Recommendations for the designation of the barley chromosomes and their arms

Ib Linde-Laursen
Department of Botany, Dendrology and Forest Genetics,
The Royal Veterinary and Agricultural University,
Rolighedsvej 21, DK-1958 Frederiksberg C, Denmark.

At the business meeting of the 7th International Barley Genetics Symposium held at the University of Saskatchewan, Saskatoon, Saskatchewan, Canada, on August 5, 1996, the following resolution on the designation of the barley chromosomes and their arms, and on the selection of a reference genome in the Triticeae was passed.

1. Each of the seven barley chromosomes is designated by a figure from 1 to 7 according to its homoeologous relationships with chromosomes of other Triticeae species. The figure is followed by the letter H, e.g. 2H.

2. The genomes of Hordeum vulgare and H. bulbosum are symbolized by the letter H.

3. The chromosome arms are designated by the letters S and L.

4. The barley genome present in the variety 'Betzes' becomes the reference genome in the Triticeae to which definitions of translocations, short arm/long arm reversals, etc. are standardized in all species.

Notes on the four points of the resolution:

1. The seven barley chromosomes should be designated as follows:

Recommendation:

1H 2H 3H 4H 5H 6H 7H

Burnham and Hagberg designations (according to Singh and Tsuchiya 1982):

5 2 3 4 7 6 1

It is not recommended to give the Burnham and Hagberg numbering together with the homoeologous numbers in publications, although authors may choose to do so, with the Triticeae nomenclature given prominence. It is MOST IMPORTANT that the genomic symbol H is included with the number, when the Triticeae system is used. Otherwise readers will tend to confuse the number with the Burnham and Hagberg numbering system.

2. The use of the genomic symbol H instead of I as proposed by Löve (1984) agrees with usage by most barley workers, but goes against a proposal for the designation of all Triticeae genomes presented in the Proceedings of the 2nd International Triticeae Symposium (Wang et al. 1996).

3. The two arms of the chromosomes of barley are designated by the upper case letters S (for the short arm) and L (for the long arm) according to the definitions by Singh and Tsuchiya (1982). The arm designation should follow the chromosome and genome designations without a space, e.g., 6HL. The arm designations given in recent molecular marker maps (e.g., Kleinhofs et al. 1993; Devos et al. 1993) should henceforth be regarded as 'names' not as referring to physically long and short arms, so no further reversals will be made.

4. The selection of the barley genome in the variety 'Betzes' as the reference genome in the Triticeae is based on the fact that barley is an excellent standard. It is diploid (unlike wheat), easy to grow and multiply under laboratory conditions worldwide (unlike Aegilops species), wheat addition lines are available for 'Betzes', and barley is economically important so that other Triticeae workers have an incentive to test their probes, etc. on barley.

Contents of the resolution had previously been discussed and passed at a workshop "Designation of the Barley Chromosomes" on August 2. The discussion was based on recommendations drawn up by a committee consisting of I. Linde-Laursen, Copenhagen, Denmark; J.S. Heslop-Harrison, Norwich, United Kingdom; K.W. Shepherd, Adelaide, Australia; and S. Taketa, Kurashiki, Japan. The committee had been established by the organizers of the 7th International Barley Genetics Symposium with the mandate to prepare recommendations for a standard nomenclature system of the barley chromosomes on the basis of a literature survey of the barley genome. The problem was last discussed at the 6th International Barley Genetics Symposium in Helsingborg in 1991 (Lundqvist and Franckowiak 1992).

On the basis of the information available in the literature, it was obvious that:

(i) the seven chromosomes of barley (H. vulgare L.) largely have the same genetic content as those in other members of the Triticeae; and

(ii) the gene loci in barley are largely collinear with the loci in other members of the Triticeae, with few ancestral translocations involving whole chromosome segments.

All available data thus show a high level of conserved synteny among the chromosomes throughout the Triticeae, and it is clear that the whole group can be treated as a single gene pool. Therefore, the committee recommended that the currently used designations of the chromosomes of barley (Burnham and Hagberg 1956; see Singh and Tsuchiya 1982) were changed to follow a system in accordance with the current wheat homoeologous group numbering (the Triticeae system). This point-of-view was adopted by the barley symposium.

References:

Burnham, C.R., and A. Hagberg. 1956. Cytogenetic notes on chromosomal interchanges in barley. Hereditas 42:467-482.

Devos, K.M., T. Miller, and M.D. Gale. 1993. Comparative RFLP maps of the homoeologous group-2 chromosomes of wheat, rye and barley. Theor. Appl. Genet. 85: 784-792.

