EFFECTS OF ULTRAVIOLET‐A RADIATION AND NUTRIENT AVAILABILITY ON THE CELLULAR COMPOSITION OF PHOTOPROTECTIVE COMPOUNDS IN GLENODINIUM FOLIACEUM (DINOPHYCEAE)1

The photoprotective response in the dinoflagellate Glenodinium foliaceum F. Stein exposed to ultraviolet‐A (UVA) radiation (320–400 nm; 1.7 W · m²) and the effect of nitrate and phosphate availability on that response have been studied. Parameters measured over a 14 d growth period in control (PAR) and experimental (PAR + UVA) cultures included cellular mycosporine‐like amino acids (MAAs), chls, carotenoids, and culture growth rates. Although there were no significant effects of UVA on growth rate, there was significant induction of MAA compounds (28 ± 2 pg · cell^?1) and a reduction in chl a (9.6 ± 0.1 pg · cell^?1) and fucoxanthin (4.4 ± 0.1 pg · cell^?1) compared to the control cultures (3 ± 1 pg · cell^?1, 13.3 ± 3.2 pg · cell^?1, and 7.4 ± 0.3 pg · cell^?1, respectively). In a second investigation, MAA concentrations in UVA‐exposed cultures were lower when nitrate was limited (P < 0.05) but were higher when phosphate was limiting. Nitrate limitation led to significant decreases (P < 0.05) in cellular concentration of chls (chl c₁, chl c₂, and chl a), but other pigments were not affected. Phosphate availability had no effect on final pigment concentrations. Results suggest that nutrient availability significantly affects cellular accumulation of photoprotective compounds in G. foliaceum exposed to UVA.     

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.     

Microphytobenthic species composition,pigment concentration,and primary production in sublittoral sediments of the Trondheimsfjord (Norway)1

In spring 2005, monthly sampling was carried out at a sublittoral site near Tautra Island. Microphytobenthic identification, abundance (ABU), and biomass (BIOM), were performed by microscopic analyses. Bacillariophyceae accounted for 67% of the total ABU, and phytoflagellates constituted 30%. The diatom floristic list consisted of 38 genera and 94 species. Intact light‐harvesting pigments chl a, chl c, and fucoxanthin and their derivatives were identified and quantified by HPLC. Photoprotective carotenoids were also observed (only as diadinoxanthin; no diatoxanthin was detected). Average fucoxanthin content was 4.57 ± 0.45 μg fucoxanthin · g sediment dry mass^?1, while the mean chl a concentration was 2.48 ± 0.15 μg · g^?1 dry mass. Both the high fucoxanthin:chl a ratio (considering nondegraded forms) and low amounts of photoprotective carotenoids indicated that the benthic microalgal community was adapted to low light. Microphytobenthic primary production was estimated in situ (MPPs, from 0.15 to 1.28 mg C · m^?2 · h^?1) and in the laboratory (MPPp, from 6.79 to 34.70 mg C · m^?2 · h^?1 under light saturation) as ¹⁴C assimilation; in April it was additionally estimated from O₂‐microelectrode studies (MPP_O2) along with the community respiration. MPP_O2 and the community respiration equaled 22.9 ± 7.0 and 7.4 ± 1.8 mg C · m^?2 · h^?1, respectively. A doubling of BIOM from April to June in parallel with a decreasing photosynthetic activity per unit chl a led us to suggest that the microphytobenthic community was sustained by heterotrophic metabolism during this period.     

