Public-Private Partnerships in Plant Genomics for Global Food Security Sara Boettigera,1*, Vivienne Anthonyb, Kayje Bookerc, and Carrie Starbuckd

aDept. of Agricultural & Resource Economics, Univ. of California, Berkeley bSenior Advisor, Syngenta Foundation for Sustainable Agriculture c,dGATD Foundation

Abstract. Advances in genomic science have ushered in a revolution in the way we approach the science of crop im-provement. These changes have major implications in the commercial sector, but also for crops important to food security and for resource-poor smallholder farmers. Both private- and public-sector organizations are not only adapt-ing their research and development (R & D), they are adapting to a new era of public-private partnerships (PPPs). The shift toward more PPPs is driven by major trends im-pacting crop improvement including: game-changing new technologies, a surge in private investment in R & D, in-creasingly complex intellectual property rights (IPRs), and decades of under-investment in public sector capacity. We provide a snapshot of the current landscape of PPPs in genomics, document progress with crop genome sequenc-ing for staple food and feed crops in developing countries; and consider specific challenges, incentives, best practices and lessons learned. For the genomics revolution to con-tribute its full potential to benefit smallholder farmers in developing countries, the genomics research agenda needs to transition its emphasis from generation of sequence data to also supporting utilization of novel genes, molecular markers and improving predictive breeding models. PPPs can realize the skills and know-how to accelerate the de-velopment of new varieties to meet agricultural productivi-ty and food nutrition challenges that can change the food security landscape. However, it remains to be seen if the incentives for the public and private sectors to partner to-

*Corresponding author. E-mail: sdb@berkeley.edu Paper commissioned by the International Development Re-search Centre

gether to achieve this goal are sufficient to overcome the inherent difficulties.

Keywords: public-private partnerships, food security, ge-nomics, crop improvement

Introduction Feeding a world population of more than 9 billion people by 2050 is a formidable chal-lenge,1,2 demanding that we produce as much food in the next 50 years as we have in the entire history of mankind.3 Success will re-quire the exploration of innovative solutions to grow more food on less land, with fewer resources. Significant progress in food secu-rity will also require a sharp focus on solu-tions for the smallholder farmer. Smallhold-ers form the backbone of our global food supply. Worldwide, there are about half a billion smallholder farms;4 80% of the food eaten in sub-Saharan Africa and Asia is grown by smallholder farmers.5 These resource-poor farmers carry a disproportionate bur-den as we collectively strive to meet the world’s future demand for food, often farm-ing on marginal land with inadequate inputs. Smallholder farmers keenly feel the costs of climate change and food price volatility.

It is increasingly evident that making head-way against global hunger will also depend on the ability to engage both public and pri-vate sectors. Developing and delivering solu-tions for smallholder farmers at scale, within the time frame needed, requires public-private partnerships (PPPs) to better lever-age the diverse resources of companies, non-profits, universities, and governments.

Against this backdrop, we study the field of plant genomics, specifically considering PPPs that impact smallholder farmers. Plant ge-nomics provides an instructive context within which to understand more about the use of PPPs to address challenges to our global food

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system. Advances in genomic science have ushered in a revolution in the way we ap-proach the science of crop improvement for food security, and the field exemplifies major changes that have brought us to the threshold of a new era of PPPs in agriculture. The trends impacting crop improvement for food security include: game-changing new tech-nologies, a surge in private investment in re-search and development (R & D), increasingly complex intellectual property right (IPR) and bio-safety considerations, and decades of un-der-investment in public sector capacity. These general trends are all integral to the recent history of plant genomics and allow us to examine our current and future ability to use PPPs to address global food security.

Methods This paper begins by surveying recent devel-opments in genomic science that have ap-plicability to benefit smallholder farmers in developing countries. It then explores the role, challenges, and importance of public-private partnerships as a vehicle to contrib-ute towards global food security.

The research approach used is three-fold. First, we reviewed the current genomics lit-erature to establish progress, critical issues, and future trends in gene-sequencing of crops pertinent to smallholder farmers. Second, we investigated the landscape of operational public-private partnerships that target their outputs for the benefit of smallholder farmers in developing countries, including their scope, direction and composition. Third, we conducted interviews with leading scientists and managers in public and private organiza-tions that are using genomics in crop breed-ing programs and are experienced in partner-ing across public and private sectors.

The primary areas of enquiry were: (1) his-torical and current developments in ge-

nomics; (2) progress with crop genome se-quencing for staple food and feed crops in developing countries; (3) characteristics of PPPs addressing genomics, including chal-lenges, incentives, best practices and lessons learned; and, (4) the need, scope and poten-tial for fostering future PPPs in this field. The outputs from each line of enquiry have been consolidated and presented as a set of con-clusions to drive further investigation.

Recent developments in plant ge-nomics for food security Genomic science provides the opportunity to understand, at a fundamental level, the driv-ers of diversity and function in living organ-isms, including plants. Plant genomics for the purpose of this research paper is defined as the investigation and advancement of new crop varieties by studying the whole genome of a plant species and sequencing nucleotides to find constituent genes. Genomics provides valuable information that can be used to dis-cover novel genes to improve crop perfor-mance, understand genetic variability, and identify genetic markers that enable targeted, faster and more successful predictive breed-ing for crop improvement.

The genomics landscape has dramatically changed in the last 10 years, driven by ad-vances in next-generation sequencing plat-forms. Sequencing technology has pro-gressed in leaps and bounds since the human genome was first sequenced in 2001. Fierce competition within the field of commercially available high-throughput sequencers has driven sequencing speeds up and sequencing costs down by orders of magnitude.6 These technical innovations were paralleled by sig-nificant advances in bioinformatics, together opening an entirely new landscape for ana-lyzing genomic information.

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The primary areas of enquiry were: (1) his-torical and current developments in ge-nomics; (2) progress with crop genome se-quencing for staple food and feed crops in developing countries; (3) characteristics of PPPs addressing genomics, including chal-lenges, incentives, best practices and lessons learned; and, (4) the need, scope and poten-tial for fostering future PPPs in this field.

1.1. Major crop genomes and high-throughput sequencing

The first vascular plant species genome se-quence to be published was Arabidopsis tha-liana (thale cress) with 157 million base pairs in 20007. The number of plant and crop ge-nomes sequenced since that time has in-creased exponentially (Figure 1).

