Chapter 39 Plant Responses to Internal and External

Chapter 39 Plant Responses to Internal and External Signals Power. Point® Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Overview: Stimuli and a Stationary Life • Linnaeus noted flowers of different species opened at different times of day/could be used by horologium florae, or floral clock – Reflects time insect pollinators active, one of environmental factors plant senses to compete successfully – Plant’s morphology/physiology tuned to surrounding by complex interactions between environmental stimuli/internal signals • Plants, being rooted to ground, must respond to environmental cues by adjusting individual pattern of growth/development – Must detect changes first – Bending of seedling toward light begins with sensing direction, quantity, and color of light Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Concept 39. 1: Signal transduction pathways link signal reception to response • All organisms received specific signals/respond to them in ways that enhance survival/reproductive success • Plants have cellular receptors that detect changes in their environment (molecule affected by stimulus) – For stimulus to elicit response, certain cells must have appropriate receptor – Stimulation of receptor initiates specific signal transduction pathway Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -2 (a) Before exposure to light Tall, spindly stem/nonexpanded leaves (morphological adaptations called etiolation enable shoots to penetrate soil, including short roots due to little need for water absorption from little water loss by shoots) Expanded leaves hindrance as shoots push through soil/chlorophyll waste of energy (underground) (b) After a week’s exposure to natural daylight Begins to resemble typical plant w/broad green leaves, short sturdy stems, long roots (transformation begins w/reception of light by specific pigment, phytochrome) by undergoing changes (de-etiolation) by reception of signal (light) which is transduced into responses (greening)

Fig. 39 -3 Review of a general model for signal transduction pathways CELL WALL 1 Reception CYTOPLASM 2 Transduction Relay proteins and second messengers Receptor Hormone or environmental stimulus Plasma membrane 3 Response Activation of cellular responses

Reception/Transduction • Internal/external signals are detected by receptors, proteins that change in response to specific stimuli – Phytochrome functioning in de-etiolation located in cytoplasm, not built into plasma membrane as most receptors • Second messengers (small molecules/ions in cell transfer and amplify signals from receptors to proteins that cause responses) activate hundreds of molecules of specific enzyme from one activated phytochrome molecule Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -4 -3 An example of signal transduction in plants: the role of phytochrome in the de-etiolation (greening) response 1 Reception 2 Transduction 3 Transcription factor 1 CYTOPLASM Plasma membrane c. GMP Second messenger produced Phytochrome activated by light Cell wall Response 2. One pathway uses c. GMP as 2 nd messenger that activates specific protein kinase. Other pathway involves increase in cytosolic level of Ca 2+, which activates different protein kinase Specific protein kinase 1 activated P Transcription factor 2 P Specific protein kinase 2 activated 3. Both pathways lead to expression of genes for proteins that function in de-etiolation (greening) response 1. Light signal detected by phytochrome receptor; phytochrome undergoes change in shape, which then activates at least two signal transduction pathways Ca 2+ channel opened Ca 2+ NUCLEUS Transcription Translation De-etiolation (greening) response proteins

Response • Signal transduction pathway leads to regulation of one or more cellular activities – In most cases, these responses to stimulation involve increased activity of enzymes – This can occur by transcriptional regulation (increases/decreases synthesis of m. RNA that codes for enzyme) or post-translational modification (activates existing enzyme molecules) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Transcriptional Regulation • Specific transcription factors bind directly to specific regions of DNA and control transcription of genes – In phytochrome-induced de-etiolation, several transcription factors activated by phosphorylation in response to appropriate light conditions • Mechanism by which signal promotes new developmental course may depend on – Positive transcription factors (activators): proteins that increase transcription of specific genes – Negative transcription factors (repressors): proteins that decrease transcription of specific genes • Mutation in Arabidopsis allows for light-grown morphology underground (except for pale color because no chlorophyll grown because no light) because genes activated by light normally blocked by repressor eliminated by mutation Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Post-Translational Modification of Proteins • Post-translational modification involves modification of existing proteins in signal response • Modification often involves phosphorylation of specific amino acids which alters protein’s hydrophobicity/ activity – Many 2 nd messengers (including some forms of phytochrome) activate protein kinases directly, often leading to kinase cascades – Many signal pathways ultimately regulate synthesis of new proteins, usually by turning specific genes on or off Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

