LIFE HISTORY OF COLPOMENIA SINUOSA (SYCTOSIPHONACEAE,PHAEOPHYCEAE) IN THE AZORES1

Colpomenia sinuosa (Mertens ex Roth) Derbès and Solier (Scytosiphonaceae, Phaeophyceae) is a common species on the rocky intertidal shores of the Azores, where reproductive gametophytes occur throughout the year. Life‐history studies of this species were carried out in culture, and both sexual and asexual reproduction were observed. Anisogamous gametes fused to form zygotes. The zygotes gave rise to a filamentous prostrate sporophyte generation bearing unilocular sporangia, under both short‐day and long‐day conditions at 15 and 22°C, and to both unilocular and plurilocular sporangia, under the lower temperature condition. Unispores developed into gametophytes, and plurispores gave rise to filamentous sporophytes. Asexual reproduction was carried out by unfused female gametes and asexual plurispores produced from the same gametophyte. Unfused gametes developed into filamentous prostrate sporophytes producing unilocular sporangia in both culture conditions, and unispores released from the sporangia gave rise to gametophytes. Asexual plurispores from field gametophytes, under both culture conditions, developed directly into new gametophytes. The species exhibited three types of life history: a heteromorphic, diplohaplontic; a heteromorphic, monophasic (both with alternation between the erect and filamentous prostrate thalli); and a monomorphic, monophasic.     

Morphology and life history of Scytosiphon canaliculatus comb. nov. (Scytosiphonales,Phaeophyceae) from Japan*

Morphology of field material and life history in culture were studied in Scytosiphon canaliculatus (Setchell et Gardner) comb. nov. from northern Japan. Erect gametophytes of S. canaliculatus are cylindrical, tubular, up to 7 mm wide and 40 cm long, and without regular constrictions. S. canaliculatus has pronounced anisogamy and ascocysts accompanied with plurilocular gametangia. The life history of S. canaliculatus showed an alternation between erect gametophytes and crustose prostrate sporophytes bearing unilocular sporangia. Since field sporophytes of S. canaliculatus were found to be identical with Hapterophycus canaliculatus Setchell et Gardner (Ralfsiaceae, Phaeophyceae), it is proposed to transfer H. canaliculatus to the genus Scytosiphon. In the field, gametophytes with plurilocular gametangia appeared in spring and disappeared in summer. Sporophytes with unilocular sporangia were collected in late autumn and winter. Unilocular sporangia were produced at 15°C in short-day culture conditions and unispores developed into erect gametophytes at 5–15°C. It is suggested that the seasonal Occurrence of gametophytes in the field is due to the seasonal formation of unilocular sporangia, which is regulated by temperature and photoperiod.     

A NEW MODE OF LIFE HISTORY IN THE FRESHWATER RED ALGAL GENUS BATRACHOSPERMUM

In a new type of Batrachospermum life history, the Batrachospermum phase does not originate from the erect filaments of a Pseudochantransia but directly from a prostrate crustose growth, the prothalle of Sirodot. It is probable that the few lowermost segments of those Batrachospermum cladomes are diploid, the remainder due to an interposed “somatic” meiosis are haploid.     

Cultivation of the red alga Chondracanthus chamissoi: sexual reproduction and seedling production in culture under controlled conditions

The red alga Chondracanthus chamissoi (Gigartinales) is endemic to the southern-central region of South America. In the Pacific Ocean, it is distributed from north-central Peru to Chiloe Island. This species is of economic importance because it is edible and used for carrageenan production. The tetrasporophyte phase was grown in the laboratory, obtaining male and female gametophytes that were incubated under different photoperiod, pH, salinity and temperature conditions. These gametophytes developed and generated reproductive structures that led to in vitro maturation. Subsequently, fertilisation occurred and formation of cystocarps was observed. Finally, carpospores were released and the formation of sporophytes completed the life history of this species under laboratory conditions. Reproductive phase growth rates were recorded for each of the different culture conditions used. Sporophytes reached the highest daily growth rate (22%), while gametophyte’s daily growth rate was slower (9%). This research confirms, in vitro, the assumption that C. chamissoi has a sexual triphasic life history Polysiphonia type with isomorphic gametophytes and tetrasporophytes. The development of the complete life history took 20 months in the laboratory.     

