UNUSUAL PHENOMENA IN THE LIFE HISTORIES OF FLORIDEAE IN CULTURE1

Several Florideae grown in natural seawater media under defined laboratory conditions have interesting and unusual life histories. Antithamnion occidentale males of one generation produced tetraspores that gave rise to nonsporangiate males. The functional females of A. pygmaeum developed spermatangia and tetrasporangia; the tetraspores formed new females. Antithamnion defectum tetrasporophytes of one generation bore spermatangia in addition to tetrasporangia; the tetraspores gave rise to typical gametophytes. Tetraspores from successive generations of Callitham-nion sp. developed into tetrasporophytes and males but no females were produced. Functional female gametophytes of Platythamnion sp. bore abortive tetrasporangia. Field-collected plants of two species of Fauchea produced tetraspores that yielded additional sporangiate plants: those of F. pygmaea being bispo-rangiate and tetrasporangiate, and those of F. lacini-ata being strictly tetrasporangiate. Male plants of Pleonosporium vancouverianum from a running seawater table bore spermatangia and polysporangia when collected. The same plants in unialgal culture produced only spermatangia.     

Deviations in the life-history of Gracilaria sp. (Rhodophyta,Gigartinales), from Coquimbo,Chile, under different culture conditions

Cystocarpic branches of a species of Gracilaria from Coquimbo, Chile, were cultured in vitro. A Polysiphonia-like life history was completed in about 6 months, but some abnormalities were observed: i. carpospores gave rise to plants producing either tetrasporangia and spermatangia, or tetrasporangia only; ii. tetraspores cultured without aeration developed into plants bearing spermatangia only; tetraspores cultured with aeration developed into 1:1 female and male gametophytes; iii. plant originated from tetraspore produced spermatangia and tetrasporangia; one of these tetraspores developed into a male gametophyte; iv. some tetraspores gave rise to spherical bodies instead of the ordinary cylindrical branches; one of them bore spermatangia after three months. The results show that environmental factors seem to be interfering with the mechanism of sex determination and induce the development of spermatangia on putative female gametophytes, or on putative tetrasporophytes. Noted added in proof: The Gracilaria sp. studied here was recently described as a new species, G. chilensis by Bird C. J., McLachlan, J & Oliveira, E. C. de, 1986. Can. J. Bot. 64: 2928–2934. Noted added in proof: The Gracilaria sp. studied here was recently described as a new species, G. chilensis by Bird C. J., McLachlan, J & Oliveira, E. C. de, 1986. Can. J. Bot. 64: 2928–2934.     

MORPHOLOGY AND DEVELOPMENT OF AHNFELTIA PLICATA (RHODOPHYTA): PROPOSAL OF AHNFELTIALES ORD. NOV.1

Ahnfeltia plicata (Hudson) Fries, the type species of Ahnfeltia Fries, is currently assigned to the Phyllophoraceae (Gigartinales). Several morphological and biochemical characters distance A. plicata from the Phyllophoraceae but, because sexual reproduction has never been demonstrated, an alternative placement has not been possible. A. plicata now is shown to have a heteromorphic sexual life history. Erect branched gametophytes are dioecious. In male sori, spermatangia are cut off transversely from spermatangial mother cells. Female sori form numerous terminal sessile carpogonia. Following fertilization, several zygotes in each sorus fuse facultatively with undifferentiated intercalary cells of the female sorus and cut off gonimoblast initials obliquely outwards. These initials give rise to branching gonimoblast filaments that fuse with apical and intercalary female sorus cells and with each other, then grow radially outward in the compound external carposporophyte and terminate in carposporangia. Carpospores develop in culture into crustose tetrasporophytes identical to Porphyrodiscus simulans Batters. Field-collected P. simulans tetraspores grew into erect A. plicata axes. Tetrasporangia are formed by division and enlargement of crust apical cells followed by sequential enlargement and maturation of tetrasporocytes in an erosive process. Monosporangia are formed in sori on male gametophytes. Pit plugs of both gametophyte and tetrasporophyte phases consist of naked plug cores without cap layers of membranes. Gametophytes exhibit both cell fusions and secondary pit connections whereas tetrasporophytes form cell fusions but lack secondary pit connections. On the basis of the unique female and postfertilization reproductive development and in conjunction with the pit plug structure which is unique among florideophytes, the order Ahnfeltiales, containing the family Ahnfeltiaceae, is proposed.     

