INTERACTIONS BETWEEN UV‐B EXPOSURE AND PHOSPHORUS NUTRITION. I. EFFECTS ON GROWTH,PHOSPHATE UPTAKE,AND CHLOROPHYLL FLUORESCENCE1

The interactive effects of P starvation and exposure to UV radiation on growth rates, quantum efficiency of PSII electron transport, and P‐uptake capacity of the chlorophyte microalga Dunaliella tertiolecta Butcher are presented. Ultraviolet radiation did not in itself cause marked changes in growth rate, though it did induce changes in the effective quantum yield of PSII. Depriving cells of phosphate resulted in significant changes in all parameters examined. The decline of growth rate and fluorescence parameters after P starvation was significantly faster in the presence of UV radiation. Ultraviolet radiation also stimulated the magnitude of the transient changes in chl fluorescence (nutrient‐induced fluorescence transient) exhibited by P‐starved cells after resupply of that nutrient.     

RAPID AMMONIUM‐ AND NITRATE‐INDUCED PERTURBATIONS TO CHL a FLUORESCENCE IN NITROGEN‐STRESSED DUNALIELLA TERTIOLECTA (CHLOROPHYTA)1

When NH₄ ⁺ or NO₃ ^? was supplied to NO₃ ^? ‐stressed cells of the microalga Dunaliella tertiolecta Butcher, immediate transient changes in chl a fluorescence were observed over several minutes that were not seen in N‐replete cells. These changes were predominantly due to nonphotochemical fluorescence quenching. Fluorescence changes were accompanied by changes in photosynthetic oxygen evolution, indicating interactions between photosynthesis and N assimilation. The magnitude of the fluorescence change showed a Michaelis‐Menten relationship with half‐saturation concentration of 0.5 μM for NO₃ ^? and 10 μM for NH₄ ⁺ . Changes in fluorescence responses were characterized in D. tertiolecta both over 5 days of N starvation and in cells cultured at a range of NO₃ ^? ‐limited growth rates. Variation in responses was more marked in starved than in limited cells. During N starvation, the timing and onset of the fluorescence responses were different for NO₃ ^? versus NH₄ ⁺ and were correlated with changes in maximum N uptake rate during N starvation. In severely N‐starved cells, the major fluorescence response to NO₃ ^? disappeared, whereas the response to NH₄ ⁺ persisted. N‐starved cells previously grown with NH₄ ⁺ alone showed fluorescence responses with NH₄ ⁺ but not NO₃ ^? additions. The distinct responses to NO₃ ^? and NH₄ ⁺ may be due to the differences between regulation of the uptake mechanisms for the two N sources during N starvation. This method offers potential for assessing the importance of NO₃ ^? or NH₄ ⁺ as an N source to phytoplankton populations and as a diagnostic tool for N limitation.     

STATE TRANSITIONS AND NONPHOTOCHEMICAL QUENCHING DURING A NUTRIENT‐INDUCED FLUORESCENCE TRANSIENT IN PHOSPHORUS‐STARVED DUNALIELLA TERTIOLECTA1

Assessments of nutrient‐limitation in microalgae using chl a fluorescence have revealed that nitrogen and phosphorus depletion can be detected as a change in chl a fluorescence signal when nutrient‐starved algae are resupplied with the limiting nutrient. This photokinetic phenomenon is known as a nutrient‐induced fluorescence transient, or NIFT. Cultures of the unicellular marine chlorophyte Dunaliella tertiolecta Butcher were grown under phosphate starvation to investigate the photophysiological mechanism behind the NIFT response. A combination of low temperature (77 K) fluorescence, photosynthetic inhibitors, and nonphotochemical quenching analyses were used to determine that the NIFT response is associated with changes in energy distribution between PSI and PSII and light‐stress‐induced nonphotochemical quenching (NPQ). Previous studies point to state transitions as the likely mechanism behind the NIFT response; however, our results show that state transitions are not solely responsible for this phenomenon. This study shows that an interaction of at least two physiological processes is involved in the rapid quenching of chl a fluorescence observed in P‐starved D. tertiolecta: (1) state transitions to provide the nutrient‐deficient cell with metabolic energy for inorganic phosphate (P_i)‐uptake and (2) energy‐dependent quenching to allow the nutrient‐stressed cell to avoid photodamage from excess light energy during nutrient uptake.     

