BIOA01H3 – Lecture 33rd Chapter 22 22.1 The Evolution of Protists Involved Endosymbiosis Likely evolved 1.5-2bya We don’t understand how exactly they’ve evolved but know that it involves endosymbiosis Protists contain mitochondria and many also contain chloroplasts 22.2 Characteristics of Protists Protists have membrane-bound nucleus, with multiple, linear chromosomes In addition to cytoplasmic organelles, incl. mitochondria and chloroplasts, also have microtubules and microfilaments, providing motility and cytoskeletal support Share characteristics of transcription and translation All organisms in eukaryotic lineages consist of protists except for animals, land plants, and fungi, which arose from protist ancestors Although some protists have characteristics resembling those of fungi, plants, or animals, several features are distinctive in only protists In contrast to fungi, most protists are motile or have motile stages in life cycle and cell walls made of cellulose not chitin Unlike plants, may photoautotrophic protists can also live as heterotrophs & some regularly combine both modes of nutrition Protists don’t retain developing embryos in parental tissue nor have highly differentiated structures such as roots, stems, leaves Photosynthetic protists also sometimes referred to as algae; generally aquatic & often unicellular and microscopic o All animals are multicellular and have features i.e. internal digestive tract & complex developmental stages o Protists also lack nerve cells, highly differentiated structures i.e. limbs o Also lack a heart, collagen, which is an extracellular support protein Most protists don’t share a common ancestor and are no more closely related to each other than they are to fungi, plants, or animals 22.3 Protists’ Diversity Is Reflected in Their Metabolism, Reproduction, Structure, and Habitat Protists are highly diverse in metabolism, reproduction, structure, and habitat 1 Habitat Live in aqueous habitats, incl. aquatic or moist terrestrial locations In bodies of water, small photosynthetic protists called phytoplankton, the organisms that capture energy of sunlight in all aquatic habitats Phototrophs provide organic substances and oxygen for heterotrophic bacteria, other protists, and small crustaceans and animal larvae that are the primary constituents of zooplankton Protists living on terrestrial, play important roles among detritus feeders that recycle matter from organic back to inorganic Structure Most protists single cells, others live in colonies in which individual cells show little or no differentiation Within colonies, individuals use cell signaling to cooperate on tasks i.e. feeding or movements Some protists are larger multicellular organisms; i.e. giant kelp of coastal waters Have complex intracellular structures, some of which are found nowhere else among living organisms These unique structures reflect key aspects of habitats in which protists live i.e. unicellular protists living in freshwater pond o Its cytoplasm hypertonic to water surrounding it, meaning water flows into cell by osmosis o Has specialized cytoplasmic organelle, contractile vacuole, which gradually fills w/ water o When vacuole reaches maximum size, moves to plasma membrane and forcibly contracts Cells of some protists supported by external cell wall or by internal/external shell build up from organic or mineral matter Instead of cell wall, other protists have pellicle, layer of supportive protein fibers located inside cell just under plasma membrane, providing strength and flexibility Sometime during live, almost all protists move Some move by amoeboid motion, in which cells extends one or more lobes of cytoplasm called pseudopodia Rest of cytoplasm and nucleus flow into pseudopodium, completing movement Others move by flagella or cilia Metabolism Almost all protists are aerobic organisms, live either as heterotrophs or photoautotrophs 2 Some heterotrophic protists obtain organic molecules by engulfing part or all of other organisms (phagocytosis) and digesting internally Others absorb small organic molecules from environment by diffusion Some live as both heterotrophs and autotrophs Reproduction May be asexual, by mitosis, or sexual, through meiotic cell division and formation of gametes Those that reproduce by both mitosis and meiosis, two modes of cell division often combined into life cycle that is highly distinctive among different protists group 22.4a Excavates Lack “Typical” Mitochondria Groups takes name from hollow (excavated) ventral feeding groove found in most members All are unicellular animal parasites lacking mitochondria and move by flagella b/c lacking mitochondria, limited production of ATP via glycolysis lack of mitochondria led biologists to consider group as most ancient