Freshwater population of Ulva flexuosa (Ulvaceae,Chlorophyta) as a food source for great pond snail: Lymnaea stagnalis (Mollusca,Lymnaeidae)

Cosmopolitan species of the genus Ulva (Ulvaceae; Chlorophyta) that populate the littoral zone of marine habitats constitute a staple diet for a variety of organisms, particularly snails, shellfish, polychaetes, and birds. Occurrence of Ulva species (e.g., U. flexuosa and U . prolifera) has also been observed in freshwater inland ecosystems that have no contact with saline water. However, the influence of the development of macroalgal mats of Ulva on indigenous organisms in limnic ecosystems has not been established. This study investigates the trophic relationships between Ulva flexuosa and one species of snail from freshwater habitats in central Europe. During the summer, the great pond snail (Lymnaea stagnalis) consumed Ulva as a source of nutrition even when other algae and plants were available. Lymnaea stagnalis consumed an average of 100 mg of Ulva thalli per day. This level of biomass exceeded the consumption of an alternative food source, the shoots of Elodea canadensis. Ulva thalli are more actively consumed by great pond snails than Elodea shoots, and this is expressed in terms of the differences of biomass consumption. It was also observed that the interior of the monostromatic tubular thalli of Ulva flexuosa serves as a protective shelter for juvenile great pond snails.     

BIOMASS AND DYNAMICS OF GROWTH OF ULVA SPECIES IN PALMONES RIVER ESTUARY1

During the last decade, the Palmones River estuary has undergone severe eutrophication followed by a green tide episode; two species of Ulva, rotundata Blid. and Ulva curvata (Kütz.) De Toni, were the main macroalgae responsible for this bloom. From November 1993 to December 1994, we followed the biomass, the growth dynamics, and tissue elemental composition (C:N:P)of Ulva species, as well as some physicochemical variables in the estuary. Maximum biomass (up to 375 g dry wt·m^?2 in some spots, corresponding to a thallus area index of nearly 17 m²Ulva·m^?2 sediment) were observed in June and December. However, the biomass varied among the sampling stations. Water nitrate, ammonia, and phosphate showed high concentrations throughout the year, with extremely high transient pulses, sustaining the high growth rates observed. Growth rates were estimated directly in the field. The rates were generally higher in Ulva discs maintained in net cages than those estimated by changes in biomass standing stock between two consecutive samplings. The difference between both estimates was used to quantify the importance of the processes causing loss of biomass, which were attributable to grazing, exported biomass, and thallus decomposition under anaerobic conditions resulting from extreme self-shading. Maximum chlorophyll content was found in winter, whereas the minimum was in spring. Atomic N:P ratios were generally higher in the algae than in the water. However, the absolute concentrations of tissue N and P were always higher than the critical levels for maximum growth, which suggests that growth was not limited by inorganic N or P availability. The results suggested that the increase in nutrient loading in the river may have triggered the massive development of green algae and that light limitation and temperature stress in summer seem to be the main factors controlling the abundance of Ulva in the estuary. In addition to light availability and thermal stress, the different loss processes may have a decisive role in the dynamics of Ulva biomass.     

Abiotic factors affecting the development of <Emphasis Type="Italic">Ulva</Emphasis> sp. (Ulvophyceae; Chlorophyta) in freshwater ecosystems

The influence of physicochemical factors on the development of Ulva species with distromatic tubular morphology was studied in three streams located in Poznan, Poland. The study evaluated key environmental factors that may influence the colonisation and growth of Ulva populations in freshwater systems. In total, nine environmental parameters were included: temperature, water depth, pH, oxygen (O₂), ammonium (NH₄ ⁺), nitrate (NO₃ ⁻), phosphate (PO₄ ³⁻), sodium chloride (NaCl) and total iron (Fe). Morphometric features of thalli (length and width, percentage of furcated and young thalli) and surface area of free-floating mats formed by the freshwater populations of Ulva were compared at all sites. Principal components analysis indicated the most important factors influencing Ulva development were sodium chloride concentrations and water depth. Two other key chemical factors affecting the freshwater form of Ulva were phosphate and nitrite concentrations. High concentrations of sodium chloride inhibited the development of Ulva, leading to a lower number of thalli in the Ulva mats. At the sites with stable and deeper water, the surface area of the mats was larger. Both phosphate and nitrite concentrations were positively correlated with an increase in the number of thalli in the mats and the thalli length.     

