v How long ago is Earth thought to have formed? v What is thought to have been the first genetic material? v Are we tetrapods?

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1 Biosc 41 Announcements 9/29 v Quick review followed by lecture quiz (history & phylogeny) v Lecture: Protists v Lab: Protozoa (animal-like protists) v Lab exam 1 is Wed! (does not cover today s lab) Review: History of Life v How long ago is Earth thought to have formed? v What is thought to have been the first genetic material? v Are we tetrapods? v Most atmospheric oxygen comes from photosynthesis Since many of the first organisms were photosynthetic (i.e. cyanobacteria), a LOT of excess oxygen accumulated (O 2 revolution) Some organisms adapted to use it (aerobic respiration) Review: History of Life v Which organelles are thought to have originated as endosymbionts? v During what event did fossils resembling modern taxa suddenly appear en masse? v How many mass extinctions seem to have occurred during Earth s history? Describe one? v When is adaptive radiation likely to occur? Review: Phylogeny v Homology is similarity due to shared ancestry v Analogy is similarity due to convergent evolution v A valid clade is monophyletic, meaning it consists of the ancestor taxon and all its descendants v A paraphyletic grouping consists of an ancestral species and some, but not all, of the descendants v A polyphyletic grouping includes distantly related species but does not include their most recent common ancestor v Maximum parsimony assumes the tree requiring the fewest evolutionary events is most likely Quiz 3 (History and Phylogeny) 1. How long ago is Earth thought to have formed? 2. Why might many organisms have evolved to use aerobic respiration? 3. Which two organelles are thought to have originated as endosymbionts? BIOSC 041 PROTISTS! Reference: Chapter is similarity due to shared ancestry, while is similarity due to convergent evolution. 5. A valid clade is, meaning it consists of the ancestor taxon and all its descendants. 1

2 Fungi Charophytes Land plants Opisthokonts Galatheammina (Xenophyophore) Benthic (deep sea) protozoan, ~10 cm in diameter Single-celled(!!) ruffled surface increases surface area for gas/nutrient exchange Red algae Chlorophytes Choanoflagellates Animals Single-celled protists can be very complex, as all biological functions are carried out by organelles in each individual cell Most protists are unicellular, and small Include unicellular, colonial, and multicellular taxa but there are many exceptions Nucleariids Brown algae Green algae Golden algae SAR clade Amoebozoans The Protists exhibit more structural and functional diversity than any other group of eukaryotes Diatoms Rhizarians Everything that is not something else But definitely not an animal The one-time kingdom of Protista has been abandoned Protists are now recognized as polyphyletic Some lineages of protists recognized as kingdoms Protist remains the informal name for highlighted taxa Protozoa are animal-like protists Alveolates General characteristics of protists Our understanding of the relationships among protist groups continues to change rapidly! One hypothesis divides all eukaryotes (including protists) into four supergroups: SAR clade Roles in Ecological Communities Producers Consumers Parasites & pathogens Diplomonads Parabasalids Stramenopiles What is a Protist? Outline Most range in size from <1-200 µm The Protists also include colonial and multicellular taxaparticularly algae Algae are considered protists because they don t meet the complex tissue criteria of true plants Range in size from 100 s um to 100 s meters (giant kelp) 2

3 Nutritional and reproductive diversity v Nutrition Photoautotrophs Contain chloroplasts Obtain energy from the sun and carbon from CO 2 Heterotrophs Obtain both energy and carbon from organic molecules, absorbed from the environment or through ingestion of other organisms Mixotrophs Combine photosynthesis and heterotrophic nutrition v Reproduction Sexual, asexual, or both! Endosymbiosis in Eukaryotic Evolution v Protists owe their diversity to endosymbiosis- a unicellular organism engulfs another cell, which becomes an endosymbiont and then an organelle in the host cell Review: Mitochondria evolved by endosymbiosis of an aerobic heterotrophic bacterium Chloroplasts evolved by endosymbiosis of photosynthetic cyanobacterium v The plastid-bearing protists evolved into red and green algae (DNA of plastid genes in red algae and green algae closely resemble DNA of cyanobacteria) v During eukaryotic evolution, red and green algae also underwent secondary endosymbiosis, in which they were ingested by a heterotrophic eukaryote Figure 28.2 Plastid Five Supergroups of Eukaryotes Diplomonads Parabasalids Membranes are represented as dark lines in the cell. Cyanobacterium Primary endosymbiosis Heterotrophic eukaryote One of these membranes was lost in red and green algal descendants. Red alga Green alga Secondary endosymbiosis Secondary endosymbiosis Secondary endosymbiosis Stramenopiles Plastid Euglenids Chlorarachniophytes v Our understanding of the relationships among protist groups continues to change rapidly v One hypothesis divides all eukaryotes (including protists) into four supergroups Stramenopiles Alveolates Rhizarians Amoebozoans Opisthokonts Green algae Diatoms Golden algae Brown algae Red algae Chlorophytes Charophytes Land plants Nucleariids Fungi Choanoflagellates Animals SAR clade Supergroup #1: Excavates v Include protists with modified mitochondria and protists with unique flagella v The clade is characterized by its cytoskeleton v Some members have a feeding groove v This diverse group includes Diplomonads Parabasalids Diplomonads Parabasalids : Diplomonads and Parabasalids v These two groups lack plastids, have modified mitochondria, and most live in anaerobic environments v Diplomonads Have modified mitochondria called mitosomes Derive energy from anaerobic biochemical pathways Have two equal-sized nuclei and multiple flagella Are often parasites, for example, Giardia intestinalis (also known as Giardia lamblia) 3

