The Protists Chapter 21 Protists • General characteristics – Unicellular, colonial, simple multicellular organisms – Eukaryotic – Some exhibit both plant and animal characteristics • Euglenoids • Slime molds Protists – Many are photosynthetic • All have chlorophyll a • Often called algae Protists • Evolutionary relationships of eukaryotes – Difficult problem • Splits between many lineages of eukaryotes are ancient • Have been events in which organisms (or parts of organisms) that are not closely related have joined to form new organisms • Not many characters are shared by all members of group • Shared characters often yield different results when analyzed cladistically Protists – At base of eukaryotic tree • Group of protists – Some lack mitochondria and live as parasites on other organisms – Rest of eukaryotes divided into two major clades • One clade – Animals, fungi, slime molds, and small group of amoeboid organisms – None are photosynthetic – Most are motile (except fungi) Protists • Other clade – Variety of protist groups – Includes both photosynthetic and nonphotosynthetic protists Photosynthetic Protists • Commonly called algae • Variety of life histories, body forms, ecological roles • Often named for distinctive colors • Unicellular, colonial, filamentous, sheetlike Amoebozoa • Clade of protists that includes amoebas and slime molds • Traits combine aspects of fungi and animals – Animal characteristics • Lack cell walls, engulf food, have motile cells at some phase of life cycle – Fungi and plant characteristics • Form sporangia and nonmotile cells with cell walls Amoebozoa – Two groups of slime molds Myxomycota • Myxomycota • Acrasiomycota Plasmodial slime molds Cellular slime molds Contain thousands of nuclei with no membranes separating them Smaller, have fewer nuclei, and do have membranes between the nuclei Acrasiomycota Alveolates • Common character – System of alveoli • Tiny membrane-enclosed sacs found beneath plasma membrane • Clade composed of four ecologically and economically significant lineages – Ciliates • • • • Found in freshwater Have cilia and engulf food Often called protozoa Example: Paramecium Alveolates – Foraminifera • Characterized by hard shell • Feeding strategies include active predation, scavenging, using sticky webs to trap food • Shells – Common fossils – Important in dating geological strata and in oil exploration Alveolates – Apicomplexa • Parasitic or pathogenic protists • Found to contain vestigial plastids • Examples – Plasmodium (causes malaria) – Toxoplasma (causes toxoplasmosis) Alveolates – Photosynthetic dinoflagellates • Most are unicellular, motile, and marine • Usually have two flagella (both emerge from same pore) – One is flat and ribbonlike, encircles cell in groove around middle, provides rotational movement – Other flagellum trails behind and provides forward movement • Contain pigments – Chlorophylls a and c – Brown pigment (fucoxanthin) Alveolates • Chloroplasts surrounded by three or four membranes – May also contain a remnant nucleus • Some are bioluminescent • Overgrowth (bloom) can cause red tide Euglenoids • Typically single-celled • Usually in freshwater but could be in salt or brackish water or soil • A few are parasitic • Lack cell wall – Flexible strips of proteins and microtubules under cell membrane • Have two flagella Euglenoids • About one third of the 1,000 species have chloroplasts – Contain chlorophylls a and b, carotenoids – Three membranes around chloroplasts • May have eyespot – Red or orange light-sensitive organelle – Pigment (astaxanthin) is a carotenoid has antioxidant properties • Extracted and sold as health supplement Euglenoids • Never been observed to reproduce sexually • Important in food chains of freshwater ecosystems • Ecological indicators of water that is rich in organic matter Heterokonts • Clade also sometimes called Stramenopiles or Chromista • All have two unequally sized flagella • Heterokonts and alveolates share common ancestor Heterokonts • Includes – Oomycota (water molds and downy mildew) – Xanthophyta (golden algae) – Chrysophyta (golden algae) – Diatoms – Brown algae Heterokonts Group Brown algae Diatoms Xanthophyta Chrysophyta Taxonomic name Phaeophyta Bacillariophyta Xanthophyta Chrysophyta Number of species Key characteristics 1,500 