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Ciencias marinas

Print version ISSN 0185-3880

Cienc. mar vol.46 n.3 Ensenada Sep. 2020  Epub Apr 16, 2021 


Chemical characterization of soluble polysaccharides in the red alga Acanthophora spicifera from La Paz Bay, Baja California Sur, Mexico

Caracterización química de los polisacáridos solubles del alga roja Acanthophora spicifera de la bahía de La Paz, Baja California Sur, México

Valérie Chantal Gabrielle Schnoller1 

Gustavo Hernández-Carmona1 

Enrique Hernández-Garibay2  * 

Juan Manuel López-Vivas3 

Mauricio Muñoz-Ochoa1 

1 Laboratorio de Química de Algas Marinas, Centro Interdisciplinario de Ciencias Marinas (CICIMAR), Instituto Politécnico Nacional (IPN), Av. Instituto Politécnico Nacional, s/n, Playa Palo de Santa Rita, CP 23096, La Paz, Baja California Sur, Mexico.

2 Centro Regional de Investigaciones Acuícolas y Pesqueras de Ensenada (CRIAP), Instituto Nacional de Pesca y Acuacultura, Carretera Tijuana-Ensenada, km 97.5, CP 22760, Ensenada, Baja California, Mexico.

3 Departamento Académico de Ciencias Marinas y Costera, Universidad Autónoma de Baja California Sur, Carretera al Sur, km 5.5, Col. Mezquitito, CP 23080, La Paz, Baja California Sur, Mexico.


Acanthophora spicifera is an invasive red alga that was recently detected in La Paz Bay, Baja California Sur, Mexico, where it has developed into a large biomass. Because it is a new species in the region, the characteristics and properties of the soluble polysaccharides (SPs) that it contains are unknown. To determine the content, chemical composition, and properties of SPs in A. spicifera, monthly samplings were carried out in 2013 at Point Roca Caimancito in La Paz Bay, and native and alkali-treated polysaccharides were extracted and characterized. The alkaline treatment produced lower yields and modified the composition of A. spicifera SPs. The polysaccharides obtained before or after the alkaline treatment had low viscosity and did not have gelling properties. In line with the obtained results, the molar ratio of components (galactose, 3,6-anhydrogalactose, sulfates [Gal:3,6-AG:sulfates]) for native (1.00:0.30:0.23) and alkali-treated (1.00:0.30:0.17) polysaccharides showed that A. spicifera SPs have a lower proportion of sulfates than that in polysaccharides belonging to carrageenans but greater than that in polysaccharides belonging to true agar. The Fourier-transform infrared spectra of SPs showed characteristic signals for sulfated galactans, with the presence of pyruvic acid; after the alkaline treatment, characteristic signals for agar-type polysaccharides (agaroid) were observed. Although A. spicifera polysaccharides have no gelling properties, additional studies are needed to clarify the structure of the SPs it contains to find appropriate uses for this resource.

Key words: Acanthophora spicifera; agar; alkaline treatment; seasonal variation; soluble polysaccharides


Acanthophora spicifera es un alga roja invasiva que recientemente fue detectada en la bahía de La Paz, Baja California Sur, México, donde se ha desarrollado en grandes biomasas. Debido a que es una nueva especie en la región, se desconocen las características y las propiedades de los polisacáridos solubles (PS) que contiene. Con el propósito de determinar el contenido, la composición química y las propiedades de los PS de A. spicifera, durante el año 2013, se realizaron muestreos mensuales en punta Roca Caimancito en la bahía de La Paz, y se extrajeron y caracterizaron los PS nativos y los PS con tratamiento alcalino. El tratamiento alcalino redujo el rendimiento y modificó la composición de los PS de A. spicifera. Los polisacáridos obtenidos antes o después del tratamiento alcalino fueron de baja viscosidad y no gelificaron. En concordancia con los resultados obtenidos, la razón molar de los componentes (galactosa, 3,6-anhidrogalactosa, sulfatos [Gal:3,6-AG: sulfatos]) para los polisacáridos nativos (1.00:0.30:0.23) y los polisacáridos con tratamiento alcalino (1.00:0.30:0.17) mostró que los PS de A. spicifera poseen una menor proporción de sulfatos que la que corresponde a los polisacáridos que pertenecen a los carragenanos, pero mayor que la que corresponde a los polisacáridos pertenecientes al agar verdadero. Los espectros obtenidos de la espectrometría infrarroja con transformada de Fourier de los PS mostraron señales características para galactanos sulfatados, con la presencia de ácido pirúvico; después del tratamiento alcalino, se observaron señales características para polisacáridos del tipo agar (agaroideo). Aunque los polisacáridos de A. spicifera no tienen propiedades gelificantes, es necesario realizar estudios adicionales para determinar la estructura de los PS que contiene esta especie con el propósito de encontrar usos apropiados para este recurso.

