GROWTH OF THE FILAMENTOUS GREEN ALGA CTENOCLADUS CIRCINNATUS (CHAETOPHORALES,CHLOROPHYCEAE) IN RELATION TO ENVIRONMENTAL SALINITY1

Clones of the filamentous green alga Ctenocladus circinnatus Borzi were isolated from algae collected at Abert Lake (Oregon) and Mono Lake (California). Stock cultures were exposed to varied salinities of natural lake water to examine the effects on growth rate, cell form, chlorophyll a, and water content. Growth rates were reduced in both clones with increased salinity over the range 25–100 g·L^?1 and were almost completely inhibited at 150 g·L^?1. Chlorophyll a increased between salinities of 25 and 100 g·L^?1, reflecting slower growth, higher proportions of akinetes, and smaller cell sizes as salinity increased. Tissue water content remained essentially constant from 25 to 100 g·L^?1 salinity. Shorter cell dimensions with increased salinity suggest that a lower surface-to-volume ratio may reduce the potential for passive loss of cell water. Prior acclimation of stock cultures to elevated salinity provided no enhancement of growth response at any salinity. The results indicate that environmental salinity can limit the productivity and distribution of Ctenocladus in nature.     

EXPERIMENTAL MESOCOSM STUDIES OF SALINITY EFFECTS ON THE BENTHIC ALGAL COMMUNITY OF A SALINE LAKE

As closed-basin systems, saline lakes are prone to fluctuate in level and salinity with climate change and hydrologic alterations. Loss of many Great Basin lakes has resulted from the diversion of tributary streams for agricultural or municipal uses. At Mono Lake, an alkaline salt lake in eastern California, salinities have risen from 50 to 100 g·L^?1 in just 50 years. Experimental mesocosms were established to simulate some of the potential ecological effects that could have accompanied this change. The influence of salinity on diatom diversity, taxonomic structure, and primary production was tested using mesocosms deployed at Mono Lake. Mesocosm tanks were 500 L in volume, 1 m square, and 0.5 m deep, with open tops covered by 1 mm mesh net. Five treatments (50, 75, 100, 125, and 150 g·L^?1) with four replicates per treatment were used over a 2-month period. The diatom-dominated benthic algae were reduced both in standing crop (from 6 to <0.1 g·m^?2) and diversity (from 30 to 12 taxa) with increased salinity, with most loss occurring in salinities ≥75 g·L^?1. Photosynthetic oxygen production also was significantly lower at salinities ≥75 g·L^?1. Diatom indicator taxa for these shifts included Denticula sp., Nitzschia frustulum, N. monoensis, N. communis, and Stephanodiscus oregonicus increasing in relative abundance in higher salinity treatments, accompanied by decreases in Achnanthes minutissima, Cymbella minuta, N. dissipata, and Rhoicosphenia abbreviata. Exhibiting dominance at moderate salinity levels (75 to 125 g·L^?1) were Nitzschia frustulum, N. communis, N. palea, and Navicula crucialis. These latter species may be limited by both physiological stress at high salinity and grazing and competition at low salinity. The filamentous chlorophyte, Ctenocladus circinnatus, and cyanobacteria (Oscillatoria spp.) occurred only in salinity treatments from 50 to 100 g·L^?1. Diversion of tributary stream flow and resulting salinity increases in this lake threaten sustained benthic primary production and algal species diversity relative to conditions prior to stream diversion. The 1994 decision of the California State Water Resources Control Board to return stream flows to Mono Lake will raise the lake level and reduce salinity to around 75 g·L^?1 and is expected to increase the diversity and productivity of the benthic algae of this ecosystem.     

Distribution and Diversity of Cyanobacteria and Eukaryotic Algae in Qinghai–Tibetan Lakes

Cyanobacteria and eukaryotic algae are important primary producers in a variety of environments, yet their distribution and response to environmental change in saline lakes are poorly understood. In this study, the community structure of cyanobacteria and eukaryotic algae in the water and surface sediments of six lakes and one river on the Qinghai–Tibetan Plateau were investigated with the 23S rRNA gene pyrosequencing approach. Our results showed that salinity was the major factor controlling the algal community composition in these aquatic water bodies and the community structures of water and surface sediment samples grouped according to salinity. In subsaline–mesosaline lakes (salinity: 0.5–50 g L^?1), Cyanobacteria (Cyanobium, Synechococcus) were highly abundant, while in hypersaline lakes (salinity: >50 g L^?1) eukaryotic algae including Chlorophyta (Chlorella, Dunaliella), Bacillariophyta (Fistulifera), Streptophyta (Chara), and Dinophyceae (Kryptoperidinium foliaceum) were the major members of the community. The relative abundance ratio of cyanobacteria to eukaryotic algae was significantly correlated with salinity. The algae detected in Qinghai–Tibetan lakes exhibited a broader salinity range than previously known, which may be a result of a gradual adaptation to the slow evolution of these lakes. In addition, the algal community structure was similar between water and surface sediment of the same lake, suggesting that sediment algal community was derived from water column.     

NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

ADAPTATION OF STYROFOAM SUBSTRATE TO BENTHIC ALGAL PRODUCTIVITY STUDIES IN LAKE TAHOE,CALIFORNIA-NEVADA1

A routine sampling technique has been developed using artificial styrofoam substrate to estimate benthic algal productivity in the littoral zone of lakes. Estimation of maximum carbon fixed in Lake Tahoe ranged from 11.1 mg C·m^?2· day^?1 at 0.5 m to 17.1 mg C·m^?2· day^?1 at 1.0 m. Estimates were made for communities composed of both diatom and green algal populations in water between 0.5 and 3.0 m. Maximum productivity occurred between 1–2 m. The technique developed can give comparable estimates of productivity if adequate replication is undertaken to decrease problems associated with periphytic heterogeneity.     

Seasonal Dynamics of Periphyton in a Large Tropical Lake

Tropical aquatic systems are generally assumed to have little seasonality in productivity patterns. However, this study indicated that there was substantial seasonal variation in epilithic productivity and biomass in tropical Lake Tanganyika, due primarily to seasonal patterns in lake hydrodynamics that influence nutrient availability. Although they support much of the lake’s biological diversity, epilithic algae made a minor contribution to the total energy budget in Lake Tanganyika. A comparison among large, oligotrophic lakes revealed no significant latitudinal trends in periphyton productivity or biomass. However, Lake Tanganyika has relatively low benthic algal biomass and is therefore more efficient at photosynthesis than the temperate lakes. The influence of wave action and consumer density and diversity may be important in moderating productivity of the epilithic community.     

OPTICAL PROPERTIES OF DENSE ALGAL CULTURES OUTDOORS AND THEIR APPLICATION TO REMOTE ESTIMATION OF BIOMASS AND PIGMENT CONCENTRATION IN SPIRULINA PLATENSIS (CYANOBACTERIA)1

Reflectance and vertical attenuation coefficient spectra from 400 to 1100 nm were investigated in detail on dense algal cultures of Spirulina in order to create algorithms for remote estimation of pigment and biomass concentration. Reflectance and the vertical attenuation coefficients were compared with biomass and pigment concentration in outdoor algal cultures. For assessing biomass concentration, the sum of reflectance above the base line from 670 to 950 nm was used. This allows the estimation of biomass with an error of less than 0.06 g·L^?1 For chlorophyll a and phycocyanin estimation, vertical attenuation coefficients at the wavelengths 440 nm (or 676 nm) and 624 nm, respectively, were employed. The developed algorithms were tested by using independent data sets in the range of chlorophyll a from 0.2 to 20mg·L^?1 and biomass from 0.15 to 1.1 g·L^?1. An error of pigment estimation of less than 0.80 mg·L^?1 was achieved. The potential use of the algorithms in algal biotechnology is further discussed.     

Abundance and spatial distribution of Artemia salina in Lake Abert,Oregon

In 1981–1982 Abert Lake had an area of 200 km², with a mean depth of 2.5 m and a total dissolved salt concentration of 82 gl^-1. The spatial distribution and abundance of the brine shrimp, Artemia salina, were monitored for 1981–1982. In 1981 during the midsummer months, with sampling primarily in the eastern side of Abert Lake, brine shrimp populations showed peak densities of 2–4 ind. 1^-1. In 1982, peak abundance occurred in early July and ranged between 5–8 ind. 1^-1. Lakewide estimates of brine shrimp derived from 14 collecting stations and assuming a uniform distribution over the lake resulted in estimates of lakewide abundance of 3 × 10¹¹ adults with a total biomass of 7 × 10⁶ kg. The major algal species in the lake was a benthic filamentous green alga (Ctenocladus circinnatus).     

