An Incubator for the Simulation of a Fluctuating Light Climate in Studies of Planktonic Primary Productivity

A laboratory system for the quantification of phytoplankton photosynthesis under fluctuating light climate conditions is described. It consists of 2 temperature-controlled incubators with a variable light supply, an algal batch culture in incubation bottles with appropriate stirrers and a set of oxygen electrodes to monitor algal photosynthesis. By the rotation of special grey filters between the incubator and the light source, a regular up and down movement in the water column is simulated in up to 7 parallel bottles. Different ratios of euphotic depth to mixing depth and different velocities can be applied. Simultaneously, 8 bottles can be incubated under constant light. The system is demonstrated in experiments with Chlamydomonas sp. Further possibilities of application are proposed.     

SEASONAL PRODUCTIVITY OF TWO RED ALGAE IN A CENTRAL CALIFORNIA KELP FOREST1

The in situ productivity of Botryocladia pseudodichotoma (Farl.) Kyl. and Rhodymenia californica var. californica Kyl., two common understory macrophytes in Macrocystis pyrifera (L.) C. Ag. forests, was determined once per month for one year. Gross productivity for B. pseudodichotoma was highest in October (1.23 mgC · g dry wt.^?1· h^?1), but rates were generally greater in spring and lower in summer. Respiration was variable throughout the year, especially in proportion to gross productivity. Gross productivity for R. californica var. californica was also highest in October (4.62 mgC · g dry wt.^?1· h^?1), and lowest in summer. Respiration was highly variable throughout the year. Deep (11 m) populations o/B. pseudodichotoma had 50% higher productivity than shallow (3 m) populations when incubated in shallow water, and 45% higher productivity when incubated in deep water. Populations of B. pseudodichotoma growing in shade at 3 m had 77% higher productivity than populations growing in sun at the same depth when incubated in a sun exposed location at 3 m. Respiration of the shade-adapted plants was only one-half that of the sun-adapted plants. In comparison with similar studies in other sub-tidal communities, net productivity of these two understory red algae is somewhat lower. Lack of strong seasonal productivity patterns and the highly variable underwater light regime suggests that understory algae may be adapted for rapid growth during short periods of high light, regardless of the time of year.     

Effects of sediment resuspension on phytoplankton production: teasing apart the influences of light, nutrients and algal entrainment

1. Wind‐induced sediment resuspension can affect planktonic primary productivity by influencing light penetration and nutrient availability, and by contributing meroplankton (algae resuspended from the lake bed) to the water column. We established relationships between sediment resuspension, light and nutrient availability to phytoplankton in a shallow lake on four occasions. 2. The effects of additions of surficial sediments and nutrients on the productivity of phytoplankton communities were measured in 300 mL gas‐tight bottles attached to rotating plankton wheels and exposed to a light gradient, in 24 h incubations at in situ temperatures. 3. While sediment resuspension always increased primary productivity, resuspension released phytoplankton from nutrient limitation in only two of the four experiments because the amount of available nitrogen and phosphorus entrained from the sediments was small compared with typical baseline levels in the water column. In contrast, chlorophyll a entrainment was substantial compared with baseline water column concentrations and the contribution of meroplankton to primary production was important at times, especially when seasonal irradiance in the lake was high. 4. Comparison of the in situ light climate with the threshold of light‐limitation of the phytoplankton indicated that phytoplankton in the lake were only likely to be light‐limited at times of extreme turbidity (e.g. >200 nephelometric turbidity units), particularly when these occur in winter. Therefore, resuspension influenced phytoplankton production mainly via effects on available nutrients and by entraining algae. The importance of each of these varied in time. 5. The partitioning of primary productivity between the water column and sediments in shallow lakes greatly influences the outcome of resuspension events for water column primary productivity.     

