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KingdomKingdom ProtistaProtista Brown

Simple

J e KingdomKingdom ProtistaProtista DivisionDivision Phaeophyta,Phaeophyta, first brown land plants evolve •Recentplants, ~500 mya

• “Standard” cells

No cells Motile cells usual • Non-standard ThingsThings wewe willwill covercover forfor brownbrown algaealgae

•• GeneralGeneral featuresfeatures -- definingdefining characteristicscharacteristics

•• MorphologicalMorphological lineageslineages –– -- useuse morphologymorphology toto anticipateanticipate evolutionevolution -- revealreveal variationsvariations inin lifelife historyhistory && gametesgametes

•• EcologyEcology –– understandingunderstanding interactionsinteractions withwith environmentenvironment andand otherother speciesspecies

•• CommercialCommercial interestsinterests –– exploitexploit ecologyecology && lifelife historyhistory

•• EvolutionEvolution –– diversitydiversity && changechange overover timetime GeneralGeneral featuresfeatures OceanOcean loverslovers 2,0002,000 speciesspecies (60(60 inin HIHI oceans)oceans) Ocean Forests 250+250+ generagenera (29(29 inin HI)HI)

MorphologicalMorphological RangeRange::

filaments to complex multicell Intertidal RecentlyRecently arrivedarrived Subtidal eukaryoteseukaryotes -- Reef algae wellwell suitedsuited toto Natives and invasives coastalcoastal habitatshabitats Natives and invasives DiversityDiversity GeneralGeneral featuresfeatures

PigmentsPigments -- StorageStorage GrowthGrowth ProductsProducts

• Chlorophyll a • Chlorophyll c • Fucoxanthin

OthersOthers • Carotenoids • UV absorbing molecules PhotosynthesisPhotosynthesis

&& PigmentsPigments sunlight • Light energy is harvested by the cell • Only specific colors are absorbed

• Other colors are reflected back to your eye Chl a,a, cc Chl a,a, cc fucoxanthin Chlorophyll a & c

Tetrapyrrole Ring

Phytol Chain of Chl a missing in Chl c GeneralGeneral featuresfeatures

PigmentsPigments -- StorageStorage GrowthGrowth photosynthesis ProductsProducts • Every cell can divide

• Chlorophyll a • Laminarin • Multicellular • Chlorophyll c Starch (C) organisms: • Fucoxanthin • Mannitol Sugar (C) Fragments regrow • True tissues

Others • Carotenoids • UV absorbing molecules CellCell wallwall structurestructure Two essential parts: • fibers of cellulose (rigid), a glucose polymer • gels of polysaccharides (flexible) as ALGINATE CellCell wallwall structurestructure Two essential parts: • fibers of cellulose (rigid), a glucose polymer • gels of polysaccharides (flexible) as ALGINATE

“Copolymer” blocks of sugars Guluronic (G) Mannuronic (M) GeneralGeneral featuresfeatures WhatWhat isis inin aa typicaltypical phaeophytephaeophyte cell?cell? • Unusual membrane system around and nucleus • large, stalked and surrounded by laminarin starch • Chloroplasts have grana EvolutionEvolution ofof eukaryoteseukaryotes XenogenousXenogenous hypothesishypothesis

Simple cells

Autotrophs - green and red algae

Mereschowsky, 1905; 1910 GeneralGeneral featuresfeatures HowHow areare phaeophytephaeophyte cellscells unusual?unusual?

•“New” First step: membranes around chloroplast & nucleus Second step:

Second eukaryote RepeatedRepeated endosymbiosesendosymbioses LifeLife HistoryHistory asas aa “Strategy“Strategy”” ZygoticZygotic MeiosisMeiosis LifeLife HistoryHistory

Gamete DOESDOES NOTNOT OCCUROCCURZygote ININ BROWNBROWNGamete ALGAEALGAE

Meiosis

MeiosisMeiosis isis associatedassociated withwith ZygoteZygote GerminationGermination MorphologicalMorphological lineageslineages

•• EvaluateEvaluate adultadult formform toto gaingain insightinsight inin possiblepossible evolutionaryevolutionary processes.processes.

•• StepStep--byby--stepstep acquisitionacquisition ofof newnew traitstraits viavia geneticgenetic change.change.

•• ExamineExamine reproductivereproductive cellscells andand otherother characterscharacters asas additionaladditional data.data.

