Interactions between nitrate and ammonium uptake: variation with growth rate,nitrogen source and species

The interaction between nitrate and ammonium uptake was examined as a function of preconditioning growth rate and nitrogen source by adding nitrate, ammonium, or both to nitrogen-sufficient,-deficient, and-starvedSkeletonema costatum (Grev.) Cleve and nitrogen-deficientChaetoceros debilis Cleve. By simultaneously measuring the internal accumulation of intermediates of nitrogen assimilation and the rates of nitrogen assimilation, the metabolic control of nitrogen uptake could be assessed. After the simultaneous addition of nitrate and ammonium to culture, both nitrate and ammonium uptake rates were decreased in comparison with the rates observed when each was added alone, although nitrate uptake was usually decreased more than ammonium uptake. Since both nitrate and ammonium uptake rates vary with time, preconditioning growth conditions, nitrogen sources present, and species, it was necessary to use several different indices to quantify inhibition. In general, ammonium inhibition of nitrate uptake inS. costatum was greatest in cultures preconditioned to ammonium and those at low growth rates, whereas ammonium uptake was inhibited most in cultures preconditioned to nitrate. In nitrogen-deficientC. debilis, nitrate uptake was more inhibited by ammonium, but uptake returned to normal rates more quickly than inS. costatum, whereas inhibition of ammonium uptake was similar. These results explain why the interaction between nitrate and ammonium uptake in the field can be so variable. Inhibition of uptake is not controlled by internal ammonium or total amino acids, nor is it related to the inability to reduce nitrate. Instead, inhibition must be determined in part by the external concentration of nitrogen compounds and in part by some intermediate(s) of nitrogen assimilation present inside the cell.Bigelow Laboratory Contribution No. 82022     

Adaptations of Gracilaria pacifica (Rhodophyta) to nitrogen procurement at different intertidal locations

Nitrogen uptake, assimilation and accumulation were studied in three populations of Gracilaria pacifica Abbott in Bamfield Inlet, British Columbia, Canada, over three summers, 1979–1981. Two of these populations were in the intertidal one high and one low, and the third was a subtidal cultured population. Nitrate uptake rates, internal nitrate content and nitrate reductase activities were highest in the low intertidal population. Time-courses of uptake and uptake kinetics were studied. Both nitrate and ammonium were taken up simultaneously. Thalli from the high-intertidal population showed enhanced nitrate and ammonium uptake following mild desiccation, and greater tolerance to desiccation in terms of maintaining nitrogen uptake after severe desiccation. Transplants were made to determine the effect of intertidal height and geographic location on responses to desiccation, nitrogen uptake, assimilation and accumulation. Nitrate and ammonium uptake rates were dependent on intertidal height and geographic location. Transplanting up the intertidal increased nitrate uptake and nitrate reductase activity, but decreased the nitrate content of the thalli. There were few significant differences in ammonium uptake rates, and ammonium, amino acid, and soluble-protein content of the various populations. All high-intertidal populations, transplanted or natural, showed enhanced nitrate uptake rates following desiccation. Enhanced ammonium uptake rates following desiccation were restricted to the high-intertidal thalli in only one geographic location. Tolerance to higher levels of desiccation also appeared to be intertidal height-dependent, but required more than five weeks to fully develop or disappear.     

Effects of host feeding and dissolved ammonium on cell division and nitrogen status of zooxanthellae in the hydroid Myrionema amboinense

There is a relationship between host feeding, nitrogen status and mitotic activity of zooxanthellae symbiotic with the marine hydroid Myrionema amboinense. Decreases in the mitotic index of zooxanthellae in starved M. amboinense, and in internal pool sizes of glutamine and glutamate, amino acids involved in ammonium assimilation via the glutamine synthetase-glutamate synthase (GS/GOGAT) pathway, were partially restored by addition of ammonium chloride to seawater in which hydroids were incubated. Levels of glutamine were more sensitive to host starvation than levels of glutamate, resulting in a decrease in the glutamine: glutamate molar ratio to that found in zooxanthellae cultured on nitrate. Hydroids starved for 5 d and then incubated in different concentrations of ammonium chloride showed a positive correlation between ammonium concentration and mitotic index of their symbiotic zooxanthellae. Host starvation caused a decrease in perturbation of levels of glutamine and glutamate during ammonium assimilation, as well as decreases in rates of assimilation of [¹⁴C]-leucine into TCA-insoluble protein, and in photosynthetic incorporation of [¹⁴C]-bicarbonate. These observations suggest that host starvation reduces nitrogen supply to the zooxanthellae, causing nitrogen stress to the symbionts and reduction in metabolic processes associated with nitrogen assimilation and photosynthesis as well as with cell division.     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Partitioning of nitrogen and carbon in cultures of the marine diatom Thalassiosira fluviatilis supplied with nitrate,ammonium, or urea

