Primary production in a tropical reservoir in Sri Lanka*

Primary production was measured every 2 weeks during 16 months (N = 33) in Tissawewa, a tropical shallow reservoir in the lowlands of south-east Sri Lanka. Results are interpreted in relation to selected environmental conditions such as oxygen concentrations, water temperature, Secchi-disc depth, wind force, conductivity, and morphoedaphic index and water level fluctuations. Because of regularly reoccurring high wind speeds the water column is well mixed. Daily gross primary production per unit area was plotted as a function of the algal biomass per unit area over the euphotic zone. Chlorophyll-a concentration in the euphotic zone was taken as measure for the algal biomass. The literature comparisons showed that the primary productivity in Tissawewa was in the same range as in 29 tropical lakes and reservoirs, of which 27 were from Africa. The productivity of these 30 tropical lakes and reservoirs was compared with: (a) 27 lakes of which 25 were temperate lakes, and (b) 49 North American temperate lakes. Firstly, comparisons were made on an annual basis for the tropical water bodies, but restricted to May–September, the growing season, for temperate water bodies. The gross primary productivity of tropical water bodies was ca. three times higher than that of temperate water bodies. These differences were even more dramatic if the two geographical regions are compared on an annual basis, i.e. the tropical systems are ca. six times more productive than their temperate counterpart.     

Seasonal Dynamics of Periphyton in a Large Tropical Lake

Tropical aquatic systems are generally assumed to have little seasonality in productivity patterns. However, this study indicated that there was substantial seasonal variation in epilithic productivity and biomass in tropical Lake Tanganyika, due primarily to seasonal patterns in lake hydrodynamics that influence nutrient availability. Although they support much of the lake’s biological diversity, epilithic algae made a minor contribution to the total energy budget in Lake Tanganyika. A comparison among large, oligotrophic lakes revealed no significant latitudinal trends in periphyton productivity or biomass. However, Lake Tanganyika has relatively low benthic algal biomass and is therefore more efficient at photosynthesis than the temperate lakes. The influence of wave action and consumer density and diversity may be important in moderating productivity of the epilithic community.     

Some aspects of photosynthetic characteristics in a set of perennial irrigation reservoirs located in five river basins in Sri Lanka

Phytoplankton primary productivity of eleven irrigation reservoirs located in five river basins in Sri Lanka was determined on a single occasion together with light climate and nutrient concentrations. Although area-based gross primary productivity (1.43–11.65 g O₂ m^–2 d^–1) falls within the range already established for tropical water bodies, net daily rate was negative in three water bodies. Light-saturated optimum rates were found in water bodies, with relatively high algal biomass, but photosynthetic efficiency or specific rates were higher in water bodies with low algal biomass, indicating nutrient limitation or physiological adaptation of phytoplankton. Concentrations of micronutrients and algal biomass in the reservoirs are largely altered by high flushing rate resulting from irrigation release. Underwater light climate and nutrient availability control the rate of photosynthesis and subsequent area-based primary production to a great extent. However, morpho-edephic index or euphotic algal biomass in the most productive stratum of the water column is not a good predictor of photosynthetic capacity or daily rate of primary production of these shallow tropical irrigation reservoirs.     

Divergent response of seasonally dry tropical vegetation to climatic variations in dry and wet seasons

Interannual variations of photosynthesis in tropical seasonally dry vegetation are one of the dominant drivers to interannual variations of atmospheric CO₂ growth rate. Yet, the seasonal differences in the response of photosynthesis to climate variations in these ecosystems remain poorly understood. Here using Normalized Difference Vegetation Index (NDVI), we explored the response of photosynthesis of seasonally dry tropical vegetation to climatic variations in the dry and the wet seasons during the past three decades. We found significant (p < 0.01) differences between dry and wet seasons in the interannual response of photosynthesis to temperature (γ^int) and to precipitation (δ^int). γ^int is ~1% °C^?1 more negative and δ^int is ~8% 100 mm^?1 more positive in the dry season than in the wet season. Further analyses show that the seasonal difference in γ^int can be explained by background moisture and temperature conditions. Positive γ^int occurred in wet season where mean temperature is lower than 27°C and precipitation is at least 60 mm larger than potential evapotranspiration. Two widely used Gross Primary Productivity (GPP) estimates (empirical modeling by machine‐learning algorithm applied to flux tower measurements, and nine process‐based carbon cycle models) were examined for the GPP–climate relationship over wet and dry seasons. The GPP derived from empirical modeling can partly reproduce the divergence of γ^int, while most process models cannot. The overestimate by process models on negative impacts by warmer temperature during the wet season highlights the shortcomings of current carbon cycle models in representing interactive impacts of temperature and moisture on photosynthesis. Improving representations on soil water uptake, leaf temperature, nitrogen cycling, and soil moisture may help improve modeling skills in reproducing seasonal differences of photosynthesis–climate relationship and thus the projection for impacts of climate change on tropical carbon cycle.     

