In situ water motion and nutrient uptake by the giant kelp Macrocystis pyrifera

Rates of NO ₃ ^- uptake by individual blades of Macrocystis pyrifera (L.) C. Agardh were measured at different flow rates in the laboratory. Dissolution rates of hemispherical, plaster buttons attached to the blade surface provided a relative measure of flow rates over blades used in uptake experiments and also over intact blades of adult kelp plants in situ (Laguna Beach, California, USA; 1981). Laboratory results indicated that uptake was saturated at a flow rate equivalent to 2.5 cm s^-1 current velocity. Flow rates over intact blades in situ always exceeded this uptake saturation level. Wave surge and movement of plant surfaces relative to the surrounding water provided sufficient flow to saturate uptake, even in a dense kelp canopy during low-current and calm sea-state conditions.     

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.     

Pigment content and photosynthetic rate of the fronds of Macrocystis pyrifera

The Macrocystis pyrifera (L.) C. Ag. frond is here described in terms of chlorophyll a, fucoxanthin, chlorophyll c and photosynthetic rate. Pigment concentrations increased back from the apical meristem reaching a maximum after 2 to 3 m. Pigment concentrations were then generally constant throughout most of the length of the frond, finally decreasing again in the oldest parts of the frond with the exception of the sporophylls. Pigment ratios remained relatively constant throughout. Maximum net photosynthetic rates on a given frond showed a decrease with tissue age on both an area basis (1040 down to 463 nmol O₂ cm^-2 h^-1) and on a chlorophyll a basis, which was shown as half-saturation constants (quantum irradiance) which dropped on an area basis from 85 mol m^-2 sec^-1 at 4.5 m above the holdfast to 26 mol m^-2 sec^-1 at 15.5 m. Young sporophytes transplanted from the sea floor to the surface (12 m) tended to decrease pigment content, while those transplanted to the bottom tended to increase all pigments, but especially fucoxanthin. Photosynthetic rates, however, changed little on a unit area basis. The results of these data are considered in the light of recent work on photosynthetic units, tissue age effects and general adaptations of the M. pyrifera frond to its light environment.     

Carbon acquisition strategies for marine macroalgae

A model system was developed to analyse differences in carbon acquisition strategies among macroalgae. During photosynthesis in a limited volume of seawater the capability of the algae to assimilate inorganic carbon as well as to change the alkalinity of the seawater was analysed. These properties were then related to the status of the carbonate equilibrium system of the seawater. The experimental system was assumed to simulate the conditions in the boundary layer during periods of low water exchange or high intensity irradiations. Fundamental differences were found between different algal classes, suggesting that capabilities to adapt to specific environmental conditions may be connected with dissimilarities in carbon acquisition strategies. In general, green algae were able to reach the highest pH (10.8 at 5°C), and thus to achieve the highest reduction in the level of inorganic carbon via a simple HCO₃ ^–/OH^– ion exchange process. For brown algae, pH increases due to carbon uptake never exceeded pH 9.7 (9.5 in a saltwater scale). In spite of this, members of the Fucaceae (littoral brown algae) were able to extract almost all of the dissolved inorganic carbon (DIC). This was achieved through a gradual decrease in the alkalinity of the enclosed water, so that the carbon assimilation could continue without any concomitant increase in pH. For red algae, the specific response was an increase in the level of inorganic carbon. Thus, for this algal class, no specific strategy for handling a shortage of inorganic carbon was documented. Within each algal class, differences in pH and DIC compensation points could be related to differences in the depths at which the algal species occurred. This paper also introduces a low cost and convenient method of analysing DIC in seawater.     

