The rate of growth of asterionella formosa Hass. in relation to its ecology

Summary In Windermere, and some other lakes in the English Lake District, the vernal increase of Asterionella formosa ceases when the concentration of silica in the water falls to approximately 0.5 mg/l. When this planktonic diatom is cultivated in water from one of these lakes, Windermere, the addition of small amounts of phosphate phosphorus permits the growth of so large a crop that all the silica present is incorporated into the cells. The intensity and duration of the illumination and the temperature of the water effect this relationship between silica, phosphorus and the growth Asterionella. Therefore, the regular, observed relationship between the concentration of silica and the vernal maximum of Asterionella in nature may be related to the amount of this substance in the water, to the amount of phosphate phosphorus present and to the rate of growth imposed upon this diatom by the light and temperature conditions in the lake during the period concerned. The possible importance of the relatively low concentration of phosphate phosphorus in certain very unproductive lakes is discussed.Written in honour of Prof. Dr. E. G. Pringsheim.     

The Isolation, Maintenance and Host Range Studies of a Chytrid Rhizophydium planktonicum Canter emend., Parasitic on Asterionella formosa Hassall

The methods of isolation and maintenance in dual clone cultureof the chytrid Rhizophydium planktonicum Canter emend, parasiticon Asterionella formosa Hass. from the plankton are described.The ability of a single fungal isolate to infect other clonesof Asterionella, Fragilaria, Tabellaria, Synedra and Cyclotella,as well as dead material was also tested. All the clones of Asterionella proved to be highly susceptibleto infection whereas only in rare instances could a sporangiumbe found which had grown and dehisced on Fragilaria and Synedra.Very few zoospores encysted on Tabellaria and Cyclotella andthose which did died without further growth. There was no evidenceto suggest that the chytrid could complete its life historyon dead material. freshwater phytoplankton, culture, parasitism, host–parasite relationships, chytridiomycetes, Rhizophydium, Zygorhizidium, diatoms, Asterionella, Fragilaria     

Determination of growth-limiting nutrients for red species of Oscillatoria and two ‘oligotrophic’ diatoms

Laboratory batch experiments with dilute phytoplankton communities were carried out to determine growth-limiting nutrients and the degree of growth limitation for Asterionella formosa, Hass., Tabellaria fenestrata (Lyngb.) Kütz. and red species of Oscillatoria under specific temperature and light conditions. Water samples from five Norwegian lakes with average epilimnetic total phosphorus concentrations ranging from 5 to 30 µg P per 1 were investigated. Both enrichment and transplant experiments were carried out. In samples from the most oligotrophic lakes (Lake Randsfjorden and Lake Tyrifjorden) phosphorus was found to be the growth-limiting nutrient. In samples from Lake Mjøsa silicate was also growth-limiting for the diatoms, while nitrogen could be limiting for both asterionella and Oscillatoria from Lake Steinsfjorden. Phosphorus was often the only limiting nutrient determined for Oscillatoria in Lake Gjersjøen (the most eutrophic of the lakes). In samples from this lake, however, Asterionella was also growth-limited directly or indirectly as a result of high (pH > 9). This was also found for Asterionella and Tabellaria from lake Mjøsa by means of transplant experiments. Oscillatoria from Lake Gjersjøen could not grow in filtered water from Lake Mjøsa but obtained maximum growth rate after addition of phosphate and chelated iron in combination.     

The Effect of Light and Darkness Upon Infection of Asterionella formosa Hassall by the Chytrid Rhizophydium planktonicum Canter emend

Observations, both experimental and microscopic, indicate thatzoospores of Rhizophydium planktonicum Canter emend., can remainalive, but rarely become adherent upon cells of Asterionellaformosa Hassall under conditions of very low light or in completedarkness. The behaviour patterns of zoospores towards host cells underdarkened conditions were compared with those which took placeunder well illuminated conditions leading to the settlementof zoospores on host cells. The differences noted may help toexplain the lack of zoospores found upon inadequately illuminateddiatom cells. Some evidence suggests that young zoospores lack the abilityto adhere to host cells. After the encystment of zoospores uponAsterionella cells in the light, their further growth and developmentcan continue in darkness. Rhizophydium planktonicum Canter emend., Asterionella formosa Hassall, chytrid, diatom, infra-red illumination, zoospore     