Small or negligible differences in growth rates, average cell size, yields in cell numbers and total cell volumes were found in cultures of Thalassiosira fluviatilis inriched with nitrate, ammonium, or urea. Intracellular pools of unassimilated nitrate, nitrate, and ammonium were found in nutrient-rich conditions, but urea was not accumlated internally. Nitrogen assimilation into organic combination rather than nitrogen nutrient uptake was a critical rate-limiting step in nitrogen utilization. The free amino acid pool, protein, lipid-associated nitrogen, pigments, and total cell nitrogen were all highest in young or mature phase cells and decreased with age in senescent cells, whereas chitan, lipid, carbohydrate, and total cellular carbon all continued to increase during senescence. Dissolved organic nitrogen compounds accumulated in the medium only during senescence. C:N and lipid:protein were sensitive indicators of nitrogen depletion and age in T. fluviatilis.     

Availability of two forms of dissolved nitrogen to the coral Pocillopora damicornis and its symbiotic zooxanthellae

The relative contribution of dissolved nitrogen (ammonium and dissolved free amino acids DFAAs) to the nitrogen budget of the reef-building coral Pocillopora damicornis was assessed for colonies growing on control and ammonium-enriched reefs at One Tree Island (southern Great Barrier Reef) during the ENCORE (Enrichment of Nutrient on Coral Reef; 1993 to 1996) project. P. damicornis acquired ammonium at rates of between 5.1 and 91.8 nmol N cm⁻² h⁻¹ which were not affected by nutrient treatment except in the case of one morph. In this case, uptake rates decreased from 80.5 to 42.8 nmol cm⁻² h⁻¹ (P < 0.05) on exposure to elevated ammonium over 12 mo. The presence or absence of light during measurement did not influence the uptake of ammonium ions. Nitrogen budgets revealed that the uptake of ammonium from concentrations of 0.11 to 0.13 μM could completely satisfy the demand of growing P. damicornis for new nitrogen. P. damicornis also took up DFAAs at rates ranging from 4.9 to 9.8 nmol N cm⁻² h⁻¹. These rates were higher in the dark than in the light (9.0 vs 5.1 nmol m⁻² h⁻¹, P < 0.001). Uptake rates were highest for the amino acids serine, arginine and alanine, and lowest for tyrosine. DFAA concentrations within the ENCORE microatolls that received ammonium were undetectable, whereas they ranged up to 100 nM within the control microatolls. The contribution of DFAAs to the nitrogen budget of P. damicornis constituted only a small fraction of the nitrogen potentially contributed by ammonium under field conditions. Even at the highest field concentrations measured during this study, DFAAs could contribute only ≃11.3% of the nitrogen demand of P.␣damicornis. This contribution, however, may be an important source of nitrogen when other sources such as ammonium are scarce or during periods when high concentrations of DFAAs become sporadically available (e.g. cell breakage during fish-grazing). Received: 22 April 1998 / Accepted: 3 November 1998     

Saturated uptake kinetics: transient response of the marine diatom Thalassiosira pseudonana to ammonium,nitrate, silicate or phosphate starvation

Changes in the saturated uptake kinetics of the limiting nutrient were followed as Thalassiosira pseudonana (Clone 3 H) batch cultures entered ammonium, nitrate, silicate and phosphate starvation. Cultures starved of ammonium or phosphate developed very high specific uptake capacities over a 24 to 48 h starvation period, due to both decreases in cell quota and increases in uptake rates per cell. In particular, the cell phosphorus quota decreased ca. 8-fold during phosphate starvation and specific uptake rates exceeded 100 d^-1. In contrast, cultures entering nitrate or silicate starvation underwent little or no further cell division, and the uptake capacity declined during starvation. After 24 to 48 h starvation, an induction requirement for uptake of nitrate or silicate was apparent. These responses are consistent with adaptation to the pattern of supply of these nutrients in the field.     

