SYNTHETIC HYBRIDS OF NEW WORLD AND OLD WORLD AGROPYRONS II. AGROPYRON RIPARIUM X AGROPYRON REPENS

Five hybrids were obtained from 12 seeds formed in 26 emasculated florets of A. riparium pollinated by A. repens. The hybrid plants were morphologically intermediate between the parents for vegetative and spike characteristics, although they resembled A. repens more closely than A. riparium. The 28-chromosome A. riparium parent behaved cytologically as an allotetraploid and formed an average of 13.98 II and 0.04 I in 94 cells at metaphase I. An average of 20.27 II and 0.36 IV were observed at metaphase I in 55 cells of A. repens, which was designated as a segmental autoallohexaploid. The hybrids contained 35 chromosomes and averaged 6.75 I, 12.49 II, 1.05 III, 0.01 IV, and 0.01 V in 162 cells interpreted at metaphase I. Bivalent chromosome pairing in the hybrids was attributed to autosyndetic pairing of 2 A. repens genomes and allosyndetic pairing between 1 A. riparium genome and 1 A. repens genome. Multivalent chromosome associations were attributed to structural hybridity. A. repens and A. riparium apparently share a genome in common, and this genome is the one responsible for rhizomes in both species. A. riparium was given a genome formula of R₂R₂SS; whereas the A. repens genome formula was written as R₁R₁X₁X₁X₂X₂, and the hybrid genome formula was designated as R₁R₂X₁X₂S. The “S” genome of A. riparium was derived from A. spicatum, and the “R” genome is the genome shared by A. repens and A. riparium. The origin and distribution of the so-called “X” genomes of A. repens remain unknown. The hybrids produced from 3 to 10% stainable pollen; however, no seed was set on the hybrids during 2 years in the field.     

THE ORIGIN OF AGROPYRON LEPTOURUM

Colchicine-induced amphiploids, 2n = 42, of diploid Agropyron libanoticum Hack., 2n = 14, X tetraploid A. caninum (L.) Beauv., 2n = 28, were morphologically and cytologically similar to A. leptourum (Nevski) Grossh., 2n = 42. Both A. leptourum and the induced amphiploids were self-fertilizing. The induced amphiploids crossed readily with A. leptourum and gave rise to partially fertile hexaploid hybrids. Chromosome pairing in the hybrids averaged 0.60^I, 18.29^II, 0.36^III, 0.58^,v, 0.02^v, and 0.22^VI in 90 diakinesis or metaphase-I cells. The genomes of the induced amphiploids are essentially homologous with those of A. leptourum except for two or more reciprocal translocations. The morphological, cytological, fertility, and crossing data provide conclusive evidence that A. libanoticum and A. caninum, or their close relatives, are the parents of A. leptourum. The genome formulas of A. libanoticum, A. caninum, and A. leptourum may be written as SS, S_xS_xH_xH_x, and SSS_xS_xH_xH_x, respectively, where S is the basic A. libanoticum genome and H is a genome derived from Hordeum.     

GENOME RELATIONS AMONG ELYMUS CANADENSIS,ELYMUS TRITICOIDES,ELYMUS DASYSTACHYS,AND AGROPYRON SMITHII

Hand-emasculated Elymus canadensis pollinated by E. triticoides, E. dasystachys, and Agropyron smithii yielded 15, 21, and 1 viable hybrid seeds from 56, 52, and 52 florets, respectively. The 28-chromosome species—E. canadensis, E. triticoides, and E. dasystachys—behaved meiotically as allotetraploids and consistently formed 14 bivalents at metaphase I. Octoploid A. smithii, 2n = 56, averaged 0.41^I, 27.72^II, and 0.03^IV in 87 metaphase-I cells. Agropyron smithii is apparently an allooctoploid or a segmental autoallooctoploid. Meiosis was similar in the E. canadensis X E. triticoides and E. dasystachys hybrids. Chromosome pairing was very low in both hybrids, about two loosely connected open-ended bivalents per cell, and may not represent genuine homologies. The genomes of E. canadensis are distinctly different from those of E. triticoides and E. dasystachys. The E. canadensis X A. smithii hybrid averaged 13.37^I and 14.31^II in 76 metaphase-I cells. More than half of the bivalents were closed at both ends. Inability to distinguish between auto- and allosyndesis resulted in two interpretations of genome relations. Either A. smithii is an alloploid with two of its four genomes similar to those of E. canadensis, or it is a segmental autoalloploid genomically unrelated to E. canadensis. The first interpretation is favored. Agropyron dasystachyum, or one of its close relatives, and E. triticoides are suggested as possible parents of A. smithii.     

