Optimal growth and maximal survival temperatures of Atlantic Laminaria species (Phaeophyta) in culture

The effects of temperature on growth rate of rapidly-growing cultured macrosporophytes of 9 isolates of Atlantic Laminaria comprising 4 species have been investigated. No significant population variation was observed within species despite wide variations in temperature between the original collecting sites. L. saccharina showed a broad temperature optimum in the 10°–15°C range, whereas L. longicruris had a sharp optimum at 10°C. L. digitata and L. hyperborea grew more slowly, with only slightly sub-optimal growth over a wide temperature range, but with peaks at 10°C (L. digitata) and 15°C (L. hyperborea). The maximum survival temperatures of individual male and female vegetatively-growing gametophytes were ascertained for these species plus the Arctic L. solidungula, and were as follows: L. saccharina and L. longicruris, 23°C; L. digitata (male), 23°C; L. digitata (female), 22°C; L. hyperborea, 21°C; L. solidungula, 18°C. The lack of within-species differences demonstrates that the success of the genus in areas with different temperature regimes is brought about by phenotypic plasticity of individuals rather than the selection of temperature races or ecotypes.     

Transversal profiles of carbon assimilation in the fronds of three Laminaria species

The brown macroalgae Laminaria digitata (Huds.) Lamour., L. hyperborea (Gunn.) Foslie, and L. saccharina (L.) Lamour. (Laminariales, Phaeophyceae) have been investigated for carbon assimilation and assimilate biosynthesis in different tissues (meristoderm, cortex, medulla) of stipes and blades. Carboxylation via ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) and phosphoenolpyruvate carboxykinase (EC 4.1.32) follows a transversal gradient from the outer to the inner tissues with maximum activity of enzymatic CO₂ fixation in the meristoderm. Such transversal profiles are also recorded for photosynthetic and dark carbon assimilation as well as for kinetics of assimilate ¹⁴C-labelling in isolated tissues. Differential chemical constitution of the blade and stipe tissues examined and results from assimilate biosynthesis are discussed with regard to a transversal translocation of photosynthates from meristodermal to cortical and medullar cells.     

Influence of ultraviolet-radiation on chlorophyll fluorescence and growth in different life-history stages of three species of Laminaria (phaeophyta)

Zoospores, gametophytes, young sporophytes and discs cut from mature sporophytes of Laminaria digitata, L. hyperborea and L. saccharina were exposed in the laboratory to UV-radiation, with a spectral composition and irradiance similar to natural sunlight, for periods ranging from 15 min to 8 d, and were then returned to white light. Germination of zoospores and the growth of gametophytes were reduced after exposures to UV longer than 1 h, whereas UV had little effect on the growth of young or mature sporophytes unless exposure continued for more than 48 h. The variable fluorescence (F _v:F_m) of all stages was strongly reduced immediately after short exposures to UV, but recovered almost completely within 24 h. However, exposure of gametophytes to UV for >4 h resulted in little or no recovery of F _v:F_m, whereas >16 h of UV were required to produce this result in young sporophytes, and >48 h in mature sporophytes. Thus, sensitivity to UV-radiation decreased from gametophytes to sporophytes, and with increasing age of sporophytes, but, in gametophytes, growth appeared to be a more sensitive indicator of UV-damage than F _v:F_m after 24 h recovery. The responses to UV of the zoospores and gametophytes of all three species were similar, but both growth and fluorescence measurements suggested that the sporophytes of L. saccharina were more sensitive to UV than those of the other two species.     

