Petria 1(2), 79-83, (1991) Lettera alla Direzione/Letter to the Editors
Plant immunization. A non pesticide control of plant disease
JOSEPH KUC´
Department of Plant Pathology, University of Kentucky, Lexington, Kentucky, 40546 USA
It is increasingly evident that the use of certain pesticides in plant production will
become restricted at the same time that the productivity, profitability and
competitiveness of agriculture must increase. Pesticides contribute to the problem of
environmental deterioration which in turn has a marked influence on the economy,
health, and the quality of life. At the same time, pesticides have been a major factor
contributing to the increased crop yields and quality attained in modern agriculture.
Aside from considerations for the environment, health and quality of life, the generation
of new pesticides is becoming more difficult and expensive. Resistant strains of
pathogens rapidly arise to many new systemic pesticides, and the difficulty of
developing "environmentally-sound" pesticides has increased their cost and reduced the
number that are available. Many recommended pesticides are being removed from the
market and others can only be used on particular crops for restricted periods of time.
Importers of agricultural products are setting strict limits for pesticide residues on food
crops, and for some pesticides there is a zero tolerance. Consumers and consumer groups
are also becoming increasingly concerned about pesticide use and residues on food
products. Advances in biotechnology, including the development and introduction of
transgenic plants, biocontrol, induced systemic resistance (plant immunization) and
enhanced development and increased use of disease resistant plants utilizing new
technologies developed by plant breeders, offer promise of providing alternative means
of disease control that are effective and economical and which would reduce the
dependence on pesticides. The newly developed biotechnology would, however, also
have the effect of increasing the effectiveness of acceptable pesticides and increase their
useful life in agriculture. Plant immunization regulates genes for defense compounds
present even in susceptible plants, is as safe for the environment as disease resistant
plants, since the same mechanisms for resistance are activated in immunized and
resistant plants, and the effect is systemic and often lasts for the life of an annual plant
(Kuc´, 1987, 1990; Dean and Kuc´, 1987a; Kuc´ and Tuzun, 1990; Tuzun and Kuc´,
1989; Madamanchi and Kuc´, 1991). Thus, existing high-yielding, high quality cultivars
can be immunized. Immunization provides protection against a broad spectrum of
pathogens including viruses, bacteria and fungi, and it does not require the introduction
of "foreign" genes. Pesticides are not available for the economic control of plant disease
caused by viruses and many bacteria. A single immunization of cucumber, muskmelon
or watermelon, utilizing Colletotrichum lagenarium (Pass.) Ell. et Halsted, tobacco
necrosis virus (TNV), immunization signal compounds or compounds which release
such signals as the immunizing agents, systemically protects the plants against disease
caused by 13 different pathogens (Kuc´, 1987, 1990; Dean and Kuc´, 1987a;
Madamanchi and Kuc´, 1991). Immunization has been successful in both the laboratory
and field (Kuc´, 1987, 1990; Kuc´ and Tuzun, 1990; Caruso and Kuc´, 1977; Tuzun et
al., 1986). Disease resistance is multicomponent and layered, as would be expected for
plants to survive the selection pressure of evolution. It includes pre-formed barriers and
antimicrobial compounds, often in external tissues, and a response phase. The response
phase includes as defense compounds: phytoalexins, chitinases, β -1,3-glucanases,
proteases, peroxidases, phenoloxidases, hydroxyproline-rich glycoproteins, lignin, and
callose. (Kuc´ 1987, 1990; Kuc´ and Tuzun, 1990; Dean and Kuc´, 1987a; Madamanchi
and Kuc´, 1991; Tuzun and Kuc´, 1989). The expression of genes for the gene products
contributing to disease resistance is regulated by compounds of plant and microbial
origin (West et al., 1985; Sharp et al., 1984a,b,c; Madamanchi and Kuc´, 1991), and
gene regulation — not the presence or absence of genes for resistance mechanisms — is
likely to be the determinant of disease resistance in plants. Thus, all of the defense
compounds noted above can be produced by resistant and susceptible plants, even by
plants reported to lack genes for resistance to a pathogen. The speed, magnitude and
timing of different elements of the response, and the activity of the gene products as
influenced by the environment, determine resistance (Kuc´, 1987, 1990). The
immunization of plants, including plants reported to lack genes for resistance, further
supports the importance of gene expression (Kuc´, 1987, 1990). Immunization is
possible by restricted inoculation with pathogens, attenuated pathogens, selected
nonpathogens, and treatment with chemical substances which are signals produced by
immunized plants or chemicals which release such signals (Kuc´, 1987, 1990).
