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Do plants feel like human? PLANT SIGNAL TRANSDUCTION an overview • The Telegraph Plant https://www.youtube.com/watch?v=J-fIKlcCbSU • What Plants Talk About (Full Documentary) https://www.youtube.com/watch?v=CrrSAc-vjG4 Oct. 15, 2015 Reception and Transduction Signal transduction pathways link signal reception to response • Plants have cellular receptors that detect changes in their environment • For a stimulus to elicit a response, certain cells must have an appropriate receptor • Stimulation of the receptor initiates a specific signal transduction pathway • A potato left growing in darkness produces shoots that look unhealthy and lacks elongated roots Etiolated potato in response to light exposure is good example of signal Transduction • These are morphological adaptations for growing in darkness, collectively called etiolation • After exposure to light, a potato undergoes changes called de-etiolation, in which shoots and roots grow normally • A potato’s response to light is an example of cell-signal processing • The stages are reception, transduction, and response • Internal and external signals are detected by receptors, proteins that change in response to specific stimuli • Second messengers transfer and amplify signals from receptors to proteins that cause responses Signal Transduction Pathways Etiolation vs De-etiolation 1. Reception 2. Transduction • Second Messengers 3. Response A. Post-translational modification: involves modification of existing proteins via the phosphorylation (Protein kinases) /dephosphorylation (Protein phosphatases) of specific amino acids (a) Before exposure to light (b) After a week’s exposure to natural daylight ??? B. Transcriptional Regulation – Specific transcription factors bind directly to specific regions of DNA and control transcription of genes 1 https://www.youtube.com/watch ?v=1R_h4itoXYc Plant Hormones in Signaling Plant Hormones • Hormones are chemical signals that coordinate different parts of an organism • In general, hormones control plant growth and development by affecting the division, elongation, and differentiation of cells • Plant hormones are produced in very low concentration, but a minute amount can greatly affect growth and development of a plant organ Plant Hormones in Phototropism Auxin (Indoleacetic acid, IAA) 1. Cell elongation Asymmetrical distribution of auxin moving down from coleoptile tip causes cells on darker side to elongate faster 2. Lateral and Adventitious Root Formation: Auxin is involved in root formation and branching 3. Auxins as Herbicides: An overdose of synthetic auxins can kill eudicots 4. Other Effects of Auxin: Auxin affects secondary growth by inducing cell division in the vascular cambium and influencing differentiation of secondary xylem 2 The Role of Auxin in Cell Elongation Cross-linking polysaccharides Cell wall–loosening enzymes – Growth regulators that stimulate cytokinesis (cell division) 3 Expansins separate microfibrils from crosslinking polysaccharides. CELL WALL 4 Cleaving allows microfibrils to slide. Cellulose microfibril H2O 2 Cell wall becomes more acidic. 1 Auxin increases proton pump activity. Plasma membrane Plasma membrane Nucleus Cell wall Cytoplasm Vacuole CYTOPLASM Cytokinins – Cytokinins are produced in actively growing tissues such as roots, embryos, and fruits Expansin 5 Cell can elongate. – Cytokinins, auxin, and other factors interact in the control of apical dominance to suppress development of axillary buds – Cytokinins work together with auxin to control cell division and differentiation – Cytokinins retard the aging of some plant organs by inhibiting protein breakdown, stimulating RNA and protein synthesis, and mobilizing nutrients from surrounding tissues Gibberellins • Gibberellins have a variety of effects, such as stem elongation, fruit growth, and seed germination • Gibberellins stimulate growth of leaves and stems Gibberellins in seed germination 1 Gibberellins (GA) send signal to aleurone. 2 Aleurone secretes -amylase and other enzymes. 3 Sugars and other nutrients are consumed. • In stems, they stimulate cell elongation and cell division Aleurone • In many plants, both auxin and gibberellins must be present for fruit to set Endosperm -amylase Sugar GA • Gibberellins are used in spraying of Thompson seedless grapes GA Water Scutellum (cotyledon) Ethylene Radicle The Triple Response to Mechanical Stress • Allows a growing shoot to avoid obstacles • The triple response consists of a slowing of stem elongation, a thickening of the stem, and horizontal growth • Plants produce ethylene in response to stresses such as drought, flooding, mechanical pressure, injury, and infection • The effects of ethylene include response to mechanical stress, leaf abscission, and fruit ripening 0.00 0.10 0.20 0.40 0.80 Ethylene concentration (parts per million) 3 ein mutant ctr mutant Ethylene in senescence • Senescence is the programmed death of plant cells or organs • A burst of ethylene is associated with senescence, the programmed destruction of cells, organs, or whole plants • Leaf Abscission A change in the balance of auxin and ethylene controls leaf abscission, the process that occurs in autumn when a leaf falls • Fruit Ripening A burst of ethylene production in a fruit triggers the ripening process (a) ein mutant (b) ctr mutant Ethylene-insensitive mutant Triple response in air but do not respond to inhibitors of ethylene synthesis ABSCISSION Abscisic Acid (ABA) 1. Slows growth • Ethylene induce leaf abscission. 