De-Etiolation (“Greening”) Proteins • Many enzymes that function in certain signal responses – Are directly involved in photosynthesis – Are involved in supplying chemical precursors for chlorophyll production – Affect levels of plant hormones that regulate growth Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Concept 39. 2: Plant hormones help coordinate growth, development, and responses to stimuli • Hormones: Hormones chemical signals that coordinate different parts of organism – Signaling molecule produced in tiny amounts by one part of organism and transported to other parts, where it binds to specific receptor and triggers responses in target cells/tissues – Many plant biologists prefer plant growth regulator to describe organic compounds that modify or control one or more specific physiological processes within plant Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

The Discovery of Plant Hormones http: //bcs. whfreeman. com/thelifewire/content/chp 38/3802001. html • Tropism: Tropism any response resulting in curvature of organs toward or away from stimulus (often caused by hormones) • In late 1800 s, Charles Darwin/son Francis conducted experiments on phototropism (plant’s response to light) – Observed that grass seedling could bend toward light only if tip of coleoptile was present (maximum exposure of leaves to light for photosynthesis) • Shoot of sprouting grass (enclosed in coleoptile) grows straight upward if seedling kept in dark/illuminated for all sides uniformly • If illuminated form one side, grows toward light (results from differential growth of cells on opposite sides of coleoptile; cells on darker side elongate faster than those on brighter side) – Postulated signal was transmitted from tip to elongating region Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Darwin/Darwin: phototropic response only when tip is illuminated: Only tip of coleoptile senses light Phototrophic bending occurred at distance from site of light perception (tip)

Fig. 39 -5 c RESULTS Boysen-Jensen: phototropic response when tip is separated by permeable barrier, but not with impermeable barrier Demonstrated that signal was mobile chemical substance Light Tip separated by gelatin (permeable) Phototrophic response occurred Tip separated by mica (impermeable) No phototrophic response occurred

Fig. 39 -6 RESULTS http: //bcs. whfreeman. com/thelifewire/content/chp 38/3802002. html Excised tip placed on agar cube In 1926, Frits Went extracted chemical messenger for phototropism, auxin by modifying earlier experiments Auxin later purified/ chemical structure found to be IAA (indoleacetic acid) acid by Kenneth Thimann Growth-promoting chemical diffuses into agar cube Control (agar cube lacking chemical) has no effect Agar cube with chemical stimulates growth Offset cubes cause curvature Off center blocks caused decapitated coleoptiles to bend away from side with agar block, as though growing toward light Concluded that agar block contained chemical produced in coleoptile tip that stimulated growth as it passed down coleoptile/ which curved toward light because higher concentration of growth-promoting chemical on darker side of coleoptile

• These studies show asymmetrical distribution of auxin moving down from coleoptile tip and causes cells on darker side to elongate faster than cells on brighter side • Studies of organs other than grass coleoptiles provide less support for this, but there is asymmetrical distribution of certain substances that may act as growth inhibitors, and these are more concentrated on lighted side of stem Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

A Survey of Plant Hormones • Plant hormones are produced in very low concentration, but minute amount can greatly affect growth/development of plant organ – Amplified in some way – Hormone may act by altering expression of genes, by affecting activity of existing enzymes, by changing properties of membranes – Any of these actins could redirect metabolism and development of cell responding to small number of hormone molecules • In general, hormones control plant growth/development by affecting division, elongation, and differentiation of cells – Some mediate shorter-term physiological responses to environmental stimuli – Each hormone has multiple effects, depending on site of action, concentration, developmental stage of plant Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Table 39 -1

Auxin • Auxin refers to any chemical that promotes elongation of coleoptiles (multiple functions in flowering plants: cell elongation/lateral root formation) – Indoleacetic acid (IAA) IAA is common natural auxin in plants – Transported directly through parenchyma tissue, from one cell to another • Polar transport: unidirectional transport of auxin (in shoot, moves only from tip to base) – Polarity of auxin movement attributable to polar distribution of auxin transport protein in cells that move hormone from basal end of one cell into apical end of neighboring cell Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