Canopy structure,photosynthetic capacity and nitrogen distribution in adjacent mixed and monospecific stands of <Emphasis Type="Italic">Phragmites australis</Emphasis> and <Emphasis Type="Italic">Typha latifolia</Emphasis>

Shifts in canopy structure associated with nonnative plant invasions may interact with species-specific patterns of canopy resource allocation to reinforce the invasion process. We documented differences in canopy light availability and canopy resource allocation in adjacent monospecific and mixed stands of Phragmites australis and Typha spp. in a Great Lakes coastal wetland presently undergoing Phragmites invasion to better understand how light availability influences leaf nitrogen content (N_mass) and photosynthetic capacity (A_max) in these species. Due to their horizontally oriented leaves, light attenuates more rapidly in monospecific stands of Phragmites than in monospecific stands of Typha, where leaves are more vertically-oriented. Whereas Typha canopies followed our prediction that patterns of N_mass and A_max should closely parallel patterns of canopy light availability, N_mass and A_max were consistent throughout Phragmites’ canopies. Moreover, we observed overall greater N_mass and lower photosynthetic nitrogen use efficiency in leaves of Phragmites than in leaves of Typha. Improved understanding of the link between N_mass and A_max in these canopies should improve our understanding of carbon and nitrogen cycling consequences of Phragmites invasion in wetland ecosystems.     

Photosynthetic response of Ulva rotundata to light and temperature during emersion on an intertidal sand flat

Summary We have investigated the diurnal response of photosynthesis and variable photosystem II (PSII) chlorophyll fluorescence at 77 K for thalli of the chlorophyte macroalga, Ulva rotundata, grown in outdoor culture and transplanted to an intertidal sand flat in different seasons. The physiological response in summer indicated synergistic effects of high PFD and aerial exposure, the latter probably attributable to temperature, which usually increased by 8 to 10° C during midday emersion. Except at extreme emersed temperatures in summer (38° C), the light-saturated photosynthesis rate (P_m) did not decline at midday. In contrast, light-limited quantum yield of photosynthetic O₂ exchange () and the ratio of variable to maximum fluorescence yield (F_v/F_m) reversibly declined during midday low tides in all seasons. Shade-grown thalli exhibited a fluorescence response suggestive of greater photodamage to PSII, whereas sun-grown thalli had greater photoprotective capacity. The fluorescence decline was smaller when high tide occurred at midday, and was delayed during morning cloudiness. These results suggest that the diurnal response to PFD in this shallow water species is modified by tidal and meteorological factors. U. rotundata has a great capacity for photoprotection which allows it to tolerate and even thrive in the harsh intertidal environment.Abbreviations F_o instantaneous yield of chlorophyll fluorescence - F_m maximum yield of fluorescence - F_v variable yield (F_m–F_o) of fluorescence - PFD photon flux density (400–700 nm) - P_m light-saturated rate of photosynthesis - PSH photosystem II - Q_A electron acceptor of PSII - light-limited quantum yield of photosynthesis     

Photosynthetic acclimation to different light levels in the brown marine macroalga, <Emphasis Type="Italic">Hizikia fusiformis</Emphasis> (Sargassaceae,Phaeophyta)

