On the causes of interspecific differences in the growth-irradiance relationship for phytoplankton. II. A general review

The causes of interspecific differences in the µ-l relationshipare examined in the context of a mechanistic model which relatesµ to irradiance in terms of six factors:, k_c photosyntheticquotient (PQ), Chl a:C, respiration and excretion. The effectof cell size on the light saturated growth rate is also considered.It is shown that photosynthetic efficiency and PQ exhibit remarkablylittle interspecific variability, and average 0.024 ±0.005 µg C(µg Chl a)^–1 h^–1 (µEm^–2 s^–1)^–1 and 1.5 ± 0.2 mol 02 molC^–1 (when NO₃^– is the nitrogen source) respectively.Two useful relationships were derived: (i) between growth efficiency,_g and Chl a:C at µ. = 0; (ii) between the compensationintensity, I_c and the Chl a-specific maintenance respirationrate. Both relationships were independent of temperature anddaylength. Species best adapted to growth at low light werefound to exhibit high Chl a:C ratios and low maintenance respirationrates. As a group, diatoms were consistently the best adaptedfor growth at low irradiance. Chiorophytes, haptophytes, chrysophytesand cryptophytes were intermediate in their performance at lowirradiance. Dinoflagellates exhibited extreme diversity, withspecies spanning the spectrum from very good performance atlow irradiance to very poor. A new µ_max-cell carbon relationshipis given based on growth rates normalized to 15°C. Evidenceis presented to show that noise in this relationship can besignificantly reduced by using only carbon-specific growth ratesand using only data for species grown at the same daylength.     

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     

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.     

The optical properties of a turbid reservoir and its phytoplankton in relation to photosynthesis and growth (Mount Bold Reservoir, South Australia)

In situ light measurements were used to obtain information oninherent and apparent optical properties. The average verticalattenuation coefficient K_d(ave) varied from 1.1 to 4.6 In unitsm^–1 During three periods the variation in K_d(ave) correlatedwith changes in chlorophyll a concentration and specific attenuationcoefficients K_s, of 0.013, 0.014 and 0.022 m² mg Chl a^–1were calculated. Chlorophyll-specific diffuse absorption coefficients(A,) for these periods were 0.012. 0.013 and 0.017 m² mg Chla^–1 and only varied significantly from estimates of K_sin the period when scattering was intense. Absorption coefficientsa(z^mid) and scattering coefficients b(z^mid) calculated for themid-point of the euphotic zone ranged between 0.45 and 2.9 mand 3.5–52.0 m respectively. Chlorophyll-specific absorptioncoefficients K_a, of 0.005, 0.006 and 0.007 m² mg Chl a^–1and scattering coefficients K_b of 0.05. 0.09 and 0.191 m² mgChl a^–1 were measured during the three periods. The highK_b value occurred when gas-vacuolate cyanobactena were dominant.Algal photosynthesis and light absorption were related throughthe maximum quantum yield _m which varied between 0.019 and 0.11mol C Einstein^–1 while average quantum yields _a, variedbetween 0.006 and 0.024 with a mean of 0.013 mol C Einstein^–1A comparison of changes in the mean irradiance of the mixedzone and chlorophyll concentration indicated that growth waslight limited below 0.04–0.05 Einsteins absorbed mg Chla^–1 day^–1.     

Temporal and vertical variation of chlorophyll a concentration, phytoplankton photosynthetic activity and light attenuation in Lake Kinneret: possibilities and limitations for simulation by remote sensing

The relationship between chlorophyll a (Chl a) and primary productivity(PP) in the uppermost water layer and the water column-based(0–15 m) integral values of those variables were examinedusing measurements taken in Lake Kinneret (Israel) from 1990to 2003. In 81% of all Chl a profiles examined, the distributionwas fairly uniform within the entire 0–15 m water column,and 12.3% of instances showed a prominent subsurface maximum,when the lake phytoplankton was dominated by the dinoflagellatePeridinium gatunense. Chl a can be reliably estimated by remotesensing techniques in the productive and turbid water of LakeKinneret, since Chl a concentration at surface layers can beextrapolated to the entire water column. Light vertical attenuationcoefficient average for wavelengths from 400 to 700 nm, K_d,ranged from 0.203 to 1.954 m^–1 and showed high degreeof temporal variation. The maximal rate of photosynthetic efficiency,P_B^opt [average 3.16 (±1.50)], ranged from 0.25 to 8.85mg C m^–3 h^–1 mg Chl a^–1. Using measured dataof Chl a, P_B^opt, and light as an input, a simple depth-integratedPP model allowed plausible simulation of PP. However, a lackof correlation between photosynthetic activity and temperature(or other variable with remotely sensed potential) renders theuse of models that require input of photosynthetic efficiencyto calculate integrated PP of little value in the case of productiveand turbid Lake Kinneret.     

