1. Signaling

2. Signaling (hormones, light, etc…) Reception Transduction Response Signal No response Receptor Relay proteins Signal Receptor Differential gene expression

3. Major signals that control plant growth and development • Environmental signals: -Light - Gravity - Temperature - Humidity - etc…

4. Major signals that control plant growth and development • Internal signals: Plant Hormones -AUXIN - CYTOKININ - ETHYLENE - ABSCISIC ACID - GIBBERELLIC ACID

5. Auxin Note: several different auxins are known to date (natural as well as synthetic). IAA is the most common natural auxin found in plants.

6. Auxin effects - promotes cell elongation - inhibits lateral meristem activity - promotes root formation

7. Auxin and differential growth:Gravitropic growth responses of Arabidopsis seedlings Cotyledons (embryonic leaves) Turn seedling 90o Hypocotyl (embryonic stem) Root Hypocotyl shows a negative gravitropic response Root shows a positive gravitropic response Areas of differential growth (one side grows faster than the other)

8. Differential growth a Rate of cell elongation is higher on the a-side of the coleoptile compared to the b-side. This leads to differential growth: increased growth rate on one side of plant organ, results in curvature of the organ. b

9. Auxin and shoot apical dominance • Decapitation of the apical bud releases the lateral buds. In the absence of auxin coming from the shoot apex, lateral buds become active leading to branching (and a more bushy shoot development)

10. Example: Auxin and lateral root formation in Arabidopsis The synthetic auxin 2,4-D promotes lateral root formation in Arabidopsis Note: 2,4-D is also used as a herbicide because it completely inhibits growth at higher concentrations.

11. Example: Auxin promotes adventitious root formation from Ilex opaca (Holly) shoots. Fig. 15-12, p. 246 Shoots form roots at their bases faster when the bases are treated with auxin. The ends of these shoots were dipped for 5 seconds in solutions containing (from left to right) 0%, 0.1% and 0.5% auxin. They were then rooted in moist vermiculite for 2 weeks.

12. Cytokinin Zeatin Zeatin is one of many natural cytokinins found in plants

13. Cytokinin effects - promotes cell division/shoot formation - promotes lateral meristem activity - controls sink/source identity of plant organs - delays senescence

14. auxin cytokinin

15. Cytokinin and shoot apical dominance • By increasing the cytokinin concentration in the shoot, lateral buds become active resulting in increased branching (and a more bushy shoot development) Cytokinin

16. The effect of cytokinin on senescence. Cytokinin applied to the right-hand primary leaf of this bean seedling inhibited its senescence. The left-hand did not get cytokinin. Fig. 15-13, p. 246

17. Gibberellin Gibberellic acid 3 Note: several different gibberellins are known to date (natural as well as synthetic). GA3 is the most common natural gibberellin found in plants.

18. Gibberellin effects - promotes stem elongation growth - promotes seed germination

19. Gibberellins promote stem elongation in many plant species Pea seedlings Pea seedlings treated with GA3

20. Gibberellins and world food production • Norman Borlaug • Nobel Peace Prize 1970 • Developed high-yielding wheat strains • Disadvantages • Strains require high levels of fertilizer (containing N, see lecture on absorption and transport of minerals) • Expensive (requires fossil fuels) • Create pollution

21. Coordination of Development via Hormone action • The major plant hormones: - Auxins - Cytokinins - Gibberellins - Abscisic acid - Ethylene Hormones that promote/control growth (direction) Survival hormones (tend to inhibit growth)

22. Ethylene

23. Ethylene effects - inhibits cell expansion - accelerates senescence - accelerates fruit ripening

24. Ethylene effects on etiolated seedlings Arabidopsis seedlings grown in the dark display an etiolated growth pattern: 1) unexpanded cotyledons 2) Apical hook 3) long thin hypocotyl Exposure to ethylene during growth in the dark results in: 1) Exagerated apical hook curvature 2) Much shorter and thicker hypocotyl

25. Ethylene and senescence Solution that contains STS, an inhibitor of ethylene action. STS delays floral senescence.

26. Ethylene and fruit ripening • Ripening of fruit stimulated by ethylene • Ethylene is THE most damaging hormone in agriculture (accelerates ripening and consequently rotting of fruits) • Involves • Conversion of starch or organic acids to sugars • Softening of cell walls to form a fleshy fruit • Rupturing of cell membrane with resulting loss of cell fluid to form dry fruit • Overripe fruit is potent source of ethylene • Promotes ripening of adjacent fruits

27. Abscisic acid

28. Abscisic acid effects - promotes stomatal closure - inhibits seed germination

29. Abscisic Acid and drought stress Abscisic acid is a signal of this emergency situation. Under drought conditions, wilted mesophyll cells of a leaf rapidly synthesize and excrete abscisic acid (ABA). This ABA diffuses to the guard cells, where an ABA receptor recognizes the presence of the hormone and acts to release K+, Cl-, and as a result H2O, thus rapidly reducing turgor pressure and closing the stomata

30. Abscisic Acid and germination Wild type (normal) Corn seeds attached . Majority of seeds are dormant: they contain ABA that prevents germination. ABA insensitive corn. Majority of seeds are already germinating while still attached to the parent plant because of a defect in ABA sensitivity.