Societal Costs of Late Blight in Potato and Prospects of Durable Resistance Through Cisgenic Modification

In the European Union almost 6 Mha of potatoes are grown representing a value of close to €6,000,000,000. Late blight caused by Phytophthora infestans causes annual losses (costs of control and damage) estimated at more than €1,000,000,000. Chemical control is under pressure as late blight becomes increasingly aggressive and there is societal resistance against the use of environmentally unfriendly chemicals. Breeding programmes have not been able to markedly increase the level of resistance of current potato varieties. New scientific approaches may yield genetically modified marker-free potato varieties (either trans- and/or cisgenic, the latter signifying the use of indigenous resistance genes) as improved variants of currently used varieties showing far greater levels of resistance. There are strong scientific investments needed to develop such improved varieties but these varieties will have great economic and environmental impact. Here we present an approach, based on (cisgenic) resistance genes that will enhance the impact. It consists of five themes: the detection of R-genes in the wild potato gene pool and their function related to the various aspects in the infection route and reproduction of the late blight causing pathogen; cloning of natural R-genes and transforming cassettes of single or multiple (cisgenic) R-genes into existing varieties with proven adaptation to improve their value for consumers; selection of true to the wild type and resistant genotypes with similar qualities as the original variety; spatial and temporal resistance management research of late blight of the cisgenic genetically modified (GM) varieties that contain different cassettes of R-genes to avoid breaking of resistance and reduce build-up of epidemics; communication and interaction with all relevant stakeholders in society and transparency in what research is doing. One of the main challenges is to explain the different nature and possible biological improvement and legislative repercussions of cisgenic GM-crops in comparison with transgenic GM-crops. It is important to realize that the present EU Directive 2001/18/EC on GM crops does not make a difference between trans- and cisgenes. These rules were developed when only transgenic GM plants were around. We present a case arguing for an updating and refinement of these rules in order to place cisgenic GM-crops in another class of GM-plants as has been done in the past with (induced) mutation breeding and the use of protoplast fusion between crossable species.     

An Ethical Assessment of Cisgenesis in Breeding Late Blight Resistant Potato

Because of objections and worries related to genetically modified organisms (GMOs), the approval of GMO crops is a long and expensive process. Recently some researchers argued that a specific form of genetic modification, cisgenesis, would be safer and ethically more acceptable and therefore require a less stringent assessment. In this paper cisgenesis, as defined in recent literature, is ethically evaluated. After some general remarks on ethics in science and technology, two different basic attitudes towards reality are sketched as an evaluative framework for interventions in nature. Combined with general characteristics of biotechnology in agriculture and a view of the role of genetic information in organisms, that framework helps to formulate an ethical distinction between and evaluation of cisgenesis and transgenesis. It is argued that there is a significant ethical difference between transgenesis and cisgenesis, but that nevertheless any form of genetic modification should be integrated in a broader normative understanding of agriculture in order to work towards a more sustainable agriculture.

Cisgenesis,a New Tool for Traditional Plant Breeding,Should be Exempted from the Regulation on Genetically Modified Organisms in a Step by Step Approach

Modern potato breeding requires over 100,000 seedlings per new variety. Main reasons are (1) the increasing number of traits that have to be combined in this tetraploid vegetatively propagated crop, and (2) an increasing number of traits (e.g., resistance to biotic stress) originates from wild species. Pre-breeding by introgression or induced translocation is an expensive way of transferring single traits (such as R-genes, coding for resistance to biotic stress) to the cultivated plant. The most important obstacle is simultaneous transfer of undesired neighbouring alien alleles as linkage drag. Stacking several genes from different wild sources is increasing this linkage drag problem tremendously. Biotechnology has enabled transformation of alien genes into the plant. Initially, transgenes were originating mainly from microorganisms, viruses or non-crossable plant species, or they were chimeric. Moreover, selection markers coding for antibiotic resistance or herbicide resistance were needed. Transgenes are a new gene source for plant breeding and, therefore, additional regulations like the EU Directive 2001/18/EC were developed. Because of a strong opposition against genetic modification of plants in Europe, the application of this Directive is strict, very expensive, hampering the introduction of genetically modified (GM) crops and the use of this technology by small and medium-sized enterprises (SMEs). Currently, GM crops are almost the exclusive domain of multinationals. Meanwhile, not only transgenes but also natural genes from the plant species itself or from crossable plant species, called cisgenes, are available and the alien selection genes can be avoided in the end product. This opens the way for cisgenic crops without alien genes. The existing EU directive for GM organisms is not designed for this new development. The cisgenes belong to the existing breeders’ gene pool. The use of this classical gene pool has been regulated already in agreements regarding breeders’ rights. We are proposing a step by step approach starting with a crop and gene specific derogation and monitoring towards a general exemption of cisgenic plants from the Directive. Two examples, i.e. development of cisgenic potato for resistance to Phytophthora infestans and cisgenic apple for resistance to Venturia inaequalis are discussed shortly for illustration of the importance of cisgenesis as a new tool for traditional plant breeding. Cisgenesis is simplifying introgression and induced translocation breeding tremendously and is highly recommended for SMEs and developing countries.     