The sequence of the first major food crop, rice, was published in 2002, and remains probably the best-understood crop genome. Rice is also being used as a model for other cereal crop species, due to its much smaller genome size (430 million base pairs). Rice is diploid with two sets of chromosomes, but many plants are polyploids with multiple sets of chromosomes and can have highly repeti-tive sequences, making analysis and assembly of genomes challenging. For example, wheat is hexaploid, with 6 sets of chromosomes that have come from 3 genomes of other grass species. Wheat has a genome size of 16 bil-lion base pairs8 (about 5 times larger than the human genome). This creates unique tech-nical challenges for determining, unraveling and managing the complexity of sequence data and constituent genes, and also genetic mapping and the identification of markers which are essential to assist acceleration of plant breeding to improve food security.

Figure 2 provides a summary of the current status of genome sequencing for the top 15 food crops of primary importance to people

in low-income countries.9 The majority has been sequenced and seven have been pub-lished in peer-reviewed journals (Appendix 2). Twenty sequences of food crops have been published in peer-reviewed journals. The number of draft crop sequences an-nounced outside of peer-reviewed journals continues to grow, together with on-line ac-cess to the genome sequence data. A recent example is Asiatic pear (Pyrus bretschneideri), published by the International Pear Genome Consortium on 6 June 2012.10

The consortium is comprised of seven pub-lic sector National and International research institutes and universities from China, USA and Japan and includes contributions from over 60 experts. Occasionally, the private sector also announces whole genome se-quence achievements such as canola (Brassi-ca napus) by Bayer Plant Sciences in 200911. This was the outcome from a public-private collaboration between Bayer and public sec-tor researchers in Australia and China, and also a private sector genomics research ser-vice provider in the Netherlands. Progress in the sequencing of food crops demonstrates significant investment in research and devel-opment by both the public and private sec-tors.

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Figure 1 Acceleration in crop genome sequencing

Source: CoGePedia data modified and updated.12

Figure 2 Current status of genome sequences of 15 major food crops in low income food deficit countries

Sequencing technology and methods have made a major step change from first genera-tion laboratory work. Now second- and third-generation technology and sequencers are available that are highly automated and

deliver increasingly accurate, high-throughput data. A new commercially com-petitive service industry of contract data provision has emerged providing vast da-tasets, faster and much cheaper. Figure 3

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shows the dramatic fall in the costs of se-quence generation.

Cost has plummeted from millions of dollars per genome to as little as US$1000-US$10,000 for simpler plant genomes and transcriptomes, depending on the sequence depth and annotation requirements. This dramatic cost transformation combined with increased throughput, reduced error rate, and increased genome read length has en-couraged re-sequencing of genomes and is shifting the research agenda from sequenc-ing individual accessions as representatives of species, to research on whole species var-iability and evolutionary phylogeny.

Sequencing is emerging as a high-resolution and inexpensive method to genotype

populations, by the public sector for fundamental research, and by the private sector to understand allelic variation and drive predictive breeding models.

In plant genomics, as in human genomics, large-scale ambitious genome projects con-tinue to push ahead the science, comple-menting the work of smaller groups. In 2008, two major plant genome sequencing programs were initiated. The 1001 Genomes Project focuses on intra-plant species varia-tion. Over 1000 geographic variants of Ara-bidopsis are being sequenced to enable de-termination of the links between genotypic and phenotypic variation. The sequences of the first 80 accessions have been published13 and more than 700 have been completed.14

Figure 3 Falling cost of DNA sequencing

Source: National Human Genome Research Institute, USA. May 2012.15

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The second program is the 1000 Plant Ge-nome Initiative, also known as 1KP or One-KP16, which is investigating inter-species se-quence variation. It is currently the largest plant genome collaborative research pro-gram and is a public-private partnership. The goal of the PPP is to sequence the tran-scriptomes of 1000 different plant species and generate sequence information on ex-pressed genes. The selection of species has been based on those producing valuable bio-products such as medicinal plants, plants adapted to extreme environmental condi-tions like salt tolerance, phylogenic research, and key plant systems such as the yield-enhancing C4 vs. C3 photosynthetic pathway that could enable improved food security. The project is an international multidiscipli-nary consortium led by the University of Al-berta, Canada, with sequencing done by the Beijing Genomics Institute (BGI) in China, and venture capital input by United States-based Musea Ventures. All sequence data are being made public through GeneBank and other open access websites.

The rush of new transcriptomes and ge-nomes is producing a trend where research groups announce their achievements and provide open-access to their data before peer review has been undertaken as part of publication process in reputable journals. This has raised questions about data stand-ards, definitions of complete and draft ge-nomes, provenance of documentation, and lack of documentation on data quality and genome assembly.17

1.2. Bioinformatics and data storage

The limiting constraint for researchers is no longer sequence creation, but data pro-cessing, management, and storage in ar-chives that enable retrieval for ongoing re-search and meta-analyses. The accessibility

of affordable contract sequencing and third–generation sequencing technology, such as the Ion Torrent Personal Genome Machine (< US$100,000 capital costs), give research-ers and even single investigators the poten-tial to generate large amounts of sequence data.18 The challenge faced by many now is how to use such large datasets.

Access to high-performance computing and bioinformatics capability is vital for re-searchers to assemble the data, do compara-tive database analysis to identify novel al-leles and genes, and convert the mass of available data into knowledge that can be applied in crop breeding programs.19 Soft-ware and algorithm development have typi-cally lagged behind advancement and break-throughs in sequencing technology. While a depth of bioinformatics expertise exists in both public and private sectors, the private sector continues to have advantages in bioin-formatics expertise in its practical applica-tion to crop breeding.

An important analytics and bioinformatics platform supporting public sector genomic researchers internationally and in develop-ing countries is iPlant.20 The US National Science Foundation is investing $50 million in iPlant to provide a cyberinfrastructure that supports the computational needs to solve major problems within plant science and bring together experts in biological and computer sciences. It provides sustainable access to high-performance computing, in-teroperable software analysis, and large data sets. A specific agreement is in place to pro-vide support to the CGIAR Generation Chal-lenge Programme (GCP) and its Integrated Breeding Platform (IBP) 21 for molecular breeding in developing countries in six sta-ple food crops: rice, maize, wheat, chickpea, cassava, and beans. We interviewed the pro-

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ject directors of iPlant and GCP as part of this research investigation (see Appendix 1).

