The potential of photosynthetic aquatic species as sources of useful cellulose fibers—a review

Photosynthetic aquatic species, i.e., micro- and macroalgae and fresh or salt water plants, contain cellulose or other fibrous materials potentially suitable for paper making. Photosynthetic aquatic species having cellulosic or fibrous characteristics necessary for paper production were reviewed. These characteristics include overall fiber content, fiber size and morphology, and fiber composition. Several species of algae and aquatic plants are reported to possess cellulose in quantities greater than 10 % of total dry weight, and in general, the cellulose content in aquatic species is lower than that of most wood species. Commercial application of these aquatic algal or plant materials has been limited to simple milling, and no commercial applications utilizing processes to isolate the cellulosic fibers from these materials have yet been found.     

Determination of biomass dry weight of marine microalgae

Total biomass dry weight (DW) and ash free dry weight (AFDW) of five species of marine microalgae, Dunaliella sp., Isochrysis galbana, Nannochloropsis sp., Nitzschiaclosterium and Porphyridium cruentum, retained on filter paper, were determined. Dunaliella and Isochrysis cells have no cell wall; Nannochloropsis, Nitzschia and Porphyridium possess a cell wall and Nitzschia and Porphyridium cells are covered by silica and mucilage coating, respectively. In all these algae, DW of non-washed samples was at least 1.2 times higher than those washed by distilled water,0.9% sodium chloride, 0.5 M ammonium formate or 0.5 M ammonium bicarbonate. DW of 0.9% sodium chloride washed samples was more than 0.8 times higher than the other three washed samples. In most of the cases, there was no significant difference between DW of samples washed by ammonium formate and ammonium bicarbonate solutions (p>0.05). The AFDW of the non-washed algal samples was about twice that washed samples, and could be accounted for by volatile component in the sea water medium. Isotonic solution of ammonium bicarbonate is a satisfactory washing agent for algal cells for dry weight determination. This revised version was published online in September 2006 with corrections to the Cover Date.     

THE EFFECT OF A HORN SNAIL ON ULVA EXPANSA (CHLOROPHYTA): CONSUMER OR FACILITATOR OF GROWTH?1

To determine whether California horn snails are more likely to be consumers or facilitators of Ulva expansa (Setch) S. & G. growth in estuaries, we conducted manipulative experiments that evaluated algal growth and the movement of N between the water column, algal tissue, and, in the second experiment, sediments. Algae grew poorly in the absence of sediments, drawing on their own sequestered N supplies (3.5% of dry weight reduced to <2%) and N released by snails and by depleting inorganic N in the water column. There was no evidence of consumption when snail densities ranged from 0 to 900.m^?2 (0, 3, 6, and 9 per aquarium), as algal growth was similar for all snail densities, and snail lengths did not increase during the 21–d experiment. when sediment was provided, N was depleted in the sediment and enhanced in the algal tissue. As in the first experiment, the water column was depleted of inorganic N and enriched with organic N, mostly in the dissolved form. Because both snails and macroalgae often dominate the shallow waters of southern California's lagoons and estuaries, our evidence that the snails are primarily facilitators of algal growth (via transfer of N from sediments to the water column) suggests that snails may play an important role in both food web and N dynamics.     

Degradation dynamics of lignocellulose materials by wood-rottingPleurotus fungi

When grown on wheat straw,Pleurotus decomposes both the lignin and the cellulose components of the substrate. The course of degradation differs during growth and fructification. The losses of dry mass during growth were about 20 %. The absolute amount of hemicelluloses, cellulose and lignin was decreasing. Hemicelluloses and lignin were degraded at a higher rate than cellulose. The total mass losses of the substrate after fructification were 32 to 45 %. Cellulose was consumed at a higher rate than lignin.     

PARTICULATE ORGANIC MATTER AS AN ALTERNATIVE NUTRIENT SOURCE FOR TROPICAL SARGASSUM SPECIES (FUCALES, PHAEOPHYCEAE)

Large Fucales are abundant on coastal coral reefs of the Great Barrier Reef, but are often limited by the availability of inorganic nutrients. Particle loads in these reef waters are high, which is generally perceived as detrimental for aquatic plants due to a reduction of light. Here, I provide evidence that several abundant Sargassum species supplement their nutrient supply with nutrients derived from the layer of particulate matter (PM) deposited on their thalli. In experiments involving removal or addition of PM, growth rates of Sargassum spp. were up to 180% higher when particles were present on the thalli. Tissue nitrogen and phosphorus levels of thalli with a surface PM layer were 10%–30% higher than those of thalli without PM. The amount of PM deposited in situ on thalli of five species of brown algae ranged from 0.6 to 0.9 g C_org·g⁻¹ dry weight alga, depending on the species' morphology. I suggest that a nutrient-rich diffusive boundary layer is created on the thallus surface by an epiphytic microbial community that remineralizes the particulate nutrients. When algal growth is nutrient limited, the use of particle-derived nutrients as a source alternative to nutrients in the water column may outweigh any potential adverse effects of the thallus particle layer.     