The Role of Auxin in Cell Elongation • One of chief functions of auxin is to stimulate elongation of cells within young developing shoots – Apical meristem of shoot major site of auxin synthesis, moving down to region of cell elongation • According to acid growth hypothesis, hypothesis auxin stimulates proton pumps in plasma membrane – Proton pumps lower p. H in cell wall, activating expansins (enzymes that loosen wall’s fabric by breaking hydrogen bonds between cellulose microfibrils and other cell wall constituents) – With the cellulose loosened, cell can elongate • Also rapidly alter gene expression, causing cells in region of elongation to produce new proteins within minutes Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Figure 39. 8 Cell elongation in response to auxin: the acid growth hypothesis 3 Expansins separate Cross-linking polysaccharides Cell wall–loosening enzymes microfibrils from crosslinking polysaccharides. Expansin CELL WALL 4 Cleaving allows microfibrils to slide. Cellulose microfibril H 2 O 2 Cell wall Plasma membrane becomes more acidic. Cell wall 1 Auxin increases proton pump activity. Plasma membrane Nucleus Cytoplasm Vacuole CYTOPLASM http: //bcs. whfreeman. com/thelifewire/content/chp 38/3802003. html 5 Cell can elongate.

Lateral and Adventitious Root Formation Auxins as Herbicides/Other Effects of Auxin • Auxin is involved in root formation and branching – Used in vegetative propagation of plants by cuttings • Overdose of synthetic auxins can kill eudicots – Monocots rapidly inactivate these synthetic auxins • Auxin affects secondary growth by inducing cell division in vascular cambium and influencing differentiation of secondary xylem – Developing seeds produce auxin which promotes fruit growth (can induce normal fruit development without pollination for commercially grown seedless tomatoes) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Cytokinins Control of Cell Division and Differentiation • Cytokinins are so named because they stimulate cytokinesis (cell division) – Produced in actively growing tissues (roots, embryos, and fruits) • Work together w/auxin to control cell division/differentiation – When concentration of both at certain levels, mass of cells continues to grow, but remains cluster of undifferentiated cells (callus) • If cytokinin levels increase, shoot buds develop • If auxin level increase, roots form Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Control of Apical Dominance • Cytokinins, auxin, and other factors interact in control of apical dominance, terminal bud’s ability to suppress development of axillary buds – Direct inhibition hypothesis says auxin/cytokinins act antagonistically in regulating axillary bud growth • Auxin transported down shoot from apical bud directly inhibits axillary buds from growing, causing stem to elongate • Cytokinins entering shoot system from roots counter action by signaling axillary buds to grow • Does not account for all experimental findings Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -9 Lateral branches “Stump” after removal of apical bud (b) Apical bud removed, enables lateral branches to grow (removes inhibition) Axillary buds (a) Apical bud intact (primary source of auxin) Inhibition of growth of axillary buds, possibly influenced by auxin from apical bud, favors elongation of shoot’s main axis (c) Auxin added to decapitated stem prevents lateral branches from growing

Anti-Aging Effects • Cytokinins retard aging of some plant organs by inhibiting protein breakdown, stimulating RNA and protein synthesis, and mobilizing nutrients from surrounding tissues – If leaves removed from plant dipped in cytokinin solution, stay greener much longer – Also slows deterioration of leaves on intact plants (used to spray on cut flowers to keep fresh) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Gibberellins Stem Elongation/Fruit Growth • Gibberellins have variety of effects, such as stem elongation, fruit growth, and seed germination • Major sites of gibberellin production are roots/young leaves – Stimulate growth of leaves/stems, little effect on roots – In stems, they stimulate cell elongation and cell division • Both auxin/gibberellins must be present for some fruit to ripen – Used in spraying of Thompson seedless grapes which makes individual grapes grow larger/ make internodes elongate, allowing more space for individual grapes (reduces yeasts/other microorganisms from infecting fruit) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -11 Mobilization of nutrients by gibberellins during the germination of grain seeds such as barley 1 After water is imbibed, gibberellins (GA) from embryo signals to aleurone to germinate 2 Aleurone secretes -amylase and other enzymes. 3 Sugars and other nutrients are consumed. Aleurone Endosperm -amylase GA GA Water Scutellum (cotyledon) Radicle Germination Sugar

