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Thought Question Plants can’t fight or hide or run away, so how do they adapt to a changing environment? 1 Lecture 7 Outline (Ch. 40, 41) I. Plant Defenses A. Methods of Attack B. Methods of Defense II. Responses to Light III. Circadian Rhythms IV. Responses to Gravity V. Responses to Touch VI. Plant Hormones A. Auxin B. Gibberellins C. Cytokinin D. Ethylene 2 E. Abscisic Acid Plant Defenses • Plants are susceptible to physical stresses Examples? • Other threats include: viruses, bacteria, fungi, animals, and other plants – Take nutrient resources of plants or use their cells – Some kill plant cells immediately, leading to necrosis • Why are nonnative invasive species especially problematic? Alfalfa plant bug 3 Plant Defenses • Dermal tissue: 1st line of defense – secrete wax: protect from water loss and attack – Dermis covered with cutin or suberin – substances to reinforce cell walls – Silica inclusions, trichomes, bark, and even thorns can also offer protection 4 Plant Defenses • Plant defenses aren’t always enough: – Mechanical wounds allow microbial entry – Parasitic worms can eat through plant cell walls • Some form tumors on roots – In some cases simply having bacteria on the leaf surface can increase damage 5 Plant Defenses • Fungi can enter through stomata Phases of fungal invasion 1. Windblown spore lands on leaves 2. Spore germinates & forms adhesion 3. Hyphae grow through cell walls and press against cell membrane 4. Hyphae differentiate 6 Toxin Defenses • Many plants produce toxins that kill herbivores, make them ill, or repel them with strong flavors or odors • Some plants have antimicrobial peptides • Secondary metabolites – Plants make defense compounds via modified metabolism – Alkaloids [Wild tobacco has elevated nicotine levels lethal to tobacco hornworms] – Tannins 7 Toxin Defenses 8 Toxin Defenses 9 Toxin Defenses • Ricin: alkaloid produced by castor bean plant – 6X more lethal than cyanide – A single seed can kill a small child – Binds ribosomes - inhibits translation 10 Immediate Plant Responses - Plants may produce protective compounds - Plants may summon “bodyguards” when attacked - Plants may warn other plants of attack - Some plants move rapidly 11 Animal “Body Guards” • Some plants “recruit” animals in mutualism • Acacia trees and ants – Small armies of ants protect Acacia trees from harmful herbivores – Plant provides ants with food and shelter Foolish katydid 12 Animal “Body Guards” – As caterpillar chews away, a wound response in the plant leads to release of a volatile compound – Female parasitoid wasp is attracted – Lays fertilized eggs in caterpillar – Eggs hatch and larvae kill caterpillar 13 Chemical Warnings • • • • Volatile chemicals released by plants boost defenses in neighbors Many virally-attacked plants produce salicylic acid – Activates an immune response Attacked plant converts salicylic acid to methyl salicylate (wintergreen) diffuses to air – Absorbed by neighboring healthy plants and reconverted to salicylic acid (aspirin) 14 Chemical Warnings • Tobacco plants produce salicylic acid to fight viral infections Salicylic acid production Salicylic acid production Methyl salicylate Salicylic acid production Virus Infected Plant Salicylic acid 15 production Touch Responses 16 Touch Responses • • • Leaves have sensory “hairs” on inside – Fly triggers hairs - generates signal Cells in outer leaf epidermis pump H+ into cell walls Enzymes activated cells absorb water Outer epidermal cells expand, close leaf • Reopening leaves takes several hours • Venus fly trap 17 Self-Check Defense Examples Secondary metabolites Recruited animals Volatile chemicals Movement 18 Sensory Systems in Plants Chapter 41 20 Plant Timekeeping/Light Detection Two major classes of light receptors: Blue-light photoreceptors • stomatal movements • phototropism Phytochromes – red/far-red receptor • shade avoidance response • photoperiodism A phytochrome consists of two identical proteins joined Photoreceptor activity. Enzyme - kinase activity. 