Demographic models to simulate the stable ratio between ecologically similar gametophytes and tetrasporophytes in populations of the Gigartinaceae (Rhodophyta)

In populations of the Gigartinaceae (Rhodophyta), gametophytes often predominate numerically over tetrasporophytes. Several hypotheses have been proposed to explain this dominance, based on the usually implicit assumption that the stable ratio between gametophytes and tetrasporophytes (G:T ratio) should be 1 if both reproductive phases are ecologically similar. We developed demographic models to test this assumption, for which we considered that both phases are ecologically similar. Defining ecologic similarity for most demographic rates is relatively straightforward, except for rates of spore output. The first set of models considered the same spore output per thallus of both phases as representing ecologic similarity. Model iterations led to stable G:T ratios of 1 for triennial and for perennial thalli, regardless of the initial G:T ratio, but not for annual thalli with initial G:T ratios different from 1. However, equal spore output may not represent ecologic similarity, due to size differences between carpospores and tetraspores. The second set of models considered the lowest possible spore output for each phase, according to the life history of this family: only one carposporangium, with one carpospore, is produced from every two gametophytes and only one tetrasporangium, with four tetraspores, is produced by every tetrasporophyte. Model iterations led to stable G:T ratios of 2.8 for most cases, a ratio of 1 being obtained only every 2 years for annual thalli with an initial G:T ratio of 1. Increasing absolute spore output, without altering the relative output between phases and incorporating density-independent mortality through a matrix model, given the same mortality rate for both phases, did not modify results. We suggest that the combination of both modeling and field research may uncover more rapidly than otherwise the most relevant ecologic differences between phases, if any, that underlie the G:T ratio observed for a given population.     

Persistence of gametophyte predominance in Chondrus crispus (Rhodophyta,Gigartinaceae) from Nova Scotia after 12 years

Gametophytes predominated clearly over tetrasporophytes in an intertidal population of Chondrus crispus at Tor Bay (Nova Scotia, Canada) in the summer of 1991. Since this species is perennial and the rocky substrate is stable at this site, we predicted that gametophyte predominance would persist after several years. We confirmed this hypothesis by re-sampling the same area in the summer of 2003. This is one of the first long-term studies of the relative abundance of life-history phases done unequivocally at the same site for the Gigartinaceae.     

GENETIC POPULATION STRUCTURE AND MATING SYSTEM IN CHONDRUS CRISPUS (RHODOPHYTA)1

Chondrus crispus Stackh. has been intensely studied, yet no study to date has elucidated its population structure or mating system despite many populations in which there was a haploid bias and lack of male gametophytes. Therefore, 12 nuclear microsatellite loci were identified in this red alga. Microsatellite markers were developed and tested against a panel of specimens collected from two shore levels at two sites in Brittany, France: Pointe de Primel and Pointe de la Jument, Concarneau. Single locus genetic determinism was verified at eight polymorphic loci, as only one band was observed for haploid genotypes, whereas one or two bands were observed for diploids. These markers enabled the detection of unique genotypes within sampled populations, indicating that very few fronds shared the same multilocus genotype. This finding suggests that asexual reproduction was not the prevailing mode of reproduction. In addition, we explored the hierarchical population structure showing that gene flow is restricted at small spatial scales (<50 m) between upper and lower Chondrus‐range populations within a shore. Sexual reproduction predominated in the populations of C. crispus studied, but probably due to fine‐scale spatial substructuring, inbreeding was also significant. In conclusion, this study reveals that fine‐scale genetic variation is of major importance in C. crispus, suggesting that differences between microhabitats should be essential in understanding evolutionary processes in this species.     

Spatial and temporal variations of Chondrus crispus (Gigartinaceae, Rhodophyta) carrageenan content in natural populations from Galicia (NW Spain)