Hizikia fusiformis thalli experience dynamic incident light conditions during the period of growth. The present study was designed to examine how changing photon irradiance affects the photosynthesis both in the short and long terms by culturing H. fusiformis under three different light levels: 35 μmol photons m^-2 s^-1 (low light, LL), 85 μmol photons m^-2 s^-1 (intermediate light, IL), and 165 μmol photons m^-2 s^-1 (high light, HL). A similar relative growth rate was observed between IL- and HL-grown algae, but the growth rate was significantly reduced in LL-grown algae. The photosynthetic rates (P _n) measured at their respective growth light levels were found to be lowest in the thalli grown at LL and highest at HL. However, LL-grown algae exhibited much higher P _n in comparison with IL- and the HL-grown thalli at the same measuring photosynthetic photon flux density, indicating the photosynthetic acclimation to low growth light in H. fusiformis. The photosynthesis–light curves showed that LL-grown algae had a highest light-saturating maximum P _n (P _max) in comparison with IL- or HL-grown algae when the photosynthetic rates were expressed on the biomass basis. However, P _max was highest in HL-grown algae compared to IL- or LL-grown algae when the rates were normalized to chlorophyll a. The photosynthesis–inorganic carbon (Ci) response curves were also significantly affected by the growth light conditions. The highest value of apparent photosynthetic conductance occurred in LL-grown algae while the lowest value in HL-grown algae. Additionally, the activity of external carbonic anhydrase (CA) tended to increase while the total CA activity inclined to decrease in H. fusiformis thalli when the growth light level altered from 35 to 165 μmol photons per square meter per second. The external CA inhibitors showed a higher inhibition in HL-grown algae compared with LL-grown algae. It was proposed that photosynthetic acclimation to low light condition in H. fusiformis was achieved through an increase in the number of reaction centers and increased capacities of electron transport and of Ci transport within cells. The ability of photosynthetic acclimation to low light confers H. fusiformis thalli to overcome the environmental low light condition as a result of the attenuation of seawater or self-shading through enhancing its photosynthetic performance and carbon assimilation necessary for growth.     

A COMPARISON OF PHOTOSYNTHETIC ELECTRON TRANSPORT RATES IN MACROALGAE MEASURED BY PULSE AMPLITUDE MODULATED CHLOROPHYLL FLUOROMETRY AND MASS SPECTROMETRY

The relationship between whole chain photosynthetic electron transport and PSII activity was investigated in Porphyra columbina (Montagne) (Rhodophyta), Ulva australis (Areschoug) (Chlorophyta), and Zonaria crenata ( J. Agardh) (Phaeophyta). Mass spectrometric measurements of gross O₂ evolution and gross O₂ uptake were combined with simultaneous measurement of pulse-modulated chl fluorescence under a range of irradiances and inorganic carbon (C_i) concentrations. At light-limiting irradiance, a good correlation between gross O₂ evolution and the electron transport rate (ETR) calculated from chl fluorescence ((F_m′− F_s)/F_m′) was found in the optically thin species (Ulva and Porphyra). The calculated ETR was equivalent to the theoretical electron requirement in these species but overestimated gross O₂ evolution in the thicker species Zonaria. In saturating light, especially when C_i availability was low, ETR overestimated gross O₂ evolution in all species. Excess electron flow could not be accounted for by an increase in gross O₂ uptake; thus neither Mehler-ascorbate-peroxidase reaction nor the photosynthetic carbon oxidation cycle were enhanced at high irradiance or low C _i. Alternative explanations for the loss of correlation include cyclic electron flow around PSII that may be engaged under these conditions or nonphotochemical energy quenching within PSII centers. The loss of correlation between ETR and linear photosynthetic electron flow as irradiance increased from limiting to saturating or at low C_i availability and in the case of optically thick thalli limits the application of this technique for measuring photosynthesis in macroalgae.     

