A flow-through system for field measurements of production by marine macroalgae

A continuous flow-through chamber for the in situ measurement of primary production in macrophytic algae (as exemplified by Ulva lactuca) is described. Rapid mixing occurred within the chamber independent of flow rate or amount of algal tissue, so that the complete-mix reactor model of sanitary engineering was closely approximated. Field experiments with the apparatus revealed a rapid response to short-term variations in production rate as well as a close correlation between production and changing light levels. The flowthrough apparatus was compared to a closed system (the traditional bell jar approach); when the water in the latter was mixed, the two methods gave similar results for incubations not longer than several hours. However, during longer incubations, the productivity in the static system was sometimes depressed, possibly due to nutrient depletion or to abnormally high levels of oxygen. Thus, the flow-through system permits reliable measurements of macroalgal production for periods of 12 h or longer.     

Growth rate and carbon affinity ofUlva lactuca under controlled levels of carbon,pH and oxygen

We examined the growth rate (µ) ofUlva lactuca L. (collected from Roskilde Fjord, Denmark in 1987) at different levels of dissolved inorganic carbon (DIC), pH and oxygen in two culture facilities. Growth was faster in Facility A (µ _max ca 0.3 d^–1) than in B (µ _max ca 0.2 d^–1), probably because of more efficient stirring and higher light intensity. The growth-DIC response curve exhibited low half-saturation constant (K _1/2) values (0.35 mM DIC in A, 0.55 mM in B) and growth rates close toµ _max at natural seawater concentration of 2 mM DIC. Growth rate showed a low sensitivity to oxygen over a wide range of DIC and oxygen concentrations. Collectively, the results demonstrated an efficient mechanism for DIC use, unaffected by acclimatization to DIC concentrations between 0.2 and 3 mM. The growth rate decreased little between pH 7.5 and 9 at 2 mM DIC, but steeply above pH 9 approaching zero just above pH 10. The decline of growth at high pH may result from direct pH effects on cell pH, reduced HCO ₃ ^- availability and impaired operation of the carbon uptake process. The growth responses ofU. lactuca to DIC, pH and oxygen resembled those observed in previous short-term photosynthetic experiments. This similarity is probably due to the fast growth ofU. lactuca which means that photosynthetic products are rapidly converted into cell growth. Based on the culture experiments we argue that field plants ofU. lactuca not exposed to stagnant water and DIC depletion are likely to be limited in growth by environmental factors other than DIC (e.g. light and nutrients). Dense mats ofU. lactuca, however, may show reduced growth as a result of DIC depletion, high pH and self-shading.     

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.     

Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

Extracellular organic carbon (EOC) in the genusCarpophyllum (Phaeophyceae): Diel release patterns and EOC lability

Release of extracellular organic carbon (EOC) by the four species of the brown alga genusCarpophyllum was investigated in light : dark incubations of 30 to 40 h duration using a¹⁴C-method. Plants were collected during 1986 in the Hauraki Gulf area, Auckland, New Zealand. InC. maschalocarpum (Turn.) Grev. andC. plumosum (A. Richard) J. Ag. rather low release rates in light (2 to 5% of the photosynthetic rates) were followed by high release rates after 4 to 5 h in darkness (10 to 15% of previous photosynthetic rates). The opposite pattern, with high release in light and low in darkness, was found in experiments withC. angustifolium J. Ag. andC. flexuosum (Esper) Grev. The pattern with substantially higher release in dark than in light is observed for the first time. EOC products were dominated by small molecules (<1000 daltons) in all four species and both in light and darkness. Decomposition experiments exhibited a preferential bacterial utilization of low molecular weight products. Thus, a relative enrichment with larger molecules took place. Bacterial conversion efficiencies of EOC in 5-d experiments averaged 0.24 and 0.09 for products released in light and dark, respectively. The rate constants for decomposition of products released in light were lower than for those released in darkness.     

