Effects of diatom mucilage on the growth and Morphology of marine algae

Germination and growth patterns were studied of germlings and sporelings of eight species of marine algae when grown on diatom mucilage. Whilst growths of green and brown algae were enhanced, growth of the red algae was inhibited. Morphologically abnormal sporelings of Ulva lactuca L. and Gigartina stellata (Stackh.) Batt. were obtained in the presence of the mucilage.     

Factors controlling the lower limits of fucoid algae on the shore

Field and culture studies on fucoid alga of the upper shore showed that their lower limits of distribution were determined largely by interspecific competition. Pelvetia canaliculata (L.) Dcne. et Thur., which is strictly confined to the highest algal zone, grew much more slowly, both in culture and on the shore, than did Fucus spiralis L., which occupies the zone immediately below Pelvetia. When transplanted to the Fucus spiralis zone, macroscopic Pelvetia thalli grew normally, but Pelvetia zygotes which germinated within this zone reached macroscopic size only when competing Fucus spiralis was repeatedly removed.Both species required high light levels to grow in culture, yet their embryos remained viable for long periods in total darkness. Thus, Pelvetia cannot grow in the shade of Fucus spiralis, but actual removal of the Pelvetia germlings from the Fucus spiralis zone must be affected by some other factor.F. spiralis grew vigorously when transplanted to M.T.L., a level at which it is normally rare. In culture, its embryos grew more slowly than those of the species found lower on the shore such as F. vesiculosus L. and F. serratus L.; hence interspecific competition may again be involvedYoung Ascophyltum nodosum (L.) Le Jol. plants grew almost as slowly as those of Pelvetia, yet Ascophyllum dominates the middle shore. In culture it showed only slightly greater tolerance of low light intensity than other fucoids but, unlike Pelvetia, persisted and grew beneath a Fucus canopy in the field. The possibility that differences in life history and palatibility to grazing molluscs contribute to the disparate competitive abilities of Pelvetia and Ascophyllum is discussed.     

An ecological study of fucus spiralis L.

An ecological study of Fucus spiralis L. has been made at Jaffrey Point, Newcastle, New Hampshire and the adjacent Great Bay Estuary System from 1972–1975. The distribution, growth, reproductive periodicity, attrition and longevity of the plants are described in relation to a variety of environmental factors. F. spiralis shows a broad but discontinuous estuarine distribution within the Great Bay Estuary System. The presence or absence of appropriate substrata is considered to be a factor determining its discontinuous distribution, since it is usually associated with metasedimentary or metavolcanic rock outcrops. The maximum growth and reproduction of F. spiralis are during the summer. The summer growth rates ranged from 1.9 to 2.8 cm/month while the average growth rate throughout the year was 1.2 cm/month.F. spiralis populations showed two periods of major attrition, namely, during the winter and summer. The average longevity of F. spiralis plants is ≈ 2 yr. The F. spiralis zone lies in the uppermost intertidal area between +2.12 and +2.31 m above M.L.W. There is a micro-stratification of biomass, stature, and reproductive activity within this zone. Average plant weight, length, and fertility tend to decrease with increasing elevation.     

Tolerance of Sargassum thunbergii germlings to thermal, osmotic and desiccation stress

The construction of artificial seaweed beds in the intertidal zone is a challenge due to extreme levels of physical stress. In order to provide a basis for the construction using the dispersal of microscopic juveniles, a three-way factorial experimental design was used to evaluate the tolerance of Sargassum thunbergii germlings shortly released from fertile thalli to temperature, salinity and desiccation in this study. Results revealed that temperature, salinity and desiccation significantly affected the growth and survival of germlings. Germlings showed rapid growth with relative growth rate (RGR, % day⁻¹) over 16% when cultured at 25 °C and full immersion in normal seawater. Although growths of germlings subjected to moderate conditions were significantly inhibited, RGRs over 13% were obtained. The RGRs of germlings below 10% were observed only at 35 °C and 9 h desiccation treatments. In comparison to growth, survival was less affected by physical stress. Germlings showed low mortalities below 10% under appropriate conditions (25 °C and 30 °C combined with full immersion), and below 60% under moderate conditions, by the end of experiment. However, the mortality rates increased to over 90% under extreme conditions (9 h desiccation and 35 °C combined with full immersion in salinity of 12). These results showed that S. thunbergii germlings had high tolerance to physical stresses. In addition to the main effects, both two-way and three-way interactions between temperature, salinity and desiccation were significant. Based on the magnitude of effect, desiccation was the predominant factor affecting both growth and survival. According to the results, construction of artificial tanks in natural habitat to minimize desiccation may be an effective strategy for S. thunbergii restoration using germlings.     

