Photosynthesis and respiration of exposed salt-marsh fucoids

Photosynthesis and respiration of the salt-marsh fucoids Ascophyllum nodosum ecad scorpioides and Fucus vesiculosus were investigated using an infrared CO₂ gas analyzer under a variety of light intensities, temperatures, and levels of desiccation while the algae were exposed to the atmosphere. Results indicated that net photosynthesis (0.5 to 2.0 mg C/g dry weight/h) saturated rapidly at light intensities (0.1 to 0.2 g cal/cm²/min) which were approximately 10 to 50% of the daily summer maximum intensities for algae found under phanerogam (Spartina alterniflora) canopies. Desiccation exhibited the most pronounced effect on photosynthesis, which increases slightly between 0 and 25% water loss, levels off, and decreases sharply at water losses greater than 50%. Dark respiration (0.1 to 0.3 mg C/g dry weight/h) is also inhibited by desiccation. Both species of algae appear to be broadly adapted to all three parameters investigated.This research was supported by research grants AG-375 and BO 38018 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).Communicated by M.R. Tripp, Newark     

Effect of diurnal variations in natural irradiance on the apical length growth and light saturation of growth in five species of benthic macroalgae

The diurnal variation in apical length growth rate of five benthic macroalgae from the Swedish West Coast (Fucus spiralis L., F. vesiculosus L., F. serratus L., Ascophyllum nodosum (L.) Le Jolis, and Chondrus crispus Stackhouse) was measured during 1.5 to 2.0-h intervals for 3 to 5 d in September 1984. All species showed a maximum in growth rate in the morning, followed by a continuous decline throughout the rest of the day (most pronounced in F. serratus, A. nodosum and C. crispus). By comparing the growth rates with the average irradiance during the same interval, saturation irradiances for length growth in natural daylight regime were estimated; 60 to 70 Wm^-2 for F. spiralis and F. vesiculosus, 90 to 100 Wm^-2 for A. nodosum, 30 to 40 Wm^-2 for F. serratus and 10 to 20 Wm^-2 for C. crispus. Average growth rates during the dark hours ranged from 33% of average day growth rates in A. nodosum to 63% in C. crispus, respectively.     

Competition and facilitation between germlings of<Emphasis Type="Italic"> Ascophyllum nodosum</Emphasis> and<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis>

Competitive interactions between germlings of Ascophyllum nodosum (L) Le Jolis and Fucus vesiculosus L. were studied both in the laboratory and on a shore of the Isle of Man, in the Irish Sea. Both intra- and interspecific competition were investigated by comparing the performance of algal germlings both in monocultures and mixed populations of the two species. The growth of germlings of both species reduced with increasing density. F. vesiculosus always grew faster than Ascophyllum and did best in mixed cultures, whereas Ascophyllum did least well when mixed with Fucus germlings. Clearly the adverse effects of F. vesiculosus on A. nodosum were greater than those of Ascophyllum cohorts. At the same total density, the survival and growth of Ascophyllum declined with an increasing proportion of Fucus germlings, implying that poor recruitment of A. nodosum results from strong competition with F. vesiculosus. However, under desiccation stress on the shore, F. vesiculosus enhanced the survival of A. nodosum at the early germling stage even though competition may occur again at the late stage. Thus, whether interactions between germlings take the form of competition or facilitation depends on the environmental conditions.Communicated by O. Kinne, Oldendorf/Luhe     

Adaptation of light-harvesting pigments to downwelling light and the consequent photosynthetic performance of the eulittoral rockweeds Ascophyllum nodosum and Fucus vesiculosus

