Avian Genetic Resources at Risk: An Assessment and Proposal for Conservation of Genetic Stocks in the USA and Canada

Clockwise from above:• White Leghorn rooster from UCD 003, a highly

inbred stock that serves as genetic backgroundto a number of congenic chicken stocks.

• Red Jungle Fowl rooster from UCD 001, a highlyinbred stock derived from the wild chickenprogenitor species.

• Hybrid rooster from cross between Red JungleFowl (UCD 001) and White Leghorn (UCD 003).Offspring from the cross of this rooster and aWhite Leghorn hen were used as a referencepopulation in developing the chicken genomemap.(Photographs courtesy of J. Clark, University ofCalifornia–Davis.)

Front cover: Chicken, turkey, and quail eggs.(Photograph courtesy of J.M. Pisenti, University ofCalifornia–Davis)

AAAAAvian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Resources at Risk:esources at Risk:esources at Risk:esources at Risk:esources at Risk:An Assessment and PAn Assessment and PAn Assessment and PAn Assessment and PAn Assessment and Proposal for Conserroposal for Conserroposal for Conserroposal for Conserroposal for Conservation ofvation ofvation ofvation ofvation ofGenetic Stocks in the USA and CanadaGenetic Stocks in the USA and CanadaGenetic Stocks in the USA and CanadaGenetic Stocks in the USA and CanadaGenetic Stocks in the USA and Canada

AuthorsJ.M. Pisenti, M.E. Delany, R.L. Taylor, Jr., U.K. Abbott, H. Abplanalp, J.A. Arthur,M.R. Bakst, C. Baxter-Jones, J.J. Bitgood, F.A. Bradley, K.M. Cheng, R.R. Dietert,J.B. Dodgson, A.M. Donoghue, A.B. Emsley, R.J. Etches, R.R. Frahm, R.J. Gerrits,P.F. Goetinck, A.A. Grunder, D.E. Harry, S.J. Lamont, G.R. Martin, P.E. McGuire,G.P. Moberg, L.J. Pierro, C.O. Qualset, M.A. Qureshi, F.T. Shultz, and B.W. Wilson

Sponsored by theUniversity of California Genetic Resources Conservation ProgramUS Dept. of AgricultureUS National Science FoundationDept. of Animal Science, University of California–Davis

Report No. 20 � September 1999

Genetic Resources Conservation ProgramDivision of Agriculture and Natural ResourcesUniversity of CaliforniaDavis, California USA

This report is one of a series published by the University of California Genetic Resources Conservation Program (technical editor:P.E. McGuire) as part of the public information function of the Program. The Program sponsors projects in the collection, inventory,maintenance, preservation, and utilization of genetic resources important for the State of California as well as research and education inconservation biology. Further information about the Program may be obtained from:

Genetic Resources Conservation ProgramUniversity of CaliforniaOne Shields Ave.Davis, CA 95616 USA

Tel: (530) 754-8501FAX: (530) 754-8505email: grcp@ucdavis.eduhttp://www.grcp.ucdavis.edu

CITATION: J.M. Pisenti, M.E. Delany, R.L. Taylor, Jr., U.K. Abbott, H. Abplanalp, J.A. Arthur, M.R. Bakst, C. Baxter-Jones,J.J. Bitgood, F.A. Bradley, K.M. Cheng, R.R. Dietert, J.B. Dodgson, A.M. Donoghue, A.B. Emsley, R.J. Etches, R.R. Frahm,R.J. Gerrits, P.F. Goetinck, A.A. Grunder, D.E. Harry, S.J. Lamont, G.R. Martin, P.E. McGuire, G.P. Moberg, L.J. Pierro,C.O. Qualset, M.A. Qureshi, F.T. Shultz, and B.W. Wilson. 1999. Avian Genetic Resources at Risk: An Assessment andProposal for Conservation of Genetic Stocks in the USA and Canada. Report No. 20. University of California Division ofAgriculture and Natural Resources, Genetic Resources Conservation Program, Davis, CA USA.

© 1999 Regents of the University of California

The University of California prohibits discrimination against or harassment of any person on the basis of race, color, national origin,religion, sex, physical or mental disability, medical condition (cancer-related or genetic characteristic), ancestry, marital status, age,sexual orientation, citizenship, or status as a covered veteran (special disabilities veteran, Vietnam-era veteran or any other veteranwho served on active duty during a war or in a campaign or expedition for which a campaign badge has been authorized). UniversityPolicy is intended to be consistent with the provisions of applicable State and Federal laws. Inquiries regarding the University’s non-discrimination policies may be directed to the Affirmative Action/Staff Personnel Services Director, University of California, Agriculturaland Natural Resources, 1111 Franklin, 6th Floor, Oakland, CA 94607-5200. Tel. (510) 987-0096.

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TTTTTable of Contentsable of Contentsable of Contentsable of Contentsable of ContentsPreface ................................................ viiExecutive Summary.............................. ixAvian Genetic Resources Task ForceMembers............................................ xviiCharge to the Task Force .................. xviiiChapter 1: Introduction .........................1Chapter 2: Avian genetic diversity:Domesticated species ............................5

Target speciesTypes of genetic stocksResearch genetic stocksCommercial stocksFancy breeds and mid-level

production stocks

Chapter 3: Avian genetic resourcesfor biological research ......................... 15

Avian reproductive biologyEmbryologyModels for human genetic diseasesImmunogeneticsCytogeneticsGenomicsDevelopmental genetics

Chapter 4: Conserving avian geneticresources ............................................. 25

Methods of conservation ofgenetic stocks

Live animalsSemen cryopreservationBlastodisc cryopreservationPrimordial germ cell cryopreservation

Chapter 5: Genetic stocks:Current resources and recent losses..... 31

Survey methodResults from the surveyLost stocksTrends in poultry genetics stock

development and conservation

Chapter 6: An Avian Genetic ResourcesSystem: Proposal ................................. 39

Task Force recommendationRationaleComponents of an Avian Genetic

Resources System (AVGRS)Facilities and organizational aspects

of AVGRSFinancing the AVGRS

References........................................... 47Appendix 1: Instructions and surveyform sent to researchers ...................... 55Appendix 2: Genetic stockcollection data..................................... 57

Description of stock categoriesNotes on specific fieldsDisclaimer

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List of boxes

1. Commercial value of the poultryindustry ............................................. 1

2. Dispersal of a genetically significantpoultry collection ............................... 2

3. Collections at risk: University ofCalifornia–Davis ................................. 2

4. Successful conservation of at-riskgenetic stocks .................................... 3

5. Parthenogenetic turkeys ..................... 6

6. Research with goose breeds inCanada .............................................. 8

7. Highly inbred stocks inimmunogenetics ................................. 9

8. Chicken chromosome rearrange-ment stocks ..................................... 10

9. Economics of sex-linked genesand chicken genetics ........................ 11

10. Development of egg-laying stocks ..... 12

11. Development of meat-producingstocks .............................................. 13

12. Small renaissance of old-stylechicken breeds ................................. 14

13. Inherited muscular dystrophyof the chicken .................................. 16

14. Deep pectoral myopathy: One priceof intensive selection ........................ 18

15. Ancona chickens give a new angleon disease resistance ....................... 19

16. Government involvement ingenome research .............................. 20

17. Patterns in vertebrate limbdevelopment ..................................... 22

18. Patterns in skin development ........... 24

19. Mosaic chickens and geneticstock preservation ............................ 28

20. Orphaned CFAR stocks preservedas frozen embryonic cells ................. 29

21. The Jackson Laboratory modelfor vertebrate genetic stocks ............. 36

22. Japanese quail at the Universityof British Columbia .......................... 37

v

List of figures

ES1 Components of the Avian GeneticResources System .............................xii

1. Red Jungle Fowl rooster fromUCD 001 ............................................ 5

2. White Leghorn rooster fromUCD 003 ............................................ 6

3. Flock with different turkey breeds ...... 6

4. Japanese quail silver mutation fromUBC SI ............................................... 7

5. Japanese quail porcupine mutationfrom UBC PC-WB ............................... 7

6. White Leghorn hen ........................... 12

7. Traditional broiler-type chickencarcass of the 1940s ........................ 13

8. Commercial Large White turkey tom .. 13

9. Silkie rooster from UCD Silkie .......... 14

10. Chromosomes from a Red JungleFowl hen from UCD 001 ................... 19

11. Wingless-2 chicken embryo fromUCD Wingless-2 X 331 ..................... 21

12. Eudiplopodia embryo foot fromUCD Eudiplopodia X 003 ................. 22

13. Hens from UCD Scaleless-Low.......... 24

14. Stage X blastodisc (at time ofoviposition)....................................... 27

15. Diagram of avian blastodisccryopreservation and recovery .......... 28

16. Components of the Avian GeneticResources System ............................ 40

ES1 Estimated costs of Avian GeneticResources System ............................. xv

1. Summary by category of existingpoultry genetic stocks (live andcryopreserved) .................................. 32

2. Number of poultry genetic stockskept as live birds in institutionsin the USA and Canada in 1998,and the number lost since 1984 ....... 34

3. Summary by category and speciesof poultry genetic stocks reportedlost since 1984 ................................. 35

List of tables

4. Estimated costs of Avian GeneticResource System .............................. 45

Appendix 2

2.0. Stock categories ............................... 57

2.1. Stock information........... Separate pagination following 60

2.2. Stocks listed by country, state orprovince, institution, and curator........... Separate pagination following 60

2.3. Curator contact information........... Separate pagination following 60

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PPPPPrefacerefacerefacerefacerefaceIN 1984, AN INTERNATIONAL SYMPOSIUM and Work-shop on Genetic Resources Conservation forCalifornia made the recommendations that pre-cipitated the establishment of the Genetic Re-sources Conservation Program within the Uni-versity of California (UC/GRCP). These recom-mendations recognized that the avian geneticstocks developed in California, primarily chickenand coturnix quail, had been valuable in re-search and the commercial poultry industry andthat continued conservation of such stocks de-pended upon long-term support for live-birdmaintenance and further research on cryopres-ervation technology. At that time, there was nonational-level program, comparable to the Na-tional Plant Germplasm System, directed to ani-mal genetic resources. By the early 1990s, theCalifornia situation was fast approaching a crisiswith the imminent retirement of the two primarydevelopers and curators of poultry research ge-netic stocks at the University of California–Davis, Ursula Abbott and Hans Abplanalp. Noplan was in place for the continued support andmaintenance of the important and widely usedgenetic stocks developed and acquired by them.Nationally, other collections were at risk as thepoultry and avian science departments in the USwere losing faculty (to retirements) and financialresources that had supported collection mainte-nance in the past. The UC Davis campus re-cently followed this trend, with the merging of itsAvian Sciences Department into the Animal Sci-ence Department in 1997. Avian genetic stocksin Canada are likewise imperiled.

Recognizing the drastic reductions in humanand financial resources for avian genetic resour-ces management in the US and Canada, UC/GRCP, in cooperation with two agencies of theUS Dept. of Agriculture (USDA), the AgriculturalResearch Service (ARS) and the CooperativeState Research, Education, and Extension Ser-

vice (CSREES), convened a binational Task Forcein 1995 to address the problem. The Task Forceincluded representatives from the many publicinstitutions that have or had poultry researchprograms, from private companies, and from re-search programs that have utilized poultry ge-netic stocks.

In support of the Task Force’s work, UC/GRCP conducted a survey of existing geneticstocks in the US and Canada to provide an in-ventory for analysis. This new inventory is pre-sented in this document. The most recent priorinventory was published in 1988 (R.G. Somes,Jr. 1988. International Registry of Poultry GeneticStocks. Storrs Agr. Exp. Sta. Bull. 476. The Uni-versity of Connecticut, Storrs). It was not sur-prising to find that many stocks had been lost inthe interval. Indeed, numerous stocks have beenlost over the course of this Task Force effort.

The majority of the Task Force convened atan initial meeting in Davis in October 1995 anda subgroup met in Washington DC in May 1996.Results of the survey and progress of the TaskForce have been presented and discussed at sev-eral meetings including the annual NE–60 Re-gional Research Project meetings and the annualmeetings of the Poultry Science Association andthe Society for Developmental Biology. An in-terim review of the issues was presented byM.E. Delany, co-chair of the Task Force, andJ.M. Pisenti, UC/GRCP’s facilitator for the TaskForce (M.E. Delany and J.M. Pisenti. 1998. Con-servation of poultry genetic research resources:Consideration of the past, present, and future.Poultry and Avian Biol. Rev. 9:25–42).

We thank Mary Delany (University of Califor-nia–Davis) and Robert L. Taylor, Jr. (Universityof New Hampshire) for serving as co-chairs of theTask Force and all the members for their contri-butions of information and editing of this report.We especially acknowledge Jacqueline Pisenti, a

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poultry developmental geneticist, who facilitatedthis effort in every way. Her great familiarity withpoultry stocks and expertise in maintaining thegenetic stocks at UC Davis was a definite assetin carrying out the charge to the Task Force. Herpersistence in pursuit of information for the sur-vey made it a successful venture. Her work inpulling together this report from componentssubmitted by many different individuals has pro-duced what we think is a document that speaksin a uniform, strong voice for the value of thesegenetic stocks and for the imperative of theircontinued conservation.

The work of the Task Force could not havebeen accomplished without financial supportfrom UC/GRCP and from grants from the Na-tional Science Foundation and the AgriculturalResearch Service. These grants both supportedthe activities of the Task Force and contributedto the maintenance of at-risk poultry geneticresource collections at the University of Califor-nia–Davis.

The Task Force has defined the types of ge-netic stocks, made an excellent summary of theextent of the use of avian genetic resources inmany different research disciplines, analyzed theinventory for trends in stock development andconservation, and finally made recommenda-tions designed to ensure the continued availabil-

ity of existing genetic resources and support thefurther development of new genetic resources.

While Congress authorized the US NationalGenetic Resources Program with the Food, Agri-culture, Conservation and Trade Act of 1990,there still is no plan for ensuring the conserva-tion of poultry and other livestock genetic re-sources critical to the US. In Canada there hasbeen a loss of federal support for poultry geneticresources and now most of their important ge-netic resources exist only as frozen embryo blas-todisc cells. With these uncertainties, it isdoubtful if new genetic stocks from current re-search will be conserved. This is a direct deter-rent to new research on critical issues in avianbiology. At the same time, we recognize thatemerging technologies of molecular biology andgenomics have increased the value of existinggenetic stocks and provided the impetus to de-velop new ones which will need conservation at-tention. The Task Force has made well consid-ered and reasonable recommendations. UC/GRCP and the USDA will work to make theserecommendations known and will support theiradoption into a viable Avian Genetic ResourcesSystem to protect and make available to all re-searchers extant avian genetic stocks and, im-portantly, to encourage new research in bio-medicine, biology, and agriculture.

Calvin O. Qualset, UC/GRCPRichard R. Frahm, USDA–CSREES

Steven Kappes, USDA–ARS

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Executive SummarExecutive SummarExecutive SummarExecutive SummarExecutive Summaryyyyy

BackgroundBackgroundBackgroundBackgroundBackgroundAMONG THE DOMESTICATED AVIAN species, tradi-tional breeds developed for meat or egg produc-tion are a tremendous source of genetic diver-sity. Such breeds result from hundreds of yearsof selection by farmers and breeders, incorporat-ing a variety of unique genes and gene combina-tions. Likewise, intense selection for odd andinteresting traits resulted in the array of fancybreeds treasured by hobbyists. The relativelysmall size of most poultry species (particularlythe chicken and Japanese quail), along withtheir phenomenal reproductive capacity and ge-netic variability, make them particularly wellsuited to multi-generational studies with rigor-ous selection or high levels of inbreeding. Thishas stimulated the development of many impor-tant genetic stocks of chicken, turkey, and quailby researchers in agricultural departments atpublic universities or in federal agricultural re-search organizations. These specialized geneticstocks have contributed widely to research inbasic, biomedical, and agricultural sciences.

To date, no formal conservation plan existsfor these stocks at any organizational level. In-stead, conservation of these important avian ge-netic resources is largely dependent upon indi-viduals, sometimes connected through breedassociations or academic circles, but largely asindependent efforts. This lack of a formal conser-vation plan is particularly perilous for researchgenetic stocks. Unlike the more widely dispersedtraditional or hobby breeds, specialized researchstocks are often kept at only one location, whichmakes them far more vulnerable to extinction. Inparticular, when such stocks can no longer bemaintained by their curating researcher, theyare often perceived as an unnecessary drain ondwindling financial resources at that institution,and may be eliminated at very short notice.

FFFFFindingsindingsindingsindingsindingsThis study documented the loss of more than200 stocks since 1984. A detailed database wasprepared for all extant stocks in the US andCanada, including 268 chicken, 20 turkey, 65Japanese quail, and 8 waterfowl or gamebirdaccessions.

1. The continuing loss of avian researchstocks is a significant concern, as in thepast 15 years more than 200 mutant,inbred, and selected avian genetic stockswere destroyed. More than one-third ofthe remaining stocks are at risk of elimi-nation within the next few years.

2. While several research institutions in theUS and Canada still maintain collections,they are at capacity and will likely facereduction in resources in the near future.There is no facility currently availablethat can accommodate the needs of allcurrently at-risk stocks.

3. Without stock conservation facilities,there will be little incentive for invest-ment in lines of research that would pro-duce new genetic stocks, particularlyhighly inbred and selected stocks. This isa direct deterrent to research on criticalissues in avian biology.

4. Many research stocks have unique ge-netic properties which would be impos-sible or very costly to recreate. Some ofthese have high potential value for futuregenetic research in the basic, biomedical,and agricultural sciences, for example:

• Uniform selected and inbred stocks aregenetically well suited for studies of the

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immune system and other complexfunctional systems.

• Strains that carry a wide variety of mu-tations affecting specific developmentalor metabolic processes provide a start-ing point for research exploring the tis-sue or cellular source of defects in em-bryonic development and metabolicfunction, leading to a better under-standing of the control of the normalprocesses.

• Mutations that closely mimic certainhuman congenital defects or disordersprovide animal models for detailed ex-perimental analysis of such conditions.

5. Information regarding the properties andavailability of research stocks is notreadily accessible to researchers whocould effectively use them.

6. Many researchers who utilize geneticstocks, especially in basic biological andmedical research areas, do not have fa-cilities for maintaining stocks and requireconsistent and reliable sources for livebirds, tissues, or eggs.

7. Avian genetic stocks can be conserved aslive animals or cryopreserved germplasm(specifically as embryonic cells, semen, orembryos).

• Live bird maintenance is expensive andlabor intensive, as many researchstocks require rigorous disease controlmeasures with specialized rearing andadult bird housing conditions, yearlymonitoring of specific performancetraits for selected stocks, and genetictesting for chromosomal-abnormalityand mutant-carrier stocks.

• Although the annual storage cost forcryopreserved germplasm is relativelylow, very little germplasm from aviangenetic stocks is currently cryopre-served, and recovery of live birds fromthe cryopreserved semen can be quitedifficult. An additional disadvantage ofsemen cryopreservation is that only themale genome is conserved. This elimi-nates or severely limits the usefulnessof this conservation technique with in-bred or selected stocks.

• Promising new methods for conserva-tion of diploid germplasm, cryopreser-

vation of dissociated early blastodiscsand primordial germ cells, are the sub-ject of recent research in Canada,Japan, and the US. These methods al-low the recovery of intact genomes inthe first generation.

8. Funding is difficult to secure for conser-vation-related research and for the main-tenance of live animal collections andgermplasm repositories.

9. It is difficult to predict which stocks willbe of value to commercial or academicresearchers at some future time. So, de-spite the uniqueness and potential use-fulness of the existing avian geneticstocks, the tenuous nature of the “value”of many stocks has precluded sustainedfinancial support for their preservationfrom any single agency.

10. There is currently no organization in theUS or Canada to monitor conservationstatus and availability of avian geneticstocks and set priorities for their conser-vation on a national or binational level.

RRRRRecommendationecommendationecommendationecommendationecommendationAn avian genetic resources management system,with strong leadership, but shared responsibil-ity, is proposed as the most efficient and secureway to conserve genetic stocks and address is-sues of risk and loss. The proposed Avian Ge-netic Resources System (AVGRS) would be com-prehensive and would require the cooperationand collaboration of scientists, funding agencies,and research institutions. The System must beoriented toward research objectives, but it couldalso support the needs of breeders, breed hobby-ists, and breed historians.

RRRRRationaleationaleationaleationaleationaleHistoric long-term collaboration between Cana-dian and US scientists provides a basis for col-laboration in the formation and operation of anAvian Genetic Resources System. The US Na-tional Plant Germplasm System and its counter-part in Canada serve as excellent models for theproposed Avian Genetic Resources System. Asystem on this binational level is necessary be-cause no dependable regional or local solutionexists.

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Components of an AComponents of an AComponents of an AComponents of an AComponents of an AvianvianvianvianvianGenetic RGenetic RGenetic RGenetic RGenetic Resources Systemesources Systemesources Systemesources Systemesources System

The Avian Genetic Resources System is envi-sioned as a multilocational organization thatwould serve the avian genetic resources needsfor the US and Canada. The AVGRS would fea-ture a central facility as a focal point for many ofthe activities of the System. The functional com-ponents are outlined in Executive Summary Fig-ure 1 and are briefly discussed here.

Avian Genetic Resources AdvisoryCommittee (AVGRAC)

The AVGRS would be advised by this binationalcommittee comprised of representatives of na-tional and state/provincial agencies, stock cura-tors, and researchers. It would consist of 12 to15 individuals who have worked with avian ge-netic resource issues, drawn from government,industry, and academic institutions in the USand Canada. Specifically, they should haveworked in close association with national, inter-national and private research-oriented organiza-tions, and be familiar with avian genetic stockconservation issues. The members should meetat least once a year and be in regular communi-cation during the year. The Committee wouldreview reports and recommendations for conser-vation of stocks received from species-orientedcommittees. It would make recommendations tothe management unit of the AVGRS.

Coordination

The various government and research institu-tions involved in avian research and conserva-tion would use the AVGRS for coordination ofinformation about genetic resources and theAVGRS would in turn be responsible for main-taining and distributing this information. Thisfunction would also include strategic planningfor conservation of particular stocks, based onadvice from advisory groups established for eachspecies. For example, imperiled stocks would beidentified to the AVGRS and a plan for their con-servation would be developed through coordi-nated analysis. International relationships wouldbe coordinated through the AVGRS, includingconservation of stocks in other countries, importand export of genetic stocks, data sharing, anddevelopment of conservation plans for landraces,wild species, and breeding populations.

Conservation

This is the cornerstone activity of the AVGRSand of critical importance. A central facility isneeded for conservation and distribution of ge-netic materials. The central facility would housethose living genetic stocks that could not bemaintained elsewhere and would serve as abackup site for important stocks that are main-tained elsewhere. This would include a securebackup repository for privately owned lines orpopulations, either as live birds or cryopreservedgermplasm at the central or secondary centerson a fee basis. The central facility would alsophysically maintain the various types of pre-served genetic resources and would coordinatethose maintained elsewhere. The cryopreserva-tion capabilities of the central facility would besupplemented by a specialized cryopreservationcenter, presently unused, at the USDA site inBeltsville, Maryland. No site for the central facil-ity is identified at this time.

The central facility would support and belinked to secondary facilities located at activeresearch centers which have the capability ofmaintaining genetic stocks for their own re-search needs. Several locations in the US andCanada would qualify as secondary sites.

Methods of conservation. Conservation meth-ods employed in the AVGRS would be live-birdmaintenance and cryopreservation.

Targets for conservation. Conservation empha-sis of the AVGRS should be on live birds, embry-onic cells, gametes, DNA, and tissues. The targetspecies for the system will be those of interest inagriculture for food production and for basic bio-logical and biomedical research. Thus, the focuswill be on chicken, turkey, and Japanese quailgenetic stocks. However, this system could alsoconsider wild turkey, jungle fowl, and gamebirds, as well as species commonly raised incaptivity. The AVGRS will emphasize:

• Genetic stocks having traits and geneticcharacteristics useful in research, suchas inbred lines, single-gene mutations,chromosome aneuploidy, and DNAmarker sequences.

• Lines and populations developed by pri-vate and public breeders by hybridizationand selection for important production-related traits.

• Domesticated mid-level production andfancy breeds held by small producers

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Binational Avian GeneticResources Advisory Commmittee

Avian Genetic Resources System

Coordination

USDAUS NIHUS NSFA&AF CanadaCollectionsNGOs

International organizations FAO IUCNScientific societiesOthers

Conservation Importation/Quarantine

Research

Central facility

Live birdsPreserved tissuesEggsBloodDNACultured tissuesCryopreserved semen, embryonic cells

Cryopreservation

Beltsville MDGuelph ONOthers

Stock colonies

US - CA, IA, NH, WI, MI, CTCanada - BC, ONOthers

GenomicsWebsitesStock registryBreed registry

Coordinated with USand Canadianorganizations

DistributionDatabase/Information

EggsSpermEmbryosLive birdsDNAPreserved tissues

CryobiologyReproductive physiologyGeneticsStock characterization

Figure ES1. Components of the Avian Genetic Resources System.

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and hobbyists in North America andEurope.

• Domesticated, but primitive, landracesexisting in Asia, Central and SouthAmerica, and elsewhere, primarily asscavenger birds.

Archival preserved specimens of birds, or-gans, skeletons, eggs, feathers, and tissues thathave been preserved as museum specimens arealso a component of the genetic resources sys-tem, since these materials provide for baselineobservations and time-course monitoring of fac-tors such as environmental toxicants.

An informal in situ system of conservation oflandraces and breeds is well established inNorth America. A monitoring and database sys-tem may be the most important need for thosegenetic resources. This could become an activityof this proposed system.

DatabasesDetailed information about all genetic stocks inthe US and Canada should be maintained andupdated by the AVGRS in a genetic resourcesinformation system, similar to the Genetic Re-sources Information Network (GRIN) developedfor the US National Plant Germplasm Systemand housed with the USDA National AgricultureLibrary. Additionally, database service would beoffered to the various breed conservancies andhobbyists groups for inventory and location ofconserved breeds, landraces, and specialtystocks. It would also be logical for the AVGRSdatabase to include DNA sequence data as theyare developed.

While GRIN currently focuses on plant infor-mation, its goal is to include information on allof the common and endangered breeds of farmanimals, including the avian genetic stocks usedprimarily in research. Collaboration with theAVGRS database would facilitate this goal.

OutreachResearchers can also be informed of the widevariety of available genetic stocks at the annualmeetings of a variety of organizations, includingthe Poultry Science Association, the Pacific Eggand Poultry Association, the Poultry BreedersRoundtable, the Society for Developmental Biol-ogy, the American Association for the Advance-ment of Science, and the American Medical As-sociation. Presentations at the commercially ori-ented meetings could be used to showcase thebenefits the different companies could derive

from supporting an avian genetic stocks conser-vation program, while the basic research or dis-ease model aspect of genetic stocks could be em-phasized for organizations promoting basic andbiomedical research. Thus, the underlying goalsof presenting genetic stocks information at suchmeetings is not only to attract the attention ofresearchers, but to engage the interest and pro-mote funding from commercial sectors that canbenefit directly from research using avian ge-netic stocks.

Another outreach option for the AVGRSwould be an independent website that wouldpromote the available avian genetic stocks to thescientific community by advertising what isavailable, and indicating those that are slated forelimination. The effectiveness of such a sitecould be multiplied by linking it with websites:the animal genetic map (Angen), the chickenmap (ChickMap), various research organizationsites (e.g., Poultry Science Association, Societyfor Developmental Biology, various commercialpoultry sites, and sites for academic institu-tions).

The AVGRS website information could be fur-ther promoted by a series of clearly written re-view articles in several of the major biologicalscience journals. In each case, a specific area ofresearch would be targeted, such as animalmodels for human diseases, limb pattern de-fects, craniofacial defects, integumentary de-fects, or immunogenetic research.

The outreach activity would also involve inter-national contacts through FAO or various coun-tries with respect to avian genetic resources. Forexample, there are genetic stocks at risk in othercountries that should be considered for rescuein the AVGRS because of their value for re-search.

Bird care and housingThe housing and care of the live birds at all cen-ters will follow American Association for Labora-tory Animal Care standards for a breedingcolony. Essentially, the breeding stock should bekept in a facility that approximates that of awell-run commercial poultry breeding farm. Thehighest degree of automation for feeding, clean-ing, watering and climate control is recom-mended. With most of the chicken geneticstocks, the adult birds will be housed in single-bird cages, and bred by artificial insemination.Other features of the facility may include: floorpens, with or without trap nests, to maintainstocks that do not perform well in cages; andseparation of males and the females (in different

xiv

rooms or cage rows), so that appropriate, sex-specific breeder diets, lighting, and feeding regi-mens can be provided for each group.

Importation and quarantine

Movement of animals introduces risks of spread-ing contagious diseases, obviously of great con-cern in long-term conservation of live birds.Movement of genetic resources as fertile eggs orsemen reduces disease transmission risk andare preferred procedures. Importation of stocksto the central facility will be done through on-site isolation and through national facilities un-der the direction of USDA Animal and PlantHealth Inspection Service. The central facilitywill have appropriate isolation and sanitationcapabilities.

Distribution

A major function of the AVGRS will be to providegenetic materials to users in the research com-munity, breeders, and others. The genetic stocksmay be transferred as live birds, semen, or eggs.These will be distributed on a cost-recovery ba-sis. Some users require a continuing supply ofgenetic stocks and these needs would be sup-plied by contractual stock reproduction pro-grams. The distribution function would supplywell-documented stocks to researchers, thuscontributing to the integrity of research projects.This functional component of the AVGRS isanalogous to services provided by the JacksonLaboratories for mouse genetic stocks.

Research

The AVGRS should have a research capabilitywithin the central facility, especially for develop-ing cryopreservation technologies. Researchwould also be done on methods for documentinggenetic integrity or diagnostics with DNA mark-ers. Other research would be done as neededand appropriate. The research activities wouldbe networked with research laboratories in theUS and Canada for collaborative work.

FFFFFacilities and organizationalacilities and organizationalacilities and organizationalacilities and organizationalacilities and organizationalaspects of Aaspects of Aaspects of Aaspects of Aaspects of AVGRSVGRSVGRSVGRSVGRS

The central facility

Ideally, the primary AVGRS facility would beconstructed de novo near or part of a major agri-

cultural institution (land-grant university) with aveterinary school that has a good avian medicineprogram, but reasonably isolated from commer-cial poultry stocks. The connection with a land-grant institution would give the center close tiesto active research laboratories and faculty, whocould benefit from such a resource and bedrawn upon in support of the center. Access tostate-of-the-art poultry disease diagnostics andveterinary care is critical, along with good, off-site quarantine facilities for newly acquired ge-netic stocks. Locating in a strong poultry pro-ducing state would also provide an existing poul-try-oriented political and commercial infrastruc-ture that could be mobilized to help support theconservation center.

This facility would include a hatchery, brood-ing and growing areas, adult bird housing, anisolation area, a cryopreservation laboratory andstorage facility, a database center, staff and ad-ministrative offices, and a laboratory to supportresearch and analytical services.

Network of secondary genetic stockcentersSecondary stock centers would be designated aspart of the AVGRS at land-grant universities andother institutions across the US and Canadathat fulfilled two criteria: (1) had adequate facili-ties and support for the genetic stocks used inits own research programs and (2) had a long-term interest in conserving genetic stocks. Suchcenters, approved as part of the AVGRS, wouldreceive funding from AVGRS for maintenance ofstocks. These secondary centers would maintainlive birds and would provide backup for at-riskstocks held in the central facility. As with thecentral facility, the secondary stock colonieswould also have a distribution function on a feebasis. Researchers could also, on a fee basis,arrange to keep research flocks at the centralfacility or one of the secondary centers, sincethey may find it difficult or impossible to keepsuch stocks at their own institutions. Secondarycenters could specialize on one or a few classesof genetic stocks.

Management

The central facility and the secondary centerswould be administered by the research institu-tions with which they are located. The centralfacility would have, at a minimum, a director ormanager (research scientist), curator, an admin-istrative assistant, database manager, cryopres-ervation research scientist, and three or four

xv

laboratory and animal care technicians. TheAVGRS would be guided by the advisory com-mittee on all management aspects.

Stock evaluation guidelines

One of the more important activities of theAVGRAC would be the evaluation of stocks forconservation, cryopreservation, or elimination.Guidelines for assessing the value of geneticstocks should be consulted.

FFFFFinancing the Ainancing the Ainancing the Ainancing the Ainancing the Avian Geneticvian Geneticvian Geneticvian Geneticvian GeneticRRRRResources Systemesources Systemesources Systemesources Systemesources System

Multiple sources of funding will be necessary tomeet all of the needs of the AVGRS. Initial costsare those to construct the central facility andupgrade the secondary stock centers. Annualcosts of the central facility would be for its per-sonnel and operations. It would also be neces-sary to support the annual activities of theAVGRAC. The central facility could also divertfunding for specific needs to the secondary cen-ters by means of annual grants.

From the US side, the biological resource pro-grams of the National Science Foundation andthe National Institutes of Health would be ex-pected to provide operational funds through di-rect grants and through grants to investigatorswho use the avian genetic resources in their re-search. The USDA’s National Genetic ResourcesSystem should participate in the AVGRSthrough the Agricultural Research Service andthe Cooperative State Research, Extension, andEducation Service. The various State Agricul-tural Experiment Stations and land-grant andother Universities should also participate. Cana-dian support and participation should be forth-coming to the extent that the AVGRS providessupport to its research and development pro-grams.

The AVGRS will be the major provider of ge-netic materials to researchers throughout thepublic and private sectors. For the most part,researchers do not have capacity to maintain livebird colonies and depend upon stock colonies fortheir research. User-fees are an appropriatemeans to recoup costs of stock maintenance.

Donation funds can be expected to supportthe perpetual maintenance of particular geneticstocks. These funds may be provided as annualgrants or through income derived from intereston endowment accounts.

Funding should be sought from the US gov-ernment for construction of the central facilityand for personnel support for operations as apart of the US National Animal Genetic Resour-ces System.

Long-term funding would be the most securefrom endowment funds. Contributors could beencouraged from the private sector, from largeintegrated commercial poultry companies to pri-vate individuals with interests in preservingpoultry stocks or willing to promote the conser-vation of stocks that can be used to study hu-man diseases.

Construction, personnel, and operationalcosts have not been established, pending furtheranalysis of potential sites for the central facilityand other considerations. For illustrative pur-poses, rough order-of-magnitude estimates aregiven in Executive Summary Table 1.

Table ES1. Estimated costs of Avian Genetic ResourcesSystem.Startup costsConstructing and equipping central facility $15,000,000Upgrading and renovating secondary centers 2,000,000Total $17,000,000Annual costsPersonnel at central facility $400,000Operating costs at central facility 100,000Grants to secondary centers (8 x $25,000) 200,000Advisory Committee 25,000Total $725,000

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AAAAAvian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Resources Tesources Tesources Tesources Tesources Task Fask Fask Fask Fask Force Membershiporce Membershiporce Membershiporce Membershiporce MembershipUrsula K. Abbott, Dept. of Animal Science,University of California, Davis, CA USA

Hans Abplanalp, Dept. of Animal Science,University of California, Davis, CA USA

James A. Arthur, Hy-Line International,Dallas Center, IA USA

Murray R. Bakst, USDA-ARS-GGPL, BARC-East,Beltsville, MD USA

Colin Baxter-Jones, Arbor Acres, Inc.,Glastonbury, CT USA

James J. Bitgood, Dept. of Poultry Science,University of Wisconsin, Madison, WI USA

Francine A. Bradley, Dept. of Animal Science,University of California, Davis, CA USA

Kimberly M. Cheng, Dept. of Animal Science,University of British Columbia, Vancouver, BCCANADA

Mary E. Delany (Co-Chair, AGRTF), Dept. ofAnimal Science, University of California,Davis, CA USA

Rodney R. Dietert, Dept. of Microbiology, Immu-nology, and Parasitology, Cornell University,Ithaca, NY USA

Jerry B. Dodgson, Dept. of Microbiology,Michigan State University, East Lansing, MI USA

Ann M. Donoghue, USDA-ARS-GGPL,BARC-East, Beltsville, MD USA

Alan B. Emsley, Poultry Consultant,Trumansburg, NY USA

Robert J. Etches, Origen Therapeutics,South San Francisco, CA USA

Richard R. Frahm, USDA-CSREES,Washington, DC USA

Allan A. Grunder, CFAR, Ottawa, ON CANADA

Roger J. Gerrits, retired, USDA-ARS, NationalGenetic Resources Program, BARC-West,Beltsville, MD USA

Paul F. Goetinck, Cutaneous Biology ResearchCenter, Dept. of Dermatology, MassachusettsGeneral Hospital, Charlestown, MA USA

David E. Harry, Nicholas Turkey BreedingFarms, Sonoma, CA USA

Susan J. Lamont, Dept. of Animal Science,Iowa State University, Ames, IA USA

Gail R. Martin, Program in Developmental Biol-ogy, Dept. of Anatomy, University of California,San Francisco, CA USA

Patrick E. McGuire, Genetic Resources Conser-vation Program, University of California, Davis,CA USA

Gary P. Moberg, Dept. of Animal Science,University of California, Davis, CA USA

Jacqueline M. Pisenti (Facilitator, AGRTF),Genetic Resources Conservation Program,University of California, Davis, CA USA

Louis J. Pierro, Storrs Agric. Experiment Station,Storrs, CT USA

Calvin O. Qualset (ex officio), Genetic ResourcesConservation Program, University of California,Davis, CA USA

Muquarrab A. Qureshi, Dept. of Poultry Science,North Carolina State University, Raleigh, NC USA

Fred T. Shultz, Animal Breeding Consultant,Sonoma, CA USA

Robert L. Taylor, Jr. (Co-Chair AGRTF), Dept. ofAnimal and Nutritional Sciences, University ofNew Hampshire, Durham, NH USA

Barry W. Wilson, Dept. of Animal Science,University of California, Davis, CA USA

xviii

Charge to the TCharge to the TCharge to the TCharge to the TCharge to the Task Fask Fask Fask Fask ForceorceorceorceorceEVALUATE THE CURRENT STATUS of avian genetic stocks maintained in the US and Canada. The assess-ment should include the following aspects:

Origin and history of avian genetic stocks.How avian genetic stocks have been utilized.Value of avian genetic stocks in agriculture, biomedical, and basic biological research.Strategies and techniques for long-term conservation.

Produce a report documenting these findings and include recommendations for organizational, fiscal,and administrative steps and personnel necessary to ensure the long-term security of avian geneticstocks for the US and Canada.

1

GENETIC DIVERSITY, IN BOTH WILD and domesticspecies, is a limited resource worth preservingfor future generations (OLDFIELD 1984; ALDERSON1990; FAO 1992; NRC 1993; BIXBY et al. 1994).While many strong advocates promote the con-servation of wild species, fewer are aware of theincreasing loss of biodiversity in our major foodspecies, particularly among domestic birds. For-tunately, breed conservation organizations havealready made some progress in encouraginghobbyists and small-scale farmers in their roleas conservators of unique and historically im-portant breeds (BIXBY et al. 1994), particularlythe less common chicken and turkey breeds(CRAWFORD and CHRISTMAN 1992). These two spe-cies are considered more at-risk than most otherlivestock species (e.g., cow, pig, sheep, goat, orhorse) due to recent and extraordinarily rapidexpansion of the commercial poultry industry.

In just 50 years, poultry production wentfrom small, individually owned and reproducedfarm flocks that formed a small but significantpart of the farm income, to the huge commercialmeat- (turkey or broiler-chicken) or egg-produc-tion ranches that are generally owned or con-trolled by large corporations (CRAWFORD 1990).In 1997 this industry generated over $21 billionin poultry products in the US (Box 1 and USDA-NASS 1998).

The intense competition engendered by therapid growth and often narrow profit marginshas served to eliminate the less competitivepoultry breeders and to consolidate the highproduction industrial bloodlines in the hands ofa dozen or so poultry breeding organizations.This has created a relatively limited genetic basefor the chicken, derived primarily from twobreeds (Leghorn and Rhode Island Red) for eggproduction, and two breeds (Plymouth Rock andCornish) for meat production (CRAWFORD 1990).A similar situation exists for the commercial tur-key. These highly selected industrial stocks con-siderably out-perform the old production breeds,given the correct feed and management. But therelentless drive to improve the meat- (and egg-)producing abilities of the commercial chickenand turkey stocks has exacted a biological cost:disease susceptibility, leg weakness, muscle de-fects, and various other inherited conditions thatinterfere with the ability of the bird to hatch,grow, and reproduce normally (CRAWFORD 1990).Despite these limitations, such stocks have al-ready displaced or diluted some of the hardy,disease-resistant indigenous farm stocks kept indeveloping countries (MASON and CRAWFORD1993).

The threat to genetic diversity extends beyondthe hobbyist, farm, and commercial poultry

IntroductionIntroductionIntroductionIntroductionIntroduction

1

ket, poultry production in Canada hasincreased slowly but surely between1988 and 1996 (the most recent pro-duction year available), from 370 to al-most 480 million broilers; from 18.6to 21.6 million turkeys; from 27.5 to28.2 million eggs. In 1995, the Cana-dian poultry industry was valued at$3.7 billion Canadian dollars (approx.$2.4 billion US) (AA-FC 1996).

Box 1. Commercial value of the poultry industry

THE CHICKEN AND TURKEY are the two mostcommercially important poultry spe-cies in the US, having steadily increasedin popularity with consumers for sev-eral decades. In 1987 these two spe-cies were the source of over $12.5 bil-lion in marketable products; in 1997,their total value was $21.6 billion, de-rived from almost 8 billion broilerchickens, 300 million turkeys, and 311

million laying hens that produced 77.4billion eggs that year (USDA-NASS 1998).This placed the 1997 poultry industry be-low the beef industry in value, but wellabove the pork or sheep industries, andabout on par with the dairy industry.

The Canadian poultry industry is alsogrowing due to both increased consumerdemand and a diverse export market.More closely regulated than the US mar-

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stocks. Many of the geneticallydiverse avian genetic poultrystocks developed, maintained,and studied by academic re-searchers have disappeared orbecome at risk in recent years(Boxes 2, 3, and 4, and Chapter5). As these specialized stocksvanish, unique opportunities forgenetic advancement of the dif-ferent species are lost, and sci-entific advancement in the agri-cultural, biomedical, and basicsciences is hampered. Loss ofthese unusual genetic stocks isnot a trivial matter, since thedifferent forms (alleles) of eachgene that underlie the observeddifferences in these stocks may be found only ina single population of birds; if that population islost, the unique alleles are also lost.

Genetic stocks of chickens, turkeys, andJapanese (Coturnix) quail have played importantroles in both basic and applied research, oftenserving as premier model organisms in the studyof fundamental questions in vertebrate biology.In the biomedical field, all three of these specieshave numerous mutant forms that have pro-vided animal models for the study of certain in-herited human disorders (Appendix 2). Theseinclude defects such as (glaucoma, macular de-generation) various limb defects, cleft palate,muscular dystrophy, and autoimmune forms ofthyroiditis, vitiligo, and scleroderma.

Agriculturally important genetic stocks in-clude those selected for various production-re-lated characteristics. These include egg produc-

tion, body shape, feed-use efficiency, legstrength, and disease resistance. Ironically, it isthese selected stocks that are particularly vul-nerable when funds become limiting, becauseimprovements in production-related traits re-quire many years and a relatively large numberof birds each year. Basic research can make useof all these types of stocks, but the varioussingle-gene mutations and genetically uniforminbred and congenic lines have proven to be par-ticularly useful.

Despite recommendations for conservation(CAST 1984; NRC 1990), no formal conservationplan exists in the US or Canada for such re-search stocks, and many have been lost or nowface extinction as curating researchers eitherretire or lose funding for stock maintenance.

The real and threatened loss of genetic diver-sity in chicken, turkey, and Japanese quail

AN EXAMPLE OF THE PROBLEMS confrontingpoultry genetic stocks is found at theUniversity of California–Davis (UCD). Aswith other organizations across thecountry, UCD has reduced resource al-locations for maintenance of geneticstocks. This has become an acute prob-lem for poultry genetic stocks oncemaintained by the Department of AvianSciences. The department itself hasbeen subsumed by the Deptartment ofAnimal Science. The retirements ofUrsula Abbott and Hans Abplanalp ofthe former Avian Sciences Deptartmenthave put at risk over 50 inbred, mu-tant, and specialty lines of chickens col-lected or developed by these two re-searchers since 1955 (see Appendix 2for descriptions of these stocks) and

Box 3. Collections at risk: University of California–Davis

formerly maintained by their researchgrants and departmental funds. Thesestocks have been distributed widely toresearchers for use in such diverse ar-eas as studies of the immune system (theeffects of the major histocompatibilitycomplex haplotypes on disease resis-tance and the characterization of thephysiological parameters of a chicken ge-netic immune-deficiency syndrome), thearchitecture of the chicken genome (twoof Abplanalp’s inbred lines were used tocreate reference backcross populationsand provide baseline DNA for the ChickenGenome Mapping Project), and the ef-fects of inbreeding on different repro-ductive traits. Some of the mutations areuseful as animal models for the study ofcertain human genetic diseases (ichthyo-

sis, muscular dystrophy, scoliosis, andscleroderma). In addition, Abbott hasmaintained and studied 14 mutationswith defined effects on craniofacialand/or limb development and two thataffect the integument. Studies withthese mutations have provided signifi-cant insights into mechanisms control-ling vertebrate morphogenesis andpattern formation. These stocks havethe potential to contribute significantlyto our understanding of a variety of cur-rent and future problems within the ba-sic, biomedical, and commercial/agri-cultural science realms of study, par-ticularly with the use of the rapidlyevolving technology that allows re-searchers to address the molecular ge-netic basis of such problems.

Box 2. Dispersal of a genetically significant poultry collection

ONE OF THE LARGEST commercially de-rived collections of chicken geneticstocks in North America was locatedat the Center for Farm Animal Research(CFAR) in Ottawa, Ontario. Many infor-mative studies were conducted withhistorical and modern commercialstocks that dated back to at least 1950.Some of the accomplishments includeproduction of unique control strainsfrom these stocks, improvement of pro-duction traits by selection with no evi-dence of plateaus, study of resistanceto diseases including Marek’s diseaseand lymphoid leukosis, comparison ofmethods of measuring eggshell qual-ity, and the development of semi-

congenic lines for meat-bird-derivedendogenous viral genes (GRUNDER et al.1995). The stocks used in these stud-ies included versions of both meat- andegg-selected stocks that had been se-questered from commercial stocks sev-eral times since the 1950s, includingcontrol strains, multitrait selectedstrains, and specialty strains selectedfor one trait such as endogenous viralgenes. Unfortunately, the facility inOttawa lost its government supportand was shut down April 1, 1997. Em-bryonic blastodermal cells from some30 CFAR stocks not transferred toother institutions were frozen at theUniversity of Guelph (Ontario, Canada).