2. Controls seed dormancy: Ratio of ABA to Gibberellins 0.5 mm 3. Drought Tolerance: ABA causes opening of potassium channels in guard cells when plant begins to wilt • Brassinosteroids slow leaf abscission. • Promote xylem differentiation similar to auxin. Protective layer Stem Strigolactones Abscission layer Petiole Responses to Light are critical for plant success • • Stimulate seed germination Establish mycorrhizal associations Help control apical dominance • • – – – • – – – • – – Light cues many key events in plant growth and development. Photomorphogenesis: Effects of light on plant Plants detect not only presence of light but also its direction, intensity, and wavelength (color) Action spectrum Relative response of a process to different wavelengths Blue and Red light most important Action spectra are useful in studying any process that depends on light Blue Light Photoreceptors Phototropism Opening of stomata Hypocotyl elongation Phytochromes (absorb red) Seed germination Shade avoidance 4 Biological Clocks and Circadian Rhythms • • • • • • 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 Circadian rhythms are cycles that are about 24 hours long and are governed by an internal “clock” Circadian rhythms can be entrained to exactly 24 hours by the day/night cycle The clock may depend on synthesis of a protein regulated through feedback control and may be common to all eukaryotes The Effect of Light on the Biological Clock: Phytochrome conversion marks sunrise and sunset, providing the biological clock with environmental cues Photoperiodism and Responses to Seasons • Photoperiod, the relative lengths of night and day, is the environmental stimulus plants use most often to detect the time of year. • Photoperiodism is a physiological response to photoperiod. • Photoperiodism and Control of Flowering. - Short-day plants: Plants that flower when a light period is shorter than a critical length. - Long-day plants: Plants that flower when a light period is longer than a certain number of - Day-neutral plants is controlled by plant maturity, not photoperiod. Phytochromes in Photoperiodism The critical night length 24 hours (a) Short-day (long-night) plant Darkness Light Critical dark period Flash of light -Short-day plants are governed by whether the critical night length sets a minimum number of hours of darkness (b) Long-day (short-night) plant -Long-day plants are governed by whether the critical night length sets a maximum number of hours of darkness Flash of light • • 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 A Flowering Hormone? • Some plants flower after only a single exposure to the required photoperiod • Other plants need several successive days of the required photoperiod • The flowering signal, not yet chemically identified, is called florigen. • Florigen may be a macromolecule governed by the CONSTANS gene. • Still others need an environmental stimulus in addition to the required photoperiod – For example, vernalization is a pretreatment with cold to induce flowering 5 Plants respond to a wide variety of stimuli other than light Meristem Transition and Flowering • For a bud to form a flower instead of a vegetative shoot, meristem identity genes must first be switched on • Researchers seek to identify the signal transduction pathways that link cues such as photoperiod and hormonal changes to the gene expression required for flowering • Because of immobility, plants must adjust to a range of environmental circumstances through developmental and physiological mechanisms • Gravitropism • Thigmomorphogenesis – Thigmotropism • Environmental Stress – Drought – Flooding – Salt – Heat – Cold MECHANICAL STIMULI • The term thigmomorphogenesis refers to changes in form that result from mechanical disturbance • Rubbing stems of young plants a couple of times daily results in plants that are shorter than controls Statoliths (a) Root gravitropic bending • Thigmotropism: Growth response to touch e.g. vines and other climbing plants 20 µm (b) Statoliths settling http://www.allposters.com/-sp/Twining-Stem-of-a-Morning-Glory-Plant-an-Example-ofThigmotropism-Ipomoea-Purpurea-Posters_i6012596_.htm Transmission of electrical impulses: action potentials • Thigmotropism is growth in response to touch • It occurs in vines and other climbing plants • Rapid leaf movements in response to mechanical stimulation are examples of transmission of electrical impulses called action potentials 6 Environmental Stresses • Environmental stresses have a potentially adverse effect on survival, growth, and reproduction • Stresses can be abiotic (nonliving) or biotic (living) • Abiotic stresses include drought, flooding, salt stress, heat stress, and cold stress ABIOTIC STRESSES http://caps.ncbs.res.in/stifdb2/help.html Drought and Flooding • During drought, plants reduce transpiration by closing stomata, slowing leaf growth, and reducing exposed surface area • Growth of shallow roots is