Biology Sylvia S. Mader Michael Windelspecht Chapter 26 Flowering Plants: Control of Growth Responses Lecture Outline See separate FlexArt PowerPoint slides for all figures and tables pre-inserted into PowerPoint without notes. 1 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Outline • 26.1 Plant Hormones • 26.2 Plant Responses 2 26.1 Plant Hormones • Flowering plants respond to environmental stimuli – Stimuli include light, gravity, carbon dioxide levels, pathogen infection, drought, and touch – Response to stimuli leads to the survival of the species. • The responses can be: – Short term • Stomata open and close in response to light levels. – Long term • The response to gravity causes downward growth of the root and the upward growth of the stem. 3 Plant Hormones • Response of plants to environmental stimuli involves signal transduction – The binding of a molecular “signal” that initiates and amplifies a response. – Signal transduction involves the following: – Receptors – proteins activated by a specific signal – Transduction pathway – a series of relay proteins or enzymes that amplify and transform the signal to one understood by the machinery of the cell – Cellular response – the result of the transduction pathway 4 Plant Hormones • Hormones – Enable plant cells to communicate – Are synthesized in one part of the plant – Travel within phloem or from cell to cell in response to the appropriate stimulus 5 Signal Transduction in Plants Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. defense hormones hormone-binding site 3 blue light signal 2 Receptor: Molecule in the plasma membrane, cytoplasm, or nucleus that receives signal and becomes activated. Response: Most often a change in gene expression or a cellular process affects plant growth and development. Transduction pathway: A series of relay proteins that amplify and convert the original signal into one that affects cellular machinery . activated phototropin auxin relay proteins Defense responses auxin carrier Responses include bending of stem 1 activated auxin receptor Cytoplasm Gene expression changes Nucleus Responses include growth of roots 6 Plant Hormones • Auxins – Produced in shoot apical meristem – Found in young leaves, flowers, and fruits • Effects of auxin on growth and development: – Apically produced auxin prevents the growth of axillary buds • Apical dominance – Promotes growth of roots and fruit – Prevents loss of leaves and fruit – Promotes positive phototropism of stems 7 Auxin and Phototropism Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 1. Coleoptile tip is intact. 2. Coleoptile tip is removed. 3. Tips are placed on agar, and auxin diffuses into the agar. 4. Agar block is placed to one side of the coleoptile. 5. Curvature occurs beneath the block. 8 Plant Hormones • How Auxins Cause Stems to Bend • When a stem is exposed to unidirectional light, auxin moves to the shady sides • Auxin binds to plasma membrane receptors; the complex leads to the activation of a proton pump • Activated proton pumps H+ out of cell – Cell wall loosens – Turgor pressure increases due to the entry of water – Cell enlarges 9 Expansion of the Cell Wall Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. cellulose fiber in cell wall auxin nucleus H+ H+ enzyme (inactive) H+ H+ chloroplast 1 Cytoplasm H+ H+ H+ active enzyme H2O H+ 2 turgor turgor 3 10 Plant Hormones • Gibberellins are growth-promoting hormones – Gibberellins cause stem elongation – There are about 70 gibberellins • Each differ slightly chemically • The most common is gibberellic acid 11 Gibberellins Cause Stem Elongation Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. b. a: © Robert E. Lyons/Visuals Unlimited; b: © Sylvan Whittwer/Visuals Unlimited 12 Plant Hormones • The cytokinins are a class of hormones that promote cell division – found in dividing tissues of roots, in seeds, and in fruits – have been used to prolong the life of flower cuttings as well as vegetables in storage – Auxin and cytokinins interact – prevent senescence (aging process) 13 Interaction of Hormones Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. b. c. d. Courtesy Alan Darvill and Stefan Eberhard, Complex Carbohydrate Research Center, University of Georgia 14 Plant Hormones • Abscisic acid (ABA) is produced by any “green tissue” (i.e., tissue containing chloroplasts) – sometimes called the stress hormone • initiates and maintains seed and bud dormancy • brings about the closure of stomata 15 Dormancy and Winter Buds 16 Dormancy and Germination 17 Abscisic Acid Promotes Closure of Stomata Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. inside outside H2O K+ K+ K+ Ca2+ ABA Open stoma Guard cell plasma membrane Closed stoma 18 Plant Hormones • Ethylene (H2C = CH2) is a gas formed from the amino acid methionine. • Effects of ethylene – Abscission • Ethylene stimulates certain enzymes, such as cellulase, which helps cause leaf, fruit, or flower drop – Ripening of fruits • Increases the activity of enzymes, such as cellulase, that soften fruits • It also promotes the activity of enzymes that produce the flavor and smell of ripened fruits. – Axillary bud inhibition – Suppression of stem and root elongation 19 Ethylene and Abscission Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. No abscission © Kingsley Stern Abscission 20 Ethylene and Fruit Ripening Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. gene for ethylene biosynthesis enzyme ripe tomatoes harvested DNA transcription mRNA translation functional enzyme for ethylene biosynthesis ethylene synthesis (in plant) green tomatoes harvested no ethylene synthesis 21 Arabidopsis Is a Model Organism • Arabidopsis thaliana – A small flowering plant related to cabbage and mustard plants – Has no commercial value – It has become a model organism for the study of plant molecular genetics, including signal transduction. • It is small, so many hundreds of plants can be grown in a small amount of space. • Generation time is short – 5-6 weeks until maturity. • It normally self-pollinates, but it can easily be crosspollinated. • The number of base pairs in its DNA is relatively small. 