line of protists; now appears ancestor of group had mitochondria nuclei of Excavates contain genes derived from mitochondria and also have organelles likely evolves from mitochondria may’ve lost mitochondria as adaptation to parasitic way of life, in which oxygen in short supply; consider two subgroups, Diplomonadida and Parabasala o Diplomonadida means double cell and organisms look like two cells together w/ two identical, functional nuclei and multiple flagella arranged symmetrically around cell’s longitudinal axis Best known diplomonad is Giardia lamblia Some are free living, many live in animal intestines; some don’t cause harm to hose, others do o Parabasala STD trichomoniasis caused by parabasalid Trichomonasvaginalis Infection usually symptomless in men, but in women can cause severe inflammation and irritation of vagina and vulva If untreated, can cause infection in uterus and fallopian tubes resulting in infertility Parabasalids take names from cytoplasmic structures associated w/ nucleus, parabasal bodies; some consider these as Golgi apparatus of cells 3 Also characterized by fin called undulating membrane formed by flagella buried in fold of cytoplasm, in addition to freely beating flagella Buried flagellum allows parabasalids to move through thick, viscous fluids 22.4b Discicristates Include the Euglenoids and Kinetoplastids: Highly Motile Protists Sometimes referred to as protozoa (proto = first; zoon = animal) b/c they’re similar to animals in that they ingest food and move by themselves Discicristates named for disk shaped mitochondrial cristae Group includes about 1800 species, almost all unicellular and highly motile, swimming by means of flagella Although most are photosynthetic, some can also live as heterotrophs and some even alternate between photosynthetic and heterotrophs Euglenoids Important primary producers in freshwater ponds, streams, etc. Most are autotrophs that carry out photosynthesis w/ same mechanisms like plants If light not available, many euglenoids also live as heterotrophs Others lack chloroplasts and live entirely as heterotrophs Name Euglena roughly translates as “eyeball organism”, reference to large eyespot which is feature of photosynthetic euglenoids o Contains pigment granules in association w/ light sensitive structure and part of sensory mechanism stimulating cells to move toward moderately bright light or away from intense ones so organisms find optimal photosynthetic conditions Also contains numerous organelles incl. contractile vacuole Rather than external cell wall, have spirally shaped pellicle formed from strips of transparent, protein-rich material underneath membrane Some euglenoids, strips arranged in spiral allowing cell to change shape in wriggling sort of motion that allows cell to change direction Can also swim by flagella; has two; one rudimentary and short, other long Kinetoplastids Sleeping sickness fatal disease, caused by various subspecies of Trypanosoma brucei that are transmitted from one host to another by bites of tsetse fly Early symptoms incl. fever, headaches, rashes, and anemia Untreaed, disease damages central nervous system, leading to sleeplike coma and eventual death 4 Disease proven difficult to control b/c present in wild mammals, providing inexhaustible reservoir for parasite Other kinetoplastids are heterotrophs and are characterized by single mitochondrion that contains large DNA-protein deposit called kinetoplast Most kinetoplastids also have leading and trailing flagellum 22.4c Alveolates Have Complex Cytoplasmic Structures and Move via Flagella or Cilia Groups named for small, membrane-bound vesicles called alveoli in layer just under plasma membrane Includes two motile, primarily free-living groups: Ciliophora and Dinoflagellata, and a nonmotile, parasitic group, Apicomplexa Ciliophora Ciliophora large group, w/ nearly 10k known species of primarily unicellular but highly complex heterotrophic organisms that swim using cilia Some live individually; some colonial Some animal parasites; others live and reproduce in hosts as mutually beneficial symbionts Have many highly developed organelles, incl. mouthlike gullet lined w/ cilia, structures that exude toxins and other defense materials from cell surface, contractile vacuoles, and complex system of food vacuoles Pellicle reinforces cell’s shape Complex cytoskeleton anchors cilia just below pellicle and coordinates ciliary beating Cilia can stop, reverse beating in synchrony, allowing ciliates to stop, back up, and turn if encounter negative stimuli Ciliates are only eukaryotes that have two types of nuclei in each cell: one or more small nuclei called micronuclei and single larger macronucleus Micronucleus is diploid nucleus contains complete complement genes. o Fncs