Modeling competition between Laurencia obtusa (Ceramiales,Rhodophyta) and Hypnea spinella (Gigartinales,Rhodophyta) at Cabo Frio Island,Rio de Janeiro,Brazil

The Lotka-Volterra competition model was used to represent the interaction between Laurencia obtusa and Hypnea spinella. A new model that considers effects of competition on algal carrying capacity is suggested. To test the models, data from field experiments conducted in an intertidal region at Cabo Frio Island, Rio de Janeiro, Brazil, were used. Both models showed that Hypnea was a stronger competitor than Laurencia. The model of interaction through the carrying capacity showed a stable coexistence between the algal populations and better represented the experimental data.     

Brominated sesquiterpene metabolites of Hypnea pannosa (Gigartinales,Rhodophyta)

A chloroform extract of freeze-dried Hypnea pannosa J. Ag., collected from the intertidal region near Karachi, Pakistan, furnished three bromine-containing secondary metabolites. These were characterised, using spectroscopic techniques, as 10-bromo-7, 12-dihydroxy-Δ^{3, 4}-laurene, filiformin and filiforminol. This is the first report of halogenated terpenoids from the genus Hypnea. author for correspondence     

Effect of ammonium on growth and carrageenan content in Kappaphycus alvarezii (Gigartinales,Rhodophyta)

Kappaphycus alvarezii, cultured in a region deficient in nitrogen with intermittent application of 10 mM ammonium at three-day intervals for 1 hour, attained a daily growth rate of 4.6%, which was considered to be the optimum growth rate. Under such conditions, the C/N ratio was 29 with a carrageenan content of 58% and gel strength of 45–70 g cm^–2. Gel strength of carrageenan was high when nitrogen content was high and vice versa.     

Effects of nitrogen and phosphorus availabilities on growth, pigment, and protein contents in Hypnea cervicornis J. Agardh (Gigartinales, Rhodophyta)

The selection of seaweed species for their use as biofilters should be based on the knowledge of their nutrient requirements and tolerance to wide variations of nutrient concentrations. Therefore, tolerance and the physiological capabilities of Hypnea cervicornis J. Agardh (Gigartinales, Rhodophyta) to growth under nitrate, ammonium, and phosphate variations and to assimilate them into soluble proteins and photosynthetic pigments were evaluated in laboratory conditions. Treatments were composed of sterilized seawater enriched with 25 % von Stosch solution (without nitrogen and phosphorus), and nitrate or ammonium and phosphate were added in combination of 100:1 and 10:1 nitrogen/phosphorus (N/P). Nitrate concentrations varied from 0 to 500 μM, and ammonium concentrations varied from 0 to 50 μM. Growth rates of H. cervicornis increased linearly with addition of ammonium, but with nitrate addition, growth varied following a saturation kinetic, and the highest growth rate (14.45 % d^?1) was observed in 200 μM of N/P ratio of 10:1. An excess of nutrients was accumulated as proteins and phycobiliproteins (mainly as allophycocyanin and phycoerythrin) at higher phosphate availability (N/P ratio of 10:1), and H. cervicornis tolerated the highest ammonium and nitrate concentrations (50 and 500 μM, respectively). These physiological responses suggest that this species could be used as biofilter for nutrient removal in eutrophicated seawater and could be cultivated in integrated multitrophic aquaculture systems.     

Effects of increased atmospheric CO2 and N supply on photosynthesis, growth and cell composition of the cyanobacterium Spirulina platensis (Arthrospira)

The consequences of the addition of CO2 (1%) in cultures of S. platensis are examined in terms of biomass yield, cell composition and external medium composition. CO2 enrichment was tested under nitrogen saturating and nitrogen limiting conditions. Increasing CO2 levels did not cause any change in maximum growth rate while it decreased maximum biomass yield. Protein and pigments were decreased and carbohydrate increased by high CO2, but the capability to store carbohydrates was saturated. C:N ratio remained unchanged while organic carbon released to the external medium was enhanced, suggesting that organic carbon release in S. platensis is an efficient mechanism for the maintenance of the metabolic integrity, balancing the cell C:N ratio in response to environmental CO2 changes. CO2 affected the pigment content: Phycocyanin, chlorophyll and carotenoids were reduced in around 50%, but the photosynthetic parameters were slightly changed. We propose that in S. platensis CO2 could act promoting degradation of pigments synthetised in excess in normal CO2 conditions, that are not necessary for light harvesting. Nitrogen assimilation was significantly not affected by CO2, and it is proposed that the inability to stimulate N assimilation by CO2 enrichment determined the lack of response in maximum growth rate. This revised version was published online in June 2006 with corrections to the Cover Date.     

