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Studies on Citrus Pollination Using Gamma-Irradiated Pollen

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Studies on Citrus Pollination Using Gamma-Irradiated Pollen s. AfT. J. Bot. , 1988, 54(3): 257-264 257 Studies on Citrus pollination using gamma-irradiated pollen J.H. de Lange* and A.P. Vincent Citrus and Subtropical Fruit Research Institute, Nelspruit, 1200 Republic of South Africa 'Present address: National Botanic Gardens, Private Bag 7, Claremont, 7735 Republic of South Africa Accepted 10 December 1987 Studies have been performed in Citrus motivated by Pandeys' theory of induced egg transformation and parthenogenetic development of embryos by pollination with irradiated pollen. Poncirus trifoliata (a Citrus relative) and shaddock pollen still germinated following 1,0 kGy irradiation and that of Palestine sweet lime following 2,0 kGy irradiation. The pollination study involved cross-pollination of Clementine mandarin using P. trifoliata or Palestine sweet lime pollen; gamma-irradiated at different doses, with or without a second pollination of the same flowers using normal Valencia sweet orange pollen. Using 0,5 or 1,0 kGy-irradiated P. trifoliata pollen, cross-pollination of Clementine, known for its limited parthenocarpic ability, resulted in the production of large numbers of seedless fruits. It was demonstrated that extensive embryo abortion took place after cross-pollination using irradiated pollen but that fertilization and/or early ovule development stimulated set and growth of fruitlets. Ovule occupation by irradiated pollen largely prevented fertilization by normal pollen. Pollination, using 0,5 or 1,0 kGy-irradiated pollen followed one day later by a second normal pollination is suggested for further studies on possible transformation of hybrids by the first pollen parent. Seed production after pollinating with pollen irradiated at high doses in a single pollination, is very low. Embryo culture to save embryos before abortion and to eliminate the second pollination, should receive attention. Studies, gemotiveer deur Pandey se teorie van geYnduseerde eiseltransformasie en partenogenetiese embrio-ontwikke­ ling deur bestuiwing met bestraalde stuifmeel, is by Citrus uitgevoer. Stuifmeel van Poncirus trifoliata Cn Citrus-verwante spesie) en pompelmoes ontkiem nog na 1,0 kGy-bestraling en die van Palestynse soetlemmetjie na 2,0 kGy-bestraling. In die bestuiwingstudie is Clementine-nartjie kruisbestuif met P. trifoliata- of Palestynse soetlemmetjie- stuifmeel; gamma­ bestraal met verskillende dosisse, met of sonder 'n tweede bestuiwing van dieselfde blomme met normale Valencia­ soetlemoenstuifmeel. Kruisbestuiwing met 0,5 of 1,0 kGy-bestraalde P. trifoliata-stuifmeel, het die produksie van baie groot getalle saadlose vrugte by Clementine, wat bekend is vir sy beperkte partenokarpiese vermoe, tot gevolg gehad. Daar is aangetoon dat kruisbestuiwing met bestraalde stuifmeel tot grootskaalse embrio-aborsie gelei het maar dat be­ vrugting en/of vroee saadknop-ontwikkeling die set en groei van vruggies gestimuleer het. Saadknop-besetting deur bestraalde stuifmeel het bevrugting deur normale stuifmeel grootliks verhoed. Bestuiwing met 0,5 of 1,0 kGy-bestraalde stuifmeel wat een dag later gevolg word deur 'n tweede normale bestuiwing, word voorgestel vir verdere stUdies met die oog op moontlike transformasie van kruisings deur die eerste stuifmeel-ouer. Enkel-bestuiwing met stuifmeel wat bestraal is teen hoe dosisse, het baie lae getalle saad tot gevolg gehad. Embrio-kulture met die oog op voorkoming van embrio­ aborsie en uitskakeling van die tweede bestuiwing, behoort aandag te ontvang. Keywords: Gamma-irradiation, parthenocarpy, pollination, seed abortion, transformation *To whom correspondence should be addressed Introduction resistance in Citrus relatives or certain Citrus species to Pandey (1975, 1981) proposed the use of irradiated pollen particular commercial cultivars would be a valuable tool in in cross-pollination to transfer only a few traits of the pollen breeding. parent to a seedling. According to him ionizing radiation of The present study was performed to investigate the pos­ tobacco pollen was employed as a 'pulverizing' agent and sible application of the approach in Citrus. not as a mutagen. This resulted in the incorporation of a few chromatin fragments in the egg genome during the Materials and Methods pseudo-fertilization by occasional association of the frag­ In vitro germination of gamma-irradiated pollen ments with their homologues among the egg chromatin. Pollen of shaddock [c. maxima (L.) Osbeck], Palestine The hybrid is identical to the female parent except for a few sweet lime [c. aurandfolia (Christm.) Swing.] and a Citrus genes from the pollen parent. relative in the subfamily Aurantioideae, Poncirus trifoliata Using this approach, promising results have also been (L.) Raf. was irradiated using a60Co source applying var­ obtained in barley (Powel et al. 1983); maize (Pandey ious doses. Pollen at a density of approximately 50 grains 1983) ; wheat (Snape et al. 1983) ; tomato (Zamir 1983) ; per mm2 was incubated on a medium containing 20% suc­ peas (Davies 