primarily in cellular reproduction, which can be asexual or sexual o Number present depends on species Macronucleus develops from micronucleus but loses all genes except those required for basic functions of cell and for synthesis of ribosomal RNA. o Contains numerous copies of these genes, allowing it to synthesize larger quantities of proteins and rRNA Ciliates abound in freshwater and marine habitats, where they feed on bacteria, algae, and each other 5 Dinoflagellata: The Dinoflagellates Red tides caused by certain dinoflagellates that make up large proportion of marine phytoplankton Typically have shell formed from cellulose plates Beating of flagella, which fit into grooves in plates, makes dinoflagellates spin like a top as they swim Most are unicellular Live as heterotrophs or auto Some lives as symbionts in tissues of other marine organisms i.e. jellyfish Apicomplexa Nonmotile parasites Take name from apical complex, group of organelles at one end of cell which helps cells attach to and invade host cells Absorb nutrients through plasma membranes and lack food vacuoles Can reproduce asexually and sexually forming gametes that fuse to produce cysts As in Giardia, when host organism engulfs cysts, they divide to produce infective cells 22.4d Heterokonts Have Two Dissimilar Flagella at Some Stage of Their Life Cycles Named for two different flagella: one smooth and a second covered w/ bristles in most heterokonts, flagella occur only on reproductive cells heterokonts include Oomycota (water moulds), Bacillariophyta (diatoms), and Chrysophyta (golden algae), and Phaeophyta (brown algae) Oomycota organism causing blight was water mould water moulds not fungi; they’re oomycetes but do share some features w/ fungi like fungi, oomycetes grow as microscopic, nonmotile filaments called hyphae forming network called mycelium also like fungi, heterotrophs, secreting enzymes that digest complex molecules of surrounding organic matter or living tissue into simpler molecules that are small enough to be absorbed into cells other features not similar to fungi are differences in nucleotide sequence, clearly indicating close evolutionary relationships to other heterokonts than fungi live almost exclusively in freshwater lakes, streams, etc. where they’re key decomposers 6 Bacillariophyta: Diatoms unicellular organisms w/ glassy silica shell substances move in and out of cell through patterned perforations in shell photoautotrophs w/ photosynthetic pathways seen in plants abundant in freshwater habitats most are free-living, some are symbionts inside other marine protists asexual reproduction in diatoms occurs by mitosis followed by form of cytoplasmic division Chrysophyta: Golden Algae nearly all are autotrophs and carry out photosynthesis similar to pathway found in plants colour due to brownish carotenoid pigment, fucoxanthin, which masks green colour of chlorophylls most chrysophytes also lives as heterotrophs if not sufficient light important in freshwater habitats and in nanoplankton community of marine phytoplankton composed of huge numbers of extremely small cells most are colonial forms, in which each cell of colony bears pair of flagella have glassy shells but in form of plates or scales Phaeophyta: Brown Algae kelp is a form of brown algae o largest and most complex of all protists o tissues differentiated into leaflike blades, stalklike stipes, and rootlike holdfasts that anchor them to bottom o hollow, gas-filled bladders give buoyancy to stipes and blades and help keep them upright and oriented toward sunlit upper layers of water o stipes of some kelps contain tubelike vessels, similar to vascular elements of plants, which rapidly distribute products of photosynthesis throughout body of alga all phaeophytes are photoautotrophs but not all are as larges as kelps nearly all of species inhabit temperate or cool coastal marine waters like golden algae, phaeophytes contain fucoxanthin, which gives them their characteristic coour cell walls contain cellulose and mucilaginous polysaccharide, alginic acid alginic acid, called algin when extracted, s essentially tasteless substance used to thicken such diverse products as ice cream, salad dressing, etc. life cycles are typically complex and consist of alternating haploid and diploid generations large structures we recognize as kelp and other brown seaweeds are diploid sporophytes called that b/c give rise to haploid spores by meiosis 7 o spores, which are flagellated swimming cells, germinate and divide by mitosis to form independent haploid gametophyte generation o gametophytes give rise to haploid gametes o most gametophytes are multicellular 22.4e Cercozoa Are Amoebas with Filamentous Pseudopods amoeba (amoibe = change) is a descriptive term for unicellular