No abstract

The effects of nitrate, phosphate, and iron starvation and resupply on photosynthetic pigments, selected photosynthetic proteins, and photosystem II (PSII) photochemistry were examined in the diatom Phaeodactylum tricornutum Bohlin (CCMP 1327). Although cell chlorophyll a (chl a) content decreased in nutrient‐starved cells, the ratios of light‐harvesting accessory pigments (chl c and fucoxanthin) to chl a were unaffected by nutrient starvation. The chl a‐specific light absorpition coefficient (a*) and the functional absorption cross‐section of PSII (σ) increased during nutrient starvation, consistent with reduction of intracellular self‐shading (i.e. a reduction of the “package effect”) as cells became chlorotic. The light‐harvesting complex proteins remained a constant proportion of total cell protein during nutrient starvation, indicating that chlorosis mirrored a general reduction in cell protein content. The ratio of the xanthophylls cycle pigments diatoxanthin and diadinoxanthin to chl a increased during nutrient starvation. These pigments are thought to play a photo‐protective role by increasing dissipation of excitation energy in the pigment bed upstream from the reaction centers. Despite the increase in diatoxanthin and diadinoxanthin, the efficiency of PSII photochemistry, as measured by the ration of variable to maximum fluorescence (Fv/Fm) of dark‐adapted cells, declined markedly under nitrate and iron starvation and moderately under phosphate starvation. Parallel to changes in Fv/Fm were decreases in abundance of the reaction center protein D1 consistent with damage of PSII reaction centers in nutrient‐starved cells. The relative abundance of the carboxylating enzyme, ribulose bisphosphate carboxylase/oxygenase (RUBISCO), decreased in response to nitrate and iron starvation but not phosphate starvation. Most marked was the decline in the abundance of the small subunit of RUBISCO in nitrate‐starved cells. The changes in pigment content and fluorescence characteristics were typically reversed within 24 h of resupply of the limiting nutrient.

No abstract

The diadinoxanthin cycle (DD-cycle) in chromophyte algae involves the interconversion of two carotenoids, diadinoxanthin (DD) and diatoxanthin (DT). We investigated the kinetics of light-induced DD-cycling in the marine diatom Phaeodactylum tricornutum and its role in dissipating excess excitation energy in PS II. Within 15 min following an increase in irradiance, DT increased and was accompanied by a stoichiometric decrease in DD. This reaction was completely blocked by dithiothreitol (DTT). A second, time-dependent, increase in DT was detected ∼ 20 min after the light shift without a concomitant decrease in DD. DT accumulation from both processes was correlated with increases in non-photochemical quenching of chlorophyll fluorescence. Stern-Volmer analyses suggests that changes in non-photochemical quenching resulted from changes in thermal dissipation in the PS II antenna and in the reaction center. The increase in non-photochemical quenching was correlated with a small decrease in the effective absorption cross section of PS II. Model calculations suggest however that the changes in cross section are not sufficiently large to significantly reduce multiple excitation of the reaction center within the turnover time of steady-state photosynthetic electron transport at light saturation. In DTT poisoned cells, the change in non-photochemical quenching appears to result from energy dissipation in the reaction center and was associated with decreased photochemical efficiency. D1 protein degradation was slightly higher in samples poisoned with DTT than in control samples. These results suggest that while DD-cycling may dynamically alter the photosynthesis-irradiance response curve, it offers limited protection against photodamage of PS II reaction centers at irradiance levels sufficient to saturate steady-state photosynthesis.

The relationship between the diadinoxanthin cycle and changes in fluorescence yield in the diatom Chaetoceros muelleri Lemm. (clone CH10, Amorient Aquafarm, Inc., Hawaii) was investigated. High‐light‐induced changes in fluorescence yield and xanthophyll de‐epoxidation occurred very rapidly (first order rate constant 1.60 min−1). The observed light‐induced changes in diatoxanthin and diadinoxanthin concentration were consistent with a two‐pool scheme for diadinoxanthin, one of which does not undergo de‐epoxidation. Changes in xanthophyll concentration correlated with changes in in vivo absorbance indicating that diadinoxanthin cycle activity in vivo can be monitored spectrophotometrically. However, changes in cell absorbance were small relative to total optical absorption cross section. Increases in the concentration of diatoxanthin were linearly correlated with increases in the rate constant for thermal de‐excitation in the antenna of photosystem II (PSII). Antenna quenching produced or mediated by diatoxanthin may, thus, protect the PSII reaction center in diatoms. Changes in the maximum fluorescence yield suggested that changes in the reaction center also contributed to nonphotochemical quenching of fluorescence. Thus, reaction center quenching affected the relationship between antenna quenching and changes in photochemical efficiency producing the effect of a decrease in fluorescence yield without a decrease in photochemical efficiency.

