Growth and production of heterotrophic nanoflagellates in a meso-eutrophic lake

The growth of heterotrophic nanoflagellates (HNF) in mesotrophicLake Constance was measured in situ during a 13 month period.Experiments were conducted with 10 µm pre-filtered lakewater incubated in diffusion chambers at 3 m water depth atthe sampling location for 24 h. Growth rates were calculatedfrom changes in cell numbers occurring during the period ofincubation. Growth rates of all dominant taxa showed pronouncedseasonal variation (–0.13 to 1.76 day^–1 and weregenerally highest in summer at high water temperatures. In situgrowth rates were well below maximum growth rates known forthe respective and similar species from laboratory experiments.While water temperature was a key parameter positively relatedto the growth of all HNF species, the effect of various potentialfood items was taxon specific and less clear. Bacterial abundancewas equally important as temperature for growth in the smallbactenvorous Spumella sp., but was insignificant for growthrates of the larger omnivorous Kathablepharis sp. In Spuniellasp., 84% of the observed seasonal variation of its growth ratecould be explained by temperature and bacterial food supply.Based on these results, a multiple linear regression equationwith temperature and bacterial concentration as dependent variableswas calculated for the growth rate of Spumella. Taxon-specificproduction rates were derived from growth rates and averagebiomass of these two species, and compared to total HNF productionestimated from previously measured community growth rates andbiomass in Lake Constance. Production peaks of Spumella sp.and Kathablepharis sp. alternated seasonally. Total HINF productionranged from –0.01 to 10 mg C m^–3 day^–1. Theaverage seasonal production varied between 1.4 and 33 mg C m^–3day^–1 over 6 consecutive years. These small protozoa thuscontribute a substantial amount to total zooplankton productionin Lake Constance.     

On the causes of interspecific differences in the growth-irradiance relationship for phytoplankton. Part I. A comparative study of the growth-irradiance relationship of three marine phytoplankton species: Skeletonema costatum, Olisthodiscus luteus and Gonyaulax tamarensis

Three marine phytoplankton species (Skeletonema costatum, Olisthodiscusluteus andGonyaulax tamarensis) were grown in batch culturesat 15°C and a 14:10 L:D cycle at irradiance levels rangingfrom 5 to 450 µEinst m^–2 s^–1. At each irradiance,during exponential growth, concurrent measurements were madeof cell division, carbon-specific growth rate, photosyntheticperformance (both O₂ and POC production), dark respiration,and cellular composition in terms of C, N and chlorophyll a.The results indicate that the three species were similar withrespect to chemical composition, C:N (atomic) = 6.9 ±0.4, photo-synthetic quotient, 1.43 ± 0.09, and photosyntheticefficiency, 2.3 ±0.1 x 10^–3 µmol O₂ (µgChl a)^–1 h^–1 (µEinst m^–2 s^–1)^–1.Differences in maximum growth rate varied as the –0.24power of cell carbon. Differences in growth efficiency, werebest explained by a power function of Chl a:C at µ = 0.Compensation intensities, ranged from 1.1 µEinst m^–2s^–1 for S. costatum to 35 forG. tamarensis and were foundto be a linear function of the maintenance respiration rate.The results indicate that interspecific differences in the µ–Irelationship can be adequately explained in terms of just threeparameters: cell carbon at maximum growth rate, the C:Chl aratio (at the limit as growth approaches zero) and the respirationrate at zero growth rate. A light-limited algal growth modelbased on these results gave an excellent fit to the experimentalµ–I curves and explained 97% of the observed interspecificvariability. ¹Present address: Lamont-Doherty Geological Observatory Columbiaof University, Palisades, NY 10964, USA     

Cyclops predation on ciliates: species.specific differences and functional responses

