Biotic stress: microbial

pathogens
HBC 206 Plant Biochemistry

Dr Fiona C Robertson
Department of Biochemistry
University of Zimbabwe
Fighting for their lives: Plants
and pathogens

Photos courtesy of CIMMYT, Scott Bauer, USDA Agricultural Research Service, IRRI, IRRI, IITA
 Plant pathogens are infectious agents
that make plants sick
Plant pathogens cause
10 – 30% yield losses
annually

Up to 25% of plant genes

respond to pathogen
infection

There is an arms race

between plants and
pathogens
Most plants are resistant to most
pathogens -
disease is the exception, not the rule
Many kinds of organisms cause
plant disease

Nematodes
are large,
multicellular Fungi and
animals oomyctes are
eukaryotes
Viruses are non-cellular,
and merely packaged
nucleic acids Bacteria are
prokaryotes

Large, E.C. 1940. Advance of the Fungi. Jonathan Cape, London.

 What makes an interaction into a
disease?
Plants are exposed to countless microbes but very
few of these interactions lead to disease.
Why?
The host plant
must be
susceptible to the
The pathogen must The disease pathogen
be able to overcome triangle
plant defenses (it takes three)

Environment
The environment must
tip the balance in favour
of the pathogen
 Strategies of pathogenicity
A successful pathogen must:
Gray mold (Botrytis
• Find the host and attach to it cinerea)
• Gain entry through the plant’s
impermeable defences
• Avoid the plant’s defence
responses
• Grow and reproduce
• Spread to other plants
Clemson University - USDA Cooperative Extension Slide Series, Bugwood.org
 Pathogens must be able to penetrate
or circumvent physical barriers
Appressorium

Appressorium

Some
pathogens
enter through
wounds or
Some pathogens
stomata
physically pierce
plant tissues or
use enzymes to
digest cell walls
 Pathogens are biotrophs, necrotrophs
or hemibiotrophs

Necrotrophs
kill cells and Hemibiotrophs
then Biotrophs live can switch from
consume the within host biotroph to
contents tissue without necrotroph
causing death

Reprinted by permission from Macmillan Publishers Ltd: Pieterse, C.M.J., Leon-Reyes, A., Van der Ent, S. and Van Wees, S.C.M.
(2009). Networking by small-molecule hormones in plant immunity. Nat Chem Biol. 5: 308-316, copyright 2009.
 Plant immune responses

Plants resist pathogens through active processes that

include recognition of the pathogen and defence
responses to fight it

Perception Signal transduction Response

 How are pathogens recognized?
Pattern recognition receptors (PRRs)
Bacterial pathogen

PRRs recognize
pathogens outside
the cell and initiate
defence responses
Fungal or
oomycete
pathogen

Perception Signal transduction Response

Pathogen recognition triggers defence
responses

Reactive
Calcium ion oxygen
influx production
This response is
Kinase cascade
called pattern
leading to
transcriptional
triggered
responses immunity (PTI)

Defense
responses

Perception Signal transduction Response

 Phytoalexins and phytoanticipins are
chemical defences
Antimicrobial compounds help Phytoanticipins and
ward off pathogens; they can be phytoalexins are secondary
preformed (phytoanticipins) or metabolites, and can be
induced (phytoalexins) highly variable amongst
species
 Callose, ROS and phytoalexins can
arrest pathogen attack
Pathogen

Callose, a
Callose
Cell wall
polysaccharide,
Plasma
acts as a
membrane
barrier, and
Perception
callose ROS and
secretion phytoalexins
are toxic to
pathogens
Signal

Mellersh, D.G. and Heath, M.C. (2001). Plasma membrane - cell wall adhesion is required for expression of plant defense responses during fungal penetration. Plant Cell. 13: 413-424.
 BUT Pathogens can produce effectors
that enhance their virulence
Plant
Microbial proteins
effectors
suppress the Reactive
oxygen
plant’s immune Calcium ion production
response influx

Kinase cascade
leading to
transcriptional
responses
 Plant resistance (R) proteins can
counter-act the effectors

R proteins recognize effector

effectors
intracellularly
R protein

Defense
R protein responses
 R protein activation leads to
enhanced triggered immunity: ETI
ETI is faster, stronger and
more prolonged than PTI

effector

R protein
Defense
responses

Transcription
responses
 Enhanced triggered immunity
Activated R proteins signal danger and trigger a
heightened defence response that includes:

