Stable carbon isotope fractionation by marine phytoplankton during photosynthesis

Abstract. In the marine environment, the range of values of carbon isotope fractionation between particulate tissue of phytoplankton and inorganic carbon can be more than 20‰ (− 35‰ < δ¹³C < − 14‰). This review considers the influence of seawater temperature, lipid content of phytoplanktonic cells, kinetic fractionation, and carbon pathway on δ¹³C values observed at sea.
In order to study the contribution of carboxylases (RUBISCO and the β-carboxylases phosphoenolpyruvate carboxylase, phosphoenoplpyruvate carboxykinase and pyruvate carboxylase) to variations of particulate δ¹³C values at sea, we present results obtained simultenously on carboxylase activities and δ¹³C in various environmental conditions. The lowest δ¹³C values are clearly associated with predominance of ribulose-1.5-bisphosphate carboxylase activity, but it was more difficult to explain the high δ¹³C values. Different hypotheses are discussed.     

Problems in estimating growth rates of marine phytoplankton from short-term14C assays

Growth rate estimates () of phytoplankton populations that were sampled from nitrogen-limited continuous cultures and then incubated for short durations in batch culture with added¹⁴C-HCO₃ ^– were significantly different than steady-state growth rates () for 3 of 5 marine phytoplankton species. Two diatoms,Thalassiosira weissflogii andChaetoceros simplex, displayed virtually identical growth rates (=) over a wide range of, whereas for a third diatom,Phaeodactylum tricornutum, was overestimated by an average of 40% compared to. In contrast, was underestimated by the¹⁴C technique for the two remaining species: up to 40% at a steady-state of 1.0 day^–1 for the chlorophyteDunaliella tertiolecta and up to 100% at of 1.4 day^–1 for the haptophytePavlova lutheri. For the latter two species the divergence between and appeared to increase with increasing steady-state. A simple model of labeled and total carbon flow between the aqueous phase and cellular biomass was constructed to demonstrate that respiration was negligible when=, but was significant when>. In the cases in which<, a rapid physiological alteration presumably took place once the steady state was disturbed and cells were placed in the incubation chambers, which perhaps was related to the nutritional state of the cultures at the time of sampling. Questions thus are raised regarding our ability to measure accurately primary productivity from shipboard experiments with confined samples of phytoplankton from nutrient-impoverished waters that probably are less hardy than the laboratory cultures used in these studies.     

Reduction of marine phytoplankton reproduction rates by copper and cadmium

The reproduction rates of 38 clones of marine phytoplankton were measured in media in which free cupric ion activity was controlled at different levels using a NTA-cupric ion buffer system. The major trend among species in their resistance to copper toxicity was a phylogenetic one, with cyanobacteria being the most sensitive, diatoms the least sensitive, and coccolithophores and dinoflagellates intermediate in sensitivity. The reproduction rates of most of the cyanobacteria were reduced at cupric ion activities above 10⁻¹² M, while most eukaryotic algae still had maximum reproduction rates at 10⁻¹¹ M. Four species, Emiliana huxleyi (Lohm.) Hay & Mohler, Skeletonema costatum (Grev.) Cleve, Thalassiosira pseudonana (Hustedt) Hasle & Heimdal and Thalassiosira oceanica (Hustedt) Hasle were particularly resistant to copper, being able to reproduce well at the highest cupric ion activities tested, 10^−9.5 M and 10^−9.2 M. There was no major difference, however, between neritic and oceanic species in their sensitivity to copper.The sensitivity of 20 species of marine phytoplankton to free cadmium ion activity was measured in a similar manner using an NTA-cadmium ion buffer system. As observed with copper, the prokaryotic cyanobacteria were the most sensitive to cadmium toxicity, diatoms were the least sensitive, and coccolithophores and dinoflagellates were intermediate. All cyanobacteria tested were dead at a cadmium ion activity of 10^−9.3 M whereas the reproduction rates of most of the eukaryotic algae were not reduced significantly until 10^−8.3 M.Comparison of these data with natural concentrations in sea water implies that cadmium is not an important ecological factor in unpolluted waters but natural copper concentrations may inhibit the reproduction of some phytoplankton species, especially cyanobacteria, in upwelled sea water. Copper may influence the seasonal succession of species.     

