Phytoplankton pigment patterns and community composition in the northern South China Sea during winter

Phytoplankton pigment patterns and community composition were investigated in the northern South China Sea using high-performance liquid chromatography and the CHEMTAX software from February 11 to 23, 2009. We recognized four different vertical distribution patterns of pigments: chlorophyll a (Chl a)-like type, divinyl chlorophyll a (DV Chl a) type, even distribution type, and surface type. The average value of ratios of accessory photo-protective pigments (APP) to accessory photo-synthetic pigments was 0.8...     

Pigment characterization for the 2011 bloom in Qinhuangdao implicated "brown tide" events in China

A large-scale bloom occurred from May to June in 2011 in sea area near Qinhuangdao of the Bohai Sea, leading to huge damage of the scallop culture industry. Similar blooms have been observed in this region for three years. The causative species of the bloom, which dominated the phytoplankton community with the maximum cell density around 109 cell/L, could not be identified with morphological features due to the small cell size (~2 mm). A pigment analytical method was then adopted to analyze the pigment profile of the phytoplankton samples collected from the blooming sea area. It was found that pico-sized (<2 mm), nano-sized (2-20 mm), and bulk phytoplankton samples had similar pigment profile, representing the pigment signature of the bloom-causative species. The major pigments detected included 19-butanoyloxyfucoxanthin (But-fuco), fucoxanthin (Fuco), diadinoxanthin (Diad) and chlorophyll a (Chl a), and high content of But-fuco was the most significant characteristics of the phytoplankton samples. Based on the pigment composition and content, the bloom-causative species could be tentatively identified as pelagophyte, "type 8" group of haptophyte, or silicoflagellate. Some unique features of the bloom, such as the extremely high cell density, small-sized and But-fuco containing cells, occurring in early summer, and the feeding-cessation effects on scallops, suggest it be a "brown tide" event similar to those reported in the east coast of the United States of America. The recurrent "brown tide" events and their dramatic impacts on the shellfish mariculture industry in Qinhuangdao need close attention in the coming years.     

Impacts of internal waves on chlorophyll a distribution in the northern portion of the South China Sea

Internal waves can bring nutrients to the upper level of water bodies and facilitate phytoplankton photosynthesis. Internal waves occur frequently in the northern portion of the South China Sea and inflict an important effect on chlorophyll a distribution. In this study, in-situ observation and satellite remote sensing data were used to study the effects of internal waves on chlorophyll a distribution. Based on the in-situ observations, lower chlorophyll a concentrations were present in the middle and bottom level in areas in which internal waves occur frequently, while the surface chlorophyll a distribution increased irregularly, and a small area with relatively higher chlorophyll a concentrations was observed in the area around the Dongsha Island. Satellite remote sensing showed that the chlorophyll a concentration increased in the area near Dongsha Island, where internal waves frequently occurred. The results of the increased chlorophyll a concentration in the surface water near Dongsha Island in the northern portion of the South China Sea indicated that internal waves could uplift phytoplankton and facilitate phytoplankton growth.     

Primary productivity and chlorophyll a in Prydz Bay and its mouth in Antarctica during the austral summer of 1999/2000

The investigation of phytoplankton standing stock, euphotic layer and photosynthesis rate were carried out in 19 - 27 of January, 2000 at three longitudinal sections (70°30＇E, 73°00＇E and 75°30＇E). The results showed that the high value of chlorophyll a concentration was in inshore bay, polynya and the continental slope of the investigated sea area. At various investigated stations, average chlorophyll a concentration at sub-surface layer (25 m) was higher than that at surface layer; its concentration at the deeper layers of over 50 m decreased with increasing depth. At anchor station, the maximum chlorophyll az concentration appeared at surface layer in Antarctic summer's afternoon while the minimum value appeared in the morning; chlorophyll a concentration at water layer of 0 - 25 m was obviously higher than that at deep water layer, being related to the releasing of ice-algae. High productivity was in inshore bay and polynya of continental shelf. Chlorophyll a concentration at surface layer is closely correlated to the dissolved oxygen concentration in seawater. The sea ares with chlorophyll a concentration of over 1.0ag/dms may be the convergence of CO2 and that of below 1.0 μg/dms may be the source of CO2.     

