Nutritional and Product Quality Innovations

Graff, Zilberman, & Bennett

Nutritional and Product Quality Innovations

Nutritional and Product Quality Innovations in Agricultural Biotechnology: A new generation of crop traits and their potential economic impacts

Gregory D. Graff 1,5 David Zilberman 2,3 Alan B. Bennett 4,5

1. Colorado State University, Agricultural and Resource Economics 2. University of California-Berkeley, Agricultural and Resource Economics 3. Giannini Foundation of Agricultural Economics 4. University of California-Davis, Plant Sciences 5. Public Intellectual Property Resource for Agriculture (PIPRA)

April 2008

Public Intellectual Property Resource for Agriculture

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Abstract

This study develops a broad and detailed picture of the range of nutritional and product quality innovations that have become technically feasible within agricultural biotechnology, focusing on those likely to be commercialized over the next five to ten years. It does this by two methods. The first is a broad survey published primary and secondary data sources, including scientific articles, public data on field trials, and public regulatory filings. This creates a dataset characterizing the broader R&D pipeline, of which we give a summary overview in Chapter 3 and a more detailed cataloguing in Chapter 4. The second method is a compendium of primary sources on plans for product commercialization, including interviews with company officials, and research administrators at university and public institutes, as well as official primary publications released by the innovating organizations. This creates a schedule of anticipated product releases for the next five to ten years, detailed in Chapter 5. What emerges is an intriguingly detailed snapshot taken at one point in time of an ever evolving and highly dynamic flow of new ideas and product development projects within a global research community and industry.

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Table of Contents List of Tables ............................................................................................................................. v List of Figures..........................................................................................................................vii 1 Introduction: Biotechnology and quality innovation in agricultural products ............... 1 1.1 Historical trends of innovation in agriculture ................................................................ 1 1.2 Qualitative improvements and biotechnology ............................................................... 2 1.3 Assessing the R&D pipeline ......................................................................................... 3 1.4 Objectives .................................................................................................................... 5 2 Agricultural markets and factors governing demand for nutritional and product quality innovations .................................................................................................................... 6 2.1 Markets in the industrialized world .............................................................................. 7 2.1.1 Food retail ............................................................................................................ 7 2.1.2 Food commodities and food processing .............................................................. 10 2.1.3 Animal feed and forage ...................................................................................... 12 2.1.4 Floriculture and nursery products ....................................................................... 13 2.1.5 Pulp and paper ................................................................................................... 14 2.2 Formal markets in developing economies ................................................................... 15 2.3 Informal and missing markets in developing countries: innovation needs for subsistence agriculture and food security ................................................................................................. 17 2.4 Regulatory approvals, perceived risks, and market demand for the products of biotechnology ....................................................................................................................... 19 3 Surveying the supply of nutritional and product quality innovations ........................... 22 3.1 The dynamics of the R&D pipeline ............................................................................ 24 3.2 Categories of nutritional and product quality traits observed in the R&D pipeline ....... 26 3.3 Crops observed in the R&D pipeline .......................................................................... 26 3.4 The sources of innovation: the organizations and countries doing research and development ......................................................................................................................... 30 4 The nutritional and product quality traits found in the survey .................................... 38 4.1 Multiple quality traits - Seed composition and feed quality ......................................... 39 4.2 Proteins and amino acids ............................................................................................ 39 4.2.1 Protein quality and level..................................................................................... 40 4.2.2 Lysine, methionine, and tryptophan – essential but deficient amino acids............ 42 4.2.3 Nutrient-enhancing enzymes ............................................................................... 44 4.2.4 Other nutritional proteins................................................................................... 45 4.2.5 Protein functional quality ................................................................................... 45 4.3 Oils and fatty acids .................................................................................................... 46 4.4 Carbohydrates ............................................................................................................ 51 4.4.1 Starches ............................................................................................................. 51 4.4.2 Fructans............................................................................................................. 57 4.4.3 Sugars ................................................................................................................ 58 4.5 Micronutrients and functional metabolites .................................................................. 59 4.5.1 Vitamins ............................................................................................................. 60 4.5.2 Minerals............................................................................................................. 61 4.5.3 Functional secondary metabolites ...................................................................... 62 4.6 Reduction or removal of non-nutrients, allergens, and toxins ...................................... 64 4.6.1 Non-nutritional, anti-nutritional, and toxic plant metabolites.............................. 64 4.6.2 Allergens ............................................................................................................ 65 4.6.3 Mycotoxins ......................................................................................................... 66 4.7 Extended shelf life ..................................................................................................... 67 4.7.1 Control of fruit ripening ..................................................................................... 67

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4.7.2 Control of leaf and flower wilting ....................................................................... 72 4.7.3 Bruising or browning ......................................................................................... 73 4.8 Esthetics and convenience .......................................................................................... 73 4.8.1 Flavor and Scent ................................................................................................ 74 4.8.2 Fruit, seed, or fiber color ................................................................................... 75 4.8.3 Flower color ...................................................................................................... 75 4.8.4 Size and morphology .......................................................................................... 77 4.8.5 Seedlessness ....................................................................................................... 77 4.8.6 Low maintenance landscaping............................................................................ 78 4.9 Fiber and wood quality............................................................................................... 78 4.9.1 Fiber quality for textiles ..................................................................................... 79 4.9.2 Fiber quality for digestibility of animal feed and forage...................................... 79 4.9.3 Wood quality for pulp......................................................................................... 81 4.10 Environmental quality traits ....................................................................................... 82 5 Commercialization of nutritional and product quality traits ........................................ 84 5.1 Crop quality innovations already commercialized....................................................... 84 5.1.1 Conventional quality innovations already commercialized .................................. 84 5.1.2 Transgenic quality innovations already commercialized...................................... 88 5.2 Transgenic nutritional and product quality innovations likely to be commercialized in the next five years ................................................................................................................. 91 5.2.1 Soy consumer traits and oil quality ..................................................................... 91 5.2.2 Feed and processing traits in corn/maize ............................................................. 92 5.2.3 Horticultural specialty products .......................................................................... 92 5.2.4 Enhanced micronutrients .................................................................................... 96 5.3 Innovations likely to be commercialized in five to ten years ....................................... 96 6 Understanding the economic impact of nutritional and product quality innovations 101 6.1 The social and economic welfare impacts of product quality innovation ................... 101 6.2 Heterogeneity and the dynamics of technology adoption .......................................... 102 6.3 The impacts of product quality innovation on risk .................................................... 103 6.4 The role of information, regulation, and marketing ................................................... 105 6.5 Price differentiation and supply channel separation .................................................. 105 7 The four mechansims of economic impact by product quality innovations ................ 107 7.1 Innovations that affect final demand: consumer traits ............................................... 107 7.2 Innovations that increase the input-use efficiency of intermediate goods: processing and feed quality traits ................................................................................................................ 109 7.3 Innovations that extend shelf life: controlled ripening traits ...................................... 110 7.4 Innovations that reduce negative externalities: environmental traits .......................... 113 7.5 Hedonic price analysis of new quality characteristics ............................................... 113 8 The distribution of the economic impacts from product quality innovation............... 116 8.1 Impacts on consumers .............................................................................................. 116 8.2 Impacts on retailers .................................................................................................. 119 8.3 Impacts on processors and manufacturers ................................................................. 119 8.4 Impacts on livestock, dairy, poultry, and aquaculture operations ............................... 119 8.5 Impacts on growers .................................................................................................. 120 8.6 Impacts on the environment ..................................................................................... 120 9 Implications and conclusions ........................................................................................ 121 9.1 The ten greatest impacts of this of technology .......................................................... 121 9.2 Implications for related areas of technical innovation and the industrial organization of agriculture........................................................................................................................... 123 9.3 Implications for consumer perceptions of biotechnology .......................................... 124 References ............................................................................................................................. 125

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List of Tables Table 2.1 Recent top selling new product innovations in U.S. food retail .................................... 9 Table 2.2 Major product categories of food manufacturing ingredients ..................................... 11 Table 2.3 U.S. harvests and uses of major feed crops in 2002 ................................................... 13 Table 3.1 The range of nutritional and product quality traits in the R&D pipeline ..................... 25 Table 3.2 Cross matrix of transgenic crops and trait categories in the R&D pipeline ................. 28 Table 3.3 Top 20 innovating organizations in quality traits, by size of project portfolio............. 31 Table 3.4 Innovations, by nationality of lead and partner R&D organizations ........................... 33 Table 4.1 Transgenic plants with multiple quality traits: improving ‘seed composition’ or ‘feed quality’ ..................................................................................................................................... 39 Table 4.2 Transgenic plants with increased protein quality and level......................................... 40 Table 4.3. Transgenic plants with increased levels of essential amino acids ............................... 42 Table 4.4. Transgenic plants with nutrient-enhancing enzymes .................................................. 44 Table 4.5 Transgenic plants with other nutritional proteins ....................................................... 45 Table 4.6. Transgenic plants with protein functional qualities .................................................... 46 Table 4.7 Transgenic plants with modified oils and fatty acids .................................................. 47 Table 4.8 Transgenic plants with general carbohydrate modifications ....................................... 51 Table 4.9 Transgenic plants with modified starch ..................................................................... 52 Table 4.10 Transgenic plants with modified fructans ................................................................ 57 Table 4.11 Transgenic plants with modified simple sugars........................................................ 58 Table 4.12 Transgenic plants with increased vitamin content .................................................... 60 Table 4.13 Transgenic plants with modified content or bioavailability of minerals .................... 61 Table 4.14 Transgenic plants with modified functional secondary metabolites, such as flavonoids, sterols, capsaicin, etc. .............................................................................................. 62 Table 4.15 Transgenic plants with removed non-nutritional, anti-nutritional, and toxic plant metabolites ................................................................................................................................ 64 Table 4.16 Transgenic plants with allergens removed, reduced, or mitigated ............................. 65 Table 4.17 Transgenic plants with mycotoxins removed, reduced, or mitigated ......................... 66 Table 4.18. Transgenic plants with controlled fruit ripening....................................................... 67 Table 4.19 Transgenic plants with control of leaf and flower wilting ........................................ 72 Table 4.20 Transgenic plants with controlled bruising or browning........................................... 73 Table 4.21 Transgenic plants with modified flavor or scent ....................................................... 74 Table 4.22 Transgenic plants with modified fruit, seed, or fiber color ....................................... 75 Table 4.23 Transgenic flowers with modified color .................................................................. 76 Table 4.24 Transgenic plants with modified size or morphology ............................................... 77 Table 4.25 Transgenic plants with seedless fruit ....................................................................... 78 Table 4.26 Transgenic landscaping plants with lower maintenance requirements ...................... 78 Table 4.27 Transgenic plants with improved fiber quality for use in texitles.............................. 79 Table 4.28 Transgenic plants with improvied fiber quality for digestability of animal feed and forage........................................................................................................................................ 79 Table 4.29 Transgenic trees with improved pulping characteristics ........................................... 81 Table 4.30 Transgenic plants for bioremediation of soil toxins .................................................. 82 Table 5.1 Recent examples of nutritional and quality-enhanced crop varieties using conventional breeding and genetic mutation, by release date .......................................................................... 86 Table 5.2 The five transgenic nutritional and quality enhanced products already commercialized, by release date........................................................................................................................... 90 Table 5.3 Nutritional and quality enhanced products likely to be commercialized in the next five years ......................................................................................................................................... 93

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Table 5.4 Nutritional and quality enhanced products likely to be commercialized in five to ten years ......................................................................................................................................... 98 Table 8.1 Mechanisms of economic impact for each category of trait ...................................... 117 Table 8.2. Expected welfare impacts on economic groups in the agricultural value chain ......... 118

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List of Figures Figure 2.1 Value added from food products and beverages as percent of total GNP for OECD countries ..................................................................................................................................... 8 Figure 2.2 Purchases by food manufacturers of food ingredients from U.S. crop agriculture...... 10 Figure 2.3 U.S. animal populations and output in 2002 .............................................................. 12 Figure 2.4 Growth in demand for meat and cereals in developing countries is expected to well exceed that in developed countries ............................................................................................ 16 Figure 3.1 Advance in the R&D pipeline of the 560 individual nutritional and product quality innovations observed by the current survey ............................................................................... 23 Figure 3.2 The number of innovations observed to have entered, exited, and remained in the R&D pipeline annually (with truncation of the dataset in recent years) ...................................... 24 Figure 3.3 Crops with active projects in nutritional and product quality traits, by year .............. 27 Figure 3.4 Share of organizations involvement in the nutritional and product quality R&D pipeline ..................................................................................................................................... 32 Figure 3.5 Global distribution of nutritional and product quality innovations and the network of cross-national collaborations, by nationality of lead and partner R&D organizations for each innovation ................................................................................................................................. 34 Figure 3.6 Share of innovating organization types at different points in the R&D pipeline ........ 35 Figure 3.7 Innovations in the R&D pipeline by type of R&D sector across four country groupings: US, Europe, other OECD countries, and developing countries .................................. 36 Figure 6.1 The market penetration of a new technology over time follows an S-curve ............ 102 Figure 7.1 The contribution of greater size to the market value of peaches (the hedonic price of ‘size’) changes with time of season. Source: (Parker and Zilberman, 1993)............................. 114

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Introduction: Biotechnology and quality innovation in agricultural products

To date, the tools of agricultural biotechnology have been commercially to confer several major crops with traits to control insect and weed pests. These powerful technologies have given growers a new kind of assurance against pest damage, improved efficiency of production under a range of conditions, thereby reducing costs and increasing the economic well being of growers. The value of these pest controlling genetic traits to growers is evidenced by their very rapid and widespread adoption where they are available, primarily in North and South America. Studies in China and India indicate that the value of pest control genetic traits may in some instances be magnified many-fold for lower income growers within the agricultural systems of less-developed economies (Pray, et al., 2001, Qaim and Zilberman, 2003). Within the ongoing debate about positive environmental impacts of these pest control biotechnologies, the most important benefits that have been pointed out include a relative shift away from pest control chemicals and an associated shift toward no-tillage soil management which reduces erosion. While these agronomic traits can improve the efficiency of agricultural production, they do not per se improve the quality of the resulting agricultural product. Indeed, the product from such genetically modified varieties is not intended to be differentiated from the product of corresponding conventional varieties. As a result any value arising from pest control biotechnologies for consumers or other industries within the agricultural value chain is indirect, potentially in the form of lower prices for the basic agricultural commodity, but is otherwise difficult to describe or ascertain in direct tangible terms. The tools of biotechnology can be used in agriculture in ways that offer rather more tangible benefits to the users and consumers of agricultural products. It has become commonplace to outline the forthcoming technological developments in agricultural biotechnology as belonging to several overlapping generations: (1) first generation ‘input’ technologies, such as pest control and other agronomic traits, (2) second generation ‘output’ technologies, such as nutritional and quality enhancements for deployment in food, feed, fiber, and ornamental value chains, and (3) third generation technologies for manufacturing novel proteins and other bioorganic materials in plants. The introduction of the second generation, quality enhancing traits is anticipated to mark a significant turning point in the economic evolution of agricultural biotechnology: a shift from being just a source of innovation for agricultural producers to being a source of innovation directly benefiting a much broader constituency including consumers and the other industries served by agriculture. This coming transition will necessarily introduce new terms into the economic relationships between innovators the users.

1.1

Historical trends of innovation in agriculture

Agriculture has long been characterized by high rates of innovation. Change in the technological base of agriculture has enabled the feeding and clothing of growing populations from a limited resource base. It has also resulted in a steady reduction in the costs of agricultural commodities. Decreased cost of food is of great benefit to consumers—especially the poorest—but it puts great pressure on agricultural production systems and farmers to compete in lowering costs and increasing efficiencies. Additionally, consumers worldwide are looking for more than just lower prices for basic food commodities. Consumers are seeking greater quality and convenience in the

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foods and consumer products they purchase. They are also increasingly aware of and responsive to the health and environmental implications of their choices. In addition, governments are increasingly active in regulating agricultural production to address public health, food safety, and the environment. Thus, the reality faced in today’s markets for food, feed, fiber, and ornamentals is one of declining costs of raw commodities, changing consumer tastes and needs, and increasingly stringent government regulation. For agricultural producers, the two primary responses to this situation have been either to increase efficiency—and thus decrease costs—or to differentiate products along particular quality characteristics. This latter approach has the potential to escape competition in undifferentiated commodities and earn a premium derived from consumers’ willingness to pay for particular quality characteristics. However, these two responses are inherently at odds with one another. Greater efficiency and lower costs are best achieved by exploiting the economies of scale of bulk commodities on international markets. Quality, however, comes at greater cost, and product differentiation involves producing on multiple smaller scales and requires more complex logistics to preserve the identity of the differentiated product as it travels from the grower to the consumer. One of the main focuses of innovation in agricultural systems today seeks to resolve this tension by achieving both greater efficiency and greater quality differentiation. Technological and organizational innovations are being pursued and implemented throughout the vertical supply chains of all agriculturally-based industries in an effort to increase efficiency while also delivering greater value to the consumer. Markets and infrastructure are adapting, with multiple sub-markets or niche categories emerging—for organic foods, ethnic foods, specialty diets, etc— taking over where single undifferentiated commodities once reigned almost exclusively. These trends are not limited to industrially developed countries but are being felt worldwide. An entire range of technologies, including farm management and agronomic practices, fertilizers and pesticides, irrigation, field machinery, computing and information technologies, inventory management control, post harvest packing and processing, and plant breeding and biotechnology are being improved and adopted. These multiple technologies work together within increasingly diverse agricultural and industrial systems to achieve both increased efficiency and improved quality. Biotechnology—given that it involves the entire scope of the inner workings of living organisms at the molecular level—is very much a ‘general purpose’ technology. It is by no means limited to solving just issues of efficiency, such as pest control, and from its very inception biotechnology has been envisioned as a way to improve the quality and nutritional value of crops.

1.2

Qualitative improvements and biotechnology

Improvement in the quality of crops for human use has been going on since the dawn of agriculture and the domestication of crop species. One simply has to consider the difference in size and palatability between wild apples and commercially grown apples, or between undomesticated teosinte and the elite hybrid corn varieties grown today. Uncounted generations of selection, replanting, and breeding have selected for size, color, texture, sugar, convenience, efficiency in processing, and the removal of unpalatable or toxic components. Indeed, nutritional content may be the newest criterion, as nutrition historically has either been poorly understood or has been merely a secondary result of selection for characteristics of color and flavor. Such changes are most often subtle and incremental. For example, over the last 20 years, levels of betacarotene in commercial lines of carrots have slowly doubled as a result of selection and breeding for a darker orange color. All of these improvements are distinct from agronomic characteristics such as yield, standability, or disease resistance which also have been notably improved through 2



the same processes of selection and breeding. Quality innovations emerging from biotechnology can thus, in an economic sense, be considered largely as an extension or continuation of the ongoing process of improving the characteristics of agricultural commodities through new means. The possibilities of improvements in nutrition and product quality using the tools of molecular biology have long been promised—and sometimes over promised. Ten years ago a quality enhanced tomato with extended shelf life, trade named as the ‘FlavrSavr®’ tomato, was in fact the first agricultural product of biotechnology. It was introduced in 1994 by Calgene of Davis, California, only to fail in the face of disappointing financial performance and stiff competition in the tomato market. In the last five years, much attention has been paid to the introduction of betacarotene—the dietary precursor of vitamin A—into staple foods such as rice, for consumption by those for whom vitamin-A deficiency is a problem. This technology, branded as ‘Golden Rice®’, was widely publicized at a very early stage, when the initial results of a team at ETH Zurich in Switzerland were published in the scientific literature and well before a real product or even a proof-of-concept with sufficient levels of beta-carotene expression to have nutritional benefit to humans had been developed. Today, in the aftermath of a decade of well-publicized yet underperforming innovations like the FlavrSavr® tomato or GoldenRice®, is there anything in the biotech R&D pipeline with nutritional or quality traits that has real substance and the potential for commercial success? Is there anything that stands out from what can be achieved through conventional breeding? Are there in fact enough such innovations coming along to characterize this as a viable ‘generation’ of agricultural biotechnologies? And, of those innovations that are forthcoming, how mature are they, and when might we expect to see them begin to penetrate agricultural markets? Who is doing the research and development work on this type of technology and where? What are the factors that make this sort of innovation economically viable? And what kinds of economic, social, and environmental impacts might they realistically be expected to have and on whom? In sum, can we in fact expect to see a transition to a new generation of agricultural biotechnology, and what would such a transition entail for all of the different parties impacted?

1.3

Assessing the R&D pipeline

Progress in the R&D pipeline of even one firm, let alone an entire industry, can be quite difficult to track. Uncertainty is the rule, with new product concepts formed only to become quickly outdated as new developments occur. Promising leads are constantly reassessed, updated, or abandoned. Progress in the early-stages of primary research can be even harder to track, especially given the scope of relevant scientific sub-fields involved: research in plant storage proteins, fatty acid biosynthesis, sugar, starch, and fructan biosynthesis, mineral accumulation and bioavailability, biosynthesis of secondary metabolites such as vitamins, isoflavones, and plant pigments, and cell wall chemistry, to name a few. Yet, an assessment of the R&D pipeline of product quality innovations is important for a wide range of stakeholders—including consumers, industry, investors, environmentalists, and policymakers—in order to form realistic expectations, make informed decisions and wise investments, and shape effective policies. Similarly, it is important for all stakeholders to understand who stands to benefit and how much from these new types of technologies, given that there will likely be a very different alignment of economic, social, and environmental impacts than there has been from the existing pest control agricultural biotechnologies.

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Typically, corporate product development teams and their public relations departments have provided most of the information available to the public on new and future products. The commercial interests of these companies potentially bias such information, at the very least by featuring only their own subsets of anticipated new products. These limitations can be overcome by looking across the full scope of R&D institutions engaged in plant genetic research and trait development globally, and conducting a thorough comparative analysis. The landscape of possible future product development candidates can be inferred by reviewing the scientific literature, as commercial product candidates will likely be drawn from the pool of transgenic plants that have proven feasible for research purposes. Other public data resources tracking R&D activity for this field exist as well, including public records of field trials and regulatory filings. This study seeks to be as thorough and broad reaching as possible in developing a picture of the range of nutritional and product quality innovations that have become technically feasible, as well as those likely to be commercialized over the next five to ten years. It does so by two methods. The first is a broad survey published primary and secondary data sources, including scientific articles, public data on field trials, and public regulatory filings. This creates a dataset characterizing the broader R&D pipeline, of which we give a summary overview in Chapter 3 and a more detailed cataloguing in Chapter 4. The second method is a compendium of primary sources on plans for product commercialization, including interviews with company officials, and research administrators at university and public institutes, as well as official primary publications released by the innovating organizations. This creates a schedule of anticipated product releases for the next five to ten years, detailed in Chapter 5. What emerges is an intriguingly detailed snapshot taken at one point in time of an ever evolving and highly dynamic flow of new ideas and product development projects within a global research community and industry. The picture that results might be better described as an R&D ‘funnel’ than an R&D ‘pipeline’. With a vibrant base of research in functional plant genomics that has been actively generating hundreds of candidate traits over the last decade or so, new genes are being discovered and their functions elucidated by researchers at a broad range of institutions—in both the public and private sectors—around the world. Yet, moving from an initial discovery to a released product entails successfully navigating a sequence of increasingly costly and difficult challenges. R&D is perhaps most accurately described as a selection or filtering process, continually testing and narrowing the field of potential candidates, with significant feedback, crosstalk, and recalculation along the way. While advances in science make product innovation possible, market demand is what makes product innovation worthwhile. In economic terms, these dual forces of ‘technology push’ and ‘demand pull’ are both required, and neither one is sufficient on its own. Many interesting genomic discoveries simply do not correspond to any viable market demand, or at least demand viable enough to make development worthwhile given the requisite costs. Likewise, simple technical feasibility is a serious issue even when there may be viable market demand. Many nutritional and quality characteristics of plants derive from highly complex and dynamic biosynthetic pathways, and these can be sufficiently complicated to prevent achieving sufficient levels of trait expression to be economically viable. Of the few products that appear to be both economically and technically viable, each must still negotiate a field of intellectual property claims, comply with regulatory requirements, and attract sufficient funding to get developed, scaled up, launched, and marketed. Then the final test comes: in the highly competitive environment of the marketplace, a new crop product must be adopted by multiple players in an often complex agricultural value chain. Out of hundreds of potentially

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interesting genetic discoveries, only a handful will ever find their way into crops being commercially cultivated. 1.4

Objectives

The specific objectives of this study are the following: •

To asses the range of markets in which there may be demand for agricultural products with new nutritional and quality characteristics.

•

To survey and describe the nutritional and quality innovations that are, or have been, in the plant biotechnology R&D pipeline, including: o Those already commercialized o Those closest to commercialization and more certain, primarily those that have already gone through a number of years of field trials or that have reached the regulatory approval phase. o Those further from commercialization and less certain, such as those recently reported in the scientific literature or that have just commenced field trials.

•

To understand the dynamics of the R&D pipeline in these agricultural biotechnology traits

•

To asses which types of organizations—such as government labs, universities, agricultural biotechnology companies, or food companies—are pursuing this kind of R&D and which countries tend to be home to this kind of R&D

•

To analyze the full range of economic, social, and environmental impacts that nutritionally and quality enhanced agricultural products can be expected to have when deployed including: o Impacts on consumers and final market demand. o Impacts on midstream activities in the food chain such as livestock, food processing, and paper goods manufacturing. o Impacts of extended shelf life on fresh produce value chains and consumers o Impacts on the environment.

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2 Agricultural markets and factors governing demand for nutritional and product quality innovations The pull of market demand—that final vote by consumers with their pocketbooks—is what ultimately drives the selection of product innovations and determines their success. A survey of the marketing literature finds demand-related factors predominate among the most important determinants of successful new product innovation (Stewart-Knox and Mitchell, 2003). Such factors include: • the uniqueness and superiority of the new product, • the innovator’s understanding of consumer wants, needs, and preferences (premarket consumer research), • the effectiveness of product launch and marketing, • the effectiveness of communication within the product development team, and • the support and involvement from top management in the innovation process. Accordingly, the two most commonly cited causes of failed new product innovation are • a lack of market knowledge and • technical problems in the new product. In the aftermath of the initial technological breakthroughs of agricultural biotechnology in the mid-1980s, it has been primarily a phenomenon of ‘technology push’ factors, ushered along by advances in scientific and technological capability, rather than as a result of immediate changes in consumer demand reverberating through the food system. It has largely been analyzed from the perspective of technology advance, with the question of demand often framed as an issue of ‘consumer acceptance’ of the new technology. In order to circumvent what has by now become a well worn discussion, we begin by considering in brief detail what the demand conditions are in the most important markets potentially affected by advances in plant biotechnology. An understanding of these conditions is all the more crucial as biotechnology shifts from delivering grower-oriented pest control traits to consumer- or user-oriented quality traits. The main uses of quality traits being developed in biotechnology are likely to be found in food retail, food processing, livestock and dairy feed and forage, cut flowers and landscaping, and pulp and paper. In considering demand conditions in these respective markets, we must keep in mind that agriculture is a global undertaking: its products are made, traded, and used world wide, and the needs of users vary widely, depending upon their country, culture, and level of economic development. Considering the ‘demand pull’ factors—including the relative sizes of these potential applications, the nature of the market and non-market channels that supply them, and the major trends within each—can provide insights into which kinds of consumer- and user-oriented biotech innovations may be successful. Several additional complexities of the agricultural value chain must be kept in mind as well. Some crop products are directly used by final consumers. In these cases, direct demand for certain quality characteristics, as expressed by consumers in the final market, is precisely what determines the uptake and success of the crop. Other products, however, are purchased by intermediaries along the value chain—such as food processors, livestock and dairy operations, or paper manufacturers—who use some components of harvested plants to produce their final products—such as prepared foods, meat and dairy products, or paper goods—which are then sold to final consumers. In these cases, demand as expressed by those intermediaries, in terms of

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ingredient specifications, quality criteria, and formulation requirements, is what determines the adoption and success of an innovation. Such quality specifications reflect what economists call derived demand when the intermediary’s choices are significantly influenced and constrained by the consumer demand they face in the final market for their output and they are, in essence, buying those ingredients on behalf of the final consumers. This is usually most pertinent for product technologies, and optimization of their final product’s quality is the primary consideration. In contrast, quality specifications may also reflect only the intermediary’s own demand when the choice does not qualitatively affect the final consumers’ choices but may impact the efficiency of production. This is most common for process technologies, and simple minimization of process costs is usually the driver for demand. Consumer market demand reflects three important components: consumer preferences (“What you need or like”), available consumer income (“What you can afford”), and other consumer constraints (such as “What you have time or energy to use”). The value created by a product innovation—and ultimately, its success or failure—will rise or fall on how well it satisfies these three components. Successful product innovations are preferred by consumers, if they are cost competitive vis-à-vis the next best available product on the market, and they work within the consumer’s constraints of time, energy, and ability to use them. Agricultural R&D serves not only a wide variety of consumer and intermediate markets, but it also serves degrees of human need beyond the reach of formal markets. Many crops in the world are grown by the very people who intend to consume them. Such subsistence growers are often eking out an existence separated economically, socially, or geographically from the commercial systems of the industrialized world. Yet, agricultural R&D projects supported by governments, foundations, and aid organizations target the needs of subsistence growers, as well as the needs of poor food consumers that may purchase a portion of subsistence growers’ harvests. 2.1 2.1.1

Markets in the industrialized world Food retail

In the industrialized world, food and beverage value added consistently make up about 3 percent of GDP and have for the last 30 years (OECD, 2003). However, food retail is one of the largest sectors of final goods manufacture and sales in the economy, with food accounting for 10% to 15% of household consumer expenditures. In the U.S. $427 billion was spent in 2002 on retail food for home use (USDA-ERS, 2003) and $417 was spent on food consumed outside of the home in restaurants and food service establishments (USDA-ERS, 2004). Major trends being observed among food consumers in North America, Europe, and other industrialized economies include: • Better taste • Convenience: precut, prepared, precooked, or ready-to-eat • Distinctiveness: varieties, flavors, texture, look, patterns, package design • Health and nutrition: vitamins, antioxidants, high-fiber, herbs, functional components • Low calorie, low fat or fat-free, low carbohydrate • More natural, environmentally friendly, or organic • Gourmet, up-market, or restaurant quality

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Figure 2.1 Value added from food products and beverages as percent of total GNP for OECD countries 4%

3%

2%

1%

0% 1970

1975

1980

1985

1990

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2000

Data source: (OECD, 2003)

There is substantial specialization within the structure of the retail food industry. Major food suppliers and manufacturers tend to focus on major brands for which supply and quality can be standardized broadly and which can be shipped, stored, and marketed in multiple countries and cultures. For example, Nestle concentrates on branded chocolates, instant coffees and teas, infant formulas, and prepared foods. Unilever, Procter & Gamble, Kraft, and PepsiCo similarly follow the formula of globally scaleable brands. Local producers, on the other hand, tend to specialize in more diversified and localized products, particularly in fresh sectors of the market like produce, dairy, or bakery products. Interestingly, there also appears to be some specialization within innovation in the retail food industry. It is often the smaller, more specialized local producers that are observed to introduce radically new product innovations that invite and affect major changes in consumer habits. Examples include Sara Lee, a Chicago bakery that innovated frozen baked goods in the 1950s and 60s. Snapple grew from a small natural foods company in New York City by innovating fresh fruit drinks in the 1970s and 80s. PowerBar was founded in California by a former marathon runner and innovated sports energy bars the 1980s and 90s. Once such innovations become widely successful, the successful ‘startup’ firm usually becomes acquired by one of the major food companies, which has the requisite financial and marketing clout to reach much wider markets with the proven product: Sara Lee was bought by Consolidated Foods in 1956; Snapple was purchased by Cadbury-Schwepps, and PowerBar was bought by Nestle. This model is also familiar in the biotechnology and high technology sectors, and in all cases, such market-changing radical innovations are rare events. The more typical pattern of product innovation is characterized by a very high churn rate in more incremental innovations. Given fairly low barriers to entry, the regional nature of many products, and the imitative nature of most food product ‘innovations’ (with estimates of just 7 to 25 percent that can be considered truly novel), a large number of innovations are attempted, but at least 7580 percent fail, (Stewart-Knox and Mitchell, 2003).

