5. Introduction
Growth
“Irreversible change in
mass, i.e. increase in
size, volume and weight
of any part of plant’s
body”
It means
quantitative increase
in body.
Cell division, Cell
enlargement.
Development
“Irreversible change in state”
It means thequalitative
change in plant body.
Seed, Seedling,
Vegetative, maturation,
Flowering.
Development is phase to
phase process.
6. Plant’s growth and development are under the control of two
sets of internal factors.
Chemical messengers
Proper utilization of these raw materials is under the control
of certain “chemical messengers” which can be classified
into hormones and vitamins.
Introduction
Nutrients
Nutritional factors such as the supply of carbohydrates,
proteins, fats and others constitute the raw materials required
for growth.
► Nutrients
► Chemical messengers
7. The site of synthesis is different
from the site of action.
Plant hormones are
physiologically active.
Vitami
n
Vitamins are used in the same
part without being transported
Vitamins by themselves are
physiologically inactive.
They act as co-factor of enzyme.
Introduction
The term Hormone is derived from a Greek root ‘hormao’ which means ‘to stimulate’
( Beylis and Starling, 1902).
Thimann (1948) suggested using the term ‘Phytohormone’ for Hormones of plant.
Hormone
8. Phytohormones are organic substances produced Naturally by theplants
which in minute/low concentration
increase,
Decrease,
modify the growth and development.
1) Plant growth hormones
2) Plant growth promoting substances
3) Plant growth regulators
Phytohormones
Other names
9. synthetic substances influence
Plant Growth regulators (PGR) refers to natural or
the growth and development.
• All plant hormone are plant growth regulators
but,
• All plant growth regulator are not plant hormones
Plant Growth Regulators
10. 1. Natural hormone
Produced by some tissues in the plant. Also called Endogenous hormones.
e.g. IAA.
2. Synthetic hormone
Produced artificially and similarto natural hormone in physiological
activity. Also called Exogenous hormones. e.g. 2,4- D, NAA etc.
3. Postulated hormone
Also produced spontaneously in theplant body, buttheir structure and
function is not discovered clearly. e.g. Florigen, Vernalin.
Classification of PGR’s
On the Basis of Origin
11. Growth promotors (Increase growth of plants):
Auxins
Gibberellins
Cytokinins
Classification of PGR’s
Stress and other hormones:
Jasmonates
Salicilic acids, etc
On the Basis of Nature of Function
Growth Inhibitors (Retard growth of plants)
ABA
Ethylene
13. JASMONIC ACID
First isolated in culture filtrate of fungi
Lasiodiplodia theobromae.
Aldridge et. al.,(1971).J.Chem.Soc.Chem. Comm. 1623-
Jasmonates are cyclopentanone compound or plant
hormones derived from α-linolenic acid.
It includes group of oxygenated fatty acids
collectively called oxylipins and Jasmonic Acid is a
precursor to different compounds to this group.
Methyl jasmonate was first isolated from the essential
oil of Jasminum grandiflorum
Demole et. al.,(1962).Helv.
Chim.Acta.45:675-695
They are ubiquitous in plant kingdom and are also produced
by
certain fungi.
14. A
E
A. Structural formula:
Jasmonic acid (JA)
A. Chemical Formula:
C12H18O3
B. Molar Mass: 210.27 g/mol
C. Boiling point: 160 °C
D. Density: 1.1 g/cm³
E. Jasmin plant:
Jasminum grandiflorum
15. BIOSYNTHESIS OF JASMONATES
Jasmonic acid synthesized from fatty acid(α-
linolenic acid)
Lipoxygenase (LOX), AOS (Allene oxide synthase),
AOC (Allene Oxide Cyclase) are key enzymes of JA
biosynthesis in Chloroplast, and they form OPDA
(oxo-phytodienoic acid).