Strategy of photo-protection in phytoplankton assemblages in the Kongsfjorden,Svalbard, Arctic

Photo-protective functions were investigated in phytoplankton assemblages at Kongsfjorden, Svalbard in spring, using their UV-absorbing compounds (mycosporine-like amino acids (MAAs)), xanthophyll pigments (diadinoxanthin (DD) and diatoxanthin (DT)) and < beta >- dimethylsulphoniopropionate (< beta >_-DMSP). The dominant phytoplankton species in the inner bay were dominated by Phaeocystis spp. and nanoflagellates, while the offshore waters were dominated by Thalassiosira spp. In the inner bay, UVabsorbing compounds and xanthophyll pigments exhibited higher ratios of MAA to chlorophyll a (MAA:chl a ratio), and both DD and DT to chlorophyll a (DD:chl a ratio and DT:chl a ratio), respectively. Thus, the photoprotective-pigments such as DD and DT appear to complement MAAs in the natural phytoplankton assemblage. However, the ratio of < beta >_-DMSP to chlorophyll a (< beta >_-DMSP:chl a ratio) did not show a distinct spatial distribution according to environmental factors or interspecies differences. In this study, we found that photoprotective compounds occurred in a manner dependent on the phytoplankton species composition in Kongsfjorden Bay, where Phaeocystis is the dominant species.     

Variation of bacteria biomass and its possible controlling factors in the East China Sea

Two surveys were performed for determining bacteria biomass (BB), temperature, salinity, chlorophyll a (chl-a) and nutrient concentrations at 11 stations with three sampling depths in the high-incidence regions of red tide in the East China Sea (ECS) in the spring of 2006. Temperature and salinity increased from nearshore to offshore region and from high latitude to low latitude in the two cruises of 2006. BB were between 0.3–5.2 mgC m⁻³ (about 2.1 mgC m⁻³ on average) and 0.2–6.0 mgC m⁻³ (about 2.7 mgC m⁻³ on average) respectively in the two cruises. BB in the surface layer decreased from the Changjiang River estuary to high sea and from low latitude to high latitude. The results showed that bacterial growth was regulated by temperature, primary production and inorganic nutrient concentrations depending on different hydrographic conditions. In the surface and middle layers where the primary production can supply enough organic substrate, temperature was the main factor to control bacteria biomass. BB showed a good correlation between the surface and middle layers in both cruises. The distribution of nutrients during both cruises showed a similar decreasing trend from nearshore region and high latitude to offshore region and low latitude. High BB values were mainly recorded from samples in the middle layer where chl-a concentrations were also high, indicating primary production being strongly correlated with temperature over the ECS shelf. In the offshore area, phosphate and silicate became limiting factors for phytoplankton growth with indirect influence on BB. Bacteria played an important role in nitrogen regeneration process turning organic nitrogen to inorganic forms such as NH₄ ⁺. The increasing ratio of NH₄ ⁺/DIN could be a proof of that.     

Decadal variability of chlorophyll a in the South China Sea: a possible mechanism

Four climatologies on a monthly scale (January, April, May and November) of chlorophyll a within the South China Sea (SCS) were calculated using a Coastal Zone Color Scanner (CZCS) (1979-1983) and the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) (1998-2002). We analyzed decadal variability of chlorophyll a by comparing the products of the two observation periods. The relationships of variability in chlorophyll a with sea surface wind speed (SSW), sea surface temperature (SST), wind stress (WS), and mixed layer depth (MLD) were determined. The results indicate that there is obvious chlorophyll a decadal variability in the SCS. The decadal chlorophyll a presents distinct seasonal variability in characteristics, which may be as a result of various different dynamic processes. The negative chlorophyll a concentration anomaly in January was associated with the warming of SST and a shallower MLD. Generally, there were higher chlorophyll a concentrations in spring during the SeaWiFS period compared with the CZCS period. However, the chlorophyll a concentration exhibits some regional differences during this season, leading to an explanation being difficult. The deepened MLD may have contributed to the positive chlorophyll a concentration anomalies from the northwestern Luzon Island to the northeastern region of Vietnam during April and May. The increases of chlorophyll a concentration in northwestern Borneo during May may be because the stronger SSW and higher WS produce a deeper mixed layer and convective mixing, leading to high levels of nutrient concentrations. The higher chlorophyll a off southeastern Vietnam may be associated with the advective transport of the colder water extending from the Karimata Strait to southeastern Vietnam.     