PHYSIOLOGICAL RESPONSES TO PHOSPHORUS LIMITATION IN BATCH AND STEADY-STATE CULTURES OF DUNALIELLA TERTIOLECTA (CHLOROPHYTA): A UNIQUE STRESS PROTEIN AS AN INDICATOR OF PHOSPHATE DEFICIENCY1

A protein unique to phosphorus stress observed in Dunaliella tertiolecta Butcher was studied in the context of phosphate-limited cell physiology and is a potential diagnostic indicator of phosphate deficiency in this alga. Cells were grown over a range of limited, steady-state growth rates and at maximum (replete) and zero (phosphate-starved) growth rates. The stress protein, absent in nutrient-replete cells, was produced under all steady-state phosphate-limited conditions and increased in abundance with increasing limitation (decreasing growth rate). Cellular carbon: phosphorus ratios and the maximum uptake rate of phosphate (V_m) increased with increasing limitation, whereas the ratio of chlorophyll a: carbon decreased. Alkaline phosphatase activity did not respond to limitation but was measurable in starved, stationary-phase cells. F_v/F_m, a measure of photochemical efficiency, was a nonlinear, saturating function of p, as commonly observed under N limitation. The maximum F_v/F_m of 0.64 was measured in nutrient-replete cells growing at μ_max, and a value of zero was measured in stationary-phase starved cells. When physiological parameters were compared, the P-stress protein abundance and F_v/F_m were the most sensitive indicators of the level of deficiency. The stress protein was not produced under N- or Fe-limited conditions. It is of high molecular weight (>200) and is associated with internal cell membranes. The stress protein has several characteristics that make it a potential diagnostic indicator: it is 1) unique to phosphorus limitation (i.e. absent under all other conditions), 2) present under limiting as well as starved conditions, 3) sensitive to the level of limitation, and 4) observable without time-course incubation of live samples.     

AMMONIUM INDUCED PHOTOSYNTHETIC SUPPRESSION IN AMMONIUM LIMITED DUNALIELLA TERTIOLECTA (CHLOROPHYTA)1

Dunaliella tertiolecta (Butcher) was grown in chemostat culture over a wide range of ammonium limited growth rates. The addition of ammonium caused a rapid temporary suppression of photosynthetic carbon fixation. The magnitude of ammonium induced photosynthetic suppression increased with the severity of ammonium limitation. Cells growing at rates greater than ca. 80%μ_max exhibited no photosynthetic suppression in response to additions of the limiting nutrient. The duration of photosynthetic suppression was related to the concentration of added ammonium. Immediately following the suppression, photosynthesis was enhanced with respect to the controls. The eventual degree of enhancement increased with the concentration of added ammonium. Steady-state cellular chlorophyll concentrations, photosynthetic rates, and assimilation numbers are reported.     

INHIBITION OF CASPASE‐LIKE ACTIVITIES PREVENTS THE APPEARANCE OF REACTIVE OXYGEN SPECIES AND DARK‐INDUCED APOPTOSIS IN THE UNICELLULAR CHLOROPHYTE DUNALIELLA TERTIOLECTA1

