stramenopiles (heterokontophyta) - univerzita karlova · stramenopiles - flagella functionally...
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Stramenopiles (Heterokontophyta)
• autotrophs – from diatoms to brown algae... – (chlorophylls a + c1 c2 c3)
• heterotrophs, parasites
• chrysolaminaran and oil, starch is not produced !!!
• usually 2 flagella, 2 rows of tubular mastigonemata (from glycoproteins, production in ER) – tripartite structure (basal part, tubular part, hairs)
Stramenopiles (Heterokontophyta)
Chrysophyceae
- Ochromonas
BR – basal part
TS – tubular part
LE – hairs
Stramenopiles - flagella
functionally differentiated
flagella - heterokont
longer anterior flagellum,
pleuronematic – with
mastigonemata
shorter posterior, usually
smooth, often reduced,
associated with stigma
• 4 membranes, thylakoides stacked in triplets, girdle lamella
nuclear membrane
2layered plastid membrane
nucleus
pyrenoid chrysolaminaran vacuole
plastid ER
girdle lamella
lamella (triplets)
oil globules
Stramenopiles - plastids
• plastids: chl a + c1 c2 c3 in different combinations
β-carotene, fukoxanthin (brown colour)
vaucheriaxanthin (Xanthophyceae, Eustigmatophycae)
Stramenopiles - plastids
• originally 3 classes (Cavalier-Smith) - Heterokontophyta:
– Chrysophyceae
– Bacillariophyceae
– Phaeophyceae
– thereafter additionall autotrophic groups aded
• David J. Patterson – Stramenopiles
– mastigonemata, tubular crists in mitochondria, open mitosis
– heterotrophic groups included
Stramenopiles - classification
Ochrophyta:
• Chrysophyceae (zlativky)
• Dictyochophyceae (dictyochy)
• Phaeothamniophyceae
• Pelagophyceae
• Bolidophyceae
• Phaeophyceae (chaluhy)
• Xanthophyceae (různobrvky)
• Eustigmatophyceae
• Bacillariophyceae (rozsivky)
• Raphidophyceae (chloromonády)
• .....
heterotrophs:
• Pseudofungi
• Bicosoecida
• Labyrinthulomycetes
• Opalinata
• Blastocystis
Stramenopiles - classification
Chromista, Stramenopiles, Ochrophyta
Yang et al. 2012
Bolidophyceae
• marine flagellates
• picoplankton (1.2 mm)
• heterokont flagella
• sister group to diatoms
• established in 1999
Bolidophyceae
diatoms probably
evolved from flagellates
• picoplanktonic marine flagellates
Bolidophyceae
Bolidomonas
Parmales • nannoplanktonic, coccoid algae
• siliceous cell wall composed of oval or three-sided cell wall
(similar to diatoms)
• cold marine waters (Antarctida)
Parmales
Parmales
Dictyochophyceae
General characteristics • flagellates
• heterokont features (flagella, plastid)
• morphologically and ecologically diverse organisms
• Dictyochales – marine, skeleton-bearing species
• Pedinellales – freshwater
• Rhizochromulinales – marine
• Florenciellales – marine
Chang et al. 2012
Dictyochales
siliceous skeleton
nucleus
flagellum
pseudopodium
mucilaginous
envelope
• siliceous skeletons, microtubule-supported pseudopodia
Dictyocha – life cycle • production of naked flagellates without skeleton
Dictyocha – recent species
Dictyocha fibula Dictyocha speculum
Dictyocha – fosil species (silicoflagellates)
Naviculopsis
Dictyocha
• found in cold waters (determination of seawater temp.)
