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TABLE OF CONTENTS
Ernest R. Sears 50-Year Celebrat ion Symposium. G. Kimber.. ..... i
North American Wheat Genetic Mapping and Cytogenetic Stocks Workshop. C. 0. Qua lse t ................................. ii
Acknowledgements.. .............................................. i ii Program.. ....................................................... i v
S u m r y Workshop Session I. A. Lukaszewski ..................... 1
Chromosomal and Chromosome-am Locations, Linkage Estimates. and Map Locations o f T r i t i c u m aestivum Gene Loci . .......................................... Gary E. Har t 3
A Proposal f o r Wheat Chromosome Band Nomenclature. Bikram S. G i l l . . ...................................... 11
Sumnary Workshop Session 11. J. P. Gustafson ................... 16
Cytogenetic mater ia ls maintained by E. R. Sears ............ 19
A l i e n species c o l l e c t i o n s a t Un ive rs i t y o f C a l i f o r n i a , Rivers ide. J. G. Waines .............................. 20
A v a i l a b i l i t y and charac te r i za t i on o f wheat genet ic .............................. stocks. Bikram S. G i l l . 25
Wheat mutant s tock l i s t , Wheat Genetics Resource Center, Kansas S ta te Un ivers i ty , Manhattan. B. S. Gill........ 26
.................... S u m r y Workshop Session 111, A. L. Rayburn.. 42
Appendix I. Proposed gu ide l ines f o r nomenclature o f biochemical l o c i i n wheat and r e l a t e d species. G. E. Har t and M. D. Gale .................................................. 44
Appendix 11. Recomnended r u l e s f o r gene symbol izat ion i n wheat.. 47
Appendix 111. Sumnary o f t he taxonomic and n menclature i conspectus o f t he T r i t i c e a e produced by s k e l l Love.. ....... 49
Workshop Attendance .............................................. 52
ERNEST ROBERT SEARS 50-YEAR CELEBRATION SYMWSIW
A p r i l 17-19, 1986
U n i v e r s i t y o f Missour i . Columbia
E rn ie Sears began h i s se rv i ce w i t h the U.S. Department o f Ag r i cu l - t u r e a t t h e U n i v e r s i t y o f Missour i on May 1, 1936. While he has now f o r m a l l y r e t i r e d from the USDA, he remains a c t i v e l y developing new stocks, new knowledge about the genet ic o rgan iza t ion o f wheat, and, as always, counsel ing and adv is ing wheat g e n e t i c i s t s i n many countr ies. I t i s through E r n i e ' s q u i e t and u n t i r i n g e f f o r t s t h a t complete se ts o f var ious aneuploids o f Chinese Spr ing wheat were developed and unse l f i sh - l y shared w i t h s c i e n t i s t s throughout the world.
It was the re fo re h i g h l y appropr ia te f o r E r n i e ' s col leagues t o have a s c i e n t i f i c meeting focusing on wheat genet ics and how h i s con t r i bu - t i o n s have been valuable t o p l a n t breeders and s c i e n t i s t s i n many d i s c i p l i n e s .
E rn ie Sears has received numerous recogn i t ions f o r h i s c o n t r i bu- t i ons , bu t t h e most p res t i g ious one was announced e a r l i e r t h i s year: The Wolf P r i z e he shared w i t h S i r Ralph R i ley . This ce leb ra t i on was a f i t t i n g venue f o r h i s col leagues t o extend congra tu la t ions t o him f o r t h i s h i g h honor.
We were pleased t o combine t h i s ce leb ra t i on symposium w i t h a wheat genet ic s tocks workshop i n which the cu r ren t s ta tus o f genet ic stocks and methodology were discussed. The emphasis was on what can be done i n t h e f u t u r e t o advance the knowledge o f genet ics o f wheat f o r t h e b e t t e r - ment o f science and the we l l -be ing o f the mu l t i t udes who depend on wheat every day f o r sustenance.
Gordon Kimber
North American Wheat Genetic Mapping and Cytogenetic Stocks Workshop
A p r i l 17-19, 1986
U n i v e r s i t y o f Missour i , Columbia
I n t h e l a s t few years, major breakthroughs i n bas ic cytogenet ics and molecular b i o l o g y have opened new approaches i n chromosome i d e n t i - f i c a t i o n and cy togenet ic mapping i n wheat. These developments w i l l a l l o w t h e cons t ruc t i on o f comprehensive molecu lar -cy to log ica l maps of wheat chromosomes. Na tu ra l l y , t h i s e n t a i l s nomenclature problems (chromosomes, bands, DNA probes, e tc . ), mapping s t ra teg ies , cy togenet ic stocks, research cooperat ion, funding avenues, and a s u i t a b l e forum f o r d iscuss ion and p u b l i c a t i o n o f in fo rmal repo r t s on an annual basis. This f i r s t workshop o f North American wheat g e n e t i c i s t s was organized t o address these and any o t h e r issues t h a t may ar ise .
We were pleased t o have the oppor tun i t y t o h o l d an in fonna l work- shop i n con junc t ion w i t h the symposium honoring Professor Sears on the 50th Anniversary o f h i s wheat genet ics research w i t h the USDA and U n i v e r s i t y o f Missour i .
The Workshop was suggested by Bikram G i l l (Kansas S ta te U n i v e r s i t y ) , who contacted Jan Dvorak (Un ive rs i t y o f C a l i f o r n i a , Davis), Perry Gustafson (USDA - U n i v e r s i t y o f Missour i ) , Gary Har t (Texas A&M U n i v e r s i t y ) , and Ca lv in Qua lse t (Un ive rs i t y o f C a l i f o r n i a , Davis) about o rgan iz ing a workshop. Gordon Kimber ( U n i v e r s i t y o f Missour i , Columbia) g rac ious l y o f f e r e d t o share the venue o f t he Sears Symposium t o h o l d t h i s workshop and completed the l o c a l arrangements.
The designated Workshop speakers in t roduced a t o p i c and presented a rev iew o f t h e c u r r e n t s ta tus o f methodology, genet ic stocks, and oppor- t u n i t i e s f o r f u t u r e work. Workshop p a r t i c i p a n t s discussed and presented t h e i r own views. The recorder suimnarized the proceedings and, w i t h t h e session c h a i r , prepared these in fo rmal proceedings f o r d i s t r i b u t i o n .
I t i s in tended t h a t the workshop w i l l be the f i r s t o f a s e r i e s o f i n fonna l meetings on wheat genet ics t h a t w i l l g r e a t l y f a c i l i t a t e t h e advancement o f knowledge and d i s t r i b u t i o n o f genet ic m a t e r i a l s i n North America. There was no i n t e n t i o n o f exc lud ing any i n d i v i d u a l o r group f rom p a r t i c i p a t i o n i n t h e Workshop. Copies o f t he proceedings may be obta ined from Cal Qualset, Chair o f the organ iz ing c o r n i t t e e , o r The Na t iona l Assoc ia t ion o f Wheat Growers Foundation.
The Sears 50-year Ce lebra t ion was organized by Gordon Kimber. Department o f Agronomy, U n i v e r s i t y o f Missouri . He received f i n a n c i a l and l o g i s t i c a l support f rom t h e Col lege o f A g r i c u l t u r e and A g r i c u l t u r a l Experiment S ta t ion .
The Workshop was organized by an ad hoc comnit tee i n c l u d i n g B i kram G i l l , Kansas Sta te Un ive rs i t y ; Jan Dvorak and Cal Qualset, U n i v e r s i t y o f C a l i f o r n i a , Davis; and Gary Hart, Texas AbM Un ive rs i t y . The cooperat ion o f a l l o f t he p a r t i c i p a n t s i s g r a t e f u l l y acknowledged, e s p e c i a l l y t he Session Recorders who developed sumnaries o f t h e key c o n t r i b u t i o n s a t each session. De ta i l ed abs t rac ts were n o t requested f rom t h e Sears Ce lebra t ion speakers. These proceedings i nc lude genet ic s tock l i s t s which represent m a t e r i a l s t h a t a r e a v a i l a b l e t o the e x t e n t poss ib le as determined by t h e holders o f t he c o l l e c t i o n s . Also present- ed a r e recomnendations from t h e Workshop which are sub jec t t o a d d i t i o n a l d iscuss ion and review; there fore , c i t a t i o n i n p u b l i c a t i o n s should be preceded by correspondence w i t h the authors.
These proceedings were prepared by the U n i v e r s i t y o f C a l i f o r n i a Genetic Resources Conservat ion Program. Reproduction and d i s t r i b u t i o n o f t h e proceedings were generously done by D r . V j r g i l Smail, o f t he Nat iona l Assoc ia t ion o f Wheat Growers Foundation, Washington, D.C.
PROGRPA
ERNEST ROBERT SEARS 50-YEAR CELEBRATION SYHPOSIW
AND
NORTH AMERICAN WHEAT GENETIC
MAPPING AND CYTOGENETIC STOCKS WORKSHOP
Univers i ty o f Missouri-Columbia, 17th - 19th o f Ap r i l 1986
Thursday 17th Apr i l . Room S204 Memorial Union
8:00 Welcome. Dean Wil l iam H. Pfander
WORKSHOP SESSION I Cal Qualset, Chair
Adam Lukaszewski , Recorder
Report 1 Current status o f gene mapping, w i th emphasis on enzymatic and non-enzymatic proteins : Gary Hart, Texas A&M University.
Report 2 Current status o f chromosome cytological map nomenclature: B i kram G i 11, Kansas State University.
1O:OO Break
Report 3 Mapping techniques - RFLPs: S. Muthukrishnan. Kansas State Universi ty.
Report 4 Mapping techniques - repeated DNAs: A. L. Rayburn, Oklahoma State Universi ty.
12:OO Lunch
FIFTY-YEAR CELEBRATION Chair: Gordon Kimber
1:30 Superoxide dismutase-1, a marker f o r T r i t iceae homoeologous- Group-2 chromosomes. Gary Hart. Texas A&M l l i i i ve rs i t y
1 : 50 Genetic analysis w i th aneuploids and der ivat ives. John Snape. Plant Breeding Ins t i t u te . Cambridge, England.
2: 10 Var iat ion i n nitrogen-use ef f ic iency, prote in and l ys ine contents i n Tr i t icum and Aegilops. Giles Waines. Univers i ty o f Cal i forn ia , Riverside.
Report 5
Report 6
10:oo
10:30
Report 7
V i t a l i t y Genes i n Wheat. S. S. Maan. Nor th Dakota Sta te Un ive rs i t y .
Break
Chair: Perry Gustafson
Chromosolnal ana lys is o f regenerated p l a n t s from anther c u l - tu re . H. C. S h a m , P. N. Mascia, S. G. Metz, T. A. Armstrong, and P. S. Baenziger. Monsanto. S t . Louis, MO.
