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Agro 641 Sources of Genetic Advance for the future

Course: SCSC 641, Fall 2009
School: Texas A&M
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: u'-.) ::Xi! fe, , '..3n'1 ..... ....:c)... _. c "-?-/1 &f--~ , ~ Sources of Genetic Advance for the Future fl1e~~-P 8o-J:LL 6.~~TI. - F )1tJle.J Donald N. Duvick Pioneer Hi-Bred International, Inc. Des Moines, Iowa dc;.~-~~I ... -' d1" JW7 We are still Victorians, in agronomy. We'are convinced of the inevitability of progress -- especially in yields of our major ,farm...

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: u'-.) ::Xi! fe, , '..3n'1 ..... ....:c)... _. c "-?-/1 &f--~ , ~ Sources of Genetic Advance for the Future fl1e~~-P 8o-J:LL 6.~~TI. - F )1tJle.J Donald N. Duvick Pioneer Hi-Bred International, Inc. Des Moines, Iowa dc;.~-~~I ... -' d1" JW7 We are still Victorians, in agronomy. We'are convinced of the inevitability of progress -- especially in yields of our major ,farm crops. Graphs of achieved'yields in the the u.s. during the past 50 or ,60 years reinforce this belief. During that time, yields in grain weight per unit are~ of maize, soybeans arid wheat have increased at a steady or even an increasing rate, especially since the mid-l950s. The average gain in maize yields for example has been about 3% per year. For wheat it has been about 2% per year, and for soybeans it has been about 1% per year. We plant breeders have pointed out that genetic improvements - better varieties and hybrids - have been responsible for at least half of the gains in yield. We do admit that agronomic improvements have also been important; better weed control, improved ~lanting a~d harvesting machinery and abdve all, plentiful applications of commercial nitrogen fertilizer have gone hand in hand with, and have often been indispensable partners to the advances due to breeding. Indeed, the fact that some yield enhancers such as nitrogen fertilizer (for example) would be of little value without modern maize varieties indicates the virtual impossibility of considering genetic improvements without taking into account other coordinated changes in farming practices, as well. - '" r:",:.:) c;,) (, '.f ~ ~. ~ '~'7 J '.. .j -2 Plant Breeding's Role in Future Productivity Gains I t i s likely that little more gain in yield can be achieved from additional non-breeding inputs in developed countries like the United States. Techniques of planting, weeding and harvesting cannot be improved much more. Plant growth regulators for yield enhancement seem as far in the future as ever. Increasingly stringent regulations and tolerances for insecticide, fungicide and herbicide applications indicate these aids to production will not be used much more widely in the . future; indeed, their use possibly will be reduced. Even nitrogen fertilizer usage has plate~ued in the past few years; economic considerations now and environmentally oriented legislation in the near future will perhaps cause .actual reduction in rates of application of this yield-enhancing input. It is probable, therefore, that yields of our major crop plants will not continue to rise at the rates we have grown accustomed to, during the past 30 years. Only genetic improvements remain as the one technique with sure promise - and ability - to raise yields and productivity of crop plants in the developing countries. And even genetic improvements of crop plants may be more difficult to produce in the future, compared to the past 100 years, in part because of doubts as to free availability of germplasm, adequacy of support for plant breeding, and proper ways for remuneration of plant breeding endeavors. , this essay. Developing countries, in contrast to developed countries, still have much room for improvement o~ yields and stability of production through addition of non-breeding inputs, as well as by means of plant breeding. These c6untries are now trying to compress into a few years the changes that were accomplished I will discuss these problems in a later section of -3 over a hundred-year time span in North America and Europe. It is clear to them - the developing countries - that plant breeding is an indispensable tool, an essential part of the total set of changes they wish to bring about. Worldwide, therefore, the importance of plant breeding - of germplasm modification - is increasingly important. Plant breeding stands on two legs: breeding methods, and sources of useful germplasm. Neither can serve in the absence of the other~ neither can be analyzed without reference to the other. In this essay I shall deal primarily with the second topic: sources of useful germplasm, with occasional reference to breeding techniques. Two Kinds of Useful Germplasm: Exotic Landraces, Elite Cultivars Useful germplasm is defined in various ways depending on the developmental stage of plant breeding in a country or region. For example, in the Great Plains region of the United States in the mid-nineteenth century, useful germplasm was defined as imported wheat varieties. Turkey wheat, an exotic variety (actually a complex of closely related varieties) was used without change as a revolutionary new wheat throughout what is now the hard red winter wheat belt of the United States, during the late 1800s and in the first years of this century. (Turkey wheat was brought to Kansas by immigrants from the Russian Crimea. It was preadapted to Kansas climate, having been selected in a Eurasian climate much like that of Kansas.) At a later stage of development, farmer selections were regarded as the best source of useful germplasm. Hundreds of such wh~atselections (derived, probably, from accidental hybridizations) were on hand at about the turn of the century in -4 the Great Plains region, although few farmers would have had broad knowledge of what was available, region wide. The many open "pollinated varieties of maize were likewise traded around, via word of mouth and the n.ewly developed farm seed companies, at about the same time, especially in the u.s. corn belt. When plant breeding as we know it was developed, starting in the early years of this century, other definitions of useful germplasm appeared. On the one hand," finished varieties were useful germplasm for breeding. They were hybridized and selfed to give F2 populations from which new lines could be selected through successive generations of selfing and selection. On the other hand, when widely used cultivars attracted epidemics of cultivar-specific disease, breeders sometimes turned to exotic varieties (usually available through Plant "Introduction Stations, or in privately developed collections) as useful sources of specific disease resistance genes. (Breeders usually find needed resistance genes in adapted germplasm. But rarely the only possible source is an unadapted exotic.) Although the exotic varieties were not used as such for grain production, they were surveyed for genes giving desired pest resistance. Backcrossing techniques then were used to transfer those useful genes (with as little linked genetic material as possible)/into elite gendtypes. Thus, highly bred, well adapted cultivars as well as exotic, poorly adapted landraces (primitive farmer varieties) were regarded as useful germplasm by the newly developed profession of crop plant breeders. These two sources, elite cultivars and exotic landraces, are still the main foundation for progress in crop plant breeding, worldwide. Methods of obtaining these two sources of useful breeding stocks have changed over the years, however. -5 Landrace Collection and storage Introduction of foreign germplasm - particularly, of landraces - was at first done primarily by means of unorganized, individual and often private