The stable level of coral primary production in a wide light range

Light adaptation and photosynthetic productivity were studied in common reef-building corals on islands of the Indian Ocean and the South China Sea. When light is attenuated, both in shade and at depth, adaptations by zooxanthellae permit maximal absorption and utilization of light. Better utilization of incident light in shade-dwelling and deep-water coral forms is reflected by higher values of gross photosynthesis on the plateau and linear portion of the photosynthesis-irradiance curve. It was shown that outer branches of reef-building corals are autotrophic in a major part of their light-range distribution and have a high and stable level of primary production.     

Nutrients,light and primary production by phytoplankton and microphytobenthos in the eutrophic,turbid Westerschelde estuary (The Netherlands)

Abiotic factors and primary production by phytoplankton and microphytobenthos was studied in the turbid Westeschelde estuary. Because of the high turbidity and high nutrient concentrations primary production by phytoplankton is light-limited. In the inner and central parts of the estuary maximum rates of primary production were therefore measured during the summer, whereas in the more marine part spring and autumn bloom were observed. Organic loading is high, causing near anaerobic conditions upstream in the river Schelde. Because of this there were no important phytoplankton grazers in this part of the estuary and hence the grazing pressure on phytoplankton was minimal. As this reduced losses, biomass is maximal in the river Schelde, despite the very low growth rates.On a number of occasions, primary production by benthic micro-algae on intertidal flats was studied. Comparison of their rates of primary production to phytoplankton production in the same period led to the conclusion that the contribution to total primary production by benthic algae was small. The main reason for this is that the photosynthetic activity declines rapidly after the flats emerged from the water. It is argued that CO₂-limitation could only be partially responsible for the noticed decrease in activity.     

In situ measurements confirm the seasonal dominance of benthic algae over phytoplankton in nearshore primary production of a large lake

1. Despite the recognition of its importance, benthic primary production is seldom reported, especially for large lakes. We measured in situ benthic net primary production by monitoring flux in dissolved inorganic carbon (DIC) concentration in benthic incubation chambers, based on continuous measurements of CO_2(aq) flux, alkalinity, and the temperature‐dependent dissociation constants of carbonic acid (K₁ and K₂). This methodology has the advantages of monitoring net primary production directly as change in carbon, maintaining continuous water recirculation, and having sufficient precision to detect change in DIC over short (i.e. 15 min) incubations, even in alkaline waters. 2. Benthic primary production on Cladophora‐dominated rocky substrata in western Lake Ontario was measured biweekly. Maximum biomass‐specific net photosynthetic rates were highest in the spring (2.39 mgC g Dry Mass^?1 h^?1), decreased to negative rates by early summer (?0.76 mgC g DM^?1 h^?1), and exhibited a regrowth in late summer (1.98 mgC g DM^?1 h^?1). 3. A Cladophora growth model (CGM), previously validated to predict Cladophora biomass accrual in Lake Ontario, successfully simulated the seasonality and magnitude of biomass‐specific primary production during the first cohort of Cladophora growth. Averaged over this growing season (May–Aug), mean areal net benthic production at the estimated depth of peak biomass (2 m) was 405 mg C m^?2 d^?1. 4. We measured planktonic primary production in proximity to the benthic study and constructed a depth‐resolved model of planktonic production. Using the CGM, benthic primary production was compared with planktonic primary production for the period May–Aug. Net benthic production from the shoreline to the 12 m contour (1–2 km offshore) equalled planktonic production. Closer to shore, benthic primary production exceeded planktonic primary production. Failure to account for benthic primary production, at least during abundant Cladophora growth, will lead to large underestimates in carbon and nutrient flows in the nearshore zone of this Great Lake.     