We compared the effect of habitat and water depth on the light-harvesting pigment content for Ascophyllum nodosum and Fucus vesiculosus at two near-shore stations in Long Island Sound (USA). Excised pieces of seaweeds were attached at depth intervals to a vertically buoyed line, and left in situ for 7 days. For comparison, fronds were collected from sun and shade habitats in the littoral zone. The three major antenna (light-harvesting) pigments increased in concentration with depth or shade. Chlorophyll c to a ratios remained stable at about 0.2. Fucoxanthin to chlorophyll a ratios decreased by 20 to 30% with depth or shade. Although pigment composition for the two rockweed species was equivalent, the maximum photosynthetic performance of F. vesiculosus exceeded that of A. nodosum by a factor of 2, while the compensation depths for 4 m-adapted A. nodosum and F. vesiculosus under natural limiting light conditions were equivalent. Plants held at 4 m had higher photosynthetic rates compared with plants held at 0 m, no matter the depth of measurement. Indirect evidence indicates that the enhanced photosynthesis of 4 m-adapted plants is due not only to higher concentrations of antenna pigments but to other physiological factors as well. We conclude that the clearly delineated vertical distribution of these two canopy species, the F. vesiculosus zone over the A. nodosum zone, is not determined by light quantity or quality, but by biotic factors as evidenced by the experiments of Menge which are cited herein.     

Food and habitat choice in floating seaweed clumps: the obligate opportunistic nature of the associated macrofauna

The species composition of macrofauna associated with floating seaweed rafts is highly variable and influenced by many factors like spatial and temporal variation, period since detachment and probably also the seaweed species. The presence of seaweed preferences was assessed by a combination of in situ seaweed samplings and multiple-choice aquarium experiments in a controlled environment, using the seaweed-associated grazing organisms Idotea baltica and Gammarus crinicornis. Results from the sampling data confirm that the seaweed composition influences macrofaunal species composition and abundance: samples dominated by Sargassum muticum displayed higher densities but lower diversities compared to samples dominated by Ascophyllum nodosum and Fucus vesiculosus. Seaweed preference was also apparent from the multiple-choice experiments, but did not exactly match the results of the community analysis: (1) I. baltica had high densities in seaweed samples (SWS) dominated by F. vesiculosus and A. nodosum, while in the experiments, this isopod was most frequently associated with Enteromorpha sp. and F. vesiculosus, and fed mostly on S. muticum, A. nodosum and Enteromorpha sp.; (2) G. crinicornis had high densities in SWS dominated by F. vesiculosus, while in the experiments, this amphipod was most frequently associated with S. muticum, but fed most on A. nodosum and F. vesiculosus. It is clear from the laboratory experiments that preference for habitat (shelter) and food can differ among seaweed species. However, food and habitat preferences are hard to assess because grazer preference may change if choices are increased or decreased, if different sizes of grazers are used, or if predators or other grazers are added to the experiments. The effects of seaweed composition may also be blurred due to the obligate opportunistic nature of a lot of the associated macrofaunal species.     

Growth and production of new recruits and adult individuals of Ascophyllum nodosum in a non-harvested population at its southern limit (Galicia,NW Spain)

Populations near the geographic distribution limits of the species are considered to live under suboptimal conditions, and hence, slight environmental changes can be critical for their survival. The potential sensitivity to disturbances of the long-living macroalga Ascophyllum nodosum was analyzed by the determination of growth, recruitment, mortality, and production of biomass of a population near its southern distribution limit. Recruitment, survival and growth rates of <2 years old individuals were determined in a new population growing in experimentally denudated squares. Demographic data for >2 years old individuals were obtained from individuals in the original population after estimating their age from the number of gas bladders in the thallus. Growth and survival were described as continuous nonlinear functions of age applied to the population and were further used to make demography-based production estimates. Recruitment of A. nodosum in denudated substrates seemed to require a previous cover of other macroalgae (as Fucus vesiculosus) as the only cohort detected during the 26-month period of the study was observed after F. vesiculosus individuals started to increase. The low production estimates (2,033 g m^?2 for a 10 year period) and poor recruitment may indicate a slow recuperation of this population to denudation. However, the large variability observed in the estimated growth curves of different populations along this southern distribution area suggests a large influence of local conditions that may help to overcome environmental changes at regional scales.     