Growth and DNA damage in young Laminaria sporophytes exposed to ultraviolet radiation: implication for depth zonation of kelps on Helgoland (North Sea)

Growth as an integrative parameter of all physiological processes was measured in young sporophytes of temperate Laminaria digitata, Laminaria saccharina and Laminaria hyperborea exposed in the laboratory to irradiance consisting of either only photosynthetically active radiation (PAR) or to a spectrum including ultraviolet radiation (UVR) (PAR+UVA+UVB) by use of cut-off glass filters. Size increment was measured every 10 min over a period of 18–21 days using growth chambers with on-line video measuring technique. In the chamber, plants were grown at 10±2°C and 16:8 h light–dark cycles with 6 h additional UVR exposure in the middle of the light period. Tissue morphology and absorption spectra were measured in untreated young sporophytes while chlorophyll a content and DNA damage were measured in treated thalli at the end of the experiment. Sensitivity of growth under UVR was found to be related to the observed upper depth distribution limit of the upper sublittoral L. digitata, upper to mid sublittoral L. saccharina and lower sublittoral L. hyperborea. Tissue DNA damage is, however, dependent on thallus thickness which minimizes UVR effect where outer cell layers shade inner cells and provide longer pathlength for UVR. Exposure to UVR causes cellular, enzymatic and molecular damage. Presence of UV-absorbing compounds further reduces effective UVR from reaching physiological targets. The cost of producing higher amount of UV-absorbing compounds and effective DNA repair mechanism can, however, divert photosynthate at the expense of growth. Tissue chlorophyll a content was not significantly different between treatments suggesting a capacity for acclimation to moderate UVR fluence. Growth acclimation to repeated UVR exposure was observed within a period of 12 days while growth inhibition was observed after a longer UVR exposure period of 21 days. The results give further insight into the effects of UVR on the cellular level and show how ecological parameters such as the upper depth distribution limit are dependent on cellular processes.     

A survey of translocation in laminariales (Phaeophyceae)

A survey of translocation of photoassimilates in 13 genera of Laminariales is presented. All showed long-distance transport of ¹⁴C-labeled products from mature source tissue to meristematic sinks (haptera and intercalary growing regions). In plants with several laminae forming one frond, older laminae may provide assimilates for the growth of younger ones, and in Macrocystis spp., where fronds of different ages and developmental stage arise from a common holdfast, mature fronds initiate and support new fronds. Translocation velocities vary from species to species but are in the range of 55 to 570 mm/h. The results strongly support the hypothesis that Laminariales in general have an effective translocation system, on which their thallus growth depends.     

Standing crop and leaf area index of the sublittoral Laminaria species near Helgoland

Standing crop values of Laminaria species sampled between 0 to 2 m depth in the sublittoral of Helgoland, a small rocky island in the southern North Sea, are similar to those reported by other workers from Norwegian or Scottish coasts. Below 2 m water depth, however, standing crop values are found to be reduced to a far greater extent with increasing depth than in the Norwegian or Scottish areas studied. Maximum standing crop values are not found in Laminaria digitata and L. saccharina growing just below the low water mark of spring tides, but in the upper L. hyperborea forest at 2 m depth. Population density and leaf area index (LAI) decrease with increasing depth (but not the average weight of individuals constituting the L. hyperborea forest). It is suggested that the depth at which LAI becomes 1 represents an important ecological threshold, since growth rates of L. hyperborea specimens appear to decrease at greater depths but not to increase above the threshold. Self-shading might equalize, to a certain extent, the quantity of underwater irradiance available to an individual phylloid at all depths at which a close vegetation of Laminaria species occurs.     

Growth of amputated and dark-exposed individuals of the brown alga Laminaria hyperborea

The seasonal growth of the brown alga Laminaria hyperborea (Gunn.) Foslie, which forms dense forests in the sublittoral zone of Helgoland, a rocky island in the Southern North Sea, was investigated by transplanting specimens of medium size onto PVC or wooden plates fixed to sub-tidal grwoth stations. The 2-year study revealed that, during the season of fast growth (January to June), young plants produce each year a new frond, larger in size than in the preceding year. This observation can be interpreted, in part, by assuming that the growth of the young frond is accomplished not only by its own assimilatory surplus, but also by reserve materials of the old frond assimilated during the preceding season of slow growth. This assumption is supported by experimental evidence: (1) Even in complete darkness Laminaria hyperborea is capable of producing a small new frond during the season of fast growth; (2) New fronds with stipe, but without old frond, grow (during the same period) considerably faster than isolated new fronds separated from the possible sources of reserve materials. Maximum growth occurs in normal plants which possess their old frond until April or May.An abbreviated version of this paper has been presented at the Sixth International Seaweed Symposium, held at Santiago de Compostela, Spain, September 9 to 13, 1968.     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Nitrogen supply,tissue composition and frond growth rates for Macrocystis pyrifera off the coast of southern California

