CYTOTAXONOMY OF TWELVE SPECIES OF HIBISCUS SECTION FURCARIA

Menzel, Margaret Y. (Florida State U., Tallahassee), and F. D. Wilson. Cytotaxonomy of twelve species of Hibiscus section Furcaria. Amer. Jour. Bot. 50(3): 262–271. Illus. 1963.—Metaphase-I chromosome numbers and pairing in 88 accessions showed that H. cannabinus, H. costatus, and H. surattensis are diploid (n = 18); and H. acetosella, H. aculeatus, H. bifurcatus, H.furcellatus, H. meeusei, H. radiatus, H. rostellatus and H. sabdariffa are tetraploid (n = 36), with similar low multivalent frequencies, hence probably allotetraploids each combining 2 well-differentiated genomes. No intraspecific variation in ploidy was found. Fertile, vigorous F₁ hybrids between H.furcellatus and H. bifurcatus showed complete chromosome pairing (n = 36), confirming a close relationship between the parents. Two African strains of H. diversifolius were octoploid (n = 72) with low multivalent frequency and hence probably contain 4 differentiated genomes. At least 4, perhaps 5 or 6, differentiated genome groups are represented in tropical Africa, and at least 2 in the American tropics.     

SOME PIECES OF THE AFRICAN GENOME PUZZLE IN HIBISCUS SECT. FURCARIA (MALVACEAE)

New chromosome counts are reported for four African diploid species of Hibiscus L. sect. Furcaria DC: H. greenwayi Bak. f., H. hiernianus Exell et Mendonça, and H. mechowii Garcke, n = 18; and H. berberidifolius A. Rich., 2n = 36. Chromosome behavior is described in seven new hybrid combinations. Hibiscus greenwayi is shown to carry an A genome. Hibiscus hiernianus and H. mastersianus have similar genomes which are not homologous with that of H. mechowii. New and earlier data are integrated to elucidate genome relationships among 13 taxa–each of the four tetraploid species has a different formula encompassing 2 of 6 mutually nonhomologous genomes, A, B, G, H, X and Y (H. acetosella Welw. ex Hiern = AABB; H. meeusei Exell = AAXX; H. sabdariffa L. = AAYY; H. rostellatus Guill. et Perr. = GGHH). Diploid donors of A, B, G, X and Y are proposed on the basis of plant, flower and pollen morphology. Diploid carriers of A (H. asper Hook. f., H. cannabinus L., H. greenwayi), B (H. surattensis L.) and G (H. sudanensis Hochr.) have been verified cytologically. Cytological confirmation of X (H. hiernianus, H. mastersianus Hiern) and Y (H. mechowii) carriers is incomplete. No putative diploid carrier of H is at hand. Genome affinities of H. berberidifolius are unknown.     

CHROMOSOME PAIRING IN NATURAL FIRST AND ADVANCED GENERATION LIATRIS HYBRIDS

Chromosome pairing was analyzed in natural F₁ and advanced generation hybrids of crosses between Liatris aspera and L. spicata, and L. aspera and L. cylindracea. The status of the hybrids was determined on the basis of morphological and chromatographic criteria. The F₁ hybrids of L. aspera × L. spicata contained pairing irregularities in 30% of their PMC's as compared to 10% in the hybrid segregates. Notably, translocation figures appeared in 7% of the F₁ PMC's as compared to 0.6% in the segregate PMC's. The F₁ hybrids of L. aspera × L. cylindracea contained irregularities in 19% of their PMC's as compared to 7% in the segregates. Translocation figures occurred in 6% of the F₁ PMC's, but only in 0.7% of the segregate PMC's. The incidence of pairing and level of structural heterozygosity was far lower in the hybrid segregates than was anticipated on the basis of F₁ pairing relationships. Thus it would appear that there is selection for structural homozygosity in the hybrid segregates.     

NEW INTERGENOMIC HYBRIDS AMONG SOME AFRICAN DIPLOID SPECIES OF HIBISCUS SECT. FURCARIA

Chromosome pairing was examined at Metaphase 1 in F₁ hybrids between Hibiscus surattensis L. (B genome) and three other diploid species, H. hiernianus Exell, H. mastersianus Hiern, and H. mechowii Garcke. The low level of chromosome association in all three types of hybrids indicated that the pollen parent of each hybrid contributed a genome that was meiotically nonhomologous with B. Earlier studies had shown that H. mastersianus and H. hiernianus have similar genomes that are nonhomologous with the genome of H. mechowii. Thus, these four African diploids with overlapping ranges encompass three different genome groups. Provisional genome designations were assigned as follows: H. hiernianus, X_hie; H. mastersianus, X_mas; H. mechowii, Y_mec.     

