Figure 28.2 ■ Excavata 5 Μm Diplomonads Excavata Parabasalids 4 Supergroups
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2/19/2015 Chapter 28: Protists 1. General Features of Protists 2. Survey of the Protista A. The Excavata B. The SAR Clade C. The Archaeplastida D. The Unikonta 1. General Features of Protists All Protists are Eukaryotes Eukaryotic organisms consist of 1 or more eukaryotic cells: Prokaryotic cell • eukaryotic cells contain a “true nucleus” and other membrane- bound organelles Nucleus Most eukaryotes are single-celled organisms, most of which are protists (i.e., not fungi, plants or animals). Eukaryotic cell Organelles 1 2/19/2015 Protists have Diverse Characteristics Protists are much more diverse that plants, animals & fungi: • most are unicellular, though some are multicellular • some are photoautotrophs (commonly called “algae”) • some are chemoheterotrophs (commonly called “protozoa”) • some have characteristics of both (mixotrophs) • protists can reproduce sexually, asexually, or both Endosymbiosis in Eukaryotic Evolution Plasma membrane The endosymbiont theory proposes Cytoplasm that mitochondria & chloroplasts are DNA derived from internalized prokaryotes. Ancestral prokaryote Nucleus Endoplasmic Photosynthetic reticulum prokaryote Mitochondrion Nuclear envelope Aerobic heterotrophic prokaryote Mitochondrion Plastid Ancestral Ancestral photosynthetic heterotrophic eukaryote eukaryote A Closer Look at Plastid Evolution Dinoflagellates Membranes Secondary are represented endosymbiosis as dark lines in the cell. Red alga Cyanobacterium Plastid 1 2 3 Primary endosymbiosis Stramenopiles Nucleus Secondary Plastid Heterotrophic endosymbiosis eukaryote One of these membranes was lost in Euglenids red and green algal descendants. Secondary endosymbiosis Green alga Chlorarachniophytes 2 2/19/2015 Protists No Longer Constitute a Single Kingdom The traditional grouping of eukaryotes involved 4 kingdoms: PROTISTA FUNGI PLANTAE ANIMALIA However this is not consistent with phylogenies based on recent molecular analysis which supports the following 4 supergroups… Figure 28.2 ■ Excavata 5 μm Diplomonads Excavata Parabasalids 4 Supergroups Euglenozoans Stramenopiles ■ “SAR” Clade 50 μm Diatoms The evolutionary Golden algae Brown algae Giardia intestinalis, history of all “SAR” “SAR” clade Alveolates Dinoflagellates eukaryotes Apicomplexans currently consists Ciliates Forams Rhizarians of 4 supergroups: Cercozoans Diatom diversity ■ Archaeplastida Radiolarians 50 μm 1. Excavata Archaeplastida Red algae Green algae Chlorophytes 2. SAR Clade Charophytes 3. Archaeplastida Land plants Amoebozoans • includes all land plants Slime molds Volvox, a colonial freshwater green alga Tubulinids ■ Unikonta 4. Unikonta Entamoebas Unikonta • includes fungi & animals Nucleariids Opisthokonts Fungi All eukaryotes that are not Choanoflagellates plants, animals or fungi are Animals 100 μm considered to be Protists. A unikont amoeba 2A. Survey of the Protista The Excavata Diplomonads Excavata Parabasalids Euglenozoans SAR clade Archaeplastida Unikonta 3 2/19/2015 General Characteristics of Excavates Below are some of the characteristics seen in Excavates: • some members have an “excavated” feeding grooves that are associated with unique cytoskeletal structures • many have modified “unclassical” mitochondria • most have 2 or more flagella • includes the following major groups: DIPOMONADS PARABASALIDS EUGLENOZOANS Diplomonads Diplomonads have reduced mitochondria called mitosomes and derive energy by anaerobic processes. They also have 2 nuclei and multiple flagella. Many are parasites such as Giardia intestinalis shown here: Parabasalids Parabasalids have reduced mitochondria called hydrogenosomes and thrive in anaerobic environments. • Trichomonas vaginalis, a sexually transmitted human pathogen shown below is one example Flagella Undulating membrane 5 μm 4 2/19/2015 Euglenozoans The Euglenozoa is a diverse clade that includes predatory heterotrophs, photosynthetic autotrophs, mixotrophs, and parasites. Flagella • a characteristic feature of this 0.2 μm clade is the crystalline rod structure within their flagella • members of this clade include: KINETOPLASTIDS 8 μm Crystalline rod (cross section) EUGLENIDS Ring of microtubules (cross section) Kinetoplastids Kinetoplastids have a single mitochondrion containing an unusual organized mass of DNA called a kinetoplast. • includes the genus Trypanosoma which contains parasites that cause “sleeping sickness” and Chaga’s disease RBC 9 μm Euglenids Euglenids have 1 or 2 flagella, and some members of this clade, such as the genus Euglena, are both autotrophic and heterotrophic (i.e., mixotrophs). Long flagellum Eyespot Light Short flagellum detector Contractile vacuole Nucleus Chloroplast Plasma 5 μm membrane Euglena (LM) Pellicle 5 2/19/2015 