The chromosomal l o c a t i o n o f a f a c t o r a f f e c t i n g megasporogene- s i s i n wheat. L. R. Joppa and N. D. Wil l iams. USDA-ARS, Nor th Dakota Sta te Un ivers i ty .
Establishment and a p p l i c a t i o n o f b lue-gra ined monoso~nics t o wheat chrmosome engineering. L i Zheng-Sheng. People's Republic o f China.
END OF SESSION
COCKTAILS AND DINNER ALUMNI CENTER
Chancel l o r Barbara S. Uehl i n g E. R. Sears w i l l speak about h i s 50 years a t UMC
Friday, 18 th A p r i l . Room 5207 Memorial Union
WORKSHOP SESSION I 1 Rosal ind Morr is , Chair
Perry Gustafson, Recorder
Mapping techniques - Physical mapping o f chromosomes: Jan DvoiHk. U n i v e r s i t y o f C a l i f o r n i a , Davis.
Genomic symbol izat ion i n the T r i t i c i n a e : Gordon Kimber, U n i v e r s i t y o f Missouri .
Break
Maintenance, construct ion, and d i s t r i b u t i o n o f genet ic stocks
Hexaploid wheat: E. R. Sears, U n i v e r s i t y o f Missour i T e t r a p l o i d wheat: L. R. Joppa, USDA. N. Dakota S ta te Univ. Al loplasmic and R f Stocks: S. S. Maan. N. Dakota S ta te Univ. A l i e n chromosomes: J. DvoFlk, U n i v e r s i t y o f Ca l i f o rn ia . Davis Species c o l l e c t i o n s : J. G. Waines, Univ. o f Cal i f . . R ivers ide
Lunch
FIFTY-YEAR CELEBRATION Chair: E. R. Sears
1:30 Notes on premeiocytes o f euploid Tr i t icum aestivum cv. Chinese Spring and mutant - Phl b . Henry Mcquade. Uni ve rs i t y of Missouri .
1:50 Frequency and o r ien ta t ion o f mul t iva lents i n the tetrasomics o f T r i t i cum a e s t i v m cv. Chinese Spring. H. Coucol i . Univer- s i t m s m , Greece.
2: 10 Weight o f Dr. Sears' research i n wheat genetics and breeding i n Rennes. G. Dossinault, J. Jahier, and F. Dosba. INRA, Domaine de l a Motte, Le Rheu, France.
2:30 Organization o f the 5s RNA fami ly i n wheat: Coevolution w i t h 18s and 26s RNA family. J. DvoFak, Univers i ty o f Cal i forn ia , Davis .
2:50 Break
Chair: John Snape
3:30 The a1 1 iance between wheat cytogeneti cs and wheat breeding . R. Morris. Univers i ty o f Nebraska.
3:50 Some information on the evo lu t ion o f t e t rap lo i d wheats. B. S. G i 11. Kansas State Universi ty.
4: 10 Transferr ing o f resistance t o E r i s i phe aminis t r i t i c i from a1 i en species t o wheat. J. Moseman. - ARS, Beltsv-irlle,
- 4:30 END OF SESSION
Saturday, 19th Apri 1 , Room 207 Cur t i s Hal 1
WORKSHOP SESSION I I I Cal Qualset, Chair
Lane Rayburn, Recorder
Report 8 Proposals f o r standardized nomenclature o f genomes, chromosome arms, and bands: Bi kram G i 11. Kansas State Universi ty.
Report 9 Proposals f o r standardized nomenclature o f genes f o r biochemical t r a i t s , RFLP's, and repeated DNA's : Gary Hart. Texas A&M Univers i ty .
Report 10 Proposals f o r genetic stocks development ( Including mu1 ti p l y marked stocks), maintenance, and d is t r ibu t ion : Perry Gustafson. USDA-ARS, Univers i ty o f Missouri.
Report 11 Proposals f o r organization o f North American wheat genet ics cooperation and fu tu re meetings: Cal Qualset. Univers i ty o f Cal i forn ia , Davis.
Prolessor Ernest R. Soar( #SFILIATIOW - U n l v e r e l t y o f M I e e o u r I . Dapar tmant o f Agronomv
Co lumb ia . MD 85211
BIRTH DATE - B o r n O c t o b e r I S . 1810 I n R l c k r e r l l . OragOn
EWCAT ION - P h . D . . H a r v a r d U n l v e r e l t y . 1838
P O S l T l a g - 1031 t o p r e e a n t . P r o f e a r o r o f Q a n * t I c a U n l v e r e l t y o f M l e e o u r l - Co lumbl8
Hl)lOlg/UIRW - W o l f F o u n d a t i o n P r l z a I n A g r l c u l t u r a f o r o u t a l a n d l n g work I n I h a a r e a o f wheat c y t o g e n a l l c a . 1888
- P r o f a r e o r E . R. Sea r8 Day I n M l a e o u r l b y D a c l a r a l l o n o f l h a G o v a r n o r , on t h e o c c l a l o n o f h l a 40 y e a r n o f a a r v l c a a t !ha M l e r o u r l Agricultural E x p r r l m e n t S t a t l o n . A p r l l 10 . 1978
- H o n o r a r y Mambar. Q e n e t i c a S o c l a t y o f Japan. 1875 - D l e t l n g u l a h e d S e r v i c e C l l a l l o n . U n l v . o f OrDgon. 1973 - H o n o r a r y D o c t o r . U n l v e r e l l y o f Q o t l n p r n . 1070 - F ~ I I O W , I n d i a n s o c ~ e t y o f Q a n a t l c r and P l a n t B r a r d l n g . ' 1088 - Mambar. N a t l o n l l Aeadmy o f S c l a n c a a . U . S . A . . 1884 - Qamna Slgma D e l l a N l t l o n l l Award. 1858 - USDA S u p e r l o r S e r v l c r Award. 1958 - H o b l l t z e l l Award. 1858 - F e l l o w . A m r r l c a n S o e l e t y o f A g r o n m y . 1958
SUlWARY WORKSHOP SESSION I
Adam Lukaszews k i , Recorder
Current Status o f Gene Mapping
Dr. Gary Har t discussed the present s ta tus of gene mapping i n wheat, w i t h emphasis on enzymatic and nonenzymatic prote ins. A t t he present t ime 272 l o c i have been loca ted on chromosome anns. These range from 1 t o 9 l o c i per chromosome o r 0 t o 6 per am. Unfortunately, there are 7 wheat chromosome anns on which no gene l o c i have been mapped. Th is i s an extremely small gene map f o r such an important species.
No. l o c i X
Prote ins 119 44 Disease 83 30 Other 70 - 26 - Tota l 272 100
Out o f t h e 119 p r o t e i n l o c i i d e n t i f i e d , 99 encode enzymatic p r o t e i n s and 20 encode nonenzymatic p ro te ins . O f these l o c i . 9 encoding f o r enzy- ma t i c and 12 encoding f o r nonenzymatic p ro te ins have been mapped. The mapping methods used by D r . H a r t inc lude the c l a s s i c a l type o f tech- nique, a l i e n t r a n s f e r techniques, d e l e t i o n techniques, t he use o f t e l o c e n t r i c chromosomes and intrachromosomal recombinants. Appendix I reproduces a paper by Har t and Gale (1986) dea l i ng w i t h naming o f biochemical l o c i and Appendix 11 reproduces t h e general gu ide l i nes f o r gene nomenclature i n wheat.
Chromosome Map Nomenclature
Dr. Bikram G i l l discussed the present s ta tus o f chromosome banding techniques and chromosome banding nomenclature. Wi th t h e N-banding technique 16 wheat chromosomes can be i d e n t i f i e d . Only chromosomes lA, 30, 40, 50, and 6D cannot be i d e n t i f i e d . I t appears t h a t a po lypyr imi - d ine DNA sequence i n wheat i s t he sequence t h a t produces bands w i t h t h e N-banding procedure. A t t h e present t ime on l y r y e chromosomes 2R, 3R, and 6R a n d i f f e r e n t i a t e d w i t h N-banding. However, t h e C-banding proce- dure produces bands on a l l o f t h e wheat and r y e chromosomes because the process a f f e c t s most heterochromatin.
The r e s t o f t h i s sec t i on was spent d iscuss ing t h e proper k i n d o f banding nomenclature t h a t should be adopted by t h e wheat genet ics workers o f North America. It was t h e general consensus o f t h e group t h a t t h e nomenclature used f o r human chromosomes s e t up by t h e Par i s Conference i n 1971 was too d e t a i l e d and should n o t be used i n wheat a t t h e present t ime. A comnit tee was s e t up t o exp lore t h e poss ib le a l t e r n a t i v e s t o present a t t h e n e x t meeting.
Mapping Techniques - RFLPs
Dr. S. Muthukrishnan discussed the advantages o f i s o l a t i n g r e s t r i c - - t i o n fragment l eng th polymorphisms (RFLPs) i n wheat. Several advantages t o us ing RFLPs were given: 1) the mutat ion need n o t be l oca ted i n the t ransc r ibed a r t o f t he gene; 2) t he t a r g e t s i z e o f t h e gene i s increased; 3 ! no in fo rma t ion regarding the na ture o f t h e gene product i s requi red; and 4) RFLPs are i n h e r i t e d as codominant Mendelian t r a i t s . To use RFLPs i n wheat we need t o consider t he f o l l o w i n g po in ts : 1) how polymorphic must each marker be i n order t o be usefu l ; 2) how many f a m i l i e s are needed i n o rder t o e s t a b l i s h l i nkage groups; 3) and how much polymorphism can we expect t o f i n d i n t he species under study. One poss ib l y s t ra tegy f o r c rea t i ng RFLPs would be t o i s o l a t e ba r ley probes and use them t o e s t a b l i s h wheat-speci f ic markers. The main problem a t t h e present t ime i n i s o l a t i n g wheat RFLPs i s t h a t there are n o t many polymorphisms i n wheat. More extensive polymorphi sms have been found by us ing ancest ra l species o f wheat. A t t h e present t ime crosses between two v a r i e t i e s o f wheat e x h i b i t i n g RFLPs a t several l o c i w i l l be used i n mapping RFLP l o c i i n wheat.
Mapping Techniques - repeated DNAs
Dr. Lane Rayburn discussed the p o t e n t i a l f o r i s o l a t i n g and s tudy ing repeated sequence DNA from wheat. A t the present t ime he has i s o l a t e d r e p e t i t i v e sequences t h a t a re n o t present i n C-banded regions, b u t a r e found i n t h e euchromatic region. A g rea t deal more work needs t o be done i n t h i s area.