Effects of solar UV-B radiation on canopy structure of Ulva communities from southern Spain

Within the sheltered creeks of Cádiz bay, Ulva thalli form extended mat-like canopies. The effect of solar ultraviolet radiation on photosynthetic activity, the composition of photosynthetic and xanthophyll cycle pigments, and the amount of RubisCO, chaperonin 60 (CPN 60), and the induction of DNA damage in Ulva aff. rotundata Bliding from southern Spain was assessed in the field. Samples collected from the natural community were covered by screening filters, generating different radiation conditions. During daily cycles, individual thalli showed photoinhibitory effects of the natural solar radiation. This inhibition was even more pronounced in samples only exposed to photosynthetically active radiation (PAR). Strongly increased heat dissipation in these samples indicated the activity of regulatory mechanisms involved in dynamic photoinhibition. Adverse effects of UV-B radiation on photosynthesis were only observed in combination with high levels of PAR, indicating the synergistic effects of the two wavelength ranges. In samples exposed either to PAR+UV-A or to UV-B+UV-A without PAR, no inhibition of photosynthetic quantum yield was found in the course of the day. At the natural site, the top layer of the mat-like canopies is generally completely bleached. Artificially designed Ulva canopies exhibited fast bleaching of the top layer under the natural solar radiation conditions, while this was not observed in canopies either shielded from UV or from PAR. The bleached first layer of the canopies acts as a selective UV-B filter, and thus prevents subcanopy thalli from exposure to harmful radiation. This was confirmed by the differences in photosynthetic activity, pigment composition, and the concentration of RubisCO in thalli with different positions within the canopy. In addition, the induction of the stress protein CPN 60 under UV exposure and the low accumulation of DNA damage indicate the presence of physiological protection mechanisms against harmful UV-B. A mechanism of UV-B-induced inhibition of photosynthesis under field conditions is proposed.     

Influence of sun irradiance and water availability on lichen photosynthetic pigments during a Mediterranean summer

This study was carried out to investigate changes in lichen photosynthetic pigments induced by different combinations of light irradiance and water availability during a Mediterranean summer. To this purpose, thalli of three epiphytic lichens with a markedly different ecology concerning photo-hygrophytism, namely Evernia prunastri (hygro-mesophytic), Flavoparmelia caperata (mesophytic) and Xanthoria parietina (xero-mesophytic), were transplanted for 30 days to N- and S-facing cardinal exposures in central Italy. To investigate the effect of thallus hydration, at each cardinal exposure 50% of thalli were hydrated daily with deionised water. The results showed that changes in the concentrations of photosynthetic pigments are species-specific and consist in a general depression of photosynthetic pigments only in the hygro-mesophytic species E. prunastri. The pattern of photosynthetic pigments was also investigated in spontaneous samples along contrasting aspects. In this case, X. parietina from S-facing slopes, adapted to direct solar influx, showed higher pigment content than N-facing thalli; F. caperata and E. prunastri avoid direct extreme solar radiation and assimilation pigments were influenced more by the shadowing of tree canopies than by the cardinal exposure. The influence of drought on lichen photosynthetic pigments in the Mediterranean area is discussed.     

Nutrient and iron limitation to <Emphasis Type="Italic">Ulva</Emphasis> blooms in a eutrophic coastal lagoon (Sacca di Goro,Italy)

Growth patterns and bloom formation of the green seaweed Ulva rigida were analysed in the eutrophic Sacca di Goro lagoon (Po River Delta, Italy). Variations of standing biomasses and elemental composition of Ulva were analysed through an annual cycle with respect to nitrogen, phosphorus and iron. Growth rates, nutrient and iron uptake and nitrate storage by macroalgal thalli were also assessed with field experiments during the formation of a spring bloom. The control of Ulva growth and the bloom formation depended on multiple factors, especially on nitrogen availability and iron deficiency. In the nitrate rich waters of the Sacca di Goro lagoon, nitrate accumulation in Ulva thalli was inversely related with Fe uptake, indicating an influence of Fe limitation on N acquisition. Since length and magnitude of nitrate luxury uptake are inversely related to the size of the intracellular nitrate pools, in nitrate rich waters the fast growing Ulva may face risk of N-limitation not only when exposed to low N concentrations or at high biomass levels, but also when exposed to pulsed dissolved nitrate concentrations at low iron availability. The potential Fe limitation could be affected by processes controlled by geochemical reactions and by macroalgal growth and decomposition. Both Fe oxidation during the active macroalgal growth and the formation of insoluble FeS and FeS₂ during bloom collapse can result in a drastic decrease of soluble iron. Thus, a potential limitation of Fe to macroalgae can occur, determining positive feedbacks and potentially controlling the extent of bloom development and persistence.     

