Distinction between light-mediated and light-independent variations in phytoplankton production rates

An array of factors simultaneously controls phytoplankton photosynthesis and hence the primary production process. Because their relative importance shifts both with depth and with season, the significance of individual factors cannot be resolved by in situ incubations, even if all relevant environmental and biotic variables are measured.Here a procedure is described by which in addition to in situ measurements, photosynthesis is simultaneously assessed in identical subsamples under constant temperature (10 °C) and light (0.66 mol m^–2 h^–1 PAR conditions, in vitro). By calculating photosynthesis per unit of chlorophyll, effects of shifting biomass on photosynthesis can be eliminated but seasonal variations of light-saturated photosynthesis generated by temperature, and vertical changes of light-requirements (e.g. by light-shade adaptation) remain obscure. Quotients of in situ photosynthetic rates divided by in vitro rates allow the quantification of light-mediated changes. Provided that photosynthesis measured in vitro is light-saturated, quotients in situ: in vitro rates should never exceed unity. They are a measure for the degree of light-limitation. In vitro rates normalized to chlorophyll give information on temporal changes caused by variations in photosynthetic capacity. In Lake Constance, mean cell size appears to control light-saturated assimilation numbers.     

Interannual variability of phytoplankton productivity and related parameters in Lake Constance: no response to decreased phosphorus loading?

In meso-eutrophic Lake Constance (Germany-Austria-Switzerland),phytoplankton bioraass, pigments and water transparency, aswell as primary productivity, have been followed between 1980and 1989. During this period, municipal phosphorus loading declinedsignificantly. Since 1981, soluble reactive phosphorus (SRP)concentrations during deep lake mixing have decreased from 3.0to currently 1 6 mmol m^–3 at a rate of 7% year^–1.Nitrate concentrations, by contrast, continued to rise. Duringthe period of maximum phosphorus loading, flushing through theoutlet and sedimentation were about equally important sinksof phosphorus from the euphotic zone. Recently, however, sedimentationand subsequent burial of P in the bottom deposits contributedabout three-quarters to the overall P-losses from the systemMain reasons for this shift are unchanged settling fluxes ofphosphorus out of the euphotic zone and decreasing concentrationsof total phosphorus in the water. Only during spring, do concentrations of soluble reactive phosphoruswithin the euphotic zone decrease in proportion to the formationof particulate organic matter. Later during the season, euphoticSRP concentrations continue to be low but are no longer matchedby high plankton biomass because phosphorus is efficiently removedby settling of particles In spite of the observed dramatic decreasein phosphorus loading since 1980, chlorophyll concentrationsand water transparency, as well as annual phytoplankton productivity(300 g C m^–2), have not shown a consistent downward trend.However, the intensity of phosphorus regeneration within theeuphoric zone, which can be used as a measure of the degreeof nutrient limitation, is likely to have increased significantlyThe most probable explanation for the insensitivity of importanttrophic state indicators to reduced nutrient loading is that,in Lake Constance, biomass accumulation to a greater extentis controlled by losses, mainly grazing by zooplankton and sedimentation,than by primary resources. This is concluded from the observationthat phytoplankton biomass always falls far short of the nutrient-dependentcarrying capacity of the system.     

Light-temperature interactions in the control of photosynthesis in Antarctic phytoplankton

Summary During October/November 1983 photosynthetic responses of natural phytoplankton from the Scotia Sea and Bransfield strait to light and temperature were examined in incubators. Both assimilation numbers at saturating light levels and the slopes of the light-limited portions of the photosynthesis versus irradiance curves were smaller than in algae from lower latitudes. However, both parameters increased significantly with rising temperatures. Light-saturated photosynthesis on the average exhibited a Q₁₀-value of ca. 4.2 between-1.5°C and +2°C. Light-limited photosynthesis between-1.5°C and +5°C rose at a rate corresponding to a Q₁₀-value of roughly 2.6. Above +5°C, temperature enhancement of both light-saturated and light-limited photosynthetic rates was minimal or absent. Our results suggest that under extremely low temperatures light-limited photosynthetic rates become temperature-dependent due to changes in maximum quantum yields.     