Variation in species light acquisition traits under fluctuating light regimes: implications for non‐equilibrium coexistence

Resource distribution heterogeneity offers niche opportunities for species with different functional traits to develop and potentially coexist. Available light (photosynthetically active radiation or PAR) for suspended algae (phytoplankton) may fluctuate greatly over time and space. Species‐specific light acquisition traits capture important aspects of the ecophysiology of phytoplankton and characterize species growth at either limiting or saturating daily PAR supply. Efforts have been made to explain phytoplankton coexistence using species‐specific light acquisition traits under constant light conditions, but not under fluctuating light regimes that should facilitate non‐equilibrium coexistence. In the well‐mixed, hypertrophic Lake TaiHu (China), we incubated the phytoplankton community in bottles placed either at fixed depths or moved vertically through the water column to mimic vertical mixing. Incubations at constant depths received only the diurnal changes in light, while the moving bottles received rapidly fluctuating light. Species‐specific light acquisition traits of dominant cyanobacteria (Anabaena flos‐aquae, Microcystis spp.) and diatom (Aulacoseira granulata, Cyclotella pseudostelligera) species were characterized from their growth–light relationships that could explain relative biomasses along the daily PAR gradient under both constant and fluctuating light. Our study demonstrates the importance of interspecific differences in affinities to limiting and saturating light for the coexistence of phytoplankton species in spatially heterogeneous light conditions. Furthermore, we observed strong intraspecific differences in light acquisition traits between incubation under constant and fluctuating light – leading to the reversal of light utilization strategies of species. This increased the niche space for acclimated species, precluding competitive exclusion. These observations could enhance our understanding of the mechanisms behind the Paradox of the Plankton.     

Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

Variation in some photosynthetic characteristics of microalgae cultured in outdoor thin-layered sloping reactors

In outdoor thin-layer sloping reactors algae are batch cultured and harvested at biomass concentrations of about 15 g (dw) I^-1 whereafter a portion is used as inoculum for the next cycle. Light saturation curves of the oxygen evolution (PII curves) of the algae were measured using diluted aliquots of suspension taken from the reactors. The maximum specific photosynthetic rates (P ^B _max) and the light intensity at the onset of saturated photosynthesis (I _k ) decreased whilst the maximum specific photosynthetic efficiency ( ^B ) increased with an increase in the biomass concentration, during the production cycle. These differences reflect transition from light- to dark-acclimated state of the algae that occurs as a result of an increase of the suspension concentration during the production cycle. During these experiments the thin-layered smooth sloping cultures (TLSS, culture depth 5–7 mm) had higher photosynthetic rates per volume than the thin-layered baffled sloping cultures (TLBS, culture depth 5–15 mm). This was ascribed to the higherP ^B _max values of the algae grown in the TLSS cultures, allowing them to utilise high incident irradiancies more effectively. However, the areal productivity of the TLBS was higher than the TLSS indicating a higher photosynthetic efficiency of the TLBS reactors. The specific productivity decreased rapidly with an increase in the biomass concentration, but the yield remained linear during the batch production cycle, even at high areal densities.     

COMPOSITION,DISTRIBUTION, AND ABUNDANCE OF DEEP‐WATER (>30 m) MACROALGAE IN CENTRAL CALIFORNIA1

The deep‐water macroalgal assemblage was described at 14 sites off the central California coast during 1999 and 2000 from SCUBA and remotely operated vehicle sampling. The stipitate kelp Pleurophycus gardneri Setchell & Gardner, previously thought to be rare in the region, was abundant from 30 to 45 m, forming kelp beds below the well‐known giant kelp forests. Macroalgae typically formed three broadly overlapping zones usually characterized by one or a few visually dominant taxa: 1) the upper “Pleurophycus zone” (30–45 m) of stipitate kelps and Desmarestia spp. with a high percent cover of corallines, low cover of uncalcified red algae, and rare green algae; 2) a middle “Maripelta zone” (40–55 m) with other uncalcified red algae and infrequent corallines and green algae; and 3) a zone (55–75 m) of infrequent patches of nongeniculate coralline algae. The green alga Palmophyllum umbracola Nelson & Ryan, not previously reported from the Northeast Pacific, was found over the entire geographical range sampled from 35 to 54 m. Year‐round profiles of water column irradiance revealed unexpectedly clear water with an average K₀ of 0.106·m ^{? 1} Received 18 January 2002. Accepted 16 December 2002. . The low percent surface irradiance found at the average lower macroalgal depth limits in this study (0.56% for brown algae, 0.12% for uncalcified red algae, and 0.01% for nongeniculate coralline algae) and lack of large grazers suggest that light controls the lower distributional limits. The ubiquitous distribution, perennial nature, and similar lower depth limits of deep‐water macroalgal assemblages at all sites suggest that these assemblages are a common persistent part of the benthic biota in this region.     

