Qualitative and quantitative analysis of carrageenan content in gametophytes of Mastocarpus stellatus (Stackhouse) Guiry along Galician coast (NW Spain)

Qualitative and quantitative differences in carrageenan composition of gametophytes of the rhodophyte Mastocarpus stellatus (Gigartinales) were observed in this study. Carrageenans in gametophytes belong to the kappa family (κ-, ι-, ν-, μ-carrageenan). The dominant fractions were κ- and ι-carrageenan (more than 80 % of the total carrageenans). Mean total carrageenan content in gametophytes was of 37.32?±?1.21 % DW. Spatial and seasonal variations were observed, mainly related to changes on environmental and oceanographic factors and the role of carrageenans in adapting the fronds to these changes. Maximum values in carrageenan content were observed for San Román (Biscay Bay) in May and for Laxe and Mougás (Atlantic coast) in June. The results of this study indicated that spatial differences in carrageenan content were due to interactions of different factors, rather than the effect of a single factor. Fronds from San Román had higher carrageenan content (43.23?±?1.87 % DW) than those collected at two sites of the Atlantic coast, Mougás and Laxe (32.20?±?1.14 % DW). San Román is exposed to the open sea, windy and oriented to the north, and the water temperature is higher in summer than in the Atlantic coast. However, seasonal variations in carrageenan content resulted to be more related to other factors directly correlated with the input of energy in the ecosystems (irradiance, sunshine hours and insolation). Thus, carrageenan content began to increase in early spring when the number of sunlight hours increased. Maximum values were reached in late spring or early summer, just before maximum values of irradiance and air temperature were achieved.     

Studies on carrageenans and effects of seawater phosphorus concentration on carrageenan content and growth ofAgardhiella subulata (C. Agardh) Kraft and Wynne (Rhodophyceae,Solieriaceae)

Gas liquid chromatography, chemical analyses, and infrared and¹³C-NMR spectroscopies indicated that phycocolloids extracted fromAgardhiella subulata had a dominant ι-carrageenan feature with less deviant ι-carrageenan and υ-carrageenan. The presence of methylated galactose and a small contamination by xylose were registered. Unattached plants were cultivated for 4 weeks in tanks receiving seawater enriched with 53.5 µM nitrate and 0 to 20 µM phosphate (P_i) week^?1. The growth was phosphorus (P)-limited up to a tissue P content of 0.14 ± 0.03% dry weight. Maximal specific growth rate and carrageenan content were observed with enrichments of 6 µM P_i and 3 µM P_i, respectively. Hence carrageenan production was promoted in the range of 3–6 µM P_i. Further P_i enrichment was useless. This phenomenon, observed with P nutrition, is comparable to the ‘Neish effect’ in nitrogen nutrition studies.     

Carrageenan patterns in the Gigartinaceae

Carrageenans of known reproductive phases of gigartinacean species are shown to differ: gelling κ (τ)-type carrageenan occurs in all instances in gametophytes, and viscous non-gelling λ-carrageenan or its variant ξ- or π-carrageenan in tetrasporophytes. Multiple analyses demonstrate that the carrageenan type is a species characteristic, with little difference between individual samples. No difference exists between the carrageenans of male and female plants of each species. Carrageenan chemistry combined with morphological, reproductive and life history feature is useful for defining the taxonomic limits of this family and these genera.     

Effect of epiphyte infection on physical and chemical properties of carrageenan produced by Kappaphycus alvarezii Doty (Soliericeae,Gigartinales, Rhodophyta)

Epiphytism of filamentous red algae is a serious problem in Kappaphycus farms in the Philippines, Indonesia, Malaysia, and Tanzania. The causative organism of epiphyte outbreak has been identified as Neosiphonia apiculata (Hollenberg) Masuda and Kogame, but its actual effect on carrageenan quality has not yet been established. Therefore, yield and quality of carrageenan from healthy and infected specimens were examined. Infected specimens showed 20.5?±?2.5 % DW lower carrageenan yield compared with the healthy seaweed (65.5?±?4.2 % DW). Infected specimens also had a higher phenolic and fatty acid content, compared with healthy specimens. The carrageenan from the infected seaweed showed 74.5?±?2.8 % lower viscosity, 52.6?±?3.6 % lower gel strength, 22.9?±?1.5 % higher syneresis, and 5 °C higher melting temperature as compared with carrageenan from healthy specimens. FTIR and ¹³C-NMR analysis of carrageenan from infected seaweed did not show any differences in their functionality or carbon atom chemical shift as compared with healthy and standard k-carrageenan. However, size exclusion chromatography showed the infected carrageenan molecular size to be 80 kDa as compared with 800 kDa for the healthy and standard k-carrageenan. These findings prove that infection of Kappaphycus by the filamentous red algae epiphyte, N. apiculata, reduces carrageenan molecular size and affects the physical properties of the carrageenan.     

