of the euphotic zone in the biofilm, light was attenuated strongly to < 5–10% of the incident surface irradiance. Strong spectral signals from chlorophyll a (440 and 675 nm) and phycobilins (phycoerythrin 540–570 nm, phycocyanin 615–625 nm) were observed as distinct maxima in the scalar irradiance attenuation spectra in the upper 0.0–0.5 mm of the biofilm. The action spectrum for photosynthesis in the cyanobacterial layer revealed peak photosynthetic activity at absorption wavelengths of phycobilins, whereas only low photosynthesis rates were induced by light absorption of carotenoids (450–550 nm). Respiration rates in light- and dark-incubated biofilms were determined using simple flux calculations on measured O₂ concentration profiles and photosynthetic rates. A significantly higher areal O₂ consumption was found in illuminated biofilms than in dark-incubated biofilms. Although photorespiration accounted for part of the increase, the enhanced areal O₂ consumption of illuminated biofilms could also be ascribed to a deeper oxygen penetration in light as well as an enhanced volumetric O₂ respiration in and below the photic zone. Gross photosynthesis was largely unaffected by increasing flow velocities, whereas the O₂ flux out of the photic zone, that is, net photosynthesis, increased with flow velocity. Consequently, the amount of produced O₂ consumed within the biofilm decreased with increasing flow velocity. Our data indicated a close coupling of photosynthesis and respiration in biofilms, where the dissolved inorganic carbon requirement of the photo-synthetic population may largely be covered by the respiration of closely associated populations of heterotrophic bacteria consuming a significant part of the photosynthetically produced oxygen and organic carbon.     

喜光榕树和耐荫榕树光适应机制的差异

100%和36%光强下生长的喜光的斜叶榕的光合能力高于耐荫的假斜叶榕,而热耗散能力与之相似,说明强光下斜叶榕主要通过光合作用利用光能和热耗散、假斜叶榕主要通过热耗散防御光破坏.100%光强下生长的两种榕树的日间光抑制程度相似,但叶表光强相同情况下各光强下生长的假斜叶榕的光抑制均比斜叶榕严重.100%光强下假斜叶榕叶片悬挂角大于斜叶榕,导致日间叶表光强低于斜叶榕,这可能是两种榕树日间光抑制程度相似的原因,表明叶片悬挂角的适应变化对假斜叶榕有重要的意义.     

Acclimation of the diatom Stephanodiscus neoastraea and the cyanobacterium Planktothrix agardhii to simulated natural light fluctuations

Functional and structural characteristics of the photosynthetic apparatus were studied in the diatom Stephanodiscus neoastraea and the cyanobacterium Planktothrix agardhii which were grown semi-continuously under constant irradiance or under simulated natural light fluctuations. The light fluctuations consisted of 24 oscillations of exponentially increasing and decreasing irradiance over a 12-h light period. Maximum irradiance was 1100 μmol photons m⁻² s⁻¹ with the ratio of maximum to minimum intensities being 100, simulating Langmuir circulations with a ratio of euphotic to mixing depth of 1. S. neoastraea acclimated to the light fluctuations by doubling the number and halving the size of photosynthetic units (PS II) while the amount of chlorophylls and carotenoids remained unchanged. The chlorophyll-specific maximum photosynthetic rate was enhanced while the slope of the photosynthesis versus irradiance curves was not influenced by the light fluctuations. Acclimation of P. agardhii was mainly characterized by an increase in chlorophyll content. Both photosystems showed only little changes in number and size. Maximum photosynthetic rate, saturating irradiance and initial slope of the photosynthesis versus irradiance curves did not vary. Although both high and low light were contained in the fluctuating light, an analogy to low or high light acclimation was not found for the diatom nor for the cyanobacterium acclimated to light fluctuations. We suggest that the acclimation to fluctuating light is a response type outside the known scheme of low and high light acclimation. This revised version was published online in June 2006 with corrections to the Cover Date.     

