NATIONAL RESEARCH SUPPORT PROPOSAL: NRSP-8
NATIONAL ANIMAL GENOME RESEARCH PROGRAM (NAGRP)
CRITICAL REVIEW
1. Summary of Accomplishments
The previously stated objectives of NRSP-8 were focused on support for the efforts of Species Coordinators who were to:
1. Serve as custodians of the species maps.
2. Serve as coordinators of the species databases
3. Serve as coordinators of shared material and information
4. Serve as coordinators for developing research priorities
Although the language of the previous NRSP-8 proposal states objectives in terms of the duties of the Coordinators, it is important to recognize that the accomplishments of NRSP-8 are actually those of the full Species Committees (along with the Technical Committee as a whole). Furthermore, it is impossible to fully separate these accomplishments from those of ARS, international, and other collaborators. This intertwining of accomplishments is inherent to the process of "coordinating" research goals. The accomplishments of each Species Committee relevant to the above objectives will be summarized.
A. Poultry
Objective 1. Poultry Genome Map(s). Two families, one developed in Compton, England (C) and one in East Lansing (EL), were designated as international reference mapping panels at the 1994 ISAG Poultry Genome Map Workshop. The EL subpanel of 52 backcross plus parental DNAs (of 400 available in the EL family) has been shipped to numerous laboratories in the UK, the Netherlands, Israel, Australia, Canada, and the US where typing is done, as well as in East Lansing. At present, the EL panel has been typed for 701 markers of which 677 are resolved into 40 linkage groups. The map coverage within linkage groups is over 3,150 cM which is about the estimated size of the chicken genome. There are 391 microsatellite markers which greatly enhance the utility of this genetic map for genome-wide QTL searches. Also, 139 Type I genes have been mapped which will aid in the development of a comparative map. Over 500 markers have been mapped on the C map, several of which are also on the EL map. This has enabled the coordinators, in collaboration with Nat Bumstead (Compton), to develop a consensus map of the EL and C families. As a result, linkage group assignments and the order of loci have been verified. Additionally, several previously separate linkage groups have been joined and chromosome assignments made. Chromosomes 1-8, 16 and Z have been identified on the genetic map; other chromosomes are microchromosomes which have not been distinguished cytologically. Independently, a large European commercial family has been used to map over 400 markers, mostly microsatellites. While this panel is not used by other labs, the resulting map is useful in improving details of the consensus map.
Objective 2. Database Development. ChickGBASE: Support was provided to Alan Hillyard on a consultant basis for the development of ChickGBASE. Dr. Hillyard also worked with Dave Burt, Alan Archibald, and others at the Roslin Institute to develop a new, more inclusive version of animal databases, Arkdb, which was recently released (discussed further below under Swine). ChickGBASE is available at http://www.ri.bbsrc.ac.uk/chickmap/chickgbase/chickgbase.html.
WWW Homepage: A WWW homepage for the Poultry Genome has been developed which links to ChickGBASE, the Roslin Institute homepage, the National Agricultural Library and many other genome mapping resources. The MapManager EL mapping database has been placed on the Homepage, so that users can immediately download and use the latest data. The consensus map has been widely distributed in hard copy and is also available on the Homepage. The Homepage also provides a list of published microsatellites, descriptions of our microsatellite kits, the latest cytogenetic map, and access to a host of other information. This site is available at http://poultry.mph.msu.edu.
Objective 3. Shared Material and Information. Reference Panel DNA: As noted above, DNA from the East Lansing international reference population has been sent to laboratories throughout the world. Microsatellite Primer Kits: Four kits of microsatellite primer pairs are now available for free distribution. The first of these is the Population Tester Kit. This contains 9 primer pairs which define microsatellites with high polymorphic information content (numerous alleles widely distributed in several populations). These primers are also fluorescent and can be multiplexed for analysis on automated DNA sequencers. Three large Comprehensive Mapping Microsatellite Kits containing a total of over 500 fluorescent primer pairs that define markers covering almost all of the chicken genome are also available. The Homepage has tables describing all kits along with a larger table with information on all published microsatellite loci (and several unpublished ones). Over 32 laboratories have been provided one or more of these kits.
