Text 3 Metabolites of Pseudomonas

Involved in the biocontrol

Of plant disease

___________________________________ David N. Dowling and Fergal O’Gara

There is increasing commercial and environmental interest in the use of microbebased agents as alternatives to, or in combination with, chemicals for controlling the spread and severity of a range of crop diseases. The identification of specific microbial metabolites that are able to control certain plant diseases has led to the development of strategies for improving the performance and predictability of the microbial strains that produce these metabolites for application in the agricultural industry. This article focuses on antimicrobial metabolites produced by fluorescent pseudomonads, discusses their role in suppressing fungal diseases of important crops and reviews the prospects of genetically manipulating the producer organisms to improve the efficacy of these biocontrol agents.

The ability of some soils to suppress plant disease has been well documented. An important example is the development of soils suppressive to Gaeumannomyces graminis var. tritici (Ggt), the causative agent of ‘take-all’ of wheat. Such natural examples of biological control have been attributed to the indigeneous beneficial rhizosphere microflora (see Glossary), especially fluorescent pseudomonads. Numerous mechanisms may account for these biocontrol properties (see Glossary), including the production of inhibitory compounds or metabolites. Microbial metabolites such as siderophores (see Glossary) and secondary metabolites with antimicrobial properties are considered to play a major role in disease suppression (see Table 1).

Metabolites with biocontrol properties have been reported from diverse members of the beneficial rhizosphere flora: however, those produced by the fluorescent pseudomonads have received the most attention. This is probably due to the abundance of this diverse group of bacteria and their obvious importance in the rhizosphere, coupled with the relative ease with which they can be genetically manipulated.

Fluorescent pseudomonads produce a variety of metabolites, many of which are inhibitory to other microorganisms and some of which are implicated in the biological control of plant pathogens.The identification of a link between a metabolite(s) and the suppression of a particular disease is an on-going goal of many research groups, which the use of recombinant DNA (rDNA) techniques has facilitated. Some of the criteria used to indicate that a particular metabolite has a primary role in biological control are shown below.

• Mutants defective in metabolite(s) are unable to show inhibition of the pathogen in the laboratory.

• The biocontrol ability of the mutants is reduced in the field.

• Complementation of the mutant with wild-type DNA sequences restores biocontrol ability.

• The purified metabolite shows fungicidal or antimicrobial properties.

• The metabolite may be detected in situ (i.e. in the rhizosphere) when producing strains are present.

The ability to fulfil some or all of these points provides good evidence for the involvement of a particular metabolite in biocontrol. A selection of plant diseases controlled by specific metabolites are summarized in Table 1.

Role of metabolites in biological control

Some of the metabolites implicated in biocontrol appear to be broad-ranging in their inhibitory action. For example, phloroglucinols and phenazines have been shown to inhibit a wide range of fungal pathogens in the laboratory. Siderophores exhibit both fungistatic and bacteriostatic

Table 1. Examples of specific microbial metabolites implicated in the control of crop diseases
Disease	Pathogen	Effective metabolite	Refs
Take-all of wheat	Gaeumannomyces graminis var. tritici (Ggt)	Phenazines c-Acetyl phloroglucinols	9 10, 11
Tan spot of wheat	Pyrenophora triticirepentis	Pyrrolnitrin	12
Pre-emergent damping-off of: cotton; sugarbeet	Pythium spp. Pythium ultimum	Oomycin A Pyoluteorin 2,4-Diacetylphloroglucinol	13 14 15
Black root-rot of tobacco	Thielabiopsis basicola	Hydrogen cyanide 2,4-Diacetylphloroglucinol	16 10
Crown gall of fruit trees	Agrobacterium tumefaciens	Agrocin 84	17-19
Flax wilt	Fusarium oxysporum	Pseudobactin B10	20
Damping-off	Pythium spp.	Ammonia	21

effects in the laboratory under conditions of low iron. In the field, these iron-chelating compounds are thought to deprive the pathogen of iron, a limiting essential nutrient.

Other metabolites are known to have very specific effects and to target particular pathogens; for example, agrocin 84, produced by Agrobacterium radiobacter, is specific for virulent strains of Agrobacterium tumefaciens. At the molecular level, agrocin 84 (a di-substituted nucleotide) is thought to act by chain termination of DNA synthesis. However, the precise mode of actions of many other metabolites is poorly understood.