Flow,flapping, and photosynthesis ofNereocystis leutkeana: a functional comparison of undulate and flat blade morphologies

A number of species of macroalagae possess a flat, strap-like blade morphology in habitats exposed to rapidly-moving water whereas those at protected sites have a wider, undulate blade shape. We have explored the functional consequences of flat, narrow vs. wide, undulate blade morphologies in the giant bull kelpNereocystis luetkeana. Our study focused on the behavior of blades in ambient water currents and the consequences of that behavior to breakage and to photosynthesis. In flowing water, the narrow, flat blades flap with lower amplitude and collapse together into a more streamlined bundle than do wide, undulate blades, and hence experience lower drag per blade area at a given flow velocity. If the algae at current-swept sites had ruffled blades, drag forces would sometimes be sufficient to break the stipes. However, flat blades in a streamlined bundle experience more self-shading than do undulate blades, which remain spread out in water currents. Thus, there is a morphological trade-off between reducing drag and reducing self-shading. Photosynthetic¹⁴C-HCO₃ uptake rates decrease in slow flow when the boundary layer along the blade surface across which diffusion takes place is relatively thick. However, blade flapping, which stirs water near the blade surface, enhances carbon uptake rates in slow water currents for both the undulate and the flat morphologies.     

Hydrodynamics,diffusion-boundary layers and photosynthesis of the seagrasses Thalassia testudinum and Cymodocea nodosa

Photosynthetic rates of aquatic plants frequently increase with increasing current velocities. This is presumably due to a reduction in the thickness of the diffusion boundary-layer which allows for a higher carbon availability on the plant surface. Blades of the seagrasses Thalassia testudinum and Cymodocea nodosa exposed to different current velocities under controlled laboratory conditions, showed increased photosynthetic rates with increasing flow only at low current velocities (expressed as blade friction velocities, u _*). Carbon saturation of photosynthetic processes occurred at a relatively low u _* level (0.25 cm s^-1) for T. testudinum collected from a calm environment compared to C. nodosa (0.64 cm s^-1) collected from a surf zone. No further enhancement of photosynthetic rates was observed at higher u _* levels, suggesting limitations in carbon diffusion through the boundary layer below critical u ^* levels and possible limitations in carbon fixation by the enzymatic system at higher u _* levels. These results, as well as those of previous theoretical studies, assumed the flow on the immediate seagrass-blade surface to be hydrodynamically smooth. The presence of epiphytes and attached debris causes the surface of in situ seagrass blades to be exposed to flows ranging from smooth to rough-turbulent. As a consequence, the boundary-layer thickness on moderately epiphytized blades under medium to high flow-conditions is not continuous, but fluctuates in time and space, enhancing carbon transport. In situ u _* levels measured directly on blades of seagrasses indicate that T. testudinum and C. nodosa can be exposed to conditions under which the boundary layer limits photosynthesis during short periods of time (milliseconds) during low-energy events. As waves cause the thickness of the diffusion boundary-layer to fluctuate constantly, carbon-limiting conditions do not persist for prolonged periods.     

Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

Ammonium excretion in a temperate-reef community by a planktivorous fish,Chromis punctipinnis (Pomacentridae), and potential uptake by young giant kelp,Macrocystis pyrifera (Laminariales)

The blacksmith Chromis punctipinnis, an abundant planktivorous damselfish off southern California, USA, shelters along rocky reefs at night. While sheltered, blacksmiths excrete ammonium that could, in turn, be utilized by nearby benthic macrophytes. Laboratory experiments during the summer and fall of 1983 and 1984 indicate that ammonium excretion at night ranged from 18.1 mol h^-1 by a 8.5 g (dry) fish, to 89.1 mol h^-1 by a 27.3 g fish; excretion rates generally declined throughout the night. Field measurements at night indicate that ammonium concentrations were significantly higher in rocky crevices occupied by blacksmiths than in unoccupied shelters, and the ammonium level in one shelter dropped after a blacksmith was experimentally removed. Young kelp plants (Macrocystis pyrifera) are capable of taking up ammonium at night. Ammonium levels in chambers containing both a blacksmith and a young kelp plant were significantly lower than in chambers containing only a fish, and ammonium levels dropped in ammoniumspiked chambers that contained kelp plants. Nighttime ammonium uptake rates by young kelp plants, which averaged 1.6 mol g^-1 (dry) h^-1, were only slightly lower than those during the day. Daytime excretion by blacksmiths occasionally results in elevated ammonium levels in the water column. On two of six days, ammonium concentrations in midwater foraging aggregations were slightly but significantly higher than in upcurrent controls; since blacksmiths typically aggregate at the incurrent margin of kelp beds, the ammonium is swept downcurrent and may be utilized by large M. pyrifera that extend through the water column. Thus, the activities of blacksmiths may results in the importation of extrinsic, inorganic nitrogen to primary producers on temperate reefs.     

