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PISATIN and LIMITATION of PEA LEAF SPOTS Trevor John Robinson

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PISATIN and LIMITATION of PEA LEAF SPOTS Trevor John Robinson PISATIN AND LIMITATION OF PEA LEAF SPOTS Trevor John Robinson ABSTRACT The two fungi Ascochyta pisi and Nycosphaerella pinodes caused limited, necrotic lesions in detached leaflets of Pisum sativum suspended above water in the light. The lesions caused by either fungus in the dark were not limited; nor were M. pinodes lesions limited in the light if the leaflets were floated on the water surface. There was no evidence from in vivo experiments that the pathogenic behaviour of these fungi was associated with production of phytotoxins. There is strong evidence that lesion limitation is associated with production by the plant of an antifungal compound, pisatin. Many fungal pathogens and a wide range of non-living agents can induce synthesis of this phytoalexin. Pea leaf tissue incubated with drops of many nutrient solutions, including several culture media and some pure sugars, produced pisatin. In the dark, metal salts did not stimulate pisatin production. However metal salts did stimulate pisatin production in the light; and in the dark, when certain sugars were present. This effect can be explained if there are two control points in pisatin biosynthesis. A different control system appears to operate in pod tissue. Solutions that were effective inducers when applied to the leaf surface were not effective when infiltrated into tissue or applied to the surface of watersoaked tissue. This could explain the rapid colonization by M. pinodes of leaflets floating on water, and the repoRrts that both pathogens are more damaging in wet weather. However Al 21.21. lesions did not spread in leaflets floating on water. This illustrates well the complexity of the interactions between peas and leaf spot pathogens. -3- CONTENTS ~ ABSTRACT 2 CONTENTS 3 INTRODUCTION AND LITERATURE REVIEW 5 Ascochyta-leaf-spot of peas 6 Toxins in Ascocbyta-leaf-spot disease 9 The Pbytoalexin Concept 10 Work by Cruickshank and Collaborators 12 "lork by Had\dger and Collaborators 18 Other '-lork on Pisatin 22 Light, Aromatic Metabolism and Disease Resistance 23 MATERIALS .AND METHODS 26 Plant Material 26 Fungi 26 Chemical Materials 27 Cul ture l-iedia 20 Preparation of Culture Fil trates 30 Inoculation of Leaflets 30 Inoculation of Leaf Disks ;1 Freparation for !1icroscopic Examination 31 Preparation of Leaf Disk Diffusates 31 ~action and Assay of Pisatin in Leaf Diffusates 33 Properties of Pisatin from Leaf Diffusates 33 Preparation and Assay of Pod Diffusates 34 Extraction of Pisatin from Leaf Tissue 35 Presentation of Pisatin Measurements 35 Limits of Pisatin Estimates 36 RESULTS 38 STIMULATION OF PISATIN PRODUCTION BY CULTURE FILTRATES 38 STIHULATION OF PISATIN PRODUCTION BY NUTRIENTS 48 EFFECT OF SURFACE STERILIZATION OF LEAF DISKS ON PISATIN 55 nmUCTION COHP011ENTS OF Sll1PLE NEDIA AS POSSIBLE INDUCERS 60 -4- STIMULATION OF PISATIN PRODUCTION BY METAL SALTS 63 CRITICAL STUDIES ON THE EXPERIl1ENTAL SYSTEM 67 Light conditions duxing plant growth 67 Production of pisatin by plants of different ages 69 Role of cut edges of disks in pisatin production 69 Pisatin content· of leaf disk tissue 71 Pisatin production by pods 73 Use of double-distilled water for nutrient solutions 77 Checks on pisatin extraction and assay procedures 78 Light condi tiona in dishes 79 Effeot of disk storage time on pisatin production 80 FURTHER EXPERn1ENTS ON STIHULATION OF PISATIN PRODUCTION 83 :BY CULTURE FILTRATES SUGAR-METAL INTERACTIONS nr PlSATIN INDUCTION 84 PISATIN PRODUCTION BY \mOLE PLANTS 94 FRODTIC TION OF PISATIN BY 'fATER-SOAKED AND INFILTRATED 96 LEAF DISKS EXTRACTIon OF PISATnI FROM INFECTED TISSUE 105 TOXIN PRODUCTION :BY CULT1JRE FILTRATES 108 ANnEX TO RESULTS 115 LESION DEVELOPN~lT 115 DISCUSSION 124 Validi ty of" Results Using the Experimental System 124 NoJtie on Use of th~ 'vord "Inductiontt 125 Induction of Pisatin 125 Role of the Cuticle 129 Water-Soaking and Pisatin Production 130 Pisatin and Ascochyta pisi and Nycospbaerella 131 pinodes Fisatin and the Phytoalexin Concept of Disease 132 Resistanoe . Retrospect 134 REFERENCES 135 ACKNO'\'l1EDGENENTS 144 INTRODUCTION AND LITERATURE REVIEW Localized necrosis of leaf tissue is a common symptom of a variety of diseases caused by fungi, bacteria and viruses. Leaf spots are discrete, small, usually (but not always) regularly shaped, and necrotic lesions; sometimes with a chlorotic halo around the central zone of dead tissue. The lesions are self-limited in the sense that they start to grow in the leaf, reach a typical size and then stop spreading. When it stops growing the lesion is usually surrounded by tissue apparently similar to that in which the pathogen had earlier been able to grow. In some diseases restriction of the parasite's growth can be attributed to preformed mechanical barriers. For example, lesions caused by ItTcosphaerella inusicola in adult banana leaves axe restricted by large vascular bundle sheaths which traverse the leaves (NcGahan and Fulton, 1965), and the symptoms produced in cucumber leaves infected with Pseudomonas