THALLUS DIFFERENTIATION OF PHOTOSYNTHESIS,GROWTH, REPRODUCTION,AND UV‐B SENSITIVITY IN THE GREEN ALGA ULVA PERTUSA (CHLOROPHYCEAE)1

The chlorophyte Ulva is perceived as a simple and uniform algal form, with little functional differentiation within a thallus. We compared morphology, pigmentation, photosynthesis, growth, reproduction, and UV‐B sensitivity between different thallus regions of Ulva pertusa Kjellman. Thallus thickness and cell size were significantly greater, whereas cell number was less in the basal region than in other regions. Photosynthetic pigment contents were lowest in the basal region and increased toward the marginal region. Photosynthetic capacity and photosynthetic efficiency normalized to fresh weight, area, volume, and cell number showed a progressive increase from the basal to marginal parts; however, on a chl basis those values were equal regardless of thallus part. Values of light saturation point were not statistically different between regions. Growth rates increased from marginal to basal and to middle parts of the thallus, whereas sporulation was highest in marginal (100%) followed by middle (30%) and basal parts (0%). Daily observation over 9 days showed that 56% of the basal cells divided once and did not produce spores, whereas every marginal cell went through its first division and 89% of the primary daughter cells also divided, resulting in 100% sporulation. A 7‐day treatment with PAR and PAR + UV‐A caused a significant decrease in the effective quantum yield of all thallus regions, followed by a recovery toward the initial values, whereas PAR + UV‐A + UV‐B irradiation led to greater photoinhibition and less recovery. Marked differences in the UV‐B sensitivity were observed, with marginal parts being more sensitive and basal parts most resistant.     

Variation in Patterns of Metal Accumulation in Thallus Parts of Lessonia trabeculata (Laminariales; Phaeophyceae): Implications for Biomonitoring

Seaweeds are well known to concentrate metals from seawater and have been employed as monitors of metal pollution in coastal waters and estuaries. However, research showing that various intrinsic and extrinsic factors can influence metal accumulation, raises doubts about the basis for using seaweeds in biomonitoring programmes. The thallus of brown seaweeds of the order Laminariales (kelps) is morphologically complex but there is limited information about the variation in metal accumulation between the different parts, which might result in erroneous conclusions being drawn if not accounted for in the biomonitoring protocol. To assess patterns of individual metals in the differentiated parts of the thallus (blade, stipe, holdfast), concentrations of a wide range of essential and non-essential metals (Fe, Cr, Cu, Zn, Mn, Pb, Cd, Ni and Al) were measured in the kelp Lessonia trabeculata. Seaweeds were collected from three sampling stations located at 5, 30 and 60 m from an illegal sewage outfall close to Ventanas, Chile and from a pristine location at Faro Curaumilla. For the majority of metals the highest concentrations in bottom sediment and seaweed samples were found at the site closest to the outfall, with concentrations decreasing with distance from the outfall and at control stations; the exception was Cd, concentrations of which were higher at control stations. The patterns of metal concentrations in different thallus parts were metal specific and independent of sampling station. These results and the available literature suggest that biomonitoring of metals using seaweeds must take account of differences in the accumulation of metals in thallus parts of complex seaweeds.     

A Comparative Study of Free Amino Acids of the Brown Alga <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> Linnaeus, 1753 from the Intertidal Zone of the Murman Shore,Barents Sea

The free amino-acid contents of the apical and middle parts of the thallus, as well as the receptacles of fertile and juvenile specimens, of the brown alga Fucus vesiculosus collected from seven different biotopes of the Murman shore, Barents Sea have been studied for the first time. The free amino-acid composition of the studied algae specimens was dominated by alanine, along with aspartic and glutamic acid. The distribution of free amino acids in the thallus of Fucus is uneven and depends on the habitat. The free aminoacid contents in the spring vary considerably between different parts of the thallus; however, certain regular patterns of their distribution are observed. A correlation has been found between the level of some individual amino acids in different parts of the thallus of fertile and juvenile algae and the salinity of the seawater.     

Environment determines nitrogen content and stable isotope composition in the sporophyte of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> (Harvey) Suringar

Nitrogen content and δ¹⁵N (nitrogen stable isotope ratio) were measured in different parts of the sporophyte of Undaria pinnatifida (Harvey) Suringar from two bays in the northeastern Japan. There were clear differences between the thalli collected in winter and in summer: high nitrogen content and low average δ¹⁵N were observed for the thalli collected in winter, while the opposite pattern was found for the thalli from summer. In addition, the pattern of internal δ¹⁵N distribution in the thallus changed with season. It is possible that the cause for these seasonal differences in the algae was the seasonal change in environmental conditions, because in winter water is normally rich in nitrogen, while in summer it is poor. U. pinnatifida sporophyte may be useful as an indicator of nitrogen sources in coastal waters, but consideration must be given to the effect of isotope fractionation on δ¹⁵N of the plant, especially in winter. Potential may exist for the use of different parts of the thallus to indicate nitrogen sources at different periods, but more investigation is necessary to accomplish this.     