Brassinosteroids • Brassinosteroids are steroids, chemically similar to sex hormones of animals – Induce cell elongation/division in stem segments and seedlings – Reduce leaf abscission (leaf drop) – Promote xylem differentiation Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Abscisic Acid • Abscisic acid (ABA) slows growth – Antagonistic to growth hormones and ratio determines final physiological outcome – Two of many effects of ABA – Seed dormancy – Drought tolerance Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Seed Dormancy • Seed dormancy ensures that seed will germinate only in optimal conditions • In some seeds, dormancy is broken when – ABA is removed by heavy rain, light, or prolonged cold – Light/prolonged exposure to cold to inactivate ABA • Precocious (early) germination is observed in maize mutants that lack transcription factor required for ABA to induce expression of certain genes Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Drought Tolerance • ABA is primary internal signal that enables plants to withstand drought – Plant begins to wilt ABA accumulates in leaves rapid closing of stomata reduces transpiration/prevents further water loss • ABA causes potassium channels in guard cells to open loss of potassium ions from cells osmotic loss of water guard cell turgor reduction closing of stomatal pores Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Ethylene (H 2 C=CH 2) The Triple Response to Mechanical Stress • Plants produce ethylene in response to stresses such as drought, flooding, mechanical pressure, injury, and infection – Effects of ethylene include response to mechanical stress, senescence, leaf abscission, fruit ripening, in response to high concentrations of externally applied auxin • Ethylene induces triple response, response which allows growing shoot to avoid obstacles (rock in ground) – Consists of slowing of stem elongation, thickening of stem (making it stronger), and horizontal growth (curvature) – Resumes vertical growth when obstacle gone Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

• Ethylene-insensitive mutants fail to undergo triple response after exposure to ethylene because they lack functional ethylene receptor (ein or ethylene-insensitive mutants) • Others undergo triple response in air without physical obstacles and have regulatory defect that causes them to produce ethylene at rates 20 x normal (eto or ethylene overproducing mutants) – Can be restored to wild-type by treating seedlings with inhibitors of ethylene synthesis • Other mutants undergo triple response in air but do not respond to inhibitors of ethylene synthesis (ctr, constitutive tripleresponse mutants) – Do not respond to inhibitors of ethylene synthesis Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -14 ein mutant ctr mutant (a) ein mutant fails to undergo triple response in presence of ethylene (b) ctr mutant undergoes triple response even in absence of ethylene

Senescence • Senescence: Senescence programmed death of plant cells or organs or entire plant – Starts w/onset of apoptosis (programmed cell death) where newly formed enzymes break down many chemical components (chlorophyll, DNA, RNA, proteins, membrane lipids) to be salvaged by plant – Burst of ethylene associated w/programmed destruction of cells, organs, entire plant Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Leaf Abscission • Change in balance of auxin and ethylene controls leaf abscission (process that occurs in autumn when leaf falls) – Essential elements salvaged/stored in stem parenchyma cells – Nutrients recycled back to developing leaves next spring – After leaf falls, protective layer of cork becomes leaf scar that prevents pathogens from invading plant Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fruit Ripening • Immature fleshy fruit (tart, hard, green) protect them from herbivores • Ripe fruit attracts animals to disperse seeds • Burst of ethylene production (positive feedback) in fruit triggers ripening process (enzymatic breakdown of cell wall softens fruit, conversion of starches/acids to sugars makes them sweet) – Moving air prevents ethylene accumulation/ carbon dioxide prevents ethylene production slows ripening of stored fruits Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Systems Biology and Hormone Interactions • Interactions between hormones and signal transduction pathways make it hard to predict how genetic manipulation will affect plant • Systems biology seeks comprehensive understanding that permits modeling of plant functions Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Concept 39. 3: Responses to light are critical for plant success • Light cues many key events in plant growth and development, photosynthesis, measuring passage of days and seasons – Photomorphogenesis: Photomorphogenesis effects of light on plant morphology • Plants detect not only presence of light but also direction, intensity, and wavelength (color) – Graph (action spectrum) depicts relative effectiveness of different wavelengths of radiation in driving particular process • Two peaks (red/blue light) for photosynthesis – Action spectra are useful in studying any process that depends on light (phototropism) – Two major classes of light receptors: blue-light photoreceptors and phytochromes Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -16 Action spectrum for blue-light-stimulated phototropism in maize coleoptiles Phototropic effectiveness 1. 0 436 nm 0. 8 0. 6 0. 4 0. 2 0 400 450 500 550 600 650 700 Wavelength (nm) (a) Action spectrum for blue-light phototropism Light Time = 0 min Time = 90 min (b) Coleoptile response to light colors Phototropic bending toward light controlled by phototropin (photoreceptor sensitive to blue/violet light, particularly blue light

