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ARTIFICIAL HYBRIDIZATION BETWEEN DIFFERENT POLYPLOID LEVELS in GLANDULARIA (VERBENACEAE)L

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ARTIFICIAL HYBRIDIZATION BETWEEN DIFFERENT POLYPLOID LEVELS in GLANDULARIA (VERBENACEAE)L Amer. J. Bot. 5.5(10): 123.5-1239. 1968. ARTIFICIAL HYBRIDIZATION BETWEEN DIFFERENT POLYPLOID LEVELS IN GLANDULARIA (VERBENACEAE)l OTTO T. SOLBRIG Department of Botany, University of Michigan, Ann Arbor and CORNELIUS PASSANI AND ROGER GLASS Gray Herbarium, Harvard University, Cambridge, Massachusetts ABST RAe T The hexaploid North American species Glandularia eleqans has been crossed with the South American diploid species G. etellaroides, G. peruviana, and G. pulchella. Analysis of the behavior of chromosomes at meiosis in the hybrid indicates that the chromosomes of G. pulchella are similar to one of the genomes of G. eleqane. Non-homologous pairing and presence of multivalents also indicates that G. eleqans is a segmental allohexaploid derived most likely from South American ancestors. THE GENUS Glandularia GmeLin is one of a the sterility of the hybrids varying between 5 large number of plant taxa with a North-South and 90%, approximately. Crosses made by Solbrig American disjunct distribution. In South America (unpub1.) have shown that all N.A. hexaploid it grows in Argentina, Uruguay, southern Brazil, species tested (5) are also interfertile and that in Paraguay, Bolivia, Chile, and Peru. In North general hybrids are highly fertile (80 % or more America it is found in northern Guatemala, pollen fertility). Mexico, and in the United States from South The present work reports the results of artificial Carolina to FLorida. west to California and north crosses between three South American diploid to Utah, and in Colorado, South Dakota, ILlinois, species and one North American hexaploid species. and Ohio. Morphologioally most species are simi­ The object of the study was to gain some infor­ lar, the major differences between taxa being leaf mation regarding the degree of affinity between shape, and floral and fruit characteristics. the species of the two continents and to investi­ The ~orth American species of the genus have gate the genomic affinity of the species involved been monographed by Perry (1933) who recog­ in the crosses. nized 18 species. No co:nplete monographic treatment exists for the South American species; MATE:UAL AND METHODs-The diploid South Moldenke (1951-64), and Troncoso (1954, 19(5) American species, Glawiuiaria pulcliella Sweet, have clarified the classification of particular G. steilaroides Covas and Schnack, and two groups of species; Schnack and Covas (1944) distinct morphological types of G. peruuiana (L.) have discussed the position of the genus and its Small, were grown in greenhouses at Cambridge, validity. It is estimated that there are approxi­ Mass., and artificial crosses attempted with the mately 30-50 species in South America. Experi­ North American hexaploid, G. elegans H.B.K. mental work by Dermen (1935), Schnack and All flowers were emasculated and the styles polli­ coworkers (Schnack and Covas, 194.5a, b; Schnack nated repeatedly for about 7 days. Most attempted and Gonzales, 1945; Schnack and Solbrig, 1953), crosses were unsuccessful, but a few of the approxi­ Solbrig (1959, 1952, 1958, and unpub1.), Lewis mately 100 attempts produced fruits. These in­ and Oliver (1961), and others has shown that the volved always G. eleqans as the pistillate parent, South American species are largely diploid with and they yielded only a very few seeds. The seeds n = 5 (only 2 tetraploids have been found out were stratified for about 3 months in moist sand of 20 species investigated), while the North and 15 plants obtained (Table 1). Meiotic prepara­ American species are mostly hexaploid with tions were made according to the squash tech­ n = L"i (only 4 tetraploid species have been nique, using aceto-carmine as a stain. All observa­ determined out of 15 species investigated). tions were made under a phase-contrast micro­ Furthermore, extensive hybridization programs scope. Pollen grains were stained with lacto­ undertaken by Schnack and coworkers and by phenol cotton-blue, and 500 grains counted to ob­ Solbrig have shown that all South American tain fertility estimates. species tested (10) are interfertile to some extent, Mature hybrid plants were cloned and some of the cloned plants treated with colchicine in an 2 I Received for publication 31 May 1968. aqueous solution at a concentration of 5 X 10- 1235 1236 AMERICAN JOURNAL OF BOTANY [Vol. 55 TABLE 1. Characteristics of the species and hybrids a elegans X peruviana elegans X Character peruviana Tetraploid Octoploid elegans pulchella pulchella Plant height 12.5 10 15 22 12 7.5 Habit Decumbent- Sprawling Sprawling Erect Sprawling Decumbent semi-erect Internode length 3.0 2.3 4.7 3.9 2.7 3.4 Leaf length 3.2 2.7 4.5 4.2 3.5 1.7 Leaf width 1..5 2.0 2.9 2.9 2.2 1.7 Leaf pubescence Short, medium