4 Parabasalids v Have reduced mitochondria called hydrogenosomes that generate some energy anaerobically v Include Trichomonas vaginalis, the pathogen that causes a type of vaginal infection in human females v Euglenozoa is a diverse clade that includes predatory heterotrophs, photosynthetic autotrophs, and parasites v Distinguishing feature A spiral or crystalline rod of unknown function inside their flagella v This clade includes the Kinetoplastids Euglenids Euglenozoa - Kinetoplastids v Kinetoplastids are distinguished from other protozoa by the kinetoplast- a single mitochondrion containing an organized mass of DNA (kdna) that comprises several copies of the mitochondrial genome v Include free-living consumers of prokaryotes in freshwater, marine, and moist terrestrial ecosystems v Also include some pathogens Trypanosoma, which causes sleeping sickness in humans Another pathogenic trypanosome causes Chagas disease More about trypanosomes v Trypanosomes evade immune responses by switching surface proteins v A cell produces millions of copies of a single protein v The new generation produces millions of copies of a different protein v These frequent changes prevent the host from developing immunity Euglenozoa - Euglenids v Euglenids have one or two flagella that emerge from a pocket at one end of the cell v Some species can be both autotrophic and heterotrophic Supergroup #2: v A highly diverse group of protists defined by DNA similarities v The SAR clade is a diverse monophyletic supergroup named for the first letters of its three major clades Stramenopiles, Alveolates, and Rhizarians v This group is the most controversial of the four supergroups (sometimes split to give 5 subgroups) v Stramenopiles covered in Biosc 42 Diatoms Golden algae Brown algae Stramenopiles Alveolates Rhizarians 4

5 Alveolata Members of the clade Alveolata have membrane-bounded sacs (alveoli) just under the plasma membrane The function of the alveoli is unknown The alveolates include have two flagella and each cell is reinforced by cellulose plates They are abundant components of both marine and freshwater phytoplankton They are a diverse group of aquatic phototrophs, mixotrophs, and heterotrophs Toxic red tides are caused by dinoflagellate blooms More about are parasites of animals, and some cause serious human diseases No movement, no free-living forms (obligate parasites) Complex intracellular structures and lifecycles (spread through host as infectious cells called sporozoites) One end, the apex, contains a complex of organelles specialized to penetrate host cells and tissues Most have sexual and asexual stages that require two or more host species for completion Figure Inside mosquito Responsible for some of the most serious human disease Malaria (Plasmodium), Toxoplasmosis Plasmodium requires both mosquitoes and humans to complete its life cycle ~900,000 people die each year from malaria Efforts are ongoing to develop malaria vaccines Inside human Merozoite Sporozoites Liver Liver cell Red blood cell Merozoite Zygote (2n) 0.5 µm Red blood cells FERTILIZATION Gametes Gametocytes Paramecium caudatum is a good example of a ciliate Have large macronuclei and small micronuclei Genetic Variation Exchange of haploid micronuclei via sexual conjugation Reproduction Asexual- binary fission Apex Oocyst MEIOSIS, a large varied group of protists, are named for their use of cilia to move and feed Key Haploid Diploid (2n) 5

6 Figure Rhizarians Contractile vacuole 50 µm Oral groove Cell mouth Cilia Micronucleus Food vacuoles Macronucleus (a) Feeding, waste removal, and water balance Key Conjugation Asexual reproduction MEIOSIS Compatible mates Diploid micronucleus Haploid micronucleus DNA evidence supports Rhizaria as a monophyletic clade (so it sometimes is split from SAR into its own supergroup) Rhizaria includes some groups of amoebas ( testate ameobas = with shells), including radiolarians, forams, and cercozoans Amoebas are protists that move and feed by pseudopodia, extensions of the cell surface Rhizarian amoebas differ from amoebas in other clades by having threadlike pseudopodia Diploid The original macronucleus micronucleus disintegrates. MICRONUCLEAR FUSION (b) Conjugation and reproduction Marine protists called radiolarians have tests fused into one delicate piece, usually made of silica use their pseudopodia to engulf microorganisms through phagocytosis The pseudopodia of radiolarians radiate from the central body Supergroup #3: include most amoeboid and flagellated protists with threadlike pseudopodia They are common in marine, freshwater, and soil ecosystems Most are heteroptrophs, including parasites and predators Foraminiferans, or forams, are named for porous, generally multichambered shells, called tests Organic material hardened with CaCO3 Pseudopodia extend through the pores in the test Foram tests in marine sediments form an extensive fossil record Many forams have endosymbiotic algae 20 cm Covered in detail in Biosc 42 Includes red algae, green algae, and land plants Dulse (Palmaria palmata) Charophytes Green algae Land plants Red algae Chlorophytes 2 cm Imbricatea (a) Ulva, sea lettuce 6