Newsletter: The Poultry Genome Newsletter was begun in April, 1995, and continues to be published quarterly. The Newsletter is also distributed through our WWW Homepage, and electronically on the angenmap email discussion group to an even larger audience. Copies are also provided to the Poultry Breeders of America. Nomenclature: Genetic nomenclature issues continue to be resolved, as led by co-coordinator Lyman Crittenden and other Technical Committee members. These issues are critical to database management and ease of access. Meetings: Technical committee chairpersons and species coordinators have worked to organize annual NAGRP meetings (Salt Lake City in '93, Minneapolis in '94 and College Park in '95) in association with appropriate Regional Research committee meetings (NC-168 for poultry). Beginning in January, 1997, the NAGRP/RR annual meetings have been joint with the new Plant and Animal Genome meeting. Coordination support for and/or attendance at numerous other international and national meetings and workshops has also improved information sharing, worldwide.
Objective 4. Setting Research Priorities. Research priorities have been set and annually revised at Species Committee meetings. Priorities set during the past NRSP-8 period include: (1) Develop a consensus map of at least 200 microsatellite markers embedded in a dense map of other markers. (2) Develop resource populations designed to use that map to identify QTL for major economic traits. (3) Integrate the genetic, physical, and classical maps. (4) Develop resources for joint use such as primer sets and large insert DNA libraries. (5) Develop rapid mapping techniques and protocols that will allow economic and efficient use of genetic markers for a variety of purposes. (6) As resources permit, extend the map to other species of poultry in the following priority order: turkeys, ducks, and Coturnix quail as a model.
Progress: Goal 1 is complete and Goal 2 is well underway at several locations in the U.S. and elsewhere. Considerable progress has been made on Goal 3, although physical and classical maps still lag behind the genetic map. Work on comparative gene mapping of chicken loci using Type I markers is underway. Goal 4 has been achieved and resources continue to expand. Large insert libraries are complete or under construction. Goal 5 has been achieved through wider availability of primers (see above), automated equipment, and private genotyping services. However, the cost per genotype still needs to decrease for maximum utilization of new technology. Progress towards Goal 6 has been limited, probably due to lack of a critical mass of investigators working with these species.
B. Cattle
Objective 1. Cattle Genome Map(s). Several maps of the bovine genome have been generated, each with unique attributes and value. The coordinator keeps track of the status and source of each and makes this knowledge available to all of the US cattle genome mapping community. Three major linkage maps, from the IBRP and USDA-MARC reference families and from the IRRF, now include approximately 1800 loci providing an average resolution of nearly 2 cM. Comparative maps now include three published ZOO-FISH painting experiments, and extensive mapping of Type I markers is on-going. Bovine chromosome nomenclature recently has been updated. H. Lewin (IL) and colleagues have initiated a bovine EST (expressed sequence tag) project that will allow expressed genes in the cow to be matched to the very large human EST database and transcript maps.
Objective 2. Database Development. Much of the genome information heretofore disseminated personally is now available in a public database. Concentrating initially on genes and subsequently on anonymous DNA markers, the coordinator and S. Kata have entered 1231 loci into BovGBase with the corresponding 229 literature references. The home page provides links to other animal databases including other cattle databases as well as sequence databases. Public access to the Cattle Genome Mapping Homepage and to BovGBase is available at http://bos.cvm.tamu.edu/bovgbase.html. Also available through the BovGBase Homepage are two comparative maps ("cow on human" and "human on cow") illustrating known conserved synteny of cow relative to humans and mice and summaries of the ZOO-FISH experiments described under Objective 1.
Objective 3. Shared Material and Information. The coordinator continues to distribute International Bovine Reference Family Panel (IBRP) DNA to US scientists for linkage mapping. Six US laboratories utilized these families in the past year. The IBRP map now includes 740 markers, 150 of which are expressed genes. Thirty laboratories have contributed to the development of this map. Microsatellite primers synthesized by Research Genetics, Inc. and targeted for modest genome coverage are available for distribution. Three sets were distributed to US laboratories in the past year. The coordinator participated in defining 410 comparative mapping anchor loci and the design of primers within most of these loci which identify comparative anchor tagged sequences (CATS). Primer sequences are published and limited quantities are being distributed by the coordinator. A panel of 30 hybrid somatic cells defining every bovine syntenic group has been assembled and characterized by the coordinator. DNA from this panel is available to US scientists. A panel of radiation hybrid cell lines has also been developed. Large insert BAC clone libraries (about 3 genome equivalents or ca. 60,000 clones) have been made in the labs of Drs. Taylor and Davis (Texas A&M) for cattle, pig, sheep and horse (see below), and these are publicly available.