efforts. Early in this century the USDA assumed responsibility for organizing and financing collection trips. Eventually the USDA collections and associated state and federal institutions and agencies were formally gathered into a single, loosely organized system, the National Plant Germplasm System (NPGS). Qualified breeders can now search through materials in the USDA collections for potentially useful breeding materials. The seed collections are maintained in medium- and long-term storage facilities _ germplasm banks. Most developed countries of the world, and some of the developing countries, now have similar germplasm storage facilities, often on a smaller scale and usually emphasizing crops important to their country. Large and well maintained germplasm banks for major crop plants of the world are also maintained by international research organizations such as CIMMYT (maize and wheat), IRRI (rice) and ICRISAT (sorghum and millets). worldwide responsibility for encouragement and assistance of germplasm banks has been assumed by an internationally financed and managed organization, the International Board for Plant Genetic Resources (IBPGR). IBPGR has a small staff and a small budget, is housed in FAO (Food and Agricultural Organization) headquarters in Rome (but does not belong to FAO), and in the past ten years has had beneficial effects on progress in collection and storage of germplasm, out of all proportion to its size. It has been an important stimulus to collecti~n of the rapidly disappearing stock of landraces. (These primitive varieties are passing rapidly into extinction as farmers in developing nations adopt new "high yield" varieties, coincident -6 with the farmers' transformation from subsistence farming to some sort of integration with the money economy). Deficiencies in the Ger~plasm Collections On the surface it appears, therefore, that plant breeders have rich sources of highly diverse germplasm, safely stored in germplasm banks in the u.s. and abroad, in plentiful amounts and representing a full sample of the world's genetic variability. In actuality~ serious problems exist. The banks are in general not well maintained. This is true of U.S. collections as well as of those abroad, in developing or developed countr;es. Understaffing and insufficient budgets not poor workers - are the chief causes of problems. Collections are not being renewed fast enough; seeds die of old age. Collections often are not well desctibed; breeders often have no way of determining what individual items might be like or how to try to use them. Some collections are overflowing the available storage space; storage banks (such as the U.S. facility at Ft. Collins, CO) simply have no more room. And in some cases, more collecting needs to be done immediately, before the primitive farmer varieties disappear forever. These problems are not universal. Some of -the internationally supported collections are in good shape, as for example that for rice, at IRRI, in the Philippines. But as a class, the germplasm collections and their storage places need much better support than they presently receive. Many of the good collections are now old enough - have been stored long enough seeds. that unless money, people and facilities are soon made available, the banks will contain only collections of dead ... ~. ~ -7 A Threat: Germplasm Embargoes An additional problem has arisen in the past ten years, particularly in regard to collections in developing countries. The idea has been developed that since germplasm diversity is critical to advances in plant breeding, and since developing nations often are rich sources of land race diversity of the major crop plants (maize in Mexico, sorghum in Ethiopia, etc.), the developing nations as a class have a valuable resource, highly desired for immediate (and future) use by the developed nations. Further, it is said that up to now, the developed nations have had free access to the germplasm of the developing nations, clearly an unjust situation. Therefore, it is now time that the .developing nations were paid a fair compensation for the products - primitive farmer varieties - that their people have spent thousands of years in developing. Since no compensation has bee~ forthcoming, the developing nations will need to withhold their resource - landrace varieties - from use, until a fair price is paid for them. A chief exponent - and promoter - of this idea has been a Canadian named Pat Mooney, working through a coalition of international organizations. The forum for implementation of the idea has been the the assembly of the Food and Agriculture Organization (FAO) at Rome. The assembly is composed, more or less like that of the United Nations, of representatives from each of the member nations, each with one vote. In 1983 the assembly of the FAO adopted an "Undertaking" (a non-binding agreement) which incorporated most of Mooney's. program for action, particularly his advocacy of FAO control of all germplasm banks coincident with abolition o~ private -or institutional - ownership of any kind of germplasm, including advanced breeding materials. -8 unspoken in the Undertaking, but implicit, is the understanding that developing nations will, by means of some kind of embargo, stop all future collecting expeditions or any kind of exportation of germplasm from their countries, until satisfactory prices can be put on material they now have or bave furnished to the developed world in previous years. (In actuality some countries, such as Ethiopia, have already imposed a de facto embargo on export of collections or other germplasm. They are acting on the principles enunciated above, without waiting for the FAO's formal imprimatur.) This simplified recounting of events of the past few years should be qualified by saying that not all representatives of all developing nations agree with the FAO undertaking, nor with all of Mooney's suggestions. And some representatives of some developed nations agree with the Undertaking and with Mooney. But by and large, lines have been drawn, and there is a possibility that collection of landrace germplasm, indeed, international movement of any kind of germplasm, may be restricted in the future as a result of a confrontation between developing and developed nations. Possible Consequences of a Germplasm Embargo Such r~striction would not be f~tal to progress (near-term, at least) in plant breeding in developed nations such as the United States, since we have large collections on hand, and bave access to collections in other developed nations, as well. But in the long eun, only haem would come to plant breeding in our country, as well as in other developed nations, foe sooner OE later we would feel the need for additional germplasm. But even greater harm would come to plant breeding in many developing countries, especially in the short run, since their immediate, critical needs are for sources of modern high yield, -9 stress resistant germplasm, to introgress into adapted materials for development of new high performance varieties. (Their breeding programs are just now getting underway and will need large infusions of germplasm from diverse sources.) Of course, any informed plant breeder knows intuitively that no country in the ~orld has all the ~ermplasm it needs, whether for pest resistance or for a myriad of other useful traits. Data summaries and analyses bear out this intuitive conclusion. For successful plant breeding to be carried out in any country, a continuing bidirectional flow of germplasm is essential. All countries are "gene poor" in some respects but "gene rich" in other respects. International restrictions on germplasm movement would cripple plant breeding in developing and developed nations, rich