Short-term effects of temperature upon the growth of intertidal fucales

The growth rate of five species of intertidal Fucales (Pelvetia canaliculata (L.) Dec. et Thur., Fucus spiralis L., Fucus vesiculosus L., Fucus serratus L., Ascophyllum nodosum (L.) Le Jolis) was measured at temperatures from 2.5 to 35 °C. An increase in temperature immediately causes a high growth rate, and during the first hour it increases linearly with temperature; at 35 °C it is 20 times the control at 7 °C. This acceleration of growth is based mainly on stored photosynthate. After the first few hours the growth rate decreases rapidly, particularly at the highest temperatures. After 2–3 weeks a temperature optimum below 17.5 °C is indicated. High temperatures, 30–35 °C, were lethal to all species, with a survival time corresponding to their vertical zonation in the natural habitat.     

Arrested development in Fucus spiralis (Phaeophyceae) germlings exposed to copper

Exposure of Fucus spiralis germlings to precise copper concentrations (0 to 844?nM?Cu²⁺) in chemically defined medium demonstrated a relationship between ultrastructural changes and growth retardation with increasing copper concentration. Electron-translucent vesicles, present in ova, which normally disappear after fertilization, accumulated in germlings exposed to Cu²⁺ above 10.6?nM, suggesting that copper may inhibit a metabolic pathway involved in cell wall formation which is initiated by fertilization. No membrane damage was observed during the exposure period. During a post-exposure period in copper-free medium, recovery occurred (rhizoid extension, apical hair formation) in germlings previously exposed to concentrations below 106?nM?Cu²⁺ and electron-translucent vesicles became granular and disappeared. It is proposed that the electron-translucent vesicles contain a cell wall precursor and that copper inhibits its incorporation into the cell wall, preventing growth and development of the zygote.     

Factors controlling the upper limits of fucoid algae on the shore

Year-round observations on the condition of intertidal seaweeds growing in situ on the shore, show that the upper limits of the zones characterized by Pelvetia canaliculata (L.) Done et Thur., Fucus spiralis L. and Ascophyllum nodosum (L.) Le Jol. were periodically pruned back by environmental conditions. The uppermost plants of each species showed clear signs of tissue damage 21 to 28 days after a time when drying conditions coincided with neap tides which exposed the plant to aerial conditions for long periods. High air temperatures aggravated the damage, but neither frost nor prolonged rain had any obvious adverse effects. On spring tides the plants were wetted every day and no damage resulted regardless of the weather.These species clearly all reach up to their physiological limits on the shore investigated, but presumably Fucus vesiculosus L. and F. serratus L. do not, for they were never observed to show signs of tissue damage attributable to exposure to air. Transplant experiments did, however, prove that F. serratus cannot survive in the F. spiralis zone and nor can F. spiralis persist in the Pelvetia canaliculata zone.Laboratory experiments also demonstrated that the ability to tolerate desiccation and then to resume photosynthesis and growth when re-submerged was greatest in P. canaliculata, the species found highest on the shore, and was progressively less in species inhabiting successively lower levels.     

