Isotopic composition (C & N) of the suspended particles and N uptake by phytoplankton in a shallow tropical coastal lagoon

The spatial distribution of the C/N ratios and variations in δ¹³C and δ¹⁵N of suspended particulate matter were used to characterise their source in Asia’s largest brackish water lagoon, Chilika, India. In addition, the significance of re-mineralised nutrients in the primary productivity of the shallow lagoon was also determined through quantification of the subsurface nitrogen uptake conditions at two relatively stable locations in the lagoon. The results indicated that the influence of terrestrial organic matter was the maximum in the northern sector and was relatively limited at the central and southern part of the lagoon. In situ ¹⁵N uptake experiments (daytime) under biogeochemically stable conditions revealed that the N uptake by phytoplankton ranged between 0.24 and 1.01?mM?m^?3?h^?1 during pre-monsoon and post-monsoon seasons. New production and regenerated production in the shallow lagoon was also estimated by calculating f-ratios (ratio of nitrate assimilation by phytoplankton to total nitrogenous nutrient assimilation, have been estimated), which varied from 0.52 in the post-monsoon to 0.38 in the pre-monsoon. Lowering of the f-ratio from post- to pre-monsoon indicated a dominance of mineralisation over the new production.     

Modeling the impact of natural and anthropogenic nutrient sources on phytoplankton dynamics in a shallow coastal domain, Western Australia

The influence of different nutrient sources on the seasonal variation of nutrients and phytoplankton was assessed in the northern area of the Perth coastal margin, south–western Australia. This nearshore area is shallow, semi-enclosed by submerged reefs, oligotrophic, nitrogen-limited and receives sewage effluent via submerged outfalls. Analysis of 14 year of field observations showed seasonal variability in the concentration of dissolved inorganic nitrogen and phytoplankton biomass, measured as chlorophyll-a. For 2007–2008, we quantified dissolved inorganic nitrogen inputs from the main nutrient sources: superficial runoff, groundwater, wastewater treatment plant effluent, atmospheric deposition and exchange with surrounding coastal waters. We validated a three-dimensional hydrodynamic-ecological model and then used it to assess nutrient-phytoplankton dynamics. The model reproduced the temporal and spatial variations of nitrate and chlorophyll-a satisfactorily. Such variations were highly influenced by exchange through the open boundaries driven by the wind field. An alongshore (south–north) flow dominated the flux through the domain, with dissolved inorganic nitrogen annual mean net-exportation. Further, when compared with the input of runoff, the contributions from atmospheric-deposition, groundwater and wastewater effluent to the domain’s inorganic nitrogen annual balance were one, two and three orders of magnitude higher, respectively. Inputs through exchange with offshore waters were considerably larger than previous estimates. When the offshore boundary was forced with remote-sensed derived data, the simulated chlorophyll-a results were closer to the field measurements. Our comprehensive analysis demonstrates the strong influence that the atmosphere–water surface interactions and the offshore dynamics have on the nearshore ecosystem. The results suggest that any additional nutrient removal at the local wastewater treatment plant is not likely to extensively affect the seasonal variations of nutrients and chlorophyll-a. The approach used proved useful for improving the understanding of the coastal ecosystem.     

Differences in photosynthetic pigment signatures between phytoneuston and phytoplankton communities in a coastal lagoon of Baja California

In order to understand the relationships between the dynamics of phytoplankton populations in the surface microlayer (MIL) and in the water column below (SSW), this study used high-performance liquid chromatography-derived pigment markers in samples from a coastal lagoon of Baja California (Estero de Punta Banda, EPB) under summer (October 2003) and winter (December 2003) conditions. Photosynthetic pigment signatures of phytoplankton at the air–sea interface (phytoneuston) and subsurface measurements were related to bottom-up (temperature, salinity, nutrient concentrations) and top-down factors (zooplankton abundance). Slicks and scum layers were observed in the inner part of the lagoon and coincided with greater stratification of layers just below the sea surface and lower wind intensities. In general, spatial variability in pigment markers and ancillary data was very high and resulted in non-significant differences between MIL and subsurface samples when different regions of EPB or sampling dates were compared. However, different patterns were found between pigments and environmental factors of MIL and SSW samples when the relative numbers of stations with positive and negative differences (ΔX = X _MIL−X _SSW) were computed. For each survey, pigment markers of phytoneuston and phytoplankton samples were not necessarily correlated. Further analysis revealed that those markers (19′-butanoyloxyfucoxanthin, prasinoxanthin, divinil-chlorophyll a) corresponded to picophytoplankton groups (haptophyte, prasinophyte, and prochlorophyte). On both dates, the MIL was enriched in 19′-hexanoyloxyfucoxanthin (a marker for a type 4 haptophyte) and fucoxanthin (marker for bacillariophytes, haptophytes, and crysophytes) and depleted in peridinin (marker for dinophytes). Different zooplankton grazers accumulated in the MIL (loricate tintinnids) and in SSW (copepod nauplii).     