Effects of light and nutrients on the net accumulation and elemental composition of epilithon in boreal lakes

1. Two experiments in the Experimental Lakes Area (ELA) in north-western Ontario, Canada examined the effects of light and two key elements on the net accumulation and elemental composition of epilithon. In Lake (L) 224, benthic algae were grown under different light intensity and phosphorus supply, while in L302S we provided three levels of two different carbon sources (bicarbonate and glucose) to algae colonizing nutrient-diffusing substrata. After 1 month of accumulation, we sampled biofilms for chlorophyll (chl), carbon (C), phosphorus (P) and algal C.
2. Increased C supply did not significantly affect algal growth (C or chl) or elemental composition (C/P ratios) in L302S. However, P enrichment increased chl and algal C, dramatically reduced the C/P ratio of epilithon, and did not affect total organic C in L224. Phosphorus enrichment also increased the proportion of algal material in the total particulate organic matter and altered the taxonomic composition of algae in L224 biofilms. Shading had no significant effect on the C/P ratio and total organic C in epilithon from the L224 experiment.
3. Our results demonstrate that P supply affects the elemental composition of organic matter that collects on rock substrata. It thus appears that low availability of P relative to C and light drives the formation and retention of high C/P organic matter on rock surfaces in oligotrophic boreal lakes.     

Effects of nitrogen,phosphorus and carbon enrichment on planktonic and periphytic algae in a softwater,oligotrophic lake in Florida,USA

The objective of this study was to investigate nutrient limitation of algal abundance in Anderson-Cue Lake, a softwater clear oligotrophic lake in north-central Florida. Nutrient diffusing clay pots and cylindrical enclosures were used in the field to test effects of different combinations of nitrogen, phosphorus, silica, and carbon on algal standing crop and composition of periphytic and planktonic algae, respectively. Effects of nutrient enrichment on periphytic algae were examined in two studies conducted 31 May – 8 July and 10 June – 15 July 1991. Nutrient effects on planktonic algae were examined in one study from 13 June – 1 July 1991. Planktonic and periphytic algal biovolume was significantly higher (p<0.05) when nitrogen and carbon were added in combination than with treatments without nitrogen, carbon, or nitrogen and carbon. Treatments with nitrogen and carbon combined resulted in lower algal diversity and dominance by coccoid green algae andScenedesmus. Results indicate that carbon and nitrogen can be limiting factors to algal growth in Anderson-Cue Lake and possibly other lakes of similar water quality.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

THE ECOLOGY OF EPIPELIC ALGAE OF FIVE WELSH LAKES,WITH SPECIAL REFERENCE TO VOLVOCALEAN GREEN FLAGELLATES (CHLOROPHYCEAE)1

The ecology of epipelic algae on the marginal sediments of five Welsh lakes was studied over an annual cycle. The lakes, Llydaw, Cwellyn, Padarn, Maelog and Coron ranged from very oligotrophic to nutrient-rich. Attention was focussed on chlamydomonad flagellates, diatoms, blue-green algae and euglenoids and the different proportions of these in algae in the epipelon of lakes of contrasting water quality. A total of 75 algal taxa was found in the five lakes, 25 were species of volvocalean flagellates. Mean annual population density of these flagellates differed by an order of magnitude between the lakes. The greatest population density was recorded for Chlamydomonas anticontata Schiller in nutrient-rich Llyn Maelog. Twenty species of pennate diatoms were recorded frequently in the epipelon. In the nutrient-rich lakes, Maelog and Coron, pennate diatoms were dominant on the sediments, where they exhibited population maxima in spring and autumn. Increase in numbers of epipelic diatoms was recorded when silica concentrations were minimum in the overlying lake waters. Navicula hungarica Grun. achieved the maximum population density, 260 000 cells · cm^?2. Euglenoids formed large epipelic populations during late-summer and autumn in these nutrient-rich lakes. Blue-green algae were more important, proportionally, in the nutrient-poor mountain lakes, which had sediments of higher organic content. Chlamydomonads were the major algal component of the epipelon in the mountain lakes, Llydaw and Cwellyn, where the sediments were characterized by larger particle size, and higher organic content. In the nutrient-rich lakes, where the sediments had higher calcium content, chlamydomonads formed significant populations only during spring and summer, when nutrient levels were minimal in the overlying lakewaters.     