Palabras clave: Acanthophora spicifera; agar; tratamiento alcalino; variación estacional; polisacáridos solubles


The wide diversity of marine environments on the costas of Mexico, from temperate oceans to tropical waters, favors the development of a wide diversity of marine algal species. Nearly 1,000 species of macroalgae can be found on these coasts (Robledo 1998). It is common for the floristic pattern of a locality to be affected by the accidental or deliberate introduction of new macroalgae species (invasive algae) (Pedroche and Sentíes 2003, Williams and Smith 2007). That is the case of the red alga Acanthophora spicifera (Børgesen 1910), a native species from the Caribbean and Florida coasts. It was recorded as an invasive species on the Central Pacific Islands (Russell 1992, Tsuda et al. 2008). Recently, A. spicifera, was found on the southwestern costas of the Gulf of California, where it has formed extensive beds with considerable biomass (Fig. 1) (Riosmena-Rodríguez et al. 2009). On the other hand, marine algae are an important source of valuable soluble polysaccharides such as agar, carrageenan, and alginates (Painter 1983, Craigie 1990, Lobban and Harrison 1994). Generally, the composition and properties of the different polysaccharides in seaweeds are related to the physiological function in the corresponding taxonomic algal classification (Kloareg and Quatrano 1988, Miller 1997, Usov and Zelinsky 2013). Red algae, in particular, are known for producing polysaccharides belonging to the family of sulfated galactans. With a few possible exceptions, the structures of all the galactans in this group have a common, simplifying feature: they use as basis linear chains of β-D-galactopyranose residues glycosidically linked through positions 1 and 3 (A units) and α-galactopyranose residues glycosidically linked through positions l and 4 (B units) in either the agar (levogyre [L] B unit) or the carrageenan (dextrogyre [D] B unit) (Painter 1983, Craigie 1990).

Figure 1 Acanthophora spicifera overgrowing on Pocillopora and Porites coral colonies off Point Roca el Caimancito, La Paz, Baja California Sur, Mexico. 

In the case of A. spicifera seaweed, which belongs to the order Ceramiales, previous studies have reported that it contains highly sulfated galactans, and studies using mainly Fourier-transform infrared (FTIR) spectroscopy suggest that it produces polysaccharides belonging to the carrageenan family, particularly the lambda type (λ) (Table 1a) (Parekh et al. 1989, Cajipe 1990, Gomaa and Elshoubaky 2016). However, this is controversial as other more complete studies have shown that A. spicifera polysaccharides belong to the agaran family (Gonҫalvez et al. 2002, Duarte et al. 2004, Seixas et al. 2007). Since A. spicifera is a new species in the La Paz Bay area, Baja California Sur, Mexico, no studies have been conducted on the characteristics of the polysaccharides it contains. The purpose of this research was to assess the seasonal content and chemical characteristics of the soluble polysaccharides in A. spicifera, native and alkali-treated, to determine the potential use for this species.

Table 1 Disaccharide repeating units of lambda (λ)-carrageenan (a), native agaran (b), and ideal agarose (c). 

Name Repetitive idealized disaccharide
Full IUPAC* name
3-linked β-D-galactopyranose 2-sulfate
(A) and 4-linked α-D-galactopyranose
2,6-disulfate (B)
Repetitive dimer in the agaran family
3-linked β-D-galactopyranose (A) and
4-linked α-L-galactopyranose 6-sulfate (B)
Repetitive dimer of agarose in the
agaran family (alkali-treated)
3 linked β-D-galactopyranose (A) and
4-linked α-L-3,6-anhydro-galactopyra
nose (B)

* International Union of Pure and Applied Chemistry.


Collection of biological material

Samples of A. spicifera were collected monthly from February to December 2013 off Point Roca El Caimancito (24º12´12.91 N, 110º18´3.43 W), in La Paz Bay, Baja California Sur, Mexico. The samples were first sun-dried and milled to Ф ≤ 2 mm, then stored in plastic bags in a cool, dry place until analysis.