Genome relationships between Hordeum procerum (6x) and H. lechleri (6x)

Investigations on the meiotic behaviour of chromosomes in interspecific hybrids (2n=6x=42) between Hordeum lechleri (6x) and H. procerum (6x) and in their component haploids have been utilized to assess the nature of pairing and the extent of genome homology between the two species. In the F₁ hybrids an average of 25 (60%) chromosomes associated at metaphase I, mostly as bivalents. A majority (60%) of the pollen mother cells (PMCs) in H. procerum haploids (2n=3x=21) displayed 21 univalents and even in the remainder, a maximum of two rod bivalents were formed resulting in an average of 0.52 bivalents per cell. In haploids of H. lechleri (2n=3x=21) however, 30% of chromosomes pair. The sum of the chromosomal associations in the component haploids represents only 17% of the complement, far below the observed frequency (60%) in the hybrids. Thus, the pairing displayed in hybrids between H. lechleri and H. procerum was mostly allosyndetic and suggestive of two genomes being common in these species.In haploid H. procerum 1/3 of the PMCs displayed a tripolar organisation of chromosomes leading to triad and hexad formation after divisions I and II respectively. The significance of hexad formation in the trihaploid H. procerum and a possible suppression of homoeologous pairing in H. procerum haploids are discussed.     

Pairing affinities of the B- and G-genome chromosomes of polyploid wheats with those of Aegilops speltoides.

Chromosome pairing at metaphase I was studied in different interspecific hybrids involving Aegilops speltoides (SS) and polyploid wheats Triticum timopheevii (AtAtGG), T. turgidum (AABB), and T. aestivum (AABBDD) to study the relationships between the S, G, and B genomes. Individual chromosomes and their arms were identified by means of C-banding. Pairing between chromosomes of the G and S genomes in T. timopheevii x Ae. speltoides (AtGS) hybrids reached a frequency much higher than pairing between chromosomes of the B and S genomes in T. turgidum x Ae. speltoides (ABS) hybrids and T. aestivum x Ae. speltoides (ABDS) hybrids, and pairing between B- and G-genome chromosomes in T. turgidum x T. timopheevii (AAtBG) hybrids or T. aestivum x T. timopheevii (AAtBGD) hybrids. These results support a higher degree of closeness of the G and S genomes to each other than to the B genome. Such relationships are consistent with independent origins of tetraploid wheats T. turgidum and T. timopheevii and with a more recent formation of the timopheevi lineage.     

Synthesis and cytological characterization of trigeneric hybrids involving durum wheat,Thinopyrum bessarabicum,and Lophopyrum elongatum

Summary In an attempt to transfer genes for salt tolerance and other desirable traits from the diploid wheatgrasses, Thinopyrum bessarabicum (2n=2x=14; JJ genome) and Lophopyrum elongatum (2n=2x=14; EE genome), into durum wheat cv Langdon (2n=4x=28; AABB genomes), trigeneric hybrids with the genomic constitution ABJE were synthesized and cytologically characterized. C-banding analysis of somatic chromosomes of the A, B, J, and E genomes in the same cellular environment revealed distinct banding patterns; each of the 28 chromosomes could be identified. They differed in the total amount of constitutive heterochromatin. Total surface area and C-banded area of each chromosome were calculated. The B genome was the largest in size, followed by the J, A, and E genomes, and its chromosomes were also the most heavily banded. Only 25.8% of the total chromosome complement in 10 ABJE hybrids showed association, with mean arm-pairing frequency (c) values from 0.123 to 0.180 and chiasma frequencies from 3.36 to 5.02 per cell. The overall mean pairing was 0.004 ring IV + 0.046 chain IV + 0.236 III + 0.21 ring II + 2.95 rod II + 20.771. This is total pairing between chromosomes of different genomes, possibly between A and B, A and J, A and E, B and J, B and E, and J and E, in the presence of apparently functional pairing regulator Ph1. Because chromosome pairing in the presence of Ph1 seldom occurs between A and B, or between J and E, it was inferred that pairing between the wheat chromosomes and alien chromosomes occurred. The trigeneric hybrids with two genomes of wheat and one each of Thinopyrum and Lophopyrum should be useful in the production of cytogenetic stocks to facilitate the transfer of alien genes into wheat.     