Size-fractionated primary production studies in the vicinity of the Subtropical Front and an adjacent warm-core eddy south of Africa in austral winter

Results are presented of size-fractionated primary productionstudies conducted in the vicinity of the Subtropical Front (STF),an adjacent warm-core eddy, and in Sub-antarctic waters duringthe third South African Antarctic Marine Ecosystem Study (SAAMESIII) in austral winter (June/July) 1993. Throughout the investigation,total chlorophyll (Chl a) biomass and production were dominatedby small nano- and picophytoplankton. No distinct patterns intotal Chl a were evident. At stations (n = 7) occupied in thevicinity of the STF, total integrated biomass values rangedfrom 31 to 53 mg Chl a m^–2. In the vicinity of the eddy,integrated biomass at the eddy edge (n = 3) ranged from 24 to54 mg Chl a m^–2 and from 32 to 43 mg Chl a m^–2 inthe eddy (n = 2). At the station occupied in the Sub-antarcticwaters, total integrated biomass was 43 mg Chl a m^–2.Total daily integrated production was highest at stations occupiedin the vicinity of the STF and at the eddy edge. Here, totalintegrated production ranged from 150 to 423 mg C m^–2day^–1 and from 244 to 326mg C m^–2 day^–1, respectively.In the eddy centre, total integrated production varied between134 and 156 mg C m^–2 day^–1. At the station occupiedin the Sub-antarctic waters, the lowest integrated production(141 mg C m^–2 day^–1) during the entire survey wasrecorded. Availability of macronutrients did not appear to limittotal production. However, the low silicate concentrations duringthe survey may account for the predominance of small nano- andpicophytoplankton. Differences in production rates between theeddy edge and eddy core were related to water column stability.In contrast, at stations occupied in the vicinity of the STF,the control of phytoplankton production appears to be relatedto several processes, including water column stability and,possibly, iron availability.     

Growth and development of Metridia pacifica (Copepoda: Calanoida) in the northern Gulf of Alaska

Juvenile growth and development rates for Metridia pacifica,one of the dominant larger copepods in the subarctic Pacific,were investigated from March through October of 2001–2004in the northern Gulf of Alaska. The relationship between prosomelength (PL, µm) and dry weight (DW, µg) was determined:log₁₀ DW = 3.29 x log₁₀ PL – 8.75. The stage durationsof copepodites ranged from 3 to 52.5 days but were 8–15days under optimal condition. Seasonally, growth rates increasedfrom March to October and typically ranged between 0.004 and0.285 day^–1, averaging 0.114 ± 0.007 day^–1(mean ± SE). After standardization to 5°C (Q₁₀ of2.7), growth rates averaged 0.083 ± 0.005 day^–1and were significantly correlated to chlorophyll a, with saturatedgrowth rates of 0.149 day^–1 for C1–C3, 0.102 day^–1for C4–C5 and 0.136 day^–1 for all stages combined.Measured juvenile growth rates were comparable with specificegg production rates in this species. The comparisons of ourrates in this study with those predicted by the global modelsof copepod growth rates suggested that further refinement ofthese models is required.     

Microzooplankton grazing in a coastal embayment off Concepcion, Chile, (~36{degrees} S) during non-upwelling conditions

The impact of grazing by natural assemblages of microzooplanktonwas estimated in an upwelling area (Concepción, Chile)during the non-upwelling season in 2003 and 2004. Seawater dilutionexperiments using chlorophyll a (Chl a) as a tracer were usedto estimate daily rates of phytoplankton growth and microzooplanktongrazing. Initial Chl a concentrations ranged from 0.4 to 1.4mg Chl a m^–3 and phytoplankton prey biomass and abundancewere numerically dominated by components <20 µm. Phytoplanktongrowth and microzooplankton grazing rates were 0.19–0.25day^–1 and 0.26–0.52 day ^–1, respectively.These results suggest that microzooplankton exert a significantremoval of primary production (>100%) during the non-upwellingperiod.     

