The challenge of estimating kelp production in a turbid marine environment

Coastal kelp forests produce substantial marine carbon due to high annual net primary production (NPP) rates, but upscaling of NPP estimates over time and space remains difficult. We investigated the impact of variable underwater photosynthetically active radiation (PAR) and photosynthetic parameters on photosynthetic oxygen production of Laminaria hyperborea, the dominant NE-Atlantic kelp species, throughout summer 2014. Collection depth of kelp had no effect on chlorophyll a content, pointing to a high photoacclimation potential of L. hyperborea towards incident light. However, chlorophyll a and photosynthesis versus irradiance parameters differed significantly along the blade gradient when normalized to fresh mass, potentially introducing large uncertainties in NPP upscaling to whole thalli. Therefore, we recommend a normalization to kelp tissue area, which is stable over the blade gradient. Continuous PAR measurements revealed a highly variable underwater light climate at our study site (Helgoland, North Sea) in summer 2014, reflected by PAR attenuation coefficients (K_d) between 0.28 and 0.87 m⁻¹. Our data highlight the importance of continuous underwater light measurements or representative average values using a weighted K_d to account for large PAR variability in NPP calculations. Strong winds in August increased turbidity, resulting in a negative carbon balance at depths >3–4 m over several weeks, considerably impacting kelp productivity. Estimated daily summer NPP over all four depths was 1.48 ± 0.97 g C · m⁻² seafloor · d⁻¹ for the Helgolandic kelp forest, which is in the range of other kelp forests along European coastlines.     

Effect of PAR irradiance intensity on Phaeocystis antarctica (Prymnesiophyceae) growth and DMSP,DMSO, and acrylate concentrations

Phaeocystis antarctica forms extensive spring blooms in the Southern Ocean that coincide with high concentrations of dimethylsulfoniopropionate (DMSP), dimethylsulfoxide (DMSO), dimethylsulfide (DMS), and acrylate. We determined how concentrations of these compounds changed during the growth of axenic P. antarctica cultures exposed to light-limiting, sub-saturating, and saturating PAR irradiances. Cellular DMSP concentrations per liter cell volume (CV) ranged between 199 and 403 mmol · L_CV⁻¹, with the highest concentrations observed under light-limiting PAR. Cellular acrylate concentrations did not change appreciably with a change in irradiance level or growth, ranging between 18 and 45 mmol · L_CV⁻¹, constituting an estimated 0.2%–2.8% of cellular carbon. Both dissolved acrylate and DMSO increased substantially with irradiance during exponential growth on a per-cell basis, ranging from 0.91 to 3.15 and 0.24 to 1.39 fmol · cell⁻¹, respectively, indicating substantial export of these compounds into the dissolved phase. Average cellular DMSO:DMSP ratios increased 7.6-fold between exponential and stationary phases of batch growth, with a 3- to 13-fold increase in cellular DMSO likely formed from abiotic reactions of DMSP and DMS with reactive oxygen species (ROS). At mM levels, cellular DMSP and acrylate are proposed to serve as de facto antioxidants in P. antarctica not regulated by oxidative stress or changes in ROS. Instead, cellular DMSP concentrations are likely controlled by other physiological processes including an overflow mechanism to remove excess carbon via acrylate, DMS, and DMSO during times of unbalanced growth brought on by physical stress or nutrient limitation. Together, these compounds should aid P. antarctica in adapting to a range of PAR irradiances by maintaining cellular functions and reducing oxidative stress.     

Photosynthetic performance,DNA damage and repair in gametes of the endemic Antarctic brown alga Ascoseira mirabilis exposed to ultraviolet radiation

