PHOTOSYNTHESIS-IRRADIANCE PATTERNS IN BENTHIC MICROALGAE: VARIATIONS AS A FUNCTION OF ASSEMBLAGE THICKNESS AND COMMUNITY STRUCTURE

Photosynthesis-irradiance (P-I) characteristics of periphyton (microphytobenthos) have been considered primarily for entire assemblages. How P-I responses vary with mat thickness and with community composition has not been considered in detail. We used a combined approach of modeling, microscale determinations of photosynthetic rate and light attenuation, and whole-assemblage O₂ flux measurements to explore P-I relationships. The modeling approach suggested that the onset of photosynthetic saturation and photoinhibition will occur at higher irradiance and that whole-mat photoinhibition (decreased photosynthesis at very high irradiance), biomass-specific maximum photosynthetic rate, and initial slope of the P-I function (α) should decrease as assemblage thickness increases or light attenuation increases. Spherical light microsensor profiles for a variety of stream algae indicated a strongly compressed photic zone with attenuation coefficients of 70–1791 m^?1 for scalar photosynthetic photon fluence density. The O₂ microelectrode measurements showed little if any photoinhibition at 2 and 4 mm depths in one filamentous green algal (Ulothrix) assemblage, with a relatively low attenuation coefficient, and no photoinhibition in a second Ulothrix community. An assemblage dominated by a unicellular cyanobacterium exhibited little photoinhibition at 2 and 4 mm, and a dense cyanobacterial (Phormidium)/xanthophyte (Vaucheria) community exhibited no photoinhibition at all. The microelectrode data revealed increases in α over several millimeters of depth (photoacclimation). These data supported the model predictions with regard to the effects of mat optical thickness on whole-assemblage values for α and photoinhibition. Whole-community O₂ flux data from 15 intact assemblages revealed positive relationships between chlorophyll a density and maximum photosynthetic rate or α expressed per unit area; the relationships with chlorophyll a were negative when photosynthetic rates were expressed per unit chlorophyll a. None of the whole assemblages exhibited photoinhibition. Thus, the data from the whole communities were consistent with model predictions.     

MODELING THE ANNUAL PRODUCTION OF INTERTIDAL BENTHIC MICROALGAE IN ESTUARINE ECOSYSTEMS1

The purpose of this study was to develop and validate a habitat-specific production simulation model to quantify annual benthic microalgal production in North Inlet estuary, South Carolina. Using hourly measurements of incident irradiance during 1990–1991 as the forcing function, the simulation model was used to obtain hourly estimates of areal benthic microalgal gross primary production in five habitat types. The model, which was validated using actual measurements of production, showed good (r²= 0.63, P < 0.001) agreement between observed and predicted production in the short Spartina alterniflora Loisel zone habitats showed the highest mean hourly production (61.1 mg C m^?2 h^?1) while intertidal mudflats had the maximum hourly rate (166.9 mg C m^?2 h^?1). Daily production was highly variable, primarily due to daily fluctuations in irradiance. Annual estimates of habitat-specific production were multiplied by the mates of habitat-specific production were multiplied by the known area of each habitat type to determine total microalgal production for the estuary (3.423 × 10⁹ g C yr^?1). Short Spartina zone habitats provided 45% of total microalgal annual production, followed by intertidal mudflats (22%), tall Spartina zones (18%), shallow subtidal (13%) and microalgal production exceeds phytoplankton and microalgal production but is less than Spartina production.     