Experiments were conducted to measure to what extent cyclopoidcopepods ingest ciliated protists. Five freshwater ciliate species,ranging in size from 22 to 120 µm diameter, were testedwith two species of cyclopoids: Cyclops abyssorum and Cyclopskolensis. Ingestion rates were measured by radiolabeling ciliateswith ¹⁴C, and from these, functional response curves (the changein ingestion rate with changing cell densities) were constructed.Cyclopoids ingest ciliates with very high estimated maximalrates of >200 cells cyclopoid^–1 h^–1 However,there are large differences in ingestion rates that are notpredictable by the size of predator or prey. One ciliate speciesof intermediate size, Coleps hirtus, is nearly immune from cyclopoidpredation at all measured ciliate densities. Three other smallciliate species that move in rapid jumps elicit Honing type3 functional responses, with very little change in ingestionrates at low ciliate densities. Thus, while cyclopoids are capableof having a very considerable impact on ciliate populations,some ciliate species appear to have behavioral, morphologicalor chemical defenses to reduce their vulnerability. This callsinto question the practice of considering ciliates a homogeneousgroup when constructing food web models.     

Biomass, feeding and production of Noctiluca scintillans in the Seto Inland Sea, Japan

The abundance and biomass of the large heterotrophic dinoflagellateNoctiluca scintillans, together with the changes in its potentialprey items, were monitored in the Seto Inland Sea, Japan, duringsummer 1997 (17 July-11 August). Growth and grazing rates ofNscintillans fed natural plankton populations were also measuredeight and seven times, respectively, during the survey period.The abundance and biomass of N scintillans averaged over thewater column (19 m) were in the range 1–345 cells 1^–1(temporalaverage = 93 cell1^–1) and 0.1–49.6 µg C l^–1(temporalaverage = 13.8 µg C l^–1; three times higher thanthat of calanoid copepods during the same period). Noctilucascintillans populations followed the changes in phytoplankton:N.scintillans biomass was increasing during the period of diatomblooms and was at a plateau or decreasing during periods oflow chlorophyll a. The growth rates of N.scintillans (µ)were also consistent with the wax and wane of the N.scintillanspopulation: N.scintillans showed highest growth rates duringdiatom blooms. A simple relationship between µ and chlorophylla concentration was established, and the production of N.scintillanswas estimated using this relationship and the measured biomass.The estimated production averaged over the water column wasin the range >0.1–5.2 µg C l^–1 day^–1(temporalaverage = 1.4 µg C l^–1 day^–1; 64% of the productionof calanoid copepods during the same period). Diatom clearancerates by N.scintillans were in the range 0.10–0.35 mlcell^–1 day^–1, and the phytoplankton population clearanceby N.scintillans was >12% day^–1. Thus, although thefeeding pressure of N.scintillans on phytoplankton standingstock was low, N.scintillans was an important member of themesozooplank-ton in terms of biomass and production in the SetoInland Sea during summer.     

Growth and development of Neocalanus flemingeri/plumchrus in the northern Gulf of Alaska: validation of the artificial-cohort method in cold waters

In situ growth and development of Neocalanus flemingeri/plumchrusstage C1–C4 copepodites were estimated by both the artificial-cohortand the single-stage incubation methods in March, April andMay of 2001–2005 at 5–6°C. Results from thesetwo methods were comparable and consistent. In the field, C1–C4stage durations ranged from 7 to >100 days, dependent ontemperature and chlorophyll a (Chl a) concentration. Averagestage durations were 12.4–14.1 days, yielding an averageof 56 days to reach C5, but under optimal conditions stage durationswere closer to 10 days, shortening the time to reach C5 (fromC1) to 46 days. Generally, growth rates decreased with increasingstage, ranging from 0.28 day^–1 to close to zero but weretypically between 0.20 and 0.05 day^–1, averaging 0.110± 0.006 day^–1 (mean ± SE) for single-stageand 0.107 ± 0.005 day^–1 (mean ± SE) forartificial-cohort methods. Growth was well described by equationsof Michaelis–Menten form, with maximum growth rates (G_max)of 0.17–0.18 day^–1 and half saturation Chl a concentrations(K_chl) of 0.45–0.46 mg m^–3 for combined C1–3,while G_max dropped to 0.08–0.09 day^–1 but K_chl remainedat 0.38–0.93 mg m^–3 for C4. In this study, in situgrowth of N. flemingeri/plumchrus was frequently food limitedto some degree, particularly during March. A comparison withglobal models of copepod growth rates suggests that these modelsstill require considerable refinement. We suggest that the artificial-cohortmethod is the most practical approach to generating the multispeciesdata required to address these deficiencies.     