Kleinhofs, A., A. Killian, M.A. Saghai Maroof, R.M. Biyashev, P. Hayes, F.Q. Chen, N. Lapitan, A. Fenwick, T.K. Blake, V. Kanazin, L. Dahleen, D. Kudrna, J. Bollinger, S.J. Knapp, B. Liu, M. Sorrells, M. Heun, J.D. Franckowiak, D. Hoffman, R. Skadsen, and B.J. Steffenson. 1993. A molecular, isozyme and morphological map of the barley (Hordeum vulgare) genome. Theor. Appl. Genet. 86:705-712.

Löve, A. 1984. Conspectus of the Triticeae. Feddes Repert. 95: 425-521.

Lundqvist, U., and J.D. Franckowiak. 1992. Committee and workshop summary. Mutations, gene mapping and nomenclature. p. 855-857. In L. Munck, K. Kirkegaard, and B. Jensen (eds.). Barley Genetics VI. Proc. Sixth Int. Barley Genet. Symp., Helsingborg, 1991. Munksgaard Int. Publ., Copenhagen.

Singh, R.J., and T. Tsuchiya. 1982. Identification and designation of telocentric chromosomes in barley by means of Giemsa N-banding technique. Theor. Appl. Genet. 64: 13-24.

Wang, R.R.-C., R. von Bothmer, J. Dvorak, G. Fedak, I. Linde-Laursen, and M. Muramatsu. 1996. Genome symbols in the Triticeae (Poaceae). p. 29-34. In R. R.-C. Wang, K.B. Jensen, and C. Jaussi (eds.). Proc. 2nd Int. Triticeae Symp., Logan, Utah, 1994. Utah State University, Logan.

Formal approval of the recommended Rules for Nomenclature and Gene Symbolization in Barley was given at the Seventh International Barley Genetics Symposium held July 30 to August 6, 1996 at the University of Saskatchewan, Saskatoon. The proposed Rules for Nomenclature and Gene Symbolization in Barley were published in BGN 2:11-14 and BGN 11:1-16. The recommendations as approved, plus a few modifications, are given below.

We were requested to use three letter locus symbols in all new and revised Barley Genetic Stock (BGS) descriptions. Also, replacement of the comma comma scheme for naming new mutants was suggested. To accomplish these tasks, the Rules for Nomenclature and Gene Symbolization in Barley were modified by adding SUPPLEMENTARY AMENDMENTS. Using a period between the locus and allele portions of the gene symbols was recommended to replace the comma comma scheme. Use of a separator to designate allele symbols was proposed by von Wettstein-Knowles (1993, Barley Newsl. 36:182). A supplementary amendment concerning the recommended names for barley chromosomes was added.

1. In naming hereditary factors, the use of languages of higher internationality should be given preference.

2. Symbols of hereditary factors, derived from their original names, should be written in Roman letters of distinctive type, preferably in italics, and be as short as possible.

AMENDMENT: The original name should be as descriptive as possible of the phenotype. All gene symbols should consist of three letters.


COMMENTS: All new gene symbols should consist of three letters. Existing gene symbols of less than three letters should be converted to the three letter system whenever symbols are revised. When appropriate, one or two letters should be added to existing symbols.

For example, add the letters "ap" to "K" to produce the symbol "Kap" to replace "K" as the symbol for Kapuze (hooded). As another example, add the letters "ud" to "n" to produce the symbol "nud" to replace "n" as the symbol for naked seed. Similarly the letter "g" can be added to "ms" to produce the symbol "msg" for genetic male sterility and the letter "e" can be added to "ds" to produce the symbol "des" for desynapsis. When inappropriate or when conflicts arise, questions should be referred to the Committee on Genetic Marker Stocks, Nomenclature, and Symbolization of the International Barley Genetics Symposium for resolution.

3. Whenever unambiguous, the name and symbol of a dominant begin with a capital letter and those of a recessive with a small letter.

AMENDMENT: When ambiguous (co-dominance, incomplete dominance, etc.) all symbols should consist of a capital letter followed by two small letters that designate the character, a number that represents a particular locus, and a letter or letters that represents a particular allele or mutational event at that particular locus.


COMMENTS: As an example, the letters "Mdh" can be used to designate the character malate dehydrogenase, "Mdh1" would represent a particular locus for malate dehydrogenase and "Mdh1a", "Mdh1b", "Mdh1c", etc. would represent particular alleles or mutational events at the "Mdh1" locus. Row number can be used as an example of symbolizing factors showing incomplete dominance. At the present time, the symbol "v" is used to represent the row number in Hordeum vulgare, "V" is used to represent the row number in Hordeum distichum, and "V^t" is used to represent the row number in Hordeum deficiens. According to the amendment to Rule 3, if row number were to be designated by the letters "Vul", the designation of the locus on chromosome 2 would then become "Vul1" and the alleles "v", "V", and "V^t" would be designated "Vul1a", "Vul1b", and "Vul1c".

