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1. A detailed study of plant- microbe interaction should provide sustainable practical solutions for the control of plant diseases in agricultural crops 2. Such studies will help in elucidate the signaling mechanism by which plant cells cope with a stress situation 3. Study can lead us to discover how organisms from different kingdom communicate with one another
Plant pathogens
Fungi
The majority of phytopathogenic fungi belong to the Ascomycetes and the Basidiomycetes. The fungi reproduce both sexually and asexually via the production of spores. These spores may be spread long distances by air or water, or they may be soil borne. Many soil borne spores, normally zoospores, are capable of living saprotrophically, carrying out the first part of their lifecycle in the soil. Fungal diseases can be controlled through the use of fungicides in agriculture, however new races of fungi often evolve that are resistant to various fungicides
Fungi
Biotrophic fungal pathogens colonize living plant tissue and obtain nutrients from living host cells. Necrotrophic fungal pathogens infect and kill host tissue and extract nutrients from the dead host cells. See Powdery Mildew and Rice Blast images below.
Powdery mildew, a Biotrophic Fungus
2.Basidiomycetes
Rhizoctonia spp. Phakospora pachyrhizi (causal agent of soybean rust) Puccinia spp. (causal agents of severe rusts of virtually all cereal grains and cultivated grasses) Cont
Cont
3. Oomycetes
The oomycetes are not true fungi but are fungal-like organisms. They include some of the most destructive plant pathogens including the genus Phytophthora which includes the causal agents of potato late blight and sudden oak death. Despite not being closely related to the fungi, the oomycetes have developed very similar infection strategies and so many plant pathologists group them with fungal pathogens. Significant oomycete plant pathogens Pythium spp. Phytophthora spp.; including the causal agent of the Great Irish Famine (18451849) late blight disease in potato-an epidemic in Ireland-leads to emigration of more than one million people to US and other countries
Bacteria
Most bacteria that are associated with plants are actually saprotrophic, and do no harm to the plant itself. However, a small number, around 100 species, are able to cause disease. Bacterial diseases are much more prevalent in sub-tropical and tropical regions of the world. Crown gall disease caused by Agrobacterium Most plant pathogenic bacteria are rod shaped (bacilli). In order to be able to colonize the plant they have specific pathogenicity factors. Five main types of bacterial pathogenicity factors are known.
Nematodes
Nematodes are small, multicellular wormlike creatures. Many live freely in the soil, but there are some species which parasitize plant roots. They are a problem in tropical and subtropical regions of the world, where they may infect crops. Potato cyst nematodes (Globodera pallida and G. rostochiensis) are widely distributed in Europe and North and South America and cause $300 million worth of damage in Europe every year. Root knot nematodes have quite a large host range, whereas cyst nematodes tend to only be able to infect a few species. Nematodes are able to cause radical changes in root cells in order to facilitate their lifestyle.
Protozoa
There are a few examples of plant diseases caused by protozoa. They are transmitted as zoospores which are very durable, and may be able to survive in a resting state in the soil for many years. They have also been shown to transmit plant viruses. When the motile zoospores come into contact with a root hair they produce a plasmodium and invade the roots.
Parasitic plants
Parasitic plants such as mistletoe and dodder are included in the study of phytopathology. Dodder, for example, is used as a conduit for the transmission of viruses or virus-like agents from a host plant to either a plant that is not typically a host or for an agent that is not graft-transmissible.
Epidemiology-Disease resistance
Plant disease resistance Management 1. Quarantine 2. Cultural 3. Plant resistance 4. Chemical 5. Biological 6. Integrated
Quarantine
Wherein a diseased patch of vegetation or individual plants are isolated from other, healthy growth. Specimens may be destroyed or relocated into a greenhouse for treatment/study. Another option is to avoid introduction of harmful non-native organisms by controlling all human traffic and activity although legislation and enforcement are key in order to ensure lasting effectiveness.
