1. Chapter 39 Plant Responses

2. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings I. Plant hormones Chemical signals that coordinate different parts of an organism. Tropism = Any response resulting in curvature of organs toward or away from a stimulus

3. What part of a grass coleoptile senses light, and how is the signal transmitted? RESULTS Control Light Illuminated side of coleoptile Shaded side of coleoptile

4. RESULTS Light Tip removed Phototropic response only when tip is illuminated Tip covered by opaque cap Tip covered by trans- parent cap Site of curvature covered by opaque shield

5. RESULTS Light Boysen-Jensen: phototropic response when tip is separated by permeable barrier, but not with impermeable barrier Tip separated by gelatin (permeable) Tip separated by mica (impermeable)

6. Question: Does asymmetric distribution of a growth- promoting chemical cause a coleoptile to grow toward the light? Excised tip placed on agar cube RESULTS Growth-promoting chemical diffuses into agar cube Agar cube with chemical stimulates growth Offset cubes cause curvature Control (agar cube lacking chemical) has no effect Control

7. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hormones control plant growth and development by affecting the division, elongation, and differentiation of cells.

8. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings A. Auxin Any chemical that promotes elongation of coleoptiles. Indoleacetic acid (IAA) is a common auxin in plants Transporter proteins move the hormone from the basal end of one cell into the apical end of the next cell.

9. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Role of Auxin in Cell Elongation Auxin stimulates proton pumps in the plasma membrane. The proton pumps H+ lower the pH in the cell wall, activating expansins, enzymes that loosen the cell wall’s fabric. With the cellulose loosened, the cell can elongate.

10. Cell elongation in response to auxin : acid growth hypothesis Cross-linking polysaccharides Cellulose microfibril Cell wall becomes more acidic. 2 1 Auxin increases proton pump activity. Cell wall–loosening enzymes Expansin Expansins separate microfibrils from cross- linking polysaccharides. 3 4 5 CELL WALL Cleaving allows microfibrils to slide. CYTOPLASM Plasma membrane H2OH2O Cell wall Plasma membrane Nucleus Cytoplasm Vacuole Cell can elongate.

11. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings B. Cytokinins Control of Cell Division and Differentiation Produced in actively growing tissues (roots, embryos, and fruits)

12. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Control of Apical Dominance Cytokinins, auxin, and other factors interact in the control of apical dominance (terminal bud’s ability to suppress axillary buds) If the terminal bud is removed, plants become bushier. Anti-Aging Effects Slow the aging of some plant organs

13. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings C. Gibberellins Stem Elongation Stimulate growth of leaves and stems. Stimulate cell elongation and cell division.

14. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fruit Growth In many plants, both auxin and gibberellins must be present for fruit to set. Germination release of gibberellins from the embryo signals seeds to germinate.

15. Effects of gibberellins on stem elongation and fruit growth (a)Gibberellin-induced stem growth (b) Gibberellin-induced fruit growth

16. Mobilization of nutrients by gibberellins during the germination of seeds such as barley Gibberellins (GA) send signal to aleurone. Aleurone secretes  -amylase and other enzymes. Sugars and other nutrients are consumed. Aleurone Endosperm Water Scutellum (cotyledon) Radicle 1 2 3 GA  -amylase Sugar

17. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Seed Dormancy Ensures that the seed will germinate only in optimal conditions. – In some seeds, dormancy is broken when ABA is removed by heavy rain, light, or prolonged cold. Drought Tolerance Primary internal signal that enables plants to withstand drought. D. Abscisic Acid

18. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings E. Ethylene Produced in response to stresses such as drought, flooding, mechanical pressure, injury, and infection. Fruit Ripening A burst of ethylene production in a fruit triggers the ripening process.

19. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Triple Response to Mechanical Stress The triple response consists of a slowing of stem elongation, a thickening of the stem, and horizontal growth.

20. ethylene-induced triple response Ethylene concentration (parts per million) 0.10 0.00 0.20 0.40 0.80

21. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Senescence Programmed death of plant cells or organs. A burst of ethylene is leads to apoptosis Leaf Abscission A change in the balance of auxin and ethylene controls leaf abscission (in autumn when a leaf falls)

22. Abscission of a maple leaf 0.5 mm Protective layer Stem Abscission layer Petiole

23. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings II. Circadian Rhythms (Biological Clocks) Many plant processes oscillate during the day. – Many legumes lower their leaves in the evening and raise them in the morning, even when kept under constant light or dark conditions.

24. Sleep movements of a bean plant Noon Midnight

25. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Circadian rhythms are cycles that are about 24 hours long and are governed by an internal “clock.” Phytochrome conversion marks sunrise and sunset, providing the biological clock with environmental cues.

26. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings III. Photoperiodism Photoperiod (the relative lengths of night and day) is the stimulus plants use most often to detect the time of year. Photoperiodism is a physiological response to photoperiod.

27. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings A. Control of Flowering Plants that flower when a light period is shorter than a critical length are called short-day plants. Plants that flower when a light period is longer than a certain number of hours are called long- day plants. Flowering in day-neutral plants is controlled by plant maturity, not photoperiod.

28. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Critical Night Length Flowering and other responses to photoperiod are actually controlled by night length, not day length. Short-day plants are governed by whether the critical night length sets a minimum number of hours of darkness. Long-day plants are governed by whether the critical night length sets a maximum number of hours of darkness.

29. Photoperiodic control of flowering 24 hours Light Critical dark period Flash of light Darkness (a) Short-day (long-night) plant Flash of light (b) Long-day (short-night) plant

30. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Red light can interrupt the nighttime portion of the photoperiod. Action spectra and photoreversibility experiments show that phytochrome is the pigment that receives red light.

31. Reversible effects of red and far-red light on photoperiodic response. 24 hours R RFR RFRR RFRRFR Critical dark period Short-day (long-night) plant Long-day (short-night) plant

32. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings IV. Tropisms Gravitropism = Response to gravity – Roots = positive; shoots = negative – Plants may detect gravity by the settling of statoliths Thigmotropism = growth in response to touch. Occurs in vines and other climbing plants.

33. Positive gravitropism in roots: the statolith hypothesis Statoliths 20 µm (b) Statoliths settling(a) Root gravitropic bending

34. Rapid turgor movements by the sensitive plant (Mimosa pudica) (a) Unstimulated state Leaflets after stimulation Pulvinus (motor organ) (c) Cross section of a leaflet pair in the stimulated state (LM) (b) Stimulated state Side of pulvinus with flaccid cells Side of pulvinus with turgid cells Vein 0.5 µm

35. Plant Control - Andersen 7 min

36. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings You should now be able to: 1.Compare the growth of a plant in darkness (etiolation) to the characteristics of greening (de-etiolation). 2.List six classes of plant hormones and describe their major functions. 3.Describe the phenomenon of phytochrome photoreversibility and explain its role in light- induced germination of lettuce seeds. 4.Explain how light entrains biological clocks.

37. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 5.Distinguish between short-day, long-day, and day-neutral plants; explain why the names are misleading. 6.Distinguish between gravitropism, thigmotropism, and thigmomorphogenesis. 7.Describe the challenges posed by, and the responses of plants to, drought, flooding, salt stress, heat stress, and cold stress.