SUPPLEMENTARY AMENDMENT: A period should be placed before the allele symbol in the complete gene symbol.

COMMENTS: Since DNA sequences similar to those of the original locus may occur at several positions in the Hordeum vulgare genome, a three letter symbol plus a number is inadequate to represent all potential loci. Also, both numbers and letters have been assigned to specific mutants and isozymes in Hordeum vulgare. The six-rowed spike locus is used as an example although the symbol Vul1 for row number in Hordeum vulgare is not recommended because the botanical classification of Hordeum spp has changed. The locus symbol vrs1 and the name six-rowed spike 1 are recommended for the v locus. Gene symbols recommended for common alleles at the vrs1 locus are vrs1.a, vrs1.b, vrs1.c, and vrs1.t for the "v", "V", "v^lr", and "V^t" genes, respectively.

4. Literal or numeral superscripts are used to represent the different members of an allelic series.

AMENDMENT: All letters and numbers used in symbolization should be written on one line; no superscripts or subscripts should be used.

5. Standard or wild type alleles are designated by the gene symbols with a + as a superscript or by a + with the gene symbol as a superscript. In formulae, the + alone may be used.

AMENDMENT: This rule will not be used in barley symbolization.

6. Two or more genes having phenotypically similar effects are designated by a common basic symbol. Non-allelic loci (mimics, polymeric genes, etc.) are distinguished by an additional letter or Arabic numeral either on the same line after a hyphen or as a subscript. Alleles of independent mutational origin may be indicated by a superscript.

AMENDMENT: Barley gene symbols should consist of three letters that designate the character, a number that represents a particular locus, and a letter or letters that represents a particular allele or mutational event at that particular locus. All letters and numbers should be written on the same line without hyphens or spaces. Alleles or mutational events that have not been assigned to a locus should be symbolized by three letters that designate the character followed by two commas used to reserve space for the locus number when determined, followed by a letter or letters representing the particular allele or mutational event. After appropriate allele testing, the correct locus number will be substituted for the commas. Where appropriate (when assigning new symbols or when revising existing symbols) letters representing alleles or mutational events should be assigned consecutively without regard to locus number or priority in discovery or publication.

COMMENTS: The use of the proposed system of symbolization can be illustrated by the desynaptic mutants. Two loci are known: lc on chromosome 1 (7H) and ds on chromosome 3 (3H). These will be resymbolized as des1a and des2b. A large number of desynaptic mutants have been collected. They will be designated des,,c, des,,d, des,,e, etc. If allele tests show that des,,c is at a different locus than des1 and des2, des,,c will become des3c. If allele tests show that des,,d is at the same locus as des2, des,,d will become des2d. In practical use, the symbol des will be used when speaking of desynapsis in general or if only one locus was known for the character. The symbol des2 will be used when speaking of that particular locus, and the symbol des2b will be used only when speaking of that particular allele or mutational event. If additional designation is needed in particular symbolization, it can be obtained by adding numbers behind the allele letters, and, if still further designation is needed, letters can be added to the symbol behind the last number. Symbolization consisting of alternation of letters and numbers written on the same line without hyphens or spaces will allow for the expansion of the symbol as future needs arise. In any work with large numbers of polymeric gene mutants, every mutant has to be given a designation not shared by any other mutant of this polymeric group and this designation should become a part of the permanent symbol representing that particular allele or mutational event. This requirement can be met by assigning allele designations in consecutive order without regard to locus number.

SUPPLEMENTARY AMENDMENT: A period should be used instead of two commas in gene symbols for mutants within a polymeric group that can not be assigned to a specific locus.

COMMENTS: The des symbol should be used when referring to desynapsis in general; des1 and des2, for specific loci; des1.a and des2.b for specific genes or alleles at their respective loci; and des.c, des.d, des.e etc., for desynaptic mutants not assigned to a specific locus.

7. Inhibitors, suppressors, and enhancers are designated by the symbols I, Su, and En, or by i, su, and en if they are recessive, followed by a hyphen and the symbol of the allele affected.

8. Whenever convenient, lethals should be designated by the letter l or L and sterility and incompatibility genes by s or S.

AMENDMENT: This rule will not be used in barley symbolization.

COMMENTS: J.G. Moseman (BGN 2:145-147) proposed that the first of the three letters for designating genes for reaction to pests should be R. The second and third letters will be the genus and species names of the pest.

