A comparative study of the cytological and reproductive behaviour of wheat and triticale subjected to two temperature regimes

Summary A comparison was made of the meiotic stability and reproductive behaviour ofTriticale hexaploide cv. Rosner andT. aestivum cv. Pitic in response to two temperature regimes (15° and 30°C) imposed continuously from premeiosis to maturity. Mean frequencies of MI open bivalents, univalents, and lagging chromosomes at AI were significantly higher in Rosner than in Pitic. Such irregularity would appear to arise from the asynchronous disjunction of chromosomes between Metaphase I and Anaphase I, resulting in meiotic disturbances and a high frequency of abnormal microspores. Despite such abnormalities, floret fertility was equivalent to that observed in Pitic, and the proportion of aneuploids recovered in the progeny, although greater than expected in wheat, was much lower than would be predicted. It is suggested that there is a strong certation effect in the transmission of aneuploid microspores and that the irregularities observed during microsporogenesis do not apply directly to megasporogenesis.No appreciable effect of temperature on meiotic chromosome behaviour during microsporogenesis was observed in either wheat or Triticale. In Triticale, high temperatures (30°C) substantially reduced spikelet fertility, and in contrast, low temperature (15°C) increased the frequency of aneuploids.Contribution No 232, Department of Plant Science, University of Manitoba, Winnipeg 19, Manitoba, Canada     

Patterns of meiosis in ABDR amphihaploids depend on the specific type of univalent chromosome division

Meiosis in wide F₁ hybrids in the tribe Triticeae is complex and irregular. Despite extensive research, the meiotic pathways in such hybrids remain largely unknown. In this study, the effect of the genotypes of disomic wheat–rye substitution lines on the regulation of meiosis in ABDR amphihaploids was analysed by comparing microsporogenesis and seed set under self pollination in hybrids of normal bread wheat (Triticum aestivum L.) and wheats with single chromosome substitutions 1R(1A), 1Rv(1A), 2R(2D)₁, 2R(2D)₂, 2R(2D)₃, 5R(5D), 5R(5A), 6R(6A) with rye (Secale cereale L.). Three types of meiotic pathways were identified in the hybrids: reductional, equational, and equational+reductional. During the reductional type division, chromosomes randomly moved toward the poles at Anaphase I (AI) and separated sister chromatids at AII. Meiosis ended with tetrad formation, and the resulting plants were sterile ((2R(2D)₁ × R, 2R(2D)₂ × R). In the equational type division, the chromosomes aligned along the equator at Metaphase I and at AI separated into chromatids that moved toward the poles. This meiotic pathway led to dyad formation and partial fertility (especially in hybrids of 1Rv(1A) and 6R(6A)). In the equational+reductional pathway, some chromosomes divided in an equational manner while others were segregated reductionally within the same microsporocyte and this led to formation of tetrads with groups of uncoiled chromosomes. Comparison of the frequencies of each of the three meiotic pathways in different hybrid combinations suggests differences in contributions of individual chromosomes to the control of the entire meiotic process.     

Relevance of sexual polyploidization for crop improvement – A review

Colchicine induced polyploids have not directly contributed for crop improvement in the past. On the other hand, the so-called natural polyploids, derived from the functioning of numerically unreduced(2n) gametes have been shown to be more relevant for crop improvement in many cases. Different types of cytological abnormalities during meiosis can give rise to 2n gametes and the genetic composition of these gametes is variable. Depending on the type meiotic abnormalities, various types of 2ngametes, such as first division restitution(FDR), second division restitution (SDR),indeterminate meiotic restitution (IMR) and post meiotic restitution (PMR) gametes,among others, have been described in recent years. For the improvement of autopolyploids such as potato, alfalfa,Vaccinium spp., and some of the fodder grasses, FDR gametes have been proved to be highly useful. However, the use of 2n gametes for the improvement of allopolyploid crops has received much less attention so far. Some of the investigations on allopolyploids, derived from Festuca-Lolium, Alstroemeria and Lilium species hybrids, have revealed that 2ngametes can be most useful for the introgression of alien genes and chromosomes into cultivars. An important feature of using sexual polyploidization in the case of allopolyploids is that introgression can be achieved through recombination due to genetic crossing-over between alien chromosomes as well as addition of alien chromosomes, which is extremely difficult or impossible to achieve in the case of colchicine induced allopolyploids. Because of the recent developments in the field of plant molecular biology, methods have become available for the analysis of 2ngametes and sexual polyploid progenies more accurately and to develop systematic breeding approaches. The methods include DNA in situ hybridization (GISH and FISH)and molecular mapping (AFLP, RFLP, RAPDs).In addition to providing basic information on the genetic and genome composition of the polyploid progenies, these methods can be potentially useful for a more efficient creation of desirable breeding material and cultivars. This revised version was published online in July 2006 with corrections to the Cover Date.     