Phosphorus Removal from Natural Waters Using Controlled Algal Production

A series of experiments designed to demonstrate the potential of using managed, attached algal production to permanently remove excess phosphorus from agricultural run-off is described. The experiments were carried out on a secondary canal in the New Hope South region of the Florida Everglades Agricultural Area from October, 1991, to May, 1992. Natural algal populations of periphyton, including species of the genera Cladophora, Spirogyra, Enteromorpha, Stigeoclonium, and a variety of filamentous diatoms such as Eunotia and Melosira, were grown on plastic screens in raceways, under a wave surge regime. Considerable biomass production of algae occurred, and the resulting algal canopy also trapped plankton and organic particulates from the water column. A seven- to eight-day harvest interval was determined to be optimal, and both hand harvesting and vacuum harvesting were employed. The vacuum device is applicable to large scale-up. In source water having total phosphorus concentrations of 0.012–0.148 ppm, mean macro-recovery dry biomass production levels of 15–27 g/m²/day were achieved. The lower rates occurred in the winter, the higher rates in the late spring. Two techniques were employed to reduce losses of fine material at harvest during the March to May period. Gravity sieving increased mean dry production levels to 33–39 g/m²/day. The mean phosphorus content of harvested biomass ranged from 0.34% to 0.43%. Total phosphorus removal rates during the spring period of average solar intensity and low nutrient supply, by methods demonstrated in this study, ranged from 104 to 139 mgTP/m²/day (380–507 kgP/ha/year). Over the incoming nutrient range studied, phosphorus removal was independent of concentration and was 16.3% of total phosphorus for 15 m of raceway. Up-stream-downstream studies of overflowing water chemistry (total P, total dissolved -P, orthophosphate -P) showed highly -significant reductions of all phosphorus species. Total phosphorus reduction closely correlated with phosphorus yield from biomass removal. Yearly, minimum phosphorus removal rates are predicted that are 100–250 times that achieved both experimentally and in long-term, large-area wetland systems. Engineering scale-up to systems of hundreds of acres is being studied.     

Increasing salinity tolerance of grain crops: Is it worthwhile?

The productivity of wheat and barley was compared in soils of different salt concentrations with a limited water supply. Productivity was assessed as total dry weight or dry weight per unit of water used (water use efficiency, WUE). Barley achieved the highest productivity because it used more of the available water and it had a greater WUE for above-ground dry weight. However, when WUE for total organic weight of roots and shoots was determined, or WUE was corrected for grain production, wheat and barley had the same productivity. In two experiments in drying soils with different salt concentrations but the same amount of soil water, wheat and barley had a higher dry weight than salt-tolerant grasses and they were more productive than C₄ halophytes and non-halophytes when adjusted for water use. In one experiment, sown at a low plant density, barley and wheat used less water than some halophytes and they completed their life cycle leaving some water behind in the soil. Their higher WUE did not compensate for their lower water use. However, when all species were sown at a high density, wheat and barley were either as productive or more productive than the most salt-tolerant species, including a C₄ halophyte, as they used all the available water and had the highest WUE. A sunflower cultivar was similary more productive than a salt-tolerant relative. The contribution that salt-tolerant relatives of wheat, barley and sunflower can make to genetically improving the productivity of these species in dry saline soils is questioned.     