SYNTHETIC HYBRIDS OF HORDEUM BOGDANII WITH ELYMUS CANADENSIS AND SITANION HYSTRIX

Seven viable hybrid seeds were obtained from 48 hand-emasculated Elymus canadensis L., 2n = 28, florets pollinated by Hordeum bogdanii Wilensky, 2n = 14. The hybrids were large, vigorous, and completely sterile plants that bore a closer morphological resemblance to E. canadensis than to H. bogdanii. Chromosome associations in the 21-chromosome hybrids averaged 9.98^I, 5.40^II, and 0.08^III in 264 metaphase-I cells. Chromosome pairing was attributed to allosyndetic pairing between E. canadensis and H. bogdanii chromosomes. The H. bogdanii genome appears to be partially homologous with one of the two E. canadensis genomes. One Sitanion hystrix (Nutt.) J. G. Smith X H. bogdanii hybrid was obtained from a cross involving 37 emasculated S. hystrix florets. This triploid, 2n = 21, hybrid was morphologically intermediate between the parents and totally sterile. Averages of 9.09^I, 5.72^II, and 0.16^III were observed in 106 cells at metaphase I. A modified form of the H. bogdanii genome appears to occur in S. hystrix as well as in E. canadensis. Many allotetraploid Agropyron, Elymus, and Sitanion species apparently contain a genome derived from Hordeum.     

THE ORIGIN OF AGROPYRON SMITHII

The hypothesis that North American octoploid Agropyron smithii Rydb., 2n = 56, originated by hybridization between tetraploid Agropyron and Elymus species, followed by chromosome doubling, was tested by observing chromosome pairing in hybrids of A. smithii with an induced amphiploid, 2n = 56, derived from E. canadensis L., 2n = 28, X E. dasystachys Trin., 2n = 28, F₁'s. Chromosome pairing in A. smithii averaged 0.52^I, 27.70^II, 0.01^III, and 0.01^IV in 184 metaphase-I cells; and the amphiploid averaged 1.13^I and 27.44^II in 195 cells. Chromosome pairing in A. smithii X amphiploid hybrids averaged 8.20^I, 23.38^II, 0.34^III, and 0.05^IV in 101 metaphase-I cells. It was concluded that A. smithii was genomically similar to the E. canadensis-E. dasystachys amphiploid. The basic genome formula of the amphiploid is SSHHJJXX, with the SSHH genomes coming from E. canadensis and the JJXX genomes coming from E. dasystachys. Consideration of the morphological, ecological, and reproductive characteristics of all North American species that contain the basic SSHH and JJXX genomes led to the conclusion that A. dasystachyum (Hook.) Scribn., SSHH, and E. triticoides Buckl., JJXX, are the probable progenitors of A. smithii.     