7 Supergroup 4: Unikonts Amoebozoans Includes animals, fungi, and some protists (that are closely related to fungi and animals) This group includes two clades: the amoebozoans and the opisthokonts (animals, fungi, and related protists) The root of the eukaryotic tree remains controversial It is unclear whether unikonts separated from other eukaryotes relatively early or late Amoebozoans are amoeba that have lobe- or tubeshaped, rather than threadlike, pseudopodia- and no shell They include slime molds, tubulinids, and entamoebas Nucleariids Fungi Choanoflagellates Animals Slime Molds (mycetozoans) once thought to be fungi Molecular systematics places slime molds in the clade Amoebozoa Two types: Plasmodial (no cell membranes, but many nuclei) Cellular (cell membranes present) Give clues to evolution of multicellularity Plasmodial Slime Molds At one point in the life cycle, plasmodial slime molds form a mass called a plasmodium (not to be confused with malarial Plasmodium) The plasmodium is not multicellular It is undivided by plasma membranes but contains many diploid nuclei Extends pseudopodia through decomposing material, engulfing food by phagocytosis Plasmodial Slime Molds Many species of plasmodial slime molds are brightly pigmented, usually yellow or orange Cellular Slime Molds Cellular slime molds form multicellular aggregates in which cells are separated by their membranes Cells feed individually, but can aggregate to form a fruiting body Spores FERTILIZATION Emerging amoeba Solitary amoebas 600 µm Fruiting bodies ASEXUAL REPRODUCTION Zygote (2n) SEXUAL REPRODUCTION MEIOSIS Amoebas Aggregated amoebas Migrating aggregate 200 µm Key Haploid Diploid (2n) 7

8 Have lobe or tube-shaped pseudopodia. Typical amoeboid movement Common unicellular amoebozoans in soil as well as freshwater and marine environments Most are heterotrophic and actively seek and consume bacteria and other protists Opisthokonts Protists are found in diverse environments Aquatic, marine and terrestrial Protists are found in all trophic levels Producer (photoautotrophs) Various types of consumer (heterotrophs feeding on photoautotrophs or other heterotrophs) Parasites (including human pathogens) Detritovore Photosynthetic Protists are parasites of vertebrates and some invertebrates Entamoeba histolytica causes amebic dysentery, the third-leading cause of human death due to eukaryotic parasites Protists play key roles in ecological communities Opisthokonts include animals, fungi, and several groups of protists Nucleariids (amoeba more similar to fungi) Choanoflagellates Nucleariids Fungi Choanoflagellates Animals Many protists are important producers that obtain energy from the sun In aquatic environments, photosynthetic protists and prokaryotes (cyanobacteria) are the main producers In aquatic environments, photosynthetic protists are limited by nutrients Primarily N and/or P, sometimes Fe These populations can explode (bloom) when limiting nutrients are added Important in recycling nutrients Symbiotic Protists Some protist symbionts benefit their hosts (zooxanthellae) photosynthesize to provide food for coral polyps that build reefs Wood-digesting protists digest cellulose in the gut of termites 8

9 Parasitic Protists v Plasmodium causes malaria v Pfiesteria shumwayae is a dinoflagellate that causes fish kills A Protist as Ecosystem Engineer? v Galatheammina (Xenophyophore- giant unicellular organism) v Can form very dense populations on undersea plains v Stirs up sediment and creates structure with slime secretions Creates habitat for other bottom-dwelling organisms Populations of crustaceans and worms much higher where Galatheammina is present Plays key role in circulating C & N back into the water column v Phytophthora ramorum causes sudden oak death Global warming and phytoplankton Figure 28.UN06 Eukaryote Supergroup Major Groups Diplomonads and parabasalids Key Morphological Characteristics Modified mitochondria Specific Examples Giardia, Trichomonas v Phytoplankton form the basis for marine and aquatic food webs (most phytoplankton are protists) v Biomass of photosynthetic protists has declined as sea surface temperature has increased v If sea surface temperature continues to warm due to global warming, this could have large effects on Marine ecosystems Fishery yields The global carbon cycle SAR Clade Kinetoplastids Euglenids Stramenopiles Diatoms Golden algae Brown algae Alveolates Rhizarians Red algae Green algae Spiral or crystalline rod in- Trypanosoma, side flagella Euglena Hairy and smooth flagella Phytophthora, Laminaria Membrane-enclosed sacs Pfiesteria, (alveoli) beneath plasma Plasmodium, membrane Paramecium Amoebas with threadlike Globigerina pseudopodia Phycoerythrin (photosyn- Porphyra thetic pigment) Plant-type chloroplasts Chlamydomonas, Ulva Land plants (See Chapters 29 and 30.) Mosses, ferns, conifers, flowering plants Amoebozoans Amoebas with lobeshaped or tube-shaped pseudopodia Amoeba, Dictyostelium Opisthokonts (Highly variable; see Chapters ) Choanoflagellates, nucleariids, animals, fungi 9

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