Formation of unreduced gametes is impeded by homologous chromosome pairing in tetraploid <Emphasis Type="Italic">Triticum turgidum</Emphasis> × <Emphasis Type="Italic">Aegilops tauschii</Emphasis> hybrids

It is believed that unreduced gametes with somatic chromosome numbers play a predominant role in natural polyploidization. Allohexaploid bread wheat originated from spontaneous hybridization of Triticum turgidum L. with Aegilops tauschii Coss. Unreduced gametes originating via meiotic restitution, including first-division restitution (FDR) and single-division meiosis (SDM), are well documented in triploid F₁ hybrids of T. turgidum with diploid Ae. tauschii (genomic constitution ABD, usually with 21 univalents in meiotic metaphase I). In this study, two T. turgidum lines known to carry genes for meiotic restitution were crossed to tetraploid Ae. tauschii. The resulting F₁ hybrids (genomes ABDD), had seven pairs of homologous chromosomes and regularly formed 14 univalents and seven bivalents at metaphase I. Neither FDR nor SDM were observed. The distribution of chromosome numbers among progeny obtained by self pollination and a backcross to T. turgidum showed the absence of unreduced gametes. These results suggest that high homologous pairing interfered with meiotic restitution and the formation of unreduced gametes. This may be related to asynchronous movement during meiosis between paired and unpaired chromosomes or to uneven distribution of chromosomes in anaphases, resulting in nonviable gametes.     

First-division restitution in hybrids of Langdon durum disomic substitution lines with rye and Aegilops squarrosa

Durum wheat ‘Langdon’(LDN) caused a high frequency of first‐division restitution (FDR) and partial fertility in hybrids with rye, Secale cereale L., and Aegilops squarrosa L. In order to determine the genetic control of FDR, a complete set of 14 Langdon durum D‐genome disomic substitution lines (LDNDS) was crossed with ‘Gazelle’ rye and one accession (RL5286) of A. squarrosa. The microsporogenesis and fertility of the hybrids were studied. The results showed that most of the hybrids expressed a high frequency of FDR and partial fertility. However, the hybrids of 2D(2A), 4D(4A), 5D(5B) and 6D(6B) crossed with both rye and A. squarrosa, as well as 1D(1A) with A. squarrosa, had either little or no FDR and were completely sterile. These hybrids had different types of first meiotic divisions compared with LDN control hybrids. The hybrids with 2D(2A), 4D(4A) and 6D(6B) had a high frequency of random segregation of chromosomes at the first division. The hybrids with 5D(5B), as expected, showed high homoeologous pairing. The hybrid of 1D(1A) with A. squarrosa had a high frequency of equational division at first division. These results suggest that the reduced or absent FDR in such hybrids might be related to the substitution of chromosomes with an FDR gene and poor compensation ability of the D‐genome chromosomes for their homoeologous A‐ or B‐ genome chromosomes. Cytological analysis suggested that chromosome 4A in LDN most likely carries a gene for high frequency of FDR in hybrids. In addition, some monads were observed at the end of meiosis in the hybrids of 3D(3A) and 6D(6A) crossed with rye. They were formed from FDR cells that failed to divide at second division, suggesting that the LDN 3A and 6A chromosomes might carry genes for normal second division of FDR cells in the rye crosses.     

Cytogenetic analysis of two interspecific Citrus allotetraploid somatic hybrids and their diploid fusion parents

Microsporogenesis of two interspecific allotetraploid somatic hybrids, i.e.‘Hamlin’ sweet orange +‘Rough Lemon’ (HR) and ‘Key’ lime +‘Valencia’ sweet orange (KV), was examined and compared with their corresponding diploid fusion parents. Meiotic analysis revealed that only 13.2 and 18.0% of the meiocytes exhibited normal tetrad formation in HR and KV somatic hybrid plants, respectively, and irregular chromosome behaviour with univalent or multivalent pairing occurred. Moreover, meiotic abnormalities such as chromosome bridges, chromosomes orientated away from the equatorial plate, especially lagging chromosomes, which resulted in different sizes of pollen grain, were frequently observed in both somatic hybrids. As expected, the percentage of chromosome abnormalities was much lower in their corresponding diploid fusion parents. HR and KV had 15.4 and 7.2% pollen germinability, respectively, intermediate between their corresponding fusion parents. The meiotic behaviour of these somatic hybrids provided valuable information for their practical utilization in a citrus breeding programme.     