The signal for immunization in cucurbits and tobacco is graft-transmissibile (Jenns and
Kuc´, 1979; Tuzun and Kuc´, 1985). It is synthesized at the site of infection or treatment
with the inducing agent (Dean and Kuc´, 1986) and is transported in the phloem (Guedes
et al., 1980; Tuzun and Kuc´, 1985). Roots and leaves are immunized by treatment of
leaves with an inducing agent (Gessler and Kuc´, 1982). The immunity signal conditions
resistance even in tissue which has not emerged from the bud (Dalisay and Kuc´, 1989;
Dean and Kuc´, 1986).
Three compounds isolated from immunized tobacco plants, β - ionone, 3-isobutyroyl-bionone and 3-n-butyroyl-b-ionone, protected tobacco in the laboratory and field against
metalaxyl resistant and susceptible strains of the blue mold
pathogen, Peronospora tabacina Adam (Salt et al., 1986, 1988; Kuc´ and Tuzun, 1990).
A group of compounds which release immunity signals may also find application for
disease control. Oxalates and di and trisodium and di and tripotassium phosphates were
recently reported to release immunity signals and systemically protect cucumber against
anthracnose (Doubrava et al., 1988; Gottstein and Kuc´, 1989). The biological spectrum
of effectiveness for oxalates and phosphates in cucumber includes as many diseases as
reported for induction of resistance utilizing microorganisms (Mucharromah and Kuc´,
unpublished).
Though immunization systemically increases the levels of some defense compounds
such as chitinase, β -1,3-glucanases and peroxidases (Metraux and Boller 1986; Tuzun et
al., 1989; Ye et al., 1989), its major effect is to sensitize plants to respond rapidly after
infection (Hammerschmidt and Kuc´, 1982; Dean and Kuc´, 1987b; Kuc´ and Tuzun,
1990; Tuzun et al., 1989; Kuc´, 1984). Thus, energy and precursors are conserved and
utilized when needed. Immunization has been demonstrated in at least 26 diverse plantpathogen interactions including pear, apple, peach, coffee, tomato, potato, cucumber,
barley, cotton, watermelon, muskmelon, wheat, green beans, soybean and sunflower
(Kuc´, 1987; Strobel, unpublished data).To obtain the maximum advantage of each
technology for disease control, it is important to integrate the technologies and to
develop technology that can be integrated. A partially resistant plant would require less
pesticide less frequently applied than would a susceptible plant. Immunization coupled
with plants bred for resistance to some diseases would not only increase the level of
resistance present but also increase the number of diseases to which the plant is resistant.
Tobacco cv Tennessee 86 is resistant to etch and chlorotic vein mottling viruses but is
highly susceptible to blue mold.
Immunization of this cultivar against blue mold rapidly produces plants which are
resistant to the three diseases and resistance is transferred to regenerants via tissue
culture (Nuckles and Kuc´, 1989; Tuzun and Kuc´, 1987, 1989). A transgenic plant with
introduced resistance to a single disease might have a high level of resistance to some
diseases bred into it and be immunized against others. To integrate technologies
successfully requires that the product of integration will "fit" different agronomic
demands. To do this will require not only the development of new technology, but will
also require a new emphasis on and direction for field-oriented research. Hopefully,
government, industry, academia and individual scientists will become increasingly aware
and responsive to the need for funds to support not only the development of new
technology but also funds to integrate and apply the technology for the practical control
of plant disease in an emerging system of sustainable agriculture world-wide.
The objectives of this new integration of technologies would be to increase crop
productivity, quality, and profitability and to reduce environmental and health hazards
associated with pesticide use. It would be as unwise and unrealistic to advocate the
elimination of pesticides from agriculture as it would be to eliminate the use of
antibiotics from medicine. It is realistic, however, to advocate the restricted use of
pesticides and their increased integration into other control practices. To an extent, this
integration is already evident, e.g. the use of disease resistant plants and appropriate
cultural practices with pesticides. At issue is the excessive use of pesticides and
reluctance to explore the possibility of alternative means for disease control and their
integration into practices where pesticides are a minor and not the sole or major control
agent.