The significance of microbial carbon in the nutrition of the deposit feeding polychaete Nereis succinea

A partial carbon budget was calculated for a population of the deposit feeding polychaete Nereis succinea (Frey and Leuckart) for a North Carolina, USA salt marsh in order to determine if the ingestion and assimilation of microbial carbon was sufficient to meet the carbon requirement. Carbon required by the population was estimated by calculating annual production, 2.1 g C m^-2, and respiration, 9.4 g. There was no net release of dissolved organic carbon. Annual consumption of microbial carbon (as determined by ATP) was estimated to be 5.2 g m^-2. Assimilation efficiency of heterotrophic, detrital microbes was estimated to be 57%. If this value is used for all the microbial carbon, then total assimilation was 3.0 g C m^-2, or about one-fourth the carbon requirement. N. succinea was able to assimilate carbon from sterile plant detritus which suggests that some of the carbon needed to balance the budget may come from direct uptake of the plant substrate. Other possible additional sources of carbon include consumption of meiofauna and uptake of dissolved organic matter.     

Influence of relative water motion on the growth,ammonium uptake and carbon and nitrogen composition of Ulva lactuca (Chlorophyta)

In a series of multifactorial laboratory experiments, Ulva lactuca discs were grown in an apparatus in which they were exposed simultaneously to 3 simulated current speeds (7.5, 15, 22.5 cm s^-1) and a still control, and either 3 ammonium concentrations (0–10, 35–45 and 115–145 M) under ample uniform light (ca 200 E m^-2 s^-1) or 3 light intensities (approximately 35, 90 and 270 E m^-2 s^-1) with uniform surplus, ammonium. Disc growth rates were determined in each experiment as well as tissue nitrogen and carbon composition and fluxes of NH₄, NO₃/NO₂ and PO₄ in media. In a supplementary series of field experiments, U. lactuca discs were simultaneously exposed to 2 different water motion regimes in adjacent chambers at several sites characterized by widely different ammonium concentrations. In field experiments, growth rates were calculated and analyzed as a function of water motion at the various sites. The application of simulated current consistently enhanced disc growth rates in the laboratory, except at the lowest light intensity. In most cases this enhancement was fully realized at the lowest applied simulated current (7.5 cm s^-1). Simulated current slightly enhanced ammonium uptake rates by U. lactuca discs, relative to rates in still water, except at the highest ammonium concentration. C:N ratios of discs generally declined with increases in simulated current, except at the highest ammonium concentration. This decline was primarily attributable to increases in per cent N and was, again, mainly realized at 7.5 cm s^-1. The results suggested that simulated current compensated for N limitation, except when light was sufficiently low to become the overriding limiting factor, but that the enhancement of growth by simulated current could not be explained in terms of N metabolism alone. Field experiments showed that the higher level of water motion consistently enhanced growth at sites with comparatively low ammonium concentrations, but not at sites with moderate or high ammonium concentrations.     

Effects of sediments on the development of Macrocystis pyrifera gametophytes

Rocky ocean bottoms are covered from time to time with small amounts of fine sediments. This material may interfere with the development of germling stages of Macrocystis pyrifera. Tests were performed by introducing sediments before and after spores were dispersed in culture dishes. 10 mg cm^-2 of sediment, enough to occlude the surface, prevents spore attachment, greatly reducing the probability of survival. Smothering of established germlings was severe at 108 mg sediment cm^-2. Water motion further reduced spore success where sediments were present, probably because of abrasive scour. Mechanisms of natural and human interference with M. pyrifera reproduction are suggested.     