Intergeneric hybrid between Brassica napus and Diplotaxis harra through ovary culture and the cytogenetic analysis of their progenies

Brassica napus (2n = 38) and Diplotaxis harra (2n = 26) were used to investigate gene transfer from D. harra to B. napus. Intergeneric F₁ hybrids (dihaploid 2n = 32 chromosomes) were obtained through ovary culture. The chromosome associations in the first meiotic division was (0–2)_III + (2–10)_II + (12–28)_I. Many seeds were harvested in the F₁ hybrid after backcrossing with B. napus, and from open pollination of the F₁ hybrid. Somatic chromosome numbers of BC₁ and hybrid plants varied from 2n = 26 to 52. In the first meiotic division, high frequencies of bivalent association and relatively low pollen fertility were observed. BC₂ plants generated from the BC₁ plants with 2n = 38 chromosomes, 69.6% showed 2n = 38 chromosomes. Many aneuploids with addition and deletion of chromosomes were also obtained. A bridge plant between B. napus and D. harra with 2n = 32 chromosomes should be valuable material for the breeding of brassica crops.     

Karyotype and meiotic analyses of six species in the subtribe Chrysantheminae

The karyotypes and meiotic behaviour of six Chrysantheminae species are investigated. Pyrethrum coccineum (Willd.) Worosch. is a tetraploid, while the other five species (Argyranthemum frutescens (L.) Sch.-Bip., Opisthopappus taihangensis (Ling) shih, Crossostephium chinense (L.) Makino, Tanacetum vulgare L. and P. parthenium (L.) Sm.) are all diploids. Their karyotypes consist mainly of median centromere and submedian centromere chromosomes, although C. chinense also has two terminal centromere chromosomes, and P. parthenium two subterminal centromere chromosomes. No satellited chromosomes are observed in any of the six species. All the species have a 2A type karyotype, except for P. coccineum which is 3A. The meiotic behaviour in the pollen mother cells (PMCs) of all six species is essentially normal, with metaphase I chromosome pairing configurations in the diploid species predominantly showing 9II, while P. coccineum shows 18II. A low frequency of quadrivalents (and trivalents) and chromosome bridges/lagging chromosomes is observed.     

Meiotic instability of common wheat strains derived from Triticum timopheevi Zhuk. crosses

Summary Meiotic instability was studied in four strains of common wheat derived from crosses involving a Triticum timopheevi derivative, C.I. 13093, and the common wheat varieties Cheyenne and Minturki. A wide range in the percentage of normal cells at six stages of meiosis, MI, AI, dyad, MII, AII, and quartet, was found. The date of sporocyte sampling influenced meiotic irregularities at six stages of meiosis. Florets from the same spikes differed in the percentage of normal cells at all the meiotic stages except MII. Plants within strains differed significantly at the MI and quartet stages only. Except at AI stage, spikes collected on the same day did not differ significantly. Apparently, environment during premeiotic phase determined the extent and the pattern of meiotic instability in spikes of these strains.Chromosome counts were taken on 213 of the highly aberrant PMC at MI. Among these, 44 different chromosome numbers ranging from 4 to 76 per cell were recorded. Cells with deviating numbers from 22 to 28 were the most frequent (30.52 percent). Chromosome numbers 12, 17, 21, 22, 24, 26 and 28 occurred in 43.9 percent of the aberrant PMC, and the number of cells with these chromosome numbers ranged from 9 to 15.The metaphase 1 chromosome pairing in deviating PMC was not related to the number of chromosomes in these PMC. Apparently, deviating PMC had a random assortment of chromosomes. In aberrant PMC and others with apparently normal chromosome numbers (but irregular meiosis) homeologous chromosomes may pair. Functional gametes from such cells may perpetuate meiotic instability that had persisted in these three advanced generation common wheat strains of hybrid origin.     

