POLYPLOIDY,DYSPLOIDY, AND CHROMOSOME PAIRING IN ECHEVERIA (CRASSULACEAE) AND ITS HYBRIDS

The 140+ species of Echeveria have more than 50 gametic chromosome numbers, including every number from 12 through 34 and polyploids to n = ca. 260. With related genera, they comprise an immense comparium of 200+ species that have been interconnected in cultivation by hybrids. Some species with as many as 34 gametic chromosomes include none that can pair with each other, indicating that they are effectively diploid, but other species with fewer chromosomes test as tetraploids. Most diploid hybrids form multivalents, indicating that many translocations have rearranged segments of the chromosomes. Small, nonessential chromosomal remnants can be lost, lowering the number and suggesting that higher diploid numbers (n = 30–34) in the long dysploid series are older. These same numbers are basic to most other genera in the comparium (Pachyphytum, Graptopetalum, Sedum section Pachysedum), and many diploid intergeneric hybrids show very substantial chromosome pairing. Most polyploid hybrids here are fertile, even where the parents belong to different genera and have very different chromosome numbers. This seems possible only if corresponding chromosomes from a polyploid parent pair with each other preferentially, strong evidence for autopolyploidy. High diploid numbers here may represent old polyploids that have become diploidized by loss, mutation, or suppression of duplicate genes, but other evidence for this is lacking. Most species occur as small populations in unstable habitats in an area with a history of many rapid climatic and geological changes, presenting a model for rapid evolution.     

THE PROBLEM OF PLOIDY IN ECHEVERIA (CRASSULACEAE) II. TETRAPLOIDY IN E. SECUNDA

Every chromosome number from n = 12 to n =34 and also many higher numbers are known in one or more of the 130+ species of Echeveria, and the numerical boundary between diploids and tetraploids is not immediately apparent. Echeveria also is extraordinary for the number and diversity of hybrids that it can produce in cultivation, both within the genus and with species of several related genera. In 42 collections studied, the morphologically and cytologically variable E. secunda of central Mexico has n = 30-32, often with one or more B-chromosomes, and some quadrivalents are formed at meiosis in nearly every cell. Twenty-four hybrids of E. secunda, with 22 species or cytotypes considered diploids, resemble the former much more closely in appearance, and at meiosis 15-16 paired elements (bivalents and multivalents) are formed, never more, regardless of the number of chromosomes, 12 to 34, that were received from the other parent. It is concluded that the 15-16 paired elements in these hybrids are formed by the 30-32 chromosomes received from E. secunda, and that most chromosomes from the other parents occur as univalents, although usually a few associate with pairs from E. secunda to produce multivalents. Hybrids of E. secunda with 11 definitely tetraploid species having n = 34 to n = 68 are nicely intermediate in morphology between their parents, form mostly or entirely bivalents at meiosis, and most, probably all, including five intergeneric hybrids, are fertile. These observations are all consistent with the conclusion that E. secunda is an autotetraploid, even though no plants of the species having n = 15 or 16 have been found, and even though some other species of Echeveria having as many as 34 gametic chromosomes appear to be effectively diploid. Observations on pollen stainability and on second-generation hybrids are all compatible with this conclusion. The high chromosome numbers in many Mexican Crassulaceae that are now effectively diploid may have originated as polyploids that have become diploidized by mutation, loss, or suppression of duplicated chromosomes, segments, and genes. Hybrids of E. secunda, with three other species that appear to be tetraploids, have less regular meiosis, apparently because all of the chromosomes from the other parents do not regularly form pairs in the hybrids. These three species may represent intermediate stages in the processes of diploidization.     

CHROMOSOMES OF GRAPTOPETALUM AND THOMPSONELLA (CRASSULACEAE)

Chromosome numbers are reported for probably all 11 species of Graptopetalum (x = 30–35) and for both species of Thompsonella (x = 26). Plants of two species of Graptopetalum have gametic numbers from about 240–275, more than have been reported in any other seed plants. In hybrids the 30–35 chromosomes in the basic genome of Graptopetalum and likewise the 26 in Thompsonella apparently do not pair among themselves, and the genomes seem to be no more potent genetically than those of other species in their subfamily having as few as 12 chromosomes. Species with these gametic numbers are therefore considered to be diploid. On the other hand, in hybrids between a diploid and a plant with a very high chromosome number the phenotype of the latter predominates, and most of its chromosomes pair with each other. Many such hybrids are fertile. These facts suggest that the high polyploids arose by autoploidy rather than by alloploidy. Nevertheless, they may store heterozygosity at some gene loci and release it in various dosages and proportions each generation.     

