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1.
Ultraviolet radiation and bio-optics in Crater Lake,Oregon   总被引:3,自引:3,他引:0  
Crater Lake, Oregon, is a mid-latitude caldera lake famous for its depth (594 m) and blue color. Recent underwater spectral measurements of solar radiation (300–800 nm) support earlier observations of unusual transparency and extend these to UV-B wavelengths. New data suggest that penetration of solar UVR into Crater Lake has a significant ecological impact. Evidence includes a correlation between water column chlorophyll-a and stratospheric ozone since 1984, the scarcity of organisms in the upper water column, and apparent UV screening pigments in phytoplankton that vary with depth. The lowest UV-B diffuse attenuation coefficients (K d,320) were similar to those reported for the clearest natural waters elsewhere, and were lower than estimates for pure water published in 1981. Optical proxies for UVR attenuation were correlated with chlorophyll-a concentration (0–30 m) during typical dry summer months from 1984 to 2002. Using all proxies and measurements of UV transparency, decadal and longer cycles were apparent but no long-term trend since the first optical measurement in 1896.  相似文献   

2.
Microalgal pigment composition, photosynthetic characteristics, single-cell absorption efficiency (Qa(λ)) spectra, and fluorescence-excitation (FE) spectra were determined for platelet ice and benthic communities underlying fast ice in Mc Murdo Sound, Antarctica, during austral spring 1988. Measurements of spectral irradiance (E(λ)) and photosynthetically active radiation (PAR) as well as samples for particulate absorption measurements were taken directly under the congelation ice, within the platelet layer, as profiles vertically through the water column, and at the benihic surface. Light attenuation by.sea ice, algal pigments, and particulates reduced PAR reaching the platelet ice layer to 3%(9–33 fimol photons m-2-?s-1) of surface values and narrowed its spectral distribution to a band between 400 and 580 nm. Attenuation by the water column further reduced PAR reaching the sea floor (28–m depth) to 0.05% of surface levels (< 1 μmol photons m-2 s-1), with a spectral distribution dominated by 470–580–nm wavelengths. The photoadaptive index (I) for platelet ice algae (5.9–12.6 μmol photons m-2.s-1) was similar to ambient PAR, indicating that algae had acclimated to their light environment (i.e. the algae were light-replete). Maximum Qa(λ) at the blue absorption peak (440 nm) was 0.63, and enhanced absorption was observed from 460–500 nm and was consistent with observed high cellular chlorophyll (chi) c:chl a and fucoxanthin: chl a molar ratios (0.4 and 1.2, respectively). Benthic algae were light-limited despite the maintenance of very low Ik values (4–11 μmol photons.m-2.s-1). Extremely high fucoxanthin: chi a ratios (1.6) in benthic algae produced enhanced green light absorption, resulting in a high degree of complementation between algal absorption and ambient spectral irradiance. Qa(λ) values for benthic algae were maximal (0.9) between 400 and 510 nm but remained >0.35 even at absorption minima. Strong spectral flattening, a characteristic of intense pigment packaging, was also apparent in the Qa(λ) spectra for benthic algae. FE and Qa(λ) spectra were similar in shape for platelet ice algae, indicating that the efficiency at which absorbed energy was transferred to photosystem II (PSII) was independent of wavelength. Fluorescence emission by benthic algae was greatest for the 500–560–nm excitation wavelengths, suggesting that most energy absorbed by accessory pigments was transferred to PSII. These results suggest that under ice algae employ complementary pigmentation and maximize absorption efficiency as adaptive strategies to low-light stress. Regulating the distribution of absorbed energy between PSI and PSII may be an adaptive response to the restricted spectral distribution of irradiance.  相似文献   

3.
Two series of lakes with increasing attenuation were examined for trends in spectral composition. They became the basis for an evaluation of the light environment at the lower boundary (LB) of Nitella meadows in three other series of lakes. Increased attenuation (K d PAR) was marked by progressive erosion of the blue window and caused primarily by humic substances. An increase in K d PAR from 0.06 to 0.81 produced, at the floor of the euphotic zone, a shift in K d min from 440 to 580 nm. Regressions of boundary depths of Nitella meadows on water clarity produced similar slope coefficients for the three series of lakes. Several trends became evident: 1, PAR irradiance at the LB increases with depth of the LB; 2, red light (E d 660) declines from richness at shallow LB to near extinction in deep water LB in clear lakes; while 3, blue light (K d 450) increases to an asymptote. Blue light appears to substitute, although less effectively, for red light irradiance in the growth regulation of charophytes. These data support an hypothesis that spectral quality is involved in the determination of lower boundary depths for benthic macro-algae.  相似文献   