Development, growth and egg production of Centropages abdominalis in the eastern subarctic Pacific

Centropages abdominalis is a neritic, omnivorous, temporallyabundant copepod present throughout the subarctic Pacific andits marginal seas. The two main objectives of this study wereto determine how temperature influences the development of C.abdominalis and whether growth rates of in situ populationsmay be limited by available food. At 6.9°C, median developmenttime from eggs laid to 50% adults was 42 days and the averageweight-specific somatic growth rate was 0.17 day^–1. At4.6°C, median development time to adult was 59 days (projected)and growth rate averaged 0.08 day^–1, suggesting that 4.6°Cmay be approaching the lower temperature for development andgrowth in this species. The functional relationship betweendevelopment time and temperature was established over the temperaturerange in which this species occurs. The in situ adult growthrates between 10 and 13°C averaged 0.14 day^–1 andwere generally lower than the laboratory-reared juvenile growthrates, which may indicate that adult C. abdominalis are foodlimited in the field during summer and autumn.     

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.     

Experimental studies into the feeding biology of rotifers in brackish water

Mass developments of rotifers of the genus Brachionus, and especiallyof B.quadridentatus, occur regularly in the largely hypertrophicchain of shallow waters (‘boddens’) south of theDarss-Zingst peninsula (Southern Baltic). Interest in the autecologyof the species is, therefore, considerable. Various food sourceswere used in laboratory experiments to ascertain the food requirementsof B.quadridentatus, determine its filtration and ingestionrates, and assess its food particle size-selection ability.Growth experiments showed that the chlorophyceans Nannochlorissp. and Chlorella vulgaris possess considerable nutritionalvalue for the species, whereas abundances declined when Microcystisfirma, detritus from Enteromorpha sp. and only bacteria (Pseudomonas),respectively, were provided as food sources. Filtration ratesvaried between 0.02 and 1.73 µl ind.^–1 h^–1,and ingestion rates between 121 and 5560 cells ind.^–1h^–1, depending on the filtration rate and algal concentration.Investigations into food particle size selection using fluorescentlatex particles revealed that particle size influences foodparticle intake. When particles of different sizes were mixed,the animals showed a preference for the larger particles andingested the smaller ones with a diameter of 1–2 µmless efficiently. The brackish water species Brachionus plicatiliswas studied besides B.quadridentatus in all experiments. Theformer species proved to be superior both in its range of utilizableparticle sizes and its growth rate. The experiments with laboratorycultures were backed up by studies performed with various rotiferspecies taken from natural populations.     

Effects of temperature and salinity on the growth of the red tide flagellates Heterocapsa circularisquama (Dinophyceae) and Chattonella verruculosa (Raphidophyceae)

Growth responses of the red tide flagellates, Heterocapsa circularisquama(Dinophyceae) and Chattonella verruculosa (Raphidophyceae),were examined with 36 different combinations of temperature(5–30°C) and salinity (10–35 PSU). Heterocapsacircularisquama did not grow at or below a temperature of 10°C.The maximum growth rate of H.circularisquama (1.3 divisionsday^–1) was obtained with a combination of 30°C and30 PSU. In contrast, C. verruculosa did not grow at 10 PSU andat temperatures of 25°C or more. The maximum growth rateof C. verruculosa (1.74 divisions day^–1) was obtainedwith a combination of 15°C and 25 PSU. A significant temperature-salinityinteraction on growth was found by factorial analysis. Basedon the physiological characteristics obtained in the presentstudy, these novel flagellates have a potential for future outbreaksof red tides in pre viously unaffected waters.     