Cisgenesis Does Not Solve the Late Blight Problem of Organic Potato Production: Alternative Breeding Strategies

Average potato yields in Dutch organic farming systems vary from 15 to 29 Mg/ha and are limited by low input of nitrogen and severe late blight attacks caused by the oomycete Phytophthora infestans. Under Dutch late blight regulations it is mandatory to kill the haulm at 7% infestation. The late blight attacks have been so early in the organic potato production of the past few years that its acreage is now gradually decreasing whereas consumer demand is increasing. Agronomic control strategies have limited success. First priority lies in breeding for highly resistant varieties to safeguard organic potato production in the Netherlands. Cisgenesis, however, is not an option for the organic sector. Although the product of cisgenesis does not contain genes from non-crossable species it is a result of a genetic engineering process which is excluded from use in organic agriculture. As the principles and standards of organic agriculture are process-based, cisgenesis does not comply with the norms and standards of organic agriculture. The arguments of the organic sector go well beyond the alleged risks of the gene technology and relate to the technology itself. Breeding at DNA-level, instead of at whole-plant level, violates the integrity of life as described in the concept of naturalness. The Dutch organic sector is now aiming at increasing the traditional breeding activities including the participation of farmer-breeders in close cooperation with the formal breeding companies. Additional selection methods need to be developed to include required traits other than late blight resistance, such as nutrient efficiency. Recently two varieties have been released with high resistance against late blight based on introgressing genes from Solanum bulbocastanum. Organic agriculture can benefit from marker assisted breeding to achieve adequate pyramiding of different, new sources of resistance.     

Debate on the Exploitation of Natural Plant Diversity to Create Late Blight Resistance in Potato

This paper reports on a debate on intriguing propositions relating to the scientific, agronomic, societal and economic impact of the BIOEXPLOIT project, focusing on late blight resistance in potato. It discusses (i) whether identifying pathogen effectors will facilitate selecting durable R genes, (ii) whether breeding for durable late blight resistance requires deploying Rpi (for Resistance to P hytophthora i nfestans) genes, (iii) whether breeding strategies and cultural practices determine the durability of new resistance genes, (iv) whether marker-assisted breeding for Phytophthora infestans resistance is already in the stage of adoption, (v) to what extent genetically-modified organism technology can advance realizing late-blight resistant potato cultivars, and (vi) whether modifying R genes will result in novel broad spectrum resistance.     

同源转基因将成为利用野生资源进行作物育种的一种有效手段

 作物育种的一个主要限制因素是各栽培种的遗传基础日趋狭窄, 迫切需要从野生种质资源中导入优异等位基因。杂交障碍和连锁累赘降低了常规育种利用这些等位基因的效率。基因组研究的发展使得越来越多的植物基因的克隆成为可能。同源转基因就是将本物种或其近缘野生种克隆的优异等位基因导入到要改良的育种材料中。这不仅可以缩短育种的周期, 而且不会有不良性状的连锁累赘。同源转基因和常规育种所要导入的目标基因的来源是相同的, 因此育成的品种同样是安全的。如果转基因条例能够将同源转基因品种视同为常规育种品种, 同源转基因将会成为人们利用野生资源进行作物改良的一种有效手段。     

Applied Biotechnology to Combat Late Blight in Potato Caused by <Emphasis Type="Italic">Phytophthora Infestans</Emphasis>

Potato is an important crop, grown worldwide. It suffers from many pests and diseases among which late blight, caused by the oomycete Phytophthora infestans, is the worst. The disease is still causing major damage in many potato production areas and control is only possible by applying fungicides frequently. The knowledge on the molecular biology and genetics of the interaction between the plant and the oomycete is developing rapidly. These are relevant fields of study, currently dominated by the discovery of many resistance genes and numerous effector proteins and the analysis of their specific mode of action. These studies may yield essential information needed for the development of durable resistance. The long-term and worldwide effort to breed for resistance so far has had little effect. A novel breeding approach may change this. It is based on cisgenic modification (CM) consisting of marker-free pyramiding of several resistance genes and their spatial and temporal deployment yielding dynamic varieties that contain potato genes only. It is envisioned that this CM approach with potato’s own genes will not only prove societally acceptable but may also result in simplifications in the legislation on use of the CM approach. Various parties in the potato research arena intend to cooperate in this novel approach in a number of developing countries where potato substantially contributes to food security. The use of resources such as land, water and energy improves when the effect of late blight is markedly reduced.