• Plant cell, tissues, organs: basic structure and function
• Transport and translocation of water and solutes • Photosynthesis and Respiration • Plant growth and development • Plant and light response • Plant hormone and plant tissues culture • Plant – environment interactions
1 OUTLINE
• Plant cell, tissues, organs: basic structure and function
• Transport and translocation of water and solutes • Photosynthesis and Respiration • Plant growth and development • Plant and light response
• Plant hormone and plant tissues culture
• Plant – environment interactions
2 Bachelor 2: Plant physiology
Lesson 7: PLANT HORMONE
Instructor: LE Thi Van Anh
le-thi-van.anh@usth.edu.vn 3 Learning outcome By the end of this course, students are able to: - Give definition of “hormone” and the general function of hormone - List major hormones representing in plant - Present the biosynthesis of hormone - Analyze the biological function of auxin, cytokinin and abscisic acid - Describe the process of auxin transport in plant - Analyze the signaling pathway of the 3 hormones: auxin, cytokinin and abscisic acid.
4 Concept Greek horman = to stimulate
Hormone = Substance or chemical that is transported and causes
specific physiological effects sinh lý Plant hormones = phytohormone (phyto = plant): hormone in plant chất điều hòa = plant growth regulators: factors that regulate the growth and development of plant
Features: đặc điểm
- Regulate growth and development - Mobile throughout plant di chuyển khắp cây - Environment and stress response phản ứng 5 Concept: major plant hormones Auxin – Greek: to grow or increase Cytokinin – cytokinesis (cell division) tác nhân gây bệnh Gibberellic acid – pathogen Gibberella Ethylene – chemical brother to ethanol Abscisic acid – abscission sự rụng lá
Brassinosteroids – derived from Brassica spp.
nguồn gốc
6 Concept: transduction pathways
7 Concept: questions to answer…
HOW were they discovered?
WHERE are they synthesized?
WHAT are their biological functions
8 AUXIN: discovery Some sorts of signal: - Are produced in the tips - Travel to elongation zone causing curvature
In 1930s: identification of Idole-3-acetic acid (IAA)
2-Methroxy- IBA 3,6-Dichlorobenzoic acid dicamba 10 AUXIN: site of synthesis
Synthesis in rapidly dividing
and growing tissues: - SAM - Young leaves - Developing fruits and seeds AUXIN: biosynthesis pathway Triptophan-dependent pathway AUXIN: biosynthesis pathway Triptophan-independent pathway AUXIN: polar transport Experiment
Polarity of transport is independent of the orientation of plant tissues:
Transport polarly from “source” to “sink” AUXIN: polar transport
The root always form at
the basal end.
Polar auxin transport is
independent of gravity AUXIN: transporter protein
Influx transport: go in to cells.
Efflux transport: go out from cells
Auxin influx carrier: AUX1 protein
Auxin efflux carrier: PIN1 protein AUXIN: signal transduction pathways AUXIN: signal transduction pathways AUXIN: Biological fuction
- Promote cell elongation
- Promote Shoot growth - Inhibit root growth (at high concentration) - Regulate tropism: phototropism, gravitropism - Regulate apical dominance - Promote formation of lateral/adventitious root - Promote fruit development - Other: delay the onset of leaf abscision; induce vascular differentiation Cell elongation in response to auxin Expansins (active at low pH) cleave cellulose microfibrils from polysaccharides. Exposed Cross-linking Cell wall polysaccharides now accessible to enzymes. cell wall enzymes polysaccharides Expansin tiếp cận CELL WALL Enzymatic Microfibril cleavage of polysaccharides Cell wall H+ allows microfibrils becomes H+ H+ to slide.Cell wall acidic. H+ can extend. Turgor H+ H+ H+ H+ causes the cell to Auxin expand. increases activity ATP H+ Plasma membrane of proton Cytoplasm pumps. 20 Cell elongation in response to auxin
Auxin cause the cellulose loosened, thus the cell can elongate lỏng ra kéo dài 21 Auxin promote shoot and inhibit root growth IAA induction on shoot growth
IAA induction on root growth
22 AUXIN: induce plant phototropism Transformed plant with DR5::GUS
A: no NPA treament B: with NPA treament