Intergeneric hybrids between Triticum crassum and Hordeum vulgare

Summary Intergeneric hybrids between Triticum crassum (2n=6x=42) and Hordeum vulgare cv. Bomi were obtained at a frequency of 15% of pollinated florets. Meiotic chromosome pairing in the hybrids was not different from that observed in a polyhaploid of T. crassum indicating negligible pairing between chromosomes of the two species and secondly that the genome of H. vulgare had no effect on intergenomic pairing in T. crassum.Contribution No. 646 Ottawa Research Station     

Genetic relationships in hibiscus sect. Furcaria

The eighteen species studied form an allopolyploid series (x=18). The morphology, crossing behavior, and geographical distribution of 6 diploid, 9 tetraploid, 2 octoploid, and 1 decaploid species were studied. From over 26,500 crosses, 19 hybrid combinations and several derived allopolyploids and three-species hybrids were obtained. Chromosome pairing in the hybrids showed that a minimum of 6 and a maximum of 14 well-differentiated genome groups exist in sect. Furcaia, at least two of which appear to be confined to the Old World. No evidence was found that New World genomes are represented in the Old World. The primary radiation of the diploid genomes probably occurred at about the same time as that of the diploid genomes of Gossypium, whereas the tetraploids and one of the octoploid species (H. furcatus Roxb., non Willd.) seem to be of later origin (late Pleistocene or Recent). Octoploid H. diversifolius Jacq., a circumtropical species, may be a relict of a much earlier round of polyploid evolution.     

Tissue-culture-facilitated production of aneupolyhaploid Thinopyrum ponticum and amphidiploid Hordeum violaceum x H. bogdanii and their uses in phylogenetic studies

Summary An aneupolyhaploid (2n = 36) of the decaploid Thinopyrum ponticum and an amphidiploid (2n = 28) of Hordeum violaceum x Hordeum bogdanii were produced through anther and inflorescence culture, respectively. Meiotic associations in pollen mother cells at metaphase I of these plants were analyzed. The aneupolyhaploid arose by direct embryogenesis from a microspore without passing through a callus phase. The mean pairing frequencies of 2.67 univalents ⁺ 0.54 rod bivalents ⁺ 8.85 ring bivalents ⁺ 2.75 trivalents ⁺ 0.17 chain quadrivalents ⁺ 0.56 ring quadrivalents ⁺ 0.65 pentavalents in the aneupolyhaploid (2n = 36) best fit the 221 model. However, the frequent multivalents (up to five trivalents, or three quadrivalents, or four pentavalents in a cell) indicated that decaploid T. ponticum has five sets of closely related genomes representable by the genome formula J₁ J₁ J₁ J₂ J₂. Colchicine treatment of inflorescence-derived H. violaceum x H. bogdanii regenerants greatly enhanced the rate of chromosome doubling, and completely doubled regenerants could be isolated. The H. violaceum x H. bogdanii amphidiploid had a mean pairing pattern of 12.53 univalents ⁺ 5.57 rod bivalents ⁺ 1.97 ring bivalents ⁺ 0.07 trivalents ⁺ 0.03 hexavalents, indicating the presence of desynaptic gene(s) in the original diploiid hybrid. Therefore, the pairing frequency in that diploid hybrid was an under-estimate of chromosome homology between the parental genomes, and additional diploid hybrids are needed to assess the genome homology between H. violaceum and H. bogdanii. These two contrasting experiments demonstrated that tissue culture techniques are useful in altering the ploidy level to produce plant materials suitable for genome analysis and phylogenetic studies.Cooperative investigation of the USDA-ARS, Forage and Range Research Laboratory, Logan, UT 84322-6300, and the Utah Agricultural Experiment Station, Utah State University, Logan, UT 84322-4810. Approved as journal paper No. 3991     

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.     

Clarification of the status of Hibiscus (sect. Furcaria) uncinellus (Malvaceae)

Hibiscus uncinellus andH. bifurcatus have been confused in the literature and in the herbarium. The morphological, ecological, and geographic differences between them are presented, and a lectotype is chosen for the former species.Hibiscus uncinellus is virtually confined to Mexico (plus one Guatemalan station), andH. bifurcatus occurs in the West Indies, South America, and Central America, as far north as Honduras.     

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.     