2B. Survey of the Protista The SAR Clade Excavata Diatoms Golden algae Stramenopiles SAR SAR clade Brown algae Dinoflagellates Apicomplexans Alveolates Ciliates Forams Cercozoans Rhizarians Radiolarians Archaeplastida Unikonta General Characteristics of the SAR Clade The SAR clade is a highly diverse monophyletic supergroup based on DNA similarities that includes the following: STRAMENOPILES ALVEOLATES RHIZARIANS Stramenopiles Most members of the Stramenopile clade have both “hairy” and smooth flagella. • this clade includes the following ecologically important groups of photosynthetic Hairy organisms: flagellum Smooth DIATOMS flagellum GOLDEN ALGAE BROWN ALGAE 5 μm 6 2/19/2015 Diatoms Diatoms have unique glass-like walls made of silicon dioxide and are a significant portion of phytoplankton in the oceans. • their remains are the main components of white sediments referred to as “diatomaceous earth” (e.g., the “white cliffs of Dover”) m μ 40 Flagellum Golden Algae Outer container • contain yellow and brown carotenoids Living cell which give them their characteristic color • typically have two flagella • most are single- celled though some are colonial 25 μm Brown Algae Brown algae are all multicellular and are the largest algae. • includes many “seaweeds” Blade such as kelp • although brown algae are Stipe not plants, many have plant-like structures shown in this image Holdfast 7 2/19/2015 Haploid (n) Alternation of Diploid (2n) Sporangia Generations • many multicellular MEIOSIS 10 cm algae have a life cycle Sporophyte similar to that of (2n) Zoospore Female plants called the Developing Gameto- “Alternation of sporophyte phytes Generations” – the Zygote (n) Mature female (2n) gametophyte Egg Male alternation between (n) FERTILIZATION multicellular haploid and diploid forms Sperm Alveolates Members of the Alveolate clade have membrane enclosed sacs called alveoli must beneath the plasma membrane. Flagellum Alveoli • this clade includes the following: DINOFLAGELLATES Alveolate APICOMPLEXANS m μ CILIATES 0.2 Dinoflagellates Flagella Dinoflagellates are also an (a) Dinoflagellate important component of flagella phytoplankton and have 2 flagella, one of which wraps around the cell and causes it to 3 μm rotate. • are responsible for the algal blooms known as “red tides” (b) Red tide in the Gulf of Carpentaria in northern Australia 8 2/19/2015 Apicomplexans The Apicomplexans are parasites of animals that have complex life cycles with multiple hosts. • the most significant genus in this clade is Plasmodium, the cause of malaria • nearly 1 million people each year die from malaria Inside mosquito Inside human Merozoite Plasmodium Sporozoites (n) Liver Life Cycle Liver cell • mosquitoes are Apex the definitive host Oocyst MEIOSIS (in which sexual Merozoite Red blood 0.5 μm cell reproduction (n) Red Zygote blood occurs) (2n) cells • humans are the FERTILIZATION intermediate host (in which reproduction Gametes Game- is asexual) tocytes Haploid (n) (n) Diploid (2n) Ciliates The Ciliates clade is a large group characterized by cilia used for locomotion and/or feeding and 2 nuclei, one macronucleus and one micronucleus. • the genus Paramecium is a good example Contractile • Ciliates also engage vacuole in a form of sexual Oral groove Cilia Cell mouth reproduction called 50 μm conjugation… Micronucleus Food Macronucleus vacuoles (a) Feeding, waste removal, and water balance. 9 2/19/2015 Conjugation in Ciliates Compatible mates Conjugation Asexual MEIOSIS reproduction Haploid Diploid micronucleus micronucleus The original macro- Diploid nucleus disintegrates. micronucleus MICRO- NUCLEAR FUSION (b) Conjugation and reproduction. Rhizarians The Rhizarians are mostly amoebas containing pseudopodia that are threadlike (unlike the Amoebozoa in the Unikonta clade. • this clade includes the following: RADIOLARIANS FORAMS CERCOZOANS Radiolarians Radiolarians have delicate, symmetrical endoskeletons made typically of silica. • the pseudopodia extend from the cell to catch prey Pseudopodia 200 μm 10 2/19/2015 Forams Foraminiferans (“hole bearers”), or forams for short, are single- celled heterotrophs that characteristically have multi-chambered shells called tests. • foram tests are found throughout the fossil record and their tests can be analyzed to estimate ocean temperatures in ancient times Cercozoans Chromatophore • most are heterotrophs, though a few such as Paulinella chromatophora are autotrophs that contain unique photosynthetic structures called chromatophores 5 μm 2C. Survey of the Protista The Archaeplastida Excavata SAR clade Archaeplastida Red algae Chlorophytes Green algae Charophytes Land plants Unikonta 11 2/19/2015 The Archaeplastida The supergroup Archaeplastida contains the following: RED ALGAE GREEN ALGAE LAND PLANTS • molecular and other data indicate the land plants are descendants