THE QUESTION OF POST-REDUCTION IN SPHAGNUM

The 19 spatially distinct chromosomal units at first meiotic metaphase in sporophytically diploid species of Sphagnum have usually been considered to be bivalents, but one investigator (Sorsa, 1956) has interpreted them as chromosomes from dissociated bivalents and meiosis as post-reductional. The present studies on diploid S. squarrosum (Pers.) Crome establish the chromosome number on the basis of the following evidence: there are in addition to m-chromosomes, 19 pairs of chromosomes in early prophase, 19 bivalents at diakinesis, 19 chromosomes in each of the two sets at second metaphase, 19 daughter chromosomes in each of the four sets at late second anaphase, and 19 chromosomes in gametophytic mitoses. The 19 bodies at first meiotic metaphase in diploid species are true bivalents in loose secondary association, which has led to their erroneous interpretation as chromosomes of dissociated bivalents. The gametic chromosome number in sporophytically diploid Sphagnum is therefore, without doubt, n = 19, and this evidence negates the claim for post-reduction in Sphagnum.     

The effect of B chromosomes on homoeologous pairing in species hybrids

The effect of B chromosomes on chromosome pairing at meiosis was investigated in the species hybrid Lolium temulentum x L. perenne at both the diploid and tetraploid level. The presence of B chromosomes drastically reduced association of homoeologous chromosomes in both the diploids and tetraploids. This was evident from the high frequency of univalents recorded in PMC's of diploid hybrids with B's and from the predominantly bivalent association of homologous chromosomes in tetraploids of this type. In the absence of B's homoeologous pairing was extensive giving a high frequency of bivalents in the diploids and multivalents as well as bivalents and univalents in the tetraploids.     

Genome relationships between Hordeum vulgare L. and H. bulbosum L.

Interrelationships between H. vulgare (2x=14) and H. bulbosum (2x=14; 4x=28) were estimated on the basis of the karyotypes and the pairing behaviour of the chromosomes in diploid, triploid and tetraploid hybrids obtained with the aid of embryo culture. — A comparison of the karyotypes of the two species revealed similarities as well as differences. It was concluded that at least 4 or more of the chromosomes were similar in morphology and probably closely related. — Diploid and tetraploid hybrids are rarely obtained and their chromosome numbers tend to be unstable whereas triploid hybrids (1 vulgare + 2 bulbosum genomes) were stable and relatively easy to produce. In the diploid hybrid only 40% of the meiotic cells contained 14 chromosomes while the numbers ranged from 7 to 16 in other cells. All hybrids exhibited pairing between the chromosomes of the two species. Diploid hybrids had a mean of 5.0 and a maximum of 7 bivalents per cell in those cells having 14 chromosomes. Triploid hybrids from crosses between 2x H. vulgare and 4x H. bulbosum exhibited a mean of 1.5 and a maximum of 5 trivalents per cell. In a hexaploid sector found following colchicine treatment of a triploid the mean frequencies of chromosome associations per cell were: 5.5_I+8.0_II+0.7_III+3.7_IV+0.3_V+0.4_VI. One unstable 27 chromosome hybrid obtained from crosses between the autotetraploid forms had a mean of 1.1 and a maximum of 4 quadrivalents per cell. The chromosome associations observed in these hybrids are consistent and are taken as evidence of homoeologous pairing between the chromosomes of the two species. Interspecific hybridization between these two species also reveals that chromosome stable hybrids are only obtained when the genomes are present in a ratio of 1 vulgare2 bulbosum. Based upon the results obtained, the possibility of transferring genetic characters from H. bulbosum into cultivated barley is discussed.     

Panicum shastense (gramineae), a sterile hybrid between P. Pacificum and P. Scribnerianum

Natural and synthetic hybrids are reported betweenPanicum, scribnerianum Nash and two other species,P. pacificum Hitchc. & Chase andP. occidentale Scribn. & Merr. The hybrids are morphologically intermediate between the parents, and those in whichP. pacificum is one parent strongly resemble the rareP. shastense Scribn. & Merr. All the parent plants studied cytologically had regular meiosis and a gametic number ofn = 9 chromosomes. The hybrids usually showed good pairing of chromosomes in meiosis, but irregularities such as univalents and lagging chromosomes were common. In the hybrids generally less than 7% of the pollen grains stained with cotton blue in lactophenol, and the plants set no seed, whereas the parent plants generally had over 90% pollen stainability and a high percentage of seed set. Pollen fromP. shastense, like that of the hybrids, was 3–4% stainable.Panicum shastense is believed to be based on a first generation sterile hybrid and the nameP. xshastense is proposed.     