Components of Growth and Dark Respiration of Kikuyu (Pennisetum clandestinum Chiov.) at Various Temperatures

Growth and dark respiration were measured in dense, miniatureswards of kikuyu grass grown at constant temperatures of 15,20, 25 and 30 °C. Total respiration over the first 12 hof darkness was very high and CO² efflux per unit surface areavaried from 2.4 to 3.9 g CO₂ m^–2 h^–1 at 15 and 30°C respectively. Such rates were consistent with the correspondinglyhigh net growth rates of 24 and 63 g d. wt m^–2 d^–1and the heavy yields of herbage. When plants were kept in thedark, CO₂ efflux subsequently declined rapidly to a lower, constantrate which was taken to be the maintenance respiration rate.The half-life of the declining phase of respiration averaged10.9 and 6.0 h at 15 and 30 °C respectively, and was curvilinearlyrelated to the specific maintenance respiration rate (m). Therapid decline in respiration was consistent with the low concentrationsof total soluble carbohydrate and starch in the herbage. Valuesof m for lamina and top growth increased with temperature witha Q₁₀ of 2.6 and 1.42 respectively, but m of stems alone wasnot affected by temperature. Using results from this study forkikuyu and from McCree (1974) for sorghum and white clover,it was noted that all three species have similar m when grownat temperatures which are near their respective optimums forgrowth. Kikuyu, Pennisetum clandestinum, growth, respiration, temperature     

Experimental studies of food and suspended sediment effects on growth and reproduction of six planktonic cladocerans

Life-long somatic growth and egg production responses of Daphniabarbata, Daphnia gibba, Daphnia laevis, Daphnia pulex, Moinamicrura and Diaphanosoma excisum were examined in relation todifferent food and suspended sediment levels, in factorial experimentsat 20°C. Food treatments involved additions of 0, 100, 500and 2500 µg C l^–1 of Selenastrum to 20 µmfiltered water, generally from source lakes. These food levelswere tested at two suspended sediment levels in all species,viz. (i) natural source-lake nephelometric turbidities of 7–65nephelometric turbidity units (NTU) ({small tilde}10–70mg l^–1 TSS) and (ii) {small tilde}3- to 4-fold enrichmentof natural levels (8-fold in D.excisum). A third sediment-freetreatment was tested in all species of Daphnia other than D.gibba.Growth in body length and reproductive output were assesseddaily for individual animals. Main and interactive effects offood and sediment on growth were assessed at selected ages fromday 3 through subsequent life using analysis of variance (ANOVA).Strong and highly significant positive food effects were apparentat all ages in all species examined, with the curious exceptionof D.excisum. The onset of maturity and fecundity were alsopositively related to food level. Sediment effects on growth,age of maturity and fecundity were variable, but tended to bemuch weaker than food effects. Both stimulatory (positive, especiallyat low food levels) and inhibitory (negative) effects of sedimenton growth were apparent, both between and within species. Performancein sediment-free water was very poor, except in D.pulex. Food-sedimentinteraction effects were variable. Generally, turbidwater specieswere influenced less adversely by sediment than clear-waterspecies, indicating the existence of environmentally appropriateadaptive responses. Overall, the findings indicate that lightlimitation of algal resources by suspended sediments, ratherthan the condition of high turbidity per se, is responsiblefor the apparently negative impact of suspended sediments frequentlyobserved on natural populations of cladoceran zooplankters.     