•• UsefulUseful meansmeans toto constructconstruct evolutionaryevolutionary hypotheseshypotheses toto testtest withwith molecularmolecular data.data. LifeLife HistoryHistory asas aa “Strategy”“Strategy”

MeiosisMeiosis isis associatedassociated withwith SporeSpore ProductionProduction GrowthGrowth && morphologymorphology Morphological Lineage #1 Ectocarpales Ectocarpus Genetic change All cells appear virtually identical - internally

Hincksia

Evolution has taken a simple shape to more complex but related forms:forms • Multi filamentous genera Ralfsia DiversityDiversity Order Ectocarpales

EctocarpusEctocarpus

Hincksia sp

RalfsiaRalfsia LifeLife historyhistory byby observationobservation

Ectocarpus

Same motile cell has 2 behaviors ? Where’sWhere’s meiosismeiosis ?? LifeLife HistoryHistory asas aa “Strategy”“Strategy” recognition zygote germling

isogametes Ectocarpus sp.

mitosis spores

germlings mitosis adults LifeLife HistoryHistory asas aa “Strategy”“Strategy”

MeiosisMeiosis isis associatedassociated withwith SporeSpore ProductionProduction ReproductionReproduction andand gametesgametes Equally sized gametes 2 Gametes – opposite strains always fuse Isogamy

Unequal gametes Anisogamy

Zygote – diploid cell via fusion of gametes & sperm gametes Oogamy egg MinimalMinimal diversitydiversity in gamete shape… Equal length flagella in gamete shape… Iso kont Swimming gametes have 2 flagella like greens • But flagella of swimming cells are unequal in length and HAIRY Unequal length flagella Hetero kont

Laterally inserted MinimalMinimal diversitydiversity inin gametegamete shape…shape…

These males gametes show the limited diversity among

Reds – spermatium

Greens – biflagellate to stephanokont males Minimal diversity in spore shape • Most brown algal spores swim •2 flagella • Settlement ? . sp. sp

Spatoglossum Hydroclathrus sp.

Rosenvingea sp.

Scytosiphon Orphans Orphans sp Diversity Tropical Diversity Tropical Colpomenia GrowthGrowth && morphologymorphology Morphological Lineage #2 Order Dictyotales

Dictyota

Evolution has taken a simpleDictyopteris filament: Multiseriate tissues create sheets or blades From apical cell or meristem areas

Padina DiversityDiversity Order Dictyotales

Dictyota sp. Padina sp. LifeLife HistoryHistory asas aa “Strategy”“Strategy” recognition meiosis

zygote tetra oogametes spores

Dictyota

spores mitosis

germlings mitosis adults LifeLife HistoryHistory asas aa “Strategy”“Strategy”

MeiosisMeiosis isis associatedassociated withwith SporeSpore ProductionProduction GrowthGrowth && morphologymorphology Morphological Lineage #3 Order Laminariales

Laminaria

Evolution has taken a simple shape: Alaria • constrained into • large sporophyte generations

Macrocystis DiversityDiversity Order Laminariales EcologyEcology

KelpKelp forestsforests sustainsustain coastalcoastal diversitydiversity epiphytic pelagic benthic understory LifeLife HistoryHistory asas aa “Strategy”“Strategy”

Macrocystis LifeLife HistoryHistory asas aa “Strategy”“Strategy”

Ecklonia cava

Spores Egg and sperm GrowthGrowth && morphologymorphology Morphological Lineage # 4 Order Fucales

Ectocarpus

Dictyopteris sp

Sargassum obtusifolium DiversityDiversity Sargassum thunbergii Gunma Japan Order Fucales

Sargassum cristaefolium Guam LifeLife HistoryHistory asas aa “Strategy”“Strategy”

MeiosisMeiosis isis associatedassociated withwith GameteGamete ProductionProduction LifeLife HistoryHistory asas aa “Strategy”“Strategy” juvenile recognition zygote

adult Sargassum sp.

oogametes

vegetative meiosis cell detail LifeLife HistoryHistory asas aa “Strategy”“Strategy” 11 freefree livingliving entityentity

CellsCells mustmust passpass throughthrough thisthis cyclecycle • to add new genetic individuals to population EcologyEcology ofof HawaiianHawaiian

ReefReef Algae.Algae. Frondose Algae of Wakiki,

B by M S. Doty 1969

i o 6000 m

a

s 5000

s

- 4000

w

e 3000 t

w 2000 t

,

g 1000 /

m

2 0 1 250 D 4 u 7 200 e ra 150 or tio 9 sh n 12 100 m o 50 fro f 13 m st 17 0 n, ud gio y re Dictyotales, , m ef o Re Fucales All browns were native representatives on reefs Most abundant biomass near shore