Kingdom Protoctista: Protozoa 1 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ Kingdom Protoctista versus Kingdom Protista General classification has seen marked changes since the mid-70s to the present. The term protista is still used in many schemes for classification. Protozoa was once considered a single phylum in the Kingdom Protista. Now considered to consist of at least 29 to 45 phyla within what used to be 3 different kingdoms (now the Kingdom Protoctista by some authors). Many taxonomists consider it best to group the different groups as different phyla. Margulis and Schwartz promote the term protoctista because protista denotes only organisms which are unicellular. The protoctista scheme contains many multi-cellular organisms with tissue specialization. The protista system places several of the protoctistan phyla into the plant and fungi kingdoms. According to the system used for classification by Lynn Margulis and Karlene Schwartz, members of the protoctista are identified primarily by exclusion from all the other kingdoms. • • • • Animals develop from a blastula. Plants develop from an embryo. Fungi develop from spores, and lack flagella and cilia Monerans (prokaryotes) lack a membrane bound nucleus. All remaining organisms are placed into the kingdom protoctista. They are aquatic, living in saltwater, freshwater, and of the watery tissues of other organisms. There are many protozoan parasites of both invertebrates and vertebrates. Depending on the source the Kingdom Protoctista is now considered to consist of at least 29 to 45 phyla within what used to be 3 different kingdoms. ¾ General Characteristics of Protoctistans 1. unicellular eukaryotes (some multinucleate, a few loosely multicellular), not all have mitochondria (microspores, many flagellates). 2. up to about 400 micrometer in size (some larger) 3. all have at least one nucleus 4. most are free living, but many parasitic forms including entire phyla 5. motile by a variety of mechanisms but also several non-motile taxa 6. Many have cyst stages secreted by trophic or spore stages Cysts/spores have four basic functions: • protect against unfavorable conditions • serve as sites for multiplication • assist in attachment to surfaces such as hosts • transmission stage from host to host 7. all types of nutrition are exhibited by the Kingdom. • autotrophs: photosynthesis • heterotrophs (holozoic vs. saprozoic) • phagocytosis: ingestion of solid particles (e.g., bacteria) • pinocytosis: same as phagocytosis but intake of liquid • saprozoic or saprotrophy: diffusion or active transport across membrane Kingdom Protoctista: Protozoa 2 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ Characteristics of Protozoa The protozoa are a diverse assemblage of unicellular eukaryotic organisms in the Kingdom Protoctista having at least two animal-like properties: (1) absence of a cell wall and (2) presence of at least one motile stage in the life cycle. All functions of life are performed within the limits of a one cell membrane. Although protozoans have no organs or tissues, there is division of labor within the cytoplasm, where various complex organelles are specialized to carry out specific tasks. These organelles tend to be more specialized than those of an average cell of a multicellular organism, functioning as skeletons, locomotory systems, sensory systems, conduction mechanisms, defense mechanisms, and contractile systems. Protozoa are often called "simple" organisms. However, many are quite complex. Ciliates, for example, are not only the most complex of protozoans but also the most elaborately organized of all known cells. Protozoans are widespread ecologically, being found in fresh, marine, and brackish water and in moist soils. Some are free-living; others live as parasites or in some other symbiotic relationship. The protozoa are an artificial assemblage of organisms placed together for convenience. Traditionally, four main groups of protozoa have been recognized: flagellates, amebas, spore formers, and ciliates. These were assembled in a single phylum, Protozoa, within the kingdom Animalia. A revision adopted by the Society of Protozoologists in 1980 recognized seven separate phyla. However, more recent DNA sequence analyses of genes have shown that the protozoa represent numerous clades of varying evolutionary relationships. For example, ameboid organisms (formerly Sarcodina) fall into numerous lineages with undetermined associations. Nevertheless, the amebas comprise recognizable morphological groups, which we collect for convenience into the informal heading "Ameobas." (or “Amebas”). ¾ Protozoa Taxonomy & Classification The following is a condensed Linnaean classification of the protozoan groups