SUPPLEMENTARY COMMENT: A motion was passed during the workshop on "Linkage Groups and Genetic Stock Collections" at the Fifth International Barley Genetics Symposium in 1986 (Barley Genetics V:1056-1058, BGN 17:1-4), that the International Committee for Nomenclature and Symbolization of Barley Genes should "recommend use of Ml as the designation of genes for resistance to powdery mildew."

9. Linkage groups and corresponding chromosomes are preferably designated by Arabic numerals.

SUPPLEMENTARY AMENDMENT: The current wheat homoeologous group numbering scheme (the Triticeae system) is recommended for Hordeum vulgare chromosomes. Arabic numerals followed by an H will indicate specific barley chromosomes. The H. vulgare chromosomes should be 7H, 2H, 3H, 4H, 1H, 6H, and 5H instead of 1, 2, 3, 4, 5, 6, and 7, respectively.

10. The letter X and Y are recommended to designate sex chromosomes.

11. Genic formulae are written as fractions with the maternal alleles given first or above. Each fraction corresponds to a single linkage group. Different linkage groups written in numerical sequence are separated by semicolons. Symbols of unlocated genes are placed within parenthesis at the end of the formula. In euploids and aneuploids, the gene symbols are repeated as many times as there are homologous loci.

12. Chromosomal aberrations should be indicated by abbreviations: Df for deficiency, Dp for duplication, In for inversion, T for translocation, Tp for transposition.

13. The zygotic number of chromosomes is indicated by 2n, the gametic number by n, and basic number by x.

14. Symbols of extra-chromosomal factors should be enclosed within brackets and precede the genic formula.

AMENDMENT:

A. Present designations for genes and mutations. - Most of the present designations should be maintained. However, new designations may be given, when additional information indicates that new designations would aid in the identification of genes and mutations.

B. New designations for genes and mutations. - New genes or mutations will be designated by characteristic, locus, allele, and then the order of identification or mutational event. Three letters will be used to identify new characteristics. Consecutive numbers will be used to identify the order of identification or mutational event. Loci will be designated by numbers and alleles by letters when they are identified. For example, des-6 indicates that this is the sixth gene or mutation identified for the characteristic des (desynaptic). des 1-6 and des 2-7 indicate that gene or mutational events 6 and 7 for the desynaptic characteristic have been shown to be at different loci and those loci are then designated 1 and 2, respectively. des 1a6 and des 1b8, indicate that the gene or mutational events 6 and 8 for the characteristic desynaptic have been shown to be at different alleles at locus 1 and those alleles are then designated a and b.

A motion was passed during the workshop of the "Nomenclature and Gene Symbolization Committee" at the Fifth International Barley Genetics Symposium in 1986 (Barley Genetics V:1056-1058) that "the recommended systems for Nomenclature and Gene Symbolization of the International Committee be published annually in the Barley Genetics Newsletter."

The last linkage map prepared for the barley chromosomes by Dr. T. Tsuchiya was published as the front cover diagram in volume 20 of the Barley Genetics Newsletter. Specific criteria, based on available linkage data and telotrisomic analyses, were used in the development of these maps. Techniques used and the loci included or excluded were questioned by some barley researchers. However, these chromosome maps and their frequent revisions are helpful in learning about barley and its genetic resources. Even though locus positions were not precise, they did provide a convenient means of finding some information about barley genes and estimating their chromosomal locations. Hopefully, the chromosome maps presented here will provide potential users some information about barley genes.

In this volume of the Barley Genetics Newsletter, new or revised Barley Genetic Stock (BGS) descriptions for many of the morphological marker loci are presented. Many BGS descriptions include information about the position of the locus in the barley genome. Based on that information and other published maps, I constructed maps for each of the seven barley chromosomes (Figures 1 to 7). Estimates of locus positions are presented in centi-Morgan units. Relative positions of loci are best guesses of their location based on, in some cases, relatively little recombination data. Since the original data are variable in accuracy and three-point data does not exist from many gene combinations, some locus positions and locus sequence estimates are no better than a guess. Maps developed by Dr. Jens Jensen are much more precise in terms of placing loci in chromosomes and should be given priority when conflicts in marker positions arise.

Names of individual chromosomes and the order in which they are presented was modified to correspond to the Triticeae scheme for naming barley chromosomes (Linde-Laursen, 1996). Revised locus symbols, which are based on a three letter code, are used in the chromosome maps (Figures 1 to 7). Even with relatively few restrictions, some genes associated with specific chromosomes could not be placed in the maps. Morphological markers for which linkage data indicate an association with a specific chromosome are listed in Table 1.