Extraction of soluble polysaccharides

Native soluble polysaccharides

Done in triplicate, 3 g of dried and milled seaweed simples were extracted by stirring at boiling point for 2 h in 90 mL of 0.1 M phosphate buffer at 6.5 pH. Diatomaceuous earth was added as a filter aid and was vacuum filtered. To increase the ionic strength in the solution, sodium chloride (NaCl) crystals were added up to 0.1 M, and soluble polysaccharides were precipitated by changing the solvent with 2 volumes of 96% ethanol. To remove the salts and water, the coagulum was first rinsed twice with 70% ethanol, then once with 96% ethanol, and then it was dried for 24 h at 60 ºC and weighed (Craigie and Leigh 1978).

Alkali-treated soluble polysaccharides

Done in triplicate, 3 g of dried and milled seaweed were treated with a solution of 6% potassium hydroxide (KOH) and then heated at 90 ºC for 60 min. The liquid was drained after cooling. To remove excess alkali, seaweed samples were first rinsed with running tap water, then with 0.05 N H2SO4 for 3 min, and finally with distilled water. The extraction was carried out following the procedure described for the native extraction (Craigie and Leigh 1978).

Physicochemical characterization

Total carbohydrates were quantified by the phenolsulfuric acid colorimetric method (DuBois et al. 1956) using galactose as standard. 3,6-anhydrogalactose content (3,6- AG) was determined by the resorcinol-acetal method of Yaphe and Arsenault (1965) modified by Craigie and Leigh (1978) using fructose as standard. To determine sulfate content, about 20 mg of dry samples of native and alkali-treated polysaccharides were hydrolyzed with 2N HCl at 100 ºC for 2 h. Sulfate content was then determined by the turbidimetric method of Tabatabai modified by Craigie and Leigh (1978) using potassium sulfate (K2SO4) as standard. The molar ratio of components, galactose, 3,6-anhydrogalactose, sulfates (Gal:3,6-AG: sulfates), was calculated from the native and alkali-treated polysaccharides specially extracted for this determination. The values of the molecular weight of each component were used in the calculation, where sulfates were calculated as sodium sulfate (SO3Na). The absolute viscosity of native and alkali-treated polysaccharides was measured in millipascals per second (mPa·s) in 3% (w/v) solutions at 20 ºC with a Brookfield Mod LVT viscometer with spindle No. 1 at 30 rpm.

Infrared spectroscopy

Fourier-transform infrared spectroscopy (FTIR) was applied. About 1 mg of dry polysaccharide samples was mixed and pressed with potassium bromide (KBr) crystals to form a translucid pellet. The spectra were then acquired in a Perkin-Elmer Spectrum 100 spectrophotometer after 10 scans at a resolution of 4 cm-1 scannings, between 400 and 4,000 cm-1 wavelengths, in transmittance mode.

Statistical analysis

All analyses were performed in triplicate. A priori tests were performed to define normality and homoscedasticity. Since the data were not normal, an analysis of variance was applied to detect significant differences among treatments (P < 0.05). A rank analysis (Kruskal-Wallis) and a posteriori Tukey tests were used with the non-parametric data (Zar 1999).


The applied alkaline treatment affected the yield and composition of the soluble polysaccharides from A. spicifera. Overall, the yields of native extractions were higher (24.7% to 39.3%) than those of extractions with the alkaline treatment (9.1% to 14.9%). In the case of native extractions, the highest yield (39.4%) was obtained in July and the lowest (26.2%) in September (Fig. 2). When using the alkaline treatment, the highest yield (14.9%) was obtained in April and the lowest (9.1%) in August. Only the yield of native polysaccharides showed significant differences throughout the year (P < 0.05) (Fig. 2).

Figure 2 Monthly variation of native and alkali-treated polysaccharide yields from Acanthophora spicifera. Vertical bars are standard deviations (n = 3). Different letters correspond to significant differences between months. 

In addition to the changes in soluble polysaccharide yields, there was an increase in the 3,6-AG content and a decrease in the sulfate contents when applying the alkaline treatment before extraction. The 3,6-AG yields for native polysaccharides ranged from 16.48% to 20.26%, and the yields for alkali-treated polysaccharides ranged from 13.80% to 25.70%. Significant differences occurred throughout the year (P < 0.05) (Table 2). Moreover, sulfate content was higher for native polysaccharides (8.80%-13.80%) than for alkali-treated polysaccharides (4.50%-8.90%). Similarly, significant differences were observed between sampling months (P < 0.05) (Table 2).

Table 2 Monthly variation of the 3,6-anhydrogalactose (3,6-AG) and sulfate contents in native and alkali-treated polysaccharides from Acanthophora spicifera. Data are presented as the mean of triplicate assays (n = 3). Different letters indicate significant differences between months. 