Maintenance and Growth Components of Carbon Dioxide Efflux from Growing Pea Fruits

Carbon dioxide efflux from 5- to 20-day-old pea fruits was measuredfor plants grown in controlled environment at 15 °C and600 µmol s^–1 m^–2 photon flux density in a16 h photoperiod. The rate of CO₂ output per fruit increasedquickly from 0.005 to 0.018 mg CO₂ min^–1 during fruitelongation and subsequently more slowly to 0.030 mg CO₂ min^–1as the fruits inflated. On a d. wt basis the rate was highest,0.175 mg CO₂ g^–1 min^–1, in the youngest fruits anddeclined curvilinearly with increasing fruit weight to 0.02mg CO₂ g^–1 min^–1. Separation of maintenance andgrowth components was achieved by starvation methods and bymultiple regression analysis. From the latter method estimatesof the maintenance coefficient declined hyperbolically from150±8.7 mg carbohydrate g^–1 d. wt day^–1 inthe very young fruits (0.05 g) to 10.4±0.36 mg carbohydrateg^–1 d. wt day^–1 in older fruits (2.0 g). On a nitrogenbasis maintenance costs decreased from 2240 to 310 mg carbohydrateg^–1 nitrogen day^–1 while nitrogen concentrationfell from 6.7 to 3 per cent d. wt. A simple linear relationshipbetween maintenance cost per unit d. wt and nitrogen concentrationwas not observed. A growth coefficient of 50±6.7 mg carbohydrate g^–1growth (equivalent to a conversion efficiency, Y_G, of 0.95)was estimated for all fruits examined. The overall efficiency, Y, increased from a mean of 0.70 to0.85 during fruit elongation and subsequently declined to 0.80.For a given fruit weight, efficiency increased asymptoticallywith relative growth rate; both asymptote and slope of the relationshipincreased as the fruits grew. Pisum sativum L., garden pea, legume fruit, carbon dioxide efflux, maintenance respiration, growth respiration     

A comparison of seasonal growth and development of the copepods Calanus marshallae and C. pacificus in the northern Gulf of Alaska

The juvenile growth rates and development times of subarcticCalanus marshallae and temperate/sub-tropical C. pacificus wereinvestigated during nine cruises (May through October, 2001–04)in the northern Gulf of Alaska. The artificial cohort methodbased on a length-weight regression was used for growth estimatesand the reciprocal of the molting rate for developmental time.The copepodite stage duration ranged from 3 to 16 days for C.marshallae (C1–C4) and 3–23 days for C. pacificus(C1–C5). Seasonally, copepodid growth rates increasedfrom May to October, ranging between 0.055 and 0.291 day^–1(mean ± SE: 0.176 ± 0.008 day^–1) for C.marshallae, while growth rates increased from August to Octoberbetween 0.018 and 0.296 day^–1 (mean ± SE: 0.142± 0.016 day^–1) for C. pacificus. After standardizationto 5°C (Q₁₀ of 2.7), growth rate averaged 0.118 ±0.007 day^–1 and 0.075 ± 0.009 day^–1 for C.marshallae and C. pacificus, respectively. Calanus marshallaegrowth rate is satisfactorily described by a Michaelis–Mentenmodel using chlorophyll-a concentration (r² = 0.33) after temperaturecorrection, but the prediction improves with a composite nonlinearmodel combining body weight into the Michaelis–Mentenfunction (r² = 0.55). Considering the limited range of dataavailable for C. pacificus, the combination of the data forboth species suggests that C. pacificus has a similar functionalresponse to growth despite the differences in the geographicand temporal distributions with C. marshallae. Measured juvenilegrowth rates of the two Calanus species in this study were comparableto other calanoid species in the same area and showed reasonableagreement to Calanus growth models but less with global copepodgrowth models.     

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.     

Community structure, biomass and productivity of size-fractionated summer phytoplankton populations in lower Narragansett Bay, Rhode Island

