The specific identity and the life history of JapaneseSyringoderma (Syringodermatales,phaeophyceae)

Observations on JapaneseSyringoderma from Rishiri Island, which had been previously identified asS. australe, were made based on newly collected fertile material. These results and comparisons with the type specimen ofSyringoderma abyssicola (=Chlanidophora abyssicola) revealed that the Rishiri Island plants should be identified asSyringoderma abyssicola. Syringoderma abyssicola from Rishiri Island formed unilocular sporangia among the paraphyses on the fan-shaped blades in winter. The first products (uni-spores) in the unilocular sporangia form flagella, and soon after form cell walls before release. Then these reduced gametophytes divide into tetrads and form swarmers, each of which contains a chloroplast with a stigma. These swarmers germinate into branched filaments, from which thicker erect filaments of apical growth tissue. At 5–10 C these erect filaments formed fan-shaped blades under long-day conditions, and unilocular sporangia under short-day conditions corresponding respectively to spring and winter at Rishiri Island. Accordingly, the seasonal growth pattern of the species is considered to be controlled by responses to photoregime and temperature. Dedicated to the memory of the late Professor Muneao Kurogi.     

The ship-fouling algaEctocarpus

Summary The unilocular reproductive stages of the brown algaEctocarpus have been examined with the electron microscope. One observation in particular, which is believed to have been made for the first time in Phaeophycean studies, is that the zoospore derived from unilocular sporangia possesses a microanatomy considerably different from that of spores formed by plurilocular sporangia. It is suggested that this difference may have an evolutionary significance and may lead to a better understanding of the life cycle ofEctocarpus.     

Parthenogenesis is rare in the reproduction of a sexual field population of the isogamous brown alga Scytosiphon (Scytosiphonaceae,Ectocarpales)

Parthenogenetic development of unfused gametes is commonly observed in laboratory cultures among various brown algal taxa. There is, however, little information on the contribution of parthenogenesis to the reproduction of field populations. In this study, we investigated whether parthenogenesis is present in a sexual population of the isogamous brown alga Scytosiphon with a 1:1 sex ratio. In culture, both female and male gametes showed higher mortality and slower development compared to zygotes. More than 90% of surviving partheno‐germlings formed parthenosporophytes irrespective of the culture conditions tested. Therefore, if parthenogenesis occurs in the field, most unfused gametes are expected to form parthenosporophytes. Contrary to this expectation, parthenosporophytes were rare in the field population. We collected 126 sporophytic thalli and isolated and cultured a unilocular sporangium from each of them. We confirmed that cultures of 120 unilocular sporangia produced both female and male gametophytes by the observation of zygotes or amplification of PCR‐based sex markers indicating that these sporangia originated from zygotic sporophytes. Only females were detected in cultures from two sporangia and only males from four sporangia suggesting that these sporangia originated from parthenosporophytes. In the Scytosiphon population, although parthenogenesis is observable in culture, our results demonstrate that the contribution of parthenogenesis to reproduction is small (≤4.8%) compared to sexual reproduction. Unfused gametes may not survive to form mature parthenosporophytes in significant numbers in the field partly due to their higher mortality and slower development compared from zygotes.     

Erythropeltidaceen (Bangiophyceae,Rhodophyta) von Helgoland

Ontogenesis and reproduction of the Helgolandian taxa of the Erythropeltidaceae have been studied. In all species monospores are only produced from differentiated sporangia. FilamentousConchocelis-like stages have not been observed. Sexual reproduction was formerly demonstrated in the heteromorphous genusErythrotrichopeltis (Kornmann, 1984). Based on these features a revised classification for the family is presented.Porphyropsis imperfecta, a new species, is a widespread epiphyte in sublittoral habitats.     

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.     