Genomics and sequencing are part of a much bigger platform of emerging “omic” sciences requiring bioinformatics to understand plant functioning and use genetic diversity to drive crop improvement.22 To fully capital-ize on the genomic data, integration with information from research on translation processes such as the range of proteins pro-duced (proteome), protein to protein inter-actions (interactome), phytohormone signal-ing (hormonome) and metabolite variation (metabolome) are also required (Figure 4). The relationship between a plant’s genotype and phenotypic properties are critical to un-derstanding yield, abiotic and biotic stresses, quality, nutritional, and other improve-ments.

Genomics and advances in plant breeding Strategies to achieve crop improvement are experiencing dramatic changes due to the transformation in sequencing and increased capacity to integrate data and knowledge from other areas of “omic” science (such as proteomics, metabolomics, and phenomics). Sequencing can be used not only to identify novel genes and molecular markers, but also to drive genomic selection and accelerate breeding programs.23 The potential to dra-matically reduce breeding times is especially critical. Greater speed will enable scientists to unlock the global diversity of crops and their wild relatives faster. This will help smallholders to meet the pressing challenges of climate change.

Figure 4 Genotype-phenotype profiling

Plant breeding for food security in develop-ing countries usually involves a wider diver-sity of germplasm, varieties, and landraces than is utilized in research and development for developed country markets. Large-scale

commercial plant breeding can target in-vestment in comparably concentrated port-folios. Public sector breeders, however, must serve the highly heterogeneous needs of smallholder farmers. These include hy-

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brids and open-pollinated varieties that grow in a huge diversity of cropping sys-tems.24 Using genomics in crops key for re-source-poor smallholder farmers offers sig-nificant potential to accelerate breeding pro-grams, study the genotype-by-environment interactions, and mine germplasm collec-tions for novel genetic material.

Molecular breeding has been estimated to have the capability to reduce the cassava breeding cycle, for example, from a typical 12-16 years to just four years. This has the potential to get new varieties to farmers faster, and also to save breeding programs hundreds of millions of dollars.25 Rice re-searchers, too, estimate that marker-assisted breeding will take 3-6 years off conventional breeding, generating economic benefits to-taling $50 to $900 million over the next few decades.26 Morrell (2012) estimates that using genomic selection models in maize could reduce the cumulative time between cycles of breeding from 5 years to 4 months.27 Full genotyping of parents would also enable selection to be done based on gene sequence data rather than presence or absence of markers.

Ultimately, understanding the genotype to phenotype correlations and the performance of allelic variation in different environmental conditions and exposure to biotic factors will create a shift in breeding effectiveness and efficiency. Development of reliable predic-tive models will enable targeted introgres-sion with very small chromosomal segments at key loci, thereby minimizing the need for back-crossing.

How we translate advances in genomics to have an impact in farmers’ fields is highly dependent on access to sophisticated bioin-formatics, statistical analysis and computa-tional resources, and the ability to contextu-

alize genomic data with phenotypic knowledge. For commercial crops with large homogenous markets, the private sector is leading the way in the use of bioinformatics, plant phenomics, and development of pre-dictive breeding models. The public sector has a broad capability for testing phenotypes in many countries under different environ-mental and climatic conditions, but ultra-high-throughput phenomic testing under controlled conditions is in its early stages. The Australian Plant Phenomics Facility is a leading example of high-throughput facilities in the public sector.28 There are important questions remaining, however, about the translation of high-throughput phenomics research tools and their impact on plant breeding for food security, where the im-portance of understanding genotype-phenotype connections within diverse envi-ronments is paramount.29 The translation of technologies used in commercial and aca-demic high-throughput phenomics to pro-duce tools that allow increases in efficiency outside the lab and in the field is likely to have a large impact on plant breeding for food security. Especially for the public sec-tor, phenotyping remains an expensive and time-consuming element in molecular breed-ing. Realizing the full potential of plant ge-nomics to improve crops for food security will require seeking knowledge and re-sources through partnerships with the pri-vate sector.

Another critical consideration when analyz-ing private and public sector roles in plant genomics for improvement of food security crops relates to the relatively limited align-ment of commercial private sector research targets and the goals of public sector re-searchers working to improve crop perfor-mance for smallholder farmers. Crops of common interest are, for example, rice, maize and wheat. Common trait targets in-

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clude: increased yield; resistance to abiotic stresses such as drought, heat, and flooding; more efficient input use, such as nitrogen-use efficiency; resistance to biotic stresses of diseases and pests; and increases in the nu-tritional value of a crop. The overlaps where private sector investments are also of benefit to smallholder farmers are important to rec-ognize as potentially fertile ground for part-nerships. Conversely, public sector invest-ment may be further targeted by identifying crops and traits important to smallholder farmers, but where incentives for private investment are lacking. Already, there are public-private collaborations on traits like drought tolerance that impact large-scale commercial farms, but also remain an enor-mous challenge for smallholder farmers (80% of the world’s crop land is rainfed, without irrigation and drought tolerance30). The Water-Efficient Maize for Africa project with partners including Monsanto, African Agricultural Technology Foundation (AATF), and the International Maize and Wheat Im-provement Centre (CIMMYT) focuses on drought tolerance, for example.

In developing crops for smallholder farmers, public institutions must serve a highly com-plex and heterogeneous market for seed, much of which is not of commercial interest to the private sector. As noted previously, a wide diversity of agro-ecological zones, bio-tic stresses, and local differences in market acceptance require new traits to be intro-duced into locally adapted germplasm. Pub-lic sector organizations, therefore, cannot take advantage of some of the economies of scale that can be utilized by the private sec-tor. Also, private sector research, develop-ment, and delivery channels are not always applicable to the markets the public sector serves. Ultimately, these differences have direct implications for the potential impact of plant genomics, and require different so-

lutions to be found. Some solutions will be unique to the context of food security crops, and others will be translations of advances used in large-scale commercial contexts.

There is little doubt that genomics will have a powerful impact on the improvement of food security crops in coming decades. Al-ready, the evidence is mounting. For exam-ple, the importance of drought-tolerance to food security has been noted above, but tra-ditional breeding has significant limitations given the complexity of the trait and the wide variety of strategies plants use for cop-ing with drought. Genomics has already al-lowed for the identification of multiple pathways that are important to understand-ing a plant’s reaction to limited water, and further progress seems likely soon.31

Clearly, new tools in genomics are already reducing costs, increasing precision, and pushing the barriers of possibilities in the science of crop improvement for food securi-ty. There is a need, however, to critically identify areas for public-private collabora-tion and targets for investment in the trans-lation of technologies used in the private sector to have impact on food security crops.