Summer bacterial populations in Mississippi River Pool 19: implications for secondary production

Bacterial populations were sampled at 37 sites in Mississippi River Pool 19. Bacterial biomass was calculated from direct epifluorescent cell counts. Bacterial production was estimated by incubating cells in situ in predator-free water inside membrane chambers and the frequency of dividing cells. Bacterial biomass in the water column ranged from 0.05 to 1.13 mg C ^-1, biomass in the vegetated areas of the pool was significantly higher than that in other habitats (P < 0.05, ANOVA). Biomass in sediments (to a depth of 10 cm) ranged from 24 to 1,073 mg C m^-2, biomass in muddy sediments was significantly higher (P < 0.05) than that in sandy sediments. Biomass on the submersed surfaces of hydrophytes was 0.06–4.90 mg bacterial C g^-1 dry weight of plant material. The vegetated habitat (water column plus vegetation) contained approximately 45 times the concentration of bacterial carbon found in nonvegetated main channel border areas and more than 100 times the concentration in the main river channel.Bacterial production rates in the water column of a vegetated section of the pool ranged from 0.03 to 3.28 g C m ^-3 s d ^-1 ; production (m ^-3) in a vegetation bed was 5.5 times that in the adjacent nonvegetated channel border areas and approximately 50 times that in the main channel. Aquatic macrophytes and associated microorganisms may be capable of providing significant inputs of carbon to secondary consumers in the pool during the summer low flow.     

Algae-associated marine molluscs in the Azores

The raolluscan fauna of a special habitat, the high intertidal algal turf, was studied at four stations on the island of Sao Miguel, Azores. The number of species found ranged from 13 to 23. However, a small group of only six species (the bivalve Lasaea adansoni and the gastropods Pisinna punctulum, Omalogyra atomus, Fossarus ambiguus, Skeneopsis planorbis and Alvania postrema) accounted for more than 80% of all the specimens collected. Mollusc density reached values of 20000 specimens per 100 g algal dry weight. The abundance and number of species of molluscs was comparable with those reported for algal zones lower on the Azorean shores.
Wave exposure and seasonality effects were found to be small, presumably because of the relatively mild abiotic conditions and the protective role of the turf. The algal substrata influenced the phytal molluscan community in two ways. First, the abundance of molluscs was significantly correlated with algal dry weight. Secondly, molluscan abundance and diversity were influenced by the algal composition of the turf. A rich algal composition, with several species of branched fleshy and coralline algae, was associated with a rich molluscan fauna. The dominance of coralline algae resulted in a abundant but species-poor fauna, while an almost monospecific turf of Gigartina was poor in both species and number of molluscan fauna.     

Cellulose, hemicelluloses, lignin and ash content of some organic materials and their suitability for use as paper pulp supplements

Freshwater algal biomass and orange and lemon peels were assessed as tissue paper pulp supplements. Cellulose and hemicellulose contents of algal biomass were 7.1% and 16.3%, respectively, whereas for citrus peels cellulose content ranged from 12.7% to 13.6% and hemicellulose from 5.3% to 6.1%. For all materials, lignin and ash content was 2% or lower, rendering them suitable for use as paper pulp supplements. The addition of algal biomass to paper pulp increased its mechanical strength significantly. However, brightness was adversely affected by chlorophyll. The addition of citrus peels in paper pulp had no effect on breaking length, increased bursting strength and decreased tearing resistance. Brightness was negatively affected at proportions of 10%, because citrus peel particles behave as coloured pigments. The cost of both materials is about 45% lower than that of conventional pulp, resulting in a 0.9-4.5% reduction in final paper price upon their addition to the pulp.     

Screening of marine microalgae for bioremediation of cadmium-polluted seawater

Twenty four strains out of 191 marine microalgal strains exhibited cadmium (Cd) resistance. They were tested for their Cd removal ability in growth media containing 50 μM Cd. Six strains out of 19 green algae and one out of five cyanobacteria removed more than 10% of total Cd from the medium. The marine green alga Chlorella sp. NKG16014 showed the highest removal of Cd 48.7% of total. Cd removal by NKG16014 was further quantitatively evaluated by measuring the amount of cell adsorption and intracellular accumulation. After 12 days incubation, 67% of the removed Cd was accumulated intracellularly and 25% of the Cd removed was adsorbed on the algal cell surface. The maximum Cd adsorption (q_max) was estimated to be 37.0 mg Cd (g dry cells)⁻¹ using the Langmuir sorption model. The Cd removal by freeze-dried NKG16014 cells was also determined. Cd was more quickly adsorbed by dried cells than that by living cells, with a q_max of 91.0 mg Cd (g dry cells)⁻¹.     