Spatial and temporal variability in filament length of a toxic cyanobacterium (Anabaena affinis)

1. This study examines the distribution of Anabaena affinis filament lengths under natural conditions as a function of depth and season, and in the laboratory as a function of growth phase. Because Anabaena affinis is only toxic when consumed, both its filament length and position in the water column are important determinants of its potential impact on zooplankton populations. 2. Star Lake (Norwich, Vermont, U.S.A.), a natural, eutrophic pond, remained thermally stratified throughout the Anabaena bloom. Filament number and length differed significantly with both sampling date and water depth. Most filaments occurred at 0.5 m, particularly at the height of the bloom. Throughout the entire water column average filament length decreased from approximately 0.53 mm in May to 0.14 mm in July. The shortest filaments occurred at the 2.5 m depth. Filament length distributions (combined for all depths) for 29 May, 12 June and 3 July, corresponding to the beginning, middle and end of the bloom, respectively, differed significantly among the three dates. These patterns most likely reflect variable growth conditions, both during the season and in the water column. 3. In the laboratory, Anabaena filament length was affected by medium composition and growth phase. Filaments were significantly longer when grown in MBL than in ASM medium. Also, the average length of Anabaena filaments grown in MBL changed significantly as cultures aged; by day 13 filament length (2.01 ± 0.38 mm, mean ± SD) was twice that on day 0 (0.97 ± 0.71 mm). As cell concentration continued to increase, mean filament length gradually decreased.     

Natural succession of macroalgal-dominated epibenthic assemblages at different water depths and after transplantation from deep to shallow water on Spitsbergen

In the current study, we investigated the primary succession of seaweeds over different time periods at different water depths. Furthermore, we followed the succession of field-grown benthic communities of different successional age, developing on ceramic tiles, prior to and after transplantation from 8 to 0.5 m water depth. The transplantation simulated changes associated with the break up of sea-ice cover, e.g. light regime or wave exposure. For this purpose, we transplanted 12 and 21-month old communities, grown at 8 m water depth, together with a set of sterile tiles, onto rafts, floating in 0.5 m water depth. Our results describe for the first time the succession of macroalgal communities in the Arctic and give important insights into the effect of disturbance of differently aged communities. The primary succession at 0.5 m water depth was mainly driven by Bacillariophyta and filamentous green algae like Urospora sp. and Ulothrix implexa. Twelve-month old communities at 8 m water depth are dominated by members of the Ectocarpales (Phaeophyceae), like Pylaiella littoralis, P. varia, and Ectocarpus siliculosus and the green alga U. implexa, whereas the 21-month old community showed a higher cover of the green algal class Ulvophyceae and sessile invertebrates. After transplantation to near surface conditions, species composition of the communities changed, but this effect was differently strong between communities of different age.     

A test of a model for estimating production in the sea

Primary productivity was measured in the sea off the coasts of Florida and Georgia with the radiocarbon method at four stations during March, 1971. Integral photosynthesis per square meter surface was determined with a mathematical model from empirical data gathered from in situ and shipboard incubation experiments. The mathematical model provided shipboard incubators to be used to estimate primary production rates while overcoming the problem of relating light quality and quantity simulated by the incubators to that found in the euphotic zone. The use of shipboard incubation can allow for measurements to be made on a moving vessel, more stations visited per day and water samples incubated at a constant light intensity. Primary productivity at the four sampling stations ranged from 0.026 to 0.042 mg C/M² hr.     