Abstract Stress physiology on the reproductive cells of Antarctic macroalgae remained unstudied. Ascoseira mirabilis is endemic to the Antarctic region, an isolated ecosystem exposed to extreme environmental conditions. Moreover, stratospheric ozone depletion leads to increasing ultraviolet radiation (280–400 nm) at the earth's surface, thus it is necessary to investigate the capacity of reproductive cells to cope with different UV irradiances. This study is aimed to investigate the impact of exposure to different spectral irradiance on the photosynthetic performance, DNA damage and gamete morphology of the A. mirabilis. Gametangia, gametes and zygotes of the upper sublittoral brown alga A. mirabilis were exposed to photosynthetically active radiation (PAR = P; 400–700 nm), P + UV‐A radiation (UV‐A, 320–400 nm) and P + UV‐A + UV‐B radiation (UV‐B, 280–320 nm). Rapid photosynthesis versus irradiance curves of freshly released propagules were measured. Photosynthetic efficiencies and DNA damage (in terms of cyclobutane pyrimidine dimers) were determined after 1, 2, 4 and 8 h exposure as well as after 2 days of recovery in dim white light. Saturation irradiance (I_k) in freshly released propagules was 52 μmol photons m⁻² s⁻¹. Exposure for 1 h under 22 μmol photons m⁻² s⁻¹ of PAR significantly reduced the optimum quantum yield (F_v/F_m), suggesting that propagules are low light adapted. Furthermore, UVR significantly contributed to the photoinhibition of photosynthesis. Increasing dose as a function of exposure time additionally exacerbated the effects of different light treatments. The amount of DNA damage increased with the UV‐B dose but an efficient repair mechanism was observed in gametes pre‐exposed to a dose lower than 5.8 × 10³ J m⁻² of UV‐B. The results of this study demonstrate the negative impact of UV‐B radiation. However, gametes of A. mirabilis are capable of photosynthetic recovery and DNA repair when the stress factor is removed. This capacity was observed to be dependent on the fitness of the parental sporophyte.     

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.     

Photosynthetic characteristics of charophytes from tropical lotic ecosystems

Responses of photosynthetic rates, determined by oxygen evolution using the light and dark bottles technique, to different temperatures, irradiances, pH, and diurnal rhythm were analyzed under laboratory conditions in four charophyte species (Chara braunii Gmelin, C. guairensis R. Bicudo, Nitella subglomerata A. Braun and Nitella sp.) from lotic habitats in southeastern Brazil. Parameters derived from the photosynthesis versus irradiance curves indicated affinity to low irradiances for all algae tested. Some degree of photoinhibition, [β= ‐(0.30–0.13) mg O2 g^?1 dry weight Ir¹ (μmol photons m^?2 s^?1)^?1], low light compensation points (I_c= 4–20 μmol photons m^?2 s^?1) were found for all species analyzed, as well as low values of light saturation parameter (I_k) and saturation (I_s) 29–130 and 92–169 μmol photons m^?2 s^?1, respectively. Photoacclimation was observed in two populations of N. subglomerata collected from sites with different irradiances, consisting of variations in photosynthetic parameters (higher values of a, and lower of I_k and maximum photosynthetic rate, P_max, in the population under lower irradiance). The highest photosynthetic rates for Chara species were observed at 10–15°C, while for Nitella the highest photosynthetic rate was observed at 20–25°C, despite the lack of significant differences among most levels tested. Rates of dark respiration significantly increase with temperature, with the highest values at 25°C. The results from pH experiments showed highest photosynthetic rates under pH 4.0 for all algae, suggesting higher affinity for inorganic carbon in the form of carbon dioxide, except in one population of N. subglomerata, with similar rates under the three levels, suggesting indistinct use of bicarbonate and carbon dioxide. Diurnal changes in photosynthetic rates revealed a general pattern for most algae tested, which was characterized by two peaks: the first (higher) during the morning (07.00–11.00) and the second (lower) in the afternoon (14.00–17.00). This suggests an endogenous rhythm determining the daily variations in photosynthetic rates.     

Annual carbon fixation in terrestrial populations of Nostoc commune (Cyanobacteria) from an Antarctic dry valley is driven by temperature regime

Nostoc commune Vaucher (a cyanobacterium) is a very conspicuous terrestrial primary producer in Victoria Land, continental Antarctica. Because polar ecosystems are considered to be especially sensitive to environmental changes, understanding the environmental constraints on net carbon (C) fixation by N. commune is necessary to determine the effects of environmental changes on the ecological functioning of ice‐free areas of the continent. A model describing net C fixation in terrestrial populations of N. commune in an Antarctic dry valley was constructed using field and laboratory measurements in which N. commune colonies were exposed to different combinations of incident irradiance (400–700 nm), temperature, and degree of desiccation. For desiccated N. commune mats with water content ≤ 30% saturation, net C fixation was highly variable between replicates and could not be modelled. However, for colonies at > 30% saturation, rates of net C fixation and dark respiration depended strongly on irradiance and temperature. Net C fixation reached a maximum rate of 21.6 μg C m^{− 2} s^{− 1} at irradiance of approximately 250 μmol m^{− 2} s^{− 1} and the optimum temperature of 20.5 °C. Agreement between predicted short‐term net C fixation and field and laboratory measurements allowed estimation of total seasonal fixation, using previously published environmental data. Annual net C fixation was estimated in the range 14.5–21.0 g C fixed m^{− 2}Nostoc mat, depending on year/season. Estimates for different seasons correlated with thermal time (accumulated hours above 0 °C during the year) rather than irradiance, in contrast to communities in local lacustrine environments, where irradiance is the main driver of primary productivity. In the terrestrial habitat, N. commune appears to compromise between an ability to capitalize on short periods of higher temperature and efficient utilization of lower irradiance at low temperature. The relationship between thermal time and net annual C fixation by N. commune is strongly linear.     