There is lateral redistribution of auxin during phototropism
AUXIN: induce gratropism
Root has positive gratropism
Shoot has negative gratropism AUXIN: induce shoot negative gratropism Lower half of agar Upper half of agar
Put horizontal tip next to
an agar block
Auxin is transported to the lower side of a horizontally
oriented oat coleoptile tip AUXIN: induce root positive gratropism Trangenic plant DR5::GFP Symmetric auxin 3h after reorientation of root accumulation in root tip
phân phối Auxin is redistribute to the lower side after reorientation of root định hướng AUXIN: induce root positive gratropism Vertical orientation
The growing apical bud inhibits the growth of lateral buds
This is regulated by auxin Auxin promote the formation lateral/adventitious root
Transgenic Arabidopsis with DR5::GUS
At the 1st stage: auxin is located in ground meristem from which lateral root primordium initiate. Then auxin is translocated toward the root tip of LR Auxin promote fruit development
Achenes produce auxin
that regulate receptacle develop into fruit GIBERRELLIN: discovery
1958: Isolation of GA1 in non infected runner bean
(Phaseolus coccineus)
1930: Bakanae disease GA in is
riceconsidered as one phytohormone caused by Giberrela fujikaroi: tall stem, less grain production
1950: Identification Giberrellin A
(GA) in culture filtrate GA3 induce internodes Separation in to 3 molecules GA1 stimulate stem growth and flowering GA1, GA2 and GA3 elongation in the in cabbage drawrf mutant 31 GIBERRELLIN: chemical structure
GAs are diterpenoid, formed from 4 isoprenoid unit
Each unit consisting of 5 carbons.
32 GIBERRELLIN: chemical structure Isoprene (C5 H8)
Terpene = (C5 H8 )n Polyisoprene_linear form
n = 1: Hemiterpenes n = 2: Monoterpenes n = 3: Sesquiterpenes n = 4: Diterpenes
Polyisoprene_ring form
33 GIBERRELLIN: Biological fuction
- Stimulate stem and root growth
- Promote seed germination - Promote transition from juvenile to adult phase - Influence floral initiation - Promote pollen development and tube growth - Promote fruit set and parthenocarpy GA stimulate stem and root growth Stimulate stem elongation - dramatic for drawrf mutant - no (a little) effect on tall (WT) plant - Dramatic for rosette plant EX: Grass family.
1 2 3 4 1: Cabage in shortday time
2: Cabage in shortday time + GA3 1: Dwarf mutant 2: Dwarf mutant + GA1 3: WT GA action depends on Auxin action 4:WT + GA1 35 Auxin vs GA
36 Auxin vs GA
37 GA promote seed germination 1. After a seed imbibe 2. Aleurone responds by 3. Sugar and other nutrient water, the embryo synthesizing and secreting absorbed from endosperm release GA as a signal digestive enzyme that by cotyledon are consumed to the aleurone, hydrolyze store nutrients in during growth of the endosperm embryo into a seedling
38 GA regulate the transition from juvenile to adult phase
Giant sequoia Giant sequoia 14-week-old
Largest tree (97m height, 8m diameter) Spray GA3 about 8 weeks earlier Oldest plant (3500 years old) Flowering at 70 – 100 years old 2 years for flower maturing 39 GA regulate the transition from juvenile to adult phase
White spruce Stem injected the previous summer with GA4/GA7 mixture in aqueous ethanol
White Spruce WT 25 m height in height Flowering at 12-30 years old 40 GA promote fruit set and parthenocarpy
GA induces growing in Thompson Seedless grapes
GA signaling pathways
WT gid1 GA3-induced elongation of Gid1: GA-insensitive dwarf1 the second leaf sheath
GID1 is GA receptor Figure 20.14 Structure of the GA3-GID1a-DELLA complex Figure 20.15 Model of GA-induced change in the GID1 protein GA signaling pathways
Degradation of the DELLA protein allows
transcriptional reprogramming to take place, and growth, or another GA response, is observed How does GA stimulate stem elongation? How does GA promote seed germination? How does GA promote seed germination? CYTOKININ: discovery 1934: Medium (mineral, vitamin, sucrose) + No hormone: - Root can be grown - Shoot can not be grown 1948: Medium + coconut milk: shoot can be grown
51 CYTOKININ: discovery 1961: First naturally occurring cytokinin found in plants (immature endosperm maize), later called zeatin
