New finds of charophytes in Iceland with an update on the distribution of the charophyte flora

Charophyte species new to the flora of Iceland are reported and an update on the distribution of the previously known species is given. Four species are new to Iceland; Chara aspera Willdenow, Chara contraria Braun ex Kützing, Tolypella canadensis Sawa and Tolypella glomerata (Desvaux) Leonhardi, while four species were previously known: Chara globularis Thuiller, Chara virgata Kützing, Nitella flexilis (L.) Agardh and Nitella opaca (Agardh ex Bruzelius) Agardh. The finding of the species new to Iceland add to the hitherto known worldwide distribution of those species, including a significant extension to the north.     

Restoration of Botshol (The Netherlands) by reduction of external nutrient load: recovery of a characean community,dominated by Chara connivens

Till about 1965 a rich characean community occurred in the shallow peat lake Botshol with six species of which the rare Nitellopsis obtusa and Chara major dominated at many sites. In the period 1980–1988 characean biomass strongly decreased and only two species, Chara globularis and C. connivens, remained in small populations at a few localities. Of the macrophyte Najas marina also some small populations remained, while the aquatic moss Fontinalis antipyretica and the filamentous alga Vaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the input of polluted water. This process of impoverishment was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca 25 µg l^-1) and a higher water transparency. Immediately after these measures the Characeae community increased in abundance and number of species. During the summer of 1990, and especially 1991, a spectacular growth occurred of Chara connivens. C. connivens was often accompanied by C. major. Other species with scattered occurrence were C. aculeolata, C. aspera, C. contraria and C. globularis. The reasons for the shift in dominance from Nitellopsis obtusa to Chara connivens are discussed. From growth experiments evidence was obtained that neither the recent higher chloride level, nor the lowered phosphate concentration were the main factors for the domination of Chara connivens.     

Restoration of Botshol (The Netherlands) by reduction of external nutrient load: Recovery of the Characean community

The formerly rich characean community in Botshol with six species of which the rareNitellopsis obtusa andChara hispida dominated at many sites, decreased to only two species,Chara globularis andC. connivens, in the period 1980–1988. The macrophyteNajas marina also remained at some sites, and the aquatic mossFontinalis antipyretica and the filamentous algaVaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the inlet of polluted water. This process of eutrophication was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca. 0.024 mg l^–1) and a higher water transparency. Immediately after these measures the Characeae community increased strongly in abundance and number of species. During the summer of 1990, and especially of 1991, a spectacular growth occurred ofChara connivens. Chara connivens was often accompanied byChara hispida. Other species with scattered occurrence wereChara aculeolata, C. aspera, C. contraria andC. Globularis. The reasons for the shift in dominance fromNitellopsis obtusa toChara connivens are discussed. One of the reasons may be the recent higher chloride content which is one of the consequences of the restoration measures.     

Chemical,physical and biological changes associated with chara succession in farm ponds

Four farm ponds in various stages of Chara vulgaris succession -from the initial invasion of the alga to its nearly complete replacement-were followed each week from ice cover to ice cover. Chemical, physical, and biological parameters were analyzed in order to determine differences between ponds and to discover the biological and physicochemical factors associated with succession in Chara vulgaris ponds. The data were analyzed statistically by intraclass (single factor) analysis of variance, the Student's t-test, and stepwise multiple regression analysis.The initial invasion of C. vulgaris appeared to be preceded by a lowering of the nutrient levels of the pond by Zygnematales growth and by the substrate-stabilizing growth of Rhizoclonium sp. Chara-dominated ponds had significantly lower free CO₂, bicarbonate alkalinity, and total hardness readings. Carbonate alkalinity, low phytoplankton productivity, high dissolved O₂, low phosphate, and high Secchi disk readings were also characteristic of ponds in which C. vulgaris was dominant.In ponds where C. vulgaris appeared in the process of being replaced, the physicochemical factors were more closely associated with phytoplankton productivity and diversity and less closely with submerged vegetation species.Succession from C. vulgaris domination to a Najas-Potamogeton community appeared to involve shading of the former by the latter and was accelerated by the smothering effect of decaying debris caused by Gloeotrichia sp.In the final stages of succession, turbid conditions resulting from the lack of substrate-stabilizing vegetation and increased phytoplankton growth appear to have discouraged C. vulgaris growth early in the season.     