Diurnal cycle and kinetics of ammonium assimilation in the green alga <Emphasis Type="Italic">Ulva pertusa</Emphasis>

The kinetics of ammonium assimilation was investigated in Ulva pertusa (Chlorophyceae, Ulvales) from northeastern New Zealand. Ammonium assimilation exhibited Michaelis–Menten kinetics with a maximum rate of assimilation (V _max) of 54 ± 5 μmol g⁻¹ dry weight h⁻¹ and half-saturation constant (K _m) of 23 ± 8 μM. In contrast, values for ammonium uptake were considerably higher with a V _max of 316 ± 59 μmol g⁻¹ dry weight h⁻¹ and K _m of 135 ± 46 μM. At environmentally relevant ammonium concentrations (5 μM), assimilation accounted for most (70%) of the ammonium taken up. Darkness decreased the maximum rate of ammonium assimilation by 83%. We investigated the hypothesis that rates of biosynthetic processes are greater in the early part of the day in Ulva. Consistent with this hypothesis, the maximum rate of ammonium assimilation in U. pertusa peaked in the morning and coincided with low levels of the photosynthetic product sucrose, which peaked in the afternoon. There was a diurnal cycle in the rate of ammonium uptake and assimilation in light and dark, but the amplitude was much greater for assimilation than uptake. Moreover, our data suggest that net ammonium assimilation only occurs during the day in U. pertusa. We suggest that two major roles for diurnal cycles are minimisation of interspecific competition for resources and metabolic costs.     

Effects of simulated current on the growth rate and nitrogen metabolism of Gracilaria tikvahiae (Rhodophyta)

In a series of multifactorial laboratory experiments, Gracilaria tikvahiae apical segments were grown in an apparatus in which they were exposed simultaneously to 3 simulated current speeds (7.5, 15, 22.5 cm s^-1) and a still control, and either 3 ammonium concentrations (<6, 37–39, and 119–136 M) under ample uniform light (ca 200 E m^-2 s^-1) or 3 light intensities (ca 35, 90, and 270 E m^-2 s^-1) with uniform surplus ammonium. Growth rates of apical segments were determined in each experiment as well as nitrogen and carbon composition of tissues and fluxes of NH₄, NO₃/NO₂, and PO₄ in media. In a supplementary series of field experiments, apical segments of G. tikvahiae weresimultaneously exposed to 2 different regimes of water motion in adjacent chambers at several sites characterized by widely different ammonium regimes. The application of simulated current significantly enhanced growth rates in all experiments which utilized recently collected plants. Generally, this enhancement was fully realized at 7.5 cm s^-1, with growth rates tending to plateau above that speed. Growth enhancement by simulated current was independent of ammonium concentration and was considerably reduced at the lowest light intensity. In experiments conducted with plants that had been maintained for several months in aquaria, simulated current failed to enhance growth rates. This suggested that growth responses were at least partly a function of prior conditioning. Growth rates were a direct function of light intensity and an inverse function of ammonium concentration, indicating ammonium inhibition at the higher applied concentrations. Simulated current slightly enhanced rates of ammonium uptake but this did not consistently result in reduced C:N ratios, suggesting that the growth-stimulating effect of relative water motion was attributable to factors other than N uptake. There was evidence of luxury consumption of ammonium. In field experiments, growth rates were not significantly related to exposure to water motion.     

Day-night differences in the emergence of demersal zooplankton from a sand substrate in a kelp forest

Day-night differences in abundance and biomass of demersal zooplankton in the water column were determined by trapping these animals as they emerged from the sand substrate in a kelp forest (Macrocystis pyrifera) ecosystem off Santa Catalina Island, California, USA. The day and night sampling periods of the 24 June 1979 new moon each lasted 12 h. Abundance and biomass of total demersal zooplankton were significantly higher in night samples. A mean of 2,425±1,168 demersal zooplankton m^-2 24 h^-1 migrated over a diel cycle; 97% of these animals were crustaceans. The mean biomass of demersal zooplankton was 94.2±27.6 mg ash-free dry wt m^-2 24 h^-1. No significant differences were found in either the abundance or biomass of demersal zooplanktion collected in low and high traps, suggesting that most animals collected 25 cm off the bottom can sustain swimming to at least 75 cm and that both traps give comparable estimates of the amount of demersal zooplankton available to planktivorous predators.     