REPRODUCTION AND LIFE HISTORY OF THE RED ALGA GALAXAURA OBLONGATA (NEMALIALES,GALAXAURACEAE)1

Culture and morphological studies showed that Galaxaura oblongata (Ellis et Solander) Lamouroux has a triphasic life history with conspicuous gametophytes and small filamentous tetrasporophytes. Development of male and female reproductive structures is very similar and both begin with the enlargement of a terminal cell of a filament branch occupying a normal vegetative position within the apical pit of a thallus branch. In male thalli this modified branch forms a conceptacle in which spermatangia are produced. In female thalli, this modified branch forms a three-celled carpogonial branch consisting of a carpogonium, hypogynous cell and basal cell. Filament branches from the basal cell form a pericarp and the gonimoblast develops directly from the carpogonium. Carposporangia are produced in conceptacles which resemble the male conceptacles. About the time the first carposporangia are produced, the carpogonium, hypogynous cell and basal cell form a large fusion cell. Released carpospores germinate in a unipolar or bipolar manner and form small filamentous thalli. Under short day conditions, cruciate tetrasporangia are produced in small clusters. Tetraspores germinate similarly to carpospores and also form small filamentous thalli. Under low nutrient conditions, small cylindrical thalli develop on the filaments and these appear similar to gametophytes collected in nature.     

SEXUAL DIFFERENTIATION OF GRIFFITHSIA (CERAMIALES, RHODOPHYTA): NUCLEAR PLOIDY LEVEL OF MIXED-PHASE PLANTS IN G. JAPONICA1

Mixed-phase plants of Griffithsia japonica Okamura spontaneously occurred in a laboratory culture. Four female plants produced tetrasporangia and spermatangia in addition to their normal female reproductive structures (bisexual/mixed-phase plants), and four male plants produced tetrasporangia as well as spermatangia (male/mixed-phase plants). To determine the nuclear ploidy level of these mixed-phase plants, relative nuclear sizes of male, female, tetrasporangial, and mixed-phase plants were measured using a microscopic image analysis system. Haploid gametophytes could be distinguished from diploid tetrasporophytes by relative nuclear sizes, with the later having nuclei twice the size of the former. Relative nuclear sizes of the mixed-phase plants were similar to those of the haploid plants. Thus, the mixed-phase plants were determined to be haploid. Haploid mixed-phase plants of G. japonica have a potential to produce male, female and tetrasporangial reproductive structures. Sex determination models are discussed to explain "haploid" mixed-phase phenomena in red algae .     

Morphology,temperature and photoperiodic responses in Audouinella botryocarpa (Harvey) Woelkerling (Acrochaetiaceae,Rhodophyta) from Ireland

Abstract

Audouinella botryocarpa is reported for the first time from the British Isles. On the west and south coasts of Ireland and in the Isle of Man it grows in the intertidal on wave-splashed limpet shells and rock surfaces. Populations examined monthly in Galway Bay formed monosporangia throughout the year and tetrasporangia from December to March and May but no gametangial plants were found. Plants isolated into culture from monospores formed monosporangia at daylengths of 8, 10, 12, 14 and 16 h at 6.5, 8, 10, 11 and 15°C; the same plants formed tetrasporangia at daylengths ≤10 h at 8, 10 and 11°C but not at 6.5 and 15°C. Spores isolated from these plants gave rise to further monosporangial plants that also formed tetrasporangia under inductive conditions. Night-breaks of 1 h in a 16 h night prevented tetrasporangial reproduction and a critical daylength of ～10 h was found at 10°C. British Isles Audouinella botryocarpa typically has 6 rounded or irregularly-shaped chloroplasts per cell, each of which has a single, centrally-placed pyrenoid. The chloroplasts grow together in mature cells, typically giving the appearance of a single plate-like chloroplast with 6 pyrenoids. The implications of these observations for the generic classification of acrochaetioid algae is discussed and it is concluded that chloroplast morphology, number and the presence or absence of pyrenoids may provide the basis for a future generic realignment of acrochaetioid algae.     

