Protozoa I Protozoans Over 50,000 known species 45 phyla (more than metazoa!) Relationship to Other Organisms Two Kingdoms – Arististotle, Linnaeus Relationship to Other Organisms Plants Plants Metaphyta Protophyta Animals Protists Animals Metazoa Protozoa Lots of problems with this scheme 3 kingdoms of Haeckle/Darwin Took care of the little stuff seen with the early microscopes Still has problems 1 Relationship to Other Organisms Copeland’s Four-Kingdom System (1938) Kingdom Kingdom Monera. Protoctista or Protista (priority?). Protozoa Red and brown algae Fungi Kingdom Plantae Green algae Fungi Kingdom Animalia Relationship to Other Organisms Whittaker – 1960’s Added Kingdom Fungi 5 Kingdoms Kingdom Animalia Plantae Eukaryotes Kingdom Fungi Kingdom Protista Kingdom Monera - Prokaryotes Kingdom Kingdoms and Domains Carl Woese - U. of Illinois (1970’s-present) Studied gene sequences of bacteria, archaea, and eukaryotes Found fundamental differences (major!) Relationship to Other Organisms Carl Woese – late 1970’s Archaea NOT Archaebacteria Biochemistry is different from bacteria More closely related to animals than they are to bacteria Briefly had six kingdoms with Archaea and Eubacteria replacing the Monera Changed to three domains Eubacteria Prokaryotes Archaea Eukarya 2 6 Kingdoms of Life 3 Domains of Life Figure 1: All organisms are connected by the passage of genes along the branches of the phylogenetic Tree of Life. Relationship to Other Organisms Protista or Protoctista Some algae (red, most green algae are not included) Protozoa Traditionally classified based on how they move: amoebae, flagellates, ciliates, sporozoans Has changed recently to also contain: Some slime molds Aquatic “molds” Figure 2: Living organisms sit like leaves at the tips of the branches of the Tree of Life. Their evolutionary history is represented by a series of ancestors which are shared hierarchically by different subsets of the organisms that are alive today. Old system classified by locomotion Ciliophora (=Ciliata, ciliates) a clade Hypotrichs, holotrichs, heterotrichs, suctorians Apicomplexa- (=Sporozoa) a likely clade Gregarina, Coccidia – includes many important parasites Mastigophora (=flagellates) a functional group Excavates, Kinetoplastids, Parabasalids, Choanoflagellates, Dinoflagellates (some are important parasites) Protozoan Phylogeny Problem Protozoa is a polyphyletic group Multiple ancestors rather than a single ancestral protozoan Aim is to establish monophyletic groups that have a single ancestor Sarcodina (=amoebas) a functional group Amoebozoa, Foraminifera, Actinopoda (Radiolaria, Heliozoa) (many are important geologically) 3 Protozoan Phylogeny Reclassification based on History Ultrastructure Biochemistry Molecular data including DNA sequencing Most trees are being constructed based on molecular data May or may not be the “best” way to go Dissenting camps: role of lateral gene transfer, choice of genes for sequencing, convergent evolution, etc. Relationship to Other Organisms Life Eukaryote classification is in flux The major clades are not yet sorted out – but there is rapid progress Based on cell structural features, there are about 60 different named eukaryote taxa according to Patterson (Tree of Life) These have been sorted into 8 clades based on molecular & structural data by Baldauf 2003 (Science 300:1703) Protista – still used as teaching tool. Grades 7-12? intro bio (with reservations) Undergrad We will not discuss Protists! Instead, note the following schemes: First - breakdown of what used to be protists. Second - taxonomic groupings of organisms covered in Ch 3 (pg 71ff). Focus on those covered in text and/or lab Kingdom Chromista The Alveolata Flattened membranous alveoli (sacs) under outer cell membrane The Alveolates Phylum Ciliophora Phylum Dinozoa (Dinoflagellata) Phylum Apicomplexa 4 The Rhizaria Very diverse group based on molecular data Tubular cristae 3 Phyla The Heterokonta Heliozoans? – probably polyphyletic Opalinata Flagellated parasites of intestines of ectothermic vertebrates Class Phylum Foraminifera Radiolaria Phylum Cercozoa Phylum Kingdom Protozoa The Amoebozoa Branching, tubular cristae Lobose pseudopodia Arcellanids Xenophyophoreans The Excavata Class Mastigamoididae Phylum Eumycetozoa Class Euglenida Order Kinetoplastea Disc-shaped mitochondrial cristae Deep ventral feeding groove (hence excavate) Phylum Parabasala – no mitochondria Class Phylum Phylum Euglenozoa Phylum Stramenophiles Trichomonadida Hypermatigia Phylum Heterolobsea The diplomonads – no mitochondria The oxymonads 5 The Opisthokonta Phylum Choanomonada Phylum Fungi Phylum Microsporidia The Myxozoans – probably degenerate cnidarians Phylum(?) Animalia Defining Characteristics of “Protozoa” 1674 Antonie van Leeuwenhoeke Nearly ubiquitous –wherever there is water Soil Water On/in plants and animals Life styles Free-living Symbioses Mutualists Parasites Antonie van Leeuwenhoeke Things that animals do (and protozoa too) Move (at some stage in the life cycle) Obtain food and digest it Obtain oxygen Maintain water and salt balance Remove metabolic wastes Reproduce Sense and react to the environment Defining Characteristics