Meiosis in different F1-hybrids of Solanum acaule Bitt. x S. bulbocastanum Dun. and its bearing on genome relationship,fertility and breeding behaviour

Summary Meiosis was studied in some triploid, tetraploid and hexaploid F₁-hybrids from Solanum acaule x S. bulbocastanum and in a triploid F₁-hybrid from S. acaule x S. tuberosum-haploid.The only anomaly found was stickiness at metaphase I, the degree of which appeared to be related to the proportion of the S. bulbocastanum chromosomes in the hybrids. No stickiness occurred at pre- and post-metaphase stages.A clear allosyndetic pairing between chromosomes of the two S. acaule genomes was observed in all triploid and tetraploid hybrids. These genomes apparently are closely related and little differentiated. The triploids S. acaule x S. bulbocastanum had 0–1 trivalent per cell, whereas 2–5 trivalents per cell were observed in the triploid S. acaule x S. tuberosum-haploid. Therefore S. acaule is more closely related to S. tuberosum L. than to S. bulbocastanum. A small amount of pairing between S. acaule and S. bulbocastanum is apparent from the occurrence of multivalents in all hybrids.Hexaploid F₁'s (2n=72) showed a nearly complete homologous pairing of chromosomes (35.2 bivalents per cell) and generally a normal separation of chromosomes at anaphase: 36-36. This offers an explanation for their high fertility. Triploid F₁'s from S. acaule x S. bulbocastanum have a high frequency of univalents leading to irregular separation of chromosomes at anaphase and consequently to unbalanced gametes and extreme sterility. In the tetraploid F₁'s (2n=48) nearly complete bivalent pairing was observed, 50% expectedly being homologous and 50% homoeologous pairing. Separation of chromosomes at anaphase was generally normal 24-24. In spite of this normal behaviour and allowing for tight stickiness at metaphase the tetraploids are very sterile. A satisfactory explanation cannot yet be given.Selfing and intercrossing hexaploid F₁'s gives normal berry set and many seeds per berry. However crosses with S. tuberosum and even those with the fertile hexaploid hybrid from 8x-S. acaule x 4x-S. tuberosum are little successful: berry set is far below normal and the berries are either parthenocarpic or contain only one or two seeds. These rather unexpected results warrant further investigation. Large-scale selfings and intercrosses of triploid and tetraploid hybrids have not been successful as yet. Among the female gametes of tetraploid hybrids a few appeared to be functional in crosses with hexaploid hybrids and in those with S. bulbocastanum.     

Production and Cytogenetical Analysis of a Rye-Cytoplasmic Tetraploid Triticale

A rye-cytoplasmic tetraploid triticale was found in F_s progenies of crosses between tetraploid rye‘No 1323’and hexaploid triticale‘KT 77′. In the tetraploid triticale, two complete rye genomes and two mixed wheat genomes, consisting of the chromosomes 1A. 2A, 4A, 7A, 3B, 5B, and 6B are present. The rye cytoplasm did not affect stability of rye chromosome pairing during metaphase 1, since rye chromosomes participated in pairing irregularities just as did wheat chramosomes, even on a larger scale. The fertility of F₀, plants ranged from 0 to 75.6 %, always associated with high grain shrivelling. The analyzed pairing behaviour of induced triploid hybrids from crosses between the tetraploid triticale and diploid rye indicates the presence of at least one wheat-rye translocation in one of the investigated triploid plants.     

The production and behaviour of hexaploid hybrids between Hordeum vulgare and H. Bulbosum

Summary To produce hexaploid (or other polyploid) hybrids, diploid or tetraploid Hordeum vulgare was crossed with hexaploid or octoploid H. bulbosum, and perennial triploid hybrids between the two species were treated with colchicine. The crosses did not yield viable plants: seedset was low, the seed aborted and embryo culture was unsuccessful. The colchicine treatments geve rise to plants in which hexaploid chromosome numbers were observed. At the hexaploid level chromosomal instability occurred, resulting in chromosome elimination.The colchicine-treated triploid hybrids showed in the first years after the treatment better fertility after open flowering than untreated plants, but the level of fertility remained very low. The offspring consisted of haploid, diploid and approximately triploid plants like H. vulgare, tetraploid and approximately tetraploid plants like H. bulbosum, and plants with hybrid morphology and unstable chromosome number, which were highly sterile. Thus the crossing barrier between H. vulgare and H. bulbosum could not be broken down at higher ploidy level.     

