Seasonal changes in the underwater light climate of two Canadian shield lakes

This study examines the seasonal variation in the underwater spectral distribution of light in a mesotrophic (Lake Cromwell) and an oligo-mesotrophic (Lake Croche) temperate lake. Gilvin is primarily responsible for the strong selective attenuation of blue light in both lakes. As a result of differing gilvin concentrations light transmission maxima of downwelling and upwelling spectra are near 615 nm in Lake Cromwell and 599 nm in Lake Croche. With increases in depth both upwelling and downwelling radiance fluxes decrease, are shifted to longer wavelengths and become more monochromatic. The greatest penetration of light occurs in the summer and spring after which a gradual decrease occurs through fall to a minimum value in winter. Under the winter cover the P ₅₀ of downwelling light shifts 10 to 20 nm towards shorter wavelengths. Seasonal changes in downwelling irradiance are related to solar altitude, concentration of suspended particles, phytoplankton populations, amount of gilvin, mixing and winter cover. The brownish colouration of these lakes is explained by reflectance of spectrally impure orangish-red light.     

Optical Properties and Light Climate in Lake Verevi

The optical properties and light climate during the ice-free period in the highly stratified Lake Verevi (Estonia) have been studied together with other lakes in same region since 1994. The upper water layer above the thermocline belongs to class “moderate” by optical classification of Estonian lakes but can turn “turbid” (concentration of chlorophyll a up to 73 mg m⁻³ and total suspended matter up to 13.2 g m⁻³) during late summer blooms. In the blue part of the spectrum, light is mainly attenuated by dissolved organic matter and in red part notably scattering but also absorption by phytoplanktonic pigments effect the spectral distribution of underwater light. Consequently, the underwater light is of greenish-yellow color (550–650 nm). Rapid change in optical properties occurs with an increase of all optically active substances close to thermocline (2.5–6 m). Optical measurements are often hampered beneath this layer so that modeling of the depth distribution of the diffuse attenuation coefficient is an useful compliment to field measurements. K_d,PAR ranges from 0.8 to 2.9 m⁻¹ in the surface layer, and model results suggest that it may be up to 5.8 m⁻¹ in the optically dense layer. This forms a barrier for light penetration into the hypolimnion.     

Modeling underwater light climate in relation to sedimentation,resuspension, water quality and autotrophic growth

The underwater light climate ultimately determines the depth distribution, abundance and primary production of autotrophs suspended within and rooted beneath the water column. This paper addresses the underwater light climate, with reference to effects of suspended solids and growth responses of autotrophs with emphasis on phytoplankton.Effects of the most important factors contributing to the absorption and scattering of light in surface waters were described. A comparison between spectral and scalar approaches to underwater light climate modeling was made and examples of linear approximations to light attenuation equations were presented. It was demonstrated that spectral and scalar photosynthesis models may converge to similar values in spectral-flat, high photon flux environments, but that scalar PAR models may overestimate biomass-specific production by 70%. Such differences can lead to serious overestimates of habitat suitability for the growth and survival of submersed macrophytes, particularly in relatively turbid, coastal waters.Relationships between physical and optical properties of suspended sediments were described theoretically, and illustrated with modeling examples and measurements. It was found that the slowly settling particulate fraction contributed substantially to the suspended solids concentration, and greatly to light attenuation within the water column. It was concluded that distinguishing particles by fall velocity and concomitant light attenuation properties in the modeling of underwater light conditions allowed the establishment of useful, although not simply linear, relationships.In eutrophic, shallow lakes, the largest contribution to light attenuation often originates from phytoplankton on a seasonal basis (months–years), but from suspended solids behavior on a shorter time scale (days–weeks), particularly when water bodies are wind-exposed. Temporal and spatial variabilities in wave height, suspended solids concentrations, and light attenuation within the water column, and their importance for autotrophic growth were described, and illustrated with a case study pertaining to Markermeer, The Netherlands. The influence of underwater light conditions on phytoplankton succession was briefly discussed and illustrated with a case study pertaining to Lake Veluwe, The Netherlands. It was concluded that modeling the underwater light climate in a water body on a few sites only can indicate how important various components are for the attenuation of light, but based on the current state of the art, it can not be expected that this will provide accurate predictions of the underwater light climate, and of phytoplankton and submersed macrophyte growth.     

