PB295E, Topics in Plant Evolution, emphasizes evolution and evolutionary mechanisms in Angiosperms. Because Paleobotany and Systematics courses are already offered at NCSU, PB295E emphasizes areas not usually contained in those two approaches to the study of Evolution in plants. The course will involve 2 class meetings per week, reading assignments and two exams. Pertinent videos will be shown during class meetings. Assignments will include readings from a Course Pac of selected articles and the book, Plant Variation and Evolution by D. Briggs and S. M. Walters, Cambridge Univ. Press. Detail of TOPICS included in PB295E, 'Topics in Plant Evolution': (1) What is Evolution? Darwin's Theory of Natural Selection Darwin's dilemma Modern synthesis Evidences of evolution (a) Artificial Selective Breeding (b) Comparative Anatomy (c) Fossil Evidence (d) Molecular (DNA) and (e) Developmental Biology Tempos of evolution: (a) Gradualism (b) Punctuated Equilibrium (2) Plant - Green Algae Connection: Plate Tectonics Early Earth Character indicators of algal/plant common ancestry Proposed mechanism for the Evolution of the Algal Zygotic meiosis life cycle to the Plant Sporic meiosis life cycle (3) Sources of the Variation necessary for evolution to occur and how a species' variation potential is a function of its Breeding System: Sources of the Variation: (a) Mutations (b) New sexual recombinations (c) Introgression Breeding Systems: (a) Out-crossing (b) Selfing and (c) Apomixis (vegetative & agamospermy) Adaptations promoting/preventing (a) and (b) above {(1) self-incompatibility genes (Gametophytic and Sporophytic) (2) Heterostyly (3) Gynodioecy (4) Temporal dioecy} (4) Types of Variation and the type that fuels evolution: Genetic vs. Environmental variation {Turesson, Clausen, Keck and Hiesey} Common Garden Experiments Ecotype concept and Challenge to Ecotype concept i.e., Clines Methods of studying presence/absence/degree of genetic variation in plant populations: (a) Isozymes (b) Chromatography (c) RFLP (d) RAPD (e) DNA Sequencing (chloroplast, mitochondria and nuclear) (5) Hardy-Weinberg Equilibrium: Population Genetics definition of Evolution Hardy-Weinberg Equilibrium - (a) the criteria necessary for no evolution to occur in a population (b) Derivation of H-W Equilibrium equation, Sample case studies illustrating usefulness of H-W (6) Deviations from Hardy-Weinberg i.e., why evolution DOES occur in natural populations: Case Studies of real plant populations illustrating reality of (a) mutations (b) gene flow (c) non-random mating (d) chance events {Genetic drift e.g., founder effects, bottlenecks} (e) natural selection {examples of stabilizing selection, directional selection, disruptive selection} (7) Species and Speciation: Species (a) definitions (b) The Species Problem (c) Incipient species (d) Species delimitation Speciation modes - Allopatric, Sympatric Examples of Pre and Post Zygotic isolating mechanisms in plants: (a) Temporal Isolation (b) Floral Isolation (c) Mechanical Isolation (d) Ecological Isolation Results of isolating Mechanisms Case Studies of Allopatric Speciation in Plants including Adaptive Radiation (8) Hybridization: Hybridization - definition Hybridization in nature: (a) Role of habitat disturbance (both nature and man caused) in hybridization (b) Hybrid Zones (c) Hybrid Swarms (d) Are all hybrids new species? (e) Views on hybrid origins Examples of Various Possible Consequences of Hybridization: (a) fully fertile hybrid (b) semi-fertile hybrid (c) asexually reproducing sterile hybrid (d) polyploidy (e) hybrid inviability Introgression - (a) definition (b) methods of detection (c) evolutionary consequences Two detailed case studies - Natural Hybridization and Introgression in the Louisiana Irises (9) Polyploidy: Polyploidy - (a) frequency in plants? (b) base numbers (c) ancient polyploid series Polyploid Detection Studies (a) Karyotypes (b) Isozymes (c) Chromatography (d) Nuclear DNA content (e) Colchicine Polyploidy Types - Allopolyploidy, Autopolyploidy, Segmental Polyploidy Case studies of polyploidy including Tragopogon, and Triticum turgidum (bread wheat) Polyploid Origins - 2 mechanisms (a) Somatic Doubling (b) Unreduced Gametes Mechanisms of Meiotic Dysfunction Persistence of Polyploids (10) Flower and Fruit Evolution: Appearance of Angiosperms Plate Tectonics - continent positions at time of Angiosperm Appearance Angiosperm Ancestry Unique Angiosperm Characteristics Old and New views of characteristics of earliest flower Evolutionary Trends of Flowers - (a) ovary position (b) floral symmetry Agents of floral evolution - pollinators Evolution of floral characters in response to selective pressure of attracting pollinators: (a) pigments (Carotenoids, Flavonoids, and Betacyanins) (b) flower shape (c) fragrances (d) floral markings, (e) nectar production Evolution of fruit characters in relation to the selective pressure of promoting fruit dispersal: (a) wind, (b) water (c) self projected and (d) animal dispersed fruit characters (11) Evolution of Secondary Metabolites - Evolution of chemical compounds in response to the selective pressures of herbivores and pathogens Alkaloids - examples of plants having evolved (a) Morphine (b) Cocaine (c) Nicotine (d) Caffeine Terpenoids - examples of plants having evolved (a) Isoprene (b) Essential oils (c) Rubber (d) Cardiac Glycosides Phenolics - examples of plants having evolved (a) Tannins (b) Salicylic Acid (c) Lignins (12) Detailed Examination of Several Case Studies of Co-evolution - identifying the selective pressures exerted by both participants Dodo bird and Sideroxylon grandiflorum tree on Mauritius- fruit and fruit disperser FACT or FALACY - a controversy Ants and Acacia trees in Central America Mistletoe and the Mistletoe birds in Australia Yucca plants and the Yucca moth in USA Sycamore Fig and Fig Wasps in Africa Goldenrod and the Gallfly in USA (13) ABC Model of Flower Development - Genetics of Floral Evolution Model research organism - Arabidopsis thaliana Stages in floral development Developmental fields of Gene Expression 5 genes specifying floral organ identity in Arabidopsis Loss of gene function: (a) Loss of A function (b) loss of B function (c) loss of C function LEAFY gene and mutation of LEAFY E function genes Unanswered questions (14) Evolution instigated by man's activities Heavy metal pollution and evolution of resistant plants Evolution for herbicide resistance in crop plants Genetically modified organisms Habitat disturbance favoring evolution by hybridization Alien Introductions and evolution by hybridization/ introgression (15) Evolutionary Stasis: Three case studies: (a) Wollemi pine (b) Ginkgo biloba (c) Metasequoia Characteristics associated with evolutionary stasis within a plant group