ON THE LIFE CYCLE,CYTOLOGY AND TAXONOMY OF CLADOPHORA CALLICOMA FROM INDIA

The present investigation deals with the cultural observations on the morphology, reproduction, life cycle, cytology and taxonomy of a freshwater Cladophora, C. callicoma (L.) Kütz, from India. There is a regular isomorphic alternation of generations between quadriflagellate zoospore-producing diploid plants and biflagellate isogamete-producing haploid plants, coupled with heterothallism in the latter. Chromosome numbers of n = 12 and 2n = 24 were determined respectively for gametophytes and sporophytes. Twelve bivalents were counted in meiosis where chiasmata varied from one to three. In light of above observations, the affinity of C. callicoma with C. glomerata, to which the former has often been assigned, has been discussed. It is proposed that the C. glomerata complex, which consists of intraspecific polyploid races, might have two distinct lines of evolution with regard to life cycle; one with forms having higher ploidy levels and lacking an alternation of generations, and the other with forms having low ploidy levels and alternation of generations.     

Sulla presenza nell'Italia meridionale di alcune alghe marine bentoniche rare per il Mediterraneo

Abstract

On the occurrence in Southern Italy of some benthic marine algae rare to the Mediterranean Sea. - The Authors report the occurrence along the coast of Southern Italy of the following species that resulted rare to the Mediterranean Sea: Antithamnion piliferum Cormaci et Furnari; Bonnemaisonia asparagoides (Woodward) C. Agardh (tetrasporophyte); Corynophlaea flaccida Kuetzing; Fosliella farinosa (Lamouroux) Howe: of this very common species, the occurrence of thalli with propagules is reported; Hypnea cervicornis J. Agardh; Jania adherens Lamouroux, new to Italy; Mesogloia lanosa P.L. et H.M. Crouan, new to Italy; Pneophyllum confervicolum (Kuetzing) Chamberlain f. minutulum (Foslie) Chamberlain, new to the Mediterranean Sea; Polystrata fosliei (Weber van Bosse) Denizot, new to Italy; Rhodymenia delicatula Dangeard; Spermothamnion johannis G. Feldmann-Mazoyer: of this common species, the occurrence of bisporocysts in monoic gametophytes is reported; Vickersia baccata (J. Agardh) Karsakoff.     

On the ecology of Bonnemaisonia hamifera in its preferred habitats on the Western coast of Brittany (France)

Sexual reproduction of Bonnemaisonia hamifera Hariot is the rule on the western coast of Brittany. Each spring the alga is luxuriant in the Molène Islands, indicating that optimum conditions for growth occur there. These conditions, which include a sandy substrate and frequently renewed clear water, seem to be essential for the maturation of the male and female organs of Bonnemaisonia hamifera on the coasts of Brittany.     

Alternate life history phases of a common seaweed have distinct microbial surface communities

Macroalgal life histories are complex, often involving the alternation of distinct free‐living life history phases that differ in morphology, longevity and ploidy. The surfaces of marine macroalgae support diverse microbial biofilms, yet the degree of microbial variation between alternate phases is unknown. We quantified bacterial (16S rRNA gene) and microeukaryote (18S rRNA gene) communities on the surface of the common intertidal seaweed, Mastocarpus spp., which alternates between gametophyte (foliose, haploid) and sporophyte (encrusting, diploid) life history phases. A large portion (97%) of bacterial taxa on the surface Mastocarpus was also present in samples from the environment, indicating that macroalgal surface communities are largely assembled from the surrounding seawater. Still, changes in the relative abundance of bacterial taxa result in significantly different communities on alternate Mastocarpus life history phases, rocky substrate and seawater at all intertidal elevations. For microeukaryote assemblages, only high intertidal samples had significant differences between life history phases although sporophytes were not different from the rocky substrate at this elevation; gametophytes and sporophytes did not differ in microeukaryote communities in the mid and low zones. By sequencing three host genes, we identified three cryptic species of Mastocarpus in our data set, which co‐occur in the mid‐to‐low intertidal zone. In these samples, M. alaskensis sporophytes harboured distinct bacterial communities compared to M. agardhii and M. intermedius sporophytes, which were not distinguishable. Conversely, microeukaryote communities did not differ among species.     