3

prompted the formation of the Avian Genetic Re-sources Task Force (AGRTF). Early in the discus-sions, Task Force members realized that therewere several major, but different, conservationissues. One is the protection of the ancestralwild populations, another is the conservation ofunique breeds and landraces of domesticatedpoultry species, and finally, a more specializedconservation effort is the preservation of uniquegenetic types developed for use in agricultural,biomedical, and basic biological research, in-cluding the various single-gene traits, highly in-bred lines, and the populations or lines underimprovement for various economic or other de-fined traits. While the preservation of wild pro-genitors of domestic species is of critical impor-tance, as is the conservation ofnondomesticated avian speciesworld-wide, the Task Force de-cided that the scope of this re-port should be restricted tospecialized genetic stocks. TheTask Force also recognized thatrare breeds conservation, aspracticed by hobbyists andothers, was already in placein North America. The NorthAmerican poultry stocks cur-rently most at-risk are theunique genetic variants andspecialty stocks held by re-search institutions in the US

and Canada. A comprehensive genetic resourcesmanagement strategy is herein proposed thatcan, in the future, provide support and servicesfor all economically important avian species.

This report discusses the history and uses ofthe different species, presents results of a surveyof the genetic resources in the targeted species,gives overviews of major research areas depen-dent on such genetic stocks, discusses the dif-ferent conservation methods currently available,and, finally, proposes a plan for a comprehen-sive Avian Genetic Resource System that wouldinsure long-term maintenance and accessibilityof these (and potentially other) endangered aviangenetic stocks.

Box 4. Successful conservation of at-risk genetic stocks

THE EXAMPLES OF THREE LINES (Trisomic,mPNU, and Triploid) serve as modelsof the traditional mode of curator trans-fer from originating-investigator/insti-tution to a new curating-researcher/in-stitution. The Trisomic and mPNU linesoriginated in the Poultry and Avian Sci-ences Department at Cornell University(S.E. Bloom) and were recentl1y trans-ferred to the University of California–Davis (M.E. Delany). The Trisomic lineis also maintained at the University ofNew Hampshire (R.L. Taylor, Jr.). TheCSIRO Triploid line was transferred

from one CSIRO facility (B. Sheldon) toanother (R. Pym). Thus, these special-ized-cytogenetic lines represent suc-cess stories not only because of theirsignificant contributions to basic andapplied research, but also in that newhomes for the lines were establishedso that their use and availability for fu-ture research are secure, at least forthe foreseeable future. Unfortunately,this model of successful transfer of cu-ratorship of genetic lines is the excep-tion today rather than the rule.

5

2

AAAAAvian genetic diversity: Domesticated speciesvian genetic diversity: Domesticated speciesvian genetic diversity: Domesticated speciesvian genetic diversity: Domesticated speciesvian genetic diversity: Domesticated species

GENETIC DIVERSITY IS CONSIDERED crucial to thecontinued survival of a species, be it wild or do-mestic. Such within-species diversity has beenthe raw material of agriculturists over millennia.In response to selective breeding and the differ-ential survival of less fit animals, preferred traitshave been accentuated and clustered to producedistinct breeds and varieties of the modern do-mesticated species (NRC 1993). In more recenttimes, researchers have deliberately isolatedvarious mutations in specialized stocks, permit-ting the systematic study of such mutations andpromoting a better understanding of the normalfunction of the affected genes.

The totality of wild and domesticated speciesform the gene pool or genetic resources basenecessary for the survival of the species. Thegenes and genotypes present in this pool repre-sent genetic resources which are accessible andcan be exploited by biologists and breeders. Inthis report, we emphasize “genetic stocks” whichhave been bred for specific traits and genes incontrast to breeds in which the individual birdshave many traits in common and can generallybe maintained with randomly breeding popula-tions. Genetic stocks are typically selected fortraits of special interest to breeders and geneti-cists. Many of them are reproductively, physi-cally, or physiologically compromised, and re-quire special care in breeding and management,even for maintenance or conservation purposes.

TTTTTarget speciesarget speciesarget speciesarget speciesarget speciesWhile the AGRTF recognizes the need for conser-vation of undomesticated avian species, this re-port primarily addresses the need for conserva-tion of specialty stocks of domesticated species,particularly chicken, turkey, and Japanesequail. A limited number of waterfowl (duck andgoose) genetic stocks and semi-domestic game-

bird stocks (ring-necked pheasant and bobwhitequail) have been developed and will be noted inthis report. Noted below are salient features ofthe most widely used domesticated species thathave the greatest need for conservation of ge-netic stocks.

Chicken

First domesticated over 6,000 years ago, thechicken (Gallus gallus or G. domesticus) presentsby far the greatest amount of genetic diversity ofthe domesticated avian species, with over 400identified genetic variations (SOMES 1988). Manyare showcased in the more-than 100 recognizedchicken breeds and commercial varieties, whichvariously integrate most of the naturally occur-ring mutations affecting size, body type, produc-tion characteristics, posture, color, featherstructure and location, comb shape, and behav-ior (see Figures 1 and 2 for wild- and domestic-type chickens). Some of the most extreme vari-ants include: the tiny, short-legged JapaneseBantam; the tall, aggressive Old English Game

Figure 1. Red Jungle Fowl rooster from UCD 001(Photo courtesy of J. Clark, University of California–Davis).

6

popular with researchers or hobbyists than thechicken or the Japanese quail, at least sixbreeds are still kept for exhibition and a fewunique research stocks have been developed(Box 5), including several commercial-type long-term selected and randombred-control lines keptat Ohio State University (see survey results, Ap-pendix 2, Tables 2.1 and 2.2). Perhaps as a con-sequence of the few researchers studying theturkey, relatively few mutations (49) have beenreported in the turkey compared to the chickenand Japanese quail (SOMES 1988).

Japanese quail

Gaining in popularity as an experimental animalin both research and education, the Japanesequail (Coturnix japonica) is a small, early matur-ing, highly efficient egg and meat producer. Untilrecently, the Japanese quail was classified as a

hatch, and grow into fully functionalmales (OLSEN 1965). The existence ofthis line has given rise to the notionthat genetic imprinting does not existin birds, although this conclusion mustbe tentative in the absence of any for-mal investigation of imprinting in theunique parthenogenetic stock. How-ever, this parthenogenetic stock existsprecariously at only two research sta-tions in the world (the University ofGuelph (Ontario, Canada) and the Uni-versity of Oman).

Fowl; the light-weight Single-comb White Leghorn hen thatcan lay more than 300 eggs inher first year of production; andthe large Rock-Cornish commer-cial meat chicken, with its phe-nomenal rate of growth andwell-fleshed carcass. These areall thought to share a commonancestor in the Red Jungle Fowl(G. gallus gallus) (Figure 1)which is still found wild in partsof India and Southeast Asia(CRAWFORD 1990; FUMIHITO et al.1994), although some poultryspecialists believe that severalother jungle fowl species (G. sonnerati, G. lafay-ettei, and G. varius) also contributed to the an-cestral gene pool (CRAWFORD 1990).

Turkey

The one commercially important avian speciesoriginating in North America, the domestic tur-key of commerce, is the product of hybridizationbetween two subspecies of turkey: the domesti-cated Meleagris gallopavo gallopavo from CentralAmerica and the wild M. g. sylvestris from theeastern United States (CRAWFORD 1990). Fromthese hybrids, birds were selected for size, tame-ness, carcass yield, and rapid growth, resultingin several distinct breeds and varieties (Figure3). The modern commercial or exhibition turkeysare large, slow-maturing birds with a muchlower reproductive potential than chicken orJapanese quail (at one generation per year forthe turkey). Although this species is far less

Box 5. Parthenogenetic turkeys

PERHAPS THE MOST SPECTACULAR use of tur-keys in experimental biology was thestudy of meiosis, fertilization, and earlyembryonic development with a strainof parthenogenetic turkeys. In the1960s and 1970s, M.W. Olsen of theUnited States Department of Agricul-ture Agricultural Research Center atBeltsville developed a line of turkeysin which an embryo would form in 30to 50% of the unfertilized eggs (par-thenogenesis). Most of these embryosdie, but a small proportion of them(about 0.5%) continue to develop,

Figure 3. Flock with different turkey breeds (Photocourtesy of F.A. Bradley, University of California–Davis).

Figure 2. White Leghorn rooster from UCD 003 (Photocourtesy of J. Clark, University of California–Davis).

7

subspecies of the common European quail (C.coturnix). It is now classified as a distinct speciesbecause of the nonhybridization of the two in thewild or in captivity (CHENG and KIMURA 1990).According to all available documentation, thedomestic Japanese quail strains used in meatand egg production (even in Europe) are descen-ded from C. japonica, which is still found insmall wild populations in Japan. While gainingpopularity as a food animal in the US, its smallsize has limited its use as a meat- or egg-pro-ducing animal to specialty markets. However,the Japanese quail has other qualities that makeit ideally suited for research. Usually reachingsexual maturity by six weeks of age, the femalesoften lay an egg a day for several months. Themales are aggressive breeders and maintain highfertility even when housed with four or morehens. The early maturity and short incubationinterval (16 to 17 days) permit as many as fivegenerations in a single year, in contrast to theslower-maturing chicken (one to two generationsper year) or the even slower maturing and lessproductive turkey (one generation per year). Thequail is sometimes called the mouse of the birdworld, since it has become extremely popular asa model species for biological research in severalfields, including toxicology, cell biology, nutri-tion, and selective animal breeding. Althoughmost researchers use unselected or randombredbirds, over 100 mutations are known in this spe-cies, including many affecting feather color andshape (Figures 4 and 5), and several causingembryo-lethal deformities (SOMES 1988; CHENGand KIMURA 1990). At present, most of these mu-tant strains are only maintained at the Univer-sity of British Columbia or by hobbyists. Twodrawbacks with this species are that the usual

productive life of an individual bird is quiteshort, frequently less than one year, and, unlikethe chicken, close inbreeding is not tolerated.Thus, only one moderately inbred line exists (atthe University of British Columbia).

Duck

Almost all of the 15 or so domestic duck breedsrecognized today are descended from the wildmallard duck (Anas platyrynchus platyrynchus),the exception being the Muscovy duck (Cairinamoschata) (LANCASTER 1990). In addition to thedifferent plumage patterns and colors, a varietyof body types and behavioral traits are foundamong the duck breeds, ranging from the boat-shaped, vocal Call ducks to the cane-shaped In-dian Runner ducks. Only 22 mutations havebeen described in the domestic duck, most in-volving feather color or pattern (LANCASTER1990). As such, these traits have been used indefining breed and variety standards, especiallyamong the more ornamental duck breeds, suchas the Call, Indian Runner, Crested, Cayuga,and Swedish. While such breeds are usuallyonly kept by hobbyists, a few are important incommercial meat production, particularly WhitePekins, Rouens, and, in some areas, Muscovy orMuscovy-domestic duck hybrids.

Goose

Six recognized domestic goose breeds were de-rived from the western Greylag goose of Europe(Anser anser anser). Several other breeds arethought to have descended from the smallerSwan goose of central Asia (A. cygnoides). TheAfrican breed is believed to be derived from a

Figure 4. Japanese quail silver mutation from UBC SI(Photo courtesy of K. Cheng, University of British Co-lumbia).

Figure 5. Japanese quail porcupine mutation fromUBC PC-WB (Photo courtesy of K. Cheng, Universityof British Columbia).

8

hybrid between these two species (HAWES 1990).Strict herbivores, geese have a long history ofdomestication, but their delayed maturity (twoyears) and low egg production rate make themless attractive as an experimental animal or as acommercially viable species (Box 6). However,due to the increasingly diverse consumer groupsin the US and Canada, formerly noncommercialspecies are becoming popular on a small scalefor specialty markets. One example is the de-mand from the Asian markets for a smaller, lessfatty meat goose. Until now, Europeans andNorth Americans have traditionally raised Emb-den geese for market purposes. This large-bod-ied, fatty bird is not well suited to the method ofcooking employed by Asian chefs. Therefore, wa-terfowl suppliers are now starting to grow thesmaller Chinese geese for this market.

Gamebirds

Several species of game birds are commonlybred commercially or by hobbyists, includingmany subspecies of the Ring-necked pheasant(Phasianus colchicus) and the Bobwhite quail(Colinus virginianus). Nine color mutations havebeen identified in the pheasant, along with sev-eral affecting skin color and feather structure,and 11 that produce biochemical polymorphisms(SOMES 1990). For most populations, very littleselective breeding or inbreeding is deliberatelypracticed, and the development of gamebirdstocks for genetic research is unusual. Excep-tions include the now-extinct inbred pheasantlines developed at the University of California–Davis (WOODARD et al. 1983) and the Bobwhite

and pheasant blood-type variants currently keptat Northern Illinois University (JARVI et al. 1996).

TTTTTypes of genetic stocksypes of genetic stocksypes of genetic stocksypes of genetic stocksypes of genetic stocksFor the purposes of this report, genetic stocksare classified into four categories that reflect thegenetic composition and type and the breedingsystem used to maintain them.

• Randombred

• Highly inbred

• Long-term selected

• Mutant (including cytogenetic variantsand transgenics)

We are primarily concerned in this reportwith conservation of genetic stocks developed forresearch purposes, which include all of thesecategories. Conservation of genetic stocks in thedifferent categories present different challengesfor successful conservation, including: high em-bryonic mortality, low viability, poor reproduc-tive traits, pronounced susceptibility to one ormore diseases, large and deleterious geneticload, poor response to specific environmentalstressors, poor recovery of cryopreserved semen,and need for a very large gene pool (more than100 birds per generation).

Randombred lines are maintained as relativelylarge populations of birds (usually over 100) inwhich little, if any, selection of breeding stock isdone by the curator. Quite simply, the numberof progeny from each male or female depends on

the reproductive success of thatbird at the time the eggs are col-lected to reproduce the popula-tion. Such randombred stocksare generally kept as closedflocks, although new bloodlinesmay be introduced to the popu-lation to improve the vigor of theflock, particularly if inbreedingdepression is observed. Thebirds may be reared and bred ina single large enclosure, with allmales having access to all fe-males. This is a common forpheasants, ducks, geese, somechickens, and naturally breed-ing turkey stocks. Alternatively,the birds may be randomly seg-regated into smaller floor pensor randomly paired or groupedin cages, as is common with

and improving egg production withcontrolled lighting and trapnesting.Early studies with Pilgrim geese (theonly goose breed that shows strongsexual dimorphism) included a demon-stration of increased egg productionwith selection (MERRITT 1962), and useof light control to increase egg produc-tion. More recently, artificial insemina-tion techniques have been improved(GRUNDER and PAWLUCZUK 1991) andgeese have been shown to lack endog-enous viruses (i.e., viral DNA integratedinto the host bird chromosomes) of theavian leukosis type (GRUNDER et al.1993). Unfortunately, with the loss offunding from the Canadian govern-ment in April of 1997, these stockswere either eliminated or dispersed.

Box 6. Research with goose breeds in Canada

WHILE GEESE ARE NOT COMMONLY used forexperimental purposes, a relativelylarge experimental population of geesewas maintained at the Center for FoodAnimal Research (CFAR) in Ottawa. Sev-eral distinct stocks of Chinese, Emb-den, and Chinese X Pilgrim hybridgeese were developed in Ottawa tostudy production traits and DNA fin-gerprinting patterns (GRUNDER et al.1994). The stocks included standardbreed control strains, stocks selectedfor multiple traits, and the unselectedreference strains (SOMES 1988). As withchicken and turkey research in the earlypart of this century, most studies withthe relatively undeveloped pure andcrossed goose varieties have been re-lated to agricultural objectives, includ-ing methods of rearing broiler geese

9

Japanese quail. A minimum of 25 pairs, usuallymore than 150 birds, is needed to keep inbreed-ing at a minimum. These stocks are often keptas a source of “normal” control birds, and alsofunction as a resource stock, from which inbredor selected stocks can be derived or qualitativemutations isolated.

Inbred lines are produced by breeding togetherclose relatives for many generations, resulting inincreasingly homozygous and homogeneousprogenies. Different types of mating schemes areused, depending on how rapidly the researcheris attempting to approach complete homozygos-ity. Disregarding parthenogenesis, the mostrapid inbreeding is produced by father-daughter,mother-son, or brother-sister (full-sib) matings.These breeding schemes can also be used to ex-pose deleterious recessive traits or to fix pre-ferred or beneficial single-gene traits in a popu-lation. Unfortunately, even in the absence of ma-jor genetic defects, the fertility and viability ofthe inbred offspring are almost always lowerthan the more outbred parent strain, a charac-teristic called inbreeding depression. If selectionand breeding strategies do not compensate forthis decline, inbreeding depression can result inthe extinction of the line within a few genera-tions. This is a particular concern in lines propa-gated by full-sib matings which also have largegenetic loads (many deleterious alleles). How-ever, once the lethal and sub-vital alleles havebeen purged from the inbred strain, it can theo-retically be bred to essentially complete homozy-gosity while maintaining reasonable reproductiveperformance traits (fertility, egg hatchability, eggproduction rates, viability, etc.). Such geneticallyuniform stocks can then be used as a standardgenetic background in the study of individualgenes and gene complexes. Par-ticularly useful inbred lines arethose which have been bred forcontrasting phenotypes due toallelic differences at single loci.These lines, having the samegenetic background for practi-cally all loci except for the alle-les of interest, are calledcongenic lines. They are used tostudy single-gene effects on pro-ductivity, for molecular charac-terization of genes affecting de-velopmental traits or diseaseresistance, and many other usesin basic biological and biomedi-cal research (ABPLANALP 1992).

Highly inbred genetic stocks are invaluable ina wide range of research fields, particularlygenomics (gene mapping) and immunogenetics(Box 7). A good example of the usefulness of in-bred strains in genomics is the mapping of clas-sical mutations. While over 80 classically identi-fied genetic mutations have been assigned to thechicken linkage map, only a few have been lo-cated on the molecular map. This is due to thelack of genetic characterization of the exhibitionbreeds and lines in which most of these muta-tions are found. Such nonuniform genetic back-grounds make them difficult to use in matingsdesigned to integrate the maps. In contrast,congenic lines, mentioned above, are uniquelyuseful for such genetic mapping. The integrationof genes in exhibition breeds into defined inbredlines would provide the necessary uniformity formolecular mapping of these traits.

Long-term selected stocks are the result ofmany generations of testing and selective breed-ing for traits governed by multiple genes (the so-called quantitative or polygenic traits). Many val-ued heritable characteristics in the poultrybreeds belong in this category. These include eggproduction rate, egg size, feed efficiency, fertility,hatchability, viability, disease resistance, bodysize and shape, and behavioral characteristics.To change the population mean for one or moreof these quantitative traits requires rigoroustesting and ranking of the individuals and familygroups for the traits-of-interest each generation,followed by selective breeding of the higher-ranked individuals and families to produce thenext generation. Many factors can affect the rateof improvement in response to selection includ-ing 1) degree of heritability of the trait or traitsinvolved, 2) selection stringency, 3) level of in-

Box 7. Highly inbred stocks in immunogenetics

BASIC INFORMATION ABOUT FACTORS control-ling disease resistance in the chickenhas been gathered largely from stud-ies with congenic strains of chickens(birds with identical, highly inbredbackgrounds but different major his-tocompatibility complex (MHC)haplotypes; ABPLANALP 1992). A num-ber of these congenic strains have beendeveloped at the University of Califor-nia–Davis, the USDA Avian Disease andOncology Laboratory in East Lansing,MI, the University of New Hampshire,and Iowa State University. Researchershave shown how each MHC-haplotypecould directly affect the resistance ofa bird to a variety of different diseases,

including coccidiosis, Newcastle’s dis-ease, and the tumor-inducing virusesthat cause Marek’s disease and lym-phoid leukosis. The congenic MHCstrains, most requiring at least ten gen-erations of back-crossing and blood-testing to develop, are key resourcesrequired for furthering our understand-ing of the way the MHC genes func-tion. Studies with these stocks have al-ready given the primary poultry breed-ers vital information to use in deter-mining the best of several alternativebreeding strategies to enhance diseaseresis-tance potential of their produc-tion stocks.

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breeding, and 4) genetic variation in the originalsource population. Selected stocks usually re-quire several generations to develop, tend to re-vert towards the original stock values if the se-lection pressure is lifted (i.e., if random or non-selected pedigree reproduction is used), andusually need to be reproduced in large numbers(several hundred birds) each generation for thebest selection differential with minimized in-breeding.

Mutant stocks incorporate one or more of themany single-gene mutations that have a majoreffect on specific morphological or physiologicaltraits. These include variants (alleles) that affecteggshell color, feather color or shape, skin color,comb shape, metabolic function, major histo-compatibility complex (MHC) haplotype identity,and pattern formation in the developing embryo.The wide array of mutations affecting feathercolor and shape are important for distinguishingbetween breeds and varieties within breeds. Inthe poultry industry, eggshell color, skin color,feather color, and feathering rate mutations,and, more recently, MHC types, have all playedimportant roles in the development of commer-cial strains and varieties. Of particular interestto biomedical researchers are those mutationsthat cause disease conditions that mimic humangenetic disorders, including muscular dystrophy,scoliosis, scleroderma, and a variety of develop-mental mutations (usually lethal) that affect thedevelopment of the face, limbs, integument, andinternal organs.

Cytogenetic variants are birds that have chro-mosomal abnormalities, such as aneuploidy,polyploidy, translocations, and large insertionsor deletions. A small number have been estab-lished in the chicken, and these have provideduseful model systems for the study of meiosis,inheritance, recombination, linkage, transcrip-tional regulation, and gene dos-age effects. Such stocks in-clude: aneuploidy for the chro-mosome encoding the MHC andnucleolar organizer region(NOR), complete triploidy (threecopies, instead of two, of allchromosomes), large deletions(the mPNU line, in which thereis segregation of an MHC/NORchromosome with a deletedNOR), and various stocks carry-ing translocations betweenmacrochromosomes (Box 8).

Transgenic stocks areformed by inserting foreign

DNA, usually containing a gene of interest, intoone of the chromosomes of germline or somaticcells. While some transgenic chickens have beenproduced in the past few years (SALTER et al.1986; 1987; SALTER and CRITTENDEN 1989), thecreation of transgenics is still very much experi-mental in chickens and other avian species.However, a number of research groups continueto develop and refine transgenic methodologies,and report promising advances in the productionof transgenic birds (SALTER et al. 1987; LOVE etal. 1994; THORAVAL et al. 1995; MARUYAMA et al.1998).

RRRRResearch genetic stocksesearch genetic stocksesearch genetic stocksesearch genetic stocksesearch genetic stocksGenetic stocks are used in three areas of re-search: agricultural, biomedical, and basic orfundamental biological research.

Agriculturally important avian genetic stocksprimarily include those selected for various pro-duction-related characteristics (egg production,body shape, feed-use efficiency, leg strength, dis-ease resistance). Another use for such stocks isto provide a flexible, rapidly responding modelsystem for testing breeding techniques and sys-tems that might also be useful with large live-stock species (e.g., pigs, sheep, and cattle). Thesestocks are particularly vulnerable to fundingcuts due to the long development period neededfor most selected stocks, and the relatively largenumbers that must be produced and monitoredannually to produce the selected population.

Biomedical research specifically uses animalmodels for the study of various human diseases.Avian models, mostly in the chicken, exist forthe autoimmune forms of vitiligo, scleroderma,and thyroiditis, as well as for various develop-mental defects, such as polydactyly, scoliosis,and cleft palate. Genetic stocks are also used inavian health research for studying the nature of

FROM THE MID-1960S TO THE 1980s, ani-mal genetics laboratories at Ohio StateUniversity, the University of Minnesota,and New Mexico State University de-veloped about 40 different chromo-some rearrangement strains in thechicken (ZARTMAN 1971; WOOSTER et al.1977; WANG et al. 1982). A number ofstudies by these laboratories made im-portant contributions to our under-standing of chromosome behavior inavian species (including recombination,chromosome segregation, identifica-tion of pseudoautosomal regions on

Box 8. Chicken chromosome rearrangement stocks

the sex chromosomes, and sources ofaneuploids). Unfortunately, with thelack of support by various agenciesover the last ten years, over thirty ofthese unique genetic resources wereirretrievably lost. The seven still in ex-istence, along with a recently isolatedspontaneous translocation, are cur-rently being maintained at the Univer-sity of Wisconsin. However, with de-partmental reorganizations and bud-getary difficulties, these stocks are alsothreatened.

11

mercial importance, such as the sex-linked genecontroling the rate of feather growth that hasbeen heavily utilized by modern chicken breed-ers (Box 9).

In marked contrast to the general perceptionthat commercial poultry stocks all have a rela-tively small and diminishing genetic base, someresearchers have reported the opposite. Specifi-cally, DUNNINGTON et al. (1994) used DNA finger-printing to measure variability among commer-cial chicken breeding populations and concludedthat a considerable reservoir of genetic diversityyet remained. IRAQI et al. (1991) reported a greatdegree of polymorphism for endogenous viral (ev)genes in five egg-type populations maintained byan Israeli commercial breeder. AARTS et al. (1991)also found variation for ev genes among andwithin six WL and four medium-heavy browneggshell lines. While none of these methods spe-cifically reflects the variation remaining in genesassociated with economically important traits,the recent substantial progress in the develop-ment of the genetic map of the chicken (CHENG etal. 1995; CHENG 1997) should soon lead to morethorough and realistic assessment of the amountof economic trait variability remaining in com-mercial poultry populations.

FFFFFancy breeds and mid-levelancy breeds and mid-levelancy breeds and mid-levelancy breeds and mid-levelancy breeds and mid-levelproduction stocksproduction stocksproduction stocksproduction stocksproduction stocks

For at least 50 years, poultry fanciers have beenthe main conservators of the majority of the

disease resistance and effects of specific geneson productivity under disease stresses.

Most of the genetic stocks are of value forstudying questions in basic biology that maylead to more applied biomedical or agriculturalresearch, or by simply contributing to the knowl-edge of how different biological systems functionin a wide variety of studies in the life sciences.

Some of the specialized stocks have beenderived directly from commercial chicken, tur-key, or Japanese quail lines, while others weredeveloped from special breeds, landraces, orwild-types.

Commercial stocksCommercial stocksCommercial stocksCommercial stocksCommercial stocksThe commercial poultry stocks have made re-markable genetic progress in the last 50 years(Boxes 9, 10, and 11). At this time, selectedstocks used in commercial egg or meat produc-tion must fit very specialized production criteria.To develop these criteria, each breeding companyhas identified particular commercial goals (eggproduction, weight gain, feed conversion, carcasscharacteristics, etc.) and seeks to meet them inthe shortest possible time (EMSLEY 1993). In thisway, the fundamental difference between basicand applied research is highlighted. While a re-searcher may have a preferred outcome for anexperiment, any result can provide useful infor-mation to that researcher or others in the re-search community. For the commercial breeder,the only outcome that is acceptable is one thatimproves the commercial product for the con-sumer, and increases final prof-itability for the producer(HUNTON 1990).

From a commercial produc-tion point of view, the loss ofunique avian germplasm has anumber of negative repercus-sions. To start with, productionobjectives and economic stan-dards are constantly changing,particularly for meat productionbirds. This means that agro-nomic industries will continueto need access to genetic diver-sity to meet future market de-mands, to adapt to adverse en-vironmental conditions, to fightnew diseases, and to meet thedemands for different nutri-tional values. Thus, an effortmust be made to identify andconserve all useful genetic re-sources that could have com-

chicks are all fast feathering. Suchchicks can be easily sexed at hatch timeby the relative feather growth by any-one with a minimum of training. Previ-ously chicks were sexed using the ventsexing method, whereby rudimentarycopulatory organs were examined todetermine sex. This was a costly pro-cedure, and at $0.03 per chick wouldcost a hatchery $3,000 for every100,000 chicks hatched. Over 600 mil-lion egg-type chicks are hatched an-nually in the US. If only half of theseare sexed by feather sexing, thechicken industry saves over $9 millionper year. Broiler breeders are incorpo-rating this gene also, as sex-separaterearing becomes more prevalent. Withover 9 billion broilers hatched in theUS each year, this also will have an eco-nomic advantage to the industry.

Box 9. Economics of sex-linked genes and chicken genetics

IN 1908, SPILLMAN REPORTED that the fe-male was the heterogametic sex inchickens (now described as ZW, as com-pared to mammals where the male isheterogametic, XY). This was based onthe finding that the barring gene wasinherited as a sex-linked gene, beingpassed from the dam to her sons. Thisearly finding has played an importantrole in commercial poultry breeding,as many lines are now sexed at hatchtime using the sex-linked rate-of-feath-ering gene. This gene influences de-velopment of the early wing feathersin the chick. If the dam carries the slowfeathering mutation, K, she passes thison to all her sons, and her W chromo-some to her daughters. If the sire ispure for the wild type gene, k+, all thedaughters receive the wild type fast-feathering gene. The male chicks areall slow feathering and the female

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poultry breeds and varieties in North America,particularly the old dual-purpose or mid-levelproduction breeds (Box 12). As the Leghornchicken, Rock-Cornish cross chicken, andbroad-breasted Large White turkey became thedominant commercial birds, commercial breed-ers could see no economic benefit to maintainingother standard breeds and varieties of poultryrecognized by the American Poultry Association(APA 1998). Today, without fanciers, it would bevery hard to find an Ancona or Silkie chicken ora Royal Palm turkey. The Lamona chicken breed,developed by the USDA, is a notable Americanexample of a once-useful old-fashioned produc-tion strain now fallen from favor.

In some cases, access to mid-level stocks canhelp small-scale producers stay in business.While they cannot compete with the Rock-Cor-nish meat cross or Leghorn egg-layer in thehighly commercial marketplaces, they can be-come financially successful by raising some ofthese heirloom birds to supply specialized nichemarkets (Box 12). There are many other positive

aspects to this practice: small parcels of landcan remain agriculturally productive; open spaceis maintained, family farmers are aided; andmoneys go into the local economy.

Biomedical researchers are starting to be-come aware of the genetic reservoir available inthe fancy breeds. They usually seek specificstandard breeds or feather patterns that can beused in exploring biological questions (see Chap-ter 3) or problems related to human medical dis-orders, e.g., a form of vitiligo in barred chickens(BOWERS et al. 1994). The Silkie breed (Figure 9)is particularly useful, with six dominant muta-tions: crest (Cr), rosecomb (R), muffs-and-beard(Mb), polydactyly (Po), ptilopody (Pt), fibromel-anosis (Fm); and one recessive mutation, hook-less (h). These mutant alleles produce: elongatedfeathers on the crown of the head (Cr) and onthe face and chin (Mb), a broad, flattened combthat is covered with small, fleshy nodules (R),extra toes (Po), feathered legs and feet (Pt), darkskin, bones, and viscera (Fm), and loose, excep-tionally fluffy body feathers (h). Not only have

Figure 6. White Leghorn hen (Photo courtesyof U.K. Abbott, University of California–Davis).

90% of all the egg-type chickens inNorth America, and probably well overhalf of the commercial egg-type chick-ens worldwide.

Crosses among lines of the WhiteLeghorn (WL) breed produce nearly allthe commercially marketed white-shellchicken eggs in North America. The WLlines in use today stem from the pure-bred stocks sold in the 1930s and1940s. Though there has been inter-crossing in many cases to develop newstrains, many of the currently usedstocks appear to have been selectedwithout intermixing for 30 years ormore. Of particular note is the com-mon use of the Mount Hope strain,which is distinguishable by its largeegg size and the B-19 and B-21 majorhistocompatibility complex blood typeswhich it carries (for an explanation ofthe major histocompatibility complex(MHC) and B-blood types, see the sec-tion on Immunogenetics in Chapter 3).

In response to regional consumerpreferences, several commercialbrown-eggshell chicken lines have alsobeen developed. Typically less efficientthan the White Leghorn strains, com-mercial brown-eggshell chicken linesare usually produced by crossingRhode Island Red males with high pro-duction White Leghorn females. Alter-natively, some high egg productionstrains of Rhode Island Red or BarredPlymouth Rock may be used.

Box 10. Development of egg-laying stocks

COMMERCIAL EGG-LAYING chickens (Figure6) have shown a substantial increasein productivity in the past 60 years.Some of this improvement has beendue to developments in the areas ofmanagement, nutrition, and diseasecontrol, but the effect of genetic im-provement is clear (ARTHUR 1986). Be-tween 1940 and 1955, the number ofeggs laid per hen in the United Statesincreased from 134 to 192 (USDA-NASS1998). By 1994, eggs per hen had in-creased to 254. The change in the

1940s and 1950s was primarily due tothe introduction of hybrid stock, utiliz-ing pure breeds which had been underdevelopment by numerous small breed-ers participating in the National PoultryImprovement Plan (NPIP). The more re-cent increase has been primarily due toselection for increased egg numbers.However, it should be remembered thatthe work of the small breeders and theformal testing parameters set by NPIPshaped the foundation stocks, paving theway for the phenomenal performance in

the modern commercialbirds.

Today, only a few largeinternational poultrybreeding companies pro-duce most of the world’scommercial egg-typechickens. Just 40 yearsago, the 1958-59 sum-mary of US random-sample-egg-productiontests (ARS 1960) listed132 breeding firms. In themost recent egg-layer teststill conducted in NorthAmerica, (NORTH CAROLINA

COOPERATIVE EXTENSION SER-VICE 1996) only five breed-ing companies were listed.These were actuallyowned by just three inter-national firms. Thesethree firms breed over

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the hobby breeders helped in supplying suchresearch birds for one-time projects, but some ofthem have participated in long-term breedingprograms for researchers.

Although the majority of exhibition and mid-level production poultry breeds have continuedto exist under the rather informal stewardship of

the hobby breeders and the different breed orga-nizations, a number of problems are associatedwith their conservation: 1) most of the amateurconservators often only keep their stocks for ashort period of time (typically just five years);2) small-scale hobby breeders who get breedingstock from a central clearing house of poultry

Box 11. Development of meat-producing stocks

IN 1950, A COMMERCIAL BROILER took 84days to grow to 1800 grams; by 1970,this was cut to 59 days, and by 1988,it was down to 43 days (from HUNTON

1990). As with the egg-type chickens,a large proportion of the improved per-formance of meat birds can be attrib-uted to developments in the areas ofmanagement, nutrition, and diseasecontrol. But choice of foundationbreeding stock and early use of breedcrosses were also important in the de-velopment of the broiler industry.

More so than the egg market, thebroiler market is strongly consumer-driven (POLLOCK 1999). Early consumerinput (chicken of tomorrow competi-tions between 1946 and 1948) gavethe broiler-breeders and growers agood picture of consumer preferences:compact, well-fleshed carcasses at af-fordable prices. In other words, thescrawny, angular cockerels (Figure 7)available in large numbers from egg-selected Single-comb White Leghornlines did not even approach the con-

sumer ideal. The broiler-breeders werefortunate to have available the Cornishbreed (derived from fighting stock im-ported from India), which had many ofthe desired carcass characteristics. Thebreeders also found that the productioncharacteristics (body type, rate of gain,feed conversion) improved rapidly in re-sponse to selection. Unfortunately, im-provement in these areas had a strongnegative effect on the already poor re-production characteristics of the Cornishlines (low egg production, low fertility,poor hatchability, reduced chick viabil-ity), and seriously impaired disease re-sistance (see section on immunogenet-ics, Chapter 3). The early breeders foundthat crossing the Cornish roosters withhens from improved dual purpose breedssolved many of these problems. These“female” line breeds, including the Ply-mouth Rock and New Hampshire, havebetter body type than Single-comb WhiteLeghorns, yet lay eggs at a relatively highrate compared to the Cornish “male”lines. Today, the commercial sire is of-

ten a cross between twopredominantly Cornishstrains, and the commer-cial dam is a cross be-tween two strains de-scended from one or more

of the dual-purpose breeds. The out-bred or crossbred parents have betterreproductive traits and general vigorthan parents from the pure-lines, andtheir offspring, a three- or four-waycross, exhibit even more hybrid vigor.Unfortunately, despite careful evalua-tion of the breeding stock, some seri-ous structural and physiological prob-lems have surfaced that appear to bethe result of the intense selection fordesirable production characteristics.These include: leg weakness, cardio-pulmonary insufficiency, breast blis-ters, increased fat deposition, andmuscle anomalies.

While the turkey industry is consid-erably smaller than the broiler chickenindustry, many of the same breedingmethods have been used, and manyof the same problems have been en-countered (HUNTON 1990). With asmaller genetic base, and a muchlarger bird to start with (Figure 8), thestructural and physiological problemsfound in chickens are often magnifiedin turkeys. Considering the small num-ber of primary breeders (three) and thescarcity of exhibition or research tur-key breeding stock, it is imperative tosafeguard the remaining genetic diver-sity of this domestic species.

Figure 8. Commercial Large White turkey tom (Photo cour-tesy of R.A. Ernst, University of California–Davis).

Figure 7. Traditional broiler-type chickencarcass of the 1940s (Photo courtesy of F.A.Bradley, University of California–Davis).

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stocks may never know their egg source or thedegree of relationship of their foundation stock;3) breeding populations are often very small,particularly for the rarer breeds, and pedigreeinformation is frequently limited or not available;4) some hobbyists deliberately inter-cross differ-ent breeds or varieties in attempts to improve ormodify exhibition traits; 5) selection for produc-tion characteristics (e.g., fertility, viability, eggproduction, or disease resistance) may be largelyignored in these small-scale breeding programs,although de facto natural selection will tend toeliminate the infertile, disease-susceptible, orleast-viable individuals; and 6) backyard breed-ers tend to have problems in controlling diseasesand may have serious endemic diseases. If therewere a formal conservation pro-gram for avian genetic resour-ces, it would be logical for it toprovide technical services tothese hobbyists who are a veryimportant component of aviangenetic resources conservation.

Figure 9. Silkie rooster from UCD Silkie (Photo cour-tesy of J. Clark, University of California–Davis).

Box 12. Small renaissance of old-style chicken breeds

WITH THE INCREASING CULTURAL diversityof our population, the white-feathered,highly selected meat- or egg-produc-ing bird no longer meets the needs ofall consumers. In response to a greatdemand by ethnic markets and themany upscale restaurants searching forthe “chicken of yesterday”, more andmore small producers are starting toraise “old fashioned” mid-level produc-tion or dual purpose breeds (those thatare reasonably efficient at producingboth meat and eggs). These produc-

ers are getting their stocks from thefew people who still maintain popula-tions of true Rhode Island Reds, Speck-led Sussex, New Hampshires, and soon. Those supplying the specialty eggmarkets are also looking for differentbreeds to produce a colored egg thatwill be distinctive (brown, tan, green,or blue), such as Orpington, Rhode Is-land Red, and Ameraucana. Unfortu-nately, most of these so-called mid-level production breeds have all butdisappeared from American farms.

15

AAAAAvian genetic resources for biological researchvian genetic resources for biological researchvian genetic resources for biological researchvian genetic resources for biological researchvian genetic resources for biological researchResearch with a variety of genetic stocks has

produced many important discoveries in the ar-eas of genetics, virology, immunology, develop-mental biology, and animal agriculture. (seeCRAWFORD 1990, for a review). These include thepioneering studies on Mendelian inheritance andsex determination of BATESON (1902), and BATE-SON and PUNNETT (1906, 1908). Not surprisingly,given its history of use in genetic studies, thefirst linkage map reported for any domestic ani-mal species was that for the chicken publishedby HUTT (1936). The turkey has provided one ofthe few known vertebrate models for partheno-genesis (embryo formation in an unfertilized egg)(Box 5), and the wide range of metabolic and de-velopmental mutations in chicken (Boxes 8, 13)and quail have allowed researchers to start in-vestigating the molecular bases of these defects,and, eventually, to determine the normal func-tion of the affected gene. In virology, chickenhosts were used by ROUS (1911) to identify thefirst tumor-causing virus, Rous sarcoma virus.Later, the genetics of host resistance to RNA vi-ruses was elucidated (CRITTENDEN et al. 1963),and the Marek’s-like herpes virus found in tur-keys was used in the first successful vaccinationprogram against a tumor-causing virus in thechicken (OKAZAKI et al. 1970). More recently,BUMSTEAD and PALYGA (1992) described one ofthe first molecular genetic maps of a domesticanimal using DNA markers (RFLPs), and thechicken genome map continues to be among thebest developed of those for agricultural animals(LEVIN et al. 1994; CHENG et al. 1995; CROOIJMANSet al. 1996; CHENG 1997). It is by far the best-developed map for any avian species or any spe-cies with ZW sex chromosome determination. Inparticular, the usefulness of the chicken genomemap in cross-species comparisons was high-lighted at the First International Workshop onComparative Genome Organization (ANDERSSON

ALTHOUGH MOST WIDELY KNOWN for their utility inthe production of food (meat and eggs) and othercommercial products (feathers, vaccines, fertil-izer, etc.), chicken, Japanese quail and otherdomesticated avian species have long been apreferred experimental organism in applied andbasic life sciences. Some features contributing totheir popularity are:

• Availability (poultry, especially, are easilyraised and readily obtained, with inex-pensive common strains providing cost-effective research animals).

• High rate of embryo production com-pared to mammals.

• Accessibility of the avian embryo (the ex-ternally incubated avian embryo is avail-able for experimental manipulation dur-ing most stages of embryonic develop-ment).

• Excellence as a higher vertebrate animalmodel.

• Ease of surgical manipulations (surgicalprocedures on birds are much less likelyto result in infections due, perhaps inpart, to their high body temperatures).

• Extensive history of use (more than acentury of research provides avian scien-tists with needed information and geneticstocks to ask specific experimental ques-tions).

• Animal welfare considerations are gener-ally less complex for domestic birds thanthey are for other vertebrates.

3

16

et al. 1996), although workshop participants alsoexpressed concern that the continuing loss ofpoultry genetic stocks will impair some impor-tant comparisons with the human and other ge-nome maps. The chicken gene library was thefirst one described for an agricultural animalspecies and, along with the human gene library,was the first constructed for any vertebrate(DODGSON et al. 1979). Critical early observationsof gene structure, organization, and expressionwere made using chicken ovalbumin (WOO et al.1978), globin (DODGSON et al. 1979), histone(ENGEL and DODGSON 1981), insulin (PERLER etal. 1980), and lysozyme (STEINER et al. 1987)genes, among others.

AAAAAvian reproductive biologyvian reproductive biologyvian reproductive biologyvian reproductive biologyvian reproductive biologyBirds such as the chicken are uniquely adaptedfor the production of large numbers of offspring,in the form of fertile eggs, over relatively longperiods of time. In the wild, birds will usually laya series of eggs on sequential days until theyhave as many eggs as they can comfortably incu-bate (a “clutch”). At that time, they will stop lay-ing eggs and start to incubate or brood the eggs.However, during their long domestication, chick-ens have been selected for large clutch size andnonbroody behavior, so that hens today will con-tinue to lay at a high rate for at least a year,

given proper conditions. In fact, chickens fromsome commercial stocks that have been selectedfor egg production may average over 280 eggs ina one-year period, with fertility of greater than95%. While genetic strains used in research areusually less productive, most produce at least100 eggs per hen per year, providing researcherswith an abundant, dependable source of indi-vidually packaged embryos of known genotype.

The functional reproductive system of mostfemale birds, including the chicken, is composedof the left ovary and the oviduct. The ovary con-tains ova in varying stages of development, withthe largest being released at regular intervals.The oviduct is a tubular organ, consisting of sev-eral parts: the funnel-shaped infundibulum,where each ovum enters the oviduct and fertili-zation occurs; the wide, thick-walled, magnum,where the albumen is produced and depositedaround each of the ova; the narrower, thinner-walled isthmus, where the shell membranes aremade; the large, pouch-like shell gland, wherethe egg is “plumped” with fluids and electrolytes,before the calcium carbonate shell is depositedon the shell membranes; and, finally, the vagina,with its specialized sperm storage sites at theutero-vaginal junction, where spermatozoa canremain viable for more than a week (VAN KREY1990; BAKST 1993). Thus, the oviduct is whereboth the fertilization and the packaging of ovum

Box 13. Inherited muscular dystrophy of the chicken

GENETIC MUSCULAR DYSTROPHY (GMD) ofthe chicken was first formally describedin 1956 in selected New Hampshirestrains at the University of California–Davis. Since then, literally hundreds ofreports have been published on the ab-normality. The disorder is caused by asingle gene; but little is known of themolecular nature of the defect (PESSAH

and SCHIEDT 1990; WILSON 1990;IANNACCONE et al. 1995).

Symptoms of the disorder typicallyappear during late embryogenesis andneonatal muscle maturation. Muscleswith fast twitch, alpha-white muscle fi-bers such as the superior pectoralisand biceps are most affected. These“white” muscle fibers exhibit irregular-shaped rounded fibers, both larger andsmaller than normal. Eventually themuscles atrophy and are replaced byfat and connective tissue. The early ap-pearance of GMD in fast twitch but notin tonic fibers suggests it is expressedin a particular myogenic lineage.

Studies have shown that the prob-lem with dystrophic muscle is not afaulty set of neural instructions. Limb

bud transplants demonstrated that dys-trophic muscles grafted to a normal hostand its nerve were phenotypically dys-trophic, and normal muscles trans-planted to a dystrophic host were nor-mal, indicating the problem was in themuscle and not directly its innervation.Thus, if neural influences are involved inGMD, it is likely it is the dystrophicmuscle that is unable to respond prop-erly to signals from its phenotypicallynormal nerve.

One of the major uses for the dystro-phic chicken has been to test therapies,including an elevated oxygen levels(ASHMORE and SOMES 1966), the drugs D-penicillamine (which affects collagen for-mation; PARK et al. 1979), cyprohepta-dine, methysergide, and diphenylhydan-toin. A Muscular Dystrophy Association-supported drug screening programachieved promising results with steroidssuch as corticosterone-21-acetate(C21A). The most effective treatments se-verely reduced the growth rate of thechicks, as if the synthesis of defectiveproteins play a role in expression of theabnormality.

The discovery of and sequencing ofdystrophin, the gene associated withDuchenne muscular dystrophy in thehuman and the mdx dystrophy in themouse, added a new dimension to thestudy of muscle abnormalities. Thedystrophin protein itself is absent inhumans with Duchenne dystrophy andin the mdx mouse. This led the Muscu-lar Dystrophy Association to focus itsefforts on the human and mouse formsof the disease. But it is still not knownwhether or not the dystrophin geneproduct is defective in GMD of thechicken. It is known that dystrophin innormal chickens is closely homologousto the mammalian gene, and that adystrophin protein is present in GMDchickens. It is not known if this pro-tein is the same as the one in normalchickens. Future research focusing onthe role of the dystrophin gene, estab-lishing expression of the abnormalityin cell culture, and studying the fac-tors that regulate maturation of nor-mal muscle are worthy of study.

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occurs, producing the neat, aseptic associationof nutrients and germplasm that will ultimatelyform the chick. Embryonic development pro-gresses as the egg moves down the oviduct, sothat when the egg is laid 24 to 27 hours afterfertilization, the embryo is a radial dome of40,000 to 60,000 cells that is already formingdistinct layers (epiblast and hypoblast). At thisstage of development, the embryo is comprisedof cells that are morphologically undifferenti-ated (EYAL-GILADI and KOCHAV 1976; WATT et al.1993) and pluripotential, i.e., able to differenti-ate into a number of different cell types(PETITTE et al. 1990).