The relationship of floods, drying, flow and light to primary production and producer biomass in a prairie stream

Factors related to autochthonous production were investigated at several sites along a prairie stream at Konza Prairie Research Natural Area. Primary production, algal biomass, litter input, and ability of floods to move native substrate were measured. Additional experiments were conducted to establish the influence of light and water velocity on primary production rates and recovery of biomass following dry periods. The study period encompassed two extreme (> 50 year calculated return time) floods, thus we were able to analyze the effects of scour on periphyton biomass and productivity. Biomass of sedimentary algae was reduced greatly by flooding and did not reach preflood amounts during the 2 months following the first flood. Rates of primary production associated with sediments recovered to levels above preflood rates within 2 weeks. Biomass of epilithic periphyton was not affected as severely as that of sedimentary algae. Little relationship was observed between water velocity and photosythetic rates. Production reached maximum rates at 25% of full sun light. Epilithic chlorophyll levels recovered within eight days following a dry period, and chl a was an order of magnitude greater on rocks than sediments 51 days after re-wetting. Estimated annual rates of primary production were 2.6 times greater in the prairie than in the forest reaches of the stream. The ratio of annual autochthonous:allochthonous carbon input was 4.81 for prairie and 0.32 for the forest. Periphyton production in prairie streams is resilient with regard to flooding and drought and represents a primary carbon source for the system.     

Changes in phytoplankton biomass and primary production between 1991 and 2001 in the Westerschelde estuary (Belgium/The Netherlands)

The Westerschelde estuary is a very polluted and turbid estuary, but the last decade the waterquality improved. Dredging activity also increased in 1997 to allow bigger ships to enter the port of Antwerpen. This could potentially decrease the light conditions for the phytoplankton. Because of all these recent changes in the estuary we studied primary productivity in 2001 and compared it to values in 1991. The results show that due to a decrease in discharge in particulate and dissolved organic carbon the oxygen concentrations in general have increased in the upstream region, although in spring and summer low oxygen concentrations (10–30% saturation) can still be found. Phosphate and ammonia concentrations have decreased and the zone of nitrification which was very large in 1991 has become very small and is now located in the uppermost upstream region of the estuary. Si-concentrations have remained the same. All nutrient concentrations are still high enough not to limit phytoplankton growth. Turbidity remained unaltered as a result of the dredging works, and as a result phytoplankton biomass in most of the estuary did not show a decrease, although there were signs that in the upstream region phytoplankton biomass decreased, possible caused by increased grazing pressure. The relationship between phytoplankton biomass and primary productivity did not change, and from the data it can be concluded that the dredging activity will not influence the gross and net primary productivity of the phytoplankton.     

Physico-chemical environment,phytoplankton biomass and production in oligotrophic,softwater lake kalgaard,Denmark

Water chemistry of Lake Kalgaard in 1976–77 was characterized by low concentrations of total-CO₂ and inorganic nutrients. The ionic composition resembled that of precipitation (Na>Ca>Mg >K and Cl>SO₄>HCO₃). The seasonal pattern of total-CO₂ and PO₄ was regulated by internal processes and maximum concentrations as a result of decomposition processes occurred during summer stagnation. NO₃ concentrations showed the opposite pattern and were relatively high from late autumn through spring and were extremely low during summer. Total-P and PO₄ increased during summer due to release from the sediment. The phytoplankton biomass of surface water was low. The water chemistry suggested a shift from N-limitation of phytoplankton during summer to P-limitation at other seasons. Maximum algal concentrations occurred at 6 m during summer, probably due to a supply of nutrients (especially NH₄) from deeper layers. Phytoplankton productivity was often bimodal, with an upper maximum at depths of 0 or 2 m and a second maximum at 6 m.     

The annual flood regime as a regulatory mechanism for phytoplankton production in Kainji lake,Nigeria

Phytoplankton production was measured in situ in Kainji lake from December 1970 to September 1972 using the oxygen light and dark bottle technique. Seasonal variations in solar radiation, transparency, temperature, and composition of subsurface light were also measured. Oxygen production per unit area varied from 220 to 4500 mg O₂ m^–2 day^–1, the maximum production rate from 95 to 400 mg O₂ m^–3 h^–1. Seasonal mixing of lake water and river water of varying turbidity changed the optical properties of the lake water and consequently affected phytoplankton production. The annual flood pattern was found to be an important factor regulating phytoplankton production in the lake.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.     

