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Chromosomal and Morphological Studies of Diploid and Polyploid Cytotypes of Stevia Rebaudiana (Bertoni) Bertoni (Eupatorieae, Asteraceae)

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Chromosomal and Morphological Studies of Diploid and Polyploid Cytotypes of Stevia Rebaudiana (Bertoni) Bertoni (Eupatorieae, Asteraceae) Genetics and Molecular Biology, 27, 2, 215-222 (2004) Copyright by the Brazilian Society of Genetics. Printed in Brazil www.sbg.org.br Research Article Chromosomal and morphological studies of diploid and polyploid cytotypes of Stevia rebaudiana (Bertoni) Bertoni (Eupatorieae, Asteraceae) Vanessa M. de Oliveira1, Eliana R. Forni-Martins1, Pedro M. Magalhães2 and Marcos N. Alves2 1Universidade Estadual de Campinas, Instituto de Biologia, Departamento de Botânica, Campinas, SP, Brazil. 2Universidade Estadual de Campinas, Centro Pluridisciplinar de Pesquisas Químicas, Biológicas e Agrícolas, Campinas, SP, Brazil. Abstract In this study, we examined the chromosome number and some morphological features of strains of Stevia rebaudiana. The chromosomes were analyzed during mitosis and diakinesis, and the tetrad normality and pollen viability were also assessed. In addition, stomata and pollen were measured and some plant features were studied morphometrically. All of the strains had 2n = 22, except for two, which had 2n = 33 and 2n = 44. Pairing at diakinesis wasn=11II for all of the diploid strains, whereas the triploid and tetraploid strains hadn=11III andn=11IV, respectively. Triploid and tetraploid plants had a lower tetrad normality rate than the diploids. All of the strains had inviable pollen. Thus, the higher the ploidy number, the greater the size of the pollen and the stomata, and the lower their number per unit area. The triploid strain produced the shortest plants and the lowest number of inflorescences, whereas the tetraploid strain had the largest leaves. Analysis of variance revealed highly significant differences among the strains, with a positive correlation between the level of ploidy and all of the morphological features examined. Key words: Asteraceae, pollen, polyploidy, Stevia rebaudiana, stevioside, stomata. Received: May 7, 2003; Accepted: November 27, 2003. Introduction species has not yet been extensively studied agronomically, The genus Stevia Cav. (Eupatorieae, Asteraceae), considerable genetic variability has been observed in plant which consists of approximately 150-200 species of herba- size, flowering period, and stevioside content, which may ceous, shrub and sub-shrub plants (Gentry, 1996), is one of vary from 2% to 10% (Magalhães, 2000). the most distinctive genera within the tribe Eupatorieae, Several of the known strains of S. rebaudiana re- mainly because of the morphological uniformity of its sulted from an experiment done at the University of New flowers and capitula, which consist of five tubular flowers Mexico designed to develop polyploids that were richer in and five involucral bracts (King and Robinson, 1987). Al- stevioside (Valois, 1992). In Brazil, plants from this experi- though geographically widespread, this genus occurs ex- ment were initially cultivated at the CENARGEN/EM- clusively in tropical and subtropical regions of the United BRAPA germplasm bank before part of the collection was States and Central and South America (Robinson and King, transferred to CPQBA/UNICAMP. The induction of poly- 1977). Randi (1980) reviewed the potential uses of Stevia ploidy to improve agronomic yields is a process commonly rebaudiana, which produces stevioside, a non-caloric used in plants of economic interest (Allard, 1960) and has sweetener that does not ferment in the human body. been applied to other species such as Nicandra physaloides Thirteen distinct strains of S. rebaudiana are cur- (Gupta and Roy, 1986), coffee (Cruz et al., 1993), Clitoria rently being domesticated and assessed agronomically by ternatea (Ghandi and Patil, 1997) and orange (Romero- the Centro Pluridisciplinar de Pesquisas Químicas, Bioló- Aranda et al., 1997). gicas e Agricolas (CPQBA/UNICAMP). Although this A better adaptability of individuals and increased or- gan and cell sizes are usually associated with polyploidy (Guerra, 1988). In Andropogon gerardii, a species with Send correspondence to E. R. Forni-Martins. Universidade Esta- dual de Campinas Instituto de Biologia, Departamento de Botânica, hexaploid (2n = 6x = 60) and enneaploid (2n = 9x = 90) Caixa Postal 6109, 13083-970 Campinas, SP, Brazil. E-mail: cytotypes, differences in plant size have been observed, and [email protected]. although seed production per unit area is the same in both 216 Oliveira et al. cytotypes, it is more efficient in hexaploids (Keeler and Da- In this study, the original designations of Valois vis, 1999). In some species of coffee with different num- (1992) for the plants resulting from each treatment have bers of chromosomes, higher levels of ploidy generally