and poor alike. The Second Major Source of Germplasm: Elite Cultivars The other major source of germplasm, essential for progress in today's plant breeding, is elite, adapted cultivars. Elite cultivars (and inbred lines of the hybrid crops), crossed in combinations that balance strengths and weaknesses, are used as parental breeding stocks. Technically speaking, such breeding is one kind of narrow-based recurrent selection. It is often called "pedigree breeding". All major u.s. crops have depended on pedigree breeding for the bulk of the advance in productivity of these crops for the past fifty years. (Broad-based population improvement, particularly in maize, has made important contributions and will be increasingly important in future years, but even this kind of breeding has as its end point the development of homozygous inbred lines which then are new sources of germplasm" for pedigree breeding.) II -10 Originators of Elite Cultivars and Inbred Lines For more than fifty years, production of new cultivars and inbreds in th~ u.s. has bee~ done primarily by breeders at public institutions, supported by the land grant universities and the USDA. Thecultivars and inbreds released by these public servants have been made available to all who wished to use them, on the assumption that since public tax monies supported the work, its products should be made available to the tax-paying public. Perhaps more important was the assumption that the public good was best served by making these products freely available to all those able to use them. An important additional source of inbred lines of mai~e and sorghum (and recently of sunflower) has been the private sector. The ability to maintain the inbred parents of hybrid crops as trade secrets attracted private enterprise to breeding and sale of hybrids, early in the developmental stages of plant breeding (as early as the 1920s, for maize). Numerous proprietary inbred lines were developed, useful as germplasm sources for further breeding as well as for use in proprietary hybrids. Kowever, because of the commercial necessity of keeping control of these privately developed inbred lines, they were not willin9ly made available to the public for breeding or other purposes. Thus, pedigree breeding with private inbred lines was, in theory at least, restricted to the pool of breeders within each company. Once hybrids made up of one or more private inbreds were offered for sale, however, the genes in the privateinbreds were available to the public. That is, proprietary hybrids, sold on the market for farmer use, were useful and legally available sources of germplasm. The heterozygous nature of the hybrid plants required somewhat different breeding methods, compared to breeding with inbred lines or with homozygous cultivacs such as wheat and soybeans, but the genes were nevertheless available for. use by all. -11 Private enterprise has more recently engaged in breeding and sale of wheat and soybean cultivars, encouraged by the Plant Variety Protection Act (PVPA), which restricts reproduction and sale of protected cultivars to owners (or assignees) of the cultivars. The entry of private enterprise into breeding these. self-pollinated crops has not changed the nature of their germplasm availability, since breeding is possible - and permitted by law - once protected cultivars are offered for sale. (Although private breeders of these crops safeguard their advanced breeding materials as trade secrets, public breeders likewise restrict exchange of. advanced breeding materials.) Thus, a diverse collection of sources of germplasm for breeding is found in the many kinds of cultivars, developed and released by public and private breeders. The extent of genetic diversity among these cultivars has major effect on the amount of future genetic gain. It also affects the stability of the naiion's crop production, to the extent that it broadens the germplasm base of crops growing o~ the farm and thereby strengthens the protection affoided by genetic diversity. To maintain and identify the diversity of germplasm in our cultivar base is the responsibility of the nation's plant breeders, public and private. Reduction in Funds for Public Plant Breeding In recent years, funds for public plant breeding have been severely restricted. The USDA officially will no longer engage in cultivar development and r~lease, except in crops where no other organizations are willing to do so - and where there is a need for cultivar development. The universities, through their . state experiment stations, have. annually had fewer funds and more needs for them outside the field of plant breeding and /,: cultivardevelopment. -12 The net result has been a reduction in plant breeding _ particularly, in cultivar development - by public breeders. This reduction has given rise to two reactions, in rather different directions. A New Role for Foundation Seed Companies Foundation seed companies, formerly dependent on public institutions for the inbred lines and pureline cultivars that they increased and sold, have instituted their own breeding and testing programs. They now offer a line of products that derives from their own breeding or that consists of products of other private breeding programs made available to the foundation seed companies through diverse arrangements. Mai~e and soybeans, in the U.S., have seen the greatest development in this direction. Services of the foundation seed companies are especially valuable to small seed companies - those with no or very little breeding of their own. A New Philosophy for Release of Publically-developed Cultivars through the state agricultural They have tried The universities - experiment stations - have instituted new programs, aimed at maintaining funds for applied plant breeding. various methods, all of them essentially mechanisms that would involve release of publically-developed cultivars to a selected group of private seed firms in return for some sort of royalty payment. To date, none-of the methods have been very successful, financially. (This may have been because of poor business organization or it may be that the university-released varieties simply were not competitive products.1 -13 A new plan, presently in progress of discussion and possible implementation by several midwestern universities, is to release certain cultivars of wheat and soybeans without restriction, that is, to make them available, for a royalty, to all who wish to reproduce and sell them - with the explicit ~nderstanding that each cultivar could be sold under a mul~iplicity of brand names, one for each different seed company. Technically, cultivars could be sold as "brand name (variety not stated)" in those states that allow this procedure. This would, theoret~cally, give individual companies incentive to advertise and sell their sample of the cultivar as a unique product, assuming the farmer did not know the same product was on the market under a diversity of other names. The net result, hopefully, would be broader promotion of the cultivar, giving rise to greater sales and more royalties to the universities. Another result would be inability of farmers to intelligently plan for genetic diversity in their individual farming operations. The genetic base of crops on u.s. farms would be much narrower than might be expected from the number of named varieties on those farms. Multip~eNames for Hybrids This potential, for confusion as to the real breadth of genetic base in u.s. farm crops, is not a new problem, however. It has long been common "trade knowledge" that certain maize and sorghum hybrids - crosses of publically available inbreds - were being sold under many different designations. It is of course possible and probable that in some