Protozoa I Protozoans Over 50,000 known species 45 phyla (more than metazoa!) Relationship to Other Organisms Two Kingdoms – Arististotle, Linnaeus Plants Metaphyta Protophyta Animals Metazoa Protozoa Lots of problems with this scheme Relationship to Other Organisms 3 kingdoms of Haeckle/Darwin Plants Animals Protists Took care of the little stuff seen with the early microscopes Still has problems Relationship to Other Organisms Copeland’s Four-Kingdom System (1938) Kingdom Monera. Kingdom 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 Kingdom Kingdom Kingdom Kingdom Animalia Plantae Eukaryotes Fungi Protista Monera - Prokaryotes Kingdoms and Domains Carl Woese - U. of Illinois (1970’spresent) Studied gene sequences of bacteria, archaea, and eukaryotes Found major fundamental differences 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 6 Kingdoms of Life 3 Domains 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” “Protists” are now distributed among all kingdoms.” pg 37 incorrect. No prokaryotic protists! 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) Sarcodina (=amoebas) a functional group Amoebozoa, Foraminifera, Actinopoda (Radiolaria, Heliozoa) (many are important geologically) 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 Protozoan Phylogeny Reclassification based on Life 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 Protista – still used as teaching tool. Grades 7-12? Undergrad intro bio (with reservations) We will not discuss Protists! Instead, note the following schemes: First - breakdown of what used to be protists. Second - taxonomic relationships of organisms covered in P. Ch 3 (pg 38). Only cover those in text and/or lab 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) 8 major eukaryote clades (mainly from Baldouf 2003) 1. 2. 3. 4. Opisthokonts includes animals, fungi Single basal flagellum on reproductive cells, flat mitochondrial cristae (most eukaryotes have tubular ones). Plants includes land plants, green algae, red algae Plastids with just two outer membranes (primary plastids). Heterokonts includes brown algae, golden algae, diatoms, water molds Unique flagellum decorated with hollow tripartite hairs (stramenopiles) and, usually, a second plain one. Plastids have 4 membranes (secondary). Cercozoa includes foraminifera (& radiolaria?) Amoebae with filose pseudopodia, often living within tests. 5. 6. 7. 8. Amoebozoa includes lobose amoebae, most slime molds Mostly naked amoebae (lacking tests), often with lobose pseudopodia. Alveolates includes ciliates, dinoflagellates, apicomplexans Have systems of cortical alveoli directly beneath their plasma membranes Discicristates includes many flagellates, some slime molds Have discoid mitochondrial cristae Amitochondrial excavates includes parabasalids, diplomonads Most have an excavated ventral feeding groove, and all lack mitochondria. 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” General characteristics highly variable Size, morphology, ultrastructure Nutritional mode, physiology Behavior, life history Importance Disease agents Model organisms in biological research Most are probably unknown 82,000 known species ½ of these are fossils (shelled forms) Defining Characteristics of “Protozoa” Importance Disease agents Model organisms Ecology Genetics Physiology Development Defining Characteristics of “Protozoa” Eukaryotic Unicellular Mostly small 5-250 μm (0.5 μm – 7 mm) Multicellularity in a few but this is a derived character 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) 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 … 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 Stylonychia 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 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=85D GyFzxvy8&feature=player_detailpage Body Structure Only one plasma membrane Everything is inside that membrane Structure is identical to the plasma membrane of all other multicellular organisms High SA/V for protist cells Membrane has fluid mosaic structure Lipids and proteins can move about laterally within the membrane Membrane has fluid mosaic structure Body Structure Internal structures Cytoskeleton Shape Rigidity Cytoskeleton Myonemes 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 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 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 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 Body Structure Special organelles Contractile vacuoles Osmoregulation in freshwater species Contractile Vacuole Osmoregulation in FW Contractile Vacuole Osmotic Regulation Body Structure Special organelles Trichocysts Defense, prey capture Gas Exchange No circulatory system All transport is by diffusion Plasma membrane must remain moist for gases to diffuse Restricts protozoa to moist habitats