PHOTOSYNTHESIS AND CELL DIVISION BY ANTARCTIC MICROALGAE: COMPARISON OF BENTHIC,PLANKTONIC AND ICE ALGAE1

Irradiance-dependent rates of photosynthesis and cell division of six species of microalgae isolated from the benthos, plankton and sea ice microbial community in McMurdo Sound, Antarctica were compared. Microalgae isolated from different photic environments had distinct photosynthetic and growth characteristics. For benthic and ice algae, photosynthesis saturated at 6 to 20 μE.m^?2.s^?1 and was photoinhibited at 10 to 80 μE.m^?2.s^?1 while for the planktonic algae, saturation irradiances were up to 13 times higher and photoinhibition was not detected. The slope of the light-limited portion of the P-I relationship was up to 50 times greater for the benthic algae than for either the ice or planktonic algae suggesting that benthic algae used the low irradiances more efficiently for carbon uptake. Cell division was dependent on the incubation irradiance for all but one microalga examined. The dependence of division rates on irradiance was however much smaller than for carbon uptake, suggesting that cell division buffers the influence of short term variations of irradiance on cellular metabolism.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

PHOTOSYNTHETIC RESPONSE OF NATURAL ASSEMBLAGES OF MARINE BENTHIC MICROALGAE TO SHORT-AND LONG-TERM VARIATIONS OF INCIDENT IRRADIANCE IN BAFFIN BAY,TEXAS1

This study was designed to understand the high variability characterizing primary production rates of microphytobenthos. The photosynthetic efficiency (α^B) and photosynthetic capacity (P^B_max) of the microphytobenthos were measured at different times of the day on two different dates (8 May and 7 July 1990). In July, unusually low light conditions were caused by the development of a brown tide (chrysophytes). Both light-limited and light-saturated photosynthesis changed at hourly and monthly scales. There was a linear relationship between α^B and P^B_max, suggesting a common response to environmental factors [α^B= 0.0075(±0.00063)·P^B_max+ 0.00097(±0.0071), R²= 0.94]. Incident irradiance at the sediment-water interface was the primary physical factor that explained variability of both α^B (84%) and P^B_max (92%). Temperature had a negative but minor effect that explained an extra 8% and 2% of the variance, respectively. There was no diel rhythm of α^B and P^B_max and incident irradiance was regulated by wind-induced currents. Therefore, microphytobenthos photosynthesis seemed to be primarily controlled by wind events in Baffin Bay.     

Marine species in the algal flora of the Anzali Swamp (Iran)

The composition of marine species in the algal flora of the Anzali Swamp, which is situated parallel to the Caspian Sea coast, is examined. Sampling was carried out at seven stations. A total of 74 species with 89 infraspecific taxa were identified: Cyanophyta, 20 species; Cryptophyta, 2 species; Bacillariophyta, 51 species (66 infraspecific taxa); and Chlorophyta, 1 species.     

PELAGIC AND BENTHIC ENVIRONMENTAL CONTROLS ON THE SPATIAL DISTRIBUTION OF A VIABLE DIATOM PROPAGULE BANK ON THE SWEDISH WEST COAST1

The distribution of viable diatom resting stages in sediments on the Swedish west coast was assessed by the most probable number (MPN) culture technique. Multivariate analyses correlated benthic and pelagic environmental factors to the observed spatial variations in the size and taxonomic composition of the propagule bank. Viable diatom resting stages were plentiful (0.2–4.8 million cells·g ^{? 1} 1 Received 11 September 2001. Accepted 12 June 2002.
dry weight) and were dominated by the genera Skeletonema, Detonula, Chaetoceros, and Thalassiosira. Size of the propagule bank was primarily related to planktonic biomass (measured as chl a) and was highest in the Orust‐Tjörn fjord system. Species composition in this fjord system was dominated by D. confervacea (Cleve) Gran and T. nordenskioeldii Cleve in contrast to stations on the outer coast, which contained more cells of T. minima Gaarder, Asterionellopsis glacialis (Castracane) Round, and Leptocylindrus danicus Cleve. These taxonomic variations were principally influenced by deep water oxygen concentrations and water column stability. Benthic resting cells of S. costatum (Greville) Cleve were abundant all along the coast but showed reduced viability in low oxygen environments. Calculations based on MPN values estimated that resuspension of sediment could provide a sizable inoculum to the plankton, although the development of planktonic blooms will also depend on forces of hydrography and weather. Although benthic resting stages may not be absolutely necessary for survival of all diatoms, these cells may be important in determining species cycles, succession, and the spatial distribution of diatoms.     