Influence of solar ultraviolet radiation on photosynthesis and motility of marine phytoplankton

Abstract: The effects of solar ultraviolet radiation (UV) on carbon uptake, oxygen evolution and motility of marine phytoplankton were investigated in coastal waters at Kristineberg Marine Research Station on the west coast of Sweden (58° 30'N, 11° 30'E). The mean irradiances at noon above the water surface during the investigation period were: photosynthetic active radiation (PAR, 400–700 nm) 1670 μmol m⁻² s⁻¹; ultraviolet-A radiation (UV-A, 320–400 nm) 35.9 W m⁻² and ultraviolet-B radiation (UV-B, 280–320 nm) 1.7 W m⁻². UV-B radiation was much more attenuated with depth in the water column than were PAR and UV-A radiation. UV-B radiation could not be detected at depths greater than 100–150 cm. Inhibition of carbon uptake by UV-A and UV-B in natural phytoplankton populations was greatest at 50 cm depth and the effects of UV-B were greater than those of UV-A. At depths greater than 50 cm there was almost no effect of ultraviolet radiation on carbon uptake. PAR, UV-A and UV-B decreased oxygen evolution by the dinoflagellate Prorocentrum minimum . Inhibition of oxygen evolution was greater after 4 h than 2 h but it was not possible to distinguish the negative effects of the different light regimes. The motility of P. minimum was not affected by PAR, UV-A and UV-B. The importance of exposure of phytoplankton to different light regimes before being exposed to natural solar radiation is discussed.     

The V-type ATPase enhances photosynthesis in marine phytoplankton and further links phagocytosis to symbiogenesis

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The effect of pH upon the photosynthesis of littoral marine algae

Summary The photosynthetic rates of a number of littoral marine algae were determined over a pH range from 8.1 to 10.3, employing the Winkler technique to measure evolved oxygen. In a large number of red, brown and green algae, the rate fell to a low value, or to complete inhibition, at pH 9.5 and higher. This indicates, in conformity with general permeability theory, that the HCO₃ ^}-ion cannot readily penetrate the cells, and is hence not available for photosynthesis.However, in two genera of coralline red algae (Bossea andCorallina) photosynthesis persists up to pH 10 or higher (though at a considerably reduced rate). This can be interpreted as due to utilization of the HCO₃ ^}-ion (though probably not the CO₃=ion). But several other non-coralline red algae (Centroceras, Botryocladia andGastroclonium), as well as the brown algaPelvetia (and possiblyFucus), and the green algaeUlva andEnteromorpha display this same ability.Thus the utilization of HCO₃ ^}-ion cannot alone be ascribed as the cause of calcium carbonate deposition, since several other non-calcareous algae have this same power. The utilization of HCO₃ ^}-at least permits calcification, however. The question might be reversed, to ask whyUlva, Enteromorpha. Pelvetia and several red algae are not calcified. Some cell wall property may be involved.     

The effect of UV-B radiation on photosynthesis and respiration of phytoplankton,benthic macroalgae and seagrasses

Several species of marine benthic algae, four species of phytoplankton and two species of seagrass have been subjected to ultraviolet B irradiation for varying lengths of time and the effects on respiration, photosynthesis and fluorescence rise kinetics studied. No effect on respiration was found. Photosynthesis was inhibited to a variable degree in all groups of plants after irradiation over periods of up to 1 h and variable fluorescence was also inhibited in a similar way. The most sensitive plants were phytoplankton and deep-water benthic algae. Intertidal benthic algae were the least sensitive to UV-B irradiation and this may be related to adaptation, through the accumulation of UV-B screening compounds, to high light/high UV-B levels. Inhibition of variable fluorescence (F_v) of the fluorescence rise curve was a fast and sensitive indicator of UV-B damage. Two plants studied, a brown alga and a seagrass, showed very poor recovery of F_v over a period of 32 h.Abbreviations F_m- fluorescence yield with reaction centres closed - F_o- fluorescence yield with reaction centres open - F_v- variable fluorescence - PAR- photosynthetically active radiation - P680- primary donor of Photosystem II - O- primary quencher of Photosystem II - Q_A- primary quinone acceptor of Photosystem II - UV-B- ultraviolet B     

The effects of continuous light and light intensity on the reproduction rates of twenty-two species of marine phytoplankton