Physiologisch-chemische Charakteristik verschiedener Thallusbereiche vonFucus serratus

Segments of five different thallus regions of the marine brown algaFucus serratus L. are physiologically and biochemically characterized with emphasis on certain aspects of their chemical composition and metabolism. Fresh weight, dry weight, and surface area values are intimately correlated. Fresh weight of the apical parts amounts to 1/3 of the basis, whereas dry weight of the same regions is only 1/5. The content of soluble and insoluble N-constituents shows characteristic differences between single thallus areas. Percentage of soluble nitrogenous compounds is least in thallus base. Pigment composition varies markedly with the age of the segments: pigment amounts are distinctly different in younger and older parts of the thallus, and even the relative proportions of the main pigments (chlorophylls and fucoxanthin) undergo measurable alterations. The content of carbohydrate constituents in different regions of the thallus is rather uniform; mean values for mannitol and alginate are 6.3 % and 28%, respectively. The content of laminarin is highest in the younger parts. All parts of theFucus thallus are photosynthetically active. A comparison of rates of photosynthesis indicates that the younger parts, especially the regions of growth and reproduction, are most active in carbon fixation. The¹⁴C-pattern of assimilates shows the general features of Phaeophyceae in all parts. In basal parts, however, remarkable amounts of¹⁴C-sucrose are additionally found. This is due to CO₂-fixation by the endophyticAcrochaete parasitica Oltm, which lives exclusively within the basal segments. The significance of these findings is discussed.     

Growth of the Brown Alga <Emphasis Type="Italic">Laminaria japonica</Emphasis> in a Laboratory Environment

The growth of an individual thallus of the brown alga Laminaria japonica was studied in a laboratory environment. It was found that daily accretion of the Laminaria thallus can remain constant for a long time. The blade grew in length at a distance from 0 to 15–25 cm from the stipe-blade border, which makes up 60% of the blade length. In width, the blade grew both in the central and in the lateral parts of the blade, 0 to 7–10 cm from the stipe-blade border, which makes up 30% of the frond length. The transposition of a perforation hole mark from the stipe-blade border toward the blade tip evidenced the formation of new tissue in the growth zone even when the thallus diminished in size due to destruction of the blade tip. Based on the results of this observation, it was concluded that both restraint of accumulation and even reduction of the algal thallus in biomass and size cannot be taken as an indication of algal growth cessation.Original Russian Text Copyright ¢ 2005 by Biologiya Morya, Skriptsova, Leletkin.     

Copper and iron concentrations in Ascophyllum nodosum (Fucales, Phaeophyta) from different sites in Ireland and after culture experiments in relation to thallus age and epiphytism

In laboratory experiments, copper concentrations in plants of Ascophyllum nodosum (L.) Le Jolis (Fucales, Phaeophyta) increased with the concentrations in the culture media and were highest in younger, meristematic thallus parts. After initial accumulation in high-copper medium and subsequent transfer to clean seawater for 5 days, no release of copper could be detected. Iron concentrations in A. nodosum tissue were not related to concentrations in the culture medium. Differences between copper concentrations in plants from different sites in areas with high yachting activity in Strangford Lough, Northern Ireland, could be explained by differences in water motion and human activity, in particular the application of copper-releasing antifouling paints to leisure boats. Iron concentrations were also highest in plants from the sheltered, polluted site but did not differ significantly between the other two sites. No differences in copper nor iron concentrations were found between different-aged thallus parts of plants from any site. X-ray microanalysis revealed that most of the iron detected was located in epiphytic pennate diatoms on the A. nodosum surface. In thallus areas without diatoms, iron levels were below the detection limit for X-ray microanalysis. Mapping for copper indicated that most of the accumulated copper was located in cells near and immediately below the thallus surface. "Epidermis"-shedding occurred in plants from the culture experiments and also in freshly-collected material and may have resulted in a loss of metal ions accumulated by surface cells and by epiphytic diatoms. The results suggest that A. nodosum could be used as a biological indicator for copper but not for iron, and that young, apical plant parts are most sensitive to changes in metal concentrations in the water.     