as presently recognized. Amebas = about 12,000 species; locomotion by pseudopodia; body naked or with external or internal test or skeleton; asexual reproduction by fission; sexuality, if present, associated with flagellated (rarely ameboid) gametes; most free-living, some parasitic. Several groups of uncertain affinities. Rhizopodans Locomotion by lobopodia, filopodia (thin pseudopodia that may branch but do not rejoin), or by cytoplasmic flow without forming discrete pseudopodia. (The rhizopodans are divided among several clades.) Examples: Amoeba, Endamoeba, Dif./lugia, Arcella, Chlamydophrys. Granuloreticulosans Locomotion by reticulopodia (thin pseudopodia that branch and often rejoin). Includes foraminiferans. Examples: Globigerina, Vertebralima. Actinopodans Locomotion by axopodia (long, slender pseudopodia). Includes radiolarians. (The actinopodans are divided among several clades.) Examples: Actinophrys, Clathrulina. Phylum Euglenozoa (yu-glen-a-zo'a) (Gr. eu-, good, true, + glime, cavity, socket, + zoon, animal). Movement by flagella; cortical microtubules. Examples: Euglena, Trypanosoma. About 7500 different species. Kingdom Protoctista: Protozoa 3 Biological Sciences 102 – Animal Biology – Notes & Vocabulary Phylum Chlorophyta (klor-of'i-ta) (Gr. chloros, green, + phyton; plant). Unicellular and multicellular algae; photosynthetic chlorophyll pigments, flagella of equal length and smooth, mostly free-living photo autotrophs. Example: Volvox. Phylum Dinoflagellata (dy'no-fla-jel-at'a) (Gr. dinos, whirling, + flagellum, little whip). Typically with two flagella, one transverse, one trailing; body usually grooved transversely and longitudinally, each groove containing a flagellum; chromoplasts bearing chlorophyll; freeliving, planktonic, parasitic, or mutualistic. Examples: Noctiluca, Ceratium, Gonyaulax. Phylum Apicomplexa (a'pi-com-plex'a) (1. apex, tip, + complex, twisted around, + a, suffix). Characteristic set of organelles (apical complex) at anterior end in some stages; cilia and flagella usually absent; all species parasitic; about 5500 species. Class Gregarinea (gre-ga -ryn' e-a) (1. gregarius, belong to a herd or flock). Mature gamete-producing individuals large, extracellular; gametes usually alike in shape and size; parasites of digestive tract or body cavity of invertebrates; life cycle with one host. Examples: Monocystis, Gregarina. Class Coccidea (kok-sid'e-a) (Gr. kokkos, kernel, grain). Mature gamete-producing individuals small, typically intracellular; parasites mostly of vertebrates. Examples: Plasmodium, Toxoplasma, Eimeria. Phylum Ciliophora (sil-i-of' o-ra) (1. cilium, eyelash, + Gr. phora, bearing). Cilia or ciliary organelles present in at least one stage of life cycle; usually two types of nuclei; binary fission across rows of cilia; budding and multiple fission also occur; sexuality involving conjugation, autogamy, and cytogamy; heterotrophic nutrition; mostly free-living; contractile vacuole typically present. (This is a very large group, now divided into three classes and numerous orders.) Examples: Paramecium, Colpoda, Tetrahymena, Stentor, Blepharisma, Epidinium, Vorticella, Euplotes, Didinium; about 9000 different species. ¾ Biological Contributions of Protozoa 1. 2. 3. 4. 5. 6. 7. 8. Intracellular specialization Division of labor between cells in colonial protozoa Asexual reproduction by mitosis appears in unicellular eukaryotes True sexual reproduction forming a zygote (occasionally found) Simplest “reflexes”/responses Shelled protozoa = first exoskeletons All types of nutrition developed Means of locomotion in aqueous media developed ¾ Types of Nutrition/Feeding Strategies Seen in Protozoans Protozoa may use one or more of the following: ¾ ¾ ¾ ¾ ¾ ¾ Autotroph Heterotroph Phagocytosis = phagotroph or holozoic feeder Ingest visible food particles Osmosis = osmotrophs or saprozoic feeders Ingest soluble food Kingdom Protoctista: Protozoa 4 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ Types of Protozoan Locomotion ¾ ¾ ¾ ¾ Flagella = flagellates Cilia = ciliates Pseudopodia = amebas Lobopodia = blunt tipped • Filipodia = thin and sharply pointed • Rhizopodia = branched filaments • Reticulopodia = branched filaments forming a • net-like structure • Axopodia = thin, pointed containing a central axial filament of microtubules ¾ FLAGELLATES (eg. Euglena) ¾ ¾ ¾ ¾ Move by means of a flagellum Formed of an axoneme = 9 + 2 arrangement of microtubules with dynein arms that use ATP to “walk” along one filament in an adjacent pair. This causes the tubules to slide past one another and bend due to the shear resistance provided by fibril spokes running to the central pair of microtubules Kinetosome (basal body) = third microtubule joins