Planktonic primary production in the German Wadden Sea

By combining weekly data of irradiance, attenuation and chlorophylla concentrations with photosynthesis (P) versus light intensity(E) curve characteristics, the annual cycle of planktonic primaryproduction in the estuarine part of the Northfrisian WaddenSea was computed for a 2 year period. Daily water column particulategross production ranged from 5 to 2200 mg C m^–2 day^–1and showed a seasonal pattern similar to chlorophyll a. Budgetcalculation yielded annual gross particulate primary productionsof 124 and 176 g C m^–2 year^–1 in 1995 and 1996,respectively. Annual amounts of phytoplankton respiration, calculatedaccording to a two-compartment model of Langdon [in Li,W.K.W.and Maestrini,S.Y. (eds), Measurement of Primary Productionfrom the Molecular to the Global Scale. International Councilfor the Exploration of the Sea, Copenhagen, 1993, pp. 20–36],and dissolved production in 1996, were both in the range of24–39 g C m^–2 year^–1. Annual total net productionwas thus very similar to particulate gross production (127 and177 g C m^–2 year^–1 in 1995 and 1996, respectively).Phytoplankton growth was low or even negative in winter. Inspring and summer, production/biomass (Pr/B) ratios varied from0.2 up to 1.7. Phytoplankton growth during the growth seasonalways surpassed average flushing time in the area, thus underliningthe potential of local phytoplankton bloom development in thispart of the Wadden Sea. The chlorophyll-specific maximum photosyntheticrate (P^B_max) ranged from 0.8 to 9.9 mg C mg^–1 Chl h^–1and was strongly correlated with water temperature (r² = 0.67).By contrast, there was no clear seasonal cycle in ^B, which rangedfrom 0.007 to 0.039 mg C mg^–1 Chl h^–1 (µmolphotons m^–2 s^–1)^–1. Its variability was muchless than P^B_max and independent of temperature. The magnitudeand part of the variability of P^B_max and ^B are presumably causedby changes in species composition, as evidenced from the rangeof these parameters found among 10 predominant diatom speciesisolated from the Wadden Sea. The ratio of average light conditionsin the water column (E_av) to the light saturation parameterE_k indicates that primary production in the Wadden Sea regionunder study is predominantly controlled by light limitationand that nutrient limitation was likely to occur for a few hoursper day only during 5 (dissolved inorganic nitrogen) to 10 (PO₄,Si) weeks in the 2 year period investigated.     

Comparative growth rates and yields of ciliates and heterotrophic dinoflagellates

Growth rates, ingestion rates and grazer yields (grazer volumeproduced/prey volume consumed) were measured for six protozoanspecies (ciliates: Favella sp., Strombidinopsis acuminatum,Uronema sp.; heterotrophic dinoflagellates: Amphidinium sp.,Gymnodinium sp., Noctiluca scintillans) in laboratory batchculture experiments. Comparative growth data indicate that theprymnesiophyte Isochrysis galbana, the prasinophyte Mantoniellasquamata, two cryptophyte species and several autotrophic dinoflagellatespecies were suitable foods for these grazers. When grown onoptimized diets at 13C, maximum ciliate growth rates (range0.77–1.01 day^–1 uniformly exceeded maximum heterotrophicdioflagellate growth rates (range 0.41–0.48 day^–1).A compilation of published data demonstrates that this growthrate difference persists across a range of ciliate and dinoflagellatetaxa and cell sizes. Comparison of volume-specific ingestionrates and yields for the six species studied here showed thatthere was no single explanation for this growth rate disparity.Heterotrophic dinoflagellates exhibited both low ingestion ratesand, in one case, low yields; ciliates were able to achievehigher growth rates via either higher ingestion rates or higheryields, depending on ciliate species. Volume yield increasedover time throughout the exponential growth phase in nearlyall experiments, suggesting variation in response to changingfood concentrations or long-term acclimation to culture conditions.Higher maximum ciliate growth rates mean that these grazershave the potential to exercise tighter control over incipientblooms of their prey than do heterotrophic dinoflagellates.     