Molecular marker loci, most isozyme loci, and many disease reaction loci are not included in these chromosome maps because time was not taken to review the papers concerning their positions relative to morphological marker loci. Hopefully, this deficiency can be addressed in future versions of these maps.

Many other maps of the barley genes have been constructed. They summarize other data, include different markers, and are often more accurate than those presented here. Those maps should be examined to learn more about specific loci and segments of the barley genome. They may aid, also, in the development future versions of these maps.

References:

Borovkova, I.G., Y. Jin, B.J. Steffenson, A. Kilian, T.K. Blake, and A. Kleinhofs. 1997. Identification and mapping of a leaf rust resistance gene in the barley line Q21861. Genome (in press).

Chicaiza, O. 1996. Genetic control of leaf rust in barley. Ph.D. Thesis. North Dakota State Univ., Fargo.

Falk, D.E. 1985. Genetic studies with proanthocyanidin-free barley. BGN 15:27-30.

Falk, D.E. 1994. Coordinator's report: Chromosome 6. BGN 23:154.

Faue, A.C., A.E. Foster, and J.D. Franckowiak. 1989. Allelism testing of an induced yellow streak mutant with the three known yellow streak mutants. BGN 19:15-16.

Fischbeck, G., and H. Häuser. 1976. Research notes. BGN 6:28-29.

Franckowiak, J.D. 1991. Association of male sterility genes with a specific chromosome using multiple marker stocks. BGN 20:31-36.

Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived lines of spring barley. BGN 24:63-70.

Gardenshire, J.H., N.A. Tuleen, and K.W. Stewart. 1973. Trisomic analysis of greenbug resistance in barley, Hordeum vulgare L. Crop Sci. 13:684-685.

Gymer, P.T. 1977. A new gene for semi-naked grain. BGN 7:33-34.

Häuser, H., and G. Fischbeck. 1979. Genetic analysis of some induced mutants. BGN 9:26-27.

Häuser, H., and G. Fischbeck. 1980. Genetic analysis of induced mutations. BGN 10:30-31.

Häuser, J., and G. Fischbeck. 1976. Untersuchungen zur Lokalisierung einiger Mutationen von Gerste (Hordeum sativum). Z. Pflanzenzücht. 77:269-280.

Häuser, J., A. Jahoor, and G. Fischbeck. 1988. Localization of induced mutants for globe shaped grains. BGN 18:54-58.

Hernandez-Soriano, J.M. 1973. Desynaptic mutants in Betzes barley. M.S. Thesis. Univ. of Arizona, Tucson.

Hockett, E.A., and R.F. Eslick. 1971. Genetic male-sterile genes useful in hybrid barley production. p. 298-307. In R.A. Nilan (ed.) Barley Genetics II. Proc. Second Int. Barley Genet. Symp., Pullman, WA, 1969. Washington State Univ. Press, Pullman.

Hor, K.S. 1924. Interrelations of genetic factors in barley. Genetics 9:151-180.

Jarvi, A.J., and R.F. Eslick. 1975. Shrunken endosperm mutants in barley. Crop Sci. 15:363-366.

Jende-Strid, B. 1993. Coordinator's report: Anthoyanin genes. BGN 22:136-137.

Jensen, J. 1979. Chromosomal location of one dominant and four recessive high-lysine genes in barley mutants. Vol. 1. p. 89-96. In Seed Protein Improvement in Cereals and Grain Legumes. Proc. Int. Symp., Neuherberg, Germany, 1978. Int. Atomic Energy Agency, Vienna.

Kasha, K.J., and G.W.R. Walker. 1960. Several recent barley mutants and their linkages. Can. J. Genet. Cytol. 2:397-415.

Linde-Laursen, I. 1997. Recommendations for the designation of the barley chromosomes and their arms. BGN 26:1-3.

Luna Villafaña, A. 1995. Mapping marker genes in chromosome 2 of barley. M.S. Thesis. North Dakota State Univ., Fargo.

Nilan, R.A. 1964. The cytology and genetics of barley, 1951-1962. Monogr. Suppl. 3, Res. Stud. Vol. 32, No. 1. Washington State Univ. Press, Pullman.

Ramage, R.T., and C.L. Crandall. 1981. Shrunken endosperm mutant seg8. BGN 11:34.

Ramage, R.T., and J.F. Scheuring. 1976. Shrunken endosperm mutants seg6 and seg7. BGN 6:59-60.

Shahla, A. and T. Tsuchiya. 1984. Additional information on the association of zb_c2 for zoned leaf with chromosome 5. BGN 14:10-11.

