A mechanistic model for describing dynamic multi-nutrient, light, temperature interactions in phytoplankton

Developed by combining components simplified from previouslydeveloped mechanistic models, acomplete dynamic model is describedfor simulating the growth of phytoplankton as functions of ammonium,nitrate, light, iron, silicon, phosphorus and temperature. Componentsmay be safely added or deleted to the base model, describingammonium–nitrate–light interactions, to suit particularmodelling scenarios. Biomass is described in terms of C andcells, while chlorophyll is also a state variable enabling thesimulation of changes in Chl a:C with photoacclimation. Themodel is capable of simulating variable silicon deposition (diatoms)and C cell^–1 with Si, Fe, or P limitations. Mechanismsfor inclusion of temperature control of nutrient transport,growth rate and cell size are given. The model is suitable forplacement in ecosystem models, containing various componentsthat can be readily modified to tune the simulation to mimicthe behaviour of specific algal groups or species. Most of thosecomponents have biological significance and can be estimatedfrom experiments or by analysis of existing data.     

Circadian variability in the photobiology of Phaeodactylum tricornutum: pigment content

Circadian variations of pigment content in the diatom Phaeodactylumtricornutum were analyzed in different light regimes. The studywas aimed at discerning the role of putative endogenous controlsfrom the constraint imposed by the alternation of light (L)and dark (D) periods. Our experiments showed that in a typicalLD cycle of illumination, pigment synthesis follows the somaticgrowth of the cell, both arresting during D periods. In particular,the diurnal increase of chlorophyll a content was proportionalto the increase in cell size and preceding cell division, occurringat night. By contrast, diadinoxanthin and ß–carotene displayed different phases, which is likely to be relatedto their involvement in photoprotection mechanisms. The experimentsalso showed that the synthesis of both photosynthetic and photoprotectivepigments was dependent not only on light availability and thephasing of somatic growth, but also responded to other internalregulation. Over the time scale of the experiments (hours todays), the removal of LD–DL triggers impaired cell physiology,whereas the circadian patterns in pigment synthesis persisted.Our results support the hypothesis that an internal regulationof cell biosynthetic machinery can improve phytoplankton fitness,even in high variable environments such as the oceanic mixedlayers. Therefore, we suggest that phytoplankton growth dependsnot only on the availability of external resources, but alsoon internal regulatory mechanisms whose unveiling would furtherour understanding of phytoplankton diversity and dynamics.     

Uptake of 13C and 15N (ammonium, nitrate and urea) by Microcystis in Lake Kasumigaura

The uptake rate of carbon and nitrogen (ammonium, nitrate andurea) by the Microcystis predominating among phytoplankton wasinvestigated in the summer of 1984 in Takahamaira Bay of LakeKasumigaura. The V_max values of Microcystis for nitrate (0.025–0.046h^–1) and ammonium (0.15–0.17 h^–1) were considerablyhigher than other natural phytoplankton. The ammonium, nitrateand urea uptake by Microcystis was light dependent and was notinhibited with nigh light intensity. The K₁ values were farlower than the I_k values. The carbon uptake was not influencedby nitrogen enrichment. Microcystis accelerated the uptake rateby changing V_max/K _s value when nitrogen versus carbon contentin cells declined. Nitrate was scarcely existent in TakahamairiBay during the summer, when Microcystis usually used ammoniumas the nitrogen source. However, the standing stock of ammoniumin the water was far lower than the daily ammonium uptake rates.Therefore, the ammonium in this water had to be supplied becauseof its rapid turn-over time (–0.7–2.6 h).     

Annual Variation of Environmental Variables, Phytoplankton Species Composition and Photosynthetic Parameters in a Coastal Lagoon

