Introduction Definitions: Fungi As Plant Pathogens Anubhav Vinayak Lisa Tofil Heather Rice • Parasite – organism that gains all or part of its nutritional requirements from the living tissues of another organism – the host. • Pathogen – organism that causes disease. A pathogen almost always a parasite but not all parasites cause diseases. Pathogens Pathogens • Pathogens – Necrotrophic – (kill the host tissues usually by directly invading them or by producing toxins or degradative enzymes and then feed on the tissues that they kill) – Biotrophic – (feed on living host tissues by producing special nutrient-absorbing structures that tap into host’s tissues) Seed rots and seedling pathogens • Athelia rolfsii – One of the most devastating seedling pathogens in the warmer parts of the world where rainfall is seasonal Pathogen Necrotroph Host-specialized Non-specialized Host-specialized Seedling pathogens (cont) • Rhizoctonia solani – Common seedling pathogen that attacks the basal stem tissues of plants – Examples Source: http://www.turffiles.ncsu.edu/diseaseID/diseasedetail.aspx?disease=RHISL Source: http://www.agroatlas.ru/content/diseases/Olee/Olee_Athelia_rolfsii/Olee_Athelia_rolfsii.jpg Biotroph Source:http://www.ctahr.hawaii.edu/nelsons/Misc/ 1_potato_sore_shin_rhizoctonia_1.jpg 1 Host-Specialized Necrotrophic Pathogens Pectic Enzymes • Degrade the middle lamella (the cementing layer) • Three major types – Pectin methyl esterase (PME) – Pectic lyase (PL) – Polygalacturonase (PG) • Necrotrophic pathogens: – Invade normal healthy tissues of plants – Overcome the defense mechanisms of their hosts • Physical barriers • Fungitoxic chemicals Botrytis cinerea Source: http://www.ccrc.uga.edu Tissue Degrading Enzymes • Necrotrophic fungi and nongrass plants: F. oxysporum and V. albo-atrum – Pectic enzymes • Gramineae (grass family): R. cerealis, F. culmorum, and T. yallundae – Arabinase, xylanase, and glucanase enzymes – Small amounts of pectic enzymes • Able to degrade typical wall components of their hosts Plant Defenses Against Necrotrophic Pathogens • Physical barriers: – Oxidative burst • Produces hydrogen peroxide at or near plant cell surface • Hydrogen peroxide strengthens cell wall against attack – Cross-links plant cell wall proteins Source: www.php.wur.nI/UK/Research/ Plant Defenses Against Necrotrophic Pathogens • Hypersensitive response: – Upon attack the infected cells die and accumulate fungitoxic chemicals • Physical barriers: – Leaf cuticle • Cutinase enzyme allows the fungus to penetrate through the cuticle – Papilla Source: staff.jccc.net/PDECELL/plants/transpiration.html • Thickening of the cell wall at the point of invasion Plant Defenses Against Necrotrophic Pathogens • Chemical defenses: – Phytoanticipins • Converted to an active compound in response to infection • S. lycopersici: – Can overcome phytoanticipins – Produce enzymes: Avenacinase and α-tomatinase 2 Plant Defenses Against Necrotrophic Pathogens • Chemical Defenses: Systemic Acquired and Induced Resistance • Systemic acquired resistance (SAR): – Phytoalexins – Increased resistance following infection • Low molecular weight • Protect plants against general attack by fungi • Produced in response to infection – Activated by an oxidative burst Hydrogen peroxide is initial signal • Mediated by salicylic acid • Induced systemic resistance (ISR): – Increased resistance following colonization of plants roots by rhizosphere bacteria • P. fluorescens • Mediated by jasmonic acid and ethylene Vascular Wilt Pathogens • Three main fungi: • Mode of infection: 1. V. albo-atrum 2. V. dahliae 3. F. oxysporum • • – Spores enter water-conducting xylem vessels More than 80 strains (“special forms”) Strains survive by producing chlamydospores • Symptoms: – – Yellowing and death of