GENOME RELATIONS OF AGROPYRON SERICEUM,HORDEUM JUBATUM AND THEIR HYBRIDS

Cytogenetic investigation of microsporogenesis in Agropyron sericeum, Hordeum jubatum, their spontaneous hybrid, Agrohordeum pilosilemma, its amphiploid, and the backcross of the amphiploid to A. sericeum, B₁, elucidated the genome relationships of A. sericeum and H. jubatum. The tetraploid parental species share a partially homologous genome which affects the pairing relationships evidenced in their hybrids. The genome formulae assigned to these plants are: A. sericeum, A“A”BB; H. jubatum, AAA'A‘; Agrohordeum pilosilemma, AA'A“B; the amphiploid, AAA'A‘A”A“BB; and B₁, AA'A”A“BB. Observed pairing configurations were compatible with the expected maximum pairing configurations predicted under the assumption of genetic control of pairing with dosage effects. This is interpreted as further support for the hypothesis that pairing in the hybrids of H. jubatum is controlled by the A genome, one dose of A allowing homeologous pairing and two doses of A promoting homeologous association.     

Cytogenetical studies on artificial hybrids amongElymus sibiricus,E. dahuricus andAgropyron tsukushiense in the tribe Triticeae,Gramineae

To explore the cytogenetical relationships ofElymus andAgropyron of the tribe Triticeae, Gramineae, two species of AsiaticElymus, E. sibiricus (2n=28) andE. dahuricus (2n=42), and a JapaneseAgropyron, A. tsukushiense (2n=42) were crossed. Pentaploid and hexaploid F₁ hybrids were vigorous. All pollen grains were aborted and none of the hybrids produced seed. For the crossE. sibiricus × A. tsukushiense, the average chromosome pairing per cell at the MI of the PMCs in the F₁ was 16.38 univalents, 8.93 bivalents, 0.25 trivalents and 0.01 quadrivalents; for the crossE. dahuricus × A. tsukushiense, it was 4.41 univalents, 17.67 bivalents, 0.32 trivalents, 0.28 quadrivalents and 0.04 quinquevalents; and for the crossE. dahuricus × E. sibiricus, it was 17.11 univalents, 8.74 bivalents, 0.04 trivalents and 0.07 quadrivalents. From the present results, it is concluded thatE. sibiricus contains one genome andE. dahuricus contains two genomes, which are homologous to those ofA. tsukushiense, and that the third genome ofE. dahuricus might be partially homologous to the remaining genome ofA. tsukushiense. This conclusion is also supported by the cytogenetical analysis ofE. dahuricus × E. sibiricus. Contribution No. 27 from the Plant Germ-plasm Institute, Faculty of Agriculture, Kyoto University, Kyoto, Japan.     

SYNTHETIC HYBRIDS OF NEW WORLD AND OLD WORLD AGROPYRONS: III. AGROPYRON REPENS × TETRAPLOID AGROPYRON SPICATUM

Three hybrids of A. repens, 2n = 42, × A. spicatum, 2n = 28, and two reciprocal hybrids were obtained from emasculated and unemasculated crosses, respectively. The 35-chromosome hybrids tended to be morphologically intermediate between the parent species but resembled A. repens more closely than A. spicatum. A. repens behaved cytologically as a segmental autoallohexaploid, and A. spicatum acted cytologically as an autotetraploid. Mean chromosome associations of 8.04 I, 12.72 II, 0.41 III, 0.06 IV, and 0.009 V were observed in 116 hybrid cells at metaphase I. Most chromosome pairing in the hybrids was attributed to autosyndesis. A. spicatum, A. repens, and their hybrids were represented by genome formulas of SSSS, R₁R₁X₁X₁X₂X₂, and SSR₁X₁X₂, respectively. Hybrid fertility ranged from 0.02 to 0.69 seeds per spikelet.     