protists that moves by means of pseudopodia several major groups of protists contain amoebas, which are similar form but aren’t closely related cercozoan amoebas produce stiff, filamentous pseudopodia, and many produce hard outer shells called tests two heterotrophic groups of cercozoan amoebas: Radiolaria and Forminifera, and a photosynthetic group, the Chlorarachniophyta Radiolaria marine organisms characterized by glassy internal skeleton and axopods slender, raylike strands of cytoplasm supported internally by long bundles of microtubules glassy skeleton is heavy, so how they keep afloat? Axopods provide buoyancy as do numerous vacuoles and lipid droplets in cytoplasm o Involved in feeding as well: pretty stick to axopods and then engulfed, brought into cell, and digested in food vacuoles Skeletons that accumulate on ocean floor become part of sediment which turns into sedimentary rock Foraminifera: Forams Organisms take name from perforations in shells through which extend long, slender strands of cytoplasm supported internally by network of needlelike spines Shells consists of organic matter reinforced by calcium carbonate Most foram shells are chambered, spiral structures Forams live in marine environments (like radiolarians) Some species planktonic but they’re most abundant on sandy bottoms and attached to rocks along coasts Forams feed in manners similar to that of radiolarians o Engulfs prey that adhere to strands, conduct them through perforation in shell into central cytoplasm, where digested in food vacuoles Some have algal symbionts that carry out photosynthesis, allowing to live as heterotrophs and autotrophs 8 Chlorarachniophyta Are amoebas that contain chloroplasts and thus are photosynthetic Combine this mode of nutrition w/ heterotrophy, engulfing food w/ many filamentous pseudopodia 22.4f Amoebozoa includes Slime Moulds and Most Amoebas Includes most of amoebas (others are in Cercozoa) as well as cellular and plasmodial slime moulds Use pseudopods for locomotion and feeding Amoebas Unicellular organisms that are abundant in marine and freshwater environments and in soil Some are parasitic Most are heterotrophs that feed on bacteria, other protists, and bits of organic matter Unlike stiff, supported pseudopodia of cercozoans, pseudopods of amoebas extend and retract at any point on body surface and are unsupported by any internal cellular organization – amoebas are thus “shaep-shifters” How can amoeba capture a fast-moving organism? o As amoeba moves, its cytoplasm doesn’t just move but also changes state, from more liquid state to more solid state and back again, allowing amoeba to send out pseudopodia in different directions very quickly o These fast-moving pseudopods can capture even fast-swimming prey such as ciliates Reproduce only asexually via binary fission Unfavourable environmental conditions, some amoebas can form cyst, essentially by rolling up and secreting protective membrane Survive as cysts until favourable conditions return Slime Moulds Unusual heterotrophic protists Exist for part of their lives as individuals that move by amoeboid motion but come together in coordinated mass that ultimately differentiates into stalked structure called fruiting body in which spores are formed Two major evolutionary lineages of slime moulds: cellular slime moulds and plasmodial slime moulds, which differ in cellular organization Both have ability to differentiate into fruiting bodies w/ stalks and sporebearing structures 9 This differentiation much simpler than complex developmental pathways of other eukaryotes, providing unique opportunity to study cell differentiation at its most fundamental level Respond to stimuli in environment, moving away from bright light and toward food Live on moist, rotting plant material Cells engulf particles of dead organic matter, along w/ bacteria, yeasts, and other microorganisms, and digest them internally Can be range ofcolours: brown, yellow, green, red, and even violet or blue Organisms exist primarily as individual cells, either separately or as coordinated mass Reproduces asexually and sexually Plasmodial Slime Moulds Exists primarily as multinucleate plasmodium, in which individual nuclei suspended in common cytoplasm surrounded by single plasma membrane o Plasmodium flows and feeds by phagocytosis like single huge amoeba – single cell that contains thousands to millions or even billions of diploid nuclei surrounded by single plasma membrane o Plasmodium, which may range in size, typically moves in thick, branching strands connected by thin sheets o Movements occur by cytoplasmic streaming, driven by actin microfilaments and myosin Often in response to unfavourable environmental conditions, fruiting bodies form on plasmodium o At tips of fruiting