Settlement, early growth and survival of Haliotis rubra in response to different algal species

Five diatom species were isolated from settlementplates at Southern Ocean Mariculture, Victoria,Australia (Navicula sp., Naviculajeffreyi, Cylindrotheca closterium, Cocconeis sp., Amphora sp.) and tested insettlement experiments with Haliotis rubralarvae. Settlement was very low on single speciesdiatom films and varied between 1%–6%. Depending onthe species combination larvae preferred to settle onfilms with mixed diatom species than single speciesfilms. The highest settlement was achieved with amixed film of Navicula sp. and Amphora sp.Five and ten-day-old germlings of Sporolithondurum induced settlement of the abalone Haliotisrubra. However, the settlement rate was significantlylower on germlings than on the whole thallus of thealga. Germlings inoculated with the diatom Navicula sp. induced higher settlement than films ofthe diatom species alone. High settlement of up to52% was also achieved with germlings of the greenalga Ulvella lens. Settlement was reduced onU. lens squares inoculated with the diatom Navicula sp. but higher than on films of the diatomalone. The settlement rate was higher if plates withU. lens were previously grazed by juvenileabalone.Post-larval growth rates were higher on monospecificdiatom films than on U. lens or on S.durum. The best growth rate was obtained with Navicula sp. U. lens and S. durum areboth good settlement inducers, but are notsufficient to support rapid growth of young H.rubra post-larvae. Survival was low on U. lensand on the diatom C. closterium. We suggest thatcommercial nursery plates seeded with U. lenswill result in high and consistent settlement, whilean inoculum with Navicula sp. will ensuresufficient food for rapid growth of the post-larvae.     

The effect of zinc on the increase in length of five species of intertidal fucales

The growth rate of Pelvetia canaliculata (L.) Dec. et Thur., Fucus spiralis L., F. vesiculosus L., F. serratus L., and Ascophyllum nodosum (L.) Le Jolis was measured in six different concentrations of zinc (0.025–14 mg/l) during a period of ten days. No significant growth response was observed at 0.1 mg/l of zinc. A significant reduction of growth during the first two to three days of exposure to 1.4 mg/l or more was followed by no or small further decline. Apices of A. nodosum from localities with different background load of zinc (0.1–0.5 mg/l and <0.01 mg/l) showed no difference in response to 2.6 mg/l of zinc.     

Carpospore early development and callus-like tissue induction of <Emphasis Type="Italic">Chrysymenia wrightii</Emphasis> (Rhodymeniaceae,Rhodophyta) under laboratory conditions

Morphological and culture studies of germlings derived from carpospores of Chrysymenia wrightii (Harvey) Yamada were carried out under various treatments combining temperature and irradiance. Basal, main, and tip branches were applied for inducing callus-like tissue. Focus was on how carpospores develop into germlings, how callus-like tissues are induced from explants, and how temperature and irradiance affect carpospore germination and discoid crust growth. Results show that carpospore development can be divided into three stages: division stage, discoid crust stage, and erect juvenile germling stage. Discoid crusts, even more than ten, might coalesce into a big discoid crust, and then developed into germlings. Filamentous fronds, formed on the rims of discoid crusts, exhibited in self-existence or co-existence form with germlings, could form spherical tufts if cultured separately. Filamentous callus-like tissues appeared on the tip branches after 13 days. PES is suitable for filament induction and culture, and filaments have potential use in germplasm preservation and vegetative propagation. Temperature (10, 15, 20, 25°C) and irradiance (8 and 36 μmol photons m⁻² s⁻¹) significantly influenced carpospore germination rate and discoid crust diameter. Carpospores germinated normally under 36 μmol photons m⁻² s⁻¹, 15~25°C, and maximum growth of discoid crusts was at 25°C, 36 μmol photons m⁻² s⁻¹; 10°C and 8 μmol photons m⁻² s⁻¹ did not favor carpospore germination or discoid crust growth.     