Public-private partnerships in ge-nomics The term “public-private partnerships” his-torically has often been used in reference to partnerships between governments and companies to deliver public services and large infrastructure projects. Here we define PPPs more broadly, as collaborations where at least one party from the public sector and one from the private sector join in a common endeavor. The contributions of the parties can be diverse, including: technology, man-agement skills, logistics capacity, infor-mation and data, access to equipment, intel-lectual property, know-how, and much else.

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We exclude, however, relationships between public and private organizations that are more arms-length, including contracted ser-vices, licensing of intellectual property and provision of only finance for research and development programs. We define “public” in the sense of working toward public inter-est goals (i.e. reducing global hunger), rather than private goals. Therefore public sector might include governments, universities, in-ternational aid agencies, philanthropic foun-dations, and civil society. Private sector is also used as a heterogeneous term to include a wide range of for-profit businesses from local small to medium enterprises (SMEs), to national companies, to multinational com-panies. The smallholder farmer is referred to throughout as the beneficiary of the goods and services produced by PPPs (better seeds, access to financial services, irrigation technology, fertilizer, market access, etc.). It is proposed that smallholder farming fami-lies should be considered as private sector entrepreneurs, with their own goals to earn enough to send their children to school and more generally increase household income and nutrition, and not as passive recipients of philanthropy.

The authors identified many research and development projects utilizing agricultural biotechnology approaches for crop im-provement in developing countries and in-volving in some way both public and private sectors. Table 1 summarizes current re-search and development PPPs with a sub-stantive genomics component that are tar-geted to benefit smallholder farmers in de-veloping countries. In each example cited, there is a public and a private sector organi-zation allocating resources, support and ex-pertise for gene sequencing and/or identify-ing molecular markers to enable molecular breeding. The table represents only part-nerships with publicly available information;

there is, no doubt, a larger field of confiden-tial contractual arrangements between pub-lic researchers and private companies where the projects may well benefit farmers in de-veloping countries, but the primary purpose is to support research directed towards commercial enterprise for larger-scale farm-ing.

Here we note some identified trends within the sample of projects reviewed. Most of the projects cannot be considered PPPs, using the above definition (i.e. they did not involve the parties’ joint investment of technology, skills, or assets). Private sector involvement was typically minimal, but some had adviso-ry input from genome sequencer hardware supplier companies and/or computational input from software companies. Collabora-tions led by public sector partners were of-ten conducted on an international basis ra-ther than a local one (e.g. Global Musa Ge-nomics Consortium, International Wheat Genome Sequencing Consortium, Interna-tional Rice Genome Sequencing Project, etc.). Generally, genomics projects led by the pub-lic sector were characterized by large con-sortia of essential experts in disciplines such as molecular biology, genetics, bioinformat-ics, computing capability and plant breeding. A current example is the collaboration an-nouncement in December 2011 by the Inter-national Centre for Tropical Agriculture (CIAT) and the BGI to sequence 5000 geno-types, varieties and wild species of cassava to support crop improvement.32

One important trend in this field has been the provision of technology by the private sector for use by public sector partners for food security goals. Agribusiness companies have provided the use of technology directly, through royalty-free licenses, or through non-assert arrangements for their intellec-tual property to enable crop improvement

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for resource-poor farmers to proceed. For example, in February 2012 Syngenta donat-ed a portion of its Maize Allelic Diversity Platform and over 1200 transgenic-free maize lines to the CGIAR Generation Chal-lenge Programme to support maize im-provement in developing countries.33 Simi-larly, in 2004 genetically modified “golden rice” seed and lines were donated to the Golden Rice Humanitarian Board to enable nutrition improvement research in rice to proceed.

Historically, both Monsanto and Syngenta have made rice genomic sequence data available for public research, but at the time did not undertake a full PPP to assist in the elucidation of the rice genome. Monsanto continues to support research for developing countries through grants to the Danforth Centre, involvement with the Water Efficient Maize for Africa PPP, provision of royalty-free licenses for enabling technology, and also through the International Cassava Con-sortium.34 Both Syngenta and the Syngenta Foundation for Sustainable Agriculture (SFSA) have a range of bilateral arrange-ments with public research institutes on ge-nomics projects (Table 1).

The most recent major genomics-related PPP initiative, specifically aiming to benefit 600 million farmers in Africa, is the African Orphan Crops Consortium. The initiative was announced in September 2011 and will combine the skills and resources of private sector companies (Mars, IBM and DuPont Pioneer-Hybrid), public sector researchers (New Partnership for Africa’s Develop-ment/NEPAD, Bioversity International, World Agroforestry Centre, African Academy of Sciences, TransFarm Africa and University of California, Davis), and an international NGO (World Wildlife Fund). The consortium is led by Mars and will focus on crops in Afri-

ca that are staple food crops but not primary targets for industry and therefore have re-ceived insufficient scientific investment.35

African scientists have identified a list of 96 crop species of relevance. The goal is for 24 species to be sequenced by the Beijing Ge-nomics Institute (BGI) in China by the end of 2014, and markers for molecular breeding to be identified. There is work, for example, on winter-thorn acacia, which has edible seeds, can be used as a soil improver due to its abil-ity to fix atmospheric nitrogen, and can help prevent soil erosion. Continuing public ac-cess to the resulting genetic information will be achieved using a website portal managed by the intellectual property organization, Public Intellectual Property Resource for Ag-riculture (PIPRA).

The consortium will also support accelera-tion of the rate of genetic improvement to increase yield and nutritional quality, by es-tablishing an African Plant Breeding Acade-my in Ghana and Kenya, as well as on the campuses of the University of California, Da-vis. This academy aims to train 250 plant breeding scientists and 500 technicians over five years. Equipment for both locations has been provided by the Life Technologies Cor-poration, a global bioprocess technology tools company.