The reproductive phenology of Delesseria sanguinea and Odonthalia dentata off the Isle of Man

Samples of individual plants of Delesseria sanguinea and Odonthalia dentata were collected from shallow subtidal populations off the south end of the Isle of Man during the reproductive season. Reproductive bladelets were measured and their state of fertility noted. In Delesseria, male bladelets appeared in early September and achieved maximum size in mid October when bladelets of all sizes became fertile; the largest of these dehisced first and all were spent by mid December. Females appeared 3 weeks later; carpogonia were fertilized during October but carpospores were not released until about February when maximum bladelet length was reached. In about half the bladelets carpogonia remained unfertilized and did not grow further and only 40% of successful fertilizations resulted in carpospore release. Tetrasporangial bladelets did not appear until November and tetraspores were released in January and February while bladelets were still growing. Gametophytes and tetrasporophytes existed in about equal numbers. In Odonthalia, all three types of bladelet appeared in early November and fertilization took place in December, but later-developing carpogonia seemed to remain unfertilized in spite of a second wave of production of spermatangia, resulting in a low fertilization success of about 7%. Both carpospores and tetraspores dehisced from January to April. In this species the gametophytes formed about 58% of the population, a proportion expected from equal survival of number of spores per parent.     

THE LIFE HISTORIES OF RHODOCHORTON PURPUREUM AND R. TENUE IN CULTURE

The life histories of the red algae Rhodochorton purpureum and R. tenue were studied in unialgal culture. Telrasporophytes produced sporangia in short day regimes (8–12 hr) with 50–200 ft-c cool white light at 10 or 15 C. Mature gametophytes were not observed in the Washington and Alaska clones of R. purpureum. Tetraspores from R. tenue and the California clones of R. purpureum give rise to unisexual gametophytes that are reproductive when smaller than the tetrasporophytes. The tetrasporophytes develop directly from the gonimoblast cells. Because of similarities in morphology and life histories, it is proposed that R. tenue be placed in synonomy with R. purpureum.     

THE LIFE HISTORY OF CALLITHAMNION BYSSOIDES IN CULTURE

The life history of Callithamnion byssoides Arnott ex Harv. in Hook. has been shown to comprise a regular sequence of gametophytic, carposporophytic, and tetrasporophytic phases in unialgal culture using supplemented seawater media. Tetraspore germlings gave rise to gametophytes bearing either antheridia or carpogonia in 13 days. Fertilization, carposporophyte development, and carpospore release took place within 5 days. Carpospore germlings produced mature tetrasporophytes in 13 days. The life history thus required approximately 1 month for completion. No deviations from this pattern were observed.     

Systematics and distribution of Thorea (Thoreaceae, Rhodophyta) from central Mexico and south-eastern Brazil