instances minor changes in one or both inbred parents, or differences in seed production ~14- practices, have produced hybrids with different performance. But theory predicts, and thorough performance trials indicate, that usually such differences are slight, perhaps not consistently distinguishabl.e from unpredictable variation among seed lots of a given hybrid. Multiple naming of wheat and soybean varieties has also be~ri practiced in the private seed trade. And "sister-lines" (nearly identical selections separated at a late stage in the selfing process) also are sold under several brand names. Thus, the private firms have pointed the way for the universities. Until recently it has seemed fruitless and scientifically not feasible to try to prove that anyone hybrid or variety, with or without minor modification, was being sold under many designations. But new technologies such as allozyme electrophoresis and reversed phase high performance liquid chromatography (RP-HPLC) are now on hand and can be used for precise comparisons of genotypes at numerous loci among maize hybrids of unknown pedigree. (Similar techniques are also being applied to other crops: wheat, barley and potatoes, for example.) Results of examination these with new techniques cannot prove identity but they can point out near identity or the strong possibility of full identity. One such study has indicated that 47% of 138 maize hybrids (the 138 hybrids represent about 75% of the total number of hybrids on the market in the u.s. today) can be put into 10 different groups of tight allozymic relationship. In the largest group, 18 hybrids were virtually indistinguishable, on the basis of combined data from use of the allozyme and RP-HPLC techniques. An even newer technology: restriction fragment length polymorphisms (RFLP), can array some 200 or more DNA fragments per variety thereby giving a test of such sensitivity that ~-- ----- -~----- c -15 virtually unequivocal tests of identity can be made. This technology now exists for maize and can be applied to the question of multiple designations for single hybrid genotypes. Thus, it will soon be possible for concerned individuals or organizations to get powerful assistance in identifying a genetically diverse set of cultivars, within any of the major field crops. Knowledge of such diversity will not (as yet) predict performance of the cultivars,but it will help farmers avoid repetitious and unintended planting of a single genotype under several brand names. How Multiple-Naming Can Reduce Genetic Diversity and Rates of Genetic Advance Of course, whether or not hybrids and self-pollinated cultivars are correctly described as to relationship has little overt effect on use of these sources of genetic advance for the future. Once on the market, all are available for breeding. But in a more subtle way the pressure to conform - to base all breeding and marketing efforts on a narrow circle of favored cultivars - will gxeatly restrict the possibilities for genetic advance in the future. If the number of genuinely different cultivars is small, -the number of breeding populations derived from their crosses will be small. If breeders generally devote their efforts only to making the smallest possible distingtiishable modifications in a few favored genotypes, or if seed companies avoid breeding effort in favor'of simply renaming cultivars or hybrids as "unique" products, the gene pool of advanced breeding stocks_ plus working hybrids and cultivars will be greatly restricted. Advances due to novel genetic combinations will depend on that small remainder of breeders whose genetic source materials are genuinely - and broadly different. -16 Legal systems have been devised that are intended to prevent duplication in naming and thus prevent "i~Visible" narrowing of the genetic base. The Federal Seed Act, for example, states that a given variety shall not be sold under more than one name. Plant variety protection (PVP~ of cultivars of the self-pollinated crops can also prevent sale {by unauthorized parties) of a protected cultivar under more than one name. Likewise, PVP for inbred lines of maize or sOrghum can add new levels of protection to that already afforded by trade secrecy, by rights of common law ownership, or by mutually understood policies of professional restraint - unwritten codes of professional ethics. Thus, unique cultivars and hybrids can stay unique, when PVP is used - and enforced. But even PVP leaves room for legal sale of nearly identical copies of highly successful cultivars or hybrids. If a candidate for protection is distinctly different from other described lines - no matter how minor the difference, or how insignificant its effect on performance - the candidate then is eligible for protection, providing other requirements such as uniformity and stability are also satisfied. Thus PVP, while helping to prevent sale of a unique cultivar under more than one name, nevertheless does allow sale of near-identical copies of protected cultivars. In summary, several examples show that diversity of a major u.S. source of useful germplasm - elite cultivars and inbreds may be restricted in the future, in part because of policy decisions by some state agricultural experiment stations and in part because of actions of some private seed firms~ The extent of the reduction in diversity will be hidden from public notice, because diversity of brand names will not be a measure of the diversity of cultivar genotypes. The genetic base of major u.s. crops: wheat, soybeans, maize and sorghum will be narrower than the numbers of brand names would suggest. .I I . , !I -17 If plant breeding's two primary sources of useful germplasm - exotic varieties and elite cultivars - do indeed have strong possibility of being restricted in availability and variety in the future, what can be dorie to alleviate or prevent such problems? Better understanding May Le~d to Continu~tion of International Germplasm Exchange Dangers of international restrictions in germplasm exchange may now be lessening. As the circle of concerned people broadens, particularly in developing countries, understanding of the true g~rmplasm needs of developing countries has deepened. More to the point, knowledgeable plant breeders in developing .countries have known all along that for indigenous plant breeding to prosper, free and two-waygermplasm exchange was essential. Their counsel is now, gradually, reaching FAD delegates and/or policy makers in their governments~ Intense debate still continues, however, complicated in many cases by arguments as to the propriety of plant variety protection in developing countries, in developed countries, or even the morality of any kind of private breeding and commercialization of seeds. Since seeds are the foundation of the world's food supply, it is said (by some) that they must be kept under public control. Such arguments come from idealogues in the developed countries, for the most part. We may some day have the interesting spectacle of a coalition of representatives of developing nations and the private seed industry standing in opposition to a coalition of representatives of social activists from the developed countries. -18 (Criticisms, recounted earlier in this paper, of plant variety protection -criticisms that are also valid in Europe have already been made in much the same form to support the case of those opposed to any private ownership of plant germplasm. This illustrates the tangled nature of the situation.) Improvements in the National Plant Germplasm System The intensity of the international debate over ownership of indigenous plant germp1asm has stirred administrators and legislators at the federal level sufficiently that better funding has been