热带雨林三种树苗叶片光合机构对光强的适应

对生长在不同光强(自然日光的8％,25％,50％)下西双版纳热带雨林3种木本植物团花(Anthocephalus chinensis)、玉蕊(Barringtonia pendala)和藤黄(Garrcinia hanburyi)幼苗光合机构的研究表明,随着生长光强的升高,植物叶片的光饱和点、补偿点、净光合速率和非光化学淬灭系数(NPQ)升高,而表现量子效率(AQY)、有效光化学量子产量(Fv／Fm)、光化学淬灭系数(qP)下降．在抗氧化系统中,超氧化物歧化酶(SOD)、抗坏血酸过氧化酶(APX)活性随着光强的升高而升高,而过氢化物酶(CAT)活性与生长光强的变化不一致．抗坏血酸(AsA)含量随着光强的升高而急剧上升。最能反映PFD的变化．可以认为,除与叶黄素循环有关的热耗散增大之外,植物叶片抗氧化系统的加强也是响应强光的一种保护措施．     

Action spectra of microalgal photosynthesis and depth distribution of spectral scalar irradiance in a coastal marine sediment of Limfjorden, Denmark

Abstract The role of complementary spectral utilization of light for the zonation of different groups of oxygenic phototrophic organisms in sediments was studied. The marine sediment was covered by a dense population of diatoms with an underlying population of cyanobacteria. Action spectra for photosynthesis and spectral scalar irradiance, E ₀, were measured directly in the sediment at a spatial resolution of 0.1 mm by the use of oxygen and light microsensors. The action spectrum for the diatoms was similar to the attenuation spectrum of the scalar irradiance, K ₀, in the diatom layer with Chl. a and carotenoids being the major photosynthetic pigments. The action spectrum of the cyanobacteria showed photosynthesis maxima at the absorption regions of Chl. a and phycocyanin. The measured depth distribution of spectral scalar irradiance and the action spectra of diatoms and cyanobacteria were used to calculate the spectral quality for photosynthesis of the 400–700 nm light to which the two populations were exposed. This spectral quality was compared to that of the light incident on the sediment surface. Due to preferential extinction of wavelengths, at which their photosynthetically active pigments had maximal absorption, the relative light quality for diatoms was reduced to 85% of the quality of incident light at a similar total quantum flux. This effect was partly due to spectral alterations of light backscattered from the underlying sediment with cyanobacteria. The cyanobacteria at the bottom of the euphotic zone, in contrast, experienced a light spectrum which was favorably altered, to 107% in quality, due to absorption by the overlying diatoms. It was concluded that these changes in spectral light quality can be considered as only one of more factors explaining the zonation of the two phototrophic populations, and that total light intensity and the chemical microenvironment are probably more important factors.     

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST AND SOUTHWEST SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. I. IRRADIANCE TOLERANCE

Deciduous and evergreen species are segregated on northeast and southwest slopes of the southern Appalachian Mountains. The segregated distributions of three ericaceous shrubs (Rhododendron maximum valley positions; Rhododendron periclymenoides on northeast slopes; Kalmia latifolia on southwest slopes) were compared to the respective irradiance environments. Growth patterns of field plants, and photosynthetic acclimation of each species to three irradiance treatments in a phytotron were studied. Rhododendron maximum, an evergreen species, was found to be most sensitive to high radiation. In phytotron experiments, quantum yield, light saturated photosynthetic capacity, photosynthesis per chlorophyll, and water use efficiency decreased at high ambient irradiance for R. maximum. These characteristics limit the growth of R. maximum on high irradiance southwestern slopes. Both K. latifolia and R. periclymenoides were able to improve their photosynthetic performance at high ambient irradiance. Rhododendron periclymenoides, a deciduous species, was found to continue increasing leaf conductance at high irradiance without an increase in photosynthesis indicating a possible limitation by water in high light environments such as southwest slopes. Kalmia latifolia, an evergreen species, had reduced photosynthetic capacity and reduced water use efficiency when grown in low irradiance conditions which coincides with the higher K. latifolia abundance on high light, southwestern slopes.     

Mathematical simulation of the diel O, S, and C biogeochemistry of a hypersaline microbial mat

The creation of a mathematical simulation model of photosynthetic microbial mats is important to our understanding of key biogeochemical cycles that may have altered the atmospheres and lithospheres of early Earth. A model is presented here as a tool to integrate empirical results from research on hypersaline mats from Baja California Sur (BCS), Mexico into a computational system that can be used to simulate biospheric inputs of trace gases to the atmosphere. The first version of our model, presented here, calculates fluxes and cycling of O(2), sulfide, and dissolved inorganic carbon (DIC) via abiotic components and via four major microbial guilds: cyanobacteria (CYA), sulfate reducing bacteria (SRB), purple sulfur bacteria (PSB) and colorless sulfur bacteria (CSB). We used generalized Monod-type equations that incorporate substrate and energy limits upon maximum rates of metabolic processes such as photosynthesis and sulfate reduction. We ran a simulation using temperature and irradiance inputs from data collected from a microbial mat in Guerrero Negro in BCS (Mexico). Model O(2), sulfide, and DIC concentration profiles and fluxes compared well with data collected in the field mats. There were some model-predicted features of biogeochemical cycling not observed in our actual measurements. For instance, large influxes and effluxes of DIC across the MBGC mat boundary may reveal previously unrecognized, but real, in situ limits on rates of biogeochemical processes. Some of the short-term variation in field-collected mat O(2) was not predicted by MBGC. This suggests a need both for more model sensitivity to small environmental fluctuations for the incorporation of a photorespiration function into the model.     