QUALITATIVE AND QUANTITATIVE OBSERVATIONS ON AQUATIC ALGAL COMMUNITIES AND RECOLONIZATION WITHIN THE BLAST ZONE OF MT. ST. HELENS, 1980 AND 19811

Despite the destruction and total rearrangement of much of the area north of Mt St. Helens, many different species of algae became established in the aquatic systems located in the blast zone within 3–4 months after the eruption of May 18, 1980. Initial sites of intense algal activity were found in benthic regions associated with warm springs and in the littoral and phytoplanktonic communities of some small lakes. In the summers of 1980–81, overall phytoplankton numbers and activity were low in the large lakes closest to the crater (e.g. Spirit Lake). However, diatoms, blue-green and green algal isolates from these lakes were obtained in laboratory cultures on a reduced inorganic medium used to enumerate metal and sulfur oxidizing bacteria. Species of Nodularia, Pseudanabaena, Anabaena, Oscillatoria, Nitzschia, Tribonema, Chlamydomonas, Chlorella, and microflagellates (predominantly cryptomonads) were among the more common forms found in preserved samples or isolated in cultures.     

Comparative population ecology of Ephydra hians Say (Diptera: Ephydridae) at Mono Lake (California) and Abert Lake (Oregon)

The population dynamics of Ephydra hians Say final instar larvae and pupae were compared over a two year period in rocky littoral habitats of two alkaline saline lakes in the western Great Basin. Relative abundance increased from 1983 to 1984 at Mono Lake (California), during dilution from ca. 90 to 80 g 1^-1 TDS (total dissolved solids). In contrast, relative abundance decreased over the same period at Abert Lake (Oregon), accompanied by a dilution of salinity from ca. 30 to 20 g l^-1 and a marked increase in the number and abundance of other benthic macroinvertebrate species. These observations are consistent with a hypothesis that proposes biotic interactions limit E. hians abundance at low salinity, and physiological stress limits abundance at high salinity.Oviposition extends from early spring to early fall. Mixed instars present throughout this period indicates multivoltine population dynamics with overlapping generations. The standing stock biomass of final instars increases exponentially in late spring and peaks in late summer or early fall. Pupae increase in proportional representation and abundance from a spring minimum to a fall maximum. The body size of adults and pupae cycle seasonally from a spring maximum to a fall minimum, and may be related to either or both food limitation, or water temperature.     

底栖藻类对消落带土壤中各形态磷的影响

为揭示底栖藻类对三峡库区消落带淹没初期土壤中不同形态磷的影响作用,开展了相关的模拟实验。结果表明,在模拟的底栖藻类影响条件下,消落带土壤中铝磷（Al-P）的含量在实验期间处于波动状态,Al-P相对于其余形态的磷更易于被藻类吸收利用,且与藻类生长存在密切相关,即底栖藻类生长状态较好时,土壤中Al-P呈现下降趋势;而生长状态较差时,土壤中Al-P又出现略微上升的趋势。在实验后期土壤中Al-P含量上升,可能是其余形态的磷转化而来。底栖藻类的生长对不同深度土壤中铁磷（Fe-P）的影响不同,其生长对2-4 cm层土壤的影响较大,而对其余层影响不大。不同深度土壤中闭蓄态磷（O-P）和钙磷（Ca-P）含量差异不明显。实验后期Fe-P、O-P和Ca-P含量一直呈现下降的趋势,只有Al-P在后期出现上升的趋势,说明在淹水条件和底栖藻类生物膜共同作用下,土壤磷的有效性提高,土壤中难溶性磷向易溶性磷转化。       

QUANTITATIVE DIFFERENCES BETWEEN BENTHIC ALGAL COMMUNITIES ALONG A DEPTH GRADIENT IN LAKE MICHIGAN1,2

This study analyzes the impact of conditions associated with depth upon benthic algal communities in Lake Michigan. Diatom abundance was greater at 9.1 and 14.6 m depths than 6.5, 22.6 and 27.4 m. Shallow (6.5 m), mid-depth (9.1 and 14.6 m) and deep (22.6 and 27.4 m) zones were distinguishable on the basis of community composition, structure and abundance. Dominance of benthic species, high diversity and low abundance in shallow communities probably resulted from substantial substrate disturbance by wave action in this productive zone. Dominance of benthic species, high diversity and high abundance characterized mid-depth communities where less wave disturbance enabled algal accumulation. Preponderance of living planktonic taxa, low diversity and low abundance delineated deep communities where planktonic algae accumulated and low light levels reduced growth of benthic species.     