Carbon metabolism and strobilation in Cassiopea andromedea (Cnidaria: Scyphozoa): Significance of endosymbiotic dinoflagellates

Scyphopolyps and scyphomedusae of Cassiopea andromeda Forskål (Cnidaria, Scyphozoa) containing dinoflagellate endosymbionts (zooxanthellae) were investigated for rates and pathways of carbon fixation. Photosynthesis by the algae, accounting for 80 and 15 mol C h^-1 on a dry weight basis in medusae and polyps, respectively, by far exceeds dark incorporation of inorganic carbon by the intact association. Photosynthetic carbon fixation is operated via C₃ pathway of carbon reduction. DCMU-treatment (1×10^-6 M and 1×10^-5 M) completely inhibits light-dependent carbon assimilation. Major photosynthates presumably involved in a metabolite flow from algal symbionts to animal tissue are glycerol and glucose. A total of 5–10% net algal photosynthate appears to be seleased in vivo to the host. This is probably less than the energy supply ultimately required for the nutrition of the polyps and medusae. The presence of zooxanthellae proved to be indispensable for strobilation in the scyphopolyps. However, photosynthesis by algal symbionts as well as photosynthate release is obviously not essential for the initiation of ephyrae as is shown by DCMU-treatment, culture in continous darkness, and aposymbiotic controls. It is therefore concluded that strobilation is supported, but not triggered by algal photosynthetic activity. The induction of strobilation thus seems to depend on a more complex system of regulation.     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Nutrients uptake and growth of Ulva lactuca (Linnaeus, 1753) in grey mullet (Mugil cephalus) wastewater versus natural estuarine water

This study investigates the capability of Ulva lactuca to grow in an integrated system, aiming to optimise the needing of resources and to decrease the ecological impact of wastewater. The nutrients uptake and the growth of U. lactuca in Mugil cephalus wastewater (WW) were evaluated and compared with U. lactuca cultivated in estuarine water (EW). Fresh thalli of U. lactuca were cultivated for 10 days in 5?L cylindrical tanks, 3 replicates per treatment. The uptake of dissolved inorganic nitrogen (DIN) and dissolved inorganic phosphorous (DIP), as well as the biomass yield and specific growth rate of U. lactuca, were assessed every two days. At the end of the experiment, U. lactuca resulted in a higher assimilation of DIN in EW (95.7?±?0.3%, mean?±?SE) than in wastewater (68.7?±?1.0%) (p?80%), as well as in the biomass yield and specific growth rate. This study demonstrates the efficiency of U. lactuca in the assimilation of DIN and DIP from M. cephalus WW, contributing to reduce the release of dissolved inorganic nutrients in the natural environment.     

Seasonal growth and composition of fronds of Macrocystis pyrifera in the Falkland Islands

Frond growth of Macrocystis pyrifera in the Falkland Islands was monitored in shallow coastal water from December 1985 to March 1987, and at a different site in deeper water from December 1985 to June 1986. Growth rates in the deeper bed were generally higher than those recorded in the coastal zone. At both sites, node initiation and elongation rate fluctuated according to the seasonal pattern of light or water temperature. In the shallow coastal area, nitrate was abundant in the winter and below detection levels during late spring and summer. Correlation analysis suggests that the production of the fronds of the giant kelp in this area was probably inhibited during the summer months by extremely low concentrations of nutrients. Internal nitrogen was exhausted approximately one month after a sharp decline in ambient nitrate concentration, and carbon reserves were formed. In the deeper bed of M. pyrifera, nitrogen was abundant all year round and the production of the fronds reflected the seasonal pattern of light or water temperature. The nitrogen content of the tissue probably did not drop below a level that limited production, and no internal carbon reserves were accumulated.     