Qualitative and quantitative differences in carrageenan composition of gametophytes and tetrasporophytes of Chondrus crispus were observed in this study. Carrageenans in gametophytes belong to the kappa family (κ-, ι-, ν- and μ-carrageenan). The dominant fractions were κ- and ι-carrageenan (more than 50% of the total carrageenans). In tetrasporophytes, the presence of λ-carrageenan was confirmed. Carrageenan content in gametophytes (37.4?±?1.68% DW) was higher than in tetrasporophytes (29.13?±?0.76% DW). Spatial and temporal variation in carrageenan content in both life cycle phases appears to be related mainly to seawater and air temperatures, insolation, water movement and desiccation. The highest values of carrageenan content were recorded in those localities where higher values of precipitation, wind speed or water movement occurred. A bimodal temporal pattern on carrageenan content was observed. Fronds showed a high carrageenan content in spring and autumn. During these seasons, the content was over 40% in gametophytes and 30% in the tetrasporophytes. In summer and winter, these values down in both life cycle phases below 30%. In general the highest carrageenan contents were related to highest seawater temperatures. On the contrary, high air temperature and high insolation appeared to be unfavourable for carrageenan production. GLM models were obtained to predict carrageenan production from natural C. crispus populations, along Galician coast.     

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.     

Taxonomic reassessment of Chondrus verrucosus (Rhodophyta,Gigartinales), with a description of Chondrus retortus sp. nov

Phylogenetic and morphological reinvestigation of Chondrus verrucosus Mikami was conducted in the Pacific coast of southeastern Honshu, Japan, including the holotype specimen and samples near the type locality. Two entities were recognized by phylogenetic analyses of plastid rbcL and nuclear EF2 sequences. In morphological analyses, the two entities were distinguished by: (i) thallus size and form, and (ii) cell shape of the medullary layer surrounding young carposporangia: one has larger thalli, is regularly canaliculated, with linear medullary cells (large‐type); the other is smaller, canaliculated with crispation and has roundish medullary cells (small‐type). The large‐type was common in southern to eastern Boso Peninsula including near the type locality; the small‐type was typical in the southwest Boso Peninsula and Sagami Bay. The large‐type was shown to be equivalent to C. verrucosus by examination of the holotype specimen. The small‐type is described as a new species, C. retortus Matsumoto et Shimada.     

Population studies on a South African carrageenophyte: Iridaea capensis (Gigartinaceae,Rhodophyta)

An intertidal population of Iridaea capensis in the south-western Cape Province of South Africa has been studied in detail for a year. The species is generally restricted to sand-inundated rocky shores, which appears not to be the case for Iridaea species in the Pacific. In early winter (June) biomass, plant density and plant size were at a minimum. A small increase in these parameters occurred in August, but there was a massive increase in plant numbers and biomass in October. The largest plants were recorded in summer (February). The population was over 70% sporophytes in April, but switched to almost 80% gametophytes following heavy recruitment between August and October, and remained significantly gametophyte-dominated through April of the following year. Carrageenan levels were high (ca 42% of dry weight) in late winter-spring, and lowest (ca 30–35%) in summer-autumn. Strong offshore wind conditions in early summer caused a reduction in biomass, and it is hypothesized that the associated upwelling of nutrient rich water may be implicated in the lower carrageenan levels at this time.     

Comparison of three techniques for identifying isomorphic phases of <Emphasis Type="BoldItalic">Chondrus crispus</Emphasis> (Gigartinaceae)

Chondrus crispus, a member of the economically important family Gigartinaceae, alternates between two free-living isomorphic life history phases. The carrageenan composition of the gametophyte and tetrasporophyte phases has been used to identify non-reproductive material, and thus provide a better understanding of the ecology of the species. The aim of this study was to compare three methods for identifying the isomorphic phases. The results obtained by the widely used resorcinol test and an iridescence method are compared with those obtained by a technique that involves the acquisition of mid-infrared absorbance spectra from dried samples by Fourier transform infrared spectroscopy (FT-IR). Both the resorcinol test and FT-IR produced similar results with a calculated gametophyte to tetrasporophyte ratio of 2.1 and 2.3:1, respectively. In contrast the iridescence method was less reliable, with a calculated gametophyte to tetrasporophyte ratio of 1.3:1 and a low predictive value (0.70) for selecting tetrasporophytes. The FT-IR technique provides a rapid assessment, does not involve the use of hazardous chemicals, has the potential to be applied to members of the Gigartinaceae for which the resorcinol and iridescence tests are less applicable and it has greater flexibility in that it can identify additional carrageenan fractions.     