A numerical investigation of the influence time distribution in a shallow coastal lagoon environment of the Gulf of California

Environmental Fluid Mechanics - The influence time distribution in Santa María La Reforma coastal lagoon, which is one of the most productive ecosystems in Mexico, has been numerically...     

Vertical distribution of phytoplankton during investigations of a natural surface film

An interdisciplinary research program (Kombiniertes Oberflächenfilmprojekt KOFF), in the North Sea area about 75 km west of the Island of Sylt, is directed towards gaining further knowledge on the biological importance of the natural surface film. The investigations include the water body below the film as well as dynamic processes at the surface. Persistence of calm weather conditions (cloudless sky) before and during the investigations permitted observations of the diurnal species-specific vertical migration of Dinophyceae. Two groups could be distinguished: (1) Prorocentrum micans and Ceratium furca; (2) C. horridum, C. macroceros and C. fusus. The active accumulation of P. micans in the slick was verified by a significant correlation coefficient—linearly correlated with the time of day. The results presented reveal continuity between surface film and underlying water column.This work was supported by the Sonderforschungsbereich (SFB) 94, Hamburg, of the Deutsche For-schungsgemeinschaft.     

Depth-dependent responses to solar ultraviolet radiation and oxidative stress in the zooxanthellate coral Acropora microphthalma

Colonies of Acropora microphthalma (Verrill 1869) were transferred from depths of 2 to 3, 10, 20, and 30 m to UV-transparent and UV-opaque respirometry chambers placed at 1 m depth at Bowl Reef, Great Barrier Reef, in March 1989. Peak rates of photosynthesis in colonies originating at 2 and 10 m were unaffected by solar ultraviolet (UV) radiation at 1 m, whereas photosynthesis showed 30 and 38% inhibition in colonies transferred from 20 and 30 m, respectively. This differential sensitivity of corals to UV radiation was consistent with the five- to tenfold higher concentrations of UV-absorbing, mycosporine-like amino acids (MAAs, putative defenses against UV) in 2- and 10-m colonies compared with 20- and 30-m colonies. Photosynthesis in zooxanthellae freshly isolated from 2- and 10-m corals, however, was inhibited by UV, indicating that the host's tissues, which contain 95% of the total MAAs in corals at these depths, are the first line of defense against solar UV and provide protection to their endosymbiotic algae. The general bathymetric decline in the activities of the antioxidant enzyme superoxide dismutase (SOD) in the host, and SOD, catalase, and ascorbate peroxidase in the zooxanthellae, is related to the decrease in potential for photooxidative stress with increasing depth.     

Nutrient enrichment of seagrass beds in a rhode island coastal lagoon

Seagrass and algal beds showed a variety of reponses when the water column was treated with low level additions of ammonium, nitrate and phosphate. The nutrients were added separately to 3 uniform seagrass beds of a temperature coastal lagoon during 1979 and 1980. (1) Ammonium caused the production of dense mats of free-floating green algae Enteromorpha plumosa and Ulva lactuca. It also stimulated growth in both the leaf and root-rhizome fractions of Zostera marina. This growth response in Z. marina was greater in the area where current reached 12 cm · s^-1 than in the area with little or no current. The concentration of nitrogen in the tissue did not change. In contrast, where current was lacking, Z. marina growth increase with ammonium was small, but the concentration of nitrogen in the tissue doubled over that in control plots. The growth of Ruppia maritima was inversely related to the growth of green algae in the same plots. The red alga Gracilaria tikvahiae did not grow better in ammonium, but its tissue reddened. (2) Nitrate additions enhanced the growth of the green seaweeds Enteromorpha spp. and U. lactuca, but not Z. marina or R. maritima. G. tikvahiae, when fertilized in isolation from other plants, showed a marginal response to this nutrient, and the tissue always reddened. (3) Phosphate enhanced growth in Z. marina and R. maritima exposed to moderate current. G. tikvahiae growing alone showed a small growth response to phosphate. The phosphate made no difference in the growth of the green seaweeds. (4) None of the nutrient supplements noticeably altered the species composition of either epiphytic or planktonic algae associated with the beds, although we did detect small increases in their numbers. The rapid and dense growth of green algae in nitrogen-enriched water probably limited growth of adjacent seagrasses and red algae. Because these seaweeds did not use the phosphate, it became available to other plant components. The overall floral response to nutrient addition in seagrass communities depends, therefore, upon the particular nutrient supplied, the ability of alternate species in the area to compete for that nutrient and the velocity of current in the specific area.     