Further experiments on larval behaviour of the tubicolous polychaete Spirorbis inornatus L'Hardy & Quiévreux

Laboratory experiments indicate that there are many ecologically isolated populations of Spirorbisinornatus L'Hardy & Quiévreux, each tending to select the algal substratum which supported their parents. In some algae certain parts of the thallus are particularly favourable for settlement and, in some areas, this involves adaptation of the local population, e.g. to settle on fronds of Laminaria in sheltered localities. Other parts are unfavourable and these are generally the parts which are most illuminated in their natural orientation (e.g. the upper sides of Himanthalia buttons and the outer sides of Laminaria holdfast branches). These parts are active in producing antibiotics, which may thus have an antifouling role, deterring settlement of larvae whether or not those larvae come from populations adapted to settling upon other parts of the thallus.Most larvae from populations on the turf-alga Chondrus settle without any pelagic phase, whereas the majority from populations on Himanthalia and Laminaria holdfasts become pelagic. The proportions choosing holdfasts in laboratory experiments are high, however, suggesting that the need for skill in such choice, by these pelagic populations, more than balances any effect from outcrossing.     

Untersuchungen über den Einfluß des hydrostatischen Druckes auf den O2-Gaswechsel mariner benthischer Algen

The after effects of non-ecological hydrostatic pressures were studied upon the rates of respiration and photosynthesis of certain benthic algae from the Baltic Sea, North Sea and the Mediterranean Sea. The influences of different pressure intensities and the different durations of pressure were analyzed. It was observed how different parts of the same thallus are effected in their gas exchange by the same pressure. The interrelationships between hydrostatic pressure and the different ecofactors e. g. temperature, salinity, pH and O₂-concentration of the external medium were also studied. Furthermore the seasonal variations, the behaviour of shallow and deep water algae, the comparison of different ecotypes and the differences in taxonomically closely related algae were observed in regard to their pressure resistance.     

Relations entre production et localisation de mycosporine et morphogenèses reproductrices chez le Pyrénomycète Gnomonia leptostyla

Relations between production and localisation of mycosporin and reproductive morphogenesis in the Pyrenomycete Gnomonia leptostyla.
The production of mycosporin (P310) has been analysed in Gnomonia leptostyla (FT.) Ces. et de Not. during mycelial growth and reproductive morphogenesis (macroconidiogenesis, microconidiogenesis and differentiation of perithecia). Conidiogenesis is induced in illuminated cultures while darkness promotes perithecial development. At 20°C, the cultures produce either macroconidia or perithecia with abortive sporophyte. Microconidia differentiation and perithecia maturation require low temperature (10°C). Mycosporin is, at all times, present in the thallus. However, the concentration of mycosporin in highest in the conidiogenous thallus, intermediate in the perithecial thallus. and lowest in the vegetative mycelium. In the conidiogenous thallus, macroconidia and microconidia are both sites of mycosporin accumulation. On the contrary, in the perithecial thallus, mycosporin levels are not higher in perithecia than in mycelia, even during their maturation period. The quantitative variations of mycosporin during the thallus development and its accumulation inside conidia suggest translocation from sites of synthesis towards reproductive cells.     

Lipids of different thallus regions of the brown algaSargassum miyabei from the Sea of Japan

The fatty acid and lipid composition of different parts of the thallus of the brown algaSargassum miyabei was studied. The upper and lower thallus regions markedly differed in amounts ofn-3 polyunsaturated fatty acids (PUFAs), monoenoic fatty acids, phospholipids, and glycolipids. The lower thallus regions were rich in monoenoic fatty acids and phospholipids, while the upper parts had higher levels ofn-3 PUFAs and glycolipids.     

Higher reproductive success for chimeras than solitary individuals in the kelp <Emphasis Type="Italic">Lessonia spicata</Emphasis> but no benefit for individual genotypes

Chimerism is a peculiar, yet widespread, type of group living in which genetically heterogeneous entities are created through fusion between conspecifics. Here we tested whether chimerism provides direct benefits to the kelp Lessonia spicata, by analyzing its consequences on reproductive investment and success, at both the genotype and thallus levels. In addition, we quantified the frequency of chimerism in two natural populations, tested if group members were close kin, and evaluated the effects of relatedness and the number of genotypes per thallus on reproduction. Chimeric thalli were frequent (>60 %) in natural populations of L. spicata. In most cases, average intragroup relatedness was not significantly different from the background population. Reproductive investment was not significantly affected by the type of thallus (chimeric versus non-chimeric), by the number of genotypes per thallus or the average relatedness within thallus. Chimerism did not result in net benefits or costs in terms of genotypic reproductive success or probability of reproducing at the genotypic level. Yet, at the thallus level, chimerism increased reproductive success and the probability of reproducing, since more than one genotype reproduced in chimeric thallus. At the population level, chimerism affects L. spicata reproductive success by allowing the coexistence of a higher density of potential reproducers and mates compared to a scenario with only non-chimeric thallus. Chimerism may then have an important effect on the effective population size and possibly in reducing selfing rates.     