long, medium long, long, long, long, medium dense medium dense dense medium dense open infrequent Leaf shape Entire Lobed Lobed Bipinnatifid Bipinnatifid Bipinnatifid Floral bract length 5mm 4mm 5mm 5mm 4mm 4mm Calyx length 11 mm 9mm 9.5mm 6mm 7mm 8mm Corolla tube 16mm 14mm 17mm 14mm 12 mm 12mm Corolla limb width 12mm 12mm 14.5 mm 10.5 mm lOmm lOmm Corolla color> Red(19) Red-purple(29) Red-purple(29) Purple(32) Purple(33) Purple(3i5) Style length 14.5 mm 13 mm 16mm 11 mm 7.5mm 8mm Anther gland absent 0.1 mm 0.25 mm 0.3mm 0.5mm 0.9mm Pollen stainability 100% 5% 70% 100% 20% 100% • All measurements are in cm unless stated otherwise. Measurements here given represent averages of several plants, b Color code number taken from the color dictionary of the Royal Horticultural Society of England. M. The colchicine solution was applied in a while six had no pairs at all. The other 29 cells cotton wad to the apical buds for a period of had from one to six pairs (Table 2). Only one cell 72 hr, while the plants were in an atmosphere had no multivalents; most cells had two (18 cells) of high humidity and at elevated temperatures or more quadrivalent associations. Of all the (ca. 30 C). chromosomes analyzed (800), 39.9 % were asso­ ciated in pairs, 59.6 % were in multivalent RESULTs-Morphology of the hybrids-The F 1 associations, and 2.5 % were not paired at all hybrids were intermediate between the parents in (Table 3). both foliar and floral characters including flower color. On the other hand they were less vigorous Glandularia elegans X G. peruviana: artificial than the parents (particularly the G. elegans octoploid-Two octoploid plants, the result of X G. stellaroides cross), and the internodes treating the corresponding hybrid with colchicine, were somewhat shorter (Table 1). The crosses of were obtained. They showed mostly 20 pairs at G. elegans with G. pulchella and G. peruviana meiotic metaphase and at diakinesis with regular bloomed normally and continuously through the separation to the poles in first anaphase, and year; the cross involving G. stellaroides on the normal development of the subsequent stages of other hand bloomed very seldom, producing only meiosis, including tetrad formation and pollen a few flowers rather than the usual showy race­ development. However, occasional univalents and mose inflorescence characteristic of the genus. multivalents (mostly trivalents and quadrivalents) were observed. Seventy-two percent of the pollen Meiotic analysis-The parental species all stained with lacto-phenol cotton-blue and was showed normal pairing and chromosome segre­ normal looking morphologically, although 24 % gation at meiosis. The plant of G. stellaroides used of the pollen grains were tetracolpate instead of had 2 supernumerary chromosomes that did not tricolpate, as is universally found in the parental align on the metaphase plate and did not divide species. but instead were usually incorporated in one of the microspores. Glandularia elegans X G. pulchella: tetraploid hybrid-Nine plants were obtained, all having Glandularia elegans X G. peruviana: tetraploid similar meiotic behavior. In this cross 42 cells hybrid-Five plants were obtained in this cross. could be analyzed exactly. A significantly larger All showed similar meiotic behavior. The most number of bivalents could be observed than in the conspicuous characteristic was the large number G. elegens X G. peruviana cross. Seventeen of of multivalents, particularly quadrivalents. Sev­ the 42 cells showed more than six pairs; no cell eral hundred cells were studied; of these 40 allowed had less than three bivalents. Most cells had at an exact and unequivocal identification of all least one quadrivalent association, although none the associations. Of the 40 cells, only five had had more than three, and a large number (10) more than six pairs of chromosomes per cell, had no multivalent associations at all. More November-December,1968] SOLBRIG ET AL.-HYBRIDIZATION IN GLANDULARIA 1237 TABLE 2. Number of cells with pairs and tetravalent assoc- hexaploid species with 15 chromosomes and a iations diploid species with 5 chromosomes, the predicted associations are a maximum of 5 pairs and 10 elegans X elegans X univalents, to a minimum association of 20 uni­ peruviana pulchella valents. Any pairing in the hybrid is interpreted Cells with: (4 X) (4 X) as indicating homology between the chromosome segments of the parental species. If multivalent OIl 6 0 III 2 0 pairing is found in the hybrid, it is assumed that 2Il 6 0 chromosomal rearrangements, specifically trans­ 3II 6 4 locations, have taken place in one or both of the 4Il 9 5 species, so that the homologous segments are 5Il 6 9 now located in more than one chromosome. 6II 10 7 Nevertheless, the general hypothesis cannot ex­ 7JI 1 3 plain the presence of more than five bivalents in 8II 3 11 the hybrid, nor the existence of multivalent asso­ 9Il 1 0 ciations involving more than five chromosomes.
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