Lake optics and depth limits for photogenesis and photosynthesis in charophyte meadows

Two series of lakes with increasing attenuation were examined for trends in spectral composition. They became the basis for an evaluation of the light environment at the lower boundary (LB) of Nitella meadows in three other series of lakes. Increased attenuation (K _d PAR) was marked by progressive erosion of the blue window and caused primarily by humic substances. An increase in K _d PAR from 0.06 to 0.81 produced, at the floor of the euphotic zone, a shift in K _d min from 440 to 580 nm. Regressions of boundary depths of Nitella meadows on water clarity produced similar slope coefficients for the three series of lakes. Several trends became evident: 1, PAR irradiance at the LB increases with depth of the LB; 2, red light (E _d 660) declines from richness at shallow LB to near extinction in deep water LB in clear lakes; while 3, blue light (K _d 450) increases to an asymptote. Blue light appears to substitute, although less effectively, for red light irradiance in the growth regulation of charophytes. These data support an hypothesis that spectral quality is involved in the determination of lower boundary depths for benthic macro-algae.     

SYNTHESIS OF MYCOSPORINE-LIKE AMINO ACIDS IN CHONDRUS CRISPUS (FLORIDEOPHYCEAE) AND THE CONSEQUENCES FOR SENSITIVITY TO ULTRAVIOLET B RADIATION

The induction and protective role of the UV-absorbing compounds known as mycosporine-like amino acids (MAAs) were examined in sublittoral Chondrus crispus Stackh. transplanted for 2 weeks in the spring and summer to shallow water under three irradiance conditions: PAR (photosynthetically active radiation; 400–700 nm), PAR + UVA (PAR + 320– 400 nm), PAR + UVA + UVB (PAR + UVA + 280– 320 nm). Sublittoral thalli collected around Helgoland, North Sea, Germany, from 6 m below the mean low water of spring tides contained less than 0.1 mg·g⁻¹ dry weight (DW) total MAAs, whereas eulittoral samples contained over 1 mg·g⁻¹ DW. Transplantation to shallow water led to the immediate synthesis of three MAAs in the following temporal order: shinorine (λ_max 334 nm), asterina (λ_max 330 nm), and palythine (λ_max 320 nm), with the shinorine content peaking and then declining after 2 days (exposure to 100 mol photons·m⁻²). Maximum total MAA content (2 mg·g⁻¹ DW) also occurred after 2 days of induction, exceeding the content normally found in eulittoral samples. Furthermore, the relative proportion of the different MAAs at this time was different than that in eulittoral samples. After 2 days the total content declined to the eulittoral value, with palythine as the principal MAA. Similar data were obtained for all treatments, indicating that MAA synthesis in C. crispus was induced by PAR and not especially stimulated by UV radiation. The ability of photosystem II (PSII) to resist damage by UVB was tested periodically during the acclimation period by exposing samples to a defined UVB dose in the lab. Changes in chlorophyll fluorescence (F_v/F_m and effective quantum yield, φ_II) indicated that PSII function was inhibited during the initial stage of acclimation but gradually improved with time. No difference among screening treatments was detected except in spring for the samples acclimating to PAR + UVA + UVB. In this treatment F_v/F_m and φ_II were significantly lower than in the other treatments. During the first week of each experiment, growth rates were also significantly reduced by UVB. The reductions occurred despite maximum MAA content, indicating an incomplete protection of photosynthetic and growth-related processes.     

Effects of light history on primary productivity in a phytoplankton community dominated by the toxic cyanobacterium Cylindrospermopsis raciborskii