八倍体小黑麦(Triticale)与六倍体小黑麦杂交若干问题的探讨

本工作以改进八倍体小黑麦与六倍体小黑麦的经济性状为目的,对60个杂交组合 F_1的田间出苗率、结实率、杂种后代的性状分离、新类型的形成以及细胞遗传的若干问题进行了探讨,观察到 F_1田间出苗率、结实率以八倍体为母本的杂交组合显著好于以六倍体为母本的杂交组合。由于杂种是普通小麦、硬粒小麦、黑麦三个物种种质的再度组     

Structural hybridity in the series Etuberosa of the genus Solanum and its bearing on crossability

Summary Three non-tuberous Solanum species of series Etuberosa-S. brevidens (brd), S. etuberosum (etb) and S. fernandezianum (frn)-upon selfing displayed compatible berry set but seed set varied and on an average was below normal. Also in crosses between species berry set was unhampered (apart from the crosses with short-styled frn as pollen plant), but the interspecific hybrid plants showed various degrees of sterility and a low average seed set upon selfing. Cytological investigations revealed that in pure species-plants semi-sterility regularly occurred and was invariably associated with multivalent associations (up to decavalents). This may account for the decreased seed set found after selfing and intercrossing. Meiosis in F₁-hybrids was even more irregular, 75% of the pmc's showing multivalents involving from 4 to as many as 22 chromosomes in various complicated configurations in which even numbers of chromosomes were predominant. However, anaphase I separation was numerically balanced in more than 75% of the cells. Pollen stainability was very low in the hybrids.It is concluded that heterozygous translocations occur within each species and that the genomes of the species are differentiated by means of gross structural changes. In view of this and of the high chromosome pairing affinity the genome symbols E ₁, E ₂ and E ₃ are proposed for etb, brd and frn, respectively.Crossability of the Etuberosa species with tuberous S. pinnatisectum (pnt) was slightly more successful (average 2.7 seeds/berry) than that of Etuberosa species hybrids with S. pinnatisectum (average 1.7 seed/berry). In the hybrids studied-etb × pnt, frn × pnt and (brd × etb) × pnt-a high frequency of univalents was the rule (range 6–16 per cell). In addition rod bivalents (4–8 per cell) and a low frequency of trivalents occurred (0–3 in the biparental hybrids and 1–4 in the trispecific hybrid).Utilization of the valuable characters of Etuberosa for potato breeding is dependent on the degree of gene exchange between Etuberosa chromosomes and chromosomes of tuberous species. In the diploid hybrids such exchange, though limited, may take place through bivalents and trivalents. As shown in this article such diploid hybrids are non-crossable because of complete male and female sterility. Only after doubling the number of chromosomes the hybrids are fertile and can be used for further crosses.     

Wheat Substitutions in Hexaploid Triticale*

Crosses between octoploid and hexaploid triticales have been made m breeding programs for several years, From an analysis of the progeny of such crosses where selections for an octoploid-like phenotype had been made, it was established that 149 out of 150 lines were hexaploid in chromosome number, C-banding and in situ hybridization demonstrated that all but five of the 62 lines analyzed in detail contained visible chromosomes or segments from the D genome. Only four lines had D-genome chromosome replacing rye chromosomes. All of the remaining 53 D-genome substitutions involved the replacing of wheat chromosomes from either the A or B genomes. This establishes the ease with which D-genome genes can be placed into triticale without the loss of rye chromosomes.     

Cytogenetic study of Helianthus laevigatus and its F1 and BC1F1 hybrids with cultivated sunflower, Helianthus annuus

Although the wild sunflower species Helianthus laevigatus has not been extensively studied it may be considered for sunflower breeding as a potential source of desirable genes for Sclerotinia stalk rot resistance and high contents of proteins and linoleic acid in the seed. A set of six H. laevigatus populations was crossed to cultivated sun~ower lines and produced nine F₁ (2-14 plants) and 66 BC₁F₁ hybrid combinations (1-13 plants). Male sterility occurred in F₁ and BC₁F₁ hybrid combinations and pollen viability was lower in the progenies than in the parents (51.6-77.2%in F₁ and in F₁ and 4.8-34.0% in BC₁F₁). Meiosis was normal in the H. laevigatus populations It was found that this tetraploid species also occurred in a hexaploid form Numerous irregularities were observed in the meiosis of the F₁ interspecific hybrids During diakinesis, quadrivalents and hexavalents were recorded in addition to bivalents Dislocated chromosomes and chromosome bridges were present in the other phases The chromosome number in F₁ was 68 (tetraploid). Irregularities in chromosome pairing were observed in the interspecific hybrids at BC₁F₁. There were many univalents, and trivalents quadrivalents and hexavalents were also present The chromosome number in the BC₁F₁ generation ranged from 34 to 60. The occurrence of meiotic irregularities in the F₁ and BC₁F₁ interspecific hybrids indicates that H. laevigatus and the cultivated sunflower differ in genome constitution.     

