Plant Responses to Signals Phytohormones (cont.)

1. Plant Responses to Signals Phytohormones (cont.) It might seem unfair to reward a person for having so much pleasure over the years, asking the maize plant to solve specific problems and then watching its responses. Barbara McClintock

2. Plant Responses to Signals III Cytokinin “cell division”

3. control explant vascular tissue Cell Division Factorsdiffusable Haberlandt (1913) showed that vascular tissue could induce cell division in quiescent tissue. potato explants cell division

4. Cytokinins • Van Overbeek discovered that coconut milk stimulated the growth (cell division) of embryos, • Carlos Miller, working in Folke Skoog’s lab in Wisconsin, systematically tried every random chemical he could get his hands on, • Autoclaved DNA (degraded) did the trick. Zeatin

5. Cytokinin Biosynthesis • Plants and bacteria make cytokinins, • cytokinin is synthesized in the root, and transported acropetally via the xylem, • bacteria infect plants, make cytokinin and cause tumors.. Don’t memorize this pathway.

6. roots [ auxin ] 0 [ cytokinin ] shoots Cytokinin / Auxin Balance Undifferentiated plant tissue (callus) can be induced to make shoots and roots when given proper levels of cytokinin and auxin.

7. Shoot (auxin production) is reduced in relation to root (cytokinin production) Shoot / Root Balanced Root (cytokinin production) is reduced in relation to Shoot (auxin production) Control of Organogenesissimplification What happens? What happens?

8. high cytokinin normal cytokinin Cytokinins …retard senescence , • Senescence, • an active, genetically controlled, developmental process, • in which cellular structures and macromolecules are broken down • and transported to growing organs. overexpressor wt

9. Concept Map Receptor/Sygnal Transduction Discovery Synthesis (where) Cytokinin Function(s) Transport auxin interaction

10. Plant Responses to Signals III Giberellins “Mendel’s dwarf”

11. Bakanae“foolish seedlings” • Rice farmers in Asia have long known of a disease which makes the rice plant grow tall, but eliminates seed production, • plants are prone to “lodging” (falling over), • Shotaro Hori (1898) demonstrated that the symptoms were induced by infection with a fungus belonging to the genus Fusarium, • Eiichi Kurosawa (1926) showed that filtrate from culture of Gibberellin fugikuroi, was an active agent, • Margaret Radley (1950s) demonstrated that Giberellin was synthesized by plant tissues. > 100 forms of gibberellin acid (GA), … 30 are active.

12. a-amylase, …hydrolyzes starch, resulting sugars nourish growing seedling. • embryo, • ...absorbs H2O, • gibberellins, • …diffuse to the aleurone layer, • GA signal transduction, • …results in the expression of a-amylase, Gibberellin …contributes to mobilization of storage products,

13. Synthesis and Transport • Gibberelins and intermediates are synthesized in young, actively growing buds and leaves, • long distance transport via the phloem, • short distance via symplastic routes, • Some synthesis in the roots, • transported acropetally via the xylem sap. Meristem and young leaves reporter gene, …firefly luciferase. GA gene promoter, expressing firefly “glow” gene.

14. dwarf pea (mutant) dwarf pea (mutant) + GA Gibberellin …stimulates cell elongation and cell division. [ GA ] rice leaf sheath bioassay

15. GA Cabbage induced to flower by GA alone. Gibberellin …contibutes to “phase changes”, • ...stimulates “bolting” in rosette habit plants, • …involved in juvenile to adult, and adult to reproductive phase changes, • may be “florigen” the flowering hormone in some species, • ... activates vegetative growth of the embryo, • weakens structures in the seed, • mobilizes stored food reserves.

16. Gibberellin …contributes to fruit development, untreated animal treated • …larger grapes, • ... longer stems, • - healthier bunches. Thompson seedless grapes

17. Concept Map Discovery Gibberellin Function(s) Transport

18. Plant Responses to Signals III Abscisic Acid “stress hormone”

19. Dormin, Abscisin II • In 1963, abscisic acid was first identified and characterized by Frederick Addicott • through biochemical analysis of cotton fruit abscission (Abscisin II), • Concurrently, a group headed by Philip Wareing was studying bud dormancy in Sycamore trees (Dormin), … Plant physiologists agreed to call the compound abscisic acid. Abscisic acid (ABA) …isomers and enantiomers occur naturally and synthetically.

20. Synthesis and Transport • ABA is synthesized in the chloroplasts, • ABA is transported in the vascular tissue, • from leaves to roots, through the phloem, • from roots to leaves through xylem. dry roots, ...send stress signals to the leaves (ABA). Result: well hydrated leaves, closed stomates.

21. Abscisic Acid…induces guard cell closure, • ABA concentration in well-watered xylem sap from sunflowers is ~ 1 - 5 nM, • ABA concentration in water stress xylem sap is as much as 3000 nM, • ABA is synthesized (or accumulates) in the roots, is mobilized for transport to the leaves.

22. Abscisic Acid…inhibits germination, • ABA (at high concentrations) in seeds inhibits germination, • ABA / GA balance often determines “internal” dormancy status, • ABA is water soluble, imbibition may serve to leach ABA from dormant seeds, • imbibition: the uptake of water by germinating seeds. ABA deficient seed

23. Concept Map Discovery Synthesis Abscisic Acid Function(s) Transport GA interaction

24. Plant Responses to Signals III Ethylene “the gaseous hormone”

25. ethylene Gas light • Egyptians gassed figs in order to stimulate ripening, • The ancient Chinese burned incense in closed rooms to enhance the ripening of pears. • In 1864, gas leaks from street lights were observed to stunt plant growth, twist plants, and abnormally thicken stems. • Dimitry Neljubow (1901) showed that the active component was ethylene. • R. Gane (1934) reported that plants synthesize ethylene.

26. Ethylene…promotes fruit ripening, • Ethylene signals the transition from unripe to ripe fruits, • cell wall components are broken down, • starches and acids are broken down resulting in “sweetening” and aromatic compounds , • pigmentation may also be induced.

27. Ethylene…promotes the “triple response”, …in etiolated seedlings, • reduced stem elongation, • thicker stem, • horizontal growth, • May provide the plant with “behavior” that will provide escape from soil impediments.

28. ctr • ein (ethylene present), • …ethylene insensitive. • ctr (ethylene absent), • …constitutive triple response. Ethylene…mutant analysis, wild type ein wild type

29. twig petiole Ethylene…contributes to apoptosis and abscission, apoptosis, …programmed cell death abscision layer, …large, thin walled cells. cork ...suberized protective layer

30. Concept Map Discovery Ethylene Function(s) Triple Response

31. Plant Responses to Signals III Brassinosteroids “steroid hormone”

32. insects plants mammalian sex hormones Steroid Hormones

33. dark light wt det2 wt det2 dwarfs, …in light. de-etiolation, ...constitutive, Brassinosteroids…de-etiolation factors and auxin-like functions,

34. Concept Map the bare essentials... Table 39.1 Brassinosteroids