SEASONAL VARIABILITY OF PHYSICOCHEMICAL AND RHEOLOGICAL PROPERTIES OF ULVAN IN TWO ULVA SPECIES (CHLOROPHYTA) FROM THE BRITTANY COAST1

The seasonal variability in the extraction yield, physicochemical characteristics, and rheological properties of ulvan from two Ulva species contributing to Brittany “green tides” has been studied. These seaweeds were collected in the water column for Ulva armoricana Dion, de Reviers et Coat and on hard substrata for Ulva rotundata Bliding. The maximum ulvan extraction efficiency was not related to the maximum ulvan content in the seaweeds, but with the active growth period of the seaweeds. Ulvan chemical structure, macromolecular characteristics, and rheological properties were affected by both species and seasons. The proportion of high‐molecular‐weight ulvan was the major factor positively correlated with the gelling properties. Characteristics of ulvan from U. rotundata subjected to tides were more affected by seasons than ulvan from U. armoricana living in a more constant environment. These results point to several useful recommendations concerning Ulva sp. biomass collected with regard to ulvan characteristics and uses.     

Photosynthetic activity and proteomic analysis highlights the utilization of atmospheric CO2 by Ulva prolifera (Chlorophyta) for rapid growth

Free‐floating Ulva prolifera is one of the causative species of green tides. When green tides occur, massive mats of floating U. prolifera thalli accumulate rapidly in surface waters with daily growth rates as high as 56%. The upper thalli of the mats experience environmental changes such as the change in carbon source, high salinity, and desiccation. In this study, the photosynthetic performances of PSI and PSII in U. prolifera thalli exposed to different atmospheric carbon dioxide (CO₂) levels were measured. Changes in photosynthesis within salinity treatments and dehydration under different CO₂ concentrations were also analyzed. The results showed that PSII activity was enhanced as CO₂ increased, suggesting that CO₂ assimilation was enhanced and U. prolifera thalli can utilize CO₂ in the atmosphere directly, even when under moderate stress. In addition, changes in the proteome of U. prolifera in response to salt stress were investigated. Stress‐tolerance proteins appeared to have an important role in the response to salinity stress, whereas the abundance of proteins related to metabolism showed no significant change under low salinity treatments. These findings may be one of the main reasons for the extremely high growth rate of free‐floating U. prolifera when green tides occur.     

INORGANIC CARBON LIMITATION OF PHOTOSYNTHESIS IN ULVA ROTUNDATA (CHLOROPHYTA)1

A computerized oxygen electrode Astern was used to make rapid and accurate measurements of photosynthetic light and dissolved inorganic carbon (DIC) response cures with a macroalga. Ulva rotundata Blid. was grown in an outdoor, continuous flow system in seawater under sunlight or 9% of sunlight at Beaufort, North Carolina. The light compensation points in the shade- and sun-grown plants, measured in seawater, were at photon flux densities (PFDs) of 16 and 27 μmol. Photons·m^?2·s^?1, respectively but the quantum yield of O₂ evolution was not significantly different. Rates of photosynthesis in seawater per unit area of thallus under saturating light and rates of dark respiration were about 1.5-fold higher in sun- than in shade-grown plants. The concentration of DIC in seawater (approximately 2 mM) limited photosynthesis at absorbed PFDs above 60–70 μmol photons·m^?2·s^?1 Addition of 20 mM inorganic carbon had no effect on quantum yield but caused about a 1.5-fold increase in the light-saturated photosynthetic rate in both shade- and sun-grown Ulva. The effect of DIC supplementation was greatest in plants grown in October and least in plants grown in June. The light- and DIC-saturated rate of photosynthesis in seawater was similar to the maximum rate obtained by exposing Ulva to 10% CO₂, in the gas phase. The carbon isotope values (δ¹³C, reflecting the ¹³C/¹²C ratio compared to a standard) of Ulva grown in the same seawater supply were dependent on light and agitation. Samples from Beaufort Inlet were more negative (δ¹³C value, ?20.03‰) than those grown in bright light with agitation (δ¹³C value, ?17.78‰ outdoors; ?17.23‰ indoors), which may indicate DIC supply limited carbon uptake in seawater.     