given - or is promised - to some parts of the NPGS. But much more is needed. Further, it appears to an outsider that thorough reorganization clarifying and strengthening of lines of authority and responsibility in the NPGS - is needed. Such reorganization could itself give rise to increased productivity. Fortunately, it appears that such reorganization has begun - a good sign that the USDA is serious about improving the NPGS. Bringing More Realism to plant variety Protection Regulations The problem of concentration of breeding on a small base of favored cultivars is a thocny one. Leaving aside the reasons . listed earlier for expecting a smaller genetic base than the number of cultivar names would suggest, one must also keep in mind the propensity of breeders to breed with only the most elite of the elite. To do this greatly increases chances of short-term success for the individual breeder, but tends to narrow the germplasm base. To stimulate a breeder to take chances with genetically different germplasm, there must be a reward for turning out -19 subs'elffl'f!f'-;1;'df-lferent cultivars. The present system of plant vart.,~~~;~i~Ctlon does not necessarily give such a reward; if anything it 't:e1Jta the breeder that copying - making near duplicates of elite cultivars through backcrossing and similar breeding techniques is an acceptable route. There is some possibility of adding an element of common sense (from a plant breeder's point of view) to plant variety protection regulations, by refining the definition of "significant difference" (often called "minimum distance") to ensure that new varieties cannot simply be near-duplicates of already existing ones. Schemes are now proposed for maize, for example, which would invoke a multi-step process in reaching a decision about significant difference. If a new line was phenotypically identical to a protected (or previously described) line for all but one or two traits, the two lines would then be compared more precisely, using new analytical tools like allozyme electrophoresis or RP-HPLC. If they again differed by less than a predetermined amount, they would be hybridized and the resulting Fl and F2 generations would be examined for degree of heterosis. An arbitrary point would be set, for each level of testing, that would be the basis for deciding whether differences were sufficient for protection of the new line. The chief difficulty with this proposed system would be to get agreement on the limits - on "how much difference is enough." Should new lines be approved only if they differ significantly in important agronomic traits? Or should an arbitrary level of genetic dissimilarity be chosen, without reference to agronomic value? by crop basis. Certainly, if the system should ever be put in place, standards would have to be set on a crop -20 And, as noted earlier, the u.s. Plant Variety Protection Act does not allow use of such a system. There is some possibility that such a system could be put in place in at least some European countries, but up to now most of them have done little more than discuss the possibility - and the problems. In one country, however (France), officials are beginning to look at new schemes - with data to support them - that would allow use of a more realistic minimum distance standard. utility Patents for Plants - Advantages, Disadvantages A new development in the United States - the result of a recent decision (ex parte Hibberd et all in the u.s. patent office - may allow use of a broader system of protection of cultivars, inbreds and hybrids. So~called "utility patents", the kind used to patent new mechanical and chemical products for nearly 200 years, can now be applied to plant varieties. The "doctrine of equivalence" in utility patents uses broader standards than those now written in the Plant Variety Protection Act, and it perhaps can effectively protect a patented cultivar against unreasonably close copies. ,/ But the possibility of utility patents for plants, plant parts and plant genes poses a new problem. Can a gene - or a plant part - be protected down through generations of breeding, for the life of the patent on that gene or plant part? If so, will no one but authorized individuals be able to lawfully breed from a cultivar which contains the patented gene or plant part? If this should be the case, the germplasm base for plant breeding could be drasticalll re~~c~? o ,-_~. /,J,_.j_) ,Ii j . / _~) f, / j ( _-> ,4 Cd j)J--j ~,. .- - - -. .1 ,"' J ~ < 1 ~l."' ..---. .,, ,;} / J , How to Fund Public Plant Breeding from Private r Sources: A Dilemma --~l /(/ . " ;-';' _ _ "'.- ; .. :" ~ ./ .: .......... ,- ' ,, -21 The problem of reduced funding for public plant breeding is perhaps even less likely to be solved. In theory, public institutions can develop basic breeding pools, introgress needed genes from exotics and increase the existing body of breeding theory; private industry can in turn build on the contributions of the public sector as it develops and markets finished cultivars. By means of such a partnership the chain of progress, from basic research to cultivar release, will be unbroken. In practice, private industry cannot afford to do breeding of all crop plants. Public breeders will need to continue breeding in some species or no one will do so. Further, to fulfill their function of teaching and training future plant breeders, the universities must continue to do at least some serious plant breeding. In practice, USDA funds increasingly are pulled away from even basic plant breeding, unless .it is related to biotechnology. And, in practice, state legislatures - the other chief source of funds for land grant institutions - are generally not interested in funding conventional plant breeding at all, although they, too, are often willing to fund biotechnology research in support of plant breeding when they believe it may expand the industrial base - and thus the economic development of their state. (Biotechnology is often described as a source of new germplasm for plant breeding . . Biotechnology will' indeed generate important and revolutionary new genetic combinations and even new genes. But such contributions will supplement, not replace, elite and exotic varieties as major sources for genetic advance.) ~22- Experiment station. administrators, understanding very well that the chain - from fundamental research through cultivar o release - cannot be broken, are trying to bring in needed funds for plant breeding, by means of commercial activity. (They consider that they are not ground breakers at the universities in instituting such activity - other colleges and departments have long had contract and royalty arrangements with business v interes~s.) But the methods devised, to date, are either unable to generate enough cash or may even be counterproductive to the ultimate end - to provide a broad base of superior cultivars to the American farmer. Recommendations for Action have now painted a very gloomy picture of the future sources of genetic advance for plant breeding, congentrating for the most part on prospects for the u.s. Perhaps I should stop at this point, but I will instead take the dangerous path of making some recommendations and predictions for the future. I Regarding the possibility of embargoes on international exchange of germplasm, I think federal administrators, land grant university officials and private industry need, cooperatively and individually, to increase their contacts with administrators and politicians in developing countries who are concerned with agriculture. We already have good communication with the developing world's plant breeding community, a byproduct o~ the education of many of them in this country, of cooperative or exchange...