Inflorescence culture of wheat-Agropyron hybrids: Callus induction,plant regeneration,and potential in overcoming sterility barriers

Segments of young inflorescences of Triticum aestivum cv. Chinese Spring (CS), its F₁ hybrids with Agropyron trachycaulum and A. scirpeum and backcross derivatives with A. yezoense, A. intermedium and A. junceum, and of a A. yezoense x T. aestivum cv. Wichita hybrid were cultured. Different parts of young spikelets of A. trachycaulum x CS F₁ and A. yezoense x Wichita F₁ 's were also cultured. Percent callus induction was lower in wheat than in the wheat-Agropyron hybrids or backcross derivatives. Percent callus induction from different organs in both hybrids was in the descending order of whole spikelet, spikelet without glumes, rachis, and glumes. No plants could be regenerated from calli of wheat and backcross derivatives except those of CS x A. intermedium combination. Callus induction in hybrids varied from 54 to 84% and plant regeneration from 14 to 31%. The regenerants required no vernalization. Variants including one with top-dense spikes and another with elongated spikelets were recovered. Out of eight A. trachycaulm x CS hybrid regenerants, one had anthers and stigma as opposed to neutral flowers of the original hybrid.     

HYBRIDIZATION AND INTROGRESSION IN THE GRASS GENUS ELYMUS

Brown , W. V., and G. A. Pratt . (U. Texas, Austin.) Hybridization and introgression in the grass genus Elymus. Amer. Jour. Bot. 47 (8) : 669–676. Illus. 1960.—The production of artificial F₁ hybrids that produce some seed between E. virginicus and E. canadensis, E. virginicus and E. interruptus, and E. canadensis and E. interruptus proves that similar plants found in nature are true hybrids. The artificial hybrids constituted the basis of comparison in an analysis of natural local populations in the Austin region. Analysis by scatter diagrams indicates that there is introgression of genes of some species into the genomes of others through the development of hybrid swarms. In the area studied the great variability of virginicus is due to the introgression of canadensis and, possibly, interruptus genes. Such hybridization and subsequent introgression have occurred many times and in many places and are still taking place. It is likely that the named varieties of canadensis and virginicus may be introgressant types that are the results of hybridizations between those species and others that occur sympatrically in Eastern United States.     

SYNTHETIC AGROPYRON-ELYMUS HYBRIDS. I. ELYMUS CANADENSIS × AGROPYRON SUBSECUNDUM

Twenty-four seeds were formed in 34 hand-emasculated E. canadensis florets exposed to A. subsecundum pollen. Fifteen of the 24 seeds germinated, and 12 plants were raised to maturity. Each of the 12 plants proved to be a hybrid. The hybrids were morphologically intermediate between the parents, although they favored A. subsecundum in general appearance. The parent plants were meiotically regular and behaved cytologically as strict allotetraploids, 2n = 28. Fourteen bivalents were formed in 12.6% of the hybrid cells interpreted at metaphase I. All other hybrid cells contained various combinations of univalents, bivalents, and multivalents. Mean chromosome associations of 0.22 univalents, 11.70 bivalents, 0.11 trivalente, 0.23 quadrivalents, 0.01 pentavalents, and 0.55 hexavalents were observed in 135 hybrid cells. All hybrid pollen examined was shriveled and non-staining. Five hybrids produced eight seeds, and seven hybrids were completely sterile. The ready cross-compatibility of E. canadensis and A. subsecundum and the relatively good chromosome pairing in their hybrids suggest a much closer relationship between the parent species than is implied by the prevailing taxonomic treatment. Structural rearrangements appear to be responsible for the relatively few differences between E. canadensis and A. subsecundum genomes and for the sterility of the hybrids. Cytological data from this and other investigations indicate a close relationship between many self-fertilizing Agropyron, Elymus, and Sitanion species. It is postulated that this composite of self-fertilizing Agropyron, Elymus, and Sitanion species originated by hybridization between A. spicatum and an unidentified Hordeum species.     

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.     