MAA Synthesis and Accumulation in Polar Macroalgae are Controlled by Abiotic Factors

Mycosporine‐like amino acids (MAAs) are regarded as powerful sunscreens protecting the algae against harmful UV radiation. The MAA protection efficiency was tested in algal samples by measuring the optimum quantum yield of photosynthesis using photosystem II fluorescence. It could be demonstrated that the recovery of photosynthesis after exposure to enhanced UV radiation is faster in individuals with high MAA content. MAAs can be synthesized in several polar macroalgae in response to different radiation conditions. Although MAA induction patterns are very species‐specific, some similarities can be found. Field studies indicate that plants from different growth habitats providing distinct radiation climate can be grouped into three physiological categories depending on their MAA content. The first group (I) includes mainly deep‐water species, typically lacking MAAs. The second group (II), algal species found in a broad range of water depths (eu‐ and sublittoral), which are able to flexibly synthesize and accumulate MAAs. The third group (III) includes supra‐ and eulittoral taxa, which always contain high MAA concentrations. In laboratory studies, we showed that taxa of group II and III responded in three different ways based on MAA accumulation when exposed to different radiation conditions (PAR, PAR + UVA, PAR + UVA + UVB). Either they: (a) exhibit highest total MAA concentration under the full artificial spectrum; (b) increase their MAA concentration after exposure to PAR and PAR + UVA or (c) MAA concentration declines after exposure to the full spectrum. Our studies have indicated that when coupled with UVR, exposure to temperature fluctuations ranging from 0 to 10 °C also affect MAA biosynthesis.     

The effects of ultraviolet radiation on photosynthetic performance,growth and sunscreen compounds in aeroterrestrial biofilm algae isolated from building facades

The effects of artificial ultraviolet radiation [UVR; 8 W m⁻² ultraviolet-A (UVA), 0.4 W m⁻² ultraviolet-B (UVB)] on photosynthetic performance, growth and the capability to synthesise mycosporine-like amino acids (MAAs) was investigated in the aeroterrestrial green algae Stichococcus sp. and Chlorella luteoviridis forming biofilms on building facades, and compared with the responses of two green algae, from soil (Myrmecia incisa) and brackish water (Desmodesmus subspicatus). All species exhibited decreasing quantum efficiency (F _v/F _m) after 1–3 days exposure to UVR. After 8–12 days treatment, however, all aeroterrestrial isolates exhibited full recovery under UVA and UVA/B. In contrast, D. subspicatus showed only 80% recovery after treatment with UVB. While Stichococcus sp. and C. luteoviridis exhibited a broad tolerance in growth under all radiation conditions tested, M. incisa showed a significant decrease in growth rate after exposure to UVA and UVA/B. Similarly D. subspicatus grew with a reduced rate under UVA, but UVA/B led to full inhibition. Using HPLC, an UV-absorbing MAA (324 nm-MAA) was identified in Stichococcus sp. and C. luteoviridis. While M. incisa contained a specific 322 nm-MAA, D. subspicatus lacked any trace of such compounds. UV-exposure experiments indicated that all MAA-containing species are capable of synthesizing and accumulating these compounds, thus supporting their function as an UV-sunscreen. All data well explain the conspicuous ecological success of aeroterrestrial green algae in biofilms on facades. Biosynthesis and accumulation of MAAs under UVR seem to result in a reduced UV-sensitivity of growth and photosynthesis, which consequently may enhance survival in the environmentally harsh habitat.     

PHYTOPLANKTON PIGMENTS IN COASTAL LAKE MICHIGAN: DISTRIBUTIONS DURING THE SPRING ISOTHERMAL PERIOD AND RELATION WITH EPISODIC SEDIMENT RESUSPENSION1

