Division: Chlorophyta (green algae) Chlorophyte Diversity 5 classes: Chlorophyceae Trebouxiophyceae Ulvophyceae Prasinophyceae Charophyceae Part II Land plants 1 2 Ulvophyceae Class Ulvophyceae: Microtubule Roots: cruciate Basal Bodies: counter- clockwise 1. How flagella are attached/constructed: • basal bodies orientation = counterclockwise • microtubule roots = cruciate Flagella Basal Bodies 2. Cell covering: • scales vs. cell wall = wall (scales in some reproductive unicells) 3. How cells actually divide: • spindle = closed • microtubule organization = phragmoplast • division by = furrow Microtubule roots Cell 3 4 1 Ulvophyceans have diverse morphologies Class Ulvophyceae: Mostly marine macroalgae, some freshwater also Haplontic, diplontic or alternation of generations No hypnozygotes Orders to know: unbranched filament branched filament • Ulotrichales • Ulvales (split off from ‘Ulotrichales’) • Cladophorales • Caulerpales (used to be ‘Codiales/Bryopsidales’) • Dasycladales 5 Division: Chlorophyta Class: Ulvophyceae Trebouxiophyceae parenchymatous coenocytic/ siphonous Order Ulotrichales –e.g.Ulothrix Haplontic with codiolum phase 1N thallus, the zygote is the only diploid stage Life -Cycle of the Ulotrichales Haplontic -Codiolum phase- cell wall is from 2n zygote with n spores inside e.g. Ulothrix Chlorophyceae Prasinophyceae syngamy Order: Ulotrichales Ulvales Cladophorales 6 Caulerpales Dasycladales • Spores have scales, not true cell walls 1N gametes 2N zygote meiosis occurs in zygote - Unicells, filaments, blades; uninucleate or multinucleate cells - Simple, haplontic life histories; all with an attached Codiolum stage formed by zygote; zygote undergoes meiosis, producing flagellated meiospores 1N gametophytes 1N codiolum stage • Cell wall of zygote (2N) stays intact to contain new meiospores (N) Focal genera: Ulothrix 1N zoospores 7 8 2 Order Ulvales e.g. Ulva Division: Chlorophyta Class: Ulvophyceae Trebouxiophyceae Order: Ulotrichales Ulvales Isomorphic Alternation of Generations: organism having a separate multicellular diploid sporophyte and haploid gametophyte stage, sporophyte & gametophyte are same sizes Chlorophyceae Prasinophyceae Cladophorales Caulerpales Dasycladales - Uninucleate cells with a single parietal chloroplast - Parenchymatous thallus; mono- or distromatic, sheets or tubes - All cells can reproduce (holocarpic reproduction) - Isomorphic alternation of generations Focal genera: Ulva “sea lettuce” 9 10 Division: Chlorophyta Class: Ulvophyceae Trebouxiophyceae Order: Ulotrichales Ulvales Ulva spp • Distromatic • Blade-like • Often ephemeral in intertidal • Fast growing, tasty to herbivores Ulva intestinalis = • Monostromati c • Tubular • Euryhaline- Tolerates freshwater, often occurs 11 near seeps Chlorophyceae Prasinophyceae Cladophorales Caulerpales Dasycladales - Large, multinucleate cells, connected end to end with cross walls - Chloroplasts reticulate or multiple discoid arrayed in reticulate network - Isomorphic alternation of generations Focal genera: Cladophora, Chaetomorpha, Dictyosphaeria, Valonia 12 3 Order Cladophorales e.g. Cladophora Isomorphic Alternation of Generations: organism having a separate multicellular diploid sporophyte and haploid gametophyte stage, sporophyte & gametophyte are same sizes -Japanese Cladophora spp: “marimo balls” or tribbles • Size: up to 30 cm •A folktale in which the hearts of a young couple who drown in the lake turn into Cladophora balls • Meditation – watch balls rising and falling 13 California Cladophorales Cladophora - “Pin cushion” alga - Branched filamentous - Harbors inverts Nutrients 14 • Protected species - three day festival – to promote conservation Tropical Cladophorales Chaetomorpha Dictyosphaeria - Unbranched filaments - Modified basal cell w/rhizoidal extensions 15 Ventricaria - Opportunistic: “Ulva of the -Made up of single central cell (big) tropics” and lots of lateral cells (small) - Can outcompete corals when there has been overfishing or storm disturbance 16 4 Division: Chlorophyta Class: Ulvophyceae Trebouxiophyceae Order Caulerpales Chlorophyceae Prasinophyceae Focal genera: Order: Ulotrichales Ulvales Cladophorales Caulerpales Dasycladales - Coencocytic, siphonous thallus