52 Figure 21.2 Structures of other aminopurines that are active as cytokinins CYTOKININ: biosynthesis
54 CYTOKININ: biosynthesis pathway
55 CYTOKININ: biosynthesis in tumor CYTOKININ: promote shoot growth Longitudinal section through SAM
AtCKX1, AtCKX2: WT overexpress of cytokinin oxidase
AtCKX1 WT AtCKX1 AtCKX2
Cytokinin promote shoot growth by increasing cell poliferation in SAM
57 CYTOKININ: inhibit root growth
WT AtCKX1 WT AtCKX1
DAPI (4,6 diamindino-2-phenylindole) staining
Cytokinin inhibit root growth by promoting the exit of cells from RAM WT AtCKX2 AtCKX2
58 Auxin/cytokinin ratio regulates morphogenesis in cultured tissues
WT AtCKX2 AtCKX2
59 Auxin/cytokinin ratio regulates morphogenesis in cultured tissues
WT AtCKX2 AtCKX2
60 Auxin/cytokinin ratio regulates morphogenesis in cultured tissues: case studies
61 Auxin/cytokinin ratio regulates morphogenesis in cultured tissues: case studies
62 CYTOKININ: modify the apical dominance
Cytokinin interact with auxin to modify apical dominance and promote the lateral bud growth
WT AtCKX1 AtCKX2
63 CYTOKININ: delay leaf senescence
1: plant expressing IPT gene
2: WT
WT AtCKX1 AtCKX2 1 2 1
64 CYTOKININ signaling pathway Simple versus phosphorelay types of two-component signaling systems CYTOKININ signaling pathway Comparison of the structures of the type-A and type-B ARRs CYTOKININ signaling pathway
WT AtCKX1 AtCKX2 1 2 1
68 ABA: discovery
1940: isolation of a substance from Sycamore leaves called Dormin
1960: confirmation of the dormancy induction by application of
Dormin
1963: discovery of a substance stimulated abscission of cotton fruits,
called Abscisin II
1964, it became evident that dormin and abscisin II were the same plant hormone. Later named abscisic acid (ABA)
69 ABA: chemical structure 15-C Sesquiterpene compound,
Physiological effect is similar to ABA ABA: biosynthesis pathway
FIGURE 23.2 ABA biosynthesis and metabolism.
In higher plants, ABA is synthesized via the terpenoid pathway ABA: transport in plants
- Externally applied ABA : Distributed in all directions
- Cell to cell transport is slow
- ABA synthesized in root cap transported to central vascular
tissues. Can be transported in both xylem and phloem ABA: signal transduction pathways
- ABA can elicit the fast or gradual responses using 3 different
receptors
- There are 3 classes:
+ Plasma membrane localized (G protein)
+ Plastid localized enzyme
+ Cytosolic ligand biding protein
ABA: core signaling pathway
V. C. Dilukshi Fernando and Dana F. Schroeder, 2016
The core ABA signal pathways ABA: Biological function - Hormone of inhibition + promote the dormancy (-GA): seed, bud + promote the abscission (+ethylene)
- Hormone of stress (drought stress)
+ Stomatal closure + Promotes root growth; inhibits shoot growth ABA/GA: Seed dormancy/germination Mutating (from GA mutant back ground) (2)
GA exogenous (1)
GA deficiency GA/ABA deficiency
GA deficiency No germination Germination Germination
Phenotype recovering
Phenotype reverting ABA: germination – GA: germination + After Koornneef et al, 1982 ABA/GA: Seed dormancy/germination
- In most plants: peak of ABA production concides with GA and
IAA levels reduction
à besides ABA, other hormones contribute to dormancy
initiation and maintenance.
- GA promotion of germination requires destruction of DELLA
family protein. ABA/GA: Seed dormancy/germination
GA Stimulates aleurone layer of A-amylase - In most plants: peak cereal grain produced of ABA production concides with GA Inhibits a-amylase mRNA and IAA levels transcription reduction
ABA
à At least 2 mechanism 1. Identify ABA gene expression protein (repress GA gene expression) 2. ABA repress GAMYB Transcriptional factor of a - amylase How does GA promote seed germination? How does GA promote seed germination? ABA closes stomata in response to water stress
ABA closes stomata to prevent water lost ABA regulate shoot/root growth in response to water stress Root development
Which hormone(s)? How do they act? How do they interact? TIR1 Hormone homeostasis: balance/interaction of hormone controlling plant growth and development