Objective 4. Setting Research Priorities. Research priorities have been set and revised at annual Species Committee meetings. These priorities include: 1. Further develop the cattle genome map, 2. Use genome maps to understand structure and organization of specific genes and gene families, and 3. Identify and determine the genomic organization and diversity of genes controlling physiologically important processes. Substantial progress has been made towards all these goals, as outlined under Objective 1 above. In addition, priorities are discussed at a variety of other meetings, some of which interface national priorities with those of the international community. For example, just in the last year, the coordinator made eight presentations on the status and future needs of genome research in agricultural species. These included talks at the HUGO Mapping Workshop, Brisbane; Iowa State University seminar series; American Genetic Association meeting, Atlanta; International Cytogenetics Conference, Zaragoza, Spain; and the International Society for Animal Genetics (ISAG), Tours, France.
C. Sheep
Objective 1. Sheep Genome Map(s). As of summer, 1997, 555 loci had been mapped to all 26 autosomes and both sex chromosomes of sheep. Of these assignments, 167 loci had been marked by physical mapping approaches and 388 via genetic linkage analyses. In the current linkage map, 57 are Type I genes and 331 are anonymous markers. The map spans nearly 2700 cM. With regard to comparative mapping, 185 markers are shared between the sheep and cattle linkage maps, and approximately 75% of bovine microsatellite primer pairs amplify a locus in sheep with about half of these also being polymorphic.
Objective 2. Database Development. SheepBase: The SheepBase informational database is now available, containing an up-to-date compilation of published data from sheep genome mapping projects, along with both physical and linkage maps of the genome, together with information on individual loci and associated references. Information is presented using the WWW interface and can be accessed through a number of nodes (e.g., http://dirk.invermay.cri.nz). The summer-'97 version of SheepBase contained 467 loci gathered from 114 publications. There were 173 Type I, 269 Type II, and 25 other loci in the database. An international editorial committee for SheepBase has been established: Grant Montgomery (New Zealand), Tom Broad (New Zealand), Noelle Cockett (Utah State) and Frank Nicholas (Australia).
Objective 3. Shared Material and Information. A collection of 223 ovine and bovine microsatellite primer pairs previously mapped to specific ovine chromosomes has been selected for distribution, to provide wide coverage of the sheep genome. To date, the primers have been distributed to 22 laboratories, worldwide. These markers are being used in projects investigating wool traits, carcass composition, parasite resistance, growth characteristics and single-gene defects. Coordination funds also supported the development of a large insert (BAC) sheep library that will be widely available for physical mapping projects. Three computer programs for genetic mapping of markers and QTL within large half-sib pedigrees have been developed for distribution. These ANIMAP programs are able to utilize paternal half-sib data in a computationally efficient manner. LODTABLE is designed to determine linkage groups by performing multiple pairwise comparisons between markers using specified recombination rates. MAKEMAP estimates recombination rates between loci within a linkage group using a maximum likelihood approach and gives the odds for different locus orders. The third program, MAPQTL, is designed to map quantitative traits to locations within genetic marker intervals. This program calculates various maximum likelihood estimators for the trait and gives the lod score at each location within the interval assuming a null hypothesis of no QTL present.
Objective 4. Setting Research Priorities. Research priorities have been set and annually revised at Species Committee meetings. Since cattle and sheep together form a joint Species Committee, the priorities are identical to those for cattle, except for the focus on sheep (1. Further develop the sheep genome map, 2. Use genome maps to understand structure and organization of specific genes and gene families, and 3. Identify and determine the genomic organization and diversity of genes controlling physiologically important processes.). Considerable progress has been made, as outlined above. In addition, these priorities have been further refined and communicated by participation of the Coordinator in a variety of meetings at which sheep genome mapping has been described. These include recent International Society for Animal Genetics meetings, the Allerton (Illinois) II Conference, and the joint NAGRP-Plant and Animal Genome meeting.
D. Swine
Objective 1. Swine Genome Map(s). A great deal of mapping data has appeared just in the past three years. This includes a rapidly expanding physical gene map and several genetic linkage maps. Three large linkage maps have been published: the USDA-ARS linkage map with over 1100 markers, the Nordic map with 236 markers and the PiGMaP map with 280 genes and markers. The last two of these maps are being revised and combined and there are over 650 genes and markers now on them (Archibald, personal communication). In total, these genetic linkage maps contain about 200 genes and 1700 other markers. The challenge is to incorporate this information into a usable database and to develop a composite map. Several groups have now produced consensus maps from the PiGMaP, Nordic and USDA maps, including maps for chromosomes 1, 5, 6, 7, 9, 13 and 14. All the maps are accessible through the WWW (http://www.public.iastate.edu/~pigmap). The physical map is also growing quickly and now includes over 600 genes and markers. Results from several labs have defined QTL locations on the genetic map and candidate gene approaches are underway to elucidate the nature of these QTL.
