NATURAL AND ARTIFICIAL HYBRIDS OF HELIANTHUS MAXIMILIANI X H. GROSSESERRATUS

Long , Robert W. (Ohio Wesleyan U., Delaware.) Natural and artificial hybrids of Helianthus Maximiliani × H. grosseserratus. Amer. Jour. Bot. 46(10): 687–692. Illus. 1959.—An investigation of the occurrence of natural hybridization in two perennial sunflowers, Helianthus Maximiliani and H. grosseserratus, was begun in 1950. Subsequently, artificial F₁, F₂, and first and second backcross generations were produced. Fertility and vigor were high in all these plants, but F₁ plants appeared to excel the others in these characteristics. Observations in the experimental garden were supplemented by examination of chromosomes in pollen mother cells, comparisons of herbarium collections, and study of wild populations. Evidence pointed to close genetic relationship of the species and to the occurrence of natural hybridization in areas of distributional overlap. In 1957 and 1958, field work in these areas resulted in the scoring of 18 natural populations, 3 of which consisted of both parental species plus putative F₁ hybrids. Two explanations are offered to account for the seeming absence of introgression. The results support the conclusion that natural hybridization leads to the establishment of F₁ hybrids and that introgression does not occur to any significant extent. Although both species display a high degree of interfertility, they are distinct morphologically. For this reason, it is advisable to maintain them as separate species.     

ADAPTIVE RADIATION IN THE HAWAIIAN SILVERSWORD ALLIANCE (COMPOSITAE-MADIINAE). II. CYTOGENETICS OF ARTIFICIAL AND NATURAL HYBRIDS

The Hawaiian silversword alliance of Argyroxiphium, Dubautia, and Wilkesia, in spite of exhibiting spectacular morphological, ecological, physiological, and chromosomal diversity, is remarkably cohesive, genetically. This is attested to by the ease of production of artificial hybrids and by the high frequency of spontaneous hybridization among such life forms as mat-forming subshrub, monocarpic rosette shrub, polycarpic shrub, cushion plant, tree, and vine. Even the least fertile of these hybrids is capable of producing backcross progeny. Moreover, first generation interspecific and intergeneric hybrids have been successfully used to produce trispecific hybrids in a number of instances. In general, the widest hybrid combinations have been as readily produced as crosses within a species. At present eight genomes or chromosome races distinguished by reciprocal translocations are recognized on the basis of meiotic analysis of artificial and spontaneous hybrids. Seven of these races are found among those species with 14 pairs of chromosomes. The eighth genome very likely characterizes all nine species of this alliance that have 13 pairs of chromosomes. The cytogenetic data indicate that redundancy of translocations involving the same chromosomes has been a recurrent theme in the chromosomal differentiation of these taxa. There appears to be little, if any, correlation between chromosomal evolution and adaptive radiation as assessed by gross habital differentiation in this group. However, within Dubautia, a novel ecophysiological trait associated with colonization of xeric habitats is restricted to species with n = 13. In contrast to the bulk of the Hawaiian flora, which is characterized by self-compatibility and chromosomal stability, it is suggested that the occurrence of self-incompatibility in the Hawaiian Madiinae may have favored selection of supergenes via chromosomal repatterning, and this may account for the diversity of chromosome structure seen in this group.     

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.     

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.     

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.     

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.     

SYSTEMATICS OF THE TRIBE ORCUTTIEAE (GRAMINEAE) AND THE DESCRIPTION OF A NEW SEGREGATE GENUS,TUCTORIA

The two genera Neostapfia and Orcuttia, recognized as being closely allied and sharing unusual features, are treated as the sole members of the tribe Orcuttieae. Neostapfia is monotypic; Orcuttia comprises eight taxa, six of which are universally treated as species, and two as varieties. Studies reported here are based on field work throughout the ranges of these two genera and examination of 550 herbarium specimens. Chromosome counts and seed protein analyses for all taxa are presented. These data indicate that each taxon is worthy of specific status and that Orcuttia, as traditionally treated, consists of discordant elements. It is concluded that a more satisfactory taxonomy results when this group is split into two genera. A new genus, Tuctoria, is described, and the combinations T. fragilis (Swallen) J. Reeder, T. greenei (Vasey) J. Reeder, and T. mucronata (Crampton) J. Reeder are proposed.     