OPDA is transported to peroxisome through ABC
transporter COMATOSE (CTS)
Reduction of cyclopentanone ring of OPDA is
catalyzed by peroxisomal OPR enzyme
Three cycles of β-oxidation occurs to
give
finally Jasmonic Acid. Enzymes involved are:
•
•
•
ACX1 (Acyl-CoA oxidase in tomato)
MFP (Multifunctional Protein)
KAT (L-3-ketoacyl CoA thiolase)
The JA and its metabolic derivatives
are
collectively called JASMONATES
Fig : Biosynthetic Pathway of Jasmonic Acid
CTS
Source: Wasternack et .al.,(2002).PNARMB.72:165-221
16. Derivatives of Jasmonic acid – THE JASMONATES
Carboxylic
conjugated
acid side chain
to ACC(1-
amino
cyclopropane-1-carboxylic acid)
Methylatedform by JA
methyltransferase
Decarboxylated to cis-jasmone
Conjugated to AA such as Ile
by JA amino acid synthase (JA-
IIe - Jasmonic Acid-Isoleucine)
Reduction
of
keto group of
pentanone ring to cucurbic acid
side
in
position
chain
C-
11
Pentenyl
hydroxylated
and C-12
Fig : Different Metabolites Produced from Jasmonic
Acid
Jasmonic Acid-Isoleucine
17. S. No Process Putative Signals Alteration/Species
1 Root Growth JA,JA-Ile Inhibition
2 Seed Germination JA Inhibition
3 Tuber Formation JA-OH,JA Induction/Potato
4 Tendril Coiling OPDA Stimulation/Bryonia
5 Nyctinasty JA Stimulation/Albizzia
6 Trichome formation JA Induction/Tomato
7 Senescence JA Stimulation
8 Flower Development
Anther Development
and dehiscence
Female Organ
Development
Filament elongation
JA
Induction/Arabidopsis
Induction/Tomato
Induction/Arabidopsis
12 Biotic Stress JA, JME,OPDA Resistance/Ubiqutitous
DIFFERENT
PHYSIOLOGICAL
ROLES
OF
JA
Source:
Wasternack, C.
(2007).Annals
of
Botany.100:681-
697
18. STRESS AND ITS TYPES
STRESS
BIOTIC ABIOTIC
Fungi
Bacteria
Virus
Nematodes
Light
Pesticide
Heavy metal
Drought Salinity Temperature
Herbivores
Stress is any change in the environmental
condition that may adversely affect the
plant’s growth, development and
adaptability
20. ROLE
OF
JA
IN
RESISTANCE
1. Increased endogenous level of JA on treatment of
Arabidopsis with necrotrophic fungus A. brassicola. Penninckx et
al.(1996). Plant Cell.8:2309-2323
2. Biosynthetic necrotrophic and SignalingArabidopsis
mutants were to attack more sensitive fungal pathogen Phytium
by mutants and necrotrophic bacteria Erwinia carotovora
(jar1,coi1) and fad3-2fad7-1fad8 (coi1 mutants). Staswick et al.,
(1998).Plant Journal 15:747-754; Vijayan et al.,(1998).PNAS
USA 95:7209-7214
3. Transgenic plants overexpressing JMT (jasmonic acid carboxyl
methyltransferase) and thus higher levels of methyl jasmonate
more resistant to necrotrophic pathogen Botrytis cinera. Walling
(2000 ). Journal of Plant Growth Regulators 19: 195-216
4. The fad3-2fad7-1fad8 mutant more sensitive to attack by
fungus Bradysia and coi1 is more sensitive to Diamond –back
moth. McConn et al.,(1997). PNAS USA.94:5473-5477; Xie et al
(1998). Science 280: 1091-1094
Source: Berger,(2002).Planta 214:497-504
21. Mutant Biochemical
Significance
Role of
Mutant
Expression of
JA responsive
genes
Effects
coi Coronatine
insensitive
JA Signaling Reduced Sensitivity to insect and necrotrophic
pathogens increased
fad Defective Fatty
Acid Desaturase
JA
Biosynthetic
Reduced Sensitivity to insect and necrotrophic
pathogens increased
cev Constitutive
Expression of VSP
JA Signaling Increased Increased sensitivity to biographic
pathogens
jar JA Resistant JA
Signaling
Reduced Increased sensitivity to necrotrophic
pathogens
opr Defective OPDA
Reductase
JA
Biosynthetic
Reduced Increased sensitivity to insects
jin JA Insensitive JA Signaling Reduced Increased sensitivity to pathogens
Some mutants of JA and Resistance in plants
22. Source:Farmer and Ryan(1992).The Plant Cell.4:129-134
Mechanism of action of JAs in biotic stress
Source:Wasternack (2007).Annals of Botany.100:681-697
23. OVERALL MECHANISM OF SYSTEMIC RESPONSE
Wounded leaves produce an 18-amino acid peptide called systemin from
carboxyl terminal of prosystemin (200 AA precursor) in PP and it elicits
production of JA in companion cell-sieve element complex.