been maintained: the letter T followed by a number refers to produce larger stomatal cells with a greater number of the concentration of colchicine used and the letter V fol- chloroplasts (Boaventura et al., 1981). lowed by a number identifies each vase (Table 1). Voucher In Gramineae such as Paspalum, diploid and tetra- specimens of each strain cultivated in the field were depos- ploid races within a species are basically distinguished by ited in the herbarium at UEC (Universidade Estadual de their reproductive system, which is sexual in diploids and Campinas) (Table 1). apomictic in polyploids (Burson, 1975). Crossbreeding be- Mitotic and meiotic studies tween individuals with different numbers of chromosomes usually leads to sterile progeny. According to Allard For the mitotic studies, root tips were obtained from (1960), in various agricultural plants with diploid and tetra- plants cultivated in vitro or from dark achenes germinated ploid individuals, the progeny are partially or completely in gerbox with gibberellic acid (50 ppm), at room tempera- sterile triploids that do not bear seeds. ture. Light achenes are sterile (Felippe et al., 1971; Mon- The main objective of this study was to determine the teiro, 1980). The root tips were pretreated with chromosome numbers of the strains of S. rebaudiana culti- paradichlorobenzene (PDB) for4hat16°C-18 °C and vated at CPQBA/UNICAMP in order to obtain information fixed in ethanol-acetic acid (3:1, v/v). Cytological prepara- that could be useful in breeding programs. The tetrad nor- tions were obtained using the Giemsa technique (Guerra, mality and pollen viability were used to estimate the regu- 1983). For meiotic analysis, floral buds were fixed in etha- larity of meiosis. Some morphological features of each nol-acetic acid (3:1, v/v) for approximately 24 h and the an- strain were also analyzed morphometrically in order to thers then squashed onto slides in acetic carmine (Medina asses the relationship with the level of ploidy. and Conagin, 1964). The cytological preparations were examined by light microscopy. All cells in which it was possible to determine Material and Methods the chromosome number, the meiotic behavior and mor- The plant material (floral buds, achenes and roots) for phological features, were included in the analysis. Chromo- the cytogenetic studies was collected from strains of S. some numbers were based on counts from an average of 10 rebaudiana cultivated in vivo or in vitro at CPQBA/UNI- cells per strain. The karyotypes were determined from a CAMP. These strains were obtained mainly from CENAR- single cell of the T0 (control) treatment. Chromosome mea- GEN/EMBRAPA, in Brasília (Table 1), and originally surements in other strains were also obtained using a cam- derived from experiments by Valois (1992), who exposed era lucida. seeds of S. rebaudiana to nine treatments by soaking them Tetrad analysis and pollen viability for 18 h in a solution of colchicine (an antimitotic agent), the concentration of which ranged from 0.5% (treatment Tetrad analysis and pollen viability were assessed as T1) to 0.001% (treatment T9). A control treatment with no described by Medina and Conagin (1964). The technique of colchicine (T0) was also done. After the treatment, all of Alexander (1980) was also used to estimate pollen viability the seeds were kept under conditions appropriate for germi- in order to allow comparison between the results obtained nation. with both techniques (Dafni, 1992). Table 1 - Chromosome number (2n), pairing at diakinesis, tetrad normality and pollen viability (%) in strains of Stevia rebaudiana (Bertoni) Bertoni. Access Origin 2n Pairing at diakinesis Tetrad normality (%) Pollen variability (%) Brasília F1 CENARGEN/EMBRAPA 22 11II 98 0 Maringá F1 Maringá-PR 22 11II 99 0 Canada F1 Canada 22 11II 96 0 T0 (control) Maringá-PR 22 11II 95 0 T5V1 CENARGEN/EMBRAPA 22 11II 96 0 T6V1 CENARGEN/EMBRAPA 22 11II 99 0 T6V3 CENARGEN/EMBRAPA 44 11IV 80 0 T7V1 CENARGEN/EMBRAPA 33 11III 64.5 0 T8V1 CENARGEN/EMBRAPA 22 11II 93 0 T8V2 CENARGEN/EMBRAPA 22 11II 97.5 0 T9V1 CENARGEN/EMBRAPA 22 11II 98 0 Chromosomes and morphology of S. rebaudiana cytotypes 217 The total number of normal and abnormal tetrads per Diakinesis was the only phase analyzed in meiosis. slide (approximately 1,000 tetrads from five floral buds) Pairing wasn=11II for all strains of S. rebaudiana, except was used to estimate tetrad normality. Pollen viability was for two, which hadn=11III andn=11IV, that corre- assessed for each strain by counting the number of viable sponded to 2n = 33 and 2n = 44, respectively (Table 1, and non-viable grains in 200 pollen grains per bud collected Figure 2). The tetrad normality rate varied from 64.5% to from five floral buds of different inflorescences. Each slide 99% (Table 1). Abnormal tetrads generally had three but was prepared with the anthers of a single bud. never four nuclei. The pollen of all strains
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