Chromosomal and Chrmsome-arm Locations. Linkage Estimates. and Map Locations o f T r i t i c u m aestivum Gene Loci
Gary E. Hart S o i l and Crop Sciences Department
Texas AM Un ive rs i t y College Stat ion, Texas 77843
Explanat ion of diagrams:
1. Chromosome-arms are designated p and q t o i nd i ca te homoeologies (Sears and Sears. 1979). except f o r the group-2 arms. which are designated S (= s h o r t ) and L (= long).
2. Mapped l o c i are l i s t e d on the r i g h t s ide o f the chromosolnes and the map p o s i t i o n s o f t he l o c i r e l a t i v e t o the centromere, expressed as percent recombination, on the l e f t s ide o f the chromosomes. L i k e l y r e l a t i v e p o s i t i o n s and order are ind ica ted f o r some l o c i f o r which map p o s i t i o n s a re not shown.
3. Unmapped l o c i f o r which chromosome-arm locat ions are known are l i s t e d t o the l e f t o f t he chromosome-arms.
4. Unmapped l o c i f o r which chromosomal locat ions, bu t not arm locat ions, a re known are l i s t e d below the chromosomes.
5. "** i n d i c a t e s genes t rans fe r red from a l i e n species, "tY i nd ica tes l o c i f o r which d i f f e r i n g and unresolved arm locat ions have been reported and, i n t h e "Linkage Estimates" sections, "I' ind ica tes genet ic independence.
The diagrams are modi f ied f rom those o f McIntosh and Cusick (1984). The f u l l names o f t h e gene l o c i l i s t e d , synonyms, and references may be found i n McIntosh (1983. 1985. 1986) and Har t and Gale (1986).
REFERENCES
Hart, G. E., and M. A. Gale. 1986. Gene symbols f o r wheat prote ins. 1986 supplement. Wheat Newsletter - 32: ( I n press).
McIntosh, R. A. 1983. Catalogue o f gene symbols fo r wheat (1983 ed i t i on ) . Proc. 6 t h I n t . Wheat Genet. Symp.. pp. 1197-1257.
McIntosh, R. A. 1985. Catalogue o f gene symbols f o r wheat, 1985 supplement. Wheat Newsletter - 31:202-212.
McIntosh, R. A. 1986. Catalogue o f gene symbols f o r wheat. 1986 supplement. Wheat Newsletter - 32: ( I n press).
McIntosh, R. A.. and J. E. Cusick. 1984. Linkage map o f wheat ( T r i t i c u m aestivum: 2n = 421, pp. 482-484. I n Genetic Maps 1984. O I B m m. Cold Spring Harbor Laboratory, N.Y.
Sears, E. R., and L. M. S. Sears. 1979. The t e l o c e n t r i c ch rmsomes of conmn wheat. Proc. 5 t h I n t . Wheat Genet. Symg., Vol. 1, pp. 389-407.
GROUP 1
rf 3 T d - 0 6 ' Per-Dl 25 - Gpi-Dl Nor-B1 Glu-B2 m - Hk-Dl Per-Bl
Linkage G I 1-A1 - Glu-Al: 66 Bt4 - R 1. 14,23 G l i - 0 1 - G i - D l * 36 Estimates: H : BFa - &I 15 m - h ~ 48 - 3 : 1.5 Bf4-m: 30 BE-m: I
m - 0 1 - rf3,Glu-02: 22 TfX - T d T 2 8
- - TfX - Nor-01: 37 CKBT - 31,39.1.66 G1U-B1 - m 6 ' : 26 Glu-81 - Nor-01: 22,23 r f 3 - o r - 3 7 7 7 - m : 34
Linkage Lr17 - Crll: I Estimates: - Vina6: 38 - -
C - Dl: 37 - : I m - a 4 : 12 m - m : I E 2 a 7: 10 El3 - r r 2 : 0 - 6.21.32.42 m -+: 12 - d: 11.15.18.42 F J T - YX6: 36
04:I k 1 17 m: &: 44 m-m-I 5- L-a: 1 m-Ra:O - 18.19.33 - 1 : e l 1JT- T.I5,21,22,28 F l 6 - 3 4 : - 25.26
< I vla
I-
Linkage Sr35 - R2: 1 - - Estimates:
Linkage Adh-A1 - Gail: 23 Sr7a - Hd: I Estimates: L r F I - -
m - F 2 7 : < 1 : 34 : 20 - W y - ~ l : I -
30 Adh-Dl
2gg P9n-DI
. Rht2/Gai2
q 50 Vrn l t - q 5 O I L r l 8 -
Aadh-A1 Aca-AZ r
Linkage Estimates:
0-Am -A2 - 01: 2 r n 1 : 37 &i-
H3-m: 9 W - w: 16 m-w: 2 w - m: 36,1 r- B ~ 3 0 . 3 3 . 3 7 , 4 1 q - zntrocnere: I q - Hn: 36 -
10 Aadh-B1 11
0-Am -A2 Aco-B2 B+-
L r l - Pm2: I 5 3 3 -TI: I mu-ViiiZ: I - -
- Nel . K?T - Hii
36.43 - - Crr
Lf nkage Estimates:
02 - Lr9: 4 82 - m: 43, 47 mi-BTC-band: 10 m r 3 : 12 KT - -1: 9.11 1 9 : very small 5l-r - m: 0.22 - -
44
IZ€X
4
Aadh-A2 Aco-Ar Z E G A l E d - Exiv sz!F -
50 -. R f 4 - - Rf5
A
P 50
. Sr8 -
20
12
0 <1
Aadh-02 19
gg EcEz
45
Srl3 q - 50'-
0 D .
. Col - P 50
- 611-82 8% F0T-m
. C-band =mw -
Lr9*.Nor-02 02- -
12
., - Sr5
a-hy-01 Aadh-02 . a-Amy-01 Azzo'r c02 EX-04 Gm!Z
- Srll/Lr3 - Ki 9 50 .. ~r29*+ -
GROUP 7
Linkage cn-A1 - 9 2 2 : 2 Estimates: cn-Ar - PiiE 40
Pml S r m 4 1 - -
A B
P
Per3 -
F - 27 ..
34.50 - 9
50 -
Ed - Yr2
P
Est-B3 Lrrn m Vrns m- -
16,23
6
g Sr22 - m - cn-A1
q 37 .. Pml Lr20/Sr15 -'- -
.. PC -
0
a-Amy-B2
- PmS
apomsal for P ~ i k r m s G i l l
mpxtma* d nar* mthdoqy Irarmas State Wvcrsity muhattan, Ranras 66506
Kimkr and Sears ( R o c I11 Int. Wheat ana t . m.) popoaed a namenclature system for the deacri*ion of anauploih in the w i t i d m e . McInto& (Proc. N Int. Whmt Genet. QUIP.) -&ad a ~ t a l o g u c of genes and gew symbols. Those standardizations qf nmenclature did much t o pranote w h e a t genetics resear&. With the wiaspread use of C-banding and N-banding techniques i n somatic chrcmoaome idantif ication, it is imperative that a standardized c h r m tand namnclature be developed for the descriptian of wheat chramallnes and chtcl~o~~mal aberrations. mst of the information given belor is taken f rm G i l l (ASA Wheat Ebrpgn*, in peee).
Since Chinese Spring is the widcly wed cultivar in q t o p m t i c anslyeis, the standard karyotype should be based on this cultivar. The ideogram of banded chranosomes i n Chinese Spring (Appendix I) was oonstrufted frcm a o~mp~sitc of C*&d and EFfan&d chramsanes and is taken from G i l l (ASA Wheat Honogra~h) . Information on chrmoeome size, cerhrcmere position, and arm ratio of indivi&l chramanms was obtained frcm the measureaent of acetocamin stained and banded chranosoloes. In addition, psi tion, Biz9 and irtensity of individvl tan& are a lm hpoctart criteria in chrammrte idantifiation and are chcm baaed on the i r relative size and position.
The designation of chrunosome arms and bands fallare the reamnerdatlane of the 1971 Pads Cbn€erence on standardization i n h w n cytogenetics. Amever, nomenclature i n hman chranogme b d n g was based on Q- and G - ~ I M ~ I , whergs in wheat it is based an C- and Nbanda mder the proposed rulea of nanenclature, each chranosome short arm ie d e s i p t e d as p and the larg a m aa q. Each p and q arm is subdivided in to regions based on chrcm~sane lanclnarke. A chrcmosome landnark is defined a s "a consistent and distinct mor#~dogic feature that is an important diagmstic aid i n identifying a chranoaane." A region is defined a s "any area d a ctvanoecrme lying between adjaoent lamkwks.* A bard is d e f h d as "a prt d a chranogrme clearly distinguishable frcm adjacent parts by v i r tue of its lighter or darker stairring ability." Each arm cimsists of a mntinwue aeries of dark bands and l i g h t bands and by definition there are no " in t e r bands. " Regions and bands a r e nunbered consecutively frcm the centranere W a r & alcng eaQ chramsant a m
The two regions aqacen t t o the -ranere are labeled "1" i n each arm, the next, more distal regions, '2", and so on. A band used a s a l a n b a r k i s considered a s belonging ent i re ly t o the region distal t o the landnark and is accorded tba tand nMbg cf "1" in that region. Ea& chrameame has a dis t inc t centramere, and the heterochramatin bisected ty the centranere is mnaidered as b o bands, each being labeled a s band 1, i n region 1, of t he appropr ia te chramoaanw ana. Scmctimee oentraneric heterochcamatin may extend i r t o en arm Md fuse with a polimal band, a s for exanple in chramgme 2A woximal Land in p arm.
In designating a p r t i c u l a r band, f ive item are required: the chraaoga~e nrmkr, genapa BPignation, the ann symbol, the regicn nrmber, and th bsnd n-r w i th in t h a t region. These items a r e given i n order without spacing or pmctmtioh For eltrmple, lBqP indiaatrs chramaane no. 1, genune B, long am, regfcn 2 and band 1.
Naenclature rules on t he subdivision of &sting 1-ks or ban& state that i n th. a n t that a band serving a s a lan-rk requires subdivision, all sub-band. &rived f r e it should retain the original regiar and tand nrmber d that landnark. Tbis rule is to be follcvad wen if subdivision should caw one or more sub-band t o l i e i n an adjaaent region. Whenwer an existing band is to be subdivided, a decimal point should be placed a f t e r the o r ig ina l band designation follaued by the nrmber aseiqied t o each &dmd. !lhe subbad are nrmbeted sequentially fran the antranere outward. For example, if the original band l q l l were subdivided in to three equal or unequal -, the w r W a d would be labeled lgF1.1, lq l l .2 , and lq11.3, w i t h eu&band ll.l being woximal and 11.3 d i s t a l t o the centranere. Finally, i f a sub-band is t o be subdivided, addtional digits but no further pmctuntian should be used; e.g., sub-tnnd lp33.1 might be further subdivided into lp33.11, 1~33.12, etc.