Physiological effects of lactulose and inulin in the caecum of rats

A model experiment was performed on rats to evaluate the effect of partial or total substitution of saccharose (S) and cellulose (C) by preparations of lactulose and inulin on the development and metabolism of the caecum. In the experimental diets given to rats for 4 weeks, the examined preparations were administered either with an equivalent amount of cellulose (each at 4% of the diet) or as sole source of dietary fibre at 8% of the diet. Compared to the saccharose group cellulose had no effect, and low doses of lactulose and inulin in the diet increased to a medium extent the weight of the caecum wall and caecal digesta. The addition of lactulose and inulin at 8% increased significantly the content of caecal digesta (4.62 and 4.11 g/100g BW, respectively) and the weight of the caecal wall (1.10 and 0.86 g/100g BW, respectively), compared to the groups with saccharose and cellulose (0.73, 0.90 and 0.24, 0.28 g/100g BW, respectively). Cellulose and cellulose partially-substituted with lactulose and inulin caused an increase in the dry matter content of caecal digesta (26.5-27.5%), compared to other groups (21.8-22.8%). The administration of lactulose and inulin preparations was accompanied by a significant drop in pH (5.47-5.81), compared to the groups with cellulose or saccharose (6.83-6.91), and a decrease in the ammonia concentration in the caecal digesta, compared to the cellulose control (0.27-0.40 and 0.62 mg/g, respectively). The group with 8% lactulose was characterized by the highest activities of microbiological alpha- and beta-galactosidase and beta-glucosidase in the caecal digesta. Cellulose and both preparations significantly decreased the activity of beta-glucuronidase, compared to the saccharose group (0.39-0.89 and 1.52 U/g, respectively). The highest concentration of VFA in the caecal digesta was observed in the saccharose group (89.2 micromol/g), and the lowest concentration in the group where cellulose was totally substituted by lactulose and inulin (55.1 and 57.5 micromol/g, respectively). The total production of VFA in the caecum was fourfold higher with 8 % lactulose and inulin (254.7 and 236.4 micromol/100g BW, respectively) than in both controls groups (65.1 and 67.8 micromol/100g BW, respectively). The high dose of inulin and lactulose increased the share of propionic acid in the VFA profile (C2:C3:C4) compared to both control groups. When 4% inulin was added to the diet a significant increase of butyrate concentration in the caecum was observed.     

Autochthonous primary production in southern Amazon headwater streams: Novel indicators of altered environmental integrity

The riparian forest reduces the amount of light entering streams, which limits autochthonous primary production. The aim of this study was to evaluate temporal variation of autochthonous primary production in pristine and altered streams, with the goal of identifying indicators of change in environmental integrity in the southern Brazilian Amazon. We evaluated free algal biomass in the water column, the presence of periphyton, and the richness and cover of aquatic herbaceous plants in 20 streams (10 pristine and 10 altered, i.e., with riparian deforestation) during the dry period, at the beginning of the rainy period, and at the end of the rainy period. In altered streams, we recorded the presence of macroscopic periphyton and the amount of algal biomass varied between the dry and flood seasons. Variations in hydrological periods did not contribute to changes in algal biomass in pristine streams; we did not observe the presence of macroscopic periphyton these streams. In altered streams, 23 aquatic herbaceous species were identified, versus only four in the pristine streams. Results showed that riparian deforestation contributes to increased autochthonous primary production, which is also influenced by different hydrological periods, with algae and aquatic herbaceous plants responding differently to dry and rainy periods. The responses of these primary producers confirm their role as important bioindicators of change in the environmental integrity of southern Amazonian streams.     

Epifauna living on sublittoral seaweeds around Cyprus

Macroepifauna living on sublittoral seaweeds were counted,identified and compared among six locations in Cyprus. Mollusks,amphipods, and polychaetes were the major taxa represented. Thedominant alga was Cystoseira barbata which was present at 4of 6 stations and ranged in wet weight from 1–2 kg m^-2 ofbottom. Percent cover of C. barbata ranged from 25 to 90%.Other dominant algae were Laurencia obtusa, Padinapavonica, and Jania rubens. The distribution of epifaunaabundance, species composition, and species richness varied amonglocations. Total epifauna abundance ranged from 1000 to 2250animals m^-2. The abundance of epifauna per gram of plant wassignificantly lower on C. barbata than on the other algalspecies. The total epifauna species richness was more related tolocal algal plant diversity than to plant biomass.     