3,6 AG (%) Sulfate (%)
Month Native Alkali-treated Native Alkali-treated
March 19.70 ± 0.37e 19.40 ± 0.14b 13.80 ± 0.15e 7.00 ± 0.12d
April 19.40 ± 0.13e 19.50 ± 0.20b 11.50 ± 0.18d 6.50 ± 0.05c
May 19.60 ± 0.11e 13.80 ± 0.02a 10.50 ± 0.07c 4.50 ± 0.07a
June 16.48 ± 0.13 a 25.70 ± 0.92f 8.80 ± 0.05a 8.90 ± 0.10e
July 20.26 ± 0.01d 22.29 ± 0.10e 13.70 ± 0.07e 6.80 ± 0.14c
August 18.19 ± 0.09c 20.64 ± 0.07c 10.30 ± 0.03c 5.40 ± 0.07b
September 17.17 ± 0.32b 19.52 ± 0.40b 10.20 ± 0.28c 6.80 ± 0.01c
October 18.23 ± 0.13c 21.03 ± 0.09d 11.10 ± 0.13d 7.80 ± 0.12d
November 19.96 ± 0.12e 20.13 ± 0.53c 10.90 ± 0.17c 6.70 ± 0.05c
December 17.20 ± 0.09b 19.71 ± 0.47b 9.92 ± 0.06b 7.60 ± 0.18d

The FTIR spectra of the soluble polysaccharides obtained from A. spicifera revealed the characteristic signals for sulfated galactans from red algae. A strong band was detected between 1,250 and 1,230 cm−1, with additional bands at 930, 890, 836, and 771 cm−1. The spectrum for the alkali-treated polysaccharide showed more and better-defined signals than the corresponding spectrum for native polysaccharide (Fig. 3).

Figure 3 Fourier-transform infrared spectra of native (N-SP) and alkali-treated (AT-SP) soluble polysaccharides from Acanthophora spicifera

Physicochemical properties

The polysaccharide obtained from A. spicifera is nongelling and produced low-viscosity solutions, which were lower than 41 mPa·s in all cases. The absolute viscosity of the native polysaccharide was significantly higher than that of the alkali-treated polysaccharide. The highest viscosity was obtained with the native process in October (41 mPa·s), while the lowest (4 mPa·s) was obtained with the alkaline treatment in December. An increase in the viscosity of the native polysaccharide was observed from mid summer to the beginning of fall; significant differences were observed mainly for the native polysaccharide (Fig. 4).

Figure 4 Absolute viscosity of native and alkali-treated polysaccharides measured with a Brookfield viscometer with spindle 1 at 30 rpm. Vertical bars are standard deviations (n = 3). Different letters indicate significant differences between months. 

The molar ratio of components (Gal:3,6-AG:sulfates) showed that the A. spicifera polysaccharide is a sulfated galactan with low proportion of 3,6-AG (Table 3). The proportion of hemiester sulfate groups in the native polysaccharide was about 1 sulfate group for every 3 disaccharide units (A-B), while with the alkaline treatment, it decreased to 1 sulfate group for almost every 4 disaccharide units (Table 3).

Table 3 Composition, molar proportion (galactose [Gal], 3,6-anhydrogalactose [3,6-AG], sulfates), and molar ratio of native and alkali-treated polysaccharides from Acanthophora spicifera

Content (%) Molar proportion Molar ratio
Treatment Gal 3,6-AG Sulfate Gal 3,6-AG Sulfate Gal 3,6-AG Sulfate
Native 70.74 18.96 10.30 0.44 0.13 0.10 1.00 0.30 0.23
Alkali-treated 72.47 19.62 7.91 0.45 0.14 0.08 1.00 0.30 0.17


Seaweeds belonging to the Rhodophyceae family are important sources of sulfated polysaccharides, either the agar or the carrageenan types (Painter 1983, Craigie 1990, Lobban and Harrison 1994). To isolate and characterize soluble polysaccharides from red seaweeds, usually in addition to obtaining the native polysaccharide, it is also necessary to apply an alkaline treatment, as it produces more chemically homogeneous molecules and, in some cases, enhances gelforming properties (Hoffmann et al. 1995, Usov 2011).