Numerous studies on avian reproductionhave been conducted since the turn of the cen-tury, particularly on the control of fertility andegg production (HUTT 1949; LERNER 1950,1958). Consequently, much is known aboutgenetic and environmental factors that influ-ence or govern reproduction in the male andfemale chicken, turkey, and Japanese quail.Some of the genetic variants affecting repro-ductive traits that were studied include: re-stricted ovulator, multiple ovulator, riboflavintransport deficient, and male infertility associ-ated with the rose-type comb.

EmbrEmbrEmbrEmbrEmbryologyyologyyologyyologyyologyThere is a long and rich history of use of theavian embryo as a model system for study ofvertebrate development (ROMANOFF 1960). Be-ginning with Aristotle (384–322 BC) whoseHistoria Animalium includes the first knownfully preserved description of a chicken em-bryo, researchers and natural science histori-ans have found bird embryos to be a readilyavailable, dependable source of embryos thatcould be manipulated and observed during thecourse of development independent of andwithout harming the female parent (a majorissue when studying mammals). Its easy ac-cess during organogenesis, relatively inexpen-sive cost, and bilaterality providing opportuni-ties for contralateral controls for microsurgeryand other perturbations have made the avianembryo one of the best characterized verte-brate models, both in classical and molecularterms (MACCABE and NOVEROSKE 1997; MORGAN1997). Another factor that has made the avianembryo the model of choice for studies of ver-tebrate development is the respectable catalogof existing mutations (listed in ABBOTT 1967;SOMES 1988; and CRAWFORD 1990). Avian em-bryos have also proven uniquely adapted tointerspecific tissue grafting because of distinct

differences between the appearance of chickenand Japanese quail cell nuclei. Selected earlyembryonic cells from quail have been graftedinto chicken blastodiscs, which, after carefulincubation, reveal exactly how the donor quailcells are distributed in the later embryos (DIE-TERLEN-LIEVRE and LEDOUARIN 1993; DIETER-LEN-LIEVRE 1997). Such studies have permittedthe development of a detailed fate map of theearly blastodisc. Additionally, avian embryoshave been used extensively in teratologicalstudies, in which their response to chemical,nutritional, hormonal, and environmentalchallenges were evaluated to provide guide-lines for identifying causes of nongenetic devel-opmental defects (LANDAUER 1967, 1973), and,eventually, for identifying the biochemicalmechanisms disturbed by such exposures.Avian embryos have also been of value in stud-ies of cell proliferation related to cancer-likecell growth or programmed cell death inherentin certain stages of embryonic development(SANDERS and WRIDE 1997).

Models for human geneticModels for human geneticModels for human geneticModels for human geneticModels for human geneticdiseasesdiseasesdiseasesdiseasesdiseases

Animal genetic defects have long been used asmodel systems in the study of human dis-eases. A few of the avian genetic disorders thathave proven especially useful include the au-toimmune forms of avian vitiligo (AUSTIN et al.1992; AUSTIN and BOISSY 1995; ERF et al.1995), scleroderma (GERSHWIN et al. 1981;ABPLANALP et al. 1990; VAN DE WATER et al.1994; SGONC et al. 1995), and thyroiditis (re-viewed by ROSE 1994), the polygenic avianscoliosis (RUCKER et al. 1986; LIEN et al. 1990;MOCHIDA et al. 1993), the autosomal recessiveform of muscular dystrophy (PESSAH andSCHIEDT 1990; WILSON 1990; IANNACCONE et al.1995) (Box 13), and a polygenic muscle defectin the broiler chicken (Box 14). Other knownavian mutations mimicking human disordersinclude: autosomal and sex-linked dwarfism,lamellar ichthyosis, poly- and hypodactyly,gouty uremia, genetic obesity, several types ofmicromelia, glaucoma, a non-autoimmuneform of vitiligo (BOWERS et al. 1994), and a vari-ety of neurological disorders (SOMES 1988;CRAWFORD 1990).

Japanese quail disease models have alsoattracted the attention of biomedical research-ers. For example, at the University of BritishColumbia, quail serve as a model in the studyof atherosclerosis, and in age-related macular

18

degeneration (AMD). This condition is the mostcommon cause of blindness in humans over theage of 65. Except for primates, the quail is cur-rently the only suitable nonhuman species inwhich to study this disease.

Despite the proven utility of existing strainsof chickens, turkeys, and Japanese quail in bio-medical research, fewer and fewer researchersare making use of these unique genetic stocks(WILSON 1990). Contributing to this decline arethe increasing costs and the more technical (andcostly) nature of many studies that have biasedresearchers towards smaller vertebrate labora-tory species (rats and mice). Whether or not it isthe best possible choice, the default conditionfor medical research continues to be the rodent,ultimately limiting the study of disorders anddiseases to those suitable to such animal mod-els. Enhanced use of exceptional avian modelsdepends on the continued accessibility of exist-ing stocks and the development and mainte-nance of new genetic stocks.

ImmunogeneticsImmunogeneticsImmunogeneticsImmunogeneticsImmunogeneticsNatural genetic variation in the major histocom-patibility complex (MHC) has been studied ex-tensively in the chicken, particularly as it relatesto disease resistance (GUILLEMOT et al. 1989; BA-CON and DIETERT 1991; HELLER et al. 1991; SUNGet al. 1993; KEAN et al. 1994; SCHAT et al. 1994;BACON and WITTER 1995; HEMENDINGER et al.1995). The different MHC haplotypes have beensystematically integrated into highly inbredstocks to control or eliminate complications inimmune response introduced by subtle differ-ences in the genetic background (ABPLANALP1992; Box 7). These studies led to a better un-

derstanding of the complexities of the vertebrateimmune system. They have also provided theframework for the development of disease-resis-tant poultry stocks (LAMONT 1994) and for under-standing the differential effectiveness of certainvaccines in chicken strains with different MHChaplotypes (BACON and WITTER 1992).

In studies which focused on cell-mediatedimmunity and macrophage function, researchersshowed that genetic differences between selectedstrains can also affect the ability of a given birdto promote or suppress Rous sarcoma tumorformation (QURESHI and TAYLOR 1993).

The relationship between MHC haplotype andimmunocompetence is currently attracting theattention of the commercial sector. This is due toa strong negative correlation between character-istics such as improved feed conversion andmore rapid growth rates with a weaker antibodyresponse to specific antigen challenges and adecreased effectiveness of cell-mediated immu-nity, both of which are related to MHC function(GAVORA 1990; HELLER et al. 1991). This trend inimmune responsiveness of commercial egg-lay-ing Leghorn strains (small, slower-growing birds)is compared with that of commercial broilerstrains. In addition, a distinct loss of immuno-competence was shown in a study that com-pared two broiler-type lines, one descended fromcommercial stocks developed in Canada in 1957,and the second from modern commercial stocks(QURESHI and HAVENSTEIN 1994). Thus, commer-cial broiler-chicken breeders are now showinginterest in improving immune system function tooffset the immune depression that has beenbred into their stocks with long-term, intensiveselection for meat production traits.

Box 14. Deep pectoral myopathy: One price of intensive selection

with better circulation and a differentbreast muscle configuration still ac-ceptable to the public which wouldprobably require outcrossing to the un-affected, unimproved genetic stocks.

DPM has its human counterpart. Itis one of a class of pathological situa-tions known as “Compartment Syn-dromes”. One of these is anterior tibi-alis syndrome or “March Gangrene” inwhich hard exercise by untrained orsusceptible individuals (such as mili-tary recruits on their first forty-milehike of basic training) results in per-manent muscle damage similar to thatfound in turkeys and large broilers.

able in as little as 15 minutes when broil-ers were induced to flap their wings forshort periods of time.

Polygenic physiological problems arenot simply resolved. Food preferences ofthe public and the economic forces ofthe poultry industry favor selection forlarge breasts maintaining a physiologi-cal situation in which deep pectoralmuscles predisposed to ischemia maycontinue to be a problem. Indeed, SILLER

(1985) said that the disorder was a “pen-alty of successful selection”; “...wild tur-keys and less intensely selected old com-mercial strains are apparently not sus-ceptible to DPM, it is obvious that thisdisease is man-made”. One solutionwould be to breed broilers and turkeys

DEEP PECTORAL MYOPATHY (DPM) is char-acterized by death of the mid-regionof the supracoracoideus muscle of thelarge broilers and marketable turkeys.It is also known as “Green Muscle Dis-ease” and “Oregon Muscle Disease”.

A series of investigations by Siller,Harper, and their colleagues (WIGHT

and SILLER 1980; WIGHT et al. 1981;HARPER et al. 1983; SILLER 1985; WILSON

et al. 1990) demonstrated that the my-opathy was due to an ischemia causedby swelling of the muscle during exer-cise. An enlarging muscle, an inelasticmuscle fascia, and a rigid sternum cre-ate a situation in which circulation iscut off to the mid-region of the muscle.Ultrastructural changes were detect-

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which includes individuals with two (normal ordisomic), three (trisomic), or four (tetrasomic)copies of chromosome 16 (a microchromosome).The biological consequences of aneuploidy andgene dosage effects on growth, development andimmunity have been the subject of many studies(DELANY et al. 1988; QURESHI et al. 1989; HEMEN-DINGER et al. 1992; LEPAGE et al. 1996) and con-tinues to be important in current research. TheTrisomic Line has been particularly useful instudying chromosome dosage effects on regula-tion of MHC expression, with tetrasomics andtrisomics expressing distinctly higher levels ofMHC-products on their cell surfaces than disom-ics (MUSCARELLA et al. 1987; DELANY et al. 1988;HEMENDINGER et al. 1992).

The mPNU line segregates for a large deletionin the nucleolar organizer region of chromosome16, giving it a reduced number of rRNA genes(DELANY et al. 1995). This line was used to estab-lish the developmental threshold (i.e., lethallimit) for rRNA gene copy number for the firsttime in a higher vertebrate (DELANY et al. 1994,1995). The Trisomic and mPNU lines were in-strumental in establishing the location of a newlocus, Rfp-Y (Box 15) on this microchromosome(MILLER et al. 1994, 1996).

The CSIRO Triploid line, maintained in Aus-tralia, is an important higher vertebrate polyp-loid model. Studies of the line have contributedinformation regarding errors of meiosis, geneticbasis for inheritance of meiotic errors, sex deter-mination, gonadal differentiation, sex reversal,and polyploidy effects on growth and develop-ment (LIN et al. 1986; THORNE et al. 1987, 1988,1991, 1997; SOLARI et al. 1991; THORNE andSHELDON 1991).

GenomicsGenomicsGenomicsGenomicsGenomicsGenomics is a rapidly evolving field of sciencethat emphasizes the study of complete genomesand chromosomes (Box 16). Genomic studieshave also changed in focus, from the general or-ganization of the genome to the identificationand localization of functional genes. Genomicresearch is well advanced with the chicken.

As more is learned about thegenetic mechanisms controllingdisease resistance and suscep-tibility, researchers will be ableto narrow their focus to specificgenes. A few of these genes arenow known, but the majorityremain to be identified. As withplant genetic stocks, some re-sistance genes may well befound in stocks not now commercially useful,e.g., wild or feral populations and breeds suchas Ancona chicken (Box 15) or other relativelyunimproved populations (GAVORA 1990).

CytogeneticsCytogeneticsCytogeneticsCytogeneticsCytogeneticsThe chicken has proven to be a good vertebratecytogenetic model, despite the complexity of itskaryotype (the majority of the 78 chromosomesare exceptionally small microchromosomes; seeFigure 10). In particular, unique cytogeneticconditions in the chicken have encouraged biolo-gists to use the chicken as a means to study al-tered chromosome constitutions (e.g., gene dos-age effects) on the biology of this and otherhigher vertebrates. The chicken is exceptionalamong the higher vertebrate animal group inthat several viable genetic strains exist whichshowcase the effects of chromosomal aneup-loidy, polyploidy, or large deletions, several ofthese occurring in microchromosomes.

The linkage and chromosomal location of themajor histocompatibility complex (MHC) with thesingle nucleolar organizer region (i.e., the rDNAlocus) were determined in studies of birds fromthe Trisomic Line (BLOOM and BACON 1985),

Figure 10. Chromosomes from a Red Jungle Fowl henfrom UCD 001 (Photo courtesy of M.E. Delany, Univer-sity of California–Davis).

AN UNUSUAL GENE CONFERRING disease re-sistance was recently discovered in therare Ancona breed of chicken. A uniquecopy of a gene outside of the knownmajor histocompatibility complex(MHC) was identified that imparts re-sistance to the commercially trouble-

Box 15. Ancona chickens give a new angle on disease resistance

some Marek’s disease virus. This gene,Rfp-Y, now also identified in otherchicken breeds, is on the same chro-mosome as the MHC, and is still beingcharacterized (BRILES et al. 1993;WAKENELL et al. 1996; MILLER et al. 1994,1996).

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Genome map development involves coordi-nated approaches (BURT et al. 1995): 1) expand-ing and improving the genetic linkage maps ofchickens and other poultry, 2) using markers,such as microsatellites, that have high utility,improving the cytogenetic maps available forbirds, and 3) making new efforts towards inte-grated physical (recombinant DNA clone-based)maps. Candidate gene research involves a vari-ety of genes of interest, including those encodingthe major histocompatibility complex genes,other immune-response genes, ribosomal RNAgenes, cytokine and other hormone-encodinggenes, and genes related to muscle and mor-phology. Finally, several efforts are underway tomap and identify anonymous genes associatedwith disease resistance, growth and reproduc-tion, and general viability of poultry.

The interdependence of genome research andgenetic resource conservation can hardly beoverstated. Inherent to all genetic experimenta-tion and breeding is the fundamental require-ment for allelic diversity. It is preferable that di-verse alleles be preserved in well-characterized,inbred (homozygous) lines. As a concrete ex-ample, it is widely agreed that one of the reasonsthat the laboratory mouse has become the mostimportant model system for biomedicine hasbeen the availability of diverse inbred lines (pio-neered most notably by the Jackson Laboratory).COPELAND and JENKINS (1991) used such inbredmouse strains in their intraspecific backcrossreference family approach to gene mapping. Asimilar strategy was followed in developing theEast Lansing chicken mappingreference population, derivedfrom the segregating progeny ofa cross between two inbredlines: Red Jungle Fowl (UCD001) and White Leghorn (UCD003), genetic stocks developedby H. Abplanalp at the Univer-sity of California, Davis (CRIT-TENDEN et al. 1993). The twolines are highly divergent, dis-playing a high degree of inter-line polymorphism. The diverseand inbred character of theselines has been a major con-tributor to their subsequentutility. In contrast, BUMSTEADand PALYGA (1992) used ani-mals from two divergent Leg-horn lines as the parents fortheir reference backcross panel.Cheng and Bacon (Avian Dis-ease and Oncology Laboratory,

personal communication) have used inbred lines(RPRL 6I2 and 7I3) for mapping of non-MHCgenes which are responsible for resistance/sus-ceptibility to Marek’s disease virus. More gener-ally, BACON’s (1987) congenic MHC lines havebeen critical for a variety of genetic analyses ofthe avian immune system. Again, the speed withwhich this work was accomplished was en-hanced and it could only have been accom-plished using the available well-characterizedgenetic lines.

One long-term objective of avian genomics isto develop an understanding of genes respon-sible for economically important traits in poul-try. These are variously known as quantitativetrait loci (QTLs) (VERRINDER GIBBINS 1993) or eco-nomic trait loci (ETLs). The ETLs are analogousto, and sometimes models of, complex genetictraits of humans and other species. In somecases, the protein product of these genes is al-ready known. In other cases, researchers seek toidentify and further characterize previously un-known ETLs (anonymous genes). Among thegenes under study are those encoding diseaseresistance traits. These include major histocom-patibility genes, cytokine genes, and anonymousgenes encoding Marek’s disease virus resistance.Other genes of interest include those whichregulate growth, reproduction and morphology(e.g., endocrine genes, collagen and other boneand connective tissue genes, anonymous ETLs),and genes which relate to general viability andcellular function (e.g., ribosomal RNA genes,

THE US GOVERNMENT IS HEAVILY investedin genome research. In response to thecall for a USDA National Genetic Re-sources Program in the 1990 Farm Bill,the USDA created the National AnimalGenome Research Program (NAGRP),administered by Cooperative State Re-search, Education, and Extension Ser-vice (CSREES) with Richard Frahm asDirecctor. The NAGRP was developedas an equal companion to the NationalAnimal Germplasm Program (NAGP),administered by Agricultural ResearchService (ARS) with Roger Gerrits as di-rector. The essential interrelationshipof genome research and germplasmresources is emphasized by the com-mon origin of and close ties betweenthe NABRP and the NAGP. Suppport foragricultural animal genomics comesprimarily from the USDA as direct sup-port to ARS laboratories, such as the

Box 16. Government involvement in genome research

Avian Disease and Oncology Laboratory(ADOL) in East Lansing, the NationalResearch Iniative Competitive GrantsProgram, and limited funding throughCSREES to Regional Research Programs(e.g., NC-168) and a National ResearchSupport Program, NRSP-8. NRSP-8 pro-vides for coordination of animal ge-nome research via the appointment ofSpecies (Poultry, Cattle, Sheep, Swine)Technical Committees and Coordina-tors. Parallel developments in animalagriculture genomics have occurredworld-wide, particularly in Europe (e.g.,Institute for Animal Health, Compton,England; Roslin Institute, Edinburgh,Scotland; University of Wageningen,The Netherlands). One of the majorgoals of the NAGRP is to cooperate withinternational efforts to enhanceprogress in animal genetics.

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anonymous viability genes, and neuropeptide/neurotransmitter genes).

Two major strategies are being used to locateand characterize genes of interest. In the candi-date gene approach, potential ETL-associatedgenes are chosen based on the physiological orgenetic effects of the proteins or RNAs they en-code as studied either in poultry or other spe-cies. The candidate gene is then isolated by re-combinant DNA techniques, further character-ized, and used to generate markers to test itsusefulness as an ETL indicator in appropriatepopulations. In the map-based approach, anony-mous ETLs are mapped in a test or resourcepopulation. Then a gene that is closely associ-ated with a particular ETL can be isolated basedon its location in the map, or by a combinationof candidate and map-based techniques. A fun-damental tool required for the latter approach(and also useful for the candidate-gene ap-proach) is a high quality, integrated genomemap. High quality means densely spaced mark-ers that are useful in many populations; inte-grated implies that the poultry genetic, cytoge-netic, and physical maps are also correlated withgenome maps of other animal species. The iden-tification of conserved syntenic groups betweenpoultry genomes and those of humans and micewill facilitate the use of genetic and physiologicaldata from these more highly studied species.

A critical point with regard to genome re-search is the availability and suitability of poul-try genetic resources. While commercial breedersmaintain populations that presumably contain awide variety of alleles, these populations are of-ten not useful for research purposes if thesebreeders are unwilling to make populations (ordata) available because of trade secrecy (propri-etary) concerns, or the commercial lines are sooutbred and therefore so heterogeneous at eco-nomic trait loci that it becomes impossible tomeasure specific effects at one or a reasonablenumber of candidate loci. Therefore, geneticmapping experiments require the availability ofdiverse populations of well-characterized, in-bred, or partially inbred (or carefully random-bred) germplasm. Public access to genetic stocksand information is essential for the advancementof science. Thus, the growing number of patentrequests on such material by private corpora-tions is a concern, as markers identified by re-searchers supported by funding from private or-ganizations, using stocks that are privatelyowned (as with recent gene mapping studies bythe Groenen group (CROOIJMANS et al. 1996) maybe restricted in their availability.

Developmental geneticsDevelopmental geneticsDevelopmental geneticsDevelopmental geneticsDevelopmental geneticsChicken developmental mutations played a sig-nificant role in the early days of the science ofgenetics. Bateson’s work at the turn of the cen-tury showed that different comb shapes in thechicken followed Mendelian inheritance pat-terns. This proved that Mendel’s principles,though discovered in plants, also applied to ani-mals (BATESON 1902; BATESON and PUNNETT1906, 1908). Today, there is increasing interestin the molecular genetic analysis of such devel-opmental or pattern mutants to determine thebasis of the developmental defects that producethese changes. The defined physical defects as-sociated with the avian developmental muta-tions, exemplified in Figures 11 and 12 and Box17, contrast strongly with many of the synthetic(knock-out) mutations in the mouse, whose phe-notype is often simply characterized by the tim-ing of early embryonic death.

Currently, developmental genetics is one ofthe most rapidly advancing areas in the biologi-

Figure 11. Wingless-2 chicken embryo from UCDWingless-2 X 331 (Photo courtesy of J.M. Pisenti,University of California–Davis).

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cal sciences (RIDDLE et al. 1993; FALLON et al.1994; MORGAN 1997; SANDERS and WRIDE 1997;CHUONG 1998). Questions that have intriguedand puzzled embryologists and geneticists fordecades can finally be addressed. The award ofthe 1995 Nobel Prize in Medicine/Physiology toE.B. Lewis, C. Nusslein-Volhard, and E.F.Wieschaus reflects the worldwide recognition ofthe fundamental importance of research in de-velopmental genetics. Their work in identifyingfruit fly (Drosophila melanogaster) developmentalmutants and determining the molecular basisfor some of these mutations provided key infor-mation for identifying a whole class of pattern-altering genes (the homeobox-containing genes)found in such diverse organisms as the fruit fly,nematode, echinoderm, fish, chicken, mouse,and human.

Avian developmental geneticists interested inpattern formation have studied an array of mor-phological mutants, most of which are recogniz-able early in development. These mutants typi-cally alter the beak, head, trunk (including thetail), limbs, and integument (skin, feathers, andscales) (ABBOTT 1967; ROMANOFF 1972; LANDAUER1967, 1973; SAWYER and GOETINCK 1988). Inmany instances, as exemplified by the wingless-2 mutation in Figure 11, the mutations have apleiotropic effect (alter more than one embryonicstructure), with combinations of defects involv-ing the limbs, trunk, beak, feathers, heart, kid-neys, and so on. Another tantalizing finding,much investigated by LANDAUER (1967, 1973),reflected the close similarity of many known mu-tant syndromes with those produced by treat-ments with a variety of chemicals or hormones,or caused by nutritional deficiencies or excesses

(the so-called phenocopy effect). A field withgreat potential then involved studies of the basisof phenocopies and of means of overcomingthem. Later, studies with pesticides provokedadditional interest, as many of these environ-mental toxins could also produce phenocopies ofdifferent mutant syndromes.

Currently, the genes affecting patterning inthe avian embryo are increasingly important indevelopmental studies (RIDDLE et al. 1993;

tremely important to bear in mind thatthe discovery that limbless encodes agene playing a critical role in limb de-velopment could not have been madebefore the molecular markers for dor-sal and ventral cell identities becameavailable. Other mutations in avianstocks may be equally valuable, buttheir importance cannot be assessedat present because of limitations in ourknowledge about limb development orbecause the appropriate markers forassaying gene function are not avail-able. If we discard stocks because wedo not appreciate their value, we maybe discarding the key to a more pro-found understanding of a particular de-velopmental or disease process.

LIMBLESS IS AN AUTOSOMAL recessive mu-tation that causes the complete ab-sence of limbs in homozygotes; het-erozygotes have normal limbs (PRAHLAD

et al. 1979). The earliest stages of limbdevelopment appear to occur normallyin the mutant homozygotes, butshortly after the limb bud forms, itceases to develop and soon regresses.Two recent studies of this mutation(GRIESHAMMER et al. 1996 and NORAMLY

et al. 1996) have provided evidencethat although the phenotype of the mu-tant is a lack of limb outgrowth, theprimary defect in limbless embryos isa lack of normal dorsal-ventral pattern-ing at a very early stage of limb devel-opment. This apparently results in aninability of the embryonic cells in the

Box 17. Patterns in vertebrate limb development

prospective limb-forming territory to re-spond appropriately to the normal limb-inducing signal, and as a consequencethe apical ectodermal ridge does notform. In the absence of this importantsignaling center, limb outgrowth doesnot occur. The importance of these stud-ies is that they have revealed a previouslyunknown link between patterning alongthe dorsal-ventral axis and the initiationof limb bud formation.

This breakthrough in understandingthe fundamental mechanism of limb de-velopment in vertebrates would not havebeen made were it not for the availabil-ity of chicken stocks carrying the limb-less mutation, because similar mutationsin other species such as mice and hu-mans are not known. However, it is ex-

Figure 12. Eudiplopodia foot from UCD EudiplopodiaX 003 (Photo courtesy of J.M. Pisenti, University ofCalifornia–Davis).

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RODRIGUEZ et al. 1996). Specifically, these muta-tions can be used to study signaling, biologicalclocks, interactions between different develop-mental controls, and the relationship of the de-fective patterns to similar syndromes producedby chemical or other experimental means. Im-portantly, these mutants can be used to investi-gate directly various hypothetical models of geneaction developed by other means.

Limb developmentFor more than 50 years, vertebrate limb develop-ment has been a major focus of developmentalbiology (SAUNDERS 1948; ABBOTT 1967; SAWYERand GOETINCK 1988; FALLON et al. 1993; LAUFERet al. 1997). Between 1972 and 1998, advancesin this field have been highlighted at six Interna-tional Conferences on Limb Development andRegeneration. The limb bud has been of interestbecause it provides an experimental paradigmfor exploring fundamental mechanisms of verte-brate tissue outgrowth and patterning (SAUNDERS1972; EDE et al. 1977; HINCHLIFFE and JOHNSON1980; CONNELLY et al. 1981), and for identifyingthe molecules that mediate these processes. Italso provides a model for studying the mecha-nism of embryonic induction (CONNELLY et al.1981). Most of what is presently known aboutvertebrate limb development has come fromstudies performed in avian species, primarily thechicken, but it is now clear that the mechanismof limb development has been highly conservedthroughout evolution, and what has beenlearned about limb development in the chick isequally informative about limb formation inmammals (FALLON et al. 1993). Indeed, it appearsthat the early steps in limb development and themolecules that perform them are virtually identi-cal in all vertebrate species (MORGAN 1997), andit is only at relatively late stages that species-specific differences become significant. A num-ber of researchers have used developmental mu-tations to test hypothetical control mechanismsgoverning limb pattern at the tissue level. Morerecently, still other researchers have returned to

these mutations to study perturbations in theexpression of developmentally regulated mol-ecules that could explain the pattern abnor-malities characteristic of such mutations(GRIESCHAMMER et al. 1996; NORAMLY et al.1996; RODRIGUEZ et al. 1996; LAUFER et al.1997). Thus, there is still much to be gainedfrom the analysis of the mutations that haveoccurred in avian stocks and which have beenmaintained in university collections. Two par-ticularly useful chicken mutations are thelimbless mutation, which has recently beenused to gain a remarkable insight into the fun-damental mechanism of limb initiation (Box17), and the eudiplopodia mutation (Figure 12),which has showcased the critical nature oftiming and location of signaling molcules inearly limb bud on the final patterning of thelimb (LAUFER et al. 1997).

Feather and scale development

The morphogenesis of cutaneous appendages,such as feathers, scales, and hair, depends ona series of reciprocal interactions between theepithelial and mesenchymal layers of the em-bryonic skin (SENGEL 1976). Such epithelialmesenchymal interactions also take place inthe development of the limb (RIDDLE et al.1993; FALLON et al. 1994; NISWANDER et al.1994), tooth (VAINIO et al. 1993), kidney (PAT-TERSON and DRESSLER 1994), lung (PETERS et al.1994), and mammary gland (CUNHA 1994). Evi-dence is accumulating which indicates that theepithelial-mesenchymal signaling that occursduring the early morphogenesis of these vari-ous organs relies on common molecular mech-anisms (Box 18 and CHUONG 1998). Thus, in-formation gained on the mechanisms for anyone of these organs systems is very likely to beapplicable to our understanding of patterningevents in other systems. A future challenge isto understand how different embryonic struc-tures acquire their identity while relying oncommon signaling mechanisms.

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opment of the feathers and scales (SAW-YER and ABBOTT 1972; SAWYER et al. 1974;SAWYER 1979). These classical studies inthe genetics, histology, and tissue andcell biology of this mutation laid essen-tial groundwork for current molecularstudies that were designed to exploremolecular control mechanisms governingfeather and scale development.

In a recent study, the role of fibro-blast growth factor (FGF) signaling wasexamined in the epithelial-mesenchymalinteractions during the initiation offeather germ formation in geneticallynormal and scaleless embryos. A spatiallyand temporally restricted pattern of tran-scription was seen for the genes that en-code FGF-2 and FGFR-1 in developingfeather germs of the normal chick em-bryo. FGF-2 expression is restricted tothe early feather epidermal placodes,whereas, FGFR-1 expression is limited tothe dermal condensations underlying theplacodes. Transcription of these genescould not be de-tected in skins ofscaleless embryosthat failed to de-velop feathers as aresult of their ecto-dermal defect. How-ever, when scalelessskin was treatedwith FGF-2, featherswould form at thesite of application ofthe growth factor,and the inducedfeathers expressFGFR-1 in their der-mal condensations .However, the re-sponse was re-stricted to a narrowwindow of time,strongest aroundday seven and eight,and gone by day 11(SONG and SAWYER

Box 18. Patterns in skin development

THE SCALELESS MUTATION has contributedsignificantly to our understanding ofboth the cellular and molecular mecha-nisms that control the formation of cu-taneous appendages, particularly thefeathers and scales in birds. Chickenshomozygous for the scaleless mutation(Figure 13), an autosomal recessivetrait, lack most feathers and scales(ABBOTT and ASMUNDSON 1957) as wellas the scleral ossicles (the precursorsto the bony eye ring) (PALMOSKI andGOETINCK 1970). These defects havemade this mutant an ideal model sys-tem for studying the early developmentof cutaneous appendages, and, by ex-trapolation, other organ systems de-pending on epithelial-mesenchymal in-teractions. In some of the earliest stud-ies in which normal and scaleless ecto-derm and mesoderm were recombined,researchers showed that the scalelessmutation specifically affects the ecto-dermal epithelium (GOETINCK andABBOTT 1963; SENGEL and ABBOTT 1963).This ectodermal defect prevents theformation of the ectodermal placodesin the skin of scaleless embryos(GOETINCK and SEKELLICK 1972), thus in-terrupting the normal sequence of tis-sue interactions. Despite the ectoder-mal defect, the mutant mesenchymeis fully capable of participating infeather and scale development whenrecombined with genetically normal ec-toderm (GOETINCK and ABBOTT 1963;SENGEL and ABBOTT 1963; SONG and SAW-YER 1996).

Interestingly, the mutant pheno-type can be greatly modified throughselection for increased or decreasednumber of feathers. The accumulatedmodifier genes for increased feather-ing alter the expression of the scale-less gene by altering the response ofthe mesodermal signals (BROTMAN

1977a,b). A number of studies also ad-dressed the cellular and subcellular(histological) differences in the devel-

1996). Based on these observations,FGF-2 was hypothesized to be an ac-tive molecular component of the earlysignaling between the dermal mesen-chyme and the overlying epitheliumduring feather morphogenesis. The ob-servation that FGF-2 can rescue the mu-tant phenotype of scaleless embryossuggests that FGF-2 either is, or isdownstream from, the signal that theectoderm of the scaleless mutant failsto generate.

The results obtained with the scale-less skins indicate that this mutant isan excellent system to elucidate sig-naling pathways by FGF and other sig-naling molecules in the morphogenesisof cutaneous appendages. Mechanismsidentified from this system may be ap-plicable to the understanding of devel-opmental mechanisms in other organsin which epithelial-mesenchymal inter-actions play a role (PETERS et al. 1994).

Figure 13. Hens from UCD Scaleless-Low (Photocourtesy of J. Clark, University of California–Davis).

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ConserConserConserConserConserving avian genetic resourcesving avian genetic resourcesving avian genetic resourcesving avian genetic resourcesving avian genetic resourcesCONSERVATION OF AVIAN WILDLIFE species receivesglobal attention because of the large numbers ofspecies presently in decline, and because birdsprovide a focus for recreational activities through-out the world. Less known are the needs for con-servation of progenitor, landrace, and feralpopulations of domesticated birds. Even lesswell publicized is the critical situation of many ofthe specialized stocks used in research, as dem-onstrated in Chapter 5.

Safeguarding existing biodiversity is a com-mon theme in the conservation of all types ofavian genetic resources, but the strategies usedto meet this goal can be quite different, depend-ing on the speicies. Wild species are generallyconserved through natural habitat protectionand take regulations (i.e., hunting or harvestinglimits for a given species), with captive breedingused in only the most severe cases of speciesdecline. Measuring the genetic diversity in wildspecies is being addressed in a relatively fewcases with DNA markers. Such data contributeto the development of rational schemes for con-serving genetic diversity. Wild progenitors of do-mesticated birds, such as the Red Jungle Fowlare treated similarly, but wild turkey conserva-tion, as another example, is enhanced by re-stocking and translocation of individuals. Thisactivity is motivated by the need for conservationof the species and its diversity, but the fact thatturkey is a recreational game bird is perhaps astronger motivation for conservation.

The goal of genetic resource conservation isthe maintenance of genetic integrity of a speciesor populations within a species. For populationsin nature, natural evolutionary processes remainactive, thus changes in gene frequency, popula-tion sizes, and geographic distribution are to beexpected. Human interventions impact speciescomposition through disruption of habitat andharvesting. In such cases, proactive conserva-

tion strategies must be applied. The situation issimilar for domesticated breeds of birds, butthere is more concern for conservation of specificcombinations of traits and genes, with minimalchanges in gene frequency. Even more restrictiveis conservation of genetic stocks where specificgenes are targeted. For these genetic entities, nochange in gene or genotype frequency can beaccepted. Thus, we recognize a distinction be-tween conservation and preservation. The formerallows utilization with minimal genetic changesover time caused by human activity and the lat-ter requires that no genetic changes occur, ex-emplified by nonliving materials, quiescent (e.g.,cryopreserved) specimens of gametes and em-bryos, or closely bred living genetic stocks.

Assays for DNA polymorphisms can be donewith nonliving biological materials, providing theopportunity for assessment of genetic diversityin historical times from museum specimens orpreserved tissues, including blood serum. Thisemphasizes the value of preserved biological ma-terials as a component of a conservation strategyfor birds. Cloned genomic or cDNA libraries canbe preserved indefinitely at low temperatures.This includes DNA clones used for preparingmolecular linkage maps and for discovery offunctional genes. Databases of DNA nucleotidesequences of genes or expressed sequence tagsare extremely rich sources of information be-cause sequences from many species are avail-able for screening and comparisons with un-known sequences.

This report focuses on the conservation ofgenetic stocks, as defined earlier. However, aholistic view of avian conservation is appropriatebecause the methods used for preserving geneticstocks and breeding populations, especially se-men cryopreservation, can be adopted or modi-fied for other species, including threatened orendangered wild species. Conservation methods

4

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need to be adopted which meet the specific re-quirements of the species or the trait (genes) be-ing conserved. An avian genetic resource conser-vation system includes several components (ac-quisition, maintenance, utilization, distribution,documentation, and health and quarantine)which can provide broader service than the con-servation and distribution of genetic stocks. Thisis elaborated in Chapter 6.

Methods of conserMethods of conserMethods of conserMethods of conserMethods of conservation ofvation ofvation ofvation ofvation ofgenetic stocksgenetic stocksgenetic stocksgenetic stocksgenetic stocks

Avian genetic materials may be preserved in theform of live birds, as cryopreserved semen anddispersed embryonic or primordial germ linecells, as preserved tissues from which DNA canbe isolated, as cloned genomic or cDNA, or asnucleotide sequence data. Of these methods,living birds and cryopreservation methods per-mit recovery of animals, while preserving DNAgives the potential of recovering selected genesor defined segments of chromosomes. However,it should be remembered that at best each DNAsequence represents only a very small portion ofa total genotype and that, while samples of totalDNA from a population may be preserved, sort-ing out the genes useful for a particular purposemay be a difficult task, particularly when geneinteractions must be taken into consideration.Cryopreservation of semen and embryonic cellsis gaining acceptance as a means of preservingendangered genetic resources, including wildspecies, rare breeds, and mutant or specialtylines used in research.

Ultimately, the preferred conservation methoddepends upon 1) the reliability of the method toyield the desired number of live animals, 2) thecharacteristics of the stock, and 3) the availabil-ity of technical support, instrumentation, andstorage facilities. Ideally, more than one methodof conservation should be used, or there shouldbe a backup site to reduce the risk of loss due tofire, equipment failure, and human errors. Beloware outlined some features of live-bird conserva-tion and three methods of cryopreservation ofsemen or embryonic cells from which live birdsmay be recovered. Semen cryopreservation hasthe longest history, although not a highly effi-cient recovery rate. Dispersed blastodisc andprimordial germ cell cryopreservation are prom-ising emerging technologies that will requirefurther research to improve recovery rates tomore acceptable levels.

Live animalsLive animalsLive animalsLive animalsLive animalsThe maintenance and periodic reproduction oflive animals is the most dependable alternativeavailable for preserving a genetic stock. This isparticularly true for the inbred and long-termselected stocks, with their integrated collectionsof quantitative trait alleles. This is also the mostreasonable way to maintain stocks that are oftenused in research. However, due to financial con-straints, most researchers cannot justify the livemaintenance of stocks that are used only spo-radically, and recent events have shown (Chap-ter 5) that many unique lines are terminated forthis reason.

The average number of birds required tomaintain a given genetic stock depends upon itscharacteristics. Randombred and selected stocksrequire a larger base population (often severalhundred birds) than those carrying single-genemutations. The latter may be realistically contin-ued with fewer than ten mutant-carrier birds if arobust outbred stock is used as one parent lineeach generation. Other factors that affect thenumber and distribution of birds needed tomaintain a viable population include bird live-ability, amount of inbreeding depression, en-demic diseases, and the risk of natural or man-made disasters, which can literally wipe out ge-netic stocks in a very short period of time.

Maintaining stocks as live animals is very ex-pensive, especially the selected lines and ran-dombred populations. Even temporary shifts ininterests of the curator or sponsoring institutioncan quickly orphan such genetic stocks, oftenleading to stock elimination and extinction ofvaluable genes or gene combinations. These re-alities are documented in Chapter 5, emphasiz-ing the need for backup sites or a permanentlydedicated conservation program.

Semen crSemen crSemen crSemen crSemen cryopreseryopreseryopreseryopreseryopreservationvationvationvationvationThe technology for semen cryopreservation(BAKST 1990; BUSS 1993; HAMMERSTEDT 1995)has been successfully adapted for the preserva-tion and propagation of chickens and certainferal and endangered species (GEE 1995). Manydifferent cryopreservation protocols have beenevaluated (SEXTON 1979, LAKE 1986, HAMMER-STEDT and GRAHAM 1992, BUSS 1993, HAMMER-STEDT 1995). According to HAMMERSTEDT (1995)semen cryopreservation is the most cost-effectivegermplasm conservation strategy. Hammerstedtoutlined the benefits of semen cryopreservationfor the commercial poultry industry, but thispreservation technique has not yet been widely

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adopted. An important feature of semen cryo-preservation for unique or endangered poultrylines is that semen can be collected frequently(twice weekly) from a small number of males foruse when females become available or in the fu-ture establishment of founder flocks. Other ben-efits of semen cryopreservation are:

• Frozen semen is not easily affected bydiseases or natural disasters.

• Semen cryopreservation from many birdsof several lines can be accomplished at arelatively low cost

• The expense of maintaining a flock isreduced or eliminated.

• Frozen semen is more easily transportednationally or internationally than livebirds or fertile eggs (HAMMERSTEDT 1995).

There are obvious limitations to the routineuse of semen cryopreservation. For some speciesand genetic stocks the method may not be reli-able or yet suitably developed. Another consider-ation is that only the male genome is preserved,which means that the complex genome of highlyinbred or selected stocks could not be preservedintact if only maintained as frozen semen, be-cause, of course, none of the corresponding in-bred or selected females would be available forinsemination with the cryopreserved material.Thus, while semen cryopreservation permits theincorporation of preserved gene complexes intodiploid animals, these gene complexes would beheterozygous in the progeny and subject to ge-netic recombination in subsequent intermatingsof progeny. On the positive side, semen cryo-preservation may prove to be the most appropri-ate method for preserving single-gene mutantstocks, particularly if the genetic background isnot a great concern in the modification of themutant gene expression.

Another problem with cryopreserved semen isthat the quality of thawed semen is currentlyquite low. This is reflected in fertility levels rarelyexceeding 70%, and more typically less than50%, due to sperm destruction and deformitycaused by semen cryopreservation and recoveryprocedures. One researcher estimated that cryo-preserved semen has less than 2% of the fecun-dity of fresh semen (WISHART 1985). This is inspite of the fact that most of the successfulcryoprotectants, diluents, and freezing tech-niques were developed with semen from highlyproductive commercial broiler chicken stocks.

Unfortunately, the semen from inbred, specialty,and mutant-carrier stocks most at risk oftendoes not respond well to cryopreservation proce-dures that worked reasonably well with commer-cial stocks. Frozen-thawed semen from theseless-robust stocks often exhibit extremely highor complete infertility associated with poor motil-ity and high numbers of freeze-damaged or killedspermatozoa. Obviously, further research isneeded, not only in the development of improvedsemen cryopreservation procedures (improveddiluents, cryoprotectants, and other supple-ments, and optimizing cooling and thawingrates), but also addressing the significance ofspecies variation, male-line variation within aspecies, and individual male variation within agiven strain. However, despite the potential util-ity of this procedure in preserving the germ-plasm of rare or endangered stocks or species,only a few researchers in Japan, England, andthe US are currently conducting research on se-men cryopreservation.

Blastodisc crBlastodisc crBlastodisc crBlastodisc crBlastodisc cryopreseryopreseryopreseryopreseryopreservationvationvationvationvationCryopreservation of the cells from an avian blas-todisc (Figure 14, the embryonic cells in an un-incubated fertile egg) is a new technique thatcan be used to preserve the intact genome ofpoultry genetic stocks (REEDY et al. 1995, Box19). While the methods are still being perfected,frozen-thawed blastodisc cells have been suc-cessfully integrated into the blastodiscs of hostchicken eggs, and developed into reproductivelycompetent adults. The long-term viability ofcryopreserved blastodisc cells is not yet known,although cryopreservation studies with semen

Figure 14. Stage X blastodisc (at time of oviposition)(Photo courtesy of M.E. Delany, University of Califor-nia–Davis).

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and other cell-types suggests that properly cryo-protected and frozen specimens will remain vi-able indefinitely if kept in liquid nitrogen.

The large size and complex structure of theavian embryo (already composed of 40,000 to50,000 cells when the egg is laid) do not permitthe application of techniques that have been de-veloped for cryopreservation of mammalian em-bryos. A different strategy is required which usesdispersed cells isolated from the relatively undif-ferentiated embryonic cells present in the newlylaid fertile egg (Box 19). Recent experimentshave demonstrated that frozen-thawed blasto-dermal cells from purebred chickens of theBarred Plymouth Rock breed can successfullyintegrate into an irradiated White Leghorn hostblastoderm to form embryos that are somatic

and germline chimeras. These chimeric chickenshatch and mature normally, and, when inter-bred, produce some offspring that are purebredBarred Rock (KINO et al. 1997). While frozen-thawed blastodermal cells form germline chime-ras less frequently than fresh blastodermal cells,a sufficient number of “reconstituted” birds oflines maintained as cryopreserved blastodermalcells can be obtained to use the technology tostore valuable genetic material (KINO et al. 1997).Thus, this technique should eventually providean inexpensive long-term solution for preservingthe intact genome of stocks (particularly inbredand selected stocks) that are considered to bevaluable genetic resources but are not currentlyrequired in an active research program (Box 20).

A NEW TECHNIQUE HAS BEEN developed atthe University of Guelph (Ontario,Canada) that can be used to reclaimboth freshly dissociated and cryopre-served chicken embryonic cells (Figure15, REEDY et al. 1995). Briefly, cells fromthe stage X embryo (EYAL-GILADI andKOCHAU 1976) can be removed, dis-persed, and injected into recipient blas-todiscs that are at the same stage ofdevelopment. The resulting chimeras(mixture of host and donor cells), whichcan be either male or female, producedonor-derived gametes at rates of upto 100% (PETITTE et al. 1990; 1993;CARSIENCE et al. 1993; THORAVAL et al.1994; KAGAMI et al. 1995). Matings ofmale and female chimeras that eachproduce donor-derived gametes yieldoffspring with the complete donorgenotype at a rate that is the productof the frequency of the production ofdonor-derived gametes by eachof thechimeric parents (KINO et al. 1997).

Box 19. Mosaic chickens and genetic stock preservation

Figure 15. Diagram of avian blastodisc cryopreservation and recovery(from REEDY et al. 1995).

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Research aimed at improvingthe rate of germline transmis-sion by cryopreserved blasto-dermal cells could increase theeffectiveness of this technologyand reduce the number of blas-toderms that are required toensure that the genetic diver-sity within a stock is main-tained in the reconstitutedpopulation. While this would bea welcome improvement in thetechnology, it should be empha-sized that the current technol-ogy can be used to conserve genetic resourcesand that it is the only fiscally realistic way tomaintain the genome of stocks in an unadulter-ated state at the present time.

In the future, the range of genetic diversitymay increase as transgenic stocks are produced.Many of these stocks will be developed for spe-cific experimental uses that have short-termutility in research. It would be prudent to storethese stocks as frozen blastodermal cells ratherthan discard the time and effort expended intheir production.

PPPPPrimordial germ cellrimordial germ cellrimordial germ cellrimordial germ cellrimordial germ cellcrcrcrcrcryopreseryopreseryopreseryopreseryopreservationvationvationvationvation

Cryopreservation of primordial germ cells (PGCs)isolated from the blood of early-stage chick em-bryos is another way of preserving the intact ge-nome of both male and female birds (NAITO et al.1994a). The PGCs are the precursors of theadult gametes (ova and spermatozoa) and form adistinct population of readily distinguishablecells even before they move from the extra-

embryonic germinal crescent into the blood ves-sels. After a short period of time being carriedthrough embryonic and extra-embryonic circula-tory systems, these cells will migrate out of theblood vessels and selectively colonize the gonad-al ridges. While in the migratory phase (mostlybetween HAMBURGER and HAMILTON (1951) stages13 and 15), the PGCs can be removed from theembryo in blood samples. Primordial germ cellscan be partially separated from the blood cellsby density gradient centrifugation and can thenbe cultured (CHANG et al. 1995; KUWANA et al.1996), cryopreserved (NAITO et al. 1994a), or im-mediately injected into a host embryo (TAJIMA etal. 1993; NAITO et al. 1994b). One of the benefitsof this method of germplasm preservation is thata highly enriched population of germline cells ofapproximately 60% PGCs (NAITO et al. 1994b)can be introduced into a host embryo. Thus,even when frozen-thawed PGCs were injected,up to 25% of the gametes were shown to be fromthe donor germline (NAITO et al. 1994a). Thisgermline recovery rate is substantially betterthan that obtained from frozen-thawed blasto-disc cells (KINO et al. 1997).