Functional divergence in heat shock response following rapid speciation of Fucus spp. in the Baltic Sea

In the Baltic Sea, the broadly distributed brown alga Fucus vesiculosus coexists in sympatry over part of its range (south west Gulf of Bothnia) with the Baltic endemic F. radicans sp. nov, while further north in colder and lower-salinity areas of the Baltic F. radicans occurs alone (north west Gulf of Bothnia). F. radicans appears to have arisen via rapid speciation from F. vesiculosus within the recent history of the Baltic (ca. 7500 BP). Possible functional divergence between the two species was investigated by comparing stress-responsive gene expression in a common-garden experiment. The experiment used two allopatric populations of Fucus vesiculosus from the Skagerrak (North Sea) and Central Baltic, as well as F. radicans from the same Central Baltic site. The two species in sympatry displayed divergent heat shock responses, while F. vesiculosus populations from allopatric sites did not. F. radicans was more sensitive to heat shock at 25°C, either alone or together with high irradiance and desiccation, than Baltic or Skagerrak F. vesiculosus. The results indicate that rapid functional divergence in the inducible heat shock response has occurred between sympatric species on a timescale of thousands of years.     

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     

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.     

Experimental studies on the effects of copper on a marine food chain

Effects of copper sulphate on a marine food chain were investigated in large tanks at Loch Ewe on the west coast of Scotland. The food chain consisted of phytoplankton, the bivalve Tellina tenuis da Costa, and O-group plaice (Pleuronectes platessa L). During initial months after settlement, T. tenuis siphons are an important food for juvenile plaice, which feed by grazing the siphon tips; these subsequently regenerate. Copper dose rates of 10, 30 and 100 μg Cu/1 were investigated; the distribution of added copper was monitored and its metabolic effects were determined. Copper levels in water, sand, in algae on tank walls, in T. tenuis shell and flesh, and in plaice muscle and viscera were measured by atomic absorption spectrophotometry. Copper accumulated in sand, T. tenuis flesh and shell, and plaice viscera. Accumulations were dose-dependent and in no case was a plateau concentration reached. Most of the added dose was taken up by the sand; accumulations in T. tenuis shell were small — less than twice background in all cases. High levels were found in T. tenuis flesh, where concentrations after 100 days were 270, 470 and 1100 μg Cu/g dry flesh for dose concentrations of 10, 30 and 100 μg Cu/1, respectively. The mean control concentration was less than 50 μg Cu/g dry flesh. There was no accumulation of copper in plaice muscle. Visceral accumulations after 100 days were 71, 147 and 567 μg Cu/g dry flesh for dose concentrations of 10, 30 and 100 μg Cu/1, respectively. The mean control level was 30 μg Cu/g dry flesh. The effect of copper on phytoplankton metabolism was investigated by measuring plant pigment levels and rates of primary photosynthetic fixation of C¹⁴-labelled bicarbonate. All dose concentrations reduced both the standing crop and the rate of photosynthesis per unit of chlorophyll a. The effects of copper on growth and condition of T. tenuis and P. platessa were investigated. All dose concentrations adversely affected T. tenuis condition (dry flesh weight for standard individual). The effect was most marked during deposition of winter reserves. In the absence of plaice predation, there was a decrease in the mean siphon weight for T. tenuis exposed to 30 and 100 μg Cu/1. For plaice, all dose concentrations of copper resulted in reduced growth, but there was no significant change in condition or biochemical composition. The ash weight of fish exposed to copper was significantly higher than for controls.     

Improved survival under heat stress in intertidal embryos (<Emphasis Type="Italic">Fucus</Emphasis> spp.) simultaneously exposed to hypersalinity and the effect of parental thermal history

Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT₅₀ (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg^–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT₅₀ under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Zonation,vesicle pressure and gas composition in Fucus vesiculosus and Ascophyllum nodosum at Kristineberg (West Coast of Sweden)