Key words: Plant immunization, Integrated control.
Parole chiave: Resistenza indotta, Lotta integrata.
Literature cited
Caruso F. and J. Kuc´, 1977. Field protection of cucumber, watermelon and muskmelon against Colletotrichum
lagenarium by Colletotrichum lagenarium. Phytopathology, 67, 1285-1289.
Dalisay R. and J. Kuc´, 1989. Effect of removing the inducer leaf on the persistence of induced systemic resistance and
enhanced peroxidase levels in cucumber. Phytopathology, 79, 1150.
Dean R. and J. Kuc´, 1986. Induced systemic protection in cucumber: The source of the "signal". Physological and
Molecular Plant Pathology, 28, 227-233.
Dean R. and J. Kuc´, 1987a. Immunization against disease: The plant fights back. In: Fungal Infection of Plants (G.
Pegg and P. Ayres, Eds) Cambridge University Press, Cambridge, 383-410.
Dean R. and J. Kuc´, 1987b. Rapid lignification in response to wounding and infection as a mechanism for induced
systemic resistance in cucumber. Physiological and Molecular Plant Pathology, 31, 69-81.
Doubrava N., R. Dean and J. Kuc´, 1988. Induction of systemic resistance to anthracnose caused by Colletotrichum
lagenarium in cucumber by oxalate and extracts from spinach and rhubarb leaves. Physiological and Molecular Plant
Pathology, 33, 69-80.
Gessler C. and J. Kuc´, 1982. Induction of resistance to Fusarium wilt in cucumber by root and foliar
pathogens. Phytopathology, 72, 1439-1441.
Gottstein H. and J. Kuc´, 1989. Induction of systemic resistance to anthracnose in cucumber by
phosphates. Phytopathology, 79, 176-179.
Guedes M., S. Richmond and S. Kuc´, 1980. Induced systemic resistance in cucumber as influenced by the location of
the inducer inoculation with Colletotrichum lagenarium and onset of flowering and fruiting. Physiological Plant
Pathology, 17, 229-233.
Hammerschmidt R. and J. Kuc´, 1982. Lignification as a mechanism for induced systemic resistance in
cucumber. Physiological Plant Pathology, 20, 61-71.
Jenns A. and J. Kuc´, 1979. Graft transmission of systemic resistance of cucumber to anthracnose induced
by Colletotrichum lagenarium and tobacco necrosis virus. Phytopathology, 69, 753-756.
Kuc´ J., 1984. Phytoalexins and disease resistance mechanisms from a perspective of evolution and adaptation. In:
Origin and Development of Adaptation. Pitman, London, 100-118.
Kuc´ J., 1987. Plant immunization and its applicability for disease control. In: Innovative Approaches to Plant Disease
Control (I. Chet, Ed.) John Wiley, New York, 255-274.
Kuc´ J., 1990. Immunization for the control of plant disease. In: Biological Control of Soil-Borne Pathogens (D.
Hornby, Ed.) CAB International Wallingford, UK, 355-373.
Kuc´ J. and S. Tuzun, 1990. Metabolic regulation of resistance genes in tobacco for the control of blue mold. In: Blue
Mold Disease of Tobacco (C. Main, H. Spurr, Eds), Tobacco Literature Service, North Carolina State University,
Raleigh, 35-46.
Madamanchi N.R. and J. Kuc´, 1991. Induced systemic resistance in plants. In: The Fungal Spore and Disease Initiation
in Plants and Animals (G. Cole and H. Hoch, Eds) Plenum Press, New York, 347-362.
Metraux J. and T. Boller, 1986. Local and systemic induction of chitinase in cucumber plants in response to viral,
bacterial and fungal infections. Physiological and Molecular Plant Pathology, 28, 161-169.
Nuckles E. and J. Kuc´, 1989. Immunization of tobacco cultivar Tn 86 against blue mold. Phytopathology, 79, 1152.
Salt S., S. Tuzun and J. Kuc´, 1986. Effects of b-ionone and abscisic acid on the growth of tobacco and resistance to
blue mold. Physiological and Molecular Plant Pathology, 28, 287-297.