Seasonal changes in silicon content of diatoms estimated from the ratio of particulate silicon to diatom volume under silicon sufficiency in diatom-rich Lake Barato

To clarify the changes in Si content of diatoms, the particulate silicon (PSi) concentration and total diatom volume (TDV) were determined in Lake Barato, Japan, from April to July 1998–2000. The soluble reactive silicon (SRSi) concentration decreased markedly with the rapid increase in TDV in May and June in all three years, although the value did not fall below that at which diatom growth might be limited. The proliferation of small discoid diatoms contributed to the decrease in SRSi concentration each year. The Si content of diatoms may not be constant as indicated by the changes in PSi:TDV ratio. The low PSi:TDV ratio and the fact that PSi concentration was lower than diatom PSi concentration (calculated from the volume of diatom species) accompanying the decrease in TDV suggests the possibility of a disturbance in the silicification in May and June 1999. These parameter changes accompanying the increase in TDV suggest that the silicification did not catch up with the cell division in early April 1998, early May 1999, and mid-June 2000. In addition, the PSi:TDV ratio increased rapidly and showed large fluctuations in July 1998 and 1999. This may have been caused by a change in dominant species from small discoid diatoms to Aulacoseira granulata because of the differences in Si content per unit cell volume.     

Interactions of Patella and macroalgae with settling Semibalanus balanoides (L.)

Biological interactions affecting densities of settling and newly-settled Semibalanus balanoides (L.) have been investigated by manipulative field experiments on the Isle of Man.The effects of sweeping by fucoid clumps of different species and Patella browsing have been compared on moderately-exposed shores. Patella allowed barnacle settlement by preventing growth of competing green algae, but reduced post-settlement densities. Small clumps of Fucus spiralis L., F. vesiculosus L, and F. serratus L. all reduced settlement considerably more so than limpets. F. serratus had the greatest sweeping effect.Interactions between macroalgae and Semibalanus balanoides have been investigated at all levels on sheltered shores and low down on more exposed shores. In the Fucus spiralis and F. vesiculosus zones, post-settlement numbers were higher than in adjacent areas where the canopy was removed. Barnacles did not settle readily in the Ascophyllum zone in either experimental or control areas. Settlement occurred in the upper part of the Fucus serratus zone in experimental areas where the canopy was removed but not in control areas. No settlement occurred in either treatment or control areas lower in the F. serratus zone. At all levels on the shore fucoid canopies seemed to reduce cyprid settlement, but the effect was greatest amongst F. serratus where there was total prevention. High on the shore the effect of enhanced post-settlement survival under the canopy outweighs reduction of cyprid settlement thus there are greater numbers in the controls. Competition with red algal turfs was shown to set the lower limit of the barnacle zone on a vertical pier face.     

COLONIZATION AND GROWTH OF ASCOPHYLLUM NODOSUM (PHAEOPHYTA) IN MAINE1

Colonization and growth of Ascophyllum nodosum (L.) Le Jol. were investigated in two Maine estuaries from 1972 to 1978. Following denudation of intertidal rock, substrata were initially colonized by Fucus vesiculosus L.; eventually, Ascophyllum supplanted Fucus, and became dominant in terms of percentage cover. Ascophyllum settled first and most densely in the low intertidal zone, but its fastest growth occurred in the mid-intertidal zone. Some, but not all, Ascophyllum germlings produced a vesicle within one year of colonization. The mean annual growth of A. nodosum was variable among sites, zones and years.     

The Effect of Elevated UV Radiation on Fucus spp. (Fucales,Phaeophyta) Zygote and Embryo Development

Abstract: This study has shown that in Fucus serratus and Fucus distichus, young zygotes and embryos are highly susceptible to elevated levels of both UVA (UVAR) and UVB radiation (UVBR). Zygotes treated with UVAR are able to polarise and germinate, but are very slow to divide; if they do, they often have skewed division planes or deformed rhizoids. Those treated with UVAR and UVBR remain spherical, they do not polarise, germinate to form rhizoids or undergo cell division. We suggest that the UVR may be affecting the cytoskeleton. Conversely, zygotes and embryos of Fucus spiralis are able to withstand these same UVR levels and, at the light microscope level, appear to develop normally. When the brown algal phenolic compound phloroglucinol was placed in a filter covering the developing embryos, normal development was seen under all treatments. Phenolic compounds protect the developing fucoids from UVR. In comparison with the other two species, Fucus spiralis grows high up on the shore and is exposed for much longer periods of time and, presumably, to higher levels of natural UVR. The failure of the juvenile stages of F. serratus and F. distichus to withstand UVR stress may have implications for the continued survival of these species in the intertidal, and may prove detrimental to the population as a whole if UVR levels increase.     