The life history of Porphyra endiviifolium from the South Shetland Islands, Antarctica

This paper describes the reproduction and life history of an intertidal species, Porphyra endiviifolium, from Antarctica. Field specimens were examined microscopically, prepared for electron microscopy and used to establish cultures. Wild populations comprised two kinds of leafy thalli, morphologically similar but distinguished by their mode of reproduction, either sexual or asexual. Carpospores from monoecious leafy gametophytes developed into conchocelis filaments in culture, and under “winter-spring” conditions these formed conchospores that germinated to produce leafy thalli. Monospores from asexual leafy thalli developed directly into two different forms of leafy thalli. Only one of the cultured morphotypes became fertile, reproducing asexually by monospores. We conclude that the phases of the life history of P. endiviifolium show different ecological strategies, the conchocelis phase reproducing in response to short days unlike the leafy thalli in which growth and reproduction respond primarily to irradiance. Received: 2 May 1997 / Accepted: 11 October 1997     

Ploidy Distribution of the Harmful Bloom Forming Macroalgae Ulva spp. in Narragansett Bay,Rhode Island,USA, Using Flow Cytometry Methods

Macroalgal blooms occur worldwide and have the potential to cause severe ecological and economic damage. Narragansett Bay, RI is a eutrophic system that experiences summer macroalgal blooms composed mostly of Ulva compressa and Ulva rigida, which have biphasic life cycles with separate haploid and diploid phases. In this study, we used flow cytometry to assess ploidy levels of U. compressa and U. rigida populations from five sites in Narragansett Bay, RI, USA, to assess the relative contribution of both phases to bloom formation. Both haploid gametophytes and diploid sporophytes were present for both species. Sites ranged from a relative overabundance of gametophytes to a relative overabundance of sporophytes, compared to the null model prediction of √2 gametophytes: 1 sporophyte. We found significant differences in cell area between ploidy levels for each species, with sporophyte cells significantly larger than gametophyte cells in U. compressa and U. rigida. We found no differences in relative growth rate between ploidy levels for each species. Our results indicate the presence of both phases of each of the two dominant bloom forming species throughout the bloom season, and represent one of the first studies of in situ Ulva life cycle dynamics.     