Spatial variation in dimethylsulfoniopropionate (DMSP) production in <Emphasis Type="Italic">Ulva lactuca</Emphasis> (Chlorophyta) from the Northeast Pacific

Although dimethylsulfoniopropionate (DMSP) has a variety of functions in marine macroalgae including that of a cryoprotectant, an osmolyte, a way to remove excess sulfur and energy, an antioxidant, and an allelopathic precursor, the latter two functions are believed to be the most important in Ulva lactuca L. (=U. fenestrata) in intertidal populations on the coast of Washington state, USA. The present study found significant variation in DMSP concentrations among U. lactuca collected in May 2005 from six sites ranging from 47°54.45′N (Possession Point, Whidbey Island, WA, USA) to 48°30.55′N (Shannon Point Beach, Anacortes, WA, USA), and also among individuals within sites, and among tissues (basal tissues near the holdfast, middle of the blades, and tips). Concentrations ranged from 37 to 224 μmol g⁻¹ fresh mass (FM). In several 10-day experiments between July 2001 and August 2004 with U. lactuca collected from several places on the coast of Washington, the effects of nutrient level (DIN), light intensity and wavelength, and grazing by the herbivorous gastropod Lacuna vincta, were examined. None of these manipulations resulted in DMSP concentrations that differed significantly from controls, and variance in DMSP concentrations within each experiment was very low. Although DMSP concentrations in U. lactuca may be affected by factors not tested in these experiments, it is also possible that the observed spatial differences reflect constitutive genotypic or phenotypic differences among geographically separated U. lactuca populations or among cryptic Ulva species. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Night metabolism of recent photosynthate by sea ice algae in the high Arctic

Changes in intracellular distribution of recent (labelled) photosynthate during light-dark incubations were consistent with overnight consumption of low molecular weight compounds and carbohydrate, but continued protein synthesis, in sea ice algae from Resolute Passage, Canada, during April and May, 1986. Synthesis of labelled protein at night varied with preceding light availability but was always less than 30% of the daytime rate. Comparisons of labelled photosynthate dynamics against oxygen consumption and net changes of particulate matter composition and concentration showed that soluble polysaccharide was not the only major metabolic substrate, and that much of the carbon lost overnight was not recent, labelled photosynthate. Total net lipid synthesis was greatly underestimated by labelled photosynthate allocation. However, overnight consumption of the labelled lipid was directly proportional to photon flux density in the preceding light period, suggesting a short-term energy storage function for the small labelled portion of the total lipid pool. Arctic ice algae appear to incorporate only limited amounts of recent photosynthate into their large lipid pools even over a full 24 h photoperiod.     

Gas-exchange studies with the staghorn coral Acropora acuminata and its zooxanthellae

Compensation point and light-saturation values were determined from oxygen-exchange experiments with branches and isolated zooxanthellae from the staghorn coral Acropora acuminata. Branches and dense suspensions of zooxanthellae showed similar lightresponse curves for oxygen exchange, with light saturation at about 23 Klux (300 W. m^-2) and compensation point occurring between 4 and 6 Klux (60–80 W. m^-2). Zooxanthellae appear to be mutually shaded in dense suspensions and coral tissues. The effects of metabolic inhibitors, including photosynthetic and respiratory inhibitiors, on oxygen exchange in coral branches and isolated zooxanthellae are presented. Bubbles formed on coral tissues and on several macroalgae under conditions of high illumination contained large amounts of oxygen, suggesting that a high oxygen tension may occur in coral tissues during the day. Photorespiration and dissolved organic carbon production by suspensions of zooxanthellae are discussed in relation to a high oxygen tension which probably occurs in coral tissues during daylight.     

Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3–28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2–1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

Interactive effects of nitrogen and dissolved inorganic carbon on photosynthesis,growth, and ammonium uptake of the macroalgae Cladophora vagabunda and Gracilaria tikvahiae

Dissolved inorganic carbon (DIC) is rarely considered limiting for macroalgae, but some research suggests that under conditions of N sufficiency, photosynthetic capacity is enhanced with DIC enrichment. During spring (April–May) and summer (July–August) 1993, we investigated the interactive effects of nitrogen (N) and DIC on photosynthetic capacity, growth, and nutrient uptake rates of the macroalgae, Cladophora vagabunda (L.) van den Hoek and Gracilaria tikvahiae (McLachlan), dominant species in a temperate eutrophic estuary (Cape Cod, Massachusetts, USA). Water-column CO₂ concentrations showed significant diurnal fluctuations, ranging from a morning CO₂ peak (21 M) to an afternoon low (13 M) during summer, probably associated with metabolic activities in a thick algal mat. Results from instantaneous photosynthesis measurements and microcosm experiments indicate that DIC limits photosynthetic capacity and growth rates of C. vagabunda during summer, perhaps related to tissue N sufficiency and low water-column CO₂ concentrations. For example, this species showed enhanced growth (F=8.69, P<0.02) under DIC but not N enrichment. G. tikvahiae showed marginal DIC enhancement of maximum photosynthetic rate, while growth was significantly stimulated by addition of N. Reduced thallus N of this species during the summer further identifies N as the primary factor limiting growth. In addition, G. tikvahiae has the ability to use DIC in its several forms, while C. vagabunda primarily uses dissolved CO₂. DIC enrichment resulted in a depression of NH₄ ⁺ uptake rates for both species, particularly during summer at saturating (60 M) ammonium levels, suggesting competition between NH₄ ⁺ uptake and DIC acquisition under conditions of N sufficiency. Dominance of C. vagabunda and G. tikvahiae in areas undergoing eutrophication has been attributed to their successful procurement and storage of N as well as to high growth rates. The present study revealed that under conditions of N sufficiency during summer, DIC may control rates of production of these opportunistic macroalgae.     