Seasonal variation in phytoplankton primary production in relation to light and nutrients in Lake Awasa,Ethiopia

The biomass and primary production of phytoplankton in Lake Awasa, Ethiopia was measured over a 14 month period, November 1983 to March 1985. The lake had a mean phytoplankton biomass of 34 mg chl a m^–3 (n = 14). The seasonal variation in phytoplankton biomass of the euphotic zone (mg chl a m^–2 h^–1) was muted with a CV (standard deviation/mean) of 31%. The vertical distribution of photosynthetic activity was of a typical pattern for phytoplankton with light inhibition on all but overcast days. The maximum specific rates of photosynthesis or photosynthetic capacity (Ø_max) for the lake approached 19 mg O₂ (mg chl a)^–1 h^–1, with high values during periods of low phytoplankton biomass. Areal rates of photosynthesis ranged between 0.30 to 0.73 g O₂ m^–2 h^–1 and 3.3 to 7.8 g O₂ m^–2 d^–1. The efficiency of utilisation of PhAR incident on the lake surface varied from 2.4 to 4.1 mmol E^–1 with the highest efficiency observed corresponding to the lowest surface radiation. Calculated on a caloric basis, the efficiency ranged between 1.7 and 2.9%. The temporal pattern of primary production by phytoplankton showed limited variability (CV = 21 %).     

Annual patterns of phytoplankton density and primary production in a large, shallow lake: the central role of light

1. We studied the seasonal dynamics of suspended particulate matter in a turbid, large shallow lake during an annual period (2005–06). We relate the patterns of seston concentration (total suspended solids), phytoplankton biomass and water transparency to the seasonal pattern of incident solar radiation (I₀). We also report the seasonal trends of phytoplankton primary production (PP) and photosynthesis photoinhibition due to photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) (I_β and UV₅₀). 2. We first collected empirical evidence that indicated the conditions of light limitation persisted during the study period. We found that the depth‐averaged irradiance estimated for the time of the day of maximum irradiance (I_mean–noon) was always lower than the measured onset of light saturation of photosynthesis (I_k). 3. We then contrasted the observations with theoretical expectations based on a light limitation scenario. The observed temporal patterns of seston concentration, both on a volume and area basis, were significantly explained by I₀ (R² = 0.39 and R² = 0.37 respectively). The vertical diffuse attenuation coefficient (kd_PAR) (R² = 0.55) and the depth‐averaged irradiance (I_mean) (R² = 0.66), significantly increased with the I₀; while the irradiance reaching the lake bottom (I_out) significantly decreased with the incident irradiance (R² = 0.49). However, phytoplankton biovolume maxima were not coincident with the time of the year of maximum irradiance. 4. A significant positive relationship was observed between PP estimated on an area basis and I₀ (R² = 0.51, P < 0.001). In addition, the parameters describing the photosynthetic responses to high irradiances displayed marked seasonal trends. The photosynthesis photoinhibition due to PAR as well as to UV were significantly related to incident solar radiation (PAR: R² = 0.73; UV: R² = 0.74). These results suggest adaptation of the phytoplankton community in response to changes in incident solar radiation.     

Distribution, biomass and primary production of an Ahnfeltia tobuchiensis (Ahnfeltiales, Rhodophyta) population in the Bay of Izmena, Kunashir Island

Regularities of distribution and primary production of an Ahnfeltia tobuchiensis (Kanno et Matsubara) Mak. population, an agar-containing red alga, were studied in the Bay ot Izmena. Experiments were conducted in a flow-through system under conditions similar to algal habitats. The population of A. tobuchiensis unattached to the ground may be from a few centimeters to as much as 1 m thick. It has been shown that only the upper part of a stratum 15–20 cm thick receives a sufficient amount ot light to realize its production potential. While 15–20% of photosynthetically active radiation (PAR) of that falling on the water surface reaches the stratum surface, only 0.1% of PAR from that falling on the water surface penetrates through stratum 15 cm thick. It has been shown for A. tobuchiensis that its photosynthetic rate curve during the daytime mainly follows the PAR intensity curve. The highest values of photosynthetic rate have been measured in the afternoon when PAR reaches its maximum. It is noted that a stratum 15–20 cm thick has peak values ot net primary production (NPP) which averages 3.2 g C m^?2 day^?1. The total area of A. tobuchiensis population was 23.4 km², and its biomass was 125 000 tons in this area. On average, the NPP of the A. tobuchiensis population made up in summer and in autumn was 46.8 and 25.0% of its biomass, respectively.     

