The Deletion Stocks of Common Wheat
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The Deletion Stocks of Common Wheat T. R. Endo and B. S. Gill Chromosomal breaks occurred in the progeny of a common wheat (Tritlcum aes- tlvum L. em Thell; 2n = 6x = 42, genome formula AABBDD) cultivar Chinese Spring with a monosomic addition of an alien chromosome from Aegllops cyllndrlca Host (2n = 4x = 28, CCDD) or A. trlunclalls L. (2n = 4x = 28, UUCC) or a chromosomal segment from A. spettoldes Tausch (2n = 2x = 14, SS). We identified 436 deletions by C-banding. The deletion chromosomes were transmitted stably to the offspring. We selected deletion homozygotes in the progeny of the deletion heterozygotes and established homozygous lines for about 80% of the deletions. We failed to establish homozygous lines for most of the deletions In the short arm of chromo- some 2A and for all deletions in the short arm of chromosome 4B, because plants homozygous for these deletions were sterile. We could not obtain any homozygotes for larger deletions In the long arms of chromosomes 4A, 5A, 5B, and 5D. The deletion stocks showed variations in morphological, physiological, and biochemi- cal traits, depending on the size of their chromosomal deficiency, and are powerful tools for physical mapping of wheat chromosomes. The aneuploid stocks developed in a com- and a chromosome fragment from A. spel- mon wheat cultivar Chinese Spring are a toides were found recently to induce chro- powerful tool for genetic and breeding mosome mutations in Chinese Spring in studies of wheat (Sears 1954, 1966; Sears the same manner as the A. cylindrica chro- and Sears 1978). These stocks are im- mosome (Endo, unpublished data). A pre- mensely useful for localization of genes on liminary attempt was made to isolate de- chromosomes and chromosomes arms letions in the progeny of wheats with mono- (Mclntosh 1988). Endo (1988) reported a somes of these chromosomes. In this arti- unique genetic system for the systematic cle, we report the details of the produc- production of even more powerful novel tion and the general features of deletion aneuploid stocks, namely, deletion stocks stocks of Chinese Spring. with various sized terminal deletions in in- dividual chromosome arms, useful for sub- Materials and Methods arm localization of genes. When a certain chromosome from Aegilops cylindrica is Wheat Stocks present in Chinese Spring in the monoso- We used two alien monosomic addition mic condition, chromosomal breaks occur lines and one alien translocation line of From the Laboratory of Genetics, Faculty of Agricul- in the gametes that lack the A. cylindrica Chinese Spring to induce deletions. One ture, Kyoto University, Kyoto 631, Japan (Endo), and the Department of Plant Pathology, Kansas State Uni- chromosome and generate various chro- alien addition line had an A. cylindrica versity, Manhattan, KS 66506-5502 (GUI). This work was mosome aberrations, including deletions. chromosome (Endo 1988). This alien chro- supported In part by a grant-ln-ald for scientific re- The broken chromosome ends, if not mosome is homoeologous to the group 2 search (no. 05660008) from the Ministry of Education, Science, and Culture (Japan); by a grant-in-ald for spe- fused to other broken ends, are stabilized chromosomes (Gill KS, unpublished data). cial scientific research on agriculture, forestry, and by the rapid gain of telomere structure The other alien addition had an A. triunci- fisheries from the Ministry of Agriculture, Forestry and Fisheries (Japan); by the Japan-U.S. Cooperative Sci- (Werner et al. 1992b). Such deletions in alis chromosome, which has a satellite ence Program of the Japan Society for the Promotion plants without the A. cylindrica chromo- and is different from the A. triuncialis chro- of Science and the National Science Foundation USA; some are transmitted regularly to the off- mosome reported previously by Endo and by USDA-NR1 Plant Genome Awards. This Is con- tribution no. 537 from the Laboratory of Genetics, Fac- spring. (1978). The alien translocation line had a ulty of Agriculture, Kyoto University, and contribution Isolation of deletion lines was conduct- modified wheat chromosome 2B with a no. 95-449-J from the Kansas Agricultural Experimental Station, Manhattan, Kansas. Address reprint requests ed in the progeny of the A. cylindrica ad- small chromosomal segment from A. spel- from the United States to Dr. B. S. Gill and from other dition line by eliminating aberrations oth- toides translocated to the end of the long countries to Dr. T. R. Endo. er than simple terminal deficiencies. Also, arm (described as T2BS-2BL-2SL hereaf- Journal of Heredity 1996:87:295-307; 0022-1503/96/J5.00 an alien chromosome from A. triuncialis L. ter). This translocation chromosome car- 295 21-+VA mulae were used to calculate the FL val- ues of the deletions in the short and long arms in the satellite chromosomes IB and / 6B, except the length of the satellite was