Two unequal lateral flagella; cell wall of algin + cellulose; chlorophylls a and c; filamentous to complex large kelps; mainly in shallow, cool, marine water 8,000 Usually no flagella; cell wall of silica + pectin; chlorophylls a and c; carbohydrates stored as oil; mainly unicellular and free-floating; prominent as freshwater or saltwater phytoplankton 400 Two flagella (various); cellulose cell wall; chlorophyll a (+ c in some), oil stored; mainly unicellular; mainly in freshwater 300 Two unequal anterior flagella; cellulose cell wall (+ silica in some), chlorophylls a and c; mainly unicellular and in freshwater Heterokonts • Oomycota – Includes egg fungi, downy mildews, and water molds – Fungal characteristics • Hyphae, produce spores, lack chlorophyll – Algal characteristics • Cellulose cell walls, swimming spores, some cellular details, some metabolic pathways Heterokonts • Oomycota – Most are decomposers – Some are pathogens of important crops • Downy mildew of grapes – Plasmopora viticola – Nearly destroyed French vineyards in nineteenth century • Potato blight – Phytophthora infestans – Changed history of Ireland in 1840s Heterokonts • Diatoms – Important members of phytoplankton – Cell wall made out of silica • Two parts (valves) to cell wall • Fit together like halves of Petri dish – Shape of cell varies – Pigments • Chlorophylls a and c • Fucoxanthin Heterokonts • Diatoms – Store food reserves as oil – Some exhibit gliding motion – Stalked diatoms grow as epiphytes on seaweeds and kelps – Can also become attached to nonliving surfaces – Create algal turfs • Coat shallow rocks in quiet freshwater or marine habitats • Have as high a daily productivity per square meter as tropical rain forest Heterokonts • Diatoms – Dominate surfaces of salt mudflats – Extensive fossil record – Indicators for petroleum exploration – Silica shells form extensive deposits Heterokonts • Brown algae – Almost exclusively marine – More abundant in cool, shallow waters – More complex brown algae kelps • • • • Chlorophylls a and c, fucoxanthin No grana in chloroplasts Store carbohydrates as mannitol or laminaran Cell walls composed of cellulose and alginates Heterokonts • Brown algae – Kelps • Example – – – – Macrocystis Largest known kelp Fast growth rate Consists of » Holdfast – anchors » Stipes – stem-like structure » Blades – leaf-like » Gas-filled air bladder - buoyancy Heterokonts • Brown algae – Kelps • Outer layer is protective and consists of cells that are also meristematic and contain chloroplasts called meristoderm • Region of cortex beneath meristoderm – Composed of parenchyma-like cells – Mucilage-secreting cells line canals through medulla • Medulla – Innermost part of stipe – Loosely packed filaments of cells Heterokonts • Brown algae – Kelps • Cells that function as sieve elements – Found in transition zone between cortex and medulla – Have sieve plates, form callose, adjoin one another to make continuous tubes – Mannitol moves through tubes • No tissue that resembles xylem The Plants • Molecular data supports idea that red algae, green algae, and land plants belong in same clade • Green algae – Not a natural monophyletic group – Gave rise to land plants The Plants • Red algae – Almost exclusively marine – Most abundant in warm water – Can grow to considerable depth – More complex forms called seaweeds • • • • Parenchyma-like tissue Holdfast for anchoring Stipes never very long Blades never have gas bladders The Plants • Red algae • Pigments – Chlorophyll a and phycobilins • Cell wall – Cellulose and sometimes agar or carrageenan • Food storage molecule – Floridean starch Comparison of Red and Brown Algae Brown algae Red algae Evolutionary Heterokonts group Plants Common name Kelp Seaweed Habitat Marine; cool, shallow water Marine; warm water; greater depths Pigments Chlorophylls a and c, fucoxanthin Chlorophyll a, phycobilins Food storage Mannitol or laminaran Floridean starch Cell wall Cellulose and alginates Cellulose; sometimes also contains agar or carrageenan The Plants • Green algae – Mainly in freshwater habitats but could be in saltwater, on snow, in hot springs, on soil, on leaves and branches of terrestrial plants – Shared characteristics • • • • • Chlorophylls a and b, carotenoid accessory pigments Food stored as starch Cellulose