Physiological effects of lactulose and inulin in the caecum of rats

A model experiment was performed on rats to evaluate the effect of partial or total substitution of saccharose (S) and cellulose (C) by preparations of lactulose and inulin on the development and metabolism of the caecum. In the experimental diets given to rats for 4 weeks, the examined preparations were administered either with an equivalent amount of cellulose (each at 4% of the diet) or as sole source of dietary fibre at 8% of the diet. Compared to the saccharose group cellulose had no effect, and low doses of lactulose and inulin in the diet increased to a medium extent the weight of the caecum wall and caecal digesta. The addition of lactulose and inulin at 8% increased significantly the content of caecal digesta (4.62 and 4.11?g/100g BW, respectively) and the weight of the caecal wall (1.10 and 0.86?g/100g BW, respectively), compared to the groups with saccharose and cellulose (0.73, 0.90 and 0.24, 0.28?g/100g BW, respectively). Cellulose and cellulose partially-substituted with lactulose and inulin caused an increase in the dry matter content of caecal digesta (26.5–27.5%), compared to other groups (21.8–22.8%). The administration of lactulose and inulin preparations was accompanied by a significant drop in pH (5.47–5.81), compared to the groups with cellulose or saccharose (6.83–6.91), and a decrease in the ammonia concentration in the caecal digesta, compared to the cellulose control (0.27–0.40 and 0.62?mg/g, respectively). The group with 8% lactulose was characterized by the highest activities of microbiological α- and β-galactosidase and β-glucosidase in the caecal digesta. Cellulose and both preparations significantly decreased the activity of β-glucuronidase, compared to the saccharose group (0.39–0.89 and 1.52?U/g, respectively). The highest concentration of VFA in the caecal digesta was observed in the saccharose group (89.2?μmol/g), and the lowest concentration in the group where cellulose was totally substituted by lactulose and inulin (55.1 and 57.5?μmol/g, respectively). The total production of VFA in the caecum was fourfold higher with 8 % lactulose and inulin (254.7 and 236.4?μmol/100g BW, respectively) than in both controls groups (65.1 and 67.8?μmol/100g BW, respectively). The high dose of inulin and lactulose increased the share of propionic acid in the VFA profile (C₂:C₃:C₄) compared to both control groups. When 4% inulin was added to the diet a significant increase of butyrate concentration in the caecum was observed.     

Analysis of the biomass quality and photosynthetic efficiency of a nitrogen-fixing cyanobacterium grown outdoors with two agitation systems

The efficiency of two different agitation systems (airlift and paddlewheel) in the biomass photoproduction of a nitrogen-fixing filamentous blue-green alga was evaluated outdoors, and the elemental and molecular composition of the cells grown with each system was analyzed. With the paddlewheel system, the productivity values achieved were over 30% higher than with the airlift system, both in summer and winter. In this last season, a conversion efficiency of total solar energy into stored biomass energy of 3.3% was estimated for the paddlewheel system. Moreover, the algal cells grown with this system exhibited a higher net protein (58.9% of dry weight) and nitrogen (11.3%) content than those grown with the airlift device, with an estimated nitrogen fixation rate of more than 2 g N m(-2) day(-1). These advantages of the paddlewheel system make this procedure more appropriate for the large-scale photoproduction of nitrogen-fixing blue-green algae outdoors.     

Microbial community structure and biomass estimates of a methanogenic Antarctic Lake ecosystem as determined by phospholipid analyses

Phospholipid analyses were performed on water column particulate and sediment samples from Ace Lake, a meromictic lake in the Vestfold Hills, Antarctica, to estimate the viable microbial biomass and community structure in the lake. In the water column, methanogenic bacterial phospholipids were present below 17 m in depth at concentrations which converted to a biomass of between 1 and 7×10⁸ cells/liter. Methanogenic biomass in the sediment ranged from 17.7×10⁹ cells/g dry weight of sediment at the surface to 0.1×10⁹ cells/g dry weight at 2 m in depth. This relatively high methanogenic biomass implies that current microbial degradation of organic carbon in Ace Lake sediments may occur at extremely slow rates. Total microbial biomass increased from 4.4×10⁸ cells/ liter at 2 m in depth to 19.4×10⁸ cells/liter at 23 m, near the bottom of the water column. Total nonarchaebacterial biomass decreased from 4.2 ×10⁹ cells/g dry weight in the surface sediment (1/4 the biomass of methanogens) to 0.06×10⁸ cells/g dry weight at 2 m in depth in the sediment. Phospholipid fatty acid profiles showed that microeukaryotes were the major microbial group present in the oxylimnion of the lake, while bacteria dominated the lower, anoxic zone. Sulfate-reducing bacteria (SRB) comprised 25% of the microbial population at 23 m in depth in the water column particulates and were present in the surface sediment but to a lesser extent. Biomass estimates and community structure of the Ace Lake eco-system are discussed in relation to previously measured metabolic rates for this and other antarctic and temperate ecosystems. This is the first instance, to our knowledge, in which the viable biomass of methanogenic and SRB have been estimated for an antarctic microbial community.     