Effects such as direct killing or pathogen inhibition exhibited by a particular metabolite in the laboratory cannot always be translated directly into biological control in the field, as the conditions experienced in the field are much more complex. However, there are a number of studies demonstrating a direct effect of particular antimicrobial metabolites on pathogen numbers in soil. One such example is the suppression of root pathogens by plant-growth-promoting rhizobacteria (PGPR) (see Glossary). In another case, the population size of antibiotic-producing Pseudomonas fluorescens 2-79 was shown to correlate positively with a reduction in the number of lesions on wheat roots caused by Ggt.

Evidence is accumulating to support the theory that metabolite production has beneficial effects on the ecological competence of the producer strain. Production of these compounds is thought to provide the producing strain with a selective advantage in the highly competitive environment of the plant rhizosphere. This idea has been substantiated by a recent report, which demonstrated that phenazine (Phz) antibiotics contributed to the persistence of the producer strains (P. fluorescens 2-79 and P. aureofaciens 30-84) compared with Phz-deficient mutants in a simulated wheat-rotation microcosm.

Glossary

Biological control – The use of living organisms to modify the agricultural ecosystem to control a crop disease or prevent the establishment of a pest. Rhizosphere – The root surface and that part of the soil directly influenced by the root system. Plant-growth-promoting rhizobacteria (PGPR) – Bacteria isolated from the rhizosphere, which have been shown to improve plant health or increase yield. They may exert their effects directly through the production of metabolites that can act as plant hormones and/or indirectly by the suppression of plant pathogens (biological control). Induced systemic acquired resistance – The ability of an agent, for example, a bacterium, chemical, virus, nematode, etc., to induce plant-defence mechanisms that lead to systemic resistance to a number of pathogens. Siderophore – Low-molecular-weight compound that is produced as a specific iron chelator by a microorganism.

Persistence may lead to improved competition and colonization of the producer strain on the plant surface, leading to niche-exclusion of the pathogen. The role of siderophores and other metabolite(s) in biocontrol may not be simply a direct antagonism of the pathogenic fungus, but may also involve more subtle ecological effects.

Alternative, more indirect, modes of action could involve stimulation of the plant’s own defence mechanisms by induced systemic acquired resistance or, possibly, by direct uptake and translocation of the metabolite within the plant.

D. N. Dowling and F. O’Gara are at the Department of

Microbiology, University College Cork, Ireland.

TIBTECH APRIL 1994 (VOL 12)

© 1994, Elsevier Science Ltd 0167 – 7799/94/S7.00

 ASSIGNMENTS

1. Read the scientific text above and present it taking into account the branch of knowledge it belongs to.

2. Analyze the text in terms of the category of informativity. Comment on how paralinguistic means of conveying information, used in the text alongside the verbal ones, add on to the factual information of the scientific text in question.

3. Comment on the lexical peculiarities of the present scientific text distinguishing the three lexical layers in its vocabulary.

4. Analyze the text in terms of the category of cohesion focusing on the lexical means conducive to the logico-semantic integrity of the scientific writing under consideration.

5. Compose and write a coherent essay summing up your observations on the linguostylistic and textlinguistic peculiarities of the scientific text “Metabolites of Pseudomonas involved in the biocontrol of plant disease”.

LIST OF LITERATURE

1. Арнольд И.В. Стилистика современного английского языка: (Стилистика декодирования). – М.: Просвещение, 1990. – 300 с.

2. Бессонова О.Л., Федорова Л.А. Методические указания по чтению газеты. – М. –1992. – 41 с.

3. Богатырева С.Т. Выделение синтагматических единиц в стиле научного изложения: Автореф... канд. филол. наук. – М.,1983 – 22 с.

4. Богатырева С.Т., Бунтина Т.А. Методические указания к аналитическому чтению «Текст как отражение особенностей различных функциональных стилей». – Донецк, 1990. – 41c.

5. Бунтина Т.А., Криводубская А.Н. Задания для самостоятельной работы по аналитическому чтению. – Донецк, 1989. – 27с.

6. Гальперин И.Р. Текст как объект лингвистического исследования. – М.: КомКнига, 2007. – 148 с.

7. Гулак А.Т. Приемы лингвистического анализа художественного текста – К., 1984. – 145 с.