Evidence of interference of Asterionella formosa with the feeding of Bosmina longirostris: a field study in a south Andes lake

Intensive sampling of a small temperate lake of the Araucanian region (Argentina), revealed changes in the demography of Bosmina longirostris when the diatom Asterionella formosa density increased (to 54 000 cells ml^–1). Bosmina population behaviour in the field was consistent with the idea that large numbers of Asterionella caused its starvation. Individuals were actually colourless and fecundity dropped to zero, as was evident when Asterionella reached 99.4% of total phytoplankton biovolume. Since cyanophytes, as a whole, are scarce in these lakes, we propose that the flagellate/diatom ratio has a direct effect on cladoceran demography.     

Physico-chemical control of the growth of a diatom, Asterionella formosa Hass., in a shallow eutrophic lake

Physico-chemical environmental control of the growth rate ofan algal population, the pennate diatom Asterionella formasaHass., was investigated for a year in natural water. This speciesoccurred in vegetative form in the water column all the time,although its population density varied by five orders of magnitudefrom a maximum of 1.4?10⁶ cells l^–1 in February to a minimumof 0.8?10¹ cells l^–1 in August. A mathematical model ofa combination of multiplicative and Liebig types suggested thatvegetative cells were severely limited in their growth rate,reaching almost 80% depression of the maximum rate between Decemberand February by low temperatures and nearly 50% depression inwarm seasons by light intensity. Nutrient limitations were onlyobserved in May, August and September, although they were notas great as by temperature and light intensity in the studylake. Population changing rates of A.formosa determined in thelake were low and agreed well with the estimated growth ratesduring winter, but those in summer were low and disagreed withthe estimated high growth rates. This suggests that populationchange of the species was highly dependent upon the specificgrowth rate in winter, but other factors became predominantin summer.     

BIOTIN LIBERATION BY THE LICHEN ALGA COCCOMYXA SP. AND BY CHLORELLA PYRENOIDOSA

The unicellular green alga Coccomyxa, a component of the lichenPeltigera aphthosa, liberated about 7.2mµg biotin permg dry weight of cells into the culture medium during a growthperiod of 15–20 days. The corresponding figure for thefree-living alga Chlorella pyrenoidosa was 0.45mµg ofbiotin. Chromatographic analysis indicated that this was freebiotin and not a bound form of the vitamin. The biotin concentrationof rinsed Coccomyxa cells was 1.88mµg per mg dry weightof cells, of which less than 0.01mµg was extractable byhot water. Cells of Chlorella contained 0.16mµg of biotinper mg dry weight, of which 0.11mµg was extractable byhot water. The biotin content of Coccomyxa, which was about12 times that of Chlorella, is thus almost entirely in the boundform. The importance of biotin in the symbiotic interactionsbetween the alga and the fungus in Peltigera is discussed. ¹Present address: University Department of Agriculture, Oxford,England. ²Present address: Institute of Marine Resources, Universityof California, La Jolla, California, U.S.A.     

Factors controlling production of phytoplankton and bacteria under ice in a humic, boreal lake

The factors controlling pelagic primary and bacterial productionof a humic, boreal lake in winter were investigated, combininglaboratory and field experiments where some of the predictedconsequences of the climate change, i.e.the increased load ofphosphate-phosphorus and dissolved organic matter (DOM), weresimulated. In situ incubations were performed in eight acrylictubes lowered underneath the ice cover for 1–4 months.In the lake, production of phytoplankton (0.03–0.33 µgC l^–1 day^–1) was lower than that of bacteria (0.2–2.3µg C l^–1 day^–1) from the end of January tomid-April. Later in spring, the light conditions improved dueto the disappearance of snow and finally ice itself, and primaryproduction was revived. The importance of light as the factorcontrolling primary production in winter was confirmed in laboratoryexperiments where additions of phosphorus and DOM did not enhancethe primary production. These same enrichments resulted in highergrowth rates and production of bacteria. However, bacterioplanktonwas simultaneously controlled by heterotrophic flagellates,as in the laboratory experiments the exclusion of flagellatesalways resulted in higher yields of bacteria of increased cellsize. There was also some evidence that the quality of substratesis of importance for microbial activities. The importance oftemperature was highlighted with the significant correlation(r² = 0.59) between bacterial production and temperature withinthe range 0.6–2.1C. Thus, on the boreal zone the possiblewarming of climate may lead to enhanced activities in winter,but under unchanged light conditions no effects on primary productionwill be expected.     