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Texas A&M - SCSC - 641
IPERSONAL PERSPECTIVE'Plant Breeding, an Evolutionary ConceptDonald N. Duvick* INTRODUCTIONRecorded history, and the an:haeologicaJ record tell us that plant culture has been an Inseparable part of human society for the past several millennia. Repeate
East Los Angeles College - LS - 72682
Protein Purification Handbook18-1132-29 Edition ACProtein PurificationHandbookHandbooks from Amersham BiosciencesAntibody PurificationHandbook 18-1037-46The Recombinant Protein HandbookProtein Amplification and Simple Purification 18-1142-75Prote
Texas A&M - SCSC - 641
, .CANOLA - A SUCCESS STORY IN BIOTECHNOLOGYWillie H. LobCargill Foods, Minneapolis,~,USAAbstract The first generation of biotechnology-derived oils have already been commercialized. Using advanced plant breeding techniques, specialty canola varieti
stonybrook.edu - MATH - 626
Numerical AnalysisLloyd N. Trefethen Oxford University May 2006AcknowledgmentsThis article will appear in the forthcoming Princeton Companion to Mathematics, edited by Timothy Gowers with June Barrow-Green, to be published by Princeton University Press
East Los Angeles College - AEI - 639
Shifting the Balance of Power?Information Society and its Impact on Trans-national and EU GovernancePaper to be presented at the Ionian Conference Facing the Challenges of the New Millennium Group 3 Citizenship and Governance May 20-22, 2000 Corfu, Gree
Colorado - MOREY - 4535
Ryan Duncan December 16, 2003Reintroduction Of Wolves: Market Failed or Market Failure?I was recently asked to conduct some research on the reintroduction of wolves into the Colorado area. Wolves used to be a normal part of the Colorado landscape but ha
East Los Angeles College - OTH - 049
Oracle DatabaseLicensing Information 10g Release 2 (10.2)B14199-06July 2006Oracle Database Licensing Information, 10g Release 2 (10.2) B14199-06 Copyright 2004, 2006, Oracle. All rights reserved. Contributors: Francisco Abedrabbo, Manmeet Ahluwalia, M
Iowa State - EE - 330
EE 330 Lecture 8IC Fabrication TechnologyQuiz 7The layout of the cascade of two CMOS inverters is shown. It has some layout errors. Identify them.And the number is .183564972And the number is .1 74938625Quiz 7 Solution:- Bulk Conne
illinoisstate.edu - ECO - 401
EC0 401Practice Exam #320091. Which of the following individuals would not be included in the civilian labor force? a) someone who is officially considered to be unemployed. b) a 32 year old man who is institutionalized. c) a student who is working par
illinoisstate.edu - ECO - 401
ECO 401Practice Exam #220091. Which of the following statements is correct? a) Economic profit is equal to the difference between total revenues and total explicit costs. b) The difference between economic profit and accounting profit is equal to the t
illinoisstate.edu - ECO - 401
ECO 401Practice Exam #120091. Which of the following is not a factor of production? a) land. b) money.c) labor. d) entrepreneurial ability.2. What is the Fundamental Premise of Economics? a) Natural resources will always be scarce. b) Individuals are
illinoisstate.edu - ECO - 105
ECO 105Study Guide for Exam #4Chapter 16 1. List and describe the principal functions of money. 2. List and define the main components of M1 and M2. What role does liquidity play in determining in which measure of the money supply a particular form of m
illinoisstate.edu - ECO - 401
ECO 401Study Guide for Exam #3Chapter 15 1. Explain how the size of the labor force is determined, i.e., who is included in the labor force, who is not, and why? 2. Given the appropriate information, be able to calculate the unemployment rate. 3. What c
illinoisstate.edu - ECO - 490
Prof. CarlsonStudy Guide Topic 5ECO 490I. MAJOR TOPICS: Listed below are the major topics in this chapter. You should have a complete understanding of each of these topics and be able to conduct analyses that utilize the corresponding concepts, princip
illinoisstate.edu - ECO - 490
Prof. CarlsonStudy Guide Topic 4ECO 490I. MAJOR TOPICS: Listed below are the major topics in this chapter. You should have a complete understanding of each of these topics and be able to conduct analyses that utilize the corresponding concepts, princip
illinoisstate.edu - ECO - 490
Prof. CarlsonStudy Guide Topic 3ECO 490I. MAJOR TOPICS: Listed below are the major topics in this chapter. You should have a complete understanding of each of these topics and be able to conduct analyses that utilize the corresponding concepts, princip
illinoisstate.edu - ECO - 490
Prof. CarlsonStudy Guide Topic 2ECO 490I. MAJOR TOPICS: Listed below are the major topics in this chapter. You should have a complete understanding of each of these topics and be able to conduct analyses that utilize the corresponding concepts, princip
illinoisstate.edu - ECO - 490
Prof. CarlsonStudy Guide Topic 1ECO 490I. MAJOR TOPICS: Listed below are the major topics in this chapter. You should have a complete understanding of each of these topics and be able to conduct analyses that utilize the corresponding concepts, princip
illinoisstate.edu - ECO - 105
ECO 105: Principles of Economic Theory Chapter 9 (Mankiw text): Application: International Trade I. The determinants of trade A. The world price and comparative advantage 1. Compare the world price of a good to the domestic price. If the world price is hi
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 11: Housing and Neighborhood DevelopmentFundamentals of housing economics A. Supply and demand revisited 1. Determinants of supply and demand 2. Measuring quantity a. &quot;Units of housing services&quot; 1. Factors that influence housing services
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 8: Issues in Economic Development PracticeBenefits from economic development A. Objectives of economic development policies 1. Job and income creation a. Focus is on the link between job creation and real incomes i. per capita income gro
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 5: Understanding Economic StructureAgglomeration economies Essentially, were talking about cost reductions as a result of economic activity that occurs in one place. As we will see, there are a number of sources of agglomeration economie
