Physiology and Cultivation of Algae and Cyanobacteria 1. Definition • Algae formal tax. standing, polyphyletic origin, artificial assemblage of O2 evolving photosynthetic organisms • Algae vs. Plants the same storage compounds, similar defence strategies against predators & parasites Plants •hi degree of differentiation •repr. organs surrounded by jacket of sterile cells •multicell. embryo remains developmentally & nutrit. independent on parents •meristems on root/shoot apices •digenetic life cycles with alterations betw. hapl. gametophyte & dipl. sporophyte Algae •don’t have roots, stems, leaves, not well defined vasc. tissue •don’t form embryo •mono- & digenetic life cycles •occur in dissimilar forms (micro algae, macro a. multicellular, colonies, branched,…) •less complexity of the thalli •hi diversity 0.2 – 60m ecology & habitats reserve & structural polysaccharides evolutionary origin •1 – 10 mil. species •½ primary production in biosphere Classification • under constant revision (Van Den Hoek et al. 1995) Occurrence & distribution Aquatic • almost everywhere (from freshwater spring to salt lakes) • tolerance of wide range of pH, temp., turbidity, O2 & CO2 conc. • planctonic » unicellular, suspended throughout lighted regions of all water (inc. polar ice) • benthic » within sediments » limited to shallow areas (because of rapid attenuation of light with depth) » attached to stones – epilithic, on mud/sand – epipelic » on other algae/plants – epiphytic, on animals – epizoic • marine benthic – after habitat – supralitoral – above high-tide level within reach wave spray – intertidal – exposed to tidal cycles – sublitoral – from extreme low-water to cca 200m deep • ocean – 71% of earth surface, more than 5000 spec. of planktonic algae – phytoplankton » base of marine food chain » produce 50% of O2 we inhale - life » death – blooms – too large populations (decrease water transparency, prod. toxins & poisons) – kelps » giant algae – temperate pelagic marine environment, till 60m submerged forests » also beneath polar ice sheet » can survive at very low depth – record of 268m u.s.l. – dark blue red algae (blue-green ligh, 0.0005% of surface intensity) » have accessory pigments, channel the energy to chl a • accessory & protective pigments – give algae wide variety of colors <> names https://doi.org/10.1007/s00338-017-1594-5 Occurrence & distribution Freshwater phytoplankton & benthic algae » base of aquatic food chain » not exhibit size range of marine relatives Subaerial » life on land » tree trunks, animal fur, snow, hot springs, desert rocks » activity – convert rock > soil − to minimize soil erosion & increase water retention & nutrient availability for plants Symbiosis • lichens, corals » to survive in environments that they could not alone Structure o thallus Unicells & unicell colonial algae • solitary cells, unicells with/w-out flagella, motile (Ochromonas)/non-motile (Nannochloris) • colony – aggregates of several single cells held together ±organized – grow – cell division – each cell can survive solely • coenobium – colony with number of cells & arrangement determined at the time of origin (e.g. Volvox – motile, Pediastrum – non-motile) Structure of thallus Filamentous algae – result from cell division in plane perpendicular to axis of filament – cell chain ─simple └branched – true/false ─uniseriate – 1 layer of cells └multiseriate – up to multiple layer Syphonous algae – siphonous/coenocytic construction of tubular filaments lacking transverse cell walls – unicellular but multinucleate (coenocytic) Structure o thallus Parenchymatous & pseudoparenchymatous algae – mostly macroscopic • parenchymatous » originated from division of primary filament (all directions) » lost filamentous structure • pseudoparenchymatous » originated from close aggregation of branched filaments, forming thallus held together with mucilages (red algae) Nutrition • algae = phototrophs • most algal divisions – contain colorless heterotrophic spec. – osmotrophy, phagotrophy – auxotrophy – cannot synthesize essential components (vitamin B12, fatty acids,..) and have to import them • algae can use wide spectrum of nutritional strategies combining: – phototrophy – heterotrophy – mixotrophy (relative contribution of photo.