Carrageenans in the gametophytic and sporophytic stages of Chondrus crispus

Summary The morphologically similar sporophytic and gametophytic plants of Chondrus crispus Stackhouse were examined and it was shown that the former contain -carrageenan. The gametophytes contain - and two additional carrageenans which are KCl-soluble and may comprise up to 25% of the total carrageenan. After alkaline modification, these KCl-soluble components were separated into a gel and a soluble carrageenan. The gel was indistinguishable from -carrageenan and presumably was derived from -carrageenan while the KCl-soluble fraction possessed a unique infrared spectrum easily distinguished from alkali-modified -carrageenan. This appears to represent a third carrageenan in the gametophytes.Our observations suggest that the biologically separate plants of C. crispus exhibit distinctive patterns of sulfation of their galactans. The sporophytes add SO₄ ^2- at C₂ of the precursor, whereas the gametophytes appear to add it principally at the available C₄ positions. Both types of plant are capable of sulfating at C₆ of the 4-linked galactose unit.Issued as NRC No. 13119.     

Controlling Carrageenan Structure Using a Novel Formylglycine-Dependent Sulfatase, an Endo-4S-iota-Carrageenan Sulfatase

Carrageenans are sulfated polysaccharides that are found in the cell walls of red algae. These polysaccharides have gelling and texturizing properties that are widely appreciated in industrial applications. However, these functional properties depend strongly on the sulfation of the moieties of the carrabiose repetition unit. Here we aimed to monitor the sulfate composition of gelling carrageenan. To do so, we screened and purified from Pseudoalteromonas atlantica a 4S-iota carrageenan sulfatase that converts ι-carrabiose into α-carrabiose units. The sequence of this protein matched the annotated Q15XH3 (Uniprot databank) formylglycine-dependent sulfatase found in the P. atlantica genome. With pure enzyme, ι-carrageenan could be transformed into a hybrid ι-/α-carrageenan or pure α-carrageenan. Analysis of the distribution of the carrabiose moieties in hybrid carrageenan chain using enzymatic degradation with Alteromonas fortis ι-carrageenase, coupled with chromatography and NMR spectroscopy experiments, showed that the sulfatase has an endo mode of action. The endo-character and the specificity of the sulfatase made it possible to prepare hybrid κ-/ι-/α-carrageenan and κ-/α-carrageenan starting from κ-/ι-carrageenan.     

Carrageenans, sulphated polysaccharides of red seaweeds, differentially affect Arabidopsis thaliana resistance to Trichoplusia ni (cabbage looper)

Carrageenans are a collective family of linear, sulphated galactans found in a number of commercially important species of marine red alga. These polysaccharides are known to elicit defense responses in plant and animals and possess anti-viral properties. We investigated the effect of foliar application of ι-, κ- and λ-carrageenans (representing various levels of sulphation) on Arabidopsis thaliana in resistance to the generalist insect Trichoplusia ni (cabbage looper) which is known to cause serious economic losses in crop plants. Plants treated with ι- and κ-carrageenan showed reduced leaf damage, whereas those treated with λ- carrageenan were similar to that of the control. In a no-choice test, larval weight was reduced by more than 20% in ι- and κ- carrageenan treatments, but unaffected by λ-carrageenan. In multiple choice tests, carrageenan treated plants attracted fewer T. ni larvae by the fourth day following infestation as compared to the control. The application of carrageenans did not affect oviposition behaviour of T. ni. Growth of T. ni feeding on an artificial diet amended with carrageenans was not different from that fed with untreated control diet. ι-carrageenan induced the expression of defense genes; PR1, PDF1.2, and TI1, but κ- and λ-carrageenans did not. Besides PR1, PDF1.2, and TI1, the indole glucosinolate biosynthesis genes CYP79B2, CYP83B1 and glucosinolate hydrolysing QTL, ESM1 were up-regulated by ι-carrageenan treatment at 48 h post infestation. Gas chromatography-mass spectrometry analysis of carrageenan treated leaves showed increased concentrations of both isothiocyanates and nitriles. Taken together, these results show that carrageenans have differential effects on Arabidopsis resistance to T. ni and that the degree of sulphation of the polysaccharide chain may well mediate this effect.     