Some quantitative aspects of seagrass ecology in a coastal lagoon of Mauritius

Seagrass distribution was recorded by snorkel dives on a grid of stations in the waterfront of Club Méditerranée at Mon Choisy-Trou Aux Biches lagoon (NW Mauritius) and subsequently mapped using SURFER 6 computer software. Above-ground (AG) and below-ground (BG) standing biomass in terms of dry weight (DW) and ash-free dry weight (AFDW) as well as shoot density and shoot length were monitored monthly from June1997 to May1998 in a mixed stand of Halodule uninervis and Syringodium isoetifolium (dominant) at a shallow, nearshore station in the lagoon. Measurements of physical and chemical parameters [water temperature, current speed, salinity, pH, dissolved oxygen (DO), nitrate and phosphate concentrations] were made simultaneously, as well as at a reference station (ORE) outside the coral reef. The bottom sediment was analysed for grain size and type composition. Variation patterns were examined and statistical correlations drawn to relate plant performance to the environmental variables measured. The SURFER 6 programme generated a satisfactory contour map of seagrass distribution in the lagoon with a cover range of 0-60%. The densest patches occurred adjacent to the shoreline experiencing weaker water currents (3-13 cm s^-1) rather than near the reef (5-35 cm s^-1), where seagrasses were absent. Sand (0.063-2 mm grain size) constituted 97.2% and 77.6% of the nearshore and near-reef sediment, respectively. The dominant grain types were derived from corals (about 80%) and mollusc shells (about 14%). The recorded range of total standing biomass for H. uninervis was 243.1-468.2 g DW m^-2 (326.9ᇛ.7 g) or 71.7-141.2 g AFDW m^-2 (96.8ᆨ.1 g) and for S. isoetifolium it was 271.7-758 g DW m^-2 (460.4끯.1 g) or 119-220.5 g AFDW m^-2 (155.1ᆮ.5 g), with a maximum biomass increase during September-December in both species. AG:BG biomass ratios were generally <1 and approximated 1 during the warmest months of December-February only. Mean shoot density (1,077-4,364 shoots m^-2 in the overall range of 998-4,428 shoots m^-2) and mean shoot length (10.9-20.8 cm in the overall range of 7-31 cm) in S. isoetifolium were higher than in H. uninervis (1,732-4,137 shoots m^-2 in the overall range of 1,522-4,327 shoots m^-2 and 7.9-13.7 cm in the overall range of 6-20 cm, respectively). Temperature showed strong positive correlations with total AFDW biomass of both species (r=0.755, P<0.01 for H. uninervis; r=0.679, P<0.02 for S. isoetifolium) and with DO (r=0.925, P<0.01). High DO levels (10.7-11.2 mg l^-1) coincided with optimum standing biomass at 27.2°C. Correlations were also strong with shoot density (r=0.881, P<0.01 for H. uninervis; r=0.952, P<0.01 for S. isoetifolium) and shoot length (r=0.752, P<0.01 for H. uninervis; r=0.797, P<0.01 for S. isoetifolium). Under optimal environmental conditions, nutrient inputs from surface run-off or underground freshwater seepage in the lagoon due to heavy rainfall may boost up seagrass biomass, as suggested by positive significant correlations between phosphate levels and AG AFDW biomass (r=0.63, P<0.05 for H. uninervis; r=0.65, P<0.05 for S. isoetifolium) and shoot density (r=0.6, P<0.05 for H. uninervis; r=0.687, P<0.02 for S. isoetifolium). The results generated in this study suggest local seagrass standing biomass is comparable to that reported in monospecific stands from elsewhere. Anthropogenic activities increasingly draw down the resilience of the seagrass beds around Mauritius, and preventative measures are indispensable to achieve coastal ecological stability.     

