Light absorption and photosynthesis of a benthic moss community: importance of spectral quality of light and implications of changing light attenuation in the water column

1. The underwater light climate and benthic moss communities of Grane Langsø were investigated in May 1997 to determine the potential effects on benthic production of changing water column attenuation and spectral quality of light.
2. A reduction in water clarity in the lake since the 1960s was manifested as a marked increase in the attenuation of blue light, relative to red light, which can be attributed to increased dissolved organic carbon.
3. The biomass of the benthic moss community ranged from a maximum of 195 gDW m⁻² at a depth of 4 m to 39 g DW m⁻² at a depth of 10 m and Drepanocladus exannulatus contributed 70% of the biomass at all depths.
4. Absorption of PAR by D. exannulatus was maximal in the highly pigmented youngest parts of the plant and these correspondingly showed the highest rates of net photosynthesis. The absolute amount of chlorophyll- a per g dry weight was greater at 10 m than 2 m, but the ratio of accessory pigments to chlorophyll- a did not change. Deep growing plants did not show adaptation to changed light quality.
5. Increased attenuation of blue light and the resultant overall decrease in water clarity is likely to impact negatively on net annual production of benthic macrophytes of Grane Langsø. Any further increase in dissolved organic carbon concentration has the potential to markedly decrease the depth to which mosses grow by reducing the length of time in a year during which net photosynthesis occurs.     

HETEROTROPHIC GROWTH OF ULVA LACTUCA (CHLOROPHYCEAE)1

The effect of external glucose (51 mM) and acetate (13 mM) on growth and photosynthetic capacity of Ulva lactuca L. was tested in laboratory cultures over 41 days in the dark and in dim light (0.9 μmol photons·m^?2·s^?1) at 7–8° C. Glucose and acetate had a significant positive effect on growth rate, chlorophyll content, and quantum yield for discs grown in the dark and in dim light. The carbon gain from heterotrophic uptake was low and only allowed U. lactuca to maintain a specific uptake was low and only allowed U. lactuca to maintain a specific growth rate of 0.005 day^?1 compared to 0.06–0.1 day^?1 at higher light intensities. However, plants with added organic substrate maintained a normal chlorophyll content and were able to photosynthesize whereas control plants lost pigmentation and photosynthetic capability after 41 days in both dim light and darkness, probably because of disorganization of the photosynthetic apparatus. This suggest that the ecological significance of heterotrophic uptake is to allow U. lactuca to survive during prolonged low light conditions with an intact photosynthetic apparatus.     

Pelagic carbon metabolism in a eutrophic lake during a clear-water phase

Dissolved and paniculate organic carbon (DOC and POC, respectively),primary production, bacterial production, bacterial carbon demandand community grazing were measured for 9 weeks in eutrophicFrederiksborg Slotssø. The period covered the declineof the spring bloom, a clear-water phase and a summer phasewith increasing phytoplankton biomass. The process rates andchanges in pools of organic carbon were combined in a carbonbudget for the epilimnion. The POC budget showed a close balancefor both the post-spring bloom and the clear-water phase, whilea surplus was found in the summer phase. Production of POC wasdominated by phytoplankton (2/3) compared to bacteria (1/3)during all phases, and there was a significant correlation betweenphytoplankton and bacterial production rates (r² = 0.48, P <0.039). Bacterial demand for DOC was balanced by productionand changes in the pool of DOC during the decline of the springbloom, but the calculated demand exceeded the supply by 81 and167%, respectively, during the other two periods. The discrepancywas most probably due to an underestimation of bacterial growthefficiency and an overestimation of in situ bacterial productionin carbon units. Production of bacterial substrate by zooplanktonactivity was estimated to be higher than the direct excretionof organic carbon from phytoplankton. The biological successionwas regulated by the balance between area primary productionand community grazing. The clear-water phase was initiated bya combination of low primary production due to low surface irradianceand high community grazing (100 mmol C m^–2 day^–1),which caused a decrease in phytoplankton biomass. However, dueto the high initial phytoplankton biomass, community grazingwas not high enough to cause a significant decrease in areaprimary production. The summer phase was initiated by a decreasein community grazing followed by an increase in phytoplanktonbiomass. Based on these observations and calculations of areaprimary production as a function of chlorophyll concentrations,we suggest that the possibility for zooplankton to regulatephytoplankton biomass in temperate lakes decreases with increasingnutrient level.     