Phytoplankton pigment distributions during the spring isothermal periods of 1998 and 1999 and their association with episodic sediment resuspension were characterized in coastal waters of southern Lake Michigan. Total and phylogenetic group chl a concentrations (derived using chemical taxonomy matrix factorization of diagnostic carotenoids) corresponded with assemblage and group biovolumes estimated from microscopic enumeration (P≤ 0.001). Diatoms and cryptophytes dominated assemblages and together typically comprised greater than 85% of relative chl a. Total chl a concentrations and both fucoxanthin·chl a ^{? 1} and alloxanthin·chl a ^{? 1} ratios were similar across depths (P> 0.05), indicating uniform distributions of and photophysiological states for assemblages and diatoms and cryptophytes, respectively, throughout the mixed water column. Total chl a concentrations were not always spatially uniform from near‐shore to offshore waters, with the greatest variability reflecting the influence of tributary inflows upon coastal assemblages. Sediment resuspension strongly influenced water column particle density and light climate; however, total and group chl a concentrations did not correspond with coefficients of K_d and suspended particulate matter concentrations (P> 0.05). The correspondence of both light attenuation and suspended particulate matter concentration with relative diatom chl a (P≤ 0.001) indicated an apparent association between sediment resuspension and diatoms. This, and the negative association (P≤ 0.0001) between relative diatom and cryptophyte chl a, corresponded with the spatial dominance of diatom and cryptophyte chl a in near‐shore and offshore waters, respectively. The presence of viable chl a and fucoxanthin within the surficial sediment layer, established this layer as a potential source of meroplanktonic diatoms for near‐shore assemblages.     

STRUCTURE AND FUNCTION OF BENTHIC ALGAL COMMUNITIES IN AN EXTREMELY ACID RIVER1

The composition of algal species and pigments and the structural and functional characteristics of the algal community were investigated in an acid stream of southwestern Spain, the Río Tinto. The algal community had low diversity and showed few seasonal differences. It was mainly made up of Klebsormidium flaccidum Kütz. (Silva, Mattox & Blackwell) that produced long greenish or purplish filaments, Pinnularia acoricola Hust. (producing brown patches) and Euglena mutabilis Schmitz. The algal filaments made up a consistent biofilm that also included fungal hyphae, iron bacterial sheaths, diatoms, and mineral particles. HPLC analyses on Río Tinto samples showed that undegraded chl accounted for 67% of the total chl in the filamentous patches but were a minority in the brown patch (2.6%). The brown patch had a concentration of carotenoids eight times lower than that observed in the green patch. When chl concentrations were weighted for the proportion of the different patches on the streambed, undegraded chl a accounted for 89.2 mg chl a·m ^{? 2} of stream surface area (5.4 g C·m ^{? 2}). This high algal biomass was supported by relatively high nutrient concentrations and by a high phosphatase activity (V_max = 137.7 nmol methylumbelliferyl substrate·cm ^{? 2}·h ^{? 1} 1 Received 15 July 2002. Accepted 17 February 2003. , K_m = 0.0045 μM). The remarkable algal biomass in Río Tinto potentially contributed to the bacterial–fungal community and to the macroinvertebrate community and emphasizes the role that the algae may have in the organic matter cycling and energy flow in extreme systems dominated by heterotrophic microorganisms.     

PHOTOSYNTHETIC ADAPTATION OF A BLOOM‐FORMING CYANOBACTERIUM MICROCYSTIS AERUGINOSA (CYANOPHYCEAE) TO PROLONGED UV‐B EXPOSURE1

The bloom‐forming cyanobacterium Microcystis aeruginosa Kütz 854 was cultured with 1.05 W·m^?2 UV‐B for 3 h every day, and its growth, pigments, and photosynthesis were investigated. The specific growth rates represented by chl a concentration and OD₇₅₀ were inhibited 8% and 9% by UV‐B exposure, respectively. Six days of UV‐B treatment significantly reduced cellular contents of phycocyanin and allophycocyanin by 32% and 62%, respectively, and markedly increased the carotenoid content by 27%, but had little effect on the chl a content. The initial values of optimal photosynthetic efficiency for UV‐B treated samples were, respectively, 52%, 87%, and 93% of controls on days 4, 7, and 10 of growth. The light‐saturated photosynthetic rates at day 6 were significantly lower than controls grown without UV‐B. The probability of electron transfer beyond Q_A decreased during UV‐B exposure, and this indicated that the acceptor side of PSII was one of main damage sites. The adaptation of M. aeruginosa 854 to UV‐B radiation could be observed from light‐saturated photosynthetic rates on day 13 and diurnal changes of chl fluorescence during the late growth phase. When both exposed to higher UV‐B, samples cultured under 1.05 W·m^?2 UV‐B for 9 days recovered faster than controls. It is suggested that M. aeruginosa 854 had at least three adaptive strategies to cope with the enhanced UV‐B: increasing the synthesis of carotenoids to counteract reactive oxidants caused by UV‐B exposure, degrading phycocyanin and allophycocyanin to avoid further damage to DNA and reaction centers, and enhancing the repair of UV‐B induced damage to the photosynthetic apparatus.     

