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Origin of Eukaryotes Prokaryotes Eukaryotes • No true nucleus • No plastids • True nucleus • Specialized plastids • Internal membrane systems are folds of plasma membrane • Internal membrane systems independent of plasma membrane Trends in Increased Prokaryote Complexity • Multicellular prokaryotes with specialized cells • Complex bacterial communities • Compartmentalization of different functions within single cells Heterocyst of Anabaena Trends in Increased Prokaryote Complexity • Multicellular prokaryotes with specialized cells • Complex bacterial communities • Compartmentalization of different functions within single cells • These trends are important because eukaryotes had to evolve from prokaryotes Origins of Eukaryotes • Earliest evidence 1.5 billion years • Acritarchs resemble cysts produced by living autotrophic protists • Development of oxygen atmosphere Electron micrograph of an acritarch. Models Proposed for the Evolution of Eukaryotes Autogenous Model - eukaryotic cells evolved from specialization of internal membranes derived from plasma membrane of prokaryotes Autogenous Model • Single-membranes organelles formed by folding of inner membrane only • Double-walled organelles by complete invagination Autogenous Model Endosymbiotic Model Predecessors of eukaryotes where symbionts, with small specialized species (endosymbionts) living within larger prokaryotes Fig. 22.12b Endosymbiotic Model Model uses chloroplasts and mitochondria as examples • Chloroplasts were photosynthesizing prokaryotes • Mitochondria evolved from aerobic heterotrophs (emphasis on role of Krebs cycle) Chloroplast Mitochondrion The Model was Controversial The endosymbiotic model differs from evolution as we discussed earlier It is a merger of evolutionary lineages giving rise to a new form of life But, Supporting evidence has strengthened validity e.g., mitochondrial and chloroplast DNA Also, symbiosis is a common phenomenon in nature Kingdom Protista • First eukaryotic organisms • Typically thought of as the unicellular eukaryotes • Some colonial and multicellular species • Addition of multicellular forms justified by similarities in cell structure and life cycles Unicellular, but Complex • Genesis of protists reveals rise of – – – – – true nucleus specialized organelles 9 + 2 flagella and cilia mitosis meiosis • Share common ancestry with multicellular eukaryotes Protistan Systematics • As would be expected, it is difficult to develop phylogenetic relationships among the protistans – Poor fossils (except those with external covering – Some features (e.g., flagella and autotrophy) arose and have been lost more than once over their evolution • Phyla are placed into supergroups (Table 28.1) Protistan Systematics • Evolutionary Relationships – Cellular structure – Gene sequences • Evolutionary relationships constantly changing • Systematics • Relationships into supergroups Informal Classification – Ecological Roles Protozoa – animal-like heterotrophic Protista Ciliate consuming diatoms Algae – autotrophic protists Informal Classification – Ecological Roles Fungus-like protists Informal Classification – Motility Ciliates Flagellates Informal Classification – Motility Amoeboid Life Processes?? Although unicellular, protistans can carry out all life processes Osmoregulation – Water Balance Vacuoles increase effective surface area in large cells. Contractile vacuoles in freshwater microbial eukaryotes such as Paramecium are used to excrete excess water. Figure 27.10 Contractile Vacuoles Bail Out Excess Water Nutrition • • • • Phagotrophy Osmotrophy Autotrophy Mixotrophy Defense • • • • Mucilage Trichocysts Bioluminescence Toxins 27.3 How Did the Microbial Eukaryotes Diversify? Food vacuoles are formed by protists when solid food particles are ingested by endocytosis. The food is digested in the vacuole. Smaller vesicles pinch off—increasing surface area for products of digestion to be absorbed by the rest of the cell. Cell surfaces Many microbial eukaryotes have diverse means of strengthening their surfaces. Cytoskeleton Internal structures that provide support and rigidity 27.3 How Did the Microbial Eukaryotes Diversify? Some amoebas make a “shell” or test from bits of sand beneath the plasma membrane. Diatoms form glassy cell walls of silica. These walls are exceptionally strong, and perhaps enhanced defense against predators. Frustule Figure 27.12 Cell Surfaces in the Microbial Eukaryotes Asexual Reproduction • All protists can reproduce asexually • Many produce cysts with thick, protective walls that remain dormant in bad conditions • Many protozoan pathogens spread from one host to another via cysts 34 Sexual Reproduction • Eukaryotic sexual reproduction with gametes and zygotes arose among the protists • Generally adaptive because it produces diverse genotypes • Zygotic and sporic life cycles 35 • Zygotic life cycles Most unicellular sexually reproducing protists Haploid cells transform into gametes + and – mating strains Thick-walled diploid zygotes Survive like cysts 36 37 • Sporic life cycle Many multicellular green and brown seaweeds Also known as alternation of generations 2 types of multicellular organisms Haploid gametophyte produces gametes Diploid sporophyte produces spores by meiosis Red seaweed variation involves 3 distinct multicellular generations 38 39 40 • Gametic life cycle All cells except the gametes are diploid Gametes produced by meiosis Diatoms Asexual reproduction reduces the size of the daughter cells Sexual reproduction restores maximal size 41 42 • Ciliate sexual reproduction – Conjugation Most complex sexual process in protists Have 2 types of nuclei (single macronucleus and one or more micronuclei) Macronuclei are the source of the information for cell function 2 cells pair and fuse – conjugation Micronuclei undergo meiosis, exchange, fusion and mitosis 43 44 Phylum Chlorophyta • Green algae, diverse group • unicellular, aggregates, colonial, multicellular • Believed to be the group that gave rise to plants – multicellular and colonial forms – alternation of generation Origins of Multicellularity • Probably arose from colonial protistan – something resembling Volvox – (Volvox is only an example!!!!!!) • Coordination and cooperation between cells • Specialized reproductive cells – Volvox - locomotion and reproduction • Ancestor probably flagellated Alternation of Generation • Life cycles that show an alternation between a multicellular haploid form and a multicellular diploid form • Sporophyte - Diploid; produces reproductive cells (haploid) called spores • Gametophyte - Haploid; produces haploid gametes. Fusion of gametes produces diploid form Fig. 28.20 Thus, the presence of alternation of generation and other similarities suggests a linkage between the Chlorophyta and the Plant Kingdom