Catenella nipae (PROSEA)

Plant Resources of South-East Asia Introduction

List of species

1, habit; 2,3, parts of vegetative thalli with haptera; 4, detail of a section with tetrasporangia in a tetrasporangial sorus; 5, longitudinal section of an apical segment showing carpogonial branches and a fusion cell producing a gonimoblast branch on one side.

Catenella nipae Zanardini

Protologue: Mem. Ist. venet. 17: 143-145, pl. 6, A (1872).

Family: Caulacanthaceae

Chromosome number: 2n= unknown

Synonyms

• Catenella opuntia (Gooden. & Woodw.) Grev. var. major J. Agardh (1876).

Origin and geographic distribution

C. nipae is restricted to the Indo-Pacific region and widely distributed in South-East Asia, Australia and New Zealand. It occurs in Burma (Myanmar), Thailand, Vietnam, Peninsular Malaysia, Singapore, Indonesia, northern Borneo, Brunei, the Philippines and Papua New Guinea. It is also recorded from West Bengal (India), Bangladesh, southern China and the Ryukyu Islands in Japan.

Uses

C. nipae is used for food and as a basis for carrageenan production, especially in Burma (Myanmar). There it is used as a salad, either fresh or after boiling water has been poured on the fresh or dried alga. Usually sesame oil and spices are used with it. For several years the "Seaweed Technology" Company (Burma) has been producing edible seaweed sheets from a combination of C. nipae and Enteromorpha spp. (green algae). C. nipae, often mixed with the closely related but smaller C. impudica (Mont.) J.Agardh, is also commonly used as a salad in the Philippines. In Thailand it is listed as an economic alga and in Malaysia as a potential source of carrageenan. In Vietnam specimens are collected in the field, probably for food purposes.

Production and international trade

It is foreseen that 1000 ha of mudflats in the Burmese mangrove swamps will be used for Catenella cultivation farms.

Properties

The structure of cell wall galactan of C. nipae is under discussion. Analyses of material from samples collected in Burma (Myanmar) showed the polysaccharide to be a previously undescribed carrageenan of the β-family, designated as α carrageenan. A repeating unit of 3-0-D-galactose (= β(1→4)-linked D-galactose) and 4-0-3,6-anhydro-D-galactose-2-0-sulphate (= α(1→3)-linked 3,6-anhydro-galactose-2-sulphate) was found in these Burmese samples. However, in samples of C. nipae collected in Australia and the Philippines, only iota carrageenan was found, consisting of mainly galactose and 3,6-anhydrogalactose, together with minor amounts (up to 1.8 mole %) of xylose and a high sulphate ester content (about 30% of dry weight as NaSO₃). Only traces of glucosyl residues were detected and no mono-0-methylgalactosyl residues. The repeating units were identified as β(1→4)-linked D-galactose-4-sulphate and α(1→3)-linked 3,6-anhydro-D-galactose-2-sulphate. The carrageenan type remains the same throughout the different reproductive phases.

Description

• Thalli forming decumbent patches up to 3 cm tall, dark red to purple, often bleached yellow-brown; fronds creeping below, subflabellately caespitose to erect above, composed of elongate ovoid, oblong or squarish, terete to strongly compressed segments (internodes), up to 6 mm long and about 2 mm wide, with constricted nodes; branching regular, repeatedly dichotomous or trichotomous; holdfasts (haptera) at terminal end of segments, bending down first ventrally to attach to the substrate, later becoming subterminal by subterminal growth of segments; new segments originating closely behind the haptera of the attached segments; segments internally composed of lacunose medulla of loosely interlacing and anastomosing longitudinal filaments from central axis soon becoming indistinct; filaments towards the periphery dichotomously branched, moniliform, forming a compact cortex of larger inner cells and smaller outer cells.
• Life cycle triphasic, diplo-haplontic and isomorphic.
• Tetrasporangia oblong, transversely zonately divided, scattered or aggregated in encircling sori below the surface of the cortex of terminal segments, 60-90(-110) μm × 40-65(-75) μm.
• Gametophytes monoecious in laboratory culture, with up to 8 mm long, narrow male segments and up to 1 mm long, round female segments; cystocarps ostiolate, solitary, sessile on shortened, obpyriform, subterminal segments with swollen cortex; carpospores ovoid, 45 μm × 25 μm, in chains; spermatangia small cell groups in cortical tissue, in ring-like spermatangial sori on male segments.

Growth and development

Apical cells of C. nipae give rise to laterals on two sides, forming two periaxial cells each. Axial cells soon transform into a network of slender filaments without a distinct central axis in the segments. In laboratory culture, tetrasporelings of 3-5 mm long developed carposporophytes which became reproductive in less than 8 months. After 5 months of growth, carpospore germlings were 3-5 mm long with 2-3 segments. At that stage most had formed viable tetraporangia. The relative abundance of vegetative, tetrasporic and carposporic phases of C. nipae in mangrove swamps has been studied in Burma (Myanmar). The reproductive phases occur during the pre-monsoon season, after which rigorous vegetative growth of the germlings contributes to the sudden occurrence of the alga in the monsoon season. Mixed phase gametophytes have been observed in C. nipae, in which also zonately-divided tetrasporangia developed on the female segments of the monoecious gametophytes.