1. The effects of instantaneous irradiance and short‐term light history on primary production were determined for samples from a subtropical water reservoir dominated by the toxic cyanobacterium Cylindrospermopsis raciborskii. ¹⁴C‐bicarbonate uptake incubations were conducted on water samples from the reservoir, for irradiance (photosynthetically active radiation) ranging from 0 to 1654 μmol quanta m⁻² s⁻¹. Prior to the ¹⁴C incubations, cells were pre‐treated at irradiance levels ranging from 0 to 1006 μmol quanta m⁻² s⁻¹. 2. The average irradiance experienced by cells during the 2–2.5 h pre‐treatment incubations affected the productivity–irradiance (P–I) parameters: exposure to high light in pre‐treatment conditions caused a substantial decrease in maximum rate of primary production P_max and the photoinhibition parameter β when compared to cells pre‐treated in the dark. 3. While the data collected in this study were not sufficient to develop a full dynamic model of C. raciborskii productivity, P_max and β were modelled as a function of pre‐treatment irradiance, and these models were applied to predict the rate of primary production as a function of both instantaneous and historical irradiance. The results indicated that while cells with a history of exposure to high irradiance will be the most productive in high irradiance, production rates will be highest overall for dark‐acclimated cells in moderate irradiance. 4. Our results may explain why optically‐deep mixing favours C. raciborskii. If the mixing depth z_m exceeds the euphotic depth z_eu, cells will be dark‐acclimated, which will increase their rate of production when they are circulated through the euphotic zone. These results also predict that production rates will be higher during morning hours than for the same irradiance in the afternoon, which is consistent with other phytoplankton studies. 5. Since the rate of production of C. raciborskii‐dominated systems cannot be described by a single P–I curve, accurate estimates of production rates will require measurements over the daily light cycle.     

The effect of elevated CO2 on the production and respiration of a Sargassum thunbergii community: A mesocosm study

Approximately one‐third of anthropogenic carbon dioxide is absorbed into the ocean and causes it to become more acidic. The Intergovernmental Panel on Climate Change (IPCC) suggested that the surface ocean pH, by the year 2100, would drop by a further 0.3 and 0.4 pH units under RCP (Representative Concentration Pathway) 6.0 and 8.5 climate scenarios. The macroalgae communities that consisted of Sargassum thunbergii and naturally attached epibionts were exposed to fluctuations of ambient and manipulated pH (0.3–0.4 units below ambient pH). The production and respiration in S. thunbergii communities were calculated from dissolved oxygen time‐series recorded with optical dissolved oxygen sensors. The pH, irradiance, and dissolved oxygen occurred in parallel with diurnal (day/night) patterns. According to net mesocosm production – photosynthetically active radiation (PAR) model, the saturation and compensation PAR, the mean maximum gross mesocosm production (GMP), and daily mesocosm respiration were higher in the CO₂ enrichment, than in the ambient condition, while the mean of photosynthetic coefficient was similar. In conclusion, elevated CO₂ stimulated oxygen production and consumption of S. thunbergii communities in the mesocosm. Furthermore, the sensitivity of the GMP of the S. thunbergii community to irradiance was reduced, and achieved maximum production rate at higher PAR. These positive responses to CO₂ enrichment suggest that S. thunbergii communities may thrive in under high CO₂ conditions.     

Model parameterization to simulate and compare the PAR absorption potential of two competing plant species

Mountain pastures dominated by the pasture grass Setaria sphacelata in the Andes of southern Ecuador are heavily infested by southern bracken (Pteridium arachnoideum), a major problem for pasture management. Field observations suggest that bracken might outcompete the grass due to its competitive strength with regard to the absorption of photosynthetically active radiation (PAR). To understand the PAR absorption potential of both species, the aims of the current paper are to (1) parameterize a radiation scheme of a two-big-leaf model by deriving structural (LAI, leaf angle parameter) and optical (leaf albedo, transmittance) plant traits for average individuals from field surveys, (2) to initialize the properly parameterized radiation scheme with realistic global irradiation conditions of the Rio San Francisco Valley in the Andes of southern Ecuador, and (3) to compare the PAR absorption capabilities of both species under typical local weather conditions. Field data show that bracken reveals a slightly higher average leaf area index (LAI) and more horizontally oriented leaves in comparison to Setaria. Spectrometer measurements reveal that bracken and Setaria are characterized by a similar average leaf absorptance. Simulations with the average diurnal course of incoming solar radiation (1998–2005) and the mean leaf–sun geometry reveal that PAR absorption is fairly equal for both species. However, the comparison of typical clear and overcast days show that two parameters, (1) the relation of incoming diffuse and direct irradiance, and (2) the leaf–sun geometry play a major role for PAR absorption in the two-big-leaf approach: Under cloudy sky conditions (mainly diffuse irradiance), PAR absorption is slightly higher for Setaria while under clear sky conditions (mainly direct irradiance), the average bracken individual is characterized by a higher PAR absorption potential. (∼74 MJ m⁻² year⁻¹). The latter situation which occurs if the maximum daily irradiance exceeds 615 W m⁻² is mainly due to the nearly orthogonal incidence of the direct solar beam onto the horizontally oriented frond area which implies a high amount of direct PAR absorption during the noon maximum of direct irradiance. Such situations of solar irradiance favoring a higher PAR absorptance of bracken occur in ∼36% of the observation period (1998–2005). By considering the annual course of PAR irradiance in the San Francisco Valley, the clear advantage of bracken on clear days (36% of all days) is completely compensated by the slight but more frequent advantage of Setaria under overcast conditions (64% of all days). This means that neither bracken nor Setaria show a distinct advantage in PAR absorption capability under the current climatic conditions of the study area.     