The annual cycle of primary productivity in a tropical estuary: the inner regions of the Golfo de Nicoya, Costa Rica

A one year cycle of primary productivity (PP) was studied using the "light and dark bottle" technique in the Golfo de Nicoya, located at 10 degrees N and 85 degrees W at the Pacific coast of Costa Rica. Samples were always incubated at 0, 1, 2, 3 and 4 m depth for 5 hrs from 8:30 till 13:30. The measurements were performed twice per month, first around high tide and one week later at low tide to account for tidal influences. This routine study was supplemented by special measurements about regional and short-term variations of primary productivity using the 14C-method, which mainly served to account for the shortcomings of the routinely employed incubation technique. The upper Golfo de Nicoya is an extremely productive, phytoplankton dominated estuarine system with an annual gross PP of 1037, a net PP of 610 and a community respiration of 427 g C m(-2) a(-1). Highest monthly PP values occurred during the dry season and at the beginning of the rainy season. Peaks in primary productivity coincided with massive blooms of red tide forming algae. Internal biological dynamics, estuarine circulation and land run-off are the most important nutrient sources. High water turbidity reduces the euphotic layer to 4-5 m depth, making the underwater light regime the rate limiting factor. On an annual basis, 41% of the organic carbon produced in the system is already consumed in the euphotic layer. Considering the entire water column (mean depth at mean tidal water level is around 7.7 m) 79% is consumed in the pelagial. Taking into account the organic material consumed and stored in the sediments the carbon budget of the upper gulf is probably balanced. Since, however, the system receives a considerable amount of organic material from its terrestrial surroundings (especially from the mangrove forests), a surplus of organic carbon is exported from the upper Golfo de Nicoya, which enhances the overall water productivity of the lower gulf and the adjacent area.     

Photoacclimation of Scenedesmus protuberans (Chlorophyceae) to fluctuating irradiances simulating vertical mixing

Photoacclimation of Scenedesmus protuberans Fritsch to fluctuatingirradiances, simulating vertical mixing, was studied in light-limitedcontinuous cultures. The algae were exposed to a simple sinusoidallight regime simulating diumal irradiance, and two fluctuatingregimes in which light fluctuations resulting from four andeight vertical circulations of the algae through the water columnwere imposed on the sinusoidal light regime. The total dailylight dose (TDLD) was kept constant. Maximum photosynthesisincreased, and cellular chlorophyll content and photosyntheticunit size decreased in response to light fluctuations. The efficiencyof photosynthesis decreased during the light period in the sinusoidallight regime, whereas it remained at the same level in the fluctuatinglight regimes. The results suggest that prolonged exposure tointermediate irradiances lowers the photosynthetic efficiencyof S.protuberans more than short exposure to high irradiancesalternated by lowlight periods, and that this species is ableto optimize its photosynthesis in fluctuating light.     

In situ study of relative electron transport rates in the marine macroalga <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> in the Baltic Sea at different depths and times of year

The brown alga Fucus vesiculous is one of the few marine species in the Baltic Sea. Fucus vesiculosus shows high morphological and physiological variability as a response to its environmental conditions. The salinity in the Baltic Sea is 4–5 psu, compared to 35 psu in the Atlantic. Photosynthesis of algae is usually measured after collection and transportation to constant culture conditions. However, in this study, relative photosynthetic electron transport rates, calculated from chlorophyll a fluorescence parameters were compared in algae collected from 1 and 4 m depths by SCUBA divers. Measurements of light response curves from the same individuals of F. vesiculosus at different depths and times of the year have, to our knowledge, not been made previously. Measurements were performed on four different occasions during the spring of 2005 (25 February, 3 and 29 April, and 26 May) in the Baltic Sea, using rapid light curves generated with a Diving PAM. In addition, samples were collected for photoinhibition studies in the laboratory. The light response curves obtained in situ at 1 and 4 m depths for F. vesiculosus showed lower values of light saturation with depth. When algae from 1 and 4 m depths were exposed to high irradiances of photosynthetically active radiation (1,400 μmol photons m⁻² s⁻¹), algae from 1 m depth showed a higher degree of photoinhibition in comparison to algae from 4 m depth.     