Nitrate uptake and assimilation in Thalassiosira weissflogii and Phaeodactylum tricornutum: interactions with photosynthesis and with the uptake of other ions

Interactions of the nitrate, phosphate, and ammonium uptake systems and the interactions of these systems with photosynthesis were investigated for Thalassiosira weissflogii and Phaeodactylum tricornutum preconditioned in continuous culture. The cultures were supplied with NO _- ³ and PO ₌ ⁴ in an N:P atomic ratio of 15:1, and residual concentrations of both nutrients in the growth chamber were very low. The rate of NO _- ³ uptake was reduced by the addition of NH ₊ ⁴ or PO ₌ ⁴ . The rate of PO ₌ ⁴ uptake by T. weissflogii was reduced by the addition of NH ₊ ⁴ . The rate of carbon fixation was reduced by NO _- ³ additions and slightly reduced by the addition of PO ₌ ⁴ . There were two components of NO _- ³ uptake, one light-dependent and one light-independent. Uptake inhibition by added PO ₌ ⁴ acted on the light-independent component. The change in the C fixation rate due to added NO _- ³ was equal to the rate of NO _- ³ uptake by the light-dependent component on a molar basis. Nitrate assimilation (reduction) rates were calculated from the time course of extracellular and intracellular NO _- ³ concentrations. The light-dark change in the assimilation rate was similar to the light-dark change in the uptake rate, suggesting close coupling between the light-dependent components of uptake and assimilation. The assimilation rate dropped upon exhaustion of extracellular NO _- ³ , implying that an uptake-coupled component of assimilation is unavailable for the assimilation of intracellular NO _- ³ . The reduction in the C fixation rate due to NO _- ³ was temporally associated with uptake rather than assimilation, but may reflect interaction with either the light-dependent uptake step or the closely coupled assimilation. Phosphate additions reduced the rate of NO _- ³ uptake, while the rate of assimilation was unaffected.     

Photosynthesis and amelioration of desiccation in the intertidal saccate alga Colpomenia peregrina

Rates of gross photosynthesis for the intertidal saccate alga Colpomenia peregrina (Sauv.) Hamel were determined under submersed and emersed conditions. Maximal photosynthetic rates were lower than for most seaweeds but comparable with other saccate members of the genus. By fitting the data to a hyperbolic tangent function, maximal photosynthetic rates were estimated to be 5.29 mmol CO₂ m^-2 h^-1 under submersed conditions and 2.06 mmol CO₂ m^-2 h^-1 under emersed conditions. I_k for submersed thalli was 69.1 E m^-2 s^-1, wherea for emersed thalli it was 149.0 E m^-2 s^-1, or 2.2 times higher. At low tide in the field and under saturating irradiance, carbon from seawater retained within the thallus cavity was assimilated at 0.9 mmol CO₂ m^-2 h^-1. In the laboratory under emersed conditions, carbon from this source was taken up at 0.6 mmol CO₂ m^-2 h^-1 at 20°C and at 0.34 mmol CO₂ m^-2 h^-1. Retained seawater also greatly reduced drying under desiccating conditions. Experimental thalli from which seawater had been removed lost thallus water continuously throughout the drying period (120 min). On the other hand, control, thalli lost water for the first 15 min, after which no further water loss occurred. At the termination of the experiment, control thalli had lost 7.2% of their water, whereas experimental thalli had lost 39.2%. Desiccation affected the emersed photosynthetic rate of experimental and control thalli. Emersed photosynthetic rates for thalli dried for 15 min were higher than for fullyhydrated thalli. However, emersed photosynthesis of thalli dried for longer than 15 min was lower than fully-hydrated rates and was directly related to percent water loss. Utilizing data from this study, a model was constructed to determine total photosynthetic production of C. peregrina over a single daylight period. From these calculations it was determined that emersed photosynthesis can increase daily photosynthetic production of C. peregrina by 50%.