Blue-Light Photoreceptors Phytochromes as Photoreceptors • Various blue-light photoreceptors control hypocotyl elongation, stomatal opening, and phototropism • Phytochromes are pigments that regulate many of plant’s responses to light throughout its life – These responses include seed germination and shade avoidance Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Phytochromes and Seed Germination • Many seeds remain dormant until light conditions change • In 1930 s, scientists at U. S. Department of Agriculture determined action spectrum for light-induced germination of lettuce seeds – Red light increased germination, while far-red light inhibited germination – Final light exposure was determining factor – Effects of red/far-red light reversible Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -18 Photoreceptor responsible for opposing effects of red/far-red light are phytochromes Two identical subunits, each consisting of polypeptide component covalently bonded to nonpolypeptide chromophore Chromophore Photoreceptor activity Kinase activity

• Phytochromes exist in two photoreversible states – Depend on color of light provided – Converts Pr (inhibits germination) to Pfr, which triggers many developmental responses (germination) – Though light contains both red and far red light, conversion to Pfr faster than conversion to Pr so ratio of Pfr to Pr increases in light, triggering germination Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -19 http: //highered. mcgraw-hill. com/sites/9834092339/student_view 0/chapter 41/animation_-_phytochrome_signaling. html Pfr Pr Red light Responses: seed germination, control of flowering, etc. Synthesis Far-red light Slow conversion in darkness (some plants) Enzymatic destruction

Phytochromes and Shade Avoidance • Phytochrome system also provides plant with information about quality of light – Sunlight contains both red/far-red radiation – During day, Pr ⇄ Pfr interconversion reaches dynamic equilibrium, with ratio of two phytochrome forms indicating relative amounts of red/far-red light – Allows plants to adapt to changes in light conditions • Shaded plants receive more far-red than red light – In “shade avoidance” response, response phytochrome ratio shifts in favor of Pr when tree is shaded, inducing tree to allocate more resources to growing taller • Direct sunlight stimulates branching/inhibits vertical growth Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Biological Clocks and Circadian Rhythms • Phytochrome helps plant keep track of passage of days/seasons • Many plant processes (transpiration, synthesis of certain enzymes) oscillate during the day – Response to changes in light levels, temperature, and relative humidity accompanying 24 -hour cycle of day/night – Many legumes lower their leaves in evening and raise them in morning, even when kept under constant light or dark conditions Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

• Circadian rhythms: rhythms cycles that are about 24 hours long and are governed by internal “clock” rather than daily responses to environmental cycle – Can be entrained (set) to exactly 24 hours by day/night cycle from environment – Deviations (free-running periods of 21 -27 hours) occur when organism kept in constant environment (not erratic because still keep perfect time, just not synchronized with outside world) – We can interfere with biological rhythm, but will reestablish position for time of day when interference removed – Clock may depend on synthesis of protein regulated through feedback control and may be common to all eukaryotes Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