A study of phosphate limitation in Lake Maarsseveen: phosphate uptake kinetics versus bioassays

In order to assess possible phosphate limitation for the phytoplankton community of Lake Maarsseveen, two techniques (phosphate uptake experiments and bioassays) were employed simultaneously in February–March 1982. In that period the ambient phosphate concentration of the lake water was less than 0.03 µM P and the diatom Asterionella formosa constituted more than 90% of the phytoplankton population. The phosphate uptake experiments showed relatively high uptake capacities and low cell phosphorus contents for the natural phytoplankton community. This suggested phosphate limitation throughout the test period. The growth stimulation of the phytoplankton after enrichment with phosphate, however, only revealed phosphate limitation from the beginning of March and bioassays may therefore be regarded as a less sensitive method.     

Ion and Glycerol Concentrations in 12 Isolates of Dunaliella

Ginzburg, M., and Ginzburg, B. Z., 1985. Ion and glycerol concentrationsin 12 isolates of Dunaliella.—J. exp. Bot. 36: 1064–1074. Twelve isolates of Dunaliella with average cell volumes rangingfrom 50 to 1400x10^–18 m³ were grown in batch culture at0.5 M or 2.0 M NaCl. Glycerol and ions (Na⁺, K⁺, Mg²⁺, CI^–,phosphate) were measured in log-phase cultures. The contentsof Mg²⁺, K⁺ and phosphate per cell were found to be a functionof cell-volume. Cell glycerol, Na⁺ and Cl^– were functionsof cell-volume and of the NaCl concentration in the medium.Solute concentrations were calculated from the measured cell-volumesand from the ³H₂O content of pellets corrected for intercellularspace using Blue Dextran. Cell glycerol was found to accountfor about one-half of the expected osmolarity, the remainderbeing largely accounted for by Na⁺ and CI^–. Key words: —Dunaliella, isolates, glycerol, ion concentrations     

The control of sub-surface maxima of diatoms in a stratified lake by physical, chemical and biological factors

The sedimentation patterns of Asterionella formosa Hass. wereobserved in two lake basins, in one of which a mid-water populationmaximum of the diatom occurred. Comparison of the alga's distributionwith vertical profiles of temperature and dissolved reactivesilicon suggested that the formation of a mid-water maximumwas associated with a sharp silicon gradient. Sinking velocitiesof the diatom were reduced at the maximum compared to thoseat lesser or greater depths. The suspension of cultures at differentdepths, in either nutrient-replete medium or exposed to theprevailing nutrient conditions, following thermal stratification,confirmed that an increase of sinking velocity occurred in silicon-or light-deficient populations. It is suggested that the mid-watermaximum of A.formosa was formed by differential colony sinkingvelocities with depth in response to variations in silicon availability.The formation of a strong silicon gradient was dependent uponthe growth of the population being restricted to the epilimnionby the onset of thermal stratification.     

Phytoplankton and nutrients in the waters north-west of Spitsbergen in the autumn of 1979

Phytoplankton biomass, primary production rates and inorganicnutrients were measured in the uppermost layer of the ice-edgeregion and in open water and compared with environmental factorsduring a three-week cruise in September – October 1979.Biomass and production values were low (maximum 2.2 µgchl a l^–1, 2.5 mg C m^–3 h^–1). A post-bloomcommunity of diatoms, consisting mainly of representatives ofChaetoceros, Leptocylindrus, Nitzschia and Thalassiosira, waspredominant. Concentrations of phosphate were quite low (maximum0.55 µM I^–1). Nitrate and silicate ranged from nomeasurable quantities to 5.7 µM l^–1 and 3.8 µMl^–1, respectively. The possibility of light and nutrientlimitation on phytoplankton growth is discussed.     

