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Lecture 5 Plant hormones and Signal transduction
PLSC 452/552 1/23/14 Department of Plant Sciences Neal Stewart Lab
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Brain teaser Little Johnny nails a sign 90 cm off the ground to a tree outside of his clubhouse that says “no girls allowed.” If the tree grows 12 cm every year, at what height will the sign be when Little Johnny returns as Big Johnny in 15 years?
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Overview Define “hormone” as a functional compound. What is the function and importance of hormones? Know the major plant hormones Understand real-world application of hormones Importance of hormones for tissue culture: introduction
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What is a hormone? Biochemical which regulates growth based on biological and environmental influences Synonyms: Plant hormones, plant growth regulators (PGRs), phytohormones Regulate growth and development Mobile throughout plant Environment and stress responsive
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Major plant hormones Auxin – Greek: auxein; to grow or increase
Cytokinin – cytokinesis (cell division) Abscisic acid – abscission Jasmonic acid – found in jasmine oil Gibberellic acid – pathogen Gibberella Ethylene – chemical brother to ethanol Brassinosteroids – derived from Brassica spp.
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Finding plant hormones
Observational: Darwin stumbles on auxin Noticed grass tips grow toward light With tip growth responded to light Without tip growth had no response Mutation screening: (aka forward genetics) Dwarf plants are can be hormone deficient
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ABRC teaching tools website
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General hormone biochemistry
Present in all cells at various levels Classes of hormones work in signal cascades Hormone-receptor interactions Respond to a host of factors and biological needs Abiotic Water stress Light Nutrient deficiency Biotic Growth Development Herbivore stress
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Hormone biosynthesis Made from four biosynthetic pathways: Terpenoids
AMP + IPP (cytokinins) Carotenoid breakdown (abscisic acid) Diterpene (gibberellic acid) Triterpene (brassinosteroids) Fatty acids (jasmonic acid) Tryptophan (auxins) Methionine (ethylene)
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KEGG database:
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Auxins Greek: auxein; to grow or increase Apical dominance growth
Cell elongation Hormone level very important 2, 4-dichlorophenoxyacetic Acid
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Taiz and Zeiger. 2002. Plant Physiology, 3rd Ed.
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Auxin: Apical dominance
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Cytokinin Cytokinesis (cell division)
Accidently added degraded DNA to medium Organization and development of xylem tissue Response to light Lateral growth of shoots Open stomata Kinetin
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Application: cotton spraying
Adding cytokinins to young cotton increase drought-resistance Arizona – 31% US cotton, highest yield acre-1
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Auxin and cytokinin ratio importance
Auxin alone = Large cells (no division) Cytokinin alone = Cells have no change Auxin + Cytokinin = Normal cell growth and division Auxin + >Cytokinin = Shoot growth >Auxin = Cytokinin = Root growth
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Auxin Cytokinin
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Abscisic acid (ABA) Originally implicated in leaf and fruit abscission
Involved in leaf senescence Maintains seed dormancy (opposed to GA) Involved in stomata regulation (closes) Single hormone unit
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Jasmonic acid First identified in jasmine oil
Response to biotic stress Wounding induces JA biosynthesis Microbial and fungal invasion Plant growth effects similar to auxin Specialty growth structures
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Jasmonic acid: Plant-insect co-evolution
Tri-trophic interactions Degenhardt (2009) Plant Physiology 149:96-102
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Jasmonic acid: Pathogen response
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Gibberellic acid Originally found in Gibberella (rice pathogen)
Responsible for ‘foolish seedling’ phenomenon Uninhibited growth until breaking Involved in cell elongation Flowering and seed germination
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Modification of GA in rice
Miyako Ueguchi-Tanaka 2005 Nature 437,
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Spraying GA for increased fruit yield
GA induces fruiting in absence of seed
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Ethylene: the cell phone of PGRs
A hydrocarbon gas Involved in fruit ripening, stress response Inhibition of growth in dark conditions Excess ethylene inhibits callus growth
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Conversational Plants?
Ethylene production increases during stress Drought Heat Perceived by neighboring plants Unstressed plants induce stress pathways
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Brassinosteroids Stress responses Stem elongation Seed germination
Switchgrass suspension cells have minor amounts of lignin (small amounts of H monolignols) Addition of brassinolide induces normal lignin formation and composition Stem elongation Seed germination Pollen tube growth Cell differentiation control Brassinolide
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Overview of hormone mutants
Cytokinin mutant Brassinosteroid mutant Cytokinin mutant closer look Phenotypes associated with the brassinosteroid, ethylene, cytokinin and abscisic acid receptor mutants. All images are shown against wild type for comparison. (A) The brassinosteroid receptor mutant bri1-301 (bottom) shows a dwarf phenotype. (B) The dark-grown ethylene receptor mutant ein4 (right) lacks the triple response (left; see main text). (C) The triple cytokinin receptor mutant cre1 ahk2 ahk3 (right) shows reduced growth. (D) The root vascular bundle normally consists of xylem, phloem and procambium cell files, whereas in wol (bottom), or in cre1 ahk2 ahk3 (not shown), there are fewer vascular cell lineages, all of which differentiate as protoxylem. The vascular bundle is encircled by a layer of pericycle cells (asterisks) (E) The abscisic acid receptor mutant fca (right) flowers later than wild type, as demonstrated by increased leaf number at flowering. Scale bars: in C, 2 mm; in D, 10 μm. [A is reprinted, with permission, from Cano-Delgado et al. (Cano-Delgado et al., 2004); B is reprinted, with permission, from Hua et al. (Hua et al., 1998); E is reprinted, with permission, from Amasino (Amasino, 2003).] Ethylene mutant grown in dark Abscisic acid mutant Bishopp A et al. Development 2006;133:
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Hormones (PGR) pathways
PGRs work in complex cascades to produce signals Synergistic Anatognistic Variety of actions in cell Gene transcription Protein degradation
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Arabidopsis histidine kinase sensing and signaling; 2) A histidine phosphotransferase protein nuclear translocation; 3) A response regulation transcription activation; and 4) a negative feedback loop through cytokinin-inducible ARR gene products.
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Introduction: the importance of manipulation of hormones for tissue culture
Auxins and cytokinins are very important Higher auxin induces root growth Equal ratio induces callus and cell enlargement Lower auxin induces shoot growth Gibberellic acid germinates difficult seeds Add brassinosteroids for cell wall induction
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Lecture summary Hormones are biochemicals that regulate plant growth based on biological and environmental cues Auxin and cytokinin are key for plant growth Abiotic and biotic stress response is regulated by hormones Cell signaling is regulated by specific receptors on cell membranes
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