Carrageenans from Cystocarpic and Sterile Plants of Chondrus Pinnulatus (Gigartinaceae, Rhodophyta) Collected from the Russian Pacific Coast

The chemical structure, gel properties and biological activity of the carrageenans isolated from cystocarpic and sterile plants of Chondrus pinnulatus were investigated. The total carrageenan content of the sterile plant was observed to be twice that of the cystocarpic plants. According to data obtained by ¹³C-NMR and FT IR, the gelling polysaccharides from cystocarpic and sterile plants of C. pinulatus have similar structures and were identified as κ/ι-carrageenans. The difference between these polysaccharides was in the ratio of the κ- and ι-segments, with a predominant content of κ-segments in cystocarpic plants (80%). Moreover, KCl-insoluble fractions possibly contain hetero-disperse μ/ν precursor: amounts of this in the polysaccharide from sterile plants were more than that extracted from the cystocarpic plants. The KCl-soluble fractions (non gelling) were λ-carrageenans with another carrageenan type that had a low amount of 3,6-anhydrogalactose. Carrageenans from cystocarpic stages showed good gelling properties, whereas those from sterile plants formed a very weak gel. Structural differences and molecular weight of carrageenans obviously determine the biological activity of the polysaccharides. Non gelling-carrageenans from both types of ιt C. pinnulatus plants showed high macrophage-phosphatase activity and κ/ι-carrageenan from cystocarpic plant possessed a potent anti-coagulant activity, which was extremely strong in a low concentration of 100 μg ml⁻¹.     

Reliability of the resorcinol method for identifying isomorphic phases in the Gigartinaceae (Rhodophyta)

The Gigartinaceae are economically important because the carrageenans in this family are used for a variety of purposes including food stabilizers, food substitutes, pharmaceutical applications and cosmetics. The resorcinol method takes advantage of the different carrageenans found in life history phases in this family to identify nonreproductive phases, and consequently, a better understanding of the biology of this economically important family is possible. This study investigates sources of variability that could affect the accuracy of the resorcinol method in identifying phases within the Gigartinaceae. Vegetative disks of Iridaea splendens from different areas of the blade and disks containing reproductive structures all reacted consistently to the resorcinol reagent. The minimum size and testing conditions required to reliably identify phases at the germling stage of I. splendens were determined. Phases of I. splendens cannot be identified if blades are stored in 3% formaldehyde in seawater. Phases of I. splendens, I. lineare, I. heterocarpa, Rhodoglossum californicum and R. affine can be identified reliably at different times of the year. Manipulations of the resorcinol method improved its accuracy in identifying phases of Iridaea cornucopiae and Gigartina exasperata but the high percent of G. exasperata misidentified demonstrates that it cannot be assumed that the current resorcinol method works for all taxa in the Gigartinaceae.     

Seasonal changes in abundance and shifts in dominance of life history stages of the carrageenophyte Sarcothalia crispata (Rhodophyta, Gigartinales) in south central Chile

The population dynamics of the carrageenophyte Sarcothalia crispatais described from subtidal beds at two localities in south-central Chile. Seasonal fluctuations in total density and biomass were not evident. Frondswere identified to phase by the presence of reproductive structures and theresorcinol reaction. The monthly changes in abundance of each kind offrond were determined. Permanent gametophytic or sporophyticdominance was not evident: the more exposed site showed a seasonal shiftfrom sporophytic dominance in summer to gametophytic dominance inwinter, whereas the more protected site showed an interannual shift fromgametophytic to sporophytic dominance. The differences between localitiesand years suggest a very local population dynamics with large contributionof self-seeding to the maintenance of the S. crispata beds.     

Patterns of gametophyte dominance ofIridaea splendens(Rhodophyta) in Vancouver Harbour,Vancouver, British Columbia,Canada

This paper presents results of field and laboratory work on one component of population structure ofIridaea splendens (Setchellet Gardner) Papenfuss (Gigartinaceae, Rhodophyta), the seasonal change in abundance of the gametophyte life-history stage. Investigations of this genus by others (using a variety of sampling and identification techniques) have shown a seasonal gametophyte dominance, sporophyte dominance, or some combination of these two. Gametophyte stages ofIridaea splendens produce kappa carrageenan and tetrasporophyte stages lambda carrageenan; a chemical colorimetric test using resorcinol can be used to identify these stages regardless of the presence of reproductive structures. In this paper we report on the proportion of gametophytes determined both by this chemical test and by the presence of reproductive structures, and on one possible determinant of the seasonal change in dominance. Analysis of field samples using the resorcinol test showed that from June–August the gametophyte stage predominates and from December–February the tetrasporaphyte stage is most common, both in reproductive and non-reproductive thalli (the latter tested as three size classes). Examination of reproductive structures gave similar results. One factor suggested in the literatures as a possible determinant of the haploid/diploid ratio is apomeiosis (in the formation of the tetraspores); our results show this to be unimportant in the Vancouver Harbor populations.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.     