Nutrient enrichment experiments were conducted m May and June of 1993 at 8 stations along a North Atlantlc transect, from Morocco to Nova Scotia, Canada. Variable fluorescence (F,/F,,) was measured in order to estimate the health or physiological state of the population as a whole. Low values across the transect indicated nutrient limited photosynthetic efficiency and probable growth rates ranging from 10 to about 50% of p,,. Where the lowest value was measured, over the Grand Banks of Newfoundland, Canada, nitrogen addition to incubated samples resulted in large, significant increases in photochemical efficiency. Numbers and cell-specific fluorescence of 3 major groups of picophytoplankton were studied using flow cytometry, in order to further quantlfy the physiological response to nutrient additions. Results ind~cated nitrogen linxtation of physiology and/or abundance of small eukaryotes, cyanobacteria, and prochlorophytes. Abundance (cell numbers) and cellular fluorescence of the 3 groups responded differently to nutrient additions. Prochlorophytes showed the greatest response to incubation in terms of cell numbers, responding especially to nitrogen addition. By contrast, cyanobacterial numbers did not change from initial values or with treatment, although cell pigment content did. Cellular fluorescence as measured by the flow cytometer reflected cell pigment content in most experiments. Increased cellular fluorescence of all groups in nitrogen-amended treatments relat~ve to unamended controls lnd~cated physiological limitation by nitrogen.

The xanthophyll cycle has been implicated as a possible photoprotective mechanism in higher plants and algae by dissipating excess excitation energy via non-photochemical q.uenching. To examine whether colonial Phaeocystis antarctica Karsten displays xanthophyll cycling, nutrient-replete c u l t u r~s were initially grown under limiting (40 pm01 quanta m -2 s~' ) and saturating (280 pm01 quanta m-2 S -' ) irradiances for photosynthesis and their responses to irradiance transitions were examined for 1 h under 4 treatments. The ~I I v~v o chl-specific absorption coefficient [aml+,(h), m2 (mg chl a)-'] for the light-limiled cultures was initially lower than the light-saturated cultures while chlorophyll (chl) anormallzed fluorrscence yields were similar for both treatments. Increases in irradiance induced increases in the diatoxanthin to diadinoxanthin ratio (DT:DD, w:w) up to 9-fold whereas parallel decreases in Irradiance similarly decreased the DT:DD ratio. Light-induced Increases in DT concentration were reduced in cultures exposed to dithiothreltol (DTT), an inhibitor of DD to DT conversion. Short-term changes in DD and DT concentrations were attributed solely to xanthophyll cycling; no de novo synthesis of DD or DT was evident based on a constant sum of DD and DT in the 1 h expenmental perturbations. It was found that DD and DT de novo synthesis required long-term acclimation; the mass ratio at steady state of (DD+DT)/chl a was 0.1 and 0.4 for the low and high light treatments, respectively. Pooled results from treatment and control cultures showed a linear relationship between light-induced changes in DT/chl a concentration and F/chl a (fluorescence to chl a ratio) and the slopes depended on the initial photoacclimated state of the culture. Cellular fluorescence changes appeared to be physiologically based; aaph()L) did not change in response to abrupt irradiance changes. Xanthophyll cycling may enable P antarctica to tolerate both high light environments and sudden changes in irradidnce, which occur during austral spring due to shallow mixed layers and intermittent shading by ice or clouds.

Abstract. Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985)(1986)(1987)(1988)(1989)) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (< 10 #m) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m -2 d -1, and integrated biomass levels reached 415 to 972 mg chlorophyll m -2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) by Chaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV= 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV=20 to 33%) than intra-annual variations (CV=85 to 120%).