DIEL PATTERNS IN LIGHT ABSORPTION AND ABSORPTION EFFICIENCY FACTORS OF ISOCHRYSIS GALBANA (PRYMNESIOPHYCEAE)1

The chl a specific absorption coefficients [a* (λ), m²·mg chl a ^{? 1}] were examined in chemostat culture of the Prymnesiophyceae Isochrysis galbana (Parke) under a 12:12‐h light:dark cycle at low light (75 μmol photons·m ^{? 2}·s ^{? 1}) and high light (500 μmol photons· m ^{? 2}·s ^{? 1}) conditions. Other associated measurements such as pigment composition, cell density, and diameter as the measure of cell size were also made at the two light regimes every 2 h for 2 days to confirm the periodicity. A distinct diel variability was observed for the a* (λ) with maxima near dawn and minima near dusk. The magnitude of diel variation in a* (440) was 15% at low light and 22% at high light. Pronounced diel patterns were observed for cell size with minima near dawn and maxima near dusk. The magnitude of diel variation in cell size was 9.3% at low light and 21% at high light. The absorption efficiency factors [Q _a (440)] were determined by reconstruction using intracellular concentrations of pigments and cell size. The Q _a (440) also showed a distinct diel variability, with minima near dawn and maxima near dusk. The diel variation in a* (λ) and Q _a (λ) was primarily caused by changes in cell size due to growth, although there was some influence from diel variations in the intracellular pigment concentrations. The results presented here indicated that diel variation in a* (λ) was an important component of the optical characterization of phytoplankton.     

RELATIONSHIP BETWEEN PHOTOSYNTHESIS AND CELL SIZE OF MARINE DIATOMS12

Three photosynthetic parameters of 7 species of marine diatoms were studied using Na₂¹⁴CO₃ at 5–8 C using log phase axenic cultures. The cell volumes of the different species varied from 70 μm³ to 40 × 10⁵μm³. The present experiment is consistent with the interpretation that the initial slope α (mg C · [mg chl a]^?1· h^?1· w^?1· m²) of photosynthesis vs. light curves is controlled by self-shading of chlorophyll a in the cell. Pm, the rate of photosynthesis at light saturation (mg C · [mg cell, C]^?1· h^?1) and R, the intercept at zero light intensity (mg C · [mg cell C]^?1· H^?1) are both dependent on the ratio of surface area to volume of cell.     

DIFFERENTIAL RESPONSES OF ANABAENA SP. PCC 7120 (CYANOPHYCEAE) CULTURED IN NITROGEN‐DEFICIENT AND NITROGEN‐ENRICHED MEDIA TO ULTRAVIOLET‐B RADIATION1

