Vertical and seasonal variation of phytoplankton photosynthesis in a brown-water lake with winter ice cover

SUMMARY. Unlike previously studied lakes with prolonged winter ice and snow cover, Lake Paajarvi, southern Finland, has a high humus content and consequently differs in both the quantity and quality of light penetration into its waters. Moreover, the range of temperature fluctuation and the degree of development of thermal stratification are greater in Paajarvi, and this increased environmental heterogeneity apparently stimulates diversity in the phytoplankton community, especially in the seasonal succession of species. Differences in the photosynthetic capacity of algae from different depths in the water column were not great. This is attributed to the extremely shallow euphotic zone, algae circulating freely through the steep light gradient and sedimenting rapidly once they pass through the thermocline into the hypolimnion. It is suggested that 'adaptation' of phytoplankton to the great seasonal changes in irradiance is achieved largely by successive growths of different species in the community, and that the adaptations and vertical migrations by individual algal species, which have been reported from polar and high alpine lakes, may be of secondary importance in Pääjärvi. The species successions in Pääjärvi produce changes in the pigment content of algae similar to those reported from polar and high alpine lakes, confirming that change in pigmentation is an important mechanism in light adaptation, whether at community or individual level. Algal pigment content was particularly high at the end of the long period of winter ice cover, indicating a degree of adaptation to the prolonged low-light conditions, which produced the extremely high photosynthetic capacities measured at this time. However, phytoplankton production at any irradiance was primarily determined by biomass.     

Seasonal and Spatial Overlap between Cladocerans in Humic Lakes

Seasonal and vertical distribution, migratory patterns and reproductive effort in coexisting cladocerans were investigated in three humic lakes with different, but low phytoplankton abundances and varying fish predation pressure. Seasonal and vertical habitat or niche overlap varied, but were high within most pairs of species in all localities. Migration was conspicuous in presence of planktivorous fish, less so in the fish free lake. Despite algal densities below incipient limiting level (30–200 μg C I⁻¹) and a low ratio (3–10) of algal to cladoceran biomass, zooplankton distribution and reproductive parameters were not clearly related to algal biomass. Bacterial biomass equalled 10–50% of phytoplankton biomass, while detritus by far was the largest of the particulate compartements. It was concluded that with a possible exception of the early summer algal bloom, additional carbon sources (bacteria, detritus) are important to cladoceran nutrition in these humic lakes. A large share of N- and P-poor detritus in the diet would give zooplankton productivity limitation by food quality in terms of elemental composition rather than food quantity. This would permit coexistence even of species with rather high food overlap, but give low production rates for all species in agreement with the observations.     

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

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

Seasonal variation in phytoplankton composition and physical-chemical features of the shallow Lake Doïrani,Macedonia, Greece

Phytoplankton species composition, seasonal dynamics and spatial distribution in the shallow Lake Doïrani were studied during the growth season of 1996 along with key physical and chemical variables of the water. Weak thermal stratification developed in the lake during the warm period of 1996. The low N:P ratio suggests that nitrogen was the potential limiting nutrient of phytoplankton in the lake. In the phytoplankton of the lake, Chlorophyceae were the most species-rich group followed by Cyanophyceae. The monthly fluctuations of the total phytoplankton biomass presented high levels of summer algal biomass resembling that of other eutrophic lakes. Dinophyceae was the group most represented in the phytoplankton followed by Cyanophyceae. Diatomophyceae dominated in spring and autumn. Nanoplankton comprised around 90% of the total biomass in early spring and less than 10% in summer. The seasonal dynamics of phytoplankton generally followed the typical pattern outlined for other eutrophic lakes. R-species (small diatoms), dominant in the early phase of succession, were replaced by S-species (Microcystis, Anabaena, Ceratium) in summer. With cooling of the water in September, the biomass of diatoms (R-species) increased. The summer algal maxima consisted of a combination of H and M species associations (sensu Reynolds). Phytoplankton development in 1996 was subject to the combined effect of the thermal regime, the small depth of mixing and the increased sediment-water interactions in the lake, which caused changes in the underwater light conditions and nutrient concentrations.     