8



Table 2.1 Recent top selling new product innovations in U.S. food retail 2000-2001

1. Boston Market frozen meals 2. Starbucks ground coffee and beans 3. Breyers Ice Cream Parlor ice cream 4. Uncle Ben’s Bowl Entrees

1st year sales* (mil $)


2001-2002

2002-2003


118.2

1. Pepsi Twist

174.6

1. Vanilla Coke

292.1

117.9

2. Mountain Dew Code Red

163.1

2. Krispy Kreme doughnuts

235.5

82.2

3. Diet Coke with Lemon

139.1

160.0

79.7

122.5

3. Sara Lee Fresh bread and rolls 4. Michelob Ultra low-carb beer 5. Berry Burst Cheerios

108.9

6. Oreo Double Delight cookies 7. Yellow Tail Australian wine 8. Gatorade Ice sports drink

78.4

9. Dannon Light 'n Fit Creamy yogurt and cookies 10. Masterfoods USA ‘Cookie&’ cookie bars

63.8

5. Frito Snack Kits

74.1

4. Marie Callenders Complete Dinners 5. Smirnoff Ice malt ale

6. Frito Ruffles flavored potato chips 7. Mini Oreos cookies

72.7

6. Sierra Mist soft drinks

69.6

95.9

8. Kraft Philadelphia Flavors flavored cream cheeses 9. Kraft Deli Delux delistyle cheeses 10. Dannon Danimals yogurt drink

66.0

7. Banquet Homestyle Bakes dinner kits 8. Marie Callenders OneDish Classics frozen entrees 9. Yoplait Whips yogurt 10. Kool Aid Jammers juice drinks

83.2

57.7 52.4

113.9

92.7 85.8

156.1 88.5

77.3 65.8

54.0

*Initial 52 weeks of sales in supermarkets, drug stores, and mass merchandisers. Source: (IRI, 2003, IRI, 2004, IRI, 2002)

The most successful product innovations in the U.S. food retail sector in recent years (see Table 2.1) are those that deliver better tasting, better quality, up-market foods (such as Boston Market meals, Starbucks® coffee, Breyer’s® ice creams, Kraft® deli quality cheeses, Marie Callenders® meals, Krispy Kreme® doughnuts, and Yellow Tail® wine), greater convenience (such as Uncle Ben’s® entrees, Frito® snack kits, Dannon® yogurt drink, and Sara Lee® fresh bread), or simple novelty—often with just a slight modification, such as a new flavor, in a familiar well-established brand (new soft drink flavors, flavored Frito’s Ruffles®, smaller and flavored Oreo® cookies, flavored Philadelphia® cream cheeses, and flavored Cheerios®). Few of the top performers deliver health attributes, and even those that appear to so (such as low-carb Michelob® beer and Dannon® light yogurt and cookies) seem to be at least as well suited for easing the consumer’s conscience as for easing the consumer’s waistline. For at least the next decade, biotechnology will likely be a challenging tool to use for direct branded product introductions in the fast and fickle world of food retail. Lessons may be learned from Calgene’s foray into fresh produce markets with the FlavrSavr® tomato in 1994-1997. Biotechnology requires long-term investment and long product development cycles, while retail food markets tend to involve rapid deployment cycles, with a high rate of failure. These low product survival rates are not unusual in consumer markets and are compatible with the relatively low cost of product introduction, particularly for consumer products that do not require government regulatory approval. In contrast, the relatively high costs of regulatory approval associated with new product introductions in the pharmaceutical and agricultural biotechnology sectors could not tolerate such a high rate of product failure. This does not mean, of course, that significant improvements in plant genetics could not be deployed in carefully selected retail brands—for example, a ‘GoldenRice®’ breakfast cereal or a high omega-3 bottled vegetable oil. However, as a general rule, we would expect to see a high rate of failure among any attempts at biotechnology-based new food products, simply given the natural rate of attrition among retail

9



food innovations and, as described above, such an anticipated failure rate is incompatible with the costs of product introduction. Those that might succeed will do so by • delivering clear and present gain to the consumer, • not compromising other existing components of quality, • adding the new characteristic to already top quality products or crop material, and • targeting a well understood segment of the food retail market.

2.1.2

Food commodities and food processing

Upstream from retail food markets, a large proportion of the output from crop production is purchased on commodity markets by food processors and used as ingredients in food manufacturing. In the U.S. the value exceeds $US 45 billion (Harris, et al., 2002). Such intermediate agricultural commodities destined for food manufacturing also account for significant shares of international trade flows, particularly from developing countries. Figure 2.2 Purchases by food manufacturers of food ingredients from U.S. crop agriculture (includes fruits & vegetables, bakery products, grain mill products, and others) 50 45

US$ billions

40 35 30 25 20 1982

1987

1992

1997

Data source: Harris, et al, USDA-ERS, 2002

The nature of the food processing industry hinges on providing retailers and food service with food products that consumers want, while depending upon ingredients sourced globally and often affected by the volatility of commodity markets or the variability of specialty sources. While following consumer trends on the demand side is crucial for food processors, they face important constraints on the supply side, and with the low-margin high-volume nature of the business, success and even survival often turns on marginal costs. Innovation and new product development for commodity food products and food ingredients varies significantly by product category (listed in Table 2.2.) Some categories are fairly mature, while others are evolving rapidly. Yet on average, the rate of innovation and introduction of new products or product categories is less frenetic than in retail food, allowing and leading food ingredient suppliers and processors to consider longer term consumer trends for opportunities to innovate.

10



Table 2.2 Major product categories of food manufacturing ingredients Baking, cereal, and snack ingredients Chocolate and confectionary ingredients Cultures, enzymes, yeasts Dairy ingredients Emulsifiers, stabilizers, hydrocolloids Fats and oils, fat substitutes Fibers, dietary fibers, prebiotics Flavors, colors, flavor enhancers

Fruit, vegetable, and nut ingredients Functional foods, neutraceuticals, and herbals Meat, poultry, and fish ingredients Preservatives and additives Proteins (non-dairy) Savory ingredients (salt, spices, seasonings) Sweeteners, sugars Others…

The shorter term consumer trends to which food processors can respond with short term innovations include things like the low-carb diet (the popularity of which has certainly peaked in the U.S.) Longer term consumer trends that guide innovation in food ingredients are those that follow major demographic shifts, such as the ageing of the Baby Boomer generation, new food habits among the younger Generations X and Y, broad public responses to major public health issues, like obesity, changes driven by regulatory requirements, such as labeling, and changes based upon new knowledge of nutritional impacts derived from new research. In one recent example, starting in 2006 labeling requirements were introduced in the U.S. for trans-fat content, based upon new understandings of the negative health affects of trans-fats, which drove significant innovation and product reformulation to substitute for trans-fats. Suppliers have also been looking for ways to meet demand inspired by increased consumer interest in health affects of omega-3 fatty acids. Another example of a long term, even chronic, need is control of allergenicity of food ingredients like soy, wheat, or peanuts, which could serve large subsegments of the population with a diagnosed or suspected allergy to one or more of these ingredients. In considering biotechnology-based innovation in the nutritional and product quality characteristics of food ingredients, an illustrative case study is the novel non-absorbing fatsubstitute ingredient, Olean®, introduced by Procter & Gamble in 1996. Olean® represents a perhaps more radical innovation, in biochemical and nutritional terms, than anything being contemplated today using biotechnology. The molecule is a combination of sucrose and fatty acids that mimics the structure and properties of fat. The initial discovery by Procter & Gamble food chemists was made in 1968. Consultations with the FDA began in 1971, and because of the unprecedented nature of the compound, it took a number of years to resolve how it should and would be regulated. FDA filings for Olean® as a novel food additive were first submitted in 1987. Primary safety issues raised were the leaching of fat-soluble nutrients, as the unabsorbed molecule passed through and out of the digestive tract, as well as some incidence of gastrointestinal problems, such as diarrhea. After extensive review Olean® was approved in 1996 as a food ingredient for salty snacks, but initially was required to bear a label warning consumers about the possible side effects. Throughout the process, before and after the product gained regulatory approval, Procter & Gamble was persistent and proactive in marketing the ingredient both to their direct customers among major firms in the food industry and to final consumers. Consumer demand for fat-free snacks proved to be significant, and today Olean is used by FritoLay in fat-free Lays®, Ruffles®, and Doritos® chips, by Nabisco in fat-free Ritz® crackers, and by Procter & Gamble in fat-free Pringles®. Biotechnology-derived food ingredients will in most cases involve a considerably less novelty than did Procter & Gamble’s Olean®. Biotechnology is likely to be used to reduce or elevate

11



levels of naturally occurring components. However, as a consequence of the innovation process of genetic engineering, issues such as safety and regulatory considerations, longer pipeline time horizons, and consumer acceptance issues may be similar. For those cases where biotechnology can introduce food processing ingredients with truly novel characteristics, a branding strategy similar to that pursued by Procter & Gamble with Olean®—or that used by J.D. Searle with NutraSweet®—may be feasible, enabling the consumer to associate the brand with a desired characteristic on order to specifically demand final products that contain that particular ingredient. Instead of taking a leading role in the public eye via branded product or ingredient at the retail level, the early quality innovations from biotechnology are more likely to increase processing efficiency and drive down costs of producing existing ingredients without being observed by the final consumer. In this context, biotechnology is more likely to play a supporting role at the level of the processing ingredient market. Regardless, such impacts could be pervasive in the food supply and be economically quite significant. 2.1.3

Animal feed and forage

The single largest use of crops, particularly within more industrialized economies, is not for human consumption but to feed the large populations of animal that produce billions of pounds of meat, dairy, eggs, and other animal products. (See Figure 2.3.) As of 2002 animal production makes up on average 1.7 percent of GDP in the industrialized economies and is on average 20 percent larger in value than crop production (OECD, 2003).

Figure 2.3 U.S. animal populations and output in 2002

50 million mature beef cattle

27 billion pounds of beef

9 million dairy cows

170 billion pounds of milk

37 million calves, heifers, bulls etc.

160 million tons of grain 9 billion broiler chickens

41 billion pounds of chicken

340 million egg producers

87 billion eggs

275 million turkeys

7 billion pounds of turkey

150 million tons of hay 60 million hogs & pigs

20 billion pounds of pork

11 million sheep & lambs.

218 million pounds of meat

12



Data sources: (USDA-ERS, 2005, USDA-ERS, 2005, USDA-ERS, 2005, USDA-ERS, 2005)

The two most important crops fed to animals are corn and hay. Of the annual U.S. corn harvest 70 percent—worth $13 billion and accounting for about 40 million acres of land in 2002—goes to feed animals. This corn makes up over 90 percent of the total grains fed to animals in the U.S. The remaining ten percent is made up by other more minor feed grains including sorghum, barley, oats, and wheat (see Table 2.3). The rest of animal feed comes from forage crops such as hay—a crop that was worth about $14 billion and used about 65 million acres of land in the US in 2002.

Table 2.3 U.S. harvests and uses of major feed crops in 2002 Crop

Total acreage harvested

Total domestic use (disappearance)

million acres

million bushels

million bushels

7935 205 248 222 1115 151 million tons

5650 160 75 150 113 151 million tons

65 Corn 7 Sorghum 4 Barley 2 Oats 46 Wheat 65 Hay Data source: (USDA-ERS, 2005)

Domestic use for animal feed as percent of total domestic use 71% 78% 30% 68% 10% 100%

In the market for animal feed and forage the biggest concerns are cost, feed efficiency, and compliance with food safety regulations and environmental impact (for instance, avoiding feed ingredients that can cause problems like Mad Cow disease). Among these, cost is the most powerful driver in regular decision making by producers. Livestock, poultry, and aquaculture operations are regularly reformulating feed rations in response to shifts in relative prices among possible feed ingredients, to take advantage of cost savings while maintaining optimal animal weight gain. Even a small difference in unit price of a feed used can make a large difference in profitability. Biotechnological innovation is poised to make some of its most significant contributions in the area of animal nutrition. Any increase in feed efficiency or nutritional content will figure into the costs faced by animal growers. Such improvements also have the potential to be environmentally beneficial. Gains in efficiency can be translated into a decrease in the vast amounts of land needed to grow feed. Better absorption of nutrients by animals means that unabsorbed nutrients such as phosphorus or nitrogen will not come out in the waste stream, reducing pollution problems from animal manure. And, there are fewer labeling and consumer acceptance issues to face when the consumers of the product are animals. 2.1.4

Floriculture and nursery products

Floriculture products include cut flowers, potted plants, and garden plants, while nursery products include landscaping trees, shrubs, groundcover, and turf grass. Rough estimates of markets for ornamental flowers and plants put global production value at around $50 billion, and retail value at around $100 - $150 billion (Chandler, 2003). More specific figures from the USDA estimate

13



the production value for floricultural and nursery products in the U.S. to be $14.3 billion in 2003 (Jerardo, 2004). Consumers look for three things in cut flowers: novelty in colors and arrangements, freshness and vase life, and convenience of purchasing and handling. Novelty is of utmost importance, for while a few traditional colors and arrangements continue to sell, cut flowers are something of a fashion good with consumers regularly responding to a new ‘look’. Freshness is also of major concern, as the duration of lifespan of the product with the consumer being a function of how much time is left after cutting, shipping, and shelf time in retail. The time-critical nature dictates the use of technologies for rapid transportation—including jet cargo—and preservation— including refrigeration and chemical preservatives. The solutions most commonly used contain silver ions such as silver thiosulfate, which are toxic and potential pollutants. Significant labor is expended changing solutions, and at least some consumers may be wary of the extent of chemical treatment. In landscaping products, low maintenance (ease of care), durability (hard to kill), and esthetic characteristics are of greatest important to consumers. Maintenance costs, however, actually represent a larger cost to the consumer than do the initial purchase and planting of landscaping and turf varieties. Preferences are also increasingly informed by the emergence of environmental awareness, out of both personal safety concerns and regulatory constraints. The market is also expanding beyond the traditional corporate landscaping and backyard gardens into smaller scale settings like window boxes. Innovation in floricultural and nursery products has been primarily in marketing and automation of production and distribution. Retail outlets have migrated from florist to supermarkets and home improvement stores as well as onto the internet, complemented by advances in just-in-time inventory management and delivery systems. Breeding has long been an important source of new product innovations, but as in any population or species, the gene pool of a given flower or turf grass only offers a limited palette of new traits and trait combinations. Biotechnology can be utilized to provide novel colors, extended shelf or vase life, modified plant architecture, and enhanced floral scent. The most advanced of these traits are in color modification and vase life. Biotechnology can be used in turf grass to make it easier and less costly to maintain, saving in costs and environmental impacts.

2.1.5

Pulp and paper

Forestry, wood, and paper products globally contribute $354 billion or 1.2 percent to global GDP annually. Of the total wood harvested globally, roughly one third is from plantation forests, consisting of planted and managed agroforestry systems, while the remainder is gathered from public and private forest lands (FAO, 2005). In the U.S. about 30 percent of wood harvested is used to make pulp and paper (EIA, 2000), and over $50 billion is spent on paper goods annually (BEA, 2004). Technology and innovation in pulp and paper have involved advances in both the mechanical and chemical processes of pulping, the recovery of chemicals from spent pulping liquor, processes for bleaching and removing lignin, and paper manufacturing, including sheet formation and finishing. Only over the last 50 years, as trees have been increasingly planted, has it become increasingly important to consider the genetics of the trees being used. It has been estimated that genetically improved fiber characteristics could save $10 per cubic meter in reduced pulping digester costs 14



and genetically reduced lignin content could save $15 per cubic meter in reduced pulping costs (Sedjo, 2005) and in addition could save significantly in energy and chemicals used and thus opportunities to reduce environmental impact. Other biotechnological innovations being developed for paper are in the level and composition of starches from potato and cassava that are used in paper manufacturing and finishing. 2.2

Formal markets in developing economies

Baseline growth in the overall demand for agricultural products in the developing world is projected to be much stronger than in the industrialized world for the foreseeable future. This is largely a function of simultaneously growing populations and incomes. Projections by the International Food Policy Research Institute, forecast demand for meat to rise by more than 55 percent between 1997 and 2020, with most of the increase occurring in developing countries. Increased meat consumption translates into greater pressure on agricultural systems, both through increased demand for feed grain and forage land and through increased animal waste volume. Developing countries are also projected to account for most of the 654 million metric ton increase in demand for cereals through 2020, with China alone accounting for 27 percent of the increased demand for cereals, India for 12 percent, all of Latin America for 11 percent, all of Sub-Saharan Africa for 11 percent. All of the world’s developed countries are projected to account for just 15 percent of the increase in cereal demand (Rosegrant, et al., 2001). In the last five to ten years, modern food supply and retail systems have expanded greatly throughout emerging and developing economies. Infrastructure and standards still vary greatly around the world, but a more formalized, centralized, and standardized food industry is being developed. Initially serving the highest income segment of consumers, entrepreneurial retailers are expanding by serving the growing urban middle class and the even larger segments of the poor, in urban as well as more rural regions of the developing world (Reardon, et al., 2003). Formal food markets in developing countries, those serving upper and middle income consumers, are thus becoming increasingly similar to those in developed countries, meaning that food quality innovations intended for developed country markets can also find a significant consumer base in the developing world. The most significant shift in demand appears to be the transition to a greater reliance on animal-based diets in highly populous developing countries. Two key factors drive the formalization of developing world markets. First is continued migration from rural to urban areas. Second, and perhaps more significant, is the emergence of a retail business model that intentionally seeks to serve local mass markets, particularly understanding and catering to the needs and buying habits of middle and low-income households. In the U.S. this kind of model has been championed by retailers like WalMart, but in developing countries it is just as often driven by local retailers. Elements of the model include contracting with suppliers over standards of timing and quality of deliveries, driving down costs and prices, increasing the number and the convenience of outlets, and exercising economies of scale in the middle to lower ranges of the market. These contracts will often dictate production technologies. Indeed, if a characteristic that meets consumer specifications is available from a biotechnology, it is likely retailer contracts that will determine whether farmers supplying that country’s market will adopt the biotech crop variety. As one or two retail chains introduce new products that meet consumer needs and lowering costs, successive rounds of imitation and adoption will follow among others.

15



Figure 2.4 Growth in demand for meat and cereals in developing countries is expected to well exceed that in developed countries Demand for meat 250

Million metric tons

200

developed countries developing countries

150

100

50

0 1974

1997

2020

Demand for cereals 1800 1600

Million metric tons

1400 1200 1000 800 600 400 200 0 1974

1997

2020

Data Source: (Rosegrant, et al., 2001)

A recent industry report on Brazil’s processed food market (Euromonitor International, 2003) illustrates these trends of market formalization, convergence of consumer demand with the developing world, market penetration by local cost cutters, and the latent demand left in the market. Key market drivers in Brazil increasingly include product innovation and packaging. Brazilian consumers are showing increased demand for convenience and time savings in preparation, light and diet products, and functional food products. While familiar multinational firms are the market leaders in processed foods—with Nestle on top with 7% of the market, followed by Parmalat and Unilever—it is the smaller regional manufacturers—offering comparable quality to the multinational companies—that are increasing share of the processed foods market, now at 24%. They are doing so by taking advantage of local market knowledge and offering lower prices. In food retailing, supermarkets and hypermarkets', following the WalMart model, have increased their share from 50% to 57% in the last five years. And, food manufacturers in Brazil are increasingly targeting low-income consumers, who consist of nearly 31 million households and represent unrealized consumption potential of 42% of the national market.

16



Trends in India are similar to these in Brazil, but the scale of underserved or latent markets is even larger, according to a recent report by Rabobank (2004). Only 1% of food in India is currently sold by organized food retailing businesses, representing about 10 million food consumers, with the rest of the Indian market currently being served by highly fractured and informal food distribution channels. In Brazil, India, and other developing countries, local companies may have significant advantages over their global competitors in introducing technologies to meet this latent local demand. On the supply side, if there are no relevant patents in force in the local country on the genetic technologies underlying appropriate product innovations, local operators may be able to copy and sell these innovations locally without paying royalties (Binenbaum, et al., 2003), while global competitors invested in the original R&D or paid royalties for access to the technology where it is patented in the major markets. On the demand side, local companies developing the retail markets are often able to pay closer attention to local conditions of demand and are more likely to see and exploit unique opportunities for using and applying innovative technologies, including local food traditions (new foods can be incorporated into the traditions and lifestyles of different cultures, but it is better done by locals), peculiarities in food and retail infrastructure available to consumers, differences in climate, humidity, and temperature, as well as local health issues and characteristics of genetic dispositions of local populations and thus the nutritional or functional food needs (e.g. such as the 'stingy gene' hypothesis of Diamond, 2003). Finally, as in the developed countries, dietary concerns among upper, middle, and now even lower income populations in the developing world are evolving rapidly from the challenge of merely getting a sufficient quantity of calories, to the challenge of getting a sufficient quality of nutritional content, particularly lower levels of saturated fats and cholesterol, and more complex carbohydrates and micronutrients, in order to help counter diet related problems like obesity, diabetes, and cardiovascular disease.

2.3

Informal and missing markets in developing countries: innovation needs for subsistence agriculture and food security

A large proportion of the world’s population fall beneath the income levels needed to access sufficient quantity or quality of food at current prices and availability on formal food markets. For those at such levels of poverty, finding regular, nutritious meals is a serious challenge, or can quickly become so given the precarious conditions under which they live. Natural calamity, political unrest, or other external factors can drive local price of food above a poor households' reach. Even when able to maintain sufficient calories from staple foods, many are not able to afford the more nutritious, non-staple foods like fresh vegetables, meats, and dairy products. They are thus unable to consistently get minimal levels of essential amino acids or micronutrients (vitamins and minerals). Such dietary imbalance or micronutrient deficiencies, described as ‘hidden hunger’ (Bouis, 2004), can lead to a range of health problems and decreased productivity, undermining ability to maintain subsistence-level production of food or income from other employment. Malnutrition and poverty can thus trap their victims in an ongoing negative cycle.

17



Food as medicine in low income and high income countries alike Better nutrition—from wide availability and lower cost of fresh fruits and vegetables, nutritionally-enhanced, and functional foods—can both make up for and supplement limited available health care for the poor and even the emerging middle class in developing countries. Because of the state of healthcare, the use of nutritionally rich, enhanced, and functional foods may be comparatively more important to consumers in the developing world. A traditional version of this is already seen: in regional street markets around the world, vendors can be found peddling a wide array of herbal remedies that are based on the region’s traditional practices or folk medicine. If a patient in a wealthy country with good quality medical care will turn to a diet of ‘heart healthy’ foods after a heart attack, all the more so will a citizen of a poor country, lacking the same quality of medical care, rely on a diet of known ‘heart healthy’ foods to increase his or her own chances of recovery and survival. However, with such diets, consumers are often left to self-diagnose and make their own partially-informed decisions. Even physicians and nurses are often not well trained in nutrition, let alone the properties of functional foods. Providing credible health information and recommendations based on clinical research about the health-promoting uses of nutritionally enhanced and functional foods is an area where health officials even in resource constrained developing countries can provide real public health benefits at minimal costs. If the availability of foods with particular nutritional or functional qualities is limited, clear information from health authorities could help create market opportunities for local growers in the region to produce them. And, as growers’ associations in the industrialized countries know, sharing information about clinically substantiated health benefits can go a long way in promoting the consumption of a product: consider the success of soy and the image that soy has gained among consumers as a health promoting food. A generation ago medical recommendations concerning the benefits of cod liver oil for patients after heart surgery drove U.S. demand for that fishy supplement into the billions.

Agricultural applications of biotechnology may be helpful in alleviating malnutrition, by making essential nutrients affordable to even the poorest. The challenge is to identify a subset of genetic traits that impart qualitative characteristics that would have wide appeal and bring real value to poor segments of the population. Deborah Delmer, who oversees biotechnology strategy for the food security programs of the Rockefeller Foundation, explains that biotechnology can be an appropriate and particularly useful tool when other technical approaches cannot deliver needed results. Such situations arise if a crop is difficult to breed (such as banana or cassava), if genes for a trait do not exist in available breeding germplasm, if multiple genes contribute to a needed trait, or if genes from outside that species need to be introduced (Delmer, 2004). Two examples of nutritional traits may fit these criteria. Researchers working on sorghum for use in Africa point out that the ability to improve on sorghum seed protein’s poor nutritional quality by classical plant breeding is limited by the low level of variation in the sorghum gene pool available for crossing. Instead, genes imported from barley and other grains can serve to make up for the lack in the sorghum gene pool (Forsyth, et al., 2003). In models that simulate the cost effectiveness of different types of interventions for delivering vitamin A to deficient populations

18



in Asia, it was found that transgenic GoldenRice strongly exceed other interventions including gardening and education programs, conventional food fortification, and the distribution of vitamin supplements (Albrecht, 2002). To maximize positive impacts for subsistence growers and to mitigate agronomic problems that can lead to yield drag, the desired traits need to be imparted to existing locally adapted crop varieties. Publicly funded R&D infrastructure and outreach systems are in place in many countries to develop and disperse improved varieties, including networks to disseminate seeds to poor farmers. These include the international Future Harvest research centers of the Consultative Group on International Agricultural Research (CGIAR), national agricultural research services, and in most countries regional private seed companies and agricultural supply vendors that act as the final link in distribution of new agricultural genetics. A transgenic crop that is appropriate for a local climate and cropping system, is proven to be safe, and is made accessible to smallholders on reasonable terms can act to complement any of the other poverty alleviation priorities advanced above. Still, the basic economics of demand and technology adoption operate just as much in informal market settings near the bottom of the socio-economic pyramid as they do in formal markets nearer the top. Subsistence growers and poor consumers have preferences, budget constraints, and other lifestyle constraints—all of the components of demand—and these determine whether or not they will embrace and use a particular innovation. Constraints of budget and time are all the greater among the poor, and preferences can often still be powerful determinants. Fundamentally, what supports technology adoption and impact is human decision making behavior. The governments, foundations, and agencies that invest in agricultural R&D seeking to generate technological public goods to impact poverty must keep such demand and adoption criteria foremost in mind, just as much as companies seeking to innovate profitable goods for market.

2.4

Regulatory approvals, perceived risks, and market demand for the products of biotechnology

An important factor determining the level of demand in the marketplace for a technology product is the public perception of that product. Careful scrutiny by public authorities can contribute to the confidence of consumers by holding innovators standards that are commensurate with the novelty of the new product and that are at least as high as those for conventional food products. In considering nutritional and product quality innovations from plant biotechnology a task force recently convened by the International Life Science Institute (ILSI) reviewed the scientific literature and the framework for food safety testing and nutritional evaluation at major national regulatory bodies (Chassy, et al., 2004). Their assessment concludes that, in general terms, the methods that are currently in place are sufficient to detect any significant differences between a genetically modified product and its conventional counterpart in terms nutritional quality of and impact on human health. When differences are identified, several testing protocols are in place that can be utilized, depending upon the needs of the case, including compositional analysis, metabolite profiling, animal feeding studies, and, under certain conditions, continued post-market monitoring. This regulatory framework is, however, operating amidst considerable public controversy about the safety of crop biotechnology that has been largely driven by environmental organizations such as Greenpeace and Friends of the Earth, in collaboration with the organic food industry and a broad coalition of interests challenging different aspects of multinational corporate capitalism, 19



trade liberalization, and economic globalization. In some countries, including many in Europe, the political context of implementing the ‘precautionary principle’ towards crop biotechnology may be, at least tacitly, condoned by established chemical, seed, and agricultural industry interests whose market share are challenged by the successes that agricultural biotechnology has achieved in pest control elsewhere in the world (Graff and Zilberman, 2004, Graff and Zilberman, 2004). An extremely important result of the public controversy is the persistence of uncertainty within significant segments of the consumer market about the technology’s safety and desirability, despite the regulatory review and approval procedures that are in place (Bonny, 2004, Bucchi and Neresini, 2004). A number of economic surveys and analyses of consumer demand have established that a significant range of consumers are not certain that consuming genetically modified foods is free of risk, assessed under a variety of conditions, and this uncertainty generally decreases their willingness to pay for products of the technology (Heiman, et al., 2000, Tegene, et al., 2003). Regardless of the source or the scientific accuracy of these consumer perceptions, the very existence of such consumer uncertainty affects consumer valuation, causing a negative shift in demand and downward pressure on price. As a result, a food product with the characteristic of being ‘genetically modified’ or ‘transgenic’ is generally regarded by analysts today, in economic terms, as being an inferior good relative to products from non-transgenic sources: simply put, consumers are willing to pay somewhat less for an otherwise identical product if it bears this characteristic. However, just because a product sells at a lower price does not prevent it from being adopted and successful in the marketplace, if it enjoys a sufficient cost advantage in production. Broad consumer uncertainty has another economically more worrisome effect. It makes individual retailers vulnerable to strategically targeted activism, thus and thus behaving in ways that are not to the economic benefit of any other group, including consumers and farmers. Indeed, it is the high level of sensitivity of food retailers to negative consumer perceptions and associations that effectively prevents greater adoption of transgenic food products at the retail level. Given that enough doubt persists in a wide enough range of the consumer population, the targeted campaigns of anti biotechnology activists can gain traction with retailers who are extremely concerned about the negative branding that can be caused by these campaigns; and thus retailers seek to avoid association with the technology. This strategy incidentally also creates an opening for other agricultural products, such as organics and health foods, to use a ‘positive’ antibiotech branding to differentiate their products as higher quality (and higher priced) alternatives in the marketplace. The adoption of the first generation of input traits from agricultural biotechnology was primarily driven by demand among farmers because of the cost savings they provided. Any role of consumers in driving demand for these traits has been indirect, through the affect of lower prices on foods produced from them. At least in the U.S., most consumers have not directly judged these products in terms of quality. The situation will necessarily be different as second generation traits with explicit user benefits are introduced. Consumers and others in the value chain, including food manufacturers, will instead actively choose these new products based on the characteristics they deliver, implying an increase in valuation, a positive shift in demand, and upward pressure on prices for those products. New quality characteristic are likely to have several affects on consumer perception. They may serve to counterbalance some consumers’ lingering negative associations with the application of biotechnology. They may serve to differentiate the market into segments of consumers who do

20



care and consumers who do not care about whether a product is genetically modified. And, third, they may actually shift some consumers’ fundamental assessment of biotechnology from a negative association to a positive association. Interestingly, when the FlavrSavr tomato was on the market and clearly labeled as ‘genetically modified’, the information was initially considered by consumers as a mark of quality, for which they were willing to pay a premium. Furthermore, in today’s marketplace, new positive quality characteristics are likely to be combined with lower prices driven by gains in efficiencies from the use of biotechnology. These two factors combined may interact to drive rapid adoption of these products, if targeted at the right niche of the market. As consumer experience accumulates over time, consumer uncertainty will continue to resolve itself, and negative associations with agricultural biotechnology can be expected to decline. Eventually the technology is likely to become ubiquitous and of little notice to the mass of consumers, as numerous other process technologies have become in other segments of the economy. Yet, it is reasonable to assume that this process may take years, and thus the release of new products should be planned for a transition period. Quality traits have the potential to affect public perceptions of biotechnology, but success in doing so will rely on a strong attentiveness to consumer opinions, the introduction of real desired qualities in the products, excellent communication in the sales and marketing of these products, and an irreproachable safety record.

21


3


Surveying the supply of nutritional and product quality innovations

New technological capability is an important source of new product innovation. Technology push is the economic term for the phenomenon whereby scientific and technical advance provides producers with an ever richer supply of innovation possibilities. It is the complex job of R&D professionals to channel the supply of scientific knowledge to effectively meet demand in the marketplace. In most cases technology imparts new or incrementally improved characteristics to existing products, so that they better meet the existing demands of consumers. On more rare occasions, a new bundle of characteristics is brought together by an innovation to create a radically new product that meets previously latent demands of consumers. We considered in Chapter 2 the scope of markets that utilize plant products—including food retail, food processing, livestock and dairy feed, ornamentals and landscaping, and pulp and paper—and some of the factors of demand in those markets for new qualities and characteristics. We turn now to consider the actual supply of innovations that have arisen from the plant sciences since the advent of plant biotechnology as a new technological capability in the mid 1980s. To provide an broad overview of the global R&D pipeline we surveyed three sources of information published between 1987 and 2004: (1) scientific articles from academic journals, scientific conferences, and institutional reports on agricultural biotechnology, (2) published online databases reporting field trials of genetically modified organisms in most of the highincome countries in the world, and (3) online databases reporting regulatory decisions from most countries in the world. The survey compiled a total of 358 published scientific articles, 2403 registered field trials (in 19 countries), and 36 reported regulatory decisions (in six countries) that involved an existing transgenic plant expressing traits affecting the nutritional or product quality of the agricultural output. We then organized these records, gathering together into individual groups all the research papers, field trials, and regulatory filings that appeared—as well as the data would allow us to determine—to point to an ‘individual product innovation candidate’, which was defined around three criteria: 1. a particular type of trait (e.g. high lysine or modified starch composition) 2. in a particular crop species (e.g. maize or potato) 3. by a particular organization (e.g. Monsanto) or set of organizations know to be in a collaborative relationship or cross-ownership (e.g. Amylogene and BASF). This technique of aggregating the different types of data allows us to trace the development of each innovation in the survey through its evolution through publication, field trials, and regulatory filings, as far as it goes towards commercialization. The technique allows for a roughly uniform treatment across the full scope of the R&D pipeline, given the widely varying levels of detail available from the different published data sources and at different phases of the R&D pipeline. However there are important limitations. Such treatment allows one ‘product innovation candidate’ to include more than one ‘genetic transformation event’ as commonly defined by regulators. In some cases, one ‘innovation candidate’ identified in this survey includes multiple events, although all of them confer the same category of trait in the same crop species. An 22



extreme example of this is canola with altered oil characteristics by Calgene and Monsanto. This is identified as a single product innovation candidate, ‘Monsanto’s modified oil canola’. Even though the data does indicates that the work involved over twenty different transgenes and at least as many events, detail in the published data was not sufficient for us to disentangle the separate oil traits. Utilizing this aggregation technique, our survey identifies 560 individual nutritional or product quality innovation candidates that have been developed since plant biotechnology first became feasible in the late 1980s. Of these, 383 succeeded in reaching initial field trials, 47 proceeded on to advanced field trials, 14 advanced to regulatory filings, five were commercialized, and, of these, two are still on the market. No product quality crop innovation, as of yet, has been a significant commercial success. The first to reach market, the FlavrSavr tomato, was in fact the first innovation of any type from agricultural biotechnology when it debuted in 1994, but it was removed from the market in 1997. By contrast, crop protection traits—including insect resistance and herbicide tolerance—introduced shortly after the FlavrSavr tomato, are now planted on millions of acres and account for billions of dollars in global sales.

Figure 3.1 Advance in the R&D pipeline of the 560 individual nutritional and product quality innovations observed by the current survey

560 have arisen in research or have been developed as proof-of-concept.

Of those, 383 have reached early field trials.

Of those, 47 have gone on to advanced field trials.

Of those, 14 have entered the regulatory approvals process.

5 have been commercialized.

And 2 are still on the market.

While this survey attempts to be as comprehensive as possible, there are natural limitations to how complete and accurate information can be on any industry’s R&D pipeline. The earlier one tries to looks in the pipeline the more sparse and dispersed is the available information. In many

23



cases, early work is simply not reported. Either it is conducted in house at companies under proprietary conditions or it is the result of an experiment at a university not deemed significant enough to warrant immediate publication in the literature. Toward the middle and later stages of the R&D pipeline, especially given the systems of field trial registration and regulatory approvals in place in most countries, accuracy in data reporting improves significantly. As innovations get closer to market more and better information is disclosed; however, at the same time, the absolute numbers of innovations in the pipeline declines significantly. The picture that emerges from this survey, while not absolute in a strict sense, certainly provides a thoroughly representative sampling from which results and trends can be drawn with confidence. 3.1

The dynamics of the R&D pipeline

The most important fact to be realized from the results of this survey is that the technology is still young. While only five products have reached the market, currently approximately 120 innovation candidates are actively being pursued, but the average age of these active projects is just 3.5 years (Figure 3.2). Since the earliest work on quality traits began in the late 1980s, a steady flow of new ideas has been forthcoming. Well into the late 1990s the rate of new candidates entering the pipeline grew more quickly than the rate of those being dropped. A balance was reached in 1999 between entry and exit of product innovation candidates in the pipeline, at which point over 150 R&D projects were in some stage of active development. Over the last five years, rates of entry and exit appear to have been roughly equal and steady, at about 50 per year, maintaining an average of well over 100 product candidates in the pipeline through the present. 180

160

Product quality innovation candidates

140

120

Active

100

Dormant Entry Exit

80

60

40

20

0 1987

1988

1989

1990

1991

1992

1993

1994

1995

1996

1997

1998

1999

2000

2001

2002

2003

2004

Year

Figure 3.2 The number of innovations observed to have entered, exited, and remained in the R&D pipeline annually (with truncation of the dataset in recent years)

24



It is also important to note the tapering off in observed activity since 2000. This is consistent with trends observed in agricultural biotechnology innovation overall, particularly following the dampening effect of the European regulatory gridlock after 1998. Yet, as can be seen in Figure 3.2, the data begins to be truncated in early 2003 1, thus reducing our confidence in the overall estimates for the recent years.

