CHEMICAL COMPOSITION AND PHOTOSYNTHETIC CHARACTERISTICS OF ETHMODISCUS REX (BACILLARIOPHYCEAE): EVIDENCE FOR VERTICAL MIGRATION1

Ethmodiscus rex (Rattray) Wiseman and Hendey cells from near surface net tows in the Southwest Atlantic Ocean and Caribbean Sea were examined for chemical composition, internal nutrient pool concentrations, and oxygen evolution characteristics. Elemental ratios indicated nitrogen limitation with C:N:P ratios of 125:9:1 (atoms), and carbon: chlorophyll (chl) ratios of 129:1 (weight). However, internal nitrate pools (1.4–27.1 mM) suggested that cells were not N-limited. Intracellular NO₃^? accounted for up to 54% (range = 3–54%) of the total N quota in some samples. Photosynthetic parameters were consistent with a high-light-adapted population and suggested an instantaneous maximum chl-specific photosynthetic rate (P^B_max) of 4.8–12.4 nmol O₂·μg chl^?1· h^?1. Respiration rates varied ten-fold and were inversely related to P^B_max Ethmodiscus chemical composition and buoyancy characteristics are similar to vertically migrating Rhizosolenia mats and the non-motile dinoflagellate Pyrocystis noctiluca Murray (Schuett). The presence of internal NO₃^? pools in Ethmodiscus suggests that this genus is also vertically migrating to exploit sub-surface nitrogen pools. Such behavior may be widespread in large, non-motile oceanic phytoplankton. Based on ascent rate data, chemical composition, and photosynthetic rates, we estimate that the entire division–migration cycle for Ethmodiscus requires at least 7–12 days.     

VERTICAL MIGRATION BY RHIZOSOLENIA SPP. (BACILLARIOPHYCEAE): IMPLICATIONS FOR FE ACQUISITION

Available data support a mechanism of buoyancy-mediated vertical migration by large-sized diatoms of Rhizosolenia spp. as a means to access "new" nitrogen from deep waters. To assess whether phytoplankton simultaneously satisfy their Fe requirements by this mechanism, field samples collected during summer 1996 at stations located along a transect through the central North Pacific gyre were assayed for the presence of flavodoxin and ferredoxin via Western blot analysis. All samples, regardless of their buoyancy status and the station from which they were collected, had accumulated flavodoxin but not ferredoxin. To understand better the significance of the field results, cultures of Rhizosolenia formosa H. Peragallo were grown in the laboratory with varying levels of total Fe (200 nM–10,000 nM). Fe had little effect on the physiological and photochemical parameters measured for each treatment. Growth rates did not exceed 0.17 d ^{− 1} and values of F_v/F_m ranged from 0.48 to 0.62. In addition, R. formosa accumulated only flavodoxin at each level of Fe addition. From these results, it appears that for some rhizosolenids, flavodoxin is constitutively expressed. The underlying basis for the constitutive nature of this flavodoxin is unclear at present, although it is likely that it is ultimately related to chronic Fe deficit incurred in natural waters.     

TEMPERATURE-DEPENDENT CHANGES IN COLONY SIZE OF THE FRESHWATER PENNATE DIATOM ASTERIONELLA FORMOSA (BACILLARIOPHYCEAE) AND THEIR POSSIBLE ECOLOGICAL IMPLICATIONS1

Changes in colony size (cell number per colony) of Asterionella Formosa Hass. were experimentally evaluated in relation to water temperature using two types of clones having colony sizes of four or eight cells. The clones were isolated from two different temperate freshwater lakes. Both clones showed the same general trend with changing temperature. Most of the colonies were normal in size at low temperatures, but colony size was twice as large at high temperatures. Variable colony sizes were present at low percentages. Colony separation occurred at the oldest connection within the colony after cell division. Culture experiments showed that the rates of specific growth and colony separation were balanced except for a rather short period of time when the temperature was changed. Optical and scanning electron micrography did not show any distinctive morphological structure at the point of connection except for porelli and mucilage pads. Seasonal changes in colony size of A. formosa observed in a freshwater lake are discussed based on these temperature results.     