Changes of algal biomass as carbon, cell number and volume, in bottles suspended in Lake Constance

Changes of algal biomass, as carbon, cell numbers and volumewere determined for phytoplankton of Lake Constance suspendedin situ in 2 l glass bottles. Phytoplankton placed at the 6%surface penetrating light level (photosynthetically availableradiation) were close to the compensation depth for growth estimatedas total particulate carbon and total cell volumes. Cell countsof individual alga] species however, showed appreciable growthof diatoms offset by the decline of flagellates. Bottles suspendedat two shallower depths in a separate experiment showed somegrowth of all species and indicated a vertical niche separationof growth of Rhodomonas minuta Skuja and R. lens Dascher andRuttner in accordance with their vertical distribution. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP) First International Workshop heldat the Limnological Institute, University of Konstanz, in April1982.     

Influence of Temperature,Photoperiod and Trophic Conditions on the Seasonal Cycles of Phytoplankton and Zooplankton in Two Deep Subarctic Lakes of Northern Canada

The densities of phytoplankton and zooplankton were determined for 12 consecutive months (1978–1979) in two deep oligotrophic lakes in northern Canada. The lakes were situated on the same river system, about 5 km from one another. While they exhibited similar temperature conditions, there were slight differences in the ionic content of the water. Overall, changing temperature was the most important factor influencing the duration of the growth cycles of phytoplankton and zooplankton. Nutrients generally controlled the intensity but not the timing of algal blooms. A similar relationship was recorded for the effects of food supply on the development of the major species of herbivorous zooplankton. Although changing photoperiod likely initiated the growth of several important algal and zooplankton species in the spring and early summer, light had no measurable impact on the development of planktonic communities throughout the remainder of the growing season. The higher ionic content of the water in one lake may have promoted the growth of protozoans during the fall and early winter.     

Vertical partitioning of the phytoplankton assemblage in ultraoligotrophic Crater Lake, Oregon, U.S.A.

SUMMARY 1. The summertime phytoplankton assemblage in abysmally deep (Z_max: 589 m) Crater Lake, Oregon, consists of over 100 species, which are variously distributed in the upper 200 m of the vertical water column. The depth distribution of the lake's three most prevalent species follows a predictabk pattern: Nitzschia gracilis in the 0–20 m stratum, Tribonema sp. at mid–depth (80–20 m), and Stephanodiscus hantzschii in the lowermost stratum (160–200 m). These major species, which account for approximately 80% or more of the lake's total phytoplankton biomass and primary production, exist under atypical temperature, light, and nutrient conditions.
2. The spatial distribution of phytoplankton in Crater Lake resembles a three-tier structure. Unlike most lakes, where the entire phytoplankton communities exist in less disparate environmental conditions, or are vertically mixed periodically by storm events and seasonal lake turnover. the Crater Lake community is partitioned into stratified environments.
5. The disparate and unusual characteristics of these environments, and the hydrological and limnological stability of the lake basin, are perhaps important factors regulating the diversity, dominance. and partitioning of the lake's phytoplankton populations.     

The development and vertical distribution of populations of gas-vacuolate bacteria in a eutrophic,monomictic lake

The role of gas vacuoles in the vertical stratification of planktonic bacteria is analysed. Measurements made with certain gas-vacuolate bacteria in laboratory culture suggest that only colonial forms could sink or float fast enough to form population maxima in lakes by vertical migration from other depths. It is suggested that in the case of individual cells the importance of the buoyancy provided by gas vacuoles is to minimise sinking rates and thereby to increase residence times of the organisms at depths where conditions support their growth.Changes in the vertical distribution of a number of gas-vacuolate bacteria were followed throughout the year in a monomictic, eutrophic lake (Crose Mere, Shropshire). All were restricted to the anaerobic hypolimnion which developed in summer. The various species formed maxima at different depths and times. With some of them (e.g. species of Thiopedia, Pelonema and Brachyarcus) growth was necessary to explain their development. In others (e. g. species of Pelodictyon and two colourless bacteria) vertical migrations might also have contributed to their development.     

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.     

Modelling the effects on phytoplankton communities of changing mixed depth and background extinction coefficient on three contrasting lakes in the English Lake District