Invasion of ancient Lake Titicaca by the globally invasive Physa acuta (Gastropoda: Pulmonata: Hygrophila)

A vital population of Physa acuta, a globally invasive species of freshwater gastropods was dicovered in ancient Lake Titicaca in April 2007. The population was found near Chucuito (Bahia de Puno) on the Peruvian side in close proximity to a boat landing of the aquaculture facility of the University of Puno. Physids occured in great abundances at this site on macrophytes in depths from 0 to 2 m. Physa acuta was associated with members of the Heleobia andicola species complex as well as with Heleobia otorni, Biomphalaria andecola, and Uncancylus crequi. Species identity was confirmed by morphological and molecular methods. We discuss the mode of introduction and raise a cautionary note on the potential impact of this global invader on the endemic gastropod fauna of ancient Lake Titicaca.     

COMPARISONS OF SALT-MARSH FUCOID PRODUCTION ESTIMATED FROM THREE DIFFERENT INDICES1

Production of Ascophyllum nodosum (L.) LeJolis ecads and Fuscus vesiculosus L. was calculated from measurements of in situ growth, seasonal variations in standing-crops and seasonal variations in photosynthetic capacity. A computer model for predicting daily, monthly and yearly net production from photosynthesis data was constructed. This model used daily irradiation, actual biomass of algae/m² contributing to production and photosynthesis vs. light intensity relationships as data inputs. Comparison of production estimated from in situ growth, standing-crops and photosynthesis indicated that both marsh fucoids turn over biomass twice per year. Total net production of both fucoids, estimated from photosynthesis data, was ca. 315 g C · m^?2· yr^?1. On the other hand, production of both fucoids calculated from standing-crop data was only 155 g C · m^?2· yr^?1.     

Coupling dissolved organic carbon,CO2 and productivity in boreal lakes

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PHOTOSYNTHESIS/PHOTON FLUX DENSITY RELATIONSHIPS AMONG COMPONENTS OF CORAL REEF ALGAL TURFS1

Photosynthesis/photon flux density (PFD) relationships were compared among some of the major components of sparse algal turfs (green endoliths, red crusts, red filaments, Crouania, Sphacelaria)/ from Tague Bay forereef St. Croix, U.S. Virgin Islands. Algal turfs grazed by the sea urchin Diadema antillarum were 2–10 times more productive per unit chl a than when not grazed by sea urchins. The maximum rate of net photosynthesis (Pnet_max) and saturation PFDs, but not the slope of the light-limited portion of the curve (α), differed significantly among the algal turf components examined. The hypothesis that increased biomass-specific primary productivity results from shifts in algal community structure was not supported because the maximum difference in photosynthesis between algal components was only a factor of two, and the less productive components were relatively more abundant under grazing. In the understory, green endoliths exhibited higher α and lower Pnet_max, suggesting shade adaptation. Photon flux density measurements taken with a fiber optic microprobe within the algal turfs indicated that photosynthesis of basal portions of algal filaments and red crusts are light-limited in ungrazed algal turfs. As self-shading changes with grazer-mediated canopy removal, algal sublayers will contribute differentially to whole turf primary productivity.     

Determining algal assemblages in oligotrophic lakes and streams: comparing information from newly developed pigment/chlorophyll a ratios with direct microscopy

1. Pigment analyses by high performance liquid chromatography (HPLC) are commonly used for determining algal groups in marine and estuarine areas but are underdeveloped in freshwaters. In this study, 15 characteristic pelagic algal species (representing five algal groups) of oligo‐ / mesotrophic lakes were cultured and pigment / Chl a ratios determined at three light intensities. 2. With the exception of cyanophytes, light treatment had little effect on pigment / Chl a ratios. This justifies the use of the same pigment / Chl a ratios during seasonal studies where light conditions may change. 3. The determined pigment / Chl a ratios were tested on seasonal samples from five oligo‐ / mesotrophic lakes and three streams using CHEMTAX software. Pigment ratios of both pelagic and benthic algal communities from the lakes and streams were analysed to determine whether the pelagic algae‐based ratios can be used for benthic algal communities. 4. HPLC combined with CHEMTAX was useful for identifying freshwater phytoplankton classes and for quantifying the abundance of phytoplankton groups. However, although correlations were significant for six of seven phytoplankton classes studied, they were weak and varied with season. 5. HPLC was valid for quantifying benthic diatom groups in stream samples, whereas for lakes more benthic algal groups were recorded with HPLC than with microscopy and correlations between the two methods were not significant. 6. The use of both HPLC and microscopy is recommended as a cost‐efficient method for analysing many samples. It is crucial, however, that the CHEMTAX software is calibrated with the correct information, and the user is aware of the limitations.     