Photosynthesis and transpiration of tomato and CO2 fluxes in a greenhouse under changing environmental conditions in winter

Responses of tomato leaves in a greenhouse to light and CO₂ were examined at the transient stage at the end of winter, when both photoperiod and irradiance gradually increase. Additionally, CO₂ fluxes were calculated for a greenhouse without supplementary lighting and without CO₂ enrichment based on CO₂ sinks (plant photosynthesis) and CO₂ sources (plant and substrate respiration). In January, tomato leaves in the greenhouse showed low photosynthesis with a maximum assimilation of 6–8 μmol CO₂ m⁻² s⁻¹, a quantum yield of 0.06 μmol CO₂ μmol⁻¹ photosynthetic active radiation (PAR) and a low light compensation point of 26 μmol PAR m⁻² s⁻¹, a combination which classifies them as shade leaves. In February, tomato leaves increased their light compensation point to 39 μmol PAR m⁻² s⁻¹ and quantum yield to 0.08, the former indicating the adaptation to increased irradiance and photoperiod. These tomato leaves increased their transpiration from 0.4 to 0.9 in January to ∼2 mmol H₂O m⁻² s⁻¹ in February. Both photosynthesis and transpiration were primarily limited by light but neither by stomatal conductivity nor by CO₂. In January, light response of photosynthesis, dark respiration and transpiration were negligibly affected by increasing CO₂ concentrations from 600 to 900 ppm CO₂ under low light conditions, indicating no benefit of CO₂ enrichment unless light intensity increased. In February, tomato leaves were photoinhibited at inherent greenhouse CO₂ concentrations on the first sunny day; this photoinhibition was further enhanced by an increased CO₂ concentration of 1000 ppm. CO₂ fluxes in the greenhouse appeared strongly dependent on solar radiation. After exceeding the light compensation point in the morning, greenhouse CO₂ concentrations decreased by 58 or by 110 ppm CO₂ h⁻¹ on a sunny day in January or February and by 23 ppm on overcast days in both months. Calculated per overall tomato canopy, plant photosynthesis contributed 42–50% to the morning CO₂ depletion in the greenhouse. Dark respiration of tomato leaves was ∼2 μmol CO₂ m⁻² s⁻¹ in January and ∼3 μmol CO₂ m⁻² s⁻¹ in February. This dark respiration resulted in rises of 15 and 17 ppm CO₂ h⁻¹ at night in the greenhouse compartment and was identified as primary source of CO₂. Respiration of the substrate used to grow the plants, which produced 7.3 ppm CO₂ h⁻¹, was identified as secondary source of CO₂. The combined plant and substrate respiration resulted in peaks of up to 900 ppm CO₂ in the greenhouse before dawn.     

SHORT‐TERM AND LONG‐TERM EFFECTS OF TEMPERATURE ON PHOTOSYNTHESIS IN THE DIATOM THALASSIOSIRA PSEUDONANA UNDER UVR EXPOSURES1

Temperature is expected to modify the effects of ultraviolet radiation (UVR) on photosynthesis by affecting the rate of repair. We studied the effect of short‐term (1 h) and long‐term (days) acclimation to temperature on UVR photoinhibition in the diatom Thalassiosira pseudonana Hasle et Heimdal. Photosynthesis was measured during 1 h exposures to varying irradiances of PAR and UVR + PAR at 15, 20, and 25°C, the latter corresponding to the upper temperature limit for optimal growth in T. pseudonana. The exposures allowed the estimation of photosynthesis–irradiance (P–E) curves and biological weighting functions (BWFs) for photoinhibition. For the growth conditions used, temperature did not affect photosynthesis under PAR. However, photoinhibition by UVR was highly affected by temperature. For cultures preacclimated to 20°C, the extent of UVR photoinhibition increased with decreasing temperature, from 63% inhibition of PAR‐only photosynthesis at 25°C to 71% at 20°C and 85% at 15°C. These effects were slightly modified after several days of acclimation: UVR photoinhibition increased from 63% to 75% at 25°C and decreased from 85% to 80% at 15°C. Time courses of photochemical efficiency (Φ_PSII) under UVR + PAR were also fitted to a model of UVR photoinhibition, allowing the estimation of the rates of damage (k) and repair (r). The r/k values obtained for each temperature treatment verified the responses observed with the BWF (R² = 0.94). The results demonstrated the relevance of temperature in determining primary productivity under UVR exposures. However, the results suggested that temperature and UVR interact mainly over short (hours) rather than long (days) timescales.     