1. The process‐based phytoplankton community model, PROTECH, was used to model the response of algal biomass to a range of mixed layer depths and extinction coefficients for three contrasting lakes: Blelham Tarn (eutrophic), Bassenthwaite Lake (mesotrophic) and Ullswater (oligotrophic). 2. As expected, in most cases biomass and diversity decreased with decreasing light availability caused by increasing the mixed depth and background extinction coefficient. The communities were generally dominated by phytoplankton tolerant of low light. Further, more novel, factors were identified, however. 3. In Blelham Tarn in the second half of the year, biomass and diversity did not generally decline with deeper mixing and the community was dominated by nitrogen‐fixing phytoplankton because that nutrient was limiting to growth. 4. In Bassenthwaite Lake, changing mixed depth influenced the retention time so that, as the mixed depth declined, the flushing rate in the mixed layer increased to the point that only fast‐growing phytoplankton could dominate. 5. In the oligotrophic Ullswater, changing the mixed depth had a greater effect through nutrient supply rather than light availability. This effect was observed when the mixed layer was relatively shallow (<5.5 m) and the driver for this was that the inflowing nutrients were added to a smaller volume of water, thus increasing nutrient concentrations and algal growth. 6. Therefore, whilst changes in mixed depth generally affect the phytoplankton via commonly recognized factors (light availability, sedimentation rate), it also affected phytoplankton growth and community composition through other important factors such as retention time and nutrient supply.     

Environmental control of phytoplankton productivity in turbulent turbid systems

Factors affecting phytoplankton productivity are analysed in turbid systems, such as shallow lakes and rivers. When resuspension from the sediment or loading from the catchment significantly increases inorganic (non-algal) turbidity and hence light attenuation potentials for high production are not realised. Energy available for phytoplankton growth is strongly regulated by underwater light availability which depends on the critical mixing depth, fluctuating light intensities and algal circulation patterns. Higher production rates in shallow waters are often compensated by greater algal respiration due to higher water temperatures when compared to deeper lakes.Total daily integral production of turbulent, turbid environments can be predicted from a combination of easily measured variables such as maximum photosynthetic rates, algal biomass, surface irradiance and some measure of underwater light attenuation.     

Erosion,phosphorus and phytoplankton response in rivers of South-Eastern Norway

The development of P fractions and phytoplankton was studied in three rivers with varying concentrations of seston.Less than 1% of the yearly TP transport may take place during periods with high algal biomass.The observation of a high growth rate of phytoplankton in the rivers coinciding with high concentrations of RP, low content of seston and high TP:Chl a ratio, indicate that the growth was often not P-limiting. During short periods with high phytoplankton biomass the ratio TP:Chl a may be low, indicating that a high fraction of TP was available.The content of P in soil samples and in samples with high seston content was about 0.1% of dry weight, and the algal availability of P often varied between 25 and 75% of TP for both types of samples.Decreasing biomass or low growth rates were observed at secchi depths less than 0.5 m and seston concentrations less than about 25 mg dry weight 1^–1. High flow rate also depressed the development of the total phytoplankton biomass. The assimilation of available P is incomplete under such conditions, i.e. under conditions of light limitation and high dilution rate.The availability of P for phytoplankton in rivers with different length, light conditions and stream velocity is discussed.     

Biotests with phytoplankton assemblages. Growth limitation along temporal and spatial gradients

The development of eight different species (populations) along temporal and vertical gradients in several lakes was studied. Many populations had an exponential growth phase and a decline phase. The growth rate was often high during the exponential phase. Some species, e.g. Oscillatoria spp. and Synedra cf. acus, often also had a long stationary phase. The growth rate and the sinking rate of these populations were often very low. Laboratory batch experiments with dilute phytoplankton populations were carried out to estimate the degree of growth limitation (L) for different populations sample from different lakes during the three growth phases. L was always low and often zero for populations initially in the exponential phase and always high for populations initially in the decline phase. The biotests also gave results that can help to explain the vertical distribution of Oscillatoria or Asterionella in three lakes investigated. The results indicate that the growth rates and the development of the populations were dependent on the external chemical and physical conditions. The transition between the different growth phases seemed often to be dependent on the external nutrient conditions. P, N, Si and Fe were probably the most growth-limiting nutrients. The growth rate of some diatoms was probably limited directly or indirectly at high pH. Laboratory biotests with natural populations may give valuable information on the growth-properties of different populations in the lakes. The biotests should, however, be carried out in combination with chemical and physical measurements and quantitative determinations of population densities.     