The Effect of Light on the Biological Clock • Light is factor that entrains biological clock to precisely 24 hours each day – Desynchronization occurs when clock on wall not synchronized w/internal clocks (jet lag) • Phytochrome conversion marks sunrise/sunset, providing biological clock w/environmental cues – Reduction of Pfr due to • No sunlight to convert Pr to Pfr • Pigment of phytochrome synthesized as Pr • Decrease of Pfr in darkness when enzymes destroy more Pfr than Pr – Increase of Pfr at dawn (light returns) resets biological clock Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Photoperiodism and Responses to Seasons • Seasonal events critical in life cycles of most plants (seed germination, flowering, onset/breaking of bud dormancy) require photoperiod – Photoperiod (relative lengths of night and day) is environmental stimulus plants use most often to detect time of year – Photoperiodism is physiological response to photoperiod, or response to change in length of night, that results in flowering in low-day and short-day plants/preparation for winter Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Photoperiodism and Control of Flowering • Some processes, including flowering in many species, require certain photoperiod • Critical length: length period of daylight, specific in length for any given species, that appears to initiate flowering in long-day plants or inhibit flowering in short-day plants – Short-day plant: plants that flower when light period is shorter than critical length/chrysanthemums, poinsettias, some soybean – Long-day plants: plants that flower when light period is longer than certain number of hours/spinach, radishes, lettuce, irises, many cereal varieties – Day-neutral plants: plants flowering controlled by plant maturity, not photoperiod/tomatoes, rice, dandelions Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Critical Night Length • In the 1940 s, researchers discovered that flowering and other responses to photoperiod are actually controlled by night length, not day length • Short-day plants (really long-night plants) governed by whether critical night length sets minimum number of hours of darkness • If daytime portion of photoperiod broken by brief exposure to darkness, no effect on flowering • If nighttime part interrupted by even few minutes of dim light, flowering will not occur • Long-day plants (really short-night plants) governed by whether critical night length sets maximum number of hours of darkness • Brief flash of light interrupting long period of darkness will induce flowering, even if not enough daylight hours Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -21 Photoperiodic control of flowering 24 hours (a) Short-day (long-night) plant, Pfr inhibits flowering Light Critical dark period Flash of light Darkness (b) Long-day (short-night) plant, Pfr induces flowering Flash of light

Fig. 39 -22 Reversible effects of red and far-red light on photoperiodic response 24 hours Red light is most effective color in interrupting nighttime portion of photoperiod Action spectra/photoreversibility experiments show that phytochrome is pigment that receives red light R flash of red light shortens dark period (night length) RFR subsequent flash of far-red light cancels red flash’s effect (restores night length) RFRRFR Critical dark period http: //bcs. whfreeman. com/thelifewire/content/chp 39/3902002. html Long-day Short-day (long-night) (short-night) plant

• Some plants flower after only single exposure to required photoperiod • Other plants need several successive days of required photoperiod • Still others need environmental stimulus in addition to required photoperiod – For example, vernalization is pretreatment with cold to induce flowering Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

A Flowering Hormone? • Only one leaf needed to cue buds to develop flowers • Florigen: Florigen flowering signal, not yet chemically identified – May be macromolecule governed by CONSTANS gene, which is expressed in leaves (as is florigen) – CONSTANS might turn on gene called FLOWERING LOCUS T (FT) in leaf/FT protein travels to shoot apical meristem and initiates flowering Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -23 Experimental evidence for a flowering hormone 24 hours Graft Short-day plant grown under short-day conditions will flower Long-day plant grafted to short-day plant Long-day plant grown under short-day induced to grow by transmission conditions doesn’t flower of florigen across grafts

Meristem Transition and Flowering • For bud to form flower instead of vegetative shoot, meristem identity genes must first be switched on – Organ identity genes that specify spatial organization of floral organs (sepals, petals, stamens, carpels) activated in appropriate regions of meristem • Researchers seek to identify signal transduction pathways that link cues such as photoperiod and hormonal changes to gene expression required for flowering Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Concept 39. 4: Plants respond to a wide variety of stimuli other than light/Gravity • Because of immobility, plants must adjust to range of environmental circumstances through developmental and physiological mechanisms • Gravitropism: Gravitropism Response to gravity/what causes roots to grow down (positive gravitropism) gravitropism and shoots up (negative gravitropism) gravitropism – Auxin plays key role in gravitropism Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