Box 20. Orphaned CFAR stocks preserved as frozenembryonic cells

THE FIRST LARGE-SCALE DEPOSITION of fro-zen blastodermal cells was completedin 1998, preserving over 30 of thestocks once kept at the Centre for FoodAnimal Research (CFAR). This had beenthe flagship research center for poul-try research for Agriculture and Agri-Foods Canada. The cryopreservedCFAR stocks were among those thatcould not be placed with other institu-tions before April 1, 1997. Blastoder-

mal cells were harvested and frozen atthe Avian Physiology and GeneticsLaboratory at the University of Guelph,(Ontario, Canada). If these stocks arerequired in the future, trained techni-cians can thaw the cells and makegermline chimeras (see Box 19), whichcan then be mated to obtain pure “re-constituted” birds from the originalCFAR lines.

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Genetic stocks: Current resources and recent lossesGenetic stocks: Current resources and recent lossesGenetic stocks: Current resources and recent lossesGenetic stocks: Current resources and recent lossesGenetic stocks: Current resources and recent losses

5

ALMOST ALL OF THE GENETIC STOCKS of chicken,turkey, Japanese quail, and other domesticatedbirds that have been used in research in the USand Canada were developed and conserved bypublic sector universities and federal govern-ment research organizations in these two coun-tries. Many in the research community havebeen aware of the gradual attrition, and occa-sional dramatic losses, of these specializedstocks over the years. Some important geneticresources were discarded before relocation planscould be developed or because a new stock cura-tor could not be located. Current threats to addi-tional abandonment of existing genetic stocksgave impetus for a study of the true situation ofexisting genetic stocks, where such stocks arelocated, and their status for conservation anduse. The situation is dynamic, for not only areestablished genetic resources at risk, but newgenetic stocks are also being developed in cur-rent research programs that will in turn requirepreservation or conservation in the years tocome. SOMES (1988) provided a benchmark withhis comprehensive, but not exhaustive, sum-mary of the state of genetic stocks up to 1988.Hence the Task Force undertook an assessmentof the current situation with the goal of produc-ing a complete database inventory of extant ge-netic stocks, along with a summary of the ge-netic stocks that had been discontinued, aban-doned, or lost since 1984. This information con-stitutes the basis for the proposal of a comprehen-sive strategy geared towards the conservation ofavian genetic stocks important to basic biologicalresearch and education. (See Chapter 6).

SurSurSurSurSurvey methodsvey methodsvey methodsvey methodsvey methodsThe study was conducted by Jacqueline Pisentiduring 1995–97 at the UC Genetic ResourcesConservation Program, with additional financial

support from the National Science Foundationand the USDA Agricultural Research Service. In1998 each curator was contacted again to pro-duce an updated report. A survey instrument(Appendix 1) was sent to all of the curators men-tioned in SOMES (1988) and the participants inthe CSREES Regional Research Project NE-60 ongenetic basis for resistance and immunity toavian diseases. Individuals listed in the PoultryScience Association 1995–96 membership direc-tory who indicated a focus on genetics or immu-nology, and individuals on the 1997 Poultry Sci-ence Resource List prepared by the Poultry Sci-ence Association and R.D. Reynells (USDA/CSREES) were also contacted (usually via emailor telephone) and queried. In all, survey formswere sent to 37 curators (11 in Canada and 26in the US) at 27 institutions listed in the SOMES(1988) registry and 72 additional individualswere contacted by phone or email. All curatorswere asked to report on the status of stocks thathad been included in the 1988 registry (i.e.,which of these stocks had been retained, trans-ferred, or eliminated), and to list any new stocksacquired since1988. They were also asked to in-dicate the prognosis for long-term security ofeach stock. Finally, respondents were asked tosend the names of any poultry genetic stock cu-rators they were aware of, either at their institu-tions or elsewhere, and these individuals notalready contacted were also asked to respond tothe survey.

RRRRResults from the suresults from the suresults from the suresults from the suresults from the surveyveyveyveyveyAccording to the survey, 361 genetic stocks arecurrently maintained (Table 1 and Appendix 2).Of the 36 curators at 27 institutions listed in the1988 registry (SOMES 1988), 31 of them (or theirsuccessors) responded; four no longer main-tained stocks, and 27 still had at least some of

32

the stocks listed in the 1988 registry. Of the 72other persons queried, 44 responded and 13 re-ported that they curated poultry genetic stocks.

By species, there are 268 chicken stocks(with 38 existing only as cryopreserved material),65 Japanese quail, 20 turkey, six waterfowl, andtwo gamebird. Table 1 shows the categorization

of the existing genetic stocks. The detailed data-base in Appendix 2, Table 2.1, lists the charac-teristics of every genetic stock that was identifiedin the survey. These were grouped into ten maincategories. The largest category is single-genemutants, with 107 stocks that display color var-iations or defects in developmental, immunologi-

Table 1. Summary by category of existing poultry genetic stocks (live and cryopreserved).Japanese Waterfowl,

Category Chicken quail Turkey Gamebird TotalNumber of stocks

Bloodtype-Gene pool 6 0 0 2 8Bloodtype-Major Histocompatibility Complex (MHC) 13 0 0 0 13Bloodtype-MHC-Inbred 51 0 0 0 51

subtotal 70 0 0 2 72

Chromosomal variant 5 0 0 0 5

Endogenous virus 1 0 0 0 1Endogenous virus-Inbred 4 0 0 0 4

subtotal 5 0 0 0 5

Inbred 27 1 0 0 28

Mutant-Color, eggshell 0 2 0 0 2Mutant-Color, feather 1 12 0 0 13Mutant-Developmental defect-Eye 5 0 1 0 6Mutant-Developmental defect-Face/limb 28 0 0 0 28Mutant-Developmental defect-Skin/feather 8 6 0 0 14Mutant-Developmental defect-Spine/tail 3 0 0 0 3Mutant-Gene pool 7 1 0 0 8Mutant-Immunological defect 5 0 0 0 5Mutant-Neurological defect 3 0 0 0 3Mutant-Physiological defect 12 0 0 0 12Mutant-Reproductive defect 3 0 1 0 4Mutant-Uncategorized 8 1 0 0 9

subtotal 83 22 2 0 107

Pure breed 12 0 5 6 23

Randombred 18 14 8 0 40

Selected-Behavioral trait 2 1 0 0 3Selected-Egg trait 11 0 2 0 13Selected-Growth trait 14 18 3 0 35Selected-Immune trait 11 1 0 0 12Selected-Physiological trait 2 8 0 0 10

subtotal 40 28 5 0 73

Transgenic 2 0 0 0 2

Uncategorized 6 0 0 0 6

Total 268 65 20 8 361

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cal, neurological, physiological, or reproductivecharacteristics. The selected stocks category,with 73 stocks, are products of long-term selec-tive breeding and are characterized by above- orbelow-normal performance in defined behavioral,growth, egg production, immunological, orphysiological traits. The next largest categorywith 72 stocks consists of the bloodtype vari-ants, representing most of the known forms ofchicken erythrocyte and leukocyte alloantigens.The inbred category, with 28 stocks, consists ofstocks approaching genetic homogeneity as aresult of crossing full sibs or other close relativesfor many generations. In addition to the inbredcategory, highly inbred stocks can be found inthe bloodtype-MHC-inbred and endogenous vi-rus-inbred categories. A few highly inbred stocksare also found among the mutant-developmentaldefect-face/limb category. The randombred cat-egory with 40 stocks and the pure breed cat-egory with 23 stocks are important as control orreference populations and represent gene poolsfrom which many of the selected and mutantstocks were derived. The uncategorized groupwith six stocks are those stocks for which nodescription was available.

Forty-one researchers at 27 institutions (24in the US and 3 in Canada) maintain 323 geneticstocks as living birds (Table 2). Six of these insti-tutions maintain 20 or more stocks (Universityof Arkansas, University of British Columbia–Vancouver, University of Connecticut–Storrs,University of California–Davis, University of Wis-consin–Madison, USDA/ADOL East Lansing,Michigan). Six maintained between ten and 19,four had six to nine, and 11 had five or fewergenetic stocks. Two institutions in the US andfour in Canada no longer keep such stocks.

In addition to the living bird collections, 87genetic stocks are maintained under semen orblastodisc cryopreservation at five institutions.Such frozen reserves are kept at the Universityof Guelph (Guelph, Ontario) (33), the Universityof California–Davis (31), the USDA Avian Diseaseand Oncology Laboratory at East Lansing, Michi-gan (17), Pennsylvania State University (4), andthe University of Wisconsin–Madison (2). Five ofthe cryopreserved stocks at University of Califor-nia–Davis are no longer maintained as live birds.The 33 cryopreserved stocks at the University ofGuelph are from the Canadian Food Animal Re-search collection and also are no longer main-tained as live birds. Thus, 51 stocks maintainedas live birds are also backed up by cryopreserva-tion, while 38 stocks now exist only under cryo-preservation.

At the time this report went to press, approxi-mately one-third of the stocks kept as live birdswere considered by their curators to be at highrisk of elimination within the next few years(30% of the chicken, 35% of the turkey, and 49%of the Japanese quail stocks). Just over 40%were considered to be in a secure situation,including 42% of the chicken, 45% of the turkey,and 37% of the Japanese quail stocks. Theremainder of the stocks (about 30%) were con-sidered to be only moderately at risk.

LLLLLost stocksost stocksost stocksost stocksost stocksIn the last 15 years, some 238 avian geneticstocks were reported as lost or eliminated byresearch institutions in the US and Canada(Table 3; this also includes those few stocks re-ported as transferred to private individuals ororganizations). The actual number of lost stocksis probably much higher. The data in Table 1show the numbers and percentages of geneticstocks lost, based on the current number of ex-isting stocks and documented numbers of loststocks. Some new stocks have been developedsince the last survey, specifically in the MHC-defined endogenous virus and selected catego-ries. These recent additions inflate the totalnumbers of existing stocks and bias downwardsthe estimated percentage of stocks lost. Even so,stock losses have been substantial: close to 40%of the reported living US stocks and over 60% ofthe Canadian stocks were discontinued since1984, 42% of all stocks (Table 2). Most notablewas the discontinuation of more than 30 geneticstocks by Agriculture and Agri-Food Canada atits facility in Ottawa. This clearly reflects a dis-turbing trend in avian genetic resources conser-vation.

Of all lost stocks, 75% were chickens, 11%were turkeys, 11% were Japanese quail, and 4%were gamebirds or waterfowl. Ironically, morethan one-third of these stocks were reported lostbetween 1995 and 1998 while this survey was inprogress, including 50 chicken stocks, 19 turkeystocks, 10 Japanese quail stocks, and four of thewaterfowl stocks. The losses were heaviest (35%of all lost stocks) among the mutant and chro-mosomal variant categories, although the se-lected (25%) and the bloodtype-MHC-inbred(21%) categories were not far behind. Some ofthese lost stocks, such as the bloodtype-MHCspecialized stocks, included genes that can berecovered by breeding and selection from exist-ing genetic resources, but at a great cost ofmoney and time. The now-extinct lethal and

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Table 2. Number of poultry genetic stocks kept as live birds in institutions in the USA and Canada in1998 and the number lost since 1984.

Kept as living birds† Lost since 1984†

Curating institution C JQ T W,G Total C JQ T W,G Total % all‡

USA Number of stocksAlabama: Auburn University 10 0 0 0 10 17 0 0 0 17 63Arkansas: University of Arkansas 12 9 0 0 21 1 2 0 0 3 13Arizona: University of Arizona 0 0 0 0 0 1 0 0 0 1 100California: University of California 46 1 0 0 47 15 7 1 4G 27 36Connecticut: University of Connecticut 20 0 0 0 20 8 1 1 0 9 31Delaware: University of Delaware 1 0 0 0 1 0 0 0 0 0 0Georgia: University of Georgia 4 9 0 0 13 2 6 0 0 8 38Illinois: Northern Illinois University 7 0 0 2G 9 0 0 0 0 0 0Illinois: University of Illinois 2 0 0 0 2 0 0 0 0 0 0Indiana: Purdue University 2 1 0 0 3 6 0 0 0 6 67Iowa: Iowa State University 17 0 0 0 17 5 0 0 0 5 23Iowa: USDA National Animal Disease Center 0 0 1 0 1 0 0 0 0 0 0Louisiana: Louisiana State University 0 3 0 0 3 2 0 0 0 2 40Maryland: University of Maryland 0 1 0 0 1 0 0 0 0 0 0Massachusetts: University of Massachusetts 4 0 1 0 5 10 0 1 0 11 69Michigan: USDA Avian Disease & Oncology Lab. 37 0 0 1W 38 2 0 0 0 2 5Minnesota: University of Minnesota 0 0 0 0 0 5 0 0 0 5 100Nebraska: University of Nebraska 1 1 0 0 2 0 0 0 0 0 0New Hampshire: University of New Hampshire 14 0 0 0 14 14 0 0 0 14 50New York: Cornell University 7 0 0 1W 8 7 1 0 0 8 50North Carolina: North Carolina State University 5 0 7 2W 14 0 0 2 0 2 13Ohio: Ohio State University 2 7 6 0 15 15 0 2 0 17 53Oregon: Oregon State University 1 0 0 0 1 0 0 17 0 17 94Pennsylvania: Pennsylvania State University 6 0 0 0 6 0 0 0 0 0 0Virginia: Virginia Polytechnic Inst. & State Univ. 4 0 0 0 4 0 3 0 0 3 43Wisconsin: University of Wisconsin 19 1 3 0 23 6 0 0 0 6 21

Total in USA 221 33 18 4W 278 116 20 23 4G 163 37 2G

CanadaBritish Columbia: Univ. of British Columbia 3 32 0 0 35 0 5 0 0 5 13Ontario: University of Guelph 3 0 1 0 4 10 0 0 0 10 71Ontario: AA-FC Center for Food Animal Research 0 0 0 0 0 30 0 0 6W 36 100Quebec: McGill University 0 0 0 0 0 4 0 0 0 4 100Quebec: University of Lavale 0 0 0 0 0 1 0 0 0 1 100Quebec: Deschambault Research Station 0 0 0 0 0 4 0 3 0 7 100Saskatchewan: University of Saskatchewan 3 0 1 2W 6 11 0 1 0 12 6 7

Total in Canada 9 32 2 2W 45 60 5 4 6W 75 63

Total 230 65 20 6W 323 176 25 27 6W 238 42 2G 4G

†C = chicken, JQ = Japanese quail, T = turkeys, W = waterfowl, G = gamebird.‡“% all” indicates the proportion of total stocks at that institution that were lost since 1984.

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Table 3. Summary by category and species of poultry genetic stocks reported lost since 1984.Japanese Waterfowl,

Category Chicken quail Turkey Gamebird† TotalNo. %‡ No. %‡ No. %‡ No. %‡ No. %‡

Bloodtype-MHC-Inbred 49 28 0 0 0 0 0 0 49 21Chromosomal variant 33 19 0 0 0 0 0 0 33 19Inbred 6 3 0 0 0 0 3G 30 9 4Mutant 25 14 11 42 19 70 0 0 55 23Pure breed 18 10 0 0 3 11 4W 40 25 10Randombred 3 2 3 12 0 0 1W 10 7 4Selected 42 24 11 46 5 19 1W,1G 20 60 5

Total 176 74§ 25 11§ 27 11§ 6W,4G 4§ 238 100§

†W=waterfowl; G=gamebird.‡This column indicates the percentage of lost stocks in a category out of the total of lost stocks for that species.§This value is the percentage of total lost stocks over all species accounted for by the total lost stocks for a givenspecies.

subvital mutant stocks, along with the largenumber of discontinued translocation stocks,appear rarely, are difficult to detect, and are of-ten discovered serendipitously. A number of theviable mutations that were kept in gene pools oras specialized stocks in research collections canstill be found among hobbyist breeds or exhibi-tion stocks, but they present problems in usebecause there is a risk of transferring diseasesor they have background genotypes that may beinappropriate for use in research.

Individual stocks and whole collections thathave been lost in recent years include:

• Inbred, specialty, and historical commer-cial stocks once maintained by Agricul-ture and Agri-Food Canada in Ottawathat were dispersed or eliminated in1997, due to loss of government support.While some of these specialized stockswere sent to other institutions or privatecompanies, 33 were retained only ascryopreserved blastodisc cells kept at theUniversity of Guelph, (Ontario, Canada)(Box 20) and 30 were eliminated. How-ever, one-half of these 30 were bloodtypestocks that are still available elsewhere.

• Five stocks (long-term inbred lines andone gene pool of dominant mutations)maintained at the University of Minne-sota, were eliminated in 1996, althoughtwo (the dominant marker stock and theinbred Rhode Island Reds) had beentransferred to the University of BritishColumbia a few years earlier.

• The collection of stocks at the Universityof Massachusetts went into dispersal in1997, due to the retirement of its cura-tor. While several important mutantstocks have found new curators, includ-ing the auto-immune vitiligo (DAM)chickens and their normal controls, thefeather color and comb-type gene pooland tester stocks have been eliminated.

• At Oregon State University (Corvallis), alarge and unique collection of 17 stockscarrying feather-color and embryo-lethalturkey mutations was eliminated in1995, due to funding difficulties. A simi-lar problem contributed to the loss of an-other collection of turkey stocks in the1960s, this one held at the University ofCalifornia–Davis.

• Fires at bird-care facilities have killed offat least two turkey stocks at the Desch-ambault Research Station and one Japa-nese quail mutation at University of Cali-fornia–Davis. Five other quail mutationssalvaged from that fire at Davis (threeunique to University of California–Davis)were subsequently eliminated due tofunding difficulties.

• At the University of California–Davis, theretirement of two researchers with largecollections of research stocks has putover 40 stocks in jeopardy (Box 3); theseinclude over 20 mutations and an arrayof MHC-defined inbred lines, many ofwhich are found nowhere else.

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been eliminated or merged with other depart-ments, leaving only eight poultry science depart-ments today (PARDUE 1997; DELANY and PISENTI1998). With reorganization of these departmentsand their resources, genetic stock losses wereinevitable as a result of loss of long-term supportfunds, reallocation of funding and resources toother areas, condemnation of poultry housingfacilities without construction of adequate re-placement housing, retirement of researcherswho served as curator of stocks, shifts in re-search interests, and new research on exoticavian species as research models or companionbirds.

The number of genetic stocks reported bySOMES (1988) is nearly the same as that detectedin the present survey. However, the present sur-vey documented the loss of nearly 200 chickenstocks, 23 turkey stocks, and at least 8 Japa-nese quail stocks, so it is evident that a consid-erable number of new stocks have been pro-duced during the past decade, possibly displac-ing some of the older ones. Most curators facedwith the need to reduce or disperse inventory tryto find alternate conservators for stocks that areslated for elimination (Boxes 2, 3, 4), althoughthis is not always successful (Box 8). Informalnetworking for new curators has worked wellenough with single-gene mutation stocks whencurating researchers had funding to support anumber of these unique stocks, but this is oftenno longer an option.

Clearly, planned and unplanned stock lossesare impacting the availability of specialized ge-netic resources to researchers. Stock elimina-tions are increasingly driven by funding prob-lems, more will be lost. Particularly at risk arethe selected lines and mutant stocks, which re-quire either larger numbers or special handlingto propagate.

TTTTTrends in poultrrends in poultrrends in poultrrends in poultrrends in poultry genetic stocky genetic stocky genetic stocky genetic stocky genetic stockdevelopment and conserdevelopment and conserdevelopment and conserdevelopment and conserdevelopment and conservationvationvationvationvation

Only a few large collections of vertebrate geneticstocks have been assembled, including themouse (Mus musculus) collection that is cur-rently kept at the Jackson Laboratory in BarHarbor, Maine (Box 21) and the Japanese quailcollection at the University of British Columbia(Box 22). But it is the chicken mutant collectionsthat probably have the longest history, startingin the 1920s with Landauer and Dunn at theUniversity of Connecticut, Storrs AgriculturalExperiment Station (a collection that still exists,in part). These early researchers in vertebrateembryology used classical embryological tech-niques to study mutant gene expression, the ef-fect of background genes on mutant phenotype,and the production of environmentally inducedphenocopies (genetically normal embryos thatresemble mutants) of specific mutant syndromes(LANDAUER 1973). Ironically, it is these areas ofstudy that attract the attention of researcherstoday, just as the remaining academic avian ge-netic stock collections have become threatened.This underscores the need for preserving theseunique gene pools and identified single-genemutations for the benefit of future researchers.Similar arguments can be made for other re-search stocks, particularly inbred and selectedstrains. Both require many years and large,pedigreed populations during their development;factors that place formidable barriers to the re-generation of such stocks because of short-termand limited funding commitments.

Traditionally, poultry stocks were maintainedat the research institutions where they were de-veloped and studied. These institutions, fre-quently in states with a large or growing poultryindustry, often had poultry oravian science departments thatwere independent of the largeranimal science departments.This served to encourage re-search on the different poultryspecies and the developmentand maintenance of uniquepoultry genetic stocks. In theearly 1960s, some 45 poultryscience departments existed inthe US; since that time and inspite of the tremendous growthin economic value of the thepoultry industry and great con-sumer interest in poultry prod-ucts, most departments have

Box 21. The Jackson Laboratory model for vertebrate geneticstocks

A GOOD EXAMPLE OF A SUCCESSFUL geneticstocks conservation program is theJackson Laboratory, a non-profit,independant research institution, in BarHarbor, Maine. Along with research andtraining, its third mission is the main-tenance and distribution of a huge in-ventory of mouse genetic stocks. Theyare maintained as either live animalsor frozen embryos and are distributedeither as live animals or DNA prepara-tions. Annually, the Laboratory sup-plies about two million mice from over1,700 stocks to universities, medicalschools, and research laboratories

worldwide. In addition to providing thegenetic stocks, the Jackson Laboratoryalso maintains and serves a computer-ized database system available any-where in the world that enables fast,efficient access to a single comprehen-sive archive on the mouse. Supportedpartly by government grants and partlyby user fees, the Jackson Laboratory isa good model for long-term preserva-tion of avian genetic stocks. Furtherinformation on the Laboratory and thedatabases can be accessed from theirweb site (URL: http://www.jax.org/).

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PRIOR TO 1980, A JAPANESE quail colonywas maintained at the Department ofAnimal Science, University of British Co-lumbia (UBC) for research and teach-ing in genetics. In 1980, the goal wasset to expand this colony and manageit as a genetic resource collection, andto make it available to users outsidethe department. In 1982, the Quail Ge-netic Resource Centre was formally es-tablished with the support of the de-partment and the Natural Sciences andEngineering Research Council ofCanada (NSERC). In the last 14 years,many new mutations and strains havebeen incorporated into the collection.At present, the quail populations at theCentre harbor 27 morphological andphysiological mutations, close to 50%of all the mutations (not includingisozyme variants) ever reported in theliterature for this species. Detailed de-scriptions of the mode of inheritance,the phenotype, and possible applica-tions of these mutations can be foundin CHENG and KIMURA (1990). In addi-tion, they also maintain 12 random-bred populations and specially devel-oped strains that are useful for biologi-cal research or as breeding stocks formeat and egg production. Stocks from

Box 22. Japanese quail at the University of British Columbia

the Centre have been widely utilized forresearch, teaching, and extension.

Areas of research include genetics,physiology, developmental biology, can-cer, atherosclerosis, eye defects, neurol-ogy, animal behavior, and animal ecol-ogy. In the past three years, the Centreassisted or collaborated in more than 35research projects. Currently, there aretwo on-going research projects usingquail as a model to study atherosclero-sis and another project using quail tostudy age-related macular degeneration(AMD). Except for monkeys, the quail isthe only suitable model to study this dis-ease which is the most common causeof blindness in humans over the age of65. The Centre has also started screen-ing chicken microsatellite DNA primersobtained from Hans Cheng to detectpolymorphism in quail.

Teaching uses include courses in bi-ology, genetics, poultry management,and animal behavior at University of Brit-ish Columbia and other universities andcolleges in the province. These stockshave also been used by students fromlocal secondary schools for their seniorscience projects. The Centre staff alsoprovides consultation for researchers andteachers on managing quail as a labora-

tory animal, and consultation for localproducers and raptor rehabilitationcenters on management of the flocksand/or marketing. In the latter cases,birds from the Centre were sometimesprovided as breeding stocks. With theincrease in consumption of quail prod-ucts in metropolitan areas in Canadaand also in the southern US, there hasbeen a significant increase in requestsfor information from other parts ofCanada and from the US.

The Centre has been partially sup-ported by an infrastructure grant fromNSERC, which provided the salary forthe full-time technician. This grant cov-ered 25% of the operating costs. Be-sides capital investments, the Univer-sity provided 40% of the operatingcosts through department funds. Theremaining 35% of the operating costswere covered by income from sales ofproducts and users’ fees. The NSERCsupport was terminated in 1995, andincome generated from egg sales andresearch contracts are not sufficient tosustain the operation. An endowmentwill be sought to generate Canada$30,000 Cdn a year to keep the opera-tion going.

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An AAn AAn AAn AAn Avian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Rvian Genetic Resources System: Pesources System: Pesources System: Pesources System: Pesources System: Proposalroposalroposalroposalroposal

TTTTTask Fask Fask Fask Fask Force recommendationorce recommendationorce recommendationorce recommendationorce recommendationIT HAS BECOME ABUNDANTLY CLEAR from the analy-sis of extant genetic stock collections and fromthe information obtained from task force mem-bers and others that many avian genetic stockshave been lost and many others are at seriousrisk of being lost. Further, the prognosis formaintaining new research stocks as they are be-ing developed in current research programs isnot good. In fact, when maintenance facilities forresearch stocks are not available, there will beno incentive for investment in lines of researchthat would produce such stocks. An avian ge-netic resources management system, with strongleadership, but shared responsibility, is pro-posed as the most efficient and secure way toconserve genetic stocks and address the con-cerns raised in this evaluation effort. The pro-posed Avian Genetic Resources System (AVGRS)would be comprehensive and would require thecooperation and collaboration of scientists, fund-ing agencies, and research institutions. The Sys-tem must be oriented toward research objec-tives, but it could also support the needs ofbreeders, breed hobbyists, and breed historians.

RRRRRationaleationaleationaleationaleationaleHistoric long-term collaboration between Cana-dian and US scientists provides a basis for col-laboration in the formation and operation of anAvian Genetic Resources System. No compre-hensive system exists in North America, nor any-where else for that matter, for the conservationand distribution of avian genetic stocks (prima-rily chicken, turkey, and Japanese quail) ofvalue for agricultural, biomedical, or biologicalresearch. The US National Plant GermplasmSystem and its counterpart in Canada serve asexcellent models for the proposed Avian Genetic

Resources System. A System on this binationallevel is necessary because no dependable re-gional or local solution exists.

Components of an AComponents of an AComponents of an AComponents of an AComponents of an Avian Geneticvian Geneticvian Geneticvian Geneticvian GeneticRRRRResources Systemesources Systemesources Systemesources Systemesources System

The Avian Genetic Resources System is envi-sioned as a multilocational organization thatwould serve the avian genetic resources needsfor the US and Canada. The AVGRS would fea-ture a central facility as a focal point for many ofthe activities of the System. The functional com-ponents are outlined in Figure 16 and are brieflydiscussed here.

Avian Genetic Resources AdvisoryCommittee (AVGRAC)

The AVGRS would be advised by this binationalcommittee comprised of representatives of na-tional and state/provincial agencies, stock cura-tors, and researchers. It would consist of 12 to15 individuals who have worked with avian ge-netic resource issues, drawn from government,industry, and academic institutions in the USand Canada. Specifically, they should haveworked in close association with national, inter-national and private research-oriented organiza-tions, and be familiar with avian genetic stockconservation issues. The members should meetat least once a year and be in regular communi-cation during the year. The Committee wouldreview reports and recommendations for conser-vation of stocks received from species-orientedcommittees. It would make recommendations tothe management unit of the AVGRS.

6

40

Binational Avian GeneticResources Advisory Commmittee

Avian Genetic Resources System

Coordination

USDAUS NIHUS NSFA&AF CanadaCollectionsNGOs

International organizations FAO IUCNScientific societiesOthers

Conservation Importation/Quarantine

Research

Central facility

Live birdsPreserved tissuesEggsBloodDNACultured tissuesCryopreserved semen, embryonic cells

Cryopreservation

Beltsville MDGuelph ONOthers

Stock colonies

US - CA, IA, NH, WI, MI, CTCanada - BC, ONOthers

GenomicsWebsitesStock registryBreed registry

Coordinated with USand Canadianorganizations

DistributionDatabase/Information

EggsSpermEmbryosLive birdsDNAPreserved tissues

CryobiologyReproductive physiologyGeneticsStock characterization

Figure 16. Components of the Avian Genetic Resources System.

41

Coordination

The various government and research institu-tions involved in avian research and conserva-tion would use the AVGRS for coordination ofinformation about genetic resources and theAVGRS would in turn be responsible for main-taining and distributing this information. Thisfunction would also include strategic planningfor conservation of particular stocks, based onadvice from advisory groups established for eachspecies. For example, imperiled stocks would beidentified to the AVGRS and a plan for their con-servation would be developed through coordi-nated analysis. International relationships wouldbe coordinated through the AVGRS, includingconservation of stocks in other countries, importand export of genetic stocks, data sharing, anddevelopment of conservation plans for landraces,wild species, and breeding populations.

Conservation

This is the cornerstone activity of the AVGRSand of critical importance. A central facility isneeded for conservation and distribution of ge-netic materials. The central facility would housethose living genetic stocks that could not bemaintained elsewhere and would serve as abackup site for important stocks that are main-tained elsewhere. This would include a securebackup repository for privately owned lines orpopulations, either as live birds or cryopreservedgermplasm at the central or secondary centerson a fee basis. The central facility would alsophysically maintain the various types of pre-served genetic resources and would coordinatethose maintained elsewhere. The cryopreserva-tion capabilities of the central facility would besupplemented by a specialized cryopreservationcenter, presently unused, at the USDA site inBeltsville, Maryland. No site for the central facil-ity is identified at this time.

The central facility would support and belinked to secondary facilities located at activeresearch centers which have the capability ofmaintaining genetic stocks for their own re-search needs. Several locations in the US andCanada would qualify as secondary sites.

Methods of conservation. Conservation meth-ods employed in the AVGRS would be live-birdmaintenance and cryopreservation.

Targets for conservation. Conservation empha-sis of the AVGRS should be on live birds, embry-onic cells, gametes, DNA, and tissues. The targetspecies for the system will be those of interest in

agriculture for food production and for basic bio-logical and biomedical research. Thus, the focuswill be on chicken, turkey, and Japanese quailgenetic stocks. However, this system could alsoconsider wild turkey, jungle fowl, and gamebirds, as well as other species commonly raisedin captivity. The AVGRS will emphasize:

• Genetic stocks having traits and geneticcharacteristics useful in research, suchas inbred lines, single-gene mutations,chromosome aneuploidy, and DNAmarker sequences.

• Lines and populations developed by pri-vate and public breeders by hybridizationand selection for important production-related traits.

• Domesticated mid-level production andfancy breeds held by small producersand hobbyists in North America and Eu-rope.

• Domesticated, but primitive, landracesexisting in Asia, Central and SouthAmerica, and elsewhere, primarily asscavenger birds.

Archival preserved specimens of birds, or-gans, skeletons, eggs, feathers, and tissues thathave been preserved as museum specimens arealso a component of the genetic resources sys-tem, since these materials provide for baselineobservations and time-course monitoring of fac-tors such as environmental toxicants.

An informal in situ system of conservation oflandraces and breeds is well established inNorth America. A monitoring and database sys-tem may be the most important need for thosegenetic resources. This could become an activityof this proposed system.

DatabasesDetailed information about all genetic stocks inthe US and Canada should be maintained andupdated by the AVGRS in a genetic resourcesinformation system, similar to the Genetic Re-sources Information Network (GRIN) developedfor the US National Plant Germplasm Systemand housed with the USDA National AgricultureLibrary. For example, all of the information in-cluded in Appendix 2 should become part of theAVGRS database. Additionally, database servicewould be offered to the various breed conservan-cies and hobbyists groups for inventory and lo-cation of conserved breeds, land-races, and spe-

42

cialty stocks. It would also be logical for theAVGRS database to include DNA sequence dataas they are developed.

While GRIN currently focuses on plant infor-mation, its goal is to include information on allof the common and endangered breeds of farmanimals, including the avian genetic stocks usedprimarily in research. Collaboration with theAVGRS database would facilitate this goal.

OutreachResearchers can also be informed of the widevariety of available genetic stocks at the annualmeetings of a variety of organizations, includingthe Poultry Science Association, the Pacific Eggand Poultry Association, the Poultry BreedersRoundtable, the Society for Developmental Biol-ogy, the American Association for the Advance-ment of Science, and the American Medical As-sociation. Presentations at the commercially ori-ented meetings could be used to showcase thebenefits the different companies could derivefrom supporting an avian genetic stocks conser-vation program, while the basic research or dis-ease model aspect of genetic stocks could be em-phasized for organizations promoting basic andbiomedical research. Thus, the underlying goalsof presenting genetic stocks information at suchmeetings is not only to attract the attention ofresearchers, but to engage the interest and pro-mote funding from commercial sectors that canbenefit directly from research using avian ge-netic stocks.

Another outreach option for the AVGRSwould be an independent website that wouldpromote the available avian genetic stocks to thescientific community by advertising what isavailable, and indicating those that are slated forelimination. The effectiveness of such a sitecould be multiplied by linking it with websites:the animal genetic map (Angen), the chickenmap (ChickMap), various research organizationsites (e.g., Poultry Science Association, Societyfor Developmental Biology, various commercialpoultry sites, and sites for academic institutions).

The AVGRS website information could be fur-ther promoted by a series of clearly written re-view articles in several of the major biologicalscience journals. In each case, a specific area ofresearch would be targeted, such as animal mod-els for human diseases, limb pattern defects,craniofacial defects, integumentary defects, or im-munogenetic research.

The outreach activity would also involve inter-national contacts through FAO or various coun-tries with respect to avian genetic resources. Forexample, there are genetic stocks at risk in other

countries that should be considered for rescue inthe AVGRS because of their value for research.

Bird care and housingThe housing and care of the live birds at all cen-ters will follow American Association for Labora-tory Animal Care (AALAC) standards for a breed-ing colony. Essentially, the breeding stockshould be kept in a facility that approximatesthat of a well-run commercial poultry breedingfarm. The highest degree of automation for feed-ing, cleaning, watering and climate control isrecommended. With most of the chicken geneticstocks, the adult birds will be housed in single-bird cages, and bred by artificial insemination.Other features of the facility may include: floorpens, with or without trap nests, to maintainstocks that do not perform well in cages; andseparation of males and the females (in differentrooms or cage rows), so that appropriate, sex-specific breeder diets, lighting, and feeding regi-mens can be provided for each group.

To minimize the disease problems in the ge-netic stocks and reduce the risk of spreadingpathogens when stocks are shipped, all conser-vation centers should follow the National PoultryImprovement Plan guidelines (USDA-APHIS1998). These procedures impact on the design ofthe facility, such as:

• All incoming stocks should be acquiredonly as fertile eggs, which are formalde-hyde-fumigated, then incubated,hatched, and reared in a facility isolatedfrom all other avian species, includingthose already shown to be free of the tar-geted disease agents. After the birds inisolation are at least six weeks old, theyshould be blood-tested for the presenceof egg-transmitted pathogens (in chick-ens, this usually includes Salmonellapullorum, S. typhimurium, Mycoplasmagallisepticum, and M. synoviae; turkeyswould also be tested for M. gallinarium),before being moved to the stock center.

• A random-sample of 10% of the adultbirds should be tested each year for evi-dence of the important poultry diseases,including the disease pathogens listedabove.

• Replacement birds should be raised instrict isolation for the first month afterhatch.

• The stock center should be physicallywell-isolated from other poultry (commer-

43

cial or hobby flocks) and from facilities inwhich other captive bird species arehoused.

• The bird houses should prevent entry ofwild birds and vermin.

• Access to the facility should be restrictedand closely monitored, complying withfull necessary sanitary restrictions.

Importation and quarantine

Movement of animals introduces risks of spread-ing contagious diseases, obviously of great con-cern in long-term conservation of live birds.Movement of genetic resources as fertile eggs orsemen reduces disease transmission risk andare preferred procedures. Importation of stocksto the central facility will be done through on-site isolation and through national facilities un-der the direction of USDA Animal and PlantHealth Inspection Service. The central facilitywill have appropriate isolation and sanitationcapabilities.

Distribution

A major function of the AVGRS will be to providegenetic materials to users in the research com-munity, breeders, and others. The genetic stocksmay be transferred as live birds, semen, or eggs.These will be distributed on a cost-recovery ba-sis. Some users require a continuing supply ofgenetic stocks and these needs would be sup-plied by contractual stock reproduction pro-grams. The distribution function would supplywell-documented stocks to researchers, thuscontributing to the integrity of research projects.This functional component of the AVGRS is an-alogous to services provided by the JacksonLaboratories for mouse genetic stocks.

Research

The AVGRS should have a research capabilitywithin the central facility, especially for develop-ing cryopreservation technologies. Researchwould also be done on methods for documentinggenetic integrity or diagnostics with DNA mark-ers. Other research would be done as neededand appropriate. The research activities wouldbe networked with research laboratories in theUS and Canada for collaborative work.

FFFFFacilities and organizationalacilities and organizationalacilities and organizationalacilities and organizationalacilities and organizationalaspects of Aaspects of Aaspects of Aaspects of Aaspects of AVGRSVGRSVGRSVGRSVGRS

The central facility

Ideally, the primary AVGRS facility would beconstructed de novo near or part of a major agri-cultural institution (land-grant university) with aveterinary school that has a good avian medicineprogram, but reasonably isolated from commer-cial poultry stocks. The connection with a land-grant institution would give the center close tiesto active research laboratories and faculty, whocould benefit from such a resource and bedrawn upon in support of the center. Access tostate-of-the-art poultry disease diagnostics andveterinary care is critical, along with good, off-site quarantine facilities for newly acquired ge-netic stocks. Locating in a strong poultry pro-ducing state would also provide an existing poul-try-oriented political and commercial infrastruc-ture that could be mobilized to help support theconservation center.

This facility would include a hatchery, brood-ing and growing areas, adult bird housing, anisolation area, a cryopreservation laboratory andstorage facility, a database center, staff and ad-ministrative offices, and a laboratory to supportresearch and analytical services.

Bird housing should be designed to minimizedisease and pest control problems, and maxi-mize caretaker efficiency and bird well-being.Initially the facility should accommodate 1,000adult birds, with options for expansion. Severaldesign schemes will be evaluated to accommo-date the genetic diversity of the stocks and theirspecial requirements and for efficiency in man-agement.

Network of secondary genetic stockcentersSecondary stock centers would be designated aspart of the AVGRS at land-grant universities andother institutions across the US and Canadathat fulfilled two criteria: (1) had adequate facili-ties and support for the genetic stocks used inits own research programs and (2) had a long-term interest in conserving genetic stocks. Suchcenters, approved as part of the AVGRS, wouldreceive funding from AVGRS for maintenance ofstocks. These secondary centers would maintainlive birds and would provide backup for at-riskstocks held in the central facility. As with thecentral facility, the secondary stock colonieswould also have a distribution function on a feebasis. Researchers could also, on a fee basis,

44

arrange to keep research flocks at the centralfacility or one of the secondary centers, sincethey may find it difficult or impossible to keepsuch stocks at their own institutions.

Secondary centers could specialize on one ora few classes of genetic stocks (e.g., inbredstrains, congenic series, randombred stocks, se-lected stocks, single-gene mutant stocks, orchromosome translocation stocks). In fact, manyof the existing poultry collections at both aca-demic and government institutions are alreadyquite specialized. The restricted number ofclasses and species of stocks at each secondarylive bird center would simplify reproduction andmaintenance for the curators, since each type ofstock often has different needs in relation topopulation sizes, selection criteria, inbreedingconsiderations, and testing techniques.

ManagementThe central facility and the secondary centerswould be administered by the research institu-tions with which they are located. The centralfacility would have, at a minimum, a director ormanager (research scientist), curator, an admin-istrative assistant, database manager, cryopres-ervation research scientist, and three or fourlaboratory and animal care technicians.

The AVGRS would be guided by the advisorycommittee on such matters as:

• Information system and websitemanagement

• Inventory control

• Conservation status determination(cryopreserved stocks only)

• Stock accessioning strategies

• Importation targets and necessary inter-national cooperation agreements

• Appropriate stock-specific conservationprotocols

• Management of grant funds for researchand preservation efforts

• Curator training programs

• Program expansion decisions

• Budgets

• Fund raising

• Species coordinating committeeorganization

• Promotion of the use of these geneticstocks

• User fees and maintenance contractguidelines

Stock evaluation guidelines

One of the more important activities of theAVGRAC would be the evaluation of stocks forconservation, cryopreservation, or elimination.Guidelines for assessing the value of geneticstocks have been outlined in a report by the Na-tional Research Council Committee on Preserva-tion of Laboratory Animal Resources (NRC1990). The report suggests that value of a givenstock should be established by considering:

1. Importance of the disease process orphysiological function exemplified by thestock (especially when used as an animalmodel for a human disorder).

2. Validity of the model and continued ge-netic integrity of the stock.

3. Replaceability of the stocks (those devel-oped after many years of selection orarising from a spontaneous mutationwould be considered relatively irreplace-able).

4. Versatility of the stock (the variety ofproblems that can be addressed with agiven stock).

5. Number of users.

The advisory committee would also formulateother criteria specifically relevant for the conser-vation of avian species.

FFFFFinancing the Ainancing the Ainancing the Ainancing the Ainancing the Avian Geneticvian Geneticvian Geneticvian Geneticvian GeneticRRRRResources Systemesources Systemesources Systemesources Systemesources System

Multiple sources of funding will be necessary tomeet all of the needs of the AVGRS. Initial costsare those to construct the central facility andupgrade the secondary stock centers. Annualcosts of the central facility would be for its per-sonnel and operations. It would also be neces-sary to support the annual activity of theAVGRAC. The central facility could also direct

45

funds for specific needs to the secondary centersby means of annual grants.

From the US side, the biological resource pro-grams of the National Science Foundation andthe National Institutes of Health would be ex-pected to provide operational funds through di-rect grants and through grants to investigatorswho use the avian genetic resources in their re-search. The USDA’s National Genetic ResourcesSystem should participate in the AVGRS throughthe Agricultural Research Service and the Coop-erative State Research, Extension, and Educa-tion Service. The various State Agricultural Ex-periment Stations and land-grant and other Uni-versities should also participate. Canadian sup-port and participation should be forthcoming tothe extent that the AVGRS provides support toits research and development programs.

The AVGRS will be the major provider of ge-netic materials to researchers throughout thepublic and private sectors. For the most part,these researchers do not have capacity to main-tain live bird colonies and depend upon stockcolonies for their research. User-fees are an ap-propriate means to recoup costs of stock mainte-nance.

Donation funds can be expected to supportthe perpetual maintenance of particular geneticstocks. These funds may be provided as annualgrants or through income derived from intereston endowment accounts.

Funding should be sought from the US gov-ernment for construction of the central facilityand for personnel support for operations as apart of the US National Animal Genetic Resour-ces System.

Long-term funding would be the most securefrom endowment funds. Contributors could beencouraged from the private sector, from largeintegrated commercial poultry companies to pri-vate individuals with interests in preservingpoultry stocks or willing to promote the conser-vation of stocks that can be used to study hu-man diseases.

Construction, personnel, and operationalcosts have not been established, pending furtheranalysis of potential sites for the central facilityand other considerations. For illustrative pur-poses, rough order-of-magnitude estimates aregiven in Table 4.

Table 4. Estimated costs of Avian Genetic Resources System.Startup costsConstructing and equipping central facility $15,000,000Upgrading and renovating secondary centers 2,000,000Total $17,000,000

Annual costsPersonnel at central facility $400,000Operating costs at central facility 100,000Grants to secondary centers (8 x $25,000) 200,000Advisory Committee 25,000Total $725,000

47

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Appendix 1. Instructions and surAppendix 1. Instructions and surAppendix 1. Instructions and surAppendix 1. Instructions and surAppendix 1. Instructions and survey form sent tovey form sent tovey form sent tovey form sent tovey form sent toresearchersresearchersresearchersresearchersresearchersINSTRUCTIONS FOR COMPLETING THE AVIAN GENETIC RESOURCES SURVEYWe appreciate your willingness to assist us in this survey. Below are some suggestions and clarifica-tions for the enclosed survey form. If you need more space to complete your catalog of genetic stocks,please feel free to copy the survey form. We would also be interested in any relevant information youwish to attach that concerns your stocks (previously published summary information, references,expanded descriptions, etc.).

1. Curating researcher or person in charge of these stocks: this is the name of the person respon-sible for the poultry stocks recorded in the survey. This person can be contacted for further informa-tion on these stocks. If more than one person is responsible for a given stock or group of stocks,please write in the other names and addresses/phone numbers at the top of the page, and indicatethe stocks that each person controls.

2. Name of Stock: how the stock is identified.

3. Species: chicken, turkey, coturnix quail or other avian species.

4. Stock Description/Characteristics: breed and any special stock characteristics, including degreeof inbreeding (for inbred stocks), selection criteria, and/or mutations or chromosomal abnormalitiespresent. If this stock is unchanged since it was listed in the 1988 International Registry, just write in“see 1988 registry” under this category. If you need more space to adequately describe a stock, pleaseattach the expanded description to the survey form.

5. Eliminated? Date/Reason: if a stock has been eliminated, please indicate date of and reason forelimination (i.e., loss of funding, housing no longer available, stock no longer needed, etc.).

6. # Birds for Maint. (M & F): how many male and female birds are needed in a core flock to maintaina viable breeding population of each stock.

7. Funding Situation: security of the funding for that stock, at present and projected five years. Ifpossible, please indicate source of funding: Government (NIH, NSF, USDA, etc.); Academic (universityor institution, Hatch funds); Private (private organizations or individuals).

8. Other Loc.? (Y/N): is the stock maintained at one or more other institutions/locations? If yes,please give the name and address of the person(s) responsible for this stock at the other location(s), ifavailable.

9. Cryo.? (Y/N): is the stock also (or only) kept as cryopreserved cells? If yes, please indicate the typeof cells that have been cryopreserved (semen, blastodiscs, etc.), and where this material is kept.

56

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Appendix 2. Genetic stock collection dataAppendix 2. Genetic stock collection dataAppendix 2. Genetic stock collection dataAppendix 2. Genetic stock collection dataAppendix 2. Genetic stock collection dataTHE INFORMATION PRESENTED HERE was collectedthrough a survey initiated in 1995 by the Uni-versity of California Genetic Resources Conser-vation Program. Many stocks developed before1988 were listed in the International Registry ofPoultry Genetic Stocks (SOMES, R.G., JR. 1988.International Registry of Poultry Genetic Stocks.Storrs Agr. Exp. Sta. Bull. 476. The University ofConnecticut, Storrs). Therefore, curators listedin the Somes registry were the starting point forthis survey. In addition, queries were sent tomembers of the NE-60 Regional Research Project(a USDA-organized group of researchers workingon the genetic basis for resistance and immunityto avian diseases), academic and governmentmembers of the Poultry Science Association(source list: PSA Membership Directory, 1995-98) who indicated a focus on genetics or immu-nology, and individuals listed by the 1997 Poul-try Science Resource List (prepared by the Poul-try Science Association and R.D. Reynnells(USDA/CSREES)) who specialized in genetics orimmunology and were affiliated with academicinstitutions with poultry, avian, or animal sci-ence departments.

