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Tetrasomic Inheritance and Isozyme Variation in Turnera Ulmifolia Vars

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Tetrasomic Inheritance and Isozyme Variation in Turnera Ulmifolia Vars 305—312 Received 18 June 1990 Heredity 66 (1991) - Genetical Society of Great Britain Tetrasomic inheritance and isozyme variation in Turnera ulmifolia vars. elegans Urb. and intermedia Urb. (Turneraceae) JOEL S. SHORE Department of Biology, York University, 4700 Keele St, North York, Ontario, Canada, M3J 1P3 Tetrasomic inheritance of three isozyme loci is demonstrated in Turnera ulmifolia vars. elegans and intermedia. The data support the occurrence of an autopolyploid origin for tetraploids of both taxonomic varieties. The extent of isozyme variation at 14 loci was determined for diploid and tetraploid populations. Tetraploid Turnera ulmifolia var. intermedia showed the lowest levels of isozyme variation, perhaps a result of founder effect, upon island colonization. In contrast, Turnera ulmifolia var. elegans showed the greatest levels of isozyme variation, suggesting that hybridization among locally differentiated diploid or tetraploid populations might have occurred. The number of independent origins of autotetraploidy in Turnera ulmifolia is unknown. Keywords:isozymevariation, tetrasomic inheritance. ulmifolia var. elegans occur in Brazil. A tetraploid Introduction population that shares characteristics of both varieties Turneraulmifolia is a species complex of perennial occurs in Columbia (Barrett, 1978; Shore & Barrett, weeds native to the Neotropics. The complex contains 198 5a), but insufficient numbers of plants were avail- diploid, tetraploid, hexaploid and octoploid cytotypes able for study. and has a base number of x =5 (Raman & Kesavan, Three lines of evidence suggest that tetraploid 1964; Barrett, 1978; Arbo & Fernandez, 1983; Shore populations of T ulmifolia vars. elegans and intermedia & Barrett, 1985a; Barrett & Shore, 1987; Fernandez, are autopolyploids. Cytological observations of tetra- 1987; Fernandez & Arbo, 1989). Diploid and tetra- ploids of both varieties have revealed the occurrence of ploid populations are distylous and individuals are quadrivalents at diakinesis and the first metaphase of strongly self-incompatible and obligately outcross, meiosis, as well as reduced pollen fertility (Raman & while hexaploids of three taxonomic varieties are Kesavan, 1964; Arbo & Fernandez, 1983; Fernandez, homostylous and self-compatible (Barrett, 1978; Shore 1987; Shore, 1991). Finally, frequencies of meiotic & Barrett, 1985b; Barrett & Shore, 1987). configurations fit the autotetraploid pairing models of Turnera ulmifolia vars. elegans and intermedia are Jackson & Hauber (1982) for four of six populations highly interfertile members of the species complex investigated (Shore, 1991). (Shore & Barrett, 1985a; Arbo & Fernandez, 1987; In this paper, the hypothesis that T ulmifolia vars. Shore, 1991) and diploid hybrids show regular bivalent elegans and intermedia have had autotetraploid origins formation at meiosis (Fernandez & Arbo, 1989). The is tested and the genetic consequences of autotetra- varieties are most easily distinguished by the presence ploidy is investigated. Genetic evidence is provided for of a purple petal spot in the cream-coloured corollas of autopolyploidy in these taxonomic varieties by demon- T ulmifolia var. elegans while var. intermedia has strating tetrasomic inheritance at three isozyme loci. To yellow flowers that lack the petal spot. Diploid popula- investigate the genetic consequences of autotetra- tions of T ulmifolia var. intermedia occur throughout ploidy, levels of isozyme variation in diploid and tetra- Central America and in portions of South America, ploid populations of T ulmifolia were compared. while tetraploid populations are apparently restricted Finally, the evidence bearing on the evolutionary origin to Puerto Rico and Hispaniola (Barrett & Shore, of the tetraploid varieties was examined. 1987). Native populations of diploid and tetraploid T 305 306 J. S. SHORE Materials and methods Dominican Republic. Note that population 131 has recently been found to be intersterile with other diploid Bulkseed samples were collected from populations populations and may represent an undescribed species found throughout much of the native range of T. of Turnera (Arbo & Fernandez, 1987; M. M. Arbo, ulmifolia vars. intermedia and elegans. Seeds were personal communication). Nevertheless, it is included germinated and population samples maintained in a in the analyses for comparisons of amounts of genetic glasshouse following the methods of Shore & Barrett variation between diploids and tetraploids (below). (1985a). The varietal status, sample size and ploidal Diploid population Ii is considered to be T. ulmifolia level of populations sampled is provided in Table 1. var. intermedia, although, it