• Plants may detect gravity by settling of statoliths, statoliths specialized plastids containing dense starch grains located in lower portions of cells (in roots, certain root cap cells) – Aggregates at lower points trigger redistribution calcium, which causes lateral transport of auxin root within – Calcium/auxin accumulate on lower side of root’s zone of elongation – At high concentrations, auxin inhibits cell elongation, slowing growth on root’s lower side – More rapid elongation on upper side causes root to curve as it grows – Still occurs in plants w/no statoliths (dense organelles, in addition to starch granules, may contribute to gravity detection) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Mechanical Stimuli • Thigmomorphogenesis: Thigmomorphogenesis changes in form that result from mechanical disturbance – Rubbing stems of young plants couple of times daily results in plants that are shorter than controls/tree growing on windy hill shorter/stockier than those in sheltered location – Mechanical stimulation signal transduction pathway increase in cytosolic Ca 2+ activation of specific genes of proteins cell wall properties • Thigmotropism is growth in response to touch – Occurs in vines/other climbing plants where coil rapidly around supports tendrils that • Grow straight until they touch something that stimulates coiling response (caused by differential growth of cells on opposite sides of tendril) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

• Rapid leaf movements in response to mechanical stimulation are examples of transmission of electrical impulses (action potentials) potentials – Rapid loss of turgor in cells within pulvini (specialized motor organs at joints of leaf) loss of potassium osmotic loss of water motor cells become flaccid regain turgor (~10 minutes) open again Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Environmental Stresses • Environmental stresses have potentially adverse effect on survival, growth, and reproduction • Stresses can be – Abiotic (nonliving) include drought, flooding, salt stress, heat stress, cold stress – Biotic (living) include pathogens, herbivores • Plants not tolerating environmental stress will succumb/be outcompeted by other plants, and become locally extinct Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Drought • During drought, plants reduce transpiration by – Closing stomata (stimulates increased synthesis/release of abscisic acid in leaf which helps keep stomata closed by acting on guard cell membranes) – Slowing leaf growth (need turgor/accumulation of abscisic acid) – Reducing exposed surface area – Also reduces photosynthesis (why drought diminishes crop growth) • Growth of shallow roots is inhibited (dries from surface down), while deeper roots continue to grow (look for deeper water) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Flooding (Oxygen deprivation) • Waterlogged soil lacks air spaces that provide oxygen for cellular respiration in roots – Oxygen deprivation ethylene causes some cells in root cortex to undergo enzymatic apoptosis air tubes provide oxygen to submerged roots – Aerial roots (mangroves) Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Salt Stress • Salt can lower water potential of soil solution – As water potential of soil solution becomes more negative, water potential gradient from soil to roots lowered, reducing water uptake • Na/certain other ions toxic to plants w/relatively high concentrations – Selectively permeable membranes of root cells impede uptake of most harmful ions, which causes problem acquiring water from solute-rich soils • Plants respond to salt stress by producing solutes tolerated at high concentrations – Keeps water potential of cells more negative than that of soil solution without admitting toxic quantities of salt • Halophytes: Halophytes special anatomical/physiological adaptations to grow best in salty soils (salt gland on leaves eliminate excess salt Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Heat Stress • Excessive heat can denature plant’s enzymes/damages metabolism • Evaporative cooling associated w/transpiration keeps temperature of leaf 3°-10°C lower than ambient air temperature – When stomata close, cooling is lost to conserve water • Most plants will begin producing heat-shock proteins (protect other proteins from heat stress) when exposed to excessive temperatures (40°C or above for temperate zone plants) – Some are chaperone proteins (chaperonins) which help other proteins fold into functional shapes – Molecules might bind to other proteins/help prevent denaturation Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Cold Stress • Cold temperatures decrease membrane fluidity – Below critical point, lipids become locked into crystalline structure which alters solute transport across membrane/ adversely affects function of membrane proteins – Altering lipid composition (unsaturated fatty acids) of membranes is response to cold stress • Freezing causes ice to form in plant’s cell walls/intercellular spaces – Generally don’t freeze because solutes lower freezing point, but water loss in cell wall can cause dehydration/high concentrations of solutes can be toxic – Many frost-resistant species increase cytoplasmic levels of specific solutes (sugars) that are well tolerated at high concentrations/help reduce loss of water Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Concept 39. 5: Plants respond to attacks by herbivores and pathogens/Defenses Against Herbivores • Plants use defense systems to deter herbivory, prevent infection, and combat pathogens • Herbivory (animals eating plants) is stress that plants face in any ecosystem – Plants counter excessive herbivory with physical defenses (thorns)/chemical defenses (distasteful/toxic compounds) • Canavanine resembles arginine replaces it in insect’s proteins disrupts protein conformation insect dies – Some plants even “recruit” predatory animals that help defend against specific herbivores Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Figure 39. 28 A maize leaf “recruiting” a parasitoid wasp as a defensive response to an armyworm caterpillar, an herbivore 4 Recruitment of parasitoid wasps that lay their eggs within caterpillars 3 Synthesis and release of volatile attractants 1 Wounding 1 Chemical in saliva 2 Signal transduction pathway