Seasonal changes in ambient NO ₃ ^– and NH ₄ ⁺ , tissue composition (N, C, and C/N ratio), and frond growth rates for Macrocystis pyrifera (L.) Agardh were examined. Ambient NO ₃ ^– showed distinct seasonal variations. Frond growth rates were variable, but showed no clear correlation to ambient NO ₃ ^– . The average N content of plant tissue did, however, show the same seasonal variations as ambient NO ₃ ^– . The longitudinal distribution of total tissue N and various components of tissue N along fronds were also analyzed. Several distinct patterns were found: high levels of protein N at growing tips and elevated levels of soluble N in lower parts of the frond. Free amino acids accounted for a major portion of the soluble N, but neither NO ₃ ^– nor NH ₄ ⁺ accumulated in the plant tissue. The longitudinal distribution of N along the fronds is compared to reported variations in C metabolism, and it is concluded that C and N sourcesink relations do not always coincide and bidirectional translocation may occur.     

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.     

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

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

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Ecotypic differentiation in light-related traits of the kelp Laminaria saccharina

Light-related traits were compared for Laminaria saccharina Lamour. collected from three habitats in Maine and New York, USA, with different ambient light regimes. Light-level, expressed as a proportion of surface irradiance (I₀), ranged from 0.04 to 0.32 I₀ in the shallow habitat, but rarely exceeded 0.04 I₀ in the deep and turbid habitats. Juvenile sporophytes collected from each habitat in April, 1985, were grown at four acclimation light-levels (0.065, 0.12, 0.26, and 0.54 I₀) in a common-garden, laboratory experiment. Photosynthesis vs irradiance (PI) parameters, light-harvesting characteristics, and rates of carbon-assimilation and growth were determined for each group of plants. The results indicated that ecotypic differentiation had occurred among the three kelp populations. Photosynthetic capacity (P_max) and photosynthetic efficiency () were generally highest for plants from the turbid habitat, lowest for deep plants, and intermediate for shallow plants. These differences were largely attributable to variations in light-harvesting characteristics. The nature and magnitude of photoacclimation responses also differed among populations. Population differences in photosynthetic parameters resulted in different rates of C-assimilation and growth by plants from shallow, deep, and turbid habitats. Predictions of in situ growth rates indicated that the severity of light-limitation and, therefore, the adaptive significance of efficient light-utilization vary among the three populations. It is concluded that ecotypic differentiation in light-related traits is important to the broad environmetal range of L. saccharina.     

Regulation of growth in Laminaria digitata: use of in-vivo nitrate reductase activities as an indicator of nitrogen limitation in field populations of Laminaria spp.