The acclimated reproduction rates of 22 species of marine phytoplankton were measured at 0.01, 0.023, 0.1, and 0.23 ly/min in continuous light and in a 14: 10 h light: dark cycle. Three species that reproduced exponentially at all four light intensities in the 14: 10 LD regime did not reproduce at all in continuous light at any of the light intensities. One species, which reproduced at the two lowest light intensities in the 14: 10 LD regime, failed to reproduce at all in continuous light at any light intensity examined. Seven species reproduced more slowly in continuous light than in the 14: 10 LD regime at most or all light intensities. Four species reproduced at roughly the same rate in both light regimes. Five species reproduced more rapidly in continuous light.

No general phylogenetic trend could be discerned from the responses of the species to the different light intensities or to continuous light. In general, species from coastal regions can reproduce as rapidly or more rapidly in continuous light than in a 14: 10 LD cycle, while most species from oceanic regions are harmed by continuous light. A phylogenetic trend in maximum potential reproduction rate is apparent, with diatoms being the fastest, dinoflagellates the slowest, and coccolithophores somewhat intermediate.     

The role of phytoplankton photosynthesis in global biogeochemical cycles

Phytoplankton biomass in the world's oceans amounts to only 1–2% of the total global plant carbon, yet these organisms fix between 30 and 50 billion metric tons of carbon annually, which is about 40% of the total. On geological time scales there is profound evidence of the importance of phytoplankton photosynthesis in biogeochemical cycles. It is generally assumed that present phytoplankton productivity is in a quasi steady-state (on the time scale of decades). However, in a global context, the stability of oceanic photosynthetic processes is dependent on the physical circulation of the upper ocean and is therefore strongly influenced by the atmosphere. The net flux of atmospheric radiation is critical to determining the depth of the upper mixed layer and the vertical fluxes of nutrients. These latter two parameters are keys to determining the intensity, and spatial and temporal distributions of phytoplankton blooms. Atmospheric radiation budgets are not in steady-state. Driven largely by anthropogenic activities in the 20th century, increased levels of IR- absorbing gases such as CO₂, CH₄ and CFC's and NO_x will potentially increase atmospheric temperatures on a global scale. The atmospheric radiation budget can affect phytoplankton photosynthesis directly and indirectly. Increased temperature differences between the continents and oceans have been implicated in higher wind stresses at the ocean margins. Increased wind speeds can lead to higher nutrient fluxes. Throughout most of the central oceans, nitrate concentrations are sub-micromolar and there is strong evidence that the quantum efficiency of Photosystem II is impaired by nutrient stress. Higher nutrient fluxes would lead to both an increase in phytoplankton biomass and higher biomass-specific rates of carbon fixation. However, in the center of the ocean gyres, increased radiative heating could reduce the vertical flux of nutrients to the euphotic zone, and hence lead to a reduction in phytoplankton carbon fixation. Increased desertification in terrestrial ecosystems can lead to increased aeolean loadings of essential micronutrients, such as iron. An increased flux of aeolean micronutrients could fertilize nutrient-replete areas of the open ocean with limiting trace elements, thereby stimulating photosynthetic rates. The factors which limit phytoplankton biomass and photosynthesis are discussed and examined with regard to potential changes in the Earth climate system which can lead the oceans away from steady-state. While it is difficult to confidently deduce changes in either phytoplankton biomass or photosynthetic rates on decadal time scales, time-series analysis of ocean transparency data suggest long-term trends have occurred in the North Pacific Ocean in the 20th century. However, calculations of net carbon uptake by the oceans resulting from phytoplankton photosynthesis suggest that without a supply of nutrients external to the ocean, carbon fixation in the open ocean is not presently a significant sink for excess atmospheric CO₂.The submitted paper has been authored under Contract No. DE-AC02-76H00016 with the US Department of Energy. Accordingly, the US Government retains a non-exclusive, royalty-free license to publish or reproduce the published form of this contribution, or allow others to do so, for US Government purposes.     

