Midterm in one week (Tuesday, 4/27) Format: Definitions Short Answer Ps/I Curve Dichotomous key Life History

istory A couple of matching sections Short Essays

Kingdom: Unikonts Phylum: Mollusca Class:Gastropoda clade: Heterobranchia informal group: Opisthobranchia clade Cephalaspidea clade Thecosomata clade Gymnosomata clade Aplysiomorpha group Acochlidiacea clade Sacoglossa clade Umbraculida clade Nudipleura clade Nudibranchia

Dichotomous Key: Cannot use names of groups, instead characteristics Example: Chlamydomonas, Dictyota, Halimeda

Or (Ochrophyta, Chromophyta, Phaeophyta)

Division: Heterokontophyta

Has Chlorophyll C

Dictyota

No Chloropyll c

Forms palmelloid stage

No palmelloid stage

Chlamydomonas

Halimeda

~ 1,500 species 99% marine

Brief history of photosynthetic organisms on earth 3.45 bya = Cyanobacteria appear and introduce photosynthesis
thought to come from invagination of plasma membrane)

Heterokontophyta Characteristics:
1) Pigments?

1.5 bya = first Eukaryotes appeared (nuclear envelope and ER 0.9 bya = first multicellular algae (Rhodophyta - Red algae) 800 mya = earliest Chlorophyta (Green algae) 400-500 mya = plants on land derived from Charophyceae 250 mya = earliest Heterokontophyta (Brown algae) 100 mya = earliest seagrasses (angiosperms)

2) Chloroplast structure?

3) Storage product?

4) Flagella?
2

Bacteria Archaea

Spirochetes Chlamydias Cyanobacteria Proteobacteria Apicomplexans Dinoflagellates Cliates Heterokontophyta Diatoms Oomycetes Haptophytes Glaucophytes Rhodophyta Chlorophytes Land Plants Charophytes Diplomonads Parabasalids Heteroloboseans Euglenids Kinetoplastids Cercozoans Foraminiferans Radiolarians Fungi Choanoflagellates Animals Loboseans Slime molds Slime molds

Division: Heterokontophyta
Chromalvaeolates

Plantae

Excavates

Eukaryotes

Rhizaria

Many classes, mostly unicellular or colony of unicells: class: Chrysophyceae class: Synurophyceae class: Dictyochophyceae class: Pelagophyceae class: Raphidophyceae class: Xanthophyceae class: Eustigmatophyceae class: Tribophyceae class: Phaeothamniophyceae class: Phaeophyceae*** *** We will focus on marine macroalgae

Unikonts

Adapted from Sadava 2007

Algal taxonomy
Hierarchical system of classification: Level: Domain Kingdom/Clade Phylum/Division Class Order Family suffix: -phyta -phyceae -ales -aceae example: Eukaryote Chromalveolates Heterokontophyta Phaeophyceae Laminariales Alariaceae

Phaeophyceae Storage Products:

laminaran and mannitol are most abundant Always sugars, never starch Storage products found in cytoplasmic vacuoles (no special name); not associated with chloroplasts or pyrenoids

Genus species

Egregia menziesii

Phaeophyceae Metabolites:
Tannins: (a.k.a. polyphenolic compounds and terpenes) - Anti-endophyte, -epiphyte, and anti-herbivory - Stored in special vesicles called physodes in the cytoplasm

Seawater is hypotonic (less saline) than most algal cells

Phaeophyceae Storage Products :

Mannitol function important in osmoregulation- algae make more mannitol to use up ions-> algal salinity decreases transporting organic material to different parts of the thallus in large species lowers freezing point

Phaeophyceae Flagella:
Heterokont flagella: Anterior flimmer flagellum used for movement long flagellum with two rows of stiff hairs (mastigoneme) directed forward Posterior whiplash flagellum used for steering short, smooth flagellum directed backward contains flavin which functions as a photoreceptor an eyespot acts a shading structure or light reflector

Phaeophyceae Morphology of Cell Walls

Two main components:
1. Cellulose microfibrils (1-10% of thallus dry weight) Function: structural support

2. Alginic acid surrounds the microfibrills (35% of thallus dry weight) Function: elasticity; flexibility; prevent desiccation; and osmoregulation (ion exchange)