Can Chara control phosphorus cycling in Lake Łuknajno (Poland)?

Waters of Lake uknajno (623 ha, 3 m max. depth postglacial basin in Masurian Lakeland) are of meso-eutrophic type in spite of an intensive land impact and substantial phosphorus loading. The reason of this peculiarity is an abundance of 4 species of charophytes (Chara aculeolata, C. aspera, C. contraria and C. tomentosa) which dominate the submerged vegetation of the lake and yielded 417 g m^-2 dry biomass in summer 1991. The plants have high potential for phosphorus uptake and accumulation as was checked in laboratory experiments. Due to this potential Chara can outcompete planktonic algae in mixed cultures at different mineral P supplies.     

Long-term changes in macrophyte vegetation after reduction of fish stock in a shallow lake

Starting in the middle of the 1970s, submerged macrophytes began to disappear from shallow Lake Warniak due to feeding pressure by grass carp (Ctenopharyngodon idella). In the middle of the 1980s, the lake was stocked with seston-feeding silver carp (Hypophthalmichthys molitrix) and bighead carp (Aristichthys nobilis). By 1993, the biomass of silverhead carp and bighead carp had declined. This allowed charophytes to recolonize the bottom of the lake. The main charophyte species at this time were Chara globularis and Chara rudis. Since then, five other stonewort species have been found: Chara contraria, C. filiformis, C. tomentosa, C. aspera and Nitellopsis obtusa. Seventeen species of aquatic angiosperms have also been found. There were distinct changes in the relative abundance and spatial distribution of particular species. C. rudis developed most intensely in the shallow parts of the lake near the southern and western shores. C. globularis gradually took over the deeper central part of the lake. In 2001, C. rudis began to retreat again. The relative abundance and spatial distribution of charophytes was correlated to water clarity (r = 0.87, p < 0.05), total phosphorus level (r = −0.78; p < 0.05), and chlorophyll a content (r = −0.79; p < 0.05).     

Allelopathic activity of Chara aspera

Allelopathic activity of Chara aspera was determined in agar diffusion assays using planktonic cyanobacteria as target organisms. Growth inhibition of cyanobacterial strains was observed in bioassays inoculated with living Chara aspera shoots as well as with 60% aqueous methanol extracts of Chara aspera. For further analysis, the methanol extract was fractionated into three parts: a lipophilic methanol – a butylmethylether-extract and a hydrophilic methanol extract. The bioassays indicated that major allelopathic activity was retained in the hydrophilic methanol – and the lipophilic butylmethylether-extract. Separation of the extracts by means of high performance liquid chromatography followed by fractionation of the eluant resulted in supplementary nine fractions, three from each part, respectively. Three fractions exhibited a strong growth inhibition of the target organism Anabaena cylindrica Lemmermann. The second and the third fraction of the lipophilic butylmethylether extract indicate the presence of novel allelopathic active compounds with lipophilic characteristics. The results lead to the suggestion that more than two chemical compounds in Chara aspera are responsible for the growth inhibition of cyanobacteria.     