Species differences in accumulation of nitrogen pools in phytoplankton

The changes in the intracellular concentrations of nitrate, ammonium, free amino acids, protein, DNA, RNA and total nitrogen were measured in batch cultures of seven species of marine phytoplankton as they progressed from being nitrogen sufficient to being nitrogen starved. After several days of nitrogen starvation, either nitrate or ammonium was added to the cultures, and the measurements were continued for 10 to 36 h. By this means it was possible to assess the long-term and short-term changes in cellular nitrogen compounds and how they relate to phytoplankton nitrogen uptake and growth. Considerable species differences were observed in the amounts and kinds of nitrogen compounds which were stored and the degree to which the utilization of these compounds could support growth if the external nitrogen supply is low or variable. Despite the species variation, the results suggest that the concentrations or ratios of a number of intracellular nitrogen compounds can be used to assess the nitrogen deficiency and/or growth rate of natural phytoplankton populations.Contribution No. 1373 from the School of Oceanography, University of Washington and Contribution No. 83006 from the Bigelow Laboratory for Ocean Sciences     

Desiccation enhanced nitrogen uptake rates in intertidal seaweeds

Desiccation increased nitrate and ammonium uptake rates upon resubmergence in late summer populations of the intertidal macroalgae Gigartina papillata (C.Ag.) J.Ag., Enteromorpha intestinalis (L.) Grev., Fucus distichus L., and Pelvetiopsis limitata (Setch) Gardn. The ratio of nitrogen uptake rates in desiccated plants to rates in hydrated plants (controls) was correlated with the position of the species in the intertidal zone. Gracilaria pacifica Abbott., the species occurring at the lowest shore level, showed no enhancement of nitrogen uptake following desiccation. The high intertidal species such as P. limitata and F. distichus showed a two-fold enhancement of nitrate and ammonium uptake following more extensive desiccation (>30%) and continued uptake even following severe desiccation (50 to 60%). After the plants had been desiccated, the increase in nitrate uptake rates upon submergence lasted much longer than a similar enhancement of ammonium uptake. The duration of the enhanced nitrate uptake was similar to the time required for total rehydration but the uptake rates were not related to the state of rehydration. The potential contribution that this enhanced nitrogen uptake following desiccation could make to total nitrogen procurement for growth is discussed. The experiments were carried out in 1979 or 1980 and repeated in 1981.This paper is dedicated to Dr. R. F. Scagel on the occasion of his retirement for his outstanding contribution to phycology     

Short-term interaction between nitrate and ammonium uptake inThalassiosira pseudonana: Effect of preconditioning nitrogen source and growth rate

The effect of preconditioning nitrogen source and growth rate on the interaction between nitrate and ammonium uptake was determined inThalassiosira pseudonana (Clone 3H). A new method, using cells on a filter (Parslow et al. 1985), allowed continuous measurement of uptake from 0.5 to 9 min after the addition of nitrate, ammonium, or both, with no variation in concentration during the course of the experiment. For each preconditioning N source and growth rate, a series of uptake experiments was conducted, including controls with only nitrate or only ammonium, and others with different combinations of concentrations of nitrate and ammonium. For the first time, preference for ammonium was separated from inhibition of nitrate uptake by ammonium. Ammonium was the preferred N source, i.e. if nitrate and ammonium were presented separately, ammonium uptake rates exceeded nitrate uptake rates. Preference for ammonium varied with both preconditioning N source and growth rate. Inhibition of nitrate uptake by ammonium, determined by comparing nitrate uptake in the presence and absence of ammonium, was observed at ammonium concentrations > 1µM, but was rarely complete. Inhibition of nitrate uptake by ammonium was less in the ammonium-limited culture than in the cultures growing on nitrate, but invariant with growth rate in the nitrate-grown cultures. Below 1µM ammonium, nitrate uptake was often stimulated and rates exceeded those in the controls without ammonium. Ammonium uptake was not inhibited by the presence of nitrate.T. pseudonana fits the classical view of the interaction between nitrate and ammonium uptake in some respects, such as preference for ammonium, and inhibition of nitrate uptake by ammonium concentrations > 1µM. However, at ammonium concentrations typical of most marine environments, nitrate uptake occurs at rapid rates. In other respects, N uptake inT. pseudonana deviates from the classical view in the following ways: (1) stimulation of nitrate uptake by low concentrations of ammonium; (2) lack of inhibition of nitrate uptake by ammonium at low nitrate concentrations; and (3) variation in preference and inhibition with preconditioning, which is markedly different for other species. Because of the apparent enormous species variation in the interaction between nitrate and ammonium uptake and the lack of detailed information for a variety of species, it is difficult to generalize about the effect of ammonium on nitrate uptake, especially in the field, where prior N availability and species composition are not usually addressed.     