Flagella attached laterally not apically

Phaeophyceae Thallus Morphology:

- Advanced forms: complex multicellular thalli - Not unicellular (except gametes and spores) - Simplest forms are branching filaments - More complex forms are parenchymatous and pseudoparenchymatous - Differentiation of cortex (outer pigmented cells) and medulla (inner non-pigmented cells) - Medullary cells primarily for storage or transport over 40 meters long - Some browns quite large

Phaeophyceae Alginates:
Alginates = salt form of alginic acid; primarily in intercellular matrix Alginates = Alginic acid + an Ion Details of how alginates are used in helping ion exchange are not well understood. One proposed mechanism is that preferentially using some ions and not others helps to balance overall ion levels. Common ions that are used to make alginates are Ca, Na and Mg
6 8

Phaeophyceae Alginates:
Human uses for alginates
- Ice-cream - Frosting prevents ice crystal formation water retaining properties; prevent drying

Algal life histories : Terminology

Sporophyte: diploid, 2n, multicellular release spores

Sporangia = structure where spores are formed Spore (mitospore, meiospore): unicellular, must settle & grow, product of mitosis(Mt) or meiosis (Me)

- Paints emulsifying agent; keeps pigments suspended and prevents brush streaking - Pharmaceuticals - Food kombu in Japan; dried and shredded laminarians Gametophyte: hapliod, 1n, multicellular release gametes

Gametangia = structure where gametes are formed Gamete: unicellular, must fuse or die, product of mitosis(Mt) or meiosis (Me) 8

Phaeophyceae Morphology of intercellular connections

Only alga to transport sugar/photosynthate in sieve elements
Plasmodesmata = connections between adjacent cells, formed during cell division - Used for cell-to-cell transport of photosynthetic products and cell communication

Phaeophyceae Reproduction:

Gametangia = structure where gametes are formed Sporangia = structure where spores are formed

uniseriate Unilocular = all spores/gametes are produced in a single compartment unangia

multiseriate

Plurilocular = divided into many small chambers (locules); one spore/gamete per chamber plurangia

10

Phaeophyceae Reproduction:
Functional Anisogamy
How do the male gametes find the female ones?

Phaeophyceae habitat:
Mostly

marine (or at least brackish water)

Intertidal and subtidal Dominate colder waters Northern hemisphere:

Pheromones = chemicals produced to elicit a specific behavioral or physiological response from another individual. Many Phaeophyceans produce sexual pheromones, all are chemically similar (unsaturated hydrocarbons). Similarity high cross-reactivity Male gametes are VERY sensitive to pheromones: very low concentration will elicit a response

# of red species > # of brown species # of red individuals << # of brown individuals Browns are less diverse but more abundant
11 13

Phaeophyceae Reproduction:
Functional Anisogamy female gamete male gamete

12 orders within class: Phaeophyceae

Our focus will be on the following 8 orders:

Division: Heterokontophyta Class: Phaeophyceae

Durvillaeales

Order: Ectocarpales
Dictyotales

Chordariales

Laminariales

Fucales

Scytosiphonales

Desmarestiales

12

14

Distinguishing among orders based on

1. Life History and Reproduction

Order: Ectocarpales:
1. Life History and Reproduction:

Isomorphic/heteromorphic alt. of gen.; diplontic Isogamous, anisogamous, oogamous

2. Macrothallus Construction:
Filamentous Parenchymatous Pseudoparenchymatous

3. Growth

2. Macrothallus Construction:

Diffuse Apical Intercalary Trichothallic Meristodermal-a surface layer of cells (epidermis) that is capable of dividing (is meristematic) 15

3. Growth:

17

Division: Heterokontophyta Class: Phaeophyceae

Life History of Ectocarpales:

Order: Ectocarpales
Dictyotales

Durvillaeales Chordariales Laminariales Fucales

- Saxicolous or epiphytic - Uniseriate filaments

Scytosiphonales

Desmarestiales

Celebrity genus: Ectocarpus

- Opportunistic spp, excellent colonizers - Female gametes releases pheromone ectocarpene