Vegetation clumping modulates global photosynthesis through adjusting canopy light environment

The spatial dispersion of photoelements within a vegetation canopy, quantified by the clumping index (CI), directly regulates the within-canopy light environment and photosynthesis rate, but is not commonly implemented in terrestrial biosphere models to estimate the ecosystem carbon cycle. A few global CI products have been developed recently with remote sensing measurements, making it possible to examine the global impacts of CI. This study deployed CI in the radiative transfer scheme of the Community Land Model version 5 (CLM5) and used the revised CLM5 to quantitatively evaluate the extent to which CI can affect canopy absorbed radiation and gross primary production (GPP), and for the first time, considering the uncertainty and seasonal variation of CI with multiple remote sensing products. Compared to the results without considering the CI impact, the revised CLM5 estimated that sunlit canopy absorbed up to 9%–15% and 23%–34% less direct and diffuse radiation, respectively, while shaded canopy absorbed 3%–18% more diffuse radiation across different biome types. The CI impacts on canopy light conditions included changes in canopy light absorption, and sunlit–shaded leaf area fraction related to nitrogen distribution and thus the maximum rate of Rubisco carboxylase activity (V_cmax), which together decreased photosynthesis in sunlit canopy by 5.9–7.2 PgC year⁻¹ while enhanced photosynthesis by 6.9–8.2 PgC year⁻¹ in shaded canopy. With higher light use efficiency of shaded leaves, shaded canopy increased photosynthesis compensated and exceeded the lost photosynthesis in sunlit canopy, resulting in 1.0 ± 0.12 PgC year⁻¹ net increase in GPP. The uncertainty of GPP due to the different input CI datasets was much larger than that caused by CI seasonal variations, and was up to 50% of the magnitude of GPP interannual variations in the tropical regions. This study highlights the necessity of considering the impacts of CI and its uncertainty in terrestrial biosphere models.     

Temporal variability in algal biomass and primary productivity in Florida lakes relative to latitudinal gradients,organic color and trophic state

Seasonal patterns in primary productivity and algal biomass in subtropical Florida lakes along increasing gradients of both dissolved organic color and phytoplankton biomass are presented. Chlorophyll a concentrations and gross primary productivity generally reached maxima during the summer and were most depressed in winter months, regardless of color or trophic classification. Primary productivity was more strongly correlated with chlorophyll a, nutrient concentrations and water clarity in clearwater (< 75 Pt units) than in colored (> 75 Pt units) systems. Amplitudes in algal biomass were considerably smaller than temperate lakes. Variability in primary production in Florida lakes was intermediate to patterns in the temperate zone and tropics, but was more closely aligned to northern latitudes. Within the Florida peninsula, variability of primary productivity decreased from north to south and corresponded to latitudinal gradients in climatic regimes.     

Temporal variability of phytoplankton in tropical lakes

Summary Temporal variability of ecological systems continues to receive theoretical and empirical attention but remains inadequately documented at low latitudes. Results of my comparative investigation of photosynthetic rates of phytoplankton in 6 equatiorial African lakes and similar information from 20 South American, Asian and African lakes studied by others provide the data for an assessment of the range of seasonal variability (expressed as coefficients of variation, CV) among tropical lakes. Sampling intervals varied from 1 week to 3 months and usually spanned at least one year. Within Africa the coefficient of variation ranged from 15% to 61%, and among all the lakes the coefficient of variation ranged from 15% to 86%. The Spearman rank correlation coefficient of the CV's of photosynthesis versus latitude is 0.24 and is not significant at the 0.05 level.Coefficients of variation of photosynthetic rates of phytoplankton in a diverse set of 45 temperate and arctic lakes ranged from 29% to 155% and were significantly different from the set of 26 tropical lakes by the Mann-Whitney U test. When all 71 lakes are compared, the Spearman rank correlation coefficient of CV's of photosynthesis versus latitude is 0.71 and is significant at the 0.0005 level.Three temporal patterns were recognized among tropical lakes. Most tropical lakes exhibit pronounced seasonal fluctuations that usually correspond with variations in rainfall, river discharges or vertical mixing. A second pattern occurs in lakes with muted fluctuations (coefficient of variation less than 20%) in which diel changes often exceed month to month changes. A third pattern is distinguished by an abrupt change from one persistent algal assemblage (i.e., extant for at least 10 generations) and level of photosynthetic activity to another persistent condition.     