Development of Eodiaptomus japonicus Burckhardt (Copepoda, Calanoida) reared on different sized fractions of natural plankton

The nutritional value of different sized fractions of naturalplankton was investigated for the growth of Eodiaptomus japonicusBurckhardt by comparing the development of its naupliar andcopepodid stages fed on differentially fractionated planktonicassemblages of a eutrophic pond, at 20°C. Water filteredthrough a 0.8 µm Nuclepore filter, containing mainly smallcoccoid bacteria (0.45–0.6 µm in cell diameter),at a concentration of 82.7 µg C 1^–1 could not supportthe development of E.japonicus. The 3 µm filtered water,containing bacteria and picoalgae. at a total concentrationof 259 µg C 1^–1, supported development but not eggproduction. The 20 µm filtered water, containing bacteria,picoalgae and large algae, at a total concentration of 2600µg C 1^–1, supported rapid development of the juvenilesand continuous egg production by the adults. The separated 3–20µm fraction, containing only large algae, could not supportthe development at concentrations of 131 and 196 µg C1^–1. However, the same rapid development of the juvenilesand continuous egg production by adults occurred at all of thetested concentrations between 261 and 3920 µg C1^–1of the large algae. The results suggest that E.japonicus favoursalgae larger than 3 µm during its complete lifespan, andthat the threshold food concentration for its development variesbetween 200 and 250 µg C 1^–1.     

Ontogeny of growth, respiration and feeding rate of the freshwater calanoid copepod Eudiaptomus graciloides

The calanoid copepod, Eudiaplomus graciloides, was reared fromegg to adult on uni-algal diets (0.1. 0.5 and 2.5 mg dry wt1^–1) using the green alga, Chlamydomonas reinhardtii,as food, or on a mixed diet consisting of Lake Esrom water filteredthrough a plankton net with pore size 45 µm and supplementedwith C. reinhardtii (2.5 mg dry wt 1^–1). On the mixeddiet at 21.0°C growth in body dry wt (W, µg dry wt)was exponential, and the growth constants were 0.21 day^–1in the early to mid juvenile stage (N1 - C4) and 0.11 day^–1in the late juvenile to early adult stage (C4-A). At 14.5°Cthe corresponding growth rate constants were 0.10 and 0.08 day^–1.Similar growth rates were found at uni-algal concentrationsof 0.5 and 2.5 mg dry wt I^–1, and it was argued that thethreshold concentration for growth in Eudiaptomus was closeto 0.1 mg dry wt I^–1. The clearance (C, ml h^–1)of copepodites was measured on the uni-algal diets. The constantsof the regression (C = aW^b) were: a = 0.125, b = 0.858 (2000C. reinhardtii ml^–1), a = 0.068, b = 0.849 (10 000), a= 0.028, b = 0.875 (50 000). Ingestion rates were calculatedfrom the clearances and the average algal concentrations. Atthe three food levels the average daily rations were 30, 67and 125% of body dry wt. The respiration rate (R, nl O₂ h^–1)was measured in individuals reared on the mixed diet. The constantsof the regression (R = aW^b) were: a = 4.82, b = 1.07 (nauplii,14.5°C), a = 4.17, b = 0.904 (copepodites and adults, 14.5°C),a = 6.87, b = 0.757 (copepodites and adults, 21.0°C). Nosignificant difference in the respiration rate of copepoditesreared on uni-algal diets and the mixed diet could be demonstrated.Energy budgets were calculated. The assimilation efficiencyand the gross growth efficiency of copepodites decreased markedlywith increasing food concentration, the net growth efficiencyvaried from an average of 0.44 at the lowest algal concentrationto 0.60 on the mixed diet. The results are discussed in relationto previous findings with both freshwater and marine copepods.     

Measurement of Yield Threshold and Cell Wal Extensibility of Intact Wheat Roots under Different Ionic, Osmotic and Temperature Treatments