A comparison of phytoplankton size-fractions in Mondsee,an alpine lake in Austria: distribution,pigment composition and primary production rates

Production rates, abundance, chlorophyll a (Chl a) concentrations and pigment composition were measured for three size classes (<2 μm, 2–11 μm and >11 μm) of phytoplankton from May to December 2000 in deep, mesotrophic, alpine lake Mondsee in Austria. The study focuses on differences among phytoplankton size fractions characterised by their surface area to volume ratio ([mm² l⁻¹: mm³l⁻¹]), pigment distribution patterns and photosynthetic rates. Particular attention was paid to autotrophic picophytoplankton (APP, fraction <2 μm) since this size fraction differed significantly from the two larger size fractions. Among the three fractions, APP showed the highest surface area to volume ratios and a high persistence in the pattern of lipophilic pigments between temporarily and spatially successive samples (about 80% similarity of pigment composition between samples over seasons and depths). The epilimnetic abundance of APP varied seasonally with an annual maximum of 180 × 10³ cells ml⁻¹ in June (at 4–9 m). The minimum (October at 12 m) was more than an order of magnitude lower (4.9 × 10³ ml⁻¹). APP peaked during autumn and contributed between 24% and 42% to the total area-integrated Chl a (10–23 mg m⁻²) and between 16% and 58% to total area-integrated production (5–64 mg m⁻²  h⁻¹) throughout seasons.     

Characterization of the light field in laboratory scale enclosures of eutrophic lake water (Lake Loosdrecht,The Netherlands)

Light conditions in laboratory scale enclosures (LSE) of shallow, eutrophic Lake Loosdrecht (The Netherlands), including a method for simulating a natural incident light course, are described. Total PAR (400–700 nm) and spectral irradiance distribution were measured at sestonic chlorophyll a and dry weight concentrations 100 mg m^–3 and 16 g m^–3, respectively. Phytoplankton was dominated by Oscillatoria spp. The euphotic depth (Z _eu) was 0.7–1.0 m. Shortly after filling the LSE with lake water, diffuse attenuation coefficients ranged from 14 m^–1 for blue to 5 m^–1 for red light. Around Z _eu, attenuation in the blue region was markedly lower and irradiance reflectance (R) continued to increase; these anomalies were caused by lateral incident light from the LSE's waterbath. Spectrophotometry indicated that absorption was mainly by particles, but dissolved humic substances (gilvin) were also important. The particles were likely to be dominated by detritus absorbing more blue relative to red light. Subsurface R in lake water in the LSE had a maximum around 705 nm and low values in the blue band, but was lower than that previously reported for measurements in situ. Wash-out of detritus, presumably both dissolved and particulate fractions, by flow-through with synthetic medium, greatly affected the spectral reflectance measured outside the LSE. The maximum value of R decreased from 0.022 to 0.009, and the peak shifted to 550 nm.     

The use of photosynthesis inhibitor (DCMU) for in situ metabolic and primary production studies on soft bottom benthos

A selective chemical photosynthesis inhibitor, DCMU (Dichorophenyl-dimethylurea), dissolved in DMSO (Dimethyl sulfoxide) was substituted for the dark incubation method commonly used to measure the oxygen consumption in metabolic and primary production studies. We compared oxygen fluxes during light incubations with DCMU and dark incubations procedure, on soft bottom benthos. For this purpose, we studied the effects of different DCMU concentrations. A concentration of 5 · 10^–5 mol l^–1 inside a clear incubation enclosure completely inhibits photosynthesis without affecting the metabolism of soft bottom benthos.     

Patterns of temporal variation in Lake Titicaca. A high altitude tropical lake. I. Background,physical and chemical processes,and primary production

A statistical analysis is presented of patterns of variation in some physical, chemical, and biological variables for a 6 year series of data from the tropical, high altitude Lake Titicaca (Peru-Bolivia). ANOVA techniques and autocorrelation analyses were used to partition the variance in Titicaca, and in some comparison tropical and temperate series, into components with repeatable annual cycles and components attributable to other kinds of patterns.In Titicaca, insolation and stratification are highly seasonal in pattern of variation, although the amount of variance relative to means is small compared to temperate lakes. However, the seasonal pattern of physical variation is only weakly imposed on chemical and biological processes, to judge from analyses of silicate, oxygen, and primary production series. Comparable temperate series of primary production and chlorophyll a are much more seasonal.     