Carbon balance during CAM: an assessment of respiratory CO2 recycling in the epiphytic bromeliads Aechmea nudicaulis and Aechmea fendleri

Abstract The regulation of crassulacean acid metabolism (CAM) under controlled environmental conditions has been investigated for two tropical epiphytes, relating plant water and carbon balance to growth form and habitat preference under natural conditions. Aechmea fendleri is restricted to wet, upper montane regions of Trinidad, while A. nudicaulis has a wider distribution extending into more arid regions of the island. Morphological characteristics of these plants are related to habitat preference in terms of leaf succulence (0.44 and 0.94 kg m^?2 for the two species respectively) and a distinct layer of water storage parenchyma in A. nudicaulis In contrast, the thinner leaves of A. fendleri contain little water-storage parenchyma and less chlorenchyma per unit area, but the plants have a more open leaf rosette. The two species differ in expression of CAM, since the proportion of respiratory CO₂ recycled as part of CAM had been found to be much lower in A. fendleri This study compared the efficiency of water use and role of respiratory CO₂ recycling under two PAR regimes (300 and 120 μnol m^?2 s^?1) and three night temperatures (12, 18 and 25 °C). Dark CO₂ uptake rates for both species were comparable to plants in the field (maximum of 2.3 ± 0.2 μmol m^?2s^?1± SD, n= 3). Total net CO₂ uptake at night increased on leaf area basis with temperature for both species under higher PAR, although under the low PAR regime CO₂ uptake was maximal at 18 °C. Water-use efficiency (WUE) increased at 18 °C and 25 °C during dark CO₂ uptake (Phase I) and also during late afternoon photosynthesis (Phase IV) in both species. For A. fendleri, dawn to dusk changes in titrable acidity (ΔH ⁺) were similar under high and low PAR, although ΔH⁺ was correlated to night temperature and PAR in A. nudicaulis. The proportion of ΔH⁺ derived from respiratory CO₂ also varied with experimental conditions. Thus percentage recycling was lower in A. fendleri under high PAR (0–10%), but was only reduced at 18 °C under low PAR. Recycling by A. nudicaulis ranged from 32–42% under high PAR, but was also reduced to 6% under low PAR at 18 °C; at 12 °C and 25 °C, recycling was 37% and 52% respectively. Previous studies have suggested a relationship between the proportion of recycling and degree of water stress. This study indicated that CAM as a CO₂ concentrating mechanism regulates both water-use efficiency and plant carbon balance in these epiphytes, in response to PAR and night temperature. However, the precise relationship between respiratory processes and the balance between external and internal sources of CO₂ is as yet unresolved.     

Phenology,irradiance and temperature characteristics of a freshwater red alga,Nemalionopsis tortuosa (Thoreales), from Kagoshima,southern Japan

Phenology, irradiance and temperature characteristics of a freshwater benthic red alga, Nemalionopsis tortuosa Yoneda et Yagi (Thoreales), were examined from Kagoshima Prefecture, southern Japan for the conservation of this endemic and endangered species. Field surveys confirmed that algae occurred in shaded habitats from winter to early summer, and disappeared during August through November. A net photosynthesis–irradiance (P–E) model revealed that net photosynthetic rate quickly increased and saturated at low irradiances, where the saturating irradiance (E_k) and compensation irradiance (E_c) were 10 (8–12, 95% credible interval (CRI)) and 8 (6–10, 95% CRI) μmol photon m^?2 s^?1, respectively. Gross photosynthesis and dark respiration was determined over a range of temperatures (8–36°C) by dissolved oxygen measurements, and revealed that the maximum gross photosynthetic rate was highest at 29.5 (27.4–32.0, 95%CRI) °C. Dark respiration also increased linearly when temperature increased from 8°C to 36°C, indicating that the increase in dark respiration at higher temperature most likely caused decreases in net photosynthesis. The maximum quantum yield (Fv/Fm) that was determined using a pulse amplitude modulated‐chlorophyll fluorometer (Imaging‐PAM) was estimated to be 0.51 (0.50–0.52, 95%CRI) and occurred at an optimal temperature of 21.7 (20.1–23.4, 95%CRI) °C. This species can be considered well‐adapted to the relatively low natural irradiance and temperature conditions of the shaded habitat examined in this study. Our findings can be applied to aid in the creation of a nature‐reserve to protect this species.     