22 Overall Appearance of Arabidopsis thaliana Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Arabidopsis thaliana Courtesy Elliot Meyerowitz/California Institute of Technology 23 26.2 Plant Responses • Tropism – Plant growth toward or away from a unidirectional stimulus • Positive tropism is growth toward the stimulus • Negative tropism is growth away from the stimulus – Gravitropism - movement in response to gravity – Phototropism - movement in response to light – Thigmotropism - movement in response to touch 24 Plant Responses • Gravitropism – When a plant is placed on its side, the stem grows upward, opposite of the pull of gravity – Stems with root caps grow downward • Response depends on sensors called statoliths – Auxin may be responsible for: • gravitropism of roots and shoots 25 Gravitropism Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. gravity b. c. 25 mm a: © Kingsley Stern; b: Courtesy Malcolm Wilkins, Glascow University; c: © BioPhot 26 Plant Responses • Phototropism – Positive phototropism of stems • Occurs because cells on the shady side of the stem elongate due to the presence of auxin • A pigment absorbing blue light initiates phototropism 27 Phototropin Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 1 cytoplasm blue light 2 3 blue light phot blue light phot ADP transduction pathway phot P plasma membrane ATP ATP 28 Plant Responses • Thigmotropism – Unusual growth due to contact with solid objects • Coiling of tendrils – Thigmomorphogenesis occurs when the entire plant responds to the presence of environmental stimuli • Wind • Rain 29 Coiling Response Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © John D. Cunningham/Visuals Unlimited 30 Plant Responses • Nastic movements: – Do not involve growth and – Are not dependent on the stimulus direction • Turgor movements result from touch, shaking, or thermal stimulation – Mimosa pudica – Venus flytrap • Sleep movements: – Occur daily in response to light and dark changes – Circadian rhythm 31 Turgor Movement Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. pulvinus Before vascular tissue cell retaining turgor cell losing turgor After © John Kaprielian/Photo Researchers, Inc.6 32 Plant Responses • Circadian rhythms: – Biological rhythms with a 24-hour cycle – Tend to be persistent • Rhythm is maintained in the absence of environmental stimuli • Caused by a biological clock 33 Sleep Movements and Circadian Rhythms Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Prayer plant (morning) a. Prayer plant (night) Morning glory (morning) b. Morning glory (night) Circadian Rhythm flowers open flowers close Period (about 24 hours) 0 12 24 36 48 Time (hours) c. (Top, both): © Tom McHugh/Photo Researchers, Inc.(Bottom left): © BIOS A. Thais/Peter Arnold, Inc.; (Bottom right): © BIOS Pierre Huguet/Peter Arnold, Inc. 34 Plant Responses • Photoperiodism: – Any physiological response prompted by changes in day or night length – influences flowering in some plants – requires participation of a biological clock and a plant photoreceptor called phytochrome 35 Plant Responses • Phytochrome is a blue-green leaf pigment that alternately exists in two forms – Phytochrome red (Pr) is inactive – Phytochrome far-red (Pfr) is active • Conversion of forms allows a plant to detect photoperiod changes • Also promotes seed germination and inhibits stem elongation 36 Phytochrome Conversion Cycle Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. lightsensitive region red light far-red light kinase inactive Pr active Pfr 37 Phytochrome Control of Shoot Elongation Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. Normal growth b. Etiolation (both): © Nigel Cattlin/Visuals Unlimited 38 Plant Responses • Flowering – Flowering plants can be divided into three groups based on their flowering status. • Short-day plants flower when the day length is shorter than a critical length • Long-day plants flower when the day length is longer than a critical length • Day-neutral plants are not dependent on day length for flowering – Some plants may require a specific sequence of day lengths in order to flower 39 Photoperiodism and Flowering Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cocklebur Clover night flash of light 24 hours critical length day flower flower 1 2 a. Short-day (long-night) plant 3 4 flower 5 6 b. Long-day (short-night) plant 40 Plant Responses • Responses to the biotic environment: • Plants are always under attack by herbivores and parasites. • Physical and Chemical Defenses – Cuticle-covered epidermis and bark – Secondary metabolites • Tannins • Alkaloids • Cyanogenic glycosides – Wound responses – proteinase inhibitors and systemin – Hypersensitive response (HR) – initiates wound response and seals wounded area – Indirect defenses – prevent egg laying in insects – Mutualistic relationships with animals - acacia tree and acacia ant 41 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Plant Predators and Parasites Alfalfa plant bug Fungus infection Monarch caterpillar and butterfly (Top left): Courtesy USDA/Agricultural Research Service, photo by Scott Bauer; (top right): © Kingsley Stern; (bottom left): © The McGraw Hill Companies, Inc./Ken Cavanagh, photographer; (bottom right): © Dwight Kuhn 42 Wound Response in Tomato Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. systemin cytoplasm lipase wounded leaf systemin release membranebound receptor proteinase inhibitors membrane lipids salicylic acid jasmonic acid transduction pathway nucleus activation of proteinase inhibitor genes 43