MORPHOLOGY AND LIFE HISTORY OF SCYTOSIPHON LOMENTARIA (SCYTOSIPHONACEAE,PHAEOPHYCEAE) FROM THE AZORES1

Morphological and culture studies of Scytosiphon lomentaria (Lyngbye) Link and Microspongium gelatinosum Reinke (Scytosiphonaceae, Phaeophyceae) were undertaken on material collected on the Island of São Miguel, Azores, where both species were commonly found. Erect thalli of S. lomentaria collected in the field were up to 33 cm long and 2.3 mm wide, tubular, hollow, and commonly constricted at intervals. The plurilocular sporangia were positioned in continuous sori on the thallus surface. Ascocysts were present. In the field, M. gelatinosum formed crustose to slightly pulvinate plants, were spongy in texture, and dark brown to black in color, which were circular or irregularly spreading over several centimeters and firmly attached to the substratum. Sessile unilocular sporangia were located in sori on the crust surface. In culture S. lomentaria plurispores developed into Microspongium‐like crustose prostrate thalli that formed unilocular sporangia. Unispores developed into new erect thalli that formed plurilocular sporangia. Sexual reproduction was not observed. In culture, M. gelatinosum unispores developed into erect thalli identical with S. lomentaria. These results are similar to those reported for other areas and suggest the occurrence in the Azorean plants of a monophasic and heteromorphic life history, involving both entities studied.     

OBSERVATIONS ON THE LIFE HISTORY OF PAPENFUSSIELLA CALLITRICHA (PHAEOPHYCEAE,CHORDARIALES) IN CULTURE1

Papenfussiella callitricha (Rosenv.) Kylin from eastern Canada was studied in culture. Zoids from unilocular sporangia develop into microscopic, filamentous, dioecious gametophytes which produce isogametes in filament cells and few-chambered plurilocular gametangia. Unfused gametes germinate to reproduce the gametophytes. Fusion takes place between a settled (“female”) and a motile (“male”) gamete. The zygote gives rise to a filamentous plethysmothallus that reproduces asexually by zoids formed in thallus cells and in few-chambered plurilocular zoidangia. Erect macrothalli are produced on the plethysmothallus, beginning with the formation of upright filaments. Later on, these filaments become the terminal assimilators of the macrothalli. Further assimilatory filaments, rhizoids, and unilocular sporangia are produced in a branching region at the base of the terminal assimilator. Zoids from unilocular sporangia formed in culture germinate to reestablish the gametophyte phase. Chromosome counts yielded n = 19 ± 3 for the gametophytes, and 32 ± 6 for the sporophyte, both plethysmothallus and macrothallus.     

Morphology and life history of Halopteris filicina (Sphacelariales, Phaeophyceae) from Korea

The vegetative morphology and life history of Halopteris filicina (Grateloup) Kutzing, collected from Korea, were examined in laboratory culture. Field plants attaining 3–5 cm in height were epilithic, tufted, yellowish-brown, and produced numerous erect axes with alternately distichous branches from compact basal discs. They were cultured under a 12:12 h LD photoperiod at 10°-C, 15°C and 20°C to observe the influence of temperature on reproduction. At 10°C plants grew only vegetatively, whereas at 15°C and 20°C they produced unilocular sporangia. Unispores released from sporangia developed into monoecious, anisogamous gametophytes that formed plurilocular female and male gametangia on the same lateral branches. The zygotes, by fusion of female macrogametes and male microgametes, developed into sporophytes bearing unilocular sporangia, whereas the unfused female gametes germinated parthenogenetically. This species was confirmed to have an isomorphic life history, basically similar to the other species of Sphacelariales.     