• Plants damaged by insects can release volatile chemicals to warn other plants of the same species – Methyljasmonic acid from spider mite-infested lima beans signals noninfested nearby plants activate expression of genes become less susceptible to spider mites and more attractive to another species of mites that preys on spider mites Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Defenses Against Pathogens http: //bcs. whfreeman. com/thelifewire/content/chp 40/40020. html • Plant’s first line of defense against infection is epidermis of primary plant body and periderm of secondary plant body • If pathogen penetrates dermal tissue, second line of defense is chemical attack that kills pathogen and prevents its spread – Is enhanced by inherited ability to recognize certain pathogens • Virulent pathogen : one plant has little specific defense against • Avirulent pathogen: one that may harm but does not kill host plant Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

• Gene-for-gene recognition involves recognition of pathogen-derived molecules by protein products of specific plant disease resistance (R) genes – Pathogen is avirulent if it has specific Avr gene corresponding to particular R allele in host plant • R protein usually recognizes single corresponding pathogen molecule encoded by one of pathogen’s Avr gene – R proteins activate plant defenses by triggering signal transduction pathways – These defenses include hypersensitive response and systemic acquired resistance Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

The Hypersensitive Response • Hypersensitivity response: response genetically programmed death of infected cells/tissues, as well as tissue reinforcement and antibiotic production, at infection site (localized, specific, containment response based on gene-for-gene recognition between host and pathogen) – Cells at infection site mount chemical defense seal off area destroy themselves – Induces production of phytoalexins (toxic compounds w/general fungicidal and bactericidal properties) and PR proteins (pathogenesis-related proteins, proteins many being enzymes that hydrolyze components in cell walls of pathogens) – Stimulates changes in cell wall that confine pathogen Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Systemic Acquired Resistance • Systemic acquired resistance: resistance long-lasting systemic response that primes plant for resisting broad spectrum of pathogens in organs distant from infection site (nonspecific, provide protection against diversity of pathogens that lasts for days) – Methylsalicylic acid most likely candidate for signaling molecule that moves from infection site to elicit systemic acquired resistance – Converted to salicylic acid in areas remote from sites of infection, which then activates signal transduction pathway that induces production of PR proteins and resistance to pathogen attack Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings

Fig. 39 -UN 4 Indicate the response to each condition by drawing a straight seedling or one with the triple response. Control Wild-type Ethylene insensitive (ein) Ethylene overproducing (eto) Constitutive triple response (ctr) Ethylene added Ethylene synthesis inhibitor

Fig. 39 -UN 5

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 5. Distinguish between short-day, long-day, and day-neutral plants; explain why the names are misleading 6. Describe how plants tell up from down 7. Distinguish between thigmotropism and thigmomorphogenesis 8. Describe the challenges posed by, and the responses of plants to, drought, flooding, salt stress, heat stress, and cold stress 9. Describe how the hypersensitive response helps a plant limit damage from a pathogen attack Copyright © 2008 Pearson Education, Inc. , publishing as Pearson Benjamin Cummings