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Polyploid Breeding in Portulaca Grandiflora L Cytologia 44: 167-174, 1979 Polyploid Breeding in Portulaca grandiflora L. A. K. Singh Plant Cytogenetics and Breeding Laboratory, B.S.N.V. Degree College, Lucknow (U.P.), India Received June 13, 1977 Portulaca grandiflora a popular annual ornamental of family Portulacaceae produces beautiful blooms during summer in a wide range of attractive colours. It is commonly known as "9 'O' clock" plant. This species was included in the ornamental breeding programme. This paper deals with the coichicine induced autoploids in pink coloured variety. Materials and methods Seeds of Portulaca grandiflora were obtained from local sources and sown in pots. As heterozygosity at diploid level could be useful in polyploid breeding no effort was made to purify the variety through selfing. Shoot tips of young seedlings were treated with 0.2% aqueous coichicine for 15 hours. Polyploids thus raised were planted in pots with suitable controls and when flowering began, buds of proper size were fixed in 1:3 acetic alcohol fortified with iron. The anthers were squashed in acetocarmine for cytological investigations. Observations Colchicine treatment checked the growth of young seedlings for 2 days. first formed leaves after treatment were thicker, longer and broader than controls. Growth of polyploids was slow and flowering was delayed by 10 days. In tetraploids there was increase in size of leaf, flower, thickness of stem, number of branches, height and spread of plant; size of stomata, anther, gynoecium and pollen grains. Fl owers of polyploids lasted longer and remained open for longer duration. The tetraploids and diploids had 25.6 and 5.2% pollen sterility. Seed setting was poor in tetraploids but seed size was bigger. The chromosome number of the species is 2n=18 (Fig. 2). Meiosis in controls was quite normal except for 2 rare univalents. Cytomixis was observed (Fig. 11). Frequency of ring and rod bivalents is similar in both diploids and tetraploids. Quadrivalent frequency was very low in tetraploids. Fig. 3 shows a PMC of tetraploid having 17 bivalents +2 univalents while Fig. 4 shows I ring quadrivalent with 16 bivalents. Chiasma frequency per chromosome in diploids and tetraploids was 0.71 and 0.68 respectively. A single tetraploid in C1 had ten petals arranged in two whorls. Selfed seeds from this tetraploid were collected and Ca generation was raised. In C1 generation segregation with regard to number of petals was observed. Different plants had 168 A. K. Singh Cytologia 44 Figs. 1-4. 1, in upper row diploid control (2 whorl), 4n single whorl, 4n double whorl, in lower row 4n 3, 4 and multiple whorls respectively. 2, a PMC of diploid showing 9 bivalents. 3, a PMC of 4n showing 17 bivalents +2 univalents. 4, a PMC of 4n showing 1 quadrivalent +16 bivalents. 1979 Polyploid Breeding in Portulaca grandiflora L. 169 Figs. 5-10. 5, androecium and gynoecium of single whorled 4n. 6-10, showing various stages of transformation of stamens and gynoecium in multiple whorled flowers. fl owers with 1, 2, 3, 4 and numrous whorls (•ä). The total number of C2 tetra ploids was 18 and out of these 5 (27.7%) had single whorl, 5 (27.7%) had two whorls, 3 (16.6%) had three whorls, 3 (16.6%) had four whorls and numerous whorls (Fig. 1). Detailed chromosomal studies of each catagory were carried out (Table 1). Chromosomal behaviour was found to be fairly similar in plants of different catagories. In one of the plants of 3 whorled type a binucleate PMC had two metaphase I plates (Fig. 13). In C2, multivalents were absent except in one plant having 2 whorls. In this plant one cell had trivalent (Fig. 12) while another cell had a chain quadrivalent. At anaphase unequal separation and stray chromosomes were noted. While there were no strays in C1 and C2 in single whorl they were present in all the other categories. PMCs showed 4, 5, 6, 7 and 8 spores in 65.5, 18.7, 4.4, 6.2 and 4.4 percent cells respectively. Table 1. Showing different chromonsomal configurations in C1 and C2 generations of autoteraploids of Portulaea grandiflora along with diploid 1979 Polyploid Breeding in Portulaca grandiflora L. 171 Figs. 11-17. 11, cytomixis in PMCs of diploid. 12, a PMC showing 1 rod trivalent +16 bivalents +1 univalent. 13, a binucleate PMC of tetraploid showing two metaphase plates. 14, style and stigma of diploid showing germination of pollen grains. 15, L. S. of the ovary of diploid. 16, L. S. of the ovary of tetraploid (multiple whorls). 