Cytogenetic Study of a Trigeneric (Triticale × Trileymus) Hybrid

Summary In this study a new trigeneric hybrid involving species from the Triticum, Secale and Leymus was produced by crossing octoploid triticale (Jinsong49) with octoploid tritileymus (950059). The chromosome constitution of the parental amphiploid, trigeneric hybrid and its progenies were studied. Genomic in situ hybridization (GISH) analysis showed that Jinsong49 and 950059 had 44 wheat chromosomes, and 12 rye chromosomes, 12 L. mollis chromosomes respectively. The mean meiotic configuration of trigeneric hybrid F₁ was 13.17 I + 20.82 II + 0.37 III + 0.02 IV. GISH results indicated the trigeneric hybrid F₁ had 6 rye chromosomes and 6 Leymus chromosomes. In the selfed derivatives of the trigeneric hybrids, while the number of selfed generation increased, the mean number of chromosomes tends to decrease gradually and slowly. GISH results revealed that most plant tested in the progeny population had 8–12 rye chromosomes, and no Leymus chromosomes were detected. The results indicated that rye chromosomes can be preferentially transmitted in the progenies of trigeneric hybrid than Leymus chromosomes.     

Production of intergeneric hybrids between Brassica juncea and Diplotaxis virgata through ovary culture, and the cytology and crossability of their progenies

The cytogenetic study was investigated in the intergeneric F₁ hybrid, F₂and backcross progenies (BC₁). The plants used were Brassica juncea(2n=36) and Diplotaxis virgata(2n=18). Three intergeneric F₁ hybrids between two species were produced through ovary culture. They showed 36 chromosomes. It might consist one genome of B. juncea and two genomes of D. virgata. The morphology of the leaves resembled that of B. juncea. The color of the petals was yellow that was like in D. virgata. The size of the petal was similar to that of B. juncea. The mean pollen fertility was15.3% and the chromosome associations in the first meiotic division were(0–1)_IV+(0–2)_III+(8–12)_II+(12–20)_I. Many F₂ and BC₁seeds were harvested after open pollination and backcross of the F₁ hybrids withB. juncea, respectively. The F₂seedlings showed different chromosome constitutions and the range was from 28 to54 chromosomes. Most seedlings had 38chromosomes followed by 36, 40 and 54. The BC₁ seedlings also showed different chromosome constitutions and the range was from 29 to 62. Most seedlings had both 40and 54 chromosomes followed by 36, 46 and52. In the first meiotic division of F₂ and BC₁ plants, a high frequency of bivalent associations was observed in all the various kinds of somatic chromosomes. Many F₃ and BC₂ seeds were obtained by self-pollination and open pollination of both F₂ and BC₁ plants, and by backcrossing both F₂ and BC₁plants with B. juncea, respectively,especially, three type progeny with 36, 40or 54 chromosomes. The somatic chromosomes of the F₃ and BC₂ plants were further investigated. The bridge plants between B. juncea and D. virgata with 36 chromosomes may be utilized for breeding of other Brassica crops as well as B. juncea. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolutionary autoploidy in the sweet potato (Ipomoea batatas (L.) Lam.) and its progenitors

Summary The role of polyploidy in the evolution of the sweet potato. I. batatas (2n=6x=90), became more clear in 1971 when wild species with 30, 60, and 90 chromosomes were discovered. These species, I. leucantha (2x), I. littoralis (4x) and I. trifida (6x), are the progenitors of the sweet potato (6x), in an autopolyploid series with doubling of the I. leucantha B genome.In the present study the hypothesis of the origin of the sweet potato was confirmed by comparative studies on some plant characters, sexual compatibility, and the behavior of artificial hexaploids produced from I. leucantha (2x) and I. littoralis (4x).Since induced hexaploid I. leucantha exhibits weak growth, a few multivalents are formed at meiosis in the sweet potato, and there are differences in morphological and physiological characteristics between the artificial hexaploids and I. batatas, raw autoploidy seems unlikely, but a balanced or diploidised autoploidy could have been achieved by genic and chromosomal changes in the course of evolution. However, there is still sufficient homology so that meiotic pairing occurs usually between regular partners but also between homoeologous chromosomes in a certain situation.     