In this study native and alkali-treated soluble polysaccharides from the red seaweed A. spicifera from La Paz Bay, Baja California Sur, Mexico, were extracted and characterized. As expected, when using the alkaline treatment on the raw material, given the high temperature and high alkali concentration, the 4-linked α-galactopyranose 6-sulfate (B unit) (Table 1b) transformed into 3,6-anhydro α-galactopyranose (Table 1c). The alkaline treatment modifies the composition and properties of polysaccharides in red seaweeds (Kloareg and Quatrano 1988, Craigie 1990, Usov 2011); however, it also entails some degradation and loss of the polysaccharide (Stanley 1987, Myslabodski 1990). Therefore, in this study, the native extraction yields (24.7% to 39.3%) were higher than the extractions with the alkaline treatment (9.1% to 14.9%) (Fig. 2); by contrast, there was an increase in the 3,6-AG content and a concurrent decrease in sulfate content (Table 2). The same trend, lower yields and increased 3,6-AG content, was observed for some alkali-treated agar-producing species (Freile-Pelegrín and Robledo 1997, Arvizu-Higuera et al. 2007, Orduña-Rojas et al. 2008) and for carrageenanproducing seaweeds (Craigie 1990, Freile-Pelegrín et al. 2006).

Using FTIR spectra, some authors have classified the polysaccharides from A. spicifera as carrageenans, particularly of the lambda (λ) type (Parekh et al. 1989, Cajipe 1990, Pickering and Mario 1999, Gomaa and Elshoubaky 2016). λ-Carrageenan is a highly sulfated galactan that yields high viscous solutions but has no gelling properties; it has been chemically characterized (data not shown) and its composition has high contents of sulfate groups (35% to 40%) and only traces of 3,6-AG. In this study, we found that A. spicifera polysaccharides also have no gelling properties, and unlike carrageenans they produce solutions with very low viscosity, even after the alkaline treatment (Fig. 4). Compared to λ-carrageenan, the chemical composition of the A. spicifera polysaccharide (native) has lower sulfate content (8.8% to 13.8%) and higher 3,6-AG content (17.0% to 20.0% for native polysaccharides) (Table 2). Futhermore, the molar ratio (G:3,6-AG:sulfates) for λ-carrageenan is 1.00:0.13:1.10 (experimental data, not shown), whereas for the A. spicifera polysaccharides it is 1.00:0.30:0.23 for the native polysaccharide and 1.00:0.30:0.17 for the alkali-treated polysaccharide (Table 3); however, the A. spicifera polysaccharide composition also differs from true agar, which has higher 3,6-AG content and lower sulfate content, with an ideal molar ratio of 1.00:1.00:0.00 (Armisen and Galatas 1987, Craigie 1990).

On the other hand, studies on A. spicifera polysaccharides have shown that the soluble polysaccharides in this species belong to the agaran family (Gonҫalvez et al. 2002, Duarte et al. 2004). They are known to be highly sulfated on C-2, with pyruvic acid linked as a cyclic ketal bridge at 4,6-O-(1′-carboxyethylidene groups) in the 1-3linked A unit (Duarte et al. 2004). This is in agreement with the fact that the polysaccharides found in other algae belonging to the order Ceramiales biosynthesize sulfated agarans (Painter 1983, Craigie 1990, Usov 2011).

In this study, the FTIR spectra of the A. spicifera polysaccharides showed that these are highly sulfated polysaccharides with pyruvic acid, as shown by the presence of a strong signal between 1,240 and 1,260 cm-1 (asymmetric stretching of O=S=O) (Stancioff and Stanley 1969) and a double signal at 1,637 and 1,417 cm-1 for the pyruvic acid carboxylic group (Prado-Fernández et al. 2003). The spectra of the alkali-treated polysaccharides showed better-defined signals between 1,000 and 700 cm-1 because of the loss of the labile sulfate in C-6 of the B unit and the concurrent 3,6-AG formation. This revealed the signal at 930 cm-1 because of 3,6-AG (Rochas et al. 1986) and a signal at 890 cm-1 for desulfated galactose (Greer and Yaphe 1984). Both signals are absent in λ-carrageenan but are characteristic of agars (Armisen and Galatas 1987, Barros et al. 2013, Gómez-Ordóñez and Rupérez 2011). There was also a strong signal at 830 cm-1, which corresponded to the equatorial sulfate in galactose 2-sulfate (Rochas et al. 1986), and signals at 790 and 717 cm-1, which are characteristic of agar (Matsuhiro and Miller 2002) (Fig. 3). Our results suggest that the soluble polysaccharides in A. spicifera belong to the agaran family; however, given their composition and properties, they must be considered deviants of the true agar (agaroids) (Craigie 1990). On the other hand, despite the deficient viscosity and gelling properties of some polyssacharides, it should be noted that sulfated galactans in red seaweeds usually show potent antiviral activity (Carlucci et al. 1997, Damonte et al. 2004), where the inhibitory effect is attributed to their negatively charged sulfated polysaccharides because of the possible conformational similarity between disaccharide repeating units of sulfated galactans and heparan sulfate, which serves as a negatively charged cell-surface target for most viruses (Chen et al. 1997, Duarte et al. 2004, Usov 2011). Considering mainly the composition and properties of the polysaccharides found in this study, A. spicifera cannot be considered a good source of agar; however, it could alternatively be used for its possible antiviral activity (Duarte et al. 2004, Bouhlal et al. 2011, Gomaa and Elshoubaky 2016) or its possible anti-inflammatory properties, as described for Acanthophora muscoides (Quinderé et al. 2013).