Effect of Sodium on Phosphate Uptake in Unicellular and Filamentous Cyanobacteria

The effect of Na⁺ on phosphate uptake was studied in four strainsof cyanobacteria: Synechococcus PCC 7942, Gloeothece PCC 6501,Phormidium sp. and Chlorogloeopsis PCC 6912. Phosphate uptakewas stimulated by Na⁺ in all cases. Li⁺ and K⁺ acted as partialanalogues for Na⁺. Half-saturation [K_1/2(Na⁺)] of phosphateuptake was reached with Na⁺ concentrations ranging from 317µM in Chlorogloeopsis to 659 µM in Phormidium. Theconcentration of phosphate required to reach half-saturationof phosphate uptake [K_1/2(P_i)]was not changed by the presenceof Na⁺. (Received April 11, 1994; Accepted July 5, 1994)     

Biochemical Limitations to Carbon Assimilation in C3 Plants--A Retrospective Analysis of the A/Ci Curves from 109 Species

Species-specific differences in the assimilation of atmosphericCO₂ depends upon differences in the capacities for the biochemicalreactions that regulate the gas-exchange process. Quantifyingthese differences for more than a few species, however, hasproven difficult. Therefore, to understand better how speciesdiffer in their capacity for CO₂ assimilation, a widely usedmodel, capable of partitioning limitations to the activity ofribulose-1,5-bisphosphate carboxylase-oxygenase, to the rateof ribulose 1,5-bisphosphate regeneration via electron transport,and to the rate of triose phosphate utilization was used toanalyse 164 previously published A/C_i, curves for 109 C₃ plantspecies. Based on this analysis, the maximum rate of carboxylation,Vc_max, ranged from 6µmol m^–2 s^–1 for the coniferousspecies Picea abies to 194µmol m^–2 s^–1 forthe agricultural species Beta vulgaris, and averaged 64µmolm^–2 s^–1 across all species. The maximum rate ofelectron transport, J_max, ranged from 17µmol m^–2s^–1 again for Picea abies to 372µmol m^–2 s^–1for the desert annual Malvastrum rotundifolium, and averaged134µmol m^–2 s^–1 across all species. A strongpositive correlation between Vc_max and J_max indicated that theassimilation of CO₂ was regulated in a co-ordinated manner bythese two component processes. Of the A/C_i curves analysed,23 showed either an insensitivity or reversed-sensitivity toincreasing CO₂ concentration, indicating that CO₂ assimilationwas limited by the utilization of triose phosphates. The rateof triose phosphate utilization ranged from 4·9 µmolm^–2 s^–1 for the tropical perennial Tabebuia roseato 20·1 µmol m^–2 s^–1 for the weedyannual Xanthium strumarium, and averaged 10·1 µmolm^–2 s^–1 across all species. Despite what at first glance would appear to be a wide rangeof estimates for the biochemical capacities that regulate CO₂assimilation, separating these species-specific results intothose of broad plant categories revealed that Vc_max and J_maxwere in general higher for herbaceous annuals than they werefor woody perennials. For annuals, Vc_max and J_max averaged 75and 154 µmol m^–2 s^–1, while for perennialsthese same two parameters averaged only 44 and 97 µmolm^–2 s^–1, respectively. Although these differencesbetween groups may be coincidental, such an observation pointsto differences between annuals and perennials in either theavailability or allocation of resources to the gas-exchangeprocess. Key words: A/C_i curve, CO₂ assimilation, internal CO₂ partial pressure, photosynthesis     

The importance of benthic recruitment to the population development of Aphanizomenon flos-aquae and internal loading in a shallow lake

The recruitment of Aphanizomenon flos-aquae out of the sedimentswas measured in the shallow, hypereutrophic Agency Lake, Oregon.There were two main episodes of recruitment, the latter of whichaccounted for 8.2% of the water column population increase duringits main growth phase. Phosphorus (P) content was higher innewly recruited cells than in water column cells. Recruitmentduring one weekly episode constituted an internal P load rateof 3.56 mg P m^–2 day^–1, accounting for 27% of thecorresponding increase in water column total phosphorus (TP).However, the majority of internal loading in the lake did notcoincide with, and therefore could not be explained by, biotranslocationdue to benthic recruitment of A.flos-aquae. ¹Present address: WES-ES-P, USAE Waterways Experiment Station,3909 Halls Ferry Road, Vicksburg, MS 39180, USA     