Plant resistance
Sophisticated agricultural developments now allow growers to choose from among systematically crossbred species to ensure the greatest hardiness in their crops, as suited for a particular region's pathological profile. Breeding practices have been perfected over centuries, but with the advent of genetic manipulation even finer control of a crop's immunity traits is possible. The engineering of foodplants may be less rewarding however, as higher output is frequently offset by popular suspicion and negative opinion about this "tampering" with nature
Chemical
Many natural and synthetic compounds exist that could be employed to combat the above threats. This method works by directly eliminating disease-causing organisms or curbing their spread; however it has been shown to have too broad an effect, typically, to be good for the local ecosystem. From an economic standpoint all but the simplest natural additives may disqualify a product from "organic" status, potentially reducing the value of the yield.
Disease development
Fungal plant pathogens use a wide range of pathogenesis strategiesNechrotrophic, enzymic attack, toxin production
e.g.Pythium,Botrytis, & Cochliobolus carbonum(maize fungal pathogen)HC-toxin inhibits histone deacetylase (activates plant defense gene) AAL toxin(PCD in tomato plants Fusicoccin toxin targets PM_H+-ATPase leads to irreversible stomata opening & plant wilting followed by PCD and necrosis Biotrophic-haustorial penetartion e.g.downy and powdery mildew ,Cladosporium fulum(-nce of haustoria)-grow out side the PCW(apoplast) survive on leaked nutrients Hemitrophic-phytophtera infestans(late blight disease in potato)
Cont.
These interactions can be conceptualized as a continuous sequence or cycle of biological events including dormancy, reproduction, dispersal, and pathogenesis. Although plant pathologists have long realized the importance of these events, Gaumann was among the first to critically evaluate the progression of events leading to disease .Gaumann called this unchanging sequence of events the infection chain, but more recently the terms disease cycle or infection cycle have come into common usage.
1.Model application
Plant disease prediction models have great potential to help meet the need for new management strategies and thus aid in maintaining global food supplies. However, the value of disease prediction models can only be fully realized when these systems are used to make decisions concerning plant disease management in production fields . Most past reviews of plant disease prediction models indicate that far more models are developed than are applied as part of operational disease management systems
Cont
Forty-eight percent of the manuscripts considered for review evaluated the deployment of prediction models as plant disease management tools. We found that the research effort expended on evaluation and application of these models is currently much greater than just a few decades ago.
Cell-wall composition
The materials in a cell wall vary between species, and in plants and fungi also differ between cell types and developmental stages. In plants, the strongest component of the complex cell wall is a carbohydrate called cellulose, which is a polymer of glucose. In bacteria, peptidoglycan forms the cell wall. Archaean cell walls have various compositions, and may be formed of glycoprotein S-layers, pseudopeptidoglycan, or polysaccharides. Fungi possess cell walls made of the glucosamine polymer chitin, and algae typically possess walls made of glycoproteins and polysaccharides. Unusually, diatoms have a cell wall composed of silicic acid. Often, other accessory molecules are found anchored to the cell wall
cell wall-structure
Up to three strata or layers may be found in plant cell walls: The middle lamella, a layer rich in pectins. This outermost layer forming the interface between adjacent plant cells and glues them together. The primary cell wall, generally a thin, flexible and extensible layer formed while the cell is growing. The secondary cell wall, a thick layer formed inside the primary cell wall after the cell is fully grown. It is not found in all cell types. In some cells, such as found xylem, the secondary wall contains lignin, which strengthens and waterproofs the wall.
In the primary (growing)) plant cell wall, the major carbohydrates are cellulose, hemicellulose and pectin. The cellulose microfibrils are linked via hemicellulosic tethers to form the cellulose-hemicellulose network, which is embedded in the pectin matrix. The most common hemicellulose in the primary cell wall is xyloglucan. In grass cell walls, xyloglucan and pectin are reduced in abundance and partially replaced by glucuronarabinoxylan, a hemicellulose. Primary cell walls characteristically extend (grow) by a mechanism called acid growth, which involves turgor-driven movement of the strong cellulose microfibrils within the weaker hemicellulose/pectin matrix, catalyzed by expansin proteins. The outer part of the primary cell wall of the plant epidermis is usually impregnated with cutin and wax, forming a permeability barrier known as the plant cuticle.-----------------------------cont.
