biotstress

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    Biotic Stress and Plant Defense

    Responses (Whos Minding the

    Fort?)

    Land plants are attacked by:

    Herbivores (insects, mammals, etc.)

    Pathogens

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    Pathogen Strategies

    1. Necrotrophic plant tissue killed and then colonized;

    broad host range

    e.g., rotting bacteria (Erwinia)

    2. Biotrophic plant cells remain alive, narrow host range

    (1 plant species)

    e.g., viruses, nematodes, fungal mildews

    3. Hemibiotrophic biotrophic phase, then necrotrophic,

    intermediate host range

    e.g., Phytophthora (potato blight disease)

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    Major Pathogens

    1) Viruses - most are RNA virusesHave a small compact genome which encodes:

    coat protein

    RNA-dependent RNA polymerase (for replication)

    movement protein(s)

    Some plant RNA viruses also encode:

    protease for processing polyprotein (Potyvirus)

    capping enzyme

    RNA helicase

    Some also exist as multiple particles, and a segmentedRNA genome (e.g., Bromoviruses, 3 particles needed forinfection).

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    An encapsulated SSRNA virus: Tobacco

    mosaic virus

    DS DNA virus,CauliflowerMosaic Virus

    Fig. 21.10, Buchanan et al.

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    Fig. 21.11 , Buchanan et al.

    Genome of CauliflowerMosaic

    Virus (CaMV)

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    1a) Viroids- naked RNA infectious

    agents

    Single strands of RNA

    Lack the protein coat of viruses Do not code for any specific protein

    Replicate in the nuclei of infected cells

    e.g., Potato spindle tuber viroid was thefirst to be identified.

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    Viroid structure

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    2) Bacteria3) Fungi - plant pathogens found among 4

    major groups

    4) Nematodes - major root parasites, alsoincrease infection by microorganisms

    Insects and other herbivores can alsotransmit plant diseases.

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    Distribution of Oak Wilt in the US

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    Hill Country Oak

    Leaves from Infected tree

    Fungus - Ceratocystis

    fagacearum

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    Modes of Transmission of Oak Wilt fungus

    Natural root grafts Sap beetle

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    Plant Defenses

    1) Physical barriers: cuticle, thorns, cell walls

    2) Constitutively produced chemicals (e.g., phytoalexins)and proteins (e.g., Ricin)

    1 and 2 together provide non-host resistance againstmost potential pathogens. However, some pathogensget around these protections (but only for certain plants),

    and generate induced responses, which work to limit thedisease.

    3) Induced responses

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    Plant Defense Response

    3 aspects of response:

    1. Hypersensitive

    2. Local

    3. Systemic

    Compatible interaction

    diseaseIncompatible interaction resistance

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    Hypersensitive response

    Rapid - within 24 h

    Not always needed for resistance

    Includes: oxidative reactions (production of hydrogen

    peroxide)

    deposition of callose (related to cellulose)

    opening of ion channels

    apoptosis (programmed cell death)

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    Disease symptoms

    (necrotic lesions

    surrounded bychlorosis) caused by

    the phytobacterium

    Pseudomonas

    syringae pv. tomato

    (Pst) strain DC3000

    on inoculated leavesof a susceptible

    Arabidopsis thaliana

    Col-0 plant. Green

    leaves were not

    inoculated. [Photo byR. Thilmony]

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    Local responses

    Cessation of cell cycle

    Induction of genes that promote resistance

    Phenylpropanoid pathway induced: products include

    salicylic acid (secondary inducer: induces otherpathogenesis-related proteins), lignins (cell wall), and

    flavonoids

    Pathogenesis-related (PR) proteins

    Phytoalexins increased

    Fortification of cell walls with lignin,

    hydroxyproline-rich glycoproteins (HRGPs), etc.

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    Systemic Response or Systemic

    Acquired Resistance (SAR)

    SAR takes 24-48 h to start, can last for months

    Involves gene activation and a transmitted signal.

    Genes induced:

    chitinases

    1,3- glucanases

    other PR proteins

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    Signaling Cascade for Defense

    Responses

    Molecular nature ofelicitors:

    1. Cell wall proteins (e.g., Harpin)

    2. Intracellular proteins (defined geneticallyin a bacterium by cloning avirulent

    loci)

    3. Peptide derived from a larger protein(from a fungus)

    4. Heptaglucan (small oligosaccharide)

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    Secondary Signals (3)

    1. Ca+2 , required for subsequent steps

    May mediate phosphorylation-dephos. events

    involved in transcriptional or post-

    transcriptional gene regulation (there are a

    number of genes whose transcription

    increases, and some decrease)

    Some defense genes also induced by

    blue-UV light or other stresses

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    2. H2O2 (hydrogen peroxide)

    Plays multiple roles:

    induces defense-related genes

    induces apoptosis

    causes cross-linking of cell wall proteins

    (more resistant to wall-degrading enzymes)

    may directly kill pathogens

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    3. Salicylic acid

    required for SAR

    levels increase locally and at distance

    from infection Systemic Signal? Probably not. Still

    unknown.

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    Model derived

    mostly fromstudies in cell

    culture using

    specific elicitors.

    However, there is

    evidence for

    induction in inact

    plants by R

    genes. Some

    aspects are also

    constitutive and

    help block mostmicrobes (non-

    host resistance).