Interactions of inorganic nitrogen in the uptake and assimilation by marine phytoplankton

The effect of ambient ammonium concentration on the nitrate uptake rate of marine phytoplankton was investigated. These studies consisted of laboratory experiments using unialgal species and field experiments using natural phytoplankton communities. In laboratory experiments, ammonium suppressed the uptake rates of nitrate and nitrite. Approximately 30 min were required for ammonium to exhibit its fully inhibitory effect on nitrate uptake. At high ammonium concentration (>3 g-at/l), a residual nitrate uptake rate of approximately 0.006 h^-1 was observed. When the ambient ammonium concentration was reduced to a value less than 1 g-at/l, the suppressed nitrate uptake rate subsequently attained a value comparable to that observed before the addition of ammonium. A range of 25 to 60% reduction in the nitrate uptake rate of natural phytoplankton communities was observed at ambient ammonium concentrations of 1.0 g-at/l. A mechanism is proposed for the suppression of nitrate uptake rate by ammonium through feedback control of the nitrate permease system and/or the nitrate reductase enzyme system. The feedback control is postulated to be regulated by the level of total amino acids in the cell.Contribution No. 936 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of a dissertation submitted to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

Development of rapid ammonium uptake during starvation of batch and chemostat cultures of the marine diatom Thalassiosira pseudonana

Cell nitrogen quotas and uptake rates following ammonium additions were measured during ammonium-limited growth transients obtained by starving batch and chemostat cultures of Thalassiosira pseudonana (Clone 3 H). During starvation, cell quotas decreased by more than 50% in batch cultures. In chemostat cultures, the drop in cell quota during starvation decreased with dilution rate, from more than 50% at 1.45 d^-1, to less than 10% at 0.22 d^-1. Minimal levels of 3 to 4×10^-2 pg-at. N cell^-1 were reached after 24 h starvation in both batch and chemostat cultures. Uptake rates over the first minute of perturbation experiments were 3 times the long-term (10 to 30 min) rates. In batch cultures, specific uptake rates increased from 4 d^-1 to 20 d^-1 after 24 h starvation. Uptake rates per cell were independent of starvation time and dilution rate in chemostat cultures, but lower in non-starved batch cultures. The implications of these data for models of phytoplankton growth are discussed: the data support models which predict a depression in average growth rates when diatoms encounter microscale patches in oligotrophic environments.     

The rapid response of the marine diatom Skeletonema costatum to changes in external and internal nutrient concentration

A nitrogen-deficient batch culture of the marine diatom Skeletonema costatum, when resupplied with a mixture of nitrate and ammonium, showed an initial enhanced nitrate uptake rate leading to a large internal concentration (pool) of nitrate. Following this initial nitrate uptake event, nitrate uptake ceased, and nitrate assimilation was inhibited until the ammonium present was used. At this point, nitrate uptake resumed and nitrate assimilation began. No internal ammonium pool was observed during nitrate utilization, but a large nitrate pool remained throughout the utilization of external nitrate. The internal nitrate pool decreased rapidly after exhaustion of nitrate from the culture medium, but growth of cellular particulate nitrogen continued for about 24 h. A mathematical simulation model was developed from these data. The model cell consisted of a nitrate pool, ammonium pool, dissolved organic nitrogen pool, and particulate nitrogen. It was found that simple Michaelis-Menten functions for uptake and assimilation gave inadequate fit to the data. Michaelis-Menten functions were modified by inclusion of inhibitory and stimulatory feedback from the internal pools to more accurately represent the observed nutrient utilization.     