OBSERVATTVNS ON THE UNDERWATER LIGHT FIELD OF TWO HWERTROPHIC,SOUTH AFRICAN IMPOUNDMENTS

SUMMARY

The spectral composition of the underwater light field was examined in two hypertrophic South African Impoundments (Hartbeespoort and Roodeplaat Dams) under a range of inorganic turbidities and chlorophyll α concentrations. The data indicated that inorganic turbidity and gilvin were dominant to chlorophyll in regulating underwater light attenuation during the present study. Under all conditions the wavelengths between 405 and 510 nm were attenuated more rapidly than near UV and the wavelengths above 510 nm and the 623 nm component penetrated deepest. Under low turbidities the 546 nm wavelength was the next most penetrating component, but its attenuation increased with increasing turbidity. This characteristic of the underwater light field may be important to the cyanobacteria which dominate in these hypertrophic lakes.     

太湖梅梁湾沿岸带水体生物学与光学特性

基于 1998～ 1999年周年 4季原位水下光场观测资料及中国科学院太湖湖泊生态系统研究站 1992～ 2 0 0 1年悬浮物、叶绿素 a、透明度长期历史观测资料分析了太湖梅梁湾沿岸带第 2号站点水体的生物学与光学特性 ,探讨了水下光合有效辐射(PAR)总量的日变化、垂直分布 ;光衰减系数的季节变化及光谱分布 ;影响光衰减系数的主要水色因子。结果表明 ,无论是 PAR还是光谱衰减系数其值都很高 ,其中 PAR衰减系数在 1.4 0～ 5 .30 / m间变化 ,均值为 2 .4 3± 0 .5 5 / m,秋季最大、夏季最小 ,真光层深度在 0 .87～ 3.2 9m间变化 ,均值为 1.98± 0 .4 1m;水下光谱在蓝光波段衰减最强烈 ,其次是红光、绿光 ,随着深度增加光谱成分出现绿移和红移现象 ,绿红光占得比例越来越大 ;光谱衰减系数随着波长的增加大致呈下降趋势 ,但在 6 70 nm附近有个峰值 ;基于线性相关分析发现在混浊的沿岸带水体中影响光衰减主要因子为水体中的悬浮物和有色可溶性有机物 ,叶绿素 a对 PAR衰减系数的贡献率只占到 1.5 9%～ 14 .2 1%。     

The underwater light-field of lakes with marked physico-chemical and biotic diversity in the water column

Errors in calculation of radiant fluxes of downwelling underwaterphotosynthetically available radiation (PAR) from force-fittingstatistical values of d to vertical broad-band PAR profiles are compounded by vertical heterogeneity in suchfactors as turbidity, gilvin (= gelbstoff) and chlorophyll.Examples are given from two Tasmanian meromictic lakes wherevertical zonation of gilvin and microorganisms in the mixolimniaand across the chemoclines produces a markedly heterogeneouswater column. Localised concentrations of gilvin produce kinkedprofiles of downwelling PAR (400–700 nm) and microbialplates in the vicinity of the chemoclines act as false, reflectivebottoms, abruptly extinguishing residual PAR by absorption andscattering.     