The diatoms ingested by freshwater snails: temporal,spatial, and interspecific variation

Seventeen species of diatoms, representing a broad range of sizes, shapes, and growth habits, were collected from rocks in rapidly-flowing sections of the Mitchell River, North Carolina. The diatoms ingested by adult Goniobasis proxima, juvenile Leptoxis carinata, and adult Physa sp. co-occurring in this habitat were indistinguishable from one another, in spite of great differences in radular morphology. All snails sampled the diatom flora almost randomly, with only one or two of the larger diatom species under-represented in the gut contents. Some snails also seemed to selectively ingest the smaller individuals of the larger diatom taxa, and larger individuals of the smaller diatom taxa. The diatoms identifiable in juvenile Goniobasis guts were somewhat more distinctive, although this seemed to be due at least partly to more mechanical breakage. The diatom flora of quiet, muddy pools was much different from that of shallow, rocky areas, but once again, Goniobasis seemed to sample the available flora randomly. Seasonal variation was also apparent in the diatom diet of Goniobasis. We suggest that in some cases, it may be reasonable to use snails to sample the diatom assemblage present in a particular habitat, if more direct methods are impractical.     

SEASONALITY OF MICROBIAL FOULING ON ASCOPHYLLUM NODOSUM (L.) LEJOL., FUCUS VESICULOSUS L., POLYSIPHONIA LANOSA (L.) TANDY AND CHONDRUS CRISPUS STACKH1

The nature and seasonal extent of microbial fouling on Ascophyllum nodosum (L.) LeJol., Fucus vesiculosus L., Polysiphonia lanosa (L.) Tandy and Chondrus crispus Stackh. were observed by scanning electron microscopy. Bacterial filaments and smaller rod and coccoid forms dominated the fouling communities on all species, with pennate diatoms constituting a minor component in contrast to results with plastic substrates on which pennate diatoms dominated and preceded bacterial colonization. The total percent microbial coverage on the surfaces of all four seaweed species was determined by monthly stereological analyses of representative composite micrographs. These showed a simultaneous decline between April and May which could represent the die-off of the cold water bacterial flora when water temperature increased past the threshold for obligate psychrophiles. Microbial colonization patterns were directly correlated (P = 0.005) with maximum coverage in April and November–December and reduced levels from May to October. Significant inverse (P < 0.041) correlations between total percent coverage and water temperature indicate distinct seasonal cycles, however, the patterns of dominance by filamentous bacteria and rod and coccoid forms were markedly different. Total coverage patterns of both rhodophytes showed no apparent seasonal cycle and were not related to water temperature. Rod and coccoid bacteria were apparently suppressed year round on P. lanosa relative to the other species. These interspecific differences in seasonal fouling patterns are discussed in light of possible modes of regulation, especially algal antibiosis.     

D-VITAMINS AND THEIR PRECURSORS AS GROWTH REGULATORS IN AXENICALLY CULTIVATED MARINE MACROALGAE1

Light petroleum extracts from thalli of Fucus vesiculosus (L.) and Laminaria digitata (Huds.) Lamour. stimulate growth of axenically cultured Fucus spiralis L. and Enteromorpha compressa (L.) Grev. in amounts per litre culture medium corresponding to one to 10 mg wet weight of the brown algae. This indicates the presence of a lipophilic, very active growth regulator in the blades. Extracts from the meristematic zone of Laminaria are much less active. Vitamins D₃, D₂ and 1,25-(OH)₂-D₃ at 1·10^?8– 1·10^?7 M increase the fresh weight of Enteromorpha compressa and Fucus spiralis up to 100% or more. The dry weight of Nemalion helminthoides (Veil, in With.) Batt. is also increased under the same conditions. Ergosterol and cholesterol are also growth-stimulating but higher concentrations (e.g. 1·10^?6 M) are necessary. The effects of the D vitamins manifest themselves without any auxins being added to the medium.     