cell walls Formation of phragmoplast during mitosis Asymmetrically attached flagella The Plants • Green algae – Groups • Chlorophyceae • Ulvophyceae • Charophytes The Plants • Green algae – Chlorophyceae • • • • Two flagella at anterior end Single, large, cup-shaped chloroplast Most have red-colored carotene eyespot Examples – Gonium – Volvox The Plants • Green algae – Ulvophyceae • • • • Sea lettuces Typically small, green seaweeds Consumed as food in many places Example – Caulerpa » Accidentally spread to areas with no natural limits to growth » Multiplied explosively » Produces toxins that are lethal to urchins and some fish » Has been discovered off coast of California The Plants • Green algae – Charophytes • Group of ancient green algae • Examples – Coleochaete » Once thought to be closest living relative to land plants – Chara » According to molecular characters → more closely related to land plants than Coleochaete Ecological and Economic Importance of Algae • Phytoplankton – Base of aquatic food chains – “grasses of the sea” – Unicellular – Produce about four times the amount of photosynthate that is produced by the Earth’s croplands each year – Bloom • Algal overgrowth Ecological and Economic Importance of Algae • Help build tropical reefs – Coralline algae • Certain red and green algae • Create carbonate exoskeleton that becomes part of reef when alga die Ecological and Economic Importance of Algae – Coralline algae • Some algae grow symbiotically with coralline animals – Mutualistic relationship » Algae produces sugar and oxygen for animal » Cells of coral contribute CO2, nitrogen, and minerals for alga – Usually dinoflagellate Symbiodinium microadriaticum – Has photosynthetic rate 10 times greater than phytoplankton » Protected and nourished by animal cytoplasm around it Ecological and Economic Importance of Algae • Medicine, food, and fertilizer – Laminaria • Harvested off coast of China as source of iodine – Porphyra (nori) • Cultivated • Supplement to Japanese diet – limu • used as food source by Polynesians in Hawaii Ecological and Economic Importance of Algae • Medicine, food, and fertilizer – Palmaria palmate • red seaweed, dulce • Used as food in British isles – Chondrus crispus • Irish moss, red algae • Used to make jelly desert called blancmange Ecological and Economic Importance of Algae • Medicine, food, and fertilizer – Good fertilizer or cattle feed supplements • • • • Compares favorably with manure as a fertilizer Enhances germination Increases uptake of nutrients Seems to give degree of resistance to frost, pathogens, and insects Ecological and Economic Importance of Algae • Uses of algal cell walls – Diatomite (diatomaceous earth) • Rich deposit near Lompoc, California • Uses of diatomite – Superior filter or clarifying material – Added to many materials to provide bulk, improve flow, increase stability » Dental impressions, grouting, paint, asphalt, pesticides – Abrasive Ecological and Economic Importance of Algae • Uses of algal cell walls – Agar • Primarily obtained from red algae, Gelidium and Gracilaria • Used as a culture medium • Substance purified from agar (agarose) used for gel electrophoresis • Used in baking industry – Added to icing to retard drying in open air or melting in cellophane packages • Used as a bulk laxative Ecological and Economic Importance of Algae • Uses of algal cell walls – Carrageenan • Reacts with proteins in milk to make stable, creamy, thick solution or gel – Used commercially in ice cream, whipped cream, fruit syrups, chocolate milk, custard, evaporated milk, bread, macaroni • Added to dietetic, low-calorie foods • Used in toothpaste, pharmaceutical jellies, and lotions • Mainly comes from Irish moss, red algae Ecological and Economic Importance of Algae • Uses of algal cell walls – Algin • Compound of brown algae • Strongly absorbs water • Used as an additive to beer, water-based paints, textile sizing, ceramic glaze, syrup, toothpaste, hand lotion Ecological and Economic Importance of Algae – Algin • Commercially harvested species – Macrocystis pyrifera – along California coast – Ascophyllum, Fucus, and Laminaria – off Maritime Canada, northeastern United States, England, China coast – Durvillea – from Australian waters Algal Reproduction • Asexual reproduction occurs