A bio-optical inversion model to retrieve absorption contributions and phytoplankton size structure from total minus water spectral absorption using genetic algorithm

We propose a bio-optical inversion model that retrieves the absorption contributions of phytoplankton and colored detrital matter(CDM),as well as the phytoplankton size classes(PSCs),from total minus water absorption spectra.The model is based on three-component separation of phytoplankton size structure and a genetic algorithm.The model performance was tested on two independent datasets(the NASA bio-Optical Marine Algorithm Dataset(NOMAD) and the northern South China Sea(NSCS) dataset).The relationships between the estimated and measured values were strongly linear,especially for aCDM(412),and the Root Mean Square Error(RMSE) of the CDM exponential slope(SCDM) was relatively low.Next,the inversion model was directly applied to in-situ total minus water absorption spectra determined by an underwater meter during a cruise in September 2008,to retrieve the phytoplankton size structure in the seawater.By comparing the measured and retrieved chlorophyll a concentrations,we demonstrated that total and size-specific chlorophyll a concentrations could be retrieved by the model with relatively high accuracy.Finally,we applied the bio-optical inversion model to investigate changes in phytoplankton size structure induced by an anti-cyclonic eddy in the NSCS.     

HPLC pigment profiles of 31 harmful algal bloom species isolated from the coastal sea areas of China

Chemotaxonomy based on diagnostic pigments is now a routine tool for macroscopic determination of the composition and abundance of phytoplankton in various aquatic environments. Since the taxonomic capability of this method depends on the relationships between diagnostic pigments and chlorophyll a of classified groups, it is critical to calibrate it by using pigment relationships obtained from representative and/or dominant species local to targeted investigation area. In this study, pigment profiles of 31 harmful algal bloom(HAB) species isolated from the coastal sea areas of China were analyzed with high performance liquid chromatography(HPLC). Pigment compositions, cellular pigment densities and ratios of pigments to chlorophyll a were determined and calculated. Among all these species, 25 kinds of pigments were detected, of which fucoxanthin, peridinin, 19'-butanoyloxyfucoxanthin, 19'-hexanoyloxyfucoxanthin, violaxanthin, and antheraxanthin were diagnostic pigments. Cellular pigment density was basically independent of species and environmental conditions, and therefore was recommended as a bridge to compare the results of HPLC-CHEMTAX technique with the traditional microscopy method. Pigment ratios of algal species isolated from the coast of China, especially the diagnostic pigment ratios, were higher than those from other locations. According to these results, pigment ratio ranges of four classes of phytoplankton common off the coast of China were summarized for using in the current chemotaxonomic method. Moreover, the differences of pigments ratios among different species under the same culturing conditions were consistent with their biological differences. Such differences have the potential to be used to classify the phytoplankton below class, which is meaningful for monitoring HABs by HPLC-CHEMTAX.     

Seasonal variations in the phytoplankton community and the relationship between environmental factors of the sea around Xiaoheishan Island in China

Seasonal variations in the phytoplankton community and the relationship between environmental factors of the sea area around Xiaoheishan Island are investigated in the present study. Xiaoheishan Island is located at 37°58′14″N and 120°38′46″E in Shandong Province, China. A total of 65 species of phytoplankton belonging to three phyla and 27 genera were identified, with Bacillariophyta having the largest number of species. The annual average chlorophyll a concentration for this area was 3.11 μg/L, and there occurs a Skeletonema costatum bloom in winter. The Shannon-Weaver indexes(log_2) of the phytoplankton from all stations were higher than 1, and the Pielou indexes were all higher than 0.3. The results of the canonical correspondence analysis(CCA) indicated that water temperature, PO_4~(3ˉ) and Cu were the environmental factors that had the greatest influence on the distribution of the phytoplankton community throughout the entire year. Although the concentration of heavy metal is well up to the state standards of the first grade of China(GB 3097-1997), these metals still have an impact on the phytoplankton community from this area.     