Stratospheric ozone depletion increases the amount of ultraviolet‐B radiation (UVBR) (280–320 nm) reaching the surface of the earth, potentially affecting phytoplankton. In this work, Anabaena sp. PCC 7120, a typically nitrogen (N)‐fixing filamentous bloom‐forming cyanobacterium in freshwater, was individually cultured in N‐deficient and N‐enriched media for long‐term acclimation before being subjected to ultraviolet‐B (UVB) exposure experiments. Results suggested that the extent of breakage in the filaments induced by UVBR increases with increasing intensity of UVB stress. In general, except for the 0.1 W · m^?2 treatment, which showed a mild increase, UVB exposure inhibits photosynthesis as evidenced by the decrease in the chl fluorescence parameters maximum photochemical efficiency of PSII (F_v/F_m) and maximum relative electron transport rate. Complementary chromatic acclimation was also observed in Anabaena under different intensities of UVB stress. Increased total carbohydrate and soluble protein may provide some protection for the culture against damaging UVB exposure. In addition, N‐deficient cultures with higher recovery capacity showed overcompensatory growth under low UVB (0.1 W · m^?2) exposure during the recovery period. Significantly increased (～830%) ATPase activity may provide enough energy to repair the damage caused by exposure to UVB.     

Nutrient limitation of algal periphyton in streams along an acid mine drainage gradient

Metal oxyhydroxide precipitates that form from acid mine drainage (AMD) may indirectly limit periphyton by sorbing nutrients, particularly P. We examined effects of nutrient addition on periphytic algal biomass (chl a), community structure, and carbon and nitrogen content along an AMD gradient. Nutrient diffusing substrata with treatments of +P, +NP and control were placed at seven stream sites. Conductivity and SO₄ concentration ranged over an order of magnitude among sites and were used to define the AMD gradient, as they best indicate mine discharge sources of metals that create oxyhydroxide precipitates. Aqueous total phosphorous (TP) ranged from 2 to 23 μg · L^?1 and significantly decreased with increasing SO₄. Mean chl a concentrations at sites ranged from 0.2 to 8.1 μg · cm^?2. Across all sites, algal biomass was significantly higher on +NP than control treatments (Co), and significantly increased with +NP. The degree of nutrient limitation was determined by the increase in chl a concentration on +NP relative to Co (response ratio), which ranged from 0.6 to 9.7. Response to nutrient addition significantly declined with increasing aqueous TP, and significantly increased with increasing SO₄. Thus, nutrient limitation of algal biomass increased with AMD impact, indicating metal oxyhydroxides associated with AMD likely decreased P availability. Algal species composition was significantly affected by site but not nutrient treatment. Percent carbon content of periphyton on the Co significantly increased with AMD impact and corresponded to an increase in the relative abundance of Chlorophytes. Changes in periphyton biomass and cellular nutrient content associated with nutrient limitation in AMD streams may affect higher trophic levels.     

A COMPARISON OF PHOTORESPONSE AMONG TEN DIFFERENT KARENIA BREVIS (DINOPHYCEAE) ISOLATES1

Many laboratories have solely used the Wilson isolate to physiologically characterize the harmful algal bloom (HAB) dinoflagellate Karenia brevis (C. C. Davis) G. Hansen et Moestrup. However, analysis of one isolate may lead to misinterpretations when extrapolating measurements to field populations. In this study, pulse‐amplitude‐modulated chlorophyll fluorometer (PAM‐FL) relative electron transport rate (ETR), F_v/F_m, and chl were compared with traditional techniques, such as ¹⁴C photosynthesis versus irradiance (P–E) curves, DCMU [3‐(3′,4′‐dichlorophenyl)‐1,1‐dimethyl urea] F_v/F_m, and extracted chl. The DCMU and PAM‐FL values of F_v/F_m (r² = 0.51) and chl (r² = 0.58) were in good agreement. There was no correlation between ¹⁴C and PAM‐FL α, P_max, and β parameters because PAM‐FL ETR was only a relative measurement. The PAM‐FL techniques were then used to investigate P–E curves, quantum yield of PSII (F_v/F_m), and chl from 10 K. brevis isolates to determine whether one or all isolates would better represent the species. Comparisons were made with a radial photosynthetron, which allowed for controlled conditions of light and temperature. Isolate α, P_max, and β varied between 0.097 and 0.204 μmol e^? · m^?2 · s^?1 · (μmol quanta · m^?2 · s^?1)^?1, 80.41 and 241 μmol e^? · m^?2 · s^?1, and 0.005 and 0.160 μmol e^? · m^?2 · s^?1 · (μmol quanta · m^?2 · s^?1)^?1, respectively. Either carbon limitation and/or bacterial negative feedback were implicated as the cause of the P–E parameter variability. Furthermore, these results directly contradicted some literature suggestions that K. brevis is a low‐light‐adapted dinoflagellate. Results showed that K. brevis was more than capable of utilizing and surviving in light conditions that may be present on cloudless days off Florida.     