When the chlorophyte alga Dunaliella tertiolecta Butcher is placed in darkness, a form of programmed cell death with many similarities to apoptosis is induced, including the induction of caspase‐like proteases. Many uncertainties about the regulation and mediators that participate in the process remain. To examine the relationship between caspase‐like activities and different apoptotic events (i.e., phosphatidylserine [PS] translocation), increases in membrane permeability and numbers of dead cells revealed by SYTOX‐green staining, and the generation of reactive oxygen species (ROS), we used the broad‐range caspase inhibitor Boc‐D‐FMK to block the activity of the whole class of caspase‐like proteins simultaneously. In the presence of the inhibitor, ROS were not produced, and cells did not die. Loss of membrane asymmetry, indicated by external labeling of PS by annexin V, was apparent at midstages of light deprivation, although it did not conform to the typical pattern for PS exposure observed in metazoans or vascular plants, which occurs at early stages of the apoptotic event. Thus, we have evidence for a link between ROS and cell death involving caspase‐like enzymes in an alga. The fact that caspase‐like inhibitors prevent not only cell death, but also ROS and loss of cell membrane integrity and asymmetry, suggests that caspase‐like proteases might have regulatory roles early in cell death, in addition to dismantling functions.     

A CHLOROPHYLL FLUORESCENCE INDEX TO ESTIMATE SHORT‐TERM RATES OF PHOTOSYNTHESIS BY INTERTIDAL MICROPHYTOBENTHOS1

An index based on chl a fluorescence quenching analysis was tested as a predictor of photosynthetic rates of undisturbed intertidal microphytobenthic assemblages. The fluorescence index, P_fluo, was derived from the combination of different chl a fluorescence parameters chosen to represent the two main sources of short‐term variability in the community‐level microphytobenthic photosynthesis: 1) the quantum yield of photosynthesis of the microalgae present in the photic zone of the sediment, φP, and 2) the community‐level efficiency of photosynthetic light absorption, ?, determined by the microalgal concentration in the photic zone. Variations in φP were traced by the fluorescence index ΔF/F_m′ (the effective quantum yield of charge separation at PSII), whereas changes in ? were followed by the fluorescence parameter F_o (dark or minimum fluorescence level). Gross photosynthetic rate, P, and fluorescence parameters were measured nondestructively and simultaneously under in situ conditions, on the same samples, using oxygen microelectrodes and pulse amplitude modulation fluorometry, respectively. Despite the large and uncorrelated hourly variability in irradiance, photosynthetic rate, and fluorescence parameters included in P_fluo, highly significant correlations between P_fluo and P were found for all the sampling periods, encompassing hourly, biweekly, and seasonal time scales. The variability in P explained by P_fluo ranged from 84.3% to 91.4% when sampling periods were considered separately and reached 81.1% when all data were pooled. The results of the study show that despite its simplicity, the index P_fluo can be used to trace short‐term variations in the photosynthetic rate of undisturbed microphytobenthic assemblages undergoing rhythmic vertical migration.     

INFLUENCE OF PHOSPHORUS LIMITATION ON TOXICITY AND PHOTOSYNTHESIS OF ALEXANDRIUM MINUTUM (DINOPHYCEAE) MONITORED BY IN‐LINE DETECTION OF VARIABLE CHLOROPHYLL FLUORESCENCE1

The effects of phosphorus (P) limitation on growth, toxicity, and variable chl fluorescence of Alexandrium minutum were examined in batch culture experiments. Cell division was greatly impaired in P‐limited cultures, but P spiking of these cultures after 9 days stimulated high levels of cell division equivalent to P‐replete cultures. The cellular concentration of paralytic shellfish toxins was consistent over the growth cycle of control cultures from lag phase into logarithmic growth phase, with toxins repeatedly lost to daughter cells during division. The low level of cell division in P‐limited cultures resulted in a 10‐fold increase of cellular toxin compared with controls, but this dropped upon P spiking due to increased rates of cell division. The history of phosphorus supply had an important effect on toxin concentration, with the P‐limited and the P‐spiked cultures showing values 2‐fold higher than the P‐replete cultures. Toxin profiles of the A. minutum strain used in these experiments were dominated by the N₁‐hydroxy toxins, gonyautoxins (GTX) GTX1 and GTX4, which were approximately 40 times more abundant than their analogues, GTX2 and GTX3, in P‐limited cultures. The dominance of the N₁‐hydroxy toxins increased significantly in control cultures as they advanced through logarithmic growth. In‐line measurements of the variable chl fluorescence of light‐adapted cells indicated consistent photochemical efficiency under P‐replete conditions. P limitation induced a drop in fluorescence‐based photochemical efficiency that was reversible by P spiking. There was an inverse linear relationship between in‐line fluorescence and cell toxin quota (r = ?0.88). Monitoring fluorescence in‐line may be valuable in managing efficient biotechnological production of toxins.     