EFFECT OF LIGHT ON VERTICAL MIGRATION AND PHOTOSYNTHESIS OF EUGLENA PROXIMA (EUGLENOPHYTA)

During daytime low tides, sediment-inhabiting populations of Euglena proxima Dangeard migrate to the surface of intertidal sand flats to form large monospecific green patches near the Duke University Marine Laboratory in Beaufort, North Carolina. The effect of incident irradiance on the upward migration of E. proxima was studied by shading the sediment surface in the field with neutral density filters to 2, 20, 55, and 100% incident irradiance. The results of these in situ experiments revealed that E. proxima 's upward migration, as measured by population density or chl a concentration, was stimulated by decreasing incident irradiance: population densities and chl a concentrations on the sediment surface at 2% incident irradiance were greater statistically than at 100% full sunlight. In situ measurements of photosynthesis were performed by incubating 3-mL aliquots of a cell suspension in scintillation vials containing ¹⁴C inorganic carbon beneath neutral density filters. These experiments revealed that E. proxima exhibited photoinhibition at 100% full sunlight, but this depression in photosynthetic performance at high light was absent when the cells were incubated in vials containing acid-cleaned sediment. This amelioration of photoinhibition in vials containing sediment is most likely due to the downward migration of the population away from photoinhibitory light levels. These results cannot be interpreted as strictly photic effects because the temperature of the vials varied with the light treatment. The results of this study suggest that the vertical migration behavior of E. proxima and its photosynthetic physiology are intimately linked. Vertical migration provides a mechanism to deal with short-term changes in irradiance and avoid photoinhibitory light levels.     

PHOTOINHIBITION OF PHOTOSYNTHESIS IN ASTERIONELLA FORMOSA (BACILLARIOPHYCEAE)1, 2

A laboratory study was made of the inhibition of photosynthesis, as measured by the radiocarbon method, of Asterionella Formosa Hass. by light from tungsten halogen lamps of irradiances up to that of full sunlight. The observed inhibitions were of total fixation of carbon and were not due to greatly increased release of extracellular products of photosynthesis. Inhibition increased width irradiance and with time of exposure. It was greater at high temperature, at high oxygen concentrations and when the cells were nutrient deficient. Recovery from exposure to high irradiances took place both in the dark and at low irradiances. The inhibition, which is similar to that observed under natural conditions, has the characteristics of photooxidative damage to both photochemical and dark reaction mechanisms.     

INTERNAL NITRATE CONCENTRATIONS IN SINGLE CELLS OF LARGE PHYTOPLANKTON FROM THE SARGASSO SEA1

Nitrate concentrations within individual cells of Ethmodiscus, Pyrocystis, and Halosphaera and chains of Rhizosolenia were determined from samples collected in the Sargasso Sea. In all cases, field populations exhibited a wide range of internal nitrate concentrations (INCs) within a single sampling date. Halosphaera INCs reached 100 mM, in contrast to diatom and dinoflagellate INCs, which did not exceed 22 mM. Sinking Rhizosolenia, Ethmodiscus and Pyrocystis had significantly lower internal NO₃- pools than did floating cells (P< 0.05). Ethmodiscus incubations in surface seawater resulted in a dramatic reduction in the proportion of high INC cells concurrent with decreases in average INCs and an increased proportion of sinking cells. Population buoyancy was inversely related to INC, and negatively buoyant cells rarely exceeded I mM INC, suggesting that a critical INC threshold may exist. The photosynthetic parameters P_max and α decreased with time as internal NO₃-_- Pools were depleted. Internal nitrate depletion rates were consistent with oxygen production rates during this time. Based on the known characteristics of Pyrocystis and Ethmodiscus, we conclude that virtually all of the > 100 μm-sized phytoplankton present in the Sargasso Sea can vertically migrate. However, the appropriate time scale for migrators such as Halosphaera that reproduce by swarmer formation is unclear and may be significantly different than the other taxa studied. Changes in the frequency distributions, buoyancy-internal pool relationships, and general P-1 photosynthesis-irradiance time series data in Ethmodiscus suggest that nutrient limitation is related to these migrations. High INC appears to be a fundamental property of the largest microalgal cells present in oligotrophic seas and suggests that nitrate transport by these nonmotile cells is widespread.     