PHOTOSYNTHESIS AND COLD ACCLIMATION: MOLECULAR EVIDENCE FROM A POLAR DIATOM1

The psychrophilic diatom Fragilariopsis cylindrus (Grunow) Krieger in Helmcke & Krieger was used to investigate photosynthesis and growth under freezing temperatures. Gene expression during a temperature shift from +5° C to ?1.8° C was studied under 3 and 35 μmol photons·m^?2·s^?1 by using a macroarray. These measurements were paralleled by determination of fluorescence induction at PSII and pigment analysis. The shift to ?1.8° C at 35 μmol photons·m^?2·s^?1 caused a marginal decrease of photosynthetic quantum yield (F_v/F_m) from 0.61 to 0.52 with fast recovery after 1 day. The ratio of chl c to chl a increased from 3.1 to 5.5, and the ratio of diatoxanthin to diadinoxanthin increased from 0.7 to 5.0. Genes encoding proteins of PSII (psbA, psbC) and for carbon fixation (rbcL) were down‐regulated, whereas genes encoding chaperons (hsp70) and genes for plastid protein synthesis and turnover (elongation factor EfTs, ribosomal protein rpS4, ftsH protease) were up‐regulated. In contrast, cold exposure at 3 μmol photons·m^?2·s^?1 induced a marginal increase in F_v/F_m from 0.61 to 0.63 and a strong increase in fucoxanthin concentrations from 0.04 up to 0.12 pg·cell^?1. This was paralleled by up‐regulation of fcp genes. The ratio of chl c to chl a also increased from 3.1 to 4.2, as did the ratio of diatoxanthin to diadinoxanthin from 0.7 to 2.2. Down‐regulation of psbA, psbC, and rbcL could also be measured but not up‐regulation of hsp70, EfTs, rpS4, and the ftsH protease. The latter genes are probably necessary to avoid cold shock photoinhibition only at higher light intensities.     

COMPARATIVE EFFECTS OF LIGHT ON PIGMENTS OF TWO STRAINS OF EMILIANIA HUXLEYI (HAPTOPHYTA)1

Calcifying and a noncalcifying strains of Emiliania huxleyi were cultured in nutrient replete turbidostats under a photon flux density (PFD) gradient from 50 to 600 μmol E·m^?2·s^?1. For both strains, growth was PFD‐saturated at 300 μmol E·m^?2·s^?1. The strains, although with clearly different physiological properties due to the presence or absence of calcification, showed the same trends and magnitude of change in their pigment compliment as a function of PFD. Light‐controlled pigment composition and the trends of change in pigment composition were identical in both strains. Fucoxanthin (Fuco) was the major carotenoid in the calcifying strain, while in the noncalcifying strain this role was assumed by 19′ hexanoyloxyfucoxanthin (19 Hex). The photoprotective pigments and 19 Hex, normalized to chl a, increased with increasing light, while chl a content per cell and chl c's and Fuco, normalized to chl a, decreased with increasing PFD. The sum of all carotenoids normalized to chl a was remarkably similar in all PFDs used. Collectively, our results suggest that 19 Hex was synthesized from Fuco with light as a modulating factor and that the total amount of carotenoids is strain‐specific and synthesized/catabolized in tandem with chl a to a genetically predefined level independent of PFD.     