BIOCHEMICAL COMPOSITION AND PHOTOSYNTHETIC CARBON METABOLISM OF NUTRIENT LIMITED CULTURES OF MERISMOPEDIA TENUISSIMA (CYANOPHYCEAE)1

Continuous cultures of Merismopedia tenuissima Lemmerman, limited by phosphorus, nitrogen, sulfur, or carbon, were compared to non limited batch cultures by two methods. The cellular content of photosynthetic pigments (chlorophyll and phycocyanin) was found to decrease in all nutrient limited cultures, except for the carbon limited culture. The ratio of carbohydrate to protein was 4- to 7-fold higher in P, N or S limited cultures than in non-limited or C limited cultures. The macromolecular products of photosynthesis were determined in samples to which NaH¹⁴CO₃ was added. Relative incorporation into protein decreased in P or N limited cultures, increased accumulation of low molecular weight compounds was found in S and P limited cultures, and little change was noted in C limited cultures as compared to non-limited cultures. Although relative incorporation into protein was significantly greater at 20μEin·m^?2·s^?1 light intensity than at 180 μEin·m^?2.s^?1 in non-limited cultures, this effect was abolished in all nutrient limited cultures. These results suggest that measurement of the cellular carbohydrate to protein ratio and the products of photosynthesis would be useful in the analysis of algal population dynamics in nature.     

不同浓度NaCl和Na2CO3处理对菊芋幼苗光合及叶绿素荧光的影响

 以砂培菊芋(Helianthus tuberosus)幼苗作为试验材料,分别进行不同浓度NaCl (50、 100、150、200、250 mmol&;#8226;L^－1)和Na₂CO₃ (25、50、 75、100、125 mmol&;#8226;L^－1)胁迫处理,以1/2全营养液作为对照,处理7 d后研究NaCl和Na₂CO₃胁迫处理对菊芋幼苗叶片光合作用及叶绿素动力学 参数的影响。结果表明：1)在NaCl处理下,当浓度小于150 mmol&;#8226;L^－1时,增加了菊芋的叶绿素含量、净光合速率(Net photosynthetic rate, P_n)和气孔导度(Stomatal conductivity, G_s),对荧光参数PSⅡ的电子传递情况( F_m/F_o)、PSⅡ原初光能转换效率(F_v/F_m)、PSⅡ量子效率 (Actual quantum yield of PSⅡ under actinic irradiation,φPSⅡ)和光化学猝灭系数(Photochemical quenching coefficient, qP)和非 光化学猝灭系 数(Non-photochemical quenching coefficient, NPQ)没有显著影响,随着浓度的增加,各项生理指标与对照相比除了NPQ显著 增加,其余均显著降低;2)在Na₂CO₃胁迫处理下,随着Na₂CO₃浓度的增加,与对照相比菊芋幼苗叶绿素含量、P_n、G_s以及叶绿素a荧光诱导动力 学参数F_m/F_o、F_v/F_m、φPSⅡ和qP均显著降低,NPQ显著增加;3)就NaCl和Na₂CO₃相比而言,在相同Na⁺浓度情况下,处于Na₂CO₃胁迫下的菊芋 幼苗的叶绿素含量、P_n、G_s以及叶绿素a荧光诱导动力学参数F_m/F_o、F_v/F_m、φPSⅡ和qP下降幅度和NPQ的增加幅度均显著大于NaCl,这说明 NaCl和Na₂CO₃胁迫均对菊芋幼苗造成不同程度的伤害,但在相同Na⁺浓度情况下,Na₂CO₃的伤害程度大于NaCl。由此说明菊芋对盐的忍耐程度高 于碱。     