Salt S., M. Reuveni and J. Kuc´, 1988. Inhibition of Peronospora tabacina (blue mold of tobacco) and related plant
pathogens in vitro and in vivo by esters of 3 (R)-hydroxy-β -ionone. 42nd Tobacco Chemists Conference Proceedings,
Tobacco Chemists Soc., Lexington, Ky, USA, 22 pp.
Sharp J., P. Albersheim, P. Ossowski, A. Pilotti, P. Garegg and B. Lindberg, 1984a. Comparison of the structures and
elicitor activities of synthetic and mycelial-wall derived hexa (B-D-glucopyranosyl) D-glucitol. Journal of Biological
Chemistry, 259, 11341-11345.
Sharp J., M. McNeil and P. Albersheim, 1984b. The primary structures of one elicitor-active and seven elicitor-inactive
hexa (B-D- glucopyranosyl) D- glucitols isolated from the mycelial walls of Phytophthora megasperma f.
sp. glycinea. Journal of Biological Chemistry, 259, 11321-11336.
Sharp J., B. Valent and P. Albersheim, 1984c. Purification and partial characterization of a V-glucan fragment that
elicits phytoalexin accumulation in soybean. Journal of Biological Chemistry, 159, 11312-11320.
Tuzun S. and J. Kuc´, 1985. Movement of a factor in tobacco infected with Peronospora tabacina which systemically
protects against blue mold. Physiological Plant Pathology, 26, 321-330.
Tuzun S. and J. Kuc´, 1987. Persistence of induced systemic resistance to blue mold in tobacco plants derived via tissue
culture. Phytopathology, 77, 1032-1035.
Tuzun S. and J. Kuc´, 1989. Induced systemic resistance in tobacco. In: Blue Mold of Tobacco (W. McKeen, Ed.)
American Phytopathological Society Press, St. Paul, Mn, USA, 177-200.
Tuzun S., W. Nesmith, R. Ferriss and J. Kuc´, 1986. Effects of stem injections with Peronospora tabacina on growth of
tobacco and protection against blue mold in the field. Phytopathology, 76, 938-941.
Tuzun S., M. Rao, U. Vogeli, C. Schardl and J. Kuc´, 1989. Induced systemic resistance to blue mold: early induction
and accumulation of β -1,3-glucanases, chitinases and other (b-proteins) in immunized tobacco. Phytopathology, 79,
979-983.
West C., P. Moesta, D. Jin, A. Lois and K. Wickham, 1985. The role of pectic fragments of the plant cell wall in the
responses to biological stress. In: Cellular and Molecular Biology of Plant Stress (J. Key and T. Kosuge, Ed.). A. Liss,
New York, 335-349.
Ye X., S. Pan and J. Kuc´, 1989. Pathogenesis-related proteins and systemic resistance to blue mold and tobacco mosaic
virus induced by tobacco mosaic virus, Peronospora tabacina and aspirin. Physiological and Molecular Plant
Pathology, 35, 161-175.
Petria 1(2), 85-98, (1991) Rassegna/Review
The role of cell injury in induced resistance
ALBERTO MATTA
Dipartimento di Valorizzazione e Protezione delle Risorse agroforestali, Sezione di Patologia vegetale - Università di
Torino, Via P. Giuria 15, I-10126 Torino
For acting in the most discontinuos and direct way the wound stress appears to be
particularly suitable for the study of the role of cell injury per se in the induction of
resistance to microbial pathogens. Such a role is emphasized by the production after
wounding of antimicrobial chemical and physical barriers becoming increasingly
efficient with time. Moreover, besides providing substitutive protective structures
against wound parasites, the wound response might confer also to intact tissues higher
resistance levels. It has been reported that wounding induces local resistance
to Cladosporium cucumerinum in cucumber, Peronospora parasitica in
radish, Fusarium lycopersici in tomato. Also cell injury caused by immersion in hot
water or exposure to chloroform vapors of the roots of tomato plants is followed by an
increased, transitory state of resistance to Fusarium and Verticillium wilt. The protection
induced by abiotic stresses against Fusarium wilt of tomato is similar to the protection
induced by treatments with avirulent fungi or fungal elicitors that injure part of the root
tissues. Moreover both types of treatment are followed by similar changes with time of
different enzyme activities and total phenols content. Cell injury is apparently involved
in the resistance induced by microrganisms also against nonvascular infections. In some
plants (cucumber, tobacco) microbial infections, but not wounds or other abiotic
stresses, induce resistance systemically. The insucces to induce resistance to fungi
systemically by wounding even in cucumber and tobacco is conflicting with the
evidence that the plants react systemically to cell injury as such in many other ways
(accumulation of phenols, increase of enzyme activities, accumulation of mRNA coded
for PRs, synthesis of proteinase inhibitors that are probably factors of resistance against
insects). The transmission of signals is clearly involved in the activation of systemic
responses to cell injury. Further research aimed to the recognition of such signals
appears to be necessary for a better understanding of the mechanisms of induction of
active resistance in plants.
