Alkanes and alkenes in marine benthic algae

Saturated and olefinic hydrocarbons were determined in additional species of benthic marine algae from the Cape Cod (Massachusetts, USA) area (see: Youngblood et al., 1971). The distribution of homologous and isomeric olefins was studied in plants of different age and in morphologically different parts of the same specimen. With two minor exceptions, only normal alkanes and alkenes are present. The methylene-interrupted C₁₉- and C₂₁-polyolefins are particularly abundant; ¹-heneicosahexaene and the corresponding pentaene are common to all brown algae, while the corresponding ³-isomers occur in green algae. The hydrocarbon concentration, the alkene-to-alkane ratio and the polyolefin content are highest in young plants or in rapidly growing tissues of older plants. This suggests a deeper involvement in cell biochemistry of straight-chain hydrocarbons than previously considered. The biosynthesis of the plant polyolefins remains to be explored; no immediately obvious precursors of the ¹-polyolefins were found among the algal fatty acids. The hydrocarbon composition of these benthic algae differs greatly from that of fossil fuels in its simplicity and predominately unsaturated nature. The separation of the isomers by gas chromatography and their structural elucidation by mass spectrometry, alone and in combination with hydrogenation and ozonolysis, are discussed.Contribution No. 3155 of the Woods Hole Oceanographic Institution.     

Hydrocarbons in benthic algae from the eastern Gulf of Mexico

Thirty-six samples of benthic algae were collected from the continental shelf along the eastern Gulf of Mexico. The algae contain 82.8±143 ppm aliphatic hydrocarbons by dry weight and 11.8±22.7 ppm aromatic and polyolefinic hydrocarbons. The aliphatic constituents of red algae are composed almost entirely of n-C₁₇ (70 to 95% of aliphatic weight); green algae have varying amounts of n-alkanes in the range of n-C₁₅ to n-C₁₉, with homologous series of odd carbon number n-alkenes. Phytadienes occur as high as 740 ppm dry weight in the green algae and do not appear to be related to time or place of collection or to presence of any petroleum pollutants. The single brown algal species contains predominantly n-C₁₅. About 30% of the samples have hydrocarbons resembling degraded petroleum residues, but no petroleum pollution of recent origin was detected in any specimen.     

Hydrocarbons of marine phytoplankton

The hydrocarbon contents of 23 species of algae (22 marine planktonic), belonging to 9 algal classes, were analyzed. The highly unsaturated 3,6,9,12,15,18-heneicosalhexaene predominates in the Bacillariophyceae, Dinophyceae, Cryptophyceae, Haptophyceae and Euglenophyceae. Rhizosolenia setigera contains n-heneicosane, presumably derived from the hexaolefin by hydrogenation. Two isomeric heptadecenes have been isolated: the double bond is located in 5-position in the bluegreen alga Synechococcus bacillaris and in 7-position in 2 green algae. Our complete analyses are discussed in the context of earlier data; some generalizations appear no longer valid. Hydrocarbon analysis of marine algae should provide a tool for the investigation of the dynamics of the marine food chain. Knowledge now available provides the background needed for distinguishing between hydrocarbons of recent biogenic origin and hydrocarbon pollutants from fossil fuels.Contribution No. 2576 of the Woods Hole Oceanographic Institution, USA.     

Petroderma and Porterinema in the New World

Two species of crustose brown algae (Ralfsiaceae) are discussed. Petroderma maculiforme is described from several Massachusetts collections and Porterinema fluviatile is described for the first time from North America. The known world-wide distribution of both species is cited, and data are given on their ecology, and vegetative and reproductive periodicities. The wide range of salinities from which plants of these species have been collected indicate that they are both euryhaline. Lithoderma lignicola Kjeliman (1883) is synonymized with Petroderma maculiforme (Wollny) Kuckuck (1897) and Porterinema marina Jaasund (1965) with. P. fluviatile (Porter) Waern (1952).Systematics-Ecology Program Contribution No. 196.     