Features of fucoid zonation (Fucus spiralis, F. vesiculosus, Ascophyllum and F. serratus) in a vertical range of 60 cm at Kristineberg (Gullmar Fjord, W. Coast of Sweden) are described. In contrast to identical species occupying areas with wider tidal ranges, these plants are smaller in size and inhabit narrow, overlapping zones. Investigation into the magnitude of pressure and gas composition in vesicles of F. vesiculosus and Ascophyllum nodosum revealed an average positive pressure equivalent to 1.7 and 7 cm of water for the 2 plants in succession. These small values compare favourably with the pressure head to which such vesicles are subjected at Kristineberg. O₂-percentage amounts to 28.5% and 32.7% in vesicles of both plants in succession. O₂-values increase in daytime and decrease at night or when plants are maintained in the dark. CO₂-content in the two plants fluctuates between 0 to 2.6% and bears no correlation to the oxygen values. It is suggested that the pressure produced in the vesicles is due mainly to oxygen. Comparison of these results with those by other authors are made whenever possible.     

Seasonal variations in salt-march macroalgae photosynthesis. II. Fucus vesiculosus and Ulva lactuca

Photosynthesis in whole plants of the salt-marsh algae Fucus vesiculosus and Ulva lactuca was evaluated by ¹⁴C-uptake under a variety of light intensities at approximately mately monthly intervals during a 15-month study. Photosynthetic capacity in both species was closely related to seasonal irradiation patterns and changes in field biomass. Maximum photosynthesis occurred in the spring and summer months. Photosynthesis on a dry weight basis was higher in U. lactuca, while photosynthesis on a chlorophyll a basis was equal in both species. Photosynthetic capacity was inversely related to pigment content. Maximum chlorophyll a concentrations occurred during the winter. Frond profile studies in F. vesiculosus indicated that apices always exhibited greatest photosynthetic capacity. Uptake of ¹⁴C into ethanol-soluble and insoluble fractions was different in each species. F. vesiculosus showed greater activity in the ethanol-soluble fraction while U. lactuca exhibited greater activity in ethanol-insoluble fractions.This research was supported by Research Grant AG-375 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Metabolism and body composition of a copepod (<Emphasis Type="Italic">Neocalanus cristatus</Emphasis>: Crustacea) from the bathypelagic zone of the Oyashio region,western subarctic Pacific

Metabolism [respiratory oxygen consumption, electron-transfer-system (ETS) activity] and body composition [water, ash, carbon (C), nitrogen (N), carbon/nitrogen (C/N) ratio] of stage C5/C6 Neocalanus cristatus from 1000 to 2000 m depth of the Oyashio region, western subarctic Pacific, were determined during the period of July 2000 through June 2003. Compared with the C5 specimens from shallow depths (<250 m), those from 1000 to 2000 m were characterized by quiescent behavior, reduced respiration rates (30% of the rates at active feeding), very low water content (61–70% of wet weight), but high C content (56–64% of dry weight) and C/N ratios (7.2–10.6, by weight). Artifacts due to the recovery of live specimens from the bathypelagic zone appeared to be unlikely in this study, as judged by the consistent results between re-compression (100 atm) and non-compression (1 atm) respiration experiments, and between ETS activities and respiration rates directly measured. In addition, the respiration rates of C6 males and females of N. cristatus from the same 1000–2000 m depth were two to three times higher than the rates of C5 individuals, but were similar to the rates of a bathypelagic copepod, Paraeuchaeta rubra. Combining these results with literature data, C budgets of: (1) diapausing C5 specimens, weighing 6–10 mg dry weight; (2) molt to C6 females; and (3) the complete the life span were established, taking into account assorted losses in respiration during diapause at stages C5 and C6, molt production and egg production. Respiratory C losses by C5 and C6 specimens were estimated on the basis of body N as adjusted metabolic rates [AMR; µl O₂ (mg body N)^–0.843 h^–1], then N budgets were also computed subtracting N lost in the form of cast molts and eggs from the initial stock. Calculations revealed that allocation of the C stock was greatest to egg production (34–57%), followed by respiration (27%) and cast molts (3%), leaving residual C of 13–36% in spent C6 females. The present results for N. cristatus from the North Pacific are compared with those of Calanus spp. in the North Atlantic.Communicated by O. Kinne, Oldendorf/Luhe     