Action spectra of microalgal photosynthesis and depth distribution of spectral scalar irradiance in a coastal marine sediment of Limfjorden, Denmark

The role of complementary spectral utilization of light for the zonation of different groups of oxygenic phototrophic organisms in sediments was studied. The marine sediment was covered by a dense population of diatoms with an underlying population of cyanobacteria. Action spectra for photosynthesis and spectral scalar irradiance, E₀, were measured directly in the sediment at a spatial resolution of 0.1 mm by the use of oxygen and light microsensors. The action spectrum for the diatoms was similar to the attenuation spectrum of the scalar irradiance, K₀, in the diatom layer with Chl.a. and carotenoids being the major photosynthetic pigments. The action spectrum of the cyanobacteria showed photosynthesis maxima at the absorption regions of Chl.a. and phycocyanin. The measured depth distribution of spectral scalar irradiance and the action spectra of diatoms and cyanobacteria were used to calculate the spectral quality for photosynthesis of the 400–700 nm light to which the two populations were exposed. This spectral quality was compared to that of the light incident on the sediment surface. Due to preferential extinction of wavelengths, at which their photosynthetically active pigments had maximal absorption, the relative light quality for diatoms was reduced to 85% of the quality of d incident light at a similar total quantum flux. This effect was partly due to spectral alterations of light backscattered from the underlying sediment with cyanobacteria. The cyanobacteria at the bottom of the euphotic zone, in contrast, experienced a light spectrum which was favorably altered, to 10% in quality, due to absorption by the overlying diatoms. It was concluded that these changes in spectral light quality can be considered as only one of more factors explaining the zonation of the two phototrophic populations, and that total light intensity and the chemical microenvironment are probably more important factors.     

Hydrogen sulfide can inhibit and enhance oxygenic photosynthesis in a cyanobacterium from sulfidic springs

We used microsensors to investigate the combinatory effect of hydrogen sulfide (H₂S) and light on oxygenic photosynthesis in biofilms formed by a cyanobacterium from sulfidic springs. We found that photosynthesis was both positively and negatively affected by H₂S: (i) H₂S accelerated the recovery of photosynthesis after prolonged exposure to darkness and anoxia. We suggest that this is possibly due to regulatory effects of H₂S on photosystem I components and/or on the Calvin cycle. (ii) H₂S concentrations of up to 210 μM temporarily enhanced the photosynthetic rates at low irradiance. Modelling showed that this enhancement is plausibly based on changes in the light‐harvesting efficiency. (iii) Above a certain light‐dependent concentration threshold H₂S also acted as an inhibitor. Intriguingly, this inhibition was not instant but occurred only after a specific time interval that decreased with increasing light intensity. That photosynthesis is most sensitive to inhibition at high light intensities suggests that H₂S inactivates an intermediate of the oxygen evolving complex that accumulates with increasing light intensity. We discuss the implications of these three effects of H₂S in the context of cyanobacterial photosynthesis under conditions with diurnally fluctuating light and H₂S concentrations, such as those occurring in microbial mats and biofilms.     

48 The effects of environmental factors on the resumption of photosynthetic activity of cyanobacterial mats following rehydration

Extensive cyanobacterial mats cover the intertidal zone near Guerrero Negro, Baja California Sur. These mats are exposed to extreme desiccation and osmotic stress between tidal flows and rains, and spend most of the time dry and metabolically inactive. Therefore, periods of hydration are extremely important for growth as well as for repair of cellular damage from desiccation and ultraviolet radiation (UVR) accrued when the mat is dry. PAM fluorometry in conjunction with carbon incorporation assays were used to determine the effects of salinity, irradiance and UVR on the recovery of photosynthetic activity in these mats after an extended period of desiccation. The mat used in our study was primary composed of Lyngbya sp. Photosynthetic activity recovery rates (using PAM fluorometry) decreased with increasing salinity. This trend was similar under high and low light intensities, but rates were significantly lower under low light. Alternatively, the carbon incorporation method showed rates increased faster in salinities of 27 and 55 ppt than in salinities of 0 or 75 ppt. The Lyngbya mat also failed to recover photosynthetic potential in the dark. Although these mats recovered faster under high intensity light, the effect of salinity on photosynthesis is more complex. UVR did not affect the recovery of photosynthetic activity, no matter which method was used. This lack of effect is most likely due to the high content of the UVR screening pigment, scytonemin, in the upper layer of the mat.     