.composition
Secondary cell walls contain a wide range of additional compounds that modify their mechanical properties and permeability. The major polymers that make up wood (largely secondary cell walls) include: 1. 2. 3. cellulose, 35-50% xylan, 20-35%, a type of hemicellulose lignin, 10-25%, a complex phenolic polymer that penetrates the spaces in the cell wall between cellulose, hemicellulose and pectin components, driving out water and strengthening the wall. Additionally, structural proteins (1-5%) are found in most plant cell walls; they are classified as hydroxyproline-rich glycoproteins (HRGP), arabinogalactan proteins (AGP), glycine-rich proteins (GRPs), and proline-rich proteins (PRPs).
Cell wall-function
The cell wall is the tough, usually flexible but sometimes fairly rigid layer that surrounds some types of cells. It is located outside the cell membrane and provides these cells with structural support and protection, and also acts as a filtering mechanism. A major function of the cell wall is to act as a pressure vessel, preventing over-expansion when water enters the cell. They are found in plants, bacteria, fungi, algae, and some archaea. Animals and protozoa do not have cell walls.
Enzymes involved
Degradation of polymers of plant cell-walls Approx.20 CWDE are reported so far. Pathogen that cause soft rot diseases in storage organs of carrot,potato tubers produce pectinase, endopolygalacturonase, pectic lyase & pectin methyl esterase There is no strong evidence that any CWDE has any role in disease specificity---they are basic compatibility factor.
Phytoalexins
Antimicrobial low mol. wt.compounds (secondary metabolites)synthesised de novo following pathogenic attack Chemically diverse,but a large number of them are products of shikimic acid (phenylpropanoid) pathway from which many other plant secondary metabolites like lignin & anthocyanins are also derived Elicitors are compounds of microbial origin that induces phytoalexin accumulation.
Formation of papillae,which are knob like structures on the inner surface of the plant CW underneath penetrating fungal spores. Increased lignifications of cell-wall is another response frequently associated with unsuccessful penetration and the hypersensitive response Lignified cells are mechanically stronger and more resistant to digestion by CWDE Lignification could physically restrict the pathogen while biochemical events are still not known Hydrogen peroxide reduction catalysed by cell-wall peroxidases generate free radicles which then spontaneously crosslink to form lignin Plants also contain a number of wall glycoproteins that are rich in proline, glycine & hydroxy prolines also called extensins -strengthen the cell-wall and may there fore resist pathogen invasion
Plant cell remains alive during infection, minimal plant cell damage
Alive only in initial stages, extensive plant cell damage in later stages
Host range
Broad
Narrow; often only a single species of plant is attacked Fungal mildews and rusts, viruses & endoparasitic nematodes, pseudomonas spp. bacteria
intermediate
Examples
Plant pathogens
Fungi
The majority of phytopathogenic fungi belong to the Ascomycetes and the Basidiomycetes. The fungi reproduce both sexually and asexually via the production of spores. These spores may be spread long distances by air or water, or they may be soil borne. Many soil borne spores, normally zoospores, are capable of living saprotrophically, carrying out the first part of their lifecycle in the soil.
Bacteria
Most bacteria that are associated with plants are actually saprotrophic, and do no harm to the plant itself. However, a small number, around 100 species, are able to cause disease. Bacterial diseases are much more prevalent in sub-tropical and tropical regions of the world.
Colonizing the apoplast to cause rots, spots,vascular wilts, cankers and blights Most are gram negative -rod shaped --e.g. pseudomonas, xanthomonas & Ervinia Characteristic features are -1.during their parasitic life, most bacteria reside within the intracellular spaces of various plant organs or in the xylem,2.many cause considerable plant tissue damage by secreting either toxins, extracellular polysaccharides (EPSs) or CWDE at some stages The secreted EPSs surround growing bacterial colony& by saturating intercellular spaces with water or by blocking the xylem to produce wilt symptoms. cont.
Cont..