Amino acid uptake by the comatulid crinoid Cenometra bella (Echinodermata) following evisceration

Changes in the rates of utilization of dissolved compounds during the period of visceral mass regeneration were examined in the crinoid Cenometra bella (Hartlaub). Rates of respiraton and incorporation of labelled amino acids increase, reaching a maximum 2 d after evisceration and returning to normal after 14 d. Rates of incorporation of radioisotope into the organic components of the arms and cirri decrease, while incorporation rate into the visceral mass increases. Incorporation rates of amino acid-derived radioactivity into skeletal carbonate and the ash-free dry weight:protein ratio of arms, cirri and oral disc decrease, reaching a minimum 2 d following evisceration.     

Survival,metabolism and growth of Ulva lactuca under winter conditions: a laboratory study of bottlenecks in the life cycle

Photosynthesis and growth in low light and survival under simulated winter conditions were studied in the freefloating green alga Ulva lactuca L., collected in Roskilde Fjord, Denmark during late autumn and maintained in stock in natural water. It adapts efficiently to low light by increasing chlorophyll concentration and light absorption and continues to grow at the lowest irradiance tested, 0.6 E m^-2 s^-1. This irradiance corresponds to minimum light requirements of deep-living marine macroalgae and phytoplankton growing under ice. The photosynthetic efficiency per unit of incident light is five-fold higher for U. lactuca grown at 1.7 E PAR m^-2 s^-1 as compared with 56.3 E m^-2 s^-1, and the efficiency per unit of light absorbed is twice as high. The maximum photosynthetic efficiency (0.051 mol C E^-1 absorbed) is similar to values for shade-adapted marine phytoplankton. U. lactuca is able to survive for two months in the dark and to resume growth immediately when transferred to light. Exposure to anoxia and sulphide gradually reduces vitality, but does not affect survival over two months. Rigorous deep freezing is detrimental to survival of U. lactuca, while field samples show that more gradual, natural freezing is not. U. lactuca is not easily fitted into one of the traditional strategy concepts. U. lactuca is a very plastic species that combines rapid growth during favourable periods (opportunism) with high survival capacity in the same type of tissue during stress periods (persistence). U. lactuca occupies a niche as a free-floating form in sedimentary coastal areas that are devoid of attached algae.     

Uptake of inorganic carbon and internal carbon cycling in symbiont-bearing benthonic foraminifera

Incorporation rates of inorganic carbon and its distribution between the organic matter and the skeleton have been measured using ¹⁴C tracer techniques on two species of symbiont-bearing benthonic foraminifera in the Gulf of Elat: Amphistegina lobifera (a perforate species) and Amphisorus hemprichii (an imperforate species). Under constant experimental conditions, incorporation rates of the radiotracer become linear with time after several hours in A. hemprichii and after one day in A. lobifera. A. lobifera showed a lag time of 24 h for skeletal incorporation, whereas in A. hemprichii uptake into the skeleton started within 2 h. Pulse-chase incubations in radioactive seawater, followed by unlabelled incubations, demonstrate transfer of photosynthetically acquired ¹⁴C into the skeleton of A. lobifera. No such transfer was found in A. hemprichii. The total ¹⁴C uptake by A. lobifera increased during the first 24 h of cold chase incubation. This increase suggests the existence of an internal inorganic carbon pool that was lost (probably evaporated) during the analysis of pulse incubations. However, during the following chase incubations, the ¹⁴C in this pool was incorporated mainly into the skeleton and retained during analysis, causing the increase in the total uptake. No such increase was found in A. hemprichii. Additional ¹⁴C uptake experiments on other species of the genera Operculina, Heterostegina and Borelis suggest that the differences in pathways for incorporation of carbon between A. lobifera and A. hemprichii can be generalized to the perforate and imperforate foraminiferal groups. In perforate species, respired carbon originally taken up through photosynthesis is partly recycled into the skeleton. In imperforate species such a transfer has not been demonstrated. Perforate species seem to have a large internal inorganic carbon pool which serves mainly for calcification and possibly also for photosynthesis, while imperforate species may take up carbon for calcification directly from seawater or have a very small inorganic carbon pool.     