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     

Seasonal variation of carrageenan yield,gel strength and viscosity in Sarcopeltis (ex Gigartina) skottsbergii from Southern Chile

Chile is one of the top carrageenan producers worldwide, and Sarcopeltis (ex Gigartina) skottsbergii one of the topmost exploited carrageenophytes from the wild in the world. Total yield, gel strength and viscosity from two contrasting environments Calbuco and Ancud (Inner and Outer Sea, Chile) were estimated monthly in approximately 2 years for this species. While carrageenan yields did not show differences between localities, gametophytes in spring–summer had 15% higher, compared to tetrasporophytes. Sizes (frond surface) normally did not affect carrageenan yields. Gametophytes showed clear differences in gel strength between seasons, but not between localities, with maximum peaks during winter–spring months in Calbuco and autumn-winter months in Ancud. Seasonal variations in viscosity were also significant. While gametophyte viscosity did not exceed 120 cPs, tetrasporophytes reached 1400 cPs in Calbuco and 1000 cPs in Ancud. More remarkably, a positive correlation between viscosity and gel strength was found in S. skottsbergii gametophytes, which is significantly different between both localities. These results suggest that selective harvesting in spring–summer should be preferred to optimize cost–benefit of harvesting activities and subsequent carrageenan productivity.     

Biomass variation and reproductive phenology of Gracilaria gracilis in a Patagonian natural bed (Chubut, Argentina)

The biomass variation and the reproduction of the natural Gracilaria gracilis bed in Bahía Bustamante (Patagonia, Argentina) were analyzed for 2 years, with the aim of determining the present situation of the population for an updated status overview; establishing the relevant features of the temporal variation in both biomass and reproductive states in relation to environmental factors, epiphytes and associated algae; and assessing carpospore availability for future spore-culture development. Field measurements and sampling were performed monthly between March 2006 and February 2008. In both years, G. gracilis biomass presented marked seasonal variations, with a minimum in winter and a maximum in late spring and in summer. During both years, coexistence of the three life-cycle phases was found, with dominance of tetrasporophytes. Two data sets from individuals originated from sexual reproduction (tetraspores and carpospores) and from asexual reproduction by thallus fragmentation were analyzed separately. In the fragmentation fraction, tetrasporophyte frequencies remained higher than those for gametophytes. However, in the spore-originated fraction, a generation ratio close to 0.5 was observed. Female gametophytes bearing cystocarps were always present, with a maximum in summer and autumn. Biological data were related to environmental factors by means of canonical correspondence analysis (CCA). The first year was characterized by higher biomass values of G. gracilis and Undaria pinnatifida, lower epiphytism, larger Gracilaria thalli and greater proportion of mature tetrasporophytes and gametophytes. The second year was characterized by a high proportion of Gracilaria vegetative thalli and high epiphyte density. The best time to obtain spores from cystocarpic thalli would be in summer and early autumn.     

Enzymatic degradation of hybrid ι-/ν-carrageenan by Alteromonas fortis ι-carrageenase

Hybrid ι-/ν-carrageenan was water-extracted from Eucheuma denticulatum and incubated with Alteromonas fortis ι-carrageenase. The degradation products were then separated by anion-exchange chromatography. The three most abundant fractions of hybrid ι-/ν-carrageenan oligosaccharides were purified and their structures were analyzed by NMR. The smallest hybrid was an octasaccharide with a ι-ι-ν-ι structure. The second fraction was composed of two decasaccharides with ι-ι-ι-ν-ι and ι-[ι/ν]-ι-ι structures. The third fraction was a mixture of dodecasaccharides which contained at least a ι-ι-ι-ι-ν-ι oligosaccharide. The carbon and proton NMR spectra of the octasaccharides were completely assigned, thereby completely attributing the ν-carrabiose moiety for the first time.     

Effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in Mazzaella parksii (Rhodophyta, Gigartinaceae) populations

We investigated the effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in populations of the intertidal red seaweed Mazzaella parksii (= M. cornucopiae, Gigartinaceae) from the Pacific coast of Canada. In July 2002, we determined the life-history phase of 864 frond samples from four populations from Barkley Sound, using the resorcinol-acetal test. For these populations, gametophytes were significantly more numerous than tetrasporophytes, and the proportion of gametophytes was significantly higher at higher elevations. The proportion of gametophytes varied directly (although less markedly) with the degree of wave exposure, but significance could not be assessed for this factor. All of these patterns were spatially consistent, as they held for two different rocky points with similar physical characteristics. One of these points had also been surveyed in 1994-1995, which allows us to conclude that the population-wise gametophyte predominance and the positive relationship between elevation and the proportion of gametophytes are stable features of this species in this area. However, the average proportion of gametophytes was similar between the studied levels of wave exposure in 1994–1995; such a difference revealed as non-significant when data were combined with those for 2002. This suggests that wave exposure is not important in determining the proportion of life-history phases for this species. Comparisons with other species are done in search of general patterns for the Gigartinaceae.     