Erwinia (have a broader host range) deploy pectic enz. Cause extensive cell death &tissue maceration--cleave plant cell-wall ploymers either by hydrolysis (polygalacturonase) or through beta- eliminations (pectate or pectin lyase) The multigenic nature of the gene families (hypersensitive responses & pathogenicity cluster) are required for bacterial pathogenesis.e.g. hrp gene seq. from plant bacteria show same virulence strategy for animals also.
Nematodes
Nematodes are small, multicellular wormlike creatures. Many live freely in the soil, but there are some species which parasitize plant roots. They are a problem in tropical and subtropical regions of the world, where they may infect crops. Potato cyst nematodes (Globodera pallida and G. rostochiensis) are widely distributed in Europe and North and South America and cause $300 million worth of damage in Europe every year. Root knot nematodes have quite a large host range, whereas cyst nematodes tend to only be able to infect a few species. Nematodes are able to cause radical changes in root cells in order to facilitate their lifestyle.
Cont..
Transport of virus particles occur by the way of intracellular (symplastic) movement through the channels between plant cells through plasmodesmata., they never cross PM like animals Plant virus movement proteins (MPs) in association with various components of cytoskeleton of the host cell facilitate transport of virus particles into adjacent cells by way of modified plasmodesmata Many viral MPs have only minimal seq. similarity-suggesting that plant viruses initially may have aquired MP function from plant genome by way of recombination. The process controlling long distance transport with in the phloem distinct from this measophyll one and still not clear challenge ahead is to identify host components in the phloem that interact with either with CP or the virus particle & their subsequent re-entry into the measophyll & root cortical cells
Cont..
Once feeding commences they loose motility & become sedentary and push stylet into cell wall & release glandular secretions & triggers partial dissolution of cell wall leads to Syncytial feeding structure---induce both symplastic connection & finally protoplast fusion mitosis uncoupled from cytokinesis & DNA endo-reduplication ( nuclei with increased DNA content) leads to abnormal cortical cell growth & formation of series of giant cells in close association with phloem Feeding tubes- associated with stylet & unlike stylet located in plant cell cytoplasm & every time when nematode feeds a new FT is formed & by the end of pathogenesis 100 FT are present in giant cells 20-40 kDa protein can pass through FT( micro -injection of fluorescent labeled dextrans of various mol. masses)
Specific/Vertical resistance
Cf genes for Cladosporium fulvum, Pto (Pathovar tomato) for Pseudomonas syringae Mi gene for Meloidogyne incognita (a nematode) Tm2a for TMV
Disease resistance is usually mediated by dominant genes, but some recessive resistance genes also exist
Based on Mendelian hypothesis resistance to plant pathogen is inherited as a single dominant or semidominant trait(1900S) Later seminal genetic studies by Harold H. Flor on flax & the flax rust pathogen proposed that inheritance of not only plant resistance but also pathogen virulence leads to gene-for- gene model (1940s) This model predicts that plant resistance will occur only when a plant has a dominant gene (R) for pathogen detoxification
Flors experiment
Flors Advance in Plant Pathology in 1942 Back in the 1940s, before DNA was shown to be responsible for inheritance,studying flax rust, in 1942, Harold Flor deduced that a single gene of this fungal pathogen was responsible for the ability of the fungus to cause disease on the flax plant. He also determined that this gene in the fungus corresponded to a particular resistance gene in the flax plant. This interaction was called the gene-for-gene theory. It applies primarily to biotrophic plant pathogens such as rust fungi organisms that require a living host to cause disease.
Cont
On the basis of the pattern of compatibility and incompatibility among potato resistance genes and races of Phytophthora infestans ,cause of late blight, one can assign a putative monogenic pathogenicity genotype to each race The validity of this concept is not applicable to classical genetics (based on molecular genetics) Concept also proposed that putative gene-for-gene interaction might have a common biochemical basis& supports possibility of the fact that many plant genes giving resistance to pathogen are allelic.,each different allele controlling resistance to a particular genotype of the pathogen
R-genes
Only few Avr genes known for pathogenic fungi ,colonizing intracellular air spaces in plants & through biochemical approach small secreted peptides can be isolated ,which work as elicitors for R gene dependent defense responses in the nce of the pathogen produces R-protein mediated plant signal perception process Most plant R- proteins have structural similarity & isolation of R-gene utilizes two strategies1.Locating R- genes on the chromosome by using populations that segregate for resistance &susceptible individuals 2. Identifying the correct seq. either by inserting a mobile genetic element ( transposon) to destroy biological activity or by using binary cosmid complementation to confer the resistance phenotype on a susceptible plant
Epigenetic mechanisms
Cont.