Yield stress threshold (Y) and volumetric extensibility () arethe rheological properties that appear to control root growth.In this study they were measured in wheat roots by means ofparallel measurement of the growth rate (r) of intact wheatroots and of the turgor pressures (P) of individual cells withinthe expansion zone. Growth and turgor pressure were manipulatedby immersion in graded osmoticum (mannitol) solutions. Turgorwas measured with a pressure probe and growth rate by visualobservation. The influence of various growth conditions on Yand was investigated; (a) At 27 °C.In 0.5 mol m^–3 CaCl₂ r, P, Y and were20.7±4.6 µm min^–1, 0.77±0.05 MPa,0.07±0.03 MPa and 26±1.9 µm min^–1MPa^–1 (expressed as increase in length), respectively.Following 24 h growth in 10 mol m^–3 KC1 these parametersbecame 12.3±3.5 µm min^–1, 0.72±0.04MPa, 0.13±0.01 MPa and 21±0.7 µm min^–1MPa^–1. After 24 h osmotic adjustment in 150 mol m^–3mannitol/0.5 mol m^–3 CaCl₂ r= 19.6±4.2 µmmin^–1, P = 0.68±0.05 MPa and Y and were 0.07±0.04MPa and 30±0.2 µm min^–1 MPa^–01, respectively.After 24 h growth in 350 mol m^–3 mannitol/0.5 mol m^–3CaCl₂ r= 13.3±4.1 µm min^–1, P= 0.58±0.07MPa, Y=0.12±0.01 MPa and ø 32±0.2 tim min^–1MPa^–1. During osmotic adjustment in 200 mol m^–3mannitol/0.5 mol m^–3 CaCl₂, with or without KCl, the recoveryof growth rate corresponded to turgor pressure recovery (t1/2approximately 3 h). (b) At 15 °C. Lowered temperature dramatically influencedthe growth parameters which became r= 8.3±2.8 um min^–1,P=0.78 MPa, r=<0.2 MPa and =15±0.1 µm min^–1MPa^–1. Therefore, Y and are influenced by 10 mol m^–3 K⁺ ionsand low temperature. In each case the effective pressure forgrowth (P-Y) was large indicating that small fluctuations ofsoil water potential will not stop root elongation. Key words: Yield threshold, cell wall extensibility, wheat root growth, temperature, turgor pressur     

Effects of nitrogen nutrition on salt-stressed Nicotiana tabacum var. Samsun in vitro plantlets

Tobacco shoots were grown in vitro for 35 d, in MS culture mediummodified to include various sources (nitrate-N, ammonium-N ora mixture) and levels (0–120 mM) of N, and in the presenceof 0–180 mM NaCI or iso-osmotic concentrations of mannitol.Growth of control plantlets was significantly inhibited whenNH₄⁺-N was the sole N source, and at high (120 mM) NO^–₃-N supply. Under conditions of salt stress (90 and 180 mM NaCI)growth was repressed, with roots being more severely affectedthan shoots. Salinity also inhibited root emergence in vitro.The only alleviation of the salt stress by nitrate nutritionobserved in this study was on shoot growth parameters of plantletsgrown on 60 mM NO^–₃-N and 90 mM NaCI. Although both weresignificantly inhibited by NaCI, nitrate reduc-tase activitywas more severely affected than nitrate uptake. When mannitolreplaced NaCI in the culture medium, similar Inhibition of growth,nutrient uptake and enzyme activity were recorded. These observations,together with the relatively low recorded values for Na⁺ andCI^– uptake, indicate that under in vitro salt stress conditionsthe negative effects of NaCI are primarily osmotic. Key words: Growth, nitrogen metabolism, osmotic stress, salinity     

Nitrogen Utilization in N-limited Barley during Vegetative and Generative Growth: I. GROWTH AND NITRATE UPTAKE KINETICS IN VEGETATIVE CULTURES GROWN AT DIFFERENT RELATIVE ADDITION RATES OF NITRATE-N