Variation in gametophyte dominance in populations of Chondrus verrucosus (Gigartinaceae,Rhodophyta)

We describe the abundance, including spatial and temporal variability, of phases of the isomorphic Chondrus verrucosus Mikami from Japan. Chondrus verrucosus occurred in a dense (～90% cover) and temporally stable bed on a small, isolated rocky outcrop (Oyakoiwa) in Shizuoka Prefecture. Small vegetative fronds were always much more abundant than large vegetative and fertile fronds over the spring to late summer periods in 1999 and 2000. Over the same period, fertile carposporophytic fronds were generally more abundant than fertile tetrasporophytic fronds, and fertile male fronds appeared infrequently at low densities. Using the resorcinol‐acetal test, we determined the proportion of gametophytes and tetrasporophytes in three populations of C. verrucosus: Oyakoiwa and Noroshi (Shizuoka) in the summers of 1999 and 2000 and Kamehana Point (Miyagi) in autumn 2000. All populations had a significantly higher proportion of gametophytes than tetrasporophytes in both years, although gametophytic proportions were lower at Noroshi (～70%) than at Oyakoiwa (～80%) and Kamehana Point (～97%). However, examination of all isolated individuals sampled on Noroshi showed equal proportions of each phase in 1999, but gametophyte dominance (74%) in 2000. Differences in dispersal and spore production between phases are discussed as mechanisms potentially contributing to variation in gametophyte dominance.     

PHYSIOLOGICAL COMPARISON OF THE ISOMORPHIC LIFE HISTORY PHASES OF THE HIGH INTERTIDAL ALGA ENDOCLADIA MURICATA (RHODOPHYTA)1

The isomorphic phases of Endocladia muricata (Post. & Rupr.) J. Ag. Were compared for photosynthetic and respiratory difference in response to a variety of environmental manipulations. Photosynthetic light response during submergence at 15° C and the pattern of respiratory recovery following prolonged emergence (3 h) at either 15° or 30° C were similar between gametophytes and tetrasporophytes. The phases showed the same ability to photosynthesize and respire during emergence at each temperature tested (15°, 25°, and 35° C, fully hydrated thalli) and at various desiccation state (measured at 25° C only). Submerged rates of photosynthesis following prolonged emergence at 15° and 30° C were, however slightly greater (17%) for tetrasporophytes as compared to gametophytes. Regardless of the life history phase, plants incubated at 15° C during emergence recovered more completely than plants incubated at 30° C. Photosynthetic recovery after 1 h in plants incubated at 15° C often “spiked” and yielded rates as great as 185% of pretreatment rates. Increased photosynthetic rates during recovery were absent for the 30° C incubations. The initial photosynthetic recovery of plants collected from the upper limits of distribution was greater than that of plants collected from the lower limits. Recovered rates of respiration were highly variable over time. Respiration often exceeded pretreatment values more then threefold, and the elevated rates were sustained for 12 h. Photosynthesis and respiration in air were comparable to rates in seawater and varied slightly with increasing temperature. Photosynthetic and respiratory rates also decreased with increasing tissue water loss. Thus, only slight differences in physiological performance were observed between phases and individuals collected from different vertical positions. Metabolic differences were transient and apparent only under experimental conditions that modeled extreme environmental conditions.     

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

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

A comparative analysis of phycocolloids produced by underutilized versus industrially utilized carrageenophytes (Gigartinales, Rhodophyta)

Carrageenan (E-407) and semi-refined carrageenan (E-407a) are some of the main additives used by the food industry for their gelling, emulsifying, thickening, and stabilizing properties. These are natural ingredients, which have been used for decades in food applications and are generally regarded as safe. Internationally, sub-tropical carrageenophytes (e.g., Kappaphycus alvarezii) are cultivated extensively as a source of raw materials for industrial extraction, and their use as potential candidates in integrated multitrophic aquaculture is tentative. We analyzed carrageenan yield (as a percentage of dry weight) and chemical composition (by Fourier transform infrared attenuated total reflection and Fourier transform-Raman) of extracts produced by several carrageenophytes (Gigartinales, Rhodophyta), from different origins, e.g., K. alvarezii (Tanzania, Indonesia, the Philippines, Panama, and Mexico), Kappaphycus striatum (Madagascar), Eucheuma denticulatum (Tanzania, the Philippines, and Madagascar), Betaphycus gelatinum (the Philippines), Chondracanthus chamissoi, and Sarcothalia crispata (Chile). For comparison, some underutilized carrageenophytes were also analyzed, e.g., Chondrus crispus, Mastocarpus stellatus, Gigartina pistillata, Chondracanthus teedei var. lusitanicus, Chondracanthus acicularis, Calliblepharis jubata, Gymnogongrus crenulatus, and Ahnfeltiopsis devoniensis (Portuguese carrageenophytes). The main findings were that the highest carrageenan yield was obtained from K. striatum (Madagascar) with 75.6 (percent dry weight (% DW)); B. gelatinum and K. alvarezii (both from the Philippines) had yields of 71.0% and 68.0% (% DW), respectively; and G. pistillata (Portugal) 65.4% (% DW). Spectroscopic analysis of the phycocolloids allowed determination of a wide range of carrageenan types, e.g., pure iota carrageenan, several kappa–iota hybrid carrageenans with different iota/kappa ratios, and kappa–beta, xi–theta, and xi–lambda hybrid carrageenans.     