The origin and meiotic behaviour of hexaploid northern wheatgrass (Agropyron dasystachyum)

The occurrence of hexaploid (2n = 6x = 42) forms in some otherwise natural tetraploid populations of Agropyron dasystachyum (2n = 4x = 28) was cytologically detected and studied. The hexaploid plants are morphologically similar to the tetraploids except for a small reduction in the anther size. The general survey of chromosome numbers of natural Northern Wheatgrass (A. dasystachyum 2n = 4x = 28) populations derived from eight different regions of Alberta indicated that the occurrence of hexaploid variants was not restricted to a single locality. A comparative study of chromosome pairing in the natural and the synthetic hexaploids revealed that the naturally occurring 42-chromosomed plants of A. dasystachyum originated as a result of fertilization between unreduced (SSHH) and the natural (SH) gametes, both coming from the tetraploid form of A. dasystachyum. Based on chromosome pairing, the genomes of the natural hexaploid A. dasystachyum have been designated as SSSHHH. The natural hexaploids appear to intercross among themselves and also with tetraploids producing euploid and aneuploid hybrids. The possible evolutionary significance of these findings is briefly discussed.     

HYBRIDIZATION IN THE GENUS GLYCINE SUBGENUS GLYCINE WILLD. (LEGUMINOSAE,PAPILIONOIDEAE)

The genus Glycine is composed of two subgenera, Glycine and Soja. Soja includes the cultivated soybean, G. max, and its wild annual counterpart G. soja, while Glycine includes seven wild perennial species. Hybridization was carried out within and between wild perennial species of the subgenus Glycine. The success rate (pods set/flowers crossed) was 11% for intraspecific and 8% for interspecific crosses. A total of 220 F₁ hybrids was examined morphologically and cytologically where possible. Hybrids within G. canescens (2n = 40) and G. latifolia (2n = 40) were fertile as expected. Glycine clandestina (2n = 40) was morphologically separable into at least three groups, which produced fertile hybrids within each group. One cross between two groups gave vegetatively vigorous but sterile hybrids. The majority of crosses within G. tabacina (2n = 80) were fertile, except that extremely narrow-leaved forms gave sterile hybrids in combination with more usual forms. Sterility was also encountered in G. tomentella when aneuploids (2n = 78) from New South Wales, Australia, were crossed with tetraploids (2n = 80) from either Queensland, Australia, or Taiwan; crosses between the latter two populations resulted in seedling lethality. Cytological behavior of sterile hybrids followed a similar pattern, whether at the diploid or tetraploid level. The frequency of chromosome pairing was approximately half that expected if genomes showed full pairing homology. Bivalent disjunction at anaphase I was usually followed by precocious division of the majority of univalents. Telophase I and II were characterized by lagging chromosomes and micronuclei, so that resulting pollen was misshapen and sterile. Chromosome pairing data from sterile intraspecific hybrids at the tetraploid level may indicate a polyphyletic origin of tetraploids, whereby different diploid populations were involved in their formation. Similarly, chromosome pairing in sterile intraspecific diploid hybrids may indicate that the various diploid groups arose independently of one another. Both 40- and 80-chromosome forms are fully diploidized, however, and if they are of ancient origin, divergence since that time could have resulted in the chromosomal differentiation which becomes apparent when intraspecific hybridization is effected. Diploid (2n = 40) interspecific hybrids G. falcata × G. canescens, and G. falcata × G. tomentella grew poorly and did not reach flowering stage. Diploid (2n = 40) crosses between G. latifolia and G. tomentella produced inviable seedlings. Tetraploid (2n = 80) hybrids between G. tomentella and G. tabacina were vegetatively vigorous but sterile owing to low chromosome pairing at meiosis, indicating little pairing homology between the two species. Diploid hybrids between G. canescens and G. clandestina, however, showed almost complete chromosome pairing at diakinesis and partial fertility. Although morphologically distinct, these two species have not diverged sufficiently to prevent hybridization and possible gene exchange through recombination. Self compatibility, perennial growth habit, and geographic isolation have favored divergence among Glycine populations to the point that gene exchange appears no longer possible in many cases. Internal isolating mechanisms have been shown to operate at various levels of plant development from hybrid lethality at seedling stage, to failure of seed-set in sterile but vegetatively vigorous hybrids.     

Elimination and duplication of particular Hordeum vulgare chromosomes in aneuploid interspecific Hordeum hybrids