Shim, J.W., A. Shahla, and T. Tsuchiya. 1984. Additional information on the primary trisomic analysis of sf (female sterile or gigas) in barley. BGN 14:42-43.

Søgaard, B. 1973. Continued linkage studies on eceriferum mutants in barley. BGN 3:57-61.

Takahashi, R., I. Hayashi, T. Inouye, I. Moriya, and C. Hirao. 1973. Studies on resistance to yellow mosaic disease in barley. I. Tests for varietal reaction and genetic analysis of resistance to the disease. Ber. Ohara Inst. landw. Biol., Okayama Univ. 16:1-17.

Takeda, K., and C.L. Zhang. 1995a. Inheritance and geographical distribution of a phenol reaction-less mutant of barley. BGN 24:32-34.

Takeda, K., and C.L. Zhang. 1995b. Inheritance of a fluorescent reaction mutant of barley. BGN 24:38-40.

Tazhin, O.T. 1982. Value of crossing-over between linked genes mo5 and n in barley BGN 12:18-21.

Tuleen, N.A., and M.E. McDonald. 1971. Location of genes Pa and Pa5. Barley Newsl. 15:106-107.

Walker, G.W.R., J. Dietrich, R. Miller, and K.J. Kasha. 1963. Recent barley mutants and their linkages II. Genetic data for further mutants. Can. J. Genet. Cytol. 5:200-219.

Figure 1. Estimated positions for morphological marker loci in chromosome 1H of barley.

Figure 2. Estimated positions for morphological marker loci in chromosome 2H of barley.

Figure 3. Estimated positions for morphological marker loci in chromosome 3H of barley.

Figure 4. Estimated positions for morphological marker loci in chromosome 4H of barley.

Figure 5. Estimated positions for morphological marker loci in chromosome 5H of barley.

Figure 6. Estimated positions for morphological marker loci in chromosome 6H of barley.

Figure 7. Estimated positions for morphological marker loci in chromosome 7H of barley.

Table 1. Other morphological markers not mapped to a relative position in the barley genome, but tentatively associated with a specific chromosome.