It am be aeen that the p o p & nonenclature pwides the neaaeeruy accuracy and f l ex ib i l i t y for &scribing banded chranoaanes and any future additians to the bands. In cannon wheat, dark bands and l igh t bands respectively represent heterochrauatic and euchranatic regions, and an attern* has been made to subdivide most wheat c h r a a ~ g m ~ s into Malogically meaningful regions, Thus, regions 1 and 2 i n p and q arms of 7B describe ~ox imal heterochramatic and dietal euehrcmatfc tands, respectively.
Sane nomenclatural modifications have been adapted t o conform to spadal si tuat ions i n wheat. Dark bands t ha t a r e not always reproducible i n a l l chranoaannes are designated by stippled bands and have not k e n assigned tand nmkrs. A good erurmple is tand SBq12. In sane unusually clear preparations, t h i s band can be subdivi&d and, if neassaly, described a s bm% 5-2.1, 5-2.2 and ao on.
In contravention t o the intended meaning i n hunan cytogenetice, p and q h e k e n used to &signate chram~sane a m hanoeologies within hanoeologotle groups. This poses problems f o r arm designation i n homoeologous groups 2 and 4 chrwacmes. 'Ihu, arm 4m whi* designntas the long ann of chromosame 4B i n our proposal, has been designated 4Bp by Bart, t o indicate hanceoloqy with 4 & ~ and 4Dp, short alms i n both cases. One so lu t ion t o t h i s problem may be the designation of pl to indfa te that i n this specific instance the p arm is actually the long arm. This wffl a t cnae ~ c w i d e information on hamoealogy a s w e l l a s arm length.
Folymor@irns i n w h e a t cultivare should be described relat ive to Chinese Spring. Ebr erne, if there are extra bands i n Wichita ( W I ) chranosane ?A, which al lau suMivisicn of a mrrespnding band i n Chinese S~cing, then it hould be designaW as 2A#22.1(WI) ,~AI&?~.~(WI) and 2Aq22.3W). Thfs indicates tha t i n c u l t i v a r Wichita, band q22 is d i v i s i b l e i n t o 3 sub-bands. If there is polymorphim f a the intensity of a tam3 (relative t o Chinese Spring), it should be described a8 q22+(WI) o r q22-(WI) + W c a t i n g a more i b e w or lcngtr lnnd and - denoting a f a in t or a snaller band i n Wichita. L i s t ing of s tandard abbreviations of wheat cul t ivars can be obtained fram OregDn Experiment Station (Spda l Rprt 719, m, OcegDn State Univeaity, Oonrallis).
The banding a n a l y s i s may provide detailed information on chranosomal aberratiars. Zhatdore, nanenclature qmbds in AFpendix If are recamended f o r t h e designat ion and description of chranosomal aberrations. The follauing examples illustrate hau sane of t h e more commonly occurr ing chrormosanal aberratiane rnsl be described.
a. Suuctural ctwgea imdv ing sin#e breaks
1. 'Ilcrmirrl d s l e t i a ~ 4 2 , a (18) (@l) A terminal delet im i n duanoaune 1 B
with a breakpint i n region 3, tam3 1 of long am
42,del(lB) (peer-q21:) Zk single calon indca tes a break a t band q21 and deletion of the lcng a m segnent distal to it.
2. Isochranpxnes 42 , i ( l 4 ) Iso&ranoaane for the lcng arm of 18. 42, i (1B) (qter-ten-ter) Break points are at or close t o the
centranere and carnot lx spedf ied. Ime B a t p t i o n i n d a t e s that both lcng anus of 1 B a re Feaent and a re seprated ty the oentranere.
b. Structural changes involving two breaks
1. Pericentdc inversions 42, h ( 4 B ) (p12q22) A p r i w n t r i c i w e r d m i n &rancecme
48 with breaks in band 12 i n the short arm and tend 22 in the long am.
42,inv(SB) (peer->p2l:: Breakage and union have occurred a t 1331-pZ1: : q 3 l A q t e r [email protected] i n t h e s h o r t a m a n d
SEq31 in the l a g arm Ime segnent lying between these tands is inverted.
2. Ring chranogmes 42 ,r(6B) (1?22@S) A centric ring 68 chrancaane, the
break points being i n region 2, band 2 of the short arm and regicm 2, band 5 of the lcng a m
42, r (68) (p22Aq25) With deletion of both termid segnents, the broken ends have joired t o form a ring. IWe the anission of the calm or &uhle mlcn
3. Diaentdc chramacmee 42,dc(lB) ( 4 2 ) A dicentric 1 B chranogme with a treak
at trand 2 i n region 1 of the l a g a m 42 ,dc(lB) (pter-42: : Breakage and M i o n have occurred a t 42-pter) band lBql2 on sister chranatids t o
form a dicentric 1B chramagme
4. Ranalocations a. r e d p o c a l translocatiana A brlanced recigrocal tranelocatim 42,rcp(2B;SB) (q211q31) between chrcm~scmre No. 28 and
No. 58, with break pi*s at tand 1 42,rcp(28:5B) (Ipter i n region 2 on the l a g a m of 2&2l::SBq31-5qter; 5-r Qranoerrne No. 28 and tand 1 5Wl: :2BqZl-+ZR~ter) regiar 3 on the lag am of
chrawgrme m. 5B.
Appendix I
~ w d i x 11. Ncmenclature symbols (after M a g o and Paris ~bnfermas)
P short arm, hmoedog3~le to Ehort arme ¶ larg am, lrmoedog~m t o lang a m fl lanq arm. hclPoelocpus to short arme
ehort arm, kmceologom to lcnq arms OM antranere ace acentric dic diaentric s satel l i te h aeaxldaly contdction r ring chraaogme i is~chraa~acme t telooentric chranogme end endoreduplication w drplicaticn i n w inwedon ins insertion i n w ins imerted insertion &l deletian T translocation r cp r e d ~ o c a l translocation rob robertsonian translocaticn (cmntric fusion) tan tandem translocation ter tenninal or end ('pter" for end of short arm) mar marker chronosane rec r e d n a r k chranosane der derivative cfirmosane mat maternal origin pat psterral origin
break (no remion, as in a terminal deletion) . . . . break and join f r m - to
7 qesticnahle identif ication of chranomne or chranogrme structure + the + and - signs should be placed before the ap~ropriate - sylFbol where thq man addition or missing whole chrauceanes.
lhq should be placed after a symbol where an increase or decrease in length d i n c t l v i b l bands or regions is meant.
WE: A l l -1s for rearrangements are to be placed &fore the &signattan d the chraa~ecmc(s) inwolved in the rearrangemnt, and the rearranged chranogme(s) should always be placed i n puenthases.
SUWARY WORKSHOP SESSION I 1
J. Perry Gustafson, Recorder
Mapping Techniques
Dr. Jan DvoFik discussed several procedures used f o r mapping genes on chromosomes. One method involves us ing i n t e r v a r i e t a l chromosome s u b s t i t u t i o n s and t e l o c e n t r i c chromosomes f o r mapping genes t o spec i f i c chromosomes. Th is i s much the same technique used by J. W. Snape, P lan t Breeding I n s t i t u t e , Cambridge, f o r mapping q u a n t i t a t i v e and q u a l i t a t i v e t r a i t s . One problem w i t h t h i s technique i s t h a t a t e l o c e n t r i c chromo- some does n o t p a i r w i t h i t s whole chromosome homologue as w e l l as do two whole homologous chromosomes; there fore , the r e s u l t s o f t e l o c e n t r i c ana lys i s can be b iased and the map d is tances d i s t o r t e d . However, the technique does have many use fu l purposes. A phys ica l map of wheat can be produced by several approaches w i t h the f i r s t one being t h e ana lys i s o f C-band polymorphisms. The v a r i a t i o n i n the r e s u l t s from one C-banding procedure t o another makes t h i s technique, a t t he present t ime, somewhat u n r e l i a b l e . I n add i t i on , no t very many C-band polymor- p h i s m ~ c u r r e n t l y e x i s t i n wheat. A second approach i s t o c rea te a phys ica l map by s tudy ing chromosomal rearrangements. Unfor tunate ly , t he re a re n o t t h a t many chromosomal rearrangements i n wheat t h a t d i f f e r - e n t i a t e v a r i e t i e s ; no t a l l chromosomes are involved. A t h i r d approach i s t o c rea te a phys ica l map by analys ing the i n s i t u h y b r i d i z a t i o n pa t te rns o f var ious DNA sequences. Th is t e c h x q f i a s the most p o t e n t i a l ; however, a t t he present t ime there a re some drawbacks: 1 ) background noise makes i t very d i f f i c u l t t o analyze accu ra te l y smal l copy o r unique DNA sequences; 2 ) there are few wheat DNA sequences a v a i l a b l e t o map; 3 ) most o f t he sequences des i rab le t o map would be very smal l and thus hard t o l abe l .
Genomic Symbol izat ion i n the T r i t i c i n a e
Dr. Gordon Kimber discussed the genome c l a s s i f i c a t i o n proposed by Love (see App. 111) and the symbols and names o f t he var ious genomes. He proposed t h a t t h e system o f nomenclature pu t f o r t h i n Japan a t t h e 6 t h I n t e r n a t i o n a l Wheat Genetics Symposium by Kimber and Sears be adopted (Table 1) . Dr. Jan DvoF6k proposed t h a t we adopt the L6ve system f o r naming perennia ls . Whi le LBve's proposals are a l a r g e departure from c u r r e n t usages, cu r ren t usage i s no t a v a l i d c r i t e r i o n under t h e Code o f Taxonon~ic Nomenclature. Lsve's proposals are d i s - cussed i n Appendix 111. F i n a l l y , D r . J. Waines suggested t h a t we change the genome o rde r i n po l yp lo ids so t h a t t he f i r s t genome named i s t he one t h a t i s supposed t o have donated cytoplasm. These two proposals were discussed and w i l l be decided on a t t h e nex t meet i r~y.