The impact of phytoplankton on spectral water transparency in the Southern Ocean: implications for primary productivity

Spectral water transparency in the Northern Weddell Sea was studied during Austral spring. The depth of the 1-% surface irradiance level (euphotic depth) varied between 35 and 109 m and was strongly influenced by phytoplankton biomass. Secchi depths were non-linearly related to euphotic depth. In phytoplankton-poor water, the most penetrating spectral region was restricted to a relatively narrow waveband in the blue (488 nm), but the range was broader, between 488 and 525 nm when phytoplankton were abundant. Water transparency in the red spectral range was always low and only to a small extent affected by phytoplankton. Two independent procedures were used to quantify the impact of phytoplankton on spectral water transparency: (1) Regression analysis of spectral in situ vertical light attenuation coefficients in the sea, against coincident chlorophyll concentrations. This method gave chlorophyll-specific light attenuation coefficients; the y-intercept could be interpreted as a measure of light attenuation by pure water plus non-algal material. (2) Spectra of in vivo light absorption derived by spectroscopy, using phytoplankton enriched to varying degrees onto filters. Thus chlorophyll-specific absorption cross-sections were determined. Estimates obtained by both procedures were in close agreement. By integrating over the spectrum of underwater irradiance, in situ chlorophyll-specific absorption cross sections of phytoplankton suspensions, related to all photosynthetically active radiation, were calculated. Light absorption by phytoplankton for photosynthesis is accomplished mainly in the blue spectral range. Also dissolved and particulate organic matter contributed to the attenuation of blue light. Because in water poor in phytoplankton, underwater irradiance was progressively restricted to blue light, chlorophyll-specific absorption cross-sections of phytoplankton, averaged over the spectrum of photosynthetically active irradiance, increased with water depth. In water with elevated phytoplankton biomass, overall light attenuation was generally enhanced. However, because the spectral composition of underwater light changed relatively little with depth, except immediately below the water surface, light absorption cross-sections of phytoplankton changed little below 10 m depth. Vertical differences in the proportions of underwater light absorbed by the phytoplankton community here were mainly dependent on biomass variations. Because of the comparatively small attenuation of blue light by non-algal matter, the efficiency of light harvesting by phytoplankton at any given concentration of chlorophyll in Antractic waters is greater than in other marine regions. At the highest phytoplankton biomass observed by us, as much as 70% of underwater light was available for phytoplankton photosynthesis. When phytoplankton were scarce, <10% of underwater light was harvested by phytoplankton.Contribution within the European Polarstern Study (EPOS), supported by the Deutsche Forschungsgemeinschaft, Grant Ti 115/16-1 to MMT, the European Science Foundation, and by the Alfred Wegener Institut für Polar-und Meeresforschung, Bremerhaven     

Diel phytoplankton periodicity in Mikołajskie lake,Poland, as determined by different methods in parallel

Photosynthetic rates as measured by the oxygen light and dark bottle method were highly correlated with estimates using the ¹⁴C technique. The high O₂/¹⁴C ratios found are explained by algal respiration and extracellular release which are included in photosynthetic measurements by the oxygen technique, while the ¹⁴C method yields values close to net photosynthesis. Separation of net- and nannoplankton using a 50 μm plankton net for filtration was not comparable to distinctions made by microscopic examination. Separation of both by filtration caused a significant decrease in the photosynthetic activity of nannoplankton in 24-hour incubations, but had no detectable effect after 4 hours of exposure. “Bottle effects” in 24-hour measurements of photosynthesis were similar using both methods. Asymmetric photosynthetic time-curves as well as vertical phytoplankton migrations were the main reasons for errors in estimates of daily photosynthetic rates from part-day incubations which were extrapolated to the entire day.     

Influence of Sediment Inflow on Phytoplankton Primary Productivity in Lake Tahoe (California-Nevada)

Light, nutrient concentrations and phytoplankton photosynthesis were studied in a Lake Tahoe sediment plume during maximum spring runoff. They were compared with conditions in clear lake waters not influenced by inlets. In the plume, nutrient concentrations increased in proportion to sediment density whereas light transmission of water was reduced with little effect on the spectral composition except for red light. Light inhibition of photosynthesis at the lake surface was less pronounced in the plume than in clear water and light limitation occurred more rapidly in deeper layers. Evidence from both lake experiments and laboratory bioassays suggests that iron had the greatest stimulatory effect on both photosynthetic activity and biomass growth at maximum sediment densities near the stream inlet. Because of less surface inhibition, photosynthetic light energy utilization efficiency was usually higher in the sediment plume which occurred in relatively shallow areas near the shore. In order to estimate overall effects of enhanced turbidity associated with nutrient loading on Lake Tahoe's primary productivity, profiles taken in shallow areas near the lakeshore were extrapolated to the maximum depth of photosynthesis. Light limitation would cause decreasing productivity, but nutrient stimulation would make this effect less pronounced. The overall effect would depend on the extent of sediment loading relative to nutrient loading.     