Other botanical information

Although the characteristics that separate C. nipae from C. impudica are quite clear (in C. impudica haptera are formed from special small and narrow segments and in C. nipae from normal segments), much confusion occurs and often both species are mixed up. This may also be complicated by the fact that these species can occur in mixed stands. Usually, however, specimens and segments of C. nipae are much larger than those of C. impudica. For that reason C. nipae is probably the only species that is used for food and as a basis for carrageenan production.

Ecology

C. nipae belongs to the so-called Bostrychia-Caloglossa association, characteristic of mangrove habitats, and grows on the stems and modified roots (stilt roots, pneumatophores) of mangrove plants. Records of other edible algae from this community indicate that C. nipae is the major species collected, the others being inadvertently included. The indirect economic importance of these algae is that they form a component of the primary production system of mangrove swamps. While nothing is yet known quantitatively, this system is nonetheless thought to be critical in maintaining inshore fish and crustacean populations.

Propagation and planting

C. nipae is propagated by spores. Experimental cultivation of C. nipae has only been undertaken in Burma (Myanmar).

Phycoculture

In Burma (Myanmar) bamboo sticks of 5 cm × 45 cm are used as spore collectors for C. nipae cultivation. The bamboo sticks are inserted to a depth of 15 cm into the mudflats of the mangrove swamps to collect the spores. Luxuriant growth of C. nipae was observed on the sides of the bamboo sticks after the rainy season.

Diseases and pests

A gall-like outgrowth on C. nipae (supposed to be C. impudica) has been described as the parasitic red alga Catenellocolax leeuwenii Weber Bosse. The hemispheric and lobed parasitic plants were sterile and there are no confirmed data on their development or taxonomic status.

Harvesting

C. nipae is harvested by hand-collection. The specially placed bamboo sticks, used for experimental cultivation of C. nipae, are harvested using a spoon.

Yield

The yield of C. nipae on the planted bamboo sticks in Burma (Myanmar) was about 40-50 g (wet weight) per bamboo stake.

Handling after harvest

C. nipae is used fresh or sun-dried for food or for carrageenan production. In Burma (Myanmar), however, carrageenan is usually mainly produced from Hypnea species, although C. nipae can also be used.

Prospects

In Burma (Myanmar) about 25 small factories are now producing strips of carrageenan for the domestic market. The general Burmese public, however, is not aware whether it is carrageenan or agar and refer to both under the name "kyaukkyaw". Their preference is for jelly desserts that are white, hard and transparent. Due to the lack of technology for producing such quality in local strips of carrageenan, agar powder imported from neighbouring countries is much more popular and may outstrip the already declining indigenous carrageenan production in Burma (Myanmar). Production of C. nipae for fresh production, as well as for sun-dried directly marketed products, is still considerable in Burma (Myanmar) and is expected to continue in the future.

Literature

• Liao, M.-L., Munro, S.L.A., Craik, D.J., Kraft, G.T. & Bacic, A., 1993. The cell wall galactan of Catenella nipae Zanardini from Southern Australia. Botanica Marina 36: 189-193.
• Post, E., 1963. Zur Verbreitung und Ökologie der Bostrychia-Caloglossa Assoziation [On the distribution and ecology of the Bostrychia-Caloglossa association]. Internationale Revue der gesamten Hydrobiologie 48: 47-152.
• Soe-Htun, U., 1998. The seaweed resources of Myanmar. In: Critchley, A.T. & Ohno, M. (Editors): Seaweed resources of the world. Japan International Cooperation Agency, Yokosuka, Japan. pp. 99-105.
• Tjon Sie Fat, L.A., 1976. Bostrychietum, plantengeografisch onderzoek over de Bostrychia-Caloglossa-gemeenschap, de algenformatie van de mangrovebossen [Bostrychietum, a plant-geographical study on the Bostrychia-Caloglossa association, the algal formation of the mangrove forests]. MSc. Thesis, Rijksherbarium, Leiden University, The Netherlands. 115 pp.
• Tseng, C.K., 1942. Marine algae of Hong Kong, II. The genus Catenella. Journal of the Washington Academy of Sciences 32: 142-146.
• Zablackis, E. & Santos, G.A., 1986. The carrageenan of Catenella nipae Zanard., a marine red alga. Botanica Marina 29: 319-322.
• Zablackis, E., West, J.A., Liao, M.-L. & Bacic, A., 1993. Reproductive biology and polysaccharide chemistry of the red alga Catenella (Caulacanthaceae, Gigartinales). Botanica Marina 36: 195-202.

Sources of illustration

Børgesen, F., 1938. Catenella nipae used as food in Burma. Journal of Botany (London) 76: Fig. 1, p. 268 (habit); Min-Thein, U. & Womersley, H.B.S., 1976. Studies on southern Australian taxa of Solieriaceae, Rhabdoniaceae and Rhodophyllidaceae (Rhodophyta). Australian Journal of Botany 24: Fig. 17, p. 52 (detail tetrasporangia, detail with carpogonial branches); Post, E., 1963. Zur Verbreitung und Ökologie der Bostrychia-Caloglossa-Assoziation [On the distribution and ecology of the Bostrychia-Caloglossa association]. Internationale Revue der gesamten Hydrobiologie 48: Fig. 8, p. 125 (details with haptera). Redrawn and adapted by P. Verheij-Hayes.

Authors

• R.J. King