Can PPPs accelerate our use of ge-nomics to deliver crop improve-ments for resource poor farmers? At their best, PPPs work to mobilize com-plementary resources and expertise from private and public sector organizations to deliver achievements that would not occur in either sector alone. In agricultural bio-technology (more generally than genomics), a defined set of market failures explains why private investment, in most cases, does not

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align with the public sector’s applications of biotechnology for the poor. The major rea-sons include: the high fixed cost of research coupled with a need for large markets to re-pay fixed costs, under-developed seed mar-kets in many poor countries, and difficulties in protecting and enforcing IPR.36 Evidence from the last two decades of crop genetics research shows that the private sector has, naturally, focused its efforts on crops and traits demanded by large-scale commercial farmers with large international kets.37,38 ,39

Public sector organizations, on the other hand, often operating within a mission to generate and apply R&D in crop genomics for the benefit of smallholder farmers, still lack many of the resources and know-how of the private sector. Specific areas that limit the public sector’s capability to use the full potential of genomics to improve food secu-rity crops include:

Genotype profiling. Creating reliable predic-tive breeding models requires detailed knowledge about the genetic profile, allelic diversity, and provenance of germplasm of large populations of parental lines of key crops. This exists in the private sector as a result of years of investment and use of “omic” technologies to understand their proprietary germplasm and breeding lines, but does not exist to the same extent in the public sector.

Bioinformatics. Public sector organizations often are constrained by insufficient access to (and the capability to use) cutting-edge bioinformatics tools and fit-for-purpose software and algorithms that would enable improvements in breeding for crops that are not of major interest to the private sector.

Data standards, storage and retrieval. Many public sector organizations lack experience

in the management of the very large datasets generated by sequencing and profiling.

Computing power. Access to the super-computing power needed to create novel analysis and maximize use of large datasets can constrain public sector genomics work.

Intellectual property. In biotechnology, more broadly than genomics, the lack of freedom to operate due to distributed IP ownership across multiple private companies on a range of critical technologies for crop im-provement can create prohibitively high transaction costs for public sector organiza-tions.40 Even though the public sector holds a great deal of valuable IP, it is dispersed through many different institutions, and much of it has been exclusively licensed to the private sector.41 One incentive for work-ing with the private sector on genetic se-quencing, analysis, and breeding, is to share access to the privately held technologies and in-house R&D of private organizations;42

Management. Sharing of management pro-cesses is an often-overlooked, but critical, asset that companies contribute to public-private genomics projects. Particularly in projects where the ultimate goal is the crea-tion of innovative products delivered to smallholder farmers’ fields, the private sec-tor has a valuable discipline of focusing on products, on the consumer (in this case the smallholder farmer), and on the market. The ability to manage even upstream genomics work with a direction for impact on the poor, and the benefits of good management in the efficient use of scarce funding resources can be of great value to the public sector partner.

The limitations outlined above provide the backdrop for our hypothesis that there are important reasons why PPPs in genomics for food security should be further analyzed and fostered. Below we examine in greater

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depth the potential for these constraints to be incentives for the public sector to seek partnerships with the private sector and we also discuss incentives the private sector may have to seek partnerships with public sector organizations.

Incentives to Partner In the past, a research institute working on crops for resource-poor farmers might have structured a genome sequencing project as a PPP to access sequencing capacity and speed. Today, genomics PPPs in crop im-provement with food security goals are formed based on different incentives. In 2009 the Global Cassava Partnership collab-orated with 454 Life Sciences for use of their FLX Platform with a goal to improve the speed and quality of the cassava genome se-quencing project.43 But just three years lat-er, this type of PPP structure would be un-likely to occur.

Today, one set of incentives for PPPs in plant genomics for food security revolves around public sector access to proprietary process-es, platforms, and know-how. We have not-ed above that top-notch bioinformatics and data management are now two key bottle-necks in public sector research programs and application of genomics for food securi-ty. Analysis and statistical models are criti-cal for genomic data to be translated in a meaningful way and expertise and know-how in the private sector is a key incentive for public organizations to engage in PPPs.

A second current trend in incentives under-lying PPPs in this field relates to intellectual property rights (including ownership of germplasm, trade secrets of processes and data, copyright over computer code, and pa-tents). Public sector organizations are the stewards of enormous collections of germplasm in their seed banks and the ge-

netic diversity represented in these collec-tions has taken on new value for the private sector.44 Some large companies with signifi-cant genomics research budgets are current-ly expanding into new markets in emerging economies and developing countries. Their historical focus on temperate germplasm is no longer sufficient and accessing high quali-ty tropical germplasm may be important to their commercial strategy. Additionally, mining the genetic diversity in public sector gene banks holds the potential for compa-nies to identify commercially valuable traits.

A third set of incentives relates to differ-ences in public and private sectors’ capaci-ties to engage in resource-intensive pheno-typing. As noted above, meaningful applica-tion of genomics to farmers’ fields requires integration of genotype and phenotype data to produce useful tools and predictive mod-els for plant breeders.

While genome sequencing projects, per se, no longer require private sector collabora-tion, there are still important public-private partnership issues in plant genomics that will determine our future ability to use these advances to provide small-scale farmers with improved crops that can positively af-fect their lives.

Despite some publications proposing that robust PPPs are based on common objec-tives, our analysis of the incentives for en-gaging in PPPs supports a hypothesis that strong PPPs can exist when partners have highly asymmetrical objectives and incen-tives for engaging in a PPP. A good under-standing of these incentives is essential for policy-making at the national and interna-tional levels, to enable more interaction be-tween public and private sectors. But it is also critical at the level of the individual PPP for prospective partners to assess the incen-

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tives, as well as the constraints of a ge-nomics-focused collaboration.

1.3. Public Sector Incentives

Particularly in R & D PPPs, access to technol-ogy is a primary incentive for public sector partners. In past decades, agricultural R&D investment trends have created a large port-folio of proprietarily-owned technologies in the private sector that have a potential to impact crop improvement for food securi-ty.45 These technologies include enabling technologies, research tools, trait genes, bio-informatics platforms, and even lab equip-ment.

In addition to accessing technology, public sector organizations find a variety of incen-tives for engaging in PPPs. Consider, for ex-ample, the market- and product-focused ex-pertise in the private sector. Public sector research organizations are often focused on initial discovery and proof-of-concept stag-es;46 the currency among scientists is journal publications rather than successful products. Private organizations, on the other hand, not only have more expertise in “downstream” functions of product development, market-ing, and market distribution, but they ap-proach “upstream” science through the lens of delivery. The rigor of assessing technical progress within the context of information about cost, time to market, regulatory hur-dles, IPR issues, market acceptance, and much else is a discipline from which public sector scientists working on crops for the poor can benefit.47 In fact, donors investing in public sector science also benefit due to improved efficiency as their investments produce adoptable varieties with impact.