Quantification of the phosphorus released by zooplankton in an oligotrophic lake (La Caldera, Spain): regulating factors and adjustment to theoretical models

Using an in situ approach, we have evaluated the phosphorusinputs from zooplankton in a high-mountain oligotrophic lake.Values of the specific gross release rate (SGRR) fluctuatedbetween 0.2 and 2.9 µg P mg^–1 dry weight h^–1,and were higher when the nauplii of Mixodiaptomus laciniatusdominated the zooplankton community. The rate of P recyclingby the zooplankton was high, reaching 1.6 µg P l^–1day^–1, and showed a highly consistent seasonal patternfrom one year to the next, with maxima in midsummer. Zooplanktonsize accounted for as much as 85% of the variance obtained inthe measurements of the specific rate of P release, while otherfactors, such as the quality or quantity of food, did not significantlyinfluence the SGRR changes. Among the models tested, only theone proposed by Peters (Int, Ver. Theor. Angew. Limnol. Verh.,19, 273–279, 1975) was useful for predictions in thissystem. The stoichiometric model of Hessen and Andersen (Arch.Hydrobiol., 35, 111–120, 1992), applied in this oligotrophicsystem, adequately predicted the phyto- and zooplankton dynamics,whereas the values of P release estimated using this model werefar higher than the excretion rates obtained experimentally.These differences were related to the type of egestion (formationof faecal pellets) of metazooplankton and to the relative importanceof the food resistance to digestion. We believe that in communitieswhere copepods constitute a substantial percentage of the zooplankton,an evaluation of the P release which is readily available [solublereactive phosphate (SRP), total dissolved phosphate (TDP)] toalgae and bacteria would not fit the predictions of generalmodels of mass balance; under these circumstances, assimilationefficiency proves to be the key parameter for predicting thereadily available P (excreted P).     

Rates of Photosynthesis in Characean Cells: I. PHOTOSYNTHETIC 14CO2 FIXATION BY NITELLA TRANSLUCENS

A study has been made of photosynthetic ¹⁴CO₂ fixation by isolated‘mature’ internodes of Nitella translucens. Experimentalconditions were similar to those used in studies of the ionicrelations of these cells. Maximum rates of photosynthesis were33–40µµmoles CO₂, fixed per cm² of surfacearea per second (equivalent to 12–15 /xmoles fixed permg chlorophyll per hour). ^l4CO₂ fixation was inhibited to thedark level by 3(3,4,dichlorophenyl)-1, 1-dimethylurea (at 0-6µM or 10µM) and by the uncoupler carbonyl cyanide-m-chlorophenylhydrazone(SµM). The presence of imidazole or ammonium sulphate(both of which uncouple ATP production in vitro) did not resultin an inhibition of 14CO2 fixation. These results are discussedin relation to published work on solute uptake by Nitella translucens.During photosynthesis there was rapid movement of ¹⁴C-labelledorganic compounds out of the chloroplasts. ¹⁴C-labelled sucrose,ammo-acids, and sugar phosphates were found in samples of vacuolarsap.     

Photoadaptation in Antarctic phytopfankton: variations in growth rate, chemical composition and P versus I curves

