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Marine – the study of aquatic and algae that live in seawater convenience term encompassing various distinctly related of the open and the littoral zone and in brackish groups of aquatic photsynthetic eukaryoes waters of estuaries

Phycology- study of algae Macroalgae - Rhodophyta, , Heterokontophyt a

alga (singular) : “I study , the intertidal alga” Microalgae () - Bacillariophyta, Dinophyta , Haptophyta algae (plural): “Algae rock my world” algal (adj.): Algal lunch, algal skirt, algal growth rate Angiosperms (wrong!) -Mangroves, Marsh Plants, “ ”

Cyanobacteria 1 2

I. What are algae? I. What are algae?

• Polyphyletic group = different ancestors, different evolutionary histories

A B C D E A B C D E A B C D E A B C D E

monophyletic polyphyletic paraphyletic or 3 4

1 Groups Groups () 1.- 2.Archae - 3. 1. Alveolates- unicellular,plasma membrane supported by - , heterokonyophyta flattened vesicles 2. Stramenopiles- two unequal flagella, 4 Excavates membranes 3. Rhizaria- unicellular amoeboids Plantae- rhodophyta, chlorophyta, seagrasses Amoebozoans 4. Excavates- unicellular Fungi 5. Plantae- most broadly defined group 6. Amoebozoans- pseudopods for movement & eating

7. Fungi- with extracellular digestion

8. Choanoflagellates- unicellular withsingle flagella

5 9. Animals- multicellular heterotrophs 6

DOMAIN Groups (Kingdom) 1.Bacteria- cyanobacteria (blue ) Defining characteristics of Algae: 2.Archae “Algae” (photoautotrophic, usually), using Chl a as 3.Eukaryotes 1. Alveolates- dinoflagellates primary pigment

2. Stramenopiles- diatoms, heterokonyophyta BUT: Limited compared to 3. Rhizaria- unicellular amoeboids terrestrial plants 4. Excavates- unicellular flagellates  No “real” vascular system

5. Plantae- rhodophyta, chlorophyta, seagrasses  organs unicellular, or all cells capable of (no sterile layer of cells 6. Amoebozoans- slimemolds surrounding sex organs aka NO ) 7. Fungi- heterotrophs with extracellular digestion  Much greater diversity of photosynthetic pigments

8. Choanoflagellates- unicellular and histories

9. Animals- multicellular heterotrophs 7 8

2 Algae show tremendous diversity of A An Alga form, , and lifestyle

sorus

blade stem photo Pete Dal Ferro

stipe leaf holdfast roots - plants that lack roots, stems & leaves 9 10

Free-living and unattached

Red = dinoflagellates like Lingulodinium polyedrum (“planktonic”)

Unattached macroalgae- (“benthic” = bottom dwelling)

photo: M organ Bond Found in all bodies of water (freshwater, marine intertidal and subtidal) as well as terrestrial systems with enough moisture11 12

3 Free-living and attached to the substrate Parasitic Epiphytic

Postelsia palmaeformis “saxicolous”, or “saxiphytic”

Smithora naiadum on Phyllospadix torreyi

Caulerpa taxifolia “psammophtyic”

Trentepohlia on 13 Monterey Cypress 14

Symbiotic and Endoymbiotic Ecological importance of algae = close association of an alga and a - ; role in interactions

- engineers: e.g. forests, beds, coral Marine: Zooxanthellae in corals, anemonies, reefs = Create structure that defines the habitat type nudibranchs,

Radiolarians, Foramaniferans = ameoba + alga

fH2O too!: Zoochlorellae in hydras, , etc.15 16

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- input into terrestrial systems (“Allochthonous input”); just Direct importance of algae to beings beginning to appreciate this

Origin’s of the world’s oil supplies (dinoflagellates,,diatoms)

Used in biological and medical research (e.g. Cyanobacteria, ; fucoids);

One product of (e.g. , Gigartina) = ; produces gel at low temperatures, used in gel electrophoresis. (HUGE in )

Eaten “as itself” (e.g. , Spirulina)

Products of algae are everywhere: (from red algae) and alginates (from , e.g. , ) from in walls, act as thickening agents

Ice cream, mayonnaise, chocolate milk, soy milk, toothpaste, salad dressings, shaving cream, fertilizers, rubber, paint, hair products 17 18

II. Algal II. Algal taxonomy

Hierarchical system of classification: Hierarchical system of classification:

Level: suffix: Level: suffix: example: Domain Domain Eukaryote Kingdom/Group Kingdom/Group Plantae /Division -phyta Phylum/Division -phyta Chlorophyta -phyceae Class -phyceae -ales Order -ales Family -aceae Family -aceae Genus Ulva species species fenestrata

• King Phillip Came Over For Good Spaghetti

• Keep Dishes Clean Or Family Gets Sick 19 20

5 II. Algal taxonomy - Nomenclature acknowledges the first and last Hierarchical system of classification: person to describe the species