Comparison of the Giemsa C-banded and N-banded karyotypes of twoElymus species,E. dentatus andE. glaucescens (Poaceae: Triticeae)

The karyotypes ofElymus dentatus from Kashmir andE. glaucescens from Tierra del Fuego, both carrying genomesS andH, were investigated by C- and N-banding. Both taxa had 2n = 4x = 28. The karyotype ofE. dentatus was symmetrical with large chromosomes. It had 18 metacentric, four submetacentric and six satellited chromosomes. The karyotype ofE. glaucescens resembled that ofE. dentatus, but a satellited chromosome pair was replaced by a morphologically similar, non-satellited pair. The C-banding patterns of both species had from one to five conspicuous and a few inconspicuous bands per chromosome. N-banding differentiated the chromosomes of the constituent genomes by producing bands in theH genome only. TheS genomes of both species were similar with five metacentric and two satellited chromosomes having most conspicuous C-bands at telomeric and distal positions. They resembled theS genome of the genusPseudoroegneria. TheH genomes had four similar metacentric and two submetacentric chromosomes. The seventhH genome chromosome ofE. dentatus was satellited, that ofE. glaucescens nonsatellited, but otherwise morphologically similar. The C-bands were distributed at no preferential positions. TheH genome ofE. dentatus resembles theH genomes of some diploidHordeum taxa.     

HYBRIDS AND INDUCED AMPHIPLOIDS OF ELYMUS CANADENSIS X AGROPYRON LIBANOTICUM

North American Elymus canadensis L., 2n = 28, and Asian Agropyron libanoticum Hack., 2n = 14, crossed with ease and yielded vigorous but sterile F₁ hybrids, 2n = 21. Chromosome pairing in the hybrids averaged 9.47^I, 5.38^II, and 0.26^III in 150 metaphase-I cells. One genome of E. canadensis is more or less homologous with the A. libanoticum genome. Treatment of the F₁ hybrids with colchicine produced 42-chromosome amphiploids, C₀, which were advanced through two seed generations, C₁ and C₂. More than half of the metaphase-I cells in the C₀ amphiploids contained 21^II; and average associations were 1.09^I, 20.16^II, 0.07^III, and 0.09^IV in 116 cells. Meiosis became increasingly irregular beyond metaphase-I; nevertheless, the C₀ amphiploids produced 68% stainable pollen and averaged 0.75 seed per spikelet. Multivalent frequencies increased in advanced generations, and the C₂ amphiploids averaged 1.11^I, 19.00^II, 0.23^III, and 0.55^IV in 100 metaphase-I cells. Meiosis was essentially regular in the C₁ and C₂ amphiploids beyond metaphase I, and the C₂ amphiploids averaged 73% stainable pollen and 2.28 seeds per spikelet. The amphiploids have an excellent chance of developing into a meiotically stable, fertile, new species. Forage characteristics of the amphiploids indicate that they have considerable economic potential as a forage grass.     

SYNTHETIC HYBRIDS OF AGROPYRON ALBICANS X A. DASYSTACHYUM,SITANION HYSTRIX,AND ELYMUS CANADENSIS

Emasculated crosses of Agropyron albicans Scribn. & Smith with A. dasystachyum (Hook.) Scribn., Sitanion hystrix (Nutt.) J. G. Smith, and Elymus canadensis L. yielded 34, 5, and 9 viable hybrid seeds from 66, 45, and 52 florets, respectively. The hybrids were for the most part morphologically intermediate between their respective parents. The parents behaved cytologically as allotetraploids, 2n = 28; but meiosis in A. albicans was somewhat more irregular than in the other three species. Chromosome pairing was good in all hybrids and indicated that the genomes of the parent species were closely homologous, but only the A. albicans × A. dasystachyum hybrids set seed. Although closely related, A. albicans and A. dasystachyum are not fully conspecific. Agropyron albicans was considered to be a subspecies of A. dasystachyum, as were A. riparium Scribn. & Smith and A. griffithsii Scribn. & Smith ex Piper.     

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.     