Culture of Hypnea musciformis (Rhodophyta,Gigartinales) on artificial substrates attached to linear ropes

Hypnea musciformis is the only species so far exploited in Brazil as raw material for the production of k-carrageenan. Due to the erratic production in space and time, increasing harvest and transportation costs, experiments have been performed in order to assess the viability of H. musciformis mariculture.In nature the species occurs as an epiphyte, and so mariculture using artificial substrates that simulated the natural host of the species was tried. These substrates were attached, at regular intervals, to linear ropes. In the sea, these ropes were stretched between cement blocks.Seeding occurs naturally, by means of spores, or detached pieces of H. musciformis scattered in the water column that get entangled on the ropes. The best yields (0.54 wet kg m^–1 month^–1) were obtained with unthreaded rope substrates maintained in a vertical position by small rafts. Production is highest in the first 18 m off the rocky shore (0–2.1 m deep), at the highest substrate density utilized (10 m^–1), 2–3 months after installing the ropes in seawater. The main factor controlling seasonal production is water movement.     

CO2和硝氮加富对龙须菜(Gracilaria lemaneiformis)生长、生化组分和营养盐吸收的影响

为了探讨大气CO2和海水硝氮浓度升高对龙须菜生理特性的影响,在实验室条件下把藻体于CoNo、C No、CoN 和C N 4种不同CO2和硝氮供应水平的处理中进行培养,15d后,分别测定各种不同培养条件下藻体的生长、色素含量、可溶性蛋白含量和营养盐吸收速率.结果表明:CO2或硝氮浓度增加都会使藻体生长加快,但当二者同时加富时对生长的促进作用没有表现出协同效应.不管是高氮还是低氮的培养条件,高浓度CO2都能使藻体的色素和可溶性蛋白的含量降低,而使硝酸还原酶活性升高;同时,在硝氮加富的条件下,高浓度CO2培养能够显著提高藻体的营养盐吸收速率.不管是高还是低的CO2浓度培养条件下,硝氮浓度增加都会使藻体的色素和可溶性蛋白含量以及营养盐吸收速率得到显著提高.这些结果说明,在大气CO2浓度升高或海水富营养化条件下,龙须菜的生长、硝酸还原酶活性和营养盐吸收速率都可能得到促进,而色素和可溶性蛋白的含量也会受到影响.     

Influences of cadmium on fine structure and metabolism of Hypnea musciformis (Rhodophyta, Gigartinales) cultivated in vitro

The in vitro effect of cadmium on apical segments of Hypnea musciformis was examined. Over a period of 7?days, the segments were cultivated with different concentrations of cadmium, ranging from 50 to 300???M. The samples were processed for microscopic and histochemical analysis of growth rates, content of photosynthetic pigments, and photosynthetic performance. Cadmium treatments increased cell wall thickness and the accumulation of plastoglobuli. Destruction of chloroplast internal organization was observed. Compared to controls, algae exposed to cadmium showed growth rate reduction, depigmentation, and blending in the lateral branches. The content of photosynthetic pigments, including chlorophyll a and phycobiliproteins, decreased after exposure to different concentrations of cadmium. These results agree with the decreased photosynthetic performance and relative electron transport rate observed after exposure of algae to cadmium. Taken together, these findings strongly indicate that cadmium negatively affects the architecture and metabolism of the carragenophyte H. musciformis, thus posing a threat to the economic vitality of this red macroalgae.     

Seasonal and spatial patterns of population density in the marine macroalga Mazzaella splendens (Gigartinales, Rhodophyta)