4.
At all seasons, the underwater light field of meso-eutrophic large (480 km2) deep (mean: 100 m) Lake Constance was studied in conjunction with the assessments of vertical distributions of phytoplankton chlorophyll concentrations. Vertical profiles of scalar, downwelling and upwelling fluxes of photosynthetically available radiation, as well as fluxes of spectral irradiance between 400 and 700 nm wavelength were measured.The overall transparency of the water for PAR is highly dependent on chlorophyll concentration. However, the spectral composition of underwater light is narrowing with water depth regardless of phytoplankton biomass.Green light is transmitted best, even at extremely low chlorophyll concentrations. This is explained by the selective absorption of blue light by dissolved organic substances and red light by the water molecules. Nevertheless, significant correlations were found between vertical attenuation coefficients of downwelling spectral irradiance and chlorophyll concentrations at all wavelengths. The slopes of the regression lines were used as estimates of chlorophyll-specific spectral vertical light attenuation coefficients (K c()).The proportions of total upwelling relative to total downwelling irradiance (reflectance) increased with water depth, even when phytoplankton were homogeneously distributed over the water column. Under such conditions, reflectance of monochromatic light remained constant. Lower reflectance of PAR in shallow water is explained by smaller bandwidths of upwelling relative to downwelling light near the water surface. In deeper water, by contrast, the spectra of both upwelling and downwelling irradiance are narrowed to the most penetrating components in the green spectral range. Reflectance of PAR was significantly correlated with chlorophyll concentration and varied from 1% and 1-% at low and high phytoplankton biomass, respectively. Over the spectrum, reflectance exhibited a maximum in the green range. Moreover, in deeper layers, a red maximum was observed which is attributed to natural fluorescence by phytoplankton chlorophyll.  相似文献   

5.
  • 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Zm) and depth of the euphoric zone (Zeu). During periods of complete mixing, Zm 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 Zm/Zeu 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.
  相似文献   

6.
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)  相似文献   

7.
Daily integrals of photosynthesis by a cyanobacterial bloom in the Baltic Sea, during the summer of 1993, were calculated from the vertical distributions of light, temperature and the organisms in the water column and from photosynthesis/irradiance curves of picoplanktonic and diazotrophic cyanobacteria isolated from the community. The distribution of chlorophyll a in size-classes <20?µm and >20?µm was monitored over 9 days that included a deep mixing event followed by calm. Picocyanobacteria formed 70% of the cyanobacterial biomass and contributed 56% of the total primary production. Of the filamentous diazotrophs that formed the other 30%, Aphanizomenon contributed 28% and a Nodularia-containing fraction 16% of the primary production. For the whole population there was little change in standardized photosynthetic O2 production, which remained at about 31?mmol?m?2 before and after the mixing event. There were differences, however, between the classes of cyanobacteria: in picocyanobacteria primary production hardly changed, while in Aphanizomenon it increased by 2.6 and in Nodularia it fell below zero. Total phytoplankton photosynthesis was strongly dependent on total daily insolation with the compensation point at a photon insolation of 22.7?mol?m?2?d?1. Similar analyses of N2 fixation showed much less dependence on depth distribution of light and biomass: Aphanizomenon fixed about twice as much N2 as Nodularia their; their fixation exceeded their own N demand by about 12%. Together, these species contributed 49% of the total N demand of the phytoplankton population. Computer models based on the measured light attenuation and photosynthetic coefficients indicate that growth of the cyanobacterial population could occur only in the summer months when the critical depth of the cyanobacteria exceeds the depth of mixing.  相似文献   