Physical–chemical variables, phytoplankton biomass, speciescomposition and photosynthesis–irradiance (P-I) parameterswere analysed during 1 year in the Santo André Lagoon,SouthwestPortugal – a land-locked coastal ecosystem withtemporary connections with the sea. When the lagoon stayed closedthe observed phytoplankton blooms were mainly caused by Prorocentrumminimum, a potentially toxic dinoflagellate. It was dominantduring most of the year but the seawater inflow to the lagoontriggered a decrease in phytoplankton biomass and an abruptshift in species composition. The maximum photosynthetic rate(P_max) ranged from 2.0 to 22.5 mg C (mg chlorophyll a)^–1h^–1 and the light saturation index (I_k), ranged from 5.2to 335.0 µE m^–2 s^–1, with winter minima andsummer maxima. P_max and I_k were both positively correlated totemperature. Abundance ofP. minimum was associated with highnitrate concentrations whereas diatoms appear when ammonium,salinity and wind velocity are high. A mathematical model todescribe photosynthetic rate as a function of irradiance andtemperature [P (I, t)] was applied to the samples in which P.minimum was the dominant species     

The influence of the diel climatic cycle on the depth-time distribution of phytoplankton and photosynthesis in a shallow equatorial lake (Lake Baringo,Kenya)

Lake Baringo is a shallow equatorial lake. This paper reports a diel study of the depth-time distribution of phytoplankton and photosynthesis at one location in Lake Baringo on 10 March 1989. The water column shows a pattern of diurnal stratification probably accentuated by the high turbidity of the water and therefore rapid attenuation of solar energy. This stratified pattern breaks down at night due to atmospheric cooling and the regular onset of winds in the early evening. The phytoplankton is dominated byMicrocystis aeruginosa with some associated epiphytes. It concentrates in the narrow euphotic zone during the diurnal period of stratification due to buoyancy of theMicrocystis; evening breakdown of the thermocline results in the phytoplankton being mixed throughout the water column. A series of measurements of photosynthesis throughout the diurnal period gives an areal rate of 3.8 g O₂ m⁻² d⁻¹. The relationship between this value and the level of fish exploitation in Lake Baringo is discussed. The diel cycle in Lake Baringo is interpreted as dominating over any seasonal limnological cycle in the lake.     

Adding Detritus to a Nutrient-Phytoplankton-Zooplankton Model:A Dynamical-Systems Approach

The dynamics of two plankton population models are investigatedto examine sensitivities to model complexity and to parametervalues. The models simulate concentrations of nutrients, phytoplankton,zooplankton and detritus in the oceanic mixed layer. In Model1, zooplankton can graze only upon phytoplankton, whereas inModel 2 they can graze upon phytoplankton and detritus. Bothfeeding strategies are employed by zooplankton in the ocean,and both are features of models in the literature. Each modelhere consists of four coupled ordinary differential equations,and can exhibit unforced oscillations (limit cycles) of thefour concentrations. By constructing diagrams that show howsteady states and oscillations persist as each parameter isvaried, a general picture of the dynamics of each model is builtup. The addition of the detritus pool to an earlier nutrient–phytoplankton–zooplanktonmodel appears to have little influence on the dynamics whenthe zooplankton cannot graze upon the detritus (Model 1), butif the zooplankton can graze upon the detritus (Model 2), thenthe dynamics are affected in a significant way. These results,obtained using the theory of dynamical systems, enhance ourknowledge of the factors governing the dynamics of planktonpopulation models.     

Light rather than iron controls photosynthate production and allocation in Southern Ocean phytoplankton populations during austral autumn

The role of iron and light in controlling photosynthate productionand allocation in phytoplankton populations of the Atlanticsector of the Southern Ocean was investigated in April–May1999. The ¹⁴C incorporation into five biochemical pools (glucan,amino acids, proteins, lipids and polysaccharides) was measuredduring iron/light perturbation experiments. The diurnal Chla-specific rates of carbon incorporation into these pools didnot change in response to iron addition, yet were decreasedat 20 µmol photons m^–2 s^–1, an irradiancecomparable with the one at 20–45 m in situ depth. Thissuggests that the low phytoplankton biomass encountered (0.1–0.6µg Chl a L^–1) was mainly caused by light limitationin the deep wind mixed layer (>40 m). Regional differencesin Chl a-specific carbon incorporation rates were not foundin spite of differences in phytoplankton species composition:at the Antarctic Polar Front, biomass was dominated by a diatompopulation of Fragilariopsis kerguelensis, whereas smaller cells,including chrysophytes, were relatively more abundant in theAntarctic Circumpolar Current beyond the influence of frontalsystems. Because mixing was often in excess of 100 m in thelatter region, diatom cells may have been unable to fulfil theircharacteristically high Fe demand at low average light conditions,and thus became co-limited by both resources. Using a modelthat describes the ¹⁴C incorporation, the consistency was shownbetween the dynamics in the glucan pool in the field experimentsand in laboratory experiments with an Antarctic diatom, Chaetocerosbrevis. The glucan respiration rate was almost twice as highduring the dark phase as during the light phase, which is consistentwith the role of glucan as a reserve supplying energy and carbonskeletons for continued protein synthesis during the night.     