Bioenergy potential of Ulva lactuca: biomass yield, methane production and combustion

The biomass production potential at temperate latitudes (56°N), and the quality of the biomass for energy production (anaerobic digestion to methane and direct combustion) were investigated for the green macroalgae, Ulva lactuca. The algae were cultivated in a land based facility demonstrating a production potential of 45 T (TS) ha⁻¹ y⁻¹. Biogas production from fresh and macerated U. lactuca yielded up to 271 ml CH₄ g⁻¹ VS, which is in the range of the methane production from cattle manure and land based energy crops, such as grass-clover. Drying of the biomass resulted in a 5-9-fold increase in weight specific methane production compared to wet biomass. Ash and alkali contents are the main challenges in the use of U. lactuca for direct combustion. Application of a bio-refinery concept could increase the economical value of the U. lactuca biomass as well as improve its suitability for production of bioenergy.     

Seasonal dynamics and bioavailability of dissolved organic matter in two contrasting temperate estuaries

Production and bioavailability of dissolved organic matter (DOM) were followed during a year in the nutrient-rich estuary, Roskilde Fjord (RF), and the more oligotrophic strait, Great Belt (GB), in Denmark. Bioavailability of dissolved organic carbon (DOC), nitrogen (DON), and phosphorous (DOP) was determined during incubations over six months. Overall, RF had three to five times larger pools of total nitrogen (TN) and total phosphorous (TP) and five to eight times higher concentrations of inorganic nutrients compared to GB. However, the allocation of carbon, nitrogen, and phosphorous into different pools were remarkably similar between the two systems. DON and DOP contributed with about equal relative fractions in the two systems: 72 ± 13% of total nitrogen and 21 ± 12% of total phosphorous. The average bioavailability of DOM was 25 ± 15, 17 ± 5.5, and 49 ± 29% for carbon, nitrogen, and phosphorous, respectively. The observed release of DIN from degradation of DON amounted to between 0.1 (RF winter) and 14 times (GB summer) the loadings from land and contributed with half of the total input of bioavailable nitrogen during summer. Hence, this study shows that nitrogen in DOM is important for the nitrogen cycling, especially during summer. The sum of inorganic nutrients, particulate organic matter, and bioavailable DOM (the dynamic pools of nutrients) accounted for 42 and 92% of nitrogen, and phosphorous, respectively, and was remarkably similar between the two systems compared to the difference in nutrient richness. It is hypothesized that the pelagic metabolism of nutrients in marine systems dictates a rather uniform distribution of the different fractions of nitrogen and phosphorous containing compounds regardless of eutrophication level.     

Water clarity and eelgrass responses to nitrogen reductions in the eutrophic Skive Fjord, Denmark

Eelgrass depth limits and water clarity in the Skive Fjord estuarine system have not improved despite nutrient input reductions of 30%. Long-term monitoring data (1989–2010) were used to investigate the underlying causes. Dissolved inorganic and organic nitrogen concentrations decreased significantly over time, whereas particulate organic nitrogen concentration, assumed to consist primarily of phytoplankton and phytoplankton detritus and calculated as a proportional factor to chlorophyll a, did not change. Total organic carbon, mostly of autochthonous origin, remained constant despite reduced nitrogen concentrations, resulting in an increasing C:N ratio of the organic material in the water column. Phytoplankton primary production also remained constant suggesting that phytoplankton growth was only limited by nitrogen to a minor degree. Alleviated grazing pressure caused by a reduction in the blue mussel standing stock and a pelagic food web dominated by jellyfish may have contributed to the constantly high phytoplankton levels. Particulate inorganic matter, likely reflecting sediment resuspension, increased over time, most probably in response to removal of blue mussels and declining eelgrass cover. The Skive Fjord estuarine system is affected by multiple pressures—nutrient enrichment, mussel dredging and climate change that must be addressed together for water clarity to improve and eelgrass to recover.     

