Novel Biotechnologies for Biocontrol Agent Enhancement and Management
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Springer Springer Springer IOS Press IOS Press
Novel Biotechnologies for Biocontrol Agent Enhancement and Management edited by
Maurizio Vurro Consiglio Nazionale delle Ricerche, Bari, Italy
and
Jonathan Gressel Weizmann Institute of Science, Rehovot, Israel
Published in cooperation with NATO Public Diplomacy Division
Proceedings of the NATO Advanced Study Institute on Novel Biotechnologies for Biocontrol Agent Enhancement and Management held in Gualdo Tadino, Italy 8–19 September 2006 A C.I.P. Catalogue record for this book is available from the Library of Congress.
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CONTENTS
Preface
ix
1. Biotechnology in Crop Protection: Towards Sustainable Insect Control Martin G. Edwards and Angharad M. R. Gatehouse
1
2. Bacteria as Biological Control Agents for Insects: Economics, Engineering, and Environmental Safety Brian A. Federici
25
3. Benefits and Risks of Using Fungal Toxins in Biological Control Maurizio Vurro
53
4. Biocontrol of Weeds with Allelopathy: Conventional and Transgenic Approaches Stephen O. Duke, Scott R. Baerson, Agnes M. Rimando, Zhiqiang Pan, Franck E. Dayan, and Regina G. Belz 5. Selecting, Monitoring, and Enhancing the Performance of Bacterial Biocontrol Agents: Principles, Pitfalls, and Progress Linda S. Thomashow, David M. Weller, Olga V. Mavrodi, and Dmitri V. Mavrodi
75
87
6. Exploiting the Interactions between Fungal Antagonists, Pathogens and the Plant for Biocontrol Sheridan L. Woo and Matteo Lorito
107
7. The Mechanisms and Applications of Symbiotic Opportunistic Plant Symbionts Gary E. Harman and Michal Shoresh
131
8. Using Strains of Fusarium oxysporum to Control Fusarium Wilts: Dream or Reality? Claude Alabouvette, Chantal Olivain, Floriane L’Haridon, S´ebastien Aim´e, and Christian Steinberg v
157
vi
CONTENTS
9. Metarhizium anisopliae as a Model for Studying Bioinsecticidal Host Pathogen Interactions Raymond J. St. Leger 10. Sclerotinia minor—Biocontrol Target or Agent? Alan Watson 11. Fusarium oxysporum f. sp. striga, Athletes Foot or Achilles Heel? Alan Watson, Jonathan Gressel, David Sands, Steven Hallett, Maurizio Vurro, and Fenton Beed 12. Control of Sclerotial Pathogens with the Mycoparasite Coniothyrium minitans John M. Whipps, Amanda Bennett, Mike Challen, John Clarkson, Emma Coventry, S. Muthumeenakshi, Ralph Noble, Chris Rogers, S. Sreenivasaprasad, and E. Eirian Jones 13. Biological Controls and the Potential of Biotechnological Controls for Vertebrate Pest Species Peter Kerr 14. Genetically Enhancing the Efficacy of Plant Pathogens for Control of Weeds Brian M. Thompson, Matthew M. Kirkpatrick, David C. Sands, and Alice L. Pilgeram 15. Interactions of Synthetic Herbicides with Plant Disease and Microbial Herbicides Stephen O. Duke, David E. Wedge, Antonio L. Cerdeira, and Marcus B. Matallo 16. Approaches to and Successes in Developing Transgenically Enhanced Mycoherbicides Jonathan Gressel, Sagit Meir, Yoav Herschkovitz, Hani Al-Ahmad, Inbar Greenspoon, Olubukola Babalola, and Ziva Amsellem 17. Functional Genomics: Functional Reconstitution of Portions of the Proteome in Insect Cell-Lines: Protein Production and Functional Genomics in Cell-lines Thomas A. Grigliatti and Tom A. Pfeifer
179
205
213
223
243
267
277
297
307
CONTENTS
18. TAC–TICS: Transposon-Based Biological Pest Management Systems Thomas A. Grigliatti, Gerald Meister, and Tom A. Pfeifer 19. Failsafe Mechanisms for Preventing Gene Flow and Organism Dispersal of Enhanced Microbial Biocontrol Agents Jonathan Gressel
vii
327
353
Epilogue—Getting from Here to Eternity David Sands
363
Index
365
PREFACE