Seventeen size-fractionation experiments were carried out duringthe summer of 1979 to compare biomass and productivity in the< 10, <8 and <5 µm size fractions with that ofthe total phytoplankton community in surface waters of NarragansettBay. Flagellates and non-motile ultra-plankton passing 8 µmpolycarbonate filters dominated early summer phytoplankton populations,while diatoms and dinoflagellates retained by 10 µm nylonnetting dominated during the late summer. A significant numberof small diatoms and dinoflagellates were found in the 10–8µm size fraction. The > 10 µm size fraction accountedfor 50% of the chlorophyll a standing crop and 38% of surfaceproduction. The <8 µm fraction accounted for 39 and18% of the surface biomass and production. Production by the< 8 µm fraction exceeded half of the total communityproduction only during a mid-summer bloom of microflagellates.Mean assimilation numbers and calculated carbon doubling ratesin the <8 µm (2.8 g C g Chl a^–1 h^–1; 0.9day^–1)and<5 µm(1.7 g C g Chl a^–1h^–1; 0.5day^–1)size fractions were consistently lower than those of the totalpopulation (4.8 g C g Chl a^–1 h^–1; 1.3 day^–1)and the <10 µm size fraction (5.8 g C g Chl a^–1h^–1; 1.4 day ^–1). The results indicate that smalldiatoms and dinoflagellates in fractionated phytoplankton populationscan influence productivity out of proportion to their numbersor biomass. ¹Present address: Australian Institute of Marine Science, P.M.B.No. 3, Townsville M.S.O., Qld. 4810, Australia.     

Respiratory ETS activity of plankton in the northwestern Alboran Sea: seasonal variability and relationship with hydrological and biological features

Respiratory electron transport system (ETS) activity was measuredin plankton samples (<200 µm) collected in the NW AlboranSea. Sampling was carried out during seasonal cruises (summerand autumn 2003 and winter and spring 2004) in 12 stations locatedin transects off the coast of Malaga (southern Spain). Thiswork reports for the first time seasonal variations of the Arrheniusactivation energy (Ea) as well as being the first study to addressCO₂ balance in the NW Alboran Sea. These variations were relatedto changes in the phytoplankton community assemblage, whichcould ultimately be caused by the seasonal variability of hydrologicalconditions. ETS activity was significantly higher in summer,coinciding with a higher chlorophyll a (Chl a) concentrationand relatively high levels of particulate organic matter. TheETS:Chl a_total ratios were low during the four seasons, suggestinga high contribution of autotrophic phytoplankton to the respiratoryactivity of planktonic community. Respiratory CO₂ production(RCP) calculated from ETS activity ranged from 4.6 to 28.1 mgC m^–3 day^–1 during the four cruises. Chl a-specificRCP was lower than the maximum photosynthetic rates reportedin the literature for the studied area, suggesting that primaryproduction (PP) and respiration in the water column might beunbalanced.     

Production of Oikopleura longicauda (Tunicata: Appendicularia) in Toyama Bay, southern Japan Sea

Oikopleura longicauda occurred throughout the year in ToyamaBay, southern Japan Sea, and analysis of its size compositionand maturity revealed that reproduction was continuous overtheyear. Somatic growth production (P_g) varied with season from0.03 to 103 mg carbon (C) m^–2day^–1 (annual P_g 4.5g C m^–2), and house production (P_e) from 0.11 to 266 mgC m^–2 day^–1 (annualP_e 11.3 g C m^–2). The annualP_g/B ratio was 176. Compared with production data of some predominantzooplankton species in Toyama Bay, it is suggested that despitetheir smaller biomass, appendicularians are an important secondaryproducer.     

Ecological studies on the phytoplankton of Boknafjorden, western Norway. II. Environmental control of photosynthesis

Both predicted (incubator) and measured (in situ) ¹⁴C-assimilationrates were studied from February to November 1981 at three stationsin Boknafjorden, a deep silled fjord of western Norway. Sampleswere taken from different light depths within the euphotic zone.A high degree of conformity was found between the two approaches.Daily values of carbon assimilation integrated over the euphoticzone varied between 0.05 and 1.4 g C m^–2. Yearly primaryproduction varied between stations from 82 to 112 g C m^–2(120–148 g C m^–2 when based on average light conditions).The light-saturation curve parameters ^B and P^B_max ranged from0.0056 to 0.0537 mg C mg Chla^–1 h^–1 µE^–1m² and from 0.7 to 8.5 mg C mg Chla^–1 h^–2 (in situassimilation numbers ranged from 0.9 to 9.3 mg C mg Chla^–1h^–1) respectively, which compare well with those publishedfrom the northwestern side of the Atlantic. The overall importanceof light in controlling photosynthesis throughout the year wasrevealed by the light utilization index , estimated to be 0.43mg C mg Chla^–1 E^–1 m². The maximum quantum yieldwas encountered on August 17, with 0.089 mol CE^–1. Chla/Cratios above and below 0.010 were found to be typical for shade-and light-adapted cells respectively. Assimilation numbers andgrowth rates were linearly related only when considering light-adaptedcells. Consistent with the findings of this study, the applicabilityof I_K, ^B and P^B_max as indicators of light-shade adaptation propertiesshould be questioned. Maximum growth rates were encounteredduring an autumn bloom of the dinoflagellate Gyrodinium aureolum(1.0 doublings day^–1), while 0.7–0.8 doublings day^–1were found for a winter bloom (water temperature of 2°C)of the diatom Skeletonema costatum. No unambiguous temperatureeffect on assimilation number and growth of phytoplankton couldbe recognized in Boknafjorden. A tendency towards increasedassimilation numbers coinciding with increased water columnstability was revealed. The highest P^B_max values were oftenencountered at almost undetectable nutrient concentrations.At least during summer this could be attributed to recyclingof nutrients by macro- and/or microzooplankton, responsiblefor a greater part of the primary production now being grazeddown. This study supports the convention that the depth of theeuphotic zone may extend considerably below the 1% light depth.     