bodies, nuclei become enclosed in separate cells o Cells undergo meiosis, forming haploid, resistant spores that are released from fruiting bodies and carried by water o wind If favourable conditions reached, continue to reproduce 22.4g Archaeplastida Include the Red and Green Algae and Land Plants Group consist of red and green algae, which are protists, and land plants, which make up kingdom Plantae The three groups of photoautotrophs share common evolutionary origin Rhodophyta: The Red Algae Small marine seaweeds Few species found in freshwater lakes and streams, or in soils Cell walls contain cellulose and mucilaginous pectins that give red algae slippery texture o Pectin widely used in industry and science 10 Extracted agar is used as culture medium in laboratory and as setting agent for jellies and deserts Some species secrete calcium carbonate into their cell walls; these coralline algae are important in building coral reefs Red algae typically multicellular w/ diverse morphologies; most have plantlike bodies composed of stalks bearing leaf-like blades Most are free-living autotrophs, some are parasites that attach to other algae or plants Most are reddish in colour, some are greenish purple or black Have complex reproductive cycles involving alternation between diploid sporophytes and haploid gametophytes No flagellated cells occur in red algae; instead, gametes released into water to be brought together by random collisions in currents Chlorophyta: The Green Algae Carries out photosynthesis using same pigments as plants o Shared pigment composition is one line of evidence that one lineage of green algae was ancestor of land plants Green algae shows more diversity than any other algal group Also have diverse morphologies, incl. unicellular, colonial, and multicellular species Multicellular forms have range of morphologies, incl. filamentous, tubular, and leaf-like forms Most live in freshwater aquatic habitats, but some are marine, or live on rocks, soil surfaces, tree bark, or snow Other organisms rely on green algae to photosynthesize for them by forming symbiotic relationships Life cycles are diverse as body forms; many reproduce sexually or asexually and some alternate between haploid and diploid generations Green algae most closely related to land plants, based on molecular, biochemical, and morphological data Which green alga might’ve been ancestor of modern land plants? o Evidence points to group known as charophytes as being most similar to algal ancestors of land plants 22.4h Opisthokonts Include the Choanoflagellates, Which May Be the Ancestors of Animals Named for single, posterior flagellum found at some stage in life cycle of these organisms Diverse group includes choanoflagellates, protists thought to be ancestors of fungi and animals Choanoflagellata (choanos = collar) named for collar surrounding flagellum that protist uses to feed and in, some species, to swim. 11 o Collar resembles upside down lampshade and is made up of small, fingerlike projections (microvilli) of plasma membrane o As flagellum moves water through collar, projections engulf bacteria and particles of organic matter in water Live in either marine or freshwater habitats Some are mobile but most are sessile Number of species are colonial w/ cluster of cells on single stalk 22.5 Some Protist Groups Arose from Primary Endosymbiosis and Others from Secondary Endosymbiosis Endosymbiosis involving photoautotrophs happened more than once, resulting in formation of wide range of photosynthetic eukaryotes First chloroplasts evolves from free-living photosynthetic prokaryotic organisms (cyanobacteria) ingested by eukaryote cells that had already acquired mitochondria In some cells, cyanobacterium not digested but instead formed symbiotic relationship w/ engulfing host cell – became an endosymbiont independent organism ling inside another organism Over evolutionary time, prokaryotic organisms lost genes no longer required for independent existence and transferred most of its genes to host’s nuclear genome In this process, endosymbiont became an organelle; moving genes from enosymbiont to nucleus would’ve given host cell better control of cell functioning Chloroplasts of red and green algae and land plants result from evolutionary divergence of photosynthetic eukaryotes formed from this primary endosymbiotic event Organisms originated from this event have chloroplasts w/ two membranes, one from plasma membrane of engulfing eukaryote and other from plasma membrane of cyanobacterium Primary endosymbiosis followed by at least three secondary endosymbiosis events, each time involving diff. heterotrophic eukaryotes engulfing photosynthetic eukaryote, producing new lineages 12
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