Effect of gibberellin and kinetin on the regeneration ability ofFucus vesiculosus L.

The influence of gibberellic acid (GA³) and kinetin (6-furfurylaminopurine) on the regeneration ability of the basal and apical thallus fragments ofFucus vesiculosus L. was examined. The naturally occurring gibberellin and kinetin-like substances in these thallus fragments were also studied. It was found that exogenously applied GA³ markedly increased the number of adventitious branches formed on the cut surface of the thallus fragments taken from the apical parts of plants. The concentration of 0.001 mg GA³ I-1 proved to be the most effective. The growth promoting effect of GA³ was increased by simultaneous action with kinetin. In experiments in which the fragments of the basal parts of the thallus were treated with GA³, as a rule a slight growth inhibition was observed. The growth responses of the investigated plant tissues to gibberellin and kinetin varied according to season. Usually their susceptibility to the applied plant hormones was greater in spring than is summer. The shifts in growth reaction were related to the seasonal changes in the content of endogenous gibberellin and kinetin-like substances in the investigated parts of the thallus. It is suggested that growth regulators of the gibberellin and cytokinin type are involved in the regeneration processes inFucus.     

Spatial organization of the three‐component lichen Peltigera aphthosa in functional terms

The cephalolichen Peltigera aphthosa (L.) Willd. is characterized by lateral heterogeneity, which manifests itself in the presence of three thallus zones, referred to as the apical, basal and medial zone. These zones differ in terms of interaction between lichen bionts and their physiological activity. The apical thallus zone is more efficient in establishing a contact with cyanobacteria, because of a higher lectin content and a larger overall thallus surface area due to the presence of numerous mycobiont hyphae. Cephalodia are formed in this zone. The interaction between the mycobiont and cyanobiont is more intense in the medial zone. However, the establishment of the contact with cyanobacteria in this zone less probable. The spatial distribution of lectins in the thallus was determined. To reveal the differences in photosynthetic activity in three thallus zones, transient analysis of chlorophyll a fluorescence and the assessment of non‐photochemical quenching of excited chlorophyll states were performed. Assimilation of absorbed light energy was more effective in the medial zone. The basal zone was characterized by decreased photosynthetic activity, lichen dissociation and thallus death.     

Developmental stages and secondary compound biosynthesis of mycobiont and whole thallus cultures of <Emphasis Type="Italic">Buellia subsororioides</Emphasis>

Lichen species have unique culture media preferences, and established cultures are known for the synthesis of secondary metabolites. This paper reports observations on the developmental stages and secondary compound biosynthesis by the mycobiont and whole thallus cultures of Buellia subsororioides. It also investigates the suitable media compositions for the culture growth, the role (nutrient or stressor) of sucrose concentrations on the growth stages, biomass, secondary compound profiles, and the quantity of biosynthesized known compounds/g of culture biomass for each treatment using mycobiont cultures. The ascospore-derived mycobiont cultures and thallus macerate-derived whole thallus cultures of B. subsororioides were established and grown using malt yeast extract (MY) medium. Mycobiont cultures were subcultured in MY medium supplemented with sucrose and its concentrations ranging from 0 to 30 % (with 2 % increment between treatments) for 120 days. The molecular identity of cultures was confirmed using nuclear ribosomal Internal transcribed spacer (ITS) DNA sequences obtained from the cultured mycobiont and from the natural thallus. The ITS DNA sequences of the mycobiont showed 99 % similarity with the sequences of the natural thallus. The mycobiont cultures under varying sucrose concentrations initiated as white cottony stages and transformed to brown compact mycelia, with optimum biomass and biosynthesis of nine secondary compounds in MY 10 %. The number of compounds (1–9) varies according to treatments. The whole thallus cultures (MY 0 %) showed a profile of secondary compounds similar to that of the natural thalli along with a trace of one unknown compound. The obtained results are encouraging for the synthesis of the desired quantities of lichen secondary compounds through cultures for relevant applications.     

Expression,subcellular localization,and <Emphasis Type="Italic">cis</Emphasis>-regulatory structure of duplicated phytoene synthase genes in melon (<Emphasis Type="Italic">Cucumis melo</Emphasis> L.)