ALTERNATION OF GENERATIONS IN ALGAE: ITS COMPLEXITY, MAINTENANCE AND EVOLUTION

Life histories of photosynthetic eukaryotes traditionally-termed algae exhibit a considerably greater degree of complexity than those of ‘higher cryptogams.’ Some algae have a so-called ‘obligate’alternation between spore-producing and gamete-producing phases, but the majority seem capable of following other pathways depending upon environmental conditions. In only four algal classes do life histories show a change in morphological and/or nuclear phases. The following basic life histories are recognized in the Chlorophyceae, Phaeophyceae and Rhodophyceae:(a) monophasic, a diploid or haploid phase, (b) two or more phases, most commonly an alternation of an isomorphic or heteromorphic haploid gametangial phase and a diploid sporangial phase, and (c) three phases (unique to florideophyte Rhodophyceae), with a diploid spore-producing phase (carposporophyte) developing on the gametangial phase, a diploid phase (tetrasporophyte if meiosis is sporic) and a haploid gametangial phase. Evidence from recent research indicates that in many algae there is an uncoupling of the morphological and nuclear phases. The dominance of one phase and suppression of another has been suggested to be due to the common occurrence in algae of apogamy, apomeiosis and parthenogenesis. Free-living morphs in heteromorphic life histories may be morphologically so dissimilar that formerly they were attributed to different genera. Evolution of the carposporangial phase in red algae is speculated to be a means of achieving zygotic amplification to compensate for the infrequency of syngamy. Such amplification allows the production of a large number of dispersible products from a single fertilization. The direct development of a free-living tetrasporangial phase is considered another mechanism for achieving amplification. In freshwater red algae the growth of an upright phase from a perennial microscopic one is considered an adaptation for maintaining their upstream position. Life history pathways in algae are controlled by subtle environmental influences (e.g. photoperiodism, temperature, light quality, nutrients). Experimental evidence is lacking to support the contention that spatial and/or temporal partitioning of the environment is a mechanism favouring the maintenance of heteromorphy. Herbivory is known to be an important selective force suppressing some morphs and accentuating the seasonal dominance of others. Differential resistance of morphs to herbivory in environments where grazing intensity is predictable may lead to the selective maintenance of heteromorphy. Algal life history patterns are unexplored in terms of evolutionary processes. Various models for the evolution of biphasic or polyphasic life histories stress the importance of the capacity for both asexual dispersal of successful genotypes and for the generation of new genotypes via meiosis and syngamy. All evidence points to the fact that many life history processes operative in algae differ significantly from those described for other cryptogams.     

Hybridization involving independent gametophytes in the Vandenboschia radicans complex (Hymenophyllaceae): a new perspective on the distribution of fern hybrids

To test our hypothesis of hybrid formation involving the ‘independent gametophyte’ phenomenon in ferns, we identified the genomic formulae and ploidy level of gametophytes of the Vandenboschia radicans complex at the periphery of a sporophyte population. We identified haploid gametophytes of V. kalamocarpa (one of the two putative parents of V.×stenosiphon) in a hybrid sporophyte population in Japan that lacks fertile non‐hybrid individuals. Furthermore, diploid sporophytes of the species were not found within a 50‐km radius. This finding supports a hypothesis of hybridization involving the ‘independent gametophyte’ phenomenon and provides a new perspective on the geographical distribution of fern hybrids.     

THE LIFE HISTORY OF COLEOCHAETE SCUTATA (CHLOROPHYCEAE) STUDIED BY A FEULGEN MICROSPECTROPHOTOMETRIC ANALYSIS OF THE DNA CYCLE1,2

The life history of Coleochaete scutata Bréb. was analyzed by Feulgen microspectrophotometry, a technique measuring DNA content in individual nuclei. By correlating nuclear DNA content with morphological structures or stages in the life history, changes in ploidy level are revealed. The microspectrophotometric study confirmed the earlier reports of a haploid vegetative thallus with mitotic division restricted primarily to the margin of the thallus. In the mitotic cycle the G₁ (pre-synthesis) phase is longer in duration than the synthesis find G₂ (post-synthesis) phases. Oogamous sexual reproduction results in resistant oospores which attain DNA levels of 2C 8C (1C being the DNA level of gamete nuclei).     

ADDITIONAL EVIDENCE FOR ECOLOGICAL DIFFERENCES AMONG ISOMORPHIC REPRODUCTIVE PHASES OF IRIDAEA LAMINARIOIDES (RHODOPHYTA: GIGARTINALES)1