Influence of Algal Photosynthesis on Biofilm Bacterial Production and Associated Glucosidase and Xylosidase Activities

Natural photosynthetic biofilms were incubated under light (100 mmol m-2 s-1) and dark conditions to elucidate the impact of photosynthesis on bacterial production, abundance, biovolume, biomass, and enzyme activities over 24 h. Use of organic carbon-free media limited carbon sources to algal photosynthesis and possibly the polysaccharides of the biofilm matrix. Bacterial production of biofilm communities was significantly higher in light incubations (p <0.001). The greatest differences in production rates between light and dark incubations occurred between 8 and 24 h. Biomass-specific a- and b-glucosidase and b-xylosidase activities were stimulated by photosynthesis, with significantly greater activities occurring at hours 16 and 24 in the light treatment (p <0.01). The results indicate that algal photosynthesis can have a significant impact on bacterial productivity, biomass, biovolume, and enzyme production over longer time periods at low photon flux densities (?100 mmol m-2 s-1).     

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.     

The upper limit of photosynthetic productivity by phytoplankton: evidence from Ethiopian soda lakes

Photosynthetic production by phytoplankton was studied in two Ethiopian soda lakes, alkalinity 51-67 m-equiv./l, with abundant blue-green algae. The deeper lake, L. Aranguadi, contained extremely dense crops composed almost entirely of SpiruUna (Oscillatoria, Arthrospira) platensis. Measurements of the spectral attenuation of light showed that the most penetrating component was displaced to the red spectral region, and estimates of the chlorophyll a content in umt area of the euphotic zone were often similar to the highest values (～200-300 mg/m²) expected on theoretical grounds. The vertical distribution of photosynthetic activity per unit water volume was of a typical pattern for phytoplankton, with light-inhibition often present, although the euphotic zones did not exceed 0.6 m in depth. From these profiles, computed rates of gross photosynthesis per unit area of lake surface reached a magnitude of 1.4–2.4 g O₂/m².h in both lakes. They were obviously severely limited by self-shading behaviour i n the algal populations. An appreciable inverse relationship between photosynthetic capacity and population density was only found in the densest populations of L. Aranguadi, with chlorophyll a content > 2000 mg/m³, where depressed rates might result from experimental artefacts in closed bottles. In this lake, two independent estimates of gross production, based on the analysis of diurnal changes in the open water, were as high as 43 and 57 g O₂/m². day. Diurnal changes of stratification in the two lakes are described and related to the controlling temperature (density) stratification. They include occasionally complete nocturnal deoxygenation in the deeper lake, evidence of heavy respiratory uptake. The high photosynthetic productivity is interpreted as dependent upon the coupling of high algal contents in the euphotic zone (^S) with high values of photosynthetic capacity (^max)- It is probably favoured by the tropical situation affecting temperature and illumination, by a surplus of dissolved inorganic phosphate, and especially by the considerable reserves of carbon dioxide in these soda lakes.     

Distinction between light-mediated and light-independent variations in phytoplankton production rates

An array of factors simultaneously controls phytoplankton photosynthesis and hence the primary production process. Because their relative importance shifts both with depth and with season, the significance of individual factors cannot be resolved by in situ incubations, even if all relevant environmental and biotic variables are measured.Here a procedure is described by which in addition to in situ measurements, photosynthesis is simultaneously assessed in identical subsamples under constant temperature (10 °C) and light (0.66 mol m^–2 h^–1 PAR conditions, in vitro). By calculating photosynthesis per unit of chlorophyll, effects of shifting biomass on photosynthesis can be eliminated but seasonal variations of light-saturated photosynthesis generated by temperature, and vertical changes of light-requirements (e.g. by light-shade adaptation) remain obscure. Quotients of in situ photosynthetic rates divided by in vitro rates allow the quantification of light-mediated changes. Provided that photosynthesis measured in vitro is light-saturated, quotients in situ: in vitro rates should never exceed unity. They are a measure for the degree of light-limitation. In vitro rates normalized to chlorophyll give information on temporal changes caused by variations in photosynthetic capacity. In Lake Constance, mean cell size appears to control light-saturated assimilation numbers.     