SPECTRAL FINGERPRINTING OF ALGAL COMMUNITIES: A NOVEL APPROACH TO BIOFILM ANALYSIS AND BIOMONITORING1

A new technique for spectral fingerprinting of major algal groups in the freshwater periphyton (i.e. cyanobacteria, green algae, and diatoms) was developed using confocal laser scanning microscopy. This technique used the differential spectral emission signatures of photosynthetic algae and allowed their spatially explicit quantification and community three‐dimensional reconstruction. Algal biovolume measurements, carried out with this technique, are superior to existing protocols involving chl and ash‐free dry mass assessments because they are nondestructive, localized, and specific at a group level. This technique can be used to generate depth profiles of the periphytic mat with various applications in aquatic ecology and biofilm analysis.     

Microcystin production by cyanobacteria in the Mwanza Gulf (Lake Victoria,Tanzania)

In order to investigate the potential for microcystin (MC) production by cyanobacteria in the Mwanza Gulf (Lake Victoria, Tanzania), nutrients, phytoplankton and microcystins were sampled inshore (3 m depth) and offshore (18 m depth) from May to August 2002. Significant differences in soluble reactive phosphorus (SRP) and nitrate concentrations between offshore and inshore indicated eutrophication via terrestrial run-off. Though the concentrations of SRP and nitrate ranged between 36–127 and 35–726 μg l ⁻¹ each, the phytoplankton biovolume was generally low. The phytoplankton community was dominated by diatoms (Nitzschia acicularis), a number of cyanobacterial species (Aphanocapsa sp., Anabaena sp., Planktolyngbya spp., Microcystis sp.) and cryptomonads. The water column was completely mixed and Nitzschiapeaked in abundance during July. All cyanobacteria were low in abundance during the entire study period (0.1–1.6 mm ³ l ⁻¹). Microcystins were analysed using high performance liquid chromatography coupled with diode array detection High Performance Liquid Chromatography with Diode Array Detection (HPLC-DAD) and in most samples no microcystins were detected. The highest concentration of [Asp ³]-MC-RR was found in open water at the surface on July 2nd, 2002 (1 μg l ⁻¹). MC concentrations did not pose a potential health risk in the Mwanza Gulf during the study period, however, it is possible that the period of higher cyanobacterial biovolumes has been missed during the sampling period of this study.     

DEVELOPMENT OF IMMUNOASSAYS FOR THE IRON‐REGULATED PROTEINS FERREDOXIN AND FLAVODOXIN IN POLAR MICROALGAE1

While the growth of Southern Ocean phytoplankton is often limited by iron availability, there are no comparable experiments on sea‐ice algae. Here we assess the use of ferredoxin and flavodoxin to investigate the iron nutritional status of sea‐ice algae and describe the development of a quantitative immunoassay for both proteins in marine diatoms. High‐affinity monoclonal antibodies toward both proteins were produced from Cylindrotheca closterium (Ehrenb.) J. M. Lewin et Reimann, and these were used to develop Western blots. Western blots run on whole protein extracts detected both proteins with little cross‐reactivity toward other proteins. The two proteins could be successfully quantitated when applied to gels at between 5 and 50 ng in a volume of 25 μL (0.2–2 μg · mL^?1). Flavodoxin and ferrodoxin expression was examined in the Antarctic diatoms Entomoneis kjellmannii (Cleve) Poulin et Cardinal, Navicula directa (W. Sm.) Ralfs, Fragilariopsis curta (Van Heurck) Hust., Pseudo‐nitzschia sp., Porosira glacialis (Grunow) E. G. Jørg., Fragilariopsis cylindrus (Grunow) Willi Krieg., Fragilariopsis sublinearis (Van Heurck) Heiden et Kolbe, C. closterium, Nitzschia lecointei Van Heurck, and the dinoflagellate Polarella glacialis Montresor, Procaccini et Stoecker. Two Arctic isolates were also examined, Nitzschia frigida (Grunow) and Fragilariopsis oceanica (Cleve) Hasle. Significant heterogeneity of protein expression was observed despite all cultures being grown in iron‐replete f/2 medium. Only one species, F. cylindrus, displayed the expected expression of ferredoxin only in iron‐replete medium. Four were observed to produce both proteins under iron‐replete conditions. Ferredoxin was not detected at all in F. curta and Pseudo‐nitzschia sp., but distinct flavodoxin bands were observed in both of these organisms. All species examined were observed to express either flavodoxin or ferredoxin or both of the proteins as determined by Western immunoblotting.     