The emission of molecular iodine (I₂) from the stipe, the meristematic area and the distal blade of the brown macroalga Laminaria digitata (Hudson) Lamouroux (Phaeophyceae) was monitored under low light and dark conditions. Photosynthetic parameters were determined to investigate both the extent of stress experienced by different thallus parts and the effects of emersion on photosynthesis. Immediately after air exposure, intense I₂ emission was detectable from all thallus parts. I₂ emission declined continuously over a period of 180 min following the initial burst, but was not affected by the light regime. The total number of mole of I₂ emitted by stipes was approximately 10 times higher than those emitted from other thallus parts. Initial I₂ emission rates (measured within 30 min of exposure to air) were highest for stipes (median values: 2,999 and 5,222 pmol g⁻¹ dw min⁻¹ in low light and dark, respectively) and lower, by one order of magnitude, for meristematic regions and distal blades. After exposure to air for between 60 and 180 min, I₂ emission rates of all thallus parts were reduced by 70–80%. Air exposure resulted in a decrease of the maximum photosystem II (PSII) efficiency (F _v/F _m) by 3%, and in a 25–55% increase of the effective PSII quantum efficiency ( \Updelta F/F^￠_\textm \Updelta F/F^{\prime}_{\text{m}} ); this was caused by a higher fraction of open reaction centres (q_P), whereas the efficiency of the latter in capturing energy ( F^￠_\textv /F^￠_\textm F^{\prime}_{\text{v}} /F^{\prime}_{\text{m}} ) remained constant. The results indicate the presence of an iodine pool which is easily volatilised and depleted due to air exposure, even under apparently low stress conditions.     

Morphology and anatomy ofCaloplaca coralligera (Teloschistaceae) as adaptation to extreme environmental conditions in the maritime Antarctic

The anatomical and morphological structure of the lichenCaloplaca coralligera (Hue.)Zahlbr. was investigated in connection with mechanisms of colonization and adaptation to the special conditions of maritime Antarctic.Caloplaca coralligera seems to be endemic to the antarctic region. The lichen is unique in its morphology, growing like a crustose lichen with a prothallus, but the very thallus being composed of many vertical, frutescent branches not higher than large isidia. Due to fusions of the distal parts of the branches the morphology is characterized by air-filled cavities between the erect thallus parts where the phycobionts are located only close to the outermost surface. But also anatomically a system of air-filled cavities is developed by a net-like structure of hyphae. This may result in an insulation from temperature exchanges with the rocky substrate. It is speculated that this peculiar thallus structure might be advantageous to a lichen growing on compact substrates by buffering the diurnal temperature extremes which are characteristic for rocks under a strong radiation exchange with the atmospheric surrounding.Dedicated to Prof. DrO. L. Lange on the occasion of his retirement and of his 65th birthday.     

Corrélations de croissance entre le mycélium et les rhizomorphes chez le Sphaerostilbe repens

Correlation of Growth between Mycelium and Rhizomorphs in Sphaerostilbe repens In Sphaerostilbe repens Berkeley and Broome, cultured on a solid medium in Petri dishes, growth of mycelium and chizomorphs depends on the thickness of the culture medium underlying the thallus. On shallow layers of agar medium, aggregated units were fewer and rhizomorph growth reduced. Vegetative mycelium which starts growth earlier than rhizomorphs contributed to inhibit the development of these latter. On deep layers of substrate, rhizomorphs were numerous, elongated rapidly, and eventually inhibited undifferentiated mycelium. – There was no evidence for the involvement of any inhibitor substance diffusing into culture medium or located in the thallus. Development of aerial mycelium and rhizomorphs, could, on the contrary, be interpreted as nutritional competition between these two parts of the colony.     

Regenerative capacity of Gracilaria fragments: effects of size,reproductive state and position along the axis

Field farming of Gracilaria chilensis in Chile is mainly based on the regenerative capacity of thallus fragments. This study evaluates experimentally the effects on growth of the few organismic determinants that the farmer is capable of effectively handling in the field. No differences were found in the length increments of ramets with and without apices, a surprising result in view of the generally accepted idea that the most important growth meristem in this group of algae is located in the apex of the thallus. Total elongation and increase in weight of fragments with an initial length of up to 20 cm are a function of the initial length of the ramet, suggesting that intercalary growth makes a significant contribution to growth. Vegetative female axes grow faster, (in terms of length and weight) and produce more laterals than fertile female axes, and these faster than fertile tetrasporic branches. No differences in the growth of ramets originating from proximal or distal parts of axes were found. Paper presented at the XVth International Seaweed Symposium.