Production,population biology and diet ofNeomysis integer (Leach) in a shallow Frisian lake (The Netherlands)

TheNeomysis integer population of a shallow Frisian freshwater lake, situated in the north of the Netherlands, was studied from May until October 1980. The results were compared with observations on populations in other Frisian lakes, and with the results reported by other authors on brackish-water and marine populations of this species.Three generations were observed: one overwintering generation and two summer generations. Reproduction stopped completely during winter. During 1980 population densities ofN. integer were relatively low in all Frisian lakes. In Slotermeer the maximum population density was only 6 individuals m^–2, production amounted to 10 mg dry wt m^–2 yr^–1. The annual P/B ratio was 4.0. The diet ofN. integer consisted, in terms of biomass, of more than 95% detritus and animal food, the latter mainly consisting ofBosmina and cyclopoid copepods. Feeding intensity reached a maximum at sunset. It was still high during first part of the night and was low in the morning just after sunrise.     

The distribution,life cycle and production of Leptophlebia vespertina (L.) (Ephemeroptera) in a lowland lake

Oak Mere, the water body studied, is a moderately productive base-poor lowland lake. It has zones of submerged marginal vegetation growing on sand at the south-west end and on peat at the north-east end. The distributions of L. vespertina and other macroinvertebrates are described in relation to these two distinct areas. Laboratory experiments on behaviour and survival are described in an attempt to explain the patterns of distribution. The seasonal distribution, variations in population density, spatial distribution variance in relation to mean density, life cycle, length-weight relationship, growth and biomass of L. vespertina are then described during two years at the sandy south-west end. The data are used to estimate annual production. Distribution, life cycle, growth and production are compared with populations in water bodies of lower conductivity in the British Isles and Scandinavia.     

Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

The bacterivory of interstitial ciliates in association with bacterial biomass and production in the hyporheic zone of a lowland stream

Rates of bacteria ingestion by interstitial ciliates were estimated and compared to bacterial biomass and production. Investigation was carried out in the hyporheic zone of a lowland stream. FISH was applied to quantitatively determine bacteria within the ciliate's food vacuoles. To estimate bacteria ingestion rates using FISH, we had to strike a new path. When numbers of bacteria in the food vacuoles remains constant with time (bacterial digestion and ingestion are at equilibrium), ingestion rate can be estimated based on the digestion time and the average number of bacteria per cell. Ciliate community was predominantly composed of bacterivorous ciliates. FISH-signals deriving from ingested bacteria were detected in Cinetochilum margaritaceum, 'other small scuticociliates', Pleuronema spp., and Vorticella spp. Ingestion rates for these taxa were 78, 150, 86, and 38 bacteria ind(-1) h(-1), respectively. The grazing impacts on bacterial biomass and carbon production were calculated based on these ingestion rates. Ciliate grazing caused a decrease in bacterial biomass of 0.024% day(-1) and in bacterial carbon production of 1.60%. These findings suggest that interstitial ciliate grazing impact on bacteria biomass and production was too low to represent an important link in the carbon flow of the hyporheic zone under study.     

Twentyfive years of changes in the distribution and biomass of eelgrass,Zostera marina,in Grevelingen lagoon,The Netherlands

The wax and wane of the eelgrass (Zostera marina L.) population in Grevelingen lagoon (East Atlantic; The Netherlands) has been documented for over 25 years, together with quantitative and semi-quantitative data on environmental variables. The population expanded after the closure of the Grevelingen estuary in 1971, but declined from 4600 ha surface area in 1978 to less than 100 ha in 1993. There is little causal evidence which factors are responsible for the observed dynamics of the population. The incomplete picture emerging from the data is that of an extremely impoverished eelgrass population, living under constant oligo-mesotrophic marine conditions. Both the sexual and the vegetative modes of reproduction are severely stressed by environmental variables, most likely a combination of low temperatures, high salinity, low dissolved silicate and low ammonium concentrations. Survival of the population asks for the restoration of moderate estuarine conditions.Contribution No. 2180 of the Netherlands Institute of Ecology, Nieuwersluis, The Netherlands.