The storage, maintenance, cons t ruc t ion , and d i s t r i b u t i o n o f genet ic s tocks housed i n Missour i , Nor th Dakota, Kansas. Nebraska, and C a l i f o r n i a were discussed. The general consensus was t h a t Drs. Sears, Kimber, Gustafson, Lukaszewski, Joppa, Mann, G i l l , Mor r is , Waines, and DvoPdk were ma in ta in ing t h e i r stocks and d i s t r i b u t i n g them t o o the r research s c i e n t i s t s on request. However, there was a general agreement
t h a t i n s u f f i c i e n t funds e x i s t f o r the storage, maintenance, and d i s t r i - b u t i o n o f these valuable stocks. Many o f the genet ic stocks w i l l n o t be a v a i l a b l e f o r d i s t r i b u t i o n u n t i l adequate funding can be obtained. I n add i t ion , there was a general consensus t h a t more c o l l e c t i o n s were . needed i n order t o secure valuable genet ic stocks and samples o f w i l d species before they are l o s t .
Table 1. Proposed genorne symbols o f t he d i p l o i d and p o l y p l o i d species o f Tr i t icum.*
Species
T . monococcum. T. u r a r t u T s i d e s T: K k S - T. m i m u m T. s e a r s i i T f r i o i d e s T: + T. comosum T. G E F E t a t u m T. d ichasians T. umbel lulatum T. turgidurn -
T. ventr icosum T. crassum ( 4 x ) T. crassum (6x1 T s m T: h T. kotschyi - T. ovatum T. m s t a t u m (4x) T. t r i a r i s t a t u m (6x) 7. macrochaetum T. columnare T. t r i u n c i a i e T. - cy l i n d r i c u n
Genomes
AG AAG ABD
- - -
Synonyms
T. boeoticum, T. monococcum - -
%: m i s t a t a z. t r i a r i s t a t a . b i u n c i a l i s , - Ae. l o r e n t i i . columnaris . t r i u n c i a l i s X. - c y l i n d r i c a
Kimber, G. and E. R. Sears. 1983. Assignment of genome symbols i n t h e T r i t i c e a e . Proc. 6 t h I n t . Wheat Genet. Symp., Kyoto, Japan. Underl ined symbols represent s u b s t a n t i a l l y mod i f i ed genomes.
CYTOGENETIC MATERIALS MAINTAINED BY E. R. SEARS
A p r i l 1986
Aneuploids Chinese lpring - Wheat
NuLt%o&, 20": lA, 6A*, 7A, 18*, 78, 4D*, 7D.
M o m b o u , 20"+11 : A1 1.
TUomiCb, 20"+ln1: 3A, 4A, 5A, 38, 48, 58, 68, I D , 2D, 4D, 5D*.
T&uuo&, ZO"+llv: A l l
Monot&dornicb, 2OM+t': A l l except 2AL. 2AS, 3AS. 4A6, 5AS. 7AS. 28L. 285, 48S, 585, 6BL, 6BS, IDS, 2DL, SDS, 6DL. 7DL.
D L t e e O b o ~ , 2OV+P: A l l except 2AL, 4A0, SAS, 285, 485, 585, 4DL (temporary), 5DS, ?DL.
Monot&di6omica, 20"+tlt1: 48s. 585, IDS, 3DL. 5DS.
Double momt&bornicb, Z0"+2t1 : A1 1 . Doubee d i thbonr i cb , Z O " + Z t n : A1 1.
Doubee dit&b phZa phfa: lA, 2A, 4A, 5A, 7A, 28, 38, 48.
Momt&o-ibo~omics, ZO"+tl+i': 1505 78LI 3DL, 3DS, 5DL.
~ h 0 n o ~ d O b 0 ~ , 201'+t"+.t' : A l l except di-7DL and -7DS.
DLt&o-mono&obomiCs, 20"+.t"+i1 : Isos 58L. IDS.
Mono-ibobomics, ZO1'*i': IAL, ZAS, 4Aa. 5AL, 6AS, 2BL. 48L, 58L. 685, 78L. 5DL, 6DS. 7DS.
DL--ibobonricb, ZOn+i": SAL*, 28L*, SBL, 685.
Double m o ~ ~ o b o n r i c , 20"+2i1 : 5A.
Compe~at ing n u U L - t W , 19"+l ' v : A1 1 except 28(2A), 2D(2A), 48(4A), 4D(4A), SA(5B).
Comp. mono-tets, 19"+l ' v + l l : 28(2A1), 20(2A'). 4B(4A8), 4D(4At). 5A(5B1), 50(5B1).
Comp. U-.i%h, 19"+1"' : 58"' (5D). 6AV'(6D).
Doubee mOnOb in&. 58, 19"+2' : A l l .
- - - -
*Seed supply low.
Intervar ieta l Substitutions
'Hope': A11 (58C's ) . he' : A l l (5BC's).
Irk-': A l l (3-5 BC's).
'~d E g w ' : A l l (2-5 BC's).
A1 ien Additions and Substi tutions
' Imperial ' Rye
Di.mnik additiou, 21"+1": A l l .
D d 0 d 2 6-M. 20n+1*: 2R(28). 2R(20). 2RL-ZAS(2A).
Haynaldia v i l losa
Udonric caddiAhn.4, 21"+111' . A l l except 3Ha.
Agropyron elongatum
3 4 ITAP67): Dismic add.. mono add.. ditelo-L add., ditelo-S add.. disomic subst. 3Ag(3D).
7 4 ( 4 ~ 1 : Disomic add.. di telo-L add., disomic subst. 7Ag(7D), d i t e l o subst. 7AgS(70), double d i t e l o subst. 2tW(7D).
Agropyron intermedium
TAFl (Caudwn) : 3Agi disomlc add.
TAFZ ( h d w n ] : 7Agi disolnic add.
Aegilops umbel1ulat.a
Chhorn. 6U(=AI: Disomic add., dimom subst. 48 (20"+1"6U+1'48), dimno subst. 68, d i m m subst. 78.
Aegilops spel toides
Trit icum timopheevil
6 1 . oddit. Uoncic.
6 T h b h t . dido-: For 6A, 60, 60.
A1 i e n Transfers - A. &ngcrbu, 3 4 to 30: M S . 1-9, 11. 14-16, 18-21; i n t e r s t l t i a l s 3, 14.
A. &ng- 3 4 & 38: NOS. 10, 12, 13.
A. &ng& 7Ag .b 70: NOS. 1-11.
Ae. r u n b d & h t a : 147 ('Transferb--6BL). T40 (68s). T41 (4A), T44 (2B), T52.
Rye: HN-1 (5RL-60). HN-2 (5RL-5BS), HN-2 motto, HN-4A (5RL-4Aa), 6B/6BL-5RL.
Ae. bpeCroideb (9~321 : 2s-2AS (adh. g l .), 25-285 (Nos. 142. 155). 2s-2DS (Nos. 132, 139, 153).
Mutants
Vhebcem2: Neatby's ( v l a ) , - Washington's (v2a), Hermsen's ( v l b v2b) -
P-: phlb, ph2a, ph2b (Cambridge 1013).
A1 1 otet rap lo ids
T. mOMCOCcLun X Ae. WkVd&th T. bowticum x Ae. WLiarLib-tatcZ T. bowticum x Ae. wnbeeeLeata T. bowticum X Ae. bqUWWba Ae. hiconnid x T. monococcrun Ae. x Ae. b ~ U W b a Ae. caudata x Ae. u m b ~ L U W 5
Ae. caudata x Ae. W & u i t Ae. caudata x Ae. bquannoda Ae. b h o n e n b h x Ae. caudata Ae. d h a h 0 n ~ i . 6 x Ae. urnbeL&dda Ae. bhakonmh x Ae. W & u i a Ae. bpeetohfes x Ae. unicvLid&uit Ae. u m b & L & & z x Ae. curiahib&th
Allohexaploids
T. dicoccwn x Ae. bqurwroba: Black glume, 'Vernal ' . T. dicoccoideb [h igh prroteinl x Ae. bquannoba T. drYucn x Ae. bqumoba: 'Carleton'. 'Pentad', ' I u m i l l o ' , 'Golden Ba l l ' . T. dicoccun x Ae. umbeeeLeata T. dicoccum x Ae. caudata T. dicoccum x Ae. bpeetodes T. dicoccum x Ae. W.ta. .Zu T. dicoccwn x Ae. b h o n m h T. cmd&am x Ae. b h m 0 ~ U U . i b
T. tbnopheevii x Ae. bqUVtA.Oba T. .thIopheevG X AA~ . mb&&l,&&Z T. tim;pheevii x Ae. caudatrr T. i hopheev i i x Ae. bpebbideb T. tbnopheevii x Ae. hiconnib T. tbnopheevii x Ae. ~~
A1 looctop lo ids
~ + i u t w of thin co l l r t ion h... k.n antarad into the Pi-c IOCroductiacl colbc~ioo of th. USD* at h l t r d l l a . Wumrcr for germplan rhould be rddrmrsed
cb wpm of chm S u l l Orsine Collectioa. USM, kl trr i l lm, m.
u0Cl.r- Q r c e Turkey I r ~ r c a u . Iraq Iran labanon U r u l Ocher rified Total
militinma orientale sinrlujrm
utiu c.ud.c. C O M .
uniarir u t a 8qIUCCOOa bicoroir lirtiu 1oIIgi.oiu ahrononoir op.lroido8 ub.l lul . t . a u r r i i cy1iodriu C t U I . ju*.lulir v0atricO.. variabil ir kotrchyi triunciolia columurir bioncialir C tiAti8taCA O V A U tureounica loroat i i tauichii v.vilovii unclarrified
Tot a1
- Elycrigia d i s t i c h
6rewp7nm buonaparti8 w orientale . triticum
1 L p l d o c r i t i c c u sardoum
lbrd.em bulbosu - coeleete " daficicas
f u r c a t u var. spontaneu - sp. wlgare
Secale af ricaaum - s n a t o l i c u " cereale " f rag i le " montsnu " segeca1e
T r i t i u l e
Only s u l l amounts of these accessions are maintained a t UC IUverside. Requests for seed should bear i n mind that seed m y have to be increased over one 8easoa to fill an order.
A v a i l a b i l i t y and Characterization of Wheat Genetic Stocks
Bikram S. G i l l Department o f Plant Pathology
Kansas State Universi ty Manhattan, Kansas 66506
Bob McIntosh, Gary Hart, Rosalind Morris, and others have done a tremendous job i n compilation o f genetic information i n wheat every year. However, i t i s my opinion tha t a l i s t o f genetic stocks t h a t are read i l y ava i lab le w i l l do much t o promote genetic research i n wheat. For over a year now, I have been working on the compilation and col lec- t i o n o f wheat genetic stocks. A genetic stock may contain a gene(s) (mutants, proteins), a DNA sequence, a polymorphic chromosome band, a chromosome deletion, o r an a l i e n t ransfer t o a wheat chromosome tha t can be scored by phenotype and/or cytology. As accepted by wheat genet ic ists, Chinese Spring wheat contains the standard a l l e l e i n every case. The genetic stocks can be maintained a t as many locations as i s desirable and i n many d i f f e r e n t countries.
