SYNTHETIC AGROPYRON-ELYMUS HYBRIDS: II. ELYMUS CANADENSIS X AGROPYRON DASYSTACHYUM

Pollination of 40 emasculated E. canadensis florets with A. dasystachyum pollen gave rise to 12 shriveled seeds, one of which was viable. The hybrid seedling developed into a vigorous plant whose vegetative and spike characteristics were for the most part intermediate between those of the parents. Both parents behaved cytologically as allotetraploids, 2n = 28, and formed 14 bivalents regularly at metaphase I. Chromosome associations in the hybrid, 2n = 28, averaged 0.42 I, 13.70 II, 0.02 III, and 0.03 IV in 142 metaphase-I cells interpreted. The hybrid produced 165 seeds under open-pollination during its second year in the field. E. canadensis and A. dasystachyum apparently contain closely homologous genomes whose major structural differences consist of several inverted segments and possibly a small reciprocal translocation. The close homology between the E. canadensis and A. dasystachyum genomes was unexpected in view of the wide morphological, ecological, and reproductive differences that separate the two species.     

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.     

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 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.     

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.     

MORPHOLOGY AND CYTOLOGY OF SYNTHETIC HYBRIDS OF AGROPYRON TRICHOPHORUM X AGROPYRON CRISTATUM

Dewey, Douglas R. (Utah State U., Logan.) Morphology and (cytology of synthetic hybrids of Agropyron trichophorum X Agropyron cristatum. Amer. Jour. Bot. 50(10): 1028–1034. Illus 1963.—Three hybrids were obtained from controlled crosses of pubescent wheatgrass, A. trichophorum (2n = 42), and hexaploid crested wheatgrass, A. cristatum (211 = 42). The hybrids were intermediate between the parent plants for all vegetative and spike characteristics observed. Under open pollination, 2 of the hybrids set 2 seeds each, and the other hybrid produced 60 seeds. Meiosis in the parent plants was basically regular. Average motaphase-I chromosome associations were 0.09 I, 20.56 II, 0.05 III, and 0.16 IV per cell in the A. trichophorum parent, which was described as a segmental autoallohexaploid. The hexaploid A. cristatum parent averaged 0.18 I, 7.44 II, 0.81 III, 2.86 IV, 0.08 V, and 2.11 VI per cell at diakinesis and was described as an autohexaploid. Chromosome pairing in the hexaploid hybrid averaged 5.08 I, 8.94 II, 4.33 III, 1.11 IV, 0.27 V, and 0.05 VI per cell. On the basis of chromosome pairing in the parent species and their hybrids, it was concluded that 1 of the A. trichophorum genomes was partially homologous with the 3 genomes of hexaploid A. cristatum. Genome formulae for hexaploid A. cristatum, A. trichophorum, and their hybrids were represented as AAAAAA, A₁A₁B₁B₁B₂B₂, and AAAA₁B₁B₂ respectively.     

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.     

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.     

SYNTHETIC HYBRIDS OF NEW WORLD AND OLD WORLD AGROPYRONS. IV. TETRAPLOID AGROPYRON SPICATUM F. INERME X TETRAPLOID AGROPYRON DESERTORUM

Emasculated and unemasculated crosses of tetraploid A. spicatum f. inerme X A. desertorum yielded four hybrids. The hybrids were morphologically intermediate between the parent species but resembled A. desertorum more closely than A. spicatum. Both parents behaved cytologically as autoploids. Mean chromosome associations of 0.04 I, 8.60 II, 0.01 III, and 2.67 IV were observed at diakinesis in the 28-chromosome A. spicatum. The A. desertorum parent contained 30 chromosomes, 2 of which were likely supernumeraries, and averaged 0.03 I, 9.85 II, and 2.57 IV at diakinesis. Three hybrids contained 30 chromosomes, and one had 29. The most common chromosome association in the 30-chromosome hybrids was 2 I and 14 II; and the average was 3.00 I, 13.40 II, 0.06 III, and 0.01 IV. A. spicatum and A. desertorum chromosomes were usually distinguishable from each other in the hybrid cell on the basis of size. All pairing in the hybrids was attributed to autosyndesis within parental genomes. A. spicatum, A. desertorum, and their hybrids were represented by genome formulas of SSSS, CCCC, and SSCC, respectively. The hybrids produced 5 to 439 seeds under open pollination. Three controlled crosses between the hybrids yielded 2, 5, and 23 seeds, respectively, on 10 maternal spikes in each cross. The prospects of developing a fertile, cytologically stable allotetraploid species from the hybrids appear favorable.     

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.     

分布于日本和中国的鹅观草及其杂种的形态学和细胞学研究

本文对分布于日本的Agropyron tsukushiense (Honda) Ohwi var. transiense (Hack.) Ohwi (2n=6x=42)和分布于中国的Roegneria kamoji Ohwi (2n=6x=42)及其杂种F_1(2n=6x=42)进行了形态学及细胞学的研究,并同时探讨了亲本种的亲缘关系。总体来看,亲本材料之间在形态上虽有差异但并不十分显著。杂种F_1的形态特征介于其父、母本之间。在减数分裂过程中,亲本种和杂种F_1的染色体配对行为均十分正常。但在检查了大量的成熟花粉和穗状花序之后,发现杂种F_1有部份不育现象。上述研究结果表明A. tsukushiense var. transiens的三个染色体组与R. kamoji的三个染色体组同源。结合形态学和育性等方面的研究资料,作者认为上述两个材料仍应属于同一分类等级。但必须指出,由于长期的地理隔离,他们之间产生了一定的形态变异和生殖障碍。按照国际植物命名法规(ICBN)上述两个材料应组合为:Roegneria tsukushiensis (Honda) B. R. Lu, Yen et J. L. Yang及其变种var. transiens (Hack.) B. R. Lu, Yen et J. L. Yang comb. nov.