20 Plant Orientation 21 Plant Responses to Light • Blue light receptor: Directional growth responses • Connect environmental signal with cellular perception of the signal, transduction into biochemical pathways, and 22 ultimately an altered growth response Plant Responses to Light • Blue light receptor: Embedded in cell membrane • When blue light detected, changes conformation, signal transduction differential elongation 23 Plant Timekeeping/Light Detection Circadian Rhythms • Cyclical responses to environmental stimuli – approximately 24 hours long – entrained to external clues of the day/night cycle • Phytochrome conversion marks sunrise and sunset – Providing the biological clock with environmental cues Many legumes – Lower their leaves in the evening and raise them in the morning 24 Noon Midnight Plant Timekeeping/Light Detection Photoperiodism • Response to time of year (seasons) • Photoperiod - relative lengths of night and day • Triggers many developmental processes – Bud break – Flowering – Leaf drop in deciduous trees • Are actually controlled by night length, not day length • that phytochrome is the pigment that receives red light, which can interrupt the nighttime portion of the photoperiod 25 Plant Timekeeping/Light Detection • Leaves detect lengths of night/day – An internal biological clock – A light-detecting phytochrome • Pigments found in leaves • Active/inactive depending on light conditions Still-unidentified chemical (florigens) travel from leaf to bud to either trigger or inhibit flowering 26 Response to Gravity • Response of a plant to the gravitational field of the Earth • Shoots exhibit negative gravitropism • Roots have a positive gravitropic response 27 Response to Gravity • Four general steps lead to a gravitropic response: 1. Gravity is perceived by the cell 2. Mechanical signal transduced into physiological signal 3. Physiological signal transduced inside cell & to other cells 4. Differential cell elongation occurs in the “up” and “down” sides of root and shoot 28 Gravity Response How Do Plants Detect Gravity? • • Starch-filled plastids – In specialized stem cells and root caps – Orient within cells toward gravity Changing plastid orientation triggers elongation root cell in root cap plastids 29 Gravity Response 30 Gravity Response 31 Thigmotropism • Thigmotropism is directional growth of a plant or plant part in response to contact • Thigmonastic responses occur in same direction independent of the stimulus • Examples of touch responses: Venus flytrap leaves Tendrils around objects Often due to differential elongation or manipulated water/turgor pressure 32 Responses to Mechanical Stimuli • Mimosa leaves have swollen structures called pulvini at base of leaflets – Stimulation triggers electrical signal – Triggers ions to outer side of pulvini – Water follows by osmosis – Decreased interior turgor pressure causes the leaf to fold 33 Responses to Mechanical Stimuli • Bean leaves – Pulvini rigid during the day – Lose turgor at night – Reduce transpiration during the night – Maximize photosynthetic surface area during the day 34 Plant Hormones (Plant) Hormone: Chemicals made in one location and transported to other locations for action Growth Reproduction Movement Water balance Dormancy 35 Plant Hormone Overview • Plants respond to stimuli and lead a stationary life • Plants, being rooted to the ground – Must respond to whatever environmental change comes their way 36 Plant Hormones Five major classes of plant hormones • Hormone effects depend on – - target cell – - developmental stage of the plant – - amount of hormone – - presence of other hormones 37 Plant Hormones 1. Auxins: • • • • • • Elongation of cells Root elongation stimulate (low concentrations) inhibit (high concentrations) Vascular tissues and fruit development Responses to light (phototropism), gravity (gravitropism), and touch (thigmotropism) 38 Cell elongation in response to auxin 3 Wedge-shaped expansins, activated by low pH, separate cellulose microfibrils from cross-linking polysaccharides. The exposed cross-linking polysaccharides are now more accessible to cell wall enzymes. Expansin 4 The enzymatic cleaving of the cross-linking CELL WALL polysaccharides allows the microfibrils to slide. The extensibility of the cell wall is increased. Turgor causes the cell to expand. Cell wall enzymes Cross-linking cell wall polysaccharides Microfibril H2O Plasma membrane H+ H+ 2 The cell wall becomes more acidic. Cell wall H+ H+ H+ H+ H+ H+ 1 Auxin increases the activity of proton pumps. Cytoplasm Nucleus Vacuole ATP H+ Plasma membrane Cytoplasm 5 With the cellulose loosened, the cell can elongate. 