Apart from th is , many other steps need t o be taken t o make sure wheat f inds i t s due place a t the cu t t ing edge o f genetics. We cannot f a i l t o do conventional mapping before we can jump i n t o molecular genetics. I not ice that, f o r example, many disease res is tan t genes remain t o be mapped and a vast major i ty o f genes are not i n the same genetic background (i.e., Chinese Spring). Simi lar ly, f o r many genes, a l l e l i c var iants need t o be discovered so that they can be useful ly employed i n genetic mapping studies. Many other challenges are read i l y apparent from a c r i t i c a l examination o f the genetic stock l i s t .
I can also see a need f o r depict ion o f genetic maps s im i la r t o corn o r tomato showing map posi t ions f o r a l l genes beginning w i th 0 ra ther than the centromere.
I n compiling a l i s t o f mutant stocks and seed stocks, I have ca l led upon many wheat genet ic is ts who have graciously responded t o my requests. The attached l i s t represents the f i r s t e f f o r t i n documenting the genetic stock and i t s a v a i l a b i l i t y . We have established a Wheat Genetics Resource Center a t Kansas State Universi ty and w i l l be pleased t o accept seed stocks, o r t o add stocks t o the l i s t , showing where the stock i s avai lable.
The K-State accession on number i s f o r our recording-keeping and i n an eventual l i s t o r i g i na l source w i l l be c i ted. I hope tha t the compila- t i o n o f t h i s information on wheat genetic stocks w i l l be useful. I am sure there are many omissions o f information and e r ro rs i n the l i s t and I w i l l appreciate i f you w i l l b r ing these t o my at tent ion.
w b a t ~ s t o & t i What Curatits Mmwa L
State Wvacdty
Bedto i 8 r l h , l9m R y r r e t s l . , 1982
r al., i s 2 mymetal . , 1982 n o & r ~cmtt , in8
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a3008
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mapa at al., 19BS
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w iwa&m Phra ZA3 998
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Afnarorth et al., 1984
Rlym et al . , 1962 Bsnito L SBlinM, 1983
D.R m t k t t in et al., 1978
nettin et al., 1978
m t i n et al . , 1978
ncttin et al., 1978
SS man Hetzger L Siltlfa@, 1970
Schick et al., 1369
Gaines et al., 1963
Starbord, 1W
UhQZ Airnorth, 1963 Chjedd & al., 1983
Qlojecki et al:, 1983
OlgjedLi et al . , 1983
Airnorth et al. , 1981
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layne et al. , 1982 b l a n d c Kerber, 1974
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R A. lCInLoeh Nemml L Bart, 1966 RA ncIntoeh eermaan L Waningo, 1972 Worland et al., 1980 Piraabb L Weleh, 1975 Seare, ncIntoab WIntorh et al . , 1982 Johnston L Rym, 1964 waer 1966 Sramey 6 Luthra, 1970
M amp mrrrf s Y2I -em S 193 waximm S
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a csSrh. 389.16 & WCim 3B
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Piatro L Bart, 1985
Barbar at al., 1968
Jaaslra, 1980 Ainan#th, 1984
ncIntodl at al., 1985 Smrr L Wangton, 1970 Smrs 5 m e , 1968 Bart
&bQL E. R Sars & R Smre E. R Sara Gate L ncIntosh, 1979 sheen L Wac, 1964 S&sn L Sny&r, 1964
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MIntoah L qdC, 1975
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llcIntoeb 6 Baker, 1968 RJ. R t z g r RJ. h g r Ilucd 6 ilcGimia, 1968
amre et al., 1.%7
RA. mntd
llczntorh at a l . , 19el
e R 9 u a W i l l * L Web, 1976
Care at al., 1967
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m o a h , 1978
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AhtbQC Aeyla L Limrs, 1953 B q r m c Livas, 1%3 Tmmwaki c Kihara, 1961 sears, 1982 -6, 1982
Pietro L Bart, 1985 Barber et al., 1%8 Jaaska, 1980 Ainarorth & al . , 1984 Airrcworth et al., 1984 Airrar~~th ct dl., 1981
Hart, 1975 prt et a l . , 1976
!all@ Am Redlad height , S
rn (semi am.) s EPiJ GI inhibitor s g y m & s
Blebnr w e r mrehali, 1976 Gale et al. , 1975
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Mworth, 1983 Aart, 1973 Drimll r Bielig, 1968
O r i e l i Bielig, 1968 Srate r Drisaoll, 1963 raikova ct al., 1980 D r i d l , 1975 Drieaall, 1975 -1- I RiWoll, 1981 0.B. Falk
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ml 1963 nETntorh.tal., 1968 Dyck L Kerbcr, 1981 Iaegering L Scers, 1966 m r e i n g L Sears, 1%6 Christopher et al., 1985
A?wnL Gale et al . , 1975 Gale et al., 1975 Hart, 1970 Rart Hart L Tuleen
Benit0 et al . , 1984 Afmorth et a l . , 1983 A$nruorth et al., 1983
Atmorth et al., 1983 Bart, 1973
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Elva a 17714 8)
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Lar at al., 1975 Airraracth e a l . , 1983
Alrv0rth.tal.I 1 9 a Alnworth et al., 1983 Abmltxth at al., 19a -
Pletro L Bart, 1985 D. R lmott IWK- 1963 Rily & (hnaun, 1967 Gallm r Futtermn, 1977 Gmllm L Pattareon, 1977 Q r l m & al., 1978 Q r l m et al. , 1978
Stock- Euws &Hairy p d n ~ l e S cs -2 Tp3 821 Sears, 1967
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li&ku Iattcrn et a l . , 1983 Briggle, 1%9 - Hart Pietro & B a t , 19BS Oleniak, 1984 Lillr et al., 1975
mJgaleY, 1972 Johnston & B q n , 1964 Rnott & &Into&, 1978
Pietro & Bart, 1985
&&lac Paynaetal., 1sa Bart Hart
kIntoah, 1972 rsy al., 1973 Bart
AwulQc -8, 1964 Work & awl, 1984 Worak r chat, 1984
Paymetal., 1983 Bart
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lay et al., 193
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lay et al., 1973 Gale et al., 1983 Bart
Cheniak, 1984 Bart, 1983
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Ilw, 1966, Uw r Walfe, 1966 - Jaaela, 1980 Gale e t al., 1983
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J O R a
SUmARY WORKSHOP SESSION I 1 I
A. Lane Rayburn. Recorder
Nomenclature Proposals
Chromosome Morphology Nomenclature
Dr. B. S. G i l l has proposed a nomenclature system f o r the designa- t i o n o f C-bands i n wheat. This proposal i s based on the human G-band nomenclature system and uses the c u l t i v a r Chinese Spr ing as a standard. Several ob jec t ions were r a i s e d t o the acceptance o f t h i s system. The two s t rongest ob jec t ions were the use o f the p and q designat ions and the designat ions o f nega t i ve l y s ta ined regions as bands. It was po in ted o u t t h a t wheat g e n e t i c i s t s have used p and q t o designate homoeology w i t h t h i s system wh i l e o thers use p and q t o designate s h o r t and l ong arms, respec t i ve l y . This w i l l have t o be resolved before any nomencla- t u r e system i s adopted. As t o the naming o f the bands, Kimber proposed an a l t e r n a t i v e system i n which the bands are designated by t h e i r d i s - tance from t h e centromere as a percent o f the arm length. Several problems were a l s o noted i n t h i s system. I n o rder t o address more s p e c i f i c a l l y these and o the r problems an ad hoc comnit tee on c h m s o m e morphology nomenclature was formed. The members i nc lude B. S. G i l l (Chair) , J. DvoFik, P. Gustafson, G. Kimber, A. L. Rayburn, and J. Snape. Th is comnit tee w i l l discuss nomenclature a l t e r n a t i v e s among themselves and present a rev i sed nomenclature a t the nex t workshop.
RFLP Nomenclature
Dr. G. Ha r t presented a nomenclature system proposal, developed i n c o n s u l t a t i o n w i t h Dr. M. Gale, mod i f ied from t h e system al ready used t o descr ibe RFLPs by human gene t i c i s t s . I n t h i s system the f i r s t code l e t t e r s r e f e r t o t h e type o f c lone used (CD-cDNA, GD-genomic DNA, etc . ) , t h e second i s a number assigned by the o r i g i n a t i n g lab, and t h i r d i s a l a b designat ion. A f t e r some discussion, Dr. E. Coe was asked t o descr ibe how t h e maize g e n e t i c i s t s were approaching t h i s problem. He s a i d t h a t w h i l e many ways e x i s t t o designate RFLPs, any system adopted should have the th ree bas ic c h a r a c t e r i s t i c s o f uniqueness, c l a r i t y , and s i m p l i c i t y . The system then described by Dr. Coe was b a s i c a l l y s i m i l a r t o Dr. H a r t ' s proposal. Discussion then proceeded t o a d d i t i o n a l prob- lems such as locus and a l l e l e designat ion. Again i t was determined t h a t a c o n n i t t e e was needed t o modi fy the proposal. The comnit tee on gene and l o c i des ignat ion cons is ts o f G. Ha r t (Chair ) , R. Appels, J. Dvokk , M. Gale, 8. G i l l , R. McIntosh, S. Muthukrishnan. J. Antoni Rafa lsky and L. Rayburn. Th is comnit tee w i l l r e p o r t a t the nex t workshop.
K u l t i p l e Genetic Stocks
It i s apparent t h a t w i t h a l l the genet ic s tocks i n ex is tence ways t o o b t a i n l i s t i n g s o f these m a t e r i a l s as w e l l as t h e i r a v a i l a b i l i t y a re needed. These m a t e r i a l s a r e m u l t i l o c a t i o n ma te r ia l w i t h l i t t l e informa- t i o n as t o what i s a v a i l a b l e and how t o go about request ing the inater l - a l . A d d i t i o n a l quest ions were 1) a r e these c o l l e c t i o n s mainta ined i n
permanent storage and 2) a re these ma te r ia l s a v a i l a b l e i n working collections? It was determined t h a t the r e s p o n s i b i l i t y t o d i s t r i b u t e the m a t e r l a l r e s t s w i t h t h e o r i g i n a t i n g labs, bu t t h a t comnon stocks cou ld be d i s t r i b u t e d f rom a c e n t r a l l oca t i on . A comnit tee was fonned f o r the reassignment o f t he c o l l e c t i o n f o r d i s t r i b u t i o n . The c o r n i t t e e c o n s i s t o f P. Gustafson (Chair ) , R. A l lan, K. Amstrong, J. DvoF6k. 0. G i l l , L. Joppa, E. Kerber, G. Kimber, D. Knott, J. Kuspira, S. S. Maan, R. Mor r is , C. Qualset, E. Sears, N. Tuleen, and G. Waines. Th is comnit- t ee i s t o con tac t labs invo lved w i t h genet ic stocks d i s t r i b u t i o n and t o determine what stocks a re ava i lab le . I n add i t i on , a quest ionna i re i s t o be sent t o these labs t o ob ta in in fo rmat ion on maintenance and d i s t r i b u - t i o n o f t h e stocks.