甘蓝型油菜亚基因组间杂种（ArAnCcCn）的细胞学和育性研究

以新型甘蓝型油菜（A^rA^rC^cC^c）与自然甘蓝型油菜品种湘油15（AⁿAⁿCⁿCⁿ）杂交，人工配制了A^rAⁿC^cCⁿ油菜亚基因组间杂种。亚基因组间杂种花粉母细胞减数分裂基本正常，中期I大多以19个二价体的构型存在，但偶尔也有单价体、三价体和四价体等异常情况出现。在减数分裂后期Ⅰ和后期Ⅱ，绝大多数细胞中的同源染色体和姊妹染色单体正常分离，仅在很小比例的花粉母细胞中观察到落后染色体。不同亚基因组间杂种的减数分裂正常程度存在一定的差别，其中Ar和An基因组之间的遗传分化可能是主要原因。产量分析表明亚基因组间杂种具有较强的杂种优势潜势，并且亚基因组间杂种的种子产量和正常花粉母细胞所占比例呈显著正相关。     

Interspecific hybridization in sexual diploid Allium senescens var. minor× apomictic tetraploid A. nutans and sexual diploid A. senescens var. minor× apomictic hexaploid A. senescens

Fourteen interspecific hybrids in sexual diploid Allium senescens var. minor× apomictic tetraploid Allium nutans L. crosses, and eight interspecific hybrids in sexual diploid A. senescens var. minor× apomictic hexaploid A. senescens L. crosses were produced. The number of chromosomes was 2n= 24 in interspecific hybrids of diploid × tetraploid, and 2n= 32 in diploid × hexaploid crosses. Triploid and tetraploid interspecific hybrids showed intermediate parental morphological characteristics. Tetraploid interspecific hybrids of A. senescens var. minor×A. senescens crosses formed two groups based on leaf colour and leaf width. Seeds were formed in 11 out of 14 triploid interspecific hybrids under natural conditions. In cytological observations of parthenogenesis, three out of 12 triploid interspecific hybrids and five out of eight interspecific tetraploid hybrids were observed. Parthenogenesis ranged from 26.0% to 86.0% in five tetraploid interspecific hybrids. Non-parthenogenesis to parthenogenesis segregated in a 3:5 ratio in A. senescens var. minor×A. senescens crosses.     

Genomic Affinity Relationships in the Genus Avena

Many interspecific hybrids between diploid, tetraploid and hexaploid Avena species were obtained without artificial embryo cultures. Most of the hybrids showed variable chromosome pairing with univalents, bivalents and multiple associations, even in PMC in the same hybrid, at first metaphase; making ii very difficult to understand the degree of genome affinity between parent species, and especially between different species. The basic or theoretical chromosome pairing consisting of bivalents and univalents is presented, in which the chromosome ratios (%) statistically agree with observations. In addition, the per cent frequency of chromosomes forming multivalents is calculated. On the bases of these numerical data, the genomic affinity relationships are discussed for present and previously reported interspecific hybrids or Avena.     

Inheritance of Cold Hardiness in Triticum aestivum × Synthetic Hexaploid Wheat Crosses

Synthetic hexaploid wheat, produced by combining tetraploid wheat (AB genome) with Triticum tauschii (D genome), was crossed to modern hexaploid wheat (Triticum aestivum ABD genome) in an attempt to introduce new cold hardiness genes into the common hexaploid wheat gene pool. The cold hardiness levels of F) hybrids ranged from similar to parental means to equal to the hardy parent, indicating that cold hardiness was controlled by both additive and dominant genes. As expected when dominant gene action is involved, differences between F₂ and parental means were smaller than comparable differences in the F., Frequency distributions of F₂—derived F₃ lines also suggested that dominant genes were involved in the control of cold hardiness in some crosses. Heritability estimates for cold hardiness ranged from 63 to 70 % indicating that selection for cold hardiness should be effective in populations arising from crosses between common and synthetic hexaploid wheat. However, high selection pressure on the progeny of crosses that included the most hardy T. aestivum, T. durum, and T. tauschii accessions as parents did not identify transgressive segregates for improved cold hardiness. These observations indicate that the close wheat relatives, sharing common genomes with T. aestivum, are not promising sources of new genes to increase the maximum cold hardiness potential of common hexaploid wheat.     