Key words: Induced resistance, Cell injury.
Il ruolo del danno cellulare nella resistenza indotta
Per le sue caratteristiche di immediatezza e discontinuità il trauma meccanico appare
particolarmente adatto allo studio del ruolo del danno cellulare in sé nel fenomeno della
resistenza indotta ai parassiti. Alcune delle principali risposte alla ferita o ad analoghe
forme di danno cellulare consistono in:
1) precoce depolarizzazione di membrana seguita da rilascio di elettroliti, digestione
enzimatica di lipidi con produzione di composti volatili fungitossici;
2) aumento di attività del ciclo respiratorio dei pentoso-fosfati e partecipazione nella
respirazione di vie alternative cianuro-resistenti di trasporto elettronico sfocianti nella
formazione di superossido;
3) ossidazione di preesistenti fenoli in chinoni fungitossici; sintesi de novo di fenoli e
loro ossidazione o polimerizzazione a suberina e lignina;
4) sintesi di fitoalessine.
Gran parte delle risposte di cui sopra concorrono nei meccanismi di resistenza ai
parassiti. Oltre a provvedere strutture protettive contro i parassiti da ferita, le risposte
alle ferite possono determinare aumento di resistenza nei tessuti intatti circostanti. Le
ferite inducono resistenza locale a Cladosporium cucumerinum in cetriolo,
a Peronosporaparasitica in ravanello, a Fusarium lycopersici in pomodoro. Non solo,
ma anche il danno cellulare prodotto da trattamenti con calore o composti tossici è
seguito da un transitorio aumento di resistenza a F. lycopersici e Verticillium dahliae in
pomodoro. La resistenza a F. lycopersici è similmente indotta dagli stress abiotici e
dall’inoculazione con funghi avirulenti o con elicitori fungini capaci di lesionare i
tessuti. Sia i trattamenti abiotici sia quelli biotici sono inoltre seguiti in pomodoro da
cambiamenti, analoghi nel tempo, delle attività perossidasica, polifenolossidasica,
chitinasica e β -1,3-glucanasica e della concentrazione fenolica. Danno cellulare è
apparentemente richiesto anche nella induzione micro-biologica di resistenza locale
verso infezione di patogeni non vascolari. Solo i microrganismi che causano necrosi
ipersensibile o necrosi estese sono efficaci nell’indurre resistenza. In alcune piante
(cetriolo, tabacco) le infezioni microbiche, ma non le ferite o altri stress abiotici,
inducono resistenza sistemicamente. L’incapacità delle ferite di indurre resistenza
sistemica in tabacco e cetriolo contrasta con l’evidenza che le piante reagiscono
sistemicamente al danno cellulare come tale in molti altri modi: con aumento di svariate
attività enzimatiche, accumulo di fenoli, accumulo di mRNA codificato per proteine di
patogenesi, sintesi di inibitori proteici di proteineasi possibilmente funzionanti da fattori
di resistenza agli insetti. La trasmissione di segnali ovviamente interviene nella
trasmissione di risposte sistemiche al danno cellulare. È stato ipotizzato che i segnali
possono essere di natura chimica (oligosaccaridi staccati dalle pareti cellulari,
traumatina, etilene, ABA, ecc.) o elettrici ed elettrochimici. Lo sviluppo di ricerche
rivolte all’individuazione del segnale trasmettitore di informazioni a tessuti distanti dalle
cellule danneggiate sarà necessario per la comprensione dei meccanismi di induzione
della resistenza attiva nelle piante.
Parole chiave: Resistenza indotta, Danno cellulare.
