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The Protists and the Origins of Eukaryotes Chapter 20 Characteristics of Protists • Eukaryotes • Primarily unicellular (some colonial and multicellular exist) • Metabolically diverse • Structurally complex • Asexual reproduction usual; sexual reproduction diverse • Basically, catch-all kingdom!! • Only real characteristics in common are: – Eukaryotes – Prefer watery environments Protist Classification • Phylogeny not well-established • Protists do not represent a monophyletic group • Some are more closely related to animals than to other protists • For convenience, protists grouped by: – Mode of nutrition (plant-like, animal-like, fungal-like) – Movement • Protists are essentially the earliest eukaryotes and 1st steps towards true multicellular organisms The Importance of Protists • Origins of eukaryotic cell • Origins of multicellularity • Origins of sexual reproduction • All of these advances are represented in various protists Figure 27.1 Major Eukaryote Groups in an Evolutionary Context (Part 1) Figure 27.1 Major Eukaryote Groups in an Evolutionary Context (Part 2) Origins of Eukaryotic Cell • Modern eukaryotic cell arose in several steps – – – – – • Flexible cell surface Cytoskeleton Nuclear envelope Digestive vesicles Endosymbiotic acquisition of certain Cytoskeleton – Extends from PM to nucleus – Probably helped form nucleus – Phagocytosis --> formation of organelles – Locomotion – Increase in cell size – Mitosis: allows for eventual meiotic division needed for sexual reproduction – Flexible cell surface: can fold inward to increase surface area • Probably lead to endocytosis methods and vesicle formation Evidence: Giardia • Have nucleus • Lack mitochondria • No other membraneenclosed organelles • Has cytoskeleton Endosymbiotic Hypothesis of Mitochondria and Chloroplasts • Eukaryotes display presence of bacterial genes encoding for energy metabolism • Aerobic bacteria gave rise to mitochondrion • Cyanobacterium gave rise to chloroplasts of green algae, plants, and red algae History of Endosymbiosis • Mitochondria derived from proteobacterium capable of aerobic metabolism • Chloroplasts appear in several distantly related protist clades – Photosynthetic pigments differ – Not all chloroplasts have a pair of membranes • Some have three • Primary endosymbiosis – All chloroplasts trace their ancestry back to engulfment of a cyanobacterium • Chlorophyll a present in all!! • One membrane from cyanobacterium, second from host – Gave rise to chloroplasts of green and red algae • Secondary and tertiary endosymbiosis – All other photosynthetic protist lineages Eukaryotes Acquired Features From Both Archaea and Bacteria Why aren’t the three rRNA genes of corn one another’s closest relatives? How would you explain the closer relationship of the mito rRNA gene of corn to the rRNA gene of E.coli than to the nuclear rRNA genes of other eukaryotes? Can you explain the relationship of the rRNA gene from the chloroplast of corn to the rRNA gene of the cyanobacterium? If you were to sequence the rRNA genes from human and yeast mito genomes, where would you expect these two sequences to fit on the gene tree? Origins of Multicellularity • Major multicellular lineages in eukaryotes: brown algae, plants, animals, fungi, red algae • Some protists form colonies or are primitive multicellular – Examples: Volvox, colonial; some algae, multicellular – True multicellular organisms - there is a division of labor among different kinds of cells and non are independent • Multicellularity – Increase in size advantageous • Less predation • Lower metabolic rate --> less food requirement Origins of Multicellularity • What is required for multicelluarity? – Cohesion • Animals = extra cellular matrix and collagen • Plants = plasmodesmata and cell walls – Communication between cells to allow for cooperation • Nervous system and endocrine system in animals • Signal transduction and hormones in plants – Developmental plan • Regulation of gene expression to guide specialization of cells – Cells must specialize!! • See distinctive organelles become more or less prevalent in different cell types – Can’t just increase size b/c SA/V ratio Origins of Multicellularity • Cells must specialize!! – See distinctive organelles become more or less prevalent in different cell types – Can’t just increase size b/c SA/V ratio • • • • Flexible membrane surface Change shape Activity near surface Vacuoles for food storage/waster removal – Food vacuoles – Contractile vacuoles (help eliminate excess water) • Multicellularity » Eventually increase cell number Origins of Sexual Reproduction • Sexual reproduction allows for adaptive radiation and lost of new species – Genetic recombination and exchange • Protists exhibit