Seasonal succession and diversity of phytoplankton in a eutrophic lagoon (Liman lake)

The seasonal succession, composition and diversity of phytoplankton in a eutrophic lagoon (Liman lake) were studied between January 2002 and November 2003. Samples were collected from surface water and deeper (1 m depth) at stations and species diversity (Shannon-Weaver, H) and evenness were calculated. Shannon Diversty was similar at Station 1 and Station 2. Minimum and maximum diversity values (0.101 and 0.765 bits. mm3) were recorded in June and July 2003 at Station 2. Cluster analysis and NMDS (Non metric multidimensial scaling) were applied to the phytoplankton community. The lagoon exhibits high conductivity (7211-10757 microScm(-1)), mean temperature of 17.7 degrees C, varying concentrations of dissolved oxygen (3.3-8.4 mg l(-1)). Light, temperature, rainfall, turbidity and salinity were expected to be the main factors affecting the seasonal succession. The seasonal succession of phytoplankton were similar at surface water and 1 m depth. A total of 130 taxa belonging to Bacillariophyta, Chlorophyta, Euglenophyta, Cyanophyta, Dinophyta, Xantophyta, Chrysophyta and Cryptophyta divisions were identified. Pseudanabaena limnetica, Kirchneriella obesa, Kirchneriella lunaris, Ankistrodesmus falcatus and Ankistrodesmus spiralis were highly represented.     

Stimulation of phytoplankton production in coastal waters by natural rainfall inputs: Nutritional and trophic implications

Recent evaluations of estuarine and coastal nutrient budgets implicate atmospheric deposition as a potentially significant (20 to 30%) source of biologically available nitrogen. We examined the potential growth stimulating impact of atmospheric nitrogen loading (ANL), as local rainfall, in representative shallow, nitrogen limited North Carolina mesohaline estuarine and euhaline coastal Atlantic Ocean habitats. From July 1988 to December 1989, using in situ bioassays, we examined natural phytoplankton growth responses, as¹⁴CO₂ assimilation and chlorophylla production, to rain additions over a range of dilutions mimicking actual input levels. Rainfall at naturally occurring dilutions (0.5 to 5%) stimulated both¹⁴CO₂ assimilation and chlorophylla production, in most cases in a highly significant manner. Parallel nutrient enrichments consistently pointed to nitrogen as the growth stimulating nutrient source. Generally, more acidic rainfall led to greater magnitudes of growth stimulation, especially at lower dilutions. Nutrient analyses of local rainfall from May 1988 to January 1990 indicated an inverse relationship between pH and NO ₃ ^- content. There have been growing concerns regarding increasing coastal and estuarine eutrophication, including ecologically and economically devastating phytoplankton blooms bordering urban and industrial regions of North America, Europe, Japan, and Korea. It appears timely, if not essential, to consider atmospheric nutrient loading in the formulation and implementation of nutrient management strategies aimed at mitigating coastal eutrophication.     

Simultaneous nitrogen and silicate deficiency of a phytoplankton community in a coastal jet-front

The nutritional status of a phytoplankton community was investigated in a coastal jet-front located in the Gulf of St. Lawrence, Canada, in 1987. During the sampling period, the frontal community was mainly composed of the diatomsChaetoceros debilis, Skeletonema costatum, Thalassiosira gravida andC. pelagicus. As previously reported for the St. Lawrence, some frontal stations were depleted both in nitrate and silicate. At stations impoverished in nitrate, internal nitrate pool concentrations were low or undetectable, suggesting that cells had not, recently, been exposed to a nitrate flux which exceeded the nitrate assimilation rate. At these impoverished stations, however, ambient and intracellular concentrations of ammonium and urea were high, suggesting that the community was not nitrogen-deficient. The comparison between the ambient silicate concentrations and the silicate requirement (K _s ) of the dominant diatoms suggests thatC. debilis andS. costatum were Si-deficient. This is further supported by the low silicate uptake rates and intracellular concentrations measured at the silicate impoverished stations. The silicate deficiency also resulted in a decrease in the seston and phytoplankton N:C ratios.Please address all correspondence and requests for reprints to Dr Levasseur at his new address: Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec G5H 3Z4, Canada     