THE RELATIVE PHOTOSYNTHETIC EFFICIENCY OF ASTERIONELLA FORMOSA HASS. (BACILLARIOPHYTA) IN NATURAL PLANKTON ASSEMBLAGES1

The relative photosynthetic efficiencies of net vs. nanoplankton and diatom vs. non-diatom plankton were estimated weekly in Woodcock Creek Reservoir, Pennsylvania, from April to November 1979, using carbon-14 in conjunction with size fractionation and germanium as a diatom inhibitor. The nanoplankton contributed a proportionally greater share of the community carbon assimilation than their share of the community biomass would indicate. Diatoms, specifically Asterionella formosa Hass. were photosynthetically inefficient and contributed significantly less than expected. The mechanism of periodic diatom dominance in lakes remains unexplained.     

RELATIONSHIPS BETWEEN NITRATE REDUCTASE ACTIVITY AND RATES OF GROWTH AND NITRATE INCORPORATION UNDER STEADY-STATE LIGHT OR NITRATE LIMITATION IN THE MARINE DIATOM THALASSIOSIRA PSEUDONANA (BACILLARIOPHYCEAE)1

Although activity of the enzyme nitrate reductase (NR) can potentially be used to predict the rate of nitrate incorporation in field assemblages of marine phytoplankton, application of this index has met with little success because the relationship between the two rates is not well established under steady-state conditions. To provide a basis for using NR activity measurements, the relationships among NR activity, growth rate, cell composition, and nitrate incorporation rate were examined in cultures of Thalassiosira pseudonana (Hustedt)Hasle and Heimdal, growing a) under steady-state light limitation, b) during transitions between low and high irradiance (15 or 90 μmol quanta.m^?2.s^?1), and c) under steady-state nitrate limitation. Using a modified assay for NR involving additions of bovine serum albumin to stabilize enzyme activity, NR activity in light-limited cultures was positively and quantitatively related to calculated rates of nitrate incorporation, even in cultures that were apparently starved of selenium. During transitions in irradiance, growth rates acclimated to new conditions within 1 day; through the transition, the relationship between NR activity and nitrate incorporation rate remained quantitative. In nitrate-limited chemostat cultures, NR activity was positively correlated with growth rate and with nitrate incorporation rates, but the relationship was not quantitative. NR activity exceeded nitrate incorporation rates at lower growth rates (<25% of nutrient-replete growth rates), but chemostats operating at such low dilution rates may not represent ecologically relevant conditions for marine diatoms. The strong relationship between NR activity and nitrate incorporation provides support for the idea that NR is rate-limiting for nitrate incorporation or is closely coupled to the rate-limiting step. In an effort to determine a suitable variable for scaling NR activity, relationships between different cell components and growth rate were examined. These relationships differed depending on the limiting factor. For example, under light limitation, cell volume and cell carbon content increased significantly with increased growth rate, while under nitrate limitation cell volume and carbon content decreased as growth rates increased. Despite the differences found between cell composition and growth rate under light and nitrate limitation, the relationships between NR activity scaled to different compositional variables and growth rate did not differ between the limitations. In field situations where cell numbers are not easily determined, scaling NR activity to particulate nitrogen content may be the best alternative. These results establish a strong basis for pursuing NR activity measurements as indices of nitrate incorporation in the field.     

RELATIONSHIP BETWEEN NUCLEOSIDE DIPHOSPHATE KINASE ACTIVITY AND LIGHT-LIMITED GROWTH RATE IN THE MARINE DIATOM THALASSIOSIRA PSEUDONANA (BACILLARIOPHYCEAE)1

Light-limited cultures of the marine diatom Thalassiosira pseudonana (Hustedt) Hasle and Heimdal (3H clone) were grown over a range of growth rates between 0.06 and 1.64 d^?1. Variations in cell volume, cell quotas of carbon, nitrogen, and protein, and maximal activity of the enzyme nucleoside diphosphate kinase (NDPK) were measured and examined as a function of growth rate. NDPK from T. pseudonana showed K_m values of 0.24 and 0.68 mM for thymidine 5′-diphosphate and adenosine 5′-triphosphate (ATP), respectively, which are similar to those found for NDPK from a variety of organisms, from bacteria to mammals. An apparent activation enthalpy of 3.52 kCal·mol^?1 was determined from Arrhenius plots. No thermodynamic transition points were noted over a temperature range from 10° to 25°C. NDPK activity was significantly correlated with growth rate but not with cell volume, carbon, nitrogen, or protein; for interspecific comparisons, normalization of enzyme activity to cell number may be most meaningful. NDPK activity per cell versus growth rate followed a U-shaped relationship, being relatively constant between 0.5 and 1.0 d^?1 and rising at higher and lower growth rates. Over this range, enzyme activity may be regulated by substrate concentration (ATP or other nucleoside triphosphates) or by adenylate energy charge. At higher growth rates where energy charge and substrate concentrations are probably high, changes in enzyme concentration appear to be required. The reasons for a rise in enzyme activity at low growth rate is unclear. Simultaneous measurement of nucleoside di- and triphosphate levels alongside NDPK measurements may help clarify the relationship, but these preliminary experiments indicate that NDPK is of limited usefulness as an index of in situ growth rate.     