The intent of the NATO Advanced Study Institute (ASI) entitled “Novel Biotechnologies for Biocontrol Agent Enhancement and Management” was to permit the meeting of the major exponents in the scientific community working with enhancing different biological control agents (fungi, bacteria, virus, nematodes, and insects) on different targets (pathogens, insects, weeds, and rodents). This multidisciplinary group, having backgrounds in the different aspects of biotechnologies (transgenic enhancement, molecular biology, formulation, genetics, risk assessment, new technology, biochemistry, and physiology), presented highly advanced lectures during the 10-day-ASI, in order to allow students to improve their capability to enhance and manage biological control agents. This approach will allow ASI attendees to bring new ideas, new approaches, or new methodologies coming from different fields of application to their own field of expertise. A further aim of the NATO ASI was to create a network of young and experienced scientists, with few geographical barriers among countries, who will develop new opportunities to collaborate in this field of science that requires a “global” collaborative approach. Forty students from twenty countries took part to the NATO ASI. In addition to the 45 lectures from the 15 lecturers, there were 25 short presentations and 8 posters on cogent research from students in this course, held between September 8- 2006 and September 19, 2006. This book represents a partial distillation of all this material together with the daily workshops on various topics, and long discussions over the excellent meals and breaks, in the very conducive environment of the Borgo Hotel Le Terre del Verde at Gualdo Tadino near Perugia, in Italy. The editors especially appreciated the efforts of the anonymous peer reviewers who expeditiously reviewed the chapters of this book. This workshop could not have been possible without the financial assistance of NATO and Valent BioSciences, as well as the lecturers who contributed their time, and in most instances their travel expenses, to assist in allowing the maximum support of students. To these all we have many thanks, along with the knowledge and collaborations engendered by this workshop. Maurizio Vurro and Jonathan Gressel Codirectors October 2006 ix
1. BIOTECHNOLOGY IN CROP PROTECTION: TOWARDS SUSTAINABLE INSECT CONTROL Martin G. Edwards and Angharad M. R. Gatehouse∗ Institute for Research on Environment and Sustainability, Division of Biology, Devonshire Building, Newcastle University, Newcastle upon Tyne, NE1 7RU, UK
Abstract. With a projected increase in world population to 10 billion over the next four decades, an immediate priority for agriculture is to achieve maximum production of food and other products in a manner that is environmentally sustainable and cost effective. Whilst insecticides are very effective in combating the immediate problem of insect attack on crops, nonspecific insecticides are harmful to beneficial organisms including predators and parasitoids of the target pest species. The concept of utilizing a transgenic approach to host plant resistance was realized in the mid 1990s with the commercial introduction of transgenic maize, potato and cotton plants expressing genes encoding the entomocidal δ-endotoxin from Bacillus thuringiensis (Bt). Other strategies based on the use of plant-derived genes (enzyme inhibitors, lectins) and those from animal sources, including insects (biotin-binding proteins, neurohormones, enzyme inhibitors), are currently being developed. The use of fusion proteins to increase the spectrum and durability of resistance is also actively being pursued. Biotechnology in crop protection is not restricted to production of transgenic crops, and has been extended to include the modification of baculoviruses for increased efficacy as biopesticides, and arthropod natural enemies (predators and parasitoids) to enhance their capacity to control insect pests, this chapter will only consider the benefits and risks of its role in the context of insect-resistant transgenic crops. Keywords: sustainability, insect-resistant transgenic crops, insect resistance genes, insecticidal proteins, fusion proteins, pests, natural enemies 1.1. Introduction 1.1.1. NEED FOR SUSTAINABLE AGRICULTURE
The dawn of agriculture occurred some 10,000 years ago with the domestication of cereals, soon to be followed by other crops (Table I). This step ∗
To whom correspondence should be addressed, e-mail:
[email protected] 1 M. Vurro and J. Gressel (eds.), Novel Biotechnologies for Biocontrol Agent Enhancement and Management, 1–23. C 2007 Springer.