海洋浮游植物的热效应

采用热效应模拟实验方法研究了冬、夏两季温度对象山港海洋浮游植物种类组成、细胞密度的影响,同时在冬季开展了温度对海水中叶绿素含量的影响研究.温度对浮游植物种类数和细胞密度的影响显著,影响程度与季节、增温幅度和实验时间有关.当夏季(自然水温为28℃)实验温度超过36℃,冬季(自然水温为12℃)实验温度超过34℃时,浮游植物种类数和细胞密度均急剧减少.温度对浮游植物细胞密度的影响显著,随着实验温度的升高,浮游植物细胞密度与温度呈峰值关系.夏季实验当温度升高至30℃时浮游植物细胞密度最高,实验至第7天时的细胞密度达到最大值为40.56 cells/dm3;冬季实验当温度升高至24℃时的浮游植物细胞密度最高,实验至第7天时的细胞密度达到最大值为625.8 cells/dm3.温度对冬季实验海水中叶绿素含量的影响显著,并且冬季海水叶绿素浓度与浮游植物细胞密度呈显著的正相关性(r=0.81,p＜0.01).实验结合自然海区海水温度的变化进行分析,并探讨人们普遍关注的热(核)电厂建设与海洋生态效应的问题,为冬、夏两季温排水对受纳海域的增温对浮游植物的热影响提供科学依据.     

Measurement of water drinking rates in marine Crustacea

Tests were made of γ-emitting compounds as potential non-absorbed reference markers for estimating water ingestion in the western rock lobster Panulirus longipus cygnus (George) and the penaeid prawn Penaeus latisulcatus (Kishinouye). The extent of any marker absorption into the tissues was measured in animals totally immersed and in rock lobsters with only the gills perfused. Distribution of marker within the body, transport into the gut, and excretion were examined following injection of the label into the blood of the rock lobster.¹²⁵I-sodium iothalamate, ⁵⁸Co-EDTA, and ⁵¹Cr-EDTA were all satisfactory reference markers with very low absorption, ⁵¹Cr-EDTA being the best. Most injected sodium iothalamate was excreted from the body over 24 h, probably via the urine, which suggests its further use in studies of antennal gland function.⁴⁶ScCl₃, ⁵¹CrCl₃, ⁵⁸CoCl₂, and ¹¹⁰AgCl were of some value for estimating drinking in penaeid prawns, although ¹¹⁰AgCl probably gave levels that were too high. ¹¹⁰AgCl was useless for the rock lobster, which absorbed much more marker than was retained in the gut.¹²⁵I-rose bengal was not practicable because it slowly precipitated in sea water.Requirements for reference markers appear to be more critical for animals with high permeability to major sea-water ions than those which extensively regulate all major ions.     

In situ growth rates of marine phytoplankton: approaches to measurement, community and species growth rates

Taxonomic structure and biomass weighting are important determinantsof measurable marine phytoplankton community growth potential.Maximal diel-averaged growth rates of communities appear tofall between 3 and 3.6 doublings day^–1. Mean net growthrates are considerably lower. Ranges of community growth ratesmeasured in tropical, sub-tropical and (summer) temperate ecosystemsare similar. There appears to be a broad dichotomy between thegrowth potential of diatom species as compared to non-diatoms.Doubling rates of small diatoms frequently exceed communitybiomass doubling rates by wide margins. Diel-averaged growthrates of large diatoms, microflagellates and non-motile ultraplanktonpopulations are lower and similar in magnitude to communitygrowth estimates. Maximum growth rates of species measured insitu are in good agreement with maximum growth rates measuredin laboratory cultures. High specific productivity of sub-dominantor rare diatom species or assemblages will be diluted in thelower specific growth rates of microflagellate and non-motileultraplankton assemblages. Specific rates of grazing upon speciesand functional groups remain to be quantified, but stabilityof community size and taxonomic structure implies close linkagebetween growth and mortality rates at the species level overtime intervals of several generations.     