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 6: Regional Growth and DevelopmentStages of growth A general view of the stages of growth can be summarized as follows: 1. An area begins by exporting one or a few products 2. the local economy becomes more complex with more diversified
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 4: Market Areas and Economic Development StrategiesDemand and market areas A. Demand in a spatial setting 1. The essential point here is that, from the customers perspective, the effective price of a good is a function of both the sellin
illinoisstate.edu - ECO - 490
ECO 490Topic 3: Business Location, Expansion and RetentionIn order to be able to influence the locational decisions of firms development practitioners need to understand the factors that influence decision makers. I. Locational Factors A. Inertia (appli
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 2: Three Fundamental and Recurring IssuesThree fundamental issues in development planning are considered in this chapter: 1. Unemployment and low wages 2. Externalities 3. Public sector decision makingII. Unemployment and low wages A. A
East Los Angeles College - LG - 603
Activity Theory and L2 learningLG 603 Applied Linguistics and SLA Research Adela Gnem Session 2Activity TheoryActivity Theory is an approach or framework for studying different kinds of human practices as development processes, with both individual and
illinoisstate.edu - ECO - 490
ECO 490 I.Topic 1: Economic Development and Market LogicHow economists view the world A. Models and assumptions 1. A good model is an abstraction/simplification of reality 2. The importance of the ceteris paribus assumption 3. A &quot;good&quot; model enables us
East Los Angeles College - LG - 603
SCT and L2 pedagogical practiceLG 603 Applied Linguistics and SLA Research Adela Gnem Session 3Dynamic Assessment (DA)Full picture of someones capability: Solo performance on a test Performance with assistance from someone else; and TransferabilityDA:
SUNY Stony Brook - ISE - 208
GUI programmingGraphical user interface-based programming1Windchill Windchill There are several formulas for calculating the windchill temperature twc The one provided by U.S. National Weather Service and is applicable for a windspeed greater than fou
Monash - SEMESTER - 7087
Local Government Law[LAW7087]Unit GuideSemester 1, 2009Faculty of LawLAW7087Unit GuideSemester 1, 2009Prepared by: Mark Hayes Monash University Law Chambers 472 Bourke Street MELBOURNE VIC 3000Produced and Published by: Faculty of Law Monash Univ
Lake County - CS - 598
Impact of Instruction Latency Architecture, Algorithms, and Programming Models: Instruction PipeliningWilliam Gropp Programming languages usually present a model in which one line (or operation) completes before the next startsThis is not what happens
Lake County - CS - 598
Final Comments on Cache Oblivious Algorithms; Parallel PerformanceWilliam GroppLimitations of the Cache Oblivious Approach Why hasn't CO taken over the world?Constant terms are important For example, the number of cache misses in the transpose algori
Lake County - CS - 598
Virtual Memory Computer Architecture and Performance: Virtual MemoryWilliam Gropp So far, weve assumed that the process is addressing memory In most systems, (user) processes use virtual addresses! Gives the process the illusion that itdirectly address
illinoisstate.edu - PSY - 138
Name _ Lab 22 Worksheet (1) A major university would like to improve its tarnished image following a large oncampus scandal. Its marketing department develops a short television commercial and tests it on a sample of n = 7 subjects. People's attitudes abo
illinoisstate.edu - PSY - 138
Ruler # 1 _ 0Ruler # 2 0 _ 1 __ 12 _ 3 __ 24 _ 5 __ 36 _ 7 __ 48 _ 9 __ 510 _ 11 __ 612 _
Villanova University - ECE - 4790
BG Mobasseri-1-Chapter2_part2BG Mobasseri-2-Chapter2_part2BG Mobasseri-3-Chapter2_part2BG Mobasseri-4-Chapter2_part2BG Mobasseri-5-Chapter2_part2BG Mobasseri-6-Chapter2_part2BG Mobasseri-7-Chapter2_part2BG Mobasseri-8-Chapter2_part2
Villanova University - ECE - 4790
1997 BG Mobasseri1August 18, 19971997 BG Mobasseri2August 18, 19971997 BG Mobasseri3August 18, 19971997 BG Mobasseri4August 18, 19971997 BG Mobasseri5August 18, 19971997 BG Mobasseri6August 18, 19971997 BG Mobasseri7August 18, 1997199
Villanova University - ECE - 4790
SPREAD SPECTRUMHiding Information in noise1999 BG Mobasseri14/20/99Origins of Spread SpectrumlMilitary communication has always been concerned with the following two issues Security Jam resistancelIn civilian communications, above issues take on
Villanova University - ECE - 4790
DIGITAL MODULATIONS (Chapter 8)Why digital modulation?lIf our goal was to design a digital baseband communication system, we have done that Problem is baseband communication won't takes us far, literally and figuratively Digital modulation to a square
Villanova University - ECE - 4790
ECE 4790 ELECTRICAL COMMUNICATIONSSPRING `99 Inst.: DR. BIJAN MOBASSERIHow is Time Spent?l3 hours of lecture per week broken up into two 75 minutes (6pm-7:15pm) The rest of the time is used for Lab presentations and discussions Lab consists of experim
Villanova University - ECE - 2409
ECE 2409 Fundamentals of MATLABSpring 20002000 Bijan Mobasseri11/20/00Course organization and policiesl l l l l l lLecture in CEER 26 - 1:00pm-2:00pm Hands-on: CEER 118 - 2:00pm-3:00pm Turn in hands-on practice when leaving Homework is due weekly b
Villanova University - ECE - 2409
X(:, [1:2:end])=[]ADVANCED MATRIX OPERATIONS - PART III1999 B. Mobasseri1How do you do this??1999 B. Mobasseri2EMPTY MATRICESlStatement x=[ ] defines a zero by zero matrix; an empty matrix. This is different from clear x which zeros x. We can us
Bethel VA - MATH - 263
Factoring Expressions and Solving Equations11. At the prompt, type the following commands and press Enter : clear syms x expr1 = (x-1)*(x-2)*(x-3)*(x-4)*(x-5) expr2 = expand(expr1) factor(expr2) solve(expr2) . . . . . . . . . . . . . . . . . . . . . . .