THE RESPONSE OF GROWTH AND BIOCHEMICAL COMPOSITION TO VARIATIONS IN DAYLENGTH, TEMPERATURE, AND IRRADIANCE IN THE MARINE DIATOM THALASSIOSIRA PSEUDONANA (BACILLARIOPHYCEAE)

The relationships between the growth rate of the marine diatom Thalassiosira pseudonana (Hustedt) and irradiance, daylength, and temperature were determined in nutrient-sufficient semicontinuous cultures. The initial slopes of the growth versus total daily irradiance curves were not affected by temperature or daylength. Growth versus irradiance was best modeled as a hyperbolic function at short daylengths and better modeled as an exponential function at longer daylengths. The maximum or light-saturated growth rates at each daylength were modeled as a hyperbolic function of daylength. This model was extended in a novel manner to include temperature dependence providing a framework that can be used to interpret other experimental data on growth rate versus daylength. The resulting model should be useful in global models of phytoplankton growth. Carbon, nitrogen, and chl a quotas were influenced by daylength, irradiance, and temperature. Both C and N quotas were positive exponential functions of irradiance, whereas N and chl a quotas were significantly greater for cells grown at the lower temperature. The ratio chl a :C quota (chl a :Q_c) was a strong negative exponential function of total daily irradiance. Cells grown at 10° C had significantly greater chl a :Q_c ratios than those grown at 18° C, and daylength also had a significant positive influence on chl a :Q_c. The apparent effect of daylength on chl a :Q_c was removed by standardizing chl a :Q_c to growth rate (μ), resulting in a temperature-dependent relationship between chl a :Q_c·μ⁻¹ and irradiance that accounted for 95% of the variation in the data.     

THE PHYSIOLOGICAL ECOLOGY OF A FRESHWATER DINOFLAGELLATE BLOOM POPULATION: VERTICAL MIGRATION,NITROGEN LIMITATION,AND NUTRIENT UPTAKE KINETICS1,2

The motile freshwater dinoflagellate Gymnodinium bogoriense Klebs., which forms dense blooms in Jezre'el Valley water reservoirs (Israel) appears to be physiologically suited to exploit stratified environments, where it outcompetes all other phytoplankton types. The dense summer blooms (“red tides”) were found to be nitrogen-limited. The algae's competitive advantage, however, cannot result from superior uptake capabilities: its K_s (μmol NH₄·L^?1) for NH₄ was higher and its V_maxμmol NH₄·mg chlorophyll a^?1·h^?1) was lower than other phytoplankton types commonly occurring in the region. The competitive advantage of G. bogoriense probably stems from other physiological capabilities: dark ammonia and phosphorus assimilation and the ability to undertake diel vertical migration cycles between the upper photic water layers during the day and nutrient-rich deeper layers at night. These findings confirm the vertical nutrient retrieval hypothesis in migrating phytoplankton.     