Dry Weight Production and Partitioning inMedicago minimaandErodium cicutariumUnder Water Stress

Medicago minimaandErodium cicutariumare two naturalized annualspecies in extensive semi-arid, temperate rangelands of centralArgentina. A field study was conducted during 1989 and 1990to evaluate the effects of different levels of soil water availabilityon above- and below-ground dry weight production and partitioningin these species. Dry weight production byM. minimawas moresensitive to water stress than that byE. cicutarium. Althoughthe response was more marked inM. minima, both species allocateda larger proportion of total plant dry weight to fruits underwater stress than under irrigated conditions during early spring.Percentage allocation of total plant dry weight into reproductiveorgans in both species, and stems and peduncles inE. cicutarium,was correlated with total dry weight of these organs; this mightindicate a correlation between sink size and strength. Persistenceof both species in the local flora appears to be associatedwith their capacity to colonize open, degraded areas and growin association with native perennial grasses, as well as totolerate severe drought periods.Copyright 1998 Annals of BotanyCompany Medicago minima(L.) Grufb. var.minima,Erodium cicutarium(L.) L'Herit, water stress, dry matter production and partitioning.     

Estimate of chitin in raw whole insects

Insects contain significant amounts of fiber as measured by crude fiber, acid detergent fiber (ADF) or neutral detergent fiber (NDF). It has always been assumed that the fiber in insects represents chitin based on the structural similarity between cellulose and chitin and the fact that the ADF fraction from insects contains nitrogen. In this study, a number of insect species that are raised commercially as food for insectivores were analyzed for moisture, crude protein (nitrogen × 6.25), fat, ash, NDF, ADF, and amino acids. Additionally, the ADF fraction was analyzed for nitrogen and amino acids to determine if proteins might be present in the ADF fraction. The ADF fraction contained a significant amount of amino acids accounting for 9.3–32.7% of the ADF (by weight). The presence of amino acids in the ADF fraction means that using ADF to estimate insect chitin results in an overestimation of insect chitin content. Using ADF adjusted for its amino acid content, the estimated chitin content of these insect species ranged from 2.7–49.8 mg/kg (as is) and 11.6–137.2 mg/kg (dry matter basis). Additionally, these data suggest that for the species measured here the amount of chitin nitrogen is quite small (as a % of total nitrogen) and that crude protein (nitrogen × 6.25) provides a reasonable estimate of the true protein for most species of insects. Zoo Biol 0:1–11, 2007. © 2007 Wiley‐Liss, Inc.     

PHYTOPLANKTON DYNAMICS IN A STABLY STRATIFIED ANTARCTIC LAKE DURING WINTER DARKNESS

Using automated overwinter sampling devices, we collected preserved phytoplankton samples from multiple depths in Lake Fryxell, a permanently ice-covered lake in southern Victoria Land, Antarctica. Photosynthetic algae were maintained in a stable water column throughout winter darkness. The algal taxa overwintered in different ways in a species-specific manner. Typical vegetative cells were the most abundant form for all species found in the water column. Populations of one chlorophyte, Stichococcus sp., were observed in winter, but the species was absent in both summers. Two cryptophyte species were more abundant in winter than in summer. We interpret the increase in algal population size as evidence of heterotrophic growth in winter and of mixotrophic behavior throughout the year. For two chlorophyte species, some portion of the population had a distinctive morphology, for example, many Chlamydomonas subcaudata Wille. formed akinetes, whereas many Chlorella sp. contained a large amount of starch or other storage material. During winter, vegetative cells of the most abundant cyanobacterial species, Phormidium angustissimum West et West, occurred at the depth of the summer maximum and at depths below the oxycline. The presence of P. angustissimum below the oxycline may have resulted from the accumulation of settling cells at a depth with a greater density. In contrast to the settling of P. angustissimum , the general absence of the other algal species below the oxycline in winter indicated that these species were not influenced by overwinter settling.