Emerson enhancement effect and quantum yield of photosynthesis for marine macroalgae in simulated underwater light fields

The contribution of enhancement to the total photosynthesis of marine macroalgae in their natural habitats was estimated by comparing the photosynthesis measured by O₂-electrode in five broad-band light fields with that predicted (on the assumption that no enhancement was occurring) from the photosynthetic action spectrum of each plant and the spectral distribution of the light fields. The excess of measured values divided by calculated values provided a measure of enhancement. Although 37% enhancement was observed for red algae in unfiltered quartz-iodine light, and 18% for green and brown algae, substantially lower values were obtained for all species in more natural light fields. In those typical of shallow coastal waters, phycoerythrin-rich red algae exhibited 15 to 20% enhancement, but little enhancement (<5%) was detected in other algae. In a green light field, representing deep coastal water, there was no significant enhancement in any species, and only green and brown algae showed any enhancement (ca 8%) in broad-band blue light, similar to that in deep oceanic waters. Quantum yields of 0.09 to 0.10 O₂ molecules per absorbed photon were recorded in most light fields for green and brown algae with thin thalli, but yields decreased in the blue light field and in species with thicker thalli. All red algae had quantum yields of about 0.08 O₂ molecules per absorbed photon, except in the blue light field, in which quantum yields were reduced by 70%.     

Benthic macrophyte communities in the Ghar El Melh lagoon (North Tunisia)

This work deals with the biodiversity and distribution of benthic macrophytes in the Ghar El Melh lagoon, a Mediterranean coastal lagoon located in the North of Tunisia. An inventory was made of the benthic flora and submerged macrophyte communities were mapped during two successive campaigns (the summer of 1999 and the winter of 2000). The following 24 macrophyte species were identified: seven red algae, two brown algae, 11 green algae, and four marine angiosperms. The results were compared with available data from the literature. Ruppia cirrhosa is the most dominant species. It is found in all lagoon parts, except in the west sector.Ruppia beds are usually associated withCladophora forming heterogeneous communities. During summerRuppia cirrhosa shows a large distribution, covering an area of ca. 21.4 km², with dense, extensive beds covering 80–100%. In winter, severalCladophora species have a very large distribution as well, covering nearly an area of 28.5 km² with an average cover of 46%. The green algaeCaulerpa prolifera is confined to the eastern part of the lagoon which is mainly affected by seawater. In comparison with previous situations, many transformations were observed in biodiversity and spatial distribution of the dominant communities. Thus,Cymodocea nodosa andZostera beds, which dominated in the 1970s, were replaced byZostera andCaulerpa prolifera in the 1980s and are currently succeeded byRuppia cirrhosa andCladophora. Restoration of the Ghar El Melh lagoon will enable an increase in the exchange with the open sea and the circulation of water, in particular in the confined zones. This should considerably improve the water quality and would positively influence the phytobenthic communities.     

A polyunsaturated hydrocarbon (3, 6, 9, 12, 15, 18-heneicosahexaene) in the marine food web

An olefinic hydrocarbon (all-cis-3,6,9,12,15,18-heneicosahexaene, HEH)_was islated from marine planktonic plants and animals. Its structure was established by ultraviolet, infrared and mass spectrometry in combination with chemical techniques. The olefin occurs in many species of marine planktonic algae and is probably derived from the corresponding docosahexaenoic acid. Rhincalanus nasutus accumulates HEH nonselectively from its algal food together with the triglyceride lipids. Other, related copepods contain little or no HEH even when grown in cultures of algae that provide R. nasutus with that olefin. The presence of HEH in marine vertebrates suggests that, within the animal lipids, hydrocarbons are remarkably stable. This work has practical implications for studies of the marine food web and of marine pollution with persistent chemicals.Contribution No. 2485 of the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, USA.     