Thirteen populations of Thorea were analyzed from central Mexico and south‐eastern Brazil. All populations were considered as belonging to a single species [Thorea hispida (Thore) Desvaux], with wide variation of morphological features. Secondary branches varying in frequency were observed in several populations with an overlapping in the range of branch density for Thorea violacea Bory and T hispida (0–9 and 11–41 per 30 mm, respectively). As this is the most distinguishing character and on the basis of the overlapping (within a same population or even a single plant), we regarded T. violacea as a synonym of T hispida.‘Chantransia’ stage in culture, as well as gametophyte and carposporophyte were described in detail. We confirmed tine coexistence of asexual monosporangia with sexual reproductive structures (carpogonia and spermatangia) and carposporangia. Size, content, arrangement and chromosome number were the most distinctive characteristics among spermatangia, carposporangia and monosporangia. Monosporangia can be promptly differentiated from spermatangia by their granulated content and larger size but they are similar to carposporangia in shape and size; however, monosporangia are not arranged in fascicles. Structures resembling bisporangia were observed in female plants of some populations. Chromosome numbers were n = 4 for spermatangia and fascicle cells, and 2n ca 8 for gonimoblast filaments, carpospores and the ‘Chantransia’ stage cells. The populations of Thorea from central Mexico and south‐eastern Brazil corroborated the known world distribution for T. hispida, consisting dommantly of tropical to subtropical rainforests, sometimes extending into warm temperate areas. Thorea hispida occurred in warm (temperature 17.6–28.O°C), neutral to alkaline (pH 7.0–8.0), high ion content (specific conductance 59–2140 μS cm^?1), moderate flowing (current velocity 17–43 cm/s) and shallow waters (depth <50cm); these data are essentially similar to previous reports.     

Life history and photoperiodic responses in Chondrus giganteus forma flabellatus (Gigartinaceae,Rhodophyta) from Japan

The life history and photoperiodic responses of the carrageenophyte Chondrus giganteus forma flabellatus (Rhodophyta) from Japan were investigated in culture. This entity has a Polysiphonia-type life history and the formation of both tetrasporangia and cystocarps is controlled by daylength. A night-break of 1 h in the middle of a 16 h night was effective in inhibiting reproduction, confirming the classical photoperiodic nature of the response. Tetrasporangia formed at daylengths of 12 h or less, whereas cystocarps only developed at daylengths of 10 h or less. The formation of spermatangia required daylengths of 12 h or less, but production was not inhibited by a night-break of 1 h in the middle of a 16 h night.     

REGENERATION AND SEXUAL DIFFERENTIATION OF GRIFFITHSIA JAPONICA (CERAMIACEAE,RHODOPHYTA) THROUGH SOMATIC CELL FUSION1

Hybrid cells were obtained from somatic cell fusion among male, female, and tetrasporangial plants in Griffithsia japonica Okamura by a wound-healing process. Isolated fusion cells regenerated new mature plants with mixed reproductive structures. The plants regenerated from hybrid cells between male and female plants developed into 1) spermatangiate, 2) carpogonial, 3) bisexual with spermatangia and carpogonial branches, 4) mixed-phase with spermatangia and tetrasporangia, or 5) bisexual/mixed-phase plants with spermatangia, carpogonial branches, and tetrasporangia. About 70% of the plants regenerated from hybrid cells between male and female plants produced tetrasporangia that were always formed with spermatangia on a single cell. Some of those tetrasporangia released tetraspores, six of which gave rise to mature plants. The plants regenerated from hybrid cells between male and tetrasporangial plants developed into spermatangiate, tetrasporangiate, or mixed-phase plants with spermatangia and tetrasporangia. The plants regenerated from hybrid cells between female and tetrasporangial plants developed into carpogonial, tetrasporangiate, or mixed-phase plants with carpogonial branches and tetrasporangia. All types of reproductive structures we re functional.     

BONNEMAISONIA HAMIFERA HARIOT IN NATURE AND IN CULTURE23

In culture tetraspores of the sporophyte (Trailliella intricata) of Sonnemaisonia hamifera gave rise to male gametophytes only. In the Gulf of St. Lawrence tetrasporangia occur annually during the autumn, but mature gametophytes have not been recorded. However, gametophytes with antheridia were found during the winter on the Atlantic coast of Nova Scotia. Both phases are capable of vegetative propagation, and this is probably the usual method of reproduction in nature.     