Altogether, a total of more than 100 individu-als were polled. Seventy-five responded and, ofthose, 40 currently maintain poultry geneticstocks as researcher-curators. These curatorsrepresent 27 different institutions (24 in the US,3 in Canada). Approximately one-half of the in-stitutions maintain 10 or fewer genetic stocksand only six maintain more than 20 stocks (Uni-versity of Arkansas–Lafayette, University of Cali-fornia–Davis, University of Connecticut–Storrs,Univerisity of Wisconsin–Madison, USDA-ADOLEast Lansing, MI, and University of British Co-lumbia–Vancouver). About one-third of the re-ported stocks are considered to be at seriousrisk of loss in the near future, and at least 94 ofthe surveyed stocks were eliminated between

1995 and 1998. Over half of these discardedstocks were not represented in collections atother institutions and thus, are lost to science.Most stocks are maintained as living birds, but38 chicken genetic stocks are maintained only inthe form of cryopreserved semen or blastodermalcells.

The survey data are presented in three tables.Table 2.1 is a listing of the 361 reported stocksgrouped first by species (chicken, Japanesequail, turkey, and waterfowl/gamebird), then bycategory of genetic stocks (the 28 categoriesused are listed in Table 2.0, then described be-

Bloodtype-Gene poolBloodtype-Major Histocompatibility Complex (MHC)

Bloodtype-MHC-InbredChromosomal variantEndogenous virusEndogenous virus-InbredInbredMutant-Color, eggshellMutant-Color, featherMutant-Developmental defect-EyeMutant-Developmental defect-Face/limbMutant-Developmental defect-Skin/featherMutant-Developmental defect-Spine/tailMutant-Gene poolMutant-Immunological defectMutant-Neurological defectMutant-Physiological defectMutant-Reproductive defectMutant-UncategorizedPure breedRandombredSelected-Behavioral traitSelected-Egg traitSelected-Growth traitSelected-Immune traitSelected-Physiological traitTransgenicUncategorized

Table 2.0. Stock categories.

58

low). Table 2.2 lists stocks grouped by institutionand curator, species, and category. Finally, Table2.3 lists curators and their contact information.Accordingly, the most important informationnecessary to look up a given genetic stock in-cludes 1) the species and type of stock or 2) theinstitution or curator who keeps that type ofstock.

Description of stock categoriesDescription of stock categoriesDescription of stock categoriesDescription of stock categoriesDescription of stock categoriesBloodtype refers to cell-surface antigens thathave been characterized on red or white bloodcells. Bloodtype-Major HistorcompatibilityComplex (MHC) consists of stocks with a de-fined MHC or B-haplotype, and may be definedfor a variety of other erythrocyte alloantigens.Bloodtype-MHC-Inbred is congenic (geneticallyidentical) to an established highly inbred line,except for the specified bloodtype. Bloodtype-Gene pool stocks contain a variety of differentbloodtypes. Leukocyte cell surface antigens arealso defined for one gene pool stock (NIU MaleBreeder Alloantigen Reservoir).

Chromosomal variant refers to stocks with al-terations in chromosome structure. These varia-tions include: insertions, deletions, transloca-tions, inversions, aneuploidy, and polysomy.Stocks with specific insertions (naturally occur-ring or as a result of genetic engineering) arelisted under the Endogenous virus or Transgeniccategories.

Endogenous virus (EV) stocks are characterizedby specific viral insertions (one or more inser-tions of a defined endogenous virus), and En-dogenous virus-Inbred stocks are congenic (ge-netically identical) to an established highly in-bred line, except for the specified viral insertion.

Inbred stocks usually have a level of inbreedingexceeding 95% (F>0.95), although one exceptionis UCD 001 (F is approximately 0.90). Other in-bred stocks include congenic stocks developedby continued back-crossing to a given inbredstock. Such congenic stocks are categorized bythe unique gene or gene complex isolated in theinbred background (e.g., Bloodtype-MHC-Inbred,Endogenous virus-Inbred, and Mutant-Develop-mental defect-Face/limb). In such stocks, theinbred ancestral stock is listed under “Originand History”.

Mutant stocks are classified into the followingcategories:

Mutant-Color, eggshellMutant-Color, featherMutant-Developmental defect-EyeMutant-Developmental defect-Face/limbMutant-Developmental defect-Skin/featherMutant-Developmental defect-Spine/tailMutant-Gene poolMutant-Immunological defectMutant-Neurological defectMutant-Physiological defectMutant-Reproductive defectMutant-Uncategorized

Color variation for feather and eggshell alsooccur in the pure breed and randombred catego-ries, particuarly among the different chickenbreeds. Researchers interested in such variantsshould look up the color genetics for thesebreeds or discuss this with the pure breed stockcurators.

Some developmental defect mutations mayactually fit into more than one category (e.g.,wingless-2 affects face, limbs, feathers, and kid-neys, but was listed by its effect on the face andlimbs). Some of the mutations listed under neu-rological, physiological, and reproductive defectscan also cause developmental defects. On theother hand, a few of the mutations or variantsdid not clearly fit into any of the above specificcategories, so were classified as Mutant-Uncategorized. A close reading of the mutantdescriptions is recommended.

Mutant-Gene pool refers to stocks that containtwo or more mutations, which may or may notaffect affect the same structures or systems.Some unique mutations are only maintained ingene-pool stocks, and so do not appear underthe more specific mutant categories. However,brief descriptions of all mutations carried withinthat stock are listed for each of the pooledstocks.

The Pure breed category includes all stocks thatare maintained as a closed flock of an identifiedbreed. It is usually not highly inbred, selected, orotherwise characterized. Some overlap occurs be-tween the Pure breed category and the Random-bred category. However, if an otherwise Purebreed stock is commonly used as a control/stan-dard for another stock, it will be listed as Ran-dombred.

Randombred stocks are usually purebred or syn-thetic (mixed commercial) stocks that are kept inlarge, closed populations (100 birds or more).

59

Typically propagated with little or no selection ofbreeding stock each generation, the number ofprogeny from each male or female usually de-pends upon their reproductive success at the timeeggs are collected to produce the next generation.

Stocks that have been selectively bred for aquantitative characteristic are termed Selected-along with the identified trait: Selected-Behav-ior trait, Selected-Egg trait, Selected-Growthtrait, Selected-Immune trait, and Selected-Physiological trait. Each stock usually has anunselected control paired with it, and two diver-gently selected stocks (e.g., for high or low bodyweight at 6 weeks) from the same source may begrouped together here.

Transgenic stocks are the product of geneticengineering. Such stocks have successfully inte-grated the introduced genetic material into theirchromosomes.

Finally there is disparate group of stocks classi-fied as Uncategorized, either for want of betterdescriptive information or because there may betoo few representatives of a type to warrant anadditional category.

While most stocks clearly belong to a single de-scriptive category, some logically could be classi-fied in more than one. In such situations, thestock was listed under the category that mostclosely fit the stock description. The distributionof all listed stocks according to these categoriesis summarized in Chapter 5 of this report.

Notes on specific fieldsNotes on specific fieldsNotes on specific fieldsNotes on specific fieldsNotes on specific fieldsTable 2.1 contains the fields described below.Stock name is also used in Table 2.2.

Stock name: The designation used usually in-corporates the name of the institution where thestock is housed or was developed, and the namethat the curator assigned to the stock. In Table2.2, the stock name is followed by the words‘Cryo. only’, if that stock is only maintained bycryopreservation. This information is given in theStatus field for Table 2.1 (see below).

Stock description: Only a brief description ofthe unique characteristics of the stocks is pro-vided; for more information, the curator shouldbe contacted.

Origin and history: Information in this field in-cludes when a stock was established or acquired,

its current and historical genetic background (ifavailable), and any other possibly useful back-ground information. In particular, for stocks thatare congenic, the specific inbred stock with whichit is essentially identical is noted.

Breed: The source breed(s) or genetic back-ground for stocks is noted here.

Genetic characteristics: For those stocks thatare characterized by a distinct mutation or set ofmutations, this field presents information on thepattern of inheritance (autosomal, sex-linked),penetrance and expressivity (dominant, co-domi-nant, recessive, incomplete penetrance, mater-nal effect, sex-limited), and effect on viability (le-thal, semi-viable; if a mutation is fully viable, nonotation is provided).

Allele symbol: Here the short notation for themutant form of the gene is listed. If the symbolis in question or has not yet been officially as-signed, the allele symbol is written in parenthe-ses (i.e., (wgr) for the wing-reduced syndrome).

Status: This field provides a subjective evalua-tion of the potential for continued financial sup-port of each stock, assessed by the curator ofthat stock. ‘Good’ indicates dependable institu-tional or government support for at least thenext two to three years. ‘Fair’ indicates less cer-tain support. ‘Poor’ describes a stock which isseriously at risk of elimination. When the spe-cific source of funding support for the stock wasprovided, that information was noted: USDA,Industry, Institutional, State Agricultural Ex-periment Station (Exp. Sta.), Hatch funds(Hatch), research funds (regional), or other(misc.). In some cases, a stock is no longer main-tained as living birds, only by germplasm cryo-preservation. For such stocks, the notation inthe status field is ‘No live birds’. Finally, someonce-distinct mutant stocks are no longer car-ried as separate accessions, but were recentlycombined with another accession, and the nameof the recipient stock is indicated in the statusfield (e.g., ‘Combined with UBC G’). For the pur-poses of Table 2.1 and this report, the originalmutant is still counted as a separate accession.

Number of birds: When available, this is thenumber of birds (males and females) kept tomaintain the stock.

Cryopreservation: If no germplasm has beencryopreserved for a given stock, the entry in thisfield is ‘No’. If germplasm has been cryopreserved,

60

then the cell type is indicated (semen, germ cells,blastodisc cells (for dissociated blastodiscs)).Stocks that are only maintained as cryopreservedgermplasm can be identified by the remark ‘Nolive birds’ in the Status field. Curators can be con-tacted for details on recovery of these stocks andcosts associated with the recovery.

Available: Many curators provide eggs or chicks toother researchers for collaborative projects; some willprovide eggs or chicks for a fee (usually to offset birdmaintenance costs), while other stocks may not beavailable because of proprietary concerns. Curatorsshould be contacted for details.

Curator: The surname of the researcher incharge of the given stock is indicated here. Con-tact information for them is listed in Table 2.3,alphabetically, by surname.

Disclaimer

The University of California Genetic ResourcesConservation Program and the Avian Genetic

Resources Task Force present this informationas a service to those interested in poultry geneticstocks that can be found at academic and gov-ernment institutions in the USA and Canada. Nocertification is implied for the stocks listedherein, either their status with regard to freedomfrom egg-borne disease agents, or their currentavailability (note that availability at the time thesurvey was conducted is indicated). Inquiriesregarding particular stocks should be directed tothe individual curator(s). In addition, readers arereminded that it is not possible to guarantee theauthenticity or scientific accuracy of the infor-mation presented herein.

Those wishing to acquire any of these stocksor genetic material derived from these stocks arereminded that they must obtain the appropriatecustoms forms and documentation, and followthe protocols governing such shipments. Eventransfer between different states in the US orprovinces in Canada may require health certifi-cation or other documentation. In the US, theUSDA Veterinary Services can provide some ofthis information.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

1

Chi

cken

Blo

odty

pe-G

ene

pool

Aubu

rn D

AXKe

pt as

clos

ed flo

ck

SCW

Lse

eFa

ir24

0No

With

Ewald

MHC:

B5;

other

eryth

rocy

te all

oanti

gens

: A/E

4, C4

, D3,

since

1965

desc

riptio

nco

llabo

ratio

nH2

, I3, K

3, P3

, “Q”

2.NI

U B

hapl

otyp

esDe

rived

from

Anc

ona s

ynthe

ticMi

xed A

ncon

ase

eFa

irN/

ANo

With

Brile

sPo

ol of

MHC

B-ha

plotyp

es: 1

, 3, 4

, 6, 8

, 10,

11, 1

2, 13

,sto

cks h

omoz

ygou

s for

mos

tde

scrip

tion

colla

bora

tion

14, 1

5, 17

, 22,

23, 2

4, 26

, 30,

31, 3

2, 33

, 24r

1, 2r

1, 2r

2,no

n-MH

C ge

nes

2r3,

21r1

, 21r

2, 2r

4, 2r

5, 24

r2, 2

4r3,

21r6

, 8r1

.NI

U B-

hapl

otyp

e Rec

ombi

nant

sDe

rived

from

Anc

ona s

ynthe

ticMi

xed A

ncon

ase

eFa

irN/

ANo

With

Brile

sRe

comb

inant

B ha

plotyp

es: R

1, R2

, R3,

R4, R

5, R6

, R7,

stock

s hom

ozyg

ous f

or no

n-MH

Cde

scrip

tion

colla

bora

tion

R8, R

9, R1

0, R1

1, R1

2.sy

stem

gene

sNI

U Ma

le Br

eede

r Allo

antig

en R

eser

voir

Segr

egati

ng fo

r eryt

hroc

yte an

dSC

WL

see

Fair

200+

NoW

ithBr

iles

Pool

of ce

ll sur

face e

rythr

ocyte

alloa

ntige

n (# a

lleles

):leu

cocy

te all

oanti

gens

desc

riptio

nco

llabo

ratio

nA(

8), B

(40)

, C(8

), D(

5), E

(10)

, H(2

), I(7

), K(

2), L

(2),

P(10

),R(

2); L

euko

cyte

alloa

ntige

ns: M

(5),

N(2)

, Q(3

), T(

3), U

(4),

W(2

), Z(

2).

NIU

Segr

egat

ing

Male

Bree

der L

ine

SCW

Lse

eFa

irN/

ANo

With

Brile

sMH

C: B

2/B5 o

r B19

/B21

; A4E

1/A5E

2; C2

/C5;

D1/D

3;de

scrip

tion

colla

bora

tion

H1/H

2; I2

/I8; K

2/K3

; L1/

L2; P

1/P4.

UNH

105

Kept

as cl

osed

flock

Ne

w Ha

mpsh

irese

eGo

od10

M/15

0FNo

Yes

Taylo

rCl

osed

flock

with

four

MHC

B ha

plotyp

es: 2

2, 23

, 24,

26.

since

1981

desc

riptio

n

Blo

odty

pe-M

HC

Aubu

rn M

Deriv

ed fr

om co

mmer

cial m

eat s

tock

Comm

ercia

lB8

Fair

35No

NoEw

aldMH

C: B

8.in

1982

meat

stock

s(p

ropr

ietar

y)Au

burn

NDe

rived

from

comm

ercia

l mea

t stoc

kCo

mmer

cial

B13

Fair

100

NoNo

Ewald

MHC:

B13

.in

1982

meat

stock

s(p

ropr

ietar

y)Au

burn

RM

Cros

s of A

ubur

n R an

d Aub

urn M

SCW

L XB2

Good

70No

With

Ewald

MHC:

B2.

stock

sco

mmer

cial

colla

bora

tion

meat

stock

sAu

burn

RMH

Deriv

ed fr

om cr

oss o

f Aub

urn R

SCW

LB3

Fair

50No

With

Ewald

MHC:

B3.

and A

ubur

n MH

stock

sco

llabo

ratio

nAu

burn

RN

Deriv

ed fr

om cr

oss o

f Aub

urn R

SCW

L XB1

3Go

od84

0No

With

Ewald

MHC:

B13

.an

d Aub

urn N

lines

in 19

88co

mmer

cial

colla

bora

tion

meat

stock

sCo

rnell

N2a

Spec

ific pa

thoge

n-fre

eSC

WL

B21,

C2Go

od23

M/10

0FNo

Yes,

feeSc

hat

MHC:

B21

; eryt

hroc

yte al

loanti

gen C

2; hig

hly re

sistan

t toMa

rek’s

dise

ase v

irus.

Corn

ell P

2aSp

ecific

patho

gen-

free

SCW

LB1

9, C2

Good

30M/

110F

NoYe

s, fee

Scha

tMH

C: B

19, e

rythr

ocyte

alloa

ntige

n C2;

highly

susc

eptib

le to

Mare

k’s di

seas

e viru

s.NI

U Fe

male

Bre

eder

Par

ent S

tock

B19

Deriv

ed fr

om A

ncon

a syn

thetic

Mi

xed A

ncon

aB1

9Fa

irN/

ANo

With

Brile

sMH

C: B

19; u

sed a

s fem

ale pa

rent

in im

muno

logica

lsto

cks h

omoz

ygou

s for

non-

MHC

colla

bora

tion

chall

enge

s.sy

stem

gene

s

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

2

Chi

cken

(con

t.)B

lood

type

-MH

C (c

ont.)

NIU

Fem

ale B

reed

er P

aren

t Sto

ck B

2De

rived

from

Anc

ona s

ynthe

ticMi

xed A

ncon

aB2

Fair

N/A

NoW

ithBr

iles

MHC:

B2;

used

as fe

male

pare

nt in

immu

nolog

ical

stock

s hom

ozyg

ous f

or no

n-MH

Cco

llabo

ratio

nch

allen

ges.

syste

m ge

nes

NIU

Fem

ale B

reed

er P

aren

t Sto

ck B

5De

rived

from

Anc

ona s

ynthe

ticMi

xed A

ncon

aB5

Fair

N/A

NoW

ithBr

iles

MHC:

B5;

used

as fe

male

pare

nt in

immu

nolog

ical

stock

s hom

ozyg

ous f

or no

n-MH

Cco

llabo

ratio

nch

allen

ges.

syste

m ge

nes

RPRL

-Cor

nell J

M-N

Deriv

ed fr

om C

orne

ll JM

rand

ombr

edSC

WL

B21

Good

:6M

/42F

Seme

nYe

sBa

con

MHC:

B21

; res

istan

t to M

arek

’s dis

ease

viru

s stra

in JM

.sto

ck; s

pecif

ic pa

thoge

n-fre

eUS

DARP

RL-C

orne

ll JM-

PDe

rived

from

Cor

nell J

M ra

ndom

bred

SCW

LB1

9Go

od:

6M/42

FSe

men

Yes

Baco

nMH

C: B

19; s

usce

ptible

to M

arek

’s dis

ease

viru

s stra

insto

ck; s

pecif

ic pa

thoge

n-fre

eUS

DAJM

.UN

H 19

2De

rived

from

cros

s of S

CWL a

ndSC

WL X

B19

Good

10M/

150F

NoYe

sTa

ylor

MHC:

B19

.An

cona

; kep

t as c

losed

flock

Anco

nasin

ce 19

88

Blo

odty

pe-M

HC

-Inbr

edIS

U 19

-13

Deriv

ed fr

om cr

osse

s of 1

920s

ISU

SCW

LB1

3Go

od2M

/16F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.98; M

HC: B

13.

inbre

d stoc

ks be

fore 1

935

colla

bora

tion

ISU

19-1

5.1De

rived

from

cros

ses o

f 192

0s IS

USC

WL

B15.

1Go

od2M

/16F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.98; M

HC: B

15.1.

inbre

d stoc

ks be

fore 1

935

colla

bora

tion

ISU

8-15

.1De

rived

from

cros

ses o

f 192

0s IS

UBa

rred L

egho

rnB1

5.1

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.98

; MHC

: B15

.1.inb

red s

tocks

befor

e 193

5co

llabo

ratio

nIS

U G-

B1Co

ngen

ic wi

th IS

U GH

SCW

LB1

3Go

od3M

/30F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.99; M

HC: B

13.

colla

bora

tion

ISU

G-B2

Cong

enic

with

ISU

GHSC

WL

B6Go

od3M

/30F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.99; M

HC: B

6.co

llabo

ratio

nIS

U GH

-1De

rived

from

a cro

ss of

Gho

stley

SCW

LB1

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.99

; MHC

: B1.

Hatch

ery (

MN) f

emale

s and

HN

colla

bora

tion

males

in 19

54IS

U GH

-13

Deriv

ed fr

om a

cross

of G

hostl

eySC

WL

B13

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.99

; MHC

: B13

.Ha

tcher

y (MN

) fem

ales a

nd H

Nco

llabo

ratio

nma

les in

1954

ISU

GH-1

5.1De

rived

from

a cro

ss of

Gho

stley

SCW

LB1

5.1

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.99

; MHC

: B15

.1.Ha

tcher

y (MN

) fem

ales a

nd H

Nco

llabo

ratio

nma

les in

1954

ISU

HN-1

2De

rived

from

a pu

re K

imbe

r line

from

SCW

LB1

2Go

od2M

/16F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.99; M

HC: B

12.

H&N

in 19

54co

llabo

ratio

nIS

U HN

-15

Deriv

ed fr

om a

pure

Kim

ber li

ne fr

omSC

WL

B15

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.99

; MHC

: B15

.H&

N in

1954

colla

bora

tion

ISU

M15.2

Impo

rted f

rom

Egyp

t in 19

54Fa

youm

iB1

5.2

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.98

; MHC

: B15

.2; or

igina

lco

llabo

ratio

nsto

ck th

ough

t to be

resis

tant to

lymp

hoid

leuko

sis.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

3

Chi

cken

(con

t.)B

lood

type

-MH

C-In

bred

(con

t.)IS

U M5

.1Im

porte

d fro

m Eg

ypt in

1954

Fayo

umi

B5.1

Good

2M/16

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.98

; MHC

: B5.

1; or

igina

lco

llabo

ratio

nsto

ck th

ough

t to be

resis

tant to

lymp

hoid

leuko

sis.

ISU

Sp21

.2Im

porte

d fro

m Sp

ain in

1954

Span

ishB2

1.2

Good

3M/30

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.98

; MHC

: B21

.2.co

llabo

ratio

nNC

SU G

B-1

Acqu

ired f

rom

Iowa S

tate U

;SC

WL

B13

Good

40M/

100F

NoYe

sQu

resh

iMH

C: B

13.

cong

enic

with

ISU

GHNC

SU G

B-2

Acqu

ired f

rom

Iowa S

tate U

;SC

WL

B6Go

od40

M/10

0FNo

Yes

Qure

shi

MHC:

B6.

cong

enic

with

ISU

GHRP

RL 15

.15-5

Cong

enic

with

RPRL

15I5;

spec

ificSC

WL

B5Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

5 fro

m RP

RL 15

I4.pa

thoge

n-fre

eUS

DARP

RL 15

.6-2

Cong

enic

with

RPRL

15I5;

spec

ificSC

WL

B2Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

2 fro

m RP

RL 6I

1.pa

thoge

n-fre

eUS

DARP

RL 15

.7-2

Cong

enic

with

RPRL

15I5;

spec

ificSC

WL

B2Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

2 fro

m RP

RL 7I

2.pa

thoge

n-fre

eUS

DARP

RL 15

.C-1

2Co

ngen

ic wi

th RP

RL 15

I5; sp

ecific

SCW

LB1

2Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

12 fr

om R

ease

heath

line C

.pa

thoge

n-fre

eUS

DARP

RL 15

.N-2

1Co

ngen

ic wi

th RP

RL 15

I5; sp

ecific

SCW

LB2

1Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

21 fr

om C

orne

ll JM-

N.pa

thoge

n-fre

eUS

DARP

RL 15

.P-1

3Co

ngen

ic wi

th RP

RL 15

I5; sp

ecific

SCW

LB1

3Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

13 fr

om C

orne

ll JM-

P.pa

thoge

n-fre

eUS

DARP

RL 15

.P-1

9Co

ngen

ic wi

th RP

RL 15

I5; sp

ecific

SCW

LB1

9Go

od:

6M/35

FSe

men

Yes

Baco

nMH

C: B

19 fr

om C

orne

ll JM-

P.pa

thoge

n-fre

eUS

DARP

RL 15

I5Sp

ecific

patho

gen-

free

SCW

LB1

5,Go

od:

9M/63

FSe

men

Yes

Baco

nInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B15

; end

ogen

ous

ALVE

1,US

DAvir

uses

ev1,

ev6,

and e

v10 o

r ev1

1; su

scep

tible

toAL

VE6,

avian

leuk

osis

virus

e A an

d B an

d Mar

ek’s

disea

seAL

VE10

,vir

us.

ALVE

11RP

RL 7I

1Sp

ecific

patho

gen-

free

SCW

LB2

, ALV

E1,

Good

:12

M/84

FSe

men

Yes

Baco

nInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B2;

endo

geno

usAL

VE3

USDA

virus

es ev

1 and

ev3;

susc

eptib

le to

avian

leuk

osis

virus

e C an

d Mar

ek’s

disea

se vi

rus.

RPRL

Lin

e 0Sp

ecific

patho

gen-

free

SCW

LB2

1Go

od:

12M/

84F

Seme

nYe

sBa

con

MHC:

B21

; free

of al

l kno

wn en

doge

nous

viru

ses;

USDA

susc

eptib

le to

avian

leuk

osis

virus

es A

, B &

C.

UCD

003

Full-s

ib cro

sses

sinc

e 195

6; on

e of

SCW

LB1

7, A4

/E7

Poor

10M/

12F

Seme

nYe

sAb

plan

alpInb

reed

ing co

effici

ent (

F)>0

.99; A

/E bl

ood t

ypes

4/7;

the re

feren

ce lin

es in

the E

ast

MHC:

B17

; sho

wn to

be re

sistan

t to R

ous s

arco

ma an

dLa

nsing

Chic

ken G

enom

e Map

ping

susc

eptib

le to

Mare

k’s di

seas

e viru

s .Pr

oject,

and f

orms

the g

eneti

cba

ckgr

ound

for a

numb

er of

cong

enic

strain

s tha

t car

ry va

rious

MHC

haplo

types

or de

velop

menta

lmu

tation

s (se

e UCD

stoc

k list

)

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

4

Chi

cken

(con

t.)B

lood

type

-MH

C-In

bred

(con

t.)UC

D 25

3Co

ngen

ic wi

th UC

D 00

3SC

WL

B18

Poor

5M/6F

Seme

nYe

sAb

plan

alpMH

C: B

18; s

hown

to be

resis

tant to

Rou

s sar

coma

and

Mare

k’s di

seas

e viru

ses.

UCD

254

Cong

enic

with

UCD

003

SCW

LB1

5Po

or5M

/6FSe

men

Yes

Abpl

analp

MHC:

B15

; sho

wn to

be su

scep

tible

to Ro

us sa

rcoma

and M

arek

’s dis

ease

viru

ses.

UCD

312

Cong

enic

with

UCD

003;

MHC

from

SCW

LB2

4Po

or5M

/6FSe

men

Yes

Abpl

analp

MHC:

B24

; sho

wn to

be su

scep

tible

to Ro

us sa

rcoma

New

Hamp

shire

chick

en st

rain

and M

arek

’s dis

ease

viru

ses.

UCD

313

Cong

enic

with

UCD

003;

died o

ut as

SCW

LB3

No liv

e0

Seme

nW

ithAb

plan

alpMH

C: B

3; sh

own t

o be r

esist

ant to

Rou

s sar

coma

viru

s.liv

e bird

s in 1

996

birds

perm

ission

UCD

330

Cong

enic

with

UCD

003;

MHC

from

SCW

LB2

1Po

or5M

/6FSe

men

Yes

Abpl

analp

MHC:

B21

; sho

wn to

be re

sistan

t to R

ous s

arco

ma an

dAu

stralo

rp in

bred

line

Mare

k’s di

seas

e viru

ses.

UCD

331

Cong

enic

with

UCD

003;

MHC

from

aSC

WL

B2Po

or5M

/6FSe

men

Yes

Abpl

analp

MHC:

B2;

show

n to b

e res

istan

t to R

ous s

arco

ma an

ddw

arf S

CWL

Mare

k’s di

seas

e viru

ses.

UCD

335

Cong

enic

with

UCD

003;

MHC

from

aSC

WL

B19

Poor

5M/6F

Seme

nYe

sAb

plan

alpMH

C: B

19; s

hown

to be

susc

eptib

le to

Rous

sarco

maco

mmer

cial R

ichar

dson

Mt. H

ope

and M

arek

’s dis

ease

viru

ses.

SCW

LUC

D 33

6Co

ngen

ic wi

th UC

D 00

3; MH

C fro

mSC

WL

BQPo

or5M

/6FSe

men

Yes

Abpl

analp

MHC:

BQ

(simi

lar to

B21

); sh

own t

o be r

esist

ant to

Rou

sRe

d Jun

gle F

owl

sarco

ma an

d Mar

ek’s

disea

se vi

ruse

s.UC

D 34

2Co

ngen

ic wi

th UC

D 00

3; MH

C fro

m a

SCW

LBO

Poor

5M/6F

Seme

nYe

sAb

plan

alpMH

C: B

O; sh

own t

o be s

usce

ptible

to R

ous s

arco

macro

ss of

Cey

lon Ju

ngle

Fowl

and

virus

.Re

d Jun

gle F

owl

UCD

361

Cong

enic

with

UCD

003

SCW

LB1

7, A2

/E7

Poor

5M/6F

NoYe

sAb

plan

alpA/

E blo

od ty

pes 2

/7.UC

D 38

0Co

ngen

ic wi

th UC

D 00

3SC

WL

B17,

A4/E

7Po

or5M

/6FNo

Yes

Abpl

analp

A/E

blood

type

s 4/7;

resis

tant to

Rou

s sar

coma

.UC

D 38

6Co

ngen

ic wi

th UC

D 00

3SC

WL

BR4/

R4Po

or5M

/6FNo

Yes

Abpl

analp

MHC:

BR4

/R4 (

B15/

B21 r

ecom

binan

t); sh

own t

o be

resis

tant to

Rou

s sar

coma

and M

arek

’s dis

ease

viru

ses.

UCD

387

Cong

enic

with

UCD

003

SCW

LBR

5/R5

Poor

5M/6F

NoYe

sAb

plan

alpMH

C: B

R5/R

5 (B2

1/B1

5 rec

ombin

ant);

show

n to b

esu

scep

tible

to Ro

us sa

rcoma

and M

arek

’s dis

ease

virus

es.

UNH

6.15-

5Co

ngen

ic wi

th UN

H 6-

1SC

WL

B5Go

od10

M/40

FNo

Yes

Taylo

rInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B5.

UNH

6.6-2

Cong

enic

with

UNH

6-1

SCW

LB2

Good

10M/

40F

NoYe

sTa

ylor

Inbre

eding

coeff

icien

t (F)

>0.99

9; MH

C: B

2.UN

H-UC

D 00

3Ac

quire

d fro

m U

Califo

rnia-

Davis

inSC

WL

B17

Good

10M/

40F

NoYe

sTa

ylor

Inbre

eding

coeff

icien

t (F)

>0.99

9; MH

C: B

17.

1986

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

5

Chi

cken

(con

t.)B

lood

type

-MH

C-In

bred

(con

t.)UN

H-UC

D 00

3.R1

Cong

enic

with

UCD

003

SCW

LBF

24-B

G23

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF2

4-BG

23 (B

comp

lex re

comb

inant)

;ba

ckcro

ssed

eigh

t time

s to U

CD-0

03.

UNH-

UCD

003.R

2Co

ngen

ic wi

th UC

D 00

3SC

WL

BF2-

BG23

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF2

-BG2

3 (B

comp

lex re

comb

inant)

; bac

kcro

ssed

eight

times

to U

CD-0

03.

UNH-

UCD

003.R

3Co

ngen

ic wi

th UC

D 00

3SC

WL

BF2-

BG23

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF2

-BG2

3 (B

comp

lex re

comb

inant)

; bac

kcro

ssed

eight

times

to U

CD-0

03.

UNH-

UCD

003.R

4Co

ngen

ic wi

th UC

D 00

3SC

WL

BF2-

BG23

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF2

-BG2

3 (B

comp

lex re

comb

inant)

; bac

kcro

ssed

eight

times

to U

CD-0

03.

UNH-

UCD

003.R

5Co

ngen

ic wi

th UC

D 00

3SC

WL

BF21

-BG1

9Go

od10

M/40

FNo

Yes

Taylo

rMH

C: B

F21-

BG19

(B co

mplex

reco

mbina

nt);

back

cross

ed ei

ght ti

mes t

o UCD

-003

.UN

H-UC

D 00

3.R6

Cong

enic

with

UCD

003

SCW

LBF

21-B

G23

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF2

1-BG

23 (B

comp

lex re

comb

inant)

;ba

ckcro

ssed

eigh

t time

s to U

CD-0

03.

UNH-

UCD

386

Acqu

ired f

rom

U Ca

liforn

ia-Da

vis in

SCW

LBF

15-B

G21

Good

10M/

40F

NoYe

sTa

ylor

MHC:

BF1

5-BG

21 (B

comp

lex re

comb

inant)

.19

95; c

onge

nic w

ith U

CD 00

3UN

H-UC

D 38

7Ac

quire

d fro

m U

Califo

rnia-

Davis

inSC

WL

BF21

-BG1

5Go

od10

M/40

FNo

Yes

Taylo

rMH

C: B

F21-

BG15

(B co

mplex

reco

mbina

nt).

1995

; con

genic

with

UCD

003

UNH-

UCD

3C.1

Cong

enic

with

UCD

003

SCW

LB1

7Go

od10

M/40

FNo

Yes

Taylo

rMH

C: B

17, w

ith re

comb

ined B

comp

lex.

Chr

omos

omal

var

iant

Otta

wa B

-19/B

-19 M

13Al

so re

ferre

d to a

s Otta

wa B

19SC

WL

B19

No liv

e0

Blas

todisc

cells

With

Etch

esMH

C: B

19; s

egre

gatin

g for

DNA

band

ing pa

ttern

s usin

gbir

dspe

rmiss

ionM1

3 pha

ge.

Otta

wa B

-21/B

-21 M

13Al

so re

ferre

d to a

s Otta

wa B

21SC

WL

B21

No liv

e0

Blas

todisc

cells

With

Etch

esMH

C: B

21; s

egre

gatin

g for

DNA

band

ing pa

ttern

s usin

gbir

dspe

rmiss

ionM1

3 pha

ge.

UCD-

Corn

ell M

ono-

PNU

Acqu

ired f

rom

Corn

ell in

1995

SCW

LGo

od12

0No

With

Delan

yEm

bryo

s hom

ozyg

ous f

or a

large

delet

ion on

the M

HCco

llabo

ratio

nch

romo

some

(16)

lack

mos

t of th

e rDN

A ge

nes a

nd di

eea

rly in

embr

yoge

nesis

; hete

rozy

gotes

deve

lop no

rmall

yan

d are

viab

le.UC

D-Co

rnell

Tris

omic

Acqu

ired f

rom

Corn

ell in

1995

SCW

LGo

od12

0No

With

Delan

yTr

isomy

or te

traso

my of

the M

HC-co

ntaini

ngco

llabo

ratio

nch

romo

some

(16)

; suc

h ane

uploi

ds ca

n hatc

h and

reac

hma

turity

, but

are o

ften s

mall a

nd ha

ve po

or pr

oduc

tion

char

acter

istics

.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

6

Chi

cken

(con

t.)C

hrom

osom

al v

aria

nt (c

ont.)

Wisc

onsin

Chr

omos

omal

Rear

rang

emen

tsKe

pt in

gene

tic ba

ckgr

ound

ofNe

w Ha

mpsh

ireFa

ir:4M

/6FNo

With

Bitg

ood

Six c

hrom

osom

al tra

nsloc

ation

s and

two i

nver

sions

Wisc

onsin

New

Ham

pshir

eins

titutio

nal

colla

bora

tion

involv

ing ch

romo

some

s 1 th

roug

h 4 an

d Z.

Endo

geno

us v

irus

RPRL

15B1

Spec

ific pa

thoge

n-fre

eSC

WL

B5, B

15,

Good

:6M

/42F

Seme

nYe

sBa

con

Non-

inbre

d; MH

C: B

5 and

B15

; sus

cepti

ble to

avian

ALVE

1US

DAleu

kosis

viru

ses A

, B &

C an

d Mar

ek’s

disea

se vi

rus;

conta

ins en

doge

nous

viru

s ev1

.

Endo

geno

us v

irus-

Inbr

edRP

RL 10

0BCo

ngen

ic wi

th RP

RL 7I

2; sp

ecific

SCW

LB2

, ALV

E1,

Good

:6M

/42F

Seme

nYe

sBa

con

Inbre

eding

coeff

icien

t (F)

>0.99

9; MH

C: B

2; en

doge

nous

patho

gen-

free

ALVE

3US

DAvir

uses

ev1 a

nd ev

3; su

scep

tible

to av

ian le

ukos

isvir

uses

B, C

, and

E, a

nd M

arek

’s dis

ease

viru

s.RP

RL 6I

3Sp

ecific

patho

gen-

free

SCW

LB2

, ALV

E1,

Good

:12

M/84

FSe

men

Yes

Baco

nInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B2;

endo

geno

usAL

VE3

USDA

virus

es ev

1 and

ev3;

susc

eptib

le to

avian

leuk

osis

virus

es A

,B, &

C, an

d res

istan

t to M

arek

’s dis

ease

viru

s.RP

RL 7I

2Sp

ecific

patho

gen-

free

SCW

LB2

, ALV

E1,

Good

:12

M/84

FSe

men

Yes

Baco

nInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B2;

endo

geno

usAL

VE3

USDA

virus

es ev

1 and

ev3;

susc

eptib

le to

avian

leuk

osis

virus

e C an

d Mar

ek’s

disea

se vi

rus.

RPRL

Rea

sehe

ath

Line

CSp

ecific

patho

gen-

free

SCW

LB1

2,Go

od:

6M/42

FSe

men

Yes

Baco

nInb

reed

ing co

effici

ent (

F)>0

.999;

MHC:

B12

; end

ogen

ous

ALVE

1,US

DAvir

uses

ev1,

ev7,

ev10

; sus

cepti

ble to

leuk

osis

B an

d C.

ALVE

7,AL

VE10

Inbr

edAD

OL 6C

.7A R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7B R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7C R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7D R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

7

Chi

cken

(con

t.)In

bred

(con

t.)AD

OL 6C

.7F R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7G R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7I R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7J R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7K R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7L R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7M R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7N R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7P R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7R R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7S R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

8

Chi

cken

(con

t.)In

bred

(con

t.)AD

OL 6C

.7T R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7V R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7W R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).AD

OL 6C

.7X R

eCon

12%

RPR

L 7I2,

88%

RPR

L 6I3

SCW

LB2

Good

:4M

/10F

NoYe

sBa

con

One o

f 19 d

iffere

nt re

comb

inant

lines

(prim

arily

cong

enic

USDA

with

RPRL

6I3)

, eac

h sub

line w

ith 12

.5% of

geno

mefro

m RP

RL 7I

2 (su

bline

s ide

ntifie

d by l

etter

s).Ot

tawa

GF

Deriv

ed fr

om O

ttawa

3SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esInb

reed

ing co

effici

ent (

F)>0

.8.bir

dspe

rmiss

ionOt

tawa

GH

Deriv

ed fr

om O

ttawa

3SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esInb

reed

ing co

effici

ent (

F)>0

.8.bir

dspe

rmiss

ionOt

tawa

M2

Deriv

ed fr

om O

ttawa

4SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esInb

reed

ing co

effici

ent (

F)>0

.7.bir

dspe

rmiss

ionOt

tawa

WG

Deriv

ed fr

om O

ttawa

8SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esInb

reed

ing co

effici

ent (

F)>0

.7.bir

dspe

rmiss

ionOt

tawa

XP

Deriv

ed fr

om O

ttawa

9SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esInb

reed

ing co

effici

ent (

F)>0

.7.bir

dspe

rmiss

ionUC

D 00

1On

e of th

e refe

renc

e line

s use

d in

Red J

ungle

Good

50M/

50F

NoW

ithDe

lany

Inbre

eding

coeff

icien

t (F)

>0.8;

from

wild

-type

jung

le fow

l.the

Eas

t Lan

sing C

hicke

n Gen

ome

Fowl

colla

bora

tion

Mapp

ing P

rojec

tW

iscon

sin A

ncon

aKe

pt as

clos

ed flo

ck fo

r An

cona

Fair:

4M/15

FNo

With

Bitg

ood

Inbre

eding

coeff

icien

t (F)

>0.9;

half-s

ibling

inbr

eedin

gmo

re th

an 20

gene

ratio

nsins

titutio

nal

colla

bora

tion

since

the m

id-19

40s.

Wisc

onsin

Leg

horn

line U

W-S

p2De

rived

from

1940

/1950

Can

adian

SCW

LFa

ir:4M

/15F

NoW

ithBi

tgoo

dInb

reed

ing co

effici

ent (

F)>0

.9; ha

lf-sibl

ing in

bree

ding

Spru

celei

gh st

rains

institu

tiona

lco

llabo

ratio

nsin

ce th

e mid-

1940

s.

Mut

ant-C

olor

, fea

ther

Wisc

onsin

Aut

osom

al Al

bino

Acqu

ired f

rom

USC

WL

Autos

omal

rece

ssive

.ca

Fair:

4M/12

FNo

With

Bitg

ood

Homo

zygo

tes ha

ve w

hite f

eathe

rs an

d red

eyes

.Ma

ssac

huse

tts-A

mher

st in

1997

institu

tiona

lco

llabo

ratio

n(U

Mass

Auto

soma

l Albi

no)

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

9

Chi

cken

(con

t.)M

utan

t-Dev

elop

men

tal d

efec

t-Eye

UBC

RCFo

und i

n Minn

esota

Rho

de Is

land

Rhod

e Isla

ndAu

tosom

al re

cess

ive.

rc or

rdPo

or12

M/40

FNo

Yes

Chen

gRe

tinal

dege

nera

tion h

omoz

ygote

s do n

ot for

m ro

ds or

Red i

n 198

0; ba

ckcro

ssed

year

ly to

Red

cone

s in t

he re

tina,

caus

ing bl

indne

ss.

UBC-

Minn

esota

Rho

de Is

land R

edUM

ass P

ink E

yeAl

so ha

s dom

inant

exten

sion o

fMi

xed

Autos

omal

rece

ssive

.pk

Disp

ersa

l25

NoYe

sSm

yth

Pink

-eye

homo

zygo

tes ha

ve pi

nk ey

es an

d a hi

ghme

lanin

(E)

in 19

98inc

idenc

e of c

atara

cts; fe

ather

color

is al

so di

luted

.W

iscon

sin B

lind/

Cata

ract

Kept

in ge

netic

back

grou

nd of

SCW

LAu

tosom

al re

cess

ivebc

Fair:

6M/12

FNo

With

Bitg

ood

Homo

zygo

us ch

icks a

re bl

ind at

hatch

, with

visib

leW

iscon

sin S

CWL

semi

-viab

le.ins

titutio

nal

colla

bora

tion

catar

acts.

Wisc

onsin

Pin

k-ey

eAc

quire

d fro

m U

Mixe

dAu

tosom

al re

cess

ive.

pkPo

or4M

/12F

NoYe

sBi

tgoo

dHo

mozy

gotes

have

pink

eyes

and a

high

incid

ence

ofMa

ssac

huse

tts-A

mher

st in

1998

catar

acts;

feath

er co

lor is

dilut

ed, e

ven w

ith do

mina

ntex

tensio

n of m

elanin

(E).

Wisc

onsin

Pop

-eye

Kept

in W

iscon

sin N

ew H

amps

hire

New

Hamp

shire

Sex-l

inked

rece

ssive

.po

pFa

ir:6M

/12F

NoW

ithBi

tgoo

dHo

mozy

gotes

have

a co

nical

protr

usion

of th

e cor

nea

gene

tic ba

ckgr

ound

institu

tiona

lco

llabo

ratio

n(ke

ratoc

onus

) firs

t see

n at 5

wk p

ostha

tch.

Mut

ant-D

evel

opm

enta

l def

ect-F

ace/

limb

Stor

rs C

hond

rody

stro

phy

Mutat

ion re

porte

d in S

CWL i

n 194

2;SC

WL

Autos

omal

rece

ssive

chPo

or5M

/19F

NoYe

sPi

erro

Homo

zygo

tes ha

ve sh

ort u

pper

beak

s, sh

orten

ed lo

ngke

pt at

Stor

rs sin

ce la

te 19

40s

embr

yonic

letha

l.bo

nes,

and b

ent ti

biae.

Stor

rs C

reep

erHe

ld at

Stor

rs sin

ce th

e 192

0s;

Mixe

d Ros

eR

and P

t are

autos

omal

Cp, R

, Pt

Poor

10M/

39F

NoYe

sPi

erro

Heter

ozyg

otes h

ave s

horte

ned l

imbs

, and

homo

zygo

tesmu

tation

pres

ent in

a few

exhib

ition

comb

WL

domi

nants

; cp i

s an

die be

fore h

atch;

stock

also

carri

es ro

se co

mb (R

) and

bree

dsau

tosom

al inc

omple

tepti

lopod

y (leg

feath

ering

: Pt).

domi

nant,

letha

l inho

mozy

gotes

.St

orrs

Dip

lopo

dia-

3Mu

tation

repo

rted i

n inb

red S

CWL i

nMi

xed S

CWL

Autos

omal

rece

ssive

dp-3

Poor

5M/22

FNo

Yes

Pier

roHo

mozy

gotes

disp

lay m

oder

ate pr

eaxia

l poly

dacty

ly,19

72; a

cquir

ed fr

om U

embr

yonic

letha

l.dw

arfin

g, ex

pose

d visc

era,

and s

horte

ned u

pper

beak

.Sa

skatc

hewa

n in t

he 19

60s

Stor

rs D

iplo

podi

a-5

Mutat

ion re

porte

d in 1

983;

acqu

ired

Mixe

d SCW

LAu

tosom

al re

cess

ivedp

-5Po

or5M

/21F

NoYe

sPi

erro

Homo

zygo

tes di

splay

mod

erate

prea

xial p

olyda

ctyly,

from

U Sa

skatc

hewa

n in l

ate 19

80s

embr

yonic

letha

l.dw

arfin

g, ex

pose

d visc

era,

and s

horte

ned u

pper

beak

.St

orrs

Lim

bles

sMu

tation

repo

rted i

n 197

9 in a

flock

Mixe

d SCW

LAu

tosom

al re

cess

ivell

Poor

5M/20

FNo

Yes

Pier

roHo

mozy

gotes

do no

t form

limb b

uds o

r limb

s, an

dof

Rhod

e Isla

nd R

eds;

acqu

ired f

rom

embr

yonic

letha

l.us

ually

have

a sh

orten

ed up

per b

eak.

U W

iscon

sin in

late

1980

sSt

orrs

Micr

omeli

a-Ab

bott

Mutat

ion re

porte

d in c

rook

ed-n

eck

Mixe

d SCW

LAu

tosom

al re

cess

ivem

m-A

Poor

5M/17

FNo

Yes

Pier

roHo

mozy

gotes

have

disti

nctly

shor

tened

long

bone

s.dw

arf s

tock f

rom

U Ca

liforn

ia-Da

visem

bryo

nic le

thal.

Stor

rs M

icrom

elia-

Hays

Mutat

ion re

porte

d in R

hode

Islan

dSC

WL

Autos

omal

rece

ssive

mm

-HPo

or5M

/20F

NoYe

sPi

erro

Homo

zygo

tes ha

ve di

stinc

tly sh

orten

ed lo

ng bo

nes.