SUMMER DECLINE OF ULVA LACTUCA (CHLOROPHYTA) IN A EUTROPHIC EMBAYMENT: INTERACTIVE EFFECTS OF TEMPERATURE AND NITROGEN AVAILABILITY?1

Throughout the summer, abundance of Ulva lactuca L. declined while biomass of Cladophora vagabunda (L.) van den Hoek and Gracilaria tikvahiae McLachlan increased in a New England embayment undergoing eu-trophication (Waquoit Bay, Massachusetts). We investigated the physiological basis for the summer dieback, focusing on temporal variations in photosynthetic performance and tissue nitrogen (N). We also compared photosynthetic and N uptake capabilities of U. lactuca with other abundant species in this eutrophic system. Photosynthetic egiciency and capacity of U. lactuca declined markedly at 25°C, compared with a spring (15°C) peak in photosynthetic performance; P_max was 4.6 ± 0.3 and 1.8 ± 0.6 μmol O₂.m^?2.s^?1 during spring and summer, respectively. Notably, summer p_max of other abundant species of the embayment was 1.5–3 × higher than that measured for U. lactuca. Ulva lactuca showed a signifciant photosynthetic response to dissolved inorganic carbon enrichment during summer, when water-column-dissolved CO₂ levels were 20% of spring values. Although ammonium uptake rates of U. lactuca were extremely high at both subsaturating (15μM) and saturating (75 μM) N concentrations, as predicted by the functional-form hypothesis, tissue N fell to 1% by late summer. We suggest that a carbon imbalance, initiated by rising water temperatures and declining water-column N; thermal stress; and biological factors (competition, grazing) all contribute to the recurrent summer decline of U. lactuca in this shallow, eutrophic embayment. Thus, while the morphology of U. lactuca might be considered a successful strategy for disturbed, or “stressed” (sensu Littler and Littler 1980), habitats, its inability to persist and flourish in this environment emphasizes the complexity of factors at work in natural systems.     

High salt stress in the upper part of floating mats of Ulva prolifera,a species that causes green tides,enhances non‐photochemical quenching

Salt stress is a major abiotic stress factor that can induce many adverse effects on photosynthetic organisms. Plants and algae have developed several mechanisms that help them respond to adverse environments. Non‐photochemical quenching (NPQ) is one of these mechanisms. The thalli of algae in the intertidal zone that are attached to rocks can be subjected to salt stress for a short period of time due to the rise and fall of the tide. Ulva prolifera causes green tides and can form floating mats when green tides occur and the upper part of the thalli is subjected to high salt stress for a long period of time. In this study, we compared the Ulva prolifera photosynthetic activities and NPQ kinetics when it is subjected to different salinities over various periods of time. Thalli exposed to a salinity of 90 for 4 d showed enhanced NPQ, and photosynthetic activities decreased from 60 min after exposure up to 4 d. This indicated that the induction of NPQ in Ulva prolifera under salt stress was closely related to the stressing extent and stressing time. The enhanced NPQ in the treated samples exposed for 4 d may explain why the upper layer of the floating mats formed by Ulva prolifera thalli were able to survive in the harsh environment. Further inhibitor experiments demonstrated that the enhanced NPQ was xanthophyll cycle and transthylakoid proton gradient‐dependent. However, photosystem II subunit S and light‐harvesting complex stress‐related protein didn't over accumulate and may not be responsible for the enhanced NPQ.     