Modeling approach to regime shifts of primary production in shallow coastal ecosystems

Pristine coastal shallow systems are usually dominated by extensive meadows of seagrass species, which are assumed to take advantage of nutrient supply from sediment. An increasing nutrient input is thought to favour phytoplankton, epiphytic microalgae, as well as opportunistic ephemeral macroalgae that coexist with seagrasses. The primary cause of shifts and succession in the macrophyte community is the increase of nutrient load to water; however, temperature plays also an important role. A competition model between rooted seagrass (Zostera marina), macroalgae (Ulva sp.), and phytoplankton has been developed to analyse the succession of primary producer communities in these systems. Successions of dominance states, with different resilience characteristics, are found when modifying the input of nutrients and the seasonal temperature and light intensity forcing.     

Irradiance reduction: Effects on standing crops of the eelgrass Zostera marina in a coastal lagoon

Abundance of the eelgrass Zostera marina L. was studied in a coastal lagoon in southern California (USA), and was found to correlate with the level of irradiance at depths greater than 0.5 m below tidal datum. Results of controlled field experiments, using canopies to reduce downwelling illuminance by 63%, confirmed that turion density is a function of the irradiance the plants receive. By Day 18 of the experiment, turion density in the shaded experimental areas had decreased compared to the density of adjacent unshaded controls. Turion densities were continually lower throughout the 9-month study in the experimental areas, which at the end of the study had a turion density only 5% that of the control areas. Flowering in the experimental areas was also inhibited by shading. The biological implications of these findings are discussed with respect to seasonal changes in incident solar radiation, water transparency, and changes in water quality due to man's increased intervention in the natural processes of coastal lagoons.Contribution No. 4 from the Center for Marine Studies, San Diego State University, San Diego, California 92182, USA.     

Cadmium uptake by a phytoplankton species in the presence of different inhibitors&

The uptake of cadmium by the Haptophyceae Hymenomonas elongata was studied as regards the energetic processes and Ca‐transport. For this purpose, experiments were carried out in the presence of an uncoupler of the phosphorylation : CCCP and of an inhibitor of calcium channels and also of Ca²⁺‐ATPase : verapamil. To reduce the number of assays, a factorial experimental design was used, in which all variables:— concentration of Cd in the medium—incubation time and—the presence of the inhibitor, are at two levels. In the absence of inhibitor, Cd uptake was found to vary as a function of metal concentration in the medium and of incubation time. CCCP significantly increased Cd uptake by H. elongata at sufficient incubation time. Therefore Cd uptake seemed linked to an energy‐dependent process, involving an ATPase. Verapamil immediately increased Cd uptake, implying an interaction between Cd and Ca.     

Relationships between nutrients and macroalgal biomass in a brazilian coastal lagoon: the impact of a lock construction

The Piratininga Lagoon is a coastal, choked, and brackish ecosystem in SE Brazil, where uncontrolled discharge of domestic sewage led to eutrophication, increasing massive proliferation of benthic macroalgae, and decrease of the lagoon mean depth. In 1992, a dam was constructed by the local Municipality at its tidal channel aiming at stabilising its water level. Main physico-chemical parameters, together with macroalgae biomass, nutrient concentrations in the water column, particulate organic carbon, and chlorophyll a were recorded monthly at 4 sampling stations within the Piratininga lagoon from April 1994 to April 1995. The data, compared with “before-lock” existing studies, show that nutrients and chlorophyll a concentrations significantly increased after the lock construction. Based on the functioning of the ecosystem, we propose to harvest the algal mats before their decomposition period in order to partly remove the nutrient stocks from the lagoon and the future ecosystem modelling to predict the impact of natural and anthropogenic eutrophication.     