Discovery of 2n gametes in tetraploid oat Avena vaviloviana

Summary Sexual polyploidization via the action of 2n gametes (gametes with the sporophytic chromosome number) has been identified as the most important evolutionary mode of polyploidization among plant genera. This study was conducted to determine whether 2n gametes are present in the tetraploid level of the genus Avena (2n=4×=28) Twenty tetraploid Avena lines, representing four species and one interspecific hybrid, were screened for pollen grain size in order to differentiate between n and 2n pollen. Avena vaviloviana (Malz.) Mordv. line PI 412767 was observed to contain large pollen grains at a 1.0% frequency. Cytogenetic analyses of pollen mother cells of PI 412767 revealed cells with double the normal chromosome number (i.e., 56 chromosomes at metaphase I and anaphase I). The mode of chromosome doubling was found to be failure of cell wall formation during the last mitotic division that preceded meiosis. The resulting binucleate cells underwent normal meiotic divisions and formed pollen grains with 28 chromosomes. Based on the formation and function of 2n gametes, three models involving diploid and tetraploid oat lines are proposed to describe possible evolutionary pathways for hexaploid oats. If stable synthetic hexaploid oat lines could be developed by utilizing 2n gametes from diploid and tetraploid oat species through bilateral sexual polyploidization, the resulting hexaploids could be used in breeding programs for transferring genes from diploids and tetraploids to cultivated hexaploids.     

Early Embryo-Sac Development and Intraspecific Variation in the Mean Endosperm Cell Cycle Time in Vicia narbonensis (L.)

The early endosperm development in seven Vicia narbonensis populations was studied under a 23/ 16 ° C day/night temperature regime. Double fertilization occurred in less than 24 h after pollination. Endosperm divisions were synchronous until the 8th division of the endosperm nuclei. Aberrant endosperm nuclei were first observed alter the 8th division and their number increased after the 10th division. Endosperm nuclei continued to divide until the 12th day after pollination. During the following days a decrease in the number of endosperm nuclei was observed together with a substantial increase in the number of aberrants. Endosperm nuclei finally disintegrated. Neither cellularization nor starch deposition was observed in the endosperm. A positive correlation (r 0.69) was recorded between early pod length and the number of endosperm nuclei found in the ovules within each pod. In the populations studied the number of endosperm nuclei increased at a steady but different rate. This was followed by embryo development as well. It was concluded that intraspecific variation in the mean endosperm cell cycle time exists in Vicia narbonensis and that populations A 174 and A 202 should be more compatible to Vicia faba ev. ‘Polycarpe’ than population A 201.     

2n Gamete formation in the genus Brachiaria (Poaceae: Paniceae)

Microsporogenesis of several Brachiaria species of the Brazilian collection at Embrapa Beef Cattle has been analyzed in detail. This paper reports abnormal cytokinesis in three accessions of three different species (Brachiaria humidicola, 2n = 4x = 36, Brachiaria decumbens, 2n = 4x = 36, and Brachiaria dura, 2n = 6x = 54). Chromosomes paired in bi-, tri-, and quadrivalents in these accessions, whereas chromosome segregation at meiosis I was characterized by exclusion of laggards as micronuclei. In a high number of meiocytes, the first sign of cytokinesis appeared only in metaphase II and did not divide the meiocyte into a dyad. Total absence of cytokinesis was also detected among meiocytes in the second division. Since in both cases the two metaphase plates were very close, they favored the rejoining of chromosome sets after anaphase II and formed a restitutional nucleus in telophase II. Second cytokinesis occurred after telophase II in most meiocytes. Monads, dyads, and triads with n or 2n nuclei were observed among meiotic products. The 2n gametes observed correspond to the first division restitution (FDR). The number of affected cells in each accession was variable, but the number of microspores with restitutional nucleus, including those scored in tetrads and the released ones, did not exceed 9%. Although polyploidy is common in the genus Brachiaria, its origin is still unclear. Current results suggest that 2n gametes may have contributed to the evolutionary history of the genus.     

黑麦与光稃野燕麦杂交后代的减数分裂研究

本研究以黑麦和光稃野燕麦为亲本，对其杂种后代的减数分裂进行了研究。结果表明：杂种后代植株染色体数减少，花粉母细胞染色体数有14，15，16，17；染色体数越多，减数分裂中期I单价体出现的频数，多价体的类型和频数也增多，二价体基本稳定在6～7。     

Production and characterization of intergeneric hybrids between Brassica oleracea and a wild relative Moricandia arvensis