EFFECT OF SUBSURFACE NUTRIENT SUPPLIES ON THE VERTICAL MIGRATION OF EUGLENA PROXIMA (EUGLENOPHYTA)1

Euglena proxima Dangeard inhabits intertidal sand flats and displays a tidal rhythm in vertical migration. During daytime low tides when the sand flat is emersed, millions of cells are visible on the sediment surface, but the population remains below the surface at all other times. An earlier study demonstrated that the extent of downward migration of E. proxima is reinforced by the presence of a subsurface layer of black sediment. The present study was designed to test the hypothesis that the higher availability of inorganic nutrients or organic substrates in or above the black layer is responsible for the enhancement of downward migration in E. proxima. This hypothesis was tested experimentally by manipulating the bottom water in 24 mesocosm containers in a tidal tank. Six replicates of each of the following nutrient treatments were tested: seawater control; deep porewater collected from 70 cm below the sediment surface; seawater enriched with ammonium, nitrate, and phosphate; and seawater enriched with acetate, glucose, and the preceding inorganic nutrients. Multivariate analysis of variance revealed that the chl a biomass and chl a‐weighted mean depth of the population at high tide were significantly greater for replicates receiving inorganic nutrients. There was no difference between those receiving only inorganic nutrients and those enriched with inorganic nutrients, acetate, and glucose. These findings represent the first experimental evidence that subsurface nutrients are an important resource that reinforces the maintenance of vertical migration behavior in benthic microalgae.     

MODULATION OF THE HEAT SHOCK UBIQUITIN POOL IN SKELETONEMA COSTATUM (BACILLARIOPHYCEAE)1

Immunoblotting experiments performed with an anti-ubiquitin antibody revealed that Skeletonema costatum (Grev.) Cleve cells contained free ubiquitin as well as ubiquitin conjugated to various endogenous proteins. A temperature shift from 18° to 30°C greatly increased the total amount of ubiquitin and particularly the ubiquitin fraction in high molecular mass conjugates. A solid-phase immunoassay indicated values of 0.031 ± 0.004 pmol·10^?6 cells for free ubiquitin and 0.046 ± 0.004 pmol·10^?6 cells for conjugated ubiquitin for cells grown at 18°C, and 0.056 ± 0.008pmol·10^?6cells and 0.21 ± 0.03 pmol·10^?6cells, respectively, after a temperature increase from 18° to 30°C. Cell-free extracts of S. costatum were equally able to form thiol ester linkages with ¹²⁵I-ubiquitin in an adenosine triphosphate–dependent manner at 18° C and at 30°C. Cell-free extracts were also able to conjugate ¹²⁵I-ubiquitin to endogenous proteins, but the ubiquitin conjugation rate at 30°C was lower than at 18°C. Incubation of S. costatum for 3 h at 30°C and then for 3 h at 18°C resulted in the formation of high amounts of ubiquitin conjugates, suggesting that partially inactive or denaturated proteins accumulate during heat stress. These denaturated proteins are then conjugated to ubiquitin very efficiently when the physiological temperature is restored. Thus, S. costatum cells contain ubiquitin and an active ubiquitin conjugation system responding to stress conditions (temperature stress). The intracellular concentration of ubiquitin conjugates is most likely limited by the availability of protein substrates to be conjugated rather than by ubiquitin-conjugating activity.     

DIEL VARIATIONS IN OPTICAL PROPERTIES OF IMANTONIA ROTUNDA (HAPTOPHYCEAE) AND THALASSIOSIRA PSEUDONANA (BACILLARIOPHYCEAE) EXPOSED TO DIFFERENT IRRADIANCE LEVELS1

Diel variations of cellular optical properties were examined for cultures of the haptophyte Imantonia rotunda N. Reynolds and the diatom Thalassiosira pseudonana (Hust.) Hasle et Heimdal grown under a 14:10 light:dark (L:D) cycle and transferred from 100 μmol photons · m^?2 · s^?1 to higher irradiances of 250 and 500 μmol photons · m^?2 · s^?1. Cell volume and abundance, phytoplankton absorption coefficients, flow‐cytometric light scattering and chl fluorescence, and pigment composition were measured every 2 h over a 24 h period. Results showed that cell division was more synchronous for I. rotunda than for T. pseudonana. Several variables exhibited diel variability with an amplitude >100%, notably mean cell volume for the haptophyte and photoprotective carotenoids for both species, while optical properties such as flow‐cytometric scattering and chl a–specific phytoplankton absorption generally showed <50% diel variability. Increased irradiance induced changes in pigments (both species) and mean cell volume (for the diatom) and amplified diel variability for most variables. This increase in amplitude is larger for pigments (factor of 2 or more, notably for cellular photoprotective carotenoid content in I. rotunda and for photosynthetic pigments in T. pseudonana) than for optical properties (a factor of 1.5 for chl a–specific absorption, at 440 nm, in I. rotunda and a factor of 2 for the absorption cross‐section and the chl a–specific scattering in T. pseudonana). Consequently, diel changes in optical properties and pigmentation associated with the L:D cycle and amplified by concurrent changes in irradiance likely contribute significantly to the variability in optical properties observed in biooptical field studies.     