Petria 1(2), 99-110, (1991) Articolo Scientifico/Scientific paper
Variazioni delle popolazioni fungine associate al "mal del piede" del frumento duro
nell'Italia meridionale
SALVATORE FRISULLO e VITTORIO ROSSI
Dipartimento di Biologia, Difesa e Biotecnologie Agro-Forestali, Università degli Studi della Basilicata, Via Nazario
Sauro, 85, I-85100 Potenza
Con una indagine, svolta nel biennio 1988/89 in 5 località dell’Italia meridionale, è stata
studiata la popolazione fungina associata alle radici ed ai culmi di frumento colpito dal
"mal del piede", come pure le variazioni delle diverse specie fungine durante il periodo
compreso fra l’accestimento e la maturazione delle spighe. Le specie isolate con
maggiore frequenza sono state: Microdochium nivale, Fusarium
culmorum, Drechslerasorokiniana, Fusarium avenaceum,Fusarium
crookwellense, Fusarium graminearum e Rhizoctonia cerealis. La frequenza di
isolamento di queste specie è variata in rapporto all’annata, alla località, allo stadio di
sviluppo delle piante ed all’organo vegetale colpito.
Parole chiave: Frumento duro, Mal del piede, Marciume radicale, Fusarium
culmorum, Microdochium nivale.
Changes of the fungal population associated with foot and root rot of durum wheat
in southern Italy
A study on the fungal population associated with foot and root rot of durum wheat, from
tillering to ripening, was carried out in five wheat-growing areas of southern Italy in
1988 and 1989. The species more frequently isolated from haulms and roots of the
diseased plants were: Microdochium nivale, Fusarium culmorum, Drechslera
sorokiniana, Fusarium avenaceum, Fusarium crookwellense, Fusarium
graminearum and Rhizoctonia cerealis. Frequency of these species changed with regard
to the year, the place, the developmental stage of plants and the part of plant from which
they were isolated.
Key words: Durum wheat, Foot-rot, Root-rot, Fusariumculmorum, Microdochium
nivale.
Petria 1(2), 111-115, (1991) Articolo Scientifico/Scientific paper
Papaver rhoeas L. ospite di due virus patogeni per le piante coltivate
IPPOLITO CAMELE1, MARIA NUZZACI1, GIAN LUIGI RANA1 e PANAIOTA E.
KYRIAKOPOULOU2
1
Dipartimento di Biologia, Difesa e Biotecnologie Agro-Forestali, Università degli Studi della Basilicata, Via Nazario
Sauro, 85, I-85100 Potenza
2
Istituto Fitopatologico Benaki, Kiphissia, Atene, Grecia
Da piante di papavero, mostranti clorosi generalizzata o nanismo e maculatura giallastra
ed infestanti rispettivamente alcuni carciofeti greci e pugliesi, sono stati isolati due virus
fitopatogeni: il virus latente italiano del carciofo (AILV) e quello del mosaico della rapa
(TuMV). I due virus sono stati identificati mediante immunomicroscopia elettronica
seguita da decorazione.
Parole chiave: Carciofo, Papavero, Virus latente italiano del carciofo, Virus del mosaico
della rapa.
Papaver rhoeas as a host for two phytopathogenic viruses
Two plant viruses, artichoke Italian latent virus (AILV) and turnip mosaic virus (TuMV)
were isolated from wild plants of Papaver rhoeas L. growing in artichoke fields in
Greece and southern Italy and showing chlorosis or yellow mottle accompained by leaf
and stem deformation, respectively. The viruses were identified by immuno-sorbent
electron microscopy followed by decoration.
Key words: Artichoke, Poppy, Artichoke Italian latent virus, Turnip mosaic virus.
Petria 1(2), 117-156, (1991)
IVth
INTERNATIONAL TRICHODERMA AND GLIOCLADIUM
WORKSHOP
Belgirate, Italy, July 17-20, 1991
Organizzato da/Organized by:
DI.VA.P.R.A. - Patologia Vegetale, Università Di Torino, Italy
Foreword
The International Trichoderma and Gliocladium Workshops are organized to promote
exchange of ideas and information among researchers working on various aspects of
these two important fungi. The first Workshop was held in 1984 at the Beltsville
Agricultural Research Center (MD, USA). At that time the group decided to meet
alternatively in the eastern and western hemispheres. The pace of research in many
areas, such as taxonomy, genetic manipulation, enzyme production, ability to act as
biological control agents, has quickened so much that the group, after the second
meeting at Salford (UK), decided to meet every other year, in order to promote better
discussion and exchange of information among researchers. During the last few years
both plant pathologists and commercial companies have shown great interest in the
potential of Trichoderma and Gliocladium as biocontrol agents and molecular biology
approaches have been successfully used to improve strains and to better understand their
mode of action. The University of Torino is very pleased to host this fourth meeting at
Belgirate (Northern Italy). The large participation of young researchers, coming from all
over the world, is particularly welcome. We hope that the discussion at this meeting will
be fruitful and helpful to further progress of research in the different areas.We gratefully
acknowledge all financial contributions received from the Ministry of Agriculture and
Forestry, the University of Torino and the Piedmont Region Government.