Dark uptake of inorganic14C in oligotrophic oceanic waters

Dark uptake of inorganic ¹⁴C by offshore plankton was measuredat two depths at 36 stations in the Atlantic Ocean from 52°Sto 26°N, mainly along 30°W. The samples were incubatedfor 2 h with and without inhibition of biological activity withHgCl₂. In addition, six time course experiments were performed.The mean dark uptake rate varied from 0.68 to 4.82 (µmolC m^–3 h^–1 over the transect and showed a significantpositive relationship with chlorophyll a. The dark uptake wasusually >5% of the maximum photosynthetic capacity (P^m),and higher values relative to P^m were associated with low valuesof P_m and not with high absolute dark values. A linear relationshipbetween dark uptake and P_m was found with a background value(y-axis intercept) of 0.51 (µmol C m^–3 h^–1and a slope of 0.77% of P^m. A major fraction of the dark signal,66–80% of the total signal, persisted in bottles treatedwith HgCl2, indicating that most of the dark signal was independentof biological activity. Time course experiments showed a lineardark uptake with time for the first hours, whereafter the uptakeceased. At stations with low concentrations of inorganic nitrogen[>1 (µmol (NH₄⁺+NO₃^–)], a second stage was observedafter 3–8 h, probably due to an increase in bacterialactivity. The results suggest three mechanisms for the darkvalue in short-term incubations in oligotrophic waters. A backgroundvalue independent of biomass and incubation time which was thedominant part of the dark signal in samples with very low phytoplanktonbiomass (>0.3 p-g Chi a 1"). Another important part was residualsof ¹⁴C associated with plankton, probably adsorbed to compoundsinside the cells. This fraction was dominant in short-term incubationsat chlorophyll concentrations >0.3 p.g Chi a H. Active uptakeby living cells (total minus ‘HgCl2 uptake‘) wasonly a minor part of the dark signal in short-term incubations,but dominated at longer incubation time (>3–9 h), probablydriven by an increase in bacterial activity. A significant enhancementof the non-photosynthetic uptake of ¹⁴C was observed in light,probably associated with a carbon-concentrating mechanism inphytoplankton or light stimulation of ß-carboxylationactivity. The results strongly suggest that dark values shouldbe subtracted from the light uptake. This correction is particularlyimportant when photosynthetic rates are low, e.g. at low lightor in short-term incubations where a time-zero background becomesa significant part of the total uptake in light. Present address: National Environmental Research Institute,Department of Marine Ecology and Microbiology, Frederiksborgvej399, PO Box 358, DK-4000 Roskilde, Denmark     

Diel changes in dark respiration in a plankton community

The dark respiration of a natural plankton community from an eutrophic lake was studied in a laboratory scale enclosure (LSE), exposed to illumination which simulated natural light conditions in the water column. The dark respiration was measured continuously for 2 hours in samples obtained from the LSE each hour for 26 hours. The relationships between dark respiration rates, carbohydrate concentrations and other parameters were investigated.The dark respiration rate showed an exponential decrease with time in the dark in all light period incubations with a time coefficient of 0.3 h^–1. The decrease in respiration rate in the dark period was much slower, reaching an approximately constant level at the end of the night. The overall dark period decline in respiration rate also exhibited an exponential pattern, but with a much lower time coefficient (0.04 h ^–1) than for the light period incubations. A linear relationship was found between dark respiration rate and carbohydrate concentration at night time but no relationship was apparent during the day. A comparison between these data and data from the literature show that this pattern of dark respiration rate decrease with time in the dark may have some general applications for dense phytoplankton communities.     

LIGHT ABSORPTION AND QUANTUM YIELD FOR GROWTH IN FIVE SPECIES OF MARINE MACROALGA1

Light utilization efficiency in five species of marine macroalgae was measured in laboratory growth experiments (13–41 days duration) at different irradiances at 7°C. All species acclimated to irradiance by changing their light absorption, resulting in a peak in light absorption between 2 and 15 μmol·m^?2.s^?1. Light absorption increased with thallus-specific chlorophyll and carbon content according to linear inverse relationships between chlorophyll content (chlarea^?1) and log[transmission] and between log[carbon content, Carea^?1] and log[transmission]. Quantum yields for light-limited growth and estimated gross photosynthesis were calculated based on incident and absorbed light. Quantum yield for photosynthesis based on light absorbed by pigments was high (mean = 114 mmol C·mol^?1 photons) and similar among the species. Quantum yield for net growth based on incident light was also high but more variable, between 22 and 75 mmol C·mol^?1 photons. Differences among species were mainly due to differences in light absorption. In conclusion, all species acclimated to low light by increasing light absorption to the maximum attainable, and growth efficiencies based on absorbed light were close to the maximum theoretically possible.