Modelling steady-state growth and photosynthesis rates of Oscillatoria rubescens continuous cultures in relation to temperature and irradiance

The lacustrine blue-green alga (= Cyanobacterium) Oscillatoriarubescens D.C. was cultured in chemostats with temperature andlight intensity as the only limiting factors. The experimentswere carried out under combinations of three temperatures 10,20 and 30°C and three irradiances 6, 18 and 30 µEm^–2 s^–1. Dilution rate, photosynthesis rate anda large number of abiotic and cellular factors were regularlymeasured to determine steady-state periods and the level ofthe associated variables. Various mathematical models were fittedwith series of data by a non-linear regression method derivedfrom Marquardt's method. Four models for calculation of specificgrowth rate and photosynthesis rate are presented. The lastone computes growth rate from calculated photosynthesis: µ= µ_max p_maxI/[K_pK_i + I(K_p + P_max)], with P_max =100 + 257Tand K₁ = 300/chl a. Values of adjusted parameters are discussed.One conclusion confirms that values of maximal growth ratesand half-saturation constants frequently considered in the literatureas absolute species characteristics should always be consideredas relative to associated prevalent growth conditions.     

Photosynthetic parameters, pigment composition and respiration rates of the marine diatom Skeletonema costatum grown in continuous light and a 12:12 h light-dark cycle

Nutrient-sufficient cultures of a Trondheimsfjord (Norway) cloneof the marine centric diatom Skeleionema costatum (Grev.) Clevewere grown at 75 µmol m^–2 s^–1 and 15C at24 and 12 h daylength to study diurnal variations and the effectof daylength on pigment and chemical composition, photosyntheticparameters, dark respiration rates and scaled fluorescence excitationspectra (F), the latter used as estimates for the absorptionof energy available to Photosystem II. Specific growth rateswere 1.06 and 0.56 day^– in 24 and 12 h daylength, respectively,while dark respiration rates were generally 85% of the net growthrate. The Chla-normalized photosynthetic coefficients P^B_m anda^B were {small tilde}20–25% higher in continuous lightthan at 12 h daylength, while the Chla:C ratio was {small tilde}15%lower (0.051 versus 0.061 w:w). Thus, the carbon-normalizedcoefficients P^c_m and a^c were <11% lower at 24 h than at 12h daylength. The maximum quantum yield _max, the Chla:C ratioand F differed negligibly, as did the light saturation indexl_k, the N:C ratio and the ratios Chlc:Chla and Fucoxanthin:Chla. P^B_m and l_k did not exhibit diurnal variations at 24 hdaylength, and varied within 23% of the daily mean at 12 h daylength.Predictions of the daily gross photosynthetic rate based ondata for a given time of the day should thus not be >10%in error relative to an integrated value based on several datasets collected through 24 h. _max was 0.084–0.117 mol O₂(mol photons)^– for gross oxygen evolution. However, ifused in mathematical models for predicting the gross and netgrowth rates (i.e. the gross and net carbon turnover rates),‘practical’ values of 0.076 and 0.040 g-at C (molphotons)^–, respectively, should be employed. Correspondingly,values for a^B and P^B_m should be adjusted pro rata. ¹Present address: College of Marine Studies, Sjmannsveien 27,N-6008 lesund, Norway     