Organizat ion o f Future Workshops
Consensus was t h a t t h i s workshop had accomplished i t s major goals and t h a t such workshops should cont inue i n t he fu tu re . Since t h e IWGS i s i n June 1988, t h e n e x t workshop was suggested t o be he ld e a r l y i n 1987 i n o rde r t o coordinate proposals t o be presented a t t h e IUGS. While several p o s s i b i l i t i e s e x i s t e d as t o the t ime and s i t e o f t h e n e x t workshop, t he most favorab le o p t i o n appeared t o be having t h e workshop imned ia te ly f o l l o w i n g the S tad le r symposium on March 18-19 a t Columbia. MO*. The comnit tee responsib le f o r t h e o rgan iza t i on o f t he workshop cons i s t s o f G. Kimber (Chair ) , P. Gustafson, G. Hart , L. Joppa, E. Kerber, J. DvoHk, and R. Mor r is . The l a s t i t em o f business was the d iscuss ion o f t he p o s s i b i l i t i e s o f a wheat genet ics pub l i ca t i on . The p o s s i b i l i t y w i t h the most support was t o have a sec t i on i n t he annual wheat news le t te r devoted t o t h i s subject . D r . Qua lse t agreed t o l ook i n t o t h i s arrangement, as w e l l as i n t o o the r a l t e r n a t i v e s .
* Deferred t o a s i m i l a r date i n 1988.
APPENDIX I Annual Wheat Newslet ter Vol. 32. June 1986
Proposed GUIDELINES FOR NOMENCLATURE OF BIOCHEMICAL LOCI I N WHEAT AND RELATED SPECIES
1 G. E. Ha r t and M. D. Gale 2
' s o i l and Crop Sciences Department, Texas A&M Un ive rs i t y , Col lege Sta t ion , Texas 77843, U.S.A.
Z ~ y t o g e n e t i c s Department, P l a n t Breeding I n s t i t u t e , , Mar is Lane, Trumpington, Cambridge, CB2 2LQ. England
(These Guidel ines were developed i n con junc t ion w i t h the prepara- t i o n o f t he 'enzymes' sec t i on o f t he 1985 supplement t o the CATALOGUE OF GENE SYMBOLS FOR WHEAT and the 'enzymes', 'non- enzymatic p r o t e i n s ' , and 'nucleolar-organizer-regions' sec t ions o f t he 1986 supplement t o the CATALOGUE. Guide1 ines f o r nomenclature f o r l o c i i d e n t i f i e d by rest r ic t ion- f ragment-polymorphism methodol- ogy are n o t inc luded b u t a re i n preparat ion. We i n v i t e readers t o c r i t i c a l l y analyze t h i s document and t o send us t h e i r conments, suggested rev is ions , etc. We p lan t o present it, a p p r o p r i a t e l y modi f ied, a t t h e 7 t h I n t e r n a t i o n a l Wheat Genetics Symposium.)
The bas ic r u l e s f o r gene symbol izat ion i n wheat, which these gu ide l i nes supplement, a r e contained i n McIntosh (1).
1. Biochemical nomenclature.
Biochemical nomenclature should be i n accord w i t h the r u l e s o f t he Comnission on Biochemical Nomenclature o f the I n t e r n a t i o n a l Union o f Biochemistry. The nomenclature recomnended by the Comnission i s pub- l i s h e d p e r i o d i c a l l y i n major i n t e r n a t i o n a l biochemical journa ls , such as
and the Biochemical Journal. Also, Nomenclature (2) m a y b e s u l ted. b o t h t r i v i a l and formal names. The
c o r r e c t fonna l name should be g iven the f i r s t t ime a substance i s mentioned i n a pub l i ca t i on ; t r i v i a l o r abbreviated names may be used subsequently. For example. ADH i s t he comnonly ysed abb rev ia t i on f o r a l coho l dehydrogenase (E.C.1. I. 1.1; Alcohol : NAD oxidoreductase).
2. Symbols f o r s t r u c t u r a l gene l o c i .
The bas ic symbol f o r a s t r u c t u r a l gene locus should t y p i c a l l y c o n s i s t o f a two-, three-, o r f o u r - l e t t e r abb rev ia t i on o f t h e o f f i c i a l C o n i s s i o n name o f t he enzyme, p ro te in , o r o t h e r substance a f fec ted . The i n i t i a l l e t t e r should be c a p i t a l i z e d and a l l characters i n t h e symbol should be i t a l i c i z e d .
3. L o c i s p e c i f y i n g the s t r u c t u r e o f s i m i l a r non-enzymatic p ro te ins , s i m i l a r enzymes, and s i m i l a r RNA molecules.
N o n a l l e l i c l o c i which spec i f y t h e s t r u c t u r e o f s i m i l a r non- enzymatic p r o t e i n ~ , o f enzymes t h a t cata lyze the same o r s i m i l a r reac- t i ons , o r o f s i m i l a r RNA molecules should be designated by the same bas ic symbol. The l o c i a re designated i n accordance w i t h one o r t he o t h e r o f two procedures:
(a ) Loc i which are members o f a homologous s e t a re designated w i t h the bas ic symbol fo l lowed by a hyphen ( ' I - ' I ) , the accepted symbol f o r t he genome i n which t h e locus i s located, and a homologous s e t number i n t h e fonn o f an Arabic numeral. For example. Adh-Al, Adh-B1, Adh-Dl, and Adh-El designate the A, 0 , D, and E g e n o m m e r - p e m y , o f the f i r s t d e s i g n a t e d homologous s e t o f a lcoho l dehydrogenase s t r u c t u r a l gene l o c i .
(b ) I n t h e absence o f evidence t h a t l o c i are members o f a homlo - gous set, they are designated i n sequent ia l se r i es by a comnon bas i c symbol f o l l owed imnediate ly by an Arabic numeral.
Rye l o c i should be designated i n accord w i t h these procedures [see(3)] . For ba r ley l o c i , procedure "a" should be used when designa- t i o n o f a locus as a member o f a homologous s e t o f T r i t i c e a e l o c i i s desired; otherwise, ba r ley locus nomenclature should be employed [see. e.g., (4 ) ] . Thus, f o r example. Adh-H1 and Adh-R1 designate t h e H and R genome members, respec t i ve l y , o f - dh-1 1 o c i .
Evidence regard ing phylogenet ic r e l a t i o n s h i p s among genes may be obta ined by comparative s tud ies o f (1 ) t he p rope r t i es o f gene products, (2) t he nuc leo t i de sequences o f genes, and (3 ) t h e intrachromosomal map p o s i t i o n s and phys ica l l oca t i ons o f l o c i i n homoeologous chromosomes o r segments. Strong evidence i s requ i red t o warrant t he conclus ion t h a t n o n a l l e l i c gene l o c i comprise a homologous set . C r i t e r i a f o r detennin- i n g whether gene l o c i t h a t encode isozymes are o r a re no t homologous and, f o r homologous gene l o c i , whether they belong the the same o r d i f f e r e n t homologous sets, a r e descr ibed i n (5 ) . Most o f t h e c r i t e r i a 1 i s t e d are e q u a l l y app l i cab le t o non-enzymatic p ro te ins . The evidence which i s t h e bas is f o r des ignat ing gene l o c i as members o f a homologous s e t should be c l e a r l y s ta ted i n t he p u b l i c a t i o n i n which symbols f o r t he l o c i a r e proposed.
4. A l l e l e s .
D i f f e r e n t a l l e l e s are designated by a lower-case Roman l e t t e r f o l l o w i n g t h e locus designat ion. For example, %-Am -Ala and @-Am -Alb a r e two a l l e l e s o f t he A genome $-Am -1 locus. ne s r a i n shou
7-57 +
designated the pro to type s t r a i n o r eac a l l e l e discovered, s ince v a r i a t i o n t h a t has n o t been detected by the methods used may be present w i t h i n each a l l e l i c c lass . Chinese Spr ing should be t h e pro to type f o r a l l e l e "a". I f an apparent ly i d e n t i c a l a l l e l e i n o t h e r s t r a i n s i s found by new methods t o be d i f f e r e n t from t h a t i n t he pro to type s t r a i n , i t should be assigned a new Romn l e t t e r and a pro to type s t r a i n designated. Th is system a l l ows t h e o r d e r l y assignment o f symbols t o newly i d e n t i f i e d
a l l e l e s and a i l ows ready comparisons o f new v a r i a n t s w i t h p rev ious l y repo r ted var ian ts .
5. Gene complexes.
Gene complexes, a l s o c a l l e d compound l o c i , c o n s i s t o f a number o f f u n c t i o n a l l y r e l a t e d genes t h a t a re genet ica l l y c lose1 y 1 inked. Whether composed o f a few o r many genes, a gene complex should be assigned one symbol, i n accord w i t h (2 ) and (3) above. Guidel ines f o r des ignat ion o f t he i n d i v i d u a l genes t h a t compose gene complexes have no t y e t been developed.
T r i t i c e a e enzyme and p r o t e i n gene l o c i a re comnonly i n i t i a l l y i d e n t i f i e d and assigned designat ions based on s tud ies of aneuploid s t r a i n s which l a c k and/or con ta in e x t r a copies o f whole chromosomes o r telosomes. Consequently, evidence may be obta ined f o r t he product ion o f two o r more s i m i l a r enzyme o r p r o t e i n protomers by one chromosome a m w i thou t genet ic evidence as t o whether o r n o t t h e protomers are t h e products o f one gene, o f d i f f e r e n t genes t h a t a r e members o f a gene complex, o r o f two o r more genes which a re n o t members of one gene complex. I n these s i t ua t i ons , on l y one locus des ignat ion should be assigned t o a chromosome a n f o r s i m i l a r p r o t e i n s o r enzymes unless s t rong biochemical evidence i nd i ca tes t h a t t he protomers a r e the products o f genes which are n o t members o f one gene complex.
6. Phenotype symbols.
The same two-, three-, o r f o u r - l e t t e r abb rev ia t i on o f the biochemi- c a l t h a t i s used as t h e bas ic genotype symbol should be used as t h e bas i c phenotype symbol bu t w i t h each l e t t e r c a p i t a l i z e d . For biochemi- c a l s encoded by the members o f a homologous s e t o f l o c i , t h e p h e n o t ~ p ; ~ ~ symbol should c o n s i s t o f t he basic symbol fo l lowed by a hyphen ("-" the same Arabic numeral as i s contained i n t he genotype symbol.