2
M. G. EDWARDS AND A. M. R. GATEHOUSE TABLE I. First attested dates of independent transition to agriculture and the main domesticates (after Olsson1 ) Region
Date
Plants
Near East Central Mexico South China North China South Central Andes Eastern United States Sub-Saharan Africa
8500 BC 8000 BC 7500 BC 6800 BC 5800 BC 3200 BC 2500 BC
Wheat barley Maize Rice Soybean Potato, manioc Sunflower Sorghum
was seen as a necessary condition for the development of civilizations. The evolution of agriculture has been divided into four discrete periods, namely the Prehistoric, Roman, Feudal and Scientific Era, with each being associated with specific advancements or developments (Table II). The Prehistoric, or Neolithic Era (10,000 BCE), was thus recognized as the era of crop domestication originating in the regions of low to middle latitude. The Roman Era (1000 BCE–500 CE) saw the introduction of metal tools, the use of animals for farm work and the development of the manipulation of watercourses for irrigation, while the Feudal Era (height, 1100 CE) saw the beginning of international trade based on exportation of crops. Interestingly, the era known as the Scientific Era started as early as the 16th century and although TABLE II. The four major eras of agriculture (www.adbio.com/science/agrihistory) Period
Date
Facts
Prehistoric (Neolithic)
10000 BCE
Domestication of crops 6000 BC: people dependent on domesticated crops
Roman
1000 BCE–500 CE
Metal tools Horses & oxen for farm work Irrigation
Feudal
1100 CE (height)
Export of crops (international trade) 8th Century: rotation
Scientific
1500 CE
15th–19th century: slave labor 16th c: First efforts in plant crop protection 17th–18th Century: Pest control World War II: Mechanization of farming; pesticides 1950s: Mutation breeding, using radioactive isotopes 1970s: Green revolution 1990s: GM crops
SUSTAINABILITY FOR CROP PROTECTION
3
there is documentary evidence for the use of pest control from ancient times, its adoption is primarily attributed to this era. Whilst mineral-based pesticides (arsenates and copper salts) had been used previously major advances in the development of synthetic insecticides did not occur until the end of the Second World War and was accompanied by the intensification of farming. Although in recent years there has been a move towards the development and use of more benign pesticides, the next major breakthrough in this area was seen with the development and commercialization of insect-resistant transgenic crops during the 1990s. In addition to advances made in crop protection, this era has also seen the development and use of mutation breeding, and in the 1970s a major landmark was achieved with the Green Revolution. 1.1.2. THE CHALLENGES AHEAD
The human population is ever increasing, with conservative estimates predicting that the population will rise to approximately 10 billion by 2050. Thus the major challenges facing the world are to feed and provide shelter for a world population that is increasing at an exponential rate (Table III; Figure 1). Furthermore, it is essential to protect human health, and ensure social and economic conditions that are conducive to the fulfillment of the human potential. Agriculture must play a major role in achieving these goals both by providing ever-increasing food yields (Figure 2) and an ever-increasing supply of natural products required by industry. The recent advent of bioethanol further confirms the constraints on agriculture.2 Thus the challenge in the forthcoming decades is to achieve maximum production of food and other products without further irreversible depletion or destruction of the natural TABLE III. World population growth World population in billions
Year
Time needed to reach this level
One Two Three Four Five Six Seven∗ Eight∗ Nine∗
1804 1927 1960 1974 1987 1999 2012 2026 2043
All of human history 123 years 33 years 14 years 13 years 12 years 13 years 14 years 17 years
Source: United Nations Populations Division, World Population Prospects. ∗ Projected population growth; medium variant.