We thank the institutions that provided support for this research (CICIMAR-IPN, CRIAP/Ensenada-INAPESCA). Thanks are also due to the National Council for Science and Technology (Mexico), IPN Comisión de Operación y Fomento de Actividades Académicas (COFAA), and Beca De Estímulo Institucional de Formación de Investigadores for granting VCGS a scholarship. GHC thanks IPN for the productivity grants provided by COFAA and Estímulo al Desempeño de los Investigadores and the economic stimulus granted by the Mexican National System of Researchers (Sistema Nacional de Investigadores).The Faculty of Marine Sciences at UABC facilitated use of the FTIR spectrometer. We thank Blanca Esther García Espinosa for her assistance and the anonymous reviewers for their valuable suggestions.


Armisen R., Galatas F. 1987. Production, properties, and uses of Agar. In: McHugh DJ. (ed.), Production and utilization of products from commercial seaweeds. FAO Fisheries Technical Paper. No. 288: Food and Agriculture Organization of the United Nations. p. 1-57. [ Links ]

Arvizu-Higuera DL., Rodríguez-Montesinos YE., Murillo-Álvarez JI., Muñoz-Ochoa M., Hernández-Carmona G. 2007. Effect of alkali treatment time and extraction time on agar from Gracilaria vermiculophylla. In: Borowitzka MA., Critchley AT., Kraan S., Peters A., Sjøtun K., Notoya M., (eds.), Nineteenth International Seaweed Symposium. Vol. 2, Developments in Applied Phycology. Dordrecht (Netherlands): Springer. p. 65-69. [ Links ]

Barros FCN., da Silva DC., Sombra VG., Maciel JS., Feitosa JPA., Freitas ALP., de Paula RCM. 2013. Structural characterization of polysaccharide obtained from red seaweed Gracilaria caudata (J Agardh). Carbohydrate Polymers. 92(1):598-603. [ Links ]

Børgesen F. 1910. Some new or little known West Indian Florideae. II. Botanisk Tidsskrift. 30:177-207, 20 figs. [ Links ]

Bouhlal R., Haslin C., Chermann J-C., Colliec-Jouault S., Sinquin C., Simon G., Cerantola S., Riadi H., Bourgougnon N. 2011. Antiviral activities of sulfated polysaccharides isolated from Sphaerococcus coronopifolius (Rhodophyta, Gigartinales) and Boergeseniella thuyoides (Rhodophyta, Ceramiales). Mar Drugs. 9(7):1187-1209. [ Links ]

Cajipe GJB. 1990. Utilization of seaweed resources. In: Dogma IJ. Jr. , Trono GC. Jr. , Tabbada RA., (eds.), Culture and use of algae in Southeast Asia: Proceedings of the Symposium on Culture and Utilization of Algae in Southeast Asia; 8-11 December 1981; Tigbauan, Iloilo, Philippines. Philippines: Aquaculture Department, Southeast Asian Fisheries Development Center. p. 77-79. [ Links ]

Carlucci MJ., Scolaro LA., Errea MI., Matulewicz MC., Damonte EB. 1997. Antiviral activity of natural sulphated galactans on herpes virus multiplication in cell culture. Planta Med. 63(5):429-432. [ Links ]

Chen Y., Maguire T., Hileman RE., Fromm JR., Esko JD., Linhardt RJ., Marks RM. 1997. Dengue virus infectivity depends on envelope protein binding to target cell heparan sulfate. Nat Med. 3(8):866-871. [ Links ]

Craigie JS. 1990. Cell walls. In: Cole KM., Sheath RG., (eds.), Biology of the Red Algae. Cambridge (UK): Cambridge University Press. p. 221-257. [ Links ]