16

17

Division: Heterokontophyta Class: Phaeophyceae

Life history of Dictyota

Reproductive structures in sori = cluster of gametangia or sporangia
Durvillaeales

Order: Ectocarpales

Dictyotales

Chordariales

Laminariales

Fucales

Tetraspores (non-flagellated) released by 2N sporophyte Gametophyte dioecious Large egg one per oogonium = female reproductive structure containing one or more eggs Sperm single hairy flagella but has second basal body 19 21

Scytosiphonales

Desmarestiales

Celebrity genera: Dictyota, Padina

- Saxicolous - Common in tropical waters; also found locally subtidal - Pheromone= dictyotene

Order: Dictyotales
1. Life History and Reproduction:

Life History of Dictyota:

2. Macrothallus Construction:

3. Growth:

20

17

A Dictyota Story:
(Stachowicz and Hay 2000)

Division: Heterokontophyta
Cape Hatteras

Local adaptation Associational defences Southern Sites + Libinia dubia, the decorator crab + Dictyota menstrualis, the chemically defended brown alga (diterpenes) + Omnivorous fishes + Crabs are specialists in decoration preference

Class: Phaeophyceae

Northern Sites + Libinia dubia, the decorator crab + No chemically noxious algae + Carnivorous fishes + Crabs are generalists in decoration preference

Order: Ectocarpales
Dictyotales

Durvillaeales Chordariales Laminariales Fucales

Scytosiphonales

Desmarestiales

Celebrity genera: Scytosiphon, Colpomenia - Saxicolous or epiphytic - Found locally in higher intertidal shallow pools - Plurangia always uniseriate (filament in single row)

21

23

The only calcified Phaeophycean:

Padina

Order Scytosiphonales:
1. Life History and Reproduction:

Increased Grazing

2. Macrothallus Construction:

3. Growth:

22

24

Division: Heterokontophyta Class: Phaeophyceae

Order: Chordariales
1. Life History and Reproduction:

Order: Ectocarpales
Dictyotales

Durvillaeales

Chordariales

Laminariales

Fucales

Scytosiphonales

Desmarestiales

2. Macrothallus Construction:

Celebrity genera: Leathesia, Ralfsia

- Saxicolous or epiphytic - Common intertidally, tropics to poles - Plurangia could be uniseriate or multiseriate

3. Growth:

26

27

Order: Scytosiphonales .life history not fully understood

Convergent evolution of form:

Gametophyte? Plurangia

Sporophyte? Unangia

Upright stage

Crust stage (Ralfsia-like)

Colpomenia = Parenchymatous; O. Scytosiphonales

25

Leathesia = Psuedoparenchymatous; O. Chordariales

28

Both genera found locally..do the crunch test!!

10

Division: Heterokontophyta Class: Phaeophyceae

Life History of Desmarestiales

Order: Ectocarpales
Dictyotales

Durvillaeales Chordariales Laminariales Fucales

Scytosiphonales

Desmarestiales

- Saxicolous - Low intertidal to subtidal - Acid weed Celebrity genus: Desmarestia 29 31

- Morphology variable within the genus

Order: Desmarestiales
1. Life History and Reproduction:

-Subtidal, but grows well in high light (very abundant if disturbance has removed other plants or canopy, e.g. El Nino, or urchin grazing) - Disappears with kelp growth due to canopy cover; shading - Acid weed: cells accumulate sulfate ions from seawater which reacts with water to produce sulfuric acid or malic acid, stored in vacuoles

2. Macrothallus Construction:

=ANTIHERBIVORY, pH 0.8-1.8, bleaches other algae

3. Growth
30 32

11

Class Phaeophyceae
Order Macrothallus Construction Macrothallus Growth
Isomorphic alternation of generations Ectocarpales Dictyotales filamentous parenchymatous diffuse apical or marginal

Heteromorphic alternation of generations Chordariales filamentous or parenchyma Desmarestiales pseudoparenchyma

variable tricothallic variable intercalary or meristodermal

Scytosiphonales parenchyma or pseudoparenchyma Laminariales parenchyma

Diplontic: 2N thallus, the gametes are the only haploid stage Fucales Durvilleales parenchyma parenchyma apical variable

12