Table 3.1 The range of nutritional and product quality traits in the R&D pipeline Trait category Proteins and amino acids Protein quality and level Lysine Methionine Tryptophan Nutrient enhancing enzymes Other nutritional proteins Protein functional qualities Oils and fatty acids Carbohydrates Starches Fructans Sugars Micronutrients and functional plant metabolites Vitamins Minerals Functional secondary metabolites Multiple quality traits: seed composition or feed quality Reduced non-nutrients, allergens, and toxins Non-nutritional, anti-nutritional, and toxic plant metabolites Allergens Mycotoxins Extended shelf life Control of fruit ripening Control of leaf and flower wilting Bruising/browning Esthetics and convenience Flavor/scent Fruit/seed color Flower color Size Seedlessness Low maintenance landscaping Fiber and wood quality Fiber quality for textiles Fiber quality for animal feed and forage digestibility Wood quality for pulp Environmental quality Bioremediation

Number of innovations 39 19 16 3 4 10 10 54 5 81 19 18 23 20 23 7 18 7 2 64 11 8 11 9 17 3 3 2 1 31 12 10

1 Such truncation in R&D data is not uncommon, as there is a lag in publication and in the updating of databases. It is not possible to know whether there has been a real drop off in R&D in 2002 and 2003 or whether the observed dip is simply a result of information not yet being available.

25


3.2


Categories of nutritional and product quality traits observed in the R&D pipeline

The genetic traits in the R&D pipeline are observed to span ten major categories, defining for us the general scope of what are today’s potentially feasible output traits2. Each category represents characteristics which—by their very nature—will be seen and valued after the crop is harvested by a user of the agricultural output or someone affected economically by its use. Table 3.1 lists the categories and the number of innovations found within each. The next chapter (beginning on page 38) provides detailed discussion and data on all 560 of the published innovation candidates surveyed, organized into these ten categories. The first five categories—proteins, oils, carbohydrates, micronutrients, and multi-trait seed composition—focus primarily on nutrition. The latter five—allergens and toxins, shelf life, esthetics and convenience, fiber quality, and environmental quality—cover a wider range of product qualities. Even though each innovation is listed under one primary category, many of the innovations observed could be considered to fit under two or more trait categories.

3.3

Crops observed in the R&D pipeline

A wide range of crop species have been given transgenic nutritional and product quality traits. This diversity is partly due to the fact that many areas of interest in fundamental research relate to such traits. The strong representation of horticultural crops is likely due to the tendency for quality enhancements to be more attractive economically in high marginal value specialty crops. In fact, during the earliest years, from 1987 until 1990, product quality innovations were observed only in horticultural crops. During the next six or seven years throughout a phase of rapid expansion, horticultural crops continued to account for two thirds of observed R&D activities. Only by the late 1990s had increasing work in field crops, particularly oilseeds, put them on a par with horticultural crops. This also resulted from the drop in R&D on horticultural crops after 1997, the year that the FlavrSavr tomato was pulled from the market. A smaller line of work also began to emerge in forage and forestry species in the mid 1990s. By 2003, the 110 active innovations candidates were about evenly split, 50-50, between horticultural and field crops, with about 10 innovations being actively pursued in forage and forestry. Cumulative totals show that the most common crops targeted for product quality innovation have been potato (89), corn/maize (65), tomato (59), canola/rape (39), tobacco (38), and soybean (36) (Table 3.2). No other single crop is in the same league as these six, and tobacco ranks in their midst only because of its dual role as a crop and as an experimental plant, serving as a model system in which to test traits that are ultimately of greater interest in other crop species. Potato takes the lead, because of three different technologies being developed for potato: starch composition modifications (for both industrial and food uses), increased protein, and reduced browning post harvest. In addition, work on potato starch has been extensive in Europe, where R&D on crop genetics tends to be more dispersed, with more organizations doing research and field testing of very similar innovations. Several trait categories are focused upon certain crops, as would be expected, such as the concentration of work on oil traits within oil crops, including the corn, cotton, and palm. Work on 2

The scope of this list—and this study—does not include novel uses of crop agriculture for the production of regulated therapeutic compounds (plant made pharmaceuticals) and industrial non-food-grade enzymes (plant made industrial products). A recent analysis of such ‘third generation’ agricultural biotechnologies can be found in Graff, G. D. "Crop biomanufacturing, Part 1: The economic opportunity." bio-era, January 2004.

26



fiber quality is concentrated in forest products (for wood fiber) as well as forage and feed crops (for fiber digestibility). Shelf life traits are being developed in fresh market produce—fruits and vegetables—as well as fresh cut flowers, with an emerging trend in tropical fruits, for harvest and handling in warm climates and shipping to distant markets. Other major trait categories are more diversified across crops. Work on protein quality is important in grains as well as forage crops, primarily for animal nutrition, but is also active in tubers and oil seeds, particularly soy, for food nutritional quality. Work in carbohydrates is found foremost in the starchy tubers, for both food and industrial starch yield, but it is also important in grains for feed quality or for ethanol fermentation feedstock. Work in carbohydrates has also encompassed fruit sugars for flavor and ripeness characteristics important to the fresh market and for soluble solids content important to processors. Levels of micronutrients and secondary metabolites have been adjusted in a broad range of plants. This is partly because the class is broad—encompassing everything from vitamins, to enzymes regulating mineral content, to functional metabolites like isoflavones, to the reduction of compounds like caffeine and nicotine.

120

100

Corn/maize

Innovation candidates

Other grains

Row Crops

80

Oil crops Cotton

60

Tubers

40 Vegetables

Horticultural crops

Fruits

20 Coffee, tea, sugar Tobacco Herbals & medicinals Floriculture & nursery

Forage Forestry

0 1987

1988

1989

1990

1991

1992

1993

1994

1995

1996

1997

1998

1999

2000

2001

Year

Figure 3.3 Crops with active projects in nutritional and product quality traits, by year

27

2002

2003



27 9 15

42 25 7 4 1 2 1

Oil Crops Canola/Oilseed Rape Soybean Oil Palm Sunflower Brown Mustard Ethiopian Mustard Mustard Linseed Flax

3

Cotton

0

0

2

2

0

0

0

Tubers Potato Cassava Sweet potato Chicory

0

13 11 1 1

0

56 51 2 1 2

6 5 1

1 1

Vegetables Legumes Pea Legumes Azuki bean Peanut Leaf vegetables Lettuce Broccoli Onion Fruit vegetables Tomato Pepper

0

2

0

13

16

1

Fruits Temperate Fruits Strawberry Apple Melon Grape Pear Raspberry Kiwi Watermelon Tropical Fruits Papaya Pineapple Banana Mango

0

2 1

29 13 8 7 1

17 8 5 2 2

3 1 1 1

4 1 3

11 4 6

2

0

4 3

9 6

1 1

1 2

1

0

2 2

92 41 36 4 4 3 1 1 1 1

1

1

0

6

14 14

0

0

0

90 82 4 2 2

29

11

0

1

73

0

1

1 1 1

2

2 1 1 1

1 1 1 2

2 1

4 2 1

1

0

0

11

12 1

6

3

1 3

1 1

2

0

25 1

10

29

8

5 4 6 1 2 2

3 2 1

1 1 4 2 2 1

28

121 65 22 20 10 4

1

2

1

Grand Total

Environmental quality

Fiber quality

Esthetics and convenience

Extended shelf life

Non-nutrients, allergens, and toxins

4 4

1

4 4

Micronutrients and secondary metabolites

Grains Corn/Maize Rice Wheat Barley Sorghum

2

50 26 7 10 5 2

Carbohydrates

Oils and fatty acids

Proteins and amino acids

Crop

Multiple quality traits: Seed composition or feed quality

Table 3.2 Cross matrix of transgenic crops and trait categories in the R&D pipeline

1

0

1

59 2

0

46 10 8 8 4 2 2 1 1 4 3 2 1



0

16 7 1 5 3

0

8

2

38

0

0

3 3

0

16 11 1 1 1 1 1

5 2 2

19 7 2 2 1 1 1 1 1 1 1 1

0

0

25 10 4 2 2 1 1 1 1 1 1 1

1 1

0

11 8 1 2

0

20 12 2 2 2 2

0

0

12 6 2 2 1 1

4 4

16 10 2 2 1 1

0

2 2

Tobacco (mostly experimental)

0

5

3

4

13

0

3

Experimental, herbal, medicinal Arabidopsis Peppermint Evening primrose Poppy Downy thorn apple Henbane

0

1 1

3 1 1 1

0

9 6

0

Floral & nursery Petunia (some experimental) Carnation Chrysanthemum Geranium Rose Begonia Lisianthus Orchid Torenia Creeping bentgrass Kentucky bluegrass

0

0

Forage Alfalfa Bahiagrass Tall Fescue Italian ryegrass Lupin

0

Forestry Poplar Aspen Pine Eucalyptus Sweetgum

0

5 2

1

1

1 1 1 0

0

1 1

0

1

2

0

1

3 3

2

1 1

1 1

1 0

0

0

29

0

0

Total

0

Coffee, tea, sugar Coffee Tea Sugar beet Sugarcane


Environmental quality

0

Fiber quality

6 5 1

Extended shelf life

7

Non-nutrients, allergens, and toxins

0

Micronutrients and secondary metabolites

Carbohydrates

0



Crop (continued)

Multiple quality traits: seed composition or feed quality

Table 3.2 - continued


3.4


The sources of innovation: the organizations and countries doing research and development

A wide range of institutions have traditionally contributed to and conducted research and development for agriculture, including government research laboratories and universities in the publicly financed sector of the economy and seed and agrochemical companies—and now increasingly biotechnology companies—in the privately financed sector. While a fundamental division of labor is generally presumed throughout the economy between public sector R&D and private sector R&D along the lines of ‘basic’ versus ‘applied’ work, this distinction does not necessarily hold fast when it comes to agricultural research. Historically, a great proportion of applied agricultural research and development has been supported by public expenditure and conducted within public institutions, with resulting technologies and crop varieties directly disseminated to farmers through cooperative extension systems or public releases. Pardey and Bientima (2001) estimate that in 1995 annual spending worldwide on agricultural R&D was $33.2 billion, of which $21.7 billion was expended by public sector sources and $11.5 billion by private sector sources. In addition, a wide range of countries contribute to and conduct agricultural R&D. While a flow of innovation and technology is generally presumed throughout the global economy from industrialized high-income countries to middle and low income developing countries, this distinction again does not necessarily hold fast when it comes to agricultural R&D. Agriculture, in fact, may be the single most globalized industry in terms of R&D activities, with Ph.D. level scientists engaged in virtually every country on Earth and R&D spending much more broadly sourced and distributed than in other industries. Pardey and Bientima (2001) estimate that of the $33.2 billion spent in 1995 over 1/3rd ($12.1 billion) was spent by developing countries. Given the relative importance of agriculture to these economies, this is not surprising. Also, given the biodiversity richness of many low income tropical countries, many of which constitute ecological centers of origin for our common crop species, the flow of certain classes of important biological assets (both genes and germplasm) in the opposite direction is significant. This survey allows for a systematic examination of where innovation in nutritional and product quality is being done, and by whom. Previous analysis of the distribution of agricultural biotechnology patents demonstrated a split of roughly 25 percent from public sector inventors and 75 percent from private sector inventors; a subset of patents of product quality traits is somewhat more evenly distributed, with 35 percent originating in the public sector and 65% in the private sector (Graff, et al., 2003). Little has been done to systematically examine the global distribution of innovation in agricultural biotechnology. And while previous analyses have been based upon discrete observations of individual patents or field trials, the aggregation here of data into higherlevel ‘product innovation candidates’ allows this study to asses the different roles of different types of institutions at different phases along the R&D pipeline. Data on innovating organizations were derived directly from information listed in field trial and regulatory filings or from the institutional affiliations of authors of published research articles. A simple tabulation of the organizations by the number of innovations with which they are involved is shown in Table 3.3. The most active organizations in nutritional and product quality innovation include corporations such as Monsanto, DuPont-Pioneer, Syngenta, Limagrain, Suntory, Bayer, and BASF and government laboratories at the USDA (U.S.), CSIRO (Australia), INRA (France), IACR (U.K.), 30



and the Max Plank Institutes (Germany). Universities enter the picture somewhat lower on the list, led by the University of California, University of Wisconsin, Wageningen University, University of Florida, University of Georgia, Iowa State University, Purdue University, Hebrew University of Jerusalem, University of Kentucky, and University of Hawaii. Of these, six universities are current members of PIPRA, an intellectual property club of public sector research organizations 3 to facilitate the application of their patented agricultural biotechnologies in specialty crops and non-commercial uses. Only one purely ‘biotech’ company makes the top twenty list, that being Exelixis Plant Sciences (formerly Agritope) of Portland, Oregon.

Table 3.3 Top 20 innovating organizations in quality traits, by size of project portfolio R&D Organization

Number of innovations 51 42 24 22 15 14 18 11 9 9 9 9 9 9 8 7 7 7 6 6 6 6 6

Monsanto DuPont-Pioneer USDA-ARS Syngenta CSIRO INRA Limagrain IACR-Rothamsted Max Planck Institutes Suntory-Florigene University of California * Bayer University of Wisconsin * Wageningen University University of Florida * University of Georgia Iowa State University * Purdue University * Hebrew University of Jerusalem University of Kentucky * BASF University of Hawaii Exelixis

Percent of all organizations’ total project involvements

6.8 % 5.6 % 3.2 % 2.9 % 2.0 % 1.8 % 1.5 % 1.5 % 1.2 % 1.2 % 1.2 % 1.2 % 1.2 % 1.2 % 1.1 % 0.9 % 0.9 % 0.9 % 0.8 % 0.8 % 0.8 % 0.8 % 0.8 %

* PIPRA member institution

Each of the R&D organizations identified in the survey was designated as belonging to the public sector or private sector. For most, the identification was straightforward, with government laboratories, universities, and non-profit research institutes in the public sector and commercial firms (both publicly traded and privately owned) in the private sector. Separating out organizations by public or private sector and then tabulating the number of innovations with which each organization is involved, the overall share of involvement by all R&D organizations is summarized in Figure 3.4. 3

PIPRA is the Public Intellectual Property Resource for Agriculture. It is a consortium of 30 leading public sector research organizations, committed to coordinating the patenting and licensing of agricultural biotechnologies invented by them to facilitate greater use of the resulting technologies in research, specialty crops, and humanitarian applications. See www.pipra.org.

31


INRA IACR-Rothamsted 1.9% 1.5%


CSIRO 2.0%

USDA-ARS 3.2% Monsanto 6.8%

Max Planck Institutes 1.2% Wageningen University 1.2%

DuPont-Pioneer 5.6% Syngenta 3.0%

PIPRA 10.9%

University of California * 1.2%

Limagrain 3.0%

University of Wisconsin * 1.2%

Suntory-Florigene 1.2%

University of Florida * 1.1%

Bayer 1.2%

Other PIPRA members* 7.4%

BASF 0.8% Rest of private sector 18.0% Rest of public sector 37.6%

Unknown 0.9%

Figure 3.4 Share of organizations involvement in the nutritional and product quality R&D pipeline

Overall there is a greater breadth of involvement in nutritional and product quality innovation by public sector organizations than private sector. The public sector has an involvement level of 59 percent, while the private sector has a penetration of 40 percent. The predominance of the public sector is in part explained by the fact that public sector is more widely dispersed across individual labs at different universities and institutes. More separate organizations are involved in the R&D process, with extensive public-public collaboration as well as a significant amount of publicprivate collaboration. (For comparison, see Figure 3.6 below, in which it is shown that public sector organizations are solely responsible for 49 percent of the 560 innovations in the sample and collaborate on another 8 percent through public-private collaboration, summing to responsibility for 57 percent counting by individual innovations.) However, it can be seen that the large players in both the public sector and the private sector are operating within the same order of magnitude. The global distribution of innovative activity is derived from the nationality of the organizations responsible for each of the 560 individual innovations surveyed, shown in Table 3.4 and Figure 3.5. Nationality was either designated as the country in which the organization identified from field trial and regulatory filings is headquartered or the countries indicated under authors’ affiliations and contact information in published research articles. For example, Syngenta registered a series of field trials in the U.S. for corn with altered seed composition, and Switzerland is credited as the nationality of the lead organization on that one innovation. And for example, in Umemoto (2002) which published results of a transgenic rice with modified amylopectin starch, the lead author, Umemoto, is affiliated with Tohoku National Agricultural Experiment Station, and thus Japan is credited as the nationality of the lead organization on this one innovation.

32



Table 3.4 Innovations, by nationality of lead and partner R&D organizations Country United States United Kingdom Germany Netherlands Japan Australia Canada France India Switzerland Spain China Israel Italy Belgium Malaysia South Africa

Lead 293 34 38 26 24 21 20 14 13 12 5 6 6 6 5 5 5

Partner 10 12 3 7 2 3 2 7 1 0 3 1 1 0 1 0 0

Total 303 46 41 33 26 24 22 21 14 12 8 7 7 6 6 5 5

Country Denmark New Zealand Sweden Philippines Argentina Brazil Finland Taiwan Indonesia Austria Egypt Hungary Mexico Poland Ukraine Portugal Sudan

Lead 3 3 3 2 2 2 2 2 1 1 1 1 1 1 1 0 0

Partner 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 1

Total 3 3 3 3 2 2 2 2 2 1 1 1 1 1 1 1 1

A ‘lead organization’ is that which is predominantly responsible for a particular innovation, either by filing for the majority of field trial registrations or being the institution with which the lead author on a research article is affiliated. A ‘partner organization’ is one filing for a minority of the field trial permits for a given innovation or is the institution with which a non-lead author on a research article is affiliated. If only one organization is responsible for an innovation, it is by default a ‘lead’ organization. If two or more organizations from a single country collaborated on an innovation, the country was only counted once, and by default as the ‘lead’. When two or more organizations from different countries collaborate, one country is counted as the home of the ‘lead organization’ and other(s) counted as the home of ‘partner organizations’. Such international research collaborations are illustrated in Figure 3.5 by lines connecting the home countries of collaborating organizations. The dominance of the United States is the most visible result from the geographic analysis. A total of 293 out of the 560 innovations, or 52 percent of the total, are led by a U.S. organization. Europe is clearly the other major center of activity, with a total of 152 out of the 560 innovations, or 27 percent of the total led by European organizations. Of the European innovations, the great majority is from five countries—the UK, Germany, Netherlands, France, and Switzerland—but a broad innovative fringe spreads across ten other European countries. The remainder of the innovations observed globally is found amongst other OECD countries and a range of developing countries. Most prominently among the other OECD countries are Japan, Australia, and Canada—with more than 20 innovations each. Most prominent among developing countries are India, China, Malaysia, and South Africa, with a handful of innovations at seven other developing countries.

33



Figure 3.5 Global distribution of nutritional and product quality innovations and the network of cross-national collaborations, by nationality of lead and partner R&D organizations for each innovation

The global network of R&D collaboration (Figure 3.5) shows the most significant set of relationships is between the U.S. and Europe, the two most active centers of R&D globally and the two primary nodes in the global network. Within Europe, there is a significant amount of collaboration amongst European organizations across European borders. Four other OECD countries make up secondary nodes in the network of international collaboration, with Japan, Australia, Canada, and Israel each maintaining collaborative links with both U.S. and European organizations. Several developing countries constitute tertiary nodes, collaborating with researchers in one of the primary or secondary nodes: India with the U.S., South Africa with Europe, and Egypt, China, Indonesia, and the Philippines with Japan. Finally, there are a set of countries that do not exhibit any collaborations in this survey. In Latin America, Brazil, Argentina, and Mexico are each independently pursuing an innovation or two. In Southeast Asia Malaysia and the Philippines are working independently as are several countries on the European fringe.

34



This geographic analysis began with the identification of each organization and then counted the number of innovations with which each was involved. It is also possible to begin with identification of individual innovations and then asses the type of R&D organization responsible for each. Using this method each of the 560 innovations was characterized as the product of public sector R&D, private sector R&D, or public-private collaboration. Innovations were then separated out into five identifiable stages of the R&D pipeline: (1) publication, (2) initial field trials, (3) mid-stage field trials, (4) late-stage field trials, and (5) regulatory filings. Of the total 560 innovations identified in the survey, 276 (or 49 percent of the total) are solely the product of a public sector R&D organization, 232 (or 41 percent) are solely the product of R&D at a firm in the private sector, and 44 (or 8 percent) are the result of public-private collaboration.

Figure 3.6 Share of innovating organization types at different points in the R&D pipeline

35



There is, however, significant shifting in roles among the public, private, and cross-sectoral collaboration at different phases in the R&D pipeline. When innovations are at the stage of research publication, two thirds (66 percent) are by public sector organizations and an additional 17 percent are by public-private collaborations. However, at the point when field trials are initiated, roles shift significantly, with just over one third (39 percent) by public sector organizations and over half (57 percent) by firms in the private sector. The role of the public sector further decreases and the involvement by the private sector increases as innovations gets closer to commercialization. The public sector’s share drops to just 25 percent of innovations in mid-stage field trials and 12 percent of innovations in late stage field trials, while the private sector’s share grows to 69 percent in mid-stage field trials and 82 percent in late stage field trials. At the stage of regulatory filings the public sector steps out completely, except for an occasional collaborative role: the private sector accounts solely for 87 percent of regulatory filings while the remaining 13 percent are by public-private collaborations. The relative importance of publicprivate collaboration is greatest at the beginning and end of the R&D pipeline but not so in between. Public-private collaborations account for 17 percent of innovations in publications and 13 percent in regulatory filings, but less than 5 percent at any stage of field trials.

Figure 3.7 Innovations in the R&D pipeline by type of R&D sector across four country groupings: US, Europe, other OECD countries, and developing countries

36



Finally, to explore the distribution of public and private sector R&D globally, the innovations— designated as the result of public, private, or public-private R&D—are separated in Figure 3.7 according to whether the lead organization is based in the U.S., Europe, another OECD country, or a developing country. Both absolute numbers of innovations by sector and share by sector is shown for each set of countries. As already noted, we again see that the lead organization on 293 innovations is in the U.S., 152 in Europe, over 70 in other OECD countries, and about 40 in developing countries. However, the share of innovations for which the private sector is responsible differs strikingly across the four groups. In the U.S. the private sector accounts for over half of innovations in the pipeline. In Europe the private sector accounts for just over a third. In other OECD countries it accounts for less than a quarter. And in the developing world the private sector is barely perceptible, taking the lead on just one innovation. Conversely the role of the public sector, while substantial in the U.S. at 42 percent, is all the more so in Europe (47 percent) and other OECD countries (67 percent)., while it is really the only type of organization leading innovation projects in developing countries (97 percent). Public private collaboration accounts for a greater share of innovations—both in absolute and relative terms—in Europe and other OECD countries than it does in the U.S. While there is some public-private collaboration on projects for the developing world, these are invariable led by companies based in Europe or the U.S. not in the developing world. Given the trend in organizational type at different stages observed in Figure 3.6 we can also deduce from Figure 3.7 that the R&D pipeline in the U.S. with a greater share of private sector involvement is most mature and closest to commercialization. The lack of any private sector involvement in developing countries does not bode well for the commercialization of even those technologies being pursued by their public sector research organizations.

37


4


The nutritional and product quality traits found in the survey

In this chapter, we visit each of the main categories of nutritional and product quality traits of the 560 ‘individual product innovation candidates’ assembled from this survey of published secondary sources. Analysis at this level of detail in each of the categories shows both technical as well as economic trends that have been followed over the 20 years of this technology’s emergence. These data support the summary in Chapter 3 and provide context for the product commercialization as discussed in Chapter 5. This chapter is divided roughly into nutritional traits and product quality traits, following the same order of trait categories as Table 3.1. Nutritional traits begin below with the broad category of multitrait seed composition and then proceeds with macronutrients (proteins, oils, and carbohydrates), micronutrients (vitamin, minerals), functional components, and the removal of antinutrients. Product quality traits then picks up with the removal of unwanted compounds and then proceeds through fruit ripening control, product esthetics and convenience, fiber and wood qualities, and environmental quality traits.

How to read the trait tables in this chapter The trait tables in this chapter are organized by nutritional or product quality trait category (seed composition, protein level, etc.), with one table for each subsection (4.1, 4.2, 4.3, etc.). Each trait table contains seperate individual product innovation candidates withinin shaded horizonal bands (defined as a given trait, expressed in a given crop, by a given organization, firm, or research team). These are organized first by crop in aphabetical order; and within each crop, by innovating organization in alphabetical order. Within each shaded band, progress in that innovation is traced in chronological order with separate entries representing different data sources. Scientific publications are the most lightly shaded, and contain citations to the papers. Biobliographic data for each can be found in the references at the end. Field trials, aggregated by year and country, are shaded more darkly. An annual total number of field trials registered for that innovation is given for each country. Regulatory actions are shaded most boldly (blue) and listed by country, agency, and year. Finally, within each line-entry, information is included, when available, on • specific trait descriptor(s) • transgene(s) expressed • the organization(s) with which authors are affiliated (for research papers) or which filed for field trials or regulatory approvals • the country or countries indicated in authors’ contact information (for research papers), or the country in which field trials were conducted or the regulatory approval was granted.

38


4.1


Multiple quality traits - Seed composition and feed quality

Advances in corn and soy genetics have begun to combine several quality traits within single varieties. These can include improved protein, oil, and/or carbohydrate compositions, as well as improved mineral bioavailability and/or digestibility of plant fiber. These combinations are primarily intended for animal feed uses. The innovations specifically listed in this category all derive from the U.S. data on field release of genetically modified plants, which simply identify the traits being field tested as ‘seed composition’ or ‘animal feed quality improved’ (USDAAPHIS, 2004). Significant activity is indicated by the fact that active field trials continue up to the present in most of these traits. The field trials here likely correspond to several products anticipated for commercialization and described later in Chapter 5.

Table 4.1 Transgenic plants with multiple quality traits: improving ‘seed composition’ or ‘feed quality’ Crop

Trait

Country

animal feed quality improved

University of Illinois

1

Action or reference field trial

Year

Corn/Maize (Zea mays)

2004

United States

seed composition altered

Monsanto Monsanto Monsanto

1 1 2

field trial field trial field trials

1998 1999 2003

United States United States United States

seed composition altered animal feed quality improved

Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer

2 2 4 2 3 2

field trials field trials field trials field trials field trials field trials

1996 2000 2001 2002 2003 2004

United States United States United States United States United States United States


Syngenta Syngenta Syngenta Syngenta

4 4 4 9

field trials field trials field trials field trials

2001 2002 2003 2004

United States United States United States United States


Agracetus Agracetus Agracetus Agracetus Monsanto Monsanto Monsanto Monsanto Monsanto Monsanto

1 2 2 2 7 2 5 21 21 12

field trial field trials field trials field trials field trials field trials field trials field trials field trials field trials

1994 1995 1996 1997 1999 2000 2001 2002 2003 2004

United States United States United States United States United States United States United States United States United States United States


Pioneer Pioneer Pioneer Pioneer

2 2 2 1

field trials field trials field trials field trial

2000 2001 2002 2003



Pioneer

2

field trials

2001

United States

Soybean (Glycine max)

Sunflower (Helianthus annus)

Genes

Institution(s)

Count

Data source: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004

4.2


Genetic engineering for quality improvement in plant proteins has been more active than for any other component or characteristic of plants. The protein quality innovations here come from a number of recent literature reviews (Atanassov, et al., 2004, Chassy, et al., 2004, Galili, et al., 2002, Mazur, et al., 1999, Monsanto, 2003) as well as a search of the primary scientific literature, combined with data on field trials of genetically modified plants (European Commission, 2005, USDA-APHIS, 2004) and regulatory actions (AgBios, 2005, European Commission, 2005,

39



European Commission, 2005) in countries around the world. Protein innovations are divided into a number of subcategories, including general increase in protein levels, enhancement of specific essential amino acids (especially lysine, methionine, and tryptophan), the introduction of nutrient enhancing enzymes and other nutritional proteins, and finally the modification of protein functional qualities such as texture and strength.

4.2.1

Protein quality and level

The most straightforward quality improvement being pursued in plant proteins is the increase in total levels of protein present in a plant’s tissues or seeds. One approach is simply to over express a gene for an existing protein or to block a gene that controls the amount of protein left in storage. Another approach transfers genes that code for storage proteins from other plants that are protein rich—like soy or amaranth—and express them in plants that are not protein rich—like corn, potato, sweet potato, or rice. The overall level of protein can, by itself, be important when protein is a limiting nutrient or a valued ingredient, such as for animal feed, for high-protein food ingredients like soy flour, or for extraction and purification of proteins or amino acids as separate ingredients.

Table 4.2 Transgenic plants with increased protein quality and level Crop

Trait

Genes

Institution(s)

Country

lignin levels decreased and protein levels increased

storage protein from soybean

University of Florida

1


Year

Bahiagrass (Paspalum notatum)

Count

2003

United States


1

field trial

2004

United States

1

field trial

1999

Canada

Brown Mustard (Brassica nigra)

modified protein level, antibiotic resistance marker

Barley (Hordeum vulgare)

storage protein altered

glutenin from wheat

ARS ARS ARS ARS

1 2 1 1

field trial field trials field trial field trial

2000 2001 2002 2003


Canola/Oilseed rape (Brassica napus)

storage protein production

storage protein gene

Agriculture Canada; University of Guelph

1

field trial

1990

Canada

altered amino acid composition

dihydrodipicolinate synthase from Corynebacterium glutamicum

Cargill

1

field trial

1998

United States

altered amino acid composition with increased oleic acid content

homomeric (HO) cytosolic acetyl-coenzyme A carboxylase (ACCase)

Michigan State University; Monsanto

(Roesler, et al., 1997)

United States

high oil content and altered amino acid composition

sn-2 acyl-transferase from yeast

National Research Council of Canada, Plant Biotechnology Institute, Saskatoon

Cassava (Manihot esculenta)

altered amino acid composition and protein altered

storage protein, synthetic

Demegen

1

field trial

2001

United States


protein altered

glutenin from wheat

ARS

1

field trial

2002

United States


storage protein

BioTechnica

1

field trial

1991

United States


Storage protein from corn

DeKalb DeKalb

1 2

field trial field trials

1992 1996

United States United States

protein quality altered and storage protein altered

dihydrodipicolinate synthase from corynebacterium glutamicum and from corn; aspartokinase from E. coli; storage protein from corn; cystathionine synthase from corn

Du Pont

5

field trials

1993

United States

41 27 4 9 14 4 2 1 2

field trials field trials field trials field trials field trials field trials field trials field trial field trials

1994 1995 1996 1997 1998 1999 2000 2001 2002

United States United States United States United States United States United States United States United States United States

1 1

field trial field trial

2001 2002


Du Pont Du Pont Du Pont Du Pont Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer protein altered

protein kinase from tobacco

40

Iowa State University Iowa State University

(Zou, et al., 1997)

Canada



protein altered

glutenin from wheat

Iowa State University Iowa State University Iowa State University

favorable amino acid profile

alpha-Lactalbumin from pig

Iowa State University

improved essential amino acid levels/oil content

Potato (Solanum tuberosum)

Rice (Oryza sativa)



1998 2000 2003


(Yang, et al., 2002)

United States

Kansas State University

(O'Quinn, et al., 2000)

United States

(Dinkins, et al., 2001)

United States

increased sulfuric amino acids

15-kD zein storage protein from corn

University of Kentucky; Ohio State University; University of Georgia

protein levels increased and seed composition altered

gene from Arabidopsis

Monsanto

1

field trial

2003

United States


storage protein from corn; Transcriptional activator from E. coli

Rutgers University

2

field trials

2001

United States

nutritional quality altered

Storage protein, synthetic

ARS ARS ARS

1 1 2


1995 1996 1998


increased protein with higher levels of essential amino acids

albumin (Ama-1) gene from amaranth

Jawaharlal Nehru University, New Delhi

(Chakraborty, et al., 2000)

India

high protein

seed protein gene (Ama1) from amaranth

Jawaharlal Nehru University, New Delhi

(Atanassov, et al., 2004)

India

increase of sulphur-containing amino acids

10 kDa zein protein gene from corn

Beijing University

increased tuber protein content

isopentenyltransferase (ipt); z11

Beijing University

high amino acid content

15 kDa zein gene

modified amino acid content

(Hu, et al., 1997, Li, et al., 2001, Lin, et al., 2004) (Atanassov, et al., 2004)

China

n.a.

(Binswanger, 1974, Xie and Liu, 2004)

China

10 kDa sulfur-rich prolamin gene from rice

n.a.