Craigie JS., Leigh C. 1978. Carrageenans and agars. In: Hellebust JA., Craigie JS., (eds.), Handbook of phycological methods: physiological and biochemical methods. New York: Cambridge University Press. p. 110-131. [ Links ]

Damonte EB., Matulewicz MC., Cerezo AS. 2004. Sulfated seaweed polysaccharides as antiviral agents. Curr Med Chem. 11(18):2399-2419. [ Links ]

Duarte MER., Cauduro JP., Noseda DG., Noseda MD., Gonçalves AG., Pujol CA., Damonte EB., Cerezo AS. 2004. The structure of the agaran sulfate from Acanthophora spicifera (Rhodomelaceae, Ceramiales) and its antiviral activity. Relation between structure and antiviral activity in agarans. Carbohydr Res. 339(2):335-347. [ Links ]

DuBois M., Gilles KA., Hamilton JK., Rebers PA., Smith F. 1956. Colorimetric method for determination of sugars and related substances. Anal Chem. 28(3):350-356. [ Links ]

Freile-Pelegrín Y., Robledo D. 1997. Influence of alkali treatment on agar from Gracilaria cornea from Yucatán, México. J App Phycol. 9(6):533-539. [ Links ]

Freile-Pelegrín Y., Robledo D., Azamar JA. 2006. Carrageenan of Eucheuma isiforme (Solieriaceae, Rhodophyta) from Yucatán, Mexico. I. Effect of extraction conditions. Bot Mar. 49: 65-71. [ Links ]

Gomaa HHA., Elshoubaky GA. 2016. Antiviral activity of sulfated polysaccharides carrageenan from some marine seaweeds. International Journal of Current Pharmaceutical Review and Research. 7(1):34-42. [ Links ]

Gómez-Ordóñez E., Rupérez P. 2011. FTIR-ATR spectroscopy as a tool for polysaccharide identification in edible brown and red seaweed. Food Hydro. 25(6):1514-1520. [ Links ]

Gonҫalvez AG., Ducatti DRB., Duarte MER., Noseda MD. 2002. Sulfated and pyruvylated disaccharide alditols obtained from a red seaweed galactan: ESIMS and NMR approaches. Carbohydr Res. 337(24):2443-2453. [ Links ]

Greer CW., Yaphe W. 1984. Characterization of hybrid (Betha- Kappa-Gamma) carrageenan from Eucheuma gelatinae J. Agardh (Rhodophyta, Solieriaceae) using carrageenases, infrared and 13C-nuclear magnetic resonance spectroscopy. Bot Mar. 27(10):473-478. [ Links ]

Hoffmann RA., Gidley MJ., Cooke D., Frith WJ. 1995. Effect of isolation procedures on the molecular composition and physical properties of Eucheuma cottonii carrageenan. Food Hydrocolloids. 9(4):281-289. [ Links ]

Kloareg B., Quatrano RS. 1988. Structure of the cell walls of marine algae and ecophysiological functions of the matrix polysaccharides. Oceanogr Mar Biol Annu Rev. 26:259-315. [ Links ]

Lobban CS., Harrison PJ. 1994. Seaweed ecology and physiology. Cambridge: Cambridge University Press. 66 p. [ Links ]

Matsuhiro B., Miller LG. 2002. Soluble polysaccharides from Rhodymenia: Characterization by FT-IR spectroscopy. Bol Soc Chil Quím. 47(3):265-271. [ Links ]

Miller IJ. 1997. The chemotaxonomic significance of the water- soluble red algal polysaccharides. Recent Res Dev Phytochem. 1:531-565. [ Links ]

Myslabodski DE. 1990. Red-algae galactans: Isolation and recovery procedures-Effects on the structure and rheology [dissertation]. [Trondheim (Norway)]: Norwegian Institute of Technology. [ Links ]

Orduña-Rojas J., Suárez-Castro R., López-Álvarez ES., Ríosmena-Rodríguez R., Pacheco-Ruiz I., Zertuche-González JA., Meling-López AE., 2008. Influence of alkali treatment on agar from Gracilariopsis longissima and Gracilaria vermiculophylla from the Gulf of California, Mexico = Influencia del tratamiento alcalino en el agar de Gracilariopsis longissima y Gracilaria vermiculophylla del Golfo de California México. Cienc Mar. 34(4):503-511. [ Links ]

Painter TJ. 1983. Algal polysaccharides. In: Aspinall GO., (ed.), The polysaccharides. New York: Academic Press. p. 195-285. [ Links ]