STRUCTURE AND FUNCTION OF BENTHIC ALGAL COMMUNITIES IN AN EXTREMELY ACID RIVER1

The composition of algal species and pigments and the structural and functional characteristics of the algal community were investigated in an acid stream of southwestern Spain, the Río Tinto. The algal community had low diversity and showed few seasonal differences. It was mainly made up of Klebsormidium flaccidum Kütz. (Silva, Mattox & Blackwell) that produced long greenish or purplish filaments, Pinnularia acoricola Hust. (producing brown patches) and Euglena mutabilis Schmitz. The algal filaments made up a consistent biofilm that also included fungal hyphae, iron bacterial sheaths, diatoms, and mineral particles. HPLC analyses on Río Tinto samples showed that undegraded chl accounted for 67% of the total chl in the filamentous patches but were a minority in the brown patch (2.6%). The brown patch had a concentration of carotenoids eight times lower than that observed in the green patch. When chl concentrations were weighted for the proportion of the different patches on the streambed, undegraded chl a accounted for 89.2 mg chl a·m ^{? 2} of stream surface area (5.4 g C·m ^{? 2}). This high algal biomass was supported by relatively high nutrient concentrations and by a high phosphatase activity (V_max = 137.7 nmol methylumbelliferyl substrate·cm ^{? 2}·h ^{? 1} 1 Received 15 July 2002. Accepted 17 February 2003. , K_m = 0.0045 μM). The remarkable algal biomass in Río Tinto potentially contributed to the bacterial–fungal community and to the macroinvertebrate community and emphasizes the role that the algae may have in the organic matter cycling and energy flow in extreme systems dominated by heterotrophic microorganisms.     

FURTHER EVIDENCE FOR ANOMALOUS SIZE SCALING OF RESPIRATION IN PHYTOPLANKTON1

Respiration per unit mass decreases as organism size increases among metazoans and heterotrophic unicells. The rate of decrease is described by a power function of organism mass; the exponent of the power function is 0.75 (Three-fourths Rule). Previously unanalyzed respiration rates for 11 species of phytoplankton ranging in size over four orders of magnitude show a size-scaling exponent of 1.13 (SE, ±0.15), which is statistically different from 0.75. This result confirms the result of an earlier study of eight phytoplankton species indicating that size scaling of respiration is absent or minimal in phytoplankton, in contrast to the pattern of heterotrophic unicells. The size-related range of respiration rates per unit mass across the full size spectrum of phytoplankton would be approximately 18–fold if respiration were scaled according to the Three-fourths Rule. If respiration does not scale with size or scales minimally with size, as suggested by present evidence, the size-related range of rates will be much smaller or negligible. The apparent anomaly of size scaling for phytoplankton respiration is potentially of great ecological and adaptive significance in unicellular algae.     