Objective 2. Database Development. PiGBASE: The Swine Coordinator has worked continuously with Alan Hillyard as a consultant and Alan Archibald (Roslin Institute) on the development of PiGBASE, the international swine mapping database. The newest version of the database (summer, 1997) is in the Arkdb format. It can be accessed at http://www.ri.bbsrc.ac.uk/pigmap/pigbase/pigbase.html or through the Pig Genome Homepage at Iowa State (http://www.public.iastate.edu/~pigmap). The new database design is constructed around the organizing concept of a locus and its position. Supporting evidence, both the mapping experiments themselves or the analyses performed to position the locus in the genome are fully documented and arranged in a modular fashion. This design concept allows the rapid addition of new types of both mapping protocols and analyses to the database without a massive redesign. Some of the mapping methods already incorporated in the design are FISH, SSCP, RAPD, SINE-PCR, PCR-RFLP, DSCP and microsatellites. Not only is the database being designed to provide the user with a complete suite of textual information, but supporting images (sequence gels, FISH, etc.) will be included as available. There are over 500 citations with over 1100 genes and markers listed. New graphical tools to manipulate and examine the data are being developed as well. These range from more efficient and new map displays to more flexible query generating tools. Lastly, electronic data submission will be strongly encouraged through the use of an online form. Also a new database (TCAGdb, The Comparative Animal Genome database) that will pull together the data from the single species databases for cross-species comparisons is nearly complete. All this work has been supported in small part by the Pig Genome Coordinator funds with the majority of effort coming from A. Archibald, Alan Hillyard, Jian Hu and Chris Mungall (BBSRC) and with funding from the UK Medical Research Council (BBSRC and the European Commission).
A World Wide Web (WWW) Homepage was established at Iowa State University early in the NRSP-8 project to support Pig Gene Mapping. This has been continuously enlarged and improved. The URL is: http://www.public.iastate.edu/~pigmap. Access via WWW allows users to obtain information such as: Coordinator Updates, addresses of pig gene mappers, genome newsletters including Pig Genome Update and the NAGRP newsletter, information on ANGENMAP, meeting schedules, available microsatellite markers, access to PiGBASE and other genome databases, USDA/ARS and international pig gene mapping information, and swine graphics. A Web page for students and producers is also available.
Objective 3. Shared Material and Information. A major effort to produce probes and obtain DNA for sharing has been made. Different panels of shared DNA primers include: 1.) 294 microsatellite marker primer pairs that are available for any lab doing gene mapping research (these were partially funded by NPPC and several swine breeding companies and have been shared with over 40 labs worldwide), 2.) approximately 150 fluorescent primer pairs (shared with over 20 labs worldwide so far), 3.) a separate set of 75 fluorescent primers for a new gene mapping technology, and 4.) a set of 30 expression primers for measuring mRNA levels by PCR of economically relevant genes. U.S. reference family DNA is available through each of the individual U.S. laboratories. PiGMaP family DNA has been imported from Edinburgh, Sweden, France, Germany and the Netherlands. This is available to individual labs that fill out a transport permit to move the DNA to their labs and has been shared with several U.S. labs. As mentioned previously, a swine BAC clone library has been constructed in the labs of Drs. Taylor and Davis (Texas A&M), and it is available from them.
Pig Genome Update is published bimonthly and distributed electronically to over 400 people. Angenmap, the gene mapping discussion group (angenmap@iastate.edu), continues to grow in activity and members (presently over 400). Several talks were given and meetings attended to discuss further coordination and cooperation. These included PiGMaP meetings, NC-210 meetings, ISAG meetings and swine industry meetings. At these meetings information on the pig gene mapping coordination effort in the U.S. has been presented and, in some cases, swine genetics research at all labs in the U.S. has been summarized for international audiences. The cooperation from PiGMaP labs and map coordinators is excellent. Two large conferences on gene mapping and quantitative trait loci, an international genetics conference and other worthwhile meetings have been attended and/or partially supported to improve information sharing. As described above, the annual NAGRP meeting has now been folded into a much larger Plant and Animal Genome meeting to enhance sharing of technology and data for all agricultural genomics.