A mechanism of propagule release from unilocular reproductive structures in brown algae

Summary The structure of unilocular sporangia inP. littoralis was investigated along with several other species of brown algae in order to study the mechanism by which propagules are released from unilocular reproductive structures. Unilocular sporangia inP. littoralis are composed of a spherical cell wall of two distinct layers and contain a number of zoospores. The mass of spores is surrounded and permeated by mucilaginous carbohydrates. It is suggested that the production of these carbohydrates generates the necessary pressure to weaken the sporangial wall. In addition, ultrastructural observations indicate that further weakening seems to occur due to digestion of the inner wall layer. Walls of sporangia were mechanically broken just prior to normal spore release in order to investigate whether internal pressure exists, and if it can effect spore discharge. Results show that an internal pressure does exist prior to normal spore discharge and that this pressure is not generated by turgor pressure of the spores themselves or by a semi-permeable wall osmoticum system. The discharge of spores seems to occur when the carbohydrate around the spores swells. The adsorption of water when plants are immersed by the incoming tide thus seems a likely mechanism of spore discharge. The similarities of unilocular reproductive structures and spore release in several brown algal species suggests common mechanisms of propagule discharge for members of thePhaeophyta.     

Untersuchungen zur Entwicklungsgeschichte der Braunalge Ectocarpus siliculosus Aus Neapel

Zusammenfassung Kulturversuche und cytologische Untersuchungen an Ectocarpus siliculosus aus dem Mittelmeer (Neapel) führten zu folgenden Ergebnissen: die haploide Chromosomenzahl liegt zwischen 18 und 31. Es sind zwei morphologisch verschiedene Wuchsformen vorhanden, die dem Freilandmaterial gleichenden haploiden Gametophyten und die nur in Kultur beobachteten Sporophyten. Letztere können sowohl in der Haplophase als auch in der Diplophase vorliegen und bilden in jedem Fall neben pluriloculären auch uniloculäre Fortpflanzungsorgane aus. In den uniloculären Sporangien der diploiden Sporophyten findet die Reduktionsteilung statt, während sie in denselben Sporangien auf haploiden Pflanzen unterbleibt. Unter den Schwärmern aus den Gametangien der Geschlechtspflanzen finden sich viergeißlige bewegliche Zygoten. Unter den Nachkommen eines Gemisches von unkopulierten Schwärmern und Zygoten wurden haploide und diploide Sporophyten gefunden. Unter den Nachkommen aus uniloculären Sporangien haploider und diploider Pflanzen sind fast stets Vertreter beider Wuchsformen zu finden. Einige der hier mitgeteilten Beobachtungen stimmen nicht mit den Angaben früherer Autoren überein.

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Studies on the life cycle of the brown alga Ectocarpus siliculosus from Naples, Italy

Summary Culture experiments and cytological studies were carried out under well-defined conditions. The haploid chromosome number is between 18 and 31. There are two morphological types of plants: the well-branched gametophyte resembling the plants found in nature, and the unbranched sporophytic form which may be haploid or diploid, found only in cultures. Meiosis takes place in the unilocular sporangia on diploid sporophytes. Haploid sporophytes form unilocular sporangia without reduction of the chromosome number. The spores from unilocular sporangia both on diploid and haploid sporophytes give rise to plants with either sporophytic or gamethophytic growth. The gametophytes are homothallic, the gametes forming motile zygotes with four flagella. Parthenogenetic development of gametes exclusively results in the formation of haploid sporophytes. Both the haploid and diploid sporophytes can propagate by means of zoids from plurilocular sporangia. Several observations reported here disagree with the findings of other workers.

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Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Experimental evidence against sexual fusions of spores from unilocular sporangia of Ectocarpus siliculosus (Phaeophyta)

Evidence that sexual fusion does not occur in zoids from unilocular sporangia of Ectocarpus siliculosus is presented as follows: (1) Zoids from various types of unilocular sporangia do not interact with genuine gametes; (2) Aggregates of two zoids, previously claimed to be zygotes, were frequently seen in cultured material but could be demonstrated to be the result of incomplete cell separation during sporogenesis; (3) Culture studies showed that there is no fusion of male and female nuclei in such aggregates.     