leaves Collapse of leaf stalks • Vascular Wilt Pathogens • Vectors • Wounds • Young root tips – Carried upwards in the water flow – Become trapped on perforated end walls • Produce additional spores – Remain restricted to the xylem vessels until the plant dies Epinasty Source: www.anbg.gov.au/cpbr/program/sc/path_vari.htm Plant Response to Vascular Wilt Fungi • Tyloses- (balloon-like swelling) • Pectic gels • Release of phenolic compounds • Accumulation of phytoalexins Smut Fungi • Members of the Basidiomycota • Over 100 species: – Loose smut of wheat: U. nuda – Maize smut: U. maydis – Stinking smut of wheat: T. caries Source: Figure 14.14, Deacon (2006) Fungal Biology, 4th ed. Source: www.uky.edu/.../wheat/disease/Ismut/Ismut1.htm 3 Smut Fungi Fungal Endophytes and Their Toxins • Mode of infection: – Germinating spores penetrate developing ovary – Resulting seeds germinate, colonize, and enter flowering spike – At maturity flowering spike produces a mass of black spores Source: www.uky.edu/.../wheat/disease/Ismut/ Ismut1.htm • Significance – produce secondary metabolites useful for commercial use – induce abiotic tolerance – insect antifeedant Source: http://cc.oulu.fi/~ampir/kotisivu-3.html Fungal Endophytes and Their Toxins Cont. Fungal Endophytes and Their Toxins Cont. • “Clavicipitaceous endophytes” • Dilemmas of endophytes – Common species is called Claviceps purpurea – These type of endophytes are found in several pasteur grasses in US, Europe and New Zealand • Lolium (ryegrass) • Festuca (fescue) • Dactylis glomerata (cocksfoot) Fungal Endophytes and Their Toxins Cont. – Produce mycotoxins – Mycotoxins harm grazing animals – endophytes increase stress tolerance – endophytes prevent insect damage over other grasses Fungal Endophytes and Their Toxins Cont. • Types of mycotoxins and effects – loliterm B- saturated aminopyrrolizidine alkaloids-“ryegrass staggers” disease of grazing sheep and cattle. Staggers are loss of coordination and/or tremors occur at times – Egrovaline- causes “fesuce foot” and “fescue toxicosis” where the animal can experience hyperthermia, weight loss, reduced pregnancy rates, decreased milk production, and for horses birth defects or abortions – Peramine- tripeptide that is repellent and toxic to insects, but not mammals Sources: http:// www.biology.ed.ac.uk/ research/groups/jdeacon/ FungalBiology/ chap14_3.htm#Fig14.18 4 Phytophthora Diseases Phytophthora Diseases Cont. • They are not fungi but are part of Oomycota • Phytophthora contains more than 50 species • Common known Phytophthora disease was the potatoe blight • Causes – Root rot – Cankers-develop near base of trees – Collar rots and crown rots- develop near base of steams of woody herbaceous plants – Aerial blights – Fruit rots Phytophthora Diseases Cont. • Phytophthora infestans – Heterothallic fungi – Originated in Central Mexico, both mating types present there (A1 and A2) – Moved over to Europe by 1840’s causing Potatoe Blight – Only A1 was dominantly present outside Central Mexico, but in 1980 began to spread Sources: http://www.ent.iastate.edu/images/ plantpath/soybean/rootrot/root_rot.jpg http://hyg.ipm.illinois.edu/photos/ canker.jpg • Phytophthora ramorum: sudden oak death – First found in Germany and Netherlands in 1993 – Described as P. ramorum in 2001 – Now found in coastal fog belt of northern California and southern Oregon – European mating type is A1 – North American mating type is A2 – Separate origin of sources http://aggie-horticulture.tamu.edu/ publications/tomatoproblemsolver/ ripe/images/photos/sour_rot.jpg Phytophthora Diseases Cont. • Phytophthora infestans Cont. – Symptoms • Black spreading lesions on foliage • Cool humid conditions allow black lesions to produce sporangiophores that emerge from leaf stomata • Potato tubers