Importance of Nutrient Competition and Allelopathic Effects in Suppression of the Green Alga Scenedesmus obliquus by the Macrophytes Chara, Elodea and Myriophyllum

Possible allelopathic effects of substances released from the macrophytes Chara globularis, Elodea canadensis, Myriophyllum spicatum on the common green alga Scenedesmus obliquus were tested in the laboratory with plastic plants and untreated medium as controls. A two-phase approach was used in which first the effects of physical presence of plants was studied (phase I) followed by the effects of plant culture filtrates (phase II). In the presence of plastic plants growth was reduced only marginally, but strong growth inhibition of Scenedesmus occurred in the physical presence of all macrophytes. In contrast, filtrates from Chara had no growth inhibitory effect on Scenedesmus. Myriophyllum filtrate reduced particle-based growth rate by 7% compared to filtration controls, while Elodea culture filtrate reduced volume-based growth by 12%, chlorophyll-based growth by 28% and particle-based growth by 15%. Photosystem II-efficiency of Scenedesmus was reduced in all three macrophyte treatments in phase I, but not in filtrates from macrophyte cultures (phase II). Thus, while enzyme activity or other physiological aspects may have been affected, the current study yielded no proof for allelopathically active compounds being directed at photosynthesis. Mean particle volume (MPV) of Scenedesmus was not influenced by macrophyte exudates and cultures remained dominated by unicells. The strong growth inhibitory effects found for Scenedesmus in the physical presence of macrophytes, but not in plastic controls, and no or weaker response in nutrient-enriched filtrates, suggest nutrient competition was a more powerful driving factor than allelochemicals. However, the experimental design does not exclude disappearance of allelochemicals during the filtration process.     

Successional events following simazine application

Successional events of aquatic vegetation in a farm pond were studied, after application of simazine. After decay of the higher plants, phytoplankton did not dominate, instead herbicide-resistant seeds and subsurface structures of Potamogeton foliosus developed. Benthic algae covered and stabilized the bottom. Following stabilization, the water cleared and Chara vulgaris growth resumed wherever the substrate was firm.Suggestions are made as to how a pond can be managed to maintain the desired pioneer vegetation of Ch. vulgaris.     

The seasonal effects on the encrustation of charophytes in two hard‐water lakes

Encrustation and element content of six charophyte species from two hard‐water lakes were investigated monthly for a period of 1 year. Seasonal patterns were analyzed for the interaction of water chemistry. Encrustation followed a seasonal pattern for Chara contraria, Chara subspinosa, and Nitellopsis obtusa in Lake Krüselin and for Chara globularis and Chara tomentosa in Lake Lützlow. However, no seasonality in the precipitated CaCO₃ was observed for C. subspinosa in Lake Lützlow and for C. tomentosa in Lake Krüselin, indicating a lake‐specific dependency. Species‐specific encrustation was found. Chara contraria and N. obtusa encrusted the most in June and August, whereas C. subspinosa and Nitella flexilis/opaca exhibited lowest encrustation in March and April. The precipitated CaCO₃ of charophytes correlated negatively to the concentration of total inorganic carbon in both lakes. Element content of plant dry weight was species‐specific for Ca and K, and lake‐specific for Mg. No specific pattern was found for the TP and Fe contents. The results showed seasonal, species, and lake‐specific influences on the encrustation of charophytes.     

Immunolocalization of myosin in rhizoids ofChara globularis Thuill

Summary Myosin-related proteins have been localized immunocytochemically in gravity-sensing rhizoids of the green algaChara globularis using a monoclonal antibody against the heavy chain of myosin from mouse 3T3 cells and a polyclonal antibody to bovine skeletal and smooth muscle myosin. In the basal zone of the rhizoids which contain a large vacuole, streaming endoplasm and stationary cortical cytoplasm, the monoclonal antibody stained myosin-related proteins as diffusely fluorescing endoplasmic strands. This pattern is similar to the arrangement of subcortical actin filament bundles. In the apical zone which contains an aggregation of ER membranes and secretory vesicles for tip growth, diffuse immunofluorescence was detected; the intensity of the signal increasing towards the apical cell wall. The most prominent myosin-staining was associated with the surface of statoliths in the apical zone. The polyclonal antibody produced a punctate staining pattern in the basal zone, caused by myosin-related proteins associated with the surface of drganelles in the streaming endoplasm and the periphery of the nucleus. In the apical zone, this antibody revealed myosin-immunofluorescence on the surface of statoliths in methacrylate-embedded rhizoids. Neither antibody revealed myosin-immunofluorescence on the surface of organelles and vesicles in the relatively stationary cytoplasm of the subapical zone. These results indicate (i) that different classes of myosin are involved in the various transport processes inChara rhizoids; (ii) that cytoplasmic streaming in rhizoids is driven by actomyosin, corresponding to the findings onChara internodal cells; (iii) that actindependent control of statolith position and active movement is mediated by myosin-related proteins associated with the statolith surfaces; and (iv) that myosin-related proteins are involved in the process of tip growth.     

