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So far one of the translocation lines, 'Weique Zuchter', has been found deviating from all,other by the complete or almost complete lack of the telomeric band of the long arm of chromosome 1B-1R, which is accompanied by an adequate reduction of length (FRIEBE 1976; MUNZER 1977). At present neither the appropriate meiotic pairing data (Table 2, data taken from ZELLER and SASTROSUMARJO 1972; BLUTHNER and METTIN 1973; BLUTHNER and METTIN 1977) nor the Giemsa analyses provide any evidence on the origin of this particular chromosome arm. Actually the data presented in Table 2 only demonstrate the capability of the deviating interchange chromosome to pair with 1RS as well as 1BL. Whether the long arm of chromosome 1B-1R of 'Weique Zuchter' is capable to pair with 1RL has not yet been investigated. And so there is at the moment no reason to speculate about the possible loss of telomeric heterochromatin effecting a bilateral pairing ability similar to the behaviour of the presumed 2D(2R) substitution in 'Rosner' as suggested by KALTSIKES et al. (1977).

On the other hand it seems very likely that the long arm of the common type of the 1B-1R interchange has derived from chromosome 1B completely or almost completely. In addition to previous results SCHLEGEL (unpubl.) by using the Giemsa banding technique was able to demonstrate pairing of chromosome 1B-1R with rye chromosome 1R being restricted to the short arms, i.e. closed bivalents were never observed.

Because of the difficulty to locate the possible breakpoint of the interchange by genetic means, we tried to get some additional evidence by the presumed mode of origin of the 1B-1R interchange. Tracing back the origin of most of the translocation lines listed in Table 1 there is no doubt about the parentage of 1R(1B) whole chromosome substitutions. So it is very likely that the substitution was prior to the translocation. Because substitutions were often crossed to karyotypically normal wheats it seems reasonable to suppose a connection between the double-monosomic state for 1B and 1R in the hybrids on the one hand and the origin of the interchange on the other. Considering 1B, 1R double-monosomic wheat hybrids there is a certain probability for centromeric misdivision of the univalents followed by fusion of the telocentrics as shown by MORRISON (1954), SEARS (1972) and SHEPHERD (1973), though casual homoeologous pairing and recombination can no longer ruled out. For that reason some of the prerequisites for either centromeric fusion or recombination have been analyzed.

Taking the amount of telosomic offspring in appropriate wheat lines and hybrids as a measure for centromeric misdivision our previous data show that 1B, 1R double-monosomics yield up to 21% telocentrics. A comparative analysis of the frequency of telosomics in the progenies of wheat hybrids being monosomic for either 1B or 1R indicated a rather high instability of rye chromosome 1R, at least in a wheat background. This suggestion was confirmed in Giemsa-stained metaphase I preparations of 1B, 1R double-monosomics. As shown in Table 3 misdivisions of rye chromosome 1R exceeded by far those of wheat chromosome 1B. Considering the low number of PMC's scored it seems worthwhile to emphasize the occurrence of simultaneous misdivisions of both chromosomes under question, which is the prerequisite for centromeric fusion.

The alternative mode of origin of the 1B-1R interchange by means of homoeologous recombination must be taken into consideration, since several authors, by using the Giemsa banding technique, have proven the occurrence of wheat-rye pairing even in the presence of wheat chromosome 5B (METTIN et al. 1976; DHALIWAL et al. 1977; POHLER and KISTNER 1977). Our additional data on a large number of bivalents and multivalents in euhaploid wheat-rye hybrids suggest a figure of about 1.5% Wheat-rye pairings of all associations (Table 4). This is, however, the average for the total of the rye chromosomes. Whether or not rye chromosome 1R, having a large amount of telomeric heterochromatin, participates proportionally in the wheat-rye associations must be left open at the moment.

Summarizing all the data available there is a lot of reason to suppose that the 1B-1R interchange derived from the whole chromosome 1R (1B) substitution by means of centromeric fusion. Therefore the breakpoint should be located at or very close to the centromere. This seems to be true, at least, for the common type of the interchange.



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