Most importantly, however, smallholder farmers benefit because the focus on the market improves the chance that they may have access to a product that delivers value

to them. PPPs offer the public sector an in-creasingly important mechanism for leverag-ing private investment to benefit the poor, focusing on applications to crops, traits, and a wide range of other technologies that are of importance to smallholder farmers in de-veloping countries. In this context, public-private partnerships are wholly consistent with the central role of public institutions as a steward of public interests, seeking to find the best ways of ensuring that scientific and technological advancement benefit the poor in developing countries.48

1.4. Private Sector Incentives

For-profit companies are unlikely to unilat-erally take on projects where the primary aim is to serve the global poor. However, PPPs may offer private companies ways of finding value to incentivize their involve-ment and investment. In agribusiness and other industries companies are increasingly approaching developing countries not in a philanthropic way, but as prospective and current business opportunities and this has driven incentives to participate far beyond the old model focused on public relations benefits.49 Common incentives for private sector organizations to engage in PPPs relat-ed to food security include: access to strong basic research skills;50 access to collections of genetic resources 51 including potential value in tropical germplasm supporting shifting markets and access to genomic data of diverse varieties; potential profit from the projects’ resulting technological discover-ies;52 benefits arising from experience with the conditions, culture, and varieties of crops used by the rural poor;53 improved employ-ee retention; positive public relations (PR) benefits; corporate social responsibility obli-gations;54 and the ability to leverage rela-tionships with extension organizations and knowledge of local seed delivery systems55.

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Incentives for companies to partner in agri-cultural PPPs hinge critically on future de-velopments in emerging market economies. These economies include vast numbers of smallholder farmers, as well as enormous commercial capacity. How these markets develop will determine our understanding of agribusiness PPPs in the future decades. Emerging market economies are more inte-grated into the global economy than ever before. In 2010, emerging market compa-nies were involved in more than half of 11,000+ cross-border mergers and acquisi-tions deals announced. 56 Populations in emerging economies continue to grow; cur-rently Asia accounts for one quarter of the world’s middle class, but expectations are for that to double by 2020.57 Predictions that emerging market economies would be engines of growth in the next decades were commonplace before the recent downturn. The World Bank reported that by 2025, six emerging market economies will account for more than half of all global growth (Brazil, Russia, India, China, South Korea, and Indo-nesia).58 Despite drastically revised figures for growth rates, the fact remains that these economies, by their sheer size and integra-tion in the world market, will remain critical to our future understanding of a new genera-tion of PPPs that looks different from our model of developed-country-based multina-tionals partnering with public sector organi-zations.

Challenges arising in PPPs with food security goals Academic work on PPPs, for the most part, has focused less on challenges faced by PPPs,59 and more on the extent to which PPPs have realized their theoretical benefits for private and social goods.60 Before clos-ing, we briefly include a section on practical challenges faced by public and private part-

ners engaging in agriculture PPPs. As in the section above, the discussion focuses on challenges generally, but these are directly applicable to the specific case of plant ge-nomics PPPs.

Some challenges are common across all PPPs, regardless of type or sector. For ex-ample, cultural differences are often cited as a cause of friction and miscommunication in partnerships between public and private or-ganizations.61 Less cited, but of critical im-portance is the governance structure of a PPP. The governance structure underpins the ultimate performance of the partnership, providing the accountability and legitimacy needed to satisfy partners as well as a larger set of stakeholders.62 Creating robust PPP governance structures that satisfy the di-verse needs of partners remains a perennial challenge in PPPs. Another challenge relates to communications. As in any partnership, success depends on good communications, both internal and external, but PPPs provide exceptional challenges in this area, due to diverse stakeholders, different working cul-tures, geographic distance, and the im-portance of challenges to reputation in part-ners engaging in PPPs.

A fourth challenge we note relates to differ-ences between how public and private sector organizations manage information. Compa-nies depend on confidentiality to protect their assets from competitors (e.g. trade se-crets), and strategic confidentiality is a core part of a company’s ability to protect its pa-tentable intellectual property. For public sector organizations, on the other hand, knowledge-sharing is often a core tenet of their work. Publishing is not only embedded in the incentive structure of many public or-ganizations, but the right to publish is often protected in institutional policies.

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Lastly, intellectual property rights (IPRs) require careful consideration. Across the public and private sectors, IPRs involve complex ethical, legal, ideological, as well as economic issues. Partners often bring wide-ly differing views of IPR that extend well be-yond how to use IPR within the partnership. Plant genomics, additionally, finds itself at the nexus of areas where public and private sectors can have divergent views, including issues related to: ethical debates surround-ing the “patenting of life;” legal debates re-garding the patentability of genes; discus-sions on the creation and stewardship of public goods; and diverse views on “anti-commons” problems and the proprietary ownership of research tools. Ideally the IPR strategy of a PPP with food security goals is tailored to support the greatest impact on the largest number of smallholders. The strategy is designed to reflect the incentives and the constraints of the partners derived from their own organizations. In practice, however, IPR issues can delay or derail the brokering of partnerships and accentuate differences in culture and institutional poli-cies.

The challenges discussed here (cultural, governance, communications, management of information, and IPRs) are not the only ones faced by PPPs in agriculture. Despite the importance of understanding incentives and constraints of PPPs in agriculture for practitioners, donors, and policy-makers, these topics tend to remain under-researched.63

Research Conclusions Our research here has focused on plant ge-nomics public-private partnerships with food security goals and our findings reflect a rapidly changing field on all fronts, and one that warrants more analysis. We close with a summary of our research conclusions.

1.5. Genome Sequencing

Over the last 10 years, there has been a ma-jor acceleration of technology progress ena-bling high-throughput genome sequencing of many crops important for food security. The cost of sequencing has plummeted in the last 3 years, which has led to new innovative strategies to investigate intra and inter-species. The scale and affordability of se-quence generation for 1000s of plant species and populations of genotypes is now a reali-ty that is also changing the research land-scape. Genome sequencing of key orphan crop species, important as staple food crops in Africa, are underway (Table 1).

The dynamics of collaborations are also changing. During the era of the early rice genome elucidation, collaboration was an imperative due to the technical constraints and the scale of the project. Now, however, high-throughput second-generation service provision contracts or bilateral arrange-ments with genomics institutes, such as with the BGI, negate the need for such large multi-partner arrangements for sequence data generation. Also, third-generation technolo-gy is arriving64 with a capital price tag that may be affordable for many research de-partments to become self-sufficient in se-quence data generation. The challenges for genomics to deliver crop improvement for smallholder farmers in developing countries now lie in the areas of bioinformatics, com-putation analysis, and data management.