Size-fractionated phytoplankton biomass in autumn of the Changjiang （Yangtze） River Estuary and its adjacent waters after the Three Gorges Dam construction

A cruise was undertaken from 3rd to 8th November 2004 in Changjiang （Yangtze） River Estuary and its adjacent waters to investigate the spatial biomass distribution and size composition of phytoplankton. Chlorophyll-a （Chl-a） concentration ranged 0.42-1.17 μg L^-1 and 0.41-10.43 μg L^-1 inside and outside the river mouth, with the mean value 0.73 μg L^-1 and 1.86 μg L^-1, respectively. Compared with the Chl-a concentration in summer of 2004, the mean value was much lower inside, and a little higher outside the river mouth. The maximal Chl-a was 10.43 μg L^-1 at station 18 （122.67°E, 31.25°N）, and the region of high Chl-a concentration was observed in the central survey area between 122.5°E and 123.0°E. In the stations located east of 122.5°E, Chl-a concentration was generally high in the upper layers above 5 m due to water stratification. In the survey area, the average Chl-a in sizes of 〉20 μm and 〈20 μm was 0.28 μg L^-1 and 1.40 μg L^-1, respectively. High Chl-a concentration of 〈20 μm size-fraction indicated that the nanophytoplankton and picophytoplankton contributed the most to the biomass of phytoplankton. Skeletonema costatum, Prorocentrum micans and Scrippsiella trochoidea were the dominant species in surface water. The spatial distribution of cell abundance of phytoplankton was patchy and did not agree well with that of Chl-a, as the cell abundance could not distinguish the differences in shape and size of phytoplankton cells. Nitrate and silicate behaved conservatively, but the former could probably be the limitation factor to algal biomass at offshore stations. The distribution of phosphate scattered considerably, and its relation to the phytoplankton biomass was complicated.     

Picoplankton distribution in different water masses of the East China Sea in autumn and winter

Picoplankton distribution was investigated in different water masses of the East China Sea in November,2006 and February,2007.The autumn and winter cruises crossed three major water masses:the coastal water mass(CWM),the mixed water mass(MWM),which forms on the continental shelf,and the Kuroshio water mass(KWM).Picoplankton composition was resolved into four main groups by flow cytometry,namely Synechococcus,Prochlorococcus,picoeukaryotes,and heterotrophic bacteria.The average abundances of Synechococcus,picoeukaryotes,and heterotrophic bacteria were(0.63±10.88)×103,(1.61±1.16)×103,(3.39±1.27)×105 cells/mL in autumn and(6.45±8.60)×103,(3.23±2.63)×103,(3.76±1.37)×105 cells/mL in winter,respectively.Prochlorococcus was not found in the CWM and seldom observed in surface samples in either season.However,Prochlorococcus was observed in the MWM and KWM(approximately 10 3 cells/mL) in both autumn and winter.Synechococcus distribution varied considerably among water masses,with the highest levels in KWM and lowest levels in CWM.The depth-averaged integrated abundance of Synechococcus was approximately 5-fold higher in KWM than in CWM,which may be due primarily to water temperature.In the MWM,Synechococcus was resolved as two subgroups;the presence of both subgroups was more common in autumn.Picoeukaryote abundance varied less among water masses than Synechococcus,and heterotrophic bacteria depth-averaged integrated abundance exhibited the smallest seasonal variations with respect to water mass.Correlation analysis showed that relationships between picoplankton abundances and environmental factors(temperature,nutrients,and chlorophyll a) differed among the three water masses,suggesting that the three water masses have different effects on picoplankton distribution(particularly Synechococcus).     

Impact of microzooplankton and Calanus sinicus (Brodsky, 1962) on phytoplankton in the Yellow Sea during early summer

Dilution incubations and Calanus sinicus addition incubations were simultaneously conducted at five stations in the Yellow Sea in June of 2004 to evaluate the impact of microzooplankton and Calanus sinicus on phytoplankton based on the Chlorophyll a (Chl-a) levels. The Chl-a growth rates (k) ranged from 0.60–1.67 d⁻¹, while microzooplankton grazed the Chl-a at rates (g) of 0.29–0.62 d^t-1. The addition of C. sinicus enhanced the Chl-a growth rate (Z) by 0.004–0.037 d⁻¹ ind.⁻¹ L. C. sinicus abundance ranged from 84.1–160.9 ind. m⁻³, which occupied 90.7%–99.1% of the copepod (>500 μm) population. The in-situ increase in phytoplankton by C. sinicus community was estimated to be 0.000 4–0.005 9 d⁻¹. These results showed that microzooplankton were the main grazers of phytoplankton, while C. sinicus induced a slight increase in the levels of phytoplankton.     

