SEMESTER-I TOPIC: RODOPHYTA (RED )

MBOT CC-1 UNIT-1(Phycology, Mycology& )

RITA KUMARI, H.O.D

UNIVERSITY DEPARTMENT OF , P.P.U. PATNA

Introduction

General Characters Occurrence Morphology

Cell structure Range of thallus Structure Cell Structure PIT-CONNECTION Reproduction classification phylogeny SUMMARY rEFERRENCES

INTRODUCTION The Rhodophyceae or the is probably the oldest groups of Eukaryotic Algae encompassing about 6500 species of mostly multicellular and predominantly marine algae. It is the only class in the division Rhodophyta. The members Rhodophyceae along with some members of and Phaeophyceae form a group of macro algae commonly called as seaweeds. They owe their red colour due to the photosynthetic pigment phycoerythrin. Despite their name, not all Rhodophyceae are red. These algae may also be purple, brown or black because of presence of the additional pigments. The pigments present are chlorophyll a, chlorophyll d, carotenes, phycoerythrins and phycocyanins. General Characters, Rhodophyta characterized by six main features:

1. The motile flagellate stages are totally absent. Even the gametes and the spores lack flagella. 2. The unique photosynthetic pigments are- chlorophyll–d, biliprotein (phycobilins) r- phycoerythrin and r-phycocyanin and the xanthophyll-taraxanthin besides, the chromatophore generally contain chlorophyll-a and the alpha and β-carotene, lutein, zeaxanthin, neoxanthin. 3. The reserve foods are and glactose florisides accumulate in cytoplasm. 4. The cell wall contains polysulphate esters of carbohydrates in addition to cellulose and pectin. 5. Adjacent cells are connected to each other by specialized pits which permit cytoplasmic connection between adjacent cells. 6. Reproduction is highly specialized; the male gamete, called the spermatium, is non motile and at time of fertilization is passively transported by the water currents and is lodged on the trichogyne of the female sex organ –carpogonium. The post fertilization changes are also unique in this group.

Occurrence: Except a few freshwater forms (Batrachospermum, , Compsopogon. thorea, Hildendrandtia) others are exclusively marine (98%) and grow mostly in the intertidal and sublittoral regions..There greater concentration in the warmer seas. A few grow in cold waters of Polar Regions. Some red algae ( , ) are calcareous and forms coral reefs in the sea. Some are parasites of macrophytic marine algae. Range of thallus Structure The Rhodophyta are the most beautiful and most showy of the marine algae. Forms such as and Delleseria are really the beautiful creation of the nature. The Subclass Bangioidea, with single order Bangiales, comprise unicellular species like Porphyridium, Rhodospora; filamentous( Goniotrichum)and parenchymatous forms Bangia, Porphyra. The thalli of possesses a single axial filament having branched laterals with or without cortication is called a uniaxial filament (Batrachospermum). Those thalli which possess a tuft of axial filaments with many laterals radiating out to the margins are called multiaxial filaments (). The growth in Bangioidea is diffuse where is in of Florideae apical. Cell Structure

Cells are eukaryotic and non motile .The cell wall of red algae is made up of an inner microfibrillar cellulosic layer and outer pectic layer. Although in Porphyra and Bangia, a xylan performs this function. The red algal cell wall contains various types of highly hydrophilic, sulfated polygalactans. agars and carrageen. Some members of (coralline red algae) deposit calcium carbonate extracellular in their cell walls. Chromatophore are single stellate, axile with a central naked in (Bangioidea) or many discoid and parietal devoid of (Florideae). Both phycoerythrin and phycocyanin are present in the phycobilisomes .Those members who do not have much phycoerythrin appear green or blue (Batrachospermum). Phycoerythrin allows red algae to live and photosynthesize at great depths below the surface of the oceans. Phycoerythrin has the ability to absorb blue light, which penetrates water at greater depths than light of longer wavelength generally used for photosynthesis. In addition to phycobilisomes, many scattered grains of floridean starch occurs in cytoplasm. Floriside is a major photosynthate in Orders Bangiales, , Rhodymeniales. In lower rhodophyceaethe cells are uninucleate e.g.; Bangiales. In others the cells are multinucleate e.g.; , Rhodymeniales.

Pit connections Pit connections are unique to red algae. These are the connections between two adjacent cells that are filled with protein. These consist of lens-shaped plugs and caps. Pit connections are the sites of structural strength in the thallus. In some algae, the plugs become dislodged in the reproductive cells hence allowing the passage of metabolites to the developing reproductive cells. The pit connections between the axial cells and the pericentral cells are called as the primary pit connections. Those between the adjacent pericentral cells are the secondary pit connections.