is polymorphic for the Seven populations of tetraploid T ulmifolia var. presence of a purple petal spot characteristic of T. elegans were sampled from Brazil. For T ulmifolia var. ulmifolia var. elegans (Shore & Barrett, 1987; Barrett, intermedia, seven diploid populations were sampled 1978). from Central and South America and 16 tetraploid Horizonal starch gel electrophoresis was used to populations were sampled from Puerto Rico and the obtain allozyme data for 10 enzyme systems. Aconitase (Aco), esterase (Est), glutamate dehydrogenase (Gd/i), glutamate oxaloacetate transaminase (Got), glucose- Table I Varietal status, population code, locality, ploidal level and sample size (n) of Turnera ulmifolia populations phosphate isomerase (Gpi), leucine aminopeptidase used (Lap), 6-phosphogluconate dehydrogenase (Pgd) and phosphoglucomutase (Pgrn), were assayed following Ploidal Shore & Barrett (1987). Malate dehydrogenase (Md/i) Population Locality level n and alcohol dehydroganse (Ad/i) were assayed follow- ing Cardy et al. (1980). Flower buds were used for the Ii Barreirinhas, Brazil 2x 18 electrophoretic assays as they provided high levels of 12 Calabozo, Venezuela 2x 12 enzyme activity. For each enzyme, the locus coding the 126 Santa Rosa, Costa Rica 2x 28 128 Santa Rosa, Costa Rica 2x 8 most anodally migrating protein was designated as 129 La Pacifica, Costa Rica 2x 26 locus 1, the next locus 2, etc., except where information 131 Juremal, Brazil 2x 40 was available on the intra-cellular location of the 135 El Salvador 2x 35 isozyme coded by the locus (Shore & Barrett, 1987). 18 Rio Piedras, P.R. 4x 14 Isozymes localized in the plastids are denoted by the 110 Parbueyon,P.R. 4x 40 letter 'p' and cytosolic isozymes by 'c'. Allozymes Ill Joyuna,P.R. 4x 26 coded by different alleles were designated by lower 112 Dorado, P.R. 4x 13 case letters, with the allele coding for the most anodally 113 Manati,P.R. 4x 11 migrating allozyme designated 'a', the next 'b', etc. For 114 Hwy2at863,P.R. 4x 28 ease of presentation, some alleles were re-coded for 115 Hwy2at67l,P.R. 4x 6 studies of inheritance (see below). 116 Tortuguera,P.R. 4x 20 The inheritance of three isozyme loci, Adh-2 for T 117 Jarabacoa,D.R. 4x 15 ulmifolia var. intermedia and Aco-2 and Gpi-c for var. 118 Santo Domingo, D.R. 1 4x 29 119 Cabrete, D.R. 4x 10 elegans was determined for tetraploids using controlled 120 San FCO de Macoris, D.R. 4x 28 crosses of parents of known electrophoretic phenotype 122 Lucas Diaz, D.R. 4x 25 and assaying the resulting progeny. For Adh-2 (a 123 BaniatPlaya,D.R. 4x 12 dimeric enzyme), a plant putatively of tetrasomic 124 St Cristobal, D.R. 4x 21 genotype ffss was crossed to four individuals homo- 125 Santo Domingo, D.R. 2 4x 27 zygous (genotype ffff) at this locus. A reciprocal cross E2 Manaus, Brazil 1 4x 23 was done for one of the individuals thus providing a E3 Belém, Brazil 4x 25 total of five families. A total of 96 progeny was scored E6 Manaus, Brazil 1 4x 21 for Adh-2 activity by assaying mature pollen in which E7 Recife, Brazil 4x 21 the locus is expressed. E8 Aracaju, Brazil 4x 28 E9 Maceio, Brazil 4x 24 For T ulmifolia var. elegans, a heterozygous plant ElO Vitória de S. Antão, Brazil 4x 18 that was putatively tetra-allelic at the Aco-2 locus was obtained. The genotype of the plant was abcd, where I =varietyintermedia, each letter refers to one of the four alleles producing E =varietyelegans, electrophoretically distinguishable monomeric allo- P.R. =PuertoRico, zymes at this locus. The plant was crossed reciprocally D.R. =DominicanRepublic, x =5. to an individual of homozygous genotype, bbbb, and AUTOTETRAPLOIDYIN TURNERA 307 85 progeny were scored. The inheritance of Gpi-c (the the diversity analysis of Nei (1973). Gene diversity cytosolic isozyme locus of the dimeric enzyme, among populations relative to total diversity was calcu- glucosephosphate isomerase) was also assessed in T lated as GST =DST/HT.In addition, genetic distances ulmifolia var. elegans. An individual with a putative tn- were calculated among all pairs of populations allelic genotype was crossed with an individual that was followed by a UPGMA cluster analysis (Nei, 1987). di-allelic as follows: finms x mmms where fm,and s, are three alleles at this locus. A total of 83 progeny was Results scored. For all three loci, goodness of fit and heterogeneity tests were done to both tetrasomic (expected for auto- Inheritance of isozymes tetraploids) and two-locus disomic (expected for alto- To test the hypothesis that tetraploid T. ulmifolia var. tetraploids) models of inheritance, using the G statistic elegans and intermedia are autopolyploids, the inheri- (Sokal & Rohlf, 1981). Tetrasomic ratios were tance of three isozyme loci was investigated. For Adh-2 predicted assuming no double reduction occurred at in T ulmifolia var. intermedia, segregation
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