INTERACTIONS BETWEEN LIGHT,NH4+, AND CO2 IN BUOYANCY REGULATION OF ANABAENA FLOS-AQUAE (CYANOPHYCEAE)1

Buoyancy of the gas-vacuolate alga Anabaena flosaquae Brébisson was measured under various levels of light, NH₄⁺, and CO₂. At high irradiance (50 μE · m^?2·^?1) the alga was non-buoyant regardless of the availability of CO₂ and NH₄⁺. At low irradiance (≤10 μE · m ^?2· s^?1) buoyancy was controlled by the availability of NH₄⁺ and CO₂. When NH₄⁺ was abundant, algal buoyancy was high over a wide range of CO₂ concentrations. In the absence of NH₄⁺, algal buoyancy was reduced at high CO₂ concentrations, however as the CO₂ concentration declined below about 5 μmol · L^?1, algal buoyancy increased. These results help explain why gas vacuolate, nitrogen-fixing blue-green algae often form surface blooms in eutrophic lakes.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

SHIFT FROM CHLOROPHYTES TO CYANOBACTERIA IN BENTHIC MACROALGAE ALONG A GRADIENT OF NITRATE DEPLETION1

A survey of the spatial distribution of benthic macroalgae in a fluvial lake of the St. Lawrence River (Lake Saint‐Pierre, Quebec, Canada) revealed a shift in composition from chlorophytes to cyanobacteria along the flow path of nutrient‐rich waters originating from tributaries draining farmlands. The link between this shift and changes in water quality characteristics was investigated by sampling at 10 sites along a 15 km transect. Conductivity, current, light extinction, total phosphorus (TP; >25 μg P · L^?1), and ammonium (8–21 μg N · L^?1) remained fairly constant along the transect in contrast to nitrate concentrations, which fell sharply. Filamentous and colonial chlorophytes [Cladophora sp. and Hydrodictyon reticulatum (L.) Bory] dominated in the first 5 km where nitrate concentrations were >240 μg N · L^?1. A mixed assemblage of chlorophytes and cyanobacteria characterized a 1 km transition zone where nitrate decreased to 40–80 μg N · L^?1. In the last section of the transect, nitrate concentrations dropped below 10 μg N · L^?1, and cyanobacteria (benthic filamentous mats of Lyngbya wollei Farl. ex Gomont and epiphytic colonies of Gloeotrichia) dominated the benthic community. The predominance of nitrogen‐fixing, potentially toxic cyanobacteria likely resulted from excessive nutrient loads and may affect nutrient and trophic dynamics in the river.     

SHORT‐ AND LONG‐TERM EFFECTS OF ELEVATED CO2 ON PHOTOSYNTHESIS AND RESPIRATION IN THE MARINE MACROALGA HIZIKIA FUSIFORMIS (SARGASSACEAE,PHAEOPHYTA) GROWN AT LOW AND HIGH N SUPPLIES1

The short‐term and long‐term effects of elevated CO₂ on photosynthesis and respiration were examined in cultures of the marine brown macroalga Hizikia fusiformis (Harv.) Okamura grown under ambient (375 μL · L^?1) and elevated (700 μL · L^?1) CO₂ concentrations and at low and high N availability. Short‐term exposure to CO₂ enrichment stimulated photosynthesis, and this stimulation was maintained with prolonged growth at elevated CO₂, regardless of the N levels in culture, indicating no down‐regulation of photosynthesis with prolonged growth at elevated CO₂. However, the photosynthetic rate of low‐N‐grown H. fusiformis was more responsive to CO₂ enrichment than that of high‐N‐grown algae. Elevation of CO₂ concentration increased the value of K_1/2(Ci) (the half‐saturation constant) for photosynthesis, whereas high N supply lowered it. Neither short‐term nor long‐term CO₂ enrichment had inhibitory effects on respiration rate, irrespective of the N supply, under which the algae were grown. Under high‐N growth, the Q₁₀ value of respiration was higher in the elevated‐CO₂‐grown algae than the ambient‐CO₂‐grown algae. Either short‐ or long‐term exposure to CO₂ enrichment decreased respiration as a proportion of gross photosynthesis (Pg) in low‐N‐grown H. fusiformis. It was proposed that in a future world of higher atmospheric CO₂ concentration and simultaneous coastal eutrophication, the respiratory carbon flux would be more sensitive to changing temperature.