inhibited, while deeper roots continue to grow • • • Salt Stress Salt can lower the water potential of the soil solution and reduce water uptake Plants respond to salt stress by producing solutes tolerated at high concentrations This process keeps the water potential of cells more negative than that of the soil solution • Enzymatic destruction of root cortex cells creates air tubes that help plants survive oxygen deprivation during flooding Raffinose family of oligosaccharides Salt-stress sensing and tolerance mechanisms ROS signal transduction pathway under salt stress Na+/H+ antiporter AtNHX1 NSCCs, nonselective cation channels; ROS, reactive oxygen species; CDPKs, calcium-dependent protein kinases; CBLs, calcineurin B-like proteins; CIPKs, CBL-interacting protein kinases; AP2/ERF, APETALA2/ETHYLENE RESPONSE FACTOR; bZIP, basic leucine zipper; NHX, Na+ /H+ exchanger; SOS, SALT OVERLY SENSITIVE. SOS3 (Ca2+ -binding protein), SOS2 (Suc nonfermenting- like) kinase SOS1, plasma membrane Na+/H+ antiporter Trends in Plant Science June 2014, Vol. 19, No. 6 7 Model for the role of signaling factors in stomatal closure and retrograde signaling during water stress Illustration of the response of plants to water stress Front. Plant Sci., 13 March 2014 | http://dx.doi.org/10.3389/fpls.2014.00086 Front. Plant Sci., 13 March 2014 | http://dx.doi.org/10.3389/fpls.2014.00086 Classes of genes that are induced by water-deficit stress Heat and Cold Stress • • • • Excessive heat can denature a plant’s enzymes Heat-shock proteins help protect other proteins from heat stress Cold temperatures decrease membrane fluidity Altering lipid composition of membranes is a response to cold stress • Freezing causes ice to form in a plant’s cell walls and intercellular spaces https://ag.purdue.edu/hla/zhulab/Pages/default.aspx http://www.intechopen.com/ Transcriptional network of abiotic stress responses BIOTIC STRESSES 8 Defenses Against Herbivores • Herbivory, animals eating plants, is a stress that plants face in any ecosystem • Plants counter excessive herbivory with physical defenses such as thorns and chemical defenses (Secondary compounds) such as distasteful or toxic compounds • • Plants damaged by insects can release volatile chemicals to warn other plants of the same species Methyljasmonic acid can activate the expression of genes involved in plant defenses 4 Recruitment of parasitoid wasps that lay their eggs within caterpillars 3 Synthesis and release of volatile attractants • Some plants even “recruit” predatory animals that help defend against specific herbivores 1 Wounding 1 Chemical in saliva 2 Signal transduction pathway Model of the signaling network for plant defense responses Green Leaf Volatile (GLV) biosynthesis Oral factors - effectors - elicitors ACS, 1-aminocyclopropane-1-carboxylate (ACC) synthase; Int. J. Mol. Sci. 2011, 12(6), 3723-3739; doi:10.3390/ijms12063723 CDPKs, Ca2+-dependent protein kinases; GOX, glucose oxidase; Chewing arthropods induce JA-dependent JAs, jasmonates; defense responses, whereas piercing-sucking MAPK, mitogen-activated protein kinase; arthropods frequently induce SA-dependent ROS, reactive oxygen species; defense responses. SA, salicylic acid. Green Leaf Volatiles (GLVs) are emitted during herbivory, pathogen infection and abiotic stress Int. J. Mol. Sci. 2013, 14(9), 17781-17811; doi:10.3390/ijms140917781 Defenses Against Pathogens • A plant’s first line of defense against infection is the epidermis and periderm • If a pathogen penetrates the dermal tissue, the second line of defense is a chemical attack that kills the pathogen and prevents its spread • This second defense system is enhanced by the inherited ability to recognize certain pathogens Int. J. Mol. Sci. 2013, 14(9), 17781-17811; doi:10.3390/ijms140917781 9 Response to attacks by pathogens 1. Plant and Microbial symbiosis 1st line of defense is periderm and epidermis 2. Host-Pathogen Coevolution Rhizobium-legume symbiosis Virulent vs Avirulent Plant-mycorrhizal symbiosis 3. Hypersensitive Response Cause cell and tissue death near infection site to restrict spread of pathogen 4. Systematic Acquired Resistance Plant wide expression of defense genes in response to salicylic acid Trends in Plant Science, March 2015, Vol. 20, No. 3 • Systemic acquired resistance causes systemic expression of defense genes and is a long-lasting response Applied Soil Ecology 85 (2015) 94–113 • Salicylic acid is synthesized around the infection site and is likely the signal that triggers systemic acquired resistance • https://www.youtube.com/watch?v=fl4NhK7AaDQ https://www.youtube.com/watch?v=QYmrOrTM-FA https://www.youtube.com/watch?v=GTH6Ja24SMw Videos • Signal Transduction Pathways https://www.youtube.com/watch?v=HEbueHx_Xag • Do Plants Respond to Music? https://www.youtube.com/watch?v=o54o71wVa2c • How Plants Communicate & Think https://www.youtube.com/watch?v=CsJkNOEwFcI Stuff They Don't Want You to Know - Plant Intelligence https://www.youtube.com/watch?v=KLhgtqd1M2g • Do Plants Respond to Pain? https://www.youtube.com/watch?v=fGLABm7jJ-Y • When Plants Talk https://www.youtube.com/watch?v=GZqB2T9zz7E Plants: Plants' solutions to life's challenges David Attenborough. https://www.youtube.com/watch?v=yOiyHp_oUKA 10