Summary Seeds formed in crosses Hordeum lechleri (6x) x H. vulgare (2x and 4x), H. arizonicum (6x) x H. v. (2x), H. parodii (6x) x H. v. (2x), and H. tetraploidum (4x) x H. v. (2x) produced plants at high or rather high frequencies through embryo rescue. Giemsa C-banding patterns were used to analyze chromosomal constitutions and chromosomal locations on the methaphase plate. Among 100 plants obtained from H. vulgare (2x) crosses, 32 plants were aneuploid with 2n=29 (1), 28 (3), 27 (13), 26 (5), 25 (4), 24 (4), or 22 (2); 50 were euploid (12 analyzed), and 18 were polyhaploid (5 analyzed). Four plants had two sectors differing in chromosome number. Two of four hybrids with H. vulgare (4x) were euploid and two were aneuploid. Parental genomes were concentrically arranged with that of H. vulgare always found closest to the metaphase centre. Many plants showed a certain level of intraplant variation in chromosome numbers. Except for one H. vulgare (4x) hybrids, this variation was restricted to peripherally located non-H. vulgare genomes. This may reflect a less firm attachment of the chromosomes from these genomes to the spindle. Interplant variation in chromosome numbers was due to the permanent elimination or, far less common, duplication of the centrally located H. vulgare chromosomes in all 34 aneuploids, and in a few also to loss/gain of non-H, vulgare chromosomes. This selective elimination of chromosomes of the centrally located genome contrasts conditions found in diploid interspecific hybrids, which eliminate the peripherally located genome. The difference is attributed to changed genomic ratios. Derivatives of various H. vulgare lines were differently distributed among euploid hybrids, aneuploids, and polyhaploids. Chromosomal constitutions of hypoploid hybrids revealed a preferential elimination of H. vulgare chromosomes 1, 5, 6, and 7, but did not support the idea that H. vulgare chromosomes should be lost in a specific order. H. vulgare SAT-chromosomes 6 and 7 showed nucleolar dominance. Aneuploidy is ascribed to the same chromosome elimination mechanism that produces haploids in cross-combinations with H. vulgare (2x). The findings have implications for the utilization of interspecific Hordeum hybrids.     

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.     

Chromosome elimination and chromosome pairing in tetraploid hybrids of Hordeum vulgare × H. bulbosum

Summary The C₀ tetraploid counterparts of diploid hybrids of Hordeum vulgare × H. bulbosum were meiotically analysed, and were found to be chromosomally less stable than the same genotypes had been as diploids. The 14 bulbosum chromosomes present in the tetraploid cytotypes were probably eliminated as pairs rather than randomly or one genome at the time. Development of the vulgare and bulbosum genomes was asynchronous in some hybrids, the bulbosum chromosomes appearing less advanced than the vulgare chromosomes in the same cell. This appeared to reduce pairing between bulbosum homologues and also suppressed homoeologous pairing.     

THE GENOME CONSTITUTION AND PHYLOGENY OF ELYMUS AMBIGUUS

Two Elymus ambiguus Vasey & Scribn. collections from Utah and Idaho were 2n = 28, and the species behaved meiotically as an allotetraploid. The E. ambiguus plants were highly self-sterile, and they hybridized readily with Asian E. junceus Fisch. (2n = 14), E. karataviensis Roshev. (2n = 28), E. multicaulis Kar. & Kir. (2n = 28), and North American E. innovatus Beal (2n = 28). Chromosome pairing at metaphase-I in the E. ambiguus X E. junceus triploid hybrids indicated that one E. ambiguus genome was closely homologous with the E. junceus genome. Chromosome pairing in the tetraploid hybrids indicated that both E. ambiguus genomes were more or less homologous with the genomes of E. karataviensis, E. multicaulis, and E. innovatus. The basic genome formula of E. ambiguus may be written as JJXX, where J is the E. junceus genome and X is a genome of unknown origin. Chromosome pairing in the hybrids indicated that E. ambiguus is more closely related to North American E. innovatus than to the Asian species. The E. ambiguus X E. innovatus hybrids were the only hybrids that set seed. Gene flow between E. ambiguus and E. innovatus is biologically possible, but geographic separation of the species precludes natural introgression.     

A novel intergeneric hybrid in the Triticeae: Triticum aestivum x Psathyrost achys juncea

Summary Two hybrid embryos of intergeneric origin between Triticum aestivum cv Fukuho (2n=6x=42, AABBDD) and Psathyrostachys juncea (2n=2x=14, NN) were successfully rescued. One hybrid plant had the expected chromosome number of 28 (ABDN), whereas the second plant had 35 chromosomes. The average meiotic chromosome pairing in the 35-chromosome hybrid was 21.87 univalents + 6.38 bivalents + 0.11 trivalents + 0.009 quadrivalents, which indicates that two copies of the N genome were present. Chromosome pairing in the 28-chromosome hybrid was low (1.35 bivalents), and pointed out the lack of homology between the wheat genomes and the P. juncea genome. These new hybrids showed some necrosis and chlorosis, which caused severe floral abortion in the plant that had 35 chromosomes. These problems became gradually less severe after 18 months.Contrib. no. 372