Cytokinins Regulators of Cell Division. Cytokinins: Other cytokinins are aminopurines. Zeatin = adenine derivative. Physiological roles of Cytokinins

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1 Cytokinins Regulators of Cell Division 1. Discovery 2. Chemical structure 3. Physiological Roles 1913 Haberlandt demonstrates water soluble substance in vascular tissue stimulates cell division in potato tubers s White et al make callus. Auxin + 10% coconut milk supported cell division of mature, differentiated cells. Cytokinins: Discovery Observations on stimulation of cell division 1950 s Skoog & Miller discover kinetin (6-furfurylaminopurine) in autoclaved herring sperm DNA Kinetin A substituted amino purine Cytokinins Discovery of a Natural Cytokinin: Zeatin 1973 Letham isolates trans-6-(4-hydroxy-3-methylbut-2- enylamino)purine from immature corn endosperm. Zeatin = adenine derivative Cytokinins: Other cytokinins are aminopurines Adenine = an amino purine Benzyladenine is a synthetic cytokinin Thidiazuron ( Freefall ) an herbicide and defoliant in cotton. Synthetic Cytokinins Physiological roles of Cytokinins Induce cell division Promote shoot formation in callus cultures (when auxin is present) Delays leaf senescence Promote expansion of dicot cotyledons In defoliants: Dropp & Freefall (diphenyurea type cytokinins) 1

2 Cytokinin oxidase rapidly degrades cytokinins Cytokinins promote cell division in shoots 1. Ck favors meristem growth 2. Localized expression of ipt gene in transgenic plants results in ectopic meristem formation These Overexpress Cytokinin oxidase Wild type Shoot meristem Isopentenyl adenine Overexpression of Cytokinin oxidase Wild type Cytokinins inhibit root growth Overexpression of CK oxidase in roots 1. Reduced CK levels result in more roots larger root apical meristems 2. Increased root development Cytokinins and Auxins regulate Morphogenesis 0.0 Roots Develop Wild Type Kinetin (mg/l) 0.2 Wild Type CK Deficient CK Deficient 1.0 Buds Develop IAA (mg/l) Cytokinin Synthesis is not restricted to plants Agrobacterium tumefaciens crown gall tumor Ti plasmid expresses IAA & CK synthesis genes 21.4 Tumor induction by Agrobacterium tumefaciens Corynbacterium fascians Witches-Broom Root Knot Disease Root-knot nematode damage to carrot Plasmid DNA has genes for 1) IAA synthesis 2) Cytokinin synthesis 3) Opine synthesis 2

3 Effect of Ti Plasmid DNA mutations on Crown Gall Tumor Development Leafy/shooty tumor has mutated tms (IAA synthesis) gene. Part of Ti Plasmid Genome ipt Cytokinins retard senescence Transgenic tobacco with ipt gene and senescence promoter remains green and photosynthetic. Promoter drives expression of isopentenyl transferase (IPT) gene IPT controls first step in CK biosynthetic pathway. Rooty tumor has a mutated ipt (cytokinin synthesis) gene IPT adds isopentenyl group to adenosine Transgenic plant Normal plant Cytokinin promotes movement of nutrients Cytokinin-induced nutrient mobilization Cytokinin promotes leaf expansion Mechanism of CK leaf expansion? Radish cotyledons 1. Involves cell wall loosening 2. No proton extrusion 3. No increase in dry matter 4. No IAA or GA effect Control 3 days 3 days + zeatin Summary Observations of Cytokinin- Overproducing Transgenic Plants END CYTOKININS Shoot apical meristems have more leaves Leaves are greener Leaf senescence is retarded Adventitious shoots form spontaneously Apical dominance is greatly reduced Plants are stunted with short internodes Rooting and root growth are reduced 3

4 Molecular basis of Cytokinin action. 1.Cytokinin increases expression of response regulator genes. 2.Phosphorylate two-component signaling system 1. CK binds to CRE1 2. Histidine kinase is activated Molecular basis of Cytokinin action 6. Genes encoding type A ARR are activated to make Type A ARR proteins. 3. Phosphate transfer to Histidine in AHP protein Hybrid sensor histidine kinase Arabidopsis histidine phosphotransfer protein 4. AHP protein phosphorylates aspartate on receiver domain of ARR proteins 5. Output domain of ARR is a transcription factor induces transcription of Type A ARR genes 7. Phosphorylated type A ARR interact with effectors for CK response Cytokinins: Metabolically inactive forms Sites of Cytokinin synthesis Ribose Here = Ribosylzeatin a ribotide Glycosides form at the 3, 7 or 9 Nitrogen of the purine ring or to the oxygen of dihydrozeatin Roots and they are transported in xylem exudate Embryos Developing leaves Young fruits Adenine 3-Methyl-2-butenal Cytokinins regulate Cell Division Cytokinins are not restricted to plants Overexpression of CK oxidase gene results in inhibited shoot growth. Elevated endogenous CK levels causes KNAT1 and STM genes overexpression leading to ectopic meristem formation. Corynbacterium fascians Witches-Broom Agrobacterium tumefaciens crown gall tumors & Ti plasmid Wild type Normal Transgenic CK-oxidase overexpression Root-knot nematode damage to carrot 4

5 Cytokinins regulate cell division STM gene 1. allows proliferation of stem cells 2. prevents stem cell conversion to organ cells Absence of stem cell proliferation Apical meristem stem cell depletion No apical meristem, no growth! Normal shoot apex stm mutant Cytokinins modify apical dominance IAA inhibits apical dominance Cytokinins initiate lateral bud growth Evidence for this comes from 1. Direct application of CK 2. CK overproducing tobacco mutants Wild type has one shoot CK mutant overproducers are bushy STM exression in SAM L1, L2, Peripheral zone & Central zone. STM normal stm mutant END Cytokinins 5

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