Foods and predators of the green sea urchin Strongylocentrotus droebachiensis in Newfoundland waters

Gut analyses of the green sea urchin Strongylocentrotus droebachiensis (O. F. Müller) demonstrated that perennial phaeophytes, mostly fucoids and Alaria esculenta, were predominant in the diet. Ephemeral species, coralline algae and animals, were consumed in smaller amounts when available. Grazing by the urchins is evidently responsible for the dearth of non-coralline sublittoral algae in Newfoundland waters. Lobsters, rock crabs, purple sea stars, other urchins, and a variety of fishes and birds feed on S. droebachiensis, but predation is apparently not effective in limiting the abundance of the urchin.Studies in Biology from the Memorial University of Newfoundland No. 234.Contribution from the Marine Sciences Research Laboratory No. 66.     

Respiration and nitrogen excretion by the squid Loligo forbesi

The amphipod Corophium volutator (Pallas) of the Bay of Fundy is a major food resource for migrating shorebirds. A relatively low total lipid content (ca 1.7%) was found in summer and fall samples of 1986. Analysis of triacylglycerides which would normally reflect an animal diet through fatty acid input, revealed high proportions of fatty acids characteristic of algae. C. volutator contains hydrocarbons (0.2% of total lipids) with a unimodal distribution in the range C₁₈ to C₃₅ with maximum at C₂₅, and a weak odd-carbon preference (carbon preference index, C.P.I. =1.22). This hydrocarbon pattern shown by C. volutator seems to be biogenic, but due to both a weak odd-carbon preference and a significant content of the isoprenoid phytane, the presence of petroleum hydrocarbons cannot be ruled out. Comparison of the hydrocarbon composition be ruled out. Comparison of the hydrocarbon composition found in C. volutator with hydrocarbons described in the literature for its most likely sources of food suggests that highly degraded material and aged detritus are components of the diet of this deposit feeder, and as important as the local diatom population. No organohalogen pesticides (DDT, dichlorodiphenyl-trichloroethane, or derivatives) or polychlorinated biphenyls (PCBs) were detected in the lipid extracts of C. volutator. Absence of these materials and low hydrocarbon levels explain the continued successful exploitation of this food resource by migrating shorebirds.     

Phylogeography of the seaweed Ishige okamurae (Phaeophyceae): evidence for glacial refugia in the northwest Pacific region

Although benthic marine algae are essential components of marine coastal systems that have been influenced profoundly by past and present climate change, our knowledge of seaweed phylogeography is limited. The brown alga Ishige okamurae Yendo occurs in the northwest Pacific, where it occupies a characteristic belt in the exposed intertidal zone. To understand the patterns of genetic diversity and the evolutionary history of this species, we analyzed mitochondrial cox3 from 14 populations (221 individuals) throughout its range. The 17 haplotypes found in this study formed five distinct clades, indicating significant genetic structure. The high differentiation and number of unique (private) haplotypes may result from the recolonization of the species from glacial refugia. Three putative refugia, each with high genetic diversity, were identified: southern Korea (including Jeju Island), northern Taiwan, and central Pacific Japan. Recolonization of I. okamurae was probably determined by ocean currents and changes in sea level during the last glacial period.     

Biogeochemistry of aromatic hydrocarbons in the benthos of microcosms†

Investigations into the fate of petroleum compounds in the marine environment were carried out using experimental microcosms of two sizes and designs. Aromatic hydrocarbons or No. 2 fuel oil were spiked to the water of a 13 m³ continuous flow system and to a 2281 recycled flow system. The transport and alteration of this oil was traced in the sediment and benthic organisms (Glycera americana, Crepidula sp., and Nephtys incisa) of these microcosms. Measurable contamination was found in both sample types. The aromatic hydrocarbon distribution, including relative isomeric distribution (e.g., C₂‐phenanthrenes) was found to be different in sediment and in organisms from that which was originally introduced to the experimental microcosm. Differences in isomer distribution between Glycera and Crepidula were also detected. Based on the experimental data: molecular weight and specific isomeric form, biochemical processes, solubility, and particle adsorption/desorption influence the fate of petroleum compounds in benthic ecosystems.     