A surgical method for accessing the plasmamembrane ofChara australis

Summary Progress in patch-clamping the plasmalemma ofChara australis has been slow because of the difficulty in obtaining a clean membrane surface upon which obtain a high resistance seal with the patch pipette. A surgical method is described here for cutting an access window in the cell wall of internodal cells fromChara australis. The method has a high success rate and also has the advantage of obviating the need for enzyme treatments. The patch-clamp technique is applied to theChara plasmalemma. Five types of channel activity are observed of which four of these are selective for Cl^– over K⁺.     

Morphological and Morphometrical Variations of Selected Rotifer Species in Response to Predation: a Seasonal Study of Selected Brachionid Species from Lake Xochimilco (Mexico)

Allelopathy of filamentous green algae (FGA) has been less studied than that of macrophytes. Little Budworth Pool, Cheshire, UK is a small, shallow, clear-water lake with high TP concentrations, very high NO_3-N concentrations, only moderate phytoplankton density, high FGA growth (mainly Spirogyra sp.) and no submerged plants. Experiments were carried out to test the possible allelopathic effects of Spirogyra on the phytoplankton of this lake and on a submerged plant Elodea nuttallii. Changes in phytoplankton growth, phytoplankton species dynamics and species composition were apparently not influenced by allelopathy of live or decaying Spirogyra. A shift from diatom (Cyclotella sp) – cryptomonad (Chroomonas acuta and Cryptomonas erosa) dominance to Chlorococcales (Micractinium pusillum, Monoraphidium contortum and Scenedesmus opoliensis) – Volvocales (Chlorogonium elongatum and Pandorina morum) dominance was recorded in both control and FGA treatments, suggesting an effect of nutrient enrichment. Nutrient concentrations and differences in competitiveness among phytoplankton species can also explain differences in their growth rates in Spirogyra filtrate. Spirogyra also did not influence apex number per plant, shoot length or growth rate of E. nuttallii. This FGA species probably cannot control phytoplankton or E. nuttallii growth in nutrient rich conditions through allelopathy.     

Effects of pH on toxicity of As,Cr, Cu,Ni and Zn to Selenastrum capricornutum Printz

Acute toxicity tests were conducted to establish the response of Selenastrum capricornutum Printz to sublethal concentrations of As, Cr, Cu, Ni and Zn at a broad range of pH levels. Cultures were incubated for a period of seven days at pH 4 in standard algal assay media containing sublethal concentrations of metals. At this low pH, growth was depressed for all metals tested. The adjustment of pH to higher levels resulted in increased growth when cultures were treated with As, Cu, or Ni and incubated for an additional 7 days. Toxicity was least at the optimum pH range for growth of the alga.The observation that the toxicity of As, Cu, and Ni to S. capricornutum decreases markedly at pH values above 4.0 may be of ecological importance in the control of acid mine pollution. If a high percentage of algae show a similar response to decreasing toxicity with increasing pH, it clearly would be of value to adopt measures which control pH as well as the levels of metals present. It was suggested that algae with a broad pH growth range, such as S. capricornutum, could benefit from the addition of highly alkaline materials to waters where certain metals are present.     