(Xie and Liu, 2004, Yu and Ao, 1997)

China

solids increased

storage protein from wheat

Monsanto

1

field trial

1996

United States


prosystemin from tomato

Washington State University

1

field trial

2002

United States

low-protein (Tsuki-no-hikari:H39) for sake brewing

AS gluterin

Japan Tabacco

2

field trials

1994

Japan

low-protein (Akihikari) for sake brewing

AS gluterin

KIK

1

field trial

1991

Japan

lepidopteran resistance and seed composition altered

seed storage protein from pea; seed storage protein from rice

Louisiana State University Louisiana State University Louisiana State University

1

field trial

1991

United States

1

field trial

1992

United States

1

field trial

1993

United States

mutant amylopullulanase (APU) gene from a thermobacterium

National Defense University; Academia Sinica

protein quality altered

aspartokinase from E. coli; dihydrodipicolinate synthase from Corynebacterium glutamicum

DeKalb

1

field trial

1996

United States

protein quality altered and oil quality altered

dihydrodipicolinate synthase from Corynebacterium glutamicum; aspartokinase from E. coli; seed storage protein from corn

Du Pont

10

field trials

1994

United States

Du Pont Du Pont Du Pont Du Pont Du Pont Du Pont - Pioneer

10 2 12 11 2 2


1995 1996 1997 1998 1999 2000


Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer

2 1 2 2

field trials field trial field trials field trials

2001 2002 2003 2004


homoserine dehydrogenase; conglycinin; cystathionine beta-lyase; cystathionine synthase; homoserine dehydrogenase; lysine ketoglutarate reductase; 10 kDa protein from corn

(Chiang, et al., 2003)

China

high-proten, low-starch rice flour



1 1 2

Taiwan

protein altered

protein kinase from tobacco

Iowa State University

1

field trial

2002

United States


storage protein from Bertholletia excelsa

Pioneer Pioneer Pioneer

1 2 5

field trial field trials field trials

1991 1992 1993


altered amino acid composition

storage protein from corn 15 kDa zein protein from corn

University of Kentucky University of Kentucky University of Kentucky; Ohio State University; University of Georgia

1 1

increased essential amino acids

field trial 1999 field trial 2001 (Dinkins, et al., 2001)



storage protein from Bertholletia excelsa

Pioneer

1

field trial

1991

United States

Pioneer Pioneer

1 1


1992 1993


41




seed storage protein from sunflower

Van der Have

Sweet potato (Opomoea batatas)

high protein storage protein altered

artificial storage protein (ASP-1) storage protein

Tuskeegee University Tuskegee University Tuskegee University

Tobacco (Nicotiana tabacum)


storage protein, synthetic

North Carolina State University

1

field trial

1995

United States

1 1

(Prakash, 2000) field trial 1997 field trial 1999


2

field trials

United States

1995

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; Table of Developing Transgenic Crop Plants in Japan (Field Tests and General Releases), Agriculture, Forestry, and Fisheries Research Council, Japanese Ministry of Agriculture, Forestry, and Fisheries, May 2003; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; and published articles cited in the body of this table.

4.2.2

Lysine, methionine, and tryptophan – essential but deficient amino acids

The composition or proportion of amino acids—the constituent building blocks that make up proteins—available in a plant based diet is important to animal and human nutrition. Only about half of the amino acids can be synthesized by humans as well as cows, pigs, and birds. Each has a profile of specific amino acids that cannot be synthesized. It is thus essential for humans and animals to get enough of these ‘essential’ amino acids in their diet. Yet certain essential amino acids are commonly deficient in grains and oilseeds. Grains—and in particular corn, the most widely used animal feed—tend to be relatively poor in lysine. Legumes including soy tend to be poor in methionine. The result is that conventional plant based diets need to be supplemented with amino acids obtained from other protein sources, or simply overfed—both of which can be inefficient, involve additional costs, and result in excess unabsorbed nitrogen passing through the animal, making manure waste more nitrogen rich and thus more potent as a pollutant.

Table 4.3. Transgenic plants with increased levels of essential amino acids a. Lysine Crop

Trait

Genes

Institution(s)

Azuki bean (Vigna angularis)

high lysine content

feedback-insensitive dihydrodipicolinate synthase (dap) from rice

Cairo University; WeNarc; NCR; Hokko Chemical; NICS

Broccoli (Brassica oleracea)

improvement of storage proteins, synthesis of lysine


lysine level increased

high lysine

dihydrodipicolinate synthase from Corynebacterium glutamicum

feedback insensitive aspartokinase (AK) and dihydrodipicolinic acid synthase (DHDPS) from Escherichia coli and Corynebacterium

increased nutritional value and synthesis of methionine/lysine rich protein


Count

Action Year or reference (El-Shemy, et al., 2004)

Country Egypt; Japan; Sudan

University of Helsinki

1

field trial

1999

Finland

Cargill

1

field trial

1996

United States

Cargill

1

field trial

1997

United States

(Falco, et al., 1995)

United States

Pioneer - DuPont

Plant Genetic Systems

1

field trial

1992

Belgium

Plant Genetic Systems Plant Genetic Systems

1 1


1992 1993

France Belguim


ARS

1

field trial

2004

United States


DeKalb DeKalb DeKalb DeKalb DeKalb Monsanto Monsanto Monsanto Monsanto Monsanto Monsanto Monsanto

field trials field trial field trials field trial field trials field trials field trials field trials field trials field trials field trials USDA deregulation pending

1994 1995 1996 1996 1997 1999 2000 2001 2002 2003 2004 2005

United States United States United States United States United States United States United States United States United States United States United States United States

(Falco, et al., 1995) (Mazur, et al., 1999)

United States



high lysine

feedback insensitive dihydrodipicolinate synthase (DHPS) from bacteria

42

Pioneer - DuPont

3 1 2 1 2 4 3 6 36 18 22 1



protein lysine level increased

high lysine

hordothionine (HT12) and barley high lysine protein 8 (BHL*)


aspartokinase from E. coli; dihydrodipicolinate synthase from corn

lysine level increased Potato (Solanum tuberosum)

Sorghum (Sorghum bicolor)



Pioneer Pioneer Pioneer Pioneer - Du Pont Pioneer - DuPont

1 2 5 8

field trial 1996 field trials 1997 field trials 1998 field trials 1999 (Jung and Falco, 2000)

United States United States United States United States United States

University of Minnesota

1

field trial

1995

United States

University of Minnesota University of Minnesota University of Minnesota

1 1 1

field trial field trial field trial

1996 1997 1998


University of Minnesota University of Minnesota

2 1

field trials field trial

1999 2000


Wilson Genetics

1

field trial

2000

United States

high lysine

threonine synthase antisense

Max-Planck-Institut für Molekulare Pflanzenphysiologie

(Zeh, et al., 2001)

high lysine

feedback insensitive dihydrodipicolinic acid synthase (DHDPS) from bacterium

Plant Research International, Wageningen University

(Sevenier, et al., 2002)

High lysine

alpha-hordothionin HT12 protein of Hordeum vulgare

Pioneer

High methionine and lysine

beta-zein from cereals; C1-2 from cereals

Council for Scientific and Industrial Research (CSIR)

high lysine

lysine-insensitive forms of AK and DHDPS from Escherichia coli and Corynebacterium


aspartokinase from E. coli; dihydrodipicolinate synthase from Corynebacterium glutamicum

Germany

Netherlands

(Zhao, et al., 2003)

United States

(Atanassov, et al., 2004, Forsyth, et al., 2003, Grootboom and O'Kennedy, 2003)

South Africa

DuPont-Pioneer

(Falco, et al., 1995, Mazur, et al., 1999)

United States

DeKalb

1

field trial

1996

United States

DeKalb DeKalb Monsanto

1 3 2

field trial field trials field trials

1997 1998 1999


high lysine, high methionine

MB1 11 kDa synthetic protein

University of Prince Edward Island

protein lysine level increased

dihydrodipicolinate synthase from E. coli

BioTechnica

1

(Beauregard, et al., 1995)

Canada

field trial

United States

1989

b. Methionine Crop

Trait

Genes

Institution(s)

Arabidopsis

overaccumulation of free methionine

threonine synthase mutant

Hokkaido University


methionine production

methionine gene

Pioneer

5

field trials

1993

Canada


2

field trials

1992

France


1

field trial

1993

Belguim

DeKalb DeKalb DeKalb DeKalb DeKalb DeKalb Monsanto Monsanto

1 1 3 3 1 1 3 1

field trial field trial field trials field trials field trial field trial field trials field trial

1992 1993 1994 1995 1997 1998 1999 2000

United States United States United States United States United States United States United States United States

increased nutritional value and synthesis of methionine/lysine rich protein


seed methionine storage increased

seed storage protein from corn

methionine level increased

storage protein from corn

high methionine

EMBRAPA

Count

Action Year or reference (Bartlem, et al., 2000)

(Atanassov, et al., 2004, Avila, et al., 2001, Vasconcelos, et al., 2003)

Japan

Brazil

high methionine

high sulfur zein (HSZ) storage proten gene from corn

DuPont


acetolactate synthase from corn; zein storage protein

Du Pont

1

field trial

1993

United States

Pioneer Pioneer Pioneer

1 2 1

field trial field trials field trial

1994 1998 1999


Rutgers University Rutgers University Rutgers University

1 1

field trial 1998 field trial 2001 (Lai and Messing, 2002)



methionine level increased


high methionine

increased expression of delta-zein protein by enhanced mRNA stability by switching Dzr1 intron

Lupin (Lupinus angustifolius)

high methionine

2S albumin from sunflower

CSIRO


high threonine and high methionine

feedback insensitive aspartokinase (AK) from bacterium

Plant Research International, Wageningen University

43

(Knowlton, et al., 1992)

Country

(Loehman, et al., 1997, Molvig, et al., 1997)


United States

Australia

Netherlands




High methionine and lysine

beta-zein from cereals; C1-2 from cereals

Council for Scientific and Industrial Research (CSIR)



seed storage protein from Bertholletia excelsa

Pioneer Pioneer Pioneer Pioneer Pioneer Pioneer Pioneer

2 3 7 3 1 1 1

field trials field trials field trials field trials field trial field trial field trial

1992 1993 1994 1995 1996 1997 1998

United States United States United States United States United States United States United States



University of Kentucky University of Kentucky

1 1


2000 2001


high methionine

15 kDa beta-zein and 10 kDa delta-zein storage proteins from corn

New Mexico State University

(Bagga, et al., 1995, Bagga, et al., 1997)

United States

high methionine

vegetative storage protein (VSP alpha) from soy

Hebrew University of Jerusalem, Rehovot


Wheat (Triticum aestivum)


ARS

(Atanassov, et al., 2004, Forsyth, et al., 2003, Grootboom and O'Kennedy, 2003)

(Guenoune, et al., 1999)

South Africa

Israel

1

field trial

1996

United States

Count

Action or reference field trial field trial field trials field trials field trials field trials field trials field trials

Year

Country

1 1 2 5 4 9 14 4

1995 1997 1998 1999 2000 2001 2003 2004


1

field trial

1999

United States

(Galili, et al., 2002)

United States

c. Tryptophan Crop

Trait


tryptophan level increased

Genes

Institution(s) DeKalb DeKalb DeKalb Monsanto Monsanto Monsanto Monsanto Monsanto

tryptophan level increased

tryptophan level increased Soybean (Glycine max)

high tryptophan protein

University of Illinois anthranilate synthase (AS) maize variant, catalyses one of the key steps in the biosynthetic pathway for tryptophan production in plants

Monsanto; Renessen

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; GM Crop Database, AgBios, accessed August 2004; and research papers cited in the body of this table.

4.2.3

Nutrient-enhancing enzymes

Enzymatically active proteins are being engineered into feed plants to enable animals to digest grain components which they have difficulty digesting. A primary example is glucanase expressed in to barley, which enables chickens to digest 50 to 75 percent of the glucan, a main form of carbohydrate in barley endosperm, and achieve weight gain equivalent to chickens on a corn fed diet. Barley glucan is fully excreted by chickens, which greatly limits the utility of conventional barley as chicken feed (von Wettstein, et al., 2000).

Table 4.4. Transgenic plants with nutrient-enhancing enzymes Crop

Trait

Genes

Institution(s)

Country

thermostable glucanase produced

Glucanase from Bacillus amyloliquefaciens; Glucanase from Bacillus macerans

Washington State University Washington State University Washington State University Washington State University

1


Year


1996

United States

1

field trial

1999

United States

2

field trials

2000

United States

(von Wettstein, et al., 2000)

United States

Applied Phytologics (Ventria Bioscience)

1

broiler feed with transgenic malt containing heat-stable (1,3–1,4)-ß-glucanase

heat stable glucanase produced and visual marker and phosphinothricin tolerance

Glucanase from Bacillus amyloliquefaciens

44

Count

field trial

1999

United States




digestibility improved

Ventria Bioscience

2

field trials

2001

United States

Ventria Bioscience

2

field trials

2001

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004, and research papers cited in the body of this table.

4.2.4

Other nutritional proteins

Some proteins have special nutritional properties for humans. For example, some soy proteins have a cholesterol reducing effect and can be expressed at higher levels in the soy plant. Other nutritional proteins, such as human lactoferrin and lysozyme, key constituent proteins naturally occurring in mother’s milk but lacking in soy or rice based infant formulas (Lonnerdal, 2002).

Table 4.5 Transgenic plants with other nutritional proteins Crop

Trait

Genes

Institution(s)

Country

value added protein for human consumption

Lactoferrin from human

Ventria Bioscience

1


Year


2004

United States


protein altered

Albumin from pig

ARS

1

field trial

2003

United States

novel protein produced

Genes from potato, rapeseed, soybean, barley, pea, rice, rye, Arabidopsis

Pioneer

2

field trials

2000

United States

Pioneer Pioneer

2 2

field trials field trials

2001 2004


(Chong, et al., 2000)

United States


human milk proteins

lactoferrin from human

Center for Molecular Biology and Gene Therapy; Loma Linda University

Rice (Oryza sativa)

human milk proteins

mature human lactoferrin

Ibaraki University; National Institute of Agrobiological Science; Gadjah Mada University

high protein and improved amino acid composition

wild-type and methionine-modified glycinin from soy

Kyoto University; National Institute of Agrobiological Resources

human milk proteins for infant formula

lactoferrin and antitrypsin from humans

value added protein for human consumption

Lactoferrin from human; Lysozyme from human; Seed storage protein antisense from rice

University of California Davis; Ventria Biosciences Ventria Bioscience


protein altered

Casein from cow

novel protein produced


human milk proteins

lactoferrin from humans

Count

(Rachmawati, et al., 2004)

(Katsube, et al., 1999, Momma, 1999)

(Chowanadisai, et al., 2003, Lonnerdal, 2002, Nandi, et al., 2002, Rodriguez, et al., 2000, Terashima, et al., 1999) 1 field trial 2002

United States

United States

1

field trial

2003

United States


1

field trial

1998

United States

Pioneer Pioneer Pioneer Pioneer

1 1 2 2

field trial field trial field trials field trials

2000 2001 2003 2004


Université des Sciences et Technologies de Lille; Meristem Therapeutics

(Salmon, et al., 1998)

Protein functional quality

Genetic modification can be used to adjust the levels of plant proteins with particular structural properties in order to achieve functional improvements, such as changes in soy protein to help soy-based products look and feel more like dairy products (Kinney, 2003) and modifications in wheat protein, particularly gluten, to adjust dough strength and baking properties of flour

45

Japan

Ventria Bioscience

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004, and research papers cited in the table.

4.2.5

Japan; Indonesia

France



(Alvarez, et al., 2001, Barro, et al., 2003, Godin and Gingras, 2000, He, et al., 2000, Howitt, et al., 2003, Pastori, et al., 2001, Shewry, et al., 2000, Vasil, et al., 2001).

Table 4.6. Transgenic plants with protein functional qualities Crop

Trait

Genes

Institution(s)


novel seed storage protein profiles with unique functional characteristics

7S or 11S protein genes silenced

DuPont

molecular design of protein for functional properties

glycinin storage proteins

University of Kyoto


glutenin from wheat wheat germ agglutinin from wheat glutenin from wheat

ARS ARS ARS ARS ARS ARS ARS ARS

1 1 1 1 3 2 3 2


increased glutenin content

Count

Action Year or reference (Fader and Kinney, 1997)

(Utsumi, et al., 1997)

Country United States

Japan

field trial field trial field trial field trial field trials field trials field trials field trials

1995 1996 1998 2000 2001 2002 2003 2004


CSIRO

1

field trial

1998

Australia



Goertzen Seed Research Goertzen Seed Research

2 1


2001 2002


Improve baking properties, high molecular weight

1Ax1 glutenins

Centro de Estudios Fotosintéticos y Bioquímicos (CEFBIO); Institute of Arable Crops Research, Long Ashton Research Station Helios Foundation; Centro de Estudios Fotosintéticos y Bioquímicos (CEFBIO)

(Alvarez, et al., 2001, Alvarez, et al., 2000)

Rothamstead Research; CSIRO; University of Bristol; DuPont Institute of Arable Crops Research, Rothamsted

(Barro, et al., 1997, Rooke, et al., 1999) 1

field trial

2001

United Kingdom; Australia United Kingdom

Instituto de Agricultura Sostenible, Consejo Superior de Investigaciones Científicas

1

field trial

1998

Spain

high glutenin (highly elastic dough)

high molecular weight (HMW) subunits of wheat glutenin; Glutenin protein (Glu-1D-1) gene encoding subunit 1Dx5 from wheat

synthesis of high molecular weight glutenin

synthesis of high molecular weight glutenin


Argentina; United Kingdom

Argentina

increasing level of gluteninic complex in wheat endosperm

Storage protein from wheat

Metapontum Agrobios

1

field trial

2004

Italy

improved bread making characteristics

puroindoline from wheat

Montana State University

1

field trial

2003

United States

MTA Mezogazdasági Kutatóintézete, Martonvásár MTA Mezogazdasági Kutatóintézete, Martonvásár MTA Mezogazdasági Kutatóintézete, Martonvásár MTA Mezogazdasági Kutatóintézete, Martonvásár

1

field trial

2000

Hungary

1

field trial

2001

Hungary

1

field trial

2002

Hungary

1

field trial

2003

Hungary

modified gluten content

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; and published articles cited in the body of this table.

4.3


Genetic engineering of plants to improve the quality of their oils by modifying the composition of fatty acids has, just as with proteins, been a highly active area in research and development. It has been so for longer than any other area of plant metabolism engineering. The specific quality innovations identified in this survey draw upon recent reviews (Chassy, et al., 2004, Galili, et al.,

46



2002, Kinney, 2003, Mazur, et al., 1999, Monsanto, 2003) and the primary literature, again combined with data on field releases and regulatory actions, in order to ascertain where each innovation is relative to reaching to market. Virtually all genetic innovations being pursued in oil quality and nutrition seek to add or increase the proportion of favorable fatty acids—such as oleic, stearic, lauric, or omega-3 (long chain polyunsaturated) fatty acids—and to remove or decrease the proportion of unfavorable fatty acids—such as linolenic or palmitic fatty acids. More sophisticated and advanced innovations are seeking to modify the levels of multiple fatty acids simultaneously, to achieve the optimal mix for conferring favorable nutritional and physical characteristics.

Table 4.7 Transgenic plants with modified oils and fatty acids Crop

Trait

Genes

Institution(s)

Arabidopsis

omega-3 and omega-6 very long chain polyunsaturated fatty acids (PUFAs)

delta-9-specific elongating enzyme from Isochrysis galbana; delta-8 desaturase from Euglena gracilis; delta-5 desaturase from Mortierella alpina

University of Bristol; Long Ashton Research Station

Brown Mustard (Brassica nigra)

oil modification

Pioneer Pioneer

oil modification


Action Year or reference (Pray, et al., 2001)

Country

2 1


1999 2000

Canada Canada

Saskatchewan Wheat Pool Saskatchewan Wheat Pool Saskatchewan Wheat Pool

1

field trial

1998

Canada

2

field trials

2000

Canada

2

field trials

2001

Canada

alteration of oil composition

Advanta Seeds

1

field trial

2001

Netherlands

alteration of oil composition

BASF

1

field trial

2003

Sweden

erucic acid altered

Biogemma

1

field trial

2002

United States

acyl-ACP desaturase antisense from Brassica rapa; acyl-aCP thioesterase from California bay laurel acyl CoA reductase from Jojoba; acyl-ACP desaturase from safflower; acyl-ACP desaturase antisense from Brassica rapa; acyl-ACP thioesterase from American elm; acyl-ACP desaturase antisense from turnip rape ketoacyl-ACP synthase from castor bean; ketoacyl-ACP synthase antisense from Brassica rapa; oleayl-ACP thioesterase from safflower fatty acid elongase from Jojoba; acyl-ACP thioesterase from cacao ACP thioesterase from California bay laurel

Calgene

4

field trials

1991

United States

Calgene

5

field trials

1992

United States

Calgene

4

field trials

1993

United States

Calgene

8

field trials

1994

United States

Calgene

1

United States

saturated fatty acid acetyltransferase from coconut and acyl-ACP thioesterase from California bay

Calgene; Monsanto

USDA 1994 deregulation; FDA approval (Knutzon, et al., 1999, Knutzon, et al., 1995)

oil profile altered

oil profile altered

increased lauric acid

acyl CoA reductase from Cuphea hookeriana; ACP acyl ACP thioesterase from Camphor; ACP acyl ACP thioesterase from Chinese tallow increased lauric acid medium chain fatty acids

acyl-ACP thioesterase from California bay thioesterase (Ch FatB2) from Cuphea hookeriana acyl-ACP thioesterase from onion

oil profile altered

ACP thioesterase from California bay laurel

delta-9 desaturase antisense from Rhizobium; delta-12 desaturase; lysophosphatidic acid acetyl transferase from coconut; ketoacyl-ACP synthase from Cuphea hookeriana; B-ketoacyl-coenzyme delta-9 desaturase antisense from rapeseed; lysophosphatidic acid acetyl transferase from Limnanthes alba; B-ketoacyl-CoA synthase; acyl-ACP thioesterase from nutmeg; acyl-ACP thioesterase from humangosteen oil profile altered high stearate oil

lauric acid acyl-acyl carrier protein (ACP) thioesterase (Garm FatA1) gene from Garcinia mangostana

increased SDA omega-3 fatty acids and gamma-linolenic (GLA) long chain

delta-6 and delta-12 desaturases from Mortierella alpina;

47

Calgene Calgene

Calgene; Monsanto Monsanto Calgene Calgene Calgene

Calgene

Calgene

Monsanto Monsanto Calgene; Monsanto Calgene Monsanto

Count

1 5

United States

1995 1995

Canada United States

(Voelker, 1997, Voelker, et al., 1996) (Dehesh, et al., 1996) 4 field trials 1996 2 field trials 1996 1 CFIA 1996 environmental approval; Health Canada approval for feed and food use 5 field trials 1997

United States United States United States Canada Canada

6


United Kingdom

1998

United States

field trials 1998 field trial 1998 (Facciotti, et al., 1999)

Canada Australia United States

4 field trials 1999 (Froman and Ursin, 2002, James, et al., 2003 576, Ursin, 2000, Ursin, 2003)


38 1

field trials

United States


polyunsaturated fatty acids


delta-15 fatty acid desaturase from canola (Brassica napus) acyl-ACP thioesterase from Cuphea hookeriana; acyl-ACP thioesterase from humangosteen

fatty acid metabolism altered and oil composition modified ACP acyl ACP thioesterase from rapeseed; ACP acyl ACP thioesterase from soybean; delta-9 desaturase from rapeseed; delta-9 desaturase from soybean; delta-12 saturase from rapeseed; delta-12 saturase antisense from rapeseed; delta-15 desaturase from rapeseed; delta-15 desaturase antisense from rapeseed

acetyl CoA carboxylase from alfalfa acyl-ACP thioesterase from soybean; acetyl CoA carboxylase from alfalfa

alteration of oil composition and increased laurate content

Monsanto

4

field trials

2000

United States

Monsanto Monsanto Monsanto Monsanto Monsanto

31 1 4 5 5

field trials field trial field trials field trials field trials

2000 2001 2002 2003 2004

Canada United States United States United States United States

Cargill

12

field trials

1995

Canada

Cargill

3

field trials

1996

United States

Cargill Cargill Cargill Cargill Cargill Cargill

13 1 16 2 32 2

field trials field trial field trials field trials field trials field trials

1996 1997 1997 1997 1998 1999

Canada United States Canada United States Canada United States

Cargill Cargill Cargill Cargill Cargill

15 2 10 2 7

field trials field trials field trials field trials field trials

1999 2000 2000 2001 2001

Canada United States Canada United States Canada

CPB Twyford Ltd

2

field trials

1997

United Kingdom

CPB Twyford Ltd

1

field trial

1998

United Kingdom

increased gamma-linolenic acid

delta-6 and delta-12 desaturases

CSIRO

alteration of oil composition and increased erucic acid and laurate content

ClFatB3 from Cuphea lanceolata; Oleate desaturase antisense from rapeseed; KAS from rapeseed; LdLPAAT from Limnanthes douglasii; BnLPAAT antisense from rapeseed

Deutsche Saatveredelung (DSV); Lippstadt Bremen

2

field trials

1997

Germany

Dow

9

field trials

2003

Canada

Du Pont

1

field trial

1993

United States

Du Pont

3

field trials

1994

United States

Pioneer

1

1996

Canada

Pioneer

2

Health Canada approval for food use field trials

1999

Canada

Federal Centre for Breeding Research on Cultivated Plants Federal Centre for Breeding Research on Cultivated Plants Federal Centre for Breeding Research on Cultivated Plants

1

field trial

1996

Germany

1

field trial

2002

Germany

1

field trial

2003

Germany

InterMountain Canola

2

field trials

1995

United States

John K King & Sons

2

field trials

1997

United Kingdom

modified oil composition and selectable marker fatty acid metabolism altered

high oleic/low linolenic acid

acyl-ACP thioesterase from soybean; delta-15 desaturase antisense from rapeseed thiolase from soybean; delta-9 desaturase from rapeseed; delta-9 desaturase antisense from rapeseed; delta-12 desaturase from rapeseed; delta-12 desaturase antisense from rapeseed; delta-15 desaturase from rapeseed; delta-15 desaturase antisense from rapeseed fatty acid desaturase (fad2) mutation

modified oil lipid metabolism, increased laurate content

fatty acid metabolism altered

acyl-ACP thioesterase (ClFatB4) from Cuphea lanceolata

delta-9 desaturase from rapeseed; delta-9 desaturase from soybean; delta-12 desaturase from rapeseed; delta-12 desaturase antisense from rapeseed; delta-15 desaturase from rapeseed; delta-15 desaturase antisense from rapeseed

alteration of oil composition and increased laurate content altered amino acid composition with increased oleic acid content

homomeric (HO) cytosolic acetyl-coenzyme A carboxylase (ACCase)

(Liu, et al., 2002)

Michigan State University; Monsanto

(Roesler, et al., 1997)

Australia

United States

oil profile altered

Michigan State University

1

field trial

1998

United States

alteration of oil composition and increased erucic acid content

Nickerson Biocem

1

field trial

1996

United Kingdom

3

field trials

1997

Germany

alteration of oil composition and increased erucic acid content

Thioesterase from Umbellularia californica; oleate desaturase (fad2) antisense from rapeseed; KAS from rapeseed; BnLPAAT antisense from rapeseed; acyl-ACP thioesterase (ClFatB3 and ClFatB4) from Cuphea lanceolata; lyso-phosphatidate acyltransferase (LPAAT) from Limnanthes douglasii; lysophosphatidic acid acyltransferase (plsC) from E. coli

Norddeutsche Pflanzenzucht (NPZ), Hans Georg Lembke

high oil content and altered amino acid composition

sn-2 acyl-transferase from yeast

Plant Biotechnology Institute (NRC)

48

(Zou, et al., 1997)

Canada



oil modification

Plant Biotechnology Institute (NRC)

2

field trials

1998

Canada

alteration of oil composition, increased laurate content, and increased stearate content

Plant Breeding International

2

field trials

1995

United Kingdom

Planta Angewandte Pflanzengenetik und Biotechnologie

2

field trials

1997

Germany

Plantech Research Institute

2

field trials

1998

Canada

Plant Science Sweden

1

field trials

2004

Sweden

Saskatchewan Wheat Pool Saskatchewan Wheat Pool Saskatchewan Wheat Pool Saskatchewan Wheat Pool

1

field trial

1998

Canada

3

field trials

1999

Canada

2

field trials

2000

Canada

2

field trials

2001

Canada

Scottish Agricultural College Scottish Agricultural College Scottish Agricultural College Scottish Agricultural College

1

field trial

1992

United Kingdom

1

field trial

1994

United Kingdom

3

field trials

1995

United Kingdom

1

field trial

1997

United Kingdom

increased lauric acid content in oil

Seedex Seedex; Monsanto

1 1


1996 1997

Australia Australia

modified oil composition

University of Manitoba

3

field trials

2003

Canada


Agriculture & Agri-Food Canada Agriculture & Agri-Food Canada Agriculture & Agri-Food Canada

2

field trials

1999

Canada

1

field trial

2000

Canada

1

field trial

2001

Canada

Du Pont Du Pont Du Pont Du Pont - Pioneer

2 1 2 5

field trials field trial field trials field trials

1994 1996 1998 1999


3 3 13 16 12 10


1999 2000 2001 2002 2003 2004



1

field trial

1996

United States

University of Minnesota University of Minnesota University of Minnesota University of Minnesota

1 1 2 1

field trial field trial field trials field trial

1997 1998 1999 2000


alteration of oil composition and increased laurate content

oleate desaturase (fad2) antisense from rapeseed; KAS from rapeseed; BnLPAAT antisense from rapeseed; acyl-ACP thioesterase (ClFatB3 and ClFatB4) from Cuphea lanceolata; lyso-phosphatidate acyltransferase (LPAAT) from Limnanthes douglasii; lysophosphatidic acid acyltransferase (plsC) from E. coli

oil modification and antibiotic resistance

oil modifications

lipid metabolic enzyme 1 from Arabidopsis, ubiquinol-cytochrome C chaperone family protein from Arabidopsis, lipid metabolic enzyme 2 from Saccharomyces cerevisiae, primary carbon metabolism enzyme from Saccharomyces cerevisiae, protein kinase from Physcomitrella patens, myb-like DNA-binding protein from Physcomitrella patens, lipid metabolic enzyme 3 from Arabidopsis DNA binding protein from Arabidopsis


modified oil composition, increased laurate content, and increased stearate content


oil quality altered

delta-12 desaturase antisense from corn

oil profile altered

oil profile altered

Cotton (Gossypium hirsutum)

high oleic and high stearic oils

Monsanto Monsanto Monsanto Monsanto Monsanto Monsanto acetyl CoA carboxylase from corn acetyl CoA carboxylase antisense from corn

identification, characterization, and silencing of delta-12 and delta-9 desaturases

modified fatty acid content in cottonseed oil

CSIRO CSIRO

(Liu, et al., 2000, Liu, et al., 2002) 1

field trial

2003

Australia Australia

high oleic acid

mutant delta-12 desaturase

University of North Texas

(Chapman, et al., 2001)

United States

Evening Primrose (Oenothera sp.)

high yield of gamma-linolenic polyunsaturated fatty acid (PUFA) in evening primrose

delta-6 desaturase from borage

Rothamsted Research

(de Gyves, et al., 2004)

United Kingdom

Ethiopian Mustard (Brassica carinata)


Flax (Linum usitatissimum)

oil modification

49

Saskatchewan Wheat Pool Saskatchewan Wheat Pool

1

field trial

1998

Canada

1

field trial

2000

Canada

University of Saskatchewan University of Saskatchewan

2

field trials

2000

Canada

4

field trials

2001

Canada


Linseed (Linum usitatissimum)



fatty acyl-desaturases and elongases

University of Hamburg; BASF; Rothamsted Research; University of Kansas; Forschungszentrum Borstel

(Abbadi, et al., 2004)

Germany; United Kingdom; United States

altered oil composition

delta-6-elongase from Physcomitrella patens delta-6-desaturase from Phaeodactylum tricornutum delta-5-desaturase from Phaeodactylum tricornutum


1

field trial

2004

Sweden

n.a.

1

field trial

2001

United States

Peppermint (Mentha piperita)

oil profile altered

Oil Palm (Elaeis oleifera and Elaeis guineensis)

high oleic palm oil

palmitoyl ACP thioesterase gene antisense from palm; KAS II gene from palm

Malaysian Palm Oil Board (MPOB)

(Parveez, 2003, Parveez, et al., 2004)

Malaysia

high stearic acid palm oil

stearoyl-ACP desaturase

Malaysian Palm Oil Board (MPOB)

(Parveez, 2003)

Malaysia

high oleic palm oil

delta-9-Stearoyl-ACP-Desaturase antisense; Palmitoyl-ACP Thioesterase (PATE) antisense from palm

National University of Malaysia

(Shah, et al., 2004, Shah and Cha, 2000)

Malaysia

low palmitic-high oleic oil

acyl-acyl carrier protein (ACP) thioesterase and beta-keto acyl ACP synthase

Palm Oil Research Institute

(Jalani, et al., 1997)

Malaysia

Rape (Brassica juncea)

decreased level of total saturated fatty acids

ADS1 gene encoding a fatty acid delta-9 desaturase from Arabidopsis

Saskatchewan Wheat Pool


reduced linolenic acid oil content

fan1 gene mutation crossed in from soybean cultivar

Agriculture & Agri-Food Canada

1

Health Canada approval for food use

2001

Canada

oil profile altered

Du Pont

3

field trials

1993

United States

Du Pont

7

field trials

1994

United States

Du Pont DuPont

19

field trials

high oleic oil

delta-9 desaturase from soybean; delta-15 desaturase antisense from soybean omega 6 desaturase omega 6 desaturase antisense from soybean delta-15 desaturase from soybean delta-12 desaturase (FAD2) gene downregulated

1995 (Kinney, 1996)


acyl-ACP thioesterase from soybean omega 3 desaturase from soybean delta-12 desaturase from soybean

Du Pont

3

field trials

1996

United States

high oleic oil

Du Pont

1

1996

United States

Du Pont

11

1997

United States

high oleic oil

omega 3 desaturase antisense from soybean acyl-ACP thioesterase from rapeseed delta-12 desaturase from soybean

FDA approval for food use field trials

Du Pont

1

1997

United States

7 1 1

1998 1999 1999

United States United States Japan

2000 2000

United States Japan

2000

Canada

2000

Australia

2001 2001

United States Japan

2002 2003 2004


1998 2000 (Dehesh, 2002) field trials 2003 field trials 2004


(Reddy and Thomas, 1996)

United States

(Anton and Yao, 1994)

high oleic oil

delta-12 desaturase from soybean

Du Pont Du Pont Du Pont

high oleic oil


Du Pont - Pioneer Du Pont

2 1

high oleic oil


Du Pont

1

high oleic oil


Du Pont

1

high oleic oil


Du Pont - Pioneer Du Pont

2 1

Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer

1 2 2

USDA deregulation field trials field trial MAFF approval for environment field trials MAFF approval for feed CIFA approval for environment, feed, and food ANZFA approval for food use field trials MHLW approval for food use field trial field trials field trials

Calgene Monsanto Monsanto Monsanto Monsanto

3 1


seed composition altered fatty acid metabolism and oil profile altered decreased saturated fatty acid levels

thioesterase beta-ketoacyl-ACP synthase (KAS)

6 8

Canada

gamma-linolenic acid oils

delta-6 desaturase from cyanobacteria

Texas A&M University

fatty acid metabolism and oil profile altered

stearoyl ACP desaturase from Rattus norvegicus; lysophosphatidate acyltransferase from yeast

University of Kentucky

4

field trials

2003

United States


3

field trials

2004

United States

DuPont; University of Nebraska University of Nebraska

(Buhr, et al., 2002, Kinney and Knowlton, 1998) field trial 2000

United States

1

University of Nebraska University of Nebraska University of Nebraska University of Nebraska University of Nebraska University of Nebraska

3 3 4 2 2 1

lower saturated fat

delta-12 desaturase (FAD2) gene downregulated

fatty acid level altered and oleic acid content altered in seed

delta-12 saturase from soybean; delta-12 desaturase antisense from soybean; palmitoyl thioesterase antisense from soybean

delta-6 desaturase from borage

low linolenic acid oil

reduce expression of omega-3 fatty-acid desaturase (fan) gene from soy

50

USDA-ARS; University of Missouri

field trials field trials field trials field trials field trials field trial

United States

2001 2002 2003 2004 2003 2004


(Bilyeu, et al., 2003)

United States




alteration of oil composition, increased stearate content

Rustica Prograin Génétique


production of unsaturated fatty acid: gamma-linolenic acid (GLA)

delta6 fatty acid desaturase from cyanobacteria

Texas A&M University

production of unsaturated fatty acid: gamma-linolenic acid (GLA)

delta6 fatty acid desaturase from borage

Long Ashton Research Station; University of Bristol


fatty acyl-desaturases and elongases

University of Hamburg; BASF; Rothamsted Research; University of Kansas; Forschungszentrum Borstel

1

field trial

1997

(Reddy and Thomas, 1996)

(Sayanova, et al., 1997)

(Abbadi, et al., 2004)

France

United States

United Kingdom

Germany; United Kingdom; United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; GM Crop Database, AgBios, accessed January 2005; and published articles cited in the body of this table.