On the mass culture of microalgae: Areal density as an important factor for achieving maximal productivity

Scenedesmus obliquus was grown in a bioreactor under constant light and at a temperature of 24°C. From measurements of biomass production, it was possible to determine the influence of areal density on productivity in open as well as closed cultures. Maximal productivity occurred at an areal density of 40 to 45 g(dw) M⁻², which was in good agreement with predicted values from a model. The optimal areal density for maximal productivity was influenced by factors such as culture depth, algal species, turbulence, available light, etc., and hence the establishment and maintenance of an optimal areal density is seen as one of the most important operational procedures for the mass culture of algae. Supersaturated concentrations of oxygen resulted in lower productivities due to photorespiration and/or oxidation.     

Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.     

Influence of changes in salinity and light intensity on growth of phytoplankton communities from the Schelde river and estuary (Belgium/The Netherlands)

In the Schelde continuum, a succession in the phytoplankton community is observed along the transition from the river to the freshwater tidal reaches of the estuary and from the freshwater to brackish reaches of the estuary. The goal of this study was to experimentally evaluate the contribution of changes in salinity and light climate to this succession. In summer 2000 and in spring 2001, phytoplankton communities from the river, the freshwater tidal reaches and the brackish reaches of the estuary were incubated under high or low light intensities and exposed to a change in salinity. HPLC analysis was used to evaluate the response of different algal groups to changes in light intensity and salinity. When incubated at a light intensity corresponding to the mean underwater light intensity of the freshwater tidal reaches, growth of phytoplankton from the river as well as from freshwater tidal reaches was significantly lower than when incubated at a light intensity corresponding to the mean underwater light intensity of the river. The phytoplankton community from the freshwater tidal reaches did not appear to be better adapted to low light intensities than the phytoplankton community from the river. Although diatoms were expected to be less sensitive to a reduction in light intensity than green algae, the opposite response was observed. Freshwater and brackish water phytoplankton were negatively affected by respectively an increase or decrease in salinity. However, the effect of salinity was not strong enough to explain the disappearance of freshwater and brackish water phytoplankton between a salinity of 0.5 and 10 psu, suggesting that other factors also play a role. In the freshwater phytoplankton communities from the river and the freshwater tidal reaches, green algae and diatoms responded in a similar way to an increase in salinity. In the brackish water phytoplankton community, fucoxanthin displayed a different response to salinity than lutein and chlorophyll a.     

The behaviour of log phase Escherichia coli at temperatures that fluctuate about the minimum for growth

AIMS: To investigate the behaviour of cold-adapted, log phase Escherichia coli exposed to temperatures that fluctuate below and above the minimum for growth. METHODS AND RESULTS: Log phase E. coli cultures were incubated at a constant temperature of 2, 4 or 6 degrees C or with temperatures allowed to increase from those temperatures for 35 min, to 10 degrees C, at 6-, 12- or 24-h intervals, as commonly occurs during retail display of chilled foods. At suitable intervals for each culture, the optical absorbance value was determined using a spectrophotometer, the forward angle light scatter was determined using a flow cytometer, and portions were spread on plate count agar for enumeration of colony forming units (CFU). Numbers of CFU decreased by 3 log units or increased by 1 log unit for cultures incubated at 6 degrees C for 17 days without or with temperatures fluctuations at < or =12-h intervals, respectively. Cells elongated when cultures were incubated at 4 or 2 degrees C with temperatures fluctuating at 6-h intervals, and at 6 degrees C at constant or fluctuating temperatures, but cells did not elongate in cultures incubated at a constant temperature of 2 or 4 degrees C. SIGNIFICANCE AND IMPACT OF THE STUDY: The minimum growth temperature of E. coli is assumed to be > or =7 degrees C. Elongated cells were able to divide when temperatures rose from 6 degrees C to above 7 degrees C for <45 min at < or =12-h intervals. Such temperature fluctuations may be experienced by chilled foods during defrosting cycles of retail display cases. The finding that cells behave differently under fluctuating than at constant temperatures may significantly affect understanding of appropriate temperatures for the safe storage of chilled foods and for predictive modelling of bacterial growth in such foods.     