Recom. symbol	BGS no.	Chr. loc.*	Locus name or phenotype	GSHO no.**	Reference
CHROMOSOME 1H
cur5 (cu5)	231	1HS	Curly 5	1710	Franckowiak, 1995
des6	215	1H	Desynapsis 6	597	Hernandez-Soriano, 1973
dsp.ab		1H?	Dense spike ab	1715	Franckowiak, 1995
dsp.ao		1HS	Dense spike ao	1723	Franckowiak, 1995
glo-e	230	1HL	Globosum-e	1755	Häuser and Fischbeck, 1980
lax.ae		1HL	Laxatum ae	1779	Franckowiak, 1995
msg20	375	1H	Male sterile genetic 20	2372	Hockett and Eslick, 1971
mtt1 (mt)	521	1HS	Mottled leaf 1	622	Kasha and Walker, 1960
pyr.aw		1HS	Pyramidatum aw	2436	Franckowiak, 1995
Sil	228	1HS	Subcrown internode length	1604	Franckowiak, 1995
yhd2 (yh2)		1H	Yellow head 2	757	Eslick (unpubl.)
zeb3 (zb3)	223	1HL	Zebra stripe 3	1451	Shahla and Tsuchiya, 1984
CHROMOSOME 2H
acr1		2HL	Accordion rachis 1	1617	Franckowiak (unpubl.)
cer-v	414	2HS	Eceriferum-v +/- ++ ++	443	Søgaard, 1973
cer-yb	445	2HL	Eceriferum-yb ++ ++ -	1535	Franckowiak, 1995
cur4 (glo-d)	460	2HL	Curly 4	1708	Häuser and Fischbeck, 1979
ert-j	90	2H	Erectoides-j	484	Franckowiak, 1995
ert-q	91	2H	Erectoides-q	1562	Franckowiak, 1995
ert-u	92	2H	Erectoides-u	496	Franckowiak, 1995
ert-zd	93	2H	Erectoides-zd	504	Franckowiak, 1995
des7	64	2H	Desynapsis 7	598	Hernandez-Soriano, 1973
fch14 (f14)	87	2HL	Chlorina seedling 14	1739	Franckowiak, 1995
gig1	463	2H?	Gigas 1	1650	Shim et al., 1984
glo-c	72	2H	Globosum-c	1329	Häuser and Fischbeck, 1976
lks.m		2HL	Short awn m	1782	Luna Villafaña, 1995
msg35	498	2HL	Male sterile genetic 35	2387	Franckowiak, 1995
msg48	520	2H	Male sterile genetic 48	2401	Franckowiak, 1991
mss1	84	2H	Midseason stripe 1	1404	Walker et al., 1963
Phr		2HL	Phenol reaction		Takeda and Zhang, 1995a
Rph1 (Pa)	70	2H	Reaction to Puccinia hordei 1	1313	Tuleen and
McDonald, 1971
Rph15		2HL	Reaction to Puccinia hordei 15	1586	Chicaiza, 1996
sdw.aw		2HL	Semidwarf aw	2446	Franckowiak, 1995
sld.d		2HS	Slender dwarf d	2479	Franckowiak, 1995
yst4	85	2HL	Yellow streak 4	2502	Faue et al., 1989
CHROMOSOME 3H
adp		3H?	Awned palea	1618	Franckowiak (unpubl.)
brh.w		3HS	Brachytic w	1687	Franckowiak, 1995
brh.ad		3HS	Brachytic ad	1671	Franckowiak, 1995
cer-ya	444	3HS	Eceriferum-ya ++ ++ -	1534	Franckowiak, 1995
cer-yd	447	3HS	Eceriferum-yd - ++ ++	1537	Franckowiak, 1995
cer-yh	451	3HS	Eceriferum-yh - ++ ++	1541	Franckowiak, 1995
des2	119	3H	Desynapsis 2	593	Hernandez-Soriano, 1973
dsp.ar		3HS	Dense spike ar	1726	Franckowiak, 1995
gra-a	131	3HL	Granum-a	1757	Fischbeck and Häuser, 1976
msg42	505	3H	Male sterile genetic 42	2395	Franckowiak, 1991
nec4	138	3H	Necrotic leaf spot 4		Fischbeck and Häuser, 1976
nec5	139	3H	Necrotic leaf spot 5		Fischbeck and Häuser, 1976
Pyr.i		3HS	Pyramidatum i	1582	Franckowiak, 1995
seg3	379	3H	Shrunken endosperm 3	752	Jarvi and Eslick, 1975
seg6	396	3HL	Shrunken endosperm 6	2467	Ramage and Scheuring, 1976
sex.j		3HL	Shrunken endosperm xenia j	2471	Franckowiak, 1995
sld.h		3HS	Slender dwarf h	2483	Franckowiak, 1995
yst3	462	3HS	Yellow streak 3	48	Franckowiak, 1995
CHROMOSOME 4H
brh.m		4H	Brachytic m	1678	Franckowiak, 1995
flo-a	182		Extra floret-a	1741	Lundqvist (pers. com.)
frp		4HL	Fluorescent reaction-pink		Takeda and Zhang, 1995b
msg25	166	4HL	Male sterile genetic 25	744	Hockett and Eslick, 1971
Rym1 (Ym)	167	4HL	Reaction to BaYMV 1		Takahashi et al., 1973
sld.e		4H	Slender dwarf e	2480	Franckowiak, 1995
sld.f		4H	Slender dwarf f	2481	Franckowiak, 1995
Zeo.h		4H	Zeocriton h	1611	Franckowiak, 1995
CHROMOSOME 5H
ari-h	329	5HL	Breviaristatum-h	1656	Franckowiak, 1995
blf2 (bb2)	337	5HL	Broad leaf 2	1667	Franckowiak, 1995
brh.j		5HL	Brachytic j	1675	Franckowiak, 1995
brh.s		5HL	Brachytic s	1684	Franckowiak, 1995
cer-za	419	5HL	Eceriferum-za ++ ++ -	1521	Franckowiak, 1995
cer-ye	448	5HL	Eceriferum-ye ++ ++ -	1538	Franckowiak, 1995
crl (cl)	325		Curly lateral	1211	Walker et al., 1963
cst (cs)		5H	Corn stalk		Nilan, 1964
flo-b	327	5HL	Extra floret-b	1742	Franckowiak, 1995
glo-f	342	5HL	Globosum-f		Häuser et al., 1988
int-b	320	5HL	Intermedium spike-b	1764	Franckowiak, 1995
Lfb		5HL	Leafy bract	1577	Walker et al., 1963
mnd4		5HL	Many-noded dwarf 4	1798	Franckowiak, 1995
msg29	466	5HL	Male sterile genetic 29	2381	Franckowiak, 1995
msg49	335	5HL	Male sterile genetic 49	2402	Franckowiak, 1991
pyr.af		5HL	Pyramidatum af	1718	Franckowiak, 1995
raw6 (r6)	334	5HL	Smooth awn 6	2437	Franckowiak, 1995
Rph2 (Pa2)	88	5HS	Reaction to Puccinia hordei 2	1593	Borovkova et
al., 1997
Rph5 (Pa5)	122	5H	Reaction to Puccinia hordei 5	1597	Borovkova et al.,
1997
sdw.ah		5HL	Semidwarf ah	2451	Franckowiak, 1995
sex.i		5HL	Shrunken endosperm xenia i	2470	Franckowiak, 1995
Zeo.f		5HL	Zeocriton f	1610	Franckowiak, 1995
CHROMOSOME 6H
ant21		6H	Anthocyanin-less 21	1634	Falk, 1994
lys6	269	6H	High lysine 6	1786	Jensen, 1979
msg28	465	6H	Male sterile genetic 28	2380	Franckowiak, 1991
msg34	471	6H	Male sterile genetic 34	2386	Franckowiak, 1991
msg39	502	6H	Male sterile genetic 39	2391	Franckowiak, 1991
msg40	503	6H	Male sterile genetic 40	2393	Franckowiak, 1991
CHROMOSOME 7H
ant18		7HL	Anthocyanin-less 18	1630	Jende-Strid, 1993
ant22		7HL	Anthocyanin-less 22	1635	Franckowiak, 1995
cer-k	405	7HS	Eceriferum-k + ++ ++	432	Franckowiak, 1995
cer-z	418	7HS	Eceriferum-z - - ++	447	Franckowiak, 1995
cer-yg	450	7HS	Eceriferum-yg - - -	1540	Franckowiak, 1995
des1	12	7H	Desynapsis 1	592	Hernandez-Soriano, 1973
des4	13	7H	Desynapsis 4	595	Hernandez-Soriano, 1973
des5	14	7H	Desynapsis 5	596	Hernandez-Soriano, 1973
mov5 (mo5)		7HS	Multi-ovary 5		Tazhin, 1982
msg22	383	7H	Male sterile genetic 22	741	Hockett and Eslick, 1971
msg26	395	7HS	Male sterile genetic 26	745	Franckowiak, 1995
msg30	467	7HL	Male sterile genetic 30	2382	Franckowiak, 1991
msg32	469	7H	Male sterile genetic 32	2384	Hockett and Eslick, 1971
pbg		7H	Pubescence of glume		Hor, 1924
Rsg1	22	7H	Reaction to Schizaphis gram 1	1317	Gardenshire et al.,
1973
seg5	381	7HS	Shrunken endosperm 5	754	Jarvi and Eslick, 1975
seg8	455	7H	Shrunken endosperm 8	2469	Ramage and Crandall, 1981
smn1 (smn)		7HS	Seminudoides 1		Franckowiak, 1995
smn2		7HS	Seminudoides 2		Gymer, 1977
snb (sb)	26	7HS	Subnodal bract	1217	Walker et al., 1963