Seasonal variation in total and soluble tissue nitrogen of Pleurophycus gardneri (Phaeophyceae: Laminariales) in relation to environmental nitrate

Seasonal variations in tissue nitrogen (ethanol soluble nitrate and ninhydrin positive substances, as well as total nitrogen) of different thallus parts of Pleurophycus gardneri Setchell and Saunders were monitored simultaneously with ambient seawater nitrate from 1982 until 1984 in Bamfield, Vancouver Island, British Columbia, Canada. A trend of low, nearly zero levels in ambient nitrate typical for the area in late spring and early summer normally contrasts with average nitrate concentrations of 10 mol NO₃ ^- l^-1 in late fall and winter. Total nitrogen content was greater in the perennial thallus parts, stipe and holdfast than in the annual blade and peaked in fall and early winter. The longitudinal thallus distribution of nitrate revealed a distinct and significant concentration of nitrate in the haptera reaching at maximum 8% nitrate-N of the internal total nitrogen. Internal nitrate concentration ranged from 20 to 5 000 times the ambient nitrate concentration in the midrib, and from 40 to 3 100 times in the wing, while the range was greatest with 400 to 14 000 times in the haptera. P. gardneri contained at most about 7 mol NO₃ ^- g fresh wt^-1 in the blade, which corresponds to about 6% of total tissue nitrogen. Ninhydrin positive substances comprised the major portion of the soluble N pool in P. gardneri and showed a pronounced seasonality. Concentrations of ninhydrin positive substances ranged from 20 to 800 g N g fresh wt^-1 in the midrib and in the wing. In the stipe, ninhydrin positive substances varied from 180 to 2 200 g N g fresh wt^-1, and from 250 to 1 200 g N g fresh wt^-1 in the haptera. Evidence is given that (1) the perennial parts, stipe and haptera of P. gardneri contain the majority of nitrogen products independent of season and ontogenetic stage; (2) ninhydrin positive substances are the most abundant internal nitrogen constituents; (3) the low N values in the blade in summer suggest a nitrogen limited growth; and (4) nitrate may not be the predominant external nitrogen source.     

Development of the ability to take up L-lysine by the diatom Phaeodactylum tricornutum

Freshly harvested cells of Phaeodactylum tricornutum Bohlin grown with nitrate, ammonium or lysine as a sole nitrogen source had a low ability to take up lysine or arginine, but this ability increased when cells were deprived, over 48 h, of either nitrogen or carbon. The effects of nitrogen and carbon deprivation were additive, and the uptake ability was greatest in cells incubated in darkness in nitrogen-free medium. Uptake ability increased in cells illuminated in the presence of 10^-5 M 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) an inhibitor of photosynthetic electron transport. An inverse relationship between rate of development of the uptake system and rate of photosynthesis was also established. Development of the uptake system was prevented by cycloheximide or by anaerobiosis. Following transfer to a normal nitrate medium, illuminated cells lost the lysineuptake system by dilution as the cells grew. There was a linear and positive correlation between the initial rate of uptake of lysine and the maximum concentration which was maintained in the cells when equilibrium was reached, indicating that transinhibition of lysine uptake may occur and that the extent of this inhibition is related to the size of the internal amino acid pool. The relevance of the findings to the growth of phytoplankton in natural waters is discussed.     

Growth of zooxanthellae in culture with two nitrogen sources

Physiological characteristics of zooxanthellae were examined under nutrient-saturated conditions created by mixing ammonium (¹⁵NH₄) with nitrate (¹⁵NO₃) to give 0.88 mM total nitrogen. Growth rate varied with the form of nitrogen provided. Ammonium alone resulted in the lowest C:N and C:chl-a ratios. Although zooxanthellae took up nitrate in the absence of ammonium, ammonium assimilation was 1.3 times higher than nitrate assimilation. Ammonium strongly inhibited nitrate assimilation. While high-ammonium treatments resulted in the highest ¹⁴C incorporation into intermediate compounds, high nitrate levels resulted in the highest ¹⁴C incorporation into protein, suggesting that the intermediate compounds are produced prior to the subsequent production of protein when ammonium is the dominant N source. The enhanced production of intermediate compounds at the expense of carbon directed to protein synthesis in the presence of ammonium might be analogous to the “host factor” observed in zooxanthellae–host symbioses, since growth rate is depressed due to low production of protein. Received: 16 March 2000 / Accepted: 26 August 2000     