Intergeneric crosses were made between Brassica oleracea and Moricandia arvensis utilizing embryo rescue. Six F₁ hybrid plants were generated in the cross‐combination of B. oleracea × M. arvensis from 64 pods by the placenta‐embryo culture technique, whereas three plants were produced in the reciprocal cross from 40 pods by the ovary culture technique. The hybrid plants were ascertained to be amphihaploid with 2n = 23 chromosomes in mitosis and a meiotic chromosome association of (0–3)II + (17–23)I at metaphase I (M I). In the backcross with B. oleracea, some of these hybrids developed sesquidiploid BC₁ plants with 2n = 32 chromosomes that predominantly exhibited a meiotic configuration of (9II + 14I) in pollen mother cells. The following backcross of BC₂ plants to B. oleracea generated 48 BC₃ progeny with somatic chromosomes from 2n = 19 to 2n = 41. The 2n = 19 plants showed a chromosomal association type of (9II + 1I) and a chromosomal distribution type of (9¹/₂ + 9¹/₂) or (9 + 10) at M I and M II, respectively. These facts might suggest that they were monosomic addition lines (MALs) of B. oleracea carrying a single chromosome of M. arvensis that could offer potential for future genetic and breeding research, together with other novel hybrid progeny developed in this intergeneric hybridization.     

The origin of unreduced microspores due to aberrant cytokinesis in the meiocytes of potato and its genetic significance

Summary In 7 aneuhaploids of Solanum tuberosum and one clone of S. phureja the mechanism of unreduced microspore production has been studied. It has been observed that the unreduced microspores are produced not only by nuclear restitution at meiosis but also by aberrant cytokinesis in the meiocyte. The size and the number of nucleoli that are present in young microspores at the sporad stage give a fairly reliable indication of the ploidy level and the type of nuclear restitution that has occurred. These two aspects are helpful to deduce the meiotic events that have probably led to the formation of unreduced microspores in some cases. It has been found that triads and dyads are formed regardless of the type of nuclear division or restitution, indicating that cytokinesis is independent of karyokinesis to some extent. Cytological basis of unreduced microspore formation in diploid forms of potato appears to be more complex than it was supposed earlier.     

Reproductive Behavior of Hexaploid/Diploid Wheat Hybrids 1

The bottleneck restricting introgression of useful genes directly from diploid into hexaploid wheats is the low number of BC₁F₁ seeds obtained. In crosses between hexaploid wheat (Triticum aestivum L.; AABBDD) and Aegilops squarrosa L. (DD) or T. urartu Thum. (AA), this bottleneck may be overcome simply by pollinating a sufficient number of F₁ spikes. However, hybrids between hexaploid wheat cultivars (T. aestivum) and T. monococcum L. (AA) generally are highly female-sterile, often having no pistils. One T. monococcum accession, PI 355520, when crossed with T. aestivum, produced hybrids with female fertility in the same range as that of T. aestivum/A. squarrosa or T. aestivum/T. urartu hybrids, ca. 0.5 to 1.0 backcross seed per spike. We found that female fertility was controlled by two duplicate genes in PI 355520, and that this accession can be used as a bridging parent to introgress genes from other T. monococcum accessions into hexaploid wheat. Pairing of homologous chromosomes was less frequent and weaker in such crosses than in T. aestivum/A. squarrosa crosses, but homoeologous bivalents occurred at a rate of almost 0.5 II per cell. Restitution division was detected in crosses involving all three diploid species and was confirmed cytologically in crosses with PI 355520. Chromosome numbers of BC₁F₁ plants ranged from 35 to 67; plants with 49 or more chromosomes occurred at frequencies of 0.09 to 0.21 among progeny of A. squarrosa and T. urartu and 0.29 in progeny of T. aestivum/T. monococcum crosses involving PI 355520. These results are consistent with those of previous studies, demonstrating the potential of direct Hexaploid/diploid crosses for rapidly introgressing useful genes into Hexaploid wheat with minimum disturbance of the background genotype.     

Transfer of resistance to race 2 of Plasmodiophora brassicae from Brassica napus to cabbage (B. Oleracea spp. Capitata). IV. A resistant 18-chromosome B1 plant and its B2 progenies

Summary The somatic karyotype and meiotic chromosome behavior were studied in an 18-chromosome B₁ plant derived from backcrossing a triploid (Brassica napus x B. oleracea ssp. capitata) F₁ hybrid to cabbage. It is considered that cabbage chromosomes no. 1 and no. 7 were substituted by two shorter B. napus chromosomes. Meiotic disturbances were more apparent during the late stages of second division. Seed fertility of this plant was largely restored in the second backcrosses with both cabbage and broccoli. 18-chromosome B₂ plants resistant to race 2 of Plasmodiophora brassicae were recovered among the progenies.Contribution no. J. 725 from the Research Station, Research Branch, Agriculture Canada, St-Jean, Québec J3B 6Z8.