Red s

tock i

n 194

4; he

ld at

Stor

rsem

bryo

nic le

thal.

since

1950

sSt

orrs

Nan

omeli

aMu

tation

repo

rted i

n SCW

L stoc

k by

Mixe

d SCW

LAu

tosom

al re

cess

ivenm

, Pt

Poor

5M/20

FNo

Yes

Pier

roHo

mozy

gotes

have

seve

rely

shor

tened

long

bone

s in t

heLa

ndau

er in

1965

; held

at S

torrs

embr

yonic

letha

l.lim

bs; m

ay al

so ca

rry th

e muta

nt all

ele fo

r leg-

feathe

ring

since

then

; also

has p

tilopo

dy(P

t).

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

10

Chi

cken

(con

t.)M

utan

t-Dev

elop

men

tal d

efec

t-Fac

e/lim

b (c

ont.)

Stor

rs P

eroc

epha

lyMu

tation

repo

rted i

n floc

k of S

CWL

Mixe

d SCW

LAu

tosom

al re

cess

ivepe

rPo

or5M

/22F

NoYe

sPi

erro

Homo

zygo

tes sh

ow va

riable

effec

ts, fr

om ab

norm

alby

Land

auer

in 19

56; h

eld at

Stor

rsem

bryo

nic le

thal w

ithup

per b

eaks

to m

icroc

epha

ly, sy

noph

thalm

ia, an

dsin

ce th

enlow

pene

tranc

e.cy

clopia

.St

orrs

Pol

ydac

tyly

Mutat

ion lo

ng kn

own i

n exh

ibitio

nMi

xed S

CWL

Autos

omal

incom

plete

PoPo

or5M

/22F

NoYe

sPi

erro

Homo

zygo

tes an

d hete

rozy

gotes

may

have

one o

r two

stock

s and

repo

rted i

n sev

eral

domi

nant.

extra

prea

xial to

es, o

r a lo

nger

-than

-nor

mal fi

rst di

git.

ancie

nt br

eeds

Stor

rs T

alpid

-2Mu

tation

repo

rted i

n SCW

L stoc

k in

SCW

LAu

tosom

al re

cess

iveta

-2Po

or2M

/10F

NoYe

sPi

erro

Homo

zygo

tes di

splay

extre

me pr

eaxia

l poly

dacty

ly (u

p19

59; a

cquir

ed fr

om U

Cali

fornia

embr

yonic

letha

l.to

10 di

gits p

er lim

b), o

vera

ll dwa

rfing,

expo

sed v

iscer

a,Da

vis in

1996

and c

left p

alate

with

a par

rot-li

ke up

per b

eak.

Stor

rs W

ingl

ess-

2Mu

tation

repo

rted i

n floc

k of S

CWL

SCW

LAu

tosom

al re

cess

ivewg

-2Po

or5M

/22F

NoYe

sPi

erro

Homo

zygo

tes la

ck w

ings,

have

redu

ced l

egs,

by La

ndau

er in

1956

; held

at S

torrs

embr

yonic

letha

l.ba

lloon

-like f

eathe

rs, sh

ort u

pper

beak

, clef

t pala

te an

dsin

ce th

ensm

all ki

dney

s.UC

D Cl

eft P

rimar

y Pala

te/S

calel

ess

Deriv

ed fr

om cr

osse

s of: S

CWL,

Mixe

dAu

tosom

al re

cess

ivecp

p, sc

Poor

5M/5F

Seme

nYe

sAb

bott

Clef

t prim

ary p

alate

(cpp

) hom

ozyg

otes l

ack m

ost o

f the

UCD

Scale

less-H

igh an

d UCD

embr

yonic

letha

l (cpp

);up

per b

eak,

may a

lso ha

ve lim

b red

uctio

ns if

Scale

less-L

ow; c

pp w

as or

igina

llyau

tosom

al re

cess

iveho

mozy

gous

for s

calel

ess;

for sc

aleles

s (sc

): sc

aleles

sca

lled e

ctrod

actyl

y whe

n fou

nd in

se

mi-vi

able

(sc).

homo

zygo

tes do

not fo

rm sp

urs,

scale

s or m

ost

UCD

Scale

less s

tocks

in 19

66fea

thers.

UCD

Colo

bom

a X 00

3Co

ngen

ic wi

th UC

D 00

3; mu

tation

SCW

LAu

tosom

al re

cess

ivecm

Poor

5MSe

men

Yes

Abbo

ttHe

mizy

gous

fema

les ar

e mod

erate

ly to

seve

rely

dwar

fedfirs

t rep

orted

in 19

70 at

Uem

bryo

nic le

thal.

with

mildl

y to s

ever

ely cl

eft pa

lates

; som

e are

lack

ingCa

liforn

ia-Da

vispr

eaxia

l digi

ts or

have

truc

ated w

ings a

nd le

gs; s

ome

are e

demi

c; the

expr

essio

n may

be hi

ghly

varia

ble, e

ven

with

the sa

me pa

rents

.UC

D Di

plop

odia-

1 X 00

3Ne

ar-co

ngen

ic wi

th UC

D 00

3;SC

WL

Autos

omal

rece

ssive

dp-1

Poor

5M/5F

Seme

nYe

sAb

bott

Homo

zygo

tes di

splay

mod

erate

prea

xial p

olyda

ctyly,

mutat

ion fir

st re

porte

d in 1

947 a

t Uem

bryo

nic le

thal.

dwar

fing,

expo

sed v

iscer

a, an

d sho

rtene

d upp

er be

ak.

Califo

rnia-

Berke

leyUC

D Di

plop

odia-

3 X 00

3Ne

ar-co

ngen

ic wi

th UC

D 00

3;SC

WL

Autos

omal

rece

ssive

dp-3

Poor

5M/5F

NoYe

sAb

bott

Homo

zygo

tes di

splay

mod

erate

prea

xial p

olyda

ctyly,

mutat

ion fir

st re

porte

d in 1

972 a

t Uem

bryo

nic le

thal.

dwar

fing,

some

with

expo

sed v

iscer

a, oc

casio

nal c

left

Califo

rnia-

Berke

leypa

late,

and s

horte

ned u

pper

beak

.UC

D Do

nald

-duc

k Bea

k/Sca

leles

sRe

porte

d in 1

967 i

n New

Mixe

dAu

tosom

al re

cess

ivedd

-2, s

c,Po

or4M

/3FSe

men

Yes

Abbo

ttDo

nald

duck

bea

k hom

ozyg

otes h

ave u

pwar

d cur

ledHa

mpsh

ire-ty

pe ch

icken

s at U

embr

yonic

letha

ls (cp

pma

y also

uppe

r bea

ks, m

ay ha

ve do

wnwa

rd cu

rled l

ower

beak

s;Ca

liforn

ia-Da

visan

d dd-

2); a

utoso

mal

includ

e cpp

some

hatch

, but

do no

t sur

vive l

ong;

no kn

own

rece

ssive

semi

-viab

leint

erac

tion w

ith sc

aleles

s; for

sc, s

ee: U

CD(sc

).Sc

aleles

s-Low

; may

also

carry

clef

t prim

ary p

alate

(cpp)

,se

e: UC

D Cl

eft P

rimar

y Pala

te/Sc

aleles

s.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

11

Chi

cken

(con

t.)M

utan

t-Dev

elop

men

tal d

efec

t-Fac

e/lim

b (c

ont.)

UCD

Eudi

plop

odia

X 00

3Co

ngen

ic wi

th UC

D 00

3; mu

tation

SCW

LAu

tosom

al re

cess

iveeu

Poor

5M/5F

Seme

nYe

sAb

bott

Homo

zygo

tes ha

ve ex

tra di

gits o

n the

dorsa

l sur

faces

offirs

t rep

orted

1959

in S

CWL f

lock a

tem

bryo

nic le

thal.

the lim

b bud

s, wh

ich de

velop

into

extra

, bido

rsal to

esU

Califo

rnia-

Berke

leyon

the f

eet a

nd oc

casio

nal d

orsa

l kno

bs or

digit

s on t

hewi

ngs.

UCD

Lim

bles

s X 00

3Ne

ar-co

ngen

ic wi

th UC

D 00

3;SC

WL

Autos

omal

rece

ssive

llPo

or5M

/5FNo

Yes

Abbo

ttHo

mozy

gotes

do no

t form

limb b

uds o

r limb

s, an

dmu

tation

first

repo

rted i

n 197

9 at U

embr

yonic

letha

l.us

ually

have

a sh

orten

ed up

per b

eak.

Wisc

onsin

UCD

Polyd

acty

ly X

003

Near

-cong

enic

with

UCD

003;

mutan

tSC

WL

Autos

omal

domi

nant

PoPo

or3M

/3FSe

men

Yes

Abbo

ttHo

mozy

gous

and h

etero

zygo

us m

utants

may

have

one

allele

comm

on in

exhib

ition

with

incom

plete

or tw

o extr

a pre

axial

toes

, or a

long

er-th

an-n

orma

l first

chick

ens

pene

tranc

e.dig

it.UC

D St

umpy

X 00

3Co

ngen

ic wi

th UC

D 00

3; mu

tation

SCW

LAu

tosom

al re

cess

ivestu

Poor

5M/5F

Seme

nYe

sAb

bott

Homo

zygo

tes ha

ve co

nical

leg bu

ds an

d a po

orly

tore

porte

d at U

Cali

fornia

-Dav

is in

embr

yonic

letha

l.no

n-va

scula

rized

allan

tois t

hat n

ever

gets

large

r tha

n the

1966

in N

ew H

amps

hire X

Cor

nish

head

; emb

ryo de

ath, ty

picall

y betw

een t

he fif

th an

dse

venth

day,

is as

socia

ted w

ith m

assiv

e mult

iple

hemo

rrhag

es.

UCD

Talp

id-2

X 00

3Co

ngen

ic wi

th UC

D 00

3; mu

tation

SCW

LAu

tosom

al re

cess

iveta

-2Po

or5M

/5FSe

men

Yes

Abbo

ttHo

mozy

gotes

disp

lay ex

treme

prea

xial p

olyda

ctyly,

first r

epor

ted in

1959

at U

embr

yonic

letha

l.dw

arfin

g, ex

pose

d visc

era,

and c

left, p

arro

t-like

uppe

rCa

liforn

ia-Da

vis in

a SC

WL f

lock

beak

.UC

D W

ing-

redu

ced

X 00

3Sy

ndro

me fo

und i

n UCD

413

SCW

LMu

ltifac

torial

, not

yet

Poor

3M/3F

NoYe

sAb

bott

A co

mplex

synd

rome

; affe

cted b

irds l

ack t

oena

ils an

d(m

uscu

lar dy

strop

hy) in

1990

;ful

ly de

fined

.so

me ph

lange

s (us

ually

on di

git 3,

some

times

digit

4)cro

ssed

seve

ral ti

mes t

o UCD

003

and m

ay ha

ve tr

unca

ted w

ings.

UCD

Win

gles

s-2 X

331

Cong

enic

with

UCD

331;

mutat

ionSC

WL

Autos

omal

rece

ssive

wg-2

Poor

5M/5F

Seme

nYe

sAb

bott

Homo

zygo

tes la

ck w

ings,

have

redu

ced l

egs,

first r

epor

ted in

1956

at U

embr

yonic

letha

l.ba

lloon

-like f

eathe

rs, sh

ort u

pper

beak

, clef

t pala

te an

dCo

nnec

ticut-

Stor

rssm

all m

etane

phric

kidn

eys.

Wisc

onsin

Am

etap

odia

Acqu

ired f

rom

UMi

xed

Autos

omal

domi

nant,

Mp

Fair:

4M/12

FNo

With

Bitg

ood

Heter

ozyg

otes l

ack m

etatar

sals

in the

legs

and

Mass

achu

setts

-Amh

erst

in 19

97let

hal in

homo

zygo

tes.

institu

tiona

lco

llabo

ratio

nme

tacar

pels

in the

wing

s; ho

mozy

gotes

usua

lly di

e(U

Mass

Ame

tapod

ia)ea

rly in

deve

lopme

nt.W

iscon

sin L

imbl

ess

Kept

in W

iscon

sin Le

ghor

n gen

etic

SCW

LAu

tosom

al re

cess

ivell

Fair:

6M/20

FNo

With

Bitg

ood

Homo

zygo

tes do

not fo

rm lim

b bud

s or li

mbs,

and

back

grou

ndem

bryo

nic le

thal.

institu

tiona

lco

llabo

ratio

nso

metim

es ha

ve a

shor

t upp

er be

ak.

Wisc

onsin

Talp

id-2

Mutat

ion fir

st re

porte

d in 1

959 a

t USC

WL

Autos

omal

rece

ssive

ta-2

Fair:

6M/12

FNo

With

Bitg

ood

Homo

zygo

tes di

splay

extre

me pr

eaxia

l poly

dacty

ly,Ca

liforn

ia-Da

vis; a

cquir

ed in

1990

s;em

bryo

nic le

thal.

institu

tiona

lco

llabo

ratio

ndw

arfin

g, ex

pose

d visc

era,

and c

left, p

arro

t-like

uppe

rke

pt in

Wisc

onsin

Legh

orn g

eneti

cbe

ak.

back

grou

ndW

iscon

sin W

ingl

ess-

2Mu

tation

first

repo

rted i

n 195

6 at U

SCW

LAu

tosom

al re

cess

ivewg

-2Fa

ir:6M

/12F

NoW

ithBi

tgoo

dHo

mozy

gotes

lack

wing

s, ha

ve re

duce

d leg

s,Co

nnec

ticut-

Stor

rs; no

w ke

pt in

embr

yonic

letha

l.ins

titutio

nal

colla

bora

tion

ballo

on-lik

e fea

thers,

shor

t upp

er be

ak, c

left p

alate

and

Wisc

onsin

Legh

orn g

eneti

csm

all ki

dney

s.ba

ckgr

ound

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

12

Chi

cken

(con

t.)M

utan

t-Dev

elop

men

tal d

efec

t-Ski

n/fe

athe

rSt

orrs

Otta

wa N

aked

Mutat

ion fir

st re

porte

d by R

. Cra

wfor

dNe

w Ha

mpsh

ireAu

tosom

al re

cess

ive.

nkPo

or3M

/17F

NoYe

sPi

erro

Homo

zygo

tes ar

e nak

ed at

hatch

with

occa

siona

lin

1982

; acq

uired

from

Usy

ndac

tyly o

f toes

2 an

d 3; s

ome f

eathe

rs ma

y for

m on

Sask

atche

wan i

n late

1980

sad

ults.

Stor

rs P

tilop

ody

Mutat

ion co

mmon

in ex

hibitio

nMi

xed S

CWL

Autos

omal

domi

nant

PtCo

mbine

dse

e: St

orrs

NoYe

sPi

erro

Homo

zygo

tes an

d hete

rozy

otes h

ave f

eathe

rs re

placin

gbr

eeds

; intro

duce

d into

Stor

rswi

th mu

ltifac

torial

with

Stor

rsNa

nome

liaso

me of

the s

cales

on lo

wer le

g, foo

t, and

toes

.Na

nome

lia w

ith B

uff C

ochin

cros

s;mo

difier

s.Na

nome

liaals

o in S

torrs

Cree

per;

and

Cree

per

Stor

rs S

calel

ess

Mutat

ion re

porte

d in N

ewMi

xed N

ewAu

tosom

al re

cess

ivesc

Poor

10M/

30F

NoYe

sPi

erro

Homo

zygo

tes do

not fo

rm sp

urs,

scale

s or m

ost fe

ather

s.Ha

mpsh

ire-ty

pe st

ock i

n 195

7;Ha

mpsh

irese

mi-vi

able.

acqu

ired f

rom

U Ca

liforn

ia-Da

vis in

1965

UCD

Ichth

yosis

X 00

3Ne

ar-co

ngen

ic wi

th UC

D 00

3;SC

WL

Autos

omal

rece

ssive

dehy

Poor

5M/5F

Seme

nYe

sAb

bott

Ichth

yosis

(deh

y) ho

mozy

gotes

deve

lop la

mella

rmu

tation

first

repo

rted i

n 195

7 at U

semi

-viab

le.ich

thyos

is: ch

icks d

ehyd

rate

rapid

ly aft

er ha

tch, a

ndCa

liforn

ia-Be

rkeley

have

char

acter

istic

string

y/gre

asy d

own f

eathe

rs; ad

ults

do no

t slou

gh le

g sca

les, a

nd de

velop

chro

nic sk

inles

ions.

UCD

Nake

d-ne

ckMu

tation

comm

on in

exhib

ition

Mixe

d SCW

LAu

tosom

al do

mina

nt.Na

No liv

e0

Seme

nW

ithAb

bott

Caus

es a

redu

ction

in si

ze of

feath

er tr

acts

on th

e hea

d,br

eeds

and i

n som

e com

merci

albir

dspe

rmiss

ionne

ck an

d ven

tral tr

unk,

usua

lly gi

ving t

he bi

rd a

bare

orstr

ains

near

ly ba

re ne

ck.

UCD

Scale

less-

High

Mutat

ion fir

st re

porte

d at U

Mixe

dAu

tosom

al re

cess

ivesc

Poor

15M/

15F

Seme

nYe

sAb

bott

Scale

less-h

igh ho

mozy

gotes

do no

t dev

elop s

cales

orCa

liforn

ia-Da

vis in

1957

; orig

inally

semi

-viab

le, w

ithsp

urs,

but s

electi

on fo

r incre

ased

feath

er de

velop

ment

non-

viable

, but

cross

bree

ding a

ndmu

ltifac

torial

mod

ifiers.

has r

esult

ed in

abun

dant

down

-like f

eathe

rs for

ming

onse

lectio

n dra

matic

ally i

ncre

ased

the bo

dy an

d spa

rse fe

ather

s for

ming

on th

e leg

s.via

bility

; bac

kgro

und b

reed

s inc

lude

Corn

ish, S

CWL,

Barre

d Roc

k, Ne

wHa

mpsh

ire, R

ed Ju

ngle

Fowl

UCD

Scale

less-

Low

Mutat

ion fir

st re

porte

d at U

Mixe

dAu

tosom

al re

cess

ivesc

Poor

15M/

15F

Seme

nYe

sAb

bott

Scale

less-l

ow ho

mozy

gotes

do no

t dev

elop s

purs,

Califo

rnia-

Davis

in 19

57; o

rigina

llyse

mi-vi

able,

with

scale

s or m

ost fe

ather

s, a c

ondit

ion th

at wa

s enh

ance

dno

n-via

ble, b

ut cro

ssbr

eedin

g and

multif

actor

ial m

odifie

rs.by

selec

tion f

or lo

w fea

ther n

umbe

r.se

lectio

n dra

matic

ally i

ncre

ased

viabil

ity; b

ackg

roun

d bre

eds i

nclud

eCo

rnish

, SCW

L, Ba

rred R

ock,

New

Hamp

shire

, Red

Jung

le Fo

wlW

iscon

sin T

ardy

Fea

ther

ing

Kept

in ge

netic

back

grou

nd of

SCW

LAu

tosom

al re

cess

ive.

tFa

ir:6M

/12F

NoW

ithBi

tgoo

dHo

mozy

gotes

have

very

slow

grow

ing fli

ght fe

ather

sW

iscon

sin Le

ghor

n cro

ssed

with

ins

titutio

nal

colla

bora

tion

after

hatch

, but

this i

s only

detec

table

if chic

ks ar

e also

Wisc

onsin

New

Ham

pshir

eex

pres

sing s

ex-lin

ked r

apid

feathe

r gro

wth (

k+).

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

13

Chi

cken

(con

t.)M

utan

t-Dev

elop

men

tal d

efec

t-Spi

ne/ta

ilSt

orrs

Dom

inan

t Rum

ples

snes

sMu

tation

repo

rted i

n 192

5; he

ld at

SCW

LAu

tosom

al do

mina

nt.Rp

Poor

6M/21

FNo

Yes

Pier

roHo

mozy

gotes

lack

pygo

style,

caud

al ve

rtebr

ae,

Stor

rs sin

ce th

enur

opyg

ial gl

and a

nd ta

il fea

thers;

fertil

ity is

usua

lly lo

w if

natur

ally m

ated.

Stor

rs R

eces

sive R

umpl

ess

Mutat

ion re

porte

d in S

CWL s

tock i

nMi

xed S

CWL

Autos

omal

rece

ssive

rp-2

Poor

10M/

67F

NoYe

sPi

erro

Homo

zygo

tes ha

ve fu

sed o

r abs

ent p

ygos

tyle a

nd19

45; h

eld at

Stor

rs sin

ce th

en; a

lsowi

th inc

omple

teca

udal

verte

brae

, and

some

kyph

osco

liosis

with

extra

has p

eroc

epha

ly (p

er);

kept

pene

tranc

e.rib

s.ho

mozy

gous

for r

p-2

UCD

Scol

iosis

Multif

actor

ial sy

ndro

me un

cove

red i

nMi

xed R

ose

At le

ast tw

o (ma

ybe

No liv

e0

Seme

nW

ithAb

bott

Affec

ted bi

rds s

how

mode

rate

to se

vere

kyph

osco

liosis

the 19

50s d

uring

inbr

eedin

g of

comb

WL

three

) auto

soma

lbir

dspe

rmiss

ionsta

rting a

t 5 w

eeks

, and

clea

rly vi

sible

in x-r

ays a

t 12

stock

s at U

Cali

fornia

-Ber

keley

rece

ssive

gene

s.we

eks;

males

are m

ore s

ever

ely af

fected

than

fema

les.

Mut

ant-G

ene

pool

OSU

Dwar

f Leg

horn

Mutat

ion ke

pt in

comm

ercia

l SCW

LSC

WL

All re

cess

ive; o

nedw

, blr,

bld,

Fair

20M/

50F

NoYe

sSa

vage

Includ

es se

ven m

utatio

ns: s

ex-lin

ked d

warf;

four

back

grou

nd, b

ut as

clos

ed flo

ckse

x-link

ed vi

able

(dw)

,ed

-a, e

d-b

autos

omal

mutat

ions t

hat c

ause

: blas

tode

rmfor

seve

n gen

erati

ons;

four a

utoso

mal le

thal

dege

nera

tion (

bld),

blood

ring

and e

arly

embr

yo de

ath

deve

loped

by P

. Ber

nier

or se

mi-le

thal

(blr)

, and

semi

-letha

l chic

k ede

ma (w

hen h

om. fo

r ed-

a(b

lr,bld,

ed-a

,ed-b

).an

d ed-

b); a

nd tw

o new

muta

tions

that

have

not y

et be

enre

porte

d: tra

nsien

t con

genit

al tre

mor a

nd ec

topia.

UBC-

Minn

esot

a Mar

ker

Acqu

ired f

rom

U Mi

nnes

ota-S

t. Pau

lMi

xed

Gene

pool

of va

rious

Po, P

t, P,

Poor

8M/40

FNo

Yes

Chen

gGe

ne po

ol wi

th 13

domi

nant

mutat

ions:

polyd

actyl

y,do

mina

nt an

dR,

U, M

,pt

ilopo

dy, p

ea an

d ros

e com

b, ur

opyg

ial b

ifurca

tion,

rece

ssive

muta

tions

;Na

, Cr,

S,m

ultipl

e sp

urs,

nake

d nec

k, cre

st, si

lver,

sex-l

inked

see d

escri

ption

.B,

I, M

b, O,

barri

ng, d

omina

nt w

hite,

muf

fs an

d bea

rd, a

nd bl

ue eg

g Cp

, pr

shell

; may

also

conta

in re

cess

ive m

utatio

ns cr

eepe

r (Cp

)an

d pro

topor

phyri

n inh

ibitor

(pr).

UCD

Dipl

opod

ia-3/S

calel

ess H

igh

A re

cent

cross

(199

7) of

UCD

Mixe

dTw

o auto

soma

ldp

-3, s

cPo

or3M

/5FSe

men

Yes

Abbo

ttDi

plopo

dia-3

(dp-

3) m

utants

disp

lay m

oder

ate pr

eaxia

lSc

aleles

s-High

with

UCD

rece

ssive

s: on

epo

lydac

tyly,

dwar

fing,

expo

sed v

iscer

a, oc

casio

nal c

left

Diplo

podia

-3 X

003

embr

yonic

letha

l (dp3

),pa

late,

and s

horte

ned u

pper

beak

; hom

ozyg

ous

one s

emi-v

iable

(sc).

scale

less m

utants

lack

spur

s, sc

ales a

nd m

ost fe

ather

s.UC

D Eu

dipl

opod

ia/Li

mbl

ess

Mildl

y inb

red,

includ

ing so

me U

CDSC

WL

Two a

utoso

mal

eu, ll

Poor

10M/

10F

Seme

nYe

sAb

bott

See U

CD E

udipl

opod

ia an

d UCD

Limb

less d

escri

ption

s.00

3; cro

ssed

with

UCD

rece

ssive

embr

yonic

Scale

less-H

igh an

d -Lo

w in

1996

letha

ls.UC

D Li

mbl

ess/S

tum

pyAn

cestr

al ba

ckgr

ound

inclu

des

SCW

LTw

o auto

soma

lll,

stuPo

or4M

/4FSe

men

Yes

Abbo

ttSe

e UCD

Limb

less a

nd U

CD S

tumpy

desc

riptio

ns.

leuko

sis-fr

ee st

ocks

kept

at U

rece

ssive

embr

yonic

Califo

rnia-

Davis

in th

e 198

0slet

hals.

UCD

Silki

e cro

ssDe

rived

from

UCD

Silk

ie; in

1995

Mixe

d Silk

ieFiv

e auto

soma

l dom

.Cr

, Fm

, h,

Poor

4M/7F

NoYe

sAb

bott

UCD

Silki

e cro

ssed

to U

CD-0

03 or

UCD

Sca

leles

s-low

tocro

ssed

with

both

UCD

003 a

nd U

CD(C

r,Po,P

t,Fm

,Mb)

, one

id, M

b, Po

,de

creas

e inb

reed

ing an

d enh

ance

egg p

rodu

ction

, then

Scale

less-L

ow to

incre

ase v

igor;

sex-l

inked

rec.

(id),

Pt, s

cre

peate

dly ba

ck-cr

osse

d to U

CD S

ilkie

(see U

CD-S

ilkie

back

cross

ed re

peate

dly to

UCD

two a

utoso

mal re

c.for

desc

iption

of m

utatio

ns).

Silki

e(h

,sc).

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

14

Chi

cken

(con

t.)M

utan

t-Gen

e po

ol (c

ont.)

UCD

Talp

id-2

/Win

gles

s-2

Mildl

y inb

red,

includ

ing so

me U

CDSC

WL

Two a

utoso

mal

ta-2

, wg-

2Po

or10

M/10

FSe

men

Yes

Abbo

ttSe

e UCD

Talp

id-2 a

nd U

CD W

ingles

s-2 de

scrip

tions

.00

3; cro

ssed

with

UCD

rece

ssive

embr

yonic

Scale

less-H

igh an

d -Lo

w in

1996

letha

ls.

Mut

ant-I

mm

unol

ogic

al d

efec

tAr

kans

as S

myt

h Li

ne B

101

Acqu

ired f

rom

J. Sm

yth at

UBr

own L

egho

rn,

Multif

actor

ial w

ithGo

od10

M/20

FNo

With

Erf

Selec

ted fo

r high

expr

essio

n of d

elaye

dMa

ssac

huse

tts-A

mher

st in

1996

;sy

ntheti

cva

riable

expr

essiv

ity.

colla

bora

tion

amela

noge

nesis

(auto

immu

ne vi

tiligo

: incre

asing

also k

nown

as D

AM lin

e (de

layed

amou

nt of

white

feath

ers w

ith ea

ch m

olt) a

lso: b

lindn

ess

amela

noge

nesis

)an

d thy

roidi

tis; a

melan

osis

now

seen

>85%

ofoff

sprin

g.Ar

kans

as S

myt

h Li

ne B

102

Acqu

ired f

rom

J. Sm

yth at

UBr

own L

egho

rn,

Multif

actor

ial w

ithGo

od10

M/20

FNo

With

Erf

Selec

ted fo

r high

expr

essio

n of d

elaye

dMa

ssac

huse

tts-A

mher

st in

1998

;sy

ntheti

cva

riable

expr

essiv

ity.

colla

bora

tion

amela

noge

nesis

(auto

immu

ne vi

tiligo

: incre

asing

also k

nown

as D

AM lin

e (de

layed

amou

nt of

white

feath

ers w

ith ea

ch m

olt) a

lso: b

lindn

ess

amela

noge

nesis

)an

d thy

roidi

tis; a

melan

osis

now

seen

>85%

ofoff

sprin

g.Co

rnell

Obe

se (B

-13)

Deriv

ed fr

om C

orne

ll C S

train,

SCW

LMu

ltifac

torial

, with

asB1

3Po

or40

M/12

0FNo

Yes

Mars

hMH

C B1

3; aff

ected

bird

s dev

elop s

ponta

neou

sse

lected

for M

HC ha

plotyp

e B13

;ma

ny as

five m

ajor

autoi

mmun

e thy

roidi

tis an

d, in

female

s, a p

ersis

tant r

ight

close

d floc

k for

over

30 ye

ars;

gene

s.Mu

lleria

n duc

t.pe

digre

ed an

nual

repr

oduc

tion

UCD

200

Mutat

ion fir

st re

porte

d at O

rego

nSC

WL

At le

ast tw

o rec

essiv

esd

Poor

10M/

12F

NoYe

sAb

plan

alpInb

reed

ing co

effici

ent (

F)>0

.80; d

evelo

ps au

toimm

une

State

U in

1961

gene

s and

scler

oder

ma (p

rogr

essiv

e sys

temic

scler

osis)

, with

multif

actor

ial m

odifie

rs.ne

crosis

and s

lough

ing of

the c

ombs

and w

attles

(self-d

ubbin

g), p

olyar

thritis

, and

some

visc

eral

lesion

s.UM

ass S

myt

h Li

neAl

so kn

ow as

the D

AM lin

e (for

Brow

n Leg

horn

,Mu

ltifac

torial

with

Disp

ersa

l20

0No

Yes

Smyt

hDe

layed

amela

nosis

(DAM

) hom

ozyg

otes d

isplay

ade

layed

amela

noge

nesis

); mu

tation

synth

etic

varia

ble ex

pres

sivity

.in

1998

grad

ual lo

ss of

feath

er pi

gmen

t with

succ

essiv

e molt

sor

igina

lly fo

und i

n the

UMa

ss B

rown

(an a

utoim

mune

cond

ition m

imick

ing hu

man v

itiligo

),Lin

ewi

th so

me re

tinal

dystr

ophy

and a

utoim

mune

thyro

iditis

.

Mut

ant-N

euro

logi

cal d

efec

tSa

skat

chew

an E

pilep

tifor

m se

izure

sSy

ntheti

c stra

in, m

ostly

Fay

oumi

,Mi

xed F

ayou

miAu

tosom

al re

cess

iveep

iGo

od10

0M/17

0FNo

With

Clas

sen

Homo

zygo

tes af

flicted

with

seizu

res o

f var

ying s

ever

ityde

rived

from

Agr

icultu

re C

anad

ase

mi-vi

able

mutat

ionco

llabo

ratio

ntho

ugho

ut life

of bi

rd, b

ut go

od vi

abilit

y with

extra

care

;(C

FAR)

stoc

ks in

1963

with

incom

plete

used

as di

seas

e mod

el in

phar

maco

logica

l and

pene

tranc

e.ph

ysiol

ogica

l rese

arch

.UN

L Pa

roxy

smAc

quire

d fro

m R.

Cole

at C

orne

ll U in

SCW

LSe

x-link

ed re

cess

ive.

pxFa

ir: Ha

tch,

5MNo

With

Beck

Post-

hatch

letha

l muta

tion,

caus

ing se

izure

s, slo

wthe

1960

sre

giona

l,co

llabo

ratio

ngr

owth

and e

ventu

al de

ath of

hemi

zygo

us fe

males

.mi

sc.

Wisc

onsin

Piro

uette

Kept

in W

iscon

sin Le

ghor

n gen

etic

SCW

LAu

tosom

al re

cess

ivepir

Fair:

6M/12

FNo

With

Bitg

ood

Homo

zgyo

us ch

icks e

xpre

ssing

this

neur

ologic

alba

ckgr

ound

semi

-viab

le.ins

titutio

nal

colla

bora

tion

mutat

ion w

ill sp

in in

circle

s for

shor

t per

iods.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

15

Chi

cken

(con

t.)M

utan

t-Phy

siol

ogic

al d

efec

tAt

hens

-Can

adian

Dwa

rfsDe

rived

from

Athe

ns-C

anad

ianCo

mmer

cial

Sex-l

inked

rece

ssive

.dw

Good

25M/

50F

NoYe

sBu

rke

Sex-l

inked

dwar

f muta

tion i

n Athe

ns-C

anad

ianRa

ndom

bred

meat

sire c

ross

back

grou

nd.

Corn

ell S

ex-li

nked

Dwa

rfDe

rived

from

the o

ld Ke

mper

SCW

LSC

WL

Sex-l

inked

rece

ssive

.Dw

, B15

Fair

120F

NoW

ithMa

rsh

Homo

zygo

us m

ales a

nd he

mizy

gous

fema

les ar

estr

ain, w

hich i

s also

was

ance

stral

colla

bora

tion

redu

ced i

n bod

y size

; muta

nts ar

e cha

racte

rized

by lo

wto

Corn

ell K

; ped

igree

d yea

rlyse

rum

T3 le

vels

and e

xpre

ssion

of a

defec

tive f

orm

ofre

prod

uctio

nthe

grow

th ho

rmon

e rec

eptor

; MHC

: B15

.Oh

io L

ow-s

core

Nor

mal

Deriv

ed fr

om S

torrs

Musc

ular

Mixe

d SCW

LNo

t yet

defin

ed.

Good

50M/

50F

NoW

ithVe

llem

anAf

fected

bird

s sho

w int

erme

diate

to low

exha

ustio

nDy

strop

hy af

ter an

outcr

oss w

ithco

llabo

ratio

nsc

ores

whe

n sub

jected

to th

e “flip

test”

used

to id

entify

comm

ercia

l SCW

L in 1

970s

musc

ular d

ystro

phy h

omoz

ygote

s; su

ch ex

pres

sion i

sas

socia

ted w

ith an

alter

ed ex

tra-ce

llular

matr

ixor

ganiz

ation

.Oh

io M

uscu

lar D

ystro

phy

Mutat

ion fir

st re

porte

d in 1

956 a

t UMi

xed

Autos

omal

rece

ssive

.am

Good

50M/

50F

NoW

ithVe

llem

anHo

mozy

gotes

deve

lop jo

int st

iffnes

s and

mus

cleCa

liforn

ia-Da

vis; a

cquir

ed fr

om U

colla

bora

tion

weak

ness

; bre

ast a

nd ot

her m

uscle

repla

ced b

y fat

and

Conn

ectic

ut-St

orrs

in 19

97; k

ept a

sco

nnec

tive t

issue

.ho

mozy

gotes

Stor

rs C

rook

ed-n

eck D

warf

Mutat

ion di

scov

ered

in N

ewMi

xed N

ewAu

tosom

al re

cess

ivecn

Poor

5M/22

FNo

Yes

Pier

roHo

mozy

gotes

are e

demi

c and

dwar

fed, w

ith cr

ooke

d,Ha

mpsh

ire-ty

pe ch

icken

stoc

k in

Hamp

shire

embr

yonic

letha

l.fra

gile n

ecks

, thin

amus

cular

legs

, and

no vo

luntar

y19

45; a

cquir

ed fr

om U

musc

le co

ntrac

tions

.Ca

liforn

ia-Da

vis in

1950

sSt

orrs

Mus

cular

Dys

troph

yMu

tation

first

repo

rted i

n 195

6 at U

Mixe

d SCW

LAu

tosom

al re

cess

ive.

amPo

or5M

/15F

NoYe

sPi

erro

Homo

zygo

tes de

velop

joint

stiffn

ess a

nd m

uscle

Califo

rnia-

Davis

; acq

uired

1958

; kep

twe

akne

ss; b

reas

t and

othe

r mus

cle re

place

d by f

at an

das

homo

zygo

tesco

nnec

tive t

issue

.UC

D 07

7Im

porte

d fro

m Sw

itzer

land i

n 198

7;SC

WL

Multif

actor

ialB1

9Po

or5M

/6FNo

Yes

Abpl

analp

Inbre

eding

coeff

iecien

t (F)

>0.95

; MHC

: B19

; high

bloo

dful

l-sib

inbre

eding

sinc

e 196

8lip

id lev

els.

UCD

413

Deriv

ed fr

om co

mmer

cial m

eat s

tock

New

Hamp

shire

Autos

omal

rece

ssive

.am

Fair

15M/

20F

NoYe

sW

ilson

Mus

cular

dystr

ophy

homo

zygo

tes de

velop

joint

stiffn

ess

displa

ying m

uscu

lar dy

strop

hy;

and p

ector

al mu

scle

weak

ness

star

ting a

bout

4 wee

ksclo

sed f

lock s

ince 1

972

post-

hatch

, a co

nditio

n pro

duce

d by t

he re

place

ment

ofthe

se m

uscle

s by f

at an

d con

necti

ve tis

sue;

some

wing

redu

ction

defec

ts no

t rela

ted to

MD

also s

een.

UCD

Croo

ked-

neck

Dwa

rfAc

quire

d fro

m U

Conn

ectic

ut-St

orrs

Mixe

d SCW

LAu

tosom

al re

cess

ivecn

Fair

15M/

20F

NoW

ithAb

bott

Homo

zygo

tes ar

e ede

mic a

nd dw

arfed

, with

croo

ked,

in 19

91; c

ross

ed to

UCD

003 s

ever

alem

bryo

nic le

thal.

perm

ission

fragil

e nec

ks, th

in am

uscu

lar le

gs, a

nd no

volun

tary

times

sinc

e 199

6mu

scle

contr

actio

ns.

UCD

Ribo

flavin

Tra

nsfe

r Def

icien

tOu

t-cro

ssed

to U

CD 00

3 in 1

996;

SCW

LAu

tosom

al re

cess

ive,

rdPo

or5M

/5FSe

men

Yes

Abbo

ttHo

mozy

gotes

can o

nly m

ake d

efecti

ve rib

oflav

in bin

ding

acqu

ired f

rom

Penn

sylva

nia S

tate U

mater

nal-e

ffect

letha

l.pr

otein;

bird

s are

viab

le, bu

t suc

h hen

s can

not p

utin

1985

enou

gh rib

oflav

in int

o the

ir egg

albu

men t

o sup

port

anem

bryo

beyo

nd 16

days

of in

cuba

tion;

such

embr

yos a

redw

arfed

and h

emor

rhag

ic.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

16

Chi

cken

(con

t.)M

utan

t-Phy

siol

ogic

al d

efec

t (co

nt.)

UCD-

Corn

ell A

utos

omal

Dwar

fMu

tation

first

repo

rted i

n Cor

nell K

SCW

LAu

tosom

al re

cess

ive.

adw

Poor

25M/

50F

NoW

ithDe

lany

Homo

zygo

tes’ b

ody s

ize re

duce

d by 3

0% , r

ecog

nizab

leSt

rain

in 19

73; tr

ansfe

rred t

o Uco

llabo

ratio

nby

six t

o eigh

t wee

ks; a

vg w

eight

of fem

ale is

1173

g,Ca

liforn

ia-Da

vis in

1998

egg w

eight

57.7g

, 245

eggs

/cycle

, gen

erall

y goo

dvia

bility

but p

oor h

atcha

bility

.UD

el Ri

bofla

vin T

rans

fer D

efici

ent

Acqu

ired f

rom

E. B

uss a

tSC

WL

Autos

omal

rece

ssive

,rd

Poor

10M/

50F

NoYe

sW

hite

Homo

zygo

tes ca

n only

mak

e defe

ctive

ribofl

avin

bindin

gPe

nnsy

lvania

Stat

e Uma

terna

l-effe

ct let

hal.

prote

in; bi

rds a

re vi

able,

but s

uch h

ens c

anno

t put

enou

gh rib

oflav

in int

o the

ir egg

albu

men t

o sup

port

anem

bryo

beyo

nd 16

days

of in

cuba

tion;

such

embr

yos d

iedw

arfed

and h

emor

rhag

ic.

Mut

ant-R

epro

duct

ive

defe

ctAr

kans

as S

d Li

neMu

tation

first

repo

rted i

n the

1960

s at

Delaw

are X

Autos

omal

rece

ssive

,sd

g or s

dGo

od25

M/15

0FNo

NoKi

rby

Males

homo

zygo

us fo

r spe

rm de

gene

ratio

n hav

eOh

io St

ate in

Dela

ware

bree

d;SC

WL

sex-l

imite

d.de

fectiv

e spe

rm du

e to e

ffere

nt du

ct ma

lfunc

tion.

out-c

ross

ed w

ith S

CWL f

rom

asin

gle m

ale in

1980

sW

iscon

sin D

oubl

e Ovid

uct L

ine

Rhod

e Isla

ndInc

omple

tely p

enetr

ant

Rov

Fair:

4M/15

FNo

With

Bitg

ood

Both

right

and l

eft ov

iduct

form

in mo

st fem

ales o

f this

Red

with

multif

actor

ialins

titutio

nal

colla

bora

tion

line.

modif

iers.

Wisc

onsin

Res

trict

ed O

vulat

orKe

pt in

Wisc

onsin

Legh

orn g

eneti

cSC

WL

Sex-l

inked

rece

ssive

.ro

Fair:

6MNo

With

Bitg

ood

Hemi

zygo

us fe

males

lay f

ew if

any e

ggs,

have

high

back

grou

ndins

titutio

nal

colla

bora

tion

blood

lipids

, and

prod

uce n

o offs

pring

; male

s do n

otap

pear

to be

affec

ted.

Mut

ant-U

ncat

egor

ized

ISU

S1-1

9HDe

rived

from

two i

nbre

d Hy-L

ineSC

WL

B19,

Go

od4M

/24F

NoW

ithLa

mon

tInb

reed

ing co

effici

ent (

F)>0

.5; M

HC: B

19; Ir

GAThig

hstr

ains i

n 196

4IrG

AThig

hco

llabo

ratio

nall

ele lin

ked t

o MHC

.IS

U S1

-19L

Deriv

ed fr

om tw

o inb

red H

y-Line

SCW

LB1

9,

Good

4M/24

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.5;

MHC

: B19

; IrGA

Tlowstr

ains i

n 196

4IrG

ATlow

colla

bora

tion

allele

linke

d to M

HC.

ISU

S1-1

HDe

rived

from

two i

nbre

d Hy-L

ineSC

WL

B1,

Good

3M/30

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.5;

MHC

: B1;

IrGAT

high

strain

s in 1

964

IrGAT

high

colla

bora

tion

allele

linke

d to M

HC.

ISU

S1-1

LDe

rived

from

two i

nbre

d Hy-L

ineSC

WL

B1,

Good

3M/30

FNo

With

Lam

ont

Inbre

eding

coeff

icien

t (F)

>0.5;

MHC

: B1;

IrGAT

lowstr

ains i

n 196

4IrG

ATlow

colla

bora

tion

allele

linke

d to M

HC.

Stor

rs R

ose C

omb

Mutat

ion lo

ng kn

own i

n exh

ibitio

nMi

xed R

ose

Autos

omal

domi

nant.

RCo

mbine

dse

e: St

orrs

NoYe

sPi

erro

Homo

zygo

tes an

d hete

rozy

gotes

have

a br

oad c

omb,

stock

; intro

duce

d into

Stor

rs Cr

eepe

rco

mb W

Lwi

th St

orrs

Cree

per

cove

red w

ith pa

pillae

, and

a sin

gle sp

ike at

the b

ack;

in 19

60s

Cree

per

has b

een a

ssoc

iated

with

male

infer

tility.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

17

Chi

cken

(con

t.)M

utan

t-Unc

ateg

oriz

ed (c

ont.)

UCD

Cres

t/Hem

oglo

bin-

DMu

tant fo

rm of

hemo

globin

repo

rted

SCW

LLin

ked a

utoso

mal

Cr, H

bDNo

live

0Se

men

With

Abbo

ttHb

D is a

hemo

globin

varia

nt, fo

und i

n both

adult

and

linke

d to m

utant

crest

allele

, foun

d at

domi

nant

(Cr)

and

birds

perm

ission

embr

yonic

hemo

globin

, whic

h is l

inked

to th

e muta

ntU

Califo

rnia-

Davis

codo

mina

nt (H

bD ).cre

st all

ele; C

rest

caus

es a

tuft o

f long

feath

ers t

o for

mon

the h

ead.

UCD

Multi

plex

Com

bCa

uses

confo

rmati

onal

faults

inMi

xed S

CWL

Multif

actor

ial.

No liv

e0

Seme

nW

ithAb

bott

Modif

ies th

e sing

le-co

mb sh

ape b

y cau

sing p

oints

comb

shap

e of s

ingle-

comb

birds

perm

ission

(flesh

y pro

jectio

ns) t

o for

m alo

ng ac

cess

ory c

omb a

xes

exhib

ition c

hicke

ns(u

p to f

ive);

these

are u

suall

y sma

ller t

han t

he pr

imar

ypo

ints o

f the s

ingle

comb

.W

iscon

sin S

ex-li

nked

Skin

Col

orKe

pt in

Wisc

onsin

Legh

orn g

eneti

cSC

WL

Sex-l

inked

rece

ssive

.y

Fair:

6M/12

FNo

With

Bitg

ood

Homo

zygo

us or

hemi

zygo

us m

utant

birds

cann

ot de

posit

back

grou

ndins

titutio

nal

colla

bora

tion

yello

w pig

ment

in the

skin.