垂穗披碱草与达乌里披碱草基因组细胞遗传学比较

采用顺序FISH-GISH技术,12个重复序列探针,包括9个三核苷酸简单重复序列、2个卫星DNA重复序列pSc119.2和pAs1以及5S rDNA,通过重复序列的物理定位对达乌里披碱草和垂穗披碱草基因组中部分重复序列的分布特征进行了比较分析,为进一步研究垂穗披碱草和达乌里披碱草的物种形成及演化提供新的分子细胞遗传学证据。结果表明:(1)所有的序列在这2个物种的染色体上都能产生可检测的杂交信号,且在2个物种中(AAC)_(10)、(ACT)_(10)、(CAT)_(10)都表现为共分布,(AAG)_(10)与(AGG)_(10)表现为近似共分布;2个物种的H基因组除5S rDNA序列外,其他序列都产生强烈且丰富的杂交位点,St与Y基因组不同重复序列探针的荧光位点数目有所差别,表现为5S rDNA、pSc119.2、(AAC)_(10)、(CAT)_(10)、(ACT)_(10)、(CAC)_(10)探针的信号位点较少或无信号,其余的探针信号位点稍多。(2)达乌里披碱草的第2对染色体上具有(AAC)_(10)、(CAT)_(10)、(ACT)_(10)的杂交位点、第6对染色体上具有(CAC)_(10)的杂交位点,而在垂穗披碱草的St基因组中未观察到上述序列杂交位点;达乌里披碱草St基因组仅有第4对染色体的端部具有pSc119.2杂交位点,而在垂穗披碱草St基因组中的pSc119.2杂交位点位于第5对染色体长臂的间隔区;相对于达乌里披碱草,垂穗披碱草St和Y基因组染色体含有更多的重复序列杂交位点。(3)达乌里披碱草的H/Y基因组间易位在不同材料间是稳定存在的,达乌里披碱草基因组相对稳定,不同材料间H基因组重复序列杂交信号多态性高于St和Y基因组;垂穗披碱草基因组的变异较大,不同材料间St和Y基因组重复序列杂交信号多态性高于H基因组。研究认为,垂穗披碱草和达乌里披碱草的H基因组均起源于布顿大麦,St基因组可能起源于不同的拟鹅观草属物种;与达乌里披碱草相比垂穗披碱草St与Y基因组可能具有更高的染色体结构变异性,而垂穗披碱草St与Y基因组变异较大的原因可能是与同区域分布的含StY基因组的物种发生了种间渗透杂交。     

Biosystematic study ofRoegneria tenuispica, R. ciliaris andR. pendulina (Poaceae:Triticeae)

Morphological and cytological studies of three tetraploidRoegneria species,R. tenuispica, R. pendulina andR. ciliaris, and their artificial hybrids were carried out.Roegneria tenuispica was morphologically similar toR. pendulina. The general appearance of the interspecific hybrids was intermediate between the parents. The hybrids showed comparatively high chromosome pairing at meiosis, but were completely or almost completely sterile. The results indicate that the three independent species share two basic genomes (S_tY) and thatR. tenuispica is more closely related toR. pendulina than toR. ciliaris. The genomes ofR. tenuispica could be designated as S _t ^t Y^t.     

Cytogenetics of a Hordeum vulgare x Elymus patagonicus hybrid (n=4x=28)

Summary Hordeum vulgare L. (2n=2x=14) was hybridized with Elymus patagonicus Speg. (2n=6x=42). The hybrid had 28 chromosomes, genomically represented as HSH₁H₂, and was perennial with a codominant phenotype. The chromosomes were meiotically associated as 19.6 univalents + 0.004 ring bivalents + 2.6 rod bivalents + 0.8 trivalents + 0.14 quadrivalents in 1,129 meiocytes, with a chiasma frequency of 4.77 per cell. The bivalent pairing presumably is an autosyndetic but modified expression of the H₁H₂ genomes of E. patagonicus, since ring bivalents were rare. This does not preclude the association of the H. vulgare H genome chromosomes with either H₁ and/or H₂ genomes of E. patagonicus to form bivalent or multivalent associations. A further evaluation of the genome homologies of H. vulgare, H. bogdanii, E. canadensis and E. patagonicus is proposed.     

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.