8.
Sunlight penetration through the water column is controlled by the amount and kind of materials dissolved and suspended in the water. Understanding UV penetration in its complexity is essential for the prediction of the impact of UV radiation on aquatic ecosystems. However, only limited data are available on the penetration of UVR into shallow waters rich in inorganic suspended solids and chromophoric dissolved organic matter (CDOM). The same is true for the specific attenuation coefficients of light-absorbing components at the UV waveband. This study analyses the role of CDOM, algal-free suspended solids and algae in the formation of underwater UVR and PAR climate in 30 water bodies from clear gravel pit lakes trough the shallow Lake Balaton to turbid soda pans. Irradiance-depth profiles were obtained at 305, 313, 320 nm (UV-B), 340, 380, 395 nm (UV-A) and 400–700 nm (PAR) with a Biospherical PUV-2500 radiometer. Vertical attenuation coefficients (K d) were calculated. Water samples were taken for the laboratory measurement of the concentration of light-absorbing components: algae as chlorophyll a (CHL), chromophoric dissolved organic matter as colour (CDOM), and algal-free suspended solids (TSS-Alg) parallel with the in situ light measurements. Specific attenuation coefficient values were calculated by multiple regression analysis (n = 140). The obtained specific UV attenuation coefficient values of CHL, CDOM and TSS-Alg made it possible to establish light attenuation at different wavelengths based on the knowledge of the concentration of these light-absorbing components.  相似文献   

9.
Light penetrance in lake kinneret   总被引:2,自引:2,他引:0  
The characteristics of light penetrance in Lake Kinneret, Israel, were observed over the years 1970 to 1973. Light measurements were made concurrently with those of algal speciation and biomass, chlorophyll concentrations and primary production. Vertical extinction coefficients of green light (filter VG9), the most penetrating spectral component, ranged from 0.15 (August 1970) to 0.93 In units m–1 (April 1970), reflecting the large differences between algal standing crops in non-bloom and bloom seasons. During the dinoflagellate bloom (Peridinium cinctum fa westii) from February through June, the increment of extinction coefficient per unit increase of chlorophyll concentration was 0.006 ln units mg–1 m2. The uneven vertical distribution of algae at this period caused irregularities in the depth curves of light penetrance. At other times, when the phytoplankton cells were more homogeneously dispersed with depth, regular light penetrance curves were observed; however, as previously noted (Rodhe, 1972), attenuation of algal photosynthetic activity often appeared to be regulated by the blue spectral component (filter BG 12). Ratios of absorbed to scattered light in the upper water column ranged from 85:15 to 75:25.  相似文献   

10.
Effects of seston on ultraviolet attenuation in Lake Biwa   总被引:3,自引:0,他引:3  
We examined the attenuation of underwater ultraviolet (UV) radiation and photosynthetically available radiation (PAR) in Lake Biwa, Japan, at offshore and inshore sites and under contrasting stratification and mixing regimes. There were large spatial differences in the water column transparency to both wavebands, despite little change in concentrations of dissolved organic carbon (DOC). The 1% of surface irradiance depth varied from 0.3 to 2.7 m at 305 nm, from 0.8 to 6.3 m at 380 nm, and from 2.3 to 12.8 m for PAR. Both PAR and UV transparency declined abruptly in the South Basin of the lake when a typhoon caused the resuspension of sediments. The water column ratio of UV to PAR increased by 30% at all stations over the course of a 3-week sampling period associated with the general increase in phytoplankton concentrations. At several sites, the diffuse attenuation coefficient for UV radiation deviated substantially from that predicted from UV-DOC models. A significantly positive linear relationship was found between UV attenuation (K d determined with a profiling UV radiometer) and the beam attenuation coefficient at 660 nm as measured by transmissometer. These results indicate that scattering and absorption by particulate matter can reduce UV transparency to below that inferred from DOC concentrations, and that current UV-exposure models should be modified to incorporate this effect. Received: March 21, 2001 / Accepted: August 17, 2001  相似文献   