JA moves throughout the plant in the phloem. Covalently modified JA (JA-x)
play important role in systemic signaling.
signal is recognized at the target cell e.g. mesophyll (leaf)
Jasmonic acid turns on defense related genes (genes for proteinase inhibitor
etc.) in target cells.
Source: Schilmiller and Howe (2005). Current Opinion in Plant Biology . 8:369-
25. Fig: Amplification of wound signaling of tomato
Wounding
Systemin
AOC
OPDA/JA
Systemin
Prosystemin
Prosystemin
Amplification of wound signaling is a major
event in systemic defense
Systemin activates
AOC
which in turn
activates Prosystemin through OPDA/JA.
OPDA/JA again activates back the AOC.
In systemic response once JA is synthesized,
its cyclic production of JA can occur by
positive feedback mechanism
JA is a systemic signal that leads to
systemic expression of genes encoding
Proteinase inhibitors(PINs)
The plant may develop some
resistance against subsequent herbivore
attack.
Thus JA signaling is necessary in systemic
response in leaf.
AMPLIFICATION OF SYSTEMIN SIGNAL
Source:Wasternack et al(2006).163: 297-
26. SYSTEMIC RESISTANCE AND ROLE OF JAs
Initial infection result in formation of necrotic lesions
SA concentrations increase and methyl-SA is released in distal plant
tissue
PR proteins in the non-invaded parts of the plants are synthesised
resulting in reduction in disease symptoms after subsequent
infection of many pathogenic species.
Source:Harman et. al.(2004) .Nature Reviews Microbiology. 2:43-56
SAR:
Systemic resistance
in plants
Systemic Acquired Resistance (SAR)
Induced Systemic Resistance (ISR)
ISR:
Non pathogenic root colonizing rhizobacteria e.g. P. fluorescens or
wounding initiates ISR
ISR does not depend on SA and PR protein
ISR requires both JA and ethylene signalling
Systemin is involved in ISR which is absent in SAR.
27. TRANSCRIPTIONAL REGULATION OF JA
Fig: Transcription factors involved in signalling pathways of JA
Source: Wasternack(2007). Annals of Botany100:681-697
Four major of interacting players of:
1.a JA signal
2.the SCF-type E3 ubiquitin ligase SCFCOI1
3.Jasmonate ZIM-domain (JAZ) repressor proteins that are
targeted by SCFCOI1 for degradation by the ubiquitin/26S
proteasome pathway ,and
4.transcription factors (TFs)
Howe et. al.(2008)Current Opinion in Plant Biology 11:428-235
Transcriptional regulation mechanism:
1.Expression of regulatory TFs for JAs
2.Cross talk with the TFs responsible for expression of
other hormones.
28. Cross Talk with Ethylene and JA
•The antagonistic action of MYC2 and ERF1
may cause independence between wound
signaling and pathogen defence signaling
Lorenzo et al (2004)Plant Cell 16:1938-1950)
•JA alone induce the expression
of AtMYC2 responsible for the activation of
wound response genes and for the repression
of pathogen response genes.
signals through the
induction
•The cooperation of ET and JA
of
ERF1 leads activation of PR genes and to
the repression of WR genes
•Therefore, the interplay between ERF1 and
AtMYC2 allows the plant selection of the
correct set of genes in response to these
two stresses
Schematic Representation in response to Pathogens and Wounding
Source: Wasternack(2007). Annals of Botany100:681-697
33. Why we study JAs? / Relevance of JAs studies
Plant lack an immune system like in animals but posses
mechanism that recognizes potential pathogens and initiate
defense responses. During their biochemical evolution, the
plants are devised with certain magic molecules of defense
(secondary metabolites) like JAs.
Recent insights into the JAs mediated plant defense cascade and
knowledge of key regulators of this will help us to design future
crops with increased biotic stress resistance and better
adaptability.
Higher crop yields might be achieved by increasing the
pathogen/insect resistance which can be achieved by manipulating
the expression of the key genes involved in JAs biosynthesis and
signaling cascades.