In situ measurement of algal growth potential in aquatic ecosystems by immobilized algae

Cells of the green algaSelenastrum capricornutum were immobilized in alginate beads. The alga was able to grow inside these beads without being grazed by zooplankton. For P-limited immobilized cells, however, a lower µ _m and initial slope of the Monod growth curve µ _m /K _s were found than for free cells.To study the feasibility of immobilized algae to estimate algal growth potentialin situ in aquatic ecosystems, a series of experiments were conducted in indoor model ecosystems (microcosms) and in a small stream. The use of immobilized algae allowed a continuous registration of algal growth potential integrated over periods with natural fluctuations in the environment. The method of encapsulation of the algae can, however, still be improved. The alginate matrix is exposed to marked degradation by microorganisms when incubated in polluted streams for a period longer than two weeks. The applicability of other types of matrices should be tested.Author for correspondence     

Seasonal variation in invertebrate grazing on Chara connivens and C. tomentosa in Kõiguste Bay, NE Baltic Sea

Charophytes are a highly endangered group of algae. In the Baltic Sea, the number of species, distribution area and biomass of charophytes have significantly decreased in recent decades. Although eutrophication triggers their initial decline, the mechanism of the final extinction of charophyte populations is not fully understood. An in situ experiment was performed to study the role of the mesoherbivores Idotea baltica, Gammarus oceanicus and Palaemon adspersus in the decline of charophytes in the north-eastern Baltic Sea. Invertebrate grazing showed a clear seasonality: grazing pressure was low in April, moderate in July, and high in October. Grazing on charophytes by P. adspersus was negligible, whereas I. baltica and G. oceanicus significantly reduced the biomass of charophytes in the field. Low photosynthetic activity (high decomposition rate) of the charophytes favoured grazing. The invertebrates studied preferred Chara tomentosa to C. connivens. Low consumption of C. connivens may reflect its non-native origin. The experiment suggests that, under moderately eutrophic conditions, grazers are not likely to control charophyte populations. However, grazers have the potential to eliminate charophytes in severely eutrophic systems under the stress of filamentous algae.Communicated by H.D. Franke     

Interaction between the macrophyte <Emphasis Type="Italic">Stratiotes aloides</Emphasis> and filamentous algae: does it indicate allelopathy?

The aquatic macrophyte Stratiotes aloides Linnaeus, which has recently received attention in studies on allelopathy, has been shown to suppress phytoplankton growth. In the Netherlands, S. aloides often co-occurs with floating filamentous algae. However, filamentous algae are generally absent in close proximity to S. aloides, resulting in gaps in filamentous algae mats. We analyzed whether those gaps may be caused by allelopathic substances excreted by S. aloides or by nutrient depletion. We studied in a field survey the colonization of natural S. aloides by filamentous algae and determined in situ nutrient concentrations in natural S. aloides stands. To analyze the relative importance of allelopathy and nutrient competition in the interaction between S. aloides and filamentous algae, we carried out field experiments. Introduction of artificial (non-allelopathic) plants in natural S. aloides stands enabled us to compare the colonization by filamentous algae of both Stratiotes sp. and artificial plants. The filamentous algae were absent in close vicinity to S. aloides. Significantly lower concentrations of ortho-phosphate and potassium were observed close to S. aloides as compared with the filamentous algae. In the field experiments the artificial plants were rapidly colonized by filamentous algae, mainly Cladophera Kützing and Spirogyra Link, while all natural plants remained free of such algae. Additionally, most nutrient concentrations did not significantly differ in the proximity of artificial or natural stands of S. aloides. The concentrations of the major growth-limiting nutrients, phosphate and nitrate, were significantly higher and nonlimiting in natural Stratiotes stands. Our main conclusion is that, although allelopathic interactions between S. aloides and filamentous algae do occur under natural conditions, nutrient competition between the two can also be an important factor.     