Survival of copepods passing through a nuclear power station on northeastern Long Island Sound,USA

About 70% of the copepods entering the cooling water system of a nuclear power plant on northeastern Long Island Sound (USA) are not returned to the Sound in the effluent. Copepod mortalities are caused by the mechanical or hydraulic stresses of passage, although our experimental design could not determine whether heat or chlorination could cause mortality in the absence of mortality induced by hydraulic stress. After passing through the power plant, copepods sink rapidly (ca. 2.5 times faster than controls). This leads to an increase in concentrations of copepods suspended in the deep water (25 to 30 m) of the effluent pond. About half of the live copepods collected at the discharge and held in situ died within 3.5 days, and 70% died within 5 days, whereas only 10% of those from the intake died in 5 days. About 60% of the copepods observed suspended in deep water in the pond were dead. The copepod mortality caused by the power plant reflects the loss in secondary production occurring below about 270×10³ m² of sea surface in Long Island Sound annually. This loss represents a reduction of about 0.1% in the annual secondary production over a 333 km² area of Long Island Sound adjacent to the power plant. Highest losses occurred during the spring (April, 1.4×10⁶ g dry weight), the lowest in autumn (November, 45.8×10³ g). If the same copepod loss rate exists for all power plants in Long Island Sound, then secondary production in 1.69×10⁶ m², or 0.05% of the total copepod production may be lost annually. A comparison of the surface outflow from Long Island to Block Island Sound with the water entrained through Millstone Unit One, and the 70% copepod loss rate in the latter area, indicates that Unit One eliminates about 0.1 to 0.3% of the copepod production in eastern Long Island Sound. This calculation compares favorably with losses computed from production data.     

A comparison of two cartographic exposure methods using Fucus vesiculosus as an indicator

Morphological variation and vertical distribution of Fucus vesiculosus were quantified at several sites in the Finnish archipelago (Baltic Sea). F. vesiculosus samples were obtained from skerries at geographical distances of 1 km or more (large scale) and at intervals of ca 100 m around a single island (small scale). The results were examined in relation to wave exposure, calculated by Baardseth and effective fetch cartographic methods. Despite the fact that the exposure indices were calculated differently they correlated strongly. Vegetative morphological characteristics of F. vesiculosus illustrate the morphological differences both within and between exposure gradients. The tallest and widest F. vesiculosus plants were found at the sheltered end of the large-scale exposure gradient. Those from equally sheltered sites of the island were smaller in all respects. Thus, the trend from small narrow plants to large wide sheltered plants was expressed differently over the different geographical scales. Consequently localities with similar exposure indices may have morphologically different F. vesiculosus populations. Shores with similar cartographic exposure indices can be different in nature. Underwater topography and shore locations, either close to the mainland or at the outermost sites of the archipelago, affect the exposure. Although a sheltered shore is indicated, the sublittoral zone may be quite exposed to the movements of water. In contrast, in an open shore environment underwater rocks, boulders and shallow water areas can provide sheltered habitats. The depth range of the F. vesiculosus belt exhibited two distinctive patterns. At sheltered sites, around islands in the outermost reaches of the archipelago F. vesiculosus can grow to a maximum depth of 5 m. In exposed habitats the belt becomes narrower, reaching a maximum depth of 3 m. Closer to the mainland F. vesiculosus is found at exposed sites to a maximum depth of 5 m; the depth range at sheltered sites is narrower, only reaching depths of 2 m or less. In conclusion, the changes in plant morphology and in the vertical belt distribution are similar to each other along both gradients at the exposed ends of the wave action spectrum; however, the two gradients diverge at the sheltered ends of the spectrum. Received: 10 August 1998 / Accepted: 11 January 1999     

Egg release and settlement patterns of dioecious and hermaphroditic fucoid algae during the tidal cycle