Pure

bre

edAr

kans

as G

iant J

ungl

e Fow

lIm

porte

d fro

m So

uthea

st As

ia in

Gian

t Jun

gleGo

od9M

/45F

NoYe

sAn

thon

yNa

turall

y high

ly re

sistan

t to R

ous s

arco

ma vi

rus;

1950

(one

male

and f

ive fe

males

);Fo

wlpr

obab

ly mo

dera

tely i

nbre

d (fou

ndati

on flo

ck of

6 bir

ds in

kept

as cl

osed

flock

1950

).Gu

elph

Silki

eDe

rived

from

exhib

ition s

tock;

Silki

eFiv

e auto

soma

lCr

, Fm

, h,

Poor

20M/

20F

NoW

ithEt

ches

Bree

d with

five m

ajor m

utatio

ns: C

r (lon

g fea

thers

onfou

ndati

on w

as on

e pair

; inbr

eedin

gdo

mina

ntsid,

Mb,

Po,

perm

ission

crown

of he

ad),

Po (o

ne or

mor

e extr

a pre

axial

toes

), Pt

minim

ized (

abou

t 35%

in 19

95)

(Cr,P

o,Pt,F

m,M

b), o

nePt

(legs

and t

oes f

eathe

red)

, Fm

(dar

k pigm

ent in

skin,

sex-l

ink re

cess

ive (id

),int

erna

l org

ans),

Mb (

long f

eathe

rs un

der c

hin),

and h

one a

utoso

mal

(feath

er de

fect: n

o hoo

klets)

.re

cess

ive (h

).NC

SU B

arre

d Pl

ymou

th R

ock

Kept

as cl

osed

flock

used

in

Barre

d Plym

outh

Fair

4M/40

FNo

Yes

Chris

tens

enre

sear

ch an

d tea

ching

Rock

NCSU

Rho

de Is

land

Red

Kept

as cl

osed

flock

used

in

Rhod

e Isla

ndFa

ir4M

/40F

NoYe

sCh

riste

nsen

rese

arch

and t

each

ingRe

dOt

tawa

New

Ham

pshi

reNe

w Ha

mpsh

ireNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

birds

perm

ission

Sask

atch

ewan

Bar

red

Plym

outh

Roc

kAc

qurie

d in 1

920s

Barre

d Plym

outh

Good

20M/

50F

NoW

ithCl

asse

nUs

ed as

a slo

w-gr

owing

chick

en m

odel

in ca

rdiac

and

Rock

colla

bora

tion

diges

tive p

hysio

logy r

esea

rch, a

nd as

sour

ce of

eggs

with

many

bloo

d and

mea

t spo

ts.Sa

skat

chew

an B

rown

Leg

horn

A str

ain of

Dan

ish B

rown

Legh

orn

Brow

n Leg

horn

Good

20M/

50F

NoW

ithCl

asse

nUs

ed as

a slo

w-gr

owing

chick

en m

odel

in ca

rdiac

and

acqu

ired f

rom

Scatt

ered

Acre

sco

llabo

ratio

ndig

estiv

e phy

siolog

y res

earch

, and

as so

urce

of fe

rtile

Hatch

ery i

n Han

over

, Onta

rio in

1965

eggs

for u

se in

rese

arch

.UB

C-Mi

nnes

ota R

hode

Islan

d Re

dAc

quire

d fro

m U

Minn

esota

-St. P

aul;

Rhod

e Isla

ndPo

or8M

/40F

NoYe

sCh

eng

From

a mo

dera

tely i

nbre

d (F

appr

ox. 0

.6) flo

ck of

ke

pt as

clos

ed flo

ck fo

r ove

r 20

Red

Rhod

e Isla

nd R

ed. B

ody w

eight

4.5 lb

.ge

nera

tions

; bac

k-cro

ss pa

rent

popu

lation

for U

BC R

C

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

18

Chi

cken

(con

t.)Pu

re b

reed

(con

t.)UC

D Si

lkie

Desc

ende

d fro

m six

exhib

ition b

irds

Silki

eFiv

e auto

soma

l dom

.Cr

, Fm

, h,

Fair

3M/3F

NoYe

sAb

bott

Bree

d with

five m

ajor m

utatio

ns: C

r (lon

g fea

thers

onac

quire

d fro

m a h

obby

-bre

eder

in(C

r,Po,P

t,Fm

,Mb)

, one

id, M

b, Po

,cro

wn of

head

), Po

(one

or m

ore e

xtra p

reax

ial to

es),

Ptce

ntral

Califo

rnia

in 19

94 (4

M/2F

)se

x-link

ed re

c. (id

),Pt

(legs

and t

oes f

eathe

red)

, Fm

(dar

k pigm

ent in

skin,

one a

utoso

mal re

c. (h

).int

erna

l org

ans),

Mb (

long f

eathe

rs un

der c

hin),

and h

(feath

er de

fect: n

o hoo

klets)

.UI

-Urb

ana C

olum

bian

Non-

exhib

ition (

meat)

varie

ty; ke

pt at

Colum

bian

Good

50M/

600F

NoW

ithPa

rson

sMe

at typ

e, wh

ite/bl

ack f

eathe

rs, sl

ow gr

owth,

long

legs

,U

Illino

is-Ur

bana

as cl

osed

flock

colla

bora

tion

narro

w br

east;

very

mild

selec

tion f

or bo

dy si

ze.

for ov

er 30

year

sUI

-Urb

ana N

ew H

amps

hire

Non-

exhib

ition (

meat)

varie

ty; at

UNe

w Ha

mpsh

ireGo

od50

M/20

0FNo

With

Pars

ons

Meat

type,

mode

rate

grow

th, go

od co

nform

ation

; new

Illino

is-Ur

bana

for o

ver 3

0 yea

rs;co

llabo

ratio

nblo

odlin

es in

trodu

ced a

ppro

x. 19

93 to

incre

ase b

ody

kept

as cl

osed

flock

exce

pt for

weigh

t.ou

tcros

s in 1

993

UMas

s Lig

ht B

rown

Leg

horn

Serve

s as c

ontro

l for U

Mass

Smy

thLig

ht Br

own

Disp

ersa

l15

M/35

FNo

Yes

Smyt

hLig

ht-br

own L

egho

rn ch

icken

that

has n

ever

show

nLin

e (no

occu

ranc

e of d

elaye

dLe

ghor

nin

1998

signs

of de

layed

amela

noge

nesis

.am

elano

gene

sis)

Ran

dom

bred

Arka

nsas

Bro

wn L

ine B

101

Acqu

ired f

rom

J. Sm

yth at

UBr

own L

egho

rneb

Good

10M/

20F

NoW

ithEr

fMH

C-ma

tched

contr

ol/pa

renta

l line

for A

rkans

as S

myth

Mass

achu

setts

-Amh

erst

in 19

96;

colla

bora

tion

line B

101.

subli

ne of

ance

stral

pare

nt sto

ck fo

rSm

yth Li

neAr

kans

as B

rown

Lin

e B10

2Ac

quire

d fro

m J.

Smyth

at U

Brow

n Leg

horn

,eb

Good

10M/

20F

NoW

ithEr

fMH

C-ma

tched

contr

ol/pa

renta

l line

for A

rkans

as S

myth

Mass

achu

setts

-Amh

erst

in 19

98;

synth

etic

colla

bora

tion

line B

102.

subli

ne of

ance

stral

pare

nt sto

ck fo

rSm

yth Li

neAr

kans

as L

ight

Bro

wn L

egho

rn B

101

Acqu

ired f

rom

J. Sm

yth at

ULig

ht Br

own

Good

10M/

20F

NoW

ithEr

fMH

C-ma

tched

contr

ol for

Arka

nsas

Smy

th lin

e B10

1;Ma

ssac

huse

tts-A

mher

st in

1996

Legh

orn

colla

bora

tion

does

not d

evelo

p dela

yed a

melan

ogen

esis

(auto

immu

ne vi

tiligo

).Ar

kans

as R

ando

mbr

edFo

rmed

by cr

ossin

g com

merci

alCo

mmer

cial

Good

16M/

48F

NoYe

sAn

thon

yRa

ndom

bred

mea

t bird

contr

ol fro

m mu

ltiple

pare

ntal

meat

(bro

iler)

pare

nt lin

es (s

even

meat

sire c

ross

strain

cros

s.ma

le lin

es an

d eigh

t fema

le lin

es)

Athe

ns R

ando

mbr

edSy

ntheti

c line

integ

ratin

gCo

mmer

cial

Good

30M/

150F

NoYe

sBu

rke

Rand

ombr

ed co

ntrol/

stand

ard;

found

ation

male

s fro

mco

mmer

cial a

nd U

GA E

xper

imen

tme

at sir

e cro

ssco

mmer

cial m

eat-t

ype (

WPR

, WCo

rnish

, NH)

; foun

datio

nSt

ation

mea

t-typ

e stoc

ks, s

tarted

infem

ales f

rom

egg-

prod

uctio

n sele

cted R

IR, B

PR, W

PR,

1956

NH, S

CWL a

nd C

ornis

h fro

m So

uther

n Reg

ional

Expe

rimen

t Stat

ions.

Athe

ns-C

anad

ian R

ando

mbr

edAc

quire

d fro

m the

Can

adian

Dep

t of

Comm

ercia

l R,

PGo

od60

M/18

0FNo

Yes

Burk

eRa

ndom

bred

contr

ol/sta

ndar

d seg

rega

ting f

or si

ngle,

Agric

ultur

e in 1

958;

includ

es W

hite

meat

sire c

ross

rose

, and

pea c

omb,

with

domi

nant

white

, som

e red

Wya

ndott

e and

thre

e syn

thetic

mea

tan

d blac

k fea

ther c

olor o

ccur

ring.

strain

s

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

19

Chi

cken

(con

t.)R

ando

mbr

ed (c

ont.)

Corn

ell C

Spe

cials

(B13

)MH

C-ma

tched

to C

orne

ll Obe

se;

SCW

LB1

3Po

or8M

/42F

NoYe

sMa

rsh

Contr

ol lin

e for

Cor

nell O

bese

; MHC

B13

; also

can

kept

as cl

osed

flock

for o

ver 3

0pr

oduc

e a pe

rsista

nt rig

ht Mu

lleria

n duc

t (oft

en le

ading

toye

ars;

deriv

ed fr

om C

orne

ll C S

train

the fo

rmati

on of

two o

viduc

ts) in

affec

ted fe

males

.NC

SU-C

orne

ll K S

train

Deriv

ed fr

om C

orne

ll K-st

rain

SCW

LB1

5Go

od40

M/10

0FNo

Yes

Qure

shi

MHC:

B15

.Ot

tawa

10No

selec

tion s

ince 1

973;

deriv

edSC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esCo

ntrol/

stand

ard m

aintai

ned w

ithou

t sele

ction

sinc

efro

m a c

ombin

ation

of N

orth

birds

perm

ission

1973

.Am

erica

n com

merci

al eg

g-lay

ersto

cks

Otta

wa 20

Form

ed by

comb

ining

nine

Comm

ercia

l No

live

0Bl

astod

isc ce

llsW

ithEt

ches

Pedig

reed

rand

ombr

ed m

eat (

broil

er) r

efere

nce s

tock f

rom

comm

ercia

l bro

iler s

ire st

ocks

inme

at sir

e cro

ssbir

dspe

rmiss

ionco

mmer

cial s

ire lin

es.

1979

Otta

wa 30

Form

ed by

cros

sing n

ineCo

mmer

cial

No liv

e0

Blas

todisc

cells

With

Etch

esPe

digre

ed ra

ndom

bred

mea

t (br

oiler

) refe

renc

e stoc

k fro

mco

mmer

cial b

roile

r dam

stoc

ks in

meat

sire c

ross

birds

perm

ission

comm

ercia

l dam

lines

.19

79Ot

tawa

5No

selec

tion s

ince 1

950

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Contr

ol/sta

ndar

d main

taine

d with

out s

electi

on si

nce

birds

perm

ission

1950

.Ot

tawa

7De

rived

from

four

comm

ercia

lSC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esCo

ntrol/

stand

ard m

aintai

ned w

ithou

t sele

ction

sinc

eeg

g-lay

er st

ocks

and k

ept w

ithou

tbir

dspe

rmiss

ion19

58.

selec

tion s

ince 1

958;

origi

nally

know

n as K

entvi

lle C

ontro

l Stra

in 7

UCD

412

Kept

as cl

osed

flock

sinc

e 197

2Ne

w Ha

mpsh

ireFa

ir15

M/20

FNo

Yes

Wils

onCo

ntrol/

stand

ard f

or U

CD-4

13 (m

uscu

lar dy

strop

hy).

UMas

s Bro

wn L

ine

Sour

ce of

delay

ed am

elano

gene

sisBr

own L

egho

rn,

Autos

omal

rece

ssive

.eb

Disp

ersa

l90

NoYe

sSm

yth

Brow

n hom

ozyg

otes a

re so

lid br

own a

s chic

ks; a

dult

mutat

ion; s

erve

s as M

HC-m

atche

dsy

ntheti

cin

1998

males

are w

ild-ty

pe, fe

males

are s

tipple

d dar

k bro

wn;

contr

ol for

UMa

ss S

myth

Line

serve

s as M

HC-m

atche

d con

trols

for U

Mass

Smy

th Lin

e.W

iscon

sin L

egho

rnKe

pt as

clos

ed flo

ck fo

r ove

r 30

SCW

LFa

ir:20

M/18

0FNo

With

Bitg

ood

Clos

ed po

pulat

ion of

Sing

le-co

mb W

hite L

egho

rns.

gene

ratio

ns; u

sed a

s gen

etic

institu

tiona

lco

llabo

ratio

nba

ckgr

ound

for s

ever

al mu

tant

stock

s at U

Wisc

onsin

-Mad

ison

Wisc

onsin

Leg

horn

line U

W-6

XFr

om cr

oss o

f com

merci

alSC

WL

Poor

Few

NoW

ithBi

tgoo

dCl

osed

rand

ombr

ed/co

ntrol

popu

lation

of 19

50s o

rigin

egg-

layer

s (Sp

ruce

leigh

X H

N) in

colla

bora

tion

egg-

type c

omme

rcial

Sing

le-co

mb W

hite L

egho

rns.

1950

s; ke

pt as

clos

ed flo

ck fo

r ove

r20

year

s; alm

ost e

xtinc

t, only

asin

gle, lo

w pr

oduc

tion f

emale

in19

98W

iscon

sin N

ew H

amps

hire

Kept

as cl

osed

flock

for o

ver 3

0Ne

w Ha

mpsh

ireFa

ir:20

M/18

0FNo

With

Bitg

ood

Clos

ed ra

ndom

bred

/contr

ol po

pulat

ion of

pure

bred

New

gene

ratio

ns; u

sed a

s gen

etic

institu

tiona

lco

llabo

ratio

nHa

mpsh

ire ch

icken

s.ba

ckgr

ound

for s

ever

al mu

tant

stock

s at U

Wisc

onsin

-Mad

ison

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

20

Chi

cken

(con

t.)Se

lect

ed-B

ehav

iora

l tra

itPu

rdue

KGB

Deriv

ed fr

om a

seve

n-wa

y, wi

deSC

WL

Fair

100M

/100F

NoW

ithMu

irSe

lected

for n

on-a

ggre

ssive

beha

vior a

nd re

sistan

ce to

comm

ercia

l cro

ss at

the N

orth

colla

bora

tion

stres

s (“K

inder

, Gen

tler B

ird”).

Centr

al Re

giona

l Pou

ltry La

bora

tory

in 19

72Pu

rdue

MBB

Deriv

ed fr

om P

urdu

e KGB

SCW

LFa

ir15

0M/25

0FNo

With

Muir

Selec

ted fo

r mor

e agg

ress

ive be

havio

r (“M

ean,

Bad

colla

bora

tion

Bird

”).

Sele

cted

-Egg

trai

tCo

rnell

K S

train

Kept

as cl

osed

flock

sinc

e 195

4; SC

WL

B15

Poor

165

NoYe

s, fee

Diet

ert

MHC:

B15

; res

istan

t to le

ukos

is co

mplex

(par

ticula

rlyse

lected

to op

timize

egg p

rodu

ction

,Ma

rek’s

dise

ase)

by se

lectio

n afte

r natu

ral e

xpos

ure t

oeg

g size

, bod

y weig

ht, ot

her

the vi

ruse

s; go

od eg

g pro

ducti

on ch

arac

terist

ics.

econ

omic

egg t

raits

until

1971

, then

rand

ombr

ed w

ith m

inima

l sele

ction

Guelp

h Ba

rred

Plym

outh

Roc

kDe

rived

from

comm

ercia

l stoc

kBa

rred P

lymou

thGo

od30

M/13

0FNo

With

Etch

esCo

mmer

cial s

tock s

electe

d for

egg p

rodu

ction

.ac

quire

d in 1

991

Rock

perm

ission

Guelp

h Co

mm

ercia

l Leg

horn

Lin

eDe

rived

from

comm

ercia

l stoc

kSC

WL

Good

30M/

130F

NoW

ithEt

ches

Comm

ercia

l stoc

k sele

cted f

or eg

g pro

ducti

on.

acqu

ired i

n 199

1pe

rmiss

ionOt

tawa

1De

rived

from

Otta

wa 5

in 19

50;

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r high

egg p

rodu

ction

and r

elated

trait

s.div

ided i

nto O

ttawa

1 an

d 3 in

1971

birds

perm

ission

Otta

wa 2

Deriv

ed fr

om se

ven C

anad

ian R

OPSC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh eg

g pro

ducti

on an

d rela

ted tr

aits.

(egg

-type

) stoc

ks in

1951

; divi

ded i

nbir

dspe

rmiss

ion19

69 in

to Ot

tawa 2

and 4

Otta

wa 3

Estab

lishe

d fro

m Ot

tawa 5

in 19

50;

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r high

egg p

rodu

ction

and r

elated

trait

s.div

ided i

nto O

ttawa

1 an

d 3 in

1971

birds

perm

ission

Otta

wa 4

Deriv

ed fr

om se

ven C

anad

ian R

OPSC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh eg

g pro

ducti

on an

d rela

ted tr

aits.

(egg

-type

) stoc

ks in

1951

; divi

ded i

nbir

dspe

rmiss

ion19

69 in

to Ot

tawa 2

and 4

Otta

wa 8

Deriv

ed fr

om O

ttawa

7 in

1969

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r high

egg p

rodu

ction

and r

elated

trait

s.bir

dspe

rmiss

ionOt

tawa

9De

rived

from

Otta

wa 7

in 19

69SC

WL

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh eg

g pro

ducti

on an

d rela

ted tr

aits.

birds

perm

ission

UCD

058

SCW

LPo

or5M

/6FNo

Yes

Abpl

analp

Inbre

eding

coeff

icien

t (F)

>=0.8

; sele

cted f

or in

creas

edeg

g pro

ducti

on.

UCD

082

SCW

LPo

or5M

/6FNo

Yes

Abpl

analp

Inbre

eding

coeff

icien

t (F)

>0.76

; sele

cted f

or eg

gpr

oduc

tion w

ith re

laxed

selec

tion i

n rec

ent g

ener

ation

s.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

21

Chi

cken

(con

t.)Se

lect

ed-G

row

th tr

ait

Aubu

rn T

ibial

Dys

chon

drop

lasia-

High

Deriv

ed fr

om a

comm

ercia

l mea

t sire

Comm

ercia

lGo

od15

M/70

FNo

Yes

Berry

One o

f two s

trains

that

were

deve

loped

from

alin

e acq

uired

in 19

88me

at sir

e line

comm

ercia

l mea

t sire

line b

y dive

rgen

t sele

ction

for

tibial

dysc

hond

ropla

sia; in

ciden

ce is

95%

for b

irds i

nthi

s line

after

11 ge

nera

tions

of se

lectio

n.Au

burn

Tib

ial D

ysch

ondr

oplas

ia-Lo

wDe

rived

from

a co

mmer

cial m

eat s

ireCo

mmer

cial

Good

15M/

70F

NoYe

sBe

rryOn

e of tw

o stra

ins th

at we

re de

velop

ed fr

om a

line a

cquir

ed in

1988

meat

sire l

ineco

mmer

cial m

eat s

ire lin

e by d

iverg

ent s

electi

on fo

rtib

ial dy

scho

ndro

plasia

; incid

ence

is 6%

for b

irds i

n this

line a

fter 1

1 gen

erati

ons o

f sele

ction

.Ot

tawa

21De

rived

from

Otta

wa 20

Comm

ercia

l No

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r high

body

weig

ht an

d low

fat.

meat

sire c

ross

birds

perm

ission

Otta

wa 23

Deriv

ed fr

om O

ttawa

20Co

mmer

cial

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh bo

dy w

eight

and f

eed e

fficien

cy.

meat

sire c

ross

birds

perm

ission

Otta

wa 25

Deriv

ed fr

om O

ttawa

20Co

mmer

cial

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh bo

dy w

eight,

low

fat, a

nd fe

edme

at sir

e cro

ssbir

dspe

rmiss

ioneff

icien

cy.

Otta

wa 31

Deriv

ed fr

om O

ttawa

30Co

mmer

cial

No liv

e0

Blas

todisc

cells

With

Etch

esSe

lected

for h

igh bo

dy w

eight,

feed

effic

iency

, and

egg

meat

sire c

ross

birds

perm

ission

prod

uctio

n.PS

U-At

hens

RB

14-4

2H (H

iK)

Deriv

ed fr

om A

thens

-Can

adian

Comm

ercia

lGo

od: E

xp.

20M/

60F

NoW

ithBa

rbat

oSe

lected

for m

axim

um gr

owth

betw

een 1

4 and

42 da

ysRa

ndom

bred

from

U G

eorg

ia-At

hens

meat

stock

X

Sta.

&co

llabo

ratio

nfro

m a s

tartin

g pop

ulatio

n of 3

50.

egg s

electe

dUS

DAfem

ale lin

esPS

U-At

hens

RB

14-4

2L (L

oK)

Deriv

ed fr

om A

thens

-Can

adian

Comm

ercia

lGo

od: E

xp.

20M/

60F

NoW

ithBa

rbat

oSe

lected

for m

inimu

m gr

owth

betw

een 1

4 and

42 da

ysRa

ndom

bred

from

U G

eorg

ia-At

hens

meat

stock

X

Sta.

&co

llabo

ratio

nfro

m a s

tartin

g pop

ulatio

n of 3

50.

egg s

electe

dUS

DAfem

ale lin

esPS

U-At

hens

RB

14H

Deriv

ed fr

om A

thens

-Can

adian

Comm

ercia

lGo

od: E

xp.

20M/

60F

Seme

n, Ge

rmW

ithBa

rbat

oOv

er ni

ne ge

nera

tions

selec

ted fo

r max

imum

grow

thRa

ndom

bred

from

U G

eorg

ia-At

hens

meat

stock

X

Sta.

&ce

llsco

llabo

ratio

nbe

twee

n zer

o and

14 da

ys fr

om a

startin

g pop

ulatio

n of

egg s

electe

dUS

DA35

0.fem

ale lin

esPS

U-At

hens

RB

14L

Deriv

ed fr

om A

thens

-Can

adian

Comm

ercia

lGo

od: E

xp.

20M/

60F

Seme

n, Ge

rmW

ithBa

rbat

oOv

er ni

ne ge

nera

tions

selec

ted fo

r mini

mum

grow

thRa

ndom

bred

from

U G

eorg

ia-At

hens

meat

stock

X

Sta.

&ce

llsco

llabo

ratio

nbe

twee

n zer

o and

14 da

ys fr

om a

startin

g pop

ulatio

n of

egg s

electe

dUS

DA35

0; ve

ry low

fertil

ity (2

5-30

%).

female

lines

PSU-

Athe

ns R

B 42

HDe

rived

from

Athe

ns-C

anad

ianCo

mmer

cial

Good

: Exp

.20

M/60

FSe

men,

Germ

With

Barb

ato

Over

nine

gene

ratio

ns se

lected

for m

axim

um gr

owth

Rand

ombr

ed fr

om U

Geo

rgia-

Athe

nsme

at sto

ck X

St

a. &

cells

colla

bora

tion

betw

een z

ero a

nd 42

days

from

a sta

rting p

opula

tion o

feg

g sele

cted

USDA

350.

female

lines

PSU-

Athe

ns R

B 42

LDe

rived

from

Athe

ns-C

anad

ianCo

mmer

cial

Good

20M/

60F

Seme

n, Ge

rmW

ithBa

rbat

oOv

er ni

ne ge

nera

tions

selec

ted fo

r mini

mum

grow

thRa

ndom

bred

from

U G

eorg

ia-At

hens

meat

stock

X

cells

colla

bora

tion

betw

een z

ero a

nd 42

days

from

a sta

rting p

opula

tion o

feg

g sele

cted

350.

female

lines

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

22

Chi

cken

(con

t.)Se

lect

ed-G

row

th tr

ait (

cont

.)Vi

rgin

ia Bo

dy W

eight

-Hig

hFo

rmed

by cr

ossin

g sev

en in

bred

Whit

e Plym

outh

Poor

100M

/100F

NoW

ithSi

egel

Selec

ted 40

gene

ratio

ns fo

r high

8-we

ek bo

dy w

eight.

lines

kept

at Vi

rgini

a Stat

e in 1

957

Rock

colla

bora

tion

Virg

inia

Body

Weig

ht-L

owFo

rmed

by cr

ossin

g sev

en in

bred

Whit

e Plym

outh

Poor

100M

/100F

NoW

ithSi

egel

Selec

ted 35

gene

ratio

ns fo

r low

8-we

ek bo

dy w

eight.

lines

kept

at Vi

rgini

a Stat

e in 1

957

Rock

colla

bora

tion

Sele

cted

-Imm

une

trai

tAr

kans

as P

rogr

esso

rDe

rived

from

SCW

L main

taine

d at U

SCW

LGo

od9M

/45F

NoYe

sEr

fSe

lected

20 ge

nera

tions

for t

umor

grow

th (p

rogr

essio

n)Ar

kans

as si

nce a

ppro

ximate

ly 19

45;

follow

ing fo

rmati

on w

ith ex

posu

re to

Rou

s sar

coma

thoug

ht to

be hi

ghly

inbre

d; als

ovir

us.

know

n as A

rkans

as R

ous S

arco

maPr

ogre

ssion

Arka

nsas

Reg

ress

orDe

rived

from

F-1

and F

-2 pr

ogen

y of

SCW

L X G

iant

Good

9M/45

FNo

Yes

Erf

Selec

ted 20

gene

ratio

ns fo

r tum

or re

gres

sion f

ollow

ingcro

sses

of S

CWL a

nd G

iant J

ungle

Jung

le Fo

wlfor

matio

n with

expo

sure

to R

ous s

arco

ma vi

rus.

Fowl

; also

know

n as A

rkans

asRo

us S

arco

ma R

egre

ssion

Athe

ns A

RDe

rived

from

a cro

ss of

Athe

nsSC

WL

Good

36M/

36F

NoYe

sBu

rke

Selec

ted fo

r res

istan

ce to

aflat

oxin.

AR2.5

and A

R3.0

in 19

97Au

burn

RKe

pt as

clos

ed flo

ck si

nce 1

948

SCW

Lse

eGo

od18

0No

With

Ewald

Resis

tant to

cocc

idios

is; M

HC: B

3; oth

er er

ythro

cyte

desc

riptio

nco

llabo

ratio

nall

oanti

gens

: A/E

2, C2

, D3,

H1, I2

, K2,

P3, “

Q”2.

Aubu

rn S

Kept

as cl

osed

flock

sinc

e 195

2SC

WL

see

Good

200

NoW

ithEw

aldSu

scep

tible

to co

ccidi

osis;

MHC

: B5;

other

eryth

rocy

tede

scrip

tion

colla

bora

tion

alloa

ntige

ns: A

/E2,

C3, D

2, H1

, I2, K

3, P2

, “Q”

1.Co

rnell

S13

Deriv

ed fr

om C

orne

ll S S

train;

SC

WL

B13

Good

28M/

106F

NoYe

s, fee

Scha

tMH

C: B

13; h

ighy s

usce

ptible

to M

arek

’s dis

ease

viru

s;sp

ecific

patho

gen-

free

free o

f mos

t avia

n viru

ses.

Otta

wa 2R

Deriv

ed fr

om O

ttawa

-2 an

d-4;

also

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r res

istan

ce to

Mar

ek’s

disea

se vi

rus.

refer

red t

o as O

ttawa

R2

birds

perm

ission

Otta

wa 3R

Deriv

ed fr

om O

ttawa

-1 an

d-3;

also

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r res

istan

ce to

Mar

ek’s

disea

se vi

rus.

refer

red t

o as O

ttawa

R3

birds

perm

ission

Otta

wa 8R

Deriv

ed fr

om O

ttawa

8 an

d-9;

also

SCW

LNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Selec

ted fo

r res

istan

ce to

Mar

ek’s

disea

se vi

rus.

refer

red t

o as O

ttawa

R8

birds

perm

ission

Virg

inia

Antib

ody L

ine-

High

Deriv

ed fr

om a

Corn

ell ra

ndom

bred

SCW

LPo

or10

0M/10

0FNo

With

Sieg

elSe

lected

over

20 ge

nera

tions

for h

igh an

tibod

y res

pons

eSC

WL,

startin

g in 1

977

colla

bora

tion

to sh

eep r

ed bl

ood c

ells.

Virg

inia

Antib

ody L

ine-

Low

Deriv

ed fr

om a

Corn

ell ra

ndom

bred

SCW

LPo

or10

0M/10

0FNo

With

Sieg

elSe

lected

over

20 ge

nera

tions

for lo

w an

tibod

y res

pons

eSC

WL,

startin

g in 1

977

colla

bora

tion

to sh

eep r

ed bl

ood c

ells.

Sele

cted

-Phy

siol

ogic

al tr

ait

Arka

nsas

Asc

ites R

esist

ant

Pedig

reed

comm

ercia

l syn

thetic

Comm

ercia

l Go

od16

M/48

FNo

NoAn

thon

ySe

lected

thre

e gen

erati

ons f

or re

sistan

ce to

ascit

essto

cks

meat

sire c

ross

unde

r high

altitu

de si

mulat

ion; fa

mily

selec

tion.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Chi

cken

gen

etic

sto

cks

Appe

ndix

2 T

able

2.1

23

Chi

cken

(con

t.)Se

lect

ed-P

hysi

olog

ical

trai

t (co

nt.)

Arka

nsas

Asc

ites S

usce

ptib

lePe

digre

ed co

mmer

cial s

ynthe

ticCo

mmer

cial

Good

16M/

48F

NoNo

Anth

ony

Selec

ted th

ree g

ener

ation

s for

susc

eptib

ility t

o asc

ites

stock

sme

at sir

e cro

ssun

der h

igh al

titude

simu

lation

; fami

ly se

lectio

n.

Tran

sgen

icAD

OL T

rans

-ALV

E6SC

WL

ALVE

6Go

od:

N/A

NoYe

sBa

con

Homo

zygo

us fo

r the

alv-6

tran

sgen

e; ex

pres

ses e

nvUS

DAgly

copr

otein

of AL

V su

bgro

up A

; res

ists i

nfecti

on w

ithAL

V su

bgro

up A

.Ot

tawa

TR

SCW

LAL

VE6

No liv

e0

Blas

todisc

cells

With

Etch

esHa

s inc

orpo

rated

the a

lv-6 t

rans

gene

.bir

dspe

rmiss

ion

Unc

ateg

oriz

edOt

tawa

6No

t rep

orted

No liv

e0

Blas

todisc

cells

With

Etch

esNo

t rep

orted

.bir

dspe

rmiss

ionOt

tawa

80No

t rep

orted

No liv

e0

Blas

todisc

cells

With

Etch

esNo

t rep

orted

.bir

dspe

rmiss

ionOt

tawa

90No

t rep

orted

No liv

e0

Blas

todisc

cells

With

Etch

esNo

t rep

orted

.bir

dspe

rmiss

ionOt

tawa

N3

Not r

epor

tedNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Not r

epor

ted.

birds

perm

ission

Otta

wa N

4No

t rep

orted

No liv

e0

Blas

todisc

cells

With

Etch

esNo

t rep

orted

.bir

dspe

rmiss

ionOt

tawa

N8

Not r

epor

tedNo

live

0Bl

astod

isc ce

llsW

ithEt

ches

Not r

epor

ted.

birds

perm

ission

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

24

Japa

nese

qua

ilIn

bred

UBC

IInb

red s

tartin

g in 1

992,

with

Poor

8M/16

FNo

Yes

Chen

gFu

ll-sibl

ing in

bree

ding f

or fo

ur ge

nera

tions

, star

ting i

nre

laxati

on of

inbr

eedin

g for

the l

ast

1992

; rela

xed i

nbre

eding

pres

sure

in 19

96 (c

ross

ing ha

lffew

year

ssib

s or le

ss cl

osely

relat

ed bi

rds).

Mut

ant-C

olor

, egg

shel

lUB

C CE

Acqu

ired f

rom

U Na

goya

(Jap

an) in

Autos

omal

rece

ssive

.ce

Comb

ined

see:

UBC

C-CE

NoYe

sCh

eng

Fema

les ho

mozy

gous

for c

elado

n lay

blue

-shell

ed eg

gs.

1988

; com

bined

with

UBC

C in

1995

with

UBC

Cin

1995

UBC

WE

Cros

s of U

Nag

oya (

Japa

n) W

E an

dAu

tosom

al re

cess

ive.

weCo

mbine

dse

e: UB

C W

-WE

NoYe

sCh

eng

Fema

les ho

mozy

gous

for w

hite-

egg l

ay w

hite-

shell

edU

Sask

atche

wan s

tock;

comb

ined

with

UBC

eggs

.wi

th UB

C W

in 19

95W

in 19

98

Mut

ant-C

olor

, fea

ther

Arka

nsas

Whi

teMu

tation

repo

rted i

n Arka

nsas

Autos

omal

rece

ssive

.Po

or36

M/36

FNo

Anth

ony

Homo

zygo

tes ha

ve pu

re w

hite f

eathe

rs wi

th da

rk ey

es,

rand

ombr

ed Ja

pane

se qu

aildif

feren

t from

Eng

lish W

hite.

Arka

nsas

Eng

lish

Whi

teAc

quire

d fro

m the

Qua

il Res

ource

Autos

omal

rece

ssive

.wh

Good

36M/

36F

NoAn

thon

yHo

mozy

gotes

have

dark

eyes

and m

ostly

whit

e fea

thers

Cente

rwi

th sp

ashe

s of w

ild-ty

pe co

lor.

UBC

BHAc

quire

d fro

m U

Nago

ya (J

apan

) inAu

tosom

al do

mina

nt.Bh

Poor

6M/12

FNo

Yes

Chen

gBl

ack-a

t-hat

ch ho

mozy

gotes

die b

y the

sixth

day o

f19

88; fr

eque

ntly b

ackc

ross

ed to

incub

ation

; hete

rozy

gotes

are v

iable,

but h

ave o

nly fa

intUB

C A

remn

ants

of the

wild

-type

yello

w str

ipes a

t hatc

h.UB

C C-

CECi

nnam

on m

utatio

n rep

orted

in U

BCAu

tosom

al re

cess

ives.

cin, c

ePo

or24

M/48

FNo

Yes

Chen

gCi

nnam

on ho

mozy

gotes

have

brigh

t ora

nge-

brow

nM

in 19

78; c

elado

n muta

tion

feathe

rs ins

tead o

f med

ium br

own,

and e

yes a

re re

d in

acqu

ired f

rom

U Na

goya

(Jap

an) in

the ch

ick; fe

males

homo

zygo

us fo

r cela

don l

ay19

88; c

ombin

ed 19

95blu

e-sh

elled

eggs

.UB

C D

Acqu

ired i

n 197

9, the

n com

bined

Two a

llelic

sex-l

inked

alD , al

Poor

24M/

48F

NoYe

sCh

eng

Dark-

eyed

dilu

te ho

mozy

gotes

have

dark

eyes

and

with

UBC

AL (f

rom

U Sa

skatc

hewa

n)re

cess

ives.

dilute

d fea

ther c

olor;

homo

zygo

tes ha

ve un

pigme

nted

in 19

84re

tinas

and n

early

whit

e fea

thers;

albin

o hem

i- and

homo

zygo

tes ar

e nea

rly w

hite.

UBC

F-SB

Fawn

muta

tion f

rom

a bre

eder

near

Autos

omal

domi

nant

Yf , sb

Poor

24M/

48F

NoYe

sCh

eng

Fawn

heter

ozyg

otes h

ave f

awn-

color

ed fe

ather

s, bu

t Yf

Vanc

ouve

r, BC

in 19

87; c

ombin

edYf is

letha

l inis

letha

l in th

e hom

ozyg

otes;

shor

t-bar

b hom

ozyg

otes

with

UBC

SBho

mozy

gotes

; sb i

s an

have

feath

er ba

rbs a

ppro

ximate

ly 75

% sh

orter

than

autos

omal

rece

ssive

.no

rmal.

UBC

HBa

ck-cr

osse

d to U

BC A

each

Autos

omal

incom

plete

EPo

or6M

/12F

NoYe

sCh

eng

Exte

nded

bro

wn ho

mozy

gotes

and,

to a l

esse

r exte

nt,ge

nera

tion

domi

nant.

heter

ozyg

otes,

have

exten

ded d

istrib

ution

of bl

ack a

ndda

rk br

own p

igmen

t in th

e fea

thers,

with

both

sexe

sap

pear

ing th

e sam

e.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

25

Japa

nese

qua

il (c

ont.)

Mut

ant-C

olor

, fea

ther

(con

t.)UB

C RH

Acqu

ired f

rom

Virg

inia P

olytec

hnic

Autos

omal

rece

ssive

.erh

Comb

ined

see:

UBC

NoYe

sCh

eng

Redh

ead h

omoz

ygote

s hav

e dist

inctly

red h

ead f

eathe

rsIns

t. in 1

977;

comb

ined w

ith U

BCwi

th UB

CRF

-RH

(same

as pa

nsy);

ruffle

d hom

ozyg

otes h

ave c

urve

d, be

ntRF

in 19

95RF

in 19

95or

ruffle

d fea

thers.

UBC

SIAc

quire

d fro

m U

Nago

ya (J

apan

) inAu

tosom

al inc

omple

teB

Poor

6M/12

FNo

Yes

Chen

gSi

lver h

omoz

ygote

s hav

e whit

e fea

thers

and a

centr

ally

1988

; bac

k-cro

ssed

to U

BC A

ever

ydo

mina

nt.de

pigme

nted r

etina

(ring

-retin

a); h

etero

zygo

tes ha

vege

nera

tion

grey

ish fe

ather

s, an

d may

have

a few

whit

e prim

aries

.UB

C W

-WE

Mutat

ion w

h fro

m flo

ck ne

arAu

tosom

al re

cess

ives.

wh, w

ePo

or24

M/48

FNo

Yes

Chen

gRe

cess

ive w

hite h

omoz

ygote

s hav

e whit

e fea

thers

and

Vanc

ouve

r, BC

(197

6); w

e fro

m a

dark

eyes

; hete

rozy

gotes

have

whit

e in v

entra

l feath

ercro

ss of

stoc

ks fr

om U

Nag

oya a

ndtra

cts an

d fac

e, wi

th a m

ixtur

e of d

ark a

nd w

hite f

eathe

rsU

Sask

atche

wan;

comb

ined 1

995

on th

e bac

k; tho

se ho

mozy

gous

for w

e lay

whit

e egg

s.UB

C W

BMu

tation

repo

rted i

n UBC

A in

1975

;Au

tosom

al re

cess

ive.

wbCo

mbine

dse

e: UB

CNo

Yes

Chen

gW

hite-

brea

sted m

utants

have

whit

e fea

thers

on fa

ce,

comb

ined w

ith U

BC P

C in

1995

with

UBC

PC-W

Bve

ntral

neck

, bre

ast, p

rimar

ies an

d mos

t sec

onda

ries.

PC in

1995

UBC

YMu

tation

repo

rted i

n UBC

D in

1983

;Au

tosom

al do

mina

nt,Y

Poor

6M/12

FNo

Yes

Chen

gYe

llow

heter

ozyg

otes h

ave y

ellow

(gold

en) f

eathe

rs wi

thba

ck-cr

osse

d to U

BC A

ever

ylet

hal in

homo

zygo

tes.

restr

icted

blac

k pigm

ent d

istrib

ution

; the m

utatio

n is

gene

ratio

nlet

hal in

homo

zygo

us fo

rm.

Mut

ant-D

evel

opm

enta

l def

ect-S

kin/

feat

her

UBC

DF-M

DFMu

tation

s rep

orted

in U

BC W

in 19

79Au

tosom

al do

mina

ntDf

, mdf

Poor

24M/

48F

NoYe

sCh

eng

Two m

utatio

ns ac

ting i

n con

cert

to pr

oduc

e defe

ctive

(Df)

and r

eces

sive

feathe

rs: sh

ort, s

parse

down

at ha

tch, a

nd de

fectiv

e(m

df) w

ith va

riable

barb

s in l

ater f

eathe

rs; th

e Df d

omina

nt mu

tation

is le

thal

expr

essiv

ity.

in ho

mozy

gotes

.UB

C PC

-WB

Pc m

utatio

n rep

orted

in U

BC W

inAu

tosom

al re

cess

ives.

pc, w

bPo

or24

M/48

FNo

Yes

Chen

gPo

rcupin

e muta

nts re

prod

uce p

oorly

and h

ave f

eathe

rs19

79; w

b muta

tion r

epor

ted in

UBC

Atha

t res

emble

porcu

pine q

uills

(the v

anes

do no

t unc

oil);

in 19

75; c

ombin

ed 19

95wh

ite-b

reas

ted m

utants

have

whit

e fea

thers

on fa

ce,

ventr

al ne

ck, a

nd br

east;

wing

fligh

t feath

ers (

prim

aries

and m

ost s

econ

darie

s) ar

e also

whit

e.UB

C RF

-RH

Rf m

utatio

n rep

orted

in U

BC B

inAu

tosom

al re

cess

ives.

rf, erh

Poor

24M/

48F

NoYe

sCh

eng

Ruffle

d hom

ozyg

otes h

ave c

urve

d, be

nt or

ruffle

d19

83; r

edhe

ad (e

rh) f

rom

Virg

inia

feathe

rs (rf

muta

tion n

ot ye

t rep

orted

in th

e lite

ratur

e).

Poly.

Inst.

in 19

77; c

ombin

ed 19

95UB

C RT

Mutat

ion re

porte

d in U

BC A

in 19

77;

Autos

omal

rece

ssive

.rt

Comb

ined

see:

UBC

MNo

Yes

Chen

gRo

ugh-

textu

red h

omoz

ygote

s hav

e fea

thers

that a

ppea

rco

mbine

d with

UBC

M in

1989

with

UBC

matte

d and

roug

h, an

d the

fema

les pr

oduc

e few

er vi

able

M in

1989

embr

yos.

UBC

SBMu

tation

repo

rted i

n UBC

A in

1982

;Au

tosom

al re

cess

ive.

sbCo

mbine

dse

e: UB

C F-

SBNo

Yes

Chen

gSh

ort d

istal

barb

muta

nts sh

ow fu

sion i

n the

dista

l bar

bspo

oled w

ith U

BC F

with

UBC

Fof

conto

ur fe

ather

s dur

ing fe

ather

grow

th; ba

rbs a

re ab

out

1/4 no

rmal

length

, mos

t visi

ble in

the c

ontou

r fea

thers

ofthe

back

.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

26

Japa

nese

qua

il (c

ont.)

Mut

ant-D

evel

opm

enta

l def

ect-S

kin/

feat

her (

cont

.)UB

C SP

Mutat

ion fo

und i

n UBC

AL i

n 198

8;Au

tosom

al re

cess

ive.

spCo

mbine

dse

e: UB

C B

NoYe

sCh

eng

Spad

e hom

ozyg

otes h

ave d

efecti

ve fe

ather

s (sp

comb

ined w

ith U

BC B

in 19

96wi

th UB

C B

mutat

ion no

t yet

repo

rted i

n the

liter

ature

).in

1997

Mut

ant-G

ene

pool

Wisc

onsin

Japa

nese

Qua

ilAc

quire

d fro

m U

Autos

omal

rece

ssive

.wh

, we

Good

:15

0M/35

0FNo

Yes

Wen

twor

thHo

mozy

gous

wh b

irds h

ave w

hite f

eathe

rs; ho

mozy

gous

Mass

achu

setts

-Amh

erst

in 19

69NA

SAwe

fema

les la

y cha

lk-wh

ite eg

gs.

Mut

ant-U

ncat

egor

ized

UBC

H5Mu

tation

repo

rted i

n UBC

WILD

inInc

omple

te do

mina

nt.Po

or48

M/96

FNo

Yes

Chen

gMu

tation

caus

es H

5 hist

ones

to fo

rm di

mers

in vit

ro.

1992

; cro

ssed

with

UBC

A in

1997

Ran

dom

bred

Arka

nsas

RBC

Acqu

ired f

rom

Easte

rn S

hore

as a

Good

36M/

36F

NoAn

thon

yRa

ndom

bred

contr

ol str

ain fr

om E

aster

n Sho

rera

ndom

bred

contr

ol in

1990

Rand

ombr

ed.

Athe

ns C

ontro

l Qua

ilKe

pt as

clos

ed flo

ck si

nce 1

963

Good

120M

/120F

NoYe

sBu

rke

Rand

ombr

ed co

ntrol,

with

whit

e egg

shell

muta

tion i

n the

gene

pool,

prop

agate

d by r

ando

m pa

ir-ma

tings

.Lo

uisia

na R

ando

mbr

ed Q

uail

Kept

at Lo

uisian

a Stat

e as c

losed

flock

Good

60M/

120F

NoYe

sSa

tterle

eUn

selec

ted, r

ando

mbre

d pop

ulatio

n with

some

color

for ov

er 20

year

smu

tation

s (tu

xedo

, red

head

, whit

e egg

).Oh

io R

1De

rived

from

a cro

ss of

Athe

nsPo

or36

M/36

FNo

Yes

Nest

orPr

opag

ated u

sing 3

6 pair

s eac

h gen

erati

on.

Rand

ombr

ed, A

thens

whit

e egg

, and

Wisc

onsin

stoc

k; ke

pt as

clos

ed flo

ckfor

38 ge

nera

tions

UBC

ADe

rived

from

a cro

ss of

UCD

Poor

48M/

96F

NoYe

sCh

eng

Rand

ombr

ed flo

ck.

Rand

ombr

ed qu

ail an

d qua

il stoc

kfro

m Ko

rea;

kept

as cl

osed

flock

for

over

70 ge

nera

tions

UBC

BAc

quire

d fro

m U

Albe

rta in

1977

,sp

Poor

24M/

48F

NoYe

sCh

eng

Exce

ption

ally n

ervo

us ra

ndom

bred

; spa

de ho

mozy

gotes

comb

ined w

ith U

BC S

P (sp

ade

have

defec

tive f

eathe

rs.mu

tation

, affe

cting

feath

ers)

by 19

98UB

C M

UBC

M fro

m a c

omme

rcial

(Mar

shAu

tosom

al re

cess

ive.

rtPo

or24

M/48

FNo

Yes

Chen

gRo

ugh-

textu

red h

omoz

ygote

s hav

e fea

thers

that a

ppea

rFa

rms)

strain

in 19

75; c

ombin

edma

tted a

nd ro

ugh,

and t

he fe

males

prod

uce f

ewer

viab

lewi

th UB

C RT

in 19

89em

bryo

s (rt

mutat

ion no

t yet

repo

rted i

n the

liter

ature

);UB

C M

have

the e

xtend

ed b

rown

allel

e, an

d are

sligh

tlyhe

avier

than

UBC

A.

UBC

NAc

quire

d fro

m U

Nago

ya (J

apan

) inPo

or24

M/48

FNo

Yes

Chen

gVe

ry do

cile r

ando

mbre

d.19

88

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

27

Japa

nese

qua

il (c

ont.)

Ran

dom

bred

(con

t.)UB

C NC

Acqu

ired f

rom

NCSU

-rand

ombr

edPo

or24

M/48

FNo

Yes

Chen

gRa

ndom

bred

that

has p

rove

n to b

e ver

y sen

sitive

toqu

ail in

1990

chan

ges i

n pho

toper

iod.

UBC

SAc

quire

d fro

m U

Sask

atche

wan i

nPo

or24

M/48

FNo

Yes

Chen

g19

83UB

C W

ILD

Foun

datio

n stoc

k con

sisted

of 12

Poor

48M/

96F

NoYe

sCh

eng

feral

birds

caug

ht in

Hawa

ii in 1

985

UCD

Rand

ombr

ed Q

uail

Deriv

ed fr

om st

ock i

mpor

ted fr

omGo

od20

0+No

Yes

Wils

onW

ild-ty

pe fe

ather

color

patte

rn, u

nsele

cted,

rand

omly

Japa

n and

Taiw

an (1

950s

and 1

972)

grou

ped i

n colo

ny ca

ges (

2M/4F

); re

prod

uced

ever

y six

to eig

ht mo

nths.