MICROALGAE AND CYANOBACTERIA: FOOD FOR THOUGHT1

In non‐Western civilizations, cyanobacteria have been part of the human diet for centuries. Today, microalgae and cyanobacteria are either produced in controlled cultivation processes or harvested from the natural habitats and marketed as food supplements around the world. Cyanobacteria produce a vast array of different biologically active compounds, some of which are expected to be used in drug development. The fact that some of the active components from cyanobacteria potentially have anticancer, antimicrobial, antiviral, anti‐inflammatory, and other effects is being used for marketing purposes. However, introduction of these products in the form of whole biomass for alimentary purposes raises concerns regarding the potential toxicity and long‐term effects on human health. Here, we review data on the use of cyanobacteria and microalgae in human nutrition and searched for available information on legislature that regulates the use of these products. We have found that, although the quality control of these products is most often self‐regulated by the manufacturers, different governmental agencies are introducing strict regulations for placing novel products, such as algae and cyanobacteria, on the market. The existing regulations require these products to be tested for the presence of toxins, such as microcystin; however, other, sometimes novel, toxins remain undetected, and their long‐term effects on human health remain unknown.     

LIGHT‐INDUCED MOTILE RESPONSES OF THE ESTUARINE BENTHIC DIATOMS NAVICULA PERMINUTA AND CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Motility of estuarine epipelic (mud‐inhabiting) diatoms is an important adaptation to living in biofilms present within fine sediments. Motility allows cells to migrate within the photic zone in response to a wide range of environmental stimuli. The motile responses of two species of benthic diatoms to photon fluence rates and spectral quality were investigated. Cultures of Navicula perminuta (Grunow) in van Heurck and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reimann both exhibited photoaccumulation at ～200 μmol · m^?2 · s^?1 and photodispersal from photon flux densities (PFDs) of ～15 μmol · m^?2 · s^?1. Photokinesis (changing cell speed) contributed toward photodispersal for both species, and red light (λ = 681–691 nm) was most effective at inducing this process. N. perminuta showed a phototactic (directional) response, with active movement in response to a light gradient. Although this response was exhibited in white light, these directional responses were only elicited by wavelengths from 430 to 510 nm. In contrast, C. closterium did not exhibit phototaxis under any light conditions used in this study. Motile benthic diatoms thus exhibit complex and sophisticated responses to light quantity and quality, involving combinations of photokinesis and phototaxis, which can contribute toward explaining the patterns of large‐scale cell movements observed in natural estuarine biofilms.     

DNA:ATP RATIOS IN MARINE MICROALGAE AND BACTERIA: IMPLICATIONS FOR GROWTH RATE ESTIMATES BASED ON RATES OF DNA SYNTHESIS1

DNA: ATP and carbon: DNA (C:DNA) ratios were measured in a total of 14 species of marine microalgae and bacteria. Comparison of several DNA assay methods with results obtained with cultures uniformly labeled with ³³P indicated that by far the most accurate results were obtained using diaminobenzoic acid (DABA) or diphen-ylamine, with DABA having the highest precision. Both the Hoechst and DAPI methods seriously underestimated DNA concentrations in algal cultures. Average DNA: ATP ratios in the algal and bacterial cultures were I7 and 34 by weight, respectively, with almost all values lying in the range of 10–40. DNA: ATP ratios in the microalgae showed no correlation with growth conditions but varied by about a factor of 3 among species. C:DNA ratios for individual species of microalgae and bacteria ranged from 21 to 155 by weight and averaged 50 for the microalgae and bacteria taken together. Growth rates of microalgal species grown in cyclostats were estimated to within 8% of dilution rates when calculated from the uptake of ³H-adenine and the DNA: ATP ratio of the species. Use of the ³H-adenine method for estimating microalgal growth rates in the field may thus be a useful tool for investigating the physiology of microalgae in nature.     