The seasonal variation in growth rate of a population of Laminaria digitata (Huds.) Lamour growing at Arbroath, Scotland was studied between August 1981 and September 1982, and was found to follow the biphasic annual cycle typical of this genus. Growth rates were maximum (0.3 cm cm^-1 mo^-1) in early June and minimum (0.05 cm cm^-1 mo^-1) between September and January. An analysis of the relationship between the seasonal changes in environmental factors (inorganic nitrogen concentrations, irradiance and temperature) with those of growth rate and the accumulation or mobilisation of cellular reserves of carbohydrates and nitrate, indicated that growth was nitrogen-limited between June and October and light-limited (with a possible co-involvement of temperature) for the remainder of the year. These conclusions were supported by the seasonal changes in the ratio of actual: potential in-vivo nitrate reductase activities in L. digitata, thus confirming the suitability of this technique for monitoring the occurrence of nitrogen limitation in Laminaria spp. The seasonal changes in blade nitrate reductase activities closely followed those of growth rate, with maximum activities [0.3 mol NO ₃ ^- reduced g^-1 (wet wt) h^-1] being present in late May and minimum levels [0.01 mol NO ₃ ^- reduced g^-1 (wet wt) h^-1] occurring between November and March. The correlation observed between nitrate reductase activities and growth rate is consistent with the ability of Laminaria spp. to store excess inorganic nitrogen, available during winter and early spring, as NO ₃ ^- , and with the requirement to conserve enzyme protein during the summer period of nitrogen limitation.     

Effects of ethyl 2-methyl acetoacetate (EMA) on the growth of Phaeodactylum tricornutum and Skeletonema costatum

We investigated the effects of ethyl 2-methyl acetoacetate (EMA) on growth of the marine diatom algae Phaeodactylum tricornutum (P. tricornutum) and Skeletonema costatum (S. costatum). Growth of P. tricornutum was significantly inhibited by the minimum concentration (3.5 mmol·L ^?1) of EMA at lower initial algal densities (IADs) (3.6×10⁴ and 3.3×10⁵ cells·mL ^?1). However, at the highest IAD, significant growth inhibition was found at above 7 mmol·L ^?1 of EMA exposure. In S. costatum, EMA concentrations of 10.5 mmol·L ^?1 or more significantly inhibited growth at lower IAD (3×10⁴ and 1.8×10⁵ cells·mL ^?1); at the highest IAD, only EMA concentrations above 14 mmol·L ^?1 obviously inhibited the growth of S. costatum. Changes in specific growth rates and pigment were consistent with algal growth, but only at higher EMA concentrations or lower IAD values was the ratio of chlorophyll a (Chla) to carotenoid significantly lower than the control. Medium effective concentration (EC ₅₀) values were in the order 4.07, 8.03 and 12.27 mmol·L ^?1 for P. tricornutum and 7.48, 11.92 and 17.22 mmol·L ^?1 for S. costatum. All these results show that the effect of EMA on the growth of algae was species specific and mainly depended on IAD, which might be an important factor to influence algal growth.     

Seasonal effects of light,temperature, nutrient concentration and salinity on the physiology and growth of Caulerpa paspaloides (Chlorophyceae)

Caulerpa paspaloides (Bory) Greville were collected during the winter and summer (1978 to 1979) from the Florida Keys, USA. Thalli collected during the winter photosynthesized more efficiently at low light intensities (I_c<1, I_k=38 Exm^-2xs^-1) than did thalli collected in the summer (I_c=13, I_k=111 Exm^-2xs^-1). Summer thalli exhibited higher P_max values (2.20 mgO₂xg^-1 dry wtxh^-1) than winter thalli (1.70 mg O₂xg^-1 dry wtxh^-1). Rates of rhizome elongation and frond initiation were strongly inhibited by winter temperatures. The maximum lethal temperature for summer thalli was 37° to 38°C as measured by both growth and photosynthesis. No evidence of nitrogen or phosphorus limitation was found. Relatively minor reductions in salinity (3S) resulted in significant increases in rhizome apex motality. Results indicate that low winter temperatures are responsible for reduced winter growth rates previously reported for the Key Largo population. Increased photosynthetic efficiency at low light intensities and preferential maintenance of rhizome elongation over frond initiation appear to allow this tropical macroalga to optimize growth and survival during the winter.     

