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The Genetics of Distyly and Homostyly in Turnera Ulmifolia L

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The Genetics of Distyly and Homostyly in Turnera Ulmifolia L Heredity 55 (1985) 167—174 The Genetical Society of Great Britain Received 10 January 1985 The genetics of distyly and homostyly in Turnera ulmifolia L. (Turneraceae) Joel S. Shore and Department of Botany, University of Toronto, Toronto, Spencer C. H. Barrett Ontario, Canada M5S 1A1. Turnera ulmifolia L. is a polymorphic complex of diploid, tetraploid and hexaploid varieties. Diploids and tetraploids are distylous and hexaploids homostylous. A controlled crossing programme demonstrated that in diploids distyly is controlled by a single locus with two alleles. Long-styled plants are ss and short-styled plants can be either Ss or SS. In tetraploids a similar pattern occurs with short-styled p!.ants usually Ssss and long-styled plants ssss. Tetrasomic inheritance was demonstrated by crosses with a SSss genotype synthesised by coichicine doubling. Double reduction could not be detected at the distyly locus. No significant deviation from a 1: 1 morph ratio was observed in surveys of nine natural populations and progeny tests of 21 open-pollinated familes from Brazil. Crosses between distylous and homostylous populations were undertaken to determine the inheritance of homostyly. The results were consistent with a model of supergene control. Clear segregation of phenotypes was observed in the F1, and the dominance relationships S> h> s, where h is an allele that determines homostyly, were obtained. INTRODUCTION Turnera ulmfolia L. is a polymorphic assem- blage of perennial weeds native throughout the Distylyoccurs in at least 23 flowering plant families Neotropics. Urban (1883) deseribed 12 varieties (Ganders, 1979) and has undoubtedly evolved within the complex, six of which are distylous and several times. The inheritance of distyly has been the remainder homostylous. Barrett (1978) determined for species in 11 genera from nine described the floral biology and breeding systems families (Ornduff, 1979). The syndrome of floral of four varieties within the complex, and a recent characters, by which the long- and short-styled survey of chromosome numbers in 43 populations morphs differ, is inherited as if controlled by a of T u1mfo1ia (Shore and Barrett, 1985; and single gene locus with two alleles. Except for two unpublished data) revealed a correlation between genera where the dominance relationships are breeding system and ploidal level. All distylous reversed (Baker, 1966; Ornduff, 1979), the long- populations are diploid or tetraploid, whereas styled morph is homozygous (ss)whilethe short- homostylous populations are hexaploid. Here we styled morph is heterozygous (Ss). Evidence from investigate the genetics of breeding system vari- the Primulaceae (Ernst 1955), and the Plum- ation in the T ulm(folia complex. Specifically we: baginaceae (Baker, 1966) suggests that distyly is (1) determine the inheritance of distyly in diploids controlled by a series of tightly linked genes com- and tetraploids; (2) provide data on morph ratios prising a supergene. Aberrant forms termed and segregation from natural populations; (3) homostyles, with anthers and stigmas at the same determine the inheritance ofhomostylyand discuss level within a flower, are interpreted as recom- evidence for supergene control of distyly. binant phenotypes that arise by crossing over within the supergene. A recombination event resulting in homostyly has, however, never been MATERIALSAND METHODS detected in heterostylous species (Charlesworth and Charlesworth, 1979a). In addition, apart from a Acrossing programme was undertaken on glass- few notable exceptions (Crosby, 1949; Ernst, 1955; house grown plants raised from seed collections Ganders, 1975), homostyles are rarely observed in or cuttings obtained from natural populations. distylous populations, although relatives of Table 1 gives the locality, chromosome number heterostylous taxa are frequently homostylous. and varietal status of accessions used in the study. 168 J. S. SHORE AND S. C. BARRETT Table IVarietal status, ploidal level, and locality of Turneraulmtfolia accessionsused in experimental studies (X =5) Ploidal Population Variety level Locality Collection No. 11 intermedia 2X Barreirinhas, Brazil Barrett 1128 13 intermedia 2X Caracas, Venezuela Barrett 1125 16 intermedia 2X Managua. Nicaragua Barrett 1342 124 intermedia 4X St. (ristohal, Dominican Rep. Barrett and Shore 1364 132 intermedia 2X Arco Verde, Brazil Barrett and Shore 1374 ES elegans 4X Crato, Brazil Barrett 222 Eli elegans 4X Salgueiro, Brazil Barrett and Shore 1372 AS angustifolia 6X Ochos Rios, Jamaica Barrett 1253 All angusti fo I ia 6X Dragons Ray, Jamaica Barrett 1259 A17 a ngusti Co Ii a 6X Pelican Lake, Grand Bahama Correll 40638 01 