each of the 9 pairs to form 9 triplets at the flagellum base near cell membrane Kinetosomes have same structure as centrioles ¾ AMEBAS (eg. Ameba proteus) ¾ ¾ ¾ ¾ Cytoplasm is not homogeneous Ectoplasm = more transparent and more rigid and gel-like (“outer” plasma) Endoplasm = more granular and contains cell organelles (“inner” plasma) ¾ Pseudopodium Formation in Amebas ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Hyaline cap appears as an extension of the ectoplasm Endoplasm begins to flow toward and into the hyaline cap Actin subunits in the endoplasm attach to regulatory actin-binding proteins that prevent actin from polymerizing Endoplasm “fountains out” to the periphery when it nears hyaline cap Interactions with phospholipids the cell membrane release the actin subunits from the regulatory proteins so that the actin can now repolymerize into microfilaments These microfilaments bind to actin binding protein to form a gel-like tube through which the endoplasm flows as the pseudopodium extends Near the trailing edge of the gel, calcium ions activate a protein that severs releasing microfilaments and allowing myosin to bind the microfilaments and pull on them This contractile force at the trailing edge forces the fluid endoplasm (and actin subunits) back toward the hyaline cap Kingdom Protoctista: Protozoa 5 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ CILIATES ¾ ¾ ¾ Have same basic structures as a flagellum, but they are shorter All typical cilia and flagella have a kinetosome at their base Ciliary movement is vital to many species for handling food, reproduction, excretion and osmoregulation ¾ Protozoan Encystment ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Some protozoa can form dormant cysts to survive harsh environmental conditions Form resistant external coverings Metabolism shuts down Can occur in some parasitic forms Can be reactivated after decades or longer? Appears to be cyclic and tied to a stage in the life cycle Cilia and flagella are resorbed Golgi apparatus secretes the cyst wall material ¾ Contractile Vacuoles ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ For osmoregulation Empty more often in freshwater protozoa than marine forms Smaller species with greater surface-to-volume ratios have more rapid filling and expulsion rates Excretion of waste (ammonia for nitrogen metabolism) is by diffusion Contractile Vacuoles Remove excess water by osmosis Proton pumps on vacuole surface actively transport Hydrogen ions (H+) and bicarbonate ions (HCO3-). These ions accumulate within a vacuole causing water to flow into the vacuole As vacuole fuses to cell membrane water is expelled to the outside of the cell These ions can be readily replaced by the action of the enzymes - carbonic anydrase – using its substrates of CO2 and H2O ¾ Protozoan Reproduction ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Binary Fission Most common type of reproduction in protozoa When parent is larger than progeny with progeny growing to adult size = budding Multiple fission = several nuclear division occur before cytokinesis so that a number of individuals are produced simultaneously Schizony (multiple fission) is common among apicomplexans and amebas Somewhat like mitosis, but nuclear membrane often persists through mitosis Sexual Reproduction If all gametes looks alike = isogametes If gametes are of two different types = anisogametes Syngamy = fertilization of an individual gamete by another Autogamy = fertilization in which gametic nucle form by meiosis and fuse to form a zygote in the same organism Conjugation = exchange of gametic nuclei between paired organisms Kingdom Protoctista: Protozoa 6 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Reproduction in Paramecium Binary fission = micronucleus divides mitotically into two daughter micronuclei and move to opposite ends of the cell. The macronucleus elongates and splits, but not by mitosis. Autogamy = “self-fertilization”; similar to conjugation with no exchange of nuclei with another Paramecium Conjugation in Paramecium Temporary union of two individuals to exchange chromosomes Macronucleus degenerates and micronucleus undergoes meiosis to make four haploid pronuclei. Three of these pronuclei degenerate One pronucleus is exchanged with the other conjugant Two pronuclei (in each Paramecium) fuse to restore diploid state Allows for genetic recombination Ciliate Reproduction Comparison Conjugation is similar to zygote formation in other animals Permits gene recombinations/variation Ciliates in clone cultures can reproduce asexually for long periods, but often usually lose vigor Conjugation restores viability Conjugation often seen with seasonal changes or a deterioration of the environment