SURVIVAL OF SCENEDESMUS ACUMINATUS (CHLOROPHYCEAE) IN DARKNESS1

Survival of the green alga Scenedesmus acuminatus Lagerh. in complete darkness was studied in axenic batch cultures at 7°C and 22°C for three months. The decrease in cell numbers was insensitive to temperature and slower than the loss of dry weight. However, the lag phase before cells began to lyse was more than twice as long at 7° C than at 22°C. The decline in cellular carbohydrates and proteins occurred in two phases. During the first 3-4 days, the decrease in cellular carbohydrate levels was significantly accelerated and temperature-sensitive. Pyrenoids disappeared within 5 days of darkness. Proteins showed 20-fold higher degradation rates at 22°C than at 7°C during the first 4 days. Thereafter, the rates of carbohydrate and protein decomposition were slow and temperature-independent. By contrast, lipids degraded only little at virtually constant and temperature-insensitive rates over the entire experimental period. After three months of dark incubation, about 40% of the remaining cells had retained their growth potential. However, the lag phase, after which cell division was resumed when exposed to light, increased with the duration of the previous dark period. The decrease in photo synthetic potential, which was more pronounced at 22° C than at 7° C, was apparent both in declining maximum assimilation numbers and maximum quantum yields. Cellular chlorophyll a concentrations in surviving cells decreased only slightly. We conclude that the primary means by which S. acuminatus survives extended dark periods is by reduction of catabolic reactions. This was suggested by the slow loss of cell weight. No evidence of significant heterotrophic acetate uptake was found. The initial temperature-dependence of most observed processes indicates that in natural environments chances for survival of algae are augmented by the prevailing low water temperatures.     

Estimation of phytoplankton loss rates from daily photosynthetic rates and observed blomass changes in Lake Constance

Potential growth rates of phytoplankton biomass were estimatedyear-round from production rates and biomass and were comparedwith observed changes in euphotic phytoplankton biomass. Potentialgrowth was always greater than observed growth. The discrepancybetween both is attributed to losses. Relative loss rates showedwide seasonal fluctuations with highest values during the springbloom and autumnal phytoplankton maximum, respectively. Lossrates of photoassimilated carbon showed one peak in late Maywhich lead to a clear-water phase. Relative loss rates werehighly correlated with potential growth rates whereas observedgrowth rates were not. This suggests that most losses occurimmediately after the production process and do not lead toincreases in biomass. During the spring bloom grazing by zooplanktonis the single most important factor leading to losses from thephytoplankton community. During that time, 80–98% of overalllosses can be accounted for by grazing, sedimentation and wash-outcombined. During brief periods in summer and autumn, sedimentationrates comprised >50% of overall losses. In autumn only 30–40%of overall losses were due to the above-mentioned processes.Residual losses can be attributed to respiration, lysis andbacterial remineralization. Grazing, respiration and lysis leadto recycling of carbon and nutrients. Sedimentation rate measurementssuggest an average euphotic carbon regeneration rate of 85%.For the transfer efficiency of carbon along the food chain therelative significance of respiratory losses in overall lossesis of fundamental importance. ¹Dedicated to Professor Elster on his 75th birthday. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP) First International Workshop heldat the Limnological Institute, University of Konstanz, in April1982.     

The water bloom of Cyanobacterial picoplankton in Lake Biwa,Japan

Unicellular autofluorescent picoplankton ranging from 0.4 to 1.5 µm in diameter were found to be a significant component of phytoplankton in the North Basin of Lake Biwa during early summer in 1989 and 1990. The abundance of these picoplankton varied seasonally by about three orders of magnitude with one maximum of up to 10⁶ cells ml^–1. Bloom-forming picoplankton were isolated by dilution and further cultivated in liquid medium. Three clones were found to be representative species of the bloom. Using epifluorescence and electron microscopy as well as absorption and fluorescence emission spectroscopy, we examined these clones according to shape and pigment composition. They have ringlike thylakoids, are photosynthetically active and have no nuclear envelope. The cyanobacterial clones isolated represent three types containing phycobilisomes with either phycocyanin or phycoerthrin as the dominant accessory pigment. They are described here as three new species, two phycoerythrin-rich types and one phycocyanin-rich type, all of them belonging to the Synechococcus group. The differences found by fluorescence emission of isolated clones are discussed with respect to in situ strain identification.