Reproduction Reproduction occurs by vegetative, asexual and sexual methods. Vegetative Reproduction: Unicellular forms reproduce by fragmentation

Fig 1 Different forms of Rhodophyceae

Asexual Reproduction Asexual reproduction takes place by means of non motile spores and are given different names depending on the nature of cells from which they are produced and their number within each sporangium. Accordingly, there are four chief types of asexual spores (1) monospores- formed singly in monosporangia, (2)neutral spores-formed by direct transformation of vegetative cells into spores(Bangioidea);(3)Carpospore-formed by division of zygote in (Bangioidea); or the diploid carpospores are formed singly in the carposporangia ( Florideae) (4)bispores, tetraspores and polyspores formed within diploid tetrasporangium in multiples of two, four and multiples of four within a sporangium of the diploid tetrasporophyte. Except for some red algae that form haploid carpospores growing into sexual , most others red algae form diploid carpospores which on germination give rise to sporophyte or diploid plants,

bearing diploid sporangia from which bispores, tetraspores and polyspores are produced after meiosis. Sexual Reproduction Sexual Reproduction is highly advanced, elaborate and oogamous. The male gametes are called spermatia and develop in spermatangia and female sex organs are called oogonia. The sexual life histories of red algae have two (Diphasic) or three multicellular phases (Triphasic). Triphasic Life Cycle Most of the red algae have three distinct phases in their life cycle- the gametophyte, carposporophyte and the tetrasporophyte. 1. Gametophyte is a free-living haploid which is concerned with sexual reproduction. It produces the gametes. 2. Carposporophyte is a diploid plant which develops from the zygote after sexual fusion is accomplished. It is parasitic on the female gametophyte and hence remains attached to it. Carposporophyte is concerned with the production of diploid spores called the carpospores. Carpospores germinate to give rise to tetrasporophyte. 3. Tetrasporophyte is an independent plant like the gametophyte. It is the site of meiosis and is concerned with the production of haploid spores called the tetraspores. Tetraspores further germinate to give rise to gametophyte of the next generation. and is concerned with the production of haploid spores called the tetraspores. Tetraspores further germinate to give rise to gametophyte of the next generation. Diphasic Life Cycle In simple red algae like Batrachospermum, the carpospores are haploid and on germination produce sexual plants. There is no asexual plant except the diploid zygote which soon undergoes meiosis. Thus there are three haploid generations the parent gametophyte concerned with the production of gametes. It is independent; - carposporophyte concerned with production of carpospores which are haploid. It is parasitic on the female gametophyte; carpospores which germinate to form the charantia stage which gives rise to the gametophytes. The life cycle of Batrachospermum thus involves alternation of successive haploid generations with the diploid zygote. Cytologically it is haplobiontic with only one kind of individuals. On the other hand, life cycle of Polysiphonia is typically triphasic with two diploid and one haploid generation. Cytologically, such life cycles are diplobiontic in which there is a regular alternation of one gametophyte phase with two sporophyte phases.

Fig 2 Triphasic Life Cycle

Carpogonium It is the female sex organ and consists of a dilated basal portion and a very narrow elongated tip, the trichogyne. The trichogyne is gelatinous and and is instrumental in receiving the male cells. A carpogonium usually possesses two nuclei; one in the trichogyne and the other in the basal part. The trichogyne nucleus degenerates soon after the carpogonium attains maturity whereas the nucleus in the basal part functions as the female gamete nucleus. Carpogonium arises in the terminal portion of a 3-4 celled carpogonial branch and the cell which gives rise to carpogonium is the supporting cell. Both the carpogonium and the carpogonial branch are generally colourless.

Fig 3.A Spermatangia producing spermatia Fig 4 A. Carpospore B. carpogonium.

Fig 5 Tetrasporangia of red algae

Spermatium A mature spermatium is a uninucleate non motile and sphericalor oblong structure with no cell wall. Spermatia are released by the gelatinization of the spermatangial wall near the apex. Fertilisation Water currents carry the spermatia to the trichogyne of the carpogonium. The walls of the male and female gametes dissolve at their point of attachment thus facilitating the movement of the male nucleus to the carpogonium. Fusion of the male and the female gametes takes place in the basal portion of the carpogonium. The fertilized carpogonium does not form a diploid zygote as seen in other groups of algae but instead puts out another long filament that carries the diploid nucleus out of the carpogonium and deposits it into a totally different cell called as the auxillary cell which is in close proximity of the carpogonium. Auxiliary cells are of two types: nutritive and generative. Nutritive cells provide nutrients for developing carposporophyte. And the generative cells give rise to gonimoblast filaments. It is the diploid tissue of fertilized carpogonium which forms the gonimoblast filaments. These gonimoblast filaments bear terminal carposporangia which form the carpospores. Carposporangia enlarge because of development of chloroplasts and vesicles containing wall precursors. Breaking of pit connections between the carposporangium and gonimoblast filaments lead to release of carpospores. Carpospores slowly swim away, eventually germinating and growing into diploid tetrasporophytes. These tetrasporophytes bear tetrasporangia and each tetrasporangium bears four tetraspores. Tetraspores germinate to give rise to male and female gametophytes. Economic Importance