The distribution and periodicity of the phytoplankton of three Ceylon lakes

Summary The phytoplankton of three Ceylon lakes, two situated in the warm lowlands and one in the cool mountains has been investigated. Certain similarities and differences in regard to growth, distribution and periodicity of the plankton algae are discussed and related to environmental conditions.The lowland lakes are dominated by Myxophyceae which form 95% and 87% of the phytoplankton respectively, while in the mountain lake, which is temperate rather than tropical, Diatoms form 65% of the phytoplankton. Of the lowland lakes that in the centre of the densely populated city of Colombo has an algal productivity which is two and a half times as great as that situated in a vast area of uncultivated land. The mountain lake, surrounded by cultivated hills, occupies an intermediate position between the low country lakes in regard to productivity.The environmental factors mainly affecting algal growth, distribution and periodicity in these lakes are temperature, rainfall, sunshine, movement of the water, amount of influx of organic matter and nature of the surrounding soil. In assessing the effect of rainfall consideration has been given not only to the total precipitation, but also to its intensity and the occurrence of periods of drought between those of heavy rainfall.Part of thesis (Ceylonese Plankton Algae) approved for the Degree of Doctor of Philosophy in the University of London and lodged in the Library of that University.     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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Phytoplankton as an Indicator of the Water Quality of the Deep Lakes South of the Alps

This paper offers a synoptic account of studies on the phytoplankton communities in the deep southern subalpine lakes (DSL) Garda, Iseo, Como, Lugano and Maggiore. The main cause of the degradation of the water quality in the DSL is eutrophication. The euphotic layers of these lakes are trophically different, ranging from the oligo-mesotrophy of lakes Maggiore and Garda to the meso-eutrophy of lakes Iseo and Lugano. The trophic status as estimated by using total phosphorus and chlorophyll a has provided consistent results in agreement with the models proposed by OECD (1982. Eutrophication of Waters. Monitoring, Assessment and Control, OECD, Paris). Though related with chlorophyll a and TP, the Secchi disk depths have significantly underestimated the trophic status of the DSL. Two trophic indices using the algal orders (PTI_orders) and species (PTI_species) were drawn up on the basis of the distribution of phytoplankton along a trophic gradient defined by the application of multivariate methods; the scores emerging from these indices were used to make a definitive ecological classification of water bodies on a scale from 1 to 5, in accordance with the Water Framework Directive. A third index (PTI_OE) was computed as the ratio between the annual mean values of the cumulative biovolumes of two groups of algal orders with opposite trophic characteristics. The three PTI indices were highly correlated, providing a consistent classification of the water bodies. The indices proposed in this work were specifically adopted for use in the DSL. However, the criteria for their implementation constitute a robust and impartial tool for assessing similar indices in other lake typologies and for evaluating the degree of specificity of the trophic indicator values assigned to the single phytoplankton orders and species.     

Nutrient and Grazer-Mediated Effects on Picoplankton and Size Structure in Phytoplankton Communities

This study examines the factors which contribute to the abundance of algal picoplankton in lakes. A three-year field study of a meso-eutrophic lake was compared with observations from oligotrophic and highly eutrophic lakes in the region. Trophic state alone (oligotrophic vs. eutrophic) was not a good predictor of the importance of picoplankton; smaller cells were relatively abundant when phosphorus was limiting other phytoplankters, but also when nitrogen was in surplus. Subsequent field experiments found that picoplankton growth was stimulated by N, but not by P additions. This relationship was strongly affected by light and grazer levels. Grazers apparently mediate the effects of nutrient deficiency, and favor the growth of larger algal size classes, especially nanoplankton. The flux of P within experimental enclosures was controlled by picoplankton abundance under low nutrient conditions, but was a function of total phytoplankton biomass under P surplus.     

Ammonia may play an important role in the succession of cyanobacterial blooms and the distribution of common algal species in shallow freshwater lakes

With the human intensification of agricultural and industrial activities, large amount of reduced nitrogen enter into the biosphere, which consequently results in the development of global eutrophication and cyanobacterial blooms. However, no research had reported the effect of ammonia toxicity on the algal succession. In this study, we investigated the ammonia toxicity to 19 algal species or strains to test the hypothesis that ammonia may regulate the succession of cyanobacterial blooms and the distribution of common algal species in freshwater lakes. The bloom‐forming cyanobacterium Microcystis aeruginosa PCC 7806 suffered from ammonia toxicity at high pH value and light intensity conditions. Low NH₄Cl concentration (0.06 mmol L^?1) resulted in the decrease of operational PSII quantum yield by 50% compared with the control exposed to 1000 μmol photons m^?2 s^?1 for 1 h at pH 9.0 ± 0.2, which can be reached in freshwater lakes. Furthermore, the tolerant abilities to NH₃ toxicity of 18 freshwater algal species or strains were as follows: hypertrophication species > eutrophication species > mesotrophication species > oligotrophication species. The different sensitivities of NH₃ toxicity in this study could well explain the distributing rule of common algal species in the freshwater lakes of different trophic states. Meanwhile, the cyanobacterial bloom (e.g. M. aeruginosa) always happened at the low concentration of ammonia in summer, and disappeared with the decrease of ammonia. This may be attributed to the toxic effect of ammonia to M. aeruginosa in spring (the average and maximum ammonia concentration were 0.08 and 0.72 mmol L^?1 in 33 Chinese lakes), and the low level of NH₃‐N in summer and fall in the lakes might be used as preferred nitrogen nutrition by M. aeruginosa, rather than with toxicity. Therefore, ammonia could be a key factor to determine the distribution of common algal species and cyanobacterial bloom in the freshwater systems.     