M. Lodovica Gullino
Presentazione
Gli incontri internazionali su "Trichoderma e Gliocladium" sono organizzati per
favorire lo scambio di informazioni e la discussione tra quanti conducono ricerche su
questi due importanti funghi. Il primo Workshop venne organizzato presso l’Agricultural
Research Center di Beltsville (Maryland, USA) nel 1984: in tale occasione, i ricercatori
operanti in questo settore decisero di incontrarsi, a intervalli regolari, alternativamente
in America e in Europa. Il ritmo assunto dalla ricerca su questi due funghi in diversi
settori (ad esempio tassonomia, manipolazione genetica, produzione di enzimi, lotta
biologica) ha spinto il gruppo, durante il secondo Workshop svoltosi a Salford (Gran
Bretagna), a incontrarsi ogni due anni, per facilitare al massimo la discussione e lo
scambio di idee. In questi ultimi anni si è osservato un crescente interesse, da parte dei
ricercatori e dell’ industria agrochimica, verso il potenziale impiego di Trichoderma e
Gliocladium come mezzi biologici di lotta: il ricorso a tecniche di biologia molecolare
sta consentendo da un lato di migliorare le prestazioni dei ceppi antagonisti e dall’altro
di chiarire i loro meccanismi di azione.
L’Università di Torino è onorata di ospitare il quarto incontro a Belgirate (Novara). E’
particolarmente apprezzata la partecipazione di giovani ricercatori, provenienti da aree
geografiche diverse. Ci auguriamo che la discussione e lo scambio di idee siano
particolarmente fruttuosi e che possano favorire l’avanzamento della ricerca nei diversi
settori. Si ringrazia vivamente il Ministero dell’Agricoltura e delle Foreste, l’Università
degli studi di Torino e la Regione Piemonte per avere generosamente contribuito alla
organizzazione di questo convegno.
M. Lodovica Gullino
Presented papers /Lavori
presentati
page/pagina
WILLIAMS M. A.J. Problems and perspectives in Trichoderma taxonomy
120
GAMS W. The stability of morphological characters in Trichoderma
depending on cultivation conditions
121
SAMUELS G., S. MANGUIN-GAGARINE., R.MEYER,
O.PETRIN. Morphological and macromolecular characterization
of Hypocrea schweinitzii and its Trichodermaanamorph
121
THRANE U. Use of HPLC-DAD for chemotaxonomic characterization
of Trichoderma and Gliocladium species
122
SAMUELS G.J, K.A SEIFERT . Reassessment of species attributed
to Gliocladium
123
PE’ER S., Z.BARAK, I. CHET. Genetic manipulation of the antagonistic
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124
TÖRRÖNEN A., T.A MYOHANEN, F. HOFER, D. BLAAS, A. HARKKI,
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JACOBS D., R.A. GEREMIA, G.H. GOLDMAN, O.KAMOEN, M. VAN
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125
HEIDENREICH E., C.P. KUBICEK. Towards cloning of biocontrol genes
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125
HAYES C.K., G.E. HARMAN, T.E.STASZ, S.L.WOO. Rapid
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126
HERRERA-ESTRELLA A., R. GEREMIA , G. GOLDMAN , M. VAN
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127
HAYES C.K., G.E. HARMAN, S.L. WOO, M.L.
GULLINO. Electrophoretic karyotyping of Trichoderma harzianum
127
GOLDMAN G.H., J. DEMOLDER, R. VILLARROEL, S. DEWAELE, M.
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128
MIGHELI Q., C. FIORETTA, L. CAVALLARIN , M.L.