4.4

Carbohydrates

Quality innovation in plant carbohydrates includes improvement in the content of simple sugars like glucose, fructose, and sucrose as well as the complex carbohydrates such as fructans and starches. The carbohydrate innovations identified in this study draw upon the some of the same general reviews of plant biotechnology and nutrition (Chassy, et al., 2004, Galili, et al., 2002, Mazur, et al., 1999, Monsanto, 2003), as well as data on field releases and regulatory actions. Carbohydrate innovations for several crops can only be identified as general ‘carbohydrate composition’ modifications because more detailed information was not available from the reporting sources.

Table 4.8 Transgenic plants with general carbohydrate modifications Crop

Trait

Country

alteration of carbohydrate composition and alteration of oil composition

Federal Centre for Breeding Research on Cultivated Plants

1


Year


2002

Germany


carbohydrate metabolism altered

Aventis Bayer

2 1


2002 2003



Genes

Institution(s)

Count


Sucrose phosphate synthase from spinach

Texas Tech University Texas Tech University Texas Tech University Texas Tech University

1 1 1 4

field trial field trial field trial field trials

1998 1999 2000 2001



Galactanase from soybean; UDP glucose glucosyltransferase from soybean

Du Pont

2

field trials

1998

United States

Produces plant-derived gums (galactomannins) useful in making food products like ice cream

mannan synthase (a cellusose synthase) from guar plant

Pioneer-Du Pont; Indian Institute of Technology

(Dhugga, et al., 2004)

United States; India

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; and published articles cited in the body of this table.

4.4.1

Starches

Modifications of starch level and composition have been extensively pursued in potato and other tubers but also in corn and the cereal grains. Genetic engineering is being used to alter the levels

51



of expression of naturally occurring starches, and is beginning to be used in the more complex tasks of modifying the structure and branching of starch polymers. The two natural forms of starch—amylose and amylopectin—have quite different properties and thus different food and industrial applications. High amylose starches are useful in confectionary foods, as well as in adhesives and paper products. High amylopectin starches (or waxy starches) are useful as thickeners, stabilizers, and emulsifiers in foods, and are an important component in paper finishing. Genetic engineering has created crops specialized for the production of either amylase or amylopectin. More advanced work involves modification in the branching patterns of amylopectin, the incorporation of phosphate groups, and the larger scale organization, size, and structure of starch granules, all of which are responsible for various starch qualities (Jobling, 2004).

Table 4.9 Transgenic plants with modified starch Crop

Trait

Genes

Institution(s)


high-amylose starch

starch synthase IIa gene (SSIIa) from barley sex6 mutants

CSIRO; INRA


starch composition


Count

Research Centre for Biotechnology, Indonesian Institute of Sciences (RCB-IIS)

Action Year or reference (Howitt, et al., 2003)

Country


Indonesia

Australia; France

starch metabolism altered

Starch synthase from cassava

University of Virgin Islands

1

field trial

2002

United States


ADP glucose pyrophosphorylase from corn

Abbott and Cobb

2

field trials

2002

United States


Starch synthase from wheat

AgReliant Genetics AgReliant Genetics

2 1


2001 2002



AgrEvo AgrEvo Aventis Aventis Bayer

7 1 1 3 2

field trials field trial field trial field trials field trials

1998 1999 2000 2001 2002


starch level increased

BASF BASF BASF BASF

1 4 6 1

field trial field trials field trials field trial

2001 2002 2003 2004


Biocem; Limagrain Limagrain Biogemma; Limagrain Limagrain Biogemma Biogemma

2 1 2 1 1 8

field trials field trial field trials field trial field trial field trials

1996 1996 1999 2000 2000 2002

France United States France United States France United States

Biogemma

2

field trials

2003

United States

Dow

1

field trial

1999

United States

Du Pont Du Pont Du Pont

5 17 25

field trials field trials field trials

1993 1994 1995


Du Pont Pioneer Du Pont - Pioneer Du Pont - Pioneer Du Pont - Pioneer

5 1 7 14 2

field trials field trial field trials field trials field trials

1996 1996 1997 1998 1999


ExSeed Genetics

2

field trials

2000

United States

ExSeed Genetics

10

field trials

2001

United States


1 1


1998 1999



1 3


2000 2001



1 2


2002 2003


2 3 2 1 3 4 12 9 7

field trials field trials field trials field trial field trials field trials field trials field trials field trials

1993 1994 1995 1995 1996 1997 1998 1999 2000

United States United States United States United States United States United States United States United States United States

starch metabolism altered Starch synthase antisense from wheat Starch synthase from wheat ADP glucose pyrophosphorylase from E. coli; Branching enzyme (TB1) from wheat

starch level increased carbohydrate metabolism altered Levansucrase from Bacillus amyloliquefaciens; Fructosyl transferase from Strep. mutans Starch branching enzyme II antisense from corn carbohydrate metabolism altered Starch branching enzyme II from corn Starch branching enzyme II from corn

starch level increased and imidazolinone tolerance


Amylase from corn Isoamylase-type starch debranching enzyme from corn Starch branching enzyme II antisense from corn; Starch branching enzyme II antisense from barley Starch debranching enzyme from corn Starch synthase from corn


Monsanto Monsanto Monsanto DeKalb Monsanto- DeKalb Monsanto - Dekalb Monsanto - DeKalb Monsanto Monsanto

52


Pea (Pisum sativum)



Monsanto Monsanto

7 5


2001 2002



Starch synthase from corn

National Starch & Chemical

1

field trial

2003

United States


Starch synthase from corn

University of Florida University of Florida

3 1


2002 2004


starch reduced

ADP glucose pyrophosphorylase from pea; ADP glucose pyrophosphorylase antisense from pea

DNA Plant Tech

1

field trial

1993

United States

alteration of starch biosynthesis

alpha-amylase

Institute of Plant Genetics and Crop Plant Research

1

field trial

1999

Germany

alteration of carbohydrate composition

starch biosynthesis; downregulation of invertase

Advanced Technologies (Cambridge) Advanced Technologies (Cambridge) Advanced Technologies (Cambridge) Advanced Technologies (Cambridge) Advanced Technologies (Cambridge)

1

field trial

1996

United Kingdom

1

field trial

2000

United Kingdom

1

field trial

2001

United Kingdom

1

field trial

2002

United Kingdom

1

field trial

2003

United Kingdom

AgrEvo

1

field trial

1996

Spain

AgrEvo AgrEvo AgrEvo

1 1 2


1996 1996 1997

United Kingdom France Germany

AgrEvo AgrEvo

2 4


1998 1999

Spain Germany

AgrEvo AgrEvo

2 3


1999 2000

Spain Germany

AgrEvo

1

field trial

2000

Spain

Aventis

1

field trial

1999

Germany

Aventis

1

field trial

2001

Germany

alteration of starch biosynthesis for improved starch quality

downregulation of amylose synthesis downregulation of granule bound starch synthase

GBSSI from potato; GBSSI antisense from potato, GBSSII from potato; GBSSII antisense from potato, SSSI antisense from potato; SSSIII antisense from potato; BE from potato; BE antisense from potato R1 from potato; R1 antisense from potato; SSIII from potato; SSIII antisense from potato; DE antisense from potato; Polyphosphatkinase (PPK) from E. coli; Citrate synthase from Saccharomyces cerevisiae; Starch phosphorylase antisense from potato; GDP-mannose-pyrophosphorylase (alpha-MPPY) antisense from potato; saccharosetransporter (SoSUT) from spinach R1 from potato; R1 antisense from potato; BEI antisense from potato; SSI from potato; SSI antisense from potato; SSIII from potato; SSIII antisense from potato; GBSSI from potato; GBSSI antisense from potato; GBSSII antisense from potato; BE from Neisseria denitrificans carbohydrate metabolism

GBSS antisense from potato

amylose-type starch

branching enzyme (BE) antisense in potato

Amylogene


downregulation of granule bound starch synthase synthesis of ADP glucose pyrophosphorylase downregulation of amylose synthesis synthesis of branching enzymes; synthesis of glucogene

Amylogene Amylogene Amylogene Amylogene

4 3 2 3

field trials field trials field trials field trials

1995 1996 1997 1998

Sweden Sweden Sweden Sweden

altered starch (no amylose)

Amylogene

1

1998

European Union

altered starch (no amylose)

Amylogene Amylogene Amylogene

3 3 1

1999 2000 2002

Sweden Sweden European Union

BASF

2

Application under Directive 90/220/EEC field trials field trials Opinion by EU Scientific Committee on Plants (SCP) field trials

2003

Germany

GBSS antisense BE1 and BE2 sense and antisense GBSS antisense BE1 and BE2 sense and antisense

BASF

2

field trials

2004

Germany

BASF

4

field trials

2004

Netherlands

granule bound starch synthase (GBSS) antisense; amylopectin; downregulation of amylose synthesis

Avebe

1

field trial

1995

Netherlands

Avebe

1

1997

European Union

Avebe

1

Application under Directive 90/220/EEC (later withdrawn) field trial

2001

Netherlands

Avebe

1

field trial

2003

Netherlands

Axis Genetics

2

field trials

1994

United Kingdom

alteration of starch biosynthesis and synthesis of starch consisting of pure amylopectin reduced amylose reduced amylose

alteration of starch biosynthesis altered starch (no amylose)

modified carbohydrate composition and improved processing quality

53

(Lilius, et al., 2000)

Sweden




Bioplant

1

field trial

1997

Germany

alteration of starch biosynthesis and improvement of starch quality

Boreal Plant Breeding

1

field trial

2004

Finland

carbohydrate level increased

Calgene

1

field trial

1990

United States

CPRO-DLO

1

field trial

1994

Netherlands

Danisco

1

field trial

1994

Denmark

Danisco Danisco Danisco Danisco

1 1 1 1

field trial field trial field trial field trial

1995 1996 1998 1999

Denmark Denmark Denmark Denmark

Frito Lay Frito Lay Frito Lay Frito Lay Frito Lay

1 5 4 4 4

field trial field trials field trials field trials field trials

1993 1994 1995 1996 1997


Gansu Agricultural University

(Hong, 2000, Xie and Liu, 2004)

China

Germicopa

2

field trials

1996

France


GBSS antisense from potato; pat from Streptomyces viridochromogenes

granule bound starch synthase gene (GBSS) antisense

alteration of starch biosynthesis and increased storage

carbohydrate metabolism altered Genes from spinach; Genes from wheat Genes from potato; Genes from yeast

improved cold sweetening

acetic invertase (AcInv) antisense from potato

alteration of starch biosynthesis, improved starch quality carbohydrate metabolism

GBSSI antisense from potato

GSF-National Research Center for Environment and Health

1

field trial

1999

Germany

alteration of starch metabolism, carbohydrate composition

granule bound starch synthase gene (GBSS) antisense

Hettema Zonen Kweekbedrijf Hettema Zonen Kweekbedrijf

2

field trials

1992

Netherlands

1

field trial

1993

Netherlands

carbohydrate metabolism

FBPase; inorganic pyrophosphatase

Institute of Plant Genetics and Crop Plant Research

1

field trial

1998

Germany


downregulation of amylose synthesis; downregulation of granule bound starch synthase (GBSS); synthesis of starch consisting of pure amylopectin

Institut für Genbiologische Forschung, Berlin

1

field trial

1992

Germany

reduced phosphate content in starch

alpha-glucan water dikinase (GWD) antisense from potato

Institut für Genbiologische Forschung Berlin; MaxPlanck-Institut für molekulare Pflanzenphysiologie

(Lorberth, et al., 1998)

Germany

modified amylopectin starch properties

starch synthases (SSII and SSIII) antisense from potato

John Innes Center; Unilever; Wageningen University

(Blumenthal and Edwards, 2000, Marshall, et al., 1996)

United Kingdom; Netherlands

alteration of carbohydrate composition and downregulation of granule bound starch synthase

granule bound starch synthase (GBSS) antisense

KARNA Vereniging

1

field trial

1991

Netherlands

KARNA Vereniging

1

field trial

1992

Netherlands

KARNA Vereniging

1

field trial

1993

Netherlands

Max Planck Institute for Plant Breeding Research, Köln Max Planck Institute for Plant Breeding Research, Köln

1

field trial

1997

Germany

1

field trial

2000

Germany

R1 antisense from potato; BE antisense from potato; GBSSI antisense from potato; GBSSII antisense from potato; RE antisense from potato; RE from Spinacia oleracea; SSSI antisense from potato; SSSIII antisense from potato; STPIIa antisense from potato; Polyphosphatekinase from E. coli; hexose phosphate translocator from E. coli; triose phosphate translocator from potato; saccharose transporter from Spinacia oleracea; phosphate transporter from yeast GBSSII; SSIII; DE; BE; GBSSI; starch binding protein R1; Polyphosphatkinase (PPK) from E. coli; citrate synthase from Saccharomyces cerevisiae; starch phosphorylase antisense from potato; GDP-Mannose-pyrophosphorylase (alpha-MPPY) antisense from potato; saccharose transporter (SoSUT) from spinach; Saccharose-Saccharose-1-Fructosyltransferase (1SST) from Cynara scolymus; Fructan-Fructan-1-Fructosyltransferase (1FFT) from Cynara scolymus

Max Planck Institute of Molecular Plant Physiology, Golm

2

field trials

1996

Germany


3

field trials

1998

Germany

STP IIa and IIb antisense from potato

Max Planck Institute of Molecular Plant Physiology, Golm Max-Planck-Institut für Molekulare

2

field trials

1999

Germany

(Regierer, et al., 2002)

Germany


alteration of carbohydrate composition, starch biosynthesis, improvement of cooking/frying characteristics

high starch level and high yield

GBSS sense and antisense from potato

plastidial adenylate kinase

54



Pflanzenphysiologie starch content of the tubers

Leghemoglobin gene from Lotus


1

field trial

2003

Germany

starch composition altered

glgC16

Meijer Seedpotatoes & Research, Reimerswaal

1

field trial

1998

Netherlands

increased starch

ADP glucose pyrophosphorylase (ADPGPP) from E. coli

Monsanto; Michigan State University Monsanto

2

field trials

1996

Canada

Monsanto

8

field trials

1997

United States

Monsanto

27

field trials

1998

United States

Monsanto

7

field trials

1999

United States

Monsanto

2

field trials

1999

Canada

Michigan State University

1

field trial

2001

United States


Nickerson Biocem

1

field trial

1995

United Kingdom


North Dakota State University North Dakota State University North Dakota State University North Dakota State University North Dakota State University North Dakota State University North Dakota State University

1

field trial

1993

United States

1

field trial

1994

United States

1

field trial

1995

United States

1

field trial

1997

United States

2

field trials

1999

United States

1

field trial

2002

United States

1

field trial

2003

United States

Plant Breeding International

1

field trial

1996

United Kingdom


1

field trial

2003

Sweden


4

field trial

2004

Sweden

Planta Angewandte Pflanzengenetik und Biotechnologie

1

field trial

1997

Germany

PlantTec Biotechnologie

1

field trial

2001

Germany

Proefstation voor de Akkerbouw en de Groenteteelt in de Vollegrond (PAGV) Proefstation voor de Akkerbouw en de Groenteteelt in de Vollegrond (PAGV)

1

field trial

1995

Netherlands

1

field trial

1996

Netherlands



ADP glucose pyrophosphorylase from E. coli



SGH1 and SGH2 sense and antisense SBE1and SBE2 antisense

increased amylopectin content increased amylose content and quality alteration of phosphate metabolism, alteration of starch biosynthesis, downregulation of granule bound starch synthase, and tolerance to glufosinate

GBSSI antisense from potato

alteration of carbohydrate composition modified starch

alteration of starch biosynthesis; granule bound starch synthase (GBSS) antisense; GBSS promoter; amylopectin

(Adams, et al., 2000)

United States

modified starch

alteration of starch biosynthesis; downregulation of granule bound starch synthase

SaKa-Ragis Pflanzenzucht

1

field trial

1995

Italy

alteration of carbohydrate composition, improved nutritional value and processing quality

S-adenosylmethionin decarboxylase (SAMDC)

Scottish Crop Research Institute

1

field trial

1995

United Kingdom

Scottish Crop Research Institute Scottish Crop Research Institute

2

field trials

1996

United Kingdom

1

field trial

2000

United Kingdom

Solavista

1

field trial

2002

Germany

Solavista

1

field trial

2003

Germany


Syngenta

1

field trial

2004

United States

alteration of starch biosynthesis, secretion of alpha-amylase, and synthesis of glucose isomerase

Tillämpad Biokemi

1

field trial

1996

Sweden

alteration of carbohydrate composition and tolerance to glufosinate

high amylose starch

inhibition of starch branching enzymes A and B (SBEI and SBEII)

Unilever; National Starch and Chemical

(Jobling, et al., 2003, Jobling, et al., 1999, Schwall, et al., 2000)

United Kingdom; United States

freeze-thaw-stable 'waxy' starch

downregulated amylopectin starch synthases (SSII and SSIII) and granule bound starch synthase (GBSS) antisense

Unilever; National Starch and Chemical Company; ProteoSys

(Jobling, et al., 2002)

United Kingdom; United States; Germany


Universidad Pública de Navarra, Dpto. Producción Agrari

1

field trial

1999

Spain

1994 1995



Sucrose phosphate synthase from corn

University of Wisconsin University of Wisconsin

1 1


amylose-free 'waxy' starch

granule-bound starch synthase (GBSS) antisense from potato granule bound starch synthase (GBSS) antisense; branching enzyme (BE) antisense

Wageningen University; University of Groningen Wageningen University

1

field trial


55

(Visser, et al., 1991) 1992

Netherlands Netherlands


downregulation of endoglucanase; synthesis of branching enzymes (BE) pVU1011 (derivative of pROK1); pBI121 (derivative of pBIN19)

Wageningen University

1

field trial

1994

Netherlands


1

field trial

1996

Netherlands

inhibition of NAD-malic enzyme

Zeneca Zeneca

1 1


1998 1999

United Kingdom United Kingdom

Zeneca Mogen Syngenta Syngenta

1 1 1


1999 2001 2002

Netherlands United States United States

Zuckerforschung Tulln

2

field trials

1996

Austria

Aventis

2

field trials

2000

United States

Aventis Aventis

2 3


2001 2002



BASF

2

field trials

2002

United States

hygromycin tolerance and starch level increased

ExSeed Genetics

1

field trial

2001

United States

alteration of starch biosynthesis, quality


Rice (Oryza sativa)


downregulation of amylose synthesis; downregulation of granule bound starch synthase (GBSS); synthesis of starch consisting of pure amylopectin

carbohydrate metabolism altered and phosphinothricin tolerance

improved starch quality for cooking and eating

QTL identification of Wx locus linked to amylose content, gel consistency, and gelatinization temperature


Sugar beet (Beta vulgaris)

Huazhong Agricultural University

Monsanto

soft grain

puroindoline (pinA and pinB) from wheat

Montana State University

high-proten, low-starch rice flour

mutant amylopullulanase (APU) pene from a thermobacterium

National Defense University; Academia Sinica

amylopectin starch properties

starch synthase (SSIIa) mutantion in rice

Tohoku National Agricultural Experiment Station; National Institute of Agrobiological Sciences; Kyushu University; Akita Prefectural University

alteration of carbohydrate composition alteration of carbohydrate composition and increased nutritional value

downregulation of granule bound starch synthase (GBSS), fructosyltransferase (FTF); fructan; glucanase; invertase

(Tan, et al., 1999)

1

field trial

China

1997

United States

(Krishnamurthy and Giroux, 2001)

United States

(Chiang, et al., 2003)

(Umemoto, et al., 2002)

Taiwan

Japan

Advanta Seeds

1

field trial

1997

Netherlands

Van der Have

1

field trial

1993

Netherlands

Van der Have Van der Have

2 2


1994 1997

Netherlands Netherlands

Sweet potato (Ipomoea batatas)

amylose-free starch

silencing of granule-bound starch synthase I (GBSSI) from sweet potato

Kyushu National Agricultural Experiment Station; Research Institute of Agricultural Resources, Ishikawa Agricultural College; Okinawa Subtropical Station, Japan International Research Center for Agricultural Sciences


starch reduced

Genes from human and mouse


1

field trial

2002

United States

Tomato (Lycopersicon esculentum)


ADP-glucose pyrophosphorylase from corn; Shrunken 2 from corn


2

field trials

1999

United States


1

field trial

2000

United States



Biogemma

1

field trial

1999

United Kingdom

Biogemma Biogemma

4 2


2003 2004


Compañía Navarra Productora de Semillas; Senasa Compañía Navarra Productora de Semillas; Senasa

1

field trial

1996

Spain

1

field trial

1997

Spain

altered starch composition

CSIRO

1

field trial

1996

Australia

alteration of starch biosynthesis and downregulation of sucrose

IACR, University of Bristol

1

field trial

2002

United Kingdom


Monsanto Monsanto Monsanto

1 1 1


1996 1997 1998



Nickerson Biocem

2

field trials

1994

United Kingdom

modified starch

Plant Biotechnology

1

field trial

1999

Canada

sucrose nonfermenting-1-related protein kinase from wheat dihydropteroate synthase ADP glucose pyrophosphorylase from wheat ATP transporter gene from Arabidpsis thaliana

alteration of starch biosynthesis and synthesis of glycogen branching enzyme

56

(Kimura, et al., 2001)

Japan



Institute (NRC)

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; GM Crop Database, AgBios, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 90/220/EEC, Belgian Biosafety Server, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 2001/18/EC, Belgian Biosafety Server, accessed January 2005; and published articles cited in the body of this table.

4.4.2

Fructans

Fructans are polysaccharides synthesized from sucrose and serving as a naturally occurring carbohydrate reserve in many plant species—including forage grasses, wheat, chicory, artichoke, asparagus, onions, garlic, leek, bananas, and agave cactus—as well as in fungi and bacteria. The two natural forms of fructans are inulins and levans. Inulins are more commonly produced by plants; levans, more commonly by fungi and bacteria. Fructans in plants have both important agronomic and nutritional properties. Extracted chicory inulin is a recognized natural food product. It and other inulin-type fructans are commonly used as sugar and fat substitutes and as stabilizers and fillers in a variety of food products. Their taste resembles that of glucose, but given their chemical structure they cannot be digested by the human gut. Instead, however they stimulate the growth of beneficial bacteria in the colon and have been associated with reduced risk of a number of diseases, including constipation, diarrhea, cancer, osteoporosis, cardiovascular disease, obesity, and diabetes (Roberfroid and Delzenne, 1998). Natural sources of inulin and levan are economically limited, and therefore a number of efforts are underway to engineering carbohydrate accumulating crops, such as chicory, potato, and sugar beet, to produce more natural inulin or to produce synthetic fructans, both inulins and levans (Cairns, 2003).

Table 4.10 Transgenic plants with modified fructans Crop

Trait

Genes

Institution(s)

Chicory (Cichorium intybus)

increased fructans

fructan fructan fructotransferase from onion

University of Utrecht

produces branched fructans

sucrose:fructan 6-fructosyltransferase (6-SFT) from barley

University of Basel; van der Have

Level and quality of inulin content

sucrose: sucrose fructosyltransferases (A33 and SST103)

Plant Research International, Wageningen


high fructan levels

levansucrase (SacB) from Bacillus amyloliquefaciens

DuPont

Italian ryegrass (Lolium multiflorum Lam.)

modified fructan metabolism

levansucrase (sacB) gene from Bacillus subtilis

Petunia (Petunia sp.)

accumulates fructans


Count

Action Year or reference (Smeekens, 1997)

(Sprenger, et al., 1997)

1

field trials

2003

Country Netherlands

Switzerland; Netherlands Netherlands

(Caimi, et al., 1996)

United States

ETH-Zurich; Monsanto; La Trobe University

(Ye, et al., 2001)

Switzerland; United States; Australia

1-sucrose:sucrose fructosyl transferase (1-FFT) and 1-fructan:fructan fructosyl transferase (1-FFT) from Jerusalem artichoke (Helianthus tuberosus)


(van der Meer, et al., 1998)



DuPont

carbohydrate metabolism

Saccharose-Saccharose-1-Fructosyltransferase (1SST) from Cynara scolymus; Fructan-Fructan-1-Fructosyltransferase (1FFT) from Cynara scolymus

Federal Biological Research Centre for Agriculture and Forestry (BBA), Institute for Integrated Plant Protection


levansucrase (SacB) from Erwinia amylovora

Institut Genbiol Forsch Berlin; Max-Plank Institute for Medical Research

57


1

field trial

Netherlands

United States

2000

Germany

(Merges, 1996)

Germany



produces full range of fructans

sucrose:sucrose 1-fructosyltransferase (1-SST) and fructan:fructan 1-fructosyltransferase (1-FFT) from globe artichoke (Cynara scolymus)

Max-Planck-Institut für Molekulare Pflanzenphysiologie, Golm

novel carbohydrate copolymer amylofructan

amyloplast-targeted bacterial fructotransferase

University of Utrecht


levansucrase (SacB) from Bacillus subtilis

University of Utrecht; Wageningen University; Advanta Seeds


levan sucrase (sacB), invertase (suc2), hexokinase (hxk), maltose binding protein (malE)

Van der Have


contains oligofructans that promote beneficial bacteria

fructosyltransferases

DuPont

Sugar beet (Beta vulgaris)

fructan producing

levansucrase (SucB) from Bacillus subtilis

Colorado State University; University of California Berkeley; van der Have

alteration of carbohydrate composition and synthesis of fructan


Sharpes International Seeds

(Hellwege, et al., 2000, Hellwege, et al., 1997)

Germany

(Smeekens, 1997)

Netherlands

(Gerrits, et al., 2001, Pilon-Smits, et al., 1996, van der Meer, et al., 1994)

Netherlands

1

1

field trial

1997

Netherlands

(Allen, et al., 2002)

United States

(Pilon-Smits, et al., 1999)

United States; Netherlands

field trial

United Kingdom

1995

fructan producing

1-sucrose:sucrose fructosyl transferase (1-SST) from Jerusalem artichoke (Helianthus tuberosus)





DuPont


sucrose:fructan 6-fructosyltransferase (6-SFT) from barley

University of Basel; van der Have

(Sprenger, et al., 1997)


levansucrase (SacB) from Bacillus subtilis

University of Utrecht; Rothamsted Experiment Station; van der Have; Advanta Seeds; University of Wageningen

(Ebskamp, et al., 1994, Gerrits, et al., 2001, Pilon-Smits, et al., 1995, Turk, et al., 1997)


Netherlands

United States

Switzerland; Netherlands Netherlands; United Kingdom

Data sources: Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; and published articles cited in the body of this table.

4.4.3

Sugars

Techniques are being applied to adjust levels of accumulation simple sugars as well as the profile of simple sugars in fruits, tomatoes, and sugarcane. In some cases sugar content has a direct bearing on fruit quality perceived by the consumer. In other cases it has an important affect on the processing quality of the crop, whether for food processing, wine making, or sugar production. The genetic methods involved include turning on and turning off the key enzymes that synthesize different sugars or that convert one type of sugar into another. For example, the enzyme invertase converts sucrose into glucose and fructose. Transgenic tomatoes with an antisense invertase gene to block the enzyme’s activity can build up much higher levels of sucrose at the expense of glucose and fructose (Klann, et al., 1996).

Table 4.11 Transgenic plants with modified simple sugars Crop

Trait

Genes

Institution(s)

Country

fruit ripening altered and fruit sugar profile altered

Sorbitol 6-phosphodehydrogenase from apple; ACC synthase from apple; ACC synthase antisense from apple; Ethylene forming enzyme from apple; Ethylene forming enzyme antisense from apple Sorbitol synthase from apple

University of California

1


Year

Apple (Malus domestica)

1997

United States

University of California University of California University of California

3 1 1

field trials field trial field trial

1999 2001 2002


sugar alcohol levels increased

Sorbitol dehydrogenase from apple

Cornell University

1

field trial

2000

United States

sugar alcohol levels increased

Sorbitol dehydrogenase from apple

Oregon State University

1

field trial

2000

United States

CSIRO

1

field trial

2002

Australia

Grape (Vitis vinifera)

expression of modified colour, sugar composition, flowering and fruit development

58

Count


Sugar accumulation and transport


Sucrose transporters (Vvsuc11, Vvsuc12, Vvsuc27)

high sucrose

Institute for Wine Biotechnology, University of Stellenbosch University of Stellenbosch

(Pretorius, et al., 2004)

South Africa


South Africa

Strawberry (Fragaria ananassa)


DNA Plant Tech

1

field trial

1999

United States

Sugarcane (Saccharum officinarum)

increased sucrose and reduced juice colour

CSIRO

1

field trial

1997

Australia

Modified sucrose metabolism and high sucrose

Sucrose metabolism genes; novel promoters

South African Sugar Experiment Station; University of Stellenbosch


fruit sugar profile altered


BHN Research BHN Research BHN Research BHN Research BHN Research BHN Research

1 1 1 1 1 1

field trial field trial field trial field trial field trial field trial

1998 1999 2000 2001 2002 2003




Calgene Calgene Calgene Calgene Calgene Gargiulo

2 1 1 2 1 1

field trials field trial field trial field trials field trial field trial

1993 1994 1994 1995 1996 1997


fruit sugar profile altered

Invertase from tomato

Campbell Campbell Campbell

1 1 1


1995 1996 1997


Harris Moran

1

field trial

1997

United States

Nestle

1

field trial

1995

United States

PetoSeed PetoSeed

1 1


1995 1996


Sunseeds

1

field trial

1996

United States

(Klann, et al., 1996)

United States

carbohydrate metabolism altered carbohydrate metabolism altered



fruit sugar profile altered fruit sugar profile altered



South Africa

Invertase antisense from tomato




1

field trial

1994

United States


University of Wisconsin University of Wisconsin University of Wisconsin

1 1 1


1994 1995 1997


Invertase from tomato; Invertase from Saccharomyces cerevisiae; Pectin esterase antisense from tomato

Zeneca

5

field trials

1996

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004,and published articles cited in the body of this table.

4.5

Micronutrients and functional metabolites

Plants are an important source of most of the micronutrients essential to human nutrition, including both vitamins and minerals, as well as many additional organic compounds that have been linked to certain health benefits, often called functional nutrients or ‘nutraceuticals’. However, these essential and functional nutrients are often available only in small concentrations, and are certainly not available in sufficient quantities in those food plants that are the most common source of macronutrients or calories in human diets. Genetic techniques are being developed to enhance the biosynthesis of a range of vitamins, increase the bioavailability of minerals, and increase the production of functional secondary metabolites. The innovations identified in this study draw upon several reviews of plant micronutrient enhancements using biotechnology (Chassy, et al., 2004, DellaPenna, 1999, Galili, et al., 2002, Grusak and DellaPenna, 1999, Kinney, 2003, Monsanto, 2003, Schijlen, et al., 2004) and to a lesser extent data on field releases.

59


4.5.1


Vitamins

Genes have been identified and introduced into a range of plants that synthesize carotenoids (including beta-carotene or pro-vitamin A, phytoene, lycopene), vitamin C, vitamin E, and folate. Also, levels of expression have been amplified in plants that already produce such vitamins.