Objective 4. Setting Research Priorities.. Research priorities set by participating scientists and stations are shared and often revised at annual Species Committee meetings. These priorities include: 1. Develop a comprehensive genetic linkage map of the pig, 2. Develop a comprehensive physical gene map of the pig, and 3. Establish procedures for preservation of genetic material in immortalized porcine cell lines. As described above, extensive progress has been made on the first two goals. Goal 3. has been made largely irrelevant as PCR-based gene mapping technologies have expanded, and has therefore been de-emphasized. There has also been a special effort to meet with individuals from swine breeding companies, the swine industry and other universities not already involved in pig gene mapping. In addition, several people from labs not previously represented were added to the NRSP-8 species committee. These efforts assist in insuring that the research prioritization process is based upon the needs of the communities that are being served.
E. Horse
The Horse Species Committee and Species Coordinator did not officially join the NAGRP until early in 1997, following an invitation to develop an addendum generated at the NAGRP meeting in October, 1995. Therefore, the progress listed below has been mostly in the last year prior to this renewal proposal.
Objective 1. Horse Genome Map(s). The collaborative effort to develop an equine gene map began in 1995 following an agreement by several research laboratories, worldwide, to cooperate in the construction of a linkage map for the horse. In connection with this program those laboratories in the U. S. applied to join the NRSP-8 in order to better coordinate the effort. DNA samples were collected from 12 horse families and distributed to laboratories for marker testing in 1996. The first linkage map, including 150 markers, should be reported in January 1998 at the annual NRSP-8/PAG VI meeting.
Microsatellite markers have been developed at the University of California (30+ markers), University of Kentucky (63+ markers), Texas A&M (11+ markers), Cornell University (50+ markers) and the University of Minnesota (16+ markers). These markers are being employed to construct a linkage map using local families as well as the international horse reference families. At the University of California a 34 member half-sib family was used for linkage studies and several new linkage relationships identified. At least 140 additional microsatellite markers are in development by participating laboratories that will contribute to the linkage map to be reported next January.
Synteny mapping is being conducted in California and Kentucky. Currently California reports 29 synteny groups employing 167 markers (microsatellites and coding genes) while the Kentucky has detected 16 synteny groups employing 100 markers (microsatellite and coding genes). Results from both labs are in agreement. Markers for coding genes developed by the National Cancer Institute (CATS, see Cattle section) were provided for synteny mapping. Workshop members from California, Kentucky and Massachusetts also led and participated in a workshop to develop a banded karyotype standard for the horse.
Several studies of individual genes (and associated Type I markers) are underway in the labs of many of the participants. Candidate gene approaches are also being carried out to identify and analyze the causes of equine disease traits. Collaborative work at the U. of Kentucky and Texas A&M in association with Washington State led to the demonstration of linkage between the gene for equine combined immunodeficiency disease and microsatellite locus HTG8. Similar approaches to other equine diseases are being carried out at the U. of Minnesota and elsewhere.
Objective 2. Database Development. Because no NAGRP funds have yet been available and because the horse genome project is in the early stages of data collection, database development has been delayed relative to those of the other species. However, during 1997 a horse database (see http://www.ri.bbsrc.ac.uk/) was established by Alan Archibald at the Roslin Institute in Edinburgh, Scotland, based on the Arkdb format discussed elsewhere in this report. The North American Editor for the database is the NRSP-8 Species Coordinator. This database will be a focal point of activity for the Equine Technical Committee if NRSP-8 is renewed.
Objective 3. Shared Material and Information. In connection with linkage map development described under Objective 1, DNA from 12 reference families was shipped to the Species Coordinator for distribution to test laboratories. The families include between 30 and 60 offspring. Each family is being tested for genetic markers, especially microsatellite markers, with the data being collated for linkage analysis. Families were provided by California, Kentucky and Minnesota. DNA from the families is being tested for markers at California, Kentucky, Minnesota, Texas A&M, Applied Biosystems and Shelterwood Laboratories. Laboratories developing markers for microsatellites are also providing PCR primers for use in synteny mapping with somatic cell hybrid panels (see Objective 1.) As mentioned previously, a horse BAC clone library has been constructed in the labs of Drs. Taylor and Davis (Texas A&M), and is available from them.
Objective 4. Setting Research Priorities. Priorities have been set at a variety of workshops prior to joining the NAGRP and at the Horse Gene Mapping workshop at the Plant and Animal Genome V Meeting (Jan. '97). They are similar to those described above for cattle and sheep: 1. Develop the horse genome map, 2. Use the horse genome map(s) to understand the structure and organization of specific genes and gene families, and 3. Identify and determine the genomic organization and diversity of genes controlling physiologically important processes. Progress is described above.