Huge amounts of data are being generated by both the public and private sectors. Pub-lications of new genome sequences occur almost monthly. In most cases, this has been a result of collaborating parties providing open access to data, samples or equipment rather than outputs having been generated together as true public-private partnerships.

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A key conclusion from this analysis is that new PPPs to generate sequence data on crops important in developing countries are not required. There are insufficient incen-tives for either the public or private to create partnerships for this purpose.

1.6. Development of genomics capability

During the genome sequencing revolution, the private sector has systematically invest-ed in all the component “omic” and breeding technologies to enable “end-driven” remu-neration imperatives, namely commerciali-zation of varieties in key business markets. This has included accessing and characteriz-ing the best germplasm in key food, feed, and biofuel crops, elucidating genetic markers and genome maps, and combining the best approaches from conventional and molecu-lar breeding. A particular emphasis has been made on defining the genetic control of complex beneficial traits such as drought tolerance, yield, and biotic stress responses to understand the performance of alleles at different loci and the effects of environmen-tal interactions.

The driver has been to enable predictive breeding and selection of parental lines for crosses to achieve competitive advantage through high-performing varieties produced by efficient use of resources and investment, and faster timescales to market. This has involved examining gene expression and phenotyping in many key business locations and having a deep knowledge about their proprietary breeding lines.

Seed companies have also placed emphasis on the need to have the most efficient meth-ods of not just generating large datasets, but also storing, retrieving, interpreting and re-lying on robust and trustworthy data. This has involved accessing high-powered com-puters, developing policy on data quality

standards and data reproducibility, optimiz-ing standard operating procedures and use of bioinformatics tools.

During this same period public sector re-search in developing countries (with the ex-ception of India and China) on subsistence food crops and especially “orphan crops” has been characterized by under investment. Consequently, energy has been channeled to try to achieve participation in the genomics revolution and data generation using new genomic technologies. Setting research data standards and all the enabling activities needed to drive science policy on data man-agement have been much lower on the agenda. Work on these topics has been done in a much more fragmented way based large-ly on individual researchers’ experience and expertise. Predictive breeding using ge-nomics capabilities has also not been given as much importance as by the private sector. Integrating molecular breeding approaches with conventional breeding and incorporat-ing the use of other genomic data has also been challenging. It requires many plant breeders to shift from a primary leadership function to being part of a multi-functional team.

1.7. PPPs and shifting the genomics agenda to benefit smallholder farmers

In order for genomics to make a real differ-ence for smallholder farmers in developing countries the research agenda needs to shift from generating sequencing data per se to utilizing newly discovered novel genes and understanding and utilizing the molecular drivers of genotype-phenotype variation. Identification of molecular markers, genomic selection and building predictive models for key crops will accelerate molecular breeding programs. Further investigation is required to determine if there are sufficient incentives

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for both the private and public to engage in partnerships for this purpose. Areas for po-tential cooperation include: the public sector benefiting from not only access to new tech-nology but also the experience and expertise within the private sector on data manage-ment, policy and implementation of data standards, documentation and data prove-nance, genome annotation and validation, bioinformatics approaches, high-throughput genotyping and integration of phenotypic data to build predictive breeding models. Potential incentives for the private sector to be explored include: gaining access to a large range of diverse environmental climatic sit-uations and abiotic stresses for field testing genotypes and also extensive climatic and environmental data65. This could enable re-finement and further training of their predic-tive breeding models and selection of paren-tal lines for specific eco-climatic zones either within areas of current business, especially in view of climate changes expected over the next 10 years or could lead to growth oppor-tunities for new business.

For potential new genomics PPPs to deliver their goals on food security and benefits for smallholder farmers, the incentives must exist to catalyze partnerships among public and private sector parties. But if PPPs in ag-riculture are ultimately going to have impact on food security, enabling the poor to have better access to improved crops, we have more to understand, also, in the manage-ment and governance of the partnerships, in managing cultural norms66, routes of com-munication, confidentiality requirements and the terms of ownership of inventions and intellectual property rights. All of these are essential areas of learning for any new genomics PPP to be successful.

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Table 1

Name and goal Research and Development Program

Scope Public Sector Private Sector

Duration Investment

African Orphan Crops Consortium.67

To improve the incomes of the 600 million Africans involved in farming.

Genetically sequence 24 ge-nomes of neglected African food crops that are important for food and income.

Educate African plant breeders in the application of genomic information to crop improve-ment (Train 250 plant breeding scientists and 500 technicians over five years).

2011-2014

2011-2016

$7.5 million

(seeking +$32 million)

New Partnership for Africa’s De-velopment (NEPAD) World Agrofor-estry Centre, Bioversity Inter-national, African Academy of Sciences, TransFarm Afri-ca, University of California, Davis

Mars, IBM, World Wild-life Fund (WWF), DuPont/Pioneer Hi-Bred

Ug99 stem rust of wheat.68 To accel-erate breeding of wheat resistant varieties.

Identifying and mapping

genetic markers for Ug99 stem rust.

2009-2011 CIMMYT, SFSA Syngenta Seeds

Disease-resistant cassava consorti-um.69

To reduce the im-pact of Cassava Mosaic Disease (CMD) and Cassava Brown Streak Dis-ease.

Disease resistance gene expres-sion improvement.

2010 KARI (Kenya), NARO (Uganda) Donald Danforth Centre (USA)

Dow Agro-sciences

Scientific know-how exchange programs

(SKEP).

To characterize rice genetic diversity, marker assisted breeding applications to ad-dress crop productivity con-straints.70

Increase the rate of yield gains and to boost the quality and diversity of hybrid rice, sharing germplasm, facilities and inter-action of scientists.71

http://www.aatf-africa.org/userfiles/DuPont-IRRI-boost-rice-yield.pdf

2009-

2009-

International Rice Research Institute (IRRI)

Syngenta

DuPont

15

Cocoa genome se-quencing consorti-um.72

To sequence the cocoa genome.

-2010 Clemson Univer-sity Genomics Institute

USDA/ARS

Hudson Alpha Institute for Bio-technology,

National Centre for Genome Re-sources, New Mexico

Indiana Universi-ty

Washington State University

PIPRA

MARS

IBM

Neem sequence.73 Sequencing the genome of Indian medicinal plant, neem.