Parekh RG., Doshi YA., Chauhan VD. 1989. Polysaccharides from marine red algae Acanthophora spicifera, Grateloupia indica and Halymenia porphyroides. Indian J Mar Sci. 18(2):139-140. [ Links ]

Pedroche FF., Sentíes AG. 2003. Ficología marina mexicana. Diversidad y Problemática actual. Hidrobiológica. 13(1):23-32. [ Links ]

Pickering T., Mario S. 1999. Survey of commercial seaweeds in South-east Viti Levu (Fiji Islands): A preliminary study on the farming potential of seaweed species present in Fiji. Suva (Fiji): Food and Agriculture Organisation of the United Nations, Fisheries and Aquaculture Department; [accessed 2000 September 1]. . [ Links ]

Prado-Fernández J., Rodríguez-Vázquez JA., Tojo E., Andrade JM. 2003. Quantitation of κ-, ι- and λ-carrageenans by mid- infrared spectroscopy and PLS regression. Anal Chim Acta. 480(1):23-37. [ Links ]

Quinderé ALG., Fontes BR., Vanderlei E. de SO., de Queiroz INL., Rodrigues JAG., de Araújo IWF., Jorge RJB., de Menezes DB., e Silva AAR., Chaves HV., et al. 2013. Peripheral antinociception and anti-edematogenic effect of a sulfated polysaccharide from Acanthophora muscoides. Pharmacol Rep. 65(3):600-613. [ Links ]

Riosmena-Rodríguez R., Ruiz G., Hernández-Kantún J. 2009. Invasión de algas exóticas en el Golfo de California: amenaza para el ambiente y la economía regionales. La Paz (Mexico): Análisis Periodísticos BCS. p. 25-26. [ Links ]

Robledo D. 1998. Seaweed resources of Mexico. In: Critchley AT., Ohno M., (eds.), Seaweed resources of the world. Tokyo: Japanese International Cooperation Agency. p. 331-342. [ Links ]

Rochas C., Lahaye M., Yaphe W. 1986. Sulfate content of carrageenan and agar determined by infrared spectroscopy. Bot Mar. 29(4):335-340. [ Links ]

Russell DJ. 1992. The ecological invasion of Hawaiian reefs by two marine red algae, Acanthophora spicifera (Vahl) Boerg. and Hypnea musciformis (Wulfen) J. Ag., and their association with two native species, Laurencia nidifica and Hypnea cervicornis J.Ag. ICES Mar Sci Symp. 194:110-125. [ Links ]

Seixas C., Barragán V., Escobar C., Fuentes J. 2007. Contenido y calidad del agar extraído de muestras de Acanthophora spicifera (Vahl) Borgesen (Rhodophyta) provenientes del caribe de Panamá. Tecnociencia. 9(1):7-13. [ Links ]

Stancioff DJ., Stanley NF. 1969. Infrared and chemical studies on algal polysaccharides. Proc Int Seaweed Symp. 6:595-609. [ Links ]

Stanley N. 1987. Production, properties and uses of carrageenan. In: McHugh DJ. (ed.), Production and Utilization of Products from Commercial Seaweeds. FAO Fisheries Technical Paper. Rome (Italy): Food and Agriculture organization of the United Nations. p. 116-146. Paper No. 288. [ Links ]

Tsuda RT., Coles SL., Guinther EB., Finlay O., Andrew R., Harris FL. 2008. Acanthophora spicifera (Rhodophyta: Rhodomelaceae) in the Marshall Islands. Micronesica. 40(1/2):245-252. [ Links ]

Usov AI. 2011. Polysaccharides of the red algae. Adv Carbohydr Chem Biochem. 65:115-217. [ Links ]

Usov AI., Zelinsky ND. 2013. Chemical structures of algal polysaccharides. In: Domínguez H., (ed.), Functional Ingredients from Algae for Foods and Nutraceuticals. Cambridge (UK): Elsevier Science. p. 45-49. [ Links ]

Williams SL., Smith JE. 2007. A global review of the distribution, taxonomy, and impacts of introduced seaweeds. Annu Rev Ecol Evol Syst. 38(1):327-359. [ Links ]

Yaphe W., Arsenault GP. 1965. Improved resorcinol reagent for the determination of fructose and 3,6-anhydrogalactose in polysaccharides. Analyt Biochem. 13(1):143-148. [ Links ]

Zar JH. 1999. Biostatistical analysis. New Jersey (USA): Prentice- Hall. 663 p. [ Links ]

Received: January 01, 2020; Accepted: May 01, 2020

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