AMMONIUM,NITRATE, PHOSPHATE,AND INORGANIC CARBON UPTAKE IN AN OLIGOTROPHIC LAKE: SEASONAL VARIATIONS AMONG LIGHT RESPONSE VARIABLES1

The dependence of substrate saturated uptake of ¹⁵NH₄⁺, ¹⁵NO₃^?, ³²PO₄^3?, and ¹⁴CO₂ on photosynthetic photon flux density (PPFD or photsynthetically active radiation, 400–700 nm) was characterized seasonally in oligotrophic Flathead Lake, Montana. PO₄^3? uptake was not dependent upon PPFD at any time of the year, whereas NH₄⁺, NO₃^?, and CO₂ uptake were consistently dependent on PPFD over all seasons. Maximal rates of NH₄⁺, NO₃^? and CO₂ uptake usually occurred near 40% of surface PPFD, which corresponded to about 5 m in the lake; inhibition was evident at PPFD levels greater than 40%. NH₄⁺, NO₃^? and PO₄^3? were incorporated in the dark at measurable rates most of the year, whereas dark CO₂ uptake was always near 0 relative to light uptake. CO₂ and NO₃^? uptake were more strongly influenced by PPFD than was NH₄^3? uptake. The PPFD dependence of PO₄^3?, NH₄⁺, NO₃^? and CO₂ uptake may affect algal growth and nutrient status by influencing the balance in diel and seasonal C:N:P uptake ratios.     

零上低温对水稻剑叶叶绿素荧光变化的影响

在不同的低温处理条件下,测定了离体水稻剑叶的叶绿素荧光诱导曲线。结果表明,与玉米、马玲薯叶片所测定的结果相类似,可变荧光(Fv)有不同程度的降低,而低温对总淬灭(P-T)影响不大。可见,在水稻叶片的碳素同化系统中的基质酶对低温不敏感。     

HEAVY METAL‐INDUCED INHIBITION OF PHOTOSYNTHESIS: TARGETS OF IN VIVO HEAVY METAL CHLOROPHYLL FORMATION1

The targets of heavy metal (here Cu^{2 +} and Zn^{2 +} ) attack on the photosynthetic apparatus of algae belonging to different phyla were investigated. Experiments with the green alga Scenedesmus quadricauda confirmed previous findings that according to the irradiance level two different phenomena occur, which were further characterized by specific changes in several photosynthetic parameters. The reaction occurring under low irradiance (shade reaction) is characterized by heavy metal substitution of Mg^{2 +} in chl molecules bound predominantly in the light harvesting complex II of Chlorophyta (LHC II). Under high irradiance (sun reaction) the LHC II chls are inaccessible to substitution and the damage occurs in the PSII reaction center instead. Algae with antenna proteins other than the LHC II did not show the two types of heavy metal attack at different irradiances. In red algae (Antithamnion plumula), low Cu^{2 +} concentrations induced the sun reaction even at very low irradiance. In brown algae (Ectocarpus siliculosus) the shade reaction occurred even in saturating irradiance. These results also indicate that despite some similarity in their features, the primary step of the sun reaction and photoinhibition is different.     

PREDICTING THE KINETICS OF DISSOLVED INORGANIC CARBON LIMITED GROWTH FROM THE SHORT-TERM KINETICS OF PHOTOSYNTHESIS IN SYNECHOCOCCUS LEOPOLIENSIS (CYANOPHYTA)1