MICROALGAL LIGHT-HARVESTING IN EXTREME LOW-LIGHT ENVIRONMENTS IN MCMURDO SOUND, ANTARCTICA1

Microalgal pigment composition, photosynthetic characteristics, single-cell absorption efficiency (Qa_(λ)) spectra, and fluorescence-excitation (FE) spectra were determined for platelet ice and benthic communities underlying fast ice in Mc Murdo Sound, Antarctica, during austral spring 1988. Measurements of spectral irradiance (E_(λ)) and photosynthetically active radiation (PAR) as well as samples for particulate absorption measurements were taken directly under the congelation ice, within the platelet layer, as profiles vertically through the water column, and at the benihic surface. Light attenuation by.sea ice, algal pigments, and particulates reduced PAR reaching the platelet ice layer to 3%(9–33 fimol photons m-²-^?s-¹) of surface values and narrowed its spectral distribution to a band between 400 and 580 nm. Attenuation by the water column further reduced PAR reaching the sea floor (28–m depth) to 0.05% of surface levels (< 1 μmol photons m-² s-¹), with a spectral distribution dominated by 470–580–nm wavelengths. The photoadaptive index (I) for platelet ice algae (5.9–12.6 μmol photons m-^2.s-¹) was similar to ambient PAR, indicating that algae had acclimated to their light environment (i.e. the algae were light-replete). Maximum Qa_(λ) at the blue absorption peak (440 nm) was 0.63, and enhanced absorption was observed from 460–500 nm and was consistent with observed high cellular chlorophyll (chi) c:chl a and fucoxanthin: chl a molar ratios (0.4 and 1.2, respectively). Benthic algae were light-limited despite the maintenance of very low I_k values (4–11 μmol photons.m-^2.s-¹). Extremely high fucoxanthin: chi a ratios (1.6) in benthic algae produced enhanced green light absorption, resulting in a high degree of complementation between algal absorption and ambient spectral irradiance. Qa_(λ) values for benthic algae were maximal (0.9) between 400 and 510 nm but remained >0.35 even at absorption minima. Strong spectral flattening, a characteristic of intense pigment packaging, was also apparent in the Qa_(λ) spectra for benthic algae. FE and Qa_(λ) spectra were similar in shape for platelet ice algae, indicating that the efficiency at which absorbed energy was transferred to photosystem II (PSII) was independent of wavelength. Fluorescence emission by benthic algae was greatest for the 500–560–nm excitation wavelengths, suggesting that most energy absorbed by accessory pigments was transferred to PSII. These results suggest that under ice algae employ complementary pigmentation and maximize absorption efficiency as adaptive strategies to low-light stress. Regulating the distribution of absorbed energy between PSI and PSII may be an adaptive response to the restricted spectral distribution of irradiance.     

Decreased ribulose-1,5-bisphosphate carboxylase/oxygenase in transgenic tobacco transformed with 'antisense'rbcS.VIII. Impact on photosynthesis and growth in tobacco growing under extreme high irradiance and high temperature

Wild-type and antisense rbcS tobacco (Nicotiana tabacum) plants were grown in a glasshouse in midsummer in Portugal with an irradiance of 1500–2000 μmol m⁻²s⁻¹ and daytime temperatures of 30–35 °C. The Rubisco content of the transformants was lower by 35, 80 and over 90% than that of the wild-type. Gas exchange was measured over three separate days. There was a near-linear relation between Rubisco content and photosynthetic rate during the period of high irradiance, allowing a flux control coefficient of 0.83–0.89 to be estimated. The relation deviated slightly from linearity, because the internal CO₂ concentration (c;) was higher in the transformants than in the wild-type (190 and 275 μmol mol⁻¹ in plants with 35 and 80% less Rubisco, respectively, compared with 175 μmol mol⁻¹ for wild-type), compensating to some extent for the decreased Rubisco content. This increase in c_i occurred because the stomatal conductance (g) remained unaltered or was even higher in plants with decreased Rubisco, despite the lower rate of CO₂ assimilation. As a consequence, water use efficiency declined. The decreased rate of photosynthesis was not accompanied by a stoichiometric decrease in apparent growth rate. These results are discussed in relation to earlier studies of the plant set in growth cabinets. It is concluded that tobacco can adjust over a wide range of growth conditions to avoid a onesided limitation by Rubisco, but that in extreme environmental conditions this capacity to adapt is exhausted.     