construction - Lots of variability in form (uniaxial, multiaxial, etc.) - Mostly biflagellate anisogamous reproduction; various life histories Codium “dead man’s fingers” Halimeda Caulerpa - Some calcified: major component of tropical sands remains of coralline greens form a layer >50 m thick at Great Barrier Reef - Synchronous spawning in tropics 17 18 Morphology Morphology Uniaxial and multiaxial construction Coenocytic/siphonous construction: = Multinucleate, no division of cytoplasm with cell walls “clotting compounds” to quickly repair wounds 19 Bryopsis Codium “ dead man’s fingers” 20 5 Order Caulerpales:e.g.Codi um Diplontic: 2N thallus, the gametes are the only haploid stage Codium fragile Morphology Utricles Codium fragile Gametangium Utricles-peripheral portions of the siphons maybe inflated & aggregated to form coherent outer surface gametangium (reproductive structure) originate from them *Codium the size of an apple = ~30 km of siphons 21 Codium and scallop interaction on East Coast 22 Codium invasion on the East Coast Alternative Stable State: Membranipora , invasive bryozoan, epiphyte on and causes mortality of native Laminaria, • Attach and grow on scallops • Weigh them down and prevent escape from predators Openings in Laminaria saccharina canopy favor establishment of… • Cast adrift in heavy storms • Results in high mortality 23 …Codium fragile invasive alga which suppresses recruitment of Laminaria sporophytes. Levin et al. 2002 24 6 Halimeda Holocarpic reproduction in many tropical members of O. Caulerpales - Thousands of individuals spawn synchronously - Water visibility drops to ~ 1m downstream from reef, but disperses within half an hour - Empty thallus disintegrates, often within hours - Anisogamous, dioecious= separate sexes - Entire thallus turns into gametes - Gametes migrate to surface overnight - In some species: gametes positively phototaxic 25 zygotes are negatively phototaxic 26 Tropical siphonous greens, timing of growth, and herbivory -Gamete release is synchronous and species and gender-specific - New segments not fully calcified, but heavily defended chemically - Major cue is amount of blue light (time from sunrise) - Chloroplasts stay in older segments until minutes before sunrise - Contrasts highly with other marine species that use lunar, tidal, temperature cues to spawn from Clifton, 1997 - New tissue put on at night (why?) 27 - Increase in calcification = decrease in chemical defenses 28 7 Synergy between calcification and secondary metabolites as defenses against herbivores Caulerpa invasion - Calcification: CaCO4 precipitated from water column, incorporated into cell wall of siphons, especially utricles (decalcify Halimeda, just a mass of enormous siphons) - Both chemical and structural defense common in tropical spp - Temporal interplay during growth and development - Old vs. young segments within an individual - Old vs. young individuals - Mark Hay’s work: true synergist effects in deterring urchins 29 Morphology • Native in Australia (but non-native strains have made it back there) • 1st introduced as a non-native in Mediterranean (1984) • In 2002 introduced at 2 sites in S.California (Carlsbad, Huntington Harbor) • Genetics reveal invasive strain different from native strain • Grows faster, larger, asexual reproduction, and resistant to cold temps • Listed as one of the top 100 invasive species on earth 30 Kleptoplastids Caulerpa = uniaxial, but with trabeculae for support trabeculae = ingrowth of wall material (Sacoglossan opistobranchs as petty theives) Siphons form rhizoidal holdfast, too 31 32 8 Division: Chlorophyta Class: Ulvophyceae - Sacoglossan radula adapted to host alga (length of tooth ~ thickness of siphon) - Common in siphonous greens - Pierce and suck out insides - Stolen chloroplasts can last up to a month or so – why not longer? Order: Trebouxiophyceae Chlorophyceae Prasinophyceae Ulotrichales Ulvales cladophorales Caulerpales Dasycladales - Extensive fossil record, group is 500 my old with 11 extant genera - Tropical, shallow water spp - Unicellular thalli with one nucleus and radial symmetry Celebrity genus: Acetabularia Elysia “mermaid’s wine glass” 33 34 1930’s - Hammerling’s experiments with Acetabularia 35 9