EFFECT OF TEMPERATURE ON LIGHT-LIMITED GROWTH AND CHEMICAL COMPOSITION OF SKELETONEMA COSTATUM (BACILLARIOPHYCEAE)1

Phytoplankton growth rate in response to irradiance can be approximated by a hyperbola defined by three coefficients: i) initial slope (α); ii) asymptote (μ_m); and, iii) X-axis intercept or compensation irradiance (I_c). To mathematically represent the interaction of temperature and irradiance on growth rate, one must describe the relationship between these constants and temperature. The marine diatom, Skeletonema costatum (Greville) Cleve, was grown in unialgal culture at different levels of irradiance and 2-3 photoperiods at 0, 5, 10, 16 and 22 C. The value of I_c is ca. 1.0 ly·day^?1 or less at all temperatures. The initial slope (div·ly^?1) is a “u-shaped” function of temperature described by the second degree polynomial, α= 0.25–0.02T+0.001T². Within the range 0–10 C, μ_m (div·day^?1) is an exponential function of temperature described by the equation, μ_m= 0.48 exp (0.126T). At each temperature and selected levels of irradiance, cell size and cellular content of C, N and chl a were determined. The C:chl a and N:chl a ratios increased with irradiance because of increases in C and decreases in chl a. At lower temperatures (0, 5, 10 C), the rate of increase of both ratios with irradiance was greater than at the higher temperatures (16, 22 C). Cellular content of N was independent of irradiance and temperature, and the C:N ratio ranged from 5 to 8 with a slight tendency to lower values at low irradiance. Cell volume was not influenced by either temperature or irradiance.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.     

PHOTOSYNTHETIC FUNCTION IN DUNALIELLA TERTIOLECTA (CHLOROPHYTA) DURING A NITROGEN STARVATION AND RECOVERY CYCLE

Phytoplankton can be exposed to periods of N starvation with episodic N resupply. N starvation in Dunaliella tertiolecta (Butcher) measured over 4 days was characterized by slow reduction in cell chl and protein content and chl/carotenoid ratio and a decline in photosynthetic capacity and maximum quantum yield of photosynthesis (F_v/F_m). In the early stages of N starvation, cell division was maintained despite reduction in cellular chl. Chl content was more sensitive than carotenoids to N deprivation, and cellular chl a was maintained preferentially over chl b under N starvation. NO₃^? resupply stimulated rapid and complete recovery of F_v/F_m (from 0.4 to 0.7) within 24 h and commencement of cell division after 10 h, although N‐replete levels of cell chl and protein were not reestablished within 24 h. Recovery of F_v/F_m was correlated with increases in cell chl and protein and was more related to increases in F_m than to changes in F₀. Recovery of F_v/F_m was biphasic with a second phase of recovery commencing 4–6 h after resupply of NO₃^?. Uptake of NO₃^? from the external medium and the recovery of F_v/F_m, cell chl, and protein were inhibited when either cytosolic or chloroplastic protein synthesis was inhibited by cycloheximide or lincomycin, respectively; a time lag observed before maximum NO₃^? uptake was consistent with synthesis of NO₃^? transporters and assimilation enzymes. When both chloroplastic and cytosolic translation was inhibited, F_v/F_m declined dramatically. Dunaliella tertiolecta demonstrated a capacity to rapidly reestablish photosynthetic function and initiate cell division after N resupply, an important strategy in competing for limiting inorganic N resources.     

THE INABILITY OF THE TEXAS “BROWN TIDE” ALGA TO USE NITRATE AND THE ROLE OF NITROGEN IN THE INITIATION OF A PERSISTENT BLOOM OF THIS ORGANISM1