Pairing and recombination at meiosis of Brassica rapa (AA) × Brassica napus (AACC) hybrids

Interspecific crosses contribute significantly to plant evolution enabling gene exchanges between species. The efficiency of interspecific crosses depends on the similarity between the implicated genomes as high levels of genome similarity are required to ensure appropriate chromosome pairing and genetic recombination. Brassica napus (AACC) is an allopolyploid, resulting from natural hybridization between Brassica rapa (AA) and Brassica oleracea (CC), both being diploid species derived from a common ancestor. To study the relationships between genomes of these Brassica species, we have determined simultaneously the pairing and recombination pattern of A and C chromosomes during meiosis of AAC triploid hybrids, which result from the interspecific cross between natural B. napus and B. rapa. Different AAC triploid hybrids and their progenies have been analysed using cytogenetic, BAC-FISH, and molecular techniques. In 71% of the pollen mother cells, homologous A chromosomes paired regularly, and usually one chromosome of each pair was transmitted to the progeny. C chromosomes remained mainly univalent, but were involved in homoeologous pairing in 21.5% of the cells, and 13% of the transmitted C chromosomes were either recombined or broken. The rate of transmission of C chromosomes depended on the identity of the particular chromosome and on the way the hybrid was crossed, as the male or as the female parent, to B. napus or to B. rapa. Gene transfers in triploid hybrids are favoured between A genomes of B. rapa and B. napus, but also occur between A and C genomes though at lower rates.     

CYTOTAXONOMIC STUDIES IN ELEOCHARIS SUBSER. PALUSTRES: CENTRAL UNITED STATES TAXA

Comparative chromosomal and morphological studies indicate that four species are present in the area surveyed. Eleocharis smallii Britt. is primarily diploid with 2n = 16, although sporadic polyploids with 2n = 36 also occur. E. macrostachya Britt. is morphologically similar with unstable polyploid numbers ranging around 2n = 38 and multivalents and univalents present in meiosis. E. xyridiformis Fern. & Brack., a species generally synonymized with E. macrostachya, is shown to be a morphologically distinct species with 2n = 18, 19, and 20. The 19-chromosome types are trisomic for one of the long chromosomes, the three homologs pairing in meiosis as a large chain trivalent. The trivalent separates equationally in the first division and preferentially in the second so that only 9- or 10-chromosome pollen grains containing an extra chromosome are formed. Trisomic cytotypes may potentially produce normal (18), reconstituted trisomies (19), or tetrasomic (20) plants, although tetrasomics have not been found. The 20-chromosome cytotype is not the expected tetrasomic, as it is karyotypically distinct from either the 18 or 19 cytotypes. In all species somatic mutations including translocations, translocation-retranslocations, and chromosome fragmentation (agmatoploidy) have been observed of which the significance, if any, has not been determined.     

EXPERIMENTAL HYBRIDS IN EPILOBIUM (INCLUDING SECT. ZAUSCHNERIA) SPECIES WITH N = 15 (ONAGRACEAE)

Experimental hybrids involving the three diploid subspecies of Epilobium sect. Zauschneria formed 15 bivalents at meiotic metaphase I, as did experimental hybrids between the three other species of Epilobium (comprising sect. Cordylophorum) with n = 15. The gametic chromosome number of E. suffruticosum Nutt., n = 15, and its relationship to the other two species are reported for the first time. Although we have not obtained hybrids between the species of these two sections, their morphological similarities are impressive and they are surely closely related.     

Study of the mitotic and meiotic chromosomes of sotol (<i>Dasylirion cedrosanum</i> Trel.)