Occurrence and biological role of light-induced chromatophore displacements in seaweeds

The occurrence of light-induced chromatophore displacements and concomitant transmittance changes in marine algae was investigated by microscope and photometrically with an automated recording microphotometer system; 16 brown, 6 green and 20 red algae were studied. In most of the brown algae, both phaeoplast displacements and transmittance changes were found. In some red algae which are frequently exposed to direct sun light during emergence at low tide, light-induced transmittance changes were measured, but they could not unequivocally be correlated with changes in the position of rhodoplasts. Among green algae, only Ulva lactuca shows chloroplast displacements which, however, follow circadian rhythms and are consequently not light-induced in the strict sense. The dose-response curves of light-induced chromatophore displacements were measured in the following Fucus and Laminaria species: F. spiralis, F. vesiculosus, F. serratus, L. digitata, L. saccharina and L. hyperborea. While in Fucus species correlations between light-induced transmittance changes and zonation of the intertidal area seem to exist, no significant differences have been found in the Laminaria species. The physiological role and ecological importance of light-induced chromatophore displacements for seaweeds living in the intertidal belt are discussed.     

A pelagic bryozoan from Antarctica

Spherical or sub-spherical bryozoan colonies were collected from the surface waters of a coastal polynya in the southeastern Weddell Sea near Halley Station in February 1992. These are the first truly pelagic marine bryozoan colones yet recorded. The collection site is the edge of the Brunt ice shelf, which is between 150 and 250 m thick in this area, and the depth of water to the seabed is -400 m. The colonies were hollow, composed of a single layer of autozooids, and appeared complete and undamaged. They were between 5.0 and 23.0 mm in diameter, were brownish in colour in life, and pale yellowish brown after preservation. Light and scanning electron microscope investigations of the colonies indicated that they belonged to the genus Alcyonidium, and they are here compared with A. flabelliforme Kirkpatrick, a known antarctic benthic species. Both a pelagic existence and hollow spherical colony form are new attributes for the phylum Bryozoa. However, because of the plasticity of form of species belonging to the genus Alcyonidium, these is not enough evidence for the introduction of a new species at this time. The colonies found may represent a previously undescribed juvenile stage of a known Alcyonidium species. It is postulated that these colonies may obtain nutrition from the often abundant populations of ice algae present in the lower layers of permanent sea-ice.     

Relative growth of different species of marine algae in wasterwater-seawater mixtures

In recent studies, we developed a combined nutrient removal-marine aquaculture process for the tertiary treatment of wastewater and the production of commercially important shellfish. Part of this process consists of an outdoor mass cultivation system for marine algae. During our previous experiments we observed that marine diatoms almost exclusively are the dominant algal species in our outdoor cultures. To better understand this phenomenon of diatom dominance we grew 16 species of marine algae in continuous monoculture under laboratory conditions simulating to some degree the conditions prevailing in our outdoor experiments. Species such as Skeletonema costatum, Monochrysis lutheri and Tetraselmis sp., which were never dominant in our outdoor cultures, grew as well in monoculture, as Phaeodactylum tricornutum, frequently, the prevalent species outdoors. However, when monocultures of Dunaliella tertiolecta and Thalassiosira pseudonana (3H) were purposely contaminated with P. tricornutum, the latter species quickly became dominant. It is suggested that a complex interaction of environmental factors is usually responsible for the dominance of a particular species; one such factor may be the nitrogen source in the growth media. Under conditions of virtually, complete nitrogen assimilation, the carbon: nitrogen ratio in the algae was high (7 to 8) when NO ₃ ^- –N was the source of nitrogen, and low (4 to 6) when NH ₄ ⁺ –N was the prime form of nitrogen. When algal growth was low, resulting in a large inorganic nitrogen residue, the carbon:nitrogen ratio was low regardless of whether NO ₃ ^- –N or NH ₄ ⁺ –N was the main nitrogen source.Contribution No. 3297 from the Woods Hole Oceanographic Institution.     