Transition from Planktonic to Benthic Algal Dominance Along a Salinity Gradient

Highly regulated salinity gradients in solar salt pond concentrating sequences provide an opportunity to investigate in situ salinity impacts on aquatic flora and fauna. The Shark Bay Salt solar ponds at Useless Inlet in Western Australia vary in salinity from seawater to four times seawater over the pond sequence. We observed a shift from planktonic to benthic primary productivity as salinity increased. Water column photosynthesis and biomass decreased markedly with increasing salinity, while benthic productivity increased as cyanobacterial mats developed. Correspondingly, productivity shifted from autotrophy to heterotrophy in the water column and from heterotrophy to autotrophy in the benthos. Both shifts occurred at intermediate salinity (S = 110 g kg⁻¹, ρ = 1.087 g cm⁻³) in the pond sequence, where there was little production by either. Within individual ponds, productivity, algal biomass and physico-chemical conditions were relatively constant over one year, with only water column photosynthesis significantly different between seasons, mostly due to greater winter production. Transitions between benthic and planktonic production and their relative magnitudes appear to be driven mostly by direct responses to salinity stress, but also by changes in nutrient availability and grazing, which are also influenced by salinity.     

Photosynthetic acclimation of Chlorella saccharophila to heat stress

Photosynthesis is one of the most important metabolic processes of algae; which is altered as a stress response. During mass cultivation of algae, temperature rise and high light are major factors that affect biomass productivity. High temperature affects photosystem II (PSII) complex irreversibly, damaging intermolecular interactions in it. However, the impact of high temperature on photosynthesis is highly variable among different algal species, depending on the prior acclimation to environmental conditions they were exposed to. The acclimation plays an important role in combating high temperature stress via regulation of photosynthetic responses. Chlorophyll a fluorescence is a highly sensitive, non‐destructive and reliable tool for such measurements of photosynthetic parameters, which provides information about algal photosynthetic performance under given conditions. To understand the effect of heat stress on the responses of high light acclimated alga Chlorella saccharophila, chlorophyll a fluorescence transients were measured after heat exposure at 40°C. Our study demonstrates that rise in temperature for short duration; during open field cultivation reversibly affects the efficiency of PSII in light acclimated alga C. saccharophila. The effects of heat stress on chlorophyll a fluorescence in this alga, grown under high light (max‐1600 μmol photons m^?2 s^?1) are presented here; they are used to infer changes in photosynthetic process during its exposure to heat, as well as their recovery after 72 h. We speculate that heat resistance may have been acquired due to prior exposures to high light.     

SEASONAL PRIMARY PRODUCTION OF UNDERSTORY RHODOPHYTA IN AN UPWELLING SYSTEM1

The in situ primary production of three common under-story members of the Rhodophyta in South African west coast kelp beds was determined monthly for a year using dissolved oxygen techniques. Strong seasonal patterns of photosynthesis and respiration were evident in all three species. Net photosynthesis of all three species was greatest in spring (October) and lowest in winter (June). Increasing photosynthesis in late winter coincided with increasing ambient irradiance and photoperiod, whereas decreasing photosynthesis in summer was not explained by changes in the environmental parameters measured. We suggest that this may he due to an innate pattern related to some other seasonal plant activity such as reproduction. Seasonal P_max and I_k values reveal that the obligate understory species, B. prolifera and E. obtusa, are shade-adapted whereas G. radula, a low intertidal and shallow subtidal dominant, is sun-adapted. Low C: X ratios consistent with a high nutrient environment and high rates of productivity were found in all three species. Net photosynthesis to respiration (Pn:R) ratios were fairly constant for B. prolifera and E. obtusa, implying that then photosynthetic processes were governed more by seasonal variations in irradiance than by instantaneous light availability. The Pn: R ratio of G. radula was variable, suggesting that this species is more responsive to rapid fluctuations in irradiance and may therefore be adapted for rapid growth during periods of high irradiance.     

Nutrient dependence of phytoplankton production in brown-water lakes with special reference to Lake Päijänne

A method for predicting the mean seasonal chlorophyll a concentration, the mean seasonal in vitro phytoplankton primary productivity per unit volume, the maximum daily production per unit volume and the seasonal integral production in brown-water lakes is presented. The production values can be calculated when the mean annual concentration of total phosphorus and the mean annual colour of the water are known. This method has been developed especially for practical water pollution studies to permit rapid and inexpensive estimates of major biological consequences of changes in effluent loads. The method can be applied for brown-water lakes where phosphorus is the limiting nutrient for primary production.