17, style and stigma of multiple whorled tetraploid showing germination of pollen grains. Pollen sterility was comparatively higher in numerous whorled plants (Table 1). Seed setting was highest in single whorled tetraploids while with increase in number of whorls, seed setting decreased and in numerous whorled (•ä) plants despite their having 67.86% fertile pollen grains there was absolutely no seed setting. They also had very high transformation of anthers into petaloid structures and in some of the owers even gynoecium was involved (Figs. 6 to 10). Fig. 5 shows the condition fl in normal 4n, single whorled plant. To encourage seed setting in this type artificial 172 A. K. Singh Cytologia 44 pollination with other types was carried out but even this could not induce any seed setting. Study of stigma squashes after artificial pollination revealed normal ger mination of pollen grains (Fig. 17). Fig. 14 shows germination of pollen grains in diploid. Longitudinal sections of overy in 2n and 4n (Figs. 15, 16), revealed quite normal ovules attached to free central placenta and when these ovules were squashed, cells of ovules showed normal stainability of nucleus in the cells. Seeds from C2 generation were collected after selfing of different categories and C3 generation was raised next year. Segregation in selfed progeny of every plant of C2 was again noted in C3 (Table 2). Also from every category segregants of all ve categories were recovered. fi In C3 also multiple whorled tetraploids were completely seed sterile. Selfed seeds from 1 to 4 whorled tetraploids have been collected for further studies. Table 2. Showing segregation in C2 progeny of single 2 whorled C1 tetraploid. Further segregation in C3 from selfed seeds of different categories of C2 plants is also given Discussion Increase in size and number of floral parts has no doubt enhanced the beauty of the tetraploids. Besides this polyploid breeding has been quite successful in other ornamentals like Torenia fournieri (Joshi and Raghuvanshi 1966), Catharanthus roseus and Tabernaemontana divaricata (Raghuvanshi and Chauhan 1969a, b, 1971 and 1972), Arrhostoxylum elegans, Tropaeolum majus and Phlox drummondii (Rag huvanshi and Pathak 1977, 1975 and 1974) and Thunbergia alata (Singh 1977). The most striking feature of tetraploids was very low multivalent frequency in C, In the present material multivalents were completely absent in C2 except in one plant. This confirms the views of Raghuvanshi and Chauhan (1969a) in Catharanthus roseus that complete bivalent formation may take place in subsequent generations in tetraploids having a very low multivalent frequency. However, in Cicer arietinum and Phlox drummondii (Raghuvanshi and Pathak 1973 and 1975) there was increase in the frequency of multivalents in C2 generation. Low mul tivalent frequency was reported by Raghuvanshi and Joshi (1966) in Foeniculum vulgare and Joshi and Raghuvanshi (1967) in Anethum graveolens. In Coriandrum sativum also high bivalent frequency was observed (Joshi and Raghuvanshi 1965). It must be mentioned that the size of chromosome in the present species under in vestigation is quite small and due to lack of availability of enough time for pairing 1979 Polyploid Breeding in Portulaca grandiflora L . 173 the multivalent frequency may be low (Raghuvanshi and Chauhan 1969a). Higher pollen sterility and variation in number of spores per sporad may have resulted from disturbed meiosis in C, which is evident by unequal separation of chromosomes at anaphase I and stray bivalent at metaphase I. In C2 also pollen sterility was maintained as in C, (Table 1). Although the tetraploids of different number of petal whorls showed slight variation in pollen sterility (Table 1) there is no direct correlation. Unlike the autoploids of Catharanthus roseus (Raghuvanshi and Chauhan 1969a), Thunbergia alata (Singh 1977) and Tropaeolum majus (Raghuvanshi and Pathak 1974) tetraploids of Portulaca grandiflora showed quite good seed setting in 1 to 4 whorled types. Only in multiple whorled tetraploids complete lack of seed setting was noted and despite intensive efforts of artificial pollination with other tetraploids, seed setting could not be induced. In multiple whorled diploids also seed setting is quite poor but complete absence of seed setting in tetraploids of this category shows presence of ploidy barrier. A ploidy barrier has also been reported in Tropaeolum majus (Raghuvanshi and Pathak 1974). Though, so far it has not been possible to work out the total number of genes involved but the segregation in C2 and C3 of different categories of flowers clearly indicate that this trait is controlled by multiple gene system and the different grades are the cumulative effect of number of genes present in them. This also shows that the material which was treated for induction of polyploids was heterozygous and due to an increase in the number of homologous chromosomes there was an increase in number of loci governing this trait. This further complicated the segregation patterns. So comparatively larger populations have to be grown before it would be possible to clearly understand the genetic nature of this character. It must be mentioned that the variation with regard to number of whorl in the diploids of same coloured flower plants is present in nature and a wide range of one to multiple row of petals in the same colour has been observed.
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