Life-cycle of <Emphasis Type="Italic">Scinaia interrupta</Emphasis> (Nemaliales,Rhodophyta)

The life-cycle of Scinaia interrupta (A.P. de Candolle) M. J. Wynne was investigated in vitro using four irradiance regimes: 4, 8, 12 and 16 μmol photons m⁻² s⁻¹. A triphasic heteromorphic life-cycle was observed. Carpospores released by cystocarps of gametophytes collected in the field developed into filamentous tetrasporophytes, which produced tetrahedral tetrasporangia. Tetrasporangial development was accelerated under higher irradiance levels. Tetraspores germinated into filamentous protonemal gametophytes, initially identical to the tetrasporophyte. Filamentous gametophytes developed apical utricles and gave rise directly to the fleshy gametophyte. Further development of the fleshy gametophyte was not observed at the lowest irradiance regime (4 μmol photons m⁻² s⁻¹). The present study reports for the first time the influence of the irradiance regime on the initial tetrasporangial development and in the development of the fleshy gametophyte, and reinforces the importance of light intensity on Scinaia life-cycle. Production of apical utricles by the filamentous gametophyte is newly reported for the genus.     

Physiological comparison between gametophytes and tetrasporophytes of Gelidium canariensis (Gelidiaceae: Rhodophyta)

The physiological performances of tetrasporophytes and gametophytes of Gelidium canariensis (Grunow) Seoane-Camba were compared to estimate whether the field predominance of tetrasporophytes is due to lower fitness of gametophytes. No significant differences between tetrasporophytes and gametophytes were detected for calorific content, protein and pigment concentrations, NADH-Diaphorase, alkaline phosphatase and glucose-6-phosphate dehydrogenase activities and photosynthesis and respiration at 15, 20 and 25 °C, and pH 6.5, 8.2 and 9.2. Our results indicate that these physiological characteristics are not responsible for the scarcity of gametophytes in the field populations of G. canariensis.Abbreviations ALP = Alkaline phosphatase - G6PDH = glucose-6-phosphate dehydrogenase - DIA = NADH-Diaphorase - TRIS = Tris[hydroxymethyl]-aminomethane - PVPP = Polyvinylpolypyrrolidone     

The life history of Ahnfeltiopsis paradoxa (Gigartinales, Rhodophyta) in laboratory culture

The life history of the red alga Ahnfeltiopsis paradoxa (Suringar) Masuda (Phyllophoraceae, Gigartinales) from Japan was completed in laboratory culture. Carpospores isolated from field-collected plants germinated to form circular crusts that were composed of a monostromatic hypothallium consisting of radiating filaments, a polystromatic perithallium consisting of tightly coalescent erect filaments, and hypobasal tissue derived from the hypothallium. The crusts were induced to sporulate by transferring them from short-day to long-day regimes at 15° and 2°C. Each crust produced several nemathecia along 1-4 concentric rings. Intercalary, cruciately or decussately divided tetrasporangia were formed in 4-6 (1-2 at the margin of the nemathecium) successive cells of a single filament of the nemathecia. Tetraspore germlings gave rise to basal discs from which upright axes developed. The upright axes first grew without branches or were sparsely branched and later bore many marginal reproductive proliferations. Procarps and spermatangia were formed in the proliferations on different individuals. Carposporophytes developed on female plants that were co-cultured with male plants. Gonimoblast filaments were formed from an auxiliary cell that fused with a carpogonium. Carposporangia developed from gonimoblast filaments and medullary cells contacted by the gonimoblast filaments. Carpospores were discharged through carpostomes formed in the thickened cortex. Tetraspores were cultured from field-collected crusts of a morphology similar to that of cultured tetrasporophytes. They gave rise to upright gametophytic axes similar in morphology to this species as seen in the field.     