4
M. G. EDWARDS AND A. M. R. GATEHOUSE
Figure 1. Predicted population growth 1950–2050. The data suggests that there has been a fourfold population increase during the last century. While the population is predicted to remain stable in developed regions of the world, based on current trends. Significant increases are predicted to occur in the least developed nations
1600
Million metric tons 432
1400 Feed
1200
Food
1000 235
1040
800 600
493
750
425
400 200 171
182
1997
2020
0
Developed countries
1997
2020
Developing countries
Figure 2. Demand for cereals for human food and animal feed, baseline scenario, 1997–2020 (personal communication A. Cockburn)
SUSTAINABILITY FOR CROP PROTECTION
5
environment, against a backdrop of climate change which not only is predicted to result in the loss of agricultural land as a consequence of rising sea levels, but is also likely to have a major impact on the dynamics of pest populations. Agriculture must become an integral part of a sustainable global society. Agricultural sustainability integrates three major goals, those of environmental health, economic profitability and social and economic equity. It thus rests on the principle that we must meet the needs of the present without compromising the ability of future generations to meet their own needs. Current figures suggest that to feed a world population of 10 billion in 2050 without allowing for additional imports of food, Africa will have to increase its food production by 300%, Latin America by 80% and Asia by 70%. Even North America, which is not usually associated with food shortages, would have to increase its food production by 30% to feed its own projected population of 348 million. Given the current scenario of some 800 million people going hungry on a daily basis and an estimated 30,000 (half of them children) dying every day due to hunger and malnutrition, it is clear that society has many major challenges to address. One step towards achieving sustainability is to identify current major constraints on crop productivity. Simply putting more land into agricultural use, thereby increasing the “agricultural footprint,” is not a viable option in the long term. Currently stress constitutes a major factor in limiting productivity; it can be classified as being either biotic (pests, pathogens and weeds) or abiotic (physical constraints, e.g., temperature, water availability, salinity) where the former can be as can be as high as 40% globally. Insecticides are effective in dealing with the immediate problem of insect attack on crops. They have been responsible for dramatic yield increases in crops that are subject to serious pest problems, but in the longer term severe drawbacks have become apparent. For example, non-specific insecticides are harmful to non-target organisms, many of which play key roles in suppressing the build up of insect populations.3 Other problems associated with high pesticide application include accumulation of toxic residues in food products and subsequent consequences for human health.4 The hypothesis that an over reliance on insecticides is nonsustainable, is further supported by the finding that many insect pests have evolved resistance to such compounds.
1.2. Role of Transgenic Crops in Agriculture Biotechnology offers many opportunities for agriculture and provides the means to address many of the constraints placed to productivity outlined above. It uses the conceptual framework and technical approaches of molecular biology and plant cell culture systems to develop commercial processes
6
M. G. EDWARDS AND A. M. R. GATEHOUSE
Figure 3. Percentage areas of genetically enhanced crops by trait and by crop. (From ISAAA James 2005)4
and products. With the rapid development of biotechnology, agriculture has moved from a resource-based to a science-based industry, with plant breeding being dramatically augmented by the introduction of recombinant DNA technology based on knowledge of gene structure and function. The concept of utilizing a transgenic approach to host plant resistance was realized in the mid 1990s with the commercial introduction of transgenic maize, potato and cotton plants expressing genes encoding the insecticidal δ-endotoxin from Bacillus thuringiensis. Similarly, the role of herbicides in agriculture entered a new era with the introduction of glyphosate-resistant soybeans in 1995. Currently the commercial area planted to transgenic crops is in excess of 90 million hectares (22 million acres) with approximately 77% expressing herbicide tolerance, 15% expressing insect resistance genes and approximately 8% expressing both traits (Figure 3; current production by country is illustrated in Table IV). Despite the increasing disquiet over the growing of such crops in Europe and Africa (at least by the media and certain NGOs) in recent years, the latest figures available demonstrate that the market is increasing, with an 11% increase between 2004 and 2005.5 Applications of transgene technology in agriculture have clearly defined benefits, not least in providing greater sustainability in terms of improved levels of crop protection resulting in higher yields and reduced pesticide application. However, a major challenge facing this new industry is in the identification of suitable genes for transfer that will confer the desired agronomic traits. In terms of insect resistance, several different classes of bacterial-, plant- and animal derived proteins have been shown to be insecticidal towards a range of economically important insect pests from different orders, with the midgut being the prime target.6 Of these the Bt toxins are the most
SUSTAINABILITY FOR CROP PROTECTION
7
TABLE IV. Major countries growing of biotech crops in 2005 Country
Million hectares
Crop
USA Argentina Brazil Canada China Paraguay India South Africa Uruguay Australia Mexico Romania Philippines Spain Colombia Iran Honduras Portugal Germany France Czech Republic
49.8 17.2 9.0 6.1 3.3 1.8 1.3 0.4 0.3 0.2 0.1 0.1 0.1 0.1