A simulation model for an Iridaea splendens (Gigartinales,Rhodophyta) population in Vancouver,Canada

Populations of Iridaea splendens at Brockton Point, Stanley Park, Vancouver, Canada were observed to alternate in dominance between the gametophytic phase in summer and tetrasporophytic phase in winter. The mechanism regulating this alternation is not clear. Using a matrix projection model to simulate population growth, we show that this alternation is possible if there are differential survival and recruitment rates of the two phases in summer and winter. Sensitivity and elasticity analyses indicate the relative importance of perennation vs. recruitment. Recruitment from tetrasporophytes and from gametophytes both contribute about 25% to the population growth. Perennation among gametophytes is more important than among tetrasporophytes. The implication of this is that if this population is to be harvested, more tetrasporophytes can be harvested than gametophytes without resulting in the depletion of the resource. This is simulated in the matrix model by comparing the relative effects on population growth of increasing the mortality rate of the perennation phase of tetrasporophyte and gametophyte by 50 to 75%, and increasing recruitment rate in either phase, from summer to winter or from winter to summer.Dedicated to Dr T. Bisalputra of the University of British Columbia on the occasion of his retirement.     

Differential ecological responses to environmental stress in the life history phases of the isomorphic red alga Gracilaria chilensis (Rhodophyta)

In order to better understand the alternation of generations that characterizes haploid–diploid life cycles, we assessed the existence of ecological differences between the two phases (haploid gametophyte and diploid tetrasporophyte) in Gracilaria chilensis, a rhodophyte with a typical Polysiphonia-type life cycle. We investigated the effect of light intensity and salinity on viability and growth of both phases at different ontogenetic stages: juveniles and adults. In our study, the survival of juvenile gametophytes (n) was higher than the survival of juvenile tetrasporophytes (2n) despite culture conditions; however, low salinity had greater effect on carpospores (2n) than on tetraspores (n). On the other hand, a complex interaction between salinity and light intensity within each life history phase generated observed differences between juvenile growth rates. Low light was shown to trigger early onset of alteration of the holdfast growing pattern. In addition, adult tetrasporophytes showed, despite the conditions, a faster vegetative growth than female and male gametophytes. These differences between phases could have led to the complete dominance of tetrasporophyte fragments of fronds observed in G. chilensis farms. We hypothesize that Chilean fishers could have unknowingly selected for tetrasporophyte thalli during domestication of the species, thus enhancing the natural trend of tetrasporophytes dominance already present in estuarine natural populations of free-floating plants.     

Cloning, expression and characterization of a new ι-carrageenase from marine bacterium, Cellulophaga sp.

Purpose of work

The purpose of this study is to report a ι-carrageenase which degrades ι-carrageenan yielding neo-ι-carratetraose as the main product in the absence of NaCl. The gene for a new ι-carrageenase, CgiB_Ce, from Cellulophaga sp. QY3 was cloned and sequenced. It comprised an ORF of 1,386 bp encoding for a protein of 461 amino acid residues. From its sequence analysis, CgiB_Ce is a new member of GH family 82 and shared the highest identity of 32 % in amino acids with ι-carrageenase CgiA2 from Zobellia galactanovorans indicating that it is a hitherto uncharacterized protein. The recombinant CgiB_Ce had maximum specific activity (1,870 U/mg) at 45 °C and pH 6.5. It was stable between pH 6.0–9.6 and below 40 °C. Although its activity was enhanced by NaCl, the enzyme was active in the absence of NaCl. CgiB_Ce is an endo-type ι-carrageenase that hydrolyzes β-1,4-linkages of ι-carrageenan, yielding neo-ι-carratetraose as the main product (more than 80 % of the total product).     

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.