Phenology of Gigartina skottsbergii (Gigartinaceae,Rhodophyta) in Ancud Bay,southern Chile

This study reports the variations in biomass and reproductive phenology in a natural bed of Gigartina skottsbergii over a period of a year and correlates these variations with the abiotic factors solar radiation, number of daylight hours, water temperature and salinity. The results obtained show an annual production cycle with maximum biomass values in spring-summer, correlated with high solar radiation and to a lesser extent with salinity. Sexual reproduction was maximum in autumn-winter, correlated with low temperature and short-day conditions. The population showed a large gametophytic dominance according to size and biomass, which suggests that there is not ecological equivalence between the life history phases of the species. Finally, based on the results, a restriction of harvesting to spring-summer is suggested.     

Chondrus crispus (Gigartinaceae,Rhodophyta) tank cultivation: optimizing carbon input by a fixed pH and use of a salt water well

The injection of exogenous carbon into intensively cultivated algal tanks is necessary to insure a maximum growth rate by stabilizing the dissolved inorganic carbon (DIC) pool, but represents the major part of the cultivation cost (ca. 73%). This study was conducted in paddle-wheel tanks ranging in size from 260 m² to 1000 m². Additional carbon was provided by carbon dioxide mixed into the incoming sea water through a tubular reactor. Production vs pH was analysed on 120 growth measurements covering two years of continuous cultivation. Whereas production peaked at pH 8.0–8.2, the economic optimum for pH regulation was in the range 8.4–8.5, where CO₂ injection was greatly reduced (–29%) for only a slight decrease in production (–4%). Expressed as a function of pH level, the specific carbon injection (g c gdw^–1 of Chondrus produced) showed an inverse exponential relationship, whereas gross photoconversion ratio (gdw mol photons^–1) varied according to a second degree equation with a low amplitude. The photoconversion ratio was not improved when the culture was maintained at a DIC concentration higher than the natural equilibrium (0.64 ± 0.11 gdw mol photons^–1 at 2.35 mM and 0.65 ± 0.15 gdw mol photons^–1 at 3.19 MM).A complementary source of carbon was found in underground salt water with a high and stable DIC concentration (10.15 ± 0.25 mmole Cl^–1). The mixing of the well water with natural sea water allowed another economy of CO₂ (–20% at pH 8.5) and nutrients (–12%), the total unitary cost of production being cut by about 17%.     

Ecology and morphological characterization of gametophyte and 'Chantransia' stages of Sirodotia huillensis (Batrachospermales, Rhodophyta) from a stream in central Mexico

The morphology and phenology of Sirodotia huillensis was evaluated seasonally in a central Mexican first‐order calcareous stream. Water temperature was constant (24–25°C) and pH circumneutral to alkaline (6.7–7.9), and calcium and sulfates were the dominant ions. The gametophyte stages were characterized by the presence of a distinctive mucilaginous layer, a marked difference in phycocyanin to phycoerythrin ratio between female and male plants, and the presence of a carpogonia with a large trichogyne (>60 µm). Occasionally three capogonia were observed on a single basal cell. The ‘Chantransia’ stages were morphologically similar to those described for the other members of Batrachospermales. A remarkable observation was the formation of dome‐shaped structures, consisting of prostrate filaments that are related with the development of new gametophytes. Chromosome numbers were n = 4 for fascicle cells, cortical filament cells and dome‐shaped cells, and 2n = 8 for gonimoblast filament cells and ‘Chantransia’ stage filaments. Gametophytes and ‘Chantransia’ stages occurred in fast current velocities (60–170 cm/s) and shaded (33.1–121 µmol photons/m²/s) stream segments. The population fluctuated throughout the study period in terms of percentage cover and frequency: the ‘Chantransia’ stages were most abundant in the rainy season, whereas gametophytic plants had the highest frequency values during the dry season. These results were most likely a result of fluctuations in rainfall and related changes in current velocity. Some characteristics of this population can be viewed as probable adaptations to high current velocities: the mucilaginous layer around plants that reduces drag; potential increase in fertilization by the elongate and plentiful trichogynes and abundant dome‐shaped structures producing several gametophytes.     