The blue-green alga (Cyanobacterium) Synechococcus leopoliensis (Racib.) Komarek was grown in dissolved inorganic carbon [DIC]-limited chemostats over the entire range of growth rates. At each growth rate, the kinetics of photosynthesis with respect to [DIC] and the maximal rate of photosynthesis (P_max) were determined. The half-saturation constant for [DIC]-limited photosynthesis (K_1/2^DIC) for cells growing below 1.7 d^?1 was constant (4.7 μM) whereas for growth rates between 1.7 d^?1 and 2.1 d^?1 (μ_max) the kinetics of photosynthesis were multiphasic with an apparent K_1/2^DIC between 1.5–2.0 mM. P_max increased in a linear fashion with growth rate for growth rates below 1.7 d^?1. No trend in P_max was apparent for growth rates greater than 1.7 d^?1. These kinetic parameters were used to predict a growth rate versus [DIC] relationship. Results show that the Monod relationship is a physiologically valid expression of growth as a function of [DIC] provided (K_1/2^DIC) remains constant. The major change in (K_1/2^DIC) as μ approaches μ_max results in the conclusion that two separate and distinct Monod equations must be used to describe growth as a function of DIC over the entire growth range. These results point to a major discontinuity in the μ vs. [DIC] curve at 1.7 d^?1 which corresponds to the change from high to low affinity photosynthetic kinetics. We believe these results account for the previously described deficiencies of the Monod equation in describing [DIC]-limited algal growth.     

INTERACTIONS AMONG IRRADIANCE,OXYGEN EVOLUTION AND NITRITE UPTAKE BY CHLAMYDOMONAS (CHLOROPHYCEAE)1,2

Nitrite uptake and oxygenic photosynthesis by cultures of Chlamydomonas sp. isolated from Lake Superior were measured at different irradiances in order to compare predictive models of nitrite uptake and to assess the proportion of photoreductant (measured as oxygen evolution, mol × 4 eq. mol^?1) that is allocated to nitrite assimilation (measured as nitrite uptake, mol × 6 eq. mol^?1). These measurements are analogous to measurements of carbon fixation (CO₂ uptake) at different irradiances and photosynthetic activities. Nitrite uptake as a function of irradiance did not follow Michaelis-Menten kinetics as proposed for nitrate by MacIsaac and Dugdale (1972) because of inhibition at high irradiances. The Haldane equation described nitrite uptake better. Nitrite uptake as a function of oxygenic photosynthesis followed Michaelis-Menten kinetics. Irradiance-dependent (Haldane) and photosynthesis-dependent models described nitrite uptake equally well. We suggest that nitrite is taken up and assimilated in response to intracellular concentrations of photoreductant that are directly proportional to photosynthetic activity and are related indirectly to irradiance. This contention is supported by photosynthesis-dependent nitrite uptake (Michaelis-Menten) at both light-limited and photoinhibited photosynthetic activities. This is consistent conceptually with deactivation of light traps at high irradiance levels. The proportion of photoreductant allocated to nitrite uptake and assimilation increased markedly at low irradiance levels. This indicates that cells synthesize important N-containing biomolecules across a broader range of irradiance levels than fixation of carbon for synthesis of energy storage and structural products.     

IN SEARCH OF A PHYSIOLOGICAL BASIS FOR COVARIATIONS IN LIGHT‐LIMITED AND LIGHT‐SATURATED PHOTOSYNTHESIS1

The photosynthesis‐irradiance (PE) relationship links indices of phytoplankton biomass (e.g. chl) to rates of primary production. The PE curve can be characterized by two variables: the light‐limited slope (α^b) and the light‐saturated rate (P^b_max) of photosynthesis. Variability in PE curves can be separated into two categories: that associated with changes in the light saturation index, E_k (=P^b_max/α^b) and that associated with parallel changes in α^band P^b_max (i.e. no change in E_k). The former group we refer to as “E_k‐dependent” variability, and it results predominantly from photoacclimation (i.e. physiological adjustments in response to changing light). The latter group we refer to as “E_k‐independent” variability, and its physiological basis is unknown. Here, we provide the first review of the sporadic field and laboratory reports of E_k‐independent variability, and then from a stepwise analysis of potential mechanisms we propose that this important yet largely neglected phenomenon results from growth rate–dependent variability in the metabolic processing of photosynthetically generated reductants (and generally not from changes in the oxygen‐evolving PSII complexes). Specifically, we suggest that as growth rates decrease (e.g. due to nutrient stress), reductants are increasingly used for simple ATP generation through a fast (<1s) respiratory pathway that skips the carbon reduction cycle altogether and is undetected by standard PE methodologies. The proposed mechanism is consistent with the field and laboratory data and involves a simple new “twist” on established metabolic pathways. Our conclusions emphasize that simple reductants, not reduced carbon compounds, are the central currency of photoautotrophs.     