Level: suffix: example: Domain Eukaryote for example: Linnaeus called this pyriferus; Kingdom/Group Stramenopiles later renamed by Carl Adardh, Phylum/Division -phyta Heterokontophyta so: Class -phyceae Phaeophyceae Order -ales Laminariales Macrocystis pyrifera (Linnaeus) Adardh Family -aceae Genus species menziesii

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1753, Linneaus divided all life into two Phyla = Division % marine ~# species Plants and Animals Cyanophyta (blue-green algae) 8 2,000 Rhodophyta (red algae) 98 6,000 Within the plants, he recognized Chlorophyta (green algae) 13 16,000 • – hidden …land plants Heterokontophyta (brown algae) 99 1,500 • Thallogams – unspecialized gametes … the algae Bacillariophyta (diatoms) 50 10,000 Dinophyta (dinoflagellates) 90 2,000

Only three genera originally recognized: Bryophyta Fucus-fleshy , liverworts 0 25,000 Ulva- membranous Conferva- filamentous Vascular plants , horsetail, club 0.1 13,018 0 722 Angiosperms 0.09 285,000

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6 -Taxonomy/ constantly under revision Division % marine ~# species # in Ca Cyanophyta (blue-green algae) 8 2,000 - Depending on who you ask, between 50,000 and Rhodophyta (red algae) 98 6,000 459 Chlorophyta (green algae) 13 16,000 72 10 million different algal spp! Heterokontophyta (brown algae) 99 1,500 137 Bacillariophyta (diatoms) 50 10,000 Dinophyta (dinoflagellates) 90 2,000 - Biological species concept? Bryophyta Mosses, liverworts 0 25,000 -Morphology? Vascular plants Ferns, horsetail, club moss 0.1 13,018

Gymnosperms 0 722 - Genetics? Angiosperms 0.09 285,000

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Mastocarpus papillatus

Petrocelis papillatus + 2N 2N ‘Petrocelis’ crust ()

1N

27 1N fronds 28 ()

7 III. Algal Endosymbiotic theory of acquisition: (L. Margolis)

Brief history of photosynthetic on - Heterotrophic eukaryote eats 3.45 bya = Cyanobacteria appear and introduce heterotrophic bacteria lead to the photosynthesis formation of mitochondria

1.5 bya = first Eukaryotes appeared ( and ER thought to come from of plasma membrane) -Heterotrophic eukaryote eats a photosynthetic bacteria

(cyanobacteria) lead to the formation 0.9 bya = first multicellular algae (Rhodophyta - Red algae) of a 800 mya = earliest Chlorophyta (Green algae) -Bacteria not digested but becomes

400-500 mya = plants on land – derived from an organelle

250 mya = earliest Heterokontophyta (Brown algae)

Support of Endosymbiotic Theory 100 mya = earliest seagrasses (angiosperms) -Genetic material of the inner membrane 29 30

Primary Endosymbiosis: Secondary endosymbiotic events

1. Heterotrophic eukaryote eats photosynthetic bacteria 1. Heterotrophic eukaryote eats (cyanobacterium). photosynthetic eukaryote

2. Results in photosynthetic eukaryote. 2. Nucleus from photosynthetic Chloroplast has 2 membranes eukaryote is lost

3. Chloroplast ends up with 4 membranes

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8 Secondary endosymbiotic events Secondary Endosymbiosis:

1. Heterotrophic eukaryote eats 1. Heterotrophic eukaryote eats photosynthetic eukaryote photosynthetic eukaryote

2. Nucleus from photosynthetic 2. Nucleus from photosynthetic eukaryote is lost eukaryote is lost

3. Chloroplast ends up with 4 3. Results in photosynthetic eukaryote. membranes Chloroplast has 4 membranes.

Tertiary endosymbiotic events in some groups

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Details of Endosymbiotic origins - Loss of What is agreed upon: e.g. Parasitic algae on : no pigments, all white • Each algal division is a ? Plocamiocolax = Parasite on Rhodophyte alga monophyletic group • Reds and Greens – 1 event-2 membranes • Browns – 2 events- 4 membranes

Plocamiocolax on Plocamium Plocamium Adapted From Palmer 2003 35 36

9 Three main divisions (phyla) of seaweeds: - Loss of plastids e.g. Heterotrophic algae Chlorophyta: = parasite in mammal muscular tissues 1 endosymbiotic event = 2 membranes - Apicomplexan, closely related to dinoflagellates ~800 mya ~16,000 species; 1,300 are marine (most are fH2O)

Heterokontophyta: 2 endosymbiotic events = 4 plastid membranes ~250 mya ~1,500 species; most are marine

Rhodophyta: 1 endosymbiotic event = 2 plastid membranes ~0.9 bya ~60% of domestic cats are infected; ~6,000 species; 5,800 marine toxoplasmosis in pregnant women… caused by an alga! 37 38

Paper Discussion on wednesday:

Lubchenco and Cubit. 1980. Heteromorphic life histories of certain marine algae as to variations in herbivory. 61(3): 676-687

Abstract Introduction Graphs & Figures Methods Results Discussion

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