Photoinhibition and the diurnal variation of phytoplankton photosynthesis--I. Development of a photosynthesis--irradiance model from studies of in situ responses

Diurnal series of fluorescence and photosynthesis assays wereconducted in high altitude (3803 m), tropical (16°), LakeTiticaca (Peru/Bolivia). Near-surface diurnal thermoclines formedon typical days of high photon flux density (PFD, {small tilde}2000 µE m^–2 s^–1). In the depth range of diurnalstratification profiles of in vivo fluorescence, both without(F_a and with (F_b DCMU, exhibited a mean decrease of 64% frommorning to mid-day, but little change (mean increase of 1.5%)through the afternoon. Three times during the day surface, mid-depth(3–5 m) and deep (15–20 m) phytoplankton sampleswere incubated with H¹⁴CO₃^– under short (<2 h) exposuresto a range of in situ PFDs. Comparison of phytoplankton in differentsamples (ANOVA) showed identical photosynthetic response insunrise (isothermal) samples but a significant drop in surfaceand mid-depth photosynthesis at all PFDs during times of diurnalstratification. Similarly, both low-light () and light-saturated(P² _max photosynthetic parameters were lower in mid-day surfacesamples compared to deep samples. In addition, previously photoinhibitedsamples had a higher threshold intensity for photoinhibition,I_T. These results, together with diurnal time series of fluorescencefrom in situ incubations, demonstrate that recovery from extendedepisodes of photoinhibition during diurnal stratification isslower than suggested by previous observations in vitro. Photosynthesisby near-surface phytoplankton is different in light increasingup to I_T than light decreasing from I_T. This effect can be modeledby reducing and P_max as a function of the maximum photoinhibitingPFD in the diurnal light history. ¹Present address: Division of Molecular Plant Biology, Universityof California, Berkeley, Berkeley, CA 94720, USA     

Seasonal uptake and regeneration of inorganic nitrogen and phosphorus in a large oligotrophic lake: size-fractionation and antibiotic treatment

Uptake and regeneration of inorganic N and P in oligotrophicFlathead Lake (Montana) were measured with ¹⁵N and ³²P incorporationand dilution experiments, six times over a seasonal cycle. Theannual mean molar N P uptake ratio at ambient concentrationswas 13 9 (range = 4 8–34.2); uptake of nitrate, ammoniumand phosphate were always below saturation indicating both Nand P deficiency Organisms >280 µm were responsiblefor 0–60% of ammonium and 0–40% of phosphate regeneration,40–100% of the ammonium and 34–98% of phosphateregeneration occurred in the <3 µm fraction The <3µm fraction accounted for 7–70% of the ammoniumand 6–64% of the phosphate uptake. Results from antibiotictreatments indicated that both prokaryotic and eukaryotic ammoniumuptake was important, and that eukaryotes accounted for 53–98%of the ammonium regeneration. During thermal stratification,heterotrophic ammonium and phosphate regeneration by organisms>3 µm supplied much of the inorganic N and P in theepilimnion. Estimated rates of allochthonous and diffusive (i.e‘new’) ammonium, nitrate and phosphate input were<5% of biotic regeneration. These results suggests that (i)both N and P dynamics should be considered when examining nutrientregulation of primary productivity of oligotrophic lakes, (ii)bacteria probably compete with phytoplankton for both ammoniumand phosphate, (iii) biotic regeneration is the main sourceof nutrients to the epilimnion during stratification, and (iv)crustacean zooplankton were relatively unimportant sources ofregenerated ammonium and phosphate.     