DEVELOPMENTAL STUDIES OF ERYTHROCLADIA ROSENVINGE IN CULTURE

Erythrocladia subintegra has been cultured for over 3 years. The life history includes reproduction by monospores, a long period of ameboid motility by monospores, their germination and growth into monostromatic disk-shaped plantlets composed of adherent, often dichotomously branched filaments. Plants become centrally polystromatic with age and produce monospores. Variations in plant size and shape, cell size and shape, spore size, shape, and motility are discussed. Variations in plants of E. subintegra grown on different substrates have been studied and discussed in relation to other species in the genus.     

Generationswechsel,Kernphasenwechsel und Sexualität der Braunalge Ectocarpus siliculosus im Kulturversuch

Zusammenfassung Der Entwicklungscyclus von Ectocarpus siliculosus wurde unter Kulturbedingungen untersucht. Gametophyten und Sporophyten können morphologisch und funktionell unterschieden werden. Die Gametophyten sind diöcisch, die Geschlechtsbestimmung erfolgt genotypisch. Sporophyten können in der haploiden, diploiden und tetraploiden Phase vorliegen. Sporophyten aller Kernphasen können unilokuläre Sporangien ausbilden, tetraploide und diploide Sporophyten führen dabei eine Reduktionsteilung durch. Schwärmer haploider Gametophyten und Sporophyten können sich spontan zu diploiden oder tetraploiden homozygoten Sporophyten entwickleln. Gametophyten können nur zusammen mit Sporophyten aus Schwärmern unilokulärer Sporangien entstehen (Heteroblastie). Aus der Reduktionsteilung tetraploider Sporophyten gingen diploide männliche Gametophyten hervor, deren Gameten mit normalen weiblichen Gameten kopulierten. Die verschiedenen Kernphasen und Wuchsformen stehen durch Reduktionsteilungen, Heteroblastie, Kopulation und spontane Aufregulierung der Chromosomenzahl miteinander in Verbindung.

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Culture experiments on life cycle, nuclear phases, and sexuality of the brown alga Ectocarpus siliculosus

Summary The life cycle of Ectocarpus siliculosus from Naples (Italy) was investigated, using well defined cultured material. Gametophytes and sporophytes differ morphologically and functionally. The gametophytes are dioecious, with genotypic determination of their sex. Sporophytes exist in the haploid, diploid and tetraploid phase. All sporophytes can form unilocular sporangia. In tetraploid and diploid sporophytes the formation of unilocular sporangia is connected with meiosis. Certain motile cells of haploid plants may spontaneously give rise to diploid or tetraploid sporophytes which are homozygous. Gametophytes can only be formed together with sporophytes form the swarmers of unilocular sporangia (heteroblasty). Meiosis in tetraploid sporophytes resulted in diploid gametophytes, the gametes of which fused with haploid female gametes. All observed nuclear phases and growth forms are connected with each other by meiosis, heteroblasty, fusion of gametes and spontaneous increase in chromosome number.

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Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

ETIOLOGY AND DEVELOPMENT OF HYPERPLASIA INDUCED BY STREBLONEMA SP. (PHAEOPHYTA) ON MEMBERS OF THE LAMINARIALES (PHAEOPHYTA)1

Filaments of Streblonema sp. isolated from hyperplasia (galls) on Nereocystis luetkeana (Mert.) Post. et Rupr. induced similar gall growths when inoculated on young sporophytes of Nereocystis luetkeana, Macrocystis integrifolia Bory, and Laminaria japonica Aresch. Filaments were irregularly branched and uniseriate with cells 5–8 μm in diameter and 10–30 μm long. Growth under varied cultured conditions produced unilocular sporangia. Zoospores released from the sporangia germinated into identical filaments which also formed unilocular sporangia. Gall tissue originated from the innermost cells of the epidermal meristematic zone. When infected these cells divided in an irregular and unorganized manner. The resulting structure was a pronounced departure from normal morphology.     