more often than sexual reproduction • Asexual methods of reproduction – Cell division (single-celled algae) – Fragmentation (filamentous algae) – Formation, liberation, germination of motile or nonmotile spores produced in sporangia Algal Reproduction • Three basic life cycles – Zygotic • Only diploid phase of life cycle is single-celled zygote – Gametic • Only haploid phase of life cycle is single-celled gamete – Sporic • Multicellular gametophytes and sporophytes Algal Reproduction • Zygotic life cycle – Example: Ulothrix – Sexual reproduction • Some of nuclei divide by mitotic divisions to produce many motile gametes inside wall of parent cell • Parent cell is gametangium • Ulothrix gamete approaches another suitable gamete • Cells fuse forming diploid zygote cell with four flagella Algal Reproduction • Zygotic life cycle • Zygote become spherical, loses flagella, enters resting stage • When conditions are right, zygote becomes metabolically active • Zygote divides by meiosis and produces + and – meiospores • Meiospores are dispersed • Each meiospore can germinate, divide by mitosis, produce a + or – haploid plant Algal Reproduction • Zygotic life cycle – Asexual reproduction • Vegetative cell becomes sporangium • 16 to 64 pear-shaped mitospores are released • After period of activity, motile mitospores settle to bottom of pond, lose flagella, produce new plant by mitosis Algal Reproduction • Gametic life cycle – Example: diatoms – Sexual reproduction • Diploid nucleus undergoes meiosis • Produces four haploid nuclei (only 1 or 2 survive to become gametes) • Gametangia near each other open, gametes emerge, fuse, form diploid zygote • Zygote increases in size • Secretes silica wall around itself, becomes vegetative diploid diatom Algal Reproduction • Gametic life cycle – Asexual reproduction • Reproduce by cell division • New cell wall forms within old one • Progeny cell that inherits small wall segment of cell wall makes wall that is even smaller • Pattern continues until critically small size is reached • Cell division stops • Cell then must reproduce sexually Algal Reproduction • Sporic life cycle with isomorphic generations – Identical looking gametophytes and sporophytes – Example: Ectocarpus • Haploid phase has gametangia on side branches • Mitosis within gametangium produces gametes • Released isogamous gametes (look alike) represent + and – mating types Algal Reproduction • Sporic life cycle with isomorphic generations • Plus and minus gametes fuse in open water → yields diploid zygote • Zygote settles to bottom, germinates, divides by mitosis, produces diploid organism • Diploid sporophyte looks identical to haploid gametophyte Algal Reproduction • Sporic life cycle with isomorphic generations • Sporophyte produces two kinds of reproductive cells – Asexual sporangia develop on side branches » Release motile cells (diploid mitospore) capable of producing new individual (sporophyte) by itself – Spherical sporangium » Produces meiospores » Meiospores germinate, producing gametophytes Algal Reproduction • Sporic life cycle with heteromorphic generations – Gametophyte and sporophyte are not identical – Most highly developed in brown algae – Example: Laminaria • Sporophyte generation with well-developed holdfast and long unbranched stipe with narrow blades Algal Reproduction • Sporic life cycle with heteromorphic generations • Sporangia form in groups just below meristoderm on blade • Single-celled sporangium undergoes meiosis • Produces 8 to 64 meiospores • Meiospores are released, swim, settle to bottom, produce gametophytes • Some gametophytes produce female gametes, some produce male gametes Algal Reproduction • Sporic life cycle with heteromorphic generations • Cells at tips of some male gametophyte filaments enlarge and function as antheridia – Nuclei undergo mitosis producing motile sperm • Cells at tips of some female gametophyte filaments enlarge and function as oogonia – Nuclei undergo mitosis producing one to several large eggs – Eggs are extruded but remain attached Algal Reproduction • Sporic life cycle with heteromorphic generations • Sperm cell fuses with egg • Produces zygote which forms sporophyte • Sporophyte separates from gametophyte, carried by currents to bottom, begins to develop into mature Laminaria sporophyte