lachrymans are confined by the veins (Williams and Keen, 1967). Cunningham (1928), who surveyed the histological changes caused by a number of leaf- spotting pathogens, discovered that in some host-parasite combinations (e.g. Cercosuora beticola on Beta vulgaris and Coccomyces prunophorae on Prunus domestica) meristematic activity in surrounding mesophyll cells caused the formation of a suberized cicatrix which isolated the infected region. In these species mechanical wounding also caused a cicatrix to form but in most species only mechanical damage and not parasitic diseases caused this response. Similarly, cork barriers are said to limit growth of the bacterium Xanthomonas pelargonii in geranium leaves (Bugbee and Anderson, 1963). Pierre and Nillar (1965) observing that the number and size of lesions produced by pjei....2hylium botryosum on alfalfa increased with exposure to high humidity suggested that dehydration of lesions in low humidity conditions reduced the flow of nutrients to the pathogen and so stopped its grw4th. Other physiological mechanisms for development of resistance in leaf spot diseases have included : induced changes in cell walls of adjacent tissues that make them less susceptible to the action of cell-vall degrading enzymes (Bateman, 1963, 1964); absence of appropriate cell-wall degrading enzymes (Williams and Keen, 1967); inactivation of such enzymes by phenolic compounds or products of their oxidation (Deverall and Wood, 1961); and production of fungitoxic or fungistatic compounds during pathogenesis. Accumulation of phenols and other aromatic compounds in invaded host tissues is common (e.g. Rohringer and Samborski, 1967), and the action of phenolic substances and their oxidation products has been demonstrated by many authors (q.v. Cruickshank, Biggs and Perrin, 1971). The role of those compounds which are present in fungistatic quantities only after infection (phytoalexins) is 'considered below. There is reason to believe that different mechanisms will be important in different diseases; and that in any one disease more than one mechanism may well be involved. This thesis is mostly concerned with one •disease : Ascochyta-leaf-spot diseases of peas (Pisum sativum L.) and especially with the production of phytoalexins in pea leaves. Ascochyta-leaf-spot of peas Three species are recognized as the causal agents of Ascochyta blight of peas (Sprague, 1929) : Ascochyta yisi Lib., Ascochyta pinodella L.K. Jones (Jones, 1927b) and Nycosphaerella pinodes (Berk. and Blox.) Stone (imperfect stage Ascochyta pinodes). Until the work of Jones (1927a) and Linford and Sprague (1927), A. pisi had been regarded as the imperfect stage of M. pinodes. The diseases produced by these pathogens can be distinguished in the field since A. pisi causes pod, stem and leaf spots but does not normally affect plant parts below soil level (Sattar, 1934). However, Hare and Walker (1944) state that in Wisconsin at least cases of foot rot have been observed.. The leaf spots, ranging from 2 - 10 mm in diameter, are typically light tan coloured with darker margins. Lesions of - 7 - M. pinodes are less clearly defined and are commonly purple-black and pin- point size, although in wet weather lesions may grow to 8 mm in diameter. N. pinodes unlike A. pisi commonly attacks the base of the stem as well as other parts of the plants and is therefore regarded as the more serious of the two pathogens. (Linford and Sprague, 1927). Penetration and development of M. pinodes in pea leaves were studied with the light microscope by Kerling (1949) and of A. pisi by Ludwig (1928), Brewer and MacNeil (1953) and Brewer (1960). The latter 2 papers reported that.limitation of A. pisi lesions reflected the development rhythm of the fungus, with vegetative growth ceasing at the onset of sporulation. Later, Brewer (1960) suggested that restriction of growth was associated with the dark rim around the lesion but he also suggested that this region forms only because vegetative growth of the fungus is retarded at sporulation. Leach (1962) and Leach and Trione (1965), studied light induced sporulation of A. pisi and found that the sensitive region seemed to be the peripheral zone (1.5 - 2.0 mm wide) of the young mycelium. The action spectrum showed a peak at 290 nm with smaller peaks at 260 and 230 nm. The effect of ultraviolet and visible light on germination, penetration and infection of A. pisi, M. pinodes and 2 other Ascochyta ago, was investigated by Blakeman and Dickinson (1967) (see below for discussion of this and other work). Work on the causes of resistance to the disease has been more limited. Gilchrist (1926) found that pea varieties showing greater resistance to foot rot caused by an unspecified Ascochyta sp. had a thicker cuticle at the base of the epicotyl. Varietal resistance to Ascochyta leaf spot was attributed by Schneider (1952) to the presence of an anthocyanin in 11 the testa of the seed and Sorgel (1956) (both references quoted by Cruickshank and Perrin, 1964) claimed that there was a correlation between an inhibitor of pycnidial formation (possibly an anthocyanin) in cotyledon decoctions and resistance to pod and leaf-spot infection.
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