PHYSICAL AND CHEMICAL CONDITIONS SURROUNDING THE DIURNAL VERTICAL MIGRATION OF CRYPTOMONAS SPP. (CRYPTOPHYCEAE) IN A SEASONALLY STRATIFIED MIDWESTERN RESERVIOR (USA)

Cross Reservoir, a small mesotrophic reservoir located at the University of Kansas Ecological Reserves (Kansas, USA), contained a dense metalimnetic community of algae and photosynthetic bacteria between early July and late October 1997–1999. Within this community, various Cryptomonas species, primarily C. erosa (Ehrenberg), C. erosa var. reflexa (Marsson), and C. rostratisformis (Skuja), diurnally migrated as indicated by in situ fluorescence monitoring and direct phytoplankton enumeration. The Cryptomonas spp. typically resided near the oxic–anoxic boundary of the water column; however, they actively migrated upward during the day and descended to lower anoxic locations at night, apparently responding to diurnal changes in their local habitat. Their nocturnal environment had moderate levels of sulfide, elevated secondary nutrients, and a community of anoxygenic phototrophic bacteria, whereas their daytime environment had higher light, lower nutrients, and no local photosynthetic bacteria. Monitoring indicated that the Cryptomonas spp. migration was generally linked to daily variations in absolute light intensity (e.g. sunny vs. cloudy days) and the level of other potentially growth-limiting resources, particularly nitrogen and phosphorus. However, further analyses showed that the primary factor that determined whether the Cryptomonas spp. migrated or not on a given day was the slope of the light gradient immediately above the Cryptomonas spp. peak.     

MORPHOLOGICAL AND PHYSIOLOGICAL EFFECTS IN PROBOSCIA ALATA (BACILLARIOPHYCEAE) GROWN UNDER DIFFERENT LIGHT AND CO2 CONDITIONS OF THE MODERN SOUTHERN OCEAN1

The combined effects of different light and aqueous CO₂ conditions were assessed for the Southern Ocean diatom Proboscia alata (Brightwell) Sundström in laboratory experiments. Selected culture conditions (light and CO_2(aq)) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) μmol photons · m^?2 · s^?1. The three CO_2(aq) conditions ranged from 8 to 34 μmol · kg^?1 CO_2(aq) (equivalent to a pCO₂ from 137 to 598 μatm, respectively). Clear morphological changes were induced by these different CO_2(aq) conditions. Cells in low [CO_2(aq)] formed spirals, while many cells in high [CO_2(aq)] disintegrated. Cell size and volume were significantly affected by the different CO_2(aq) concentrations. Increasing CO_2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO_2(aq) treatments. This included growth rates, chlorophyll a concentration and photosynthetic yield (F_V/F_M). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO₂ conditions co‐determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO_2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.     

LIGHT DEPENDENCE OF GROWTH AND PHOTOSYNTHESIS IN PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

Phaeodactylum tricornutum Bohlin was maintained in exponential growth over a range of photon flux densities (PFD) from 7 to 230 μmol·m^?2s^?1. The chlorophyll a-specific light absorption coefficient, maximum quantum yield of photosynthesis, and C:N atom ratio were all independent of the PFD to which cells were acclimated. Carbon- and cell-specific, light-satuated, gross photosynthesis rates and dark respiration rates were largely independent of acclimation PFD. Decreases in the chlorophyll a-specific, gross photosynthesis rate and the carbon: chlorophyll ratio and increases of cell- or carbon-specific absorption coefficients were associated with an increase in cell chlorophyll a in cultures acclimated to low PFDs. The compensation PFD for growth was calculated to be 0.5 μmol·m^?2s^?1. The maintenance metabolic rate (2 × 10^?7s^?1), calculated on the basis of the compensation PFD, is an order of magnitude lower than the measured dark respiration rate(2.7 × 10^?6mol O₂·mol C^?1s^?1). Maintenance of high carbon-specific, light-saturated photosynthesis rates in cells acclimated to low PFDs may allow effective use of short exposures to high PFDs in a temporally variable light environment.     