MICROPHOTOMETRIC ASSESSMENT OF SPECTRAL ABSORPTION AND ITS POTENTIAL APPLICATION FOR CHARACTERIZATION OF HARMFUL ALGAL SPECIES

A comparison was made of microphotometric measurements and spectrophotometric measurements of particulate spectral absorption of four algal species, including the chlorophyte Dunaliella tertiolecta Butcher; a nontoxic dinoflagellate, Amphidinium carterae Hulburt; a diatom, Chaetoceros gracilis Schütt; and a toxic dinoflagellate, Gymnodinium breve Steidinger. Particulate spectral absorption of monospecies cultures was estimated as the product of the average absorption efficiency factor, Q _a, determined by microphotometry, the cellular cross-sectional area, and the cell number density. Estimates of particulate spectral absorption from microphotometric measurements were, in most cases, within one standard deviation of values determined from spectrophotometric measurements of algal suspensions. Estimates of Q_a(675) were shown to be consistent with values reported in previous studies for cells of similar size and pigmentation and were consistent with theoretical predictions. Absorption spectra of mixtures of C. gracilis and G. breve were numerically decomposed into contributions by absorption signatures of monospecies cultures using either spectrophotometric or microphotometric measurements as the basis for end members. Modeled contributions assigned to either species displayed trends consistent with the actual proportions contributed to the spectrum by each algal culture. However, the technique was sensitive to measurement variability, which reduced the level of agreement between modeled and actual contributions. The utility of this approach for identification of algal taxa will depend on the degree to which algal spectral absorption signatures differ and the capabilities for acquiring high-resolution data with low signal-to-noise ratios.     

热带淡水产油微藻的分离筛选与鉴定

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WATER COLUMN PRODUCTIVITY ATTRIBUTABLE TO DISPLACED BENTHIC DIATOMS IN WELL-MIXED SHALLOW ESTUARIES1

We attempted to determine the extent to which benthic diatoms contribute to water column primary productivity in shallow-water estuaries and to elucidate the primary mechanisms responsible for suspending the diatoms. A perliminary study conducted in Mugu Lagoon, California indicated that productivity of ocean water entering the lagoon during flood tides was often several orders of magnitude less than that of the same water mass about 3 h later. Benthic pennate diatoms displaced from the sediments into the water column accounted for the increase. A more detailed study was conducted in Barataria Estuary, Louisiana where, for one month, daily measurements were made of benthic and water column productivity and several other environmental variables. During the month, the relationship between water column and benthic primary productivity varied from strongly negative to weakly negative to positive. K-systems analysis indicated that factors comprised of wave height, meteorological tides, astronomical tides, and benthic productivity and standing crop accounted for the full range of variation in water column productivity. Benthic pennate diatoms, represented an average of 74% of the diatom taxa in water column samples. We conclude that the primary productivity of well-mixed shallow estuarine waters is often greatly aumented by displaced benthic algae.     

FOURIER TRANSFORM INFRARED SPECTROSCOPY OF MICROALGAE AS A NOVEL TOOL FOR BIODIVERSITY STUDIES,SPECIES IDENTIFICATION,AND THE ASSESSMENT OF WATER QUALITY1

Fourier transform infrared (FTIR) spectrometry was used to study the spectral features of 12 eukaryotic and two prokaryotic species of microalgae. The algae were cultured in liquid media containing either NO₃^? or NH₄⁺ as the sole N‐source; for the NH₄⁺ treatment, the algae were subjected to short‐term (24 h) or long‐term (1 month) incubations; for the hypersaline species, cells were also grown in the presence of 2 M NaCl. Over 500 spectra, acquired from at least three distinct cultures for each species, in each growth regime, were subjected to hierarchical cluster analysis (HCA) and were successfully separated according to their taxonomy, showing that the overall spectra were characteristic of each species and that this technique could be fruitfully employed to separate microalgal species living in a similar condition (as would be the case for a natural assemblage). In addition, in most cases, it was possible to differentiate between algae subjected to different growth treatments although belonging to the same species. We also demonstrated that it is possible to accurately identify species and determine the nutritional status of their environment of origin (e.g., N‐source), provided that suitable FTIR spectral libraries are available. This study aims to provide the basis for the development of rapid, easy, and inexpensive methods for the evaluation of biodiversity in natural phytoplankton samples and to monitor the water quality of natural environments.