CofC - CS - 129
An Integer Programming Model for the Sudoku ProblemAndrew C. Bartlett Amy N. Langville March 18, 2006Abstract Sudoku is the new craze in logic puzzles. Players must fill in an n n matrix, which contains some given entries, so that each row, column, and
Lake County - CONF - 134
Price and Profit: Investigating a Conundrum by Carl R. Zulauf, Gary Schnitkey, and Carl T. NordenSuggested citation format: Zulauf, C. R., G. Schnitkey, and C. T. Norden. 2006. Price and Profit: Investigating a Conundrum. Proceedings of the NCCC-134 Conf
Lake County - CONF - 134
Empirical Confidence Intervals for WASDE Forecasts of Corn, Soybean and Wheat Prices by Olga Isengildina, Scott H. Irwin, and Darrel L. GoodSuggested citation format: Isengildina, O., S. H. Irwin, and D. L. Good. 2006. Empirical Confidence Intervals for
Lake County - CONF - 134
Value of Single Source and Backgrounded Cattle as Measured by Health and Feedlot Profitability by Babatunde Abidoye and John D. LawrenceSuggested citation format: Abidoye, B., and J. D. Lawrence. 2006. Value of Single Source and Backgrounded Cattle as Me
Lake County - CONF - 134
An Assessment of the Livestock Mandatory Reporting Act by Clement E. WardSuggested citation format: Ward, C. E. 2006. An Assessment of the Livestock Mandatory Reporting Act. Proceedings of the NCCC-134 Conference on Applied Commodity Price Analysis, Fore
Lake County - CONF - 134
The Value of Information Provision at Iowa Feeder Cattle Auctions by Harun Bulut and John D. LawrenceSuggested citation format: Bulut, H. and J. D. Lawrence. 2006. The Value of Information Provision at Iowa Feeder Cattle Auctions. Proceedings of the NCCC
Lake County - CONF - 134
Farmers' Subjective Perceptions of Yield and Yield Risk by Thorsten M. Egelkraut, Bruce J. Sherrick, Philip Garcia and Joost M. E. PenningsSuggested citation format: Egelkraut, T. M., B. J. Sherrick, P. Garcia and J. M. E. Pennings. 2006. &quot;Farmers' Subje
Southern Illinois University, Carbondale - ECON - 113
Welcome to Professor Mitchell's Economics 113 e-Resource for &quot;Comparative Advantage and Trade&quot;!Absolute Costs (in hours) Fish Bread Richland 1 1.5 Poorland 3 2 Payoffs Without Trade: Richland $8 Yes! Poorland $4 Yes! The relative price of fish in: Richla
Southern Illinois University, Carbondale - ECON - 113
Welcome to Professor Mitchell's Human Resources Helper, Scenario B!Scenario B: The business can hire zero, one, two, three, or four workers. This scenario parallels that of Session 3 in Experiment 5 in that the firm may hire up to four workers. Step 1 Sp
Southern Illinois University, Carbondale - ECON - 113
Professor Mitchell's Economics 113 &quot;Shrinkage Calculator&quot;Specify a Seller Cost for each unit produced (a positive, whole number please) Specify an amount for the fine if this is an illegal good; zero if it is a legal good. Specify an amount for the suppl
Southern Illinois University, Carbondale - ECON - 113
Welcome to Professor Mitchell's Human Resources Helper, Scenario A! Scenario A: The business can hire zero, one, or two workers.This scenario parallels that of Sessions 1 and 2 in Experiment 5 in that the firm may hire only zero, one, or two workers. Ste
Southern Illinois University, Carbondale - ECON - 113
Professor Mitchell's &quot;Practice with [Sales] Taxes&quot; WorksheetDistribution of Demanders Buyer Number in Value MarketSupply, Demand and Competitive Equilibriu$50Profits of demanders Profits of suppliers Demand Supply C.E. without a tax ERR [ no tax yet ]
Southern Illinois University, Carbondale - ECON - 113
Econ 113, Spring 2007 May 9, 2007 Final Exam d, 125 pointsNeatly print your name . . . . . . . . . . . . . . . . . . . . . .You have 100 minutes to complete the exam; there are 8 problems. Points for each problem are given in parentheses with the proble
Southern Illinois University, Carbondale - ECON - 113
Econ 113, Spring 2007 May 9, 2007 Final Exam c, 125 pointsNeatly print your name . . . . . . . . . . . . . . . . . . . . . .You have 100 minutes to complete the exam; there are 8 problems. Points for each problem are given in parentheses with the proble
Southern Illinois University, Carbondale - ECON - 113
Econ 113, Spring 2007 May 8, 2007 Final Exam b, 125 pointsNeatly print your name . . . . . . . . . . . . . . . . . . . . . .You have 100 minutes to complete the exam; there are 8 problems. Points for each problem are given in parentheses with the proble
Southern Illinois University, Carbondale - ECON - 113
Econ 113, Spring 2007 May 8, 2007 Final Exam a, 125 pointsNeatly print your name . . . . . . . . . . . . . . . . . . . . . .You have 100 minutes to complete the exam; there are 8 problems. Points for each problem are given in parentheses with the proble
Southern Illinois University, Carbondale - ECON - 113
Econ 1132, Spring 2007 April 20, 2007 Exam 2d, 100 pointsNeatly print your name . . . . . . . . . . . . . . . . . . . . . . . . . .The exam ends at 11:50 a.m.; there are 5 problems. Points for each problem are given in parentheses with the problem numbe