Heterokontophyta (Ochrophyta)
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• Ochrophytes are algae of diverse organization and include unicellular, colonial, filamentous and parenchematous thalli. • They are characterized by the presence of chlorophylls a and c in their plastids as well as xanthophylls (e.g. fucoxanthin) and other carotenoids that mask the chlorophylls (Table 2). • Due to the presence of these pigments, many ochrophytes have a yellowish-green, gold or brown appearance. • As storage products, they accumulate oils and Heterokontophyta chrysolaminarin (C) in cytoplasmic vesicles, but never starch. (Ochrophyta) • Cell walls contain cellulose, and in certain species they contain silica. • Cells possess one or more plastids, each with an envelope formed by two membranes of chloroplast and two membranes of chloroplast endoplasmic reticulum. • Thylacoids, in stacks of three, in most ochrophytes are Fig. Semidiagrammatic drawing of a light surrounded by a band of thylakoids, girdle lamella, just and electron microscopical view of the basic organization of a cell of the beneath the innermost plastid membrane. Chrysophyceae. (C) Chrysolaminarin • Ochrophytes have heterokont flagellated cells with two vesicle; (CE) chloroplast envelope; different flagella, an anterior tinsel (mastigonemes) and (CER) chloroplast endoplasmic reticulum; (CV) contractile vacuole; (E) posterior whiplash (smooth) flagellum (Fig. ). eyespot; (FS) flagellar swelling; (G) Golgi • In this group of algae, mastigonemes consist of three body; (H) hair of the anterior flagellum; parts, a basal, tubular and apical part formed by fibrils. (MB) muciferous body; (MR) microtubular root of flagellum; (N) nucleus. Chrysophyceae and related groups The following classes are commonly recognized in this division and will be discussed here: • Chrysophyceans (Golden-brown algae) are mostly unicellular 1. Chrysophyceae (golden-brown algae) flagellate organisms. 2. Synurophyceae • Some are amoeboid or coccoid. 3. Eustigmatophyceae • Most of the species in the Chrysophyceae are freshwater, occur in 4. Pinguiophyceae unpolluted and soft waters (low in calcium). 5. Dictyochophyceae (silicoflagellates) • Some are strictly marine algae and part of the nanoplankton. 6. Pelagophyceae • Some have a single functional flagellum like Chromulina, while others 7. Bolidophyceae can form colonies like the freshwater genus Uroglena. 8. Bacillariophyceae (diatoms) 9. Raphidophyceae (chloromonads) 10.Xanthophyceae (yellow-green algae) 11.Phaeothamniophyceae 12.Phaeophyceae (brown algae) • Phylogenetic analysis using rDNA nucleotide sequences of the 16S subunit have shown that the classes Chrysophyceae, Synurophyceae, Eustigmatophyceae, Raphidophyceae, Pelagophyceae and Dictyochophyceae are evolutionarily close. Phaeophyceae, Xanthophyceae, Phaeothamniophyceae, Pinguiophyceae and Chrysomerophyceae are also related, and Bacillariophyceae and Chromulina Uroglena Bolidophyceae form an isolated group. • The chloroplasts are parietal and • Many of the Chrysophyceae have a usually only a few in number, often tinsel flagellum that is inserted at the only one or two. anterior end of the cell parallel to the • Chlorophylls a, c1, and c2 are cell axis and a whiplash flagellum that present, with the main carotenoid is inserted approximately perpendicular being fucoxanthin which give them a golden color. to the tinsel flagellum. • The chloroplasts are surrounded by • The posterior whiplash flagellum is two membranes of chloroplast E.R., usually the shorter flagellum and has a the outer membrane of which is swelling at its base on the side toward usually continuous with the outer the cell contains an electron-dense membrane of the nuclear envelope. area referred to as the photoreceptor. • The thylakoids are usually grouped • The flagellar swelling fits into a three to a band. depression of the cell immediately • Pyrenoids are common in beneath which, inside the chloroplast, chloroplasts of the Chrysophyceae. is the eyespot. • The storage product is chrysolaminarin (leucosin), a β- • The eyespot consists of lipid globules 1,3 linked glucan, supposedly found inside the anterior portion of the in a posterior vesicle (C). chloroplast, between the chloroplast • Contractile vacuoles in the anterior envelope and the first band of portion of the cell (CV). Semidiagrammatic drawing of a light and electron thylakoids. Semidiagrammatic drawing of a light and electron microscopical view of the basic organization of a cell of microscopical view of the basic organization of a cell of the Chrysophyceae. (C) Chrysolaminarin vesicle; (CE) the Chrysophyceae. (C) Chrysolaminarin vesicle; (CE) chloroplast envelope; (CER) chloroplast endoplasmic chloroplast envelope; (CER) chloroplast endoplasmic reticulum; (CV) contractile