Decomposition of urea associated with photosynthesis of phytoplankton in coastal waters

Decomposition of urea in seawater was studied in Mikawa Bay, a shallow eutrophic bay on the southern coast of central Japan. The urea concentration in seawater ranged from 1.3 to 5.9 μg-at. N/1 and comprised 12 to 40% of the dissolved organic nitrogen. Using ¹⁴C labelled urea, the rate of CO₂ liberation from urea and the incorporation rate of urea carbon into the particulate organic matter were determined. For the surface samples, high rates of CO₂ liberation from urea as well as the incorporation of urea carbon into the particulate organic matter were observed in the light, while much lower rates were obtained in the dark. Incubation experiments with exposure to different light intensities revealed that the rate of CO₂ liberation from urea and the incorporation of urea carbon into particulate organic matter changed with light intensity, showing a pattern similar to that of photosynthesis. The highest liberation and incorporation rates were observed at 12,000 lux. Incubation in light and in dark produced marked decreases and increases, respectively, in urea and ammonia, while no appreciable changes were observed for nitrate and nitrite. It is suggested that urea decomposition associated with photosynthetic activity of phytoplankton is one of the major processes of urea decomposition, and that it plays a significant role in the nitrogen supply for phytoplankton in coastal waters.     

Phytoplankton exudate release as an energy source for the growth of pelagic bacteria

The release of dissolved organic carbon (DOC) from phytolankton during photosynthesis, and the utilization of this carbon by planktonic bacteria, was studied using ¹⁴CO₂ and selective filtration. Natural sea water samples from a coastal area of the Northern Baltic Sea were incubated in the laboratory for detailed studies, and in situ for estimation of annual dynamics. In a laboratory incubation (at +1°C) the concentration of ¹⁴C-labelled dissolved organic carbon increased for about 2 h and then reached a steady state, representing about 0. 1% of the total DOC. Labelled organic carbon in the phytoplankton and bacterial fractions continued to increase almost linearly. The continuous increase in the bacterial fraction is thought to represent almost instantaneous utilization of the DOC released from the phytoplankton during photosynthesis. As an annual average, in 4 h in situ incubations, about 65% of the labelled organic carbon was found in the phytoplankton fraction (>3 m), about 27% in the bacterial fraction (0.2 to 3 m) and the remaining 8% as DOC (<0.2 m). Large variations in these percentages were recorded. The measured annual primary production was 93 g C m^-2 (March to December), and the estimated bacterial production due to phytoplankton exudates 29 g C m^-2. This represents a release of DOC of about 45% of the corrected annual primary production of 110 g C m^-2 (assuming a bacterial growth efficiency of 0.6).     

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.     

Primary productivity and particulate organic matter flow in an estuarine mangrove-wetland in Hong Kong

Net primary productivity and organic matter flow of a mangrove-dominated wetland was estimated by following production and detritus dynamics in a tidal pond in north west Hong Kong in 1986–1988 (9.1 ha). Total productivity was 12.47 t dry wt ha^–1 yr^–1, of which >90% was from emergent macrophytes (the mangroveKandelia candel and the reedPhragmites communis). High turbidity and high summer temperatures probably limited respective production by phytoplankton and benthic macroalgae (dominated byEnteromorpha crinata). Despite the high total productivity, little detritus was exported from the emergent macrophyte stands, due to the low inundation frequency. This created a net water column carbon deficit which was provided for by the high organic matter import (mean = 4.42 g ash free dry wt m^–2 d^–1) from the incoming water. This same sediment and particulate organic carbon input giving a high accretion rate of 1.7 cm yr^–1 was probably also the force behind progressive dis-coupling of emergent macrophyte production from water column consumers. This resulted in a tendency to retain production in the emergent macrophyte stands while the water column community increasingly relied on allochthonous carbon. This shift from a net exporter to a net importer of carbon in landward wetlands is probably characteristic of the transition into nutrient-conservative terrestrial systems.     