The genus Dasylirion is a group of plants typically present in the Chihuahuan Desert, perennial, with a dioecious sexual behavior and commonly called sotoles. This genus has been little studied from the biological point of view, and the bases of its reproductive response remain unknown. In this work we studied the chromosome number and meiotic response of Dasylirion cedrosanum in the county of Saltillo, Coahuila, located at the North East of Mexico. For the preparation of mitotic chromosomes, we used a technique based on enzymatic treatment with pectolyase and cellulase, as well as staining with acetocarmin dye. For the study of meiosis, male flower buds were collected, fixed and stained for analysis with the same dye. As a result, the gametic (n = x = 19) and somatic chromosome (2n = 38) numbers of D. cedrosanum are reported for the first time, being consistent with previous findings in other Dasylirion species, which points to a constant ploidy level across the genus. Variation was observed in the morphology and size of the somatic chromosomes, with types ranging from submetacentric to subtelocentric, and sizes oscillating in a range of 4.43 µm, with an average total length of 112.38 µm for the diploid chromosome complement. This shows that the chromosome complement of D. cedrosanom would belong to a 3B classification of Stebins, with a medium variation between chromosome lengths and low chromosome asymmetry. This variation indicates the feasibility of constructing a chromosome ideotype for this species. The meiotic chromosome pairing showed a chromosome behavior consistent with a disomic inheritance characteristic of a diploid species, with prevalence of ring and chain bivalents, typically without pairing abnormalities. Bivalent configurations in all cases were symmetrical.The normal and symmetrical meiotic pairing indicates a balanced production of gametes, and suggests the absence of heteromorphic sex determination.     

CHROMOSOME PAIRING AND POLLEN FERTILITY IN INTERSPECIFIC HYBRIDS OF SPECIES OF PARTHENIUM (ASTERACEAE)

Meiotic analyses and pollen viability tests were performed on F, hybrids between diploid guayule (Parthenium argentatum Gray 2n = 36), P. rollinsianum Rzedowski (2n = 36), P. alpinum var. tetraneuris Barneby (2n = 36), and P. alpinum var. alpinum Nutt. (2n = 36). Parthenium chromosomes are small and karyomorphologically similar, and meiotic analysis is difficult because of chromosome clumping. However, cytogenetic studies at metaphase I indicated univalents can be seen in a lateral view of the metaphase plate. Chromosome pairing and the number of univalents varied within and between the interspecific hybrids, with an average univalent number of 1.54 for the P. rollinsianum hybrids, 2.36 for the P. alpinum var. tetraneuris hybrids, and 2.46 for the P. alpinum var. alpinum hybrids. Pollen viability tests for the parental species and the hybrids were conducted by germination of pollen grains on stigmas. The percent of viable pollen recorded for the diploid guayule hybrids with P. rollinsianum, P. alpinum var. tetraneuris, and P. alpinum var. alpinum are 21.94, 13.47, and 11.17, respectively. The degree of chromosome pairing and pollen viability is striking because there are many morphological differences between the parents. The chromosome homology of these species based on their pairing behavior allows for the design of a backcross breeding program that would permit the transfer of the desirable characteristics from these species into diploid guayule.     

Meiosis and pollen mitosis in diploid and triploid Colocasia antiquorum Schott.

The relationship between diploid and triploid forms of Colocasia antiquorum Schott. was assessed through comparative meiotic and pollen mitotic studies. Owing to poor spreading of the chromosomes of both materials, karyological observations on pachytene nuclei were limited to a few chromosomes. Among the two nucleolar chromosomes and a metacentric, telochromomere-bearing chromosome of the diploid, the latter and one of the nucleolar chromosomes characterized by a heteropycnotic short arm were identified in both bivalent and trivalent associations in the triploid. The homologues in these cases were homomorphic and intimately paired. Two types of heteromorphic bivalents exhibiting partial pairing of homomorphic segments were also recorded in the triploid. Among the 14 bivalents of the diploid at diakinesis, two were nucleolus-associated. In the triploid, chromosomal associations at diakinesis included trivalents (2 to 9), bivalents and univalents, and the chiasma frequency per paired chromosome was lower than in the diploids. In 21.6 percent of the PMCs at this stage intragenomic pairing of one or two chromosomes was observed. Post-diakinesis stages in the diploid were regular while in the triploid they were marked by various irregularities in a majority of the cells. However, fertility (stainability), size and divisional frequency of pollen in both materials were remarkably similar. Chromosome numbers in pollen nuclei in the triploid ranged from 8 to 25. Based on these data an autopolyploid origin for the triploid Colocasia and a lower base number than the gametic chromosome number for this genus are advanced.     

Chromosome relationships between Lolium and Festuca (Gramineae)

With a view to eclucidating chromosome relationships between Lolium perenne (Lp), L. multiflorum (Lm) and Festuca pratensis (Fp), chromosome pairing in different diploid (2n=14), auto-allotriploid (2n=3x=21), trispecific (2n=3x=21), amphidiploid (2n=4x=28) and auto-allohexaploid (2n=6x=42) hybrids between them was analysed. At all these levels of ploidy there was very good chiasmate pairing between the chromosomes of the three species and, on the whole, there was little evidence of preferential pairing of the chromosomes of a particular species in the triploid, tetraploid and hexaploid hybrids. A critical test for this also came from the synaptic ability of the chromosomes of the single genome with those of the duplicated genome in the auto-allotriploids which formed predominantly trivalents with 2, 3 or even 4 chiasmata. Moreover, the homology between the Lp and Lm chromosomes seems strong enough to pass the discrimination limits of the B-chromosomes which do not suppress homoeologous pairing in the Lp LmLm triploid and LpLm diploid hybrids. — The triploids having two genomes of a Lolium species and one of F. pratensis had some male and female fertility which suggested genetic compatibility of the parental chromosomes resulting, presumably, in compensation at the gametic level. Also, the occurrence of comparable chiasma frequencies in the auto-allotriploids and trispecific hybrids showed that they were not markedly affected whether two doses of one genome and one of the other or all the three different genomes from the three species were present. From the trend of chromosome pairing in all these hybrids it is concluded that there is little structural differentiation between the chromosomes of the three species, no effective isolation barrier to gene-flow between them, and that they are closely related phylogenetically, having possibly evolved from a common progenitor. Taxonomic revision of the two Lolium species is suggested.     

CHROMOSOMES,HYBRIDS AND PLOIDY OF SEDUM CREMNOPHILA AND ECHEVERIA LINGUIFOLIA (CRASSULACEAE)

Sedum cremnophila and Echeveria linguifolia have generally been placed in different genera on the basis of their flowers—largely because the petals are spreading in one and erect in the other—and the genera have been placed in different subfamilies. However, they are very similar vegetatively and in their unusual inflorescence, their karyotypes are similar (n = 33), and they readily hybridize to produce fertile F₁ hybrids. Study of hybrids of these two species with numerous others leads to the conclusion that each of the two is effectively diploid, with a genome consisting of 33 chromosomes that are all different and that do not pair with each other. Therefore, the good chromosome pairing and the fertility of the hybrid between them are the result of close structural and genetic homology between the corresponding chromosomes of the two species. Taxonomic revision to reflect their very close relationship is desirable. Some other species of Sedum and Echeveria also may need to be reclassified.     

Autosyndetic pairing in Gibasis (Commelinaceae) hybrids revealed by C-banding

Two closely related species of Gibasis, G. karwinskyana and G. consobrina, and their F₁ hybrids were studied cytologically at the diploid and tetraploid level. Despite similarity in their basic karyotype, pairing was extremely limited in the diploid hybrid and almost exclusively autosyndetic in the tetraploid, except for multivalent formation due to interchange heterozygosity. The analysis was considerably facilitated by the use of C-banding techniques at meiosis, by which the chromosomes of each species could be readily identified. In the parents, quadrivalents were formed between homologous but non-identical chromosomes, which also formed autosyndetic bivalents in the hybrids. Meiotic pairing in the hybrids was unaffected by polytypy for C-bands among different populations of the parental species.     

Somatic hybridization between potato and Nicotiana plumbaginifolia

Summary Electrofusion was carried out between mesophyll protoplasts from the transformed diploid S. tuberosum clone 413 (2n=2x=24) which contains various genetic markers (hormone autotrophy, opine synthesis, kanamycin resistance, -glucuronidase activity) and mesophyll protoplasts of a diploid wild-type clone of N. plumbaginifolia (2n=2x=20). Hybrid calli were obtained after continuous culture on selection medium containing kanamycin. Parental chromosome numbers, determined at 2 months after fusion, revealed hybrid-specific differences between the individual calli. On the basis of these differences three categories of hybrids were distinguished. Category I hybrids contained between 8 and 24 potato chromosomes and more than 20 N. plumbaginifolia chromosomes; category II hybrids had between 1 and 20 N. plumbaginifolia chromosomes and more than 24 potato chromosomes; category III hybrids contained diploid or subdiploid numbers of chromosomes from both parents. The hybrids were evenly distributed over the three categories. After a 1-year culture of 24 representative hybrid callus lines on selection medium the karyotype of 10 hybrids remained stable, whereas 8 hybrids showed polyploidization of the genome of one parent, together with no or minor changes of the chromosome numbers of the other parent. Six hybrids showed slight changes in the hybrid karyotype. The elimination of chromosomes of a particular parent was not correlated to their metaphase location. The processes of spontaneous biparental chromosome elimination leading to the production of asymmetric hybrids of different categories are discussed.     

INTERSPECIFIC HYBRIDIZATION IN HAPLOPAPPUS AND ITS BEARING ON CHROMOSOME EVOLUTION IN THE BLEPHARODON SECTION

Jackson , R. C. (U. Kansas, Lawrence.) Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. Jour. Bot. 49 (2) : 119–132. Illus. 1962.—Cytological analyses of interspecific hybrids between H. gracilis (n = 2) and H. spinulosus ssp. australis (n = 8) indicate that ssp. australis is a segmental allotetraploid, derived from past hybridization between 2 taxa with chromosome numbers of n = 4. Analysis of hybrids between H. gracilis (n = 2) and H. ravenii (n = 4), a previously undescribed species, has shown that the chromosome segments of these 2 species are almost completely homologous. Differential contraction is suggested as the explanation for the disappearance in late pachytene of presumed non-homologous segments which were evident in some cells at early pachytene. The pairing relationship of gracilis and ravenii chromosomes at pachytene and later prophase I stages of meiosis indicates that gracilis has evolved from ravenii by an aneuploid reduction process similar to that described for Crepis. The close morphological relationship of the 2 species adds further support to this proposition. Data from the cytological analysis of both interspecific hybrids indicate that x = 4 is the basic chromosome number for the Blepharodon section of Haplopappus.     

Multidisciplinary approach to genome analysis in the diploid species,Thinopyrum bessarabicum and Th. elongatum (Lophopyrum elongatum), of the Triticeae

Summary The J and E genome species of the Triticeae are invaluable sources of salt tolerance. The evidence concerning the phyletic relatedness of the J genome of diploid Thinopyrum bessarabicum and the E genome of diploid Th. elongatum (=Lophopyrum elongatum) is discussed. Low level of chromosome pairing between J and E at different ploidy levels, suppression of J-E pairing by the Ph1 pairing regulator that inhibits homoeologous pairing, complete sterility of the diploid hybrids (JE), karyotypic divergence of the two genomes, differences in total content and distribution of heterochromatin along their chromosomes, and marked differences in gliadin proteins, isozymes, 5S DNA, and rDNA indicate that J and E are distinct genomes. Well-defined biochemical markers have been identified in the two genomes and may be useful in plant breeding. The level of distinction between J and E is comparable to that among the universally accepted homoeologous genomes A, B, and D of wheat. Therefore, the J and E genomes are homoeologous and not homologous, although some workers continue to call them homologous. The previous workers' data on chromosome pairing in diploid hybrids and/ or karyotypic differences in the conventionally stained chromosomes do not provide sufficient evidence for the proposed merger of J and E genomes (and, hence, of the genera Thinopyrum and Lophopyrum) specifically and for establishing genome relationships generally. Extra precautions should be exercised before changing the designation of an established genome and before merging two genera. A uniform, standardized system of genomic nomenclature for the entire Triticeae is proposed, which should benefit cytogeneticists, plant breeders, taxonomists, and evolutionists.Cooperative investigations of the USDA-Agricultural Research Service and the Utah Agricultural Experiment Station, Logan, UT 84322, USA. Approved as Journal Paper no. 3832     

Introgression of chromosomes of Festuca arundinacea var. glaucescens into Lolium multiflorum revealed by genomic in situ hybridisation (GISH)

An F₁ hybrid (n=4x=28) between the tetraploid species Festuca arundinacea var. glaucescens (GGG′G′) and a synthetic tetraploid Lolium multiflorum (LmLmLmLm) was backcrossed to diploid L. multiflorum to produce triploid (2n=3x=21) BC₁ hybrids (LmLmG). At metaphase I of meiosis the triploids had a preponderance of ring bivalents and univalents with some linear and frying-pan trivalents. Genomic in situ hybridisation (GISH) differentiated the Festuca chromosomes from Lolium and revealed that the bivalents were exclusively between Lolium homologues, while the univalents were Festuca. Despite the limited amount of homoeologous chiasmata pairing in the triploids, some recombinant chromosomes were recovered in the second backcross when the hybrids were further crossed to diploid L. multiflorum. The progeny from the second backcross was predominantly diploid. Genotypes with recombinant chromosomes and chromosome additions involving an extra Festuca chromosome were identified using GISH. Changes in plant phenotype were related to the presence of Festuca chromatin. Received: 20 September 2000 / Accepted: 05 January 2001