Filtration rate,using a new indirect technique,in thirteen species of suspension-feeding bivalves

A method for determining filtration rates in undisturbed suspension-feeding bivalves is described. Concentrations of particulate matter in the water collected in the inhalant (C _i) and exhalant (C _e) currents were estimated with an electronic particle counter. The clearance was calculated as , where Fl=flow rate through the tube collecting exhalant water. Only above critical levels of water flow (Fl) were clearances representative of filtration rates. At 10° to 13°C, the filtration rates (F, 1 h^-1) within one or two orders of magnitude of dry weight (w, g), in Cardium echinatum L., C. edule L., Mytilus edulis L., Modiolus modiolus (L.) and Arctica islandica (L.) followed the allometric equations: 4.22w ^0.62, 11.60w ^0.70, 7.45w ^0.66, 6.00w ^0.75 and 5.55w ^0.62, respectively. Five species of bivalves [Spisula subtruncata (da Costa), Hiatella striata (Fleuriau), Cultellus pellucidus (Pennant), Mya arenaria L. and Venerupis pullastra (Montagu)] filtered with the same rates as individuals of Cardium echinatum and A. islandica of equivalent soft weight. In Pecten furtivus and P. opercularis filtration rates were about twice the rates measured in individuals of Mytilus edulis of comparable body weight. The gill area in M. edulis increases with size at the same rate as the filtration rate.     

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.     

The influence of macrofauna on estuarine benthic community metabolism: a microcosm study

Effects of benthic macrofauna (Corophium volutator, Hydrobia sp., Nereis virens) on benthic community metabolism were studied over a 65-d period in microcosms kept in either light/dark cycle (L/D-system) or in continuous darkness (D-system). Sediment and animals were collected in January 1986 in the shallow mesohaline estuary, Norsminde Fjord, Denmark. The primary production in the L/D-system after 10 d acted as a stabilizing agent on the O₂ and CO₂ flux rates, whereas the D-system showed decreasing O₂ and CO₂ flux throughout the period. Mean O₂ uptake over the experimental period ranged from 0.38 to 1.24 mmol m^–2 h^–1 and CO₂ release varied from 0.80 to 1.63 mmol m^–2 h^–1 in both systems. The presence of macrofauna stimulated community respiration rates measured in darknes, 1.4 to 3.0 and 0.9 to 2.0 times for O₂ and CO₂, respectively. In contrast, macrofauna lowered primary production. Gross primary production varied from 1.06 to 2.26 mmol O₂ m^–2 h^–1 and from 1.26 to 2.62 mmol CO₂ m^–2 h^–1. The community respiratory quotient (CRQ, CO₂/O₂) was generally higher in the begining of the experiment (0–20 d, mean 1.89) than in the period from Days 20 to 65 (mean 1.38). The L/D-system exhibited lower CRQ (ca. 1) than the D-system. The community photosynthetic quotient varied for both net and gross primary production from 0.64 to 1.03, mean 0.81. The heterotrophic D-system revealed a sharp decrease in the sediment content of chlorophyll a as compared to the initial content. In the autotrophic L/D-system, a significant increase in chlorophyll a concentration was observed in cores lacking animals and cores with C. volutator (The latter species died during the experiment). Due to grazing and other macrofauna activities other cores of the L/D-system exhibited no significant change in chlorophyll a concentration. Community primary production was linearly correlated to the chlorophyll a content in the 0 to 0.5 cm layer. Fluxes of DIN (NH₄ ⁺+NO₂ ^–+NO₃ ^–) did not reveal significant temporal changes during the experiment. Highest rates were found for the cores containing animals, mainly because of an increased NH₄ ⁺ flux. The release of DIN decreased significantly due to uptake by benthic microalgae in the L/D-system. No effects of the added macrofauna were found on particulate organic carbon (POC), particulate organic nitrogen (PON), total carbon dioxide (TCO₂) and NH₄ ⁺ in the sediment. The ratio between POC and PON was nearly constant (9.69) in all sediment dephts. The relationship between TCO₂ and NH₄ ⁺ was more complex, with ratios below 2 cm depth similar to those for POC/PON, but with low ratios (3.46) at the sediment surface.