Table 4.12 Transgenic plants with increased vitamin content Crop

Trait

Genes

Institution(s)

Arabidopsis

increased vitamin E

gamma-Tocopherol C-methyltransferase from Arabidopsis

University of Nevada, Reno

incresased folate levels

GTP cyclohydrolase-1 (folE) from E. coli

Donald Danforth Plant Science Center; Tufts University; DuPont


high alpha- and beta-carotene

phytoene synthase (crtB) gene from bacterium

Monsanto


carotenoid metabolism altered

Mustard (Brassica juncea)

Monsanto

increased vitamin C

dehydroascorbate reductase (DHAR) from wheat

University of California, Riverside

high vitamin E

gamma-tocopherol methyl transferase


high vitamin E

homogentisic acid geranylgeranyl transferases (HGGT)

USDA-ARS; Donald Danforth Plant Science Center

high beta-carotene

phytoene synthase

Tata Energy Research Institute Tata Energy Research Institute

phytoene synthase (Ssu psy) from corn; phytoene desaturase (Ssu-tpCrt1) Potato (Solanum tuberosum)

beta-carotene increased

beta-carotene hydroxylase antisense from potato

increased carotenoid content and increased food quality Rice (Oryza sativa)

(Hossain, et al., 2004)

United States

(Shewmaker, et al., 1999)

United States

field trial

2001

United States

(Dechenaux, et al., 2003)

United States

(Rocheford, 2002)

United States

(Cahoon, et al., 2003)

United States

(Dhawan, 2001, Shewmaker, et al., 1999) (Sharma, et al., 2003)

India India

2002

United States

Technische Universität München

1

field trial

2002

Germany

ETH-Zurich

phytoene synthase (psy) from daffodil; phytoene desaturase (crt1); lycopene cyclase (lyC)

ETH-Zurich; International Rice Research Institute PhilRice Syngenta Syngenta


high vitamin C

D-galacturonic acid to ascorbic acid enzyme in strawberry fruit

University of Malaga; Universidad de Córdoba


astaxanthin (a carotenoid that occurs in the natural diet of many aquatic animals and that is supplemented in feed - creates the characteristic pink color to salmon, trout, and shrimp), produced in flowers

beta-carotene ketolase (CrtO) from the algae Haematococcus pluvialis

Hebrew University of Jerusalem


United States

field trial

phytoene synthase from daffodil

carotenoid content altered

1

Country

1

high beta-carotene ‘Golden Rice’

phytoene synthase (psy) from corn; phytoene desaturase (crt1)

Action Year or reference (Shintani and DellaPenna, 1998)

Boyce Thompson Institute

phytoene expressed

‘Golden Rice’ cross breeding to transfer trait from japonica to local indica varieties seed composition altered 4 ‘Golden Rice 2’ with increased betacarotene content

Count

(Burkhardt, et al., 1997) (Byerlee and Fisher, 2000, Datta, et al., 2003) (Atanassov, et al., 2004) 1

field trial 2004 (Paine, et al., 2005)

Switzerland Switzerland; Philippines Philippines United States United Kingdom

(Agius, et al., 2003, Medina-Escobar, et al., 1997)

Spain

(Mann, et al., 2000)

Israel

R J Reynolds

1

field trial

1994

United States

1

field trial

2003

United States

vitamin C content increased

L-gulono-gamma-lactone oxidase from Rattus norvegicus

Virginia Tech

high beta-carotene and lycopene levels

lycopene beta-cyclase (beta-Lcy) from tomato up and down-regulated

ENEA; CNRS

(Rosati, et al., 2000)

Italy; France

lycopene biosynthesis

phytoene desaturase (Pds) gene from tomato isolated


(Mann, et al., 1994)

Isreal

increased beta-carotene

map-based cloning of beta and old-gold color mutations


(Ronen, et al., 2000)

Isreal

carotenoid metabolism altered, high lycopene

lycopene cyclase (tLcy) from tomato

Metapontum Agrobios

1

high lycopene and high total soluble solids content

QTL marker assisted breeding

Pennsylvania State University

(Buschena and Zilberman, 1999)

field trial

2004

4 “Composition” here refers to beta-carotene content. The field trial was conduced in Louisiana, according to the U.S. field trial database. Several accounts have been made to a field trial of Golden Rice at the Louisiana State University AgCenter in collaboration with the Golden Rice Humanitarian Board, of which Syngenta is a member Shultz, B. (2004 ) Golden Staple. Crowley, LA..

60

Italy

United States



carotenoid metabolism altered

phytoene synthase from tomato

PetoSeed

1

field trial

1995

United States

high lycopene and beta-carotene levels

phytoene desaturase (crtl) and phytoene synthase (crtB) from bacterium Erwinia uredovora

University of London; Syngenta

carotenoid levels increased


University of Nottingham

carotenoid content altered


Zeneca

7

field trials

1996

United States

phytoene destaurase from Erwinina uredova pentenlypyrophosphate isomerase from tomato

Zeneca

3

field trials

1998

United States

phytoene synthase from Erwinia spp.

Zeneca

6

field trials

1999

United States

(Fraser, et al., 2001, Fraser, et al., 2002, Nelson and Romer, 1996)

United Kingdom

(Fray, 1995, Fray and Grierson, 1993)

United Kingdom

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004, and published articles cited in the body of this table.

4.5.2

Minerals

The most common genetic strategy to increasing the availability of mineral nutrients involves reduction in the level of phytic acid (or phytate) a mineral-binding compound that naturally occurs in plants. Phytic acid combines with the absorbed phosphorus, iron, zinc, magnesium, or calcium in plants and makes these minerals very difficult for humans or animals to absorb in the gut. In other words the presence of phytic acid reduces the bioavailability of minerals already present in plant tissues. The minerals instead pass through animals or humans as part of the solid waste and are excreted, which along with excess nitrogen from undigested proteins contributes to making animal waste a potent ‘nutrient rich’ pollutant. The formation of phytic acid is knocked out or reduced directly in ‘low phytic acid’ varieties. Without the phytic acid binding up these minerals, they are left in soluble form and are therefore nutritionally available. Another strategy breaks down the phytic acid complex by introducing the enzyme phytase, found in fungi and bacteria. ‘High phytase’ plant varieties are engineered with the phytase gene. The phytase enzyme is active in the plant. The level of phytic acid is reduced. As a result, phosphorus and the other minerals are soluble and nutritionally available, and the phosphorus pollutant is removed from the waste stream. In addition to solving the problem of mineral bioavailability, levels of mineral present in the tissues of crop plants often need to be increased in the first place. Several genetic methods have been invented to increase plants’ uptake of iron, zinc, and calcium.

Table 4.13 Transgenic plants with modified content or bioavailability of minerals Crop

Trait

Genes

Institution(s)

Alfalfa (Medicago sativa)

increased phytase

phytase gene from the fungus Aspergillus niger

U.S. Dairy Forage Research Center; University of Wisconsin


high zinc levels

zinc transporter gene from Arabidopsis

Donald Danforth Plant Science Center; CSIRO


phytate reduced

gene from ice plant

Dow

1

field trial

2000

United States

Dow

1

field trial

2001

United States

Dow

1

field trial

2002

United States

Du Pont - Pioneer

2

field trials

1998

United States

Du Pont - Pioneer

3

field trials

1999

United States

Monsanto

3

field trials

1999

United States

Monsanto

2

field trials

2000

United States

Monsanto

1

field trial

2001

United States

increased phosphorus

phytate reduced

Lettuce

high iron levels

ferratin from soybean

Central Research Institute

61

Count

Action Year or reference (Austin-Phillips and Zeigelhoffer, 2001, Koegel, et al., 1996)

(Venkatramesh, et al., 2000)

(Goto, et al., 2000)


United States; Australia

Japan



(Lactuca sativa)

Rice (Oryza sativa)




of Electric Power Industry iron levels increased

ferritin from soybean

Harris Moran

1

field trial

2002

high iron and zinc

ferratin from soybean

Central Research Institute Electric Power Industry; National Institute of Agrobiololgy Resources; International Rice Research Institute

(Goto, et al., 1999, Vasconcelos, et al., 2003)

Japan; Phillipines

high iron, with absorption enhanced

ferritin from Phaseolus vulgaris; phytase gene from Aspergillus fumigatus; metallothionein-like gene from rice

ETH-Zurich

(Gura, 1999, Lucca, et al., 2002, Lucca, et al., 2001, Lucca, et al., 2000, Potrykus, 1999)

Switzerland

high iron

nicotianamine aminotransferase gene from barley

University of Tokyo

(Takahashi, et al., 2001)

Japan

high phytase

phytase gene from yeast

Ajinomoto

(Nakamura, et al., 2000)

Japan

high phytase

phytase gene from Aspergillus niger

Virginia Tech

phytate reduced

inositol hexaphosphate phosphohydrolyase from soybean

Virginia Tech

high calcium levels

Ca/H antiporter (CAX1) gene from Arabidopsis

Baylor College of Medicine

enhanced phosphorus uptake

citrate synthase gene from Pseudomonas aeruginosa

Centro de Investigación y de Estudios Avanzados del IPN, Unidad Irapuato

(López-Bucio, et al., 2000)

high iron

ferratin gene from soybean

Central Research Institute of Electric Power Industry

(Goto, et al., 1998)

high iron

ferratin gene from soybean

CNRS; INRA

high phytase

phytase from Aspergillus niger

Mogen International

high iron

iron reductase (FRE 1 and FRE2) genes from yeast

Oregon State University

(Samuelsen, et al., 1998)

high phytase

two phytase genes from Aspergillus niger

Danish Institute of Agricultural Sciences; Royal Veterinary and Agricultural University; Riso National University

(Brinch-Pedersen, et al., 2000)

(Denbow, et al., 1998, Li, et al., 1997) 1

field trial

United States

United States

2002

United States

(Hirschi, 1999)

United States

(van Wuytswinkel, et al., 1999)

(Pen, et al., 1993)

Mexico

Japan

France

Netherlands

United States Denmark


4.5.3

Functional secondary metabolites

Most of the secondary plant metabolites with ‘functional’ properties being genetically engineered involve the flavonoid pathway. These include anthocyanins (also discussed in section 4.8.3 below regarding their role in the modification of flower color), stilbenes (such as resveratrol, found in grape skins and red wine, and associated with a reduced risk of heart disease, popularized as the ‘French Paradox’), and isoflavonoids (found in soybean and other legumes, often called phytoestrogens, and associated with a number of health benefits including relief of menopause, reduced osteoporosis, improved blood cholesterol, lowered risk of coronary disease, and lower risk of certain cancers). Other secondary metabolites being engineered include plant sterols, stanols, and capsaicin (the ‘heat’ in hot peppers).

Table 4.14 Transgenic plants with modified functional secondary metabolites, such as flavonoids, sterols, capsaicin, etc. Crop

Trait

Genes

Institution(s)


secondary metabolite increased

resveratrol synthase from peanut

Samuel Roberts Noble Foundation Samuel Roberts Noble Foundation Samuel Roberts Noble Foundation Samuel Roberts Noble Foundation

produces resveratrol

resveratrol glucoside

62

Count


Year

Country

1

1998

United States

1

field trial

1999

United States

(Hipskind and Paiva, 2000)

United States

1

field trial

2001

United States



secondary metabolite increased

resveratrol synthase from peanut



produces stilbenes

stilbene synthase gene from grape

University of Hannover; Bundesforsch Anstalt Ernahrung, Institut Ernahrungsphysiol, Karlsruhe

Arabidopsis

produces soy isoflavones

isoflavone synthase (IFS) from soy transcription factor C1 and R (CRC) from corn chalcone reductase (CHR) from soy

DuPont

increased level of isoflavones

isoflavone synthase (IFS) from soy chalcone isomerase from alfalfa

Samuel Roberts Nobel Foundation

accumulation of anthocyanins

transcriptional activators R and C1 from corn

Stanford University


produces stilbene resveratrol

stilbene synthase (STS) gene from grape

University of Hamburg


accumulation of sterol compounds and tocopherols

3-hydroxysteroid oxidases steroid5a-reductases tocopherol biosynthetic enzyme

Monsanto


production of soy isoflavones

isoflavone synthase (IFS) from soy

DuPont

Downy thorn apple (Datura metel)

high levels of scopolamine

hyoscyamus H6H tropinone reductase 2

Indian Institute of Science; Madurai Kamraj University

(Department of Biotechnology, 2003)

India

Henbane (Hyoscyamus niger)

high scopolamine levels

hyoscyamus H6H tropinone reductase 2

Indian Institute of Science; Madurai Kamraj University


India

Kiwi (Actinidia deliciosa)


stilbene synthase (STS) from grape

National Institute of Fruit Tree Science; Zhejiang Academy of Agricultural Science

(Kobayashi, et al., 2000)

Pepper (Capsicum annuum)

hotter peppers, with higher capsaicin levels

capsaicin biosynthesis enzymatic regulation genes

CFTRI



antioxidant flavonid anthocyanin and steroid alkaloid glycoside production

chalcone synthase (CHS) chalcone isomerase (CHI) dihydroflavonol reductase (DFR)

Institute of Bioorganic Chemistry, Polish Academy of Science; University of Wroclaw; Mickiewicz University; Wroclaw Medical University

(Lukaszewicz, et al., 2004, Stobiecki, et al., 2003)

accumulation of anthocyanins

transcription factor genes LC and C1 from corn

Plant Research International, Wageningen; Unilever

Rice (Oryza sativa)


stilbene synthase (STS) from grape

Bayer; Max-PlankInstitut fur Zuchtungsforshcung; University of Hamburg


high isoflavones secondary metabolite increased increased isoflavone levels

isoflavone synthase from soy

DuPont Pioneer DuPont

2

(Jung, et al., 2000) field trials 2001 (Yu, et al., 2003)


Monsanto Monsanto

2 6


1999 2000


(Gustine, 1995, Hain, et al., 1993)

Germany; United States

(Yu, et al., 2000)

United States

Cl and R transcription factors from corn

sterols and stanols increased



produces resveratrol

phytoalexin stilbene synthase (STS) gene from grape

Bayer; USDA-ARS

production of soy isoflavones

isoflavone synthase from soy

DuPont

high chlorogenic acid antioxidant levels

hydroxycinnamoyl-CoA quinate hydroxycinnamoyl transferase (HQT) encoding chlorogenic acid (CGA)

John Innes Centre; Institute of Food Research

antioxidant enzyme increased

Lipton Lipton Lipton

high flavonol antioxidants

chalcone isomerase from petunia

high flavonol antioxidants

transcription factor genes LC and C1 from corn

sterols increased Wheat (Triticum aestivum)

produces stilbene resveratrol produces stilbene resveratrol

Plant Research International Wageneigen; Unilever Research; University of Essex; Plant Research International, Wageningen; Unilever; University of Exeter; University of Salamanca; Institute of Food Research, Norwich Texas Tech University

stilbene synthase (STS) gene from grape stilbene synthase (STS) gene

63

University of Hamburg University of Hohenheim

1 1


2002 2004


(Szankowski, et al., 2003)

(Jung, et al., 2000, Yu, et al., 2000)

United States

(Liu, et al., 2002)

United States

(Lloyd, et al., 1992)

United States

(Leckband, et al., 2002)

(Venkatramesh, et al., 2000)

United States

(Yu, et al., 2000)

United States

(Aharoni, et al., 2000)

(Stark-Lorenzen, et al., 1997)

(Niggeweg, et al., 2004)

1 1 1

Germany


1998 1999 2000

Japan; China

Poland

Netherlands; United Kingdom

Germany

United Kingdom


(Bovy, et al., 2002, Colliver, et al., 2002, Muir, et al., 2001)

United Kingdom; Netherlands

(Bovy, et al., 2002, Le Gall, et al., 2003)

Netherlands; United Kingdom; Spain

1

field trial

1996

(Leckband, et al., 2002) (Fettig and Hess, 1999)

United States Germany Germany




4.6

Reduction or removal of non-nutrients, allergens, and toxins

A wide range of compounds naturally occurring in plants, involving both integral constituents such as proteins and secondary metabolites as well as the byproducts of common pathogens, can be removed or mitigated through genetic interventions. The innovations identified in this study draw upon several reviews discussing the prevention and removal of unwanted components (Chassy, et al., 2004, Galili, et al., 2002, Kinney, 2003, Monsanto, 2003), our reading of the primary scientific literature, and, to a lesser extent, data on field releases of genetically modified plants. 4.6.1

Non-nutritional, anti-nutritional, and toxic plant metabolites

While some secondary metabolites, such as vitamins or functional compounds, can deliver nutritional and health benefits, others can have negative affects on human health. The problems can range from mild inconveniences caused by ‘non-nutritional’ compounds to life threatening poisons. Some individuals suffer from health impacts through over reaction to otherwise desired stimulants such as caffeine and nicotine. Raffinose, a compound found in soy can cause gastrointestinal discomfort and gas in many individuals. Among fatal toxins that can occur in crop plants is the steroidal glycoalkaloid solanine produced in the green eyes of potatoes.

Table 4.15 Transgenic plants with removed non-nutritional, anti-nutritional, and toxic plant metabolites Crop

Trait

Genes

Institution(s)

Arabidopsis

reduced glucosinolates

cyanogenic CYP79A1 gene from Sorghum bicolor; cytochrome P450 CYP79A2 from Arabidopsis

Royal Veterinary and Agricultural University


low indole glucosinolates

tryptophan decarboxylase gene from Catharanthus roseus

Universite de Montreal

reduced glucosinolates

National Research Centre on Plant Biotechnology


reduced or eliminated cyanogen content

cytochrome P450 genes ( CYP79D1 and CYP79D2) antisense

Ohio State University

Coffee (Coffea arabica)

decaffinated

CS2B antisense

CFTRI

decaffinated

theobromine synthase RNA interference

decaffinated

altered caffeine content

Count

Action Year or reference (Bak, et al., 1999, Wittstock and Halkier, 2000) (Chavadej, et al., 1994)

(Vageeshbabu and Chopra, 1997)

(Siritunga and Sayre, 2003)

Country Denmark

Canada India

United States


India

Nara Institute for Science and Technology

(Ogita, et al., 2003)

Japan

N7-methyltransferase genes from coffee (Coffea robusta)

SPIC Science Foundation


India

xanthosine-N7-methyltransferase (XMT) from coffee (Coffea arabica) Xanthosine-N7-methyltransferase antisense from coffee

University of Hawaii

characterization of caffeine biosynthesis enzyme

N-methyltransferase from coffee (Coffea arabica)

University of Zurich

Legumes (Fabaceae)

reduced raffinose saccharides

alpha-galacosidase from bacterium (Thermotoga neapolitana)

International Institute of Cell Biology

Poppy (Papaver somniferum)

altered alkaloid production pathway


steroidal glycoalkaloids reduced

caffeine levels reduced

UDP glucose glucosyltransferase from potato

64


1

(Moisyadi, et al., 1998)

United States

field trial

United States

1999

(Waldhauser, et al., 1997)

(Kuchuk, 2000)

Switzerland

Ukraine

CSIRO

1

field trial

2002

Australia

USDA-ARS USDA-ARS USDA-ARS USDA-ARS

2 2 1 3

field trials field trials field trial field trials

1997 1998 1999 2000



reduced levels of steroidal glycoalkaloids


solanidine glucosyltransferase (Sgt) antisense

USDA-ARS USDA-ARS USDA-ARS USDA-ARS USDA-ARS


reduced raffinose saccharides reduces flatulance and improves growth performance

inhibiting galactinol synthase and myo-inositol 1phosphate synthase

DuPont

Tea (Camellia sinensis)

reduced caffine

caffeine synthase gene from tea

Ochanomizu University; University of Tsukuba; University of Glasgow


visual marker and nicotine levels reduced nicotine levels reduced

Quinolinate phosphoribosyl transferase from tobacco; Quinolinate phosphoribosyl transferase antisense from tobacco Aquaporin from cucumber

nicotine levels reduced

Quinolinate phosphoribosyl transferase antisense from tobacco

alkaloids reduced

Formamidopyrimidine DNA glycosylase from tobacco

North Carolina State University Vector Tobacco

3 2

field trials 2001 field trials 2002 (McCue, et al., 2003) field trial 2003 field trial 2004


(Hitz, et al., 2002, Hitz and Sebastian, 1998, Kerr, 1997)

United States

(Kato, 2000)

Japan; United Kingdom

1 1

1

field trial

1999

United States

15

field trials

2001

United States

Vector Tobacco

7

field trials

2002

United States

Vector Tobacco

1

2002

United States

Vector Tobacco

4

USDA deregulation field trials

2002

Canada

Vector Tobacco

1

field trial

2003

United States

Vector Tobacco

1

field trial

2003

Canada

Vector Tobacco

4

field trials

2004

United States

Southern Piedmont AREC

1

field trial

2002

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; GM Crop Database, AgBios, accessed January 2005; and published articles cited in the body of this table.

4.6.2

Allergens

Three general strategies have been taken to reduce allergenic effects of plants. The most straightforward is the attempt to knock out an allergenic protein, whether using an antisense gene, reducing the level of expression of those proteins, or targeting the allergens with monoclonal antibodies expressed in plants. Another strategy involves the transgenic expression of the compound thioredoxin, which reduces the allergenic properties of problematic proteins and increases their digestibility. Finally, attempts are being made to induce an allergen-like immune response stimulus using ‘weakened’ transgenic versions of allergenic proteins, in order to suppress development of asthma.

Table 4.16 Transgenic plants with allergens removed, reduced, or mitigated Crop

Trait

Genes

Institution(s)

Italian or perennial ryegrass (Lolium multiflorum)

pollen allergen reduced

Pollen allergen (Lolp) antisense from Italian ryegrass

Samuel Roberts Noble Foundation

Lupin (Lupinus angustifolius)

allergen-like immune response stimulus in order to suppress development of asthma

gene for a potential allergen sunflower seed albumin (SSA-lupin)

Australian National University

Peanut (Arachis hypogae)

reduced allergens

Ara h 1, Ara h 2, and Ara h 3 proteins from peanut

University of Arkansas for Medical Sciences and Arkansas Children's Hospital Research Institute

(Bannon, et al., 2001, Bannon, et al., 1999, Burks, et al., 1999, Rabjohn, et al., 2002)

United States


reduced allergen

IgE binding epitopes of patatin protein (Sol t 1) from potato

Monsanto; Mount Sainai Hospital

(Alibhai, et al., 2000, Astwood, et al., 2000)

United States

Rice (Oryza sativa)

low allergenicity (Kinuhikari)

AS albumin

Mitsui Toatsu

hypoallergenic

16 kDa allergen protein antisense from rice

Nagoya University; Mitsui Toatsu


hypoallergenic

cysteine protease P34 from soy silenced

hypoallergenic

P34/Gly m Bd 30K protein from soybean

USDA-ARS; Donald Danforth Center; University of Arkansas; Pioneer DuPont University of Arkansas for Medical Sciences

65

Count 2

Action or reference field trials

Year

Country

2004

United States

(Smart, et al., 2003)

1

field trial

Australia

1991

Japan

(Tada, et al., 1996)

Japan

(Herman, 2002, Herman, et al., 2003)

United States

(Helm, et al., 2000)

United States




hypoallergenic

Gly m Bd 30K (soybean vacuolar protein P34), Gly m Bd 28K, alpha-subunit of beta-conglycinin, KSTI, Gly m2, Gly m IA, Gly m IB, rGLY m3 and glycinin G1 from soybean

Pioneer-DuPont

reduced allergen

thioredoxin

University of California, Berkeley

(Jung and Kinney, 2001)

United States

(Buchanan and Yoon, 2000)

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Table of Developing Transgenic Crop Plants in Japan (Field Tests and General Releases), Agriculture, Forestry, and Fisheries Research Council, Japanese Ministry of Agriculture, Forestry, and Fisheries, May 2003; and published articles cited in the body of this table.

4.6.3

Mycotoxins

Contamination by mycotoxins (fumonisins, aflotoxins, vomitoxins) resulting from post-harvest fungal growth (largely Fusarium and Aspergillus) can severely compromise grain quality for both animal feed and human food uses, leading to significant economic losses and health risks including carcinogenicity. Reduction or elimination of mycotoxin contamination shares economic mechanisms similar to other quality improvements in this category, including the elimination of plant derived toxins or antinutrient components. The results are greater feed efficiency (given reduced sickness and digestion problems in animals), higher value of product to food processors, and greater confidence among consumers of food safety. Three general transgenic approaches are being pursued to reduce mycotoxins. The first approach is to break down the mycotoxin itself. The second approach is to combat infecting fungus through disease or fungal resistance traits. The third is to reduce susceptibility to fungal contamination in the first place by reducing insect damage through insect resistance traits, including Bt. Because the bulk of Bt and other insect resistance traits lie outside of the scope of this study and only indirectly confer the benefit of mycotoxin reduction, we include here only on the first and second approaches, those which specifically target mycotoxins or the fungi that produce them.

Table 4.17 Transgenic plants with mycotoxins removed, reduced, or mitigated Crop

Trait

Genes

Institution(s)

Country

mycotoxin degradation

Amino polyol amine oxidase

Pioneer Pioneer

1 9

Action or reference field trial field trials

Year


1996 1997


Pioneer

6

field trials

1998

United States

Pioneer

3

field trials

1999

United States

(Duvick, 2001)

United States

decreased fuminosin

fuminosin esterase; fuminosin deaminase

Count

Pioneer Pioneer

1

field trial

2001

United States

Pioneer

1

field trial

2004

United States

Pioneer

2

field trials

2000

United States


fumonisin degradation


Fusarium fungal resistance

FRG gene from fungus

Instituto de Agricoltura Sostenibile, Consejo Superior de Investigaciones Cientificas

1

field trial

2004

Spain

fumonisin resistance


Syngenta

1

field trial

2003

Germany

fumonisin resistance tolerance to Fusarium


Syngenta Syngenta

1 2


2003 2004

United Kingdom Germany


66


4.7


Extended shelf life

The biology involved when fruit ripens and leaves wilt and fall off includes a complex set of enzymatically mediated chemical reactions that remain a subfield of considerable interest in the plant sciences. Transgenic approaches have proven useful both in elucidating some of the cellular and molecular mechanisms involved as well as producing some promising candidates for commercial produce and ornamentals, particularly in tomato and carnation. In addition the chemical processes that cause browning of fruits and vegetables when cut or bruised are of interest to the fresh produce industry as significant growth continues in prepared and pre-cut segments of the market. 4.7.1

Control of fruit ripening

To major approaches have been taken to control fruit ripening. One approach is to modify the expression of proteins that affect components of the cell wall during ripening and fruit softening, proteins such as polygalacturonase (PG), glucanases, and pectin methyltransferase (Brummell and Harpster, 2001). The second approach is to modify expression of the plant hormone ethylene, which plays a central role in signaling fruit ripening, mostly concentrating on regulating levels of several key intermediates and enzymes involved in the biosynthesis of ethylene, including Sadenosylmethionine (SAM) transferase, S-adenosylmethionine (SAM) hydrolase, ACC synthase, and ACC oxidase, ACC deaminase, and ethylene forming enzyme (Stearns and Glick, 2003). But fruit ripening and softening is a complex process, and other aspects of fruit quality have been addressed in a range of ways. A significant amount of work was done in the early 90s to bring controlled fruit ripening traits to market. Six of the innovations identified here (one melon and five tomatoes) reached the phase of regulatory filings, and two, Calgene’s FlavrSavr tomato and Zeneca’s processing tomato, are among the few quality traits that have been commercialized to date. However, both of these products were discontinued, and currently nothing with controlled ripening traits is on the market.

Table 4.18. Transgenic plants with controlled fruit ripening Crop

Trait

Genes

Institution(s)

Country

fruit ripening altered

S-adenosylmethionine (SAM) transferase from bacteriophage T3; S-adenosylmethionine (SAM) transferase from E. coli

Agritope

2


Year


2000

United States

Exelixis

1

field trial

2001

United States

Cornell University

1

field trial

1995

United States

Cornell University

1

field trial

1999

United States

Cornell University

1

field trial

2003

United States

(Hrazdina, et al., 2003)

United States


Banana (Musa acuminata)

ACC synthase from apple; ACC synthase antisense from apple; Attacin E from Hyalophora cecropia; Cecropin B from Hyalophora cecropia; Polygalacturonase from apple

controlled ripening

ACC synthase from apple, antisense

Cornell University

fruit ripening altered and fruit sugar profile altered

ACC synthase from apple; ACC synthase antisense from apple; Ethylene forming enzyme from apple; Ethylene forming enzyme antisense from apple; Sorbitol 6-phosphodehydrogenase from apple; Sorbitol 6-phosphodehydrogenas

University of California, Berkeley

controlled ripening

ACS2 expression by producing antisense RNA production

Bose Institute

controlled ripening

ACC synthase

Indian Agricultural Research Institute, New

67

Count

1

field trial

1997

United States


India

(Sharma, et al., 2003)

India



Delhi Broccoli (Brassica oleracea L. var. italica)

delayed post-harvest wilting

ACC oxidase antisense from tomato

Crop & Food Research Ltd; Lincoln University

Coffee (Coffea arabica)

ethylene production reduced

ACC oxidase antisense from coffee; ACC synthase antisense from coffee


2

field trials

1999

United States

Monsanto

1

field trial

1994

United States

Monsanto

1

field trial

1997

United States

1

field trial

1999

Italy

fruit ripening delayed

(Henzi, et al., 1999)

New Zealand


improved processing characteristics

tryptophan-2-monoxygenase

Università degli studi di Ancona, Dip. Biotec. Agrarie

Mango (Mangifera indica)

Delayed ripening

ACC synthase antisense; ACC oxidase antisense; other enzymes involved in ripening

Institute of Plant Breeding-University of the Philippines at Los Baños (IPB-UPLB)

Melon (Cucumis melo)


S-adenosylmethionine (SAM) transferase from E. coli; S-adenosylmethione (SAM) hydrolase from bacteriophage T3

Agritope

1

field trial

1996

United States

Agritope

5

field trials

1997

United States

Agritope

6

field trials

1998

United States

Agritope

5

field trials

1999

United States

Agritope

1

1999

United States

Agritope

4

USDA deregulation pending an environmental assessment field trials

2000

United States

Exelixis

1

field trial

2001

United States

Asgrow

1

field trial

1995

United States

Asgrow

2

field trials

1996

United States

Harris Moran

1

field trial

1998

United States

Harris Moran

1

field trial

1999

United States

delayed ripening

S-adenosylmethionine (SAM) hydrolase (sam-k) gene from bacteriophage T3



Papaya (Carica papaya)

Pepper (Capsicum annuum)

Pineapple (Ananas comosus)

Philippines

ripening control

ACC oxidase antisense from melon

INRA

ripening control

ACC oxidase from apple antisense

Universidade Federal de Pelotas


gene from melon

University of California, Davis

Delayed ripening

ACC synthase antisense ACC oxidase antisense

Institute of Plant Breeding-University of the Philippines at Los Baños (IPB-UPLB)


Philippines

Delayed fruit ripening, extended shelf life

ACC oxidase gene from Eksotika papaya

Malaysian Agricultural Research and Development Institute (MARDI)


Malaysia

ethylene production reduced

Recombinase from bacteriophage P1 ACC synthase from papaya


1

field trial

1998

United States

University of Queensland

1

field trial

1999

Australia


1

field trial

2002

Australia

Agritope

2

field trials

2000

United States

Exelixis

1

field trial

2001

United States

(Gao, et al., 2003)

Japan; United States

delayed ripening

Pear (Pyrus communis)



S-adenosylmethionine (SAM) transferase from bacteriophage T3 S-adenosylmethionine (SAM) transferase from E. coli

1

(Ayub, et al., 1993)

France

(Fonseca, et al., 2001)

Brazil

field trial

1999

United States

ripening control

ACC synthase and ACC oxidase from pear, sense and antisense

Yamagata University; University of California, Davis


B-1,4-endoglucanase antisense from pepper

DNA Plant Tech

1

field trial

1994

United States

B-1,3-glucanase antisense from pepper glucanase from pepper

DNA Plant Tech

1

field trial

1995

United States

DNA Plant Tech

1

field trial

1996

United States

hemicellulase from pepper

DNA Plant Tech

1

field trial

1997

United States

flower and fruit set altered

habinlin from Capparis masaikai ACC synthase


1

field trial

1997

United States

ethylene synthesis reduced

ACC synthase from pineapple cysteine proteinase inhibitors from rice


1

field trial

2003

United States


1

field trial

1998

Australia

flowering & ripening qualities

68


controlled ripening Potato (Solanum tuberosum)


ACC synthase from pineapple, sense and antisense

solids increased

controlled ripening

Frito Lay Frito Lay ACC synthase

proteinase inhibitor I tryptophan monooxygenase from Agrobacterium




(Botella, et al., 2000)

Australia



1991 1992


India

Monsanto

1

field trial

1991

United States

Monsanto

3

field trials

1992

United States

Monsanto

3

field trials

1993

United States

Monsanto

6

field trials

1994

United States

Monsanto

5

field trials

1995

United States

Monsanto

11

field trials

1996

United States

Monsanto

8

field trials

1999

United States

Monsanto

1

field trial

1999

Canada

Monsanto

5

field trials

2000

United States

North Dakota State University

1

field trial

1993

United States

ethylene metabolism altered

ACC oxidase antisense from potato

Pennsylvania State University

1

field trial

1999

United States

alteration of ethylene biosynthesis

rol gene, S-adenosylmethionine (SAM) decarboxylase

Scottish Crop Research Institute

1

field trial

1998

United Kingdom

ethylene metabolism altered

ethylene receptor protein from Arabidopsis

University of Idaho

1

field trial

2001

United States


S-adenosylmethionine (SAM) transferase from bacteriophage T3 S-adenosylmethione (SAM) hydrolase from E. coli; S-adenosylmethionine (SAM) transferase from E. coli; polygalacturonase inhibitor protein

Agritope

1

field trial

1995

United States

Agritope

2

field trials

1998

United States



Università degli studi di Ancona

1

field trial

1999

Italy


S-adenosylmethionine (SAM) transferase from E. coli

Agritope

1

field trial

1996

United States

Agritope

1

field trial

1998

United States

Calgene

1

field trial

1996

United States

DNA Plant Tech

2

field trials

1997

United States

Monsanto

1

field trial

1994

United States

1

field trial

1999

Italy

fruit ripening altered fruit ripening delayed

glucanase from strawberry polygalacturonase inhibitor protein from bean



1 2

Indian Agricultural Research Institute, New Delhi

tuber solids increased

Raspberry (Rubus idaeus)




Università degli studi di Ancona

delayed ripening by ethylene regulation

ACC deaminase

Agricultural Research Council - Vegetable and Ornamental Plant Institute (ARC-VOPI); University of Cape Town


S-adenosylmethione (SAM) hydrolase from E. coli S-adenosylmethione (SAM) hydrolase from bacteriophage T3; S-adenosylmethionine (SAM) transferase from E. coli S-adenosylmethionine hydrolase (SAMase) from bacteriophage T3