Nitrogen inputs into the euphotic zone by vertically migrating Rhizosolenia mats

Rhizosolenia mats conduct extensive vertical migrations in theoligotrophic central North Pacific (cNP) gyre that permit thesediatoms to acquire nitrate at depth and return to the surfacefor photosynthesis. The ultimate fate of this N within the ecosystemis unknown, but may include remineralization by grazing, lossto depth by sinking biomass, or N excretion by Rhizosoleniamats. Direct release of N by mats into the mixed layer wouldrepresent an upward biological pump that circumvents the diffusionbarriers and nutrient sinks at the base of the oceanic euphoticzone. We examined Rhizosolenia mat N release along a transect(28–31° N) in the summer of 2002 (Hawaii to California)and 2003 (Hawaii to west of Midway Island) using sensitive fluorometricand chemiluminescence methods. Nitrate, NO₂^– and NH₄⁺release was determined. Nitrate and NH₄⁺ release by the matsoccurred in both 2002 (22.84 ± 6.04 and 3.69 ±1.74 nmol N µg^–1 Chl a h^–1, respectively)and 2003 (23.74 ± 3.54 and 3.60 ± 0.74 nmol Nµg^–1 Chl a h^–1, respectively). Nitrite releaseonly occurred in the 2003 summer period but occurred in bothyears when Fe chelators were added. Fv/Fm values decreased westwardin 2003 suggesting a gradient of increasing physiological stresstowards the west. The various physiological measures are consistentwith concurrent Fe stress; however, other possibilities exist.Nitrate excretion was the dominant form of N release in bothyears and provided a substantial addition to the ambient nitratepool in the mixed layer. Rhizosolenia mat nitrate release suppliesat least 4–7% of the nitrate pool on daily basis, andpossibly as much as 27%. Rhizosolenia mats are part of a largephytoplankton community that appears to migrate, and rates couldbe significantly higher. Literature reports suggest little orno nitrification in the upper euphotic zone, and thus biologicaltransport and release of nitrate may be a major source to thisregion. This N release is uncoupled from upward CO₂ transportand, like N₂ fixation, provides a component of the N pool availablefor net carbon removal.     

Effect of cell flotation on growth of Anabaena circinalis under diurnally stratified conditions

We tested the hypothesis that the growth rate of Anabaena circinalis,under diurnally stratified conditions, would increase as flotationvelocity increased owing to higher light availability. An insitu experiment compared the growth of diurnally stratifiedpopulations of A. circinalis with flotation velocities of 0.5and 1.0 m h^–1, with neutrally buoyant populations thatwere exposed to either mixed or persistently stratified conditions.The experiment was conducted in the turbid lower Murray Riverin South Australia (vertical attenuation coefficient = 4.52± 0.36 m^–1). To represent the mixing patterns,A. circinalis was contained in diffusion chambers that weremoved to different positions in the water column throughoutthe day. Diurnal populations with flotation velocities of 1.0and 0.5 m h^–1 grew at 0.23 ± 0.01 and 0.15 ±0.01 day^–1, respectively. Mixed populations grew at 0.19± 0.01 day^–1, whereas persistently stratified populationsgrew at 0.43 ± 0.01 day^–1. Results were used toextend a model that predicts growth of A. circinalis when exposedto the different mixing patterns. The model showed that bloomsare unlikely to be formed when the period of diurnal stratificationis <1 week, regardless of flotation velocity. When the diurnallystratified period is >1 week, flotation velocity is importantand a bloom may form depending on values assigned to the growthperiod and maximum mixed depth (Z_m).     

Physiological evaluation of temperature effect on the growth processes of the mysid, Neomysis intermedia Czerniawsky

Ingestion, respiration, and molting loss rates were measuredover the 3 – 29°C range in Neomysis intermedia. Weightspecific rates of these physiological processes ranged from2 to 140% body C day^–1 for ingestion, from 2 to 15% bodyC day^–1 for respiration, and from 0.1 to 5% body C day^–1for molting loss. All weight-specific rates showed a logarithmicdecrease with a logarithmic increase in body weight, and a logarithmicincrease with a linear increase in temperature below 20 or 25°C.The effect of temperature, however, was different between thephysiological rates, with a large temperature dependency foringestion (Q₁₀ = 2.6 –3.9) and molting loss (Q¹⁰ = 2.9– 3.6) and a moderate temperature dependency for respiration(Q₁₀ = 1.9 – 2.1). Calculated assimilation efficiencychanged with body size, but was constant over the temperaturerange examined. Allocation of assimilated materials varied witha change in temperature, reflecting the different temperaturedependence between physiological processes. It was deduced thatthe strong temperature dependency of the growth rate in N. intermediaobserved in the previous studies resulted from the large temperatureeffect on ingestion and assimilation rates, superimposed bythe different allocation of assimilated materials. ¹Present address: Department of Botany, University of Tokyo,Hongo, Tokyo 113, Japan