Phytoplankton blooms and fish recruitment rate

The relationships between phytoplankton and zooplankton productionand fish larval survival to recruitment are examined by linkingtwo generic models. It is first demonstrated that the phytoplankton–zooplanktonmodels can be appropriately combined with a zooplankton–larvae–recruitmentmodel. The combined model reveals some general principles. Recruitmenttends to be a domed-shaped function of initial fish egg production.‘Bloom’ phytoplankton conditions are important forhigh recruitments. The timing and duration of fish egg productionis important in determining recruitment through their impacton the phytoplankton bloom. It is argued that optimal recruitmentwould be obtained if the duration of larval feeding was lessthan the duration of the phytoplankton bloom; a hypothesis whichis testable.     

Response of Sargasso Sea phytoplankton biomass, growth rates and primary production to seasonally varying physical forcing

The response of phytoplankton biomass, growth rates and primaryproduction to seasonally varying physical forcing was studiedat a station southeast of Bermuda over an 18 month period. Phytoplanktongrowth rates and primary production were measured using thepigment-labeling method, and phytoplankton biomass was calculatedfrom these measurements. Phytoplankton carbon biomass variedsystematically over the year. Highest values were observed duringthe winter and spring. Seasonal variations of chlorophyll (Chi)a in the surface layer could primarily be attributed to variationsin phytoplankton biomass and secondarily to photoacclimation.During the summer period, average values of carbon (C)/Chl ratios(g C g^–1 Chi) ranged from 160 at the surface to 33 atthe 1.6% light level, changes attributed to photoacclimationof the phytoplankton, consistent with the observation that phytoplanktonbiomass did not vary as a function of depth. Phytoplankton growthrates in the surface layer did not vary systematically overthe year, ranging from 0.15 to 0.45 day^–1, in spite ofseasonally varying concentrations of nitrate. Growth rates variedas a function of depth from average values of 0.3 day^–1in the surface layer to <0.1 day¹ at the 1.6% light level.Thus, the primary response of the phytoplankton community tonutrient enrichment during the winter period was an increasein phytoplankton biomass rather than an increase in growth rates.A simple nutrient-phyto-plankton-zooplankton model was usedto explore this phenomenon. The model demonstrated that theobserved response of the phytoplankton to nutrient enrichmentis only possible when phytoplankton growth is not severely limitedby nutrients.     

Diurnal and seasonal variation in the contributions of autotrophic pico-, nano- and microplankton to the primary production of an upland lake

An investigation of the diurnal variation in productivity andcontribution to production of populations of autotrophic picoplankton(0.2–2.0 µm), nanoplankton (>2 <20 µm)and microplankton (>20 µm) was carried out at monthlyintervals, from May to October 1989, in Llyn Padarn a mesotrophicupland lake in North Wales. Maximum rates and contributionsto production of the lake by autotrophic picoplankton occurredduring mid-late summer, with the highest average daily contributionfrom picoplankton (64%) recorded in September at 4 m depth.Diurnal variation in contributions from picoplankton was pronounced,with greatest input, recorded at the end of the day, duringthe period of picoplankton dominance in mid-late summer. Maximumcontribution from picoplantkon (86% of total, 9.2 mg C m^–3h^–1) was recorded in September. Nanoplankton primary productionwas of greatest significance in June and July, although levelswere lower than for picoplankton in subsequent months. Contributionsvia nanoplankton increased with depth in the lake at this time,reaching a maximum of 78% of the total at the end of the dayat 9 m depth in early July. At this time, diurnal variationin contributions via nanoplankton was considerable, with maximumphotosynthesis generally at the end of the photoperiod at depthsof 4 and 9 m. Microplankton made the greatest impact on primaryproduction during the mixed water conditions of spring and autumn,and at these times did variation in production was less thanthose of both pico and nanoplankton during summer thermal stratification.Photosynthetic capacity was lower for picoplankton than fornanoplankton and microplankton; the highest values were 5, 33and 51 mg C (mg chl a)^–1) h^–1) for pico-, nano-and microplankton, respectively. The photosynthetic efficiencyof all three size categories of phytoplankton increased withdepth. Maximum values were similar for all phytoplankton groups,between 75 and 131 mg C (mg chl a)^–1) E^–1) m² butmean levels of photosynthetic efficiency for the 6 months werelower for picoplankton than for nano- or microplankton. Ratesof carbon fixation per cell for picoplankton spanned three ordersof magnitude, varied considerably diurnally and reached maximumvalues of 484 fg C(cell)^–1) h^–1) in the afternoonin near-surface waters in the early stages of exponential populationgrowth in July. During the population maximum of picoplanktonin August and September, maximum daily values of carbon fixationper cell, assimilation number and photosynthetic efficiencywere all recorded at the end of the day. The seasonal and diurnalpatterns of production of the three size categories of planktonicalgae in Llyn Padarn were distinct. During spring, microplankton(mainly diatoms) were the dominant primary producers. As thermalstratification developed, nanoplankton were the major contributorsto phytoplanktonic production, particularly in the deeper regionsof the euphotic zone. Picoplankton made the greatest contributionto production in August and September, exhibiting maximum inputtowards the end of the light cycle. Diatoms became the majorphotosynthetic plankton in the mixed water conditions prevalentin Uyn Padarn in October.     