Agritope Agritope

controlled ripening


S-adenosylmethionine (SAM) transferase from bacteriophage T3


1 3

Agritope


South Africa

1992 1993


(Busquin, et al., 2004)

United States

Agritope

3

field trials

1994

United States

Agritope

2

field trials

1995

United States

Agritope

2

field trials

1996

United States

Agritope

1

USDA deregulation

1996

United States

Agritope

6

field trials

1997

United States

Agritope

2

field trials

1998

United States

Agritope

2

field trials

1999

United States

field trial

1995

United States


polygalacturonase antisense from tomato

Asgrow

1


ACC synthase antisense from tomato polygalacturonase antisense from tomato

Gargiulo

1

field trial

1995

United States

BHN Research

2

field trials

1996

United States

BHN Research

1

field trial

1997

United States

BHN Research

1

field trial

1998

United States

BHN Research

1

field trial

1999

United States

BHN Research

1

field trial

2000

United States

BHN Research

1

field trial

2001

United States

BHN Research

1

field trial

2003

United States

solids increased

controlled ripening

ethylene inducible promoters, both natural and synthetic, to express ACS2 by producing antisense RNA during fruit ripening

69

Bose Institute


India


controlled ripening fruit ripening delayed


polygalacturonase from tomato, antisense polygalacturonase from tomato; polygalacturonase antisense from tomato

Calgene Calgene

2

Calgene

1

field trial

1989

United States

Calgene

3

field trials

1990

United States

Calgene

3

field trials

1991

United States

ACC deaminase from Pseudomonas

Calgene

3

field trials

1992

United States



Calgene

1

controlled ripening

pectin methylesterase and polygalacturonase, antisense sucrose phosphate synthase from tobacco sucrose phosphate synthase from corn ACC deaminase from tomato ACC synthase antisense from tomato S-adenosylmethione hydrolase

Calgene

USDA 1992 deregulation (Lichtenberg, et al., 1993)

United States

isopentenyl transferase from Agrobacterium; pectin esterase antisense from tomato


Polygalacturonase antisense from tomato

(Shewmaker, et al., 1999) field trials 1988


United States

Calgene

5

field trials

1993

United States

Calgene

10

field trials

1994

United States

Calgene

2

1994

United States

Calgene

8

USDA deregulations field trials

1995

United States

USDA deregulations USDA deregulation

1995

United States

1996

United States



Calgene

2



Calgene

1



Campbell

1

field trial

1991

United States

Campbell

1

field trial

1992

United States

Campbell

2

field trials

1993

United States

Pectin methylesterase antisense from tomato Polygalacturonase from tomato

Campbell Campbell

1 1


1994 1995


Pectin methylesterase antisense from tomato

Campbell

2

field trials

1996

United States

Campbell

2

field trials

1997

United States


ACC oxidase; ethylene forming enzyme (EFE)

Central China Agricultural University (CCAU)


ACC synthase from tomato

DNA Plant Tech

2

field trials

1992

United States

DNA Plant Tech

6

field trials

1993

United States

DNA Plant Tech

10

field trials

1994

United States


DNA Plant Tech

1

1994

United States

Glucanase from pepper; Transcriptional activator from petunia

DNA Plant Tech

8


1995

United States


DNA Plant Tech

1

United States

5


1995

DNA Plant Tech

1996

United States

DNA Plant Tech

4

field trials

1997

United States

DNA Plant Tech

3

field trials

1998

United States

DNA Plant Tech

2

field trials

1999

United States

DNA Plant Tech

1

Health Canada approval for food use

1999

Canada

Hispareco

1

field trial

1993

Spain



reduced ethylene accumulation and delayed ripening


alteration of ripening characteristics

(Atanassov, et al., 2004, Zhu, et al., 2003)

China


pectin methylesterase from tomato

Heinz

1

field trial

1992

United States

fruit solids increased

pectin methylesterase from tomato; Pectin methylesterase antisense from tomato polygalacturonase

Heinz

3

field trials

1993

United States

IDAL; Heinz

1

field trial

1993

Portugal


Heinz

1

field trial

1994

United States

downregulation of pectin methylesterase

IDAL; Heinz

1

field trial

1994

Portugal

Heinz

1

field trial

1995

United States

fruit solids increased solids increased

polygalacturonase from tomato

Heinz

1

field trial

1999

United States


Pectin methylesterase antisense from tomato; Polygalacturonase from tomato; Polygalacturonase antisense from tomato Pectin methylesterase from tomato

Hunt-Wesson

3

field trials

1994

United States

Hunt-Wesson

5

field trials

1995

United States

Hunt-Wesson

1

field trial

1998

United States

controlled ripening

ACC synthase

Indian Agricultural Research Institute, New Delhi

controlled ripening

ACC deaminase from soil bacteria Pseudomonas chloraphis ACC deaminase from Pseudomonas chlororaphis

Monsanto

fruit ripening altered and solids increased

Monsanto


(Klee, et al., 1991, Reed, et al., 1995)

India

United States

2

field trials

1991

United States

Monsanto

4

field trials

1992

United States


Monsanto

10

field trials

1993

United States

ACC synthase antisense from tomato

Monsanto

18

field trials

1994

United States

Monsanto

10

field trials

1995

United States

Monsanto

1

1995

United States


Monsanto

1

USDA deregulation field trial

1998

United States

solids increased

Monsanto

1

field trial

1999

United States

fruit ripening altered by reduced ethylene

ACC deaminase from Pseudomonas chlororaphis

70





Pectin methylesterase from tomato; Polygalacturonase antisense from tomato; ACC synthase antisense from tomato Pectin methylesterase antisense from tomato; Polygalacturonase from tomato

PetoSeed

1

field trial

1991

United States

PetoSeed

3

field trials

1992

United States


PetoSeed

1

field trial

1993

United States

Invertase antisense from tomato; Polygalacturonase from tomato; Polygalacturonase antisense from tomato; Pectin esterase from tomato; Pectin esterase antisense from tomato ACC oxidase antisense from tomato

PetoSeed

9

field trials

1994

United States

PetoSeed Ibérica

3

field trials

1994

Spain

Invertase from Saccharomyces cerevisiae

PetoSeed

4

field trials

1995

United States

PetoSeed

4

field trials

1996

United States

Seminis; Zeneca

1

field trial

1998

Spain

Seminis

1

field trial

1999

United States

(Tieman, et al., 1992, Tieman, et al., 1995) 1 field trial 1992

United States

controlled ripening

pectin methylesterase, sense and antisense

Purdue University


pectin methylesterase from tomato

Purdue University

pectin methylesterase antisense from tomato

Purdue University

1

field trial

1993

United States

polygalacturonase from tomato; polygalacturonase antisense from tomato; pectin methylesterase from tomato; pectin methylesterase antisense from tomato

Purdue University

4

field trials

1994

United States

Purdue University

4

field trials

1995

United States

Purdue University

3

field trials

1996

United States

Purdue University

1

field trial

1997

United States

Purdue University

1

field trial

2001

United States

(Mattoo, et al., 2002, Mehta, et al., 2002) 1 field trial 2002

United States

fruit solids and firmness increased

high lycopene levels, improved juice quality and vine life

delayed ripening

S-adenosylmethionine decarboxylase gene (ySAMdc; Spe2) from yeast Catalase antisense from tomato; S-adenosylmethione decarboxylase from Saccharomyces cerevisiae

Purdue University; USDA-ARS Purdue University

Polygalacturonase antisense

Unifoods

1

field trial

1992

Australia

Unifoods

1

field trial

1993

Australia

controlled ripening

polygalacturonase from tomato

controlled ripening

ACC synthase, antisense

controlled ripening

E8 from tomato, antisense

fruit ripening altered Polygalacturonase antisense from tomato; Polygalacturonase from tomato; Glucanase from tomato Polygalacturonase antisense from tomato


United States

Ethylene receptor protein from tomato; Ethylene receptor protein antisense from tomato

Universiyt of California, Berkeley; University of California, Davis University of California, Berkeley; USDA-ARS University of California, Berkeley; Lawrence Berkeley Laboratory University of California, Davis University of California, Davis

United States

(Giovannoni, et al., 1989)

United States

(Oeller, et al., 1991)

United States

(Peñarrubia, et al., 1992)

United States

1

field trial

1999

United States

1

field trial

2003

United States

University of California, Davis

1

field trial

2004

United States


1

field trial

1996

United States

University of Florida University of Florida University of Florida

2 2 3

field trials field trials field trials

1997 1998 1999



1

field trial

2000

United States

1

field trial

1997

seed set reduced and fruit solids increased

Phytochrome A from oat

University of Georgia

controlled ripening controlled ripening

ACC oxidase from tomato ethylene-forming enzyme (EFE) from tomato, antisense

University of Nottingham University of Nottingham

(Hamilton, et al., 1990) (Malla and Gray, 2001)

United Kingdom United Kingdom

slower ripening, better tasting

Rin gene silenced

USDA-ARS; Boyce Thompson Institute; Syngenta

(Vrebalov, et al., 2002)

United States


polygalacturonase from tomato; polygalacturonase antisense from tomato

Van den Bergh Foods

controlled ripening

polygalacturonase antisense and sense from tomato

University of Nottingham; ICI Seeds

solids increased and fruit ripening delayed, processing characeristics altered fruit ripening altered

polygalacturonase sense and antisense from tomato

Zeneca

5

field trials

1994

United States


Hunt-Wesson

2

field trials

1994

United States

supressed polygalacturonase activity


Zeneca; Petoseed; HuntWesson Zeneca

1


1994

United States

1995

United States

1995

United States

1996

United States

1

field trial

1994

(Smith, et al., 1990, Smith, et al., 1990, Smith, et al., 1988, Taylor, et al., 1990)

3

United States

United States

United Kingdom

polygalacturonase level decreased


Zeneca; Petoseed

1


galactanase from tomato

Zeneca

2


Zeneca

1

field trial

1997

United States

Zeneca

1

1997

European Union

Zeneca

2

Approval under Directive 90/220/EEC field trials

1998

United States

Zeneca

1

ACNFP opinion as safe for processed food use

1998

United Kingdom

Improved processing characteristics

improved processing quality by reducing levels of polygalacturonase

partial polygalacturonase (PG) gene from tomato

71



decreased polygalacturonase activity

truncated partial sense polygalacturonase (PG) gene results in downregulation of endogenous PG gene

Zeneca

1

downregulated amount of polygalacturonase (PG) produced

truncated partial sense polygalacturonase (PG) gene results in downregulation of endogenous PG gene

Zeneca

1

Health Canada approval for food use SCF opinion opinion as safe for processed food use

1999

Canada

1999

European Union

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; Confined Research Field Trials, Plant Biosafety Office, Plant Products Directorate, Canadian Food Inspection Agency, accessed August 2004; GM Crop Database, AgBios, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 90/220/EEC, Belgian Biosafety Server, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 2001/18/EC, Belgian Biosafety Server, accessed January 2005; and published articles cited in the body of this table.

4.7.2

Control of leaf and flower wilting

The wilting of leaves and flowers follows some of the same biological mechanisms as does ripening and softening of fruit. In particular strategies have been pursued to control the production of ethylene or the sensitivity of the plant to ethylene.

Table 4.19 Transgenic plants with control of leaf and flower wilting Crop

Trait

Genes

Institution(s)

Country

leaf senescence delayed

Isopentenyl transferase from Agrobacterium

Cal West Seeds

1


Year


1999

United States

Cal West Seeds

1

field trial

2000

United States

Cal West Seeds

1

field trial

2001

United States

Cal West Seeds

1

field trial

2002

United States

Calgene Pacific

1

field trial

1992

Australia

Calgene Pacific

2

field trials

1994

Australia

Florigene

1

field trial

1994

Netherlands

Suntory; Florigene

1

field trial

1994

Carnation (Dianthus caryophyllatus)

increased vase life

ACC oxidase antisense from carnation

Florigene

1

field trial

1995

Netherlands

increased shelf life

ACC synthase

Florigene

1

1996

Australia

increased shelf life

ACC synthase

Florigene

1

GMAC environmental approval Approval under Directive 90/220/EEC

1998

European Union

improved vase-life

ACC; UDP-glucosyl transferase (ert1); phytoene desaturase (crt1); floral binding protein (fbp1) promoter

Hilverda Plant Technology; P. Kooij & Zonen; Van Staaveren

3

field trial

1997

Netherlands

improved vase-life

ACC; UDP-glucosyl transferase (ert1); phytoene desaturase (crt1); floral binding protein (fbp1) promoter

StaWest Carnation Group; West Select

2

field trial

1998

Netherlands

Sanford Scientific

1

field trial

1997

United States

Harris Moran

2

field trials

2001

United States

Harris Moran

2

field trials

2003

United States

senescence altered

Seminis

1

field trial

2000

United States

extended flower life

Monsanto

1

field trial

1997

United States

Monsanto

2

field trials

2000

United States

extended flower life

Lettuce (Lactuca sativa)

senescence altered


Translation initiation factor 5A from lettuce

Florigene

(Savin, et al., 1995)

Japan

delayed wilting

Geranium (Pelargonium)


ethylene gene antisense; ACC synthase gene; ACC oxydase gene

Count

long shelf life plants

cytokinin

University of Leeds


G1158; G1163

Ball Helix

1

field trial

2000

United States

ripening altered

Polygalacturonase antisense from tomato; Trypsin inhibitor from cowpea

Calgene

1

field trial

1989

United States


Isopentenyl transferase from Arabidopsis

University of Wisconsin

1

field trial

2000

United States

72

(Meyer, et al., 2001)

Australia

United Kingdom



Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; GM Crop Database, AgBios, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 90/220/EEC, Belgian Biosafety Server, accessed January 2005; and published articles cited in the body of this table.

4.7.3

Bruising or browning

The chemical reaction that causing browning in apples or potatoes is caused by the enzyme polyphenol oxidase. It appears to be fairly straightforward to eliminate such browning by reducing or eliminating expression of the gene that codes for that enzyme.

Table 4.20 Transgenic plants with controlled bruising or browning Crop

Trait

Genes

Institution(s)

Country

brown spot resisitant and polyphenol oxidase levels altered

Polyphenol oxidase from apple; Polyphenol oxidase antisense from apple

n.a.

2


Year


2002

United States

n.a.

1

field trial

2003

United States

n.a.

1

field trial

2004

United States

Polyphenol oxidase antisense from potato

ARS

1

field trial

1994

United States

Polyphenol oxidase from potato

ARS

2

field trials

1995

United States

ARS

2

field trials

1996

United States

ARS

2

field trials

1997

United States

ARS

2

field trials

1998

United States

ARS

2

field trials

1999

United States

ARS

1

field trial

2000

United States

CSIRO

1

field trial

1995

Australia

J.R. Simplot Company

1

field trial

2004

United States


blackspot bruise resistance

reduced browning

Polyphenol oxydase antisense

capable of growth on defined synthetic media and bruising reduced reduced polyphenol oxidase

polyphenol oxidase (PPO) antisense

bruising reduced Polyphenol oxidase from potato

bruising reduced

Polyphenol oxidase from potato; Polyphenol oxidase from tomato

bruising reduced

Count

Keygene; Wageningen University

(Bachem, et al., 1994)

Netherlands

Monsanto

2

field trials

1997

United States

Monsanto

34

field trials

1998

United States

Monsanto

11

field trials

2000

United States

Rutgers University

1

field trial

1997

United States

Rutgers University

1

field trial

1998

United States

University of Idaho University of Idaho University of Idaho University of Idaho

1 1 1 1


2001 2002 2003 2004



4.8


When thinking of improved product qualities, some of the first that come to mind include esthetic characteristics of a product, such as taste, scent, color, or size, as well as characteristics that lend to the convenience of using the product, such as fruit without seed or—as in one case listed here—grass that requires less chemical lawn care and less mowing.

73


4.8.1


Flavor and Scent

While much of flavor in plants products results from the sugars present and their balance with acids, most of the more distinctive flavor and aroma characteristics result from the complex mixture of volatile secondary metabolites present. Given the complexity of human sensory responses and the complexity of the chemistry involved, this arena of breeding and genetic modification is very much in its infancy; however, in a number of instances a single compound or class of compounds does present itself as the key to enhancing or eliminating a sensory response. Two areas of notable work have been in terpenoids, particularly the important flavor compound linalool, and in lipid-derived volatiles (Galili, et al., 2002). An example of the first is a tomato with increased linalool (Lewinsohn, et al., 2001) and an example of the second is a tomato with enhanced aroma from increased fatty acids (Binswanger, 1974). There are certainly others. For example, Japanese researchers have identified and successfully regulated the gene responsible for the chemical irritant that makes one ‘cry’ when cutting onions, thereby paving the way to a milder and more convenient to use onion (Imai, et al., 2002). In another example, researchers have been successful in identifying the compounds responsible for the astringent, beany flavor of soybeans, opening the way for more palatable and dairy-like flavors in soy products (Kinney, 2003).

Table 4.21 Transgenic plants with modified flavor or scent Crop

Trait

Country

disulfides reduced in endosperm, improves flavor in brewed beer

Coors Brewing

1


Year


1998

United States


phosphinothricin tolerance and flavor enhancer

Sunseeds

1

field trial

1996

United States

Onion (Allium cepa)

tearless onions

lachrymatory-factor synthase removed, responsible for producing propanthial S-oxide, the chemical irritant.

House Foods Corporation

Pineapple (Ananas comosus)

fruit sweetness increased

Mabinlin from Capparis masaikai


1

1997

United States


reduced grassy and beany off-flavors

down regulate lipoxygenases and hydroperoxide lyases from soybean

DuPont

(Kinney, 2003, Kitamura, 1995)

United States


improved flavor and ripening

array of ripening related genes from strawberry

Horticultural Research International

improved flavor and ripening

acyl carrier protein; caffeoyl-CoA3-O-methyltransferase; sesquiterpene cyclase; major latex protein; cystathionine synthase; dehydrin; an auxin-induced gene; a metallothionein-like protein from strawberry

INRA

(Lanjouw, et al., 1996)

France

key enzyme in production of fruit flavor

strawberry alcohol acyltransferase (SAAT) gene

Plant Research International, Wageningen

(Aharoni, et al., 2000)

Netherlands

enhanced aroma from increased Linalool

linalool synthase (LIS) gene from Clarkia breweri

Agricultural Research Organization

enhanced aroma from modified levels of flavor aldehydes and alcohols

alcohol dehydrogenase

CSIRO; Zeneca

enhanced aroma from increased fatty acids and fatty acid-derived flavor compounds

delta-9 desaturase from yeast

Rutgers University


Genes

Institution(s)

Count

(Imai, et al., 2002)

field trial

(Manning, 1998)

(Lewinsohn, et al., 2001)

(Prestage, et al., 1999, Speirs, et al., 1998) (Binswanger, 1974)


74

Japan

United Kingdom

Isreal

Australia; United Kingdom United States


4.8.2


Fruit, seed, or fiber color

Pigmentation of plants is often closely associated with important nutrients. The orange of carrots is caused by carotenoids and the reds of tomato by carotenoids and lycopene. Novel colors of fruit and vegetables, such as yellow watermelon and red bell peppers, have been introduced through the years from conventional breeding. This has not been an area of significant commercial interest in transgenic research, although a few early investigations were made in altering the pigmentation of cotton fibers and clarifying the juice of sugarcane. In some cases transgenic alteration of color can provide a simple means of identifying plants that contain the transgene.

Table 4.22 Transgenic plants with modified fruit, seed, or fiber color Crop

Trait


alteration of pigment production, insect resistance, and tolerance to glufosinate

Genes

Institution(s)

color altered Cotton (Gossypium hirsutum)

melanin produced in cotton fibers

copper transfer protein, apotyrosinase, copper binding protein and tyrosinase from Streptomyces antibioticus

Country

1


Year

Ciba-Geigy; INRA

Count

1995

France

Ciba-Geigy

1

field trial

1996

France

Monsanto

1

field trial

2002

United States

Calgene

1

field trial

1997

United States

Calgene

1

field trial

1998

United States


modified colour, sugar composition, flowering and fruit development

CSIRO

1

field trial

2002

Australia

Sugarcane (Saccharum officinarum)

increased sucrose content and reduced juice colour

CSIRO

1

field trial

1997

Australia


pigment composition altered and dwarfed

Transcription factors for DET1, COP, and HY5 genes from tomato silenced; Campesterol synthesis (DIM) gene from tomato

n.a.

8

field trials

2002

United States

enhanced pigmentation

delila (DEL) gene from Antiffhinum

John Innes Centre

pigment metabolism altered

seed color altered and fruit ripening altered

(Mooney, et al., 1995)

PetoSeed

2

field trials

1995

United States

PetoSeed

11

field trials

1996

United States

Seminis

2

field trials

1997

United States

Seminis

1

field trial

1998

United States

Purdue University

1

field trial

2001

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; and published articles cited in the body of this table.

4.8.3

Flower color

While altering or enhancing the color of other plant products may not be a great priority, it is in flowers. In particular, biotechnology has been used to develop colors in flowers that are not possible given the gene pool of those species. The most active area of research and development has been with anthocyanin biosynthesis, to produce blue shades in carnation and rose, other work has involved the accumulation of calcones to produce yellow colors (Mol, et al., 1999). Several transgenic blue mauve carnations had reached the regulatory stage in several countries and have been commercialized.

75

United Kingdom



Table 4.23 Transgenic flowers with modified color Crop

Trait

Begonia (Begonia semperflorens)

flower color altered

Carnation (Dianthus caryophyllatus)

altered flower colour, pigment production, violet color

modified flower color

Chrysanthemum (Chrysanthemum morifolium Ramat)

Genes

Institution(s)

Country

1


Year

Scotts

Count

2003

United States

Scotts

1

field trial

2004

United States

delphinidin gene

Calgene Pacific

2

field trials

1994

Australia

Delphinidin gene, dihydroflavonol reductase (DFR), lacZ, t-mas, D8 3', hf1, p-chsA flavonoid 3’, 5’-hydroxylase (F3’, 5’H) NADPHcytochrome P450 reductase; dihydroflavonol reductase (DFR)

Florigene Europe

1

field trial

1995

Netherlands

Florigene

1

GMAC approval

1995

Australia

Suntory; Florigene

2

field trials

1996

Japan

delphinidin gene

Suntory; Florigene

6

field trials

1997

Japan


flavonoid 3’, 5’-hydroxylase (F3’, 5’H) NADPHcytochrome P450 reductase; dihydroflavonol reductase (DFR)

Florigene

1


flavonoid 3’, 5’-hydroxylase (F3’, 5’H) NADPHcytochrome P450 reductase; dihydroflavonol reductase (DFR)

Florigene

1

flavonoid 3 ',5 '-hydroxylase

Florigene Suntory; Toyo University

1

novel flower color novel flower color and increased fragrance

flavanone 3-hydroxylase antisense

altered flower color

flower pigment modification

novel flower color

Hebrew University of Jerusalem; Technical University Munchen Weihenstephan Florigene

1

pigment biosythesis genes

Calgene Pacific

CHS and DFR sense and antisense

Florigene Europe

chalcone synthase (CHS) sense and antisense

DNA Plant Technology

Geranium (Pelargonium)

glyphosate tolerance and color altered

Lisianthus (Eustoma grandiflorum)

new flower colours/patterns and evaluation of phenotype and field performance novel flower color

NOS flavonoid gene

Orchid (Dendrobium)

flower color altered


Approval 1997 under Directive 90/220/EEC Approval 1998 under Directive 90/220/EEC field trial 2002 (Fukui, et al., 2003)

European Union

(Zuker, et al., 2002)

Isreal; Germany

European Union

Australia Japan

1992

Netherlands

1

Application under Directive 90/220/EEC (withdrawn) field trial

1993

Australia

1

field trial

1994

Netherlands

(Courtney-Gutterson, et al., 1994)

United States

Scotts Scotts

1 1


2001 2002


New Zealand Institute of Crop and Food Research New Zealand Institute of Crop and Food Research

1

field trial

1997

New Zealand

(Deroles, et al., 1998)

New Zealand

Flavonol 3-hydroxylase from petunia

University of Hawaii, Manoa

1

novel flower color

cytochrome P450 flavonoid 3',5'-hydroxylase from petunia

Calgene Pacific (Floragene); Suntory; INRA

novel flower color

dihydroflavonol 4-reductase (DFR) A1 gene from corn

Max-Plank-Institut fur Zuchtungsforschung; University of Tubingen Max Planck Institute for Plant Breeding Research, Köln Max Planck Institute for Plant Breeding Research, Köln Max Planck Institute for Plant Breeding Research, Köln

chalcone synthase (Chs) antisense

alteration of pigment production and flower color

A1 dihydroflavonol-4-reductase from corn

novel flower color

chalcone reductase from Medicago sativa

altered plant form or pigmentation

pigment metabolism altered

Dihydrofolate reductase from corn

glyphosate tolerance and color altered

field trial

1999

(Holton, et al., 1993)

(Meyer and Heidmann, 1994, Meyer, et al., 1987)

United States

Australia; Japan; France

Germany

1

field trial

1991

Germany

1

field trial

1996

Germany

1

field trial

1997

Germany

1

field trial

1999

New Zealand

Rogers

1

field trial

1992

United States

Scotts

2

field trials

2000

United States

Scotts

1

field trial

2001

United States

Scotts

2

field trials

2002

United States

Scotts

3

field trials

2003

United States

Scotts

2

field trials

2004

United States

New Zealand Institute of Crop and Food Research New Zealand Institute of Crop and Food Research

(Davies, et al., 1998)

novel flower color

malonyl-coenzyme A : anthocyanidin 3-Oglucoside-6 ''-O-malonyltransferase from dahlia flowers

Tohoku University; Institute Phys & Chem Research; Suntory; Minami Kyushu University

altered flower pigmentation

chalcone synthase antisense

Vrije University Amsterdam

(van der Krol, et al., 1988)

Netherlands

novel flower color

cytochrome b5 enzyme flavonoid 3',5'-hydroxylase from petunia

Vrije University Amsterdam

(de Vetten, et al., 1999)

Netherlands

76

(Nandi, et al., 2002)

New Zealand

Japan



Rose (Rosa hybrida)

altered flower colour

chalcone synthase gene; flavonoid 3'5'hydroxylase gene

Florigene

Torenia (Torenia fournieri Lind.)

novel flower color

chalcone synthase (CHS) or the dihydroflavonol-4reductase (DFR) gene in sense or antisense

National Research Institute for Vegetables Ornamental Plants & Tea; Suntory

2

field trials

1994

(Aida, et al., 2000, Aida, et al., 2000)

Australia

Japan

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; Table of Developing Transgenic Crop Plants in Japan (Field Tests and General Releases), Agriculture, Forestry, and Fisheries Research Council, Japanese Ministry of Agriculture, Forestry, and Fisheries, May 2003; GM Crop Database, AgBios, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 90/220/EEC, Belgian Biosafety Server, accessed January 2005; Notifications for placing transgenic plants on the EU Market under Directive 2001/18/EC, Belgian Biosafety Server, accessed January 2005; and published articles cited in the body of this table.

4.8.4

Size and morphology

For ages the size and morphology of crop products have been important breeding goals. So much has been achieved through conventional breeding that there is apparently little that can be added through transgenic approaches. Most of the innovations that mention changes in size or weight appear to be side effects from experimentation. This may, however, be an area of innovation that has simply not yet seen its time come.

Table 4.24 Transgenic plants with modified size or morphology Crop

Trait

Genes

Institution(s)

Country

seed size and weight increased

ADP glucose pyrophosphorylase from corn

University of Florida University of Florida

1 1

Action or reference field trial field trial

Year

Corn/Maize (Zea Mays)

1999 2001



2

field trials

2002

United States


Count

dwarfed and pigment composition altered

Transcription factor for DET1 gene silenced from tomato; Transcription factor for HY5 gene silenced from tomato; Transcription factor for COP gene silenced from tomato; Campesterol synthesis (DIM) gene from tomato

n.a.

8

field trials

2002

United States

seed size increased and larger fruit

Agamous-like gene 8 from Arabidopsis


1

field trial

1999

United States


1

field trial

2000

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004.

4.8.5

Seedlessness

Likewise, seedlessness or parthenocarpy, the formation of fruits devoid of embryos or endosperm (and thus seeds), has been a goal of much conventional breeding. It has been successfully achieved in several commercially important fruits and vegetables. Transgenic approaches have been taken in melons and tomatoes, perhaps simply to better understand the underlying genetic mechanisms, as little appears to be in development for market.

77



Table 4.25 Transgenic plants with seedless fruit Crop

Trait

Genes

Institution(s)

Country

parthenocarpic character

Tryptophan-2-monoxygenase synthesis

Istituto Sperimentale per l'Orticoltura, Sezione Ascoli Piceno

1


Year


Count

1999

Italy


glyphosate tolerance and parthenocarpy

Seminis Seminis Seminis

1 1 1


2000 2001 2002


Watermelon (Citrullus lanatus)

parthenocarpy

Seminis Seminis Seminis Seminis

1 1 1 1


2001 2002 2003 2004


Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004, and Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004.

4.8.6

Low maintenance landscaping

While this survey has, as a rule, excluded agronomic or input traits, such as herbicide tolerance, intended to reduce cost to farmers, the exception is made for turf grass because of the economics of this landscaping plant. Homeowners everywhere are, in a very real sense, grass farmers, employing tractors, fertilizers, and herbicides to cultivate and maintain millions of acres grass. Maintenance of herbicide resistant turf grass is expected to be significantly easier, in addition to using more environmentally benign and inexpensive lawn care chemicals to control weeds. Altered plant development and morphology, such as dwarfism, would help to reduce or even eliminate the need for mowing, with its attendant costs, labor requirements, fuel use, and pollution. Together these traits promise an increased level of convenience in maintaining landscaping by homeowners, as well as at schools, parks, and businesses with landscaped grounds, and, in particular, golf courses.

Table 4.26 Transgenic landscaping plants with lower maintenance requirements Crop

Trait

Creeping bentgrass (Agrostis stolonifera)

altered plant development and glyphosate tolerance tolerance to the herbicide glyphosate

Kentucky bluegrass (Poa pratensis)

altered plant development, morphology, and glyphosate tolerance

Genes

Institution(s)

5-enolpyruvylshikimate-3-phosphate synthase (epsps) gene from Agrobacterium

Count


Year

Country

2003

United States

1

FDA approval for incidental animal forage

2003

United States

Scotts

4

field trials

2003

United States

Scotts

1

field trial

2004

United States

Scotts

2

Scotts Seeds

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004, and GM Crop Database, AgBios, accessed January 2005.

4.9

Fiber and wood quality

Another entire subfield of plant science concerns itself with the chemical and physical properties of plant fibers and wood. As in other areas, transgenic analysis has proven a powerful tool for understanding the formation and composition of these economically important materials. At the same time it has led to strategies for modifying the chemical and physical characteristic,

78



particularly through the modification of lignin in the cell wall (Baucher, et al., 2003, CAI, 2004, McCaslin, 2001). 4.9.1

Fiber quality for textiles

An early innovation in cotton created a polyesterized fiber structure that was intended to improve cotton textiles. The work was, however, discontinued in the late 90s.

Table 4.27 Transgenic plants with improved fiber quality for use in texitles Crop

Trait


fiber strength and quality altered polyesterized fiber structure

Genes

Institution(s)

Parathion hydrolase from Flavobacterium

Agracetus Agracetus

enzymes for production of polyhydroxybutyrate (PHB) (PhaB and phaC genes) from Alcaligenes eutrophus

Count 1 6

Agracetus

Action or reference field trial field trials

Year

Country

1994 1995


(John and Keller, 1996)

United States

Agracetus

7

field trials

1996

United States

Agracetus

1

field trial

1997

United States

Monsanto

1

field trial

1998

United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004 and published articles cited in the body of this table.

4.9.2

Fiber quality for digestibility of animal feed and forage

A rapidly expanding area of work being pursued by a range of researchers in public sector organizations and small biotech firms is the improvement in the digestibility of animal feed and forage. A common strategy is reduction or modification of lignin in the fibers of the plant. A number of these innovations are in model plants, Arabidopsis and tobacco, although some field trials have been conducted in corn, alfalfa, bahiagrass, and tall fescue.