PHOSPHORUS LIMITATION AND CARBON METABOLISM IN A UNICELLULAR ALGA: INTERACTION BETWEEN GROWTH RATE AND THE MEASUREMENT OF NET AND GROSS PHOTOSYNTHESIS1

Chlamydomonas reinhardii Dangeard was grown in continuous culture under P limitation at a range of dilution rates. Carbon uptake measurements were performed using double isotope (¹²C/¹⁴C) techniques and the fluxes of carbon in the light and dark were analysed over the range of growth rates. ¹⁴C uptake was shown to be equal to gross photosynthesis only at maximum relative growth rates; at low relative growth rates ¹⁴C uptake approximated net photosynthesis. The altered pattern of C uptake was found to be due to the suppression of dark respiration in the light and the release of ¹⁴C0₂ from respiratory pathways at low relative growth rates. Metabolic channelling of ¹⁴C from photosynthetic pathways to respiratory pathways occurred at low growth rates as the specific activity of the respired CO₂ reached 45% of the input gas mixture. These data are discussed in the light of the controversy concerning the measurement of gross and net photosynthesis in natural populations and in the light of models of ¹⁴C uptake in single celled algae. Existing models are shown to be adequate for high relative growth rates but not for low relative growth rates under P limitation.     

KINETICS OF NUTRIENT UPTAKE AND GROWTH IN PHYTOPLANKTON1

Microscopic algae ran grow rapidly in natural waters that are extremely low in essential macro and micro nutrients. Yet, their nutrient uptake systems exhibit only mediocre nutrient affinities, the saturation constants being often 10–1000 times the (estimated) ambient concentrations. The large difference which exists between the saturation constants for growth (K_μ) and short term uptake (K_ρ) are due to the acclimation capabilities of the organisms. Over the acclimation range, K_μ to K_ρ, the algae can maintain maximum growth rate by modulating both their internal nutrient quotas (Q) and their maximum short term nutrient uptake rates (ρ_max) in response to variations in external nutrient concentrations. The commonly assumed hyperbolic relationships for steady growth and uptake (viz “chemostat theory”) are coherent with a hyperbolic expression for short term uptake including a variable maximum (ρ_max). The ratio of the saturation constants for growth and uptake is then directly related to the extreme in quotas and maximum uptake rates: K_μ/K_ρ= Q_min/Q_max·ρ^lo_max/ρ^hi_max. This result is applicable even when the exact hyperbolic laws are not. Published data on Fe, Mn, P and N limitation in algae are generally in accord with the theory and demonstrate a wider acclimation range for trace than for major nutrients.     

KINETICS OF NUTRIENT UPTAKE AND GROWTH IN PHYTOPLANKTON1

Microscopic algae can grow rapidly in natural waters that are extremely low in essential macro and micro nutrients. Yet, their nutrient uptake systems exhibit only mediocre nutrient affinities, the saturation constants being often 10–1000 times the (estimated) ambient concentrations. The large difference which exists between the saturation constants for growth (K_u) and short term uptake (K_p) are due to the acclimation capabilities of the organisms. Over the acclimation range, K_u, to K_p the algae can maintain maximum growth rate by modulating both their internal nutrient quotas (Q) and their maximum short term nutrient uptake rates (P_max) in response to variations in external nutrient concentrations. The commonly assumed hyperbolic relationships for steady growth and uptake (viz “chemostat theory”) are coherent with a hyperbolic expression for short term uptake including a variable maximum (P_max). The ratio of the saturation constants for growth and uptake is then directly related to the extreme in quotas and maximum uptake rates: K^μ/K^ρ= Q^min/Q^maxρ^max/ρQ^max. This result is applicable even when the exact hyperbolic laws are not. Published data on Fe, Mn, P and N limitation in algae are generally in accord with the theory and demonstrate a wider acclimation range for trace than for major nutrients.