Integrate-and-fire models of insolation-driven entrainment of broadcast spawning in corals

The circa-annual cycle of gametogenesis produces mature gametes at the spawning “season” for successful mass spawning of broadcast corals. We develop a bioenergetic integrate-and-fire model that reveals how annual insolation rhythms can entrain the gametogenetic cycles in tropical hermatypic corals to the appropriate spawning season, since photosynthate is their primary source of energy. In the presence of short-term fluctuations in the energy input, a feedback regulatory mechanism is likely required to achieve coherence of spawning times to within one lunar cycle, in order for subsequent signals such as lunar and diurnal light cycles to unambiguously determine the “correct” night of spawning. The feedback mechanism can also provide robustness against population heterogeneity that may arise due to genetic and environmental effects. We solve the integrate-and-fire bioenergetic model numerically using the Fokker–Planck equation and use analytical tools such as rotation number to study entrainment.     

Phytoplankton community structure derived from HPLC analysis of pigments in the Faroe-Shetland Channel during summer 1999: the distribution of taxonomic groups in relation to physical/chemical conditions in the photic zone

A study of the phytoplankton community in the Faroe-ShetlandChannel was conducted in July 1999. Samples were collected atvarious depths in the photic zone along three transects (thenorthern entrance, the center and the southern entrance). Exceptfor a few easterly stations where nitrate and silicate werebelow 1 µM, all nutrients (phosphate, silicate, ammonium,nitrite and nitrate) were non-limiting for phytoplankton growth.HPLC pigment analysis revealed a pronounced (>50%) dominanceof Prymnesiophyceae at all stations. Their pigment ratio ofdiatoxanthin + diadinoxanthin/Chl a (DDX/Chl a) indicated thatthe phytoplankton community was controlled by light. Primaryproduction in the delayed spring bloom varied from 1.2 to 1.8g C m^–2 day^–1 along the northern transect. Alongthe other two transects, primary production ranged from 1.6to 3.8 g C m^–2 day^–1. Associated with the characteristicsindicating the establishment of a bloom, the relative contributionof diatoms and Prymnesiophyceae increased, whereas that of Prasinophyceae,Cryptophyceae, Chrysophyceae and Cyanobacteriaceae decreased.With respect to their vertical distribution, Cyanobacteriaceae,Chrysophyceae and Dinophyceae tended to have a higher abundance,relative to other taxonomic groups, in the surface layers. Therelative abundance of diatoms and Chlorophyceae increased withdepth. The DDX/Chl a ratio of the Prymnesiophyceae decreasedwith depth, indicating that vertical mixing in the upper 30m of the photic zone occurred less frequently than the timespan of physiological acclimation of cellular pigment composition.     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Bottom-up excitable models of phytoplankton blooms

A simple nutrient-phytoplankton model is used to explore the dynamics of phytoplankton blooms. The model exhibits excitable behaviour in the sense that a large scale outbreak can only be triggered when a critical nutrient threshold is exceeded. The model takes into account several features often neglected but whose combined effect proves very important: (i) rapid nutrient recycling associated with the microbial loop and patch formation; (ii) self-shading; and (iii) a bottom-up approach, whereby nutrient levels are responsible for both the triggering and the demise of the bloom. Although the literature is replete with studies of ‘top-down’ models in which zooplankton grazing control the triggering and demise of the bloom, bottom-up models are nevertheless appropriate in many circumstances. We provide a full mathematical investigation of the effects of these three different features in an excitable system framework.     