Pedinellales
• mixotrophic and heterotrophic species
• in freshwater and brackish waters
• rare
Pedinellales autotrophic
Pedinella
Apedinella Pseudopedinella
Pedinellales heterotrophic
Pteridomonas
Actinomonas
Pedinellales heterotrophic
Actinosphaerium Actinophrys
Rhizochromulinales
• autotrophic
• amoeboid cells with pseudopods
• marine (puddles in tidal zones)
Florenciellales • Florenciella • Pseudochatonella
Pelagophyceae
General characteristics
• class described in 1993
• marine organisms of various thalli types
• tiny flagellates, as well as large
seaweed-like organisms
• Pelagomonadales, Sarcinochrysidales
Pelagomonadales • picoplanktonic algae
• paraflagellar bar, reduced system of flagella roots
Pelagomonas – Pacific
Pelagococcus – free-living of symbiotic in foraminifera
Sarcinochrysidales • benthic, filamentous or thallose organisms
• sometimes very abundant, and considered as a pest
Sarcinochrysidales Chrysocystis (noodle-alga)
Sarcinochrysidales
Chrysophaeum
Yang et al. 2012
PX clade
• includes Phaeophyceae and Xanthophyceae
• presence of electron-opaque vesicles beneath the plasma
membrane (= physodes in Phaeophyceae) – probably
associated with the production of unique cell wall materials
Xanthophyceae
General characteristics
• ca 100 genera, 600 species
• mostly coccoid or filamentous
• phytoplankton of freshwater bodies
• soil algae
General characteristics
• majority of species rare, in a small number
Chloroplasts • yellow-green color
• chrysolaminaran as storage product
Flagellates
plastid ER
2layered plastid membrane
girdle lamella
lamella (triplets)
nuclear membrane
• subapical flagella, acronema in small flagellum
• stigma in chloroplast, photoreceptor photoreceptor
Cysts
• only in some species
• endogenous, impregnated by silica
Cell wall
• often composed by two pieces, impregnated by silica
Bumilleriopsis Tribonema
Reproduction
• asexual – vegetative division, zospores, aplanospores
• sexual – only Vaucheria
Evolutionary parallelism
Characiopsis
Chloridella
Bracteacoccus Botrydiopsis
Characium
Chlorella
Evolutionary parallelism
• chromatography, ultrastructure
Xanthophyceae Chlorophyceae
668 nm – chl a 668 nm
668 nm
668 nm
634 nm – chl c
634 nm 634 nm
Traditional classification
• 7 orders (Bailey & Anderson 1998)
Rhizochloridales
• amoeboid thallus, pseudopodia
• very rare
Myxochloris – multinuclear plasmodia in hyalocytes, Krkonoše Mts.
Mischococcales
• coccoid species, rigid CW (in some species in two pieces)
• metaphyton, soil, rarely in phytoplankton
Mischococcales Goniochloris – two-pieced CW, plankton
Mischococcales Characiopsis – epiphyte
Mischococcales Botrydiopsis – up to 60 mm in diameter
Mischococcales Borydiopsis – up to 60 mm in diameter
Mischococcales Bumilleriopsis
Mischococcales Ophiocytium – dendroidal, colony-like clusters
Mischococcales Mischococcus – dendroid colonies in submerged plants
Tribonematales • branched or simple filamentous thallus
Tribonematales Tribonema – frequently in clear, slightly acidic waters
Tribonematales Xanthonema – frequent soil alga
Tribonematales Bumilleria
Tribonematales Heterococcus – branched filaments, in soil
Tribonematales Heterococcus caespitosus – photobiont of lichen Verrucaria
Botrydiales Botrydium granulatum – multinuclear, siphonal thallus
• saclike vesicles+ rhizoids
• grows on damp soil
• zoospores, aplanospores
Vaucheriales Vaucheria – branched, coenocytic thallus; freshwater, brackish, soil
Vaucheriales Vaucheria – cell organisation
apical
zone
subapical
zone
vacuolized
zone
sporangium synzoospore
asexual
reproduction
sexual
reproduction antheridium
oogonium
spermatozoides
hypnozygote
Vaucheriales
Vaucheriales Asterosiphon – enigmatic macroscopic alga
Cambra-Sánchez, 2011
Phylogeny • cell wall organisation
Phylogeny
Chlorellidium
Botrydiopsis
Vaucheria
Tribonema
Maistro et al. 2009
• 4 major clades
• Botrydiopsis
pyrenoidosa is not a
Xanthophyte
• Pseudobumilleriopsis
as a separte genus
Pleurochloridella botrydiopsis
• differs from Xanthophytes by possessing fucoxanthin
Phaeothamniophyceae
Phaeothamnion – freshwater bodies, pools and ponds
• unbranched or branched filaments
• formally in Chrysophyceae
• class established in 1998
(analysis of molecular data)
Phaeothamniophyceae Stichogloea – mucilaginous colonies, in alpine lakes
• Aurearena cruciata – sandy beaches of Japan
• coccoid algae, flagella retained underneath the cell wall
Aurearenophyceae
Tetrasporopsis fuscescens
• cool and slow- or moderately fast-flowing water
• traditionally classified in Chrysophyceae
Chrysomerophyceae
Giraudyopsis
• marine benthic alga
Chrysomerophyceae
• branched filaments
• related to brown algae
Schizocladiophyceae
Stramenopiles age • radiation in Ordovician
• impact to the evolution/diversity of green algae?