Growth and nitrate uptake kinetics in vegetatively growing barley(Hordeum vulgare L., cvs Laevigatum, Golf, and Mette) were investigatedin solution culture under long-term limitations of externalnitrogen availability. Nitrate was fed to the cultures at relativeaddition rates (RA) ranging from 0.02 to 0.2 d^–1. Therelative growth rate (RG, calculated for total plant dry weight)correlated well with RA in the range 0.02 to 0.07 d^–1.In the RA range from 0.07 to 0.2 d^–1 RG continued to increase,but an increasing fraction of nitrogen, added and absorbed,was apparently stored rather than used for structural growth.The RG of the roots was less affected by RA. V_max, for net nitrateuptake increased with RA up to 0.11 d^–1, but decreasedat higher RA. The decline in V_max coincided with a build-upof nitrate stores in both roots and shoots. V_max, expressedper unit nitrogen in the plants (the relative V_max, was higherthan required for maintenance of growth (up to 30-fold) at lowRA, whereas at higher RA the relative V_max decreased. Kineticpredictions of steady-state external nitrate concentrationsduring N-limited growth ranged from 0.2 to 5.0 mmol m^–3over the RG range 0.02 to 0.11 d_–1. It is suggested thatthe nitrate uptake system is not under specific regulation atlow RA, but co-ordinated with root protein synthesis and growthin general. At RA higher than 0.11 d_–1, however, specificregulation of nitrate uptake, possibly via root nitrate pools,become important. The three cultivars showed very similar growthand nitrate uptake characteristics. Key words: Barley, growth, nitrogen limitation, nitrate uptake, kinetics     

Density and distribution of bacterioplankton and planktonic ciliates in the Bering Sea and North Pacific

The total number of planktonic bacteria in the upper mixed layerof the Bering Sea during the late spring-early summer periodranged between 1 and {small tilde}4 x 10⁶ ml^–1 (biomass10–40mg C m^–3). In the northern Pacific, along 47–52⁶N,the corresponding characteristics of the bacterioplankton densityin the upper mixed water layer were: total number 1–2x 10⁶ cells ml^–1 and biomass 15–46mg C m^–3Below the thermocline at 50–100 m, the density of bacterioplanktonrapidly decreased. At 300 m depth, it stabilized at 0.1–0.2x 106 cells ml^–1. The integrated biomass of bacterioplanktonin the open Bering Sea ranged between 1.2 and 3.6 g C m^–2(wet biomass 6–18 g m^–2) Its production per dayvaried from 2 to 23 mg C m^–3 days^–1 in the upper0–100 m. The numerical abundance of planktonic ciliatesin this layer was estimated to be from 3 to l0 x 10³ cells l^–1,and in the northern Pacific from 0.4 to 4.5 x 10³ l^–2.Their populations were dominated by naked forms of Strombidium,Strombilidium and Tontonia. In some shelf areas, up to 40% ofthe total ciliate population was represented by the symbioticciliate Mesodinium rubrum. The data on the integrated biomassof basic groups of planktonic microheterotrophs are also presented,and their importance in the trophic relationships in pelagiccommunities of subarctic seas is discussed.     

Plant Growth in Relation to the Supply and Uptake of NO3-: A Comparison Between Relative Addition Rate and External Concentration as Driving Variables

Two approaches to quantifying relationships between nutrientsupply and plant growth were compared with respect to growth,partitioning, uptake and assimilation of NO₃^– by non-nodulatedpea (Pisum sativum L. cv. Marma). Plants grown in flowing solutionculture were supplied with NO₃^– at relative addition rates(RAR) of 0·03, 0·06, 0·12, and 0·18d^–1, or constant external concentrations ([NO₃^–)of 3, 10, 20, and 100 mmol m^–3 over 19 d. Following acclimation,relative growth rates (RGR)approached the corresponding RARbetween 0·03–0.12 d^-1, although growth was notlimited by N supply at RAR =0.18 d^-1. Growth rates showed littlechange with [NO₃–] between 10–100 mmol m^–3(RGR=0·15 –0·16 d^-1). The absence of growthlimitation over this range was suggested by high unit absorptionrates of NO₃^–, accumulation of NO₃^– in tissues andprogressive increases in shoot: root ratio. Rates of net uptakeof NO₃^– from 1 mol m^–3 solutions were assessed relativeto the growth-related requirement for NO₃^–, showing thatthe relative uptake capacity increased with RGR between 0·03–0·06d^–1 , but decreased thereafter to a theoretical minimumvalue at RGR