2009 2011

(completed)

Department of Informatics (DIT/MCIT); Institute for Bio-informatics and Applied Biotech-nology

Stand Ge-nomics Pri-vate Ltd

20

Appendix 1 List of experts interviewed

Name Interview Position/role Organization

Lawrence Kent 9 December 2011 Agricultural development

Senior Program Officer

Bill and Melinda Gates Foun-dation

Dr Claude Fauquet 26 September 2011 Co-chair Global Cassava Partnership

Director International Laboratory for Trop-ical Agricultural Biotechnology (ILTAB)

Donald Danforth Plant Science Centre,

Dr Kevin Pixley 14 November 2011 Director Genetic Resources Programme

Maize Leader Harvest Plus Programme

CIMMYT, Mexico

Dr Alan B Bennett 15 November 2012 Executive Director

Professor UC Davis

PIPRA

Dr Homer Caton 27 April 2012 Head Corn Genetics/molecular breeding program

Syngenta, SBI North Carolina

Dr Jean-Marcel Ribaut 23 April 2012 Executive Director Generation Challenge Pro-gramme, Mexico

Dr Erik Legg 8 May 2012 Group Leader Omics/Functional Genomics Syngenta, Slater, Iowa

Dr Stephen Goff 15 May 2012 Project Director and Principal Investigator - iplant

University of Arizona

Dr. Mathew Reynolds 13 July 2012 Head of Wheat Physiology CIMMYT, Mexico

21

Appendix 2 List of plant genomes published in scientific peer-reviewed journals

Plant name Latin Name Type of crop Journal Year Date Web source

Arabidopsis Arabidopsis thaliana Model Nature 2000 December 14th

http://dx.doi.org/10.1038/35048692

Rice Oryza sativa L. ssp. japonica

Food Science 2002 April 5th http://dx.doi.org/10.1126/science.1068275

Rice Oryza sativa L. ssp. indica

Food Science 2002 April 5th http://dx.doi.org/10.1126/science.1068037

Poplar Populus trichocarpa Industrial Science 2006 September 15th

http://dx.doi.org/10.1126/science.1128691

Grape Vitis vinifera Food Nature 2007 September 27th

http://dx.doi.org/10.1038/nature06148

Moss Physcomitrella patens Ancient lower plant with small genome

Science 2008 January 4th http://dx.doi.org/10.1126/science.1150646

Papaya Carica papaya Food Nature 2008 April 24th http://dx.doi.org/10.1038/nature06856

Lotus Lotus japonicus Model for nitrogen-fixation

DNA Re-search

2008 May 28th http://dx.doi.org/10.1093/dnares/dsn008

Sorghum Sorghum bicolor Food, feed Nature 2009 January 28th http://dx.doi.org/10.1038/nature07723

Cucumber Cucumis sativus Food Nature Genetics

2009 November 1st http://dx.doi.org/10.1038/ng.475

Maize Zea mays Food, feed Science 2009 November 20th

http://dx.doi.org/10.1126/science.1178534

Soybean Glycine max Food Nature 2010 January 14th 10.1038/nature08670

Brachy Brachypodium dis-tachyon

Model mono-cotyledon grass

Nature 2010 February 11th http://dx.doi.org/10.1038/nature08747

Apple Malus × domestica Food Nature Genetics

2010 August 3rd http://dx.doi.org/10.1038/ng.654

22

Castor Bean Ricinus communis Food, industri-al

Nature Biotech-nology

2010 August 22nd http://dx.doi.org/10.1038/nbt.1674

Jatropha Jatropha curcas Oil DNA Re-search

2010 December 13th

10.1093/dnares/dsq030

Strawberry Fragaria vesca Food Nature Genetics

2010 December 26th

http://dx.doi.org/10.1038/ng.740

Cocoa chocolate Theobroma cacao Food Nature Genetics

2010 December 26th

http://dx.doi.org/10.1038/ng.736

Arabidopsis lyrata Arabidopsis lyrata Model Nature Genetics

2011 April 10th http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3083492/

Date Palm Phoenix dactylifera Food, oil Nature Biotech-nology

2011 May 29th http://dx.doi.org/10.1038/nbt.1860

Selaginella Selaginella moellen-dorffii

Ancient lower plant with small genome

Science 2011 May 20th http://dx.doi.org/10.1126/science.1203810

Potato Solanum tuberosum Food, industri-al

Nature 2011 July 14th http://dx.doi.org/10.1038/nature10158

Thellungiella par-vula

Thellungiella parvula Model plant with tolerance to abiotic stresses e.g. salt and cold

Nature Genetics

2011 August 7th http://dx.doi.org/10.1038/ng.889

Various Brassica rapa Food, oil Nature Genetics

2011 August 28th http://www.nature.com/ng/journal/v43/n10/abs/ng.919.html

Cannabis Cannabis sativa Fiber, medici-nal

Genome Biology

2011 October 20th http://dx.doi.org/10.1186/gb-2011-12-10-r102

Pigeon pea Cajanus cajan Food Nature Biotech-nology

2011 November 6th http://dx.doi.org/10.1038/nbt.2022

Medicago Medicago truncatula Model legume

Nodule for-mation and nitrogen-fixing

Nature 2011 December 22nd

10.1038/nature10625

23

Field bean

Field pea

Vicia faba

Pisum sativum

Food, feed BMC Ge-nomics

2012 March 20th www.biomedcentral.com/1471-2164/13/104

Foxtail millet Setaria italica Food Nature biotech-nology

2012 May 13th http://dx.doi.org/10.1038/nbt.2196

Tomato Solanum lycopersicum

Solanum pimpinelli-folium

Food Nature 2012 May 30th http://www.nature.com/nature/journal/v485/n7400/full/nature11119.html.

Source: CoGepedia Plant genome statistics (updated to end May 2012). 74

24

Acknowledgements GATD was commissioned by Canada’s Inter-national Development Research Centre (IDRC) to prepare this research paper. Grate-ful thanks are also given to the expert scien-tists and research managers who participated as part of the interview process, and to the Syngenta Foundation for Sustainable Agricul-ture for its scientific and advisory input.

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69http://www.dowagro.com/newsroom/corporate/2010/20100112a.htm

http://www.dowagro.com/newsroom/corporate/2011/20110715a.htm

70http://www.syngenta.com/global/corporate/en/news-center/news-releases/Pages/en-090907.aspx

71http://www.aatf-africa.org/userfiles/DuPont-IRRI-boost-rice-yield.pdf

72http://www.cacaogenomedb.org/main

73http://www.biomedcentral.com/1471-2164/13/464, http://www.biocon.com/docs/strandls_neem.pdf

74http://genomevolution.org/wiki/index.php/Plant_Genome_Statistics

27