The molecular basis of epigenetics is complex. It involves modifications of the activation of certain genes, but not the basic structure of DNA. Additionally, the chromatin proteins associated with DNA may be activated or silenced. This accounts for why the differentiated cells in a multi-cellular organism express only the genes that are necessary for their own activity. Epigenetic changes are preserved when cells divide. Most epigenetic changes only occur within the course of one individual organism's lifetime, but, if a mutation in the DNA has been caused in sperm or egg cell that results in fertilization, then some epigenetic changes are inherited from one generation to the next
Applications of epigenetics
Epigenetic research uses a wide range of molecular biologic techniques to enhance our understanding of epigenetic phenomena, including chromatin immunoprecipitation (together with its large-scale variants ChIP-on-chip and ChIP-seq), fluorescent in situ hybridization, methylationsensitive restriction enzymes, DNA adenine methyltransferase identification (DamID) and bisulfite sequencing. Furthermore, the use of bioinformatic methods is playing an increasing role (computational epigenetics).
- Fungus
The filamentous fungus Neurospora crassa is a prominent model system for understanding the control and function of cytosine methylation. In this organisms, DNA methylation is associated with relics of a genome defense system called RIP (repeatinduced point mutation) and silences gene expression by inhibiting transcription elongation.
- Yeast
The yeast prion PSI is generated by a conformational change of a translation termination factor, which is then inherited by daughter cells. This can provide a survival advantage under adverse conditions. This is an example of epigenetic regulation enabling unicellular organisms to respond rapidly to environmental stress. Prions can be viewed as epigenetic agents capable of inducing a phenotypic change without modification of the genome.
:
2.local response and gene inactivation:
Alterations in secondary metabolites/pathways Cessation of cell cycle Synthesis of pathogen related proteins Accumulation of benzoic and salicylic acid Production of ethylene and jasmonic acid Fortification of cell-walls( lignin,PGIPs,HRGPs)
ROS-defense-mechanism
H2O2 is directly toxic to pathogen & in presence of iron give rise to highly reactive hydroxyl radicle or may contribute to structural reinforcement of plant cell walls either by crosslinking various hydroxy-proline and proline- rich glycoproteins to the polysaccharide matrix or by increasing the rate of lignin polymerization by peroxidase enzyme activity and both would make the plant cell wall more resistant to microbial penetration and enzymatic degradation Signaling role of ROS: H2O2 induces benzoic acid 2hydroxylase(BA 2-H) enzyme activity for SA biosynthesis H2O2 also induces genes for protein involved in certain cell protection mechanism(e.g.glutathione s- reductase. Cont
Cont..
ROS production also alter the redox balance in responding cells (e.g.-specific plant transcription factors are redox regulated
Freezing stress
Some plants can acclimate to subfreezing temperatures A primary function of freeze- tolerance mechanism is membrane stabilization Roles of the osmolytes and antifreeze proteins that accumulate in promoting freezing tolerance remain poorly understood Freezing tolerance involves changes in gene expression
Oxidative stress
Tropospheric ozone is linked to oxidative stress in plants Ozone causes oxidative damage to biomolecules Chloroplasts are susceptible to ozone induced damage Increased synthesis of antioxidants and antioxidant enzymes can improve tolerance to oxidative stress Oxidative stress or ozone can interact with plant hormones such as SA and ethylene to produce plant responses
Heat Stress
Heat stress alters cellular functions Plants can acclimate to heat stress HSPs are conserved among different organisms Five classes of HSPs are defined according to size Expression of many HSPs is controlled by a transcription factor that recognizes a conserved promotor sequences