11.
Depth limits and minimum light requirements of freshwater macrophytes   总被引:25,自引:1,他引:25  
1. Data for maximum colonization depth (Zc) of five groups of submerged macrophytes and light attenuation were collected for forty-five Danish lakes and 108 non-Danish lakes. The macrophyte groups were bryophytes, charophytes, caulescent angiosperms, rosette-type angiosperms and Isoetes spp. 2. The data showed systematic differences among the groups in the relationship of Zc to water transparency. In lakes with low transparency (Secchi disc transparency (Zs) less than 7 m) caulescent angiosperms and charophytes penetrated deepest followed by bryophytes and Isoetes spp. In more transparent lakes bryophytes grew deepest, followed by charophytes, caulescent angiosperms and Isoetes spp. Rosette-type angiosperms had the lowest Zc in all types of lakes. Charophytes and caulescent angiosperms had similar depth limits in lakes with Zs < 4 m but charophytes grew deeper in more transparent lakes. The depth limits of both groups were independent of light penetration in lakes with very low transparency (Zs < 1 m). The annual light exposure for the deepest growing macrophytes (bryophytes) was 20–95 mol photons m–2. 3. The relationship between Zc, macrophyte type and lake transparency could be explained by three distinct processes regulating Zc. In lakes with low transparency (Zs < 1 m), tall macrophytes (caulescent angiosperms and charophytes) compensate for light limitation by shoot growth towards the water surface and Zc is therefore independent of transparency. In lakes with medium transparency (1 m < Zs < 4 m) Zc for angiosperms, charophytes and Isoetes spp. is constrained by light attenuation in the water column, corresponding to a linear relationship between Zc and Zs. This pattern also applies to bryophytes, despite lake transparency. In transparent lakes, the minimum light requirement at Zc increased with increasing transparency for angiosperms, charophytes and Isoetes spp. 4. The minimum light requirements among submersed macrophytes (including marine macroalgae) depend on their plant-specific carbon value (plant biomass per unit of light-absorbing surface area) for the species/group, indicating that the light requirements of submersed plants are tightly coupled to the plants’ possibility to harvest light and hence to the growth form. 5. The light requirements increased on average 0.04% surface irradiance per degree increase in latitude corresponding to an average decrease in Zc of 0.12 m per degree latitude.  相似文献   

12.
The impact of UVB on the Antarctic phytoplankton photosystem II repair cycle, involving the rapidly cycled D1 protein, was studied during summer 2002. On sunny and overcast days, phytoplankton (from 1-m depth) were exposed to natural light (+UVB) and Mylar-screened (–UVB) conditions. Half of the samples from each treatment were inoculated with lincomycin, an inhibitor of synthesis of chloroplast-encoded proteins including the D1 protein. Blocking D1 repair caused significant Fv/Fm depressions on sunny days but had not effect on the overcast day. Most of the Fv/Fm depression was caused by PAR and UVA with a non-significant contribution from UVB. In the presence of D1 repair, suppressing UVB had no effect on Fv/Fm when the samples originated from a weakly stratified water column with no defined upper mixed layer (UML) while it alleviated Fv/Fm depression when the phytoplankton samples originated from within an UML deeper than the depth of UVB penetration. These results suggest that UVB had more effect on the D1 repair process than on the damage process itself but that phytoplankton sensitivity to surface UVB exposure was influenced by their previous light history, partly determined by the vertical structure of the water column.  相似文献   

13.
Rod spectral sensitivity data (λmax), measured by microspectrophotometry, were compiled for 403 species of ray-finned fishes in order to examine four hypothesized predictors of rod spectral sensitivity (depth, habitat, diet and temperature). From this database, a subset of species that were known to be adults and available on a published phylogeny (n = 210) were included in analysis, indicating rod λmax values averaging 503 nm and ranging from 477 to 541 nm. Linear models that corrected for phylogenetic relatedness showed that variation in rod sensitivity was best predicted by habitat and depth, with shorter wavelength λmax values occurring in fishes found offshore or in the deep sea. Neither diet, nor the interaction of diet and habitat, had significant explanatory power. Although temperature significantly correlated with rod sensitivity, in that fishes in temperate latitudes had longer wavelength rod λmax values than those in tropical latitudes, sampling inequity and other confounds require the role of the temperature to be studied further. Together, these findings indicate that fish rod λmax is influenced by several ecological factors, suggesting that selection can act on even small differences in fish spectral sensitivity.  相似文献   

14.
H. M. Dale 《Hydrobiologia》1986,133(1):73-77
In stratified lakes with high light penetration, the maximum depth at which macrophytes occur is frequently limited by temperature. At this depth a variety of species may be found. On the other hand, when the clarity of water limits the light penetration and the temperature at depth is sufficient for good plant growth, the plants occurring at greatest depth are ones that do not require photosynthetic oxygen for root growth. Such plants include the Charales, Isoetes, Utricularia and Ceratophyllum.  相似文献   