8. Задорнова В.Я. Стилистика английского языка. Методические указания для студентов факультета романо-германской филологии. – М., 1986. – 32с.

9. Зильберман Л.И. Структурно-семантический анализ научного текста. М.: Наука, 1982. – 133 с.

10. Кожина М.Н. О речевой системе научного стиля сравнительно с некоторыми другими. – Пермь: Изд-во Перм. ун–та, 1972. – 395 с.

11. Кочан І. М. Лінгвістичний аналіз тексту: Навчальний посібник. – Київ: «Знання», 2008. – 423 с.

12. Кузнецова Л.А. Методическая разработка по курсу «Лингвистика текста». – Львов, 1981. – 28 с.

13. Кухаренко В.А. Практикум з стилiстики англiйської мови: (англ. мовою): Пiдруч. для студ. ф-тiв iнозем. мов вузiв освiти. – Вид 3-тє. – Вiнниця: Нова книга, 2003. – 160 с.

14. Лебедева М.С., Фролова Г.М. Язык средств массовой информации Великобритании и США. – Москва: МГЛУ, 2002.

15. Литературный энциклопедический словарь / Под ред. В.М.Кожевникова, П.А. Николаева. – М., 1987.

16. Мараховский А.Н. Некоторые основные понятия стилистики и лингвистики текста. Лингвистика текста и методика преподавания языков. К., 1981. – c. 6-13.

17. Мараховский А.Н., Воробьева О.П. и др. Стилистика английского языка. – К.: Вища школа, 1984. – 247 с.

18. Минделли Е.И. Словарный состав научного текста как предмет тематического, морфологического, и морфосинтаксического изучения: Автореф... канд. филол. наук. – М., 1982. – 22 с.

19. Мосткова С.Я., Смыкалова Л.А. English Literary Terms. – Leningrad, 1967. – 109с.

20. Николаева Т.М. Лингвистика текста / Современное состояние и перспективы. – Новое в зарубежной лингвистике. – М.: Прогресс, 1978. – Вып. 8. – С. 5-43.

21. Новиков Л.А. Лингвистическое толкование художественного текста. - М.: Русский язык, 1979. - 251 с .

22. Разинкина Н.М. Развитие языка английской научной литературы. Лингвостилистические исследования. М.: Наука, 1978. – 211 с.

23. Сосновская В.Б. Аналитическое чтение: Учеб. пособие для ин-тов и фак. иностр. яз. – М. : Высш. шк., 1974. – 180 с.

24. Crystal D. The Cambridge Encyclopedia of the English Language. – CUP, 1995. – 327p.

25. Galperin I.R. Stylistics. M. Higher School, 1997. – 395p.

26. Gilyanova A.G., Ossovskaya M.I., Turaeva Z. Ya. Analytical Reading: Учебник английского языка для 3 курса институтов и факультетов иностранных языков. – Москва: Высшая школа, 1971. – 224 с.

27. Grundy P. Newspapers. – Oxford: Oxford University Press, 1999. – 134 p.

28. Hamilton J. Straight to the source. Newspapers. – Minnesota: ABDO Publishing Company, 2005. – 32 p.

29. Johansson S. Some aspects of the Vocabulary of Learned and Scientific English. – Goteborg, 1978. – 57 p.

30. John E. Richardson. Analysing Newspapers: An Approach from Critical Discourse Analysis. – New York: Palgrave Macmillan, 2007. – 260 p.

31. Kukharenko V.A. Seminars in Style. M., 1991. – 184p.

32. Martinet A. A functional View of Language. – London-Oxford, 1968. – 163 p.

33. Savory Th. H. The language of Science. Its Growth, Character and Usage. – London, 1953.- 184 p.

34. Siegal Allan M. and Connolly William G. The New York Times Manual of Style and Usage.– New York: Times Books/Random House, 1999. – 365 pp.

35. Starikova E.N. Modern English. – Kiev, 1987. – 270p.

36. Wernep A. A short guide to English style. – London: Oxf. Univ. Press. – 1961. – 198 p.

1⁾ Mind that the Russian word ‘науки’ may refer both to science subjects such as chemistry, biology, physics, etc. and to humanities subjects such as history, literature, philosophy, linguistics, etc., while in English there are two different words: sciences vs humanities.