Insight into demographic processes that operate at larger spatial scales can be achieved through studying local populations when a particular species of interest is examined over time, by many investigators, in a variety of locations. On the west coast of North America, Mazzaella splendens (Setchell et Gardner) Fredericq is such a species of interest. A synthesis of local demographic studies of M. splendens from the late 1960s to the present reveals a pattern that is potentially common to the larger natural populations. This is the pattern: population density is high in summer and low in winter for both alternate free‐living life history phases of M. splendens. The magnitude of this seasonal change decreases in increasingly wave‐exposed habitats. In wave‐sheltered habitats there is a seasonal alternation from summer haploid to winter diploid dominance. This alternation gradually changes to constant diploid dominance as wave exposure in the habitat increases. Changes in population density are primarily a function of appearances and disappearances of perennating basal crusts (genets), as modules are produced or lost, rather than differential module production by genets of one phase over those of the other. To test the generality of this pattern, we examined seasonal changes in density, in local populations of M. splendens, in both a wave‐sheltered and a wave‐exposed habitat at Second Beach, Barkley Sound. Greater seasonal fluctuation in population density at wave‐sheltered, compared to wave‐exposed habitats is supported as a pattern potentially common to the natural populations of M. splendens. A change from summer haploid dominance in wave‐sheltered areas to summer diploid dominance in wave‐exposed areas is similarly supported. All changes in population density were the result of appearances and disappearances of genets rather than differential module production by haploid versus diploid basal crusts, also consistent with previous observations. A seasonal alternation in phase dominance, however, was absent from the wave‐sheltered site at Second Beach, Barkley Sound for 3 consecutive years. Seasonal alternation in phase dominance of M. splendens appears dependent on local conditions and is not common to all natural populations.     

Management of natural beds and standing stock evaluation of Hypnea musciformis (Gigartinales, Rhodophyta) in south-eastern Brazil

Hypnea musciformis, a source of-carrageenan, is abundant along the coast ofBrazil. Although the species has been exploitedsporadically on the north-east coast, this has beendone in the absence of data on seasonal fluctuationsin stock and on management of the natural beds. Thispaper evaluates the stock of a population from theUbatuba Bay, S-E Brazilian Coast, and how thepopulation recovers from monthly harvests. Dry monthlyyield ranged from 4.9 to 201.2 g m^-2 and showedno obvious seasonal pattern. The mean recovery rateafter harvesting is estimated as 35 days, and thestanding stock in the Bay (1.5 km) is approximately3.68 t y^-1 dry wt.     

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.     

A taxonomic revision of a Hypnea charoides-valentiae complex (Rhodophyta, Gigartinales) in Japan, with a description of Hypnea flexicaulis sp. nov.

Morphology of a Hypnea charoides‐valentiae complex (Rhodophyta, Gigartinales) in Japan is investigated on the basis of field‐collected and cultured plants. We can recognize two species, Hypnea charoides Lamouroux and H. flexicaulis Yamagishi et Masuda, sp. nov., which are distinguished by: (i) conspicuous percurrent main axes growing straight in the former versus less conspicuous percurrent main axes growing flexuosely in the latter; (ii) narrow branching angles (less than 90°) in the former versus wide branching angles (up to 150°) in the latter; (iii) the presence of branches and branchlets showing abrupt adaxial bending in the former versus those showing pronounced abaxial bending, some growing into hooked branchlets in the latter; and (iv) the absence (in the former) or presence (in the latter) of antler‐like branches. Nucleotide sequences of the plastid‐encoded large subunit of the ribulose‐1,5‐bisphosphate carboxylase/oxygenase gene (rbcL) support the differentiation of these two entities: the pairwise distance between them was 83 b.p. (6.2%).     

Effects of plant growth regulators on growth and morphogenesis in tissue culture of Chondracanthus chamissoi (Gigartinales,Rhodophyta)

Chondracanthus chamissoi (C. Agardh) Kützing (Gigartinales, Rhodophyta) is an edible species and commercialised for carrageenan production in Chile. Investigations on growth and development are needed to improve its cultivation; therefore, this study aims to evaluate the effects of plant growth regulators (PGRs) on its growth and morphogenesis. PGRs tested were two auxins [indole-3-acetic acid (IAA) and 2,4-dichlorophenoxyacetic acid (2,4-D)] and one cytokinin (benzylaminopurine (BA)) in concentrations of 0.5, 5.0 and 50.0 μM. These PGRs were added to seawater enriched with half strength of von Stosch solution and were gelled with 0.6 % agar, and treatment control (without PGR addition) was conducted simultaneously. Apical and intercalary segments were used as initial explants. Each treatment was tested with six replicates of five axenic explants, and statistical analyses were performed. After culturing in a solid medium for 10 weeks to induce growth and callus formation, explants were cultured in liquid medium with the same experimental conditions for 10 weeks. Effects of auxins (IAA and 2,4-D) and the cytokinin BA on growth rates of apical segments of C. chamissoi were not significant, while low concentration of IAA stimulated the growth of intercalary segments. On the other hand, high concentrations of BA and IAA stimulated the callus formation in apical and intercalary segments, respectively. In liquid medium, PGR did not have a significant effect on growth rates of apical segments, while 2,4-D in concentrations from 0.5 to 50.0 μM stimulated growth of intercalary segments, and formation of lateral branches was stimulated by low 2,4-D in apical segments. These results suggest that PGRs have a regulatory role on callus formation and growth of specific explants of C. chamissoi. Furthermore, the formation of lateral branches stimulated by auxin could be used for seedling production under controlled conditions and could improve the micropropagation and cultivation of C. chamissoi in the Chilean coast.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