39 Other Auxin Stimulated Responses: • Lateral / branching root formation • Promote fruit growth (tomato sprays) • As herbicide, overdose kills eudicots Auxin is produced: • At the shoot apex, seeds, other actively growing tissues. 40 Plant Hormones 2. Gibberellins: • Stem elongation, flowering, and fruit development • Seed germination and bud sprouting 41 Gibberellins stimulate germination • After water is imbibed, the release of gibberellins from the embryo signals the seeds to break dormancy and germinate Responds by synthesizing and secreting digestive enzymes that hydrolyze stored nutrients in the endosperm. embryo releases gibberellin as a signal Aleurone Nutrients absorbed from the endosperm by the cotyledon are consumed during growth of the embryo into a seedling. Endosperm Embryo GA amylase Sugar GA Water cotyledon 42 Plant Hormones 3. Cytokinins: • Stimulate cell division and differentiation • Produced in actively growing tissues such as roots, embryos, and fruits Anti- aging effects. • Inhibit protein breakdown • Stimulate RNA and protein synthesis • Mobilize nutrients from surrounding tissues (florist sprays) 58 day old cutting: Genetically engineered to express more cytokinin on right 43 44 Control of Apical Dominance • Cytokinins and auxins interact in the control of apical dominance – The ability of a terminal bud to suppress development of axillary buds • If the terminal bud is removed – Plants become bushier “Stump” after removal of apical bud Axillary buds 44 Lateral branches Plant Hormones 4. Ethylene: • • Gas at room temperature Promotes abscission (falling off) of fruits, flowers, and leaves • Required (with auxin) for fruit development 45 46 Self-Check Why will these ripe bananas help the green avocados ripen faster? 46 Plant Hormones 5. Abscisic Acid: • • Initiates closing stomata in water-stressed plants Induces and maintains dormancy in buds and seeds – (inhibits gibberellins) 47 48 Abscisic Acid Two of the many effects of abscisic acid (ABA) are • Seed dormancy – Ensures seeds germinate only when conditions are optimal • Drought tolerance – Closes stomata, decreases shoot growth Coleoptile K+ K+ K+ Why is that one kernel (seed) germinating prematurely? 48 Self-Check Hormone Name Functions Auxin Gibberellin Cytokinin Ethylene Abscisic Acid 49 Plant Orientation Sprouts know where to go • Auxin controls direction of sprouting seedling • Distribution of auxin within shoot and root cells is influenced by gravity and light 50 Plant Orientation Opaque cap over tip. 51 Plant Orientation Clear cap over tip. Opaque sleeve over bending region. 52 Plant Orientation Cells elongate rapidly. Cells elongate slowly. 53 Plant Orientation Shoot Elongation • In shoot, light and gravity cause auxin movement to the lower side Auxin stimulates elongation of stem cells Stem bends away from gravity & toward light Root Growth Due to gravity, auxin builds up on the lower side of the root Auxin retards elongation of root cells, and the root bends toward gravity 54 Senescence • Process by which leaves, fruits, and flowers age rapidly – Promoted by changes in hormone levels • Cytokinin and auxin production decreases • Ethylene production increases 55 Senescence • Proteins, starches, and chlorophyll broken down – Products stored in roots and other permanent tissues Abscission Ethylene stimulates production of enzyme that digests cell walls at base of petiole Leaf falls when cells are sufficiently weakened 56 Senescence bud leaf petiole abscission layer 57 Dormancy • Period of reduced metabolic activity in which the plant does not grow and develop Maintained by abscisic acid Dormancy broken by: increased temperature, longer day length occur in the spring 58 Lecture 7 Summary 1. Plant Physical & Biological Stresses (Ch. 40) 2. Methods of Defense (Ch. 40) Toxins / volatiles Animals Movement 3. Responses to Light (Ch. 41) - photoreceptors - circadian rhythms 4. Responses to Gravity (Ch. 41) - stems - roots 5. Responses to Touch (Ch. 41) 6. Plant Hormones (Ch. 41) - general functions - role in cell elongation - senescence - dormancy