A re-examination of the morphology and reproduction of Nothocladus lindaueri (Batrachospermales, Rhodophyta)

The vegetative morphology and reproduction of the freshwater rhodophyte Nothocladus lindaueri Skuja [=Batrachospermum lindaueri (Skuja) Necchi et Entwisle] were examined by light and electron microscopy. It was confirmed that this alga has a typical batrachospermalean pit plug with two cap layers, the outer one of which is domed. During elongation of hair cells, the primary wall is broken, forming a basal collar. Hair cells have a single nucleus and abundant Golgi bodies, en-doplasmic reticula (ER) and vesicles. Dividing apical cells of the fascicles have a nucleus with art adjacent zone of exclusion, the latter containing a single polar ring. Branched trichogynes and fertilized carpogonia are shown for the first time in this species. Carpogonial branch and involucral cells contain a prominent axial nucleus, proplastids, ER and vesicles. The pit plugs disintegrate among these cells leaving open pit connections. Carpogonia have plentiful mitochondria and vesicles. The wall at the trichogyne apex is thickened and densely stained. The carposporophyte centre consists of a mass of fusion cells with open pit connections, and indeterminate gonimoblast filaments arise from this mass. The combination of a symmetrical carpogonial base, a carposporophyte centre consisting of a mass of fusion cells, and exclusively indeterminate gonimoblast filaments appears to be unique among the members of the Batrachospermaceae. The specimen of N. lindaueri contains epiphytic filaments of Audouinella meiospora producing both spermatangia and monosporangia. Spermatium formation in N. lindaueri remains unknown.     

OBSERVATIONS ON LIAGOROPHILA ENDOPHYTICA,A RARE SPECIES IN THE ACROCHAETIACEAE (RHODOPHYCEAE)1

Liagorophila, a monotypic genus in the Acrochaetiaceae (Nemalionales) from the zuestern Pacific, is redescribed, particularly with reference to its sexual reproductive structures. The fertilized carpogonium shows a longitudinal first division; the spermatangia may be borne directly on a normal vegetative cell. Monosporangia are not known. For these reasons, and although vegetatively resembling some species of Acrochaetium, the genus should be recognized as distinct from other genera of Acrochaetiaceae.     

IS THERE AN ECOPHYSIOLOGICAL EXPLANATION FOR THE GAMETOPHYTE–TETRASPOROPHYTE RATIO IN GELIDIUM SESQUIPEDALE (RHODOPHYTA)?1

In the fall, when 61% of the fronds of the Gelidium sesquipedale (Clem.) Born. et Thur. population located in Albufeira (southern Portugal) were reproductive, about 90% of these fronds were tetrasporophytes, whereas an equal percentage of female and male gametophytes was found (5%). The comparison of physiological performances of the reproductive phases (males, females and tetrasporophytes) did not reveal a physiological advantage of tetrasporic fronds. There were no significant differences either in the photosynthesis, nitrogen uptake, nitrate reductase activity, or biochemical composition of adult fronds. On the other hand, vegetative recruitment and spore production in the laboratory were significantly different. The re‐attachment to calcareous substrate and the subsequent rhizoidal growth were faster in tetrasporophytes. Particular levels of temperature, rather than irradiance, had an important effect on the phase differences in the spore release, attachment, and germination rates. Significant results were the higher release of carpospores at all irradiances at 17°C, and the higher attachment percentage of carpospores at 13°C versus tetraspores. Under higher temperatures (21°C), tetraspores showed higher attachment rates while carpospores germinated more. G. sesquipedale cystocarps released carpospores for 2 months, while tetrasporangia stopped shedding tetraspores after 1 month, resulting in a 3‐fold higher production of carpospores than tetraspores. Results showed that vegetative and spore recruitment may explain the low gametophyte–tetrasporophyte ratio of the studied population of G. sesquipedale as opposed to the physiological performance of phases.     

Differential responses of tetrasporophytes and gametophytes of Mazzaella laminarioides (Gigartinales,Rhodophyta) under solar UV radiation

The effects of solar UV radiation on mycosporine‐like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.