Reconstruction of the Time Series of the Underwater Light Climate in a Shallow Turbid Lake

Lake ülemiste is a shallow, eutrophic lake which has served the city of Tallinn as a water reservoir for many centuries. Its light climate was studied by combining a routinely measured data set with a modelling approach. For 26 years (1978–2004), data was collected on such optically active substances (OAS) and water parameters as water colour, turbidity and phytoplankton biomass. Simple modelling enabled the quantification of long-term time-series data and the subsequent calculation of the diffuse attenuation coefficient, euphotic depth and average light of the mixed layer. Several changes in the hydrological cycle have taken place during the period under study, among which are an increase in the water level of about 0.5 m and a decrease in the external water load from 108 million m³ year⁻¹ to about 25 million m³ year⁻¹. At the same time euphotic depth has shown a distinct trend towards increasing since the early 1990s. The euphotic depth also showed an increase (from 1.1 to 1.4 m) due to an improvement in underwater light conditions – mainly in the spring (April and May) and autumn (October and November) because of the lower amount of dissolved organic matter in the lake. The average light availability in the mixed layer has increased, but this has not affected the phytoplankton biomass as the latter is not light-limited during the summer period.     

Summer depth selection in crustacean zooplankton in nutrient‐poor boreal lakes is affected by recent residential development

1. Strong vertical gradients in light, water temperature, oxygen, algal concentration and predator encounters during summer in stratified lakes may influence patterns of depth selection in crustacean zooplankton, especially Daphnia species. 2. To test how crustacean depth selection varies among lakes along a gradient of catchment disturbance by recent residential development and land use change, we calculated the weighted mean depth distribution of the biomass of crustaceans by day and night in eight nutrient‐poor boreal lakes. 3. Generally, the greatest biomass of crustaceans was located at the metalimnion or at the lower boundary of the euphotic zone during thermal stratification in July. The crustacean zooplankton avoided warm surface layers and tended to stay in colder deep waters by both day and night. They also remained at greater depths in lakes with a more extensive euphotic zone. 4. There was some evidence of upward nocturnal migrations of large Daphnia and copepods in some lakes, and one case of downward migration in a lake inhabited by chaoborid larvae. 5. Multivariate regression trees (MRT) were used to cluster crustaceans and Daphnia species in homogeneous groups based on lake natural and disturbance factors. For crustaceans, the depth of the euphotic zone, the sampling depth (epilimnion, metalimnion and hypolimnion), time (day or night) of sampling and the biomass of chlorophyll a were the main driving factors. For Daphnia species, the drainage area, the sampling depth, the cleared land surface area within the catchment and the concentration of total dissolved phosphorus were the main factors.     

Effects of gilvin on the composition and dynamics of metalimnetic communities of phototrophic bacteria in freshwater North-American lakes

The spectral distribution of light reaching the populations of phototrophic bacteria in the metalimnion of stratified lakes is a selective factor determining the community composition. At deep metalimnia, light spectra are enriched in photons of the central part of the spectrum (500-600 nm) and benefit Chromatiaceae, brown-coloured Chlorobiaceae and phyco-erythrine-containing cyanobacteria. Their carotenoids (okenone, spiriloxanthine, isorenieratene) and phycoerythrines allow these phototrophic bacteria to use light from the narrow central spectral wavebands. Otherwise, shallow metalimnetic communities receive light from a wide range (400-800 nm) and their composition is more diverse and usually enriched in green-coloured Chlorobiaceae, which are unable to take advantage of the central part of the spectrum. Gilvin compounds (humic substances dissolved in water), have strong effects on light absorption, especially at shorter wavelengths. Therefore, light spectra in lakes with high gilvin contents are enriched in photons of long wavelengths (> 600 nm). Several Wisconsin lakes with different gilvin contents were studied during the period of summer stratification in 1994. Spectral distribution of light reaching their metalimnia changed with increasing gilvin contents (measured as g(440) ). In the latter, phototrophic metalimnetic bacterial communities were absolutely dominated by green-coloured Chlorobiaceae. Intermediate lakes could experiment changes on their community composition depending on variations in gilvin content, as happened in Little Long lake. The dynamics of this lake was studied during summer 1995. The ratio of green-coloured species in respect to brown-coloured species increased after a sudden increase of gilvin due to strong rainfall. These results agree with the photosynthetic advantage of green-coloured Chlorobiaceae under red-light illumination, inferred from laboratory experiments, and suggest a bacteriochlorophyll-dependent, light-harvesting strategy of these phototrophic sulphur bacteria.     

Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

Secchi disk — chlorophyll relationships in a lake with highly variable phytoplankton biomass

In meseutrophic Lake Constance mean euphotic phytoplankton chlorophyll concentrations vary about 100-fold over the year. Concomitant fluctuations in euphotic depth (Z_eu) and Secchi depth (Z_s) are related to each other in a non-linear fashion that as a rough approximation can be expressed by Z_eu 5 Z_s.Secchi depth is to a great extent a function of beam attenuation of light which depends on the inherent optical properties of the water and is highly sensitive to light scattering from particles. Euphotic depth, by contrast, is a function of the vertical light attenuation coefficient which also depends on absorption and scattering, but is less sensitive to the latter than beam attenuation. Algal cells both absorb and scatter light and therefore influence Secchi depth and euphotic depth, however, in different fashions.Whenever the lake is clear due to scarce phytoplankton, scattering is small and beam attenuation only exceeds vertical light attenuation by a relatively small factor. As a consequence, the ratio of euphotic depth to Secchi depth is small (1.5–2.5). When the lake is turbid due to high algal density, enhanced scattering from algal cells and detrital particles causes beam attenuation to rise more than vertical light attenuation, thus leading to high ratios of euphotic depth to Secchi depth (3–5). The relatively close relationships between Secchi depth and chlorophyll in Lake Constance are due to (1) high influence of chlorophyll concentration on water transparency, (2) co-variation of phytoplankton and other suspended particles, and (3) limited variation of cellular chlorophyll contents.     

Oxygen Productivity of Planktonic Algae under Fluctuating and Constant Light Conditions

Net oxygen productivity in cultures of Monoraphidium minutum, Cryptomonas sp. and Planktothrix agardhii exposed to fluctuating and constant light conditions was measured in a laboratory incubator. The fluctuating light climate simulated a linear up and down movement in a 2 m water column at 4 different ratios of euphotic depth to mixing depth. In addition, cultures were kept at a constant light climate simulating static incubation at 0, 0.5, 1 and 2 m depth and at the depth of the mean irradiance, respectively. Integral productivity in the simulated water column was lowest when algae were incubated at constant light in different depths, highest when the algae were incubated at constant mean photon flux density (PFD) and intermediate under fluctuating light. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Nutrients versus physical factors in determining the primary productivity of waters with high inorganic turbidity

Primary production was studied in two reservoirs of the Modder River, which are polluted by two cities over a period that included the tail end of a drought and severe flooding. It was shown that suspended inorganic materials affected the underwater light climate and consequently had a marked influence on primary productivity. The ratio of euphotic to mixing depth was shown to be the most important factor affecting overall productivity and that nutrients are of secondary importance only. Control measures which are based on bottom-up relations, would most probably be inapplicable to the turbid waters of the Modder River.     