Reduced digestibility of UV-B stressed and nutrient-limited algae by Daphnia magna

Daphnia magna was fed the green alga Selenastrum capricornutum, cultured under four different growth conditions: (1) phosphorus limitation, (2) nitrogen limitation, (3) UV-B irradiation, and (4) no nutrient limitation, no UV-B irradiation. Contrary to non-limited algal cells, nutrient-limited cells were not efficiently assimilated. Especially, P-limited cells passed through the gut mostly intact, while N-limited cells were partly assimilated. Also, algae exposed to moderate doses of UV-B radiation (0.3 mW cm^–2 of UV₃₁₂) were less efficiently assimilated after being grazed. Digestibility of the algae decreased with increased UV-B exposure time. Nutrient-limited and UV-B stressed algal cells increased in volume and became granular in appearance. These changes in the algal cells, combined with changed cell wall properties, most probably reduced their digestibility.     

Hypergravity can reduce but not enhance the gravitropic response ofChara globularis protonemata

Summary The relationship between the position of the statoliths and the direction and rate of tip growth in negatively gravitropic protonemata ofChara globularis was studied with a centrifuge video microscope. Cells placed perpendicularly to the acceleration vector (stimulation angle 90 °) showed a gradual reduction of the gravitropic curvature with increasing accelerations from 1g to 8g despite complete sedimentation of all statoliths on the centrifugal cell flank. It is argued that the increased weight of the statoliths in hypergravity impairs their acropetal transport which is induced when the cell axis deviates from the normal upright orientation. When the statoliths were centrifuged deep into the apical dome at 6g and a stimulation angle of 170 ° the gravitropic curvature after 1 h was identical to that determined for the same cells at 1g and the same stimulation angle. This indicates that gravitropism in Chara protonemata is either independent of the pressure exerted by the statoliths on an underlying structure or is already saturated at 1g. When the statoliths were moved along the apical cell wall at 8g and the stimulation angle was gradually increased from 170 ° to 220 ° the gravitropic curvature reverted sharply when the cluster of statoliths passed over the cell pole. This experiment supports the hypothesis that in Chara protonemata asymmetrically distributed statoliths inside the apical dome displace the Spitzenkörper and thus the centre of growth, resulting in gravitropic bending. In contrast to the positively gravitropic Chara rhizoids, no modifications either in the transport of statoliths during basipetal acceleration (6g, stimulation angle 0 °, 5 h) or in the subsequent gravitropic response could be detected in the protonemata. The different effects of centrifugation on the positioning of statoliths in Chara protonemata and rhizoids indicate subtle differences in the function of the cytoskeleton in both types of cells.Dedicated to Prof. Dr. Zygmunt Hejnowicz on the occasion of his 70th birthday     

Ultrastructure and phylogeny of the spermatozoid ofChara vulgaris (Charophyceae)

At maturity, spermatozoids of the green algaChara vulgaris are biflagellated, contain little cytoplasm, and coil for approximately 2 1/2 gyres within the mother cell wall. The anterior of the cell contains an ovoid headpiece anchoring two slightly staggered basal bodies that are positioned above and directly in front of approximately 30 linearly arranged mitochondria. An elongated stellate pattern occupies the transition zone between the BBs and axonemes. Flagella emerge from the cell just in front of the nucleus and encircle the full length of the spermatozoid. The spline comprises a maximum of 38 microtubules surrounding the anterior mitochondria and gradually decreases posteriorly to a minimum of 11. The dense nucleus is narrow, cylindrical, and occupies the central revolution of the cell. Six starch-laden plastids and associated mitochondria are linearly arranged at the cell posterior. Phylogenetic analyses of charalean taxa and archegoniates based on spermatogenesis strongly support the orderCharales, withNitella as the sister group toChara. Diagnostic features ofChara spermatozoids include absence of a lamellar strip and axonemes embedded in the cell for almost the entire length of the anterior mitochondria. Potential relationships amongCharales, Coleochaetales and archegoniates are evaluated in regards to the probable course of evolution of streamlined biflagellated gametes.