become infected later (spread of motile zoospores) • Tubers rot then destroyed by secondary bacteria invaders • May also be spread by wind born sporangia or splashed sporangia onto leaves • Phytophthora Diseases Cont. Sources Cont. http://plant-disease.ippc.orst.edu/ disease.cfm?RecordID=1434 Sources: – http://www.biology.ed.ac.uk/research/groups/jdeacon/microbes/blight.htm ( fig 14.19a ) Phytophthora Diseases Cont. • Phytophthora ramorum: sudden oak death cont. – Plants affected and symptoms • Rhododendron branches, Vibrurnum bushes, ornamental Rhododendron and Vibrurnum (Europe), and oaks-- “live and tan oaks” (US) • Produces lesions on leaves and terminal shoots of shrubs (shoot dieback) • Cankers at base of trees (destroys cambium) • Cracked bark • Seepage of dark, viscous sap from bark near base of tree (gummosis) • Sudden death • Slow progressive building of infection under bark 5 Phytophthora Diseases Cont. • Phytophthora ramorum: sudden oak death cont. Sources: http://www.biology.ed.ac.uk/research/groups/jdeacon/ FungalBiology/pramorum.htm (Fig 14.20 a-e) Biotrophic Plant Pathogens Cont. Biotrophic Plant Pathogens • Characterized by an extended nutritional relationship with living host cells as opposed to necrotrophic pathogens which kills plant tissues • Success depends upon – ability to avoid eliciting host cell death – securing a continuous nutrient supply from living host tissues Biotrophic Plant Pathogens Cont. • Example Fulvia (Cladosporium) fulvum – One strain only feeds from the host if it avoids inducing hypersensitive response – Governed by avirulence genes of the pathogen and corresponding resistance genes in host • gene for gene relationship is common for biotrophic plant pathogens – Specific type of biotroph covered is Haustorial biotrophs Source: http://www.biology.ed.ac.uk/research/groups/ jdeacon/FungalBiology/chap14_3.htm#Fig14.21 Biotrophic Plant Pathogens Cont. • Rust Fungi – Black stem rust of wheat (Puccinia graminis) • Life cycle fig 14.22 Sources: http://www.biology.ed.ac.uk/research/groups/ jdeacon/FungalBiology/chap14_3.htm#Fig14.22 Biotrophic Plant Pathogens Cont. • Powdery Mildew Fungi – Part of Ascomycota cause problems throughout all seasons, especially when dry and hot and if not treated with fungicides – Common types • powdery mildew of roses (Sphaerotheca pannosa) • powdery mildew of gooseberries (S. mors-uvae) • powdery mildew of hawthorn bushes (Podosphaera clandestina) – This species undergoes multiple cycles of infection in one season like rust fungi 6 Question 1 Biotrophic Plant Pathogens Cont. • Downy mildews – Part of Oomycota – haustorial biotroph act like rust fungi and powdery mildew fungi, but need humid conditions for infection – Examples • Blemia lactucae of lettuce • Plasmopara viticola on grape vines • Pseudoperonospora humuli on hops • What structure do pectic enzymes degrade in plant cells? A. B. C. D. Sources: http://www.hort.cornell.edu/ department/facilities/lihrec/vegpath/ photos/downymildew_lettuce.htm Question 2 • What is the name of the signal molecule that mediates systemic acquired resistance? A. B. C. D. Jasmonic acid Xylanase Ethylene Salicylic acid Question 4 • What type of fungi has multiple life cycles including uredospore, teliospores, basidiospores, and aeciospores? A. Powdery Mildew Fungi B. Rust Fungi C. Blemia lactucae D. Phytophthora ramorum Primary wall Middle lamella Secondary wall Plasmodesmata Question 3 • Which of the following is not a defense mechanism used by plants to prevent invasion by necrotrophic pathogens? A. B. C. D. Phytoanticipins Cutinase enzyme Oxidative burst Phytoalexins Question 5 • Which type of fungi has two types found in the US (A1 and A2) A. Phytophthora infestans B. Phytophthora ramorum C. Blemia lactucae D. Pseudoperonospora humuli 7