The vertical distribution of phytoplankton in stratified water columns

What determines the vertical distribution of phytoplankton in different aquatic environments remains an open question. To address this question, we develop a model to explore how phytoplankton respond through growth and movement to opposing resource gradients and different mixing conditions. We assume stratification creates a well-mixed surface layer on top of a poorly mixed deep layer and nutrients are supplied from multiple depth-dependent sources. Intraspecific competition leads to a unique strategic equilibrium for phytoplankton, which allows us to classify the distinct vertical distributions that can exist. Biomass can occur as a benthic layer (BL), a deep chlorophyll maximum (DCM), or in the mixed layer (ML), or as a combination of BL+ML or DCM+ML. The ML biomass can be limited by nutrients, light, or both. We predict how the vertical distribution, relative resource limitation, and biomass of phytoplankton will change across environmental gradients. We parameterized our model to represent potentially light and phosphorus limited freshwater lakes, but the model is applicable to a broad range of vertically stratified systems. Increasing nutrient input from the sediments or to the mixed layer increases light limitation, shifts phytoplankton towards the surface, and increases total biomass. Increasing background light attenuation increases light limitation, shifts the phytoplankton towards the surface, and generally decreases total biomass. Increasing mixed layer depth increases, decreases, or has no effect on light limitation and total biomass. Our model is able to replicate the diverse vertical distributions observed in nature and explain what underlying mechanisms drive these distributions.     

ANTARCTIC AQUATIC ECOSYSTEMS AS HABITATS FOR PHYTOPLANKTON

1. The Southern Ocean is a large-scale, relatively homogeneous upwelling ecosystem whose phytoplankton apparently grows suboptimally over much of its area. By contrast there is a wide variety of freshwater habitats in the Antarctic and in some of these phytoplankton growth efficiency is very high. The two habitats share similar temperature and irradiance regimes, but differ markedly in availability of inorganic nutrients, in grazing pressure and in the time- and space-scales on which various physical processes act. 2. Concentrations of inorganic nutrients in the marine ecosystem have been represented as being in excess of phytoplankton requirements, but the ionic composition of some nutrient pools may not conform to phytoplankton preferences. 3. Nutrient-limitation determines phytoplankton production in Antarctic lakes and gives rise to gross differences between lakes. 4. Irradiance in the water column varies greatly over the year in both marine and freshwater ecosystems. Most algae are shade-adapted, with the ability to utilize low irradiance but with sub-optimal response to high irradiance. However, local phytoplankton maxima may attain very high carbon fixation and growth rates. 5. Consistently low temperatures characterize both systems. Their effects on photo-synthetic carbon uptake mirror shade-adaptation. Division rates of marine phytoplankton may however be very much higher than predicted for ambient temperatures. 6. Vertical mixing is important in both ecosystems and influences the environment experienced by phytoplankton cells. This appears to have little effect on the average performance of phytoplankton in the strongly mixed surface water column of the Southern Ocean, where the mixed depth may exceed 100 m. This can be related partly to the shade-adapted photosynthetic response. Euphotic depths range from 20 to 100 m. 7. Strong vertical mixing under ice-free conditions in lakes may maximize photosynthetic efficiency, whilst distinct vertical stratification in permanently ice-covered lakes gives rise to segregation of nutrient uptake and regeneration. 8. Physical removal of phytoplankton biomass by grazing is locally important in the Southern Ocean, in contrast to the estimated mean mesoscale impact of grazing. Vertical sedimentation losses appear important in the context of mixing depth and generation time, and may be modified by vertical circulation of water. 9. Loss of phytoplankton biomass from lakes during the ice-free period is dominated by physical removal via the lake outflow. Grazing is generally unimportant, except where larvae of otherwise nektobenthic zooplankton hatch in synchrony with a phytoplankton maximum. Sedimentation is important under ice-cover.