Tracing the origin of green macroalgal blooms based on the large scale spatio-temporal distribution of Ulva microscopic propagules and settled mature Ulva vegetative thalli in coastal regions of the Yellow Sea,China

From 2008 to 2016, massive floating green macroalgal blooms occurred annually during the summer months in the Yellow Sea. The original source of these blooms was traced based on the spatio-temporal distribution and species composition of Ulva microscopic propagules and settled Ulva vegetative thalli monthly from December 2012 to May 2013 in the Yellow Sea. High quantities of Ulva microscopic propagules in both the water column and sediments were found in the Pyropia aquaculture area along the Jiangsu coast before a green macroalgal bloom appeared in the Yellow Sea. The abundance of Ulva microscopic propagules was significantly lower in outer areas compared to in Pyropia aquaculture areas. A molecular phylogenetic analysis suggested that Ulva prolifera microscopic propagules were the dominant microscopic propagules present during the study period. The extremely low biomass of settled Ulva vegetative thalli along the coast indicated that somatic cells of settled Ulva vegetative thalli did not provide a propagule bank for the green macroalgal blooms in the Yellow Sea. The results of this study provide further supporting evidence that the floating green macroalgal blooms originate from green macroalgae attached to Pyropia aquaculture rafts along the Jiangsu coastline of the southern Yellow Sea.     

PHOTOSYNTHETIC PROPERTIES OF PROTOPLASTS,AS COMPARED WITH THALLI,OF ULVA FASCIATA (CHLOROPHYTA)1

Protoplasts were prepared from Ulva fasciata Delile, and their photosynthetic performance was measured and compared with that of thalli discs. These protoplasts maintained maximal rates of photosynthesis as high as those of thalli (up to 300 μmol O₂·mg chlorophyll^?1·h^?1) for several hours after preparation and were therefore considered suitable for kinetic studies of inorganic carbon utilization. The photosynthetic K_1/2(inorganic carbon) at pH 6.1 was 3.8 μM and increased to 67, 158, and 1410 μM at the pH values 7.0, 7.9, and 8.9, respectively. Compared with these protoplasts, thalli had a much lower affinity for CO₂ but approximately the same affinity for HCO₃^?. Comparisons between rates of photosynthesis and the spontaneous dehydration of HCO₃^? (at 50 μM inorganic carbon) revealed that photosynthesis of both protoplasts (which lacked apparent activity of extracellular/surface-bound carbonic anhydrase) and thalli (which were only 25% inhibited by the external carbonic anhydrase inhibitor acetazolamide) could not be supported by CO₂ formation in the medium at the higher pH values, indicating HCO₃^? uptake. Since both protoplasts and thalli were sensitive to 4,4′-diisothiocyanostilbene-2,2′-disulfonate, we suggest that HCO₃^? transport was facilitated by the membrane-located anion exchange protein recently reported to function in certain Ulva thalli. These findings suggest that the presence of a cell wall may constitute a diffusion barrier for CO₂, but not for HCO₃^?, utilization under natural seawater conditions.     

The effect of water flow on photosynthetic processes of the alga Ulva lactuca L.

Photosynthetic rates of aquatic macrophytes are affected by the diffusion of carbon through the boundary layer which decreases with increasing flow velocities. This is shown by a significant increase in photosynthetic rates of Ulva lactuca fronds exposed to friction velocities (u _*) between 0.0 and 0.3 cm s^–1. No further increase in photosynthetic rates is observed at u _* values between 0.3 and 1.0 cm s^–1. Stagnating flow conditions at saturating light levels caused a 29% reduction in photosynthetic rates. Experiments in a closed system indicate that Ulva may utilize internal carbon sources during periods of stagnation. On a subtidal rock directly exposed to wave action, Ulva lactuca is not exposed to such conditions while in a wave flushed intertidal pool stagnation occurs during very short time intervals based on in situ measurements of u _* levels on fronds of this species using a microprocessor-controlled, autonomous recorder and hot-film sensors.     