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.     

Homeology Between the Chromosomes of Avena macrostachya and the Avena C Genome

To gain more information about the widely unknown genome constitution of the taxonomically and agronomically interesting wild oat Avena macrostachya, this tetraploid species was crossed (as the pollinator) with several diploid Avena species. Hybridization failed with A. strigosa (A_s A_s), A. hirtula (A_s A_s), and A. longiglumis (A₁ A₁), but succeeded with A. atlantica (A_s A_s), A. prostrata (A_p A_p), and A. pilosa (C_p C_p). The output of hybrids and the vitality of the F: plants were highest in the combination A. pilosa×A. macrostachya. Hybrids from this cross relatively frequently showed trivalents at metaphase I as compared to the nearly consistent formation of seven univalents and seven bivalents in hybrids with the A genome species A. atlantica and A. prostrata. Altogether, the results confirm an auto-ploid origin of A. macrostachya and demonstrate that among the two known basic genomes A and C of the genus Avena only the latter exhibits considerable homeology to the A. macrostachya genome.     

The successful transfer of primocane fruiting expression from raspberry to Rubus hybrid berry

This paper describes the successful transfer of the primocane fruiting (PF) character from raspberry (Rubus idaeus) to Rubus hybrid berries following crosses between raspberry and blackberry. The transfer was achieved by doubling the chromosome complement of PF raspberry selections in culture using colchicine, and using regenerant tetraploid PF plants as the female parents in crosses with tetraploid or octoploid blackberries, or with hexaploid hybrid berries (all summer fruiting). The results show that the PF character is expressed more readily in hybrid plants where at least 50% of the chromosomes in the hybrid genome are derived from raspberry. In vitro germination of cut seeds overcame the poor germination often encountered in Rubus and ensured the rapid recovery of hybrid seedlings. This revised version was published online in July 2006 with corrections to the Cover Date.     

Hexaploid triticales from hybrids of 'Langdon' durum D-genome substitutions with 'Gazelle' rye

The formation of unreduced gametes in some hybrids between disomic D‐genome substitutions (DS) of durum wheat cv.‘Langdon’ and rye provides a convenient approach for the rapid introduction of D‐genome chromosomes into hexaploid triticale. Meiotic pairing at metaphase I and seed fertility in spontaneous and colchicine‐induced amphidiploids derived from F₁ hybrids between a set of ‘Langdon’ DS and ‘Gazelle’ rye were analysed. The purpose was to determine the effects of the substitution of D‐genome chromosomes for their A‐ and B‐genome homoeologues on hexaploid triticale and to select stable disomic D‐genome substitutions of hexaploid triticale. The results showed that the disomic substitutions with D‐genome slightly increased the frequency of univalents (1.0‐3.13) compared with the ‘Langdon’ control primary hexaploid triticale (0.76). Substitutions 2D(2A) and 3D(3B) were partly desynaptic. The substitutions 1D(1A), 1D(1B) and 7D(7B) exhibited high seed fertility but the others had decreased fertility. Except for 2D(2A), 5D(5A), 3D(3B) and 5D(5B), 10 of the 14 possible hexaploid triticale D‐genome disomic substitutions have been obtained. The results suggest that the poor compensation ability of some D‐genome chromosomes for their homoeologous A‐ and B‐genome chromosomes is a major factor affecting meiotic stability and fertility in the hexaploid triticale D‐genome substitutions.     

Rapid Incorporation of D-genome Chromosomes into A and/or B Genomes of Hexaploid Triticale

Two series of progenies were developed from hybrids between octoploid (AABBDDRR) and tetraploid triticale ((AB)(AB)RR). One arose from the successive selfing of the F₁s, while the second was established after one backcross of the F₁ hybrids with the respective 8 × triticale parent. Altogether, 250 F₃ and BC₁F₂ lines were developed, of which 112 were karyotyped in the F₄/F₅ or in BC₁F₃/BC₁F₄ generations using C-banding and SDS-PAGE. The 112 lines represented 61 different karyotypes, of which 39 appeared to be stabilized, having pairs of homologous wheat chromosomes only, while 22 karyotypes exhibited 1—3 heterologous pairs. The frequency of karyotypically stabilized lines originating from the series with one backcrossing was much higher (79.5 %) than those derived from the successive selfing of the F₁ hybrids (51.7%). Six lines had the pure hexaploid triticale chromosome constitution. The frequency of disomic substitutions of D genome chromosomes for their homoeologous A and/or B genome chromosomes ranged from one to six per line with an average of 1.7. Except for 3B and 6B all possible D(A/B) substitutions were obtained. Chromosomes ID and 3D substituted for their homoeologues with the highest frequency, while the substitution of chromosome 4D for 4A or for 4B was the least frequent. D(R) substitutions were found in eight lines only. A complete set of 6x triticale lines was established in which chromosome ID was present in all possible combinations, i.e. single 1D(1A/1B/1R) disomic substitutions as well as disomic ID addition.     