Gametophytes are more abundant thou sporophytes in wave exposed rocky intertidal populations of Iridaea laminarioides Bory in Central Chile. In this study we experimental tested the differential effects of selected ecological factors on karyologically different life history phases. In the field, gametophytes dominated at higher elevation and during summer; tetrasporophytes were most abundant low in the intertidal and during the fall. Laboratory responses correlated with these patterns. Gametophytes exhibited greater desiccation tolerance than tetrasporophytes. Optimum growth of gametophytes occurred at higher temperatures (20°C) and longer photoperiods (16:8 h LD) than sporophytes (15°C and 12:12 h LD). Grazing preferences changed with the developmental stage of the alga, but all herbivores tested had increased preference for diploid tissues as compared to haploid. Number of spores produced with respect to total plant surface, or total rocky surface, or settlement of spores and their germination rate did not show significant differences between phases but showed great variability in space and time. Spontaneous spore release, however, was always higher in cystocarpic than in tetrasporangial thalli. Such a combination of results suggests that some real ecological differences exist between the two life history phases of I. laminarioides. Such ecological differences permit a prediction of vertical and temporal patterns of distribution for both phases. Horizontal patterns of distribution cannot be explained because the several selection factors probably interact differently in various habitats.     

PHOTOSYNTHETIC RESPONSES TO LIGHT AND TEMPERATURE OF THE HETEROMORPHIC MARINE ALGA MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Physiological differentiation of the heteromorphic life-history phases of the red alga Mastocarpus papillatus (C. Agardh) Kützing was assessed. Photosynthetic responses to light and temperature of the erect, foliose gametophyte were compared to those of the crustose tetrasporophyte. Plants of both phases were collected from four locations on the Pacific coast of Baja California, Mexico, and California, USA, between 32–4l° N latitude. Within each location, the chlorophyll-specific, light-saturated photosynthetic rates of gametophytes were generally five times greater than those of tetrasporophytes. Initial slopes of photosynthesis: irradiance curves were greater for gametophytes than for tetrasporophytes. The crust and the blade from each location were similar with respect to dark respiration rates. For tetrasporophytes from all locations, the photosynthetic temperature optima were between 12–15° C. The photosynthetic temperature optima for gametophytes ranged from 15–17° C for plants from Trinidad Head (41° N) to 22–25° C for plants from Punta Descanso (32° N). Both gametophytes and tetrasporophytes from the northernmost location had significantly higher photosynthetic rates than the same phase from the other three locations. The photosynthetic responses to light support models for the life history in which life history phases have different ecological roles. The gametophyte, thought to be specialized for rapid growth and competition, may allocate more resources to photosynthetic machinery, hence the higher maximum photosynthetic rates. The tetrasporophyte, thought to be specialized for resistance to herbivores, may allocate more resources to structural or chemical defenses in preference to photosynthetic machinery. Consequently, the tetrasporophyte has lower photosynthetic capacity.     

Evolution and maintenance of haploid–diploid life cycles in natural populations: The case of the marine brown alga Ectocarpus

The evolutionary stability of haploid–diploid life cycles is still controversial. Mathematical models indicate that niche differences between ploidy phases may be a necessary condition for the evolution and maintenance of these life cycles. Nevertheless, experimental support for this prediction remains elusive. In the present work, we explored this hypothesis in natural populations of the brown alga Ectocarpus. Consistent with the life cycle described in culture, Ectocarpus crouaniorum in NW France and E. siliculosus in SW Italy exhibited an alternation between haploid gametophytes and diploid sporophytes. Our field data invalidated, however, the long‐standing view of an isomorphic alternation of generations. Gametophytes and sporophytes displayed marked differences in size and, conforming to theoretical predictions, occupied different spatiotemporal niches. Gametophytes were found almost exclusively on the alga Scytosiphon lomentaria during spring whereas sporophytes were present year‐round on abiotic substrata. Paradoxically, E. siliculosus in NW France exhibited similar habitat usage despite the absence of alternation of ploidy phases. Diploid sporophytes grew both epilithically and epiphytically, and this mainly asexual population gained the same ecological advantage postulated for haploid–diploid populations. Consequently, an ecological interpretation of the niche differences between haploid and diploid individuals does not seem to satisfactorily explain the evolution of the Ectocarpus life cycle.     