BIOLOGICAL AND CHEMICAL CHARACTERISTICS OF THE GIANT DIATOM ETHMODISCUS (BACILLARIOPHYCEAE) IN THE CENTRAL NORTH PACIFIC GYRE

Cells of the giant diatom Ethmodiscus Castr. gathered from the upper 15 m were examined for O₂ evolution, nitrate reductase activity (NRA), C and N composition, internal NO concentrations, , and ¹⁵NO, ¹⁵NH , and ³²Si uptake in a series of cruises in the central N. Pacific gyre. The δ¹⁵N (2.56–5.09 ‰), internal NO concentrations (0.0– 11.5 mM NO⁻), and NRA (6.7 ± 4.7 × 10⁻⁴μM NO cell ⁻¹·h⁻¹) were consistent with recent exposure to elevated nitrate concentrations and utilization of deep NO as a primary N source. These results are similar to other diatoms that migrate vertically to the nutricline as part of their life cycle. Rate measures (Si[OH]₄ uptake, NRA, and O₂ evolution) indicated surface doubling times from 45 h to 75 h. Both NO and NH uptake in surface waters were low and inadequate to supply N needs at surface NO and NH concentrations. Our results suggest a partitioning in nutrient acquisition, with N acquired at depth and C and Si acquired at the surface. Doubling rates were two to three times higher than predicted from cell volume and C content models. These data are consistent with the observed elemental content being lower than expected because of the dominance of cell volume by the vacuole. Our calculations suggest that Ethmodiscus contributes little to the biogeochemistry of the upper water column via upward nutrient transport. Although reported as a paleo-upwelling indicator, thisevidence suggests that Ethmodiscus has adapted to the nutrient-poor open ocean by a vertical migration strategy and has biological characteristics inconsistent with a upwelling indicator.     

EFFECT OF TEMPERATURE ON LIGHT-LIMITED GROWTH AND CHEMICAL COMPOSITION OF SKELETONEMA COSTATUM (BACILLARIOPHYCEAE)1

Phytoplankton growth rate in response to irradiance can be approximated by a hyperbola defined by three coefficients: i) initial slope (α); ii) asymptote (μ_m); and, iii) X-axis intercept or compensation irradiance (I_c). To mathematically represent the interaction of temperature and irradiance on growth rate, one must describe the relationship between these constants and temperature. The marine diatom, Skeletonema costatum (Greville) Cleve, was grown in unialgal culture at different levels of irradiance and 2-3 photoperiods at 0, 5, 10, 16 and 22 C. The value of I_c is ca. 1.0 ly·day^?1 or less at all temperatures. The initial slope (div·ly^?1) is a “u-shaped” function of temperature described by the second degree polynomial, α= 0.25–0.02T+0.001T². Within the range 0–10 C, μ_m (div·day^?1) is an exponential function of temperature described by the equation, μ_m= 0.48 exp (0.126T). At each temperature and selected levels of irradiance, cell size and cellular content of C, N and chl a were determined. The C:chl a and N:chl a ratios increased with irradiance because of increases in C and decreases in chl a. At lower temperatures (0, 5, 10 C), the rate of increase of both ratios with irradiance was greater than at the higher temperatures (16, 22 C). Cellular content of N was independent of irradiance and temperature, and the C:N ratio ranged from 5 to 8 with a slight tendency to lower values at low irradiance. Cell volume was not influenced by either temperature or irradiance.     

CORRELATED EVOLUTION OF GENOME SIZE AND CELL VOLUME IN DIATOMS (BACILLARIOPHYCEAE)1

A correlation between genome size and cell volume has been observed across diverse assemblages of eukaryotes. We examined this relationship in diatoms (Bacillariophyceae), a phylum in which cell volume is of critical ecological and biogeochemical importance. In addition to testing whether there is a predictive relationship across extant species, we tested whether evolutionary divergences in genome size were correlated with evolutionary divergences in cell size (using independent contrasts). We estimated total DNA content for 16 diatom species using a flow cytometer and estimated cell volumes using critical dimensions with scaling equations. Our independent contrast analyses indicated a significant correlated evolution between genome size and cell volume. We then explored the evolutionary and ecological implications of this evolutionary relationship. Diatom cell volume is an important component of the global carbon cycle; therefore, understanding the mechanisms that drive diatom genome evolution has both evolutionary and ecological importance.