WATER-BLOOMS

1. Peculiarities in the ecology of planktonic blue-green algae are reviewed in relation to recent advances in understanding their physiological characteristics. 2. Dense water-blooms are always the result of buoyant migration of existing populations to the lake surface under calm weather conditions. The size of the population is the direct result of photoautotrophic growth, and is dependent upon light and the availability of inorganic nutrients; it is apparently enhanced by moderately high water temperatures, high pH, low oxygen tensions and possibly, the presence of organic solutes. The relative effectiveness of these factors is untested. 3. Buoyancy is imparted by gas vacuoles whose principal function is to regulate the position of the alga in the water column. Control is effected by two mechanisms: (i) ‘dilution’ of newly produced vacuoles during active cell division; (ii) changes in cell turgor-pressure acting on the gas-vacuole structure. Gas-vacuole production is greatest at low light intensities and the alga becomes more buoyant; at higher light intensities, increased turgor-pressure collapses the weaker vacuoles causing the alga to lose buoyancy. 4. Potentially, algae are able to poise themselves at an optimum point in the light gradient, usually towards the bottom of the euphotic zone, where the algae are likely to encounter the conditions most favouring their growth. 5. Different species of blue-green algae differ in the typical sizes of their colonies and, hence, in their rates of controlled movement. These differences are interpreted as hydrodynamic adaptations to the variations in turbulent water movements to which the algae are subject. 6. Populations of single-filamentous Oscillatoria agardhii and O. rubescens come to occupy the stable metalimnia of stratified lakes, provided that they are located within the euphotic zone. 7. The large stream-lined colonial forms occur mainly in polymictic lakes and in the unstable epilimnia of stratified lakes where light penetration is restricted to the superficial layers. These algae are adapted to sink or float rapidly to the optimum depth when turbulence subsides. Because of their potentially high rates of movement, it is the large colonial forms that commonly form blooms. 8. Bloom formation can occur when most of the algae possess excess buoyancy. Excess buoyancy is acquired when the photosynthetic rate is insufficient to develop the necessary turgor-pressure to cause collapse of the vacuoles. Photosynthesis may be sufficiently impaired under four circumstances: (i) during turbulent circulation of the population over a depth that significantly exceeds the euphotic depth; (ii) in the absence of light (e.g. at night): (iii) at limiting concentrations of carbon dioxide: and (iv) when the algal population is senescent. 9. Because bloom-formation depends upon the coincidence of persistent algal overbuoyancy with calm weather, its occurrence is incidental, and serves no vital function in the biology of blue-green algae. 10. Some possible causes for the occurrence of blue-green algal blooms in a relatively restricted range of water bodies are discussed. Large bloom-forming populations are probably restricted to moderately rich, mildly alkaline, thermally unstable lakes in all regions, except those which are permanently cold. Extremes of poverty or richness of nutrients, short water-retention times and low pH seem to be factors which select against planktonic blue-green algae.     

An Incubator for the Simulation of a Fluctuating Light Climate in Studies of Planktonic Primary Productivity

A laboratory system for the quantification of phytoplankton photosynthesis under fluctuating light climate conditions is described. It consists of 2 temperature-controlled incubators with a variable light supply, an algal batch culture in incubation bottles with appropriate stirrers and a set of oxygen electrodes to monitor algal photosynthesis. By the rotation of special grey filters between the incubator and the light source, a regular up and down movement in the water column is simulated in up to 7 parallel bottles. Different ratios of euphotic depth to mixing depth and different velocities can be applied. Simultaneously, 8 bottles can be incubated under constant light. The system is demonstrated in experiments with Chlamydomonas sp. Further possibilities of application are proposed.     

Role of terrestrial carbon in aquatic UV exposure and photoprotective pigmentation of meiofauna in subarctic lakes

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Lake Chisholm,a polyhumic forest lake in Tasmania

Lake Chisholm is a polyhumic, warm monomictic forest lake in western Tasmania. Its large relative depth and sheltering forest result in nine months stratification. The high humic content is a dominant feature, producing a sharp, shallow thermocline, a shallow euphotic depth (< 1 m) and an underwater light climate dominated by red wavelenghts. The hypolimnion is anoxic and sulphide-laden and even in winter circulation is sluggish. For much of the year the lake resembles a biogenically meromictic lake, and though there is only slight chemical enrichment of the hypolimnion there is nonetheless considerable vertical structure in the water column. Chromophyte flagellates are the dominant algae, a few species often forming monospecific blooms in a sporadic manner. Lake Chisholm is seen as an oceanic, mid-latitude counterpoint to dimictic, polyhumic, flagellate haunts in Scandinavia.     