Tidal Dependence of Photoinhibition of Photosynthesis in Marine Macrophytes of the South China Sea*

During an expedition in spring 1992 to Hainandao, an island in the tropical zone of the South China Sea, the daily courses of photoinhibition of different brown algal species and of the seagrass Thalassia hemprichii were investigated. Experiments were carried out with the new portable chlorophyll fluorometer PAM 2000 (Walz, Germany). As a measure of photoinhibition Fv/Fm was used and as a measure of the photosynthetic yield ΔF/Fm'. Photoinhibition occurred in all algae floating near the water surface and reached its maximum between noon and the early afternoon. In the evening photosynthesis was always fully recovered. The extent of photoinhibition depended on both the depth of the algae and the course of the irradiance during the day. Algae of the sublittoral zone showed only a low degree of photoinhibition at high fluence rates when they were covered by a water column of more than 1 m, even if the water was clear. The seagrass Thalassia hemprichii grew in the middle and upper intertidal zone and showed a significant photoinhibition at low tide only when it was not water-covered. Apparently, it is able to cope with extreme high light conditions without downregulation of photosynthesis caused by photoinhibition.     

ZONATION OF DEEP WATER BENTHIC ALGAE IN THE GULF OF MAINE1

Algal community structure is described for a deep-water rock pinnacle in the Gulf of Maine. Three depth zones of algal dominance were apparent consisting of 1) leathery macrophytes (to 40 m), 2) foliose red algae (to 50 m) and 3) crustose algae (fleshy crusts to 55 m and coralline crusts to 63 m). Microscopic filamentous and erect calcareous algae were also present but inconspicuous. Upright macroscopic filamentous and thin sheet-like forms were not observed on the pinnacle. Sea anemones (Metridium senile) dominated some vertical faces and abrupt prominences in the shallowest regions of the pinnacle (to 24 m) and locally appeared to set the upper vertical limits of kelp and possibly foliose reds. Laminaria sp. formed an open park-like canopy from 24 to 30 m whereas Agarum cribrosum, the deepest kelp, grew as isolated individuals to 40 m. Peyssonnelia sp. and Leptophytum laeve were the deepest occurring fleshy (to 55 m) and calcareous crusts (to 63 m), respectively. The occurrence of these algae at record depths for the Gulf of Maine and for cold water marine environments may be the result of an absence of large herbivores and the high productivity potential of the benthos in these relatively clear waters. By compiling data on depth distribution patterns world-wide, it is evident that the three zone structure of algal morphologies observed in the Gulf of Maine is a global phenomenon.     

Annual productivity of Spirulina (Arthrospira) and nutrient removal in a pig wastewater recycling process under tropical conditions

An evaluation was made of the annual productivity of Spirulina (Arthrospira) and its ability to remove nutrients in outdoor raceways treating anaerobic effluents from pig wastewater under tropical conditions. The study was based at a pilot plant at La Mancha beach, State of Veracruz, Mexico. Batch or semi-continuous cultures were established at different seasons during four consecutive years. The protein content of the harvested biomass and the N and P removal from the ponds were also evaluated. Anaerobic effluents from digested pig waste were added in a proportion of 2% (v/v) to untreated sea-water diluted 1:4 with fresh water supplemented with 2 g L^–1 sodium bicarbonate, at days 0, 3 and 5. A straight filament strain of Spirulina adapted to grow in this complex medium was utilized. A pH value 9.5 ± 0.2 was maintained. The productivity of batch cultures during summer 1998 was significantly more with a pond depth of 0.10 m than with a depth 0.065 m. The average productivity of semi-continuous cultures during summer 1999 was 14.4 g m^–2 d^–1 with a pond depth of 0.15 m and 15.1 g m^–2 d^–1 with a depth of 0.20 m. The average annual productivity for semi-continuous cultures operating with depths of 0.10 m for winter and 0.15 and 0.25 m for the rest of the year, was 11.8 g m^–2 d^–1. This is the highest value reported for a Spirulina cultivation system utilising sea-water. The average protein content of the semi-continuous cultures was 48.9% ash-free dry weight. NH₄-N removal was in the range 84–96% and P removal in the range of 72–87%, depending on the depth of the culture and the season.