The effect of temperature on leaching and subsequent decomposition of dissolved carbon from inundated floodplain litter: implications for the generation of hypoxic blackwater in lowland floodplain rivers

Dissolved organic carbon export from floodplain litter during flood events is an important energy subsidy for lowland rivers. Temperature affects both the release and decomposition of dissolved organic carbon from floodplain plant litter. Unseasonally warm flood conditions have resulted in the release and consumption of carbon at a rate so rapid that water column oxygen has been depleted, causing the death of aquatic organisms upon exposure to this ‘hypoxic blackwater’. To date, there has not been a systematic investigation of the role of temperature on hypoxic blackwater dynamics. In a series of laboratory experiments, we investigated the temperature dependence of both leaching and decomposition of carbon from a common floodplain litter component in south-eastern Australia – the leaves of the river red gum Eucalyptus camaldulensis. The leaching rate increased with temperature and approximated Arrhenius kinetics. Additionally, the solubility of the leaf carbon increased substantially at temperatures above ～25°C. The rate of organic carbon respiration also increased with temperature, and the relationship with temperature was approximately linear. These data can be used to improve models for the prediction of hypoxic blackwater risk.     

Release of dissolved organic carbon from the estuarine intertidal macroalga Enteromorpha prolifera

The estuarine macroalga Enteromorpha prolifera was collected from Coos Bay, Oregon, USA during 1981, and its release of photosynthate as dissolved organic carbon (DOC) was studied using ¹⁴C as a tracer. During photosynthesis in 30 S sea water, with a fixation rate averaging 7.37 mg C g^-1 dry wt h^-1, release ranged from 0.13 to 0.57 mg C g^-1 dry wt h^-1 and from 1.65 to 6.23% of total fixed carbon. Release of DOC appears to be linear with time over 3 h. As exposed algae become increasingly desiccated, their photosynthetic rates decline dramatically, but upon reimmersion the highly desiccated algae lose a larger fraction of their fixed carbon than the slightly desiccated algae. This loss comes in a pulse release of DOC over the initial 15 min, followed by declining release rates. The pulse loss due to rainfall is 5 times greater than that due to tidal resubmergence, and may briefly exceed the prior photosynthetic rate. Although lowering the salinity from 30 to 5 does not substantially alter photosynthetic rates, it does increase the DOC release range up to 1.02 mg C g^-1 dry wt h^-1 and 16.10% of fixed carbon. Heterotrophic microbes from the algal habitat readily use the available DOC at about 15% h^-1.     

Photosynthetic characteristics of giant kelp (Macrocystis pyrifera) determined in situ

Rates of net photosynthesis and nocturnal respiration by individual blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, were determined in situ by measuring oxygen production in polyethylene bags during spring/summer of 1983. Mature blades from different depths in the water column exhibited different photosynthetic characteristics. Blades from the surface canopy (0 to 1 m depth) exhibited higher photosynthetic capacity under saturating irradiance and higher photosynthetic efficiency at low irradiances than blades from 3 to 5 or 7 to 9 m depths. Saturating irradiance was lower for canopy blades than for deeper blades. Canopy blades showed no short-term photoinhibition, but photosynthetic rates of deeper blades were significantly reduced during 1 to 2 h incubations at high irradiances. Results of 1 to 2 wk acclimation experiments indicated that differences between photosynthetic characteristics of blades from different depths were primarily attributable to acclimation light conditions. Vertical displacement of blades within the kelp canopy occurred on a time-scale of 1 min to 1 h. Blades continually moved between the unshaded surface layer and deeper, shaded layers. Vertical movement did not maximize photosynthesis by individual blades; only a small proportion of blades making up a dense surface canopy maintained light-saturated photosynthetic rates during midday incubations. The relatively high photosynthetic rates exhibited by canopy blades over the entire range of light conditions probably resulted from acclimation to intermittent high and low irradiances, a consequence of vertical displacement. Vertical displacement also reduced the afternoon depression in photosynthesis of individual canopy blades. The overall effect of vertical displacement was optimization of total net photosynthesis by the kelp canopy and, therefore, optimization of whole-plant production.