MULTIPOLAR MEIOSIS IN DIPLOID CRESTED WHEATGRASS,AGROPYRON CRISTATUM

A new model of spindle organizers is proposed: The spindle organizer in a higher plant is similar to the centriole of animal cells. It is a unit cell organelle which follows regular cell division cycles and is genome specific. Each genome carries its own spindle organizer. During fertilization, a male spindle organizer enters the egg cell. It may fuse with the female spindle organizer, or either one may degenerate. In a hybrid, both male and female spindle organizers may exist, and multipolar divisions separate different genomes into different groups. The same mechanism can be used to explain the formation of a polyhaploid from a polyploid. The chromosome behavior of an individual is believed to be an interaction of chromosome homology and the homology between chromosomes and their spindle organizers. This model is based on observations of multipolar meiosis which occurred in two cultures of diploid crested wheatgrass, Agropyron cristatum (L.) Gaertn. In these two cultures multipolar meiosis occurred at every stage after late diakinesis. The seven bivalents were separated into groups at late diakinesis. More than one equatorial plate was formed at metaphase I. Each micrometaphase plate behaved as an independent unit and had its own anaphase movement. Usually the chromosome complement separated into two groups with (4–3), (5–2), and (6–1) separations observed in about an equal number of cells. Cells with chromosomes divided into three or four groups were found less frequently. Multipolar meiosis may take place at either first or second division. Cell plates were formed across each spindle apparatus, cleaving each group of chromosomes into smaller micro-cells. At the “quartet” stage, 4- to 12-celled “quartets” were observed. Pollen stainability was measured at above 75% in both cultures. Stained pollen grains could be classified into two distinct size classes. Darkly stained, small pollen grains represented the result of multipolar meiosis and may have been viable. Multipolar cell divisions provided a mechanism which polyploids might reduce their ploidy level.     

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.     

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.     

异育银鲫及其人工杂合种外源遗传物质的检测分析

采用 DNA 杂交的方法,对异育银鲫及其人工杂合种进行了外源 DNA 的检测分析,发现两个雌核发育系的异育银鲫及其人工杂合种的 DNA 与红鲤 DNA 片段间的分子杂交均有阳性斑点出现,这表明,在异育银鲫及其人工杂合种产生过程中,父本(红鲤)的 DNA 片段的确可以随机地掺入到母本细胞的 DNA 中,从而产生了异精效应,使其子代生长加快,并出现父本性状。     

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.     

须芒草族植物花粉形态的观察

在光学显微镜下和扫描电镜下对禾本科须芒草族（Ａｎｄｒｏｐｏｇｏｎｅａｅ）中分隶于８个亚族３４个属的３６种植物的花粉进行形态面容和比较研究。结果显示,本族植物花粉形态较为一致,花粉近形或扁球形,单萌发孔,孔,周围加厚,具盖,外壁表面散布有颗粒。这表明其是一个自然类群。总体来讲,芬烨大的演化分异,只是表面纹饰的和芬烨大小有一;定的差异。纹饰可分为三种类型粗糙型,不明显疠状突起才明显疠状突起型。花粉开矿     

AN ANATOMIC-SYSTEMATIC STUDY OF THE CLASSICAL TRIBE FESTUCEAE (GRAMINEAE)

Using the classical tribe Festuceae as a frame of reference, 135 grass genera have been examined in an effort to determine natural generic relationships which often have been obscured by parallel and convergent evolution. Studies of embryo and leaf structure, ligules, lodicules, and lemmas have greatly enlarged, verified, and nearly completed existing evidence that the classical concept of the Festuceae is highly artificial and that the tribe contains many genera which are only remotely related. A more nearly phylogenetic system is achieved when more than two-thirds of these genera are distributed among 4 major groups, which may be worthy of subfamily rank: 55 genera are retained in the festucoid group, 46 are moved to the eragrostoid, 2 to the bambusoid, 13 to the arundoid, and 8 to a recently designated group labelled the centothecoid. Three genera remain unplaced; and material was unavailable or insufficient to make accurate determinations for 8 genera.