&hetero. can vary) » often in extreme environment (limiting light,…) – after nutritional strategies: – obligate heterotrophic algae – primarily heterotrophs, but capable phototrophy in limiting prey concentration (Gymnodium gracilentum - Dinophyta) – obligate phototrophic algae – primarily phototrophs, but capable phagotrophy/osmotrophy when light is limiting (Dinobryon divergens - Heterocontophyta) – facultative mixotrophic algae – can equally well grow as photo-/heterotrophs (Fragilidinium subglobosum - Dinophyta) – obligate mixotrophic algae – primary mode is phototrophy & phago-&/osmotrophy provides essential substances (e.g.photoauxotrphs, Euglena gracilis - Euglenophyta) Reproduction • vegetative by division of single cell or fragmentation of colony • asexual by production of motile spores • sexual by union of gametes – vegetative & asexual » allow stability of adapted genotypes from generation to the next » fast & economical increase of number of individual » lack genetic variability – sexual » involves - plasmogamy (union of cells) - karyogamy (union of nuclei) - chromosome/gene association & meiosis >> genetic recombination » allow variation, but is more costly Vegetative & Asexual reproduction • Binary fission & Cellular bisection – simplest form – parent org. divides into two equal parts of the same hereditary info as parent – unicellular a. - longitudinal - transverse – growth of population - lag > exponential > log > stationary (plateau) phase – in multicellular a. & colonies - leads to the growth of individual • Zoospore, Aplanospore & Autospore – zoospores - flagelate motile spores that may be produced within parental vegetative cell (Clamydomonas - Chlorophyta) – aplanospores - aflagelate spores that begin their development within parent cell wall before being released - can dvelop into zoospores – autospores - aflagelate daughter cells released from ruptured cell wall of parental cell, - replicas of vegetative cells that produce them & lack the capacity to develop into zoospore (Nannochloropsis - Heterocontophyta, Chlorella - Chlorophyta) spores - may be produced within - ordinary cells - specialized sporangia Vegetative & Asexual reproduction • Autocolony formation – coenobium/colony - each cell can produce new colony similar to parent. – cell division produce multicellular group (not the unicellular individuals) > differs from the parent in cell size not in number e.g. Volvox (Chlorophyta) – gonidia - series of cells which produce a hollow sphere within the hollow of parental colony (released after its rupture) • Fragmentation – ± random process whereby non-coenobic colonies/filaments break into two/several fragments having capacity to develop into new individual Vegetative & Asexual reproduction • Resting stages – under unfavourable conditions (desiccation) – thick-walled cells • hypnospores & hypnozygotes – thick-walled, produced ex novo from cells previosly separated from parent cells » hypnospores - Ulothrix spp., Chlorococcum (Chlorophyceae) » hypnozygotes - Spyrogyra spp. (Chlorophyceae), Dinophyta – enables algae to survive temporary drying out of small water bodies & allow transport to another (e.g. via birds) • statospores – endogenous cysts formed within vegetative cells by members of Chrysophyceae e.g. Ochromonas spp. » cyst walls consist of silica >> preserved as fossils – spherical, ellipsoidal, often ornamented with spines or other projections – wall - with pores sealed by unsilicified bung – within cysts lie nucleus, chloroplasts, reserve material – after dormancy - germination - form one/several flagellate cells • akinetes occurrence in blue-green algae – enlarged vegetative cells that develop thickened wall in response to limiting env. nutrients or light (e.g. Anabaena cylindrica - Cyanophyta) – extremely resistant to drying & freezing – long-term anaerobic storage of genetic material, remain viable in sediments for many years in hard conditions – in suitable conditions > germination into new vegetative cells Sexual reproduction Gametes – morphologically identical/different with/from vegetative cells (a. group speciphic sign) – haploid DNA content – possible different gamete types • isogamy - both gametes types motile & indistinguishable • heterogamy - gametes differ in size » anisogamy - both gametes are motile, 1. small - sperm 2. large - egg » oogamy - 1. motile, small - sperm 2. non-motile, very large - egg Algae exhibit 3 different life cycles with variation within different groups • main difference - where meiosis occur & type of cells it produces & whether there is more than one free-living stages Sexual reproduction Haplontic or zygotic life cycle – single predominant haploid vegetative phase, with meiosis after germination of zygote » Chlamydomonas (Chlorophyta) Diplontic or gametic life cycle – single predominant diploid vegetative phase – meiosis gives rise to haploid gametes » Fucus (Heterocontophyta), Diatoms Haplontic or zygotic life cycle Diplontic or gametic life cycle Sexual reproduction Diplohaplontic or sporic life cycle – present alternations of generation between two different phases consisting of haploide gametophyte & diploid sporophyte • gametophyte - produce gamete by mitosis • sporophyte - produce spore by meiosis – alternation of generations can be • isomorphic - both phases morphologicaly identical » Ulva (Chlorophyta) • heteromorphic - with predominance of – sporophyte - Laminaria (Heterocontophyta) – gametophyte - Porhyra (Rodophyta) Diplohaplontic or sporic life cycle -isomorphic alternation of generations Diplohaplontic or sporic life cycle -heteromorphic alternation of generations