Effects of environmental factors on carrageenan fromGymnogongrus griffithsiae (Gigartinales,Rhodophyta)

The effects of environmental factors on substratum coverage, thallus length, production of iota carrageenan and gelling behavior were investigated in field populations of the red algaGymnogongrus griffithsiae (Turner) Martius. Irradiance and water temperature affected thallus substratum coverage and water gel strength of the carrageenan. The significant monthly differences in thallus length, carrageenan content and carrageenan gelling properties were not attributable to seasonal variations in irradiance and water temperature. The effect of temperature and N loading on biomass productivity, carrageenan yield and gelling characteristics were also investigated using cultures. N-enriched cultures showed decreased gel yields, higher viscosities and higher water gel strengths in the carrageenans extracted from these cultures. Milk reactivity tests were correlated with thallus nitrogen and water dessert gel strength, but neither culture temperature nor N loading showed a significant effect. The field and culture studies indicate how temperature, irradiance and N availability affect growth and hydrocolloid production.Author for correspondence     

Phylogeography of the Hypnea musciformis species complex (Gigartinales,Rhodophyta) with the recognition of cryptic species in the western Atlantic Ocean

Populations of the marine benthic red macroalgae Hypnea musciformis and Hypnea pseudomusciformis along the Atlantic and Pacific Oceans were tested for phylogeographic structure using the DNA barcode COI‐5P combined with rbcL for the construction of the phylogenetic tree. Strong patterns of genetic structure were detected across 210 COI‐5P DNA sequences, and 37 COI‐5P haplotypes were found, using multiple statistical approaches. Hypnea musciformis was found in the Northeast and Northwest Atlantic, the Mediterrean Sea, Namibia, and along the Pacific coast of Mexico. Two new putative species were detected, Hypnea sp. 1 in the Caribbean Sea and Hypnea sp. 2 in the Dominican Republic. Three distinct marine phylogeographic provinces were recognized in the Southern Hemisphere for H. pseudomusciformis: Uruguay, South‐Southeast Brazil, and Northeast Brazil. The degree of genetic isolation and distinctness among these provinces varied considerably. The Uruguay province was the most genetically distinct, as characterized by four unique haplotypes not shared with any of the Brazilian populations. Statistically significant results support both, isolation by distance and isolation by environment hypotheses, explaining the formation and mantainance of phylogeographic structuring along the Uruguay‐Brazil coast. Geographic, taxonomic and molecular marker concordances were found between our H. pseudomusciformis results and published studies. Furthermore, our data indicate that the Hawaiian introduced populations of H. musciformis contain Hypnea sp. 1 haplotypes, the current known distribution of which is restricted to the Caribbean.     

Influence of tissue source and growth rates on dry weight and carrageenan composition of Chondrus crispus (Gigartinales,Rhodophyta)

Carrageenan analyses were conducted on vegetative female clones of Chondrus crispus that were cultured to provide tissues with differing growth rates. Tissue dry weights increased from apex to base of fronds. Total carrageenan contents were lower in apical 1 to 2 cm segments than elsewhere in the frond, except when the alga was grown at high photon irradiances. Clone 373A contained more carrageenan than clone G8. The proportion of 0.3 M KCl-soluble polymers in the total native carrageenans varied from 44 to 92%, being highest in older tissues of fronds cultured at high photon irradiances. The apical 1 cm segments contained less KCl-soluble carrageenans than other tissues from the corresponding fronds. The KCl-soluble carrageenans, when alkali-modified and refractionated, afforded the expected kappa-iota carrageenan in > 79% yields. The remainder consisted of a polymer containing 23.1% SO₃Na and 8.4% 3,6-anhydrogalactose. Lambda carrageenan was not detected. Variations in carrageenan distribution between the apical region and other parts of the frond may reflect the increasing influence of medullary tissue developed as the immature cells differentiate.     