orientalis 6X Corrientes, Argentina Arho 1538 Vi velutina 6X Tuxtla Gutierrez, Mexico Koch & 1-ryxell 7834i Each individual was given a code indicating its and a short-styled plant were selected for accession, or family if applicable, followed by a reciprocal crosses. To determine the inheritance of numeric value specifying the particular individual homostyly, three homostylous varieties, from five within the accession, and by a letter indicating its accessions of T. ulmijhlia, were crosed to diploid phenotype (L =long-styled,S =short-styled,H = long- and short-styled plants of known genotype. homostyled).All pollinations were performed in To examine the equilibrium frequency in T. a pollinator-free glasshouse using fine forceps. u1mifilia we extend the morph frequency data of Flowers were emasculated prior to cross-pollina- Barrett (1978) by sampling a total of 1211 plants tion. Pollinated flowers were individually marked from nine natural populations of the tetraploid T. and maturing capsules were wrapped with parafllm ulmifolia var. elegans in N.E. Brazil. Random or to prevent loss of seeds during dehiscence. As complete samples of populations were taken with distylous members of the complex are strongly at least 90 plants sampled per population. Further, self-incompatible two different methods were used progeny tests of 21 open-pollinated families, col- to obtain seeds from self- or intra-morph crosses: lected from population El I in Brazil were per- (1) flower buds were opened one day prior to formed to test for heterogeneity among families anthesis and pollinated from an open flower of the and among maternal morphs. On average 62 pro- same individual (hud-selfing), (2) plants were geny were scored for each family. screened for the presence of a weakened incom- All goodness-of-fit and heterogeneity tests were patibility reaction (pseudo-compatibility) and performed using the G statistic (Sokal and Rohif, those yielding more than three seeds per pollina- 1981). tion were later used in the crossing programme. Progenies obtained from crosses were grown to flowering in individual pots. RESULTS To investigate the inheritance of distyly at diploid and tetraploid levels accessions comprising Neitherof the two methods employed to obtain different varieties were employed (table 1). seed from selfing long- or short-styled plants was Reciprocal crosses were generally performed and completely successful. Bud-selfing in T. ulm(fo!ia the results pooled. Preliminary results suggested results in seed set more often in short- than long- that short-styled morphs were of genotype Ss. To styled individuals; only one of many long-styled detect the occurrence of SS genotypes, 19 short- plants yielded seed. In both morphs the yield of styled progeny derived from the self of 132-1 S were seed from bud-selfing is less than 75 per cent of selfed and at least 17 progeny from each family that from legitimate pollinations. Screening plants were grown to flowering and scored for floral for pseudo-compatibility resulted in the recovery morph. To investigate the possibility of tetrasomic of a few individuals that produced sufficient seed inheritance and double reduction, the chromosome on self-pollination to he useful for genetic study. number of seedlings derived from the cross 132-IS x Two plants were identified among diploids (1311- Il-IOL, was doubled. A drop of 0l per cent col- 3L and 132-iS) and two among tetraploids (ES-IL chicine applied to cotyledons resulted in the and ES-los). Progeny (>80 per cross) derived by recovery of several tetraploid plants and a long- selfing or crossing plants ES-IL and ES-lOS were GENETICS OF HOMOSTYLY 169 screened for self-compatibility. Six progeny Four of 19 short-styled plants, derived by yielded sufficient seed for further study. selfing plant 132-iS, yielded no long-styled pro- geny upon selfing. At least 34 selfed progeny were Inheritance of distyly in diploids scored for each of these four plants. The probabil- (i) ity of observing no long-styled plants among the Theinheritance of distyly in diploid T. ulmifolia progeny of a heterozygous parent for a sample size var. intermedia was determined using three long- of 34 is less than 6 x i05. Hence these individuals and six short-styled plants, from four different are likely to be short-styled plants of genotype SS. accessions, and one long-styled plant (1311-3L), Among the segregating progenies, 105 long-styled from the cross 13-3Sxll-1OL. Bud pollinations and 326 short-styled plants were obtained and no were used for all illegitimate crosses except those deviation from 1: 3 or heterogeneity, was observed involving individuals 1311-3L and 132-iS which (Gdev= 009,P =076; Ghet = 178,P =0.22).The yielded sufficient seed on selfing. The data presen- occurrence of four non-segregating plants as well ted in table 2 are consistent with the common
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