UMar

yland

Ran

dom

bred

Qua

ilAc

quire

d fro

m U

Wisc

onsin

/USD

A in

Good

100

NoW

ithOt

tinge

rthe

1970

s; ke

pt as

clos

ed flo

ck fo

rco

llabo

ratio

nap

prox

imate

ly 20

year

sUN

L W

ild-ty

pe C

otur

nix

Acqu

ired f

rom

U Ge

orgia

-Athe

nsFa

ir: Ha

tch,

30-6

0 pair

sNo

With

Beck

regio

nal,

colla

bora

tion

misc

.

Sele

cted

-Beh

avio

ral t

rait

Purd

ue C

otur

nix K

GBDe

rived

from

Athe

ns C

ontro

l qua

il (U

Good

:10

,000 b

irds

NoW

ithMu

irSe

lected

for n

on-a

ggre

ssive

beha

vior s

tartin

g in 1

988.

Geor

gia-A

thens

)Ind

ustry

colla

bora

tion

Sele

cted

-Gro

wth

trai

tAr

kans

as H

10De

rived

from

Arka

nsas

RBC

Good

36M/

36F

NoAn

thon

ySe

lected

18 ge

nera

tions

for h

igh 10

-day

body

weig

ht.Ar

kans

as H

17De

rived

from

Arka

nsas

RBC

Good

36M/

36F

NoAn

thon

ySe

lected

18 ge

nera

tions

for h

igh 17

-day

body

weig

ht.Ar

kans

as H

28De

rived

from

Arka

nsas

RBC

Good

36M/

36F

NoAn

thon

ySe

lected

18 ge

nera

tions

for h

igh 28

-day

body

weig

ht.Ar

kans

as H

40De

rived

from

Arka

nsas

RBC

Good

36M/

36F

NoAn

thon

ySe

lected

18 ge

nera

tions

for h

igh 40

-day

body

weig

ht.Ar

kans

as H

LGo

od36

M/36

FNo

Anth

ony

Selec

ted fo

r high

early

body

weig

ht ga

in (1

0-17

days

),low

later

body

weig

ht ga

in (1

7-28

days

).Ar

kans

as L

HGo

od36

M/36

FNo

Anth

ony

Selec

ted fo

r low

early

body

weig

ht ga

in (1

0-17

days

),hig

h late

r bod

y weig

ht ga

in (1

7-28

days

).At

hens

52De

rived

from

a cro

ss of

Athe

ns 51

Good

36M/

36F

NoYe

sBu

rke

High

body

weig

ht se

lected

.an

d 53 (

both

selec

ted 38

gene

ratio

nsfor

high

4-we

ek bo

dy w

eight)

Athe

ns 54

Deriv

ed fr

om a

cross

of tw

o stoc

ks,

Good

36M/

36F

NoYe

sBu

rke

Low

body

weig

ht se

lected

.bo

th se

lected

38 ge

nera

tions

for lo

w4-

week

body

weig

ht

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

28

Japa

nese

qua

il (c

ont.)

Sele

cted

-Gro

wth

trai

tAt

hens

56De

rived

from

a cro

ss of

Athe

ns P

-,Go

od36

M/36

FNo

Yes

Burk

eInt

erme

diate

body

weig

ht sto

ck re

sultin

g fro

m a c

ross

ofT-

, and

S-lin

eslon

g-ter

m se

lected

high

and l

ow bo

dy w

eight

stock

s.At

hens

P-li

neDe

rived

from

Athe

ns C

ontro

l qua

ilGo

od60

M/60

FNo

Yes

Burk

eSe

lected

100 g

ener

ation

s for

high

4-we

ek bo

dy w

eight

on a

28%

prote

in die

t; at 7

0 gen

erati

ons,

the ad

ult si

zewa

s ove

r 150

% ab

ove t

he co

ntrol-

stand

ard p

opula

tion.

Athe

ns T

-line

Deriv

ed fr

om A

thens

Con

trol q

uail

Good

60M/

60F

NoYe

sBu

rke

Selec

ted 10

0 gen

erati

ons f

or hi

gh 4-

week

body

weig

htwh

ile un

der a

low

prote

in an

d thio

urac

il stre

ss di

et;re

sists

grow

th de

pres

sion o

n diet

s with

up to

0.2%

thiou

racil

.Oh

io H

WDe

rived

from

Ohio

R1

Poor

48M/

48F

NoYe

sNe

stor

Inbre

eding

coeff

icien

t (F)

=0.41

7; se

lected

30 ge

nera

tions

for in

creas

ed 4-

week

body

weig

ht.Oh

io H

W-H

PDe

rived

from

Ohio

HW

Poor

36M/

36F

NoYe

sNe

stor

Inbre

eding

coeff

icien

t (F)

=0.37

5; se

lected

for m

aleinc

reas

ed 4-

week

body

weig

ht, fo

r fem

ale in

creas

edpla

sma p

hosp

horu

s (ind

icator

of yo

lk pr

ecur

sors)

.Oh

io H

W-L

PDe

rived

from

Ohio

HW

Poor

36M/

36F

NoYe

sNe

stor

Inbre

eding

coeff

icien

t (F)

=0.35

9; se

lected

for m

aleinc

reas

ed 4-

week

body

weig

ht, fo

r fem

ale de

creas

edpla

sma p

hosp

horu

s two

wee

ks af

ter st

art o

f lay.

Ohio

LW

Deriv

ed fr

om O

hio R

1Po

or48

M/48

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F)=0

.357;

selec

ted 30

gene

ratio

nsfor

decre

ased

4-we

ek bo

dy w

eight.

UBC

G-QM

UBC

G de

rived

from

a co

mmer

cial

Poor

24M/

48F

NoYe

sCh

eng

Selec

ted fo

r ave

rage

fema

le bo

dy w

eight

of 28

0 g at

6str

ain (M

arsh

Far

ms);

comb

ined w

ithwe

eks.

UBC

QM in

1992

UBC

QFAc

quire

d fro

m De

scha

mbau

lt in 1

990

Poor

48M/

96F

NoYe

sCh

eng

Selec

ted fo

r ave

rage

fema

le bo

dy w

eight

of 28

0 g at

6we

eks.

UBC

QMAc

quire

d fro

m De

scha

mbau

lt in

Comb

ined

see:

UBC

G-QM

NoNo

Chen

gSe

lected

for a

vera

ge m

ale bo

dy w

eight

of 28

0 g at

619

90; c

ombin

ed w

ith U

BC G

in 19

92wi

th UB

Cwe

eks.

G in

1992

Sele

cted

-Imm

une

trai

tAt

hens

AR3

.0Go

od36

M/36

FNo

Yes

Burk

eSe

lected

for r

esist

ance

to af

latox

in.

Sele

cted

-Phy

siol

ogic

al tr

ait

Athe

ns H

-PCH

OLDe

rived

from

Athe

ns C

ontro

l qua

ilGo

od30

M/30

FNo

Yes

Burk

eSe

lected

for h

igh bl

ood p

lasma

chole

stero

l for 3

7ge

nera

tions

; cur

rentl

y rela

xed s

electi

on.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Japa

nese

qua

il ge

netic

sto

cks

Appe

ndix

2 T

able

2.1

29

Japa

nese

qua

il (c

ont.)

Sele

cted

-Phy

siol

ogic

al tr

ait (

cont

.)At

hens

L-P

CHOL

Deriv

ed fr

om A

thens

Con

trol q

uail

Good

30M/

30F

NoYe

sBu

rke

Selec

ted fo

r low

blood

plas

ma ch

oleste

rol fo

r 37

gene

ratio

ns; c

urre

ntly r

elaxe

d sele

ction

.Lo

uisia

na H

igh

Stre

ss R

espo

nse

Deriv

ed fr

om Lo

uisian

a Ran

domb

red

Good

60M/

120F

NoYe

sSa

tterle

eSe

lected

over

ten y

ears

for hi

gh bl

ood c

ortic

oster

oidQu

aillev

els in

resp

onse

to st

ress

.Lo

uisia

na L

ow S

tress

Res

pons

eDe

rived

from

Louis

iana R

ando

mbre

dGo

od60

M/12

0FNo

Yes

Satte

rlee

Selec

ted ov

er te

n yea

rs for

low

blood

cortic

oster

oidQu

aillev

els in

resp

onse

to st

ress

.Oh

io H

PDe

rived

from

Ohio

R1

Poor

36M/

36F

NoYe

sNe

stor

Inbre

eding

coeff

icien

t (F)

=0.36

; sele

cted 3

0 gen

erati

ons

for in

creas

ed pl

asma

phos

phor

us in

the f

emale

two

week

s afte

r star

t of la

y.Oh

io L

PDe

rived

from

Ohio

R1

Poor

36M/

36F

NoYe

sNe

stor

Inbre

eding

coeff

icien

t (F)

=0.39

4; se

lected

30 ge

nera

tions

for de

creas

ing le

vels

of pla

sma p

hosp

horu

s in t

hefem

ale tw

o wee

ks af

ter st

art o

f lay.

UBC

RES

Acqu

ired f

rom

NCSU

in 19

88Po

or24

M/48

FNo

Yes

Chen

gSe

lected

for r

esist

ance

to at

hero

scler

osis

when

fed h

ighch

oleste

rol d

iets.

UBC

SUS

Acqu

ired f

rom

NCSU

in 19

88Po

or48

M/96

FNo

Yes

Chen

gSe

lected

for s

usce

ptibil

ity to

athe

rosc

leros

is wh

en fe

dhig

h cho

lester

ol die

ts.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Turk

ey g

enet

ic s

tock

sAp

pend

ix 2

Tab

le 2

.130

Turk

eyM

utan

t-Dev

elop

men

tal d

efec

t-Eye

UMas

s Glau

com

aNa

turall

y mati

ng, e

xhibi

tion-

type

Slate

gaDi

sper

sal

10M/

25F

NoYe

sSm

yth

Glau

coma

homo

zygo

tes te

nd to

deve

lop gl

auco

ma an

dtur

key;

in the

proc

ess o

f tran

sfer t

oin

1998

beco

me bl

ind.

priva

te cu

rator

.

Mut

ant-R

epro

duct

ive

defe

ctGu

elph

Parth

enog

enet

ic Tu

rkey

Origi

nally

deve

loped

at U

SDA

Belts

ville

Small

multif

actor

ialPo

or10

M/30

FNo

Yes

Etch

esSe

lected

for s

ponta

neou

s dev

elopm

ent o

f 40%

labor

atorie

s in B

eltsv

ille, M

D, by

Whit

eun

fertili

zed e

ggs (

parth

enog

eneti

c dev

elopm

ent),

with

Olse

n in t

he 19

70s

0.5%

of de

velop

ing em

bryo

s (all

male

) hatc

hing.

Pure

bre

edNC

SU B

lack S

pani

shAc

quire

d fro

m Oh

io St

ate U

in 19

80;

Blac

k Spa

nish

Fair

5M/10

FNo

Yes

Chris

tens

enke

pt as

clos

ed flo

ck us

ed in

rese

arch

and t

each

ingNC

SU B

ronz

eAc

quire

d fro

m Oh

io St

ate U

in 19

80;

Bron

zeFa

ir5M

/15F

NoYe

sCh

riste

nsen

kept

as cl

osed

flock

used

inre

sear

ch an

d tea

ching

NCSU

Slat

eAc

quire

d fro

m Oh

io St

ate U

in 19

80;

Slate

Fair

5M/15

FNo

Yes

Chris

tens

enke

pt as

clos

ed flo

ck us

ed in

rese

arch

and t

each

ingSa

skat

chew

an B

ronz

e Tur

key

Acqu

ired f

rom

Ridle

y (a b

reed

er in

Bron

zeGo

od30

M/80

FNo

With

Clas

sen

Rese

mbles

comm

ercia

l turke

ys fr

om th

e 194

0s (c

arca

ssSa

skato

on ar

ea) in

the 1

980s

;co

llabo

ratio

nan

d pro

ducti

on ch

arac

terist

ics) a

nd is

able

to br

eed

rese

mbles

old c

omme

rcial

stock

sna

turall

y; us

ed in

card

iac, d

igesti

ve, a

nd re

prod

uctiv

eph

ysiol

ogy t

each

ing an

d res

earch

.W

iscon

sin M

idge

t Whi

teDe

velop

ed by

J. S

myth

(UMi

dget

Whit

eGo

od12

0No

Yes

Wen

twor

thMi

dget

white

turke

y, ma

les=1

2 lb,

female

s=8lb

.Ma

ssac

usett

s); ac

quire

d in 1

972

and k

ept a

s clos

ed flo

ck

Ran

dom

bred

NADC

Tur

key

Origi

nally

from

USD

A Be

ltsvil

leBe

ltsvil

le Sm

allGo

od70

-90

NoYe

sRi

mler

Natio

nal A

nimal

disea

se C

enter

rese

arch

stoc

k.tur

key s

tocks

; kep

t as c

losed

flock

Whit

efor

over

30 ye

ars;

rece

ntly s

ent

to So

uthea

st Po

ultry

Res.

Lab.

NCSU

Ohi

o-RB

C1Ac

quire

d fro

m Oh

io St

ate U

in 19

80;

Larg

e Whit

eGo

od10

M/60

FNo

Yes

Chris

tens

enke

pt as

clos

ed flo

ckNC

SU O

hio-

RBC2

Acqu

ired f

rom

Ohio

State

U in

1980

;La

rge W

hite

Good

10M/

60F

NoYe

sCh

riste

nsen

kept

as cl

osed

flock

Ohio

RBC

1De

rived

from

a cro

ss of

four

Larg

e Whit

ePo

or36

M/36

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F) ap

prox

imate

ly 0.1

48.

comm

ercia

l turke

y stra

ins in

1957

Ohio

RBC

2De

rived

from

a cro

ss of

two

Larg

e Whit

ePo

or36

M/36

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F) ap

prox

imate

ly 0.1

2.co

mmer

cial tu

rkey s

trains

in 19

66

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Turk

ey g

enet

ic s

tock

sAp

pend

ix 2

Tab

le 2

.131

Turk

ey (c

ont.)

Ran

dom

bred

(con

t.)Oh

io R

BC3

Deriv

ed fr

om a

cross

of O

hio F

Line

Larg

e Whit

ePo

or36

M/36

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F) ap

prox

imate

ly 0.0

33.

with

comm

ercia

l turke

y stra

ins in

1986

Wisc

onsin

Bro

ad-b

reas

ted

Bron

zeDe

rived

from

Tryl

or B

ronz

e of

Broa

d-br

easte

dGo

od30

M/20

0FSe

men,

PGC

Yes

Wen

twor

thOl

d com

merci

al-typ

e tur

key.

Jero

me O

rgan

izatio

n in N

. Wisc

.;Br

onze

kept

as cl

osed

flock

sinc

e 196

9W

iscon

sin B

road

-bre

aste

d W

hite

Deriv

ed fr

om N

ichola

s X R

alston

Broa

d-br

easte

dGo

od30

M/20

0FSe

men,

PGC

Yes

Wen

twor

thOl

d com

merci

al-typ

e tur

key.

synth

etic m

eat c

ross

deve

loped

atW

hite

Was

hingto

n Stat

e U; k

ept a

sclo

sed f

lock s

ince 1

969

Sele

cted

-Egg

trai

tNC

SU O

hio-

EDe

rived

from

Ohio

-E; k

ept a

s La

rge W

hite

Good

10M/

60F

NoYe

sCh

riste

nsen

Selec

ted fo

r incre

ased

egg p

rodu

ction

.clo

sed f

lock

Ohio

E L

ine

Deriv

ed fr

om O

hio R

BC1

Larg

e Whit

ePo

or72

M/72

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F)=0

.478;

selec

ted 38

gene

ratio

nsfor

incre

ased

egg p

rodu

ction

.

Sele

cted

-Gro

wth

trai

tNC

SU O

hio-

FDe

rived

from

Ohio

-F; k

ept a

s La

rge W

hite

Good

10M/

60F

NoYe

sCh

riste

nsen

Selec

ted fo

r incre

ased

body

weig

ht at

16 w

eeks

.clo

sed f

lock

Ohio

F L

ine

Deriv

ed fr

om O

hio R

BC2

Larg

e Whit

ePo

or36

M/72

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F)=0

.26; s

electe

d 30 g

ener

ation

sfor

incre

ased

16-w

eek b

ody w

eight.

Ohio

FL

Line

Deriv

ed fr

om O

hio F

line

Larg

e Whit

ePo

or36

M/54

FNo

Yes

Nest

orInb

reed

ing co

effici

ent (

F)=0

.274;

selec

ted 17

gene

ratio

nsfor

incre

ased

shan

k widt

h.

Tabl

e 2.

1 St

ock

info

rmat

ion

Cat

egor

yGe

netic

Allel

eNu

mbe

rCr

yo-

Stoc

k nam

e and

des

crip

tion

Orig

in an

d hi

stor

yBr

eed

char

acte

ristic

ssy

mbo

lSt

atus

of b

irds

pres

erv.

Avail

able

Cura

tor

Wat

erfo

wl/G

ameb

ird g

enet

ic s

tock

sAp

pend

ix 2

Tab

le 2

.132

Wat

erfo

wl/G

ameb

irdB

lood

type

-Gen

e po

olNI

U Bo

bwhi

tes

Acqu

ired f

rom

Miss

issipp

i No

rther

nse

eFa

ir50

+No

With

Brile

sVa

rious

eryth

rocy

te all

oanti

gens

in qu

ail.

State

U in

1992

Bobw

hite

desc

riptio

nco

llabo

ratio

nNI

U Ph

easa

nts

Acqu

ired f

rom

state

game

bird

Ring

-nec

ked

see

Fair

50+

NoW

ithBr

iles

Popu

lation

segr

egati

ng fo

r a va

riety

of MH

C B

hatch

eries

in W

iscon

sin an

d Ph

easa

ntde

scrip

tion

colla

bora

tion

haplo

types

.Illi

nois

in 19

86

Pure

bre

edAD

OL P

ekin

Duc

kKe

pt as

clos

ed flo

ck si

nce

Pekin

Duc

kGo

od:

8M/35

FNo

Yes

Baco

nSp

ecific

patho

gen f

ree,

prob

ably

mode

ratel

y inb

red,

the 19

60s

USDA

repr

oduc

ed 2X

/year

.Co

rnell

Whi

te P

ekin

Spec

ific pa

thoge

n-fre

ePe

kin D

uck

Good

9M/39

FNo

Yes,

feeSc

hat

Comm

ercia

l duc

k stoc

k.NC

SU B

rown

Chi

naKe

pt as

clos

ed flo

ck; u

sed i

n Br

own C

hines

eFa

ir10

M/15

FNo

Yes

Chris

tens

enre

sear

ch an

d tea

ching

Goos

eNC

SU P

ilgrim

Kept

as cl

osed

flock

; use

d in

Pilgr

im G

oose

Fair

10M/

15F

NoYe

sCh

riste

nsen

rese

arch

and t

each

ingSa

skat

chew

an P

ilgrim

Goo

seAc

quire

d fro

m Ag

ricult

ure-

Cana

daPi

lgrim

Goo

seGo

od9M

/18F

NoW

ithCl

asse

nGo

ose b

reed

with

sexu

ally d

imor

phic

color

s; us

ed in

(CFA

R) in

1965

colla

bora

tion

teach

ing bi

rd ha

ndlin

g in a

gricu

lture

and v

eterin

ary

medic

ine cl

asse

s.Sa

skat

chew

an R

ouen

Duc

kAc

quire

d fro

m Mi

ller H

atche

ries i

nRo

uen D

uck

Good

12M/

24F

NoW

ithCl

asse

nUs

ed in

teac

hing b

ird ha

ndlin

g in a

gricu

lture

and

Sask

atoon

in 19

65co

llabo

ratio

nve

terina

ry me

dicine

clas

ses.

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

CANADA—BC University of British ColumbiaCurator: Kimberly Cheng

ChickenMutant-Developmental defect-Eye

UBC RCMutant-Gene pool

UBC-Minnesota MarkerPure breed

UBC-Minnesota Rhode Island Red

Japanese quailInbred

UBC IMutant-Color, eggshell

UBC CEUBC WE

Mutant-Color, featherUBC BHUBC C-CEUBC DUBC F-SBUBC HUBC RHUBC SIUBC W-WEUBC WBUBC Y

Mutant-Developmental defect-Skin/featherUBC DF-MDFUBC PC-WBUBC RF-RHUBC RTUBC SBUBC SP

Mutant-UncategorizedUBC H5

RandombredUBC AUBC BUBC MUBC NUBC NCUBC SUBC WILD

Selected-Growth traitUBC G-QMUBC QFUBC QM

Selected-Physiological traitUBC RESUBC SUS

CANADA—ON University of GuelphCurator: Robert J. Etches

ChickenChromosomal variant

Ottawa B-19/B-19 M13 Cryo. onlyOttawa B-21/B-21 M13 Cryo. only

InbredOttawa GF Cryo. onlyOttawa GH Cryo. onlyOttawa M2 Cryo. onlyOttawa WG Cryo. onlyOttawa XP Cryo. only

Pure breedGuelph SilkieOttawa New Hampshire Cryo. only

RandombredOttawa 10 Cryo. onlyOttawa 20 Cryo. onlyOttawa 30 Cryo. onlyOttawa 5 Cryo. onlyOttawa 7 Cryo. only

Selected-Egg traitGuelph Barred Plymouth RockGuelph Commercial Leghorn LineOttawa 1 Cryo. onlyOttawa 2 Cryo. onlyOttawa 3 Cryo. onlyOttawa 4 Cryo. onlyOttawa 8 Cryo. onlyOttawa 9 Cryo. only

Selected-Growth traitOttawa 21 Cryo. onlyOttawa 23 Cryo. onlyOttawa 25 Cryo. onlyOttawa 31 Cryo. only

Selected-Immune traitOttawa 2R Cryo. onlyOttawa 3R Cryo. onlyOttawa 8R Cryo. only

TransgenicOttawa TR Cryo. only

UncategorizedOttawa 6 Cryo. onlyOttawa 80 Cryo. onlyOttawa 90 Cryo. onlyOttawa N3 Cryo. onlyOttawa N4 Cryo. onlyOttawa N8 Cryo. only

TurkeyMutant-Reproductive defect

Guelph Parthenogenetic Turkey

Appendix 2 Table 2.2 1

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

CANADA—SK University of SaskatchewanCurator: Henry L. Classen

ChickenMutant-Neurological defect

Saskatchewan Epileptiform seizuresPure breed

Saskatchewan Barred Plymouth RockSaskatchewan Brown Leghorn

TurkeyPure breed

Saskatchewan Bronze Turkey

Waterfowl/GamebirdPure breed

Saskatchewan Pilgrim GooseSaskatchewan Rouen Duck

USA—AL Auburn UniversityCurator: Wallace Berry Other contact: Gayner R. McDaniel

ChickenSelected-Growth trait

Auburn Tibial Dyschondroplasia-HighAuburn Tibial Dyschondroplasia-Low

USA—AL Auburn UniversityCurator: Sandra Ewald

ChickenBloodtype-Gene pool

Auburn DAXBloodtype-MHC

Auburn MAuburn NAuburn RMAuburn RMHAuburn RN

Selected-Immune traitAuburn RAuburn S

USA—AR University of Arkansas–FayettevilleCurator: N.B. Anthony

ChickenPure breed

Arkansas Giant Jungle FowlRandombred

Arkansas RandombredSelected-Physiological trait

Arkansas Ascites ResistantArkansas Ascites Susceptible

Japanese quailMutant-Color, feather

Arkansas English WhiteArkansas White

RandombredArkansas RBC

USA—AR University of Arkansas–FayettevilleCurator: N.B. Anthony

Selected-Growth traitArkansas H10Arkansas H17Arkansas H28Arkansas H40Arkansas HLArkansas LH

USA—AR University of Arkansas–FayettevilleCurator: Gisela Erf

ChickenMutant-Immunological defect

Arkansas Smyth Line B101Arkansas Smyth Line B102

RandombredArkansas Brown Line B101Arkansas Brown Line B102Arkansas Light Brown Leghorn B101

Selected-Immune traitArkansas ProgressorArkansas Regressor

USA—AR University of Arkansas–FayettevilleCurator: John Kirby

ChickenMutant-Reproductive defect

Arkansas Sd Line

USA—CA University of California–DavisCurator: Ursula K. Abbott Other contact: J.M. Pisenti

ChickenMutant-Developmental defect-Face/limb

UCD Cleft Primary Palate/ScalelessUCD Coloboma X 003UCD Diplopodia-1 X 003UCD Diplopodia-3 X 003UCD Donald-duck Beak/ScalelessUCD Eudiplopodia X 003UCD Limbless X 003UCD Polydactyly X 003UCD Stumpy X 003UCD Talpid-2 X 003UCD Wing-reduced X 003UCD Wingless-2 X 331

Mutant-Developmental defect-Skin/featherUCD Ichthyosis X 003UCD Naked-neck Cryo. onlyUCD Scaleless-HighUCD Scaleless-Low

Mutant-Developmental defect-Spine/tailUCD Scoliosis Cryo. only

Appendix 2 Table 2.2 2

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

USA—CA University of California–DavisCurator: Ursula K. Abbott Other contact: J.M. Pisenti

Mutant-Gene poolUCD Diplopodia-3/Scaleless HighUCD Eudiplopodia/LimblessUCD Limbless/StumpyUCD Silkie crossUCD Talpid-2/Wingless-2

Mutant-Physiological defectUCD Crooked-neck DwarfUCD Riboflavin Transfer Deficient

Mutant-UncategorizedUCD Crest/Hemoglobin-D Cryo. onlyUCD Multiplex Comb Cryo. only

Pure breedUCD Silkie

USA—CA University of California–DavisCurator: Hans Abplanalp

ChickenBloodtype-MHC-Inbred

UCD 003UCD 253UCD 254UCD 312UCD 313 Cryo. onlyUCD 330UCD 331UCD 335UCD 336UCD 342UCD 361UCD 380UCD 386UCD 387

Mutant-Immunological defectUCD 200

Mutant-Physiological defectUCD 077

Selected-Egg traitUCD 058UCD 082

USA—CA University of California–DavisCurator: Mary E. Delany

ChickenChromosomal variant

UCD-Cornell Mono-PNUUCD-Cornell Trisomic

InbredUCD 001

Mutant-Physiological defectUCD-Cornell Autosomal Dwarf

USA—CA University of California–DavisCurator: Barry Wilson

ChickenMutant-Physiological defect

UCD 413Randombred

UCD 412

Japanese quailRandombred

UCD Randombred Quail

USA—CT University of ConnecticutCurator: Louis J. Pierro

ChickenMutant-Developmental defect-Face/limb

Storrs ChondrodystrophyStorrs CreeperStorrs Diplopodia-3Storrs Diplopodia-5Storrs LimblessStorrs Micromelia-AbbottStorrs Micromelia-HaysStorrs NanomeliaStorrs PerocephalyStorrs PolydactylyStorrs Talpid-2Storrs Wingless-2

Mutant-Developmental defect-Skin/featherStorrs Ottawa NakedStorrs PtilopodyStorrs Scaleless

Mutant-Developmental defect-Spine/tailStorrs Dominant RumplessnessStorrs Recessive Rumpless

Mutant-Physiological defectStorrs Crooked-neck DwarfStorrs Muscular Dystrophy

Mutant-UncategorizedStorrs Rose Comb

USA—DE University of DelawareCurator: Harold B. White III

ChickenMutant-Physiological defect

UDel Riboflavin Transfer Deficient

USA—GA University of GeorgiaCurator: William H. Burke

ChickenMutant-Physiological defect

Athens-Canadian DwarfsRandombred

Athens RandombredAthens-Canadian Randombred

Appendix 2 Table 2.2 3

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

USA—GA University of GeorgiaCurator: William H. Burke

Selected-Immune traitAthens AR

Japanese quailRandombred

Athens Control QuailSelected-Growth trait

Athens 52Athens 54Athens 56Athens P-lineAthens T-line

Selected-Immune traitAthens AR3.0

Selected-Physiological traitAthens H-PCHOLAthens L-PCHOL

USA—IA Iowa State UniversityCurator: Susan J. Lamont

ChickenBloodtype-MHC-Inbred

ISU 19-13ISU 19-15.1ISU 8-15.1ISU G-B1ISU G-B2ISU GH-1ISU GH-13ISU GH-15.1ISU HN-12ISU HN-15ISU M15.2ISU M5.1ISU Sp21.2

Mutant-UncategorizedISU S1-19HISU S1-19LISU S1-1HISU S1-1L

USA—IA National Animal Disease CenterCurator: Richard Rimler

TurkeyRandombred

NADC Turkey

USA—IL Northern Illinois UniversityCurator: W. Elwood Briles

ChickenBloodtype-Gene pool

NIU B haplotypesNIU B-haplotype RecombinantsNIU Male Breeder Alloantigen ReservoirNIU Segregating Male Breeder Line

Bloodtype-MHCNIU Female Breeder Parent Stock B19NIU Female Breeder Parent Stock B2NIU Female Breeder Parent Stock B5

Waterfowl/GamebirdBloodtype-Gene pool

NIU BobwhitesNIU Pheasants

USA—IL University of Illinois–UrbanaCurator: Carl Parsons Other contact: Bob Leeper

ChickenPure breed

UI-Urbana ColumbianUI-Urbana New Hampshire

USA—IN Purdue UniversityCurator: W.M. Muir

ChickenSelected-Behavioral trait

Purdue KGBPurdue MBB

Japanese quailSelected-Behavioral trait

Purdue Coturnix KGB

USA—LA Louisiana State University–Baton RougeCurator: Daniel G. Satterlee

Japanese quailRandombred

Louisiana Randombred QuailSelected-Physiological trait

Louisiana High Stress ResponseLouisiana Low Stress Response

USA—MA University of Massachusetts–AmherstCurator: J. Robert Smyth Jr

ChickenMutant-Developmental defect-Eye

UMass Pink EyeMutant-Immunological defect

UMass Smyth LinePure breed

UMass Light Brown LeghornRandombred

UMass Brown Line

Appendix 2 Table 2.2 4

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

USA—MA University of Massachusetts–AmherstCurator: J. Robert Smyth Jr

TurkeyMutant-Developmental defect-Eye

UMass Glaucoma

USA—MD University of Maryland–College ParkCurator: Mary Ann Ottinger

Japanese quailRandombred

UMaryland Randombred Quail

USA—MI USDA-ARS-Avian Disease and Oncology LaboratoryCurator: Larry D. Bacon Other contact: Laura Parks

ChickenBloodtype-MHC

RPRL-Cornell JM-NRPRL-Cornell JM-P

Bloodtype-MHC-InbredRPRL 15.15-5RPRL 15.6-2RPRL 15.7-2RPRL 15.C-12RPRL 15.N-21RPRL 15.P-13RPRL 15.P-19RPRL 15I5RPRL 7I1RPRL Line 0

Endogenous virusRPRL 15B1

Endogenous virus-InbredRPRL 100BRPRL 6I3RPRL 7I2RPRL Reaseheath Line C

USA—MI USDA-ARS-Avian Disease and Oncology LaboratoryCurator: Larry D. Bacon Other contact: Laura Parks

InbredADOL 6C.7A ReConADOL 6C.7B ReConADOL 6C.7C ReConADOL 6C.7D ReConADOL 6C.7F ReConADOL 6C.7G ReConADOL 6C.7I ReConADOL 6C.7J ReConADOL 6C.7K ReConADOL 6C.7L ReConADOL 6C.7M ReConADOL 6C.7N ReConADOL 6C.7P ReConADOL 6C.7R ReConADOL 6C.7S ReConADOL 6C.7T ReConADOL 6C.7V ReConADOL 6C.7W ReConADOL 6C.7X ReCon

TransgenicADOL Trans-ALVE6

Waterfowl/GamebirdPure breed

ADOL Pekin Duck

USA—NC North Carolina State University–RaleighCurator: Vern L. Christensen

ChickenPure breed

NCSU Barred Plymouth RockNCSU Rhode Island Red

TurkeyPure breed

NCSU Black SpanishNCSU BronzeNCSU Slate

RandombredNCSU Ohio-RBC1NCSU Ohio-RBC2

Selected-Egg traitNCSU Ohio-E

Selected-Growth traitNCSU Ohio-F

Waterfowl/GamebirdPure breed

NCSU Brown ChinaNCSU Pilgrim

Appendix 2 Table 2.2 5

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

USA—NC North Carolina State University–RaleighCurator: Muquarrab Qureshi

ChickenBloodtype-MHC-Inbred

NCSU GB-1NCSU GB-2

RandombredNCSU-Cornell K Strain

USA—NE University of Nebraska–LincolnCurator: Mary Beck

ChickenMutant-Neurological defect

UNL Paroxysm

Japanese quailRandombred

UNL Wild-type Coturnix

USA—NH University of New HampshireCurator: Robert Taylor Jr

ChickenBloodtype-Gene pool

UNH 105Bloodtype-MHC

UNH 192Bloodtype-MHC-Inbred

UNH 6.15-5UNH 6.6-2UNH-UCD 003UNH-UCD 003.R1UNH-UCD 003.R2UNH-UCD 003.R3UNH-UCD 003.R4UNH-UCD 003.R5UNH-UCD 003.R6UNH-UCD 386UNH-UCD 387UNH-UCD 3C.1

USA—NY Cornell UniversityCurator: Rodney R. Dietert

ChickenSelected-Egg trait

Cornell K Strain

USA—NY Cornell UniversityCurator: James Marsh

ChickenMutant-Immunological defect

Cornell Obese (B-13)Mutant-Physiological defect

Cornell Sex-linked DwarfRandombred

Cornell C Specials (B13)

USA—NY Cornell UniversityCurator: K.A. Schat

ChickenBloodtype-MHC

Cornell N2aCornell P2a

Selected-Immune traitCornell S13

Waterfowl/GamebirdPure breed

Cornell White Pekin

USA—OH Ohio State University–OARDCCurator: Karl E. Nestor

Japanese quailRandombred

Ohio R1Selected-Growth trait

Ohio HWOhio HW-HPOhio HW-LPOhio LW

Selected-Physiological traitOhio HPOhio LP

TurkeyRandombred

Ohio RBC1Ohio RBC2Ohio RBC3

Selected-Egg traitOhio E Line

Selected-Growth traitOhio F LineOhio FL Line

USA—OH Ohio State University–OARDCCurator: Sandra G. Velleman

ChickenMutant-Physiological defect

Ohio Low-score NormalOhio Muscular Dystrophy

USA—OR Oregon State University–CorvallisCurator: Thomas F. Savage

ChickenMutant-Gene pool

OSU Dwarf Leghorn

Appendix 2 Table 2.2 6

Species/Category/Stock nameCountry—State/Province InstitutionTable 2.2. Stocks listed by country, state or province, institution, and curator.

Species/Category/Stock nameCountry—State/Province Institution

USA—PA Pennsylvania State UniversityCurator: Guy F. Barbato

ChickenSelected-Growth trait

PSU-Athens RB 14-42H (HiK)PSU-Athens RB 14-42L (LoK)PSU-Athens RB 14HPSU-Athens RB 14LPSU-Athens RB 42HPSU-Athens RB 42L

USA—VA Virginia Polytechnic Institute and State UniversityCurator: Paul B. Siegel Other contact: E. Ann Dunnington

ChickenSelected-Growth trait

Virginia Body Weight-HighVirginia Body Weight-Low

Selected-Immune traitVirginia Antibody Line-HighVirginia Antibody Line-Low

USA—WI University of Wisconsin–MadisonCurator: James J. Bitgood Other contact: John F. Fallon

ChickenChromosomal variant

Wisconsin Chromosomal RearrangementsInbred

Wisconsin AnconaWisconsin Leghorn line UW-Sp2

Mutant-Color, featherWisconsin Autosomal Albino

Mutant-Developmental defect-EyeWisconsin Blind/CataractWisconsin Pink-eyeWisconsin Pop-eye

Mutant-Developmental defect-Face/limbWisconsin AmetapodiaWisconsin LimblessWisconsin Talpid-2Wisconsin Wingless-2

Mutant-Developmental defect-Skin/featherWisconsin Tardy Feathering

Mutant-Neurological defectWisconsin Pirouette

Mutant-Reproductive defectWisconsin Double Oviduct LineWisconsin Restricted Ovulator

Mutant-UncategorizedWisconsin Sex-linked Skin Color

RandombredWisconsin LeghornWisconsin Leghorn line UW-6XWisconsin New Hampshire

USA—WI University of Wisconsin–MadisonCurator: Bernard C. Wentworth

Japanese quailMutant-Gene pool

Wisconsin Japanese Quail

TurkeyPure breed

Wisconsin Midget WhiteRandombred

Wisconsin Broad-breasted BronzeWisconsin Broad-breasted White

Appendix 2 Table 2.2 7

Table 2.3. Curator contact information.

Appendix 2 Table 2.3 1

Ursula K. AbbottDept. of Animal ScienceUniversity of CaliforniaOne Shields AvenueDavis CA 95616 USA

Tel: (530) 752-7325Alt tel: 752-1300

FAX: (530) 752-8960ukabbott@ucdavis.edu

Hans AbplanalpDept. of Animal ScienceUniversity of CaliforniaOne Shields AvenueDavis CA 95616 USA

Tel: (530) 752-1300FAX: (530) 752-8960

N.B. AnthonyDept. of Poultry ScienceCenter for Poultry ScienceUniversity of ArkansasFayetteville AR 72701 USA

Tel: (501) 575-4833FAX: (501) 575-3026

Larry D. BaconAvian Disease and Oncology Laboratory3606 East Mount Hope RoadEast Lansing MI 48823 USA

Tel: (517) 337-6828FAX: (517) 337-6776baconld@pilot.msu.edu

Guy F. BarbatoDept. of Poultry SciencePennsylvania State UniversityUniversity Park PA 16802 USA

Tel: (814) 865-4481FAX: (814) 865-5691gfb1@email.psu.edu

Mary BeckDept. of Animal ScienceUniversity of NebraskaLincoln NE 68583 USA

Tel: (402) 472-6439FAX: (402) 472-6362mbeck@unlinfo.unl.edu

Wallace BerryDept. of Poultry ScienceAuburn UniversityAuburn University AL 36849 USA

Tel: (334) 844-2600FAX: (334) 844-2641

James J. BitgoodDept. of Animal Sciences1675 Observatory Dr.University of WisconsinMadison WI 53706 USA

Tel: (608) 263-4300FAX: (608) 262-5157bitgood@calshp.cals.wisc.edu

W. Elwood BrilesDept. of Biological SciencesNorthern Illinois UniversityDekalb IL 60115-2861 USA

Tel: (815) 753-0433FAX: (815) 753-0461

William H. BurkeDept. of Poultry Science120 Livestock-Poultry Bldg.University of GeorgiaAthens GA 30602-2772 USA

Tel: (706) 542-1358FAX: (706) 542-1827bburke@uga.cc.uga.edu

Kimberly ChengDept. of Animal ScienceUniversity of British ColumbiaVancouver BC V6T 1Z4 CANADA

Tel: (604) 822-2480FAX: (604) 822-4400kmtc@unixg.ubc.ca

Vern L. ChristensenDept. of Poultry ScienceNorth Carolina State UniversityBox 7608Raleigh NC 27695-7608 USA

Tel: (919) 515-5385Alt tel: 515-2626

FAX: (919) 515-2625vern_christensen@ncsu.edu

Henry L. ClassenDept. of Animal/Poultry ScienceUniversity of SaskatchewanSaskatoon SK S7N 5B5 CANADA

Tel: (306) 966-6600Alt tel: 966-4128

FAX: (306) 966-4560hank.classen@skyway.usask.ca

Mary E. DelanyDept. of Animal ScienceUniversity of CaliforniaOne Shields AvenueDavis CA 95616 USA

Tel: (530) 754-9343Alt tel: 752-1300

FAX: (530) 752-8960medelany@ucdavis.edu

Rodney R. DietertDept. of Microbiology/ImmunologyRice HallCornell UniversityIthaca NY 14853 USA

Tel: (607) 253-4015FAX: (607) 253-3369rrd1@cornell.edu

E. Ann DunningtonDept. of Animal/Poultry SciencesVirginia Polytechnic Institute and StateUniversityBlacksburg VA 24061-0306 USA

Tel: (540) 231-9179FAX: (540) 231-3713

Gisela ErfDept. of Poultry ScienceCenter for Poultry ScienceUniversity of ArkansasFayetteville AR 72701 USA

Tel: (501) 575-8664FAX: (501) 575-3026gferf@comp.uark.edu

Robert J. EtchesOrigen Therapeutics651 Gateway Blvd., Ste. 980South San Francisco, CA 94080 USA

Tel: (650) 827-0200FAX: (650) 827-0300retches@origentherapeutics.com

Sandra EwaldDept. of Poultry Science236 Animal Sciences Bldg.Auburn UniversityAuburn University AL 36859 USA

Tel: (334) 844-2647Alt tel: 844-2722

FAX: (334) 844-2652ewaldsj@vetmed.auburn.edu

John F. FallonDept. of Anatomy1300 University Ave.University of WisconsinMadison WI 53706-1532 USA

Tel: (608) 262-3775FAX: (608) 262-2327

John KirbyDept. of Poultry ScienceCenter for Poultry ScienceUniversity of ArkansasFayetteville AR 72701 USA

Tel: (501) 575-8623FAX: (501) 575-3026jkirby@comp.uark.edu

Table 2.3. Curator contact information.

Appendix 2 Table 2.3 2

Susan J. LamontDept. of Animal Science201 Kildee HallIowa State UniversityAmes IA 50011-3150 USA

Tel: (515) 294-4100FAX: (515) 294-2401sjlamont@iastate.edu

Bob Leeper284 Animal Science LaboratoryUniversity of Illinois1207 W. Gregory Dr.Urbana IL 61801 USA

FAX: (217) 333-7861James MarshDept. of Microbiology/Immunology C5-103College of Veterinary MedicineIthaca NY 14853 USA

Tel: (607) 253-4010FAX: (607) 253-3384jam24@cornell.edu

W.M. MuirDept. of Animal SciencePurdue UniversityWest Lafayette IN 479071151 USA

Tel: (765) 494-8032FAX: (765) 494-9346bmuir@hub.ansc.purdue.edu

Karl E. NestorDept. of Animal SciencesOhio Agricultural Reasearch andDevelopment Ctr.Ohio State UniversityWooster OH 44691 USA

Tel: (330) 263-3757FAX: (330) 263-3949nestor.1@osu.edu

Mary Ann OttingerDept. of Poultry ScienceUniversity of MarylandCollege Park MD 20742 USA

Tel: (301) 405-5780FAX: (301) 314-9059mo20@umail.umd.edu

Laura ParksAvian Disease and Oncology Laboratory3606 East Mount Hope RoadEast Lansing MI 48823 USA

Tel: (517) 337-6828FAX: (517) 337-6776parksl@pilot.msu.edu

Carl ParsonsDept. of Animal Sciences284 Animal Science Laboratory1207 W. Gregory Dr.Urbana IL 61801 USA

Tel: (217) 333-8774FAX: (217) 333-7861

Louis J. PierroStorrs Agr. Exp. Sta.University of Connecticut1376 Storrs Rd.Storrs CT 06269-4010 USA

Tel: (860) 486-0334FAX: (860) 486-4375lpierro@canrl.cag.uconn.edu

Jacqueline M. PisentiGenetic Resources Conservation ProgramUniversity of CaliforniaOne Shields AvenueDavis CA 95616 USA

Tel: (530) 754-8508FAX: (530) 754-8505jmpisenti@ucdavis.edu

Muquarrab QureshiDept. of Poultry ScienceNorth Carolina State UniversityBox 7608Raleigh NC 27695-7608 USA

Tel: (919) 515-5388FAX: (919) 515-2625m_qureshi@ncsu.edu

Richard RimlerNational Animal Disease CenterMWA/USDAP.O. Box 70Ames IA 50010 USA

Tel: (515) 239-8534Alt tel: 239-8255

FAX: (515) 239-8458

Daniel G. SatterleeDept. of Poultry ScienceLouisiana State University206 Clyde Ingram HallBaton Rouge LA 70803 USA

Tel: (504) 388-4406FAX: (504) 388-1259dsatterlee@agctr.LSU.edu

Thomas F. SavageDept. of Animal ScienceWithycombe Hall 112Oregon State UniversityCorvallis OR 97331-6702 USA

Tel: (541) 737-5066FAX: (541) 737-4174thomas.savage@orst.edu

K.A. SchatDept. of Microbiology/ImmunologyCollege of Veterinary MedicineCornell UniversityIthaca NY 14853-6401 USA

Tel: (607) 253-3365FAX: (607) 253-3369kas24@cornell.edu

Paul B. SiegelDept. of Animal/Poultry SciencesVirginia Polytechnic Institute and StateUniversityBlacksburg VA 24061-0306 USA

Tel: (540) 231-6472FAX: (540) 231-3713jacksons@vt.edu

J. Robert Smyth JrVeterinary/Animal Science307 Stockbridge HallUniversity of MassachusettsAmherst MA 01002 USA

Tel: (413) 545-2137FAX: (413) 545-6326

Robert Taylor JrDept. of Animal/Nutr. ScienceKendall HallUniversity of New HampshireDurham NH 03824 USA

Tel: (603) 862-2130FAX: (603) 862-3758bob.taylor@unh.edu

Sandra G. VellemanDept. of Animal ScienceOhio State University/OARDC1680 Madison Ave.Wooster OH 44691-4096 USA

Tel: (330) 263-3905FAX: (330) 263-3949velleman.1@osu.edu

Bernard C. WentworthDept. of Poultry Science1675 Observatory Dr., Rm. 260University of WisconsinMadison WI 53706-1284 USA

Tel: (608) 262-8945FAX: (608) 262-6005wentwort@calshp.cals.wisc.edu

Harold B. White IIIDept. of Chemistry/BiochemistryUniversity of DelawareNewark DE 19716 USA

Tel: (302) 831-2908FAX: (302) 831-6335halwhite@udel.edu

Barry WilsonDept. of Animal ScienceUniversity of CaliforniaOne Shields AvenueDavis CA 95616 USA

Tel: (530) 752-3519FAX: (530) 752-8960bwwilson@ucdavis.edu

Top right: Wild-type Japanesequail (UCD Randombred Quail)used in a wide range of experi-mental studies. Female (right) andmale (left). Photograph courtesy ofJ. Clark, University of California–Davis)Center left: Fawn mutation in theJapanese quail (UBC F-SB).(Photograph courtesy of K. Cheng,University of British Columbia)Center right: Male giant Japanesequail (UBC G-QM). More thandouble the size of unselectedquail, this line was developed byintensive selection for increasedsix-week body size in the females.(Photograph courtesy of K. Cheng,University of British Columbia)Right: Modern commercial LargeWhite turkeys. (Photographcourtesy of R.A. Ernst, Universityof California–Davis.)