Table 4.28 Transgenic plants with improvied fiber quality for digestability of animal feed and forage Crop

Trait

Genes

Institution(s)

Arabidopsis

altered lignin

ferulate-5-hydroxylase or coniferaldehyde 5hydroxylase (F5H or Cald5H) from poplar

reduced lignin

cinnamoyl CoA reductase (CCR) antisense from Arabidopsis sinapyl alcohol dehydrogenase coded by Atcad-D (CAD) gene in Arabidopsis

INRA; Free University of Brussels; State University of Ghent INRA; CNRS

reduced lignin


reduced lignin

4-coumarate:coenzyme A ligase (4CL) antisense from Arabidopsis

altered lignin

ferulate-5-hydroxylase (F5H) enzyme (a cytochrome P450-dependent monooxygenase) in Arabidopsis

reduced lignin

reduced activity of p-coumarate 3-hydroxylase (C3H) enzyme (a cytochrome P450-dependent monooxygenase) encoded by reduced epidermal fluorescence (REF8) gene in Arabidopsis

reduced lignin

altered lignin biosynthesis

Count

Action Year or reference (Sibout, et al., 2002)

{Goujon, 2003 #1004

INRA; Natural Resources Canada

Country France; Belgium

France

(Sibout, et al., 2003)

France; Canada

(Lee, et al., 1997)

Canada; United States

University of British Columbia; Purdue University Purdue University; USDA-ARS, US Dairy Forage Research Center; University of Wisconsin Purdue University

(Marita, et al., 1999, Meyer, et al., 1998)

United States

(Franke, et al., 2002)

United States

cinnamoyl-CoA reductase (CCR) gene alteration in Arabidopsis

University of Manchester

(Jones, et al., 2001)

Caffeate O-methyltransferase from alfalfa

Forage Genetics International

2

field trials

2001

United States


1

field trial

2003

United States


1

field trial

2004

United States

79

United Kingdom



improved digestability - avoid cow bloating

Chalcone synthetase (ORF CHs A and CHS2Alf) sense and antisense

Instituto Nacional de Tecnología Agropecuaria (INTA)

alteration of lignin biosynthesis and improvement of digestibility

downregulation of cinnamoyl CoA reductase


reduced lignin

caffeic acid O-methyltransferase (COMT) encoding brown midrib (Bmr) gene

Purdue University

altered lignin

caffeic acid 3-O-methyltransferase (COMT) caffeoyl coenzyme A 3-O-methyltransferase (CCoAOMT)

Samuel Roberts Noble Foundation; USDA-ARS, US Dairy Forage Research Center; University of Wisconsin

cell wall altered cell wall altered altered lignin biosynthesis

Bahiagrass (Paspalum notatum)

lignin levels decreased


digestibility improved


alteration of lignin biosynthesis and tolerance to glufosinate

Tall Fescue (Festuca arundinacea)

field trials

Belguim

1994

(Bout and Vermerris, 2003)

United States

(Guo, et al., 2001, Marita, et al., 2003)

United States


1

field trial

1999

United States

University of Wisconsin

2

field trials

1998

United States

W-L Research

1

field trial

1998

United States

W-L Research

1

field trial

2000

United States

O-methyltransferase from sorghum

University of Florida University of Florida University of Florida

1 1 1


2001 2003 2004


Ventria Bioscience Ventria Bioscience

1 1


2002 2003


Biogemma

1

field trial

2002

France

Biogemma

1

field trial

2003

France

Du Pont - Pioneer

2

field trials

1999

United States

(Chabbert, et al., 1994, Vignols, et al., 1995)

France; Spain

F5H ferulate-5-hydroxylase antisense from Zea mays caffeoyl-coenzymeA O-methyl transferase antisense from Zea mays

increased digestability from modified lignin content and composition

caffeic acid 3-O-methyltransferase (COMT) brown midrib 3 (bm3) mutation

Institut National Agronomique; INRA; Centro de Investigacion y Desarrollo, Consejo Superior de Investigaciones Cientificas

alteration of forage quality, alteration of lignin biosynthesis, and tolerance to glufosinate

downregulation of cinnamoyl CoA reductase

SES Seeds

2

field trials

1996

Belgium

SES Seeds

2

field trials

1997

Belgium

increased digestability from modified lignin content and composition

cinnamyl alcohol dehydrogenase (CAD) expression of brown midrib 1 (bm1) mutation

University of Dundee; Zeneca; Jealotts Hill Research Station; INRA; IACR, Long Ashton Research Station

lignin levels decreased and lignin levels decreased

O-methyltransferase from sorghum


modified lignin for improved digestability

cinnamyl-alcohol dehydrogenase (CAD)

Dr. R. Maag Ltd.(Roche); FOM Institute of Atomic and Molecular Physics, Unit for Mass Spectrometry of Macromolecular Systems

modified lignin for improved digestability

caffeic acid-O-methyltransferase (COMT)

Purdue University

hypolignified


Cinnamyl alcohol dehydrogenase from Tall Fescue; Caffeate O-methyltransferase antisense from Tall Fescue

improved forage digestability

down-regulation of cinnamyl alcohol dehydrogenase

alteration of lignin biosynthesis and tolerance to glufosinate


Argentina

Caffeate O-methyltransferase antisense from alfalfa; Caffeoyl CoA O-methyltransferase antisense from alfalfa Caffeate O-methyltransferase from alfalfa

cell wall altered


2


1

field trial

1998

(Pillonel, et al., 1991)

(Bout and Vermerris, 2003)

United Kingdom; France

United States

Switzerland; Netherlands

United States

Biogemma

1

field trials

2003

France

Samuel Roberts Nobel Foundation

1

field trial

2001

United States

Samuel Roberts Nobel Foundation Samuel Roberts Noble Foundation Samuel Roberts Nobel Foundation

1

field trial

2002

United States

(Dechenaux, et al., 2003)

United States

2

field trials

2003

United States

RAGT (Rouergue Auvergne - Gévaudan Tarnais)

1

field trial

2000

France

altered lignin content and composition

phenylalanine ammonia-lyase (PAL) from tobacco

Salk Institute; Hebrew University of Jerusalem; Samuel Roberts Nobel Foundation; John Innes Center

reduced lignin

caffeic acid O-methyltransferase (COMT)

University of Strasbourg; INRA; Samuel Roberts Nobel Foundation

80

(Halpin, et al., 1998)

(Elkind, et al., 1990, Korth, et al., 2001, Mertz, et al., 1964, Sewalt, et al., 1997)

United States; Isreal; United Kingdom

(Atanassova, et al., 1995, Ni, et al., 1994)

United States




cinnamyl alcohol dehydrogenase (CAD)

Zeneca; Jealotts Hill Research Station; University of Toulouse


cinnamate 4-hydroxylase (C4H)

Samuel Roberts Nobel Foundation; Salk Institue; University of London; Washington State University; Zeneca


caffeoyl-coenzyme A O-methyltransferase (CCoAOMT) antisense

University of Georgia; USDA-ARS; INRA; University of Strasbourg

altered lignin

ferulate 5-hydroxylase (F5H) from Arabidopsis

Purdue University


4-coumarate:coenzyme (4CL) antisense

Mitsubishi Paper; Tokyo University; Japan Forestry & Forest Products Research Institute


cinnamoyl-CoA reductase (CCR) sense and antisense

alteration of lignin biosynthesis

downregulation of cinnamoyl CoA reductase (CCR)

CNRS; University of Toulouse; INRA; Zeneca; USDA-ARS, US Dairy Forage Research Center; University of Wisconsin; University of Strasbourg; University of Dundee; Jealotts Hill Research Station; UPS; Ctr Tech Ind Papiers Cartons & Celluloses Centre de biologie et physiologie végétales Centre de biologie et physiologie végétales

(Halpin, et al., 1994)

United Kingdom; France

(Blee, et al., 2001, Sewalt, et al., 1997)

United States

(Pincon, et al., 2001, Zhong, et al., 1998)

United States; France


United States

(Kajita, et al., 1997, Kajita, et al., 1996)

(Aharoni, et al., 2000, Pincon, et al., 2001, Piquemal, et al., 2002, Ralph, et al., 1998)

Japan

France; United Kingdom

1

field trial

1996

France

1

field trial

1997

France

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; BioTrack Database of Field Trials, OECD, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; and published articles cited in the body of this table.

4.9.3

Wood quality for pulp

The same biosynthetic pathway for lignin that is being modified to improve fiber digestibility for animals is also being tested to improve fiber digestibility for a very different animal: paper pulping plants. Lower lignin levels could reduce the energy expended for mechanical shredding and the amount of chemicals used to separate lignin from cellulose in the process of pulping for paper manufacture.

Table 4.29 Transgenic trees with improved pulping characteristics Crop Aspen (Populus tremuloides)

Trait low lignin levels and modified reactivity (S/G ratio)

Genes 4-coumarate-CoA ligase (4CL) antisense and coniferaldehyde 5-hydroxylase (CAld5H)

Institution(s) Michigan Technological University; USDA-ARS, US Dairy Forage Research Center; University of Wisconsin; Iowa State University

Count Action Year (Hu, et al., 1999, Li, et al., 2003)


better quality cellulose

celluose synthase

Michigan Technological University

(Joshi, et al., 2003)

United States

Eucalyptus (Eucalyptus grandis)


Pine (Pinus sp.)

reduced lignin

ArborGen ArborGen cinnamyl alcohol dehydrogenase (CAD) mutation


North Carolina State University; USDA-ARS, US Dairy Forage Research Center; University of Wisconsin ArborGen ArborGen

81

3 2

2003 2004


(MacKay, et al., 1997, Ralph, et al., 1997)

United States

1 4



2003 2004



Poplar (Populus sp.)


cinnamyl alcohol dehydrogenase (CAD); caffeic acid O-methyltransferase (COMT)


downregulation of cinnamoyl CoA reductase (CCR); downregulation of caffeic acid o-methyl transferase (COMT) cinnamyl alcohol dehydrogenase (CAD) cinnamoyl CoA reductase (CCR) caffeic acid o-methyl transferase (COMT)


Sweetgum (Liquidambar styraciflua)


Institut National Agronomique; Centre Technique du Papier; Centro de Investigacion de Empresa Nacional Celulosas; INRA; University of Gent; Syngenta INRA

INRA

(Baucher, et al., 1996, Jouanin, et al., 2000, Lapierre, et al., 1999, Pilate, et al., 2002)

France; Spain; Belgium; United Kingdom

1

field trial

1999

France

1

field trial

2003

France

reduced lignin content and altered composition

ferulate 5-hydroxylase from Arabidopsis

Purdue University


United States

altered lignin biosynthesis and male sterility/fertility

downregulation of cinnamoyl CoA reductase (CCR)

Station d'Amélioration des Arbres Forestiers (SAAF) Station d'Amélioration des Arbres Forestiers (SAAF) Station d'Amélioration des Arbres Forestiers (SAAF)

1

field trial

1995

France

1

field trial

1996

France

1

field trial

1997

France

1

field trial

1998

United States

cell wall altered

Cellulose binding protein from Clostridium cellulovorans

Union Camp

reduced lignin content and altered composition

caffeoyl-coenzyme A O-methyltransferase (CCoAOMT)

University of Ghent; INRA; Katholieke Univ Leuven; USDA-ARS, US Dairy Forage Research Center; University of Georgia; USDA-ARS, Richard B. Russell Agricultural Research Center

alteration of lignin biosynthesis

downregulation of cinnamoyl CoA reductase (CCR); downregulation of caffeic acid o-methyl transferase (COMT)

Zeneca

1

field trial

1995

United Kingdom

Zeneca

1

field trial

1996

United Kingdom

ArborGen

1

field trial

2003

United States


(Meyermans, et al., 2000, Zhong, et al., 2000)

Belgium; France; United States

Data sources: USDA-APHIS Environmental Releases Database, Information Systems for Biotechnology (ISB), accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 90/220/EEC, Joint Research Centre, European Commission, accessed August 2004; Deliberate releases into the envornment of GMOs under Directive 2001/18/EC, Joint Research Centre, European Commission, accessed August 2004; and published articles cited in the body of this table.

4.10 Environmental quality traits While a number of the traits listed previously have beneficial environmental attributes, such as reduced phosphorus levels in animal waste or less toxic chemicals for lawn maintenance, those attributes are really happy side effects of the primary trait characteristic. However, at least one class of transgenic plants is entirely intended for environmental cleanup. A range of genetic mechanisms have been found to have the effect of increasing plants’ ability to absorb toxic materials from soil. Alterations in the production of the plant hormone ethylene enable absorption of arsenic and heavy metals. Enzymes that capture and modify mercury ions and other metals enable plants to sequester of heavy metals. This area is also quite young and is primarily a venue of public sector researchers and small biotech firms. Table 4.30 Transgenic plants for bioremediation of soil toxins Crop

Trait

Brassica

heavy metal bioremediation


phytoremediation of arsenate

Genes

Institution(s) ARS

ACC deaminase from bacterium E. cloacae

82

Alberta Research Council; University of Waterloo

Count 1


Year

Country

2003

United States

(Nie, et al., 2002)

Canada


Poplar (Populus)


phytoremediation of soils and removal of heavy metals phytoremediation of soils and removal of heavy metals

Albert Ludwigs University, Freiburg Albert Ludwigs University, Freiburg

1

field trial

2002

Germany

1

field trial

2003

Germany


Applied PhytoGenetics

2

field trials

2003

United States


Mercuric ion reductase from E. coli


1

field trial

2001

United States

halogenated hydrocarbons metabolized

Cytochrome P450 from human

University of Washington

1

field trial

1999

United States

Rice (Oryza sativa)


Mercuric ion reductase from E. coli; Organomercury lyase from E. coli

Applied PhytoGenetics

1

field trial

2003

United States



Mercuric ion reductase from E. coli


1

field trial

2001

United States

heavy metals sequestered

Metallothionein from mouse

University of Kentucky University of Kentucky

1 1


1989 1990


heavy metal accumulating

ACC deaminase from bacteria E. cloacae

University of Waterloo


(Grichko, et al., 2000)


83

Canada


5


Commercialization of nutritional and product quality traits

The previous chapters have provided a snapshot of the entire R&D pipeline, from early stage research through regulatory filings. We turn now to examine the end of the pipeline that leads to commercialization, identifying quality traits that have been commercialized and pointing to others that various organizations are preparing for commercial release within the foreseeable future. As we have explained, the future output of any industry’s R&D pipeline can be quite uncertain; thus, we proceed here in three parts. First we look at crop quality innovations resulting both from conventional breeding and biotechnology that have already been commercialized. We then look at those transgenic nutritional and quality innovations likely to be released within the next five years. Third we take a longer view, and with greater margin for error, take some educated guesses at transgenic product innovations that may be ready for the market in five to ten years. Because of the additional level of insights needed, this analysis is based upon in depth investigation of public announcements, company reports, and in person interviews with researchers at some of the leading public and private sector R&D organization in agricultural biotechnology. Because the level of detail provided by these sources was often significantly different than the level of detail achieved in Chapter 4, there is not a one-to-one correspondence in every case between an ‘individual product innovation candidate’ identified in Chapter 4 and an actual product slated for commercialization in this chapter. Some of the single lines of innovation identified in Chapter 4 we see yielding multiple products.

5.1

Crop quality innovations already commercialized

Transgenic crops with improved quality characteristics can be expected to follow many of the same patterns and precedents set by crop quality innovations achieved through breeding programs. And, in some cases crops with improved quality characteristics will in fact compete directly with those conventionally bred crop quality innovations. For both reasons it is helpful to have some sense of context and what kinds of quality innovation have been going on more generally. We then proceed by considering brief case studies of the five transgenic product quality innovations that have already been commercialized, in order to establish any precedents or lessons learned for future product quality innovation. 5.1.1

Conventional quality innovations already commercialized

Some of the most recent manifestations of post harvest quality improvements from conventional breeding programs can be found in Table 5.1, which lists several recently released varieties— primarily affecting changes in soybean oils and corn carbohydrates. These varieties were developed through conventional breeding methods to cross in either naturally occurring or induced gene mutations. Most of the innovations listed in Table 5.1 represent improvements in specific characteristics that are likewise being developed or extended through transgenic approaches, as we have seen in the previous chapter. As such, they demonstrate both the feasibility and the challenges of introducing such traits into today’s commodity markets. They

84



can even represent, in some markets, potential competition to the transgenic varieties being designed to deliver similar characteristics. Several public sector breeding programs—in Ontario, Ohio, and Iowa—and at least two commercial partnerships—Pioneer/Bunge and Monsanto/Cargill—have introduced soy varieties with low levels of linolenic fatty acid—reduced from an average of about 7 percent in conventional soybean oil to between 1 and 3.5 percent (Table 5.1). The linolenic fatty acid constituent of vegetable oils is a source of instability and is thus one of the primary reasons for the partial hydrogenation of such oils prior to their use in manufactured food products. While hydrogenation has allowed vegetable oils like soy to substitute for saturated fats, hydrogenation causes the formation of trans fats, which are at least as bad for cardiovascular health as saturated fats. As a result, the FDA is changing food nutritional content labels starting in 2006, requiring that trans fats be listed in the same way as saturated fats. These labeling requirements are expected to induce a major shift by food manufacturers to alternative oils, in order not to list a high trans fat content, which consumers are expected to avoid. As a result, efforts have been underway at both agricultural research universities and the major seed companies to provide new varieties that will help soy growers to meet these impending changes in oil demand. Low linolenic soy is expected to be a demand-shifting product quality innovation (discussed in section 7.1) with users distinctly valuing the new characteristic over existing alternatives in the market. A suite of hybrid corn varieties introduced under the IndustrySelect™ program 5 by Pioneer in 2003 primarily offer improved carbohydrate characteristics, including higher energy—either in animal metabolism with high available energy (HAE) hybrids or in ethanol fermentation with high total fermentables (HTF) hybrids—and higher starch components—either high extractable starch (HES) hybrids or waxy (WX) hybrids with higher amylopectin content. The company’s plan is to use each of these as a starting point for a line of new varieties that will offer increasing quality improvements, eventually combining transgenic innovations with these conventionally bred traits. HAE can clearly be useful on farm by operations that grow their own feed and would not fully require special handling at the elevator to capture the additional value. The other varieties, however, require more extensive supply chain management. Being optimized to specific off farm uses, they require that the grower be able to locate a customer who will be willing to pay an associated premium for the harvest. Growers of these varieties need to be linked into an identity preservation program, to grow under contract for such a user. In addition, to support the market for these value added traits, Pioneer has also made a testing tool available —a whole-grain Near Infrared (NIR) rapid assay—for grain handlers and processors to be able to readily identify and quantify these traits. High energy and high starch are input-use efficiency enhancing product innovations: users in livestock production or corn processing value these new characteristic because they enhance efficiency, reduce costs, or increasing revenues. While final consumers will not see any qualitative difference in the final products that result from these inputs, they may see reduced prices, depending on market conditions.

5

See http://www.pioneer.com/media/industryselect/ .

85



Table 5.1 Recent examples of nutritional and quality-enhanced crop varieties using conventional breeding and genetic mutation, by release date Product

Qualities/Traits

Benefits

Uses

Organization

Release year

Low linolenic OT96-15 soy variety

Conventionally bred variety with lower proportion of linolenic fatty acid (~3.4%).

A more stable soybean oil that has less need for hydrogenation and thus reduces or eliminates trans fats.

Substitute for partially hydrogenated soybean oil in food products and claim of ‘low/no trans-fats’ on label

Agri-Food Canada and Guelph University 6, (distributed by SeCan)

2001

Low linolenic HS96-3818 soy variety

Conventionally bred variety with half the usual proportion of linolenic fatty acid (~3.5%).



Ohio State University 7

2002

IndustrySelect™ High Available Energy (HAE) Corn, version 1.0 IndustrySelect™ High-Total Fermentables (HTF) Corn

Conventional hybrid with higher metabolizable energy profile.

Higher nutritional and caloric value per bushel.

Monogastric animal feed

Pioneer-DuPont 8

2003

Conventional hybrid with greater carbohydrate content

Up to 4% greater ethanol yield.

Dry-grind ethanol production

Pioneer-DuPont 9

2003

IndustrySelect™ High Extractable Starch (HES) Corn for Wet Milling

Conventional hybrid with higher starch content

2% greater yield of extactable starch.

Wet milling for food ingredients such as high fructose corn syrup

Pioneer-DuPont 10

2003

IndustrySelect™ Waxy (WX) Corn for Wet Milling

‘Waxy’ mutant variety

Yields almost 100% amylopectin starch .

Wet milling for food ingredients such as thickeners, stabilizers, and emulsifiers

Pioneer-DuPont 11

2003

6

http://www.agbios.com/dbase.php?action=ShowProd&data=OT96-15 http://www.agriculture.purdue.edu/AgAnswers/story.asp?storyID=3056 8 http://www.pioneer.com/media/industryselect/products/hae.pdf 9 http://www.pioneer.com/media/industryselect/products/htf.pdf 10 http://www.pioneer.com/media/industryselect/products/hes.pdf 11 http://www.pioneer.com/media/industryselect/products/hes.pdf 7

86



1% linolenic soybean varieties IA2064 and IA3017

Conventional bred varieties that combine fan1, fan2, and fan3 genes for reduced proportion (1%) of linolenic fatty acids.



Iowa State University 12

2003-2004

Vistive™ low linolenic soy variety

Conventionally bred variety with lower proportion of linolenic fatty acid (< 3%).


Monsanto and Cargill 13

2004

Low linolenic soy variety 93M20

Conventionally bred variety with lower proportion of linolenic fatty acid (< 3%).


Substitute for partially hydrogenated soybean oil in food products and claim ‘low/no trans fats’ on label Substitute for partially hydrogenated soybean oil in food products and claim ‘low/no trans fats’ on label

Pioneer-DuPont and Bunge 14

2005

12

http://www.notrans.iastate.edu/ http://www.monsanto.com/monsanto/us_ag/content/enhanced_value/vistive/ 14 http://www.pioneer.com/media/knowhow/soybeans/soybean_oil.htm 13

87


5.1.2


Transgenic quality innovations already commercialized

In contrast to these breeding innovations in major commodities, the five transgenic quality traits that have already been commercialized to date (Table 5.2) have targeted specialty products— fresh and processed tomatoes, specialty oils, cut flowers, and a specialty cigarette tobacco. Only one of these five products was introduced by a firm that could even begin to be considered an incumbent player in the market for the new product. Yet even Vector Tobacco (Liggett) has only a 3 percent market position behind the leading firms in tobacco products (Davis, 2003). The common theme running through these five transgenic product introductions is that of a visionary entrepreneur seeking to take advantage of a significantly novel product characteristic, resulting from a fairly straightforward single-gene intervention, to attempt penetrating a high margin market. Such ‘low hanging fruit’ have included the FlavrSavr™ tomato by Calgene (now owned by Monsanto) and a processing tomato developed by Zeneca Plant Sciences (now merged into Syngenta), that were released in 1994 and 1995 respectively. Both of these innovations blocked expression of polygalacturonase (PG), an enzyme involved in the breakdown of pectin in the fruit cell wall and thus instrumental in the softening of the fruit. (See section 4.7.1 and Table 4.18.) The intention of these innovations was to use the slower softening characteristic to drive entry in high quality fresh and processing tomatoes, to capture market share as well as eventually to extend markets by meeting unmet latent demand. These two firms differed in the target market they were seeking to influence given the dual characteristic of the trait. Calgene was targeting final consumers with the demand enhancing characteristics of a fresher, better looking, better tasting tomato while Zeneca was targeting processors with the input efficiency enhancing aspect of the trait, being primarily less loss to waste and a more optimal profile of fruit characteristics, including pectin levels, for processing. Yet, neither Calgene nor Zeneca had experience in the food industry, and many of the problems that led to the eventual withdrawal of these two products can be attributed to this fact. These markets are highly dynamic with multiple factors constantly shifting the competitive landscape and many suppliers jostling for position. Extended shelf life innovations of this type have a complex impact on the value chain, and thus require considerable value chain coordination. In fresh markets, where Calgene’s tomato was sold, extended shelf life can have both a demand effect, as consumers encounter a more attractive and flavorful product, and an input-use efficiency effect, as less of the product is lost to waste in shipping and retail. In processing markets, where Zeneca’s tomato was sold, extended shelf life has an input-use efficiency effect for processors and the potential for translating into a more competitively priced final product for the consumer. Calgene also introduced an oilseed rape variety in 1994, called Laurical™, with a modified oil composition that suited it for uses in confectionary foods and personal care products. This variety was developed by inserting a gene for a thioesterase enzyme, which increased the biosynthesis of lauric and myristic fatty acids. The variety was inherited by Monsanto upon its acquisition of Calgene in the late 1990s. After running large numbers of field trials with modified oils in 1998 and 2000, Monsanto has reduced its attention to this trait in the years since. (See section 4.3.) This high laurate vegetable oil reportedly remained on the market for some time but did not enjoy success and was subsequently discontinued. Since it largely substituted for other oils such as palm and cocoa butter, it may be characterized as a demand affecting innovation, to the extent that it enables consumers to avoid saturated fats. It may also have an input-use efficiency effect, but only to the extent that it provides a more reliable or lower cost supply to food manufacturers.

88



The first genetically modified flower to be marketed was the mauve colored Moondust™ carnation, commercialized in Australia in 1996 by Florigene, a company founded in Melbourne in 1986 to apply the emerging tools of plant biotechnology to floriculture. A darker colored Moonshadow™ followed in 1997. Four more shades have since been added to Florigene’s Moonseries™; all achieved by adding a gene for production of the blue pigment delphinidin to an otherwise colorless or white variety of carnation. (See section 4.8.3.) The earliest field trials on record were done in Australia in 1994, in the Netherlands in 1995, and in Japan in 1996. Regulatory approvals were obtained in Australia in 1995, and in the EU in 1997 and 1998 (Table 4.23.) It may indeed be the most successful quality trait innovation from plant biotechnology to date, with unit sales near 10 million carnations in 2003 (Trends in Japan, 2004) generating revenues of $2.1 million (CNN, 2003). Demand for novel colored flowers seems particularly strong in Japan, perhaps explaining Suntory’s acquisition of Florigene in 2003. Novel flower color is a clear example of an innovation with a demand effect: consumers see and respond to the new characteristic of the product. Marginal cost of supplying the product is essentially unchanged. The tobacco market’s equivalent to decaf coffee, called Quest™, was introduced by Vector Tobacco in 2003 to test markets in several Great Lakes states and later in Arizona. The technology, which suppresses a key step in nicotine biosynthesis in tobacco, arose out of research at North Carolina State University. Executives at Liggett first took an interest in the technology in 1997 and funded further research at the university, which resulted in a technical breakthrough in 2000 that virtually eliminated nicotine from the tobacco plant (Davis, 2003). Vector Tobacco Inc. had meanwhile been formed as a subsidiary of Liggett in 1999, and initiated its own field trials of the reduced nicotine tobacco in 2001, and Vector’s transgenic tobacco was approved and deregulated by the USDA in 2002. (See Table 4.15.) Vector launched the Quest™ product in 2003. Specifically, Quest™ offers three decreasing levels of nicotine content, the third of which is essentially zero. The firm claims that this allows a smoker to reduce or eliminate their exposure to nicotine while continuing the familiar social ritual of smoking. However, until Vector is able to get FDA approval, no health claims can be made concerning any potential role of this cigarette as a smoking-cessation product. The reduction of nicotine is, in theory, a demand affecting innovation, with consumers willing to pay to continue smoking while reducing some of its negative consequences. Others are likely to value the product precisely as a tool to quit, possibly in combination with smoking cessation products like the nicotine patch, in a manner that would allow them to phase out the addiction before giving up the familiar ritual.

89



Table 5.2 The five transgenic nutritional and quality enhanced products already commercialized, by release date Commercial launch 1994 in the US (discontinued in 1997)

Product

Qualities/Traits

Benefits

Uses

Organization

FlavrSavr™ fresh tomato

Decreased level of polygalacturonase, an enzyme that breaks down fruit pectin. This slows fruit softening and prolongs shelf life while other ripening characteristics, such as color and flavor, continue to develop.

Prolonged shelf life allows later harvest and thus more time to ripen in the field, and longer window for ripening after harvest, resulting in improved appearance and flavor of fresh tomatoes.

Fresh market tomato.

Calgene

1994 (discontinued)

Laurical™ high laurate rapeseed oil

Fatty acid content modified to increase levels of lauric and myristic acids

Expands the range of uses of canola oil and replaces some imports of tropical lauric oil with domestic production

Manufacture of soaps and detergents; substitute for cocoa butter in confectionary foods.

Calgene/ Monsanto

1995 in the UK (discontinued in 1999)

Processing tomato

Decreased level of polygalacturonase, enzyme that breaks down fruit pectin. This increases the ratio of solid to liquid contents in the fruit.

Improved quality for processors. Transporting and processing more ‘tomato’ and less water.

Canned tomato paste.

Zeneca

1996 in AsiaPacific

Moonseries™ colored carnations

Blue-mauve series of colors

Novel colors for carnation. Offers new choices to customers and new uses for carnation.

Ornamental, including cut flowers, centerpieces, and bouquets.

Florigene/ Suntory

2003 in regional U.S. test markets

Quest™ nicotine-free and reduced nicotine cigarettes

Reduced nicotine content by blocking production of a key tobacco enzyme that produces nicotine; three grades offer nicotine levels that range from moderate, at 0.6 mg, to essentially zero, at less than 0.05 mg (compared to standard cigarettes at 1.0 mg)

Reduces or removes the addictive component of cigarettes, also reduces levels of tobacco-specific nitrosamines (TSNA’s) a potent carcinogen in tobacco smoke.

Allows smokers to reduce nicotine intake while “still enjoying the ritual of smoking”; may be allowed to be promoted as a smoking-cessation product.

Vector Tobacco/ Liggett

90


5.2

Transgenic nutritional and product quality innovations likely to be commercialized in the next five years

Looking forward five years we see a convergence in the use of biotechnology and other breeding methods to advance quality innovation in both major market products like feed corn and food ingredients as well as specialty market products like fresh fruit and cut flowers. The tools of plant biotechnology are being increasingly employed to continue those trends already observed. In addition, beta-carotene enhanced GoldenRice, a first example of the use of biotechnology for humanitarian interventions against malnutrition, may be forthcoming within this timeframe. Table 5.3 provides a schedule, in expected chronological order, of nutritional and quality innovations slated to be released over the next five years. These have been identified through our interviews with company representatives, various industry publications, as well as our survey of innovations in the previous chapters. Three criteria were used to place innovations in this final phase of the R&D pipeline: (1) if a qualified representative or a recent official publication of the innovating organization identified that product as forthcoming within the next five years; (2) if informed secondary sources indicated the product to be forthcoming, as cited in the table. (3) if applications have been submitted for regulatory approval. 5.2.1

Soy consumer traits and oil quality

The largest area of advance in genetic quality likely to be noticed by food consumers will be in soy. Almost a dozen different quality enhanced soy products will be forthcoming in the next five years. These include varieties with high sucrose and low rafinosaccharides, varieties with reduced volatiles and ‘off’ flavors, and varieties with functional protein characteristics like improved solubility and texture. Such soybeans are aimed at enhancing the appeal of soy products to mainstream food consumers who may be substituting away from dairy products and into soybased products for health reasons. (For details, see Table 5.3.) One new soy variety specifically provides increased levels of specific soy proteins associated with some of the established positive health effects of soy, such as the lowering of blood cholesterol. Other forthcoming transgenic soy varieties have improved oil profiles—reducing the levels of unstable fatty acids like linolenic acid, increasing levels of stable fatty acids like oleic acid, or both. Greater stability reduces or eliminates the need for hydrogenation of soybean oils, the process that creates trans-fats. Such soybean oils with greater stability are intended to give food processors an option for eliminating trans-fats from a wide range of snack foods and baked goods. The two key innovators of these soy products are DuPont-Pioneer and Monsanto, in collaboration respectively with commodity handlers Solae (a joint venture of DuPont and Bunge) and Renessen (a joint venture of Monsanto and Cargill). These alliances are preparing to manage the identity preservation of these quality added soybean products, from seed production, to grower relations, to handling, processing, and distribution to major customers. Consumer acceptance, regulatory, and trade issues of such quality differentiated soy products should be minimal. Some of these traits are the result of conventional breeding and mutations, which require no additionally

91


regulatory approvals. Other, transgenic traits will be combined with the Roundup Ready trait, which is already popular among soybean growers, and thus the varieties will already be designated as genetically engineered crops. The new quality traits, once themselves approved, thus should not fundamentally affect regulatory status and thus the management of resulting product. On the other hand, the new quality traits should have increased appeal among food manufacturers and consumers. Of course, being identity preserved to capture their value, these varieties and their products will find it relatively easy to comply with traceability and labeling requirements in jurisdictions such as Europe. 5.2.2

Feed and processing traits in corn/maize

The other major market quality innovations forthcoming in the next five years are in corn. Up to six quality enhanced varieties are expected in the next five years. These include transgenic traits that improve efficiency in animal nutrition, food processing, and ethanol production. Monsanto has filed for regulatory clearance in the U.S. of a high lysine variety, and EMBRAPA, the Brazilian agricultural research agency, is getting close on a high methionine variety, with both of these providing higher levels of an essential amino acid for improved animal nutrition (also see Table 4.3). Syngenta reports plans to commercialize varieties that express the enzyme phytase, used as feed nutrition supplement that improves animals’ absorption of minerals like phosphorus, and the enzyme amylase, used in ethanol production 15. DuPont-Pioneer has plans to release next versions in their IndustrySelect™ product lines within two to five years, including a corn hybrid improved for wet milling, one with higher yields of extractable starch, one with higher yields of an oil with high oleic acid composition, and another HAE variety for animal feed with both higher oil content and improved fiber digestibility. (For more details see Table 5.3.) 5.2.3

Horticultural specialty products

The other innovations likely to be forthcoming in the next five years are in horticultural crops providing specialty products—fresh and processed fruits, cut flowers, a turf grass for landscaping, and a potato starch for paper manufacturing. These thus follow more closely after the pattern of transgenic product quality innovations already commercialized, seeking to enter smaller markets with higher margins, and attracting consumers with a particularly distinct change in an important quality characteristic (discussed on page 88). Improvements in freshness and shelf life are underway for several types of fruit:

15 16

•

A small biotechnology firm in British Columbia, called Okanagan Biotechnology Inc. (OBI), reports its readiness to commercialize a range of apple varieties in which the browning that quickly follows when apples are cut or bruised is eliminated 16. This characteristic enables ready-cut apple snacks and other processing uses, as well as improving the shelf life and appearance of fresh market whole apples.

•

Syngenta is preparing to commercialize a long shelf life banana that will be able to last up to five days longer than conventional bananas.

http://www.syngenta.com/en/ar2003/main/science.aspx http://www.okanaganbiotechnology.com/non-browning-apple.php

92


Table 5.3 Nutritional and quality enhanced products likely to be commercialized in the next five years Estimated launch or release date 2005-2006 *

Product

Qualities/Traits

Benefits

Uses

Organization

High sucrose soy

A non-transgenic product based on a gene mutation resulting in high sucrose content and reduced rafinosaccharides

Improves flavor and functionality; reduces flatulence; could encourage wider use of soy (with associated health benefits like lower heart disease risk and healthy weight maintenance)

Ingredients for current soy based products

Pioneer-DuPont / Solae

2005 *

Non-browning apple

Silencing of the polyphenol oxidase (PPO) gene in the apple fruit

Slows or stops the browning of apple fruit. Fruit will not go brown when bruised or cut. Replaces need for preservatives to prevent browning. Reduced wastage of damaged and discolored fruit. More attractive apples on market shelf.

Fresh fruit. Ready-cut fresh fruit.

Okanagan Biotechnology Inc

2006 *

Improved flavor soy

A non-transgenic product developed through molecular breeding, with reduced off-flavor volatiles and flavor binding components

Improves flavor; could encourage wider use of soy (with associated health benefits like lower heart disease risk and healthy weight maintenance)

Could increase the spectrum of soy based products marketed to mainstream consumers.

Monsanto

2006 *

Low linolenic soy oil

Lower proportion of linolenic fatty acids (

Nutritional and Product Quality Innovations

Nutritional and Product Quality Innovations

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