References
1. McIntosh, R. A. 1983. A catalogue o f gene symbols f o r wheat (1983 e d i t i o n ) . Proc. 6 t h I n t e r . Wheat Genetics Symp., pp. 1197-1254.
2. Enzyme Nomenclature. Recomnendations (1984) o f t he Nomenclature Conni t tee o f the I n t e r n a t i o n a l Union o f Biochemistry. 1984. Academic Press, N.Y.
3. Sybenga, J. 1983. Rye chromosome nomenclature and homoeology r e l a t i o n s h i p s . Z. Pflanzenzcchtg. - 90:297-304.
4. Brown, A. H. D. 1983. Bar ley. I n : I soz mes i n P l a n t Genetics and Breedin S. D. Tanksley and T. J r t o n (ms . ) . E l +:
-u-Y- s e v i e r m e n c e u i she rs B.V., Amsterdam.
5. Hart . G. E. 1986. Genetic and biochemical s tud ies o f e n z w s . I n : .wheat. E. G. Heyne (Ed.). American Soc ie ty o f ~ ~ r o & m y , M a d i s r ( I n press).
APPENDIX I1
I-3 R.lr lor G u a S y m b d h d a h What (Myad fram the Inmrudorul Rda d G d N a m m c b ~ )
I. Inlvm~yMiarytctorrtbci lrd-ofbi ( l la incaruciacul i tyM klivm-.
2. S y m M ot W i u y facton, derived fmm their or+d n~na, lbould iniul#, a in Ronvn k t m d d h i n d w typ.
3. W&ncver unambiguous, h e rume and lymbd of a d o m h t lbould with c a p i d l e t c a d chaeofarrceive w i t h a d l e t t c r (rc &spcd.lruLakr ymbdiziq hat: pathogen wtcrm).
4. ~ A U d n ~ k n u r d i n r y m b d i u ~ ~ k w r i t t c n o l r c l u ~ ; U ~ u pdbk no uperrrip or subrripa rbould be urd.
5. The plu ("+") + will not k uwd in r y m b d a o d hardiuy helam in rbaL 6. T w n a m o r e # e n a h . ~ ~ ~ ~ r b o u l d b ~ ~ h
a ranmm buic lymbd. Noa-rlklic &i (mi* pdymaic W) dl b d a h t e d in .ccord.na with nro pardurn: (i) in rguadJ, pdymcric aaia w k UI Anbic a d hmdhtdy blia tk
gem Iymbd; e.6.. S d . (ii) i n h o ~ ~ w h m t h e b v i c y m b d h r o l b n d b y a ~ ( " - " )
followed by tbc laur daipation uLiql the fanr dthe .oscpcsd g a w ~ qnhl d a h o m a d q w r t n u m b a ~ t c d b y . a ~ n u n m l ; c g . . r l M l d a i @ . c a t k A ~ m e m b a d t h e d n c A A + c .
Beauwtheletca"I"dnumbcr"I"nuy kcwhrcd,thcfumerrbouldbeavoidd utbelucletterinabuicymbd. DilTerrnt.Ue14or.UJsdhkpda~mu- uliolvl~,arrd~tdbyak-R-*nu~rhbcunumkr daipation; e.6.. Srik, A B A 1..
6.. Tanpwuy symbol daignrtioar. Whre dam ue not avdabk, pmvish hu ban made for tanpony symbob. lime rh31 wnnim d the bdc ymbd f d l o w e d b y m . b b n L ~ f o r t h c L i a r o r s t o & d l o A n b i c n u m b a ~ t o the p; e.g.. SrFrl, SrFr2, etc. rdcr to twn g a m for & to PPrLd. F.r*t in cu&iw Fedei.tiaa It is mammadcd th.1 ofhiJ rrcorb of temporary daig- rutiom be kept, but it is not a n r i d th.t subquent numben from other h h m r i u (e.~., SrFrS) be checked a & u t urlier n u m h eitha phenotypialty or g u d d l y .
7. Inhibiton, s u p p r a ~ , md mhuvm uc d-ted by the rymbdr I, Sr. d h, o r b y i , n , ~ a X t h y u c r m r i v c , f d l o w d b y a ~ p . a . I l d c h e l y m b d d t b c allele aRutcd.
8. WILL NOT APPLY. Whenever convenient, lettub should k dai(putd by th ktca I a L, and sterility and incompatibility gcna by J or S.
9. I s what 4d rchtd apecia, linkage poup and amupond'ing dvomaome u e daigmrcd by .a Anbic numml (1-7) followed by genome d d p u e d by a a p i d R- letter; i.e., for hauploid what d p u p wriora (Momh and QM
0, I A - l a Thh m*, ah a d & d * RmMm -&I-XXI. - * L m r L u i . t - c L . c d J c b u . -- - ~ ( w * b a u L - d r i w r i r r ) w ~ L ~ d u d ( ~ ) . . W L r r e b . I # = W r n ~ . a n d C . . ~ ~ ~ ( b C ~ ~ t o t L I ~ --L--kLd.d ~ ) d ~ h v e b a p o p o m d .
la W111.NOTAPH.Y. T b . b n e n X d Y u c ~ t o A j . j y l . ~ - I I . O r L ~ ~ \ * r i a e . a u ~ w i t h t l m ~ . I * b ~ O n t w _ F
& I L & ( p l - m 8 4 k k 4 w p o u p 12 Q ~ r b m c i o c r u b - b b L I - . .
Mbr-. Dpbrdqbdm,InfRL*ria,TfRmrbcda..dTpbr I.rbutb.l.~.n~d~dai*cdIhmnhcdrl*dobyiaDrpir, S r P L ~ i a d i a n u i r r P a a d d e ~ ~ b e u i 0 . r -*lab mtmmsadd. Ocbshariaril lh " InPupiQrnluirrcL. -rPdud W h a a g r d o r a a d i s i ~ r v l v a ~ p d c P C t L c m v c h r a P o v r ~ m y b D ~ i D b a c l . a h l b r i q c L , ~ ~ ; e a . * 4 r n ( T P ) n h r r n . L t s ~ t L . ~ " b . i l d ' ~ m c b n m r a m c Q ) S a l d 4 A w & h L o L - m ~
IS. l%erlpdc.rabrdc&amammihAi.rrAbl)4kill.YI*db. dhb&;byz
14. Syd&b-bbmWhaPI .rdwidhhuhd p s r r d c c b w -
APPENDIX I11
Summary of the taxonomic and nomenclatural conspectus of the Triticeae advocated by bkell Lave (1984. Conspectus of the
Triticeae. Feddes Repertorium 95:425-521.)
This system divides the tribe into thirty-seven genera based on twenty-three single-haplome taxa (Lave. 1982. Generic evolution of the wheatgrasses. Biologisches Zentralblatt 101:199- 212.). Thus a genus consists of all species which are found by genome analysis to possess s single given haplome or autoploid combinations of that haplome. For example, the genus Crfthodlur, consists of the two diploid species with the A haplome, C. monococcum and C. urartu. For alloploid genera, one genus consists of species with combinations of given haplomes. Bor example, the genus Aegilops consists of the four tetraploid species and single hexaploid species which possess the M and U haplomes .
This system is dependent upon knowledge derived from genome analysis. Future work may reveal that some genera now defined with one haplome actually involve an additional haplome. This is especially likely in large genera such as Crlteslon which contains many species for which genome analysis has not been made. As such information accumulates, the system can accomodate it with appropriate taxonomic changes without changing the foundation of the system.
Readers are urged to consult the original papers cited above for further information. Acceptance of the system is growing. For example, D. R. Dewey has accepted it in principle and in practice. with some exceptions, in his recent treatment of North American wheatgrasses (The genomic system of classification as a guide to intergeneric hybridization with the perennial Triticeae. p. 209-279 in J. P. Gustafson, ed. 1984. Gene Manipulation in Plant Improvement. Plenum Publishing Corp.)
Synonvm
T r f ti cum ronococcum
T . bfcorne
T. d i c h a s i a n s
T . t a u s c h i f
T . t r f p s a c o f des
T . comosum
T . u n i a r f s t a t u m
T . u n b e l l u l a t u m
T . t u r g f d u m
T . a e s t f v u m
T . kotschyi
T . c y l i n d r i c u m
T . t r i u n c i a l e
T . ven t r f cosum
T . j u v e n a l e
S p e c i e s name a s Genome symbol advoca ted by Lave ~ a v e O l d
C r i thodi um monococcum A A
S f t o p s f s bicornfs B sb
Orrhopygi um cauda tum C C
Pa t ropyrum t a u s c h i i D D
~ m b l yopyrum m u t i cum z nt Comopyrum comosum M M
Chennapyrum u n f a r f s t a tum L U n
Kfharapyrum umbe l lu la tum U U
G fgach f I o n p o l onf cum AB AB
T r i t i c u m a e s t i v u m ABD A B D
~ e g i I emma k o t s c h y i BU us1
C y l f n d r o p y r u m c y l f n d r f cum C D C D
Aegf l o p o d e s t r f u n c f e l f s CU UC
Gas tropyrum v e n t r i c o s ~ m DM Dun
Aegf l onearum j u v e n a l e DMU DMU
A e g i l o p s g e n f c u l a t a MU UM
Examules of other taxa in the Triticeae
Species name as LOve ' s svnonvm advocated bv LOve Genome
Secal e cereal e Secale cereale R
Dasypyrur vf llosum Dasypyrum vil losum V
Breropyrur orf en tal e Eremopyrum orientale FF
Heterantheliur piliferum Heteranthelium piliferum Q
Henrardia persica Henrardia persica 0
Hordeum vulgar8 Hordeum vulgare I
H. secalinur Cri tesion brevisubulatum HH
Crf thopsis delileana Cri thopsis del ileana K
Taenia therum Taenia therum capu t-medusae capu t-medusae
Horde1 ymus europaeus Hordelymus europaeus HT
Festucopsf s serpen ti ni Fes tucopsis serpen tini G
Agropyron crf sta tum Agropyron cris ta tum PP
A. elongatur Lophopyrum elonga tum E
A. junceum
A. strigosum
Thinopyrum junceum JJJ
Pseudoroegneria s trigosa
A. repens Blytrigia repens EJS
A. smf thff Pascopyrum smi thi i HJNS
El ymus sf bi ri cus Elymus sibiricus HS
E. lanugfnosur
B. arenarius
Psa thyros tachys N lanuginosa
Leymus arenarius JN
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