Nitrogen uptake and growth of the germlings and mature thalli of Fucus distichus

Fucus distichus L. was collected near Vancouver, Canada, in late fall and early winter, 1981. The effects of the forms of nitrogen (nitrate, ammonium or urea) and periodic exposure to air on growth, rhizoid development and nitrogen uptake in germlings was investigated. Gamete release, fertilization, germination and germling growth had no requirement for a specific form of nitrogen. Periodic exposure to air increased secondary rhizoid development twofold. Nitrate and ammonium uptake rates of the germlings were higher than for the mature thalli (20 to 40 times for nitrate and 8 times for ammonium), while the halfsaturation constant (K _s) values for nitrate were similar (1 to 5 M). The germlings showed saturable uptake kinetics but the mature thalli did not. When germlings were exposed to air it caused a 70% decrease in nitrate uptake, but not change in ammonium uptake. Ammonium uptake in the mature thalli was proportional to the ambient ammonium concentration. Nitrate uptake in the mature thalli appeared to follow saturation kinetics at low nitrate concentrations, but showed a non-saturable component at concentrations greater than 10 M. Presence of ammonium inhibited nitrate uptake by the mature plants but not by the germlings.     

不同氮肥对油菜B与Mo吸收的影响

利用盆栽试验研究了潮土中不同氮肥对油菜硼钼吸收的影响，结果表明，前期(5周)施氮并没有促进油菜的生长，而且大部分氮肥对油菜生长表现抑制作用，氯化铵处理的作用最大，硫酸铵和尿素处理次之，硝酸铵处理再次之；不同氮肥均有抑制油菜生长前期(5周)硼吸收和累积的作用，其中以氯化铵、硝酸铵和尿素的作用为甚；不同氮肥同样抑制油菜生长前期(5周)对钼的吸收和累积，以硫酸铵处理作用最突出，氯化铵处理次之。但随着时间的推移，10、15周不同氮肥大多促进油菜的生长，增加油菜对硼钼的吸收和累积，且各氮肥处理间的差异逐步减小。     

Effect of seawater velocity on inorganic nitrogen uptake by morphologically distinct forms of Macrocystis integrifolia from wave-sheltered and exposed sites

In conditions of low water motion (<0.06 ms^–1), the availability of essential nutrients to macroalgae, and thus their potential productivity, may be limited by thick diffusion boundary-layers at the thallus surface. The ability of macroalgae to take up nutrients in slow moving water may be related to how their blade morphology affects diffusion boundarylayer thickness. For the giant kelp, Macrocystis integrifolia Bory, morphological measurements indicate that blades of plants from a site exposed to wave action are thick, narrow and have a heavily corrugated surface. In contrast, blades from a site with a low degree of water motion are relatively thin, with few surface corrugations and large undulations along their edges. The aim of our work was to test the hypothesis that morphological features of M. integrifolia blades from a sheltered site allow enhanced inorganic nitrogen uptake at low seawater velocities compared to blades with a wave-exposed morphology. The rate of nitrate and ammonium uptake by morphologically distinct blades of M. integrifolia, from sites that were sheltered from and exposed to wave action, were measured in the laboratory at a range of seawater velocities (0.01 to 0.16 ms^–1), between March and May 1993. For both sheltered and exposed blade morphologies, nitrate and ammonium uptake rates increased with increasing seawater velocity, reaching a maximum rate at 0.04 to 0.06 ms^–1. Uptake parameters V _max (maximum uptake rate) and U _0.37 (the velocity at which the uptake rate is 37% of the maximum rate) were estimated using an exponential decay formula. These parameters were similar for both blade morphologies, at all seawater velocities tested. Additional measurements suggest that the nitrogen status of M. integrifolia blades from wavesheltered and exposed sites were similar throughout the experimental period, and thus nitrogen status did not affect the rate of nitrogen uptake in these experiments. on the basis of these results, we conclude that blade morphology does not enhance nitrogen uptake by M. integrifolia in conditions of low water motion. Potential effects of diffusion boundary-layers on kelp productivity are discussed.