Life history of Japanese Stypocaulon durum (Sphacelariales, Phaeophyceae)

Phenology, morphology, life history and responses to different temperature and photoperiod conditions were studied in Japanese Stypocaulon durum (Ruprecht) Okamura. Erect thalli of the species were collected year-round, but the mature thalli forming either uniloc-ular sporangia or two different types of plurilocular structures (evidently gametangia) on separate thalli were found only in winter. ln culture, an isomorphic life history is suggested for the species, alternating between a sporophyte forming unilocular sporangia and gam-etophytes forming plurilocular macro- (female) and micro- (male) gametangia. Contents of unilocular sporangia were not released, but germinated in situ, developing into erect thalli forming plurilocular gametangia. Macrogametangia released aplanogametes (oospores), but male gametangia appeared to be non-functional, although flagellated cells were once formed in the loc-uli. This is the first report of plurilocular gametangia in the species. Although the species grew well and matured under considerably lower temperature conditions than European Stypocaulon scoparium (L.) Sauvageau, its temperature requirements showed similarity to northwestern Atlantic Stypocaulon species. This supports the notion that northwestern Atlantic Stypocaulon is conspecific with S. durum.     

Zur Synonymie vonLitosiphon pusillus auf der Basis von Kulturversuchen

In June 1988,L. pusillus was collected for the first time in the Helgoland area. On the basis of observations in cultures, Pedersen's hypothesis that the microthalli ofL. pusillus are identical with those of threePilocladus species is discussed. The general morphology as well as the shape and dimensions of the unilocular sporangia are clearly different inPilocladus andLitosiphon microthalli. Therefore, Pedersen's opinion cannot be accepted.      

Reproduction in Halopteris (Sphacelariales)

Oogamy is recorded in five southern species of Halopteris. Ineach the gametangia and asexual sporangia occur in similar sori,although situated on separate plants. The plurilocular antheridiaand the unilocular oogonia occur within one and the aame sorus.In both the dehiscence is apical. In H. congesta fertilization has been observed in cultures.The polarity of the zygote is established by the direction ofthe first-formed wall. Early divisions convert it into a nodulecomposed of small cells. First rhizoids appear and then cylindricalshoots arise from the opposite pole. Sporelings of five NewZealand species of Halopteris have been raised in culture. Theconditions in the culture greatly affected the character ofthe sporelings, the simplest of which at the end of 2 months'growth were only uniseriate filaments. The best developed sporelingshad a compact basal disc with up to four upright shoots showingthe beinnings of branching.     

Life history and systematic position of the rare brown alga Climacosorus mediterraneus

Cultures of Climacosorus mediterraneus were established with material from Spitsbergen. This species has previously been reported from three widely separated geographic areas: the west Mediterranean, north-west France, and north Norway. Although adapted to temperatures near 0°C, the northern isolate grew well at 15°C which approximates the summer temperature of the southern habitats. The life history proved to be entirely asexual with successive generations bearing unilocular sporangia on the erect filaments and plurilocular sporangia infrequently on the prostrate system. Swarmers of the two reproductive organs gave rise to similar progeny. Chromosome counts revealed equal numbers in somatic cells and unilocular sporangia which substantiates the development observed. The absence of true hairs and the few longitudinal divisions, which only occur in connection with formation of unilocular sporangia and branches, suggest that C. mediterraneus should be assigned to the Pilayellaceae.     

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.     

RELATION OF TEMPERATURE TO DEVELOPMENT OF THE MACROTHALLUS OF DESMOTRICHUM UNDULATUM1,2

Desmotrichum undulatum (J. Agardh) Reinke (Phaeophyta) was found in the winter months in the littoral zone of a tidal marsh in Virginia. When field collections were maintained in culture media at 21 C, most cells of the thallus developed into plurilocular sporangia. Released zoospores developed into microthalli from which clonal cultures were established. At 21 C microthalli produced sporangia which released zoospores, subsequently developing into more microthalli. However, microthalli at 6 C formed filaments which subsequently developed into thalloid plants. When the thalloid plants of Desmotrichum formed in culture were subjected to 21 C, plurilocular sporangia were formed. These results support the proposition that the seasonal periodicity of Desmotrichum is due to a temperature-regulated phenomenon in the development of macrothalli.