Etiology of infectious diseases in cultivated Chondrus crispus (Gigartinales,Rhodophyta)

The appearance of cavities and holes in fronds of commercially cultivated Chondrus crispus is described. These ultimately arise from the ravages of a green spot or green rot disease system in which several biotic agents can participate. Nematodes capable of bacterial grazing were recovered from necrotic lesions and we suggest that the nematodes can facilitate wound healing in diseased tissues. Bacteria isolated from disease lesions and from the surfaces of healthy fronds were screened for pathogenic strains. A particularly virulent one, the DOR isolate, was purified from small dark orange colored colonies grown on dilution plates. It was present in necrotic tissue and also recovered from surface scrapings of old healthy fronds, but not from their apical regions. Growth of the DOR isolate appeared to be inhibited by other bacteria colonizing the algal surface. It was shown to be a facultative pathogen, the virulence of which depended on the availability of ammonium or constituents easily metabolized to ammonium. It induced green rot disease in healthy C. crispus and was recoverable in pathogenic form from experimentally infected frond apices. A wound, disease and recovery cycle is discussed to illustrate potential interrelationships involving animal grazers, algal endophytes, Petersenia pollagaster, bacteria and nematodes.     

Ammonium uptake by Chondrus crispus Stackhouse (Gigartinales,Rhodophyta) in culture

Cultivated Chondrus crispus was used in N-NH₄ uptake experiments in the laboratory. An elevation of temperature increased the apparent rate of uptake, especially up to 11 °C. Uptake in the dark was found to be 83 % of that in the light. The apparent uptake decreased with increasing internal N pool; rates were 26.5, 22.2 and 20.2 µg N g dry wt^–1 min^–1 for internal N pools of 2.7, 3.5 and 4.6%, respectively. Apparent uptake increased with the substrate N concentration. The resulting curve has two components: an active uptake and a diffusion component at high (> 5000 µg N L^–1) external N levels. Ks and V _max were calculated by deducting the diffusion component from the uptake curve: these were of 497 µg N L ^–1 and 14.4 µg N g dry wt^–1 min^–1. respectively, and reflect a low substrate affinity. This could be the result of 10 years of continuous culture of C. crispus. Uptake was similarly followed in the culture tanks and showed comparable results; nighttime would be the most appropriate time to supply nutrients.     

Regeneration and reproduction of Mazzaella cornucopiae (Rhodophyta, Gigartinaceae) after frond harvesting

The effects that different intensities of frond harvesting have on frond regeneration and subsequent production of reproductive structures were investigated for the red intertidal alga Mazzaella cornucopiae (Postels & Ruprecht) Hommersand from British Columbia, Canada. Harvesting was done by pruning fronds in the late spring (when stand biomass is highest) of 1993 at two intensities: total and partial collection of fronds, in this second case leaving all frond biomass less than 1 cm high in place. Holdfasts were not damaged. Total percent cover of thalli, frond density, mean frond length, and stand biomass for these experimental quadrats were statistically similar to values for control quadrats in the spring of 1994. These results suggest that one total harvest of fronds per year, done in late spring without damaging holdfasts, may give the highest sustainable yield of biomass. The effects of harvesting intensity on reproduction were variable and difficult to explain. Neither the appearance nor the abundance of cystocarpic fronds were affected by frond pruning, compared with control areas, but pruning did affect the appearance and the abundance of tetrasporic fronds. Partial pruning resulted in a longer presence of tetrasporic fronds, whereas total pruning was associated with their complete absence. Results are compared with those for the few other species of the Gigartinaceae for which experimental harvesting has been done. This revised version was published online in August 2006 with corrections to the Cover Date.     

Relationship between cover of Chondrus crispus (Gigartinales,Rhodophyta) and Phymatolithon (Corallinales,Rhodophyta) on friable rock substrata

Field observations in sublittoral Prince Edward Island, Canada, indicated that on a friable sandstone substratum Chondrus crispus was more commonly associated with Phymatolithon sp. than with bare rock. Thus, a substantial proportion of the population of Irish moss along the coast of Prince Edward Island occurs on this encrusting coralline. These observation may be explained on the basis of the relative stability of the substratum in contrast to other studies in which sloughing of epithallial cells by species of both Phymatolithon and Lithothamnium has been reported to limit epiphytism by fleshy macrophytes.Issued as NRCC 31426.