• Production of the stress hormone salicylic acid

(SA)
• Production of reactive oxygen species (ROS)
• The hypersensitive cell death response (HR)
• Expression of pathogenesis-related (PR) proteins
• Systemic signals and systemic acquired resistance
(SAR)
 Sterol-binding activity of PR-1
contributes to its antimicrobial activity
• PATHOGENESIS-RELATED 1 (PR-1) protein was identified
50 years ago as a small protein induced in response to
pathogens
• Its mode of action had remained obscure until last year
(2017)
• Researchers figured out that PR-1 could bind sterols
• Phytophthora oomycetes are unable to synthesize
sterols and rely on their uptake from their growing
media
• PR-1 can inhibit growth of Phythophthora brassicae
through direct removal of sterols from pathogens
 Systemic acquired resistance (SAR)
involves a mobile signal
Systemic
Uninfected tissues show response

enhanced resistance to SAR

subsequent pathogen
challenge. The nature of
the mobile signals are still

Signals
being debated
Defense
responses

Local response
Summary - PTI is suppressed by effectors,
which sometimes trigger ETI
PAMP-triggered Effector triggered Effector-triggered
immunity suppression immunity

Defense Defense
Responses Defense
Responses Responses

R proteins
Effectors suppress recognize
PTI effectors and
induce ETI

Adapted from Macmillan Publishers Ltd: Pieterse, C.M.J., Leon-Reyes, A., Van der Ent, S. and Van Wees, S.C.M. (2009). Networking by small-molecule hormones in plant immunity. Nat Chem Biol. 5: 308-316,
 Breeding Pathogen resistant crops
If we can discover the R genes that lead to resistance
we can breed/engineer crops that are resistant
 Plant responses to necrotrophic
pathogens
There are no known R genes that confer resistance to
necrotrophic pathogens, therefore breeding resistance to
necrotrophs is more challenging than to biotrophs

Arabidopsis plants infected with the

fungus Botrytis cinerea

The oomycete Pythium

on cucumber
Photo credits: Florida Division of Plant Industry Archive, Florida Department of Agriculture and Consumer Services, David B. Langston, University of Georgia, Bugwood.org; Gary Loake
Plant responses to Plant Viruses
Helical Rigid Rod - TMV
Twin virions – Cassava Mosaic Virus
 Plant responses to viruses are largely
mediated by the hypersensitive response
and by siRNAs
Plant defences against viruses
largely occur at the level of
RNA-mediated silencing, and
through the hypersensitive
response to kill infected cells

Waterhouse, P.M. and Fusaro, A.F. (2006). Viruses face a double defense by plant small RNAs. Science. 313: 54-55; Reprinted by permission from Macmillan Publishers Ltd Lam, E. (2004) Controlled cell death, plant
survival and development. Nat. Rev. Mol. Cell Biol. 5: 305 – 315.
RNA silencing of a virus
viral dsRNA

Dicer

siRNA

RISC
Viral RNA silencing suppressor proteins
Just as bacteria and fungi have effector proteins
viral dsRNA
that can block plant defence responses
Viruses have silencing suppressor protiens that
can block RNA silencing

Dicer
Viral suppressor protein

Viral suppressor protein

siRNA

RISC
Viral suppressor protein
 Genetically engineered resistance to
papaya ringspot virus
A gene encoding a viral coat
protein was introduced into
papaya. The modified plants are
resistant to viral infection. This was
the first commercialized transgenic
fruit

Papaya ringspot virus symptoms

In the 1980s, PRSV nearly

wiped out papaya production in
Hawaii. Now most of the
papaya produced in Hawaii are Susceptible Resistant
genetically resistant

Gonsalves, D., S. Tripathi, J. B. Carr, and J. Y. Suzuki. 2010. Papaya Ringspot virus. The Plant Health Instructor. DOI: 10.1094/PHI-I-2010-1004-01;Photo credits: S. Ferreira.
 Summary - Plant immune responses
Plants have strong and vigorous immune responses

Plants can recognize Defence responses

• pathogen- include
associated • production of
molecular antimicrobial
patterns compounds
• some effector • the hypersensitive
molecules Defense response
• Viral dsRNA Responses • systemic acquired
resistance
• Viral RNA silencing
 Strategies to prevent and manage
disease

Make the plant

resistant through
genetic or other
Avoid or eliminate the The disease methods
pathogen triangle
(it takes three)

Environment
Manipulate the
environment to favor the
plant