LIFE HISTORIES OF BLIDINGIA MINIMA (CHLOROPHYCEAE), ESPECIALLY SEXUAL REPRODUCTION1

Blidingia minima (Näg. ex Kütz.) Kylin from Muroran, Hokkaido, Japan, has been shown to exhibit four patterns of life history in culture. Sexual reproduction, reported fully for the first time in the genus, occurs in two of them. I. An isomorphic-heteromorphic complex in which erect, tubular sporophytes alternate with dioecious gametophytes of the same forms, with the irregular production by both phases of discoid or pulvinate (pincushion-like) microthalli capable of both sexual and asexual reproduction. II. An alternation of heteromorphic phases in which an erect, tubular sporophyte alternated irregularly with a gametophytic microthallus. III. An asexual alternation of heteromorphic phases in which an erect, tubular frond alternates irregularly with a microthallus by means of quadriflagellate zoospores. IV. An asexual, monophasic life cycle in which erect, tubular fronds are perpetuated by quadriflagellate zoospores. The first pattern occurred in spring populations from one of three sites. The second occurred in populations from two sites. The third and fourth occurred in all populations tested. The life history of Blidingia minima from Muroran is similar to that of Kornmannia zostericola from Muroran.     

Life history of Chnoospora implexa (Chnoosporaceae, Phaeophyceae) in culture

The life history of the brown alga Chnoospora implexa J. Agardh (Chnoosporaceae, Scytosiphonales) from Japan was studied in laboratory cultures. This species showed a heteromorphic and diphasic life history, alternating between erect gametophytes and discoid sporophytes. The gametophytes were dioecious and produced isogametes. The zygotes developed into sporophytes at 20°C under long‐day conditions, which formed plurilocular zoidangia. Zoids released from the plurilocular zoidangia developed again into sporophytes that always formed plurilocular zoidangia at 20°C and 25°C in long‐day conditions, and mainly unilocular zoidangia at 25°C in short‐day conditions. Zoids released from unilocular zoidangia developed into dioecious gametophytes. At 15°C zygotic erect thalli were formed and were revealed to be diploid by microspectrofluorometric measurements of nuclear DNA contents. The development and reproduction of unfused gametes were similar to those of zygotes. Some strains showed a direct‐type life history; gametophytic thalli were produced, but not via a sporophytic phase.     

The roles of the Chantransia phase of Lemanea (Lemaneaceae,Batrachospermales, Rhodophyta) and of the ‘Mushroom’ phase of Himanthalia (Himanthaliaceae,Fucales, Phaeophyta)

Summary

The life cycle of the Batrachospermales (freshwater florideophyte red such as Batrachospermum, Lemanea) is a shortened variant of the ‘normal’ marine florideophyte life cycle. The perennial Chantransia diploid phase is microscopic and encrusting. Each winter it produces one or more semi-erect haploid gametophytes by vegetative meiosis. Gamete production and fertilization is followed by production of diploid carposporophytes; these produce diploid carpospores which disperse, and regenerate the Chantransia phase. The question of the extent to which the Chantransia phase contributes resources to the gametophyte was approached by physiological-hydrodynamic modelling. These computations suggest that the photosynthetic rates in situ on an area basis are 20 times greater for the gametophyte than the Chantransia phase; this agrees with the observed ratios of peak biomass. The conclusion is that the Chantransia phase has a negligible role in provisioning the growing gametophyte, and that the role of the Chantransia phase is to occupy space with living biomass throughout the year, including exposure at summer drawdown, and (perhaps) by dispersal via production of monospores. A similar conclusion is arrived at on the basis of biomass data for the role of the perennial ‘mushroom’ phase of the semelparous marine Fucalean brown alga Himanthalia elongata in relation to the short-lived but much larger reproductive receptacles.     

The male gametophyte of Bonnemaisonia hamifera Hariot in Norway

The male gametophyte of Bonnemaisonia hamifera Hariot is recorded for the first time in Norway. The plants were collected in the Bj?r?ya archipelago situated between 10° 46′-10° 52′ E and 64° 35·5′-64° 37′ N. Numerous large antheridia were present on the plants which were attached to the rock at a depth of 150–200 cm below E.L.W.S.