Diurnal variations in the contributions of autotrophic picoalgae and heterotrophic bacteria to planktonic production in an upland lake

An investigation of the diurnal variation in contributions toproduction of the autotrophic and heterotrophic components ofthe picoplankton community was carried out during August andSeptember in Llyn Padarn, a mesotrophic upland lake in NorthWales. The picoplankton was separated using 1 µm pore-sizedfilters into the smaller cell sized fraction (<1 µm),the majority of the bacteria and the larger cell sized picoalgae(<3>1 µm), together with some bacteria. The distributionof bacterial heterotrophic activity between these two fractionsof picoplankton was assessed by uptake of [¹⁴C]glucose and differentialfiltration. Thus, the absolute autotrophic production by picoalgaeand the heterotrophic contribution by bacteria to picoplanktoncommunity production via uptake of extracellular organic carbon(EOC) were determined. Rates of picoplankton community productionexhibited diurnal variation with maximum rates of 19.1 mg Cm^–3 h^–1 recorded at 18.00 h at 4 m depth in September.The bacterial contribution to picoplankton community productionincreased markedly between 15.00 and 18.00 h. Rates of absoluteautotrophic production varied less over 24 h than rates of accumulationin bacteria of ¹⁴C-labelled EOC released from the entire phytoplanktoncommunity. Bacteria contributed up to 86–98% of the neworganic carbon within the picoplankton community at the endof the day. The maximum rate of absolute autotrophic productionin the picoplankton was 1.6 mg C m^–3 h^–1 at 18.00h at 1 m in August, and the maximum rate of bacterial accumulationof new organic carbon was 18.5 mg C m^–3 h^–1 at 18.00h in September at 4 m depth. The diurnal pattern of picoplanktoncommunity production involved increasing rates during the daywith a maximum at 18.00 h. Autotrophic processes were dominantin the first 3–6 h of the light cycle and heterotrophicuptake of ¹⁴C-labelled EOC was the major component from 15.00h onwards. Bacterial uptake of newly released EOC by phytoplanktonwas rapid, comprised the majority of picoplankton production,particularly later in the day, and contributed a maximum of60% of the total pariculate primary production in plankton between15.00 and 18.00 h at 4 m in September with a mean contributionof between 6 and 26% over 24 h in these investigations. Theimportance of autotrophic processes in picoplankton communityproduction has been overestimated in previous investigations.Bacteria play a major role in transferring newly produced EOCrapidly from phytoplankton to the picoplankton community. Atthe end of the day, the majority of newly produced organic carbonis in bacterial cells and this production is significant inthe dynamics of carbon production within the entire planktoniccommunity.     

Ammonium Uptake by Phytoplankton Regulates Nitrification in the Sunlit Ocean

Nitrification, the microbial oxidation of ammonium to nitrate, is a central part of the nitrogen cycle. In the ocean’s surface layer, the process alters the distribution of inorganic nitrogen species available to phytoplankton and produces nitrous oxide. A widely held idea among oceanographers is that nitrification is inhibited by light in the ocean. However, recent evidence that the primary organisms involved in nitrification, the ammonia-oxidizing archaea (AOA), are present and active throughout the surface ocean has challenged this idea. Here we show, through field experiments coupling molecular genetic and biogeochemical approaches, that competition for ammonium with phytoplankton is the strongest regulator of nitrification in the photic zone. During multiday experiments at high irradiance a single ecotype of AOA remained active in the presence of rapidly growing phytoplankton. Over the course of this three day experiment, variability in the intensity of competition with phytoplankton caused nitrification rates to decline from those typical of the lower photic zone (60 nmol L⁻¹ d⁻¹) to those in well-lit layers (<1 nmol L⁻¹ d⁻¹). During another set of experiments, nitrification rates exhibited a diel periodicity throughout much of the photic zone, with the highest rates occurring at night when competition with phytoplankton is lowest. Together, the results of our experiments indicate that nitrification rates in the photic zone are more strongly regulated by competition with phytoplankton for ammonium than they are by light itself. This finding advances our ability to model the impact of nitrification on estimates of new primary production, and emphasizes the need to more strongly consider the effects of organismal interactions on nutrient standing stocks and biogeochemical cycling in the surface of the ocean.