it not included. For estimating the FL value 21' 21'+1'A of deletions in the satellite regions of IB and 6B, the formula gamete with chromosomal normal aberrations gamete deletion satellite FL = x2V S arm satellite Figure 1. C-bandlng of (a) A. cyiindnca, (b) A triun- normal homologue cialis, and (c) T2BS-2BI^2SL chromosomes. Deletion _ S arm x drtelosomlc heterozygote or was used. Because of their indirect mea- ries the Gclb (gametocidal) gene (Tsuji- nullisomic- surements, the FL values do not necessar- moto and Tsunewaki 1988). Figure 1 shows tetrasomic ily indicate the precise breakpoints of the self deletions and, therefore, were used to or- the C-banding patterns of the Aegilops and r translocation chromosomes. We isolated der only those deletions whose break- most of the deletions from the A. cylindri- Deletion Deletion points were neither in nor close to ca addition line. A small number of dele- homozygote hem Izy goto C-bands. We analyzed several of the dele- tions were isolated from the A. cylindrica tions by in situ hybridization using the Figure 2. A scheme showing the production of the addition line with the cytoplasm of A. kot- deletion stocks In common wheat. "A" stands for the 18S-26S rDNA probe, as described by Mu- schyi (Mukai and Endo 1992) and from the Aegilops chromosome or chromosome segment caus- kai et al. (1991). A. triuncialis addition and T2BS-2BL-2SL ing chromosomal aberrations. lines. We used the ditelosomic (Sears and Results Sears 1978) and nullisomic-tetrasomic length (FL) of the breakpoints from the (Sears 1966) lines of Chinese Spring to iso- centromere for the short-arm deletions ac- We studied a total of 475 plants of the first late some of the deletion chromosomes in progeny, backcrossed or selfed, of the cording to the formula the hemizygous condition. A.cylindrica monosomlc addition line. deletions arm About half of the progeny had one or more FL = deletion chromosome Crosses and Cytology L arm chromosomal structural changes and most of the plants with the chromosomal We first backcrossed the alien addition Sarm and translocation lines to Chinese Spring, normal homologue aberrations did not carry the A. cylindrica analyzed the chromosome constitutions Larm chromosome, as reported previously of the progeny by C-banding (the modified The FL of long-arm deletions was calculat- (Endo 1988). Of the 52 backcrossed prog- C-banding; Gill et al. 1991), and grew ed by the formula eny of the A. triuncialis addition line, 44 plants that had a deletion or deletions and plants did not have the alien chromosome, Sarm and 32 out of the 44 plants had some chro- no alien chromosome (Figure 2). We either FL = normal homologue self-pollinated or backcrossed those plants Larm mosomal aberrations. Of the 31 back- crossed progeny of the T2BS-2BL-2SL line, once to Chinese Spring, followed by self- Sarm ing. We cytologically screened the selfed deletion chromosome. 12 plants that did not bear the transloca- deletion L arm progeny for homozygous plants with de- tion chromosome had some chromosomal letion chromosomes and the least degree The arm ratios were used to compensate aberrations. For the production of dele- of aberrations in the other chromosomes, for the difference in contraction of chro- tion stocks, we selected 156, 12, and 4 that is, translocations and aneuploidy. mosomes between cells. The arm ratios of plants with one or more deletions and no When two or more deletions occurred in the normal wheat chromosomes were Aegilops chromosome or alien transloca- a single plant in the homozygous condi- taken from Gill et al. (1991). The same for- tion chromosome from the progeny of the tion, we made no further effort to separate them. Table 1. Distribution of deletion breakpoints in different genomes, chromosome arms, and For some deletions, when homozygotes homoeologous groups In Chinese Spring wheat were not obtained after the screening of the selfed progeny, we crossed the appro- No. of deletion breakpoints priate ditelosomic or nullisomic-tetraso- Homoeo- Genomes: A B D logous mic lines to the deletion heterozygotes to group Arms: S L S L S L Total obtain the deletion hemizygotes in the F, progeny. The F progeny then were 1 5 g 22 18 5 8 64 2 2 9 13 11 6 12 57 screened for deletion homozygotes. 3 4 #: 1Q 12 9 3 46 We photographed 5-10 chromosomes 4 4 13 9 14 5 15 60 5 U 23 9 ts 4 12 77 for each of the deletions and measured 6 5 8 11 15 7 11 57 the lengths of the short (excluding the sat- 7 13 25 6 16 6 9 75 ellites for chromosomes IB and 6B) and Subtotal 51 89 80 104 42 70 Total 140 184 112 436 long arms. We calculated the fraction 296 The Journal of Heredity 199687(4) )llilll"l> ' 1A S-3 S-4 S-l S-2 S-5 L-3 L-6 L-4 L-5 L-2 L-l IB S-4 S-ll S-18 S-19 S-23 S-22 S-5 S-6 S-8 S-13 S-16 S-2 S-9 S-7 S-3 S-20 S-10 S-21 S-15 S-l S-12 S-14 L-4 L-10 L-13 L-3 L-5 L-12 L-15 L-17 L-8 L-9 L-2 L-14 L-l L-7 L-16 L-18 L-6 L-ll 1111(11limit ID S-5 S-4 S-l S-2 S-3 I-9 -? I -R -R -7 i _1 i ..A Figure 3.