The timing of gamete release by fucoid algae, although known to be restricted to calm days is not clearly understood within a circadian time scale. The need for externally fertilizing species to avoid gamete dilution suggests that in wave-exposed areas spawning may occur during particular tidal phases. However, this may differ between mating systems, as selfing species may be less affected by gamete dilution. In this study, two different approaches were used to determine when egg release occurs during the tidal cycle in two sister species with different mating systems. First, egg settlement of Fucus vesiculosus (dioecious) and Fucus spiralis (selfing hermaphrodite) was quantified on removable substrates (egg settlement disks) every day for 2 months and settlement patterns were used to statistically estimate the radius of a circle that would encompass 99% of each patch of settled eggs (the egg dispersal radius). Also, egg release was quantified every 2 h during the tidal cycle. A significantly larger egg dispersal radius (P < 0.02) was found for F. spiralis than F. vesiculosus, and this difference was somewhat site dependent with a greater difference between species in exposed sites. The egg dispersal radius was negatively correlated with significant wave height and positively correlated with sea surface temperature for both the species (P < 0.05), with a greater effect of both the factors for F. spiralis than for F. vesiculosus. Egg release during the tidal cycle was variable between species and experiments, with F. vesiculosus releasing more eggs, later in the day, and at a lower tide, than F. spiralis, which released fewer eggs, throughout the day and at all tides. The dioecious species, F. vesiculosus, may have developed a specific adaptation for timing the egg release to periods when emersed in exposed habitats to avoid rapid dilution of gametes that require outcrossing for fertilization. On the other hand, egg release for F. spiralis, which can self-fertilize, occurred both when emersed and immersed, suggesting this species has developed less synchrony with specific environmental factors. Site dependence also suggests local wave conditions can modulate timing of release. The data are consistent with the relaxation of the selective constraints of water motion on fertilization success in a selfing hermaphrodite, relative to an obligate outcrossing species. Results support the idea that species with different mating systems evolve different sensitivities to environmental cues for gamete release with specific implications for inbreeding and successful external fertilization in the ocean.     

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).     

A mathematical model for the uptake of heavy metals in benthic algae

A mathematical model for the uptake of heavy metals in the benthic algae Ascophyllum nodosum has been developed. The model allows assignment of age-dependent growth parameters which also may be a function of external factors. Mortality curves for A. nodosum corresponding to a highly polluted and a nearly unpolluted area have been used in the study of metal uptake. Uptake has been simulated for different values of growth parameters and different concentrations of heavy metals in seawater. Emphasis has been on the uptake of zinc. The simulations showed that the concentration of zinc in the algae changed much less (6%) than the biomass (30%) with changes in intrinsic growth rate, mortality and carrying capacity.The concentration in the algae was found to be an approximately linear function of the mean concentration in seawater up to about 100 ppb. At very high concentrations, associated with high mortality, the deviation from linearity may be significant (about 13% at 162 ppb at one locality). The calculation indicates further that variations in the zinc concentration in the seawater are considerably damped in the algae.Simulations have also been performed to study the effects of sampling different parts of the algae, and some initial simulations have been performed to study the effects of heavy metal exchange between algae and sea water.     

Determination of photosynthetic rates for the marine algae Fucus vesiculosus and Laminaria digitata

A continuously recording, flow-through oxygen electrode system for the measurement of oxygen exchange is described and applied to an investigation of photosynthetic rates in the marine algae Fucus vesiculosus L. and Laminaria digitata (Huds.) Lam. The photosynthetic rate (mg O₂.g dry weight^-1.h^-1) at 15°C and 21.5 mW.cm^-2 (usually just saturating) ranges in F. vesiculosus from 1.20 in basal portions of the thallus to 9.27 at the apices and in L. digitata from 1.19 mg O₂ at the thallus base to 3.97 mg O₂ at distances of several centimetres behind the upper thallus margin. This variation is reduced when the photosynthetic rate is expressed in terms of fresh weight or surface area.This research was carried out while one of us (R.J.K.) was an Alexander von Humboldt fellow at the University of Kiel, and is part of the programme Sonderforschungsbereich 95, Wechselwirkung Meer-Meeresboden, Universität Kiel.