Relationships between nutrients and macroalgal biomass in a brazilian coastal lagoon: the impact of a lock construction

The Piratininga Lagoon is a coastal, choked, and brackish ecosystem in SE Brazil, where uncontrolled discharge of domestic sewage led to eutrophication, increasing massive proliferation of benthic macroalgae, and decrease of the lagoon mean depth. In 1992, a dam was constructed by the local Municipality at its tidal channel aiming at stabilising its water level. Main physico-chemical parameters, together with macroalgae biomass, nutrient concentrations in the water column, particulate organic carbon, and chlorophyll a were recorded monthly at 4 sampling stations within the Piratininga lagoon from April 1994 to April 1995. The data, compared with “before-lock” existing studies, show that nutrients and chlorophyll a concentrations significantly increased after the lock construction. Based on the functioning of the ecosystem, we propose to harvest the algal mats before their decomposition period in order to partly remove the nutrient stocks from the lagoon and the future ecosystem modelling to predict the impact of natural and anthropogenic eutrophication.     

Fishery resource management in Chilika lagoon: a study on coastal conservation in the Eastern Coast of India

A significant proportion of the fishing population comprises small-scale fishermen and many studies illustrate that these people are exploited by middlemen in the process of fish marketing combined with money lending. The negative dependency gives rise to poverty and triggers indiscriminate fish catch that threatens fishery resources depletion. This article explores the root causes of failures in resource-led development from the viewpoint of coastal resource conservation. The study presents a case study of Chilika lagoon, India and focuses on the interaction between small-scale fishermen and middlemen. The findings reveal that most of the small-scale fishermen have been exploited by specific middlemen and the underlying causes of the present fish marketing structure stem from (i) indebtedness and (ii) the unstable situation because of perpetual conflicts over fishery resources among the fishers across Chilika lagoon. Based on these observations, this article presents some recommendations on fishery resource conservation from the perspective of a fish marketing structure.

Natural attenuation of MTBE and BTEX compounds in a petroleum contaminated shallow coastal aquifer

Natural attenuation of MTBE and BTEX compounds in a petroleum contaminated coastal aquifer in the Tel-Aviv area was investigated. Significant decrease in MTBE concentration and complete disappearance of BTEX compounds occur within 100 m groundwater flow. Highly anaerobic conditions were determined in the close vicinity to the spill source. In order to examine the contribution of microbial degradation to the attenuation process at anaerobic conditions, compound-specific carbon isotope ratio analysis was employed. Carbon isotope enrichment of toluene up to 2.4% along with the drop in its concentration up to 80% was observed within 20 cm below the water table. The lack of isotope fractionation despite the significant concentration decrease for other studied compounds indicates significant contribution of abiotic processes to the natural attenuation.     

The nature of the photosynthate in natural phytoplankton populations in relation to light quality

The relative concentrations of a variety of organic compounds in phytoplankton from different depths of the photic zone of Saanich Inlet and Indian Arm, British Columbia, Canada, were determined. In surface phytoplankton the greatest proportion of ¹⁴C in newly-formed organic compounds was in the ethanol-soluble fraction irrespective of the light intensity. The proportion of the label in the insoluble fraction increased with depth. Within the ethanol-soluble fraction the relative importance of carbohydrates decreased and amino acids increased with depth. Experiments in Indian Arm indicated that these observed changes were in response to light quality rather than intensity. The percentage release of dissolved organic carbon as a percentage of total fixation decreased with depth and was directly related to the size of the ethanol-soluble fraction.     

Biochemical fractionation of primary production by phytoplankton in Belgian coastal waters during short- and long-term incubations with 14C-bicarbonate

Short- and long-term time course studies of radiocarbon accumulation in the intracellular end-products of photosynthesis (proteins, polysaccharides, lipids, small metabolites) and extracellular monomers and polymers were conducted at natural light intensity during a 24-h period in Belgian coastal waters dominated by large diatoms species in September, 1983. It is shown that carbon losses observed during the long-term incubation are due to the catabolism of reserve products (polysaccharides and lipids), which occurs both during the light and dark periods and provides carbon and energy for pursuing protein synthesis during the dark. Catabolism rates, as calculated by means of a simple mathematical model, indicate reduced rates of lipid catabolism (1–2% h^-1, respectively for the light and dark periods), although polysaccharide catabolism proceeds at much higher rates, namely 20% h^-1 during the light and 8% h^-1 during the dark period. Assuming that protein synthesis proceeds at a constant rate during the 24-h period and that 1–3 glucan constitutes the main storage product of this diatom population, it is shown that at least 65% of the gross primary production is catabolized by the cells. From this, only 16% are mobilized for dark protein synthesis. The remaining is respired, especially during the light period.