vacuole; (E) eyespot; (FS) reticulum; (CV) contractile vacuole; (E) eyespot; (FS) flagellar swelling; (G) Golgi body; (H) hair of the flagellar swelling; (G) Golgi body; (H) hair of the anterior flagellum; (MB) muciferous body; (MR) anterior flagellum; (MB) muciferous body; (MR) microtubular root of flagellum; (N) nucleus. microtubular root of flagellum; (N) nucleus. ١ • Most of the Chrysophyceae are sensitive to changes in the environment and • The presence of cellulose in cell walls is common, and some species survive the unfavorable periods as statospores. are covered with scales or protected by an organic sheath called the • The formation of a cyst or statospore or resting spore is one character by lorica (L), an envelope around the protoplast, but not generally which a member of the Chrysophyceae or Synurophyceae may be recognized. attached to the protoplast as a wall is. • Statospores, are shaped like a small, externally ornamented bottle enclosed in a silicified wall with a terminal pore closed by nonsilicified plug (P). • Scales (if they are present) are made of silica and are radially or • A vegetative cell forms a statospore internally. biradially symmetrical • Very few chrysophyceans are naked. • When a statospore germinates, there is a dissolution of the plug or separation of it from the spore wall. The protoplast then moves out of the statospore by amoeboid motion, forming flagella as it moves out. Chrysococcus rufescens. (a) Whole cell. (b) Cell Formation of a statospore or cyst Statospore of Ochromonas sphaerocystis. undergoing reproduction. (c) Ultrastructure of in Ochromonas tuberculata. (a)– vegetative cell. (B) Branched cytoplasmic process; (E) (c) The formation of the eyespot; (G) Golgi; (L) lorica; (LV) leucosin vesicle; (LF) statospore. (d) A mature long flagellum; (N) nucleus; (SF) short flagellum; (V) statospore as seen from the collar contractile vacuole. end. (C) Chloroplast; (Co) collar of statospore; (Cr) chrysolaminarin vesicle; (CV) contractile vacuole; (D) discobolocyst; (N) nucleus; (P) plug in pore of statospore; (S) statospore wall; (SDV) silica deposition vesicle; (Sp) spine. • Two different types of projectiles occur in the • Nutrition in the Chrysophyceae can be either phototrophic, Chrysophyceae, muciferous bodies and phagotrophic, or mixotrophic (photosynthetic organism capable of discobolocysts. taking up particles and molecules from the medium). • On discharge the contents of the vesicle in • Food particles include both living (bacteria, small algae, or even muciferous bodies (MB) often form a fibrous network cells of its own kind) and non-living (detritus, fecal material). outside the cell. • The mixotrophic chrysophytes, Epipyxis pulchra have the ability to • The discobolocysts (D) are in the outer layer of cytoplasm and consist of a single membrane select or reject specific food item. bounded vesicle with a hollow disc in the outward facing part of the vesicles. The discharge is explosive, taking place by the expansion of the projectile into a thin thread 6 to 11 µm long. Semidiagrammatic drawing of a light and electron microscopical view of the basic organization of a cell of the Chrysophyceae. (MB) muciferous body. Phagotrophy in Epipyxia pulchra. The cell has a posterior stalk by which it is attached to a lorica. (a) The long tinsel flagellum beats in such a way that water and suspended particles are drawn to the cell. (b) A particle is seized by the long tinsel flagellum. (c) The particle is maneuvered between the (a) Ochromonas tuberculatus. (D) discobolocyst; (b) two flagella. (d) A feeding cap from the cell envelopes the particle. (e) The particle is enclosed within Charged and discharged discobolocysts. a food vacuole within the cytoplasm. The stalk has pulled the cell into the lorica. • Chrysophytes are notorious for their production of fishy or rancid smells, reflecting release of unsaturated aldehydes derived from the high cell content of polyunsaturated acids. • Mitotic division is the most • These chemicals are classified as algal volatile organic common mechanism of compounds (AVOCs). asexual reproduction. • Sexual reproduction is rare, but when it occurs, gametes are anisogamous. • Vegetative cells are haploid, and meiosis is the first division of the zygote. • In some cases, zygotes are statospores, representing a resting phase. Three unsaturated fatty-acid derivatives produced by chrysophytes that result in rancid or fishy odors. The life cycle of Dinobryon. ٢ • Eustimatophytes are yellow-green unicells that • Synurophyceae are flagellate algae occur in freshwater, brackish water, and seawater as covered with silica-scales (S). well as in the soil. • They are closely related to the Chrysophyceae. • They produce naked zoospores. • The Synurophyceae differ from chrysophyceans in • Most bear a single pleuronematic