Action spectra and spectral quantum yield of photosynthesis in marine macroalgae with thin and thick thalli

Net photosynthesis at 10mol photons m^-2 s^-1 in each of 24 wavelengths was measured in absolute units by an O₂-electrode and corrected for dark respiration to construct action spectra for gross photosynthesis in nine species of algae, which included plants with thin and thick thalli from each of four major pigment groups. The photosynthesis of green and brown algae with thin thalli decreased in green light, but species with thick thalli from these two groups had action spectra which were almost flat, and matched the optical blackness of the thalli but did not reflect the pigment differences between the species. Among the red algae, on the other hand, there was little difference between the action spectra for thin and thick algae. Only wavelengths absorbed by the phycobilin pigments were effective in photosynthesis, even in species (e.g. Chondrus, Phyllophora) which absorbed all visible wavelengths strongly. Maximal quantum yields of 0.10 to 0.12 O₂ molecules per absorbed photon were recorded for thin green and brown algae, but thicker algae in these two groups had lower values. Red algae exhibited maximal values close to 0.10 O₂ molecules per absorbed photon, irrespective of thallus thickness or phycocyanin content, but the quantum yields of phycoerythrin-rich species in the 600 to 650 nm waveband were lower than those of phycocyanin-rich species.     

Investigation of the sources of aliphatic hydrocarbons in the mussel Mytilus edulis from north sea oil production platforms by Capillary glc and CGCMS†

Aliphatic hydrocarbons isolated from mussels collected over a 20 month period from three North Sea Forties field oil production platforms have been examined by capillary gas chromatography and gas chromatography‐mass spectrometry.

The biological hydrocarbons consist predominantly (300–700 μh g^‐1 lipid) of C₃₁ and C₃₃ n‐alkenes with 2, 3 and 4 double bonds probably derived from a dietary intake of Emiliania huxleyi, a microscopic coccolithophorid alga. In some of the mussels C₁₈, C₂₀ and C₂₂ n‐alkanes are present in unusually high abundance compared to their odd carbon number homologues. Possible reasons for this distribution are discussed.

The concentrations of fossil fuel hydrocarbons (mainly unresolved complex mixtures) in the mussels ranged from 330 to 5,298 μg g^‐1 of lipid. No significant increase in values was detected two months after the start of discharge of treated Forties production water from the Forties D platform.

To determine the pollutant hydrocarbon sources, the sterane and terpane distributions of hydrocarbon fractions isolated from the mussels and from a number of fossil fuels, including Forties crude oil, were examined by mass fragmentography. The steranes in the mussels from the Forties C and D platforms contained higher proportions of regular 14α(H), 17α(H) components than Forties oil in which the steranes were mainly 13ß(H), 17α(H) diasteranes. In addition, the terpane distributions of the mussel fractions differed markedly from that of Forties crude oil, particularly in the relative abundance of diterpanes to triterpanes, which was higher in the mussel fractions than in the oil, and in the absence of 17α(H), 18α(H), 21ß(H)‐18, 30‐bisnorhopane, a known component of Forties crude. Furthermore, the ratio of C₂₉ to C₃₀ triterpanes was greater than unity in the mussels but much less than unity in the Forties oil. These data preclude Forties or other similar North Sea crudes as the major source of pollutant aliphatic hydrocarbons in the mussels. The relative concentrations and distributions of steranes and triterpanes suggest that the most likely source of pollutants is a Middle Eastern based oil derived either from rig activities, or from background pollution in the North Sea. The low concentration of ≥ C₂₇ steranes in the more polluted mussels (e.g. Forties B; UCM > 200 ppm dry wt.) suggests that gas oils used on the platforms may be the major source of petrogenic hydrocarbons in these samples.     