ABSORPTION AND UTILIZATION OF IRRADIANCE BY CYANOBACTERIAL MATS IN TWO ICE-COVERED ANTARCTIC LAKES WITH CONTRASTING LIGHT CLIMATES

We investigated the under-ice light climate and the efficiency with which light was absorbed and utilized by benthic algal mats in Lakes Hoare and Vanda, two perennially ice-covered lakes in the McMurdo Dry Valleys area of Southern Victoria Land, Antarctica. The ice cover and water column of Lake Vanda were much more transparent than those of Lake Hoare (18% vs. 2% transmission though ice and attenuation coefficients for downwelling irradiance of 0.05 vs. 0.12 m ^{− 1}, respectively). In both lakes the under-ice spectra were dominated by blue-green wavelengths. The benthic flora under perennial ice covers of both lakes comprised thick mucilaginous mats, dominated by cyanobacteria. The mats were well suited to absorb the dominant blue-green wavelengths of the under-ice light, with phycoerythrin being present at high concentrations. The pigment systems of the benthic mats absorbed 30%–50% of the light that reached them, varying with depth and lake. There was a tendency for the percentage of absorption to increase as ambient irradiance decreased. The efficiency of utilization of absorbed irradiance was examined by constructing absorbed irradiance/oxygen evolution curves to estimate community quantum yield. Mats from 13 m in Lake Hoare showed the highest quantum yields, approaching 1 mol of carbon fixed for every 8 mol quanta absorbed under light-limiting conditions. Lake Vanda mats had lower quantum yields, but these increased with depth. Calculated in situ irradiance occasionally exceeded the measured saturating irradiance for oxygen evolution in both lakes, thus efficiency in situ was below the maximum at times. As in other environments, optimization strategies allowed efficient capture and utilization of the lower and middle ranges of experienced irradiance but led to a compromised capacity to use the highest irradiances encountered at each depth.     

Metabolic shifts in hypersaline microbial mats upon addition of organic substrates

The responses of hypersaline microbial mats to the addition of acetate, glycolate or glucose were investigated using oxygen, pH and sulphide microsensors. Changes in community structure were investigated with molecular techniques. Acetate addition inhibited respiration in the photic zone, stimulated respiration in the aphotic zone and had no effect on gross photosynthesis. Glycolate addition strongly increased both respiration and gross photosynthesis in the photic zone. Thus, glycolate and acetate were probably consumed in those regions of the mat where these substrates are usually formed. Moreover, photosynthesis was only stimulated by increased respiration and concomitant CO2 production in the photic zone which indicates that the photosynthetic and respiratory populations must be present in close proximity to each other. Glucose addition had an unexpected negative effect on the microbial population, strongly inhibiting both respiration and gross photosynthesis within hours. After four days, oxygen profiles in the light were equal to those measured in the dark. After replacing the water phase with unamended water, photosynthesis and respiration recovered within a week. None of the physiological changes were accompanied by detectable shifts in the cyanobacterial or the overall microbial community. The mechanism of inhibition of photosynthesis by glucose requires further investigation.     

SEASONAL VARIATIONS IN PHOTOSYNTHESIS-IRRADIANCE RESPONSES BY BIOFILMS IN MEDITERRANEAN STREAMS

The relationships between primary production and irradiance were analyzed over an annual cycle in natural biofilms of two undisturbed streams: La Solana (LS), an open calcareous stream, and Riera Major (RM), a shaded siliceous stream. In LS, low photosynthetic efficiency (α^chl and α^area) and high values of both the light saturation parameter (I_k) and the carotenoid / chlorophyll ratio indicated adaptation to high light regimes. On the other hand, higher photosynthetic efficiency and lower I_k as well as photoinhibition at high irradiance found in the biofilms of RM indicated shade adaptation. However, the lack of correlation between light availability in nature and the photosynthetic parameters studied in the laboratory suggested that light was not the most important factor in determining seasonal changes in the photosynthetic behavior in this stream. Both in the open and shaded streams, algal patches collected simultaneously exhibited different photosynthesis-irradiance (P-I) curues, showing that community composition influenced the P-I parameters. In the open stream (LS), however, significant negative correlations between α^area and chlorophyll a and between P _max^chl and chlorophyll a ( r = -0.994 , P < 0.001, and r = -0.929 , P < 0.05, respectively) suggested that photosynthesis was affected by self-shading. Due to the absence of photoinhibition in the biofilms of LS, high photosynthetic rates were maintained at the ambient high light environment, thus compensating for low photosynthesis at low irradiance. In the shaded stream (RM), because photosynthesis was saturated at low irradiances, primary production was relatively high given the low light conditions .     

Flexible and reversible responses to different irradiance levels during photosynthetic acclimation of Cypripedium guttatum

Cypripedium guttatum can be found both in open and shady habitats. Photosynthetic acclimation of C. guttatum to different light availabilities was detected using measurements of chlorophyll fluorescence, photosynthesis and leaf traits. When growing under low light conditions, C. guttatum exhibited a greater efficiency in photochemical utilization of absorbed light energy, and a lower ability for non-photochemical dissipation of excess light energy, as compared to the plants growing under high light conditions. Under intermediate light conditions, C. guttatum exhibited higher photosynthetic capacity (A(max)) than those under both low light or high light conditions. The differences in A(max) among three light environments was linked to the differences in biochemical efficiency, leaf N content (LNC) and leaf dry mass per unit area (LMA), but not to the differences of chlorophyll content. However, there were no significant differences in the light compensation points (LCP) and light saturation points (LSP) for photosynthesis for the plants growing under the three light conditions. These results indicate that the photosynthetic capacity of C. guttatum leaves allows for flexible and reversible responses to different irradiance levels. Photosynthetic acclimation in C. guttatum was affected by biochemical changes, the changes in LMA and ratio of Chl a/b. Successful acclimation of C. guttatum to a broad range of light levels likely allows for its wide geographical distribution. A level of about 45% sunlight appears to be optimal for photosynthesis.     

Acclimation of photosynthesis to high irradiance in rice: gene expression and interactions with leaf development

Rice (Oryza sativa L.) has been used to study the long-term responses of photosynthesis to high irradiance focusing on the composition of the photosynthetic apparatus and leaf morphology. Typical sun/shade differences in chloroplast composition are seen in the fifth leaf following growth in high irradiance compared with low irradiance (1000 and 200 micromol m(-2) s(-1), respectively): higher light-saturated rates of photosynthesis (P(max)), higher amounts of Rubisco protein, and a lower chlorophyll a:b ratio. In addition, leaves were thicker under high light compared with low light. However, responses appear more complex when leaf developmental stage is considered. Using a system of transferring plants from low to high light in the laboratory responses that occur before and after full leaf extension have been studied. Acclimation of photosynthesis is limited by leaf age: the transfer to high light, post-leaf extension, is characterized by alterations in chlorophyll a:b but not in Rubisco protein, which may be limited by leaf morphology. Microarray analysis of gene expression was carried out on plants that were transferred to high light post-leaf extension. A down-regulation of light-harvesting genes was seen. No change in the expression level of Rubisco genes was observed. Up-regulation of genes involved in photoprotection was observed. It was also shown that high-light leaf morphology is established prior to formation of the zone of cellular elongation and division. The endogenous and environmental factors which establish the characteristics of high light acclimation may be important for attaining high rates of assimilation in leaves and crop canopies, and the fifth leaf in rice provides a convenient model system for the determination of the mechanisms involved.     

A METHOD TO IMPROVE THE SPATIAL RESOLUTION OF PHOTOSYNTHETIC RATES OBTAINED BY OXYGEN MICROSENSORS

To determine gross photosynthesis in benthic microalgal communities, oxygen microelectrodes were used to measure the rate of decrease within the first 4 s after extinction of light. Photosynthetic rates calculated from third-order polynomial fits to the curve of decreasing O₂ concentration were compared to the rates obtained by the traditional method, where rates were estimated from linear regression. When photosynthesis was calculated for the fitted initial rates of O₂ decrease, maximum rates in microbial mats were up to 32% higher, and the depth-integrated gross photosynthesis was 5%–10% higher than the rates determined by the traditional method. The determinations from fitted initial rates also resulted in a more detailed profile of photosynthetic rate than that normally obtained. Computer simulation based on diffusion models, where the estimated initial rates of O₂ decrease were assumed to represent actual photosynthesis rates, verified the validity of the curve-fitting procedure for obtaining high-resolution photosynthesis profiles.