Annual patterns of phytoplankton density and primary production in a large, shallow lake: the central role of light

1. We studied the seasonal dynamics of suspended particulate matter in a turbid, large shallow lake during an annual period (2005–06). We relate the patterns of seston concentration (total suspended solids), phytoplankton biomass and water transparency to the seasonal pattern of incident solar radiation (I₀). We also report the seasonal trends of phytoplankton primary production (PP) and photosynthesis photoinhibition due to photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) (I_β and UV₅₀). 2. We first collected empirical evidence that indicated the conditions of light limitation persisted during the study period. We found that the depth‐averaged irradiance estimated for the time of the day of maximum irradiance (I_mean–noon) was always lower than the measured onset of light saturation of photosynthesis (I_k). 3. We then contrasted the observations with theoretical expectations based on a light limitation scenario. The observed temporal patterns of seston concentration, both on a volume and area basis, were significantly explained by I₀ (R² = 0.39 and R² = 0.37 respectively). The vertical diffuse attenuation coefficient (kd_PAR) (R² = 0.55) and the depth‐averaged irradiance (I_mean) (R² = 0.66), significantly increased with the I₀; while the irradiance reaching the lake bottom (I_out) significantly decreased with the incident irradiance (R² = 0.49). However, phytoplankton biovolume maxima were not coincident with the time of the year of maximum irradiance. 4. A significant positive relationship was observed between PP estimated on an area basis and I₀ (R² = 0.51, P < 0.001). In addition, the parameters describing the photosynthetic responses to high irradiances displayed marked seasonal trends. The photosynthesis photoinhibition due to PAR as well as to UV were significantly related to incident solar radiation (PAR: R² = 0.73; UV: R² = 0.74). These results suggest adaptation of the phytoplankton community in response to changes in incident solar radiation.     

The effect of photosynthetically active radiation and temperature on the photosynthesis of two Vietnamese species of Sargassum,S. mcclurei and S. oligocystum,based on the field and laboratory measurements

SUMMARY The effects of photosynthetically active radiation (PAR) and temperature on the photosynthesis of two Vietnamese brown algae, Sargassum mcclurei and S. oligocystum (Fucales), were determined by field and laboratory measurements. Dissolved oxygen sensors and pulse‐amplitude modulated (PAM) fluorometry were used for the measurements of photosynthetic efficiency. A Diving‐PAM revealed that underwater measurements of the effective quantum yield (Φ _PSII ) of both species declined with increasing incident PAR, with minimum Φ _PSII occurring during noon to early afternoon. Φ _PSII recovered in the evening, indicating photo‐adaptation to excessive PAR. In laboratory experiments, Φ _PSII also decreased under continuous exposure to 1000 μmol photons m^?2 s^?1; and full recovery occurred after 12 h of dark acclimatization. The net photosynthesis – PAR experiments of S. mcclurei and S. oligocystum conducted at 28°C revealed that the net photosynthetic rate quickly increased at PAR below the saturation irradiance of 361 and 301 μmol photons m^?2 s^?1 and nearly saturated to maximum net photosynthetic rates of 385 and 292 μg O₂ g_ww ^{? 1} min^?1 without photoinhibition, respectively. Gross photosynthesis and dark respiration experiments determined over a range of temperatures (12–40°C), revealed that the maximum gross photosynthetic rates of 201 and 147 μg O₂ g_ww ^{? 1} min^?1 occurred at 32.9 and 30.7°C for S. mcclurei and S. oligocystum, respectively. The dark respiration rates increased exponentially over the temperature ranges examined. The estimated maximum value of the maximum quantum yield occurred at 19.3 and 20.0°C and was 0.76 and 0.74, respectively. Similar to the natural habitat of the study site, these two species tolerated the relatively high temperatures and broad range of PAR. The ability of these species to recover from exposure to high PAR is one of the mechanisms that allow them to flourish in the shallow water environment.     

Propagation of Hedera helix: Influence of irradiance to stock plants, length of internode and topophysis of cutting

Abstract Stock plants of Hedera helix cv. Pittsburgh were grown in controlled environment rooms at four different irradiances (light intensities) 10, 22, 37 or 46 W m^-2 PAR (photosynthetically active radiation). The root formation on single internode cuttings from these stock plants was observed in relation to length of the internodes and the position on the vine (topophysis). The analysis indicated that the root number was primarily dependent on internode length which in turn was dependent on irradiance to stock plants and topophysis. The irradiance of 37 W m^-2 PAR was optimal for internode length and root number and the basal internode was the one which produced the greatest number of roots.