15.
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.  相似文献   

16.
Certain factors influencing phytoplankton productivity are accentuated in turbid waters. They include mixing, spectral quality shifts, scattering, light fluctuations, and overall light attenuation. Measurements of productivity is influenced by the presence of inorganic turbidity. Together with the above factors high turbidity causes difficulties to assess and model phytoplankton productivity. Estimations of B, Pm B, Ik and m only reflect on the physiological condition of the phytoplankton, which differs little between water types of temperate regions. Measurement of integral vertical productivity, efficiency and fractional absorption by the phytoplankton of light energy conversion, however, are greatly influenced by inorganic turbidity. Because of high ratios of mixing to euphotic depth, the critical mixing depth is one of the most important factors influencing overall productivity in turbid waters.  相似文献   

17.
Harding  William R. 《Hydrobiologia》1997,344(1-3):87-102
This paper reports on a two-year analysis of the wind climateand its effect on phytoplankton primary production in ashallow (mean depth = 1.9 m), hypertrophic South Africancoastal lake, Zeekoevlei. The lake is subject to continuousmixing of the euphotic zone (Z eu = 0.8 m), andcomplete mixing of the water column to the mean depth on adaily basis. Median annual wind speeds, prevailing fromeither the north or the south, were 6.4 m s–1. There wasan almost total absence of calms, measured as hourly meanwind speeds of <1 m s–1. Notwithstanding the highfrequency of mixing, the lake supports a dense population ofphytoplankton, dominated by Cyanophyte and Chlorophytespecies. Mean concentrations of chlorophyll-a were240 g l–1. The attenuation of photosyntheticallyavailable radiation, PAR, was high, with mean K dvalues of 6.4 m–1 and water transparencies of <0.5 m.Levels of primary productivity, determined using the lightand dark bottle oxygen method, were very high, comparable toor exceeding that of the most productive systems yet studied.Maximum volumetric productivity ranged from 525 to 1524 mg Cm–3 h–1, and was confined to the upper 0.5 m of thewater column. Daily areal productivity, P d,varied between 1.2 and 4.3 g C m–2 d–1, and that ofthe maximum chlorophyll-a specific photosynthetic rate,P B max, between 1.6 and 7.9 mg C (mgChl-a)–1 h–1. Primary production was limited bywater temperature and the attenuation of PAR. The highfrequency of wind-induced mixing resulted in regular mixingof the phytoplankton through the euphotic zone, and reducedthe overall importance of P max at a single layer inthe depth profile. Similarly, the regularity of mixing wasrecognized as a limitation of the incubation of bottle chainsto determine primary production levels.  相似文献   

18.
The turbulence correction terms for the photosynthetic oxygen production of phytoplankton given by BAUMERT (1974) are used for the case that the variations in velocity are exponentially autocorrelated. In case of the P(I)-relation by STEEMANN NIELSEN one can show that the value of the relative correction term is — in the worst case — less than, or equal to 1% if the depth of epilimnion (mixing depth) and the light intensity at the water surface are high. The demand for weak turbulence and light extinction — necessary for validity of the equations — is met for some real objects, especially for oligotrophic seas and lakes.  相似文献   

19.
20.
This paper shows that the sub-surface light regime in the offshore North Sea varies spatially and seasonally between different ecohydrodynamic regions, which is likely to have important implications for primary production and carbon and nutrient fluxes in different areas of the North Sea. Measurements of downward irradiance were collected using different instruments (i.e. water column-profiling instruments, semi-autonomous moorings, and remote sensing) at three ecohydrodynamically distinct sites in the North Sea: in the southern Bight (SB), at the Oyster Grounds (OG) and north of the Dogger Bank (ND). The ND site was the deepest, and had the lowest and least variable light attenuation coefficients (mean Kd(PAR) = 0.11 m?1). The onset of the phytoplankton spring bloom was earlier than at the other two sites. In summer, ND had low Kd(PAR) ~ 0.07 m?1 and light penetration was shifted towards blue-green wavelengths (490–560 nm), with water itself being one of the strongest contributors to overall attenuation. In contrast, the SB site was characterised by the highest and most variable values of Kd(PAR) (mean = 0.54 m?1), comparable to near-coastal waters, and the spring bloom started almost a month later than at the ND site. The vertical variability of the attenuation coefficient and the strong PAR attenuation in the blue region of the spectrum were the result of higher concentrations of phytoplankton, CDOM and SPM, due to riverine inputs, shallow depth and strong tidal mixing. The OG site showed intermediate conditions between the ND and SB sites with a mean Kd(PAR) = 0.23 m?1, and deepest penetration of irradiance in the green region of the spectrum at 560 nm. The implications of these results for phytoplankton growth and ecosystem modelling are discussed.  相似文献   

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