both asexual and sexual reproduction – Asexual: binary fission, budding, spores – Sexual: • Conjugation • Haploid life cycle, alternation of generation life cycle – Gametes haploid – Both diploid and haploid cells undergo mitosis • Diploid life cycle – Gametes haploid – Only diploid cells undergo mitosis • Eukaryotic reproduction requires existence of mitosis and meiosis!!!! – Both processes dependent on linear chromosomes and spindle fibers (microtubules of cytoskeleton) Figure 27.13 Alternation of Generations Life Cycles Plants • Isogamy vs oogamy – Gametes look alike; gametes from each gender distinct • Sporophyte vs gametophyte – Gives rise to spores; gives rise to gametes • Isomorphic vs heteromorphic – Haploid and diploid individuals distinct; haploid and diploid individuals appear alike Animals Haploid Life Cycle: Chlamydomonas • Individual is haploid • Gametes form by mitosis • Gametes fuse to form zygote – Zygote undergoes meiosis to form spores that grow by mitosis to form new individual – NO DIPLOID INDIVIDUAL!!!! Figure 27.15 A Haplontic Life Cycle Alternation of generations: Ulva and Laminaria • Ulva is green algae – Isogamic • – Isomorphic • – • Both generations phenotypically similar but differ in ploidy Alternation of generations Laminaria: brown algae – Oogamy • – – Fertilization of egg by sperm (gametes phenotypically distinguished, generally egg predominant) Heteromorphic • • Phenotypically indistinguishable gametes Generations phenotypically distinct Alternation of generations Plants – – – Oogamic (most) Heteromorphic Alternation of generations Figure 27.14 An Isomorphic Life Cycle Diploid Life Cycle: Fucus • Fucus: Brown Algae • All animals • Only individual is diploid • Gametes by meiosis Survey of the Protists • Cell wall or test • Absence of cell wall – Contractile vacuoles for maintaining osmotic pressure • Type of nutrition – Photoautotroph – Heterotroph by ingestion – Heterotroph by absorption • Locomotion – – – – • Non-motile Flagella Cilia pseudopodia Note - this arrangement does not follow phylogeny - grouped in many textbooks by convenience! Animal-like Protists • In general – – – – Motile Unicellular or colonial Wall-less Heterotrophic • Protists with Pseudopodia • Rhizopodia (amoebas) – Pseudopodia – Entamoeba histolytica - amoebic dysentery • Foraminifera (forams) – Tests made of CaCO3 – Long-hairlike pseudopodia that poke out thru holes of test • Actinopoda (radiolarians) – Silicon test – Long-hairlike pseudopodia that poke out thru holes of test Animal-like Protists • Ciliophora (cilliates) – Complex organization – Asexual reproduction; sexual by conjugation involving micronuclei Animal-like Protists • Zoomastigophora (zooflagellates) – Generally symbionts or parasites – Giardia – Trypanosoma • Chagas • African sleeping sickness – Choanoflagellates • Closest relative to Animals!!!!!! (rRNA analysis) Animal-like Protists • Apicomplexa (sporozoans) – Nonmotile, parasitic, spore-forming – Complicated life-cycles – Toxoplasma – Plasmodium: malaria Figure 27.27 A Link to the Animals Figure 27.1 Major Eukaryote Groups in an Evolutionary Context (Part 1) Plant-like Protists • Pyrrophyta (dinoflagellates) – 2 flagella; one wraps around middle of cell – Cell protected by celluose/silica plates – Chlorophylls a and c, carotenoids – Red-tide • Chrysophyta (golden-brown algae; diatoms) – Diatoms formally called Bacillariophyta • Diatoms have cell wall of silica; major component of phytoplankton – Chlorophyll a and c, fucoxathin • Euglenophyta (eugleniods) – 1/3 have chloroplasts, rest do not – Chloroplasts like those of green algae • Chlorophyll a, b and carotenoid – 2 flagella – No cell wall – Eyespot to detect light Plant-Like Protists • Chlorophyta (green algae) – Closest relatives to plants • Chlorophyll a, b, carotenoids • Store food as starch • Walls of cellulose – Lichens: green algae + fungi • Rhodophyta (red algae) – Unicellular to multicellular – Chlorophyll a, phycobilins – Food stored as floridian starch • Phaeophyta (brown algae) – All multicellular – Chlorophylls a and c, fucoxanthin – Store good as laminarin Figure 27.23 Chlorophytes Figure 27.22 Red Algae Figure 27.20 Brown Algae Importance of Protists • Link to eukaryotic origins • Impact on Human Health – Plasmodium (malaria) – Trypanosoma (African sleeping sickness; Chagas) • Ecological Importance as Primary Producers – Dinoflagellates • Marine phytoplankton • Endosymbiotic with corals • Ride tides and algal blooms – Diatoms • Marine phytoplankton • Common in fresh water • Diatomaceous earth – Chlorophytes (green algae) • Links to origins of animal kingdom - choanoflagellates Figure 27.8 Dinoflagellate Endosymbionts are Photosynthesizers Figure 27.10 Chromalveolates Can Bloom in the Oceans Figure 27.19 Diatom Diversity