of “Protozoa” Haeckel coined the termed Protista Protist was anything that wasn’t clearly plant or animal Now it’s anything that isn’t plant, animal, fungi or bacteria Most are probably unknown Over 84,000 species of these are fossils (shelled forms) No formal taxonomic category called Protista ½ 6 Defining Characteristics of “Protozoa” General characteristics highly variable Size, morphology, ultrastructure Nutritional mode, physiology Behavior, life history Defining Characteristics of “Protozoa” Disease Model Ecology Genetics Physiology Development Ancestral to multicellular animals and plants (polyphyletic): ex. Choanoflagellates and sponges Mutualists in other inverts: ex. dinoflag. in corals Disease agents: ex. Plasmodium in malaria Eukaryotic Unicellular Mostly small 5-250 μm (0.5 μm – 7 mm) in a few but this is a derived character Multicellularity No collagen or chitin in cell walls Heterotrophic Ancestral state is non-photosynthetic Photosynthesis in a few groups is a derived character Most are motile (except Apicomplexa) agents organisms Importance Defining Characteristics of “Protozoa” Importance Protist Bauplan - On Being Unicellular Strategies and constraints of a Unicellular Bauplan Size limitations Body structure Excretion Gas exchange Support and locomotion Nutrition Reproduction Activity and sensitivity On Being Unicellular Protozoa vs metazoa Protozoa are unicellular Thought BUT to be simplest form of life Urotricha globosa Cothurnia annulata Pseudodifflugia sp. … Protozoa are not simple! A protozoan is more complex than any single metazoan cell Very complex internal structure Specialized organelles take the place of organs in metazoans Halteria grandinella Ctedoctema acanthocryptum Vorticella aquadulcis 7 Stylonychia Defining Characteristics of “Protozoa” Similarity of structures does NOT imply relatedness Various types of evidence are used in testing hypotheses regarding evolutionary relatedness Could Gene be convergent evolution sequencing ultratructure Cellular Size limitations Surface area to volume ratio SA V increases as radius squared increases as radius cubed As cell becomes larger, diffusion becomes more and more difficult (~1 mm limit) Need to have lots of complex projections etc. to increase SA Need large SA/V 8 Body Structure Cytoplasm has two regions Ectoplasm Next to cell membrane Clear, stiff, gelatinous Endoplasm Inner portion More fluid in nature http://www.youtube.com/watch?v=85DGyFzxvy8 &feature=player_detailpage Body Structure Membrane has fluid mosaic structure Only one plasma membrane Everything is inside that membrane is identical to the plasma membrane of all other multicellular organisms High SA/V for protist cells Structure Membrane has fluid mosaic structure Lipids and proteins can move about laterally within the membrane 9 10 Body Structure Internal structures Cytoskeleton Cytoskeleton Myonemes Shape Rigidity Cytopharynx Cytopharynx Macronucleus 250 µm Cirri Body Structure Outer coverings - give shape/support/protection Pellicle Interlocking strips of protein below plasma membrane in Euglenids Gives cell shape and stability while permitting flexibility Eugloid movement = metaboly 11 Body Structure Lorica Vase-shaped protective shell Body Structure Test External “shell” Usually many parts Plates are below plasma membrane (cellulose plates in dinoflagellates, various materials in testate amoebas) CaCO3 foraminiferans Silica in radiolarians Body Structure The usual eukaryotic organelles May be more than one macronucleus Runs everyday activities of cell May be one or more micronuclei Used for sexual reproduction Some are anaerobes Most have no mitochondria or cytochromes, and have an incomplete TCA cycle (tricarboxylic acid cycle) Some contain hydrogenosomes-small membranedelimited organelles containing a unique electron transfer system that uses protons as terminal electron acceptors to form molecular hydrogen Trichonympha lives in the gut of termites Giardia is an intestinal parasite Mitochondria clockwise Paramecium Cryptomonad Euglena 12 Body Structure Some protozoa are anaerobes. Trichonympha lives in the gut of termites Giardia is an intestinal parasite Most have no mitochondria or cytochromes, and have an incomplete TCA cycle (tricarboxylic acid cycle). Some contain hydrogenosomes-small membranedelimited organelles containing a unique electron transfer system that uses protons as terminal electron acceptors to form molecular hydrogen. Some have symbiotic aerobic bacteria that do the job of TCA cycle for the host. Body Structure Special organelles Contractile Body Structure Defense against predation Change shape to become harder to eat. Euplotes detects presence of predator Chemically Physically Euplotes swells in middle and becomes too big for Lembadion to swallow Contractile Vacuole Osmoregulation in FW vacuoles Osmoregulation in freshwater species Contractile Vacuole Osmotic Regulation 13 Body Structure Special organelles Trichocysts Defense, prey capture Mucus Toxins Adherence to prey (Didinium) Gas Exchange No circulatory system All transport is by diffusion Plasma membrane must remain moist for gases to diffuse Restricts protozoa to moist habitats 14