* Chromosome numbers and arm designations are based on the Triticeae system. Utilization of this system for barley chromosomes was recommended at the business meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan, Canada, on August 5, 1996. The Burnham and Hagberg (1956) designations of barley chromosomes were 1 2 3 4 5 6 and 7 while new designations based on the Triticeae system are 7H 2H 3H 4H 1H 6H and 5H, respectively.

** The seed stock associated with each BGS number is held as a GSHO stock number in the Barley Genetics Stock Collection at the USDA-ARS National Small Grains Germplasm Research Facility, Aberdeen, Idaho, USA.

Table 1. A listing of Barley Genetic Stock (BGS) descriptions in Barley Genetics Newsletter volume 26, recommended locus symbols and names, and stock location information.

*Recommended locus symbols are based on utilization of a three letter code for barley genes as approved at the business meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan, Canada, on August 5, 1996.

+ Chromosome numbers and arm designations are based on a resolution passed at the business meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan, Canada, on August 5, 1996. The Burnham and Hagberg (1956) designations of barley chromosomes were 1 2 3 4 5 6 and 7 while new designations based on the Triticeae system are 7H 2H 3H 4H 1H 6H and 5H, respectively.

** The seed stock associated with each BGS number is held as a GSHO stock number in the Barley Genetics Stock Collection at the USDA-ARS National Small Grains Germplasm Research Facility, Aberdeen, Idaho, USA.

Table 2. An alphabetic listing of new and revised Barley Genetic Stock (BGS) descriptions for loci in barley (Hordeum vulgare), including information on chromosomal locations, recommended locus names, and original cultivars.

* Chromosome numbers and arm designations are based on the Triticeae system. Utilization of this system for barley chromosomes was recommended at the business meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan, Canada, on August 5, 1996. The Burnham and Hagberg (1956) designations of barley chromosomes were 1 2 3 4 5 6 and 7 while new designations based on the Triticeae system are 7H 2H 3H 4H 1H 6H and 5H, respectively.