The effect of water column mixing on phytoplankton succession, diversity and similarity

The lake number was used to describe the mixing condition forthree consecutive years (1992–1994) in Lake Tegel, Berlin,and compared to the successions of diatoms, dinoflagellatesand cyanobacteria, the main phytoplankton groups in the lake,as well as to diversity and similarity indices. Using both diversityand similarity indices in juxtaposition provides an indicationof the growth type of prevailing species (r- or K-strategists)and the degree of competition in the aqua-ecological system.A genera] pattern of these indices can be recognized as threephases: (i) high diversity—during spring, summer and autumn,interrupted by (ii) phases of low diversity during the latespring clear-water phase as the number of spring species plummeted,and (iii) during the late summer, climax populations of K-strategists.On a smaller time scale, similarity and diversity proved toreact sensitively to disturbances at frequencies intermediatein relation to the generation times of the phytoplankton. Thissupports the ‘intermediate disturbance hypothesis’,as proposed for phytoplankton by Padisak et al. [(eds) IntermediateDisturbance Hypothesis in Phytoplankton Ecology. Kluwer Academic,1993]. Diversity may remain quite high even for extended periodsduring summer climax situations, in conjunction with a highdegree of similarity, if deeper mixing of the epilimnion occursat time intervals of 2–3 weeks, as during the summer of1993. This enables the prevalence of ‘ruderal’ species,together with some motile K-strategists who actively seek optimaldepths for photosynthesis. During such summer situations describedby frequent occurrences of lower lake numbers, the epilimnionof Lake Tegel is mixed deeply enough to support ruderals, butnot too vigorously to counteract competitive advantages of motilespecies. Thus, vertical niche separation enhances diversity.     

The influence of turbidity on net phytoplankton photosynthesis in some Irish lakes

SUMMARY. An important influence on gross photosynthetic rates per unit area is the success with which phytoplankton competes for the available light with non-algal suspended material and dissolved organic compounds. Using a range of Irish lakes, with euphotic zones varying between 0.7 and 20 m and with chlorophyll-α values between 1 and 860 mg m⁻³, the effect on gross rates of photosynthesis is analysed for changes in the balance between the factors contributing to light attenuation. Net values per unit area are also likely to be modified in well-mixed systems as the ratio of light to dark regions in the water column are altered with changes in light penetration. Depth gradients in dark respiration are reported for L. Neagh which vary according to previous light history and nutrient stress. Possible cases of restraint on phytoplankton growth are discussed for optically deep situations.     

Stimulation of phytoplankton photosynthesis by bottom-ice extracts in the Arctic

Chlorophyll-specific photosynthetic rates of marine phytoplankton collected under landfast sea ice in the Canadian Arctic were stimulated by additions of a chelator, ethylenediamine tetra-acetic acid (EDTA), and trace metals. This stimulation was imitated by filtered extracts of bottom ice colonized by sea-ice algae. Compared to controls, the assimilation rates for experimental additions averaged 166%, 184%, and 119% for ETDA, trace metals, and ice extracts, respectively. All experimental treatments displayed similar oscillations consistent with tidal forcing where mixing and photosynthetic performance are enhanced during spring tides. These results suggest that some bioactive soluble material(s) produced within the bottom-ice algal layer acts as a "conditioning" agent that enhances the growth of phytoplankton in arctic waters. The bioactive agent(s) remains unidentified.     

The effect of water stress on photosynthesis and related parameters in Pinus halepensis

Net photosynthesis, transpiration, dark respiration rates and stomatal and mesophyll resistances were studied in young potted seedlings of Pinus halepensis Mill. under gradually decreasing soil and leaf water potentials. Stomatal resistance under non-limiting xylem water potentials was 6–7 times higher than mesophyll resistance. Stomata started to close at threshold xylem water potentials of −0.8 MPa, whereas mesophyll resistance started to increase at about −1.4 MPa. Decreasing xylem water potentials increased the CO₂ compensation point and decreased the water use efficiency (expressed by the photosynthesis to transpiration ratio) and dark respiration rate. It is concluded that at least part of the drought resistance characteristics of P. halepensis are associated with a sensitive stomatal mechanism which enables an efficient control of water loss.     

Effect of heavy metals on marine phytoplankton

Phytoplankton community responses were used in a heavy metals (HM) marine pollution monitoring biological system. A multifactorial analysis was used in in situ experiments in the White Sea to study HM toxicity to a planktonik algae population in their copresence. It was ascertained that algae with a high rate of increase in population size, producing metabolites, restricting development of phytoplankton community associates, gain advantages in isolation conditions. The algae responded differently to addition of individual metals and their combinations in the environment. Change in the number of dominants resulted in disturbance of the phytoplankton community structure. It has been shown that there are cardinal points in phytoplankton development, determining the growth of individual algae populations and planktonic community on the whole, against a background of competition for biogenic elements and space in conditions of TM environment pollution.