1. Food:

A number of red algae are edible, e.g., Porphyra (Laver), (Dulse), (Irish ). Rhodymenia (also called sheep’s weed) is also used as fodder.. Porphyra (popular Japanese name is nori )is a popular flavouring agent, an ingredient of soups and salads, a wrapper for sushi and is eaten with rice and fish.

2. Phycocolloids:

A number of phycocolloids are extracted for commercial use. They include agar, carrageenin and funori. Agar is used in solidifying laboratory culture media and is added as stabilizer or thickener in the preparation of jellies, puddings, creams, cheese, bakery, etc. Agar is obtained from cell wall of and . Carrageenin is used as a clearing agent in liquors, leather finishing and as emulsifier in chocolates, ice-creams, and tooth pastes, paints, etc. It is extracted from Chondrus. Funori is glue used as adhesive and in sizing textiles, papers, etc. 3. Bromine: Bromine is obtained from some of the red algae, e.g., Rhodomela Polysiphonia.

4. Medicines: Corallina is capable of curing worm infections. Polysiphonia has antibacterial properties. Agar is laxative. Carrageenin can coagulate blood.

Classification

Fritsch 1935 divided the class into two orders- Class. Rhodophyceae Sub-class Bangioideae in which there is intercalary growth of the thallus and there is direct division of the zygote into carpospores .

Order. 1. Bangiales

Sub-class Florideae - in which the growth of the thallus is terminal and - the carpospores are formed indirectly from a zygote.

Order. 1. Nemalionales

2.

3. Cryptonemiales

4.

5. Rhodymeniales

6. Ceramiales

Phylogeny

Rhodophyta apparently shows a much closer relationship with Cyanophyta than with any other algal phyla. the feature shared by the two phyla are;

(i) Flagellated or motile cells are absent in both algae.

(ii) The formation of similar phycobilisomes blue (phycocyanin) and red (phycoerythrin) as the accessory photosynthetic pigments .

(iii) Both the groups have a common pattern of fatty acid formation which differs from other plants in that the lipid content does not increase as the thallus grows.

(iv) In both case, photosynthetic thylakoids occur singly and widely separated.

(v) In both case, the principle constituents of mucilage are sulfated galactoses, uronic acid, glucose, and xylose.

(vi)The presence of Pit connections.

(vii) The accumulation of fairly similar food reserves, e.g. floridean starch in the Rhodophyta and cynophycean starch in Cyanophyta.

Regarding the ancestry of the class, two alternatives have been suggested (Klein and Cronquist, 1967): (1) they originated from some simple prokaryotic blue ; and (2) they were derived from some archaic eukaryotic algae which themselves originated from the blue green algae.

Summary

The Rhodophyta (red algae) owe their colour to the presence of excess of r-phycoerythrin in their chromatophores and this red pigment masks the colour of other photosynthetic pigments. The Rhodophyta are characterized by six main features: (1) the flagellated motile stages are totally absent; (2) the sexuality is highly specialized; the male gamete, called spermatium, is motionless and at the time of fertilization is passively transported to and lodged on the trichogyne of the female carpogonium; also, there are distinct post-fertilization developments not found in any other algal phyla; (3) chlorophyll-d, biliproteins (r-phycoerythrin and r-phycocyanin) and the xanthophyll taraxanthin are the characteristic pigments; besides, the chromatophores generally contain chlorophyll-a, α and β carotenes, lutein, zeaxanthin, neoxanthin and rarely other xanthophylls; (4) the reserve foods are floridean starch and galactoside floridosides and these do not accumulate within the chromatophore but outside it, in the cytoplasm; (5) the cell wall contains polysulphate esters of carbohydrates in addition to cellulose and pectin; and (6) the transverse walls in multicellular forms are generally provided with pits which permit cytoplasmic connection between adjacent cells.

References:

1. A TEXT BOOK OF ALGAE H.D. KUMAR AND H.N SINGH EWP 1995

2. TEXT BOOK OF ALGAE O.P. SHARMA; Tata McGraw-Hill 2008.