Longitudinal profiles of carbon dioxide fixation capacities in marine macroalgae

Fucus serratus L., Fucus spiralis L., and Fucus vesiculosus L. (Fucales, Phaeophyceae) as well as Laminaria digitata (Huds.) Lamour., Laminaria hyperborea (Gunn.) Fosl., and Laminaria saccharina (L.) Lamour. (Laminariales, Phaeophyceae) have been investigated for the distribution of enzymic CO₂ fixation capacities via phosphoenolpyruvate carboxykinase (EC 4.1.1.32) (PEP-CK) and via ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) (RubP-C) in different regions of the thalli. The maximum of PEP-CK activity is found to be confined to the growing regions of the algae, while the activity of RubP-C achieves its highest values in the entirely differentiated parts of the fronds. These findings are confirmed by the results of photosynthetic and light-independent (dark) carbon assimilation as determined by in vivo ¹⁴CO₂ fixation. The physiological significance of these differential patterns of carboxylation patterns is discussed with respect to the ontogenetic stage and the chemical constitution of the different thallus parts.     

Natural chelators in sea water: Detoxification of Zn2+ by brown algal polyphenols

We have examined growth responses of several species of marine phytoplankton, cultured with and without heavy metal stress, to supplements of polymeric polyphenols from the brown algae Ascophyllum nodosum (L.) Le Jol. and Fucus vesiculosus L. In the absence of additional heavy metals. supplements of up to 4000 μg. 1 of polyphenols had a small effect on initial growth rates for three of these microalgae and had no effect on maximum cell densities for four species. One very common, heavy metal-sensitive diatom, Skeletonema costatum (Grev.) Cl., showed significantly increasing maximum cell density in the cultures, with increasing addition of polyphenols to the medium. The toxicity of Zn²⁺ (0.5–2.0 mg·1 ¹) to cultures of the diatom Phaeodactylum tricornutum Bohlin was relieved by supplements (100–200 μg·1 ¹) of the brown algal polyphenols. Exudation of these polyphenols from brown seaweeds may contribute to the natural chelating capacity of inshore sea water.     

Recovery and development of parasexual fusion products inEnteromorpha linza (L.) J. Ag. (Ulvales,Chlorophyta)

Newly released zoospores fromEnteromorpha linza (L.) J. Ag. lack significant cellulose cell wall material and are suitable for treatment as protoplasts in a parasexual fusion process using high pH-Ca^{+ +}, PEG and centrifugation. Treated zoospores settled on glass cover slips within 3 h and were examined microscopically at 1000 ×. Presumptive fusion products were identified by their larger size and presence of twin chloroplasts and eyespots. Unfused zoospores adjacent to fusion cells were killed by 2–3 min exposure to blue light (410–490 nm) from a high pressure mercury illuminator. Unexposed fusion cells developed into uniseriate germlings within 10 days at which stage they could be readily identified at 60 × with a dissecting microscope and isolated by micropipette. Ten-day germlings from both unfused zoospores and fusion cells were stained with the DNA-localizing fluorochrome hydroethidine and relative nuclear DNA content determined with epi-(incident) UV illumination. All germlings were found to be uninucleate. Germlings from unfused zoospores had haploid nuclei with 1N = 10 and 1C and 2C levels of DNA, while germlings from fusion cells had diploid nuclei with 2N = 20 and 2C and 4C levels of DNA. These result are interpreted as evidence of karyogamy following parasexual zoospore fusions. Isolated diploid germlings, cultured for 10 weeks were found to conserve their 2N chromosome complements and elevated levels of nuclear DNA. Although most diploid germlings were morphologically similar to haploid control plants, some exhibited ‘gigas’ characteristics, including larger cells, chloroplasts, and nuclei. These results are discussed in terms of unique phenotypes that result when nuclear and organellar genes are combined in different ways.