1984) and chillies (Daskalov 1984) . rose and 0,6% Difco Bacto agar at pH 7 and 26°C. The An obstacle in the application of this technique is the incubation period was 24 h and all treatments were replica­ production of small numbers of seed (Pandey 1981). The in ted three times. In the case of Palestine sweet lime, pollen vitro culture of embryos before abortion may serve to over­ tube length was also recorded. come this problem. Transformed seed may also be produced and in larger Cross-pollination of Clementine mandarin (C. reticulata numbers by a normal fertilization by unirradiated pollen Blanco) preceded by the pollination using irradiated pollen. In such Flowers of Clementine, a mono-embryonic cultivar, were cases gene contribution in the hybrid is predominantly cross-pollinated using P. trifoliata or Palestine sweet lime from the two parents involved in the normal fertilization pollen gamma-irradiated at different doses. In additional with a minor contribution from the 'irradiated pollen' treatments these pollinations were followed by a second parent (Pandey 1981). cross-pollination on the same flowers 1 or 3 days following These findings have important implications in Citrus the first pollination, using untreated pollen of Valencia breeding. Hybrids between commercial citrus cultivars and sweet orange [c. sinensis (L.) Osbeck]. In three additional related species such as Poncirus trifoliata and Microcitrus treatments Clementine flowers were cross-pollinated using australasica produce inedible fruits . The transfer of only only untreated Valencia pollen at (a) anthesis (b) 1 day and certain favourable traits such as cold tolerance or disease (c) 3 days following anthesis. 258 S.-Afr. Tydskr. Plantk., 1988,54(3) The treatments were not replicated for practical reasons. On the seventh month following pollination (at ma­ Each of the 33 treatments shown in Table 1 constituted turity) the fruits were measured finally, fruit set deter­ flowers on a single tree. In most treatments 100 flowers mined in each treatment and number of seeds in individual were used. In some treatments somewhat smaller numbers fruits determined. of flowers were involved depending on the availability of At 3 months following pollination a few fruits resulting flowers within 1 day prior to anthesis. The total number of from cross-pollination with P. trifoliata pollen irradiated at flowers treated was 2 609. 1,0 kGy were cut and the seeds observed macroscopically. Each of the treated flowers was identified by tying a short cotton string loosely around the flower peduncle and Observations on seedlings resulting from cross-pollination labelling the branch concerned. Records were taken of A total of 17 104 seeds were planted. Records were taken each fruit's cross-diameter and of fruit set in each treatment on germination percentages of seed and on leaf shape of on the second, third, fourth, fifth and seventh month fol­ the Clementine (x P. trifoliata) hybrids. lowing pollination. Tracing back the growth history of each individual fruit Results and Discussion was made possible by the labelling. This allowed data on In vitro germination of gamma-irradiated pollen the growth of fruit having the same seed content at ma­ The results are shown in Figure 1. Pollen of Palestine sweet turity, to be grouped together and averaged as described lime was still found to exhibit considerable germination at before (de Lange & Vincent 1972). the relatively high dose of 1,0 kGy. Even at 1,5 and 2,0 kGy it showed germination percentages of 4,7 and 1,3 re­ spectively compared with the 19,0% germination of normal pollen. The average lengths of Palestine sweet lime pollen Table 1 Pollination treatments applied to Clementine flowers tubes at 1,0; 1,5 and 2,0 kGy were 432 J.tm, 360 J.tm and 248 J.tm respectively compared with 576 J.tm in the control. Pollen parent in cross-pollination at anthesis In one season shaddock pollen irradiated at 1,0 kGy ex­ hibited 42,0% germination compared with 79,3% in the P. trifoliata Palestine sweet lime control (Figure 1B) while there was no germination at 1,0 kGy kGy kGy in another season. Germination of P. trifoliata pollen irradiated at 1,0 kGy was 2,0% compared to 47% in the o 0,05 0,2 0,5 1,0 0,05 0,2 0,5 1,0 o control. Control 2 345 6 7 8 9 10 Van den Boom & den Nijs (1983) demonstrated germi­ Valencia 11 12 13 14 15 16 17 18 19 20 nation of Cucumis pollen at even higher doses and in the pollen 1 day case of C. metuliferus considerable germination and pollen following first tube growth at 4,0 kGy. pollination Valencia 21 22 23 24 25 26 27 28 29 30 Cross-pollination of Clementine mandarin pollination Seed abortion 3 days following Macroscopic observation on a few fruitlets from flowers first pollinated by normal and 1,0 kGy-irradiated pollen at 2 pollination months following anthesis, showed extensive abortion of 31 : Valencia pollen at anthesis ovules in the latter treatment (Figure 2). At maturity large 32 : Valencia pollen 1 day following anthesis numbers of small abortive seeds were found in fruits from 33 : Valencia pollen 3 days following anthesis flowers pollinated with 0,5 or 1,0 kGy-irradiated pollen.
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