The response of phytoplankton to variations in the light regimewas studied during the VULCAN and ACDA cruises in the Antarctic.Unenriched batch cultures of 12–19 days' duration reachedchl concentrations of 10–50 µg^–1 and exhibitedexponential growth rates, with the maximal rate being 0.41 doubl,day^–1. Ice edge algae exhibited maximum growth rates atphoton flux densities (PFD) of 30–100 µE m^–2S^–1and the growth rate was reduced by about 30% at 500–1000µE m^–2S^–1 The chl/C ratio ranged between 0.004and 0.018, with the lowest ratios at PFDs above 500 µEm^–2S^–1 chl/C ratios were also below maximum at PFDsbelow 40–50 µE m^–2S^–1 The C:N:P ratioswere close to the Redfield ratios; the Si/C ratio averaged 0.16(atoms), and the ATP/C ratio averaged from 0.0024 to 0.0050in different culture senes. When thawed after having been frozenfor 10 days, shade-adapted cultures were in a much better conditionthan sun-adapted ones. P versus I data showed that the maximumassimilation number varied from 0.75 to 4.4 µg C (µgchl)^–1h^–1. It varied inversely with the chl/C ratio;therefore the maximum carbon turnover rate varied little betweensamples (0.024/0.035 h^–1). Low biomass communities exhibitedrelatively high values for (the initial slope of P versus Icurves), low values for 1_sat (160–330 µE m^–2S^–1),and they were susceptible to photoinhibition. In contrast, communitiesdominated by Odontella weissflogii exhibited low values for, a high value for I_sat (560 µE m^–2S^–1 andthey tolerated high PFDs. The photo-adaptational status of thephytoplankton in natural water samples is discussed relativeto the profile of water column stability and mixing processes.     

Feeding and growth rates of the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

The goal of this research is to enhance our knowledge of thecontributions of doliolids to the planktonic community as consumersand secondary producers. The objectives are to quantify feedingand growth rates of Dolioletta gegenbauri gonozooids at fourfood concentrations and four temperatures in order to determinetheir impact as grazers throughout the water column. Althoughdoliolids are abundant in numerous regions of the coastal ocean,and are considered to be major planktonic grazers, data on ratesof feeding and growth are scarce. Laboratory experiments wereconducted at 16.5, 20, 23.5 and 26.5°C to quantify removalof a 50:50 volumetric concentration of Thalassiosira weissflogiiand Rhodomonas sp. at four different food concentrations of20, 60, 160 and 390 µg C l^–1. Results from theseexperiments suggest that clearance rates are similar at concentrationsfrom 20 to 60 µg C l ^–1, and decrease as the foodconcentrations increase to 160 and 390 µg C l ^–1.The ingestion rates increase over a range of phytoplankton concentrationsfrom 20 to 160 µg C l ^–1, then decrease when abnormallyhigh concentrations of 390 µg C l ^–1 are offered.Clearance and ingestion rates increase as temperature increasesfrom 16.5 to 26.5°C. The exponential growth rates rangefrom k = 0.2–0.7, with the lowest rates occurring at thehighest food concentration. Growth rates increase with increasingtemperature from K = 0.1–0.3 at 16.5°C to 0.45–0.7at 26.5°C. In each case, the small- and medium-sized zooidshad higher growth rates than the larger gonozooids. These resultssuggest that doliolid feeding and growth rates are a functionof environmental food concentrations and temperatures, and implythat they can be important consumers in a changing neritic environment.     

Subcellular Distribution of Inorganic Phosphate, and Levels of Nucleoside Triphosphate, in Mature Maize Roots at Low External Phosphate Concentrations: Measurements with 31P-NMR

Maize plants were grown in nutrient solution without phosphate,or in which inorganic phosphate (Pi) was maintained at nearlyconstant concentrations of 1 µM, 10µM or 0·5mM. In vivo ³¹P-NMR measurements showed that there was no discernibledifference in the cytoplasmic Pi content (µmol cm^–3root volume) of the mature roots of plants exposed to 1 µM,10µM or 0·5 mM external phosphate for up to 12d. However, the vacuolar Pi content of the mature roots variedabout 10-fold between these three groups. The cytoplasmic Pi content of roots receiving no external phosphatedecreased significantly after about 7 d total growth, and atabout this time the vacuolar pool of Pi became too small foraccurate measurement. The presence of 1 µM Pi in the nutrientsolution completely prevented this decline in cytoplasmic Pi,and there was some evidence that it also raised the Pi contentof the root vacuoles above the almost undetectable level foundin the totally P-starved roots. During the first 7–9 d of growth, the nucleoside triphosphatecontent of the mature roots was unaffected by the concentrationof phosphate in the nutrient solution. The results highlight the close control of cytoplasmic concentrationsof certain important phosphorus metabolites in roots growingin soil of normal agricultural fertility. Key words: Vacuole, cytoplasm, intracellular compartmentation, NTP, P-nutrition