Photosynthetic and fatty acid acclimation of four phytoplankton species in response to light intensity and phosphorus availability

Photosynthetic acclimation of phytoplankton to lower irradiation can be met by several strategies such as increasing the affinity for light or increasing antenna size and stacking of the thylakoids. The latter is reflected by a higher proportion of polyunsaturated fatty acids (PUFAs). Additionally, photosynthetic capacity (P_max), respiratory losses, and proton leakage can be reduced under low light. Here we consider the effect of light intensity and phosphorus availability simultaneously on the photosynthetic acclimation and fatty acid composition of four phytoplankters. We studied representatives of the Chlorophyceae, Cryptophyceae and Mediophyceae, all of which are important components of plankton communities in temperate lakes. In our analysis, excluding fatty acid composition, we found different acclimation strategies in the chlorophytes Scenedesmus quadricauda, Chlamydomonas globosa, cryptophyte Cryptomonas ovata and ochrophyte Cyclotella meneghiniana. We observed interactive effects of light and phosphorus conditions on photosynthetic capacity in S. quadricauda and Cry. ovata. Cry. ovata can be characterized as a low light-acclimated species, whereas S. quadricauda and Cyc. meneghiniana can cope best with a combination of high light intensities and low phosphorus supply. Principal component analyses (PCA), including fatty acid composition, showed further species-specific patterns in their regulation of P_max with PUFAs and light. In S. quadricauda and Cyc. meneghiniana, PUFAs negatively affected the relationship between P_max and light. In Chl. globosa, lower light coincided with higher PUFAs and lower P_max, but PCA also indicated that PUFAs had no direct influence on P_max. PUFAs and P_max were unaffected by light in Cry. ovata. We did not observe a general trend in the four species tested and concluded that, in particular, the interactive effects highlight the importance of taking into account more than one environmental factor when assessing photosynthetic acclimation to lower irradiation.     

Growth of the seaweed Ulva rigida C. Agardh in relation to biomass densities, internal nutrient pools and external nutrient supply in the Sacca di Goro lagoon (Northern Italy)

Growth of the seaweed Ulva rigida C. Agardh was investigated in relation to biomass densities, internal nutrient pools and external nutrient supply. Research was carried out from 23 March to 5 July 1994 in the Sacca di Goro (Po Delta, Northern Italy), whose south-eastern part was covered by extensive mats of Ulva rigida. Two types of field experiments were conducted by incubating Ulva thalli inside large cages. In the first experiment, beginning on 23 March, 100 g of wet thalli were placed into the cages, allowed to grow for two weeks, then collected and replaced. This procedure was repeated 8 times over the study period. In the second experiment, Ulva thalli were left inside the cages and collected at selected time intervals (14, 27, 41, 64 and 76 days) in order to simulate the effects of increased density on growth and nutrient storage.We recorded specific growth rates (NGR) ranging from 0.025 to 0.081 d^–1 for a period up to two months in the repeated short-term experiments performed at relatively low initial algal densities (300–500 g AFDW m^–3). These NGR resulted significantly related to dissolved inorganic nitrogen (DIN) in the water column. Tissue concentrations of total Kjeldahl nitrogen (TN) were almost constant, while extractable nitrate decreased in a similar manner to DIN in the water column. Total phosphorus showed considerable variation, probably linked to pulsed freshwater inflow.In the long-term incubation experiment, NGR of Ulva was inversely related to density. Internal concentrations of both total P and TN reached maximum values after one month; thereafter P concentration remained almost constant, while TN decreased below 2% w/w (by dry weight). The TN decrease was also accompanied by an abrupt decrease in nitrate tissue concentration. The biomass incubated over the two month period suffered a progressive N limitation as shown by a decreasing NY ratio (49.4 to 14.6). The reciprocal control of Ulva against biogeochemical environment and viceversa is a key factor in explaining both resource competition and successional stages in primary producer communities dominated by Ulva. However, when the biomass exceeds a critical threshold level, approximately 1 kg AFDW m^–3, the macroalgal community switches from active production to rapid decomposition, probably as a result of selfshading, biomass density and development of anaerobic conditions within the macroalgal beds.