Variation in downwelling diffuse attenuation coefficient in the northern South China Sea

The diffuse attenuation coefficient for downwelling irradiance （Kd（λ）） is an important parameter for ocean studies. Based on the optical profile data measured during three cruises in the northern South China Sea in autumn from 2003 to 2005, variations in the Kd（λ） spectra were analyzed. The variability of Kd（λ） shows much distinct features in both magnitude and spectra pattern, it is much higher in coastal waters than that of open oceanic waters; and the blue-to-green （443/555） ratio of Kd（λ） tends to increase with chlorophyll a concentration （[Chl-a]） from open ocean to coastal waters. These characteristics can be explained most by the increase of aw＋p（443）/aw＋p（555） with [Chl-a]. In short waveband, the relation between Kd（λ）-Kw（λ） and [Chl-a] can be well described by a power law function, indicating the large contribution of phytoplankton to the variations in Kd（λ）. As for the spectral model of the diffuse attenuation coefficient, there are good linear relationships between Kd（490） and Kd（λ） in other wavelengths with own slope and intercept of a linear functions in the spectral range 412-555 nm. Kd（490） is well correlated with the spectral ratio of remote sensing reflectance; and should enough measurement data are given, this empirical algorithm would be used in the Kd（λ） retrieval from ocean color satellite data. The variation in Kd（λ） provides much useful information for us to study the bio-optical property in the northern South China Sea.     

Response of phytoplankton community structure and size-fractionated Chlorophyll <Emphasis Type="Italic">a</Emphasis> in an upwelling simulation experiment in the western South China Sea

The South China Sea (SCS), which is the largest marginal sea in the western tropical Pacific, plays an important role in regional climate change. However, the research on the phytoplankton community structure (PCS) response to the upwelling remains inadequate. In January 2014, the upwelling simulation experiment was performed in the western SCS. Results indicate that the nutrient-rich bottom water not only increased the total Chlorophyll a (Chl a) concentrations, but would potentially altered the PCS. Due to new nutrients added, microphytoplankton had more sensitivity response to nutrient uptake than other phytoplankton groups. The variation of nutrients induced by formation, weakening and disappearance of upwelling resulted in phytoplankton species succession from cyanophyta to bacillariophyta. It may be the leading factor of the changes in PCS and size-fractionated Chl a. The initial concentration of DIP less than 0.1 μmolL^?1 could not sustain the phytoplankton growth. This indicates that phosphorus may be the limiting factor in the western SCS.     

Records of bulk organic matter and plant pigments in sediment of the “red-tide zone” adjacent to the Changjiang River estuary

Cultural eutrophication caused by nutrient over-enrichment in coastal waters will lead to a cascading set of ecosystem changes and deleterious ecological consequences,such as harmful algal blooms(HABs) and hypoxia.During the past two decades since the late 1990s,recurrent large-scale HABs(red tides)and an extensive hypoxic zone have been reported in the coastal waters adjacent to the Changjiang River estuary.To retrieve the history of eutrophication and its associated ecosystem changes,a sediment core was collected from the "red-tide zone" adjacent to the Changjiang River estuary.The core was dated using the~(210)Pb radioisotope and examined for multiple proxies,including organic carbon(OC),total nitrogen(TN),stable isotopes of C and N,and plant pigments.An apparent up-core increase of OC content was observed after the 1970s,accompanied by a rapid increase of TN.The concurrent enrichment of δ~(13)C and increase of the C/N ratio suggested the accumulation of organic matter derived from marine primary production during this stage.The accumulation of OC after the 1970 s well reflected the significant increase of primary production in the red-tide zone and probably the intensification of hypoxia as well.Plant pigments,including chlorophyll a,p-carotene,and diatoxanthin,showed similar patterns of variation to OC throughout the core,which further confirmed the important contribution of microalgae,particularly diatoms,to the deposited organic matter.Based on the variant profiles of the pigments representative of different microalgal groups,the potential changes of the phytoplankton community since the 1970s were discussed.     