Light quality in relation to growth,photosynthetic rates and carbon metabolism in two species of marine plankton algae

The effect of light quality on growth, photosynthesis and carbon metabolism in two species of marine algae,Cyclotella nana (Hustedt) andDunaliella tertiolecta (Butcher), was examined. Relative growth constants forC. nana were 0.37, 0.29 and 0.25 in blue, white and green light, respectively. Corresponding constants were 0.41, 0.31 and 0.29 forD. tertiolecta. Photosynthetic rates in both species were higher in blue light and lower in green light compared with white light of the same intensity. More than 60% of¹⁴C assimilated byC. nana orD. tertiolecta grown in blue or green light was incorporated into the ethanol-insoluble fraction, compared with 10 to 30% in this fraction in white light. The relative importance of the various components within this fraction was independent of light quality. Although less¹⁴C was assimilated into the ethanol-soluble fraction in blue or green light, there was a relative increase in some amino acids and organic acids in this fraction and a decrease in sugars and sugar phosphates relative to white light of the same intensity. These differences were independent of light intensity, photosynthetic rate and cell density in the cultures.     

(<Emphasis Type="Italic">Z</Emphasis>)-9-Pentacosene − contact sex pheromone of the locust borer, <Emphasis Type="Italic">Megacyllene robiniae</Emphasis>

Summary. Male locust borers, Megacyllene robiniae (Förster), responded to females only after contacting them with their antennae, indicating that mate recognition was mediated by a contact sex pheromone. GC-MS analyses of whole-body extracts of males and females determined that the profiles of compounds in the extracts were qualitatively similar, but differed considerably in the ratios of compounds between sexes. Biological activities of reconstructed blends of the most abundant straight-chain (nC₂₃, nC₂₄, nC₂₅, nC₂₆), methyl-branched (3me-C₂₃, 3me-C₂₅), and unsaturated (Z9:C₂₃, Z9:C₂₅, Z9:C₂₇ compounds in extracts from females were assessed in arena bioassays, assessing four distinct steps in the mating behavior sequence of males (orientation, arrestment, body alignment, mounting and attempting to couple the genitalia). Males were unresponsive to freeze-killed, solventwashed females treated with blends of straight-chain and methyl-branched alkanes, but responded strongly to females treated with the blend of alkenes. Further trials determined that the complete sequence of mating behaviors, up to and including coupling the genitalia, was elicited by Z9:C₂₅ alone. Z9:C₂₅ comprised 16.4 ± 1.3% of the total hydrocarbons in whole-body hexane extracts of females and was co-dominant with two other hydrocarbons that were not active. In contrast, in solid phase microextraction (SPME) wipe samples from several areas of the cuticle, Z9:C₂₅ appeared as the single dominant peak, comprising 34.6 – 37.8% of the sampled hydrocarbons. Our data indicate that Z9:C₂₅ is a contact sex pheromone of M. robiniae, being the most abundant hydrocarbon on the surface of the cuticular wax layer of females where it is readily accessible to the antennae of males.     

Release of dissolved organic matter by some marine macrophytes

The amount of photoassimilated carbon released as dissolved organic carbon was investigated for 5 species of benthic marine macrophytes (Thalassia testudinum, Diplanthera wrightii, Acanthophora spicifera, Chondria dasyphylla, and Dictyota dichotoma) and 1 species of pelagic marine macrophyte (Sargassum natans). Release rates ranged between 0.006 and 0.053 mg C g^-1 h^-1. Percent release values ranged from 1.1 to 3.8%. Spermatophytes had slightly lower percent release values than algae. Investigations performed on the ability of release products to be utilized by heterotrophic organisms showed that 20 to 30% of the released organic carbon was assimilated within a 2-h period.     

Ecological investigations of the marine nematode Metoncholaimus scissus

Significant concentrations of the large oncholaimid Metoncholaimus scissus Wieser and Hopper (1967) (>7000 animals/24 cm² area or 2.68x10⁶ specimens/m²) have been noted in soft-bottom turtle grass communities. Activities of the nematode frequently show an indirect relationship to blooms of the benthic diatom Pleurosigma balticum. Analysis of biomass data for M. scissus (as much as 28 g wet weight/m²) demonstrate the importance of microsite activity and sensitivity of the species to small significant changes in the benthic environment.This work was supported by grant GM 12824 from the National Institutes of Health to the Institute of Marine and Atmospheric Sciences, University of Miami. Contribution No. 1188 from the Institute of Marine and Atmospheric Sciences, University of Miami and from the Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa.