THE PROBLEM OF PLOIDY IN ECHEVERIA (CRASSULACEAE) I. DIPLOIDY IN E. CILIATA

The largely Mexican genus Echeveria is characterized by an extensive series of dysploid chromosome numbers, with every gametic number from 12 to 34 known in at least one species. Within this nearly three-fold range of numbers, the boundary between diploidy and tetraploidy is not immediately apparent. However, species of Echeveria can be hybridized in an extraordinary number of combinations, both among themselves and with related genera, and study of the morphology of the hybrids and the pairing of their chromosomes provides information that helps to identify the ploidy of the parents. This paper reports observations from study of 80 hybrids between E. ciliata (n = 25) and 73 other species and/or cytotypes. Hybrids between E. ciliata and definite diploids are all nicely intermediate morphologically, whatever the chromosome numbers. In these same hybrids, most chromosomes become involved in pairing at meiosis, and the number of paired elements (bivalents and multivalents) approaches or equals, but never exceeds, the number of chromosomes received from the lower-numbered parent. In most cells, relatively few univalents are present, sometimes none. These observations are considered to indicate that all paired elements include at least one chromosome from each parent and therefore that pairing occurs between chromosomes of different parents only (allosyndesis). Since none of the 25 gametic chromosomes of E. ciliata is able to pair with any other, although they do pair very extensively with chromosomes from many other species having a wide range of numbers, E. ciliata is considered to be diploid in spite of its relatively high chromosome number. On the other hand, hybrids of E. ciliata with definite polyploids resemble the latter much more closely in their morphology, and at meiosis most or all pairing occurs by autosyndesis between chromosomes received from the polyploid parent, while the chromosomes from E. ciliata generally remain unpaired. In these respects most, but not all, species of Echeveria having as many as 34 gametic chromosomes have the same properties as E. ciliata and also are considered to be diploid. The ancestral chromosome number in the genus is not clear, but it is probably near the upper end of the series of dysploid numbers.     

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.     

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 CHROMOSOMES OF PACHYPHYTUM (CRASSULACEAE)

Eleven of the 12 species of Pachyphytum, all that are available, have n = 31–33 standard chromosomes, or a multiple. Accessory chromosomes were found in some or all collections of four species; some cells of one plant have more than 50 of them. Accessory chromosomes often occur in groups at metaphase I, corresponding to their origin from one to several chromocenters of prophase I. Intraspecific polyploidy occurs within five species, with diploids to 12-ploids (n = ca. 186) in P. compactum and diploids to decaploids (n = ca. 160) in P. hookeri. Although the basic chromosome number is high, evidence from meiosis in certain hybrids shows that the basic 31–33 chromosomes are probably all different: they do not pair with each other and they do not duplicate each other. Polyploids, with 62 or more chromosomes, are probably autopolyploids: they form multivalents, and the chromosomes they contribute to hybrids pair with each other. Three different probable hybrids have been found in the wild, and more than 300 hybrids have been produced in cultivation.     

CHROMOSOME RELATIONSHIPS OF DIPLOID SPECIES OF GRINDELIA (COMPOSITAE) FROM COLORADO,NEW MEXICO,AND ADJACENT AREAS

Previous studies of chromosome relationships of Grindelia species recognized three basic genomes designated Oxylepis, Hallii, and Havardii. Differences are based on different end arrangements of the chromosomes resulting from reciprocal translocations. This report will review and give additional information about the genomes and interrelationships of 17 species. All of the species are diploids (2n = 12) and show six bivalents at meiosis. Species in this study that have the Oxylepis genome are G. oxylepis var. eligulata, G. fastigiata, G. inornata, G. revoluta, and G. squarrosa. Species that have the Havardii genome included G. havardii, G. grandiflora, G. lanceolata, G. littoralis, and G. texana. The Hallii genome is present in G. camporum var. davyi and G. procera. Hybrids of species with the same genome have six bivalents at meiosis. Hybrids between species with the Oxylepis genome and those having the Havardii genome have four bivalents and one quadrivalent at meiosis. Likewise for Oxylepis x Hallii hybrids. A new genome is presented for G. subalpina which would explain the configurations of two bivalents and two quadrivalents observed in G. subalpina x G. havardii and G. subalpina x G. fastigiata hybrids. This is designated the Subalpina genome. Species tested but with genomes as yet undetermined are G. acutifolia, G. arizonica, G. nana, and G. scabra.     

A CYTOGENETIC ANALYSIS OF CERTAIN POLYPLOIDS IN GRINDELIA (COMPOSITAE)

Two diploid taxa, Grindelia procera and G. camporum, and 3 tetraploid ones, G. camporum, G. hirsutula, and G. stricta, have been studied to ascertain their interrelationships. Meiosis in diploid parental strains was regular, the common chromosome configuration being 5 rod bivalents and 1 ring bivalent. The average chiasmata frequency per chromosome was 0.60. Pollen fertility was about 90% in all strains examined. Diploid interspecific hybrids had normal meiosis with an average chiasmata frequency of 0.56 per chromosome. No heterozygosity for inversions or interchanges was detected, and pollen fertility was above 85%. Meiosis in parental tetraploid strains was characterized by the presence of quadrivalents in addition to a complementary number of bivalents. The average chiasmata frequency per chromosome was 0.59 and pollen fertility was generally about 80%. Tetraploid interspecific hybrids also had quadrivalents, normal meiosis, and high pollen fertility. Close genetic relationships between the diploids and between the tetraploids are indicated, and geographical, ecological, and seasonal barriers to gene exchange exist. Attempts to obtain hybrids between diploids and tetraploids were successful in a few cases. The hybrids were tetraploid and had normal meiosis and fertility similar to parental and F₁ tetraploids. Their origin was by the union of unreduced gametes of the diploid female parent and normal pollen from the tetraploid parent. On the basis of chromosome homology, normal meiosis, plus high fertility exhibited in the diploid, tetraploid, and diploid X tetraploid interspecific hybrids, these species of Grindelia are considered to be a part of an autopolyploid complex. Gene exchange between diploids and diploids, tetraploids and tetraploids, and diploids and tetraploids is possible. Tetraploid G. camporum may have originated by hybridization between G. procera and diploid G. camporum with subsequent doubling of chromosomes and selection for the combined characteristics of the diploids.     

Synaptonemal complex formation in hybrids of Lolium temulentum x Lolium perenne (L.)

Comparisons were made between two kinds of tetraploids derived from the hybrid Lolium temulentum x L. perenne. One hybrid behaves like an autotetraploid with multivalents at first metaphase of meiosis in pollen mother cells. The other behaves like an allotetraploid, in which pairing at first metaphase is restricted to bivalents comprised of strictly homologous chromosomes. The diploidisation of the latter form is controlled by determinants located on both the normal, A chromosomes and on supernumary B chromosomes. Reconstruction of synaptonemal complexes and their elements, from serial sections through pollen mother cell nuclei examined under the electron microscope, reveals that at zygotene pairing in both forms results in multivalent formation involving non-homologous as well as homologous chromosomes. The mechanism responsible for the diploidisation is, therefore, not based on a restriction of pairing at early meiosis to homologous chromosomes but on a correction or transformation of the multivalent chromosome associations to bivalents subsequent to zygotene. The transformation is not completed until late pachytene. In the multivalent-forming tetraploid a maximum of four chromosomes are associated at first metaphase. Yet configurations of a higher valency are found at zygotene. There is, therefore, a partial transformation of multivalents even in this autotetraploid form which restricts configurations at metaphase I to homologous and homoeologous chromosomes only. In both hybrids some homologous bivalents are not the product of resolution of multivalents but result from two-by-two pairing from the beginning of zygotene.     

The fate of multivalents during meiotic prophase in the hybrid Gibasis consobrina x G. karwinskyana Rafin. (Commelinaceae)

The chromosomes of the two closely related diploid species, Gibasis consobrina and G. karwinskyana (Commelinaceae; 2n=2x=10), are morphologically alike, yet form few chiasmate associations at metaphase I in the f₁ hybrid. During meiotic prophase, however, synaptonemal complexes join the majority of the chromosomes of the complement in complex multiple pairing configurations. The F₁ hybrid between different tetraploid genotypes of the same two species similarly forms multivalents during meiotic prophase, which are subsequently eliminated in favour of strictly homologous bivalents before metaphase I. One quadrivalent comprising interchange chromosomes inherited from one of the parents, usually persists to first metaphase. Evidently the resolution of multivalents to bivalents at first metaphase, which accounts for diploidisation, is not attributable to the elimination of multivalents per se, but of multivalents comprising chromosomes of limited homology.     

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.     

SEMISTERILITY IN THE INTERSPECIFIC HYBRID MELILOTUS POLONICA × M. ALBA

Jaranowski , J. K. (Coll. of Agriculture, ul. Wojska Polskiego 71c, Poznan, Poland.) Semisterility in the interspecific hybrid Melilotus polonica × M. alba. Amer. Jour. Bot. 48(1): 28–35. Illus. 1961.—Interspecific hybrids between Melilotus polonica (n = 8) and M. alba (n = 8) are readily secured. The F₁ hybrids are intermediate between the parents and partially sterile with a mean percentage of 58.8 (ranging from 46.8 to 72.6) defective pollen grains. Six bivalents and a chain or ring of 4 chromosomes occur at diakinesis and metaphase-I of microsporogenesis. A crossshaped configuration characteristic of a reciprocal translocation is present at pachytene, indicating that one of the parents is homozygous for an interchange of relatively large section between two of the members of the chromosome complex. Chromosome bridges, lagging chromosomes, movement of the univalents to the same pole and precocious division of the univalents lead to aberrant chromosome distribution during the course of meiosis. Reduction in self-fertility indicates a corresponding aberrant distribution of chromosomes during megasporogenesis. Pollen sterility in the F₂ generation ranged from 24.8% to 72.5% with a mean value of 54.6%. Two plants in the F₂ generation which had relatively low pollen sterility proved to be aneuploids (2n + 1). Meiotic irregularities in the F₂ plants were comparable to those exhibited by the F₁ plants.     

Autogamy and hybridization as evolutionary mechanisms in Panicum subgenus Dicanthelium (Gramineae)

A predominantly autogamous breeding system is described forP. occidentale Scribn.,P. pacificum Hitchc. & Chase, andP. thermale Boland. Five synthetic hybrids and three generations of progeny are also described for crosses among these three species. All parents haven = 9 chromosomes and normal meiosis. Hybrids exhibit various irregularities during meiosis, but nine bivalents are usually formed. One population ofP. occidentale and one ofP. thermale were found to have chromosomes with interchanges larger than those of other populations. The F₂ and F₃ generations had markedly increased fertility, as determined by pollen stainability. It is proposed that autogamy and hybridization are a common means of maintaining and stabilizing genetic variability in these grasses and that these means account for much of the taxonomic difficulty posed by the subgenus. The possibility of a low rate of introgression between populations is also proposed.     

CYTOGENETICS OF AGROPYRON FERGANENSE AND ITS HYBRIDS WITH SIX SPECIES OF AGROPYRON,ELYMUS, AND SITANION

Meiosis and mode of reproduction are described in Agropyron ferganense Drob., a perennial forage grass from Central Asia. This species is diploid (2n = 14); it exhibits normal meiosis and reproduces by cross-pollination. Hybrids were produced between A. ferganense and six species with known genome formulas: 1) North American A. spicatum (Pursh) Scribn. & Smith, an SS diploid (2n = 14), 2) Middle Eastern A. libanoticum Hack., an SS diploid (2n = 14), 3) North American A. dasystachyum (Hook.) Scribn., an SSHH tetraploid (2n = 28), 4) Eurasian A. caninum (L.) Beauv., an SSHH tetraploid (2n = 28), 5) North American Sitation hystrix (Nutt.) J. G. Smith, an SSHH tetraploid (2n = 28), and 6) South American Elymus patagonicus Speg., an SSHHHH hexaploid (2n = 42). Almost complete chromosome pairing in the A. ferganense x A. spicatum and A. libanoticum hybrids demonstrated that A. fergenanse is an SS diploid, but it is genetically isolated from the other SS diploids because of high sterility in the F₁ hybrids. S-genome diploids form a network of species that extend from the Middle East through Central Asia to western North America. Frequent occurrence of seven univalents and seven bivalents at metaphase I in the triploid hybrids of A. ferganense x A. dasystachyum, A. caninum and S. hystrix was consistent with the proposed genome formulas of SS for A. ferganense, SSHH for the three tetraploid species, and SSH for the hybrids. Chromosome pairing was highly variable in the A. ferganense x E. patagonicus hybrids; however, some cells had almost complete bivalent pairing, an expected observation in an SSHH hybrid from a cross between an SS diploid (A. ferganense) and an SSHHHH hexaploid (E. patagonicus). Various options were considered concerning the appropriate generic classification of the S-genome diploids, which are now commonly placed in Agropyron. The inclusion of these species in the genus Eiytrigia, as advocated by some Soviet taxonomists, appears to be a reasonable decision.     

CHROMOSOME DIFFERENTIATION IN SEVERAL SPECIES OF MELILOTUS

Shastry , Sishta V. S., William K. Smith , and Delmer C. Cooper . (U. Wisconsin, Madison.) Chromosome differentiation in several species of Melilotus. Amer. Jour. Bot. 47(8) : 613–621. Illus. 1960.—Two species of the section Eumelilotus (M. alba and M. officinalis), 2 of Micromelilotus (M. messanensis and M. segetalis), 2 F₁ hybrids (M. officinalis × M. alba and M. messanensis × M. segetalis), 2 autotetraploids (M. alba and M. officinalis), and 1 allotetraploid (M. officinalis × M. alba), were utilized during the course of this investigation. The 4 species and F₁ hybrids have 16 somatic chromosomes and the tetraploids have twice that number (32). The 2 Eumelilotus species are completely isolated because of seed failure after cross pollination. The F₁ hybrid (M. officinalis × M. alba), obtained elsewhere by the use of embryo-culture techniques, was intermediate between the parents in certain morphological characters and was ca. 75% pollenfertile whereas the parents approached complete fertility. No structural differences between the chromosomes were evident at pachytene. Disturbances which led to the reduced fertility occurred at later stages of meiosis. The Micromelilotus species are cross compatible, but the F₁ hybrid (M. messanensis × M. segetalis) is highly sterile. Despite chromosome structural differences of various types evident at pachytene, bivalents regularly occur at metaphase I. Irregular distribution of the chromosomes at later stages of meiosis leads to high sterility. Species which readily cross but produce a hybrid of very low fertility are likely to compound chromosomal structural differences, because of abnormalities in meiosis, in contrast with species that are completely incompatible.     

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.     

Microsporogenesis in some Triploid Dactylorchid Hybrids

During meiosis in naturally occurring triploid hybrids betweenthe diploid Orchis fuchsii Druce (2n = 4O) and the two tetraploids,O. purpurella Steph. and O. praetermissa Druce (2n = 8O), thereis a regular formation of 20 bivalents and 20 bivalents. Sincethe two tetraploid species themselves show typical ‘diploid’behaviour in synapsis and fertility, they are considered tobe allopolyploids, and the hybrid pairing to be allosyndetic.The implication is therefore that both tetraploids are amphidiploidsof which O. fuchsii has been one progenitor. It is suggestedthat varieties of the polytypic diploid O. latifolia L. sec.Pugsl. may have been the other progenitors. A feature of interestin the microsporogenesis of both parents and hybrids is theclose synchronization of nuclear events in the pollen massulae,which behave as physiological units throughout meiosis and pollen-mitosis.In the triploids, although numerous dysploid nuclei are produced,none dies prematurely, probably because of mutual compensationwithin what is, in effect, a common cytoplasmic matrix.     

NATURAL HYBRIDS BETWEEN ORYZOPSIS AND STIPA. III. ORYZOPSIS HYMENOIDES × STIPA PINETORUM

Johnson, B. Lennart. (U. California, Los Angeles.) Natural hybrids between Oryzopsis and Stipa. III. Oryzopsis hymenoides × Stipa pinetorum. Amer. Jour. Bot. 50(3): 228–234. Illus. 1963.—On the basis of morphological characters, the spontaneous hybrid Oryzopsis hymenoides × Stipa pinetorum is included in 0. bloomeri which consists of a number of sterile, natural hybrids between O. hymenoides (2n = 48) and various American species of Stipa (2n = 32-82). While intermediate between its parents in many attributes, the pinetorum hybrid is different from the other hybrids included in O. bloomeri with respect to lemma-hair length and other characters which are diagnostic for S. pinetorum (2n = 32). The hybrid has 2n = 40 chromosomes, but some plants of the hybrid and some of O. hymenoides had small supernumerary, somatic chromosomes. The parents formed only bivalents at meiosis. The hybrid showed some affinity among 14 chromosomes per sporocyte. This affinity is nearly as great as that shown by other Stiporyzopsis hybrids with 56 or 58 chromosomes, and is consistent with the earlier suggestion that bivalent formation in polyploid species of Stipa may be gene-controlled.     

Cytogenetic analysis of Lycopersicon esculentum (+) Solanum etuberosum somatic hybrids and their androgenetic regenerants

The aim of the study was to characterize genomic relationships among cultivated tomato (Lycopersicon esculentum Mill.) (2n=2x=24) and diploid (2n=2x=24) non-tuberous wild Solanum species (S. etuberosum Lindl.). Using genomic in situ hybridization (GISH) of mitotic and meiotic chromosomes, we analyzed intergeneric somatic hybrids between tomato and S. etuberosum. Of the five somatic hybrids, two plants were amphidiploids (2n=4x=48) mostly forming intragenomic bivalents in their microsporocytes, with a very low frequency of multivalents involving the chromosomes of tomato and S. etuberosum (less than 0.2 per meiocyte). Tomato chromosomes showed preferential elimination during subsequent meiotic divisions of the amphidiploids. Transmission of the parental chromosomes into microspores was also evaluated by GISH analysis of androgenic plants produced by direct embryogenesis from the amphidiploid somatic hybrids. Of the four androgenic regenerants, three were diploids (2n=2x=24 or 2n=2x+1=25) derived from reduced male gametes of the somatic hybrids, and one plant was a hypertetraploid (2n=4x+4=52). GISH revealed that each anther-derived plant had a unique chromosome composition. The prospects for introgression of desirable traits from S. etuberosum into the gene pool of cultivated tomato are discussed. Received: 2 August 2000 / Accepted: 4 December 2000     

Meiosis in gray voles of the subgenus <Emphasis Type="Italic">Microtus</Emphasis> (Rodentia,Arvicolinae) and in their hybrids

The results of light and electron microscopic (EM) studies of meiosis in Microtus arvalis males of the karyoform “arvalis” (2n = 46, NFa = 80), in hybrids between the chromosomal forms arvalis and obscurus (2n = 46, NFa = 68), in M. rossiaemeridionalis voles (2n = 54, NFa = 54), and in a hybrid between the species M. rossiaemeridionalis and kermanensis (2n = 54, NFa = 54) are presented. SC (synaptonemal complex) karyotypes of the parental forms and the hybrids were constructed on the basis of measurements of the length of autosomal SCs revealed by the EM analysis in spermatocytes at the stage of middle pachytene. The SC karyotypes of M. arvalis and the hybrids ♀ obscurus × ♂ arvalis consist of 22 synaptonemal complexes of autosomal bivalents and the axial elements of the synaptonemal complexes of the sex chromosomes X and Y. The SC karyotypes of M. rossiaemeridionalis and the hybrid M. rossiaemeridionalis × M. kermanensis consist of 26 synaptonemal complexes of autosomal bivalents and a sex bivalent; they differ only in the length of the Y chromosome axis (Y chromosome in the hybrid was inherited from M. kermanensis). Asynaptic configurations of the autosomal SCs were not observed in the hybrids. The SC axial elements of the X and Y chromosomes in the parental forms and in the hybrids were located close to each other throughout pachytene, but they did not form a synaptic region. The normal synapsis in sterile hybrids (M. rossiaemeridionalis × M. kermanensis) and the behavior of the sex chromosomes in meiosis in fertile and sterile hybrids are discussed in the context of specific features of meiosis and reproductive isolation.     

Cytological studies on F1 hybrids of Solanum integrifolium with S. melongena and S. melongena var. insanum

S. integrifolium (2n = 24) can easily be crossed as the pistillate parent with S. melongena (2n = 24) and S. melongena var. insanum (2n = 24). However, crosses in the other direction do not succeed. Both hybrids are vigorous. Chromosome association at diakinesis and metaphase I was studied. Chromosome associations higher than bivalents were observed in the hybrids indicating structural repatterning of chromosomes. The modal chromosome association in hybrids was twelve bivalents per PMC. This is suggestive of the retention of ancestral chromosome homeologies by the taxa concerned. Despite regular meiosis both hybrids were highly pollen-sterile (about 95%), which was attributed to segregational events of the recombined chromosomes.     

The genetics of meiotic chromosome pairing in Lolium temulentum x Lolium perenne tetraploids

Summary The degree of preferential pairing of homologous chromosomes was estimated in a series of tetraploid hybrids of Lolium temulentum x Lolium perenne by means of cytological and genetic analyses. The correlations between the frequency of bivalents at first metaphase of meiosis in the hybrid tetraploids and the degree of preferential pairing calculated from the segregation pattern of isozyme alleles in a test cross was extremely high. The results showed clearly that suppression of heterogenetic pairing in these Lolium tetraploids is achieved by a genetic system involving the A chromosomes as well as the B chromosome system which has been known for some time. Certain similarities with the genetic system controlling pairing in polyploid wheats are discussed.