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128
HOWELL C. R., R.D. STIPANOVIC. Antibiotic production by Gliocladium
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129
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130
GEREMIA R.A., G.H. GOLDMAN, D. JACOBS, M. VAN MONTAGU, A.
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131
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132
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root rot of cucumber in peat following substrate amendment with oatmeal
132
MCKENZIE L. I., D. BENZI, D. DELLAVALLE , M.L. GULLINO Survival
on the phylloplane of strains of Trichoderma spp. antagonistic to Botrytis
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133
SURICO G. Observation by SEM of the attachment of bacterial plant
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134
JIN X., G.E. HARMAN, G.PERUZZOTTI. Production of quality biomass
of Trichoderma harzianum for biocontrol using liquid fermentation
135
ARTECONI M., P. BERGONZONI, L. PERRONE, C.
136
MALLEGNI. Trichoderma and Gliocladium production in submerged
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enhance biocontrol efficacy of Trichoderma harzianum
137
JEYARAJAN R., G. RAMAKRISHNAN. Efficacy
of Trichoderma formulation against root rot disease of grain legumes
137
LUMSDEN R.D., J.C.LOCKE, J.F. WALTER. Approval of Gliocladium
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of Pythium and Rhizoctonia damping-off
138
WILCOX W.F., G.E. HARMAN. Control of Phytophthora root rots of
soybeans and raspberries with Trichoderma and Gliocladium spp.
139
LI HANGING. A study of application of Trichoderma against Sclerotinia
sclerotiorium in soy-beans
140
VANNACCI G., S. PECCHIA, C. MALLEGNI, W. CORTELLINI, F.
FACCINI. Biocontrol of Sclerotinia lettuce drop
140
MOHAN L., N. SHUNMUGAM. Biological control of bulb-rot of garlic
142
LODHA B.C., K. MATHUR , J. WEBSTER. Management of rhizome rot
of ginger using Gliocladium and Trichoderma species
142
JEYARAJAN R., G. RAMAKRISHNAN, B. RAJAMANICKAM,
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biocontrol agent for root rot disease of grain legumes and oilseeds
143
RATTINK H. Possibilities of biocontrol of soilborne fungi by Trichoderma
harzianum in dutch glasshouse crops
144
SREENIVASAPRASAD S., K. MANIBHUSHANRAO. Potential
of Gliocladium virens and Trichoderma longibrachiatum as biocontrol
agents of fungal pathogens
145
MUKHOPADHYAY A.N., P.K.MUKHERJEE. Innovative approaches in
biological control of soilborne diseases in chickpea
146
XU T., J.PZHONG , D.B. LI. Antagonism of Trichoderma harzianum T82
and Trichoderma sp. NF9 against soil-borne fungous pathogens
146
KÖHL J., W.M.L MOLHOEK, N.J. FOKKEMA. Biological control
of Botrytis aclada and B. squamosa in onions
147
ELAD Y., A. COHEN. Biological control and combination of the
biocontrol agent Trichoderma with fungicides for the control of gray mold
148
GULLINO M.L., C. ALOI, D. BENZI, A. GARIBALDI. Biological and
integrated control of grey mould of vegetable crops
149
ANSELMI N., M. NEGRI, G. NICOLOTTI, G. SANGUINETI. Biological
control with Trichoderma spp. against basidiomycetes agents of wood
decay and root rots in forest trees
150
NICOLOTTI G., N. ANSELMI, M.L. GULLINO. Selection of strains
of Trichoderma spp. active against Heterobasidion annosum
151
DE MELO I.S., A.C. DA SILVA. Resistance of UV induced mutants
of Trichoderma harzianum to benzimidazole and dicarboximide fungicides
151
BABY U.I., K. MANIBHUSHANRAO. Biological control of rice sheath
blight with Gliocladium virens and Trichoderma longibrachiatum
152
PROKKOLA S. Biological control of liquorice rot (Mycocentrospora
acerina) with Trichoderma spp.
153
GERMEIER CH., H. FEHRMANN. Sclerotium rolfsii on cereals: prospects
to biological control
154
PRATELLA G.C., M- MARI. Trichoderma and Gliocladium in biological
control of postharvest diseases
155
TAMIETTI G., D. BENZI, L. FERRARIS. Studies on the antagonistic
activity of Gliocladium virens against Sclerotium cepivorum
155
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