Effects of CO(2) enrichment on photosynthesis, growth, and biochemical composition of seagrass Thalassia hemprichii (Ehrenb.) Aschers

The effects of CO2 enrichment on various ecophysiological parameters of tropical seagrass Thalassia hemprichii(Ehrenb.)Aschers were tested.T.hemprichii,collected from a seagrass bed in Xincun Bay,Hainan island of Southern China,was cultured at 4 CO2(aq)concentrations in flow-through seawater aquaria bubbled with CO2.CO2 enrichment considerably enhanced the relative maximum electron transport rate(RETRmax)and minimum saturating irradiance(Ek)of T.hemprichii.Leaf growth rate of CO2enriched plants was significantly higher than that in unenriched treatment.Nonstructural carbohydrates(NSC)of T.hemprichii,especially in belowground tissues,increased strongly with elevated CO2(aq),suggesting a translocation of photosynthate from aboveground to belowground tissues.Carbon content in belowground tissues showed a similar response with NSC,while in aboveground tissues,carbon content was not affected by CO2 treatments.In contrast,with increasing CO2(aq),nitrogen content in aboveground tissues markedly decreased,but nitrogen content in belowground was nearly constant.Carbon: nitrogen ratio in both tissues were obviously enhanced by increasing CO2(aq).Thus,these results indicate that T.hemprichii may respond positively to CO2-induced acidification of the coastal ocean.Moreover,the CO2-stimulated improvement of photosynthesis and NSC content may partially offset negative effects of severe environmental disturbance such as underwater light reduction.     

Metabolic and cellular alterations induced by diesel oil in Hypnea musciformis (Wulfen) J. V. Lamour. (Gigartinales,Rhodophyta)

Understanding the mechanisms by which marine organisms cope to environmental stressful conditions is a fundamental question for ecotoxicology. We examined biochemical and cellular responses of Hypnea musciformis exposed in vitro to four concentrations of diesel oil (0 (control), 0.001, 0.01, 0.1, and 1 % v/v) for 0, 30 min, 1, 12, and 24 h. Chl a content decreased after exposure for 1, 12, and 24 h. Carotenoids increased after 30 min and 12 h and decreased after 1 and 24 h. Phenolic content was lower in samples exposed to 0.1 and 1 % of diesel oil. After treatment, samples showed a decrease in floridean starch grains, cell volume and autofluorescence intensity, changes in chloroplast morphology, and on the surface topography of the cell wall. These findings provide preliminary but important results on biochemical and cellular responses of H. musciformis when exposed to diesel oil and could be considered as tools for monitoring oil spills.     

Effects of auxins and cytokinins on tissue culture of Grateloupia dichotoma (Gigartinales,Rhodophyta)

The role played by plant growth regulators in algae is poorly known. In order to increase the knowledge about the function of auxins and cytokinins in seaweeds, explants such as apical and intercalary segments and callus-like structures (CLS) of Grateloupia dichotoma were cultured in semi-solid or liquid artificial media ASP 12-NTA. Two auxins, indole-3-acetic acid (IAA) and 2,4-dichlorophenoxyacetic acid (2,4-D), and one cytokinin, 6-benzylaminopurine (BA), at concentrations of 0.5 and 5.0 mg l^–1 were tested. Moreover, IAA and BA were tested together at concentrations of 1:5 and 5:1 mg l^–1. All treatments promoted the growth of CLS in intercalary segments; CLS from apical segments were significantly higher in treatments with 2,4-D or IAA:BA (1:5 mg 1^–1). The morphogenetic responses for auxins and BA were opposite, auxins inhibited while BA promoted the formation of lateral branches; however, auxins promoted the elongation of such branches. The process of plant regeneration observed on CLS was stimulated significantly by treatment with high concentration of BA or IAA:BA (1:5 mg 1^–1) in semi-solid and liquid media. The growth of upright axes was stimulated significantly by treatment with 2,4-D in semi-solid medium, and IAA:BA (1:5 mg l^–1) in liquid medium. These results show the importance that plant growth regulators could have in the control of growth, morphogenetic processes and micropropagation in red algae.This paper is part of the PhD thesis of NSY.