A test of Tyler’s Line – response of chironomids to a pH gradient in Tasmania and their potential as a proxy to infer past changes in pH

1. Tyler’s Line delimits two distinct limnological provinces that reflect differences in climate, geology and vegetation in Tasmania. Lakes west of Tyler’s Line are typically acidic and dystrophic with relatively shallow euphotic zones, whereas eastern lakes are circumneutral and oligotrophic or ultra‐oligotrophic, allowing deeper penetration of light. Consequently, Tyler’s Line defines a boundary where species assemblages change over a relatively short distance. 2. A survey of 48 Tasmanian lakes was undertaken to identify indicator taxa of the two limnological provinces and breakpoints along the pH gradient where shifts in taxa occur. Chironomidae (Diptera) were used because they are ideal candidates for lake classification. 3. Three independent methods (geographical position, piecewise linear regression, two‐way indicator species analysis) verified that chironomids accurately reflect the environmental variables defining Tyler’s Line at lake and catchment scales. Chironomid genera are often speciose, and members of the same genus can have markedly different responses to a given environmental variable. Although the types of taxa changed along the pH gradient, richness did not. This finding contrasts with many studies from the northern hemisphere but accords with other studies from Australia. 4. Models of pH, developed using both partial least squares and weighted averaging partial least squares, can be used to understand past natural variability of pH in Tasmania and to test hypotheses regarding the timing, magnitude and source of contamination in impacted aquatic ecosystems.     

The impact of phytoplankton on spectral water transparency in the Southern Ocean: implications for primary productivity

Spectral water transparency in the Northern Weddell Sea was studied during Austral spring. The depth of the 1-% surface irradiance level (euphotic depth) varied between 35 and 109 m and was strongly influenced by phytoplankton biomass. Secchi depths were non-linearly related to euphotic depth. In phytoplankton-poor water, the most penetrating spectral region was restricted to a relatively narrow waveband in the blue (488 nm), but the range was broader, between 488 and 525 nm when phytoplankton were abundant. Water transparency in the red spectral range was always low and only to a small extent affected by phytoplankton. Two independent procedures were used to quantify the impact of phytoplankton on spectral water transparency: (1) Regression analysis of spectral in situ vertical light attenuation coefficients in the sea, against coincident chlorophyll concentrations. This method gave chlorophyll-specific light attenuation coefficients; the y-intercept could be interpreted as a measure of light attenuation by pure water plus non-algal material. (2) Spectra of in vivo light absorption derived by spectroscopy, using phytoplankton enriched to varying degrees onto filters. Thus chlorophyll-specific absorption cross-sections were determined. Estimates obtained by both procedures were in close agreement. By integrating over the spectrum of underwater irradiance, in situ chlorophyll-specific absorption cross sections of phytoplankton suspensions, related to all photosynthetically active radiation, were calculated. Light absorption by phytoplankton for photosynthesis is accomplished mainly in the blue spectral range. Also dissolved and particulate organic matter contributed to the attenuation of blue light. Because in water poor in phytoplankton, underwater irradiance was progressively restricted to blue light, chlorophyll-specific absorption cross-sections of phytoplankton, averaged over the spectrum of photosynthetically active irradiance, increased with water depth. In water with elevated phytoplankton biomass, overall light attenuation was generally enhanced. However, because the spectral composition of underwater light changed relatively little with depth, except immediately below the water surface, light absorption cross-sections of phytoplankton changed little below 10 m depth. Vertical differences in the proportions of underwater light absorbed by the phytoplankton community here were mainly dependent on biomass variations. Because of the comparatively small attenuation of blue light by non-algal matter, the efficiency of light harvesting by phytoplankton at any given concentration of chlorophyll in Antractic waters is greater than in other marine regions. At the highest phytoplankton biomass observed by us, as much as 70% of underwater light was available for phytoplankton photosynthesis. When phytoplankton were scarce, <10% of underwater light was harvested by phytoplankton.Contribution within the European Polarstern Study (EPOS), supported by the Deutsche Forschungsgemeinschaft, Grant Ti 115/16-1 to MMT, the European Science Foundation, and by the Alfred Wegener Institut für Polar-und Meeresforschung, Bremerhaven