Selection for seed set in crosses of Avena sativa L. × A. Abyssinica Hochst.

Summary Precoz oat line, discovered in a mass planting of wild oats in Colombia, is a tetraploid belonging to Avena abyssinica. It crossed easily with Kyto, but not with Bacata, both hexaploid oat cultivars. Pentaploid hybrid plants of Precoz × Kyto and Precoz × Bacata had 35 chromosomes, tended to have seven bivalents and 21 univalents at meiosis, and were completely sterile, Colchicine treatment of pentaploid F₁ plants produced decaploid tillers from which five amphiploid seeds were obtained. Selection for high percentage of seed set in the third and fourth amphiploid generations of Precoz × Kyto gave fifth generation plants with 80–100% fertility. The chromosome numbers of these plants approximated the octoploid level. Some derivatives from crosses of third generation amphiploids with hexaploid cultivars had the hexaploid chromosome number and good fertility.Pentaploid hybrid plants and amphiploid derivatives from them were as early in heading as Precoz when tested at either 12-or 15-hour photoperiods. This suggests that the photoperiod response from Precoz is dominant and has good penetrance.Journal Paper no J-7408, Project 1752.     

Transfer of high kernel weight and high protein from wild tetraploid wheat (Triticum turgidum dicoccoides) to bread wheat (T. aestivum) using homologous and homoeologous recombination

Summary Wheat pentaploids were produced by hybridizing a high kernel weight (1000 grain wt=56 g), high protein (25.4%) line of wild tetraploid wheat (Triticum turgidum dicoccoides) as male parent, with the three hexaploids (T. aestivum) — normal Chinese Spring and its two homoeologous pairing mutants, ph _1b and ph ₂. The pentaploids were crossed as female parents to the two commercial hexaploid cultivars Warigal and Barkaee and 42-chromosome stable plants selected from the F₁ of the pentaploid x hexaploid crosses.Mean protein content of certain F₃ lines from all six pentaploid x hexaploid crosses was significantly higher than Chinese Spring and the respective commercial hexaploid parent (p<0.005) indicating high protein had been transferred from the tetraploid to the hexaploid level.Kernel weight amongst certain F₃ lines of the three pentaploids x Barkaee was significantly (p<0.0005) higher than either Chinese Spring or Barkaee, indicating the transfer also of high kernel weight from the tetraploid to the hexaploid level. However kernel weight was not significantly increased over Warigal in any F₃ lines of its crosses with the three pentaploids.High levels of homoeologous chromosome pairing in the ph-mutant pentaploids, plus evidence for significant modification of the composition of high-molecular weight (HMW) glutenin subunits of grain protein in certain F₃ derivatives of the ph-mutant pentaploid x hexaploid, crosses indicates that the ph-mutant-derived lines may possess novel (intergenome) genetic recombination, at least for high protein, and possibly kernel weight.     

中间偃麦草染色体组型研究

通过系统对比分析 ABD、AB、AG 等类型小麦与中间偃麦草杂交后 F_1的减数分裂资料确认中间偃麦草没有与小麦 B 组同源的染色体组。减数分裂染色体 N 带分析进一步证实了这一点。以目前研究来看,中间偃麦草染色体组型以有两组彼此部分同源,另一组与它们远缘为宜,建议以字母 NNiX 代之。     

The production of multispecific octoploids from Fragaria species and the cultivated strawberry

Summary A method of incorporating genetic material from five species of Fragaria and the cultivated strawberry into fertile octoploids is described. A synthetic octoploid derived from a hexaploid x diploid hybrid was crossed to octoploid cultivar breeding lines until a BC₂ hybrid was produced. A second synthetic octoploid in which two diploid species and a tetraploid species were combined was crossed to a cultivated strawberry to produce a hybrid breeding clone. The two breeding clones were crossed and 222 seedlings were produced. The seedlings were generally vigorous and fruitful, some having commercial potential.