Picophytoplankton abundance and community structure in the Philippine Sea, western Pacific

Flow cytometric determinations of the abundance distribution and community structure of picophytoplankton (i.e., Prochlorococcus spp., orange fluorescence Synechococcus spp. and picoeukaryotes) were used for samples taken from the Philippine Sea in the western tropical Pacific Ocean from September to October of 2004. A fluorescence probe was employed to detect Chlorophyll a (Chl a). Abundances of Prochlorococcus spp., orange fluorescence Synechococcus spp. and picoeukaryotes ranged from 0.1 to 58×10³ cells ml^?1, 0.38 to 17×10² cells ml^?1 and 0.42 to 26×10² cells ml^?1, respectively. Synechococcus spp. and picoeukaryotes co-occurred in relatively shallow water with the maximum abundance observed at 50 to 70 m depth, while Prochlorococcus spp. only occurred in the 70 to 200 m layer. Prochlorococcus spp. was the dominant picophytoplankton population in terms of abundance and biomass. The cell size and carbon biomass content were estimated for the three picophytoplankton groups. In addition, among the three groups of picophytoplankton, the relative contribution of red fluorescence to the total red fluorescence varied with depth. The fluorescence and light scatter properties of individual cells indicated that in the upper 100 m layer, picoeukaryotes were a major contributor to total red fluorescence, while at the depth below 100 m, Prochlorococcus spp. and Synechococcus spp. made an important contribution to the total red fluorescence.     

Distributions of picophytoplankton and phytoplankton pigments along a salinity gradient in the Changjiang River Estuary,China

We investigated the abundance of different picophytoplankton groups and the phytoplankton pigment ratio in relation to environmental factors such as nutrients and suspended solids along a salinity gradient in the Changjiang River Estuary. The average numbers of Synechococcus spp.(Syn) and picoeukaryotes(Euk) were(2.7 ±5.1) ×103 and(1.1±1.4) ×103 cells m L-1, respectively. Prochlorococcus spp.(Pro) was only found in the high-salinity brackish water with the concentration of 3.0×103 cells m L-1. Syn and Euk numbers both tended to increase offshore and Syn showed a larger variation in cell abundance than Euk. The contribution of picophytoplankton to total phytoplankton biomass increased with increasing salinity and decreasing nutrient concentrations from the estuary to the open ocean. The response of different picophytoplankton groups to environmental variables was different. Water temperature was more important in its control over Euk than over Syn, while nutrients were more important in their influence over Syn than over Euk. Phytoplankton pigment ratios were different in the three different ecological zones along the salinity gradient(i.e., freshwater zone with 0-5 range, fresh and saline water mixing zone with 5-20 range, and high-salinity brackish water zone with 20-32 range), where three different phytoplankton communities were discovered, suggesting that phytoplankton pigment ratios can be considered as a complementary indicator of phytoplankton community structure in the Changjiang River Estuary.     

THE BIOGEOCHEMISTRY OF PHOTOSYNTHETIC PIGMENTS IN THE JIULONG RIVER ESTUARY AND WESTERN XIAMEN BAY

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Tempo-spatial variation of nutrient and chlorophyll-α concentrations from summer to winter in the Zhangzi Island Area (Northern Yellow Sea)

Nutrient and Chlorophyll-a (Chl-a) concentrations were investigated monthly along three transects extending from a mariculture area to open waters around the Zhangzi Island area from July to December 2009. The objective of this study is to illustrate food availability to the bottom-sowed scallop Patinopecten yessoensis under the influences of the Yellow Sea Cold Water Mass (YSCWM), freshwater input and feedbacks of cultivated scallops. Significant thermal stratification was present in open waters from July to October, and salinity decreased in July and August in surface layers in the mariculture area. Nutrient concentrations increased with depth in both areas in summer, but were similar through water column in November and December. On average, nutrient increased from summer to autumn in all components except ammonia. Nutrient concentrations lower than the minimum thresholds for phytoplankton growth were present only in upper layers in summer, but stoichiometric nitrogen limitation existed in the entire investigation period. Column-averaged Chl-a concentration was lower in open waters than in mariculture area in all months. It increased significantly in mariculture area in August and October, and was less variable in open waters. Our results show that nutrients limitation to phytoplankton growth is present mainly in upper layer in association with stratification caused by YSCWM in summer. Freshwater input and upwelling of nutrients accumulated in YSCWM can stimulate phytoplankton production in mariculture area. Farming activities may change stoichiometric nutrient ratios but have less influence on Chl-a concentration.