A planktonic alga similar in general morphology and pigments to Aureococcus anophagefferens Hargraves and Sieburth has caused persistent and ecologically damaging blooms along the south Texas coast. Experiments using 100 μM NO₃^?, NO₂^?, and NH₄⁺ demonstrated that the alga could not use NO₃^? for growth but could use NO₂^? and NH₄⁺. Doubling iron or trace metal concentrations did not permit growth on NO₃^?. Chemical composition data for cultures grown in excess NO₃^? or NH₄⁺, respectively, were as follows: N·cell^?1 (0.88 vs. 1.3 pg), C:N ratio (25:1 vs. 6.4:1), C:chlorophyll a (chl a) (560:1 vs. 44:1), and chl a·cell^?1 (0.033 vs. 0.16 pg). These data imply that cells supplied with NO₃^? were N-starved. Culture addition of 10 mM final concentration chlorate (a nitrate analog) did not affect the Texas isolate while NO₃^? utilizing A. anophagefferens was lysed, suggesting that the NO₃^? reductase of the Texas isolate is nonfunctional. Rates of primary productivity determined during a dense bloom indicated that light-saturated growth rates were ca. 0.45 d^?1, which is similar to maximum rates determined in laboratory experiments (0.58 d^?1± 0.16). However, chemical composition data were consistent with the growth rate of these cells being limited by N availability (C:N 28, C:chl a 176, chl a·cell^?1 0.019). Calculations based on a mass balance for nitrogen suggest that the bloom was triggered by an input of ca. 69 μM NH₄⁺ that resulted from an extensive die-off of benthos and fish.     

BIO-OPTICAL CHARACTERISTICS AND PHOTOADAPTIVE RESPONSES IN THE TOXIC AND BLOOM-FORMING DINOFLAGELLATES GYRODINIUM AUREOLUM,GYMNODINIUM GALATHEANUM,AND TWO STRAINS OF PROROCENTRUM MINIMUM1

Photoadaptive responses in the toxic and bloom-forming dinoflagellates Gyrodinium aureolum Hulbert, Gymnodinium galatheanum Braarud, and two strains of Prorocentrum minimum (Pavillard)Schiller were evaluated with respect to pigment composition, light-harvesting characteristics, carbon and nitrogen contents, and growth rates in shade- and light-adapted cells. The two former species were grown at scalar irradiances of 30 and 170 μmol · m ^?2 at a 12-h daylength at 20° C. The two strains of P. minimum were grown at 35 and 500 μmol. m^?2· s^?1 at a 2-h daylength at 20° C. For the first time, chlorophyll (chl) c₃, characteristic of several bloom-forming prymnesiophytes, was detected in G. aureolum and G. galatheanum. Photoadaptional status affected the pigment composition strongly, and the interpretation of the variation depended on whether the pigment composition was normalized per cell, carbon, or chl a. Species-specific and photoadaptional differences in chl a-specific absorption (°a_c, 400–700 nm) and chl a-normalized fluorescence excitation spectra of photosystem II fluorescence with or without addition of DCMU (°F and °F_DCMU 400–700 nm) were evident. Gyrodinium aureolum and G. galatheanum exhibited in vivo spectral characteristics similar to chl c₃-containing prymnesiophytes in accordance with their similar pigmentation. Prorocentrum minimum had in vivo absorption and fluorescence characteristics typical for peridinin-containing dinoflagellates. Species-specific differences in in vivo absorption were also observed as a function of package effect vs. growth irradiance. This effect could be explained by differences in intracellular pigment content, cell size/shape, and chloroplast morphology/numbers. Light- and shade-adapted cells of P. minimum contained 43 and 17% of photoprotective carotenoids (diadino + diatoxanthin) relative to chl a, respectively. The photoprotective function of these carotenoids was clearly observed as a reduction in °F and °F ^DCMU at 400–540 nm compared to °ac in light-adapted cells of P. minimum. Spectrally weighted light absorption (normalized to chl a and carbon, 400–700 nm) varied with species and growth conditions. The use of quantum-corrected and normalized fluorescence excitation spectra with or without DCMU-treated cells to estimate photosynthetically usable light is discussed. The usefulness of in vitro absorption and fluorescence excitation spectra for estimation of the degradation status of chl a and the ratio of chl a to total pigments is also discussed.