2. Degree of Accomplishments
The speed with which the fields of genomics and bioinformatics have advanced makes it difficult to compare the present state of animal genomics and that which existed at the time NRSP-8 was first proposed, approximately 5 years ago. At that time, genome maps for most of the species involved rarely exceeded 100 markers or so, there were no established reference mapping panels (so linkage relationships between markers used in one cross could not be accurately compared to those in another), few, if any, published microsatellites were available for these species, and, not only were there no genome databases, there was not even a public World Wide Web! Perhaps most important, it was nearly inconceivable that one could reasonably locate and hope to isolate a gene encoding a QTL, and there were no DNA-based tests in use for marker assisted selection.
In other words, most of what is now taken for granted (discussed at length above) was available only for humans or model organisms at that time, if, in fact, it were available in any species. Taken in this context, the overall rate of progress in mapping the genomes of agricultural animals has been truly impressive, although it is impossible to fully differentiate the accomplishments of NAGRP participants from other animal geneticists, both here and abroad. Perhaps most important, cattle, sheep, swine and chickens all have one or more well established reference linkage maps, typically with 1,000-2,000 total markers, most of these being highly polymorphic microsatellites. Internationally accepted reference populations, databases, and nomenclature have been established for all these species. Linkage maps have been correlated with one another and, in many cases, with cytogenetic maps. Several whole genome scans for QTL are underway and are beginning to bear fruit. Candidate gene searches to pin down these QTL have been successful or are well along the way. In some cases, the animal industries are already applying the fruits of this research to their breeding programs. Of course, our maps pale in comparison to those available for human and mouse, species whose genomes have received approximately two orders of magnitude more financial support. The good news is that the human and mouse genomes will be an immensely rich source of information for us through the mechanism of comparative genomics, with large blocks of conserved linkage extending from the human even out to the chicken genome. It is worth noting here that the NAGRP recently received one of the USDA 51st Annual Honor Awards for its contribution to this worldwide effort.
3. Areas of Further Investigation
There is no question that agricultural animal genomics will continue to progress rapidly based on the success of and support available to genome studies of human, mouse, and other model organisms. At least three critical features of the existing human and mouse maps are presently lacking: 1.) adequate marker density for fine structure mapping, 2.) physical maps based on ordered contigs assembled from large insert DNA libraries, and 3.) close alignment of conserved syntenic groups throughout our maps to those of the "information-rich" mouse and human genomes. These are among the primary goals for the future of the NAGRP and animal genomics. Animal geneticists also must keep pace with the rapid developments in the field of bioinformatics. All of animal science (indeed, all of biology) has entered a new age in which available information vastly exceeds the ability of any single scientist to comprehend and remember everything that is relevant. The goal of bioinformatics is to store that information in a form that is both regularly (and critically) updated, but at the same time is easily and immediately accessible. Further advances will surely be made in bioinformatics, some of which we may not yet be able to comprehend (as we could not have comprehended the present influence of the Web five years ago). It is critical to cost-effective progress that animal geneticists have access to and use these advances to help design and communicate the results of their research.
The areas of further investigation for NRSP-8 are outlined in detail in the proposal for its renewal. These objectives have been refined to take into account the rapid advance of the relevant sciences and to avoid duplication with efforts of Regional Research Committees and other groups. The objectives also take into account those of our foreign collaborators so that they will be mutually beneficial. These objectives are to:
1. Develop high resolution comparative genome maps aligned across species that link agricultural animal maps to those of the human and mouse genomes.
2. Increase the marker density of existing linkage maps used in QTL mapping and integrate them with physical maps of animal chromosomes.
3. Expand and enhance internationally shared species genome databases and provide other common resources that facilitate genome mapping.
With the help of cost-effective support through a renewed NRSP-8, collaborative efforts to reach these objectives over the next 5 years will be greatly enhanced. The development of high quality (integrative and comparative) animal genome maps can be viewed as a fundamental tool for animal breeding in the same way that the understanding of basic biochemical pathways is fundamental to the understanding of animal